Cigüeñas, Ibis, Espatulas.pdf

STORKS, IBISES

and SPOONBILLS of the World

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STORKS, IBISES and SPOONBILLS of the World JAMES A. HANCOCK JAMES A. KUSHLAN and M. PHILIP KAHL Illustrated by

ALAN HARRIS and DAVID QUINN

Published with the support of the Brehm Foundation

ACADEMIC PRESS Harcourt Brace Jovanovich, Publishers London * San Diego * New York * Boston * Sydney * Tokyo * Toronto

First published 1992 by Academic Press. Digital editions published 2011 by Christopher Helm Publishers, an imprint of A&C Black Publishers Ltd, 36 Soho Square, London W1D3QY Copyright © 2001 text Academic Press Limited Copyright © 2001 illustrations Alan Harris and David Quinn ISBN (epub) 978-1-4081-3500-6 ISBN (e-pdf) 978-1-4081-3499-3 All rights reserved. No part of this publication may be reproduced or used in any form or by any means - photographic, electronic or mechanical, including photocopying, recording, taping or information storage or retrieval systems — without permission of the publishers.

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Contents INTRODUCTION

1

CLASSIFICATION

5

CONSERVATION

15

C O U R T S H I P AND R E P R O D U C T I O N

20

F E E D I N G B E H A V I O U R AND ECOLOGY

29

SPECIESD E S C R I P T I O N S American Wood Stork (Mycteria americana)

35

Milky Stork (Mycteria cinered)

43

Yellowbilled Stork (Mycteria ibis]

49

Painted Stork (Mycteria leucocephala]

53

Asian Openbill Stork (Anastomus oscitans)

59

African Openbill Stork (Anastomus lamelligerus)

63

Black Stork (Ciconia nigra)

69

Abdim's Stork (Ciconia abdimii)

75

Woollynecked Stork (Ciconia episcopus)

81

Storm's Stork (Ciconia stormi)

87

Maguari Stork (Ciconia maguari)

91

White Stork (Ciconia ciconia)

97

Oriental White Stork (Ciconia boyciana)

103

Blacknecked Stork (Ephippiorhynchus asiaticus)

109

Saddlebill Stork (Ephippiorhynchus senegalensis)

115

Jabiru Stork (Jabiru mycteria)

119

Lesser Adjutant Stork (Leptoptilos javanicus)

125

Greater Adjutant Stork (Leptoptilos dubius)

129

Marabou Stork (Leptoptilos crumenifems)

133

Shoebill (Balaeniceps rex)

139

American White Ibis (Eudocimus ruber)

147

Barefaced Ibis (Phimosus infuscatus)

157

Glossy Ibis (Plegadis falcinellus)

163

Whitefaced Ibis (Plegadis chihi)

169

Puna Ibis (Plegadis ridgwayi)

173

Sharptailed Ibis (Cercibis oxycerca)

111

Plumbeous Ibis (Harpiprion caerulescens)

179

Contents

Buffnecked Ibis (Theristicus caudatus)

183

Green Ibis (Mesembrinibis cayennensis)

189

Hadada Ibis (Bostrychia hagedasti)

193

Wattled Ibis (Bostrychia carunculatd)

197

Olive Ibis (Bostrychia olivacea)

199

Spotbreasted Ibis (Bostrychia rard)

203

Madagascar Crested Ibis (Lophotibis cristatd)

207

Sacred Ibis (Threskiornis aethiopicus]

211

Oriental White Ibis (Threskiornis melanocephalus}

217

Australian White Ibis (Threskiornis moluccd)

221

Strawnecked Ibis (Threskiornis spinicollis]

227

Waldrapp Ibis (Geronticus eremitd)

231

Bald Ibis (Geronticus calvus)

237

Black Ibis (Pseudibis papillosa)

241

Giant Ibis (Thaumatibis giganted)

245

Oriental Crested Ibis (Nipponia nippon)

249

Eurasian Spoonbill (Platalea leucorodid)

253

Royal Spoonbill (Platalea regia)

259

Blackfaced Spoonbill (Platalea minor)

263

African Spoonbill (Platalea alba)

267

Yellowbilled Spoonbill (Plataleaflavipes]

271

Roseate Spoonbill (Platalea ajajd)

275

APPENDIX

283

BIBLIOGRAPHY

329

INDEX

383

Introduction

T

I here is no one word or concept that can describe fairly the diverse group of birds numbered among the storks, the ibises and the spoonbills. Compared to the majority of birds, they are among the larger and all have relatively long legs, necks and bills as adaptations to their fundamental way of life—walking about in shallow water. However, storks, ibises and spoonbills range in appearance from the elegant to the ungainly, in size from the moderate to the huge, and in behaviour from the flamboyant to the secretive. Their large bills may be massive or thin, straight or curved, tubular or flattened. Some species are found in large spectacular assemblies and others in crepuscular isolation. Some of the species were once widespread and common, but are now at or near to extinction. Others have always been rare, with limited ranges and highly specialized ecological niches about which little is known. Each has its own mysteries yet to be revealed, and each has its own claim on our fascination. We have approached the task of preparing this book with some trepidation, not only because of its magnitude but also because of the growing certainty that, unlike most heron species, an unacceptable number of storks, ibises and spoonbills are facing extinction. Realization amongst the general public that much of our avian population is becoming dangerously depleted in many, if not all, parts of the world, has been slow to crystallize. Even amongst ornithologists, the true, and in many cases, horrifying situation of these species is only now becoming apparent as information pours in from remote portions of the globe. Information on rare storks, ibises and spoonbills is arriving at a rapid rate, and much of it is not encouraging. We have attempted to include as much of this up-to-date information on the changing status of the species as possible. But again and again in preparing this book, we find ourselves emphasizing how these birds are in competition with humans for space and resources. With the world's human population expanding, the space left for large waterbirds has shrunk so rapidly in so few years that we have barely yet come to grips with the devastation that has been wrought. Two particular scenarios tend to repeat themselves among the species accounts. One is that of shy elusive species living as far away from humans as they can in areas where they have had, until recent years, little contact with people, but where they now find little habitat left. The other is of species which have lived in close proximity to humans and have very often become dependent upon them for their very survival. An example of the first scenario is Storm's Stork, about which we have learnt very little but which, because of deforestation, might well have disappeared before we had fully realized that it existed at all. The second group is epitomized by the Waldrapp Ibis which has been the recipient of every type of persecution which humans are capable of inflicting. Today we have reached a vital stage in the relationship between humans and large wading birds. The task of conserving our environment and the birds within it can only be tackled if sufficient is known about the needs of individual species within the increasingly hostile globe upon which we live. We hope that we have managed to present as clear a picture as is possible of the precarious position of the various species of storks, ibises and spoonbills around the world and, where possible, we have suggested appropriate action to help their cause. A singular message from our review is: what is so desperately needed is protection and management of the wetland habitats that are essential to most of the species we discuss. We have endeavoured in this book to share with the reader our own fascination and concern for these birds. We are fortunate to have been able to draw upon our own experience and that of our editors and publisher to present what we hope is a readable and authentic account of most of what is known about these species. We have been most fortunate, too, in obtaining the services of two highly talented and dedicated artists, Alan Harris and David Quinn, who have provided portraits of each of the species described. We hope this original artwork will contribute to both the enjoyment and usefulness of this monograph.

2

Introduction

Figure 1. A Maguari Stork in full breeding plumage with two nestlings approximately 2-3 weeks old (northeastern Argentina).

Figure 2. tralia).

In today's financial climate, meeting the expenses of production and publication of a book of this nature, containing original plates, is thwart with difficulties. Vital to the project was the support of the Brehm Fond and Mr Wolf W. Brehm. We also are quick to acknowledge the seminal contribution to the project made by Mr Charles Luthin, working on behalf of the Brehm Fond, Vogelpark Walsrode, and the Specialist Group on Storks, Ibises, and Spoonbills. A volume such as this would have been impossible to publish at other than an astronomical price without this generous support. We are fortunate to have been able to draw much of our information and our insight from our own experience. The authors have been privileged to travel extensively, observing these birds on each of the continents they inhabit. Between us, we have seen in the wild all but 8 of the 49 species we discuss. Two of us have spent many years studying one of the sister groups of these birds, the herons (Ardeidae), having previously summarized our knowledge in a companion volume, The Herons Handbook (1984). Two of us have spent many years studying the storks, spoonbills and ibises themselves, the results of which have been published in numerous papers addressed to our scientific colleagues; one

having spent over 30 years studying the storks and more recently spoonbills and providing a fundamental perspective on their behaviour, ecology and taxonomy, the other, over 20 years studying populations and communities of storks, ibises and spoonbills on two continents, providing a comparative perspective on their ecology, habitat and conservation. To supplement our own observations we have consulted an abundant although very uneven literature in an attempt to synthesize what ornithologists before us have found about the biology and conservation needs of these species. Much of this literature is excellent; and, for present and future students, we hope the extensive, yet selected, bibliography in this book will be an entrance into this literature. The bibliography is a collaborative effort, assembled by a team led by Dr Malcolm Coulter (see below). One caveat: we would urge caution and selectivity in the use of this literature, some of which goes back many years. We have attempted to be selective and to present the conflicting views, but undoubtedly will have erred in our judgement in those cases where facts we gleaned from the literature are really quite debatable. We have also sought advice from a wide-ranging and knowledgeable host of experts. The response to our requests has been enormous, and we are deeply indebted to a very

A Glossy Ibis near its nest (southeastern Aus-

Introduction 3

large number of collaborators both amateur and professional who have been so supportive, encouraging and generous in sharing their insight and understandings. Some of our trepidation is also due to our certain knowledge that no matter how thorough our own efforts, the literature, or our correspondence, a book such as this will be incomplete and in some areas will be proved incorrect. Our limitations can, and undoubtedly will, be quickly recognized. The available knowledge of the biology of these birds is highly erratic. Some species are very well known, whilst others have hardly been studied at all. Our species accounts reflect this discrepancy. Over and over again we bring to the readers' attention significant gaps in our knowledge about the most basic aspects of biology. It is the nature of such a work to summarize, synthesize, pose hypotheses, and encourage further study and testing of these hypotheses. We hope this book will inspire others to fill in the important gaps in the knowledge and to test some of the hypotheses we propose, so that the business of conservation can continue more effectively. If the book accomplishes an increased interest and study, we will be satisfied to have accomplished our primary goaL Range maps are always troublesome. We are conscious of their summary value but have found, like many authors before us, that illustrating the ill-known and changing distribution of many species is well nigh impossible. We have, therefore, sought to show general ranges of each species, reserving for the text a more detailed description. Even here we have been handicapped by lack of precise information on the range of a surprisingly large number of species, about which little is recorded, and not least by the often unreliable recordings by collectors of the past whose poorly labelled specimens lack credibility. Information on basic nesting behaviour is missing for some species; and, for many, information on colour changes during courtship are lacking or conflicting. Soft-part colours of museum specimens are especially unreliable guides. In the field, information on courtship colours are seldom recorded in all phases of their development. Such knowledge as we have presented should be subjected to most careful further study by field-workers who are fortunate enough to come across them. Yet for species that change the colour of their skin during courtship, this information is vital. Voices and other noises are of importance to most of these species. Yet our understanding of their vocalizations is primitive compared to that of other birds. However, in some cases the only reliable identification of a species that can be made is by voice. These are large birds, with mostly gruff and non-melodic voices. We have rendered their calls in English inventing such words as 'Cha-cha', or 'Hunk-hunk*. These are recognizable by those who know them, but not necessarily by those who do not (in some cases, one simply had to be there). With three authors combining their respective strengths and weaknesses, it is unfortunate that there has to be an order of authorship. Such an order does not imply the relative contribution of each. This was a true collaboration of experiences and ideas. As an Anglo-American team, we express the hope that the individual and joint efforts here

Figure 3. A Eurasian Spoonbill head-scratching (northern India). presented will have brought together the sort of information which will provide encouragement and a structure for future researchers and observers. It is our hope, too, that the emphasis we have placed on the dire need for conservation of many of these species, and the wetland habitats on which they depend, will encourage responsible authorities to act on their behalf before it is too late. ACKNOWLEDGEMENTS We are extremely grateful to all those who have, lent their assistance in the production of this book. As noted in the Introduction, we especially acknowledge the support of Mr Wolf W. Brehm, who has made it possible for original plates to be produced. We thank, too, our long-time colleague Mr Charles Luthin, who nurtured the project, and because of whose efforts much of the information now available on the status of these species has been gathered by field-workers throughout the world. We thank too the excellent artists, Messrs Alan Harris and David Quinn, for their outstanding contribution to the quality and usefulness of the book. In actual production, we thank Dr Andrew Richford, our editor, Penny Robinson and Debra Kruse at Academic Press for their considerable help. We also thank especially Mrs Jane Ratliff and Mrs Bonnie Knight for their typing assistance and Mr Melvin Seid for bibliographic assistance. We thank our colleagues who worked on the bibliography. This was a collaboration lead by Dr Malcolm Coulter to assemble a bibliography for this group of species. The present reference section is derived from the first edition of this bibliography, which is available on computer disk from Dr Coulter; the compilers of that bibliography are

4 Introduction

Figure 4.

Strawnecked Ibis. Huge flocks fly north from southern Australia after breeding.

Malcolm Coulter, Koen Brouwer, Albert L. Bryan, Jr., M. Philip Kahl, Catherine King, James A. Kushlan, Charles S. Luthin, Marcel van Wieringen, and David P. Young, Jr. We are very grateful to our many correspondents who generously provided their unpublished information, observations, data and insight, and many of whom were kind enough to review preliminary drafts of various chapters. We take great pleasure in acknowledging their contributions: Eduardo Aguilera, Bubphar Amget, J. Ash, George Archibald, P.C. Bhattacharjee, William Belton, Bas Van Balen, Hilde Bult, Navancet Bhatt, A. Brosset, Linda Birch, Keith Bildstein, Walter Bock, Koen Brouwer, Benjamin Busto, Nigel Collar, Miles Coverdale, Malcolm Coulter, Juan P. Cueilo, Montserrat Carbonell, Finn Danielsen, J.H. Desai, Peter Davey, Andrea Dinep, Catherine Deulofeu, Jon Fjeldsa, Peter Frederick, Mercedes Foster, Paula Frohring, Michael Gochfeld, Juan Carlos Guix, Alfredo Guillet, D.A. Holmes, U. Hirsch, David Holyoak, Peter Hubbell, Cathy King, Richard Klein, P. Kennerley, Kim Lowe, O. Langrand, Charles Luthin, Catrina Martin and the Texas Parks and Wildlife and Texas Colonial Waterbird Survey for the use of survey data, Nigel Mathews, Burt Monroe, David Manry, D. Melville, Adam Mrugasiewicz, Gonzalo Morales, Samuel Narosky, Eva Neumier, R. M. Naik, John Ogden, Fabio Olmos, Storrs Olson, Raul Lopez Pacheco, Duncan Parish, Kenneth Parkes, Jan Pinowski, D.

Pomeroy, E. Poorter, the late Peter Pugh, Crawford Prentice, Ian Parker, Philip Round, Christina Ramo, James Rodgers, Asad Rahmani, W.B. Robertson, Tom Roberts, R.A. Ryder, Roberto Schlatter, Charles Sibley, the late Helmut Sick, Marcel J. Silvius, Henrick Skov, W.R. Siegfried, Sandy Sprunt, D. Scott, Holger Schulz, the late Ernst Schiiz, Gary Stiles, P. Saikia, C. Sivasubramanian, S. Smirenski, Betsy Thomas, Hashim Tyalji, Daniel M. Taylor, Tao Yu, Victor Terera, Emil Urban, AJ. Urfi, J. vanTets, David Wells, Scott Wood, Marcel V. Wieringen, S.W. Winter, M.D. Williams, Josef Witkowski, W.J.M. Verheugt. We gratefully acknowledge the University of Mississippi for its support for one of us (J.A.K.) and of this project, the MacArthur Foundation for the award of a MacArthur Fellowship to one of us (M.P.K.) allowing him to devote his time to this project, and the staff of Cox and Kings Travel Ltd for organizing travel arrangements in Asia for one of us (J.A.H.). NOTE ON RANGE MAPS The following colour coding has been adopted. Areas where the species concerned is a breeding resident are shown in blue. Where a species is completely or partially migratory, the breeding range is shown in red and the non-breeding range in green. Arrows indicate principal migration routes and ? areas where the status is doubtful.

Classification A S Y S T E M FOR C L A S S I F I C A T I O N We are able to understand the world around us in part through our ability to classify objects and events into manageable units and by establishing, in our mind at least, logical relationships among them. This is especially true of the natural world; the classification of living beings is a specialized science called Taxonomy or Systematics. As in our everyday efforts at pigeon-holing objects and ideas, the scientific system of classifying organisms also has two goals, to identify and name the entities of the natural world and also to describe the relationships among them. In developing a classification, taxonomists seek to place together organisms that are related through descent from a common ancestor. Such organisms share a common genetic inheritance that we expect to be expressed in various characteristics that we can see or measure. Of course, characteristics may in fact, be similar without being derived from genes shared with a common ancestor, and, conversely, there is no reason to suspect that all genetic similarity should be expressed in characteristics we can easily observe or measure. An increasing ability to discern biochemical differences, including characteristics of the DNA itself, provides additional measures of genetic similarity. The basic unit of taxonomy for organisms such as birds is the species. For sexually reproducing organisms, a species consists of the various populations of individuals that are potentially capable of breeding with each other and producing offspring that are not at such a disadvantage that they in turn cannot breed successfully. The species itself is identified by two latinized names, the genus name followed by the specific epithet, as for example in the scientific name for the Sacred Ibis, Threskiornis aethiopicus. Acceptability of names is governed by codified rules, especially the rules of priority—the first valid name proposed for an organism is the name everyone is expected to use. Determining the right name to use is sometimes but not always straightforward. Sometimes what constitutes the original description is debatable; species repeatedly have been reassigned among genera as new information and opinions about their relationships emerge. So most species have gone by various names since their discovery. In each species account, the scientific name and information on the source of the original description are given, as is the identity of the taxonomist credited with naming the bird. This attribution is placed in parentheses if the genus to which the species is assigned has changed since description. Unlike scientific names, common names vary from place to place and from language to language. There has been some effort to standardize common names in various geographic regions, where technical committees deliberate and provide lists of recommended usages. In each species account, we make an effort to provide the reader with some of these names. There is, however, no 'right' or 'wrong' common name, and some species have a score of names in several languages. The English names we use, we recommend for their clarity within context of the stork-ibis-spoonbill group worldwide. Over wide geographic areas populations of some species may diverge into recognizably different kinds of organisms but yet be fully capable of interbreeding. These distinguishable geographic entities are often called races or subspecies. Subspecies may be named, should it be useful to do so. For example the Sacred Ibis of Africa is Threskiornis aethiopicus aethiopicus, whereas the isolated birds of Madagascar are Threskiornis aethiopicus bernieri. In most cases, populations are not named, even though we can expect variation to occur among populations scattered over wide areas of the globe. What population should be considered a species and what a subspecies is often unclear, and so may be contentious. Interbreeding potential is what marks a species. Members of two different species would not interbreed if they occurred together; members of the same species or two subspecies of the same

6 Classification

species would; however, since they do not occur together, proof of their compatibility is usually lacking. To be considered a subspecies, a population must be distinctive yet potentially interfertile with other subspecies and capable of producing offspring that are not at a disadvantage in nature. (A species having more than one subspecies is called polytypic.) One should keep in mind that the more fundamental question of interest is not whether a population has a subspecies name, but rather what are the biological implications of the variation of characteristics among populations. Species that are similar, and obviously closely related, that are found in disjunct ranges are called allospecies and considered members of a superspecies. The Sacred Ibis, Oriental White Ibis, and Australian White Ibis are members of a superspecies, in our classification scheme. The reproductive isolation of individuals of two species results from differences in morphology, behaviour, biochemistry and, ultimately, genetics. These all provide criteria by which we too can identify various species, and also the criteria upon which are based the higher classifications, which attempt to express the genetic similarities that derive from sharing common ancestors. The genus name is a low-level way to provide an indication of relationships among species. Species that share the same genus name are more closely related to each other than they are to species in other genera. That is, species sharing a genus name also share a common ancestor, and a genetic similarity, not shared with other species.

Figure 5. An American White Ibis in breeding plumage, with gular-sac inflated (southeastern USA). Figure 6. American White Ibis. Immature (left) and adult (southeastern USA).

Classification

Higher-level classification proceeds in the same manner, with related genera being grouped into families, families into orders, orders into classes, classes into phyla, and phyla into kingdoms. To show finer relationships, primary taxa can be further subdivided into higher and lower groups, such as suborder (below order), subfamily and tribe (below family) or superspecies (above the species). In some detailed classification schemes, there may be many more taxa used to show details of relationships. Above the species level, classifications become increasingly a matter of professional opinion; although modern techniques of biochemical genetics and cluster analysis can make the process of seeking similarities more objective, questions of convergence nevertheless remain. STORKS, IBISES AND SPOONBILLS Who, then, are the storks, ibises and spoonbills? According to currently accepted working classifications, storks, ibises and spoonbills share a common classification down to the ordinal level. They are animals (kingdom Animalia), chordates (phylum Chordata), birds (class Aves), and wading birds (order Ciconiiformes). Which birds and which groups are additionally included in the order Ciconiiformes is, however, a matter of considerable debate in the pages of the scientific literature. The storks, ibises, spoonbills, herons and others are identifiable in part by their long legs and long necks, both adaptations for foraging by wading about in shallow water. These few characteristics of obvious ecological importance may seem to be rather slim branches on which to hang a taxonomic relationship. However, this intuitive grouping has held up to repeated scrutiny since Linnaeus in 1758, and we can be reasonably confident that the characteristics represent a fairly robust adaptive strategy. Other characteristics of the storks, ibises and spoonbills relate to the adaptive strategy of being a 'wading bird'. Their long necks have 15-20 vertebrae, and are capable of considerable mobility, as would be expected to be able to feed. Their tibiae and tarsi are unfeathered, and toes are elongated, often slightly but never fully webbed, with the lengthened back toe (hallux) also expected of a wading bird. Some groups sport toenails specialized for feather maintenance. They also have other characters in common (T.H. Huxley 1867, Baird et al. 1884, Goues 1903, Witherby et al. 1939), such as similar sterna, palates and syringes, minute intestinal caeca, elongated bills, long, broad and rounded wings, and short tails. But other birds also have some of these characteristics. They also share such common morphological features as particular conformations of carotid arteries and bony palate, which however are not unique to these birds. The sexes are identical in plumage; but females are, to a greater or lesser degree, smaller than males. Many species nest in colonies and all have helpless (altricial) young that adults feed in the nest. In addition to the similarities, the groups of birds traditionally included in the order differ in aspects of behaviour and morphology. It is, in fact, not possible to enumerate obvious morphological characteristics that definitively sep-

7

arate these species from other birds, some of which may or may not be included in the order. This deficiency has led to considerable debate, over the past 50 years, as to which birds should be included in the Ciconiiformes (see Sibley and Ahlquist 1972 and 1990 for further discussion). Storrs Olson (1979) concluded that it was an artificial assemblage. Alan Feduccia (1980) has suggested that resolution of the phylogenies of the groups currently included in the Ciconiiformes remains one of the major challenges of contemporary evolutionary ornithology. Either way, the use of biochemical characteristics has gone a long way to increasing our confidence in the close relationships among birds in the traditional ciconiiform groups. The concept of a ciconiiform taxon has a long and distinguished history. T.H. Huxley (1867) suggested a group, which he called Pelargomorphae, that included the ibises and spoonbills, storks, herons and flamingos. This was the basic view of Gadow and of Wetmore, who produced the classification scheme that for almost a century has been the basis for the modern consensus of avian classification. Wetmore's (1960) Ciconiiformes was subdivided into four suborders for the herons, the Shoebill, the storks, ibises and spoonbills, and the flamingos; these are the 'traditional ciconiiforms'. The herons were further divided into two

Figure 7. A Royal Spoonbill with crown feathers fully erect (southeastern Australia).

8

Classification

families, the typical herons (Ardeidae) and the Boatbilled Heron (Cochleariidae). The storks (Ciconiidae) and the ibises/spoonbills (Threskiornithidae) were in their own individual families, as was the distinctive Hamerkop (Scopidae). Wetmore's classification, minus the flamingos, was similar to that proposed about the same time by Verheyen (1959) and was congruent with one of the first applications of biochemical genetic data by Sibley (1960a,b). Since then, other studies have been conducted, including a proposed revision of the classification of the world's birds, based on biochemical and other evidence, by Charles Sibley and his associates (Sibley et al. 1988, Sibley and Ahlquist 1990, Sibley and Monroe 1990). These studies, in one sense, produced a radically different view of the Ciconiiformes, a more encompassing view that includes within the order other groups. Many of these have previously been suggested to be related in one way or another, but they were generally retained in other orders. At the same time, taxa have been removed from previously recognized orders, implying that these were not natural assemblages. This expanded ciconiiform order included, in addition to our groups, such groups as the waders (i.e. shorebirds), gulls and terns, hawks, grebes, tropicbirds, boobies, anhingas, cormorants, herons, the Hamerkop, flamingos, pelicans, New World vultures, frigatebirds, penguins, loons, petrels, and albatrosses.

In another sense, these studies have been comforting in that nestled within the expanded Ciconiiformes are the herons, storks, ibises and spoonbills, which continue to be perceived as being related at some level between family and order. It is useful to draw a distinction between a classification and a species sequence (Bock 1990). A classification is a three-dimensional tree of relationships that ideally reflects the phylogeny (i.e. the evolutionary relationships) of the group (Sibley and Ahlquist 1990). A sequence is an ordered list of taxa, derived from a classification schema. A standard sequence is a heuristic device that facilitates communication, and is not tampered with lightly (Mayr 1989, Bock 1990). In this monograph we have followed the standard species sequence of Peters' (1931) Check-list (Kahl I979a,b, J. Steinbacher 1979), to the fullest extent possible given the new understandings of species limits adopted in this book.

Figure 8. A female Yellowbilled Spoonbill performing the Greeting Display (southeastern Australia).

Figure 9. A Marabou Stork yawning at its nest (northwestern Kenya).

Questions about Ciconiiform Taxonomy All classification systems are really the expression of a series of hypotheses that provide the basis for new questions and new hypotheses (Sibley et al. 1988). Some of the principal taxonomic questions exposed by proposed classifications include: the relationships of the flamingos; the relationship

Classification

of the herons to the other groups; the most appropriate classification of the Hamerkop; the most appropriate classification of the Shoebill; the boundaries of the order with respect to possibly related taxa such as pelicans and their relatives, New World vultures, and hawks; and the family and generic relationships among the various species within the constituent groups of the traditional ciconiiforms. The relationship of the flamingos has been an especially controversial question, particularly because they show morphological similarities to ciconiiforms, waterfowl and waders. The question of interest in our context is whether the similarities they show to other long-legged wading birds might be due to convergence or to shared ancestry. Sibley and Ahlquist (1972) concluded that flamingos were indeed ciconiiforms and that flamingos linked the wading birds and the waterfowl. Feduccia and Olson (Feduccia 1978, 1980, Olson 1979, Olson and Feduccia 1980), on the other hand, argue from palaeontological and other evidence that shorebirds constitute the ancestral group of both the flamingos and the waterfowl, and that none of these groups is particularly closely related to the ciconiiforms. On the basis of their studies of DNA, Sibley and Ahlquist (1990), however, conclude that flamingos are indeed relatively closely related to other members of the heron-stork-ibis cluster of birds. The herons are, by all accounts, closely related to each other (Payne and Risley 1976). They have distinctive, narrow feather tracts and patches of powder-down feathers, a completely feathered head, seasonal development of elongated display plumes, a narrow skull, a long, straight bill, a modified 6th cervical vertebra that allows the neck to be held in an S-shape in flight, a comb-like (pectinate) middle toenail, and young with a single down coat. Hancock and Kushlan (1984), in a companion volume to the present monograph, provided a synthesis of the biology and classification of the group, the latter mostly derived from the studies of Payne and Risley (1976) and Payne (1979), although retaining several traditionally generic allocations pending further study. One of the longstanding issues regarding the heron group relates to the distinctive Boatbilled Heron (Cochlearius cochlearius). Hancock and Kushlan (1984) suggested that it should be grouped with the night herons. Sheldon (1987) more recently provided biochemical evidence for a subfamilial relationship between the Boatbilled Heron and tigerherons, and no distinction between day and night herons. Despite these finding, the linear arrangement of taxa by Payne and Risley (1976) and the overall integrity of the heron family continues to remain intact. If anything, recent studies have tended to find the Boatbilled Heron to be less distinctive from the other herons than previously thought. However, the relationships of herons to other wading birds has not been so well settled. Olson (1979) suggested that they show few morphological similarities to the other traditional ciconiiforms and are the only currently successful representatives of an early radiation of crane-like birds not clearly related to modern storks or ibises. He considers the family to be one of uncertain affinity (Olson 1985a,b). Biochemical genetic evidence continues to support their ciconiiform heritage and a relationship at some level within

9

Figure 10. Abdim's Storks at their nest on a rocky island in Lake Shala (southern Ethiopia). the order to ibises and storks (Sheldon 1987). More specifically, this evidence suggests that herons, the Hamerkop, flamingos, ibises, storks, the Shoebill and pelicans are descended from a common ancestor (Sibley and Ahlquist 1990). The Hamerkop (Scopus umbrettd) is a small bird, lacking the long legs and neck characteristic of the other traditional ciconiiforms. We have found that it flies, perches, feeds, nests, and engages in courtship, all in distinctive ways (Kahl 1967b, 1979c). As noted above, biochemical evidence (Sibley et al. 1988) suggests that it is related to the herons, storks and ibises, but it certainly is not a true stork. Because of its distinctiveness, we have chosen not to treat the Hamerkop in the present monograph. The Shoebill (Balaeniceps rex) shares anatomical characteristics with the true storks, including a distinctive tubular ear bone (Feduccia 1977) and a Bill-clattering Display (Kahl 1972e). It also differs in other respects. For example, we found that it did not excrete on its legs for cooling (Kahl 1967a) and its chromosomes differ from those of the true storks (De Boer and Van Brink 1982). It shows many similarities to pelicans (Cottam 1957, Feduccia 1977, Saiff 1978, Olson 1979, 1985b). Sibiey and Ahlquist (1972) considered the species to be a ciconiiform, and Feduccia (1977) considered it to be a stork. Kahl (1979a) and Olson (1985b) assigned it to its own family adjacent to the true storks. Sibley and Ahlquist (1990) proposed that, within their much expanded Giconiiformes, it is most closely related to pelicans, and with the pelicans is placed in a superfamily between the superfamily that includes the ibises and the one that includes the storks and New World vultures. Because of

10 Classification

Sibley and Ahlquist (1990) describe well the probable history of this group. Each is a result of a radiation exploiting an unoccupied niche, each showing its own way of life. Each flourished, and spread widely, retaining characteristics of their common ancestry in their morphological and genetic similarities. In this monograph, continuing our evaluation of the biology of these groups that began with The Herons Handbook (Hancock and Kushlan 1984), we summarize the biology of the storks, the ibises, the spoonbills and the Shoebill.

Figure 11. A Lesser Adjutant Stork feeding its young (northeastern India). its historical identification with storks, we have chosen to include it in this monograph. Although the boundaries of the order Ciconiiformes continue to be debated, there emerges a clear consensus regarding the storks, ibises and spoonbills. From the morphologybased classifications of Huxley and Wetmore, through the various behavioural studies such as some of our own, to the biochemical studies of Sibley and others, these three groups of birds are repeatedly found to be related. Analysis of DNA hybridization similarities indicate that herons, the Hamerkop, flamingos, ibises and spoonbills, storks, New World vultures, pelicans, and the Shoebill are more closely related among themselves than they are to other groups of birds and so are descended from a common ancestor.

Relationships within the Storks The typical storks are a distinctive group sharing a number of characteristics, including long legs with half of the tibia bare, relatively short toes with small webs, 12 tail feathers, 12 primary feathers (the 12th being minute and/or lacking in the wood storks and openbills), bare portions on the head, feathered oil gland, a common musculature of the pectoral associated with soaring flight (Vanden Berge 1970), a stout bill, predominantly simple black and white plumage colours, young with two down coats, large air sacs under the neck skin, and a lack of powder down. They also have behaviours in common, including cooling their legs by defecating on them (Kahl 1963b), a trait they share with the New World vultures (Ligon 1967). Olson (1985b) and Sibley et al. (1988), and others before them, suggested that they are closely related to the New World vultures, and Sibley and Ahlquist (1990) included the vultures in the same family as the storks, remarking that they had been taxonomically misplaced for two centuries. The most recent classifications of stork species are based on behaviour (Kahl 1979b), anatomy (D.S. Wood 1983), and a combination of the two (D.S. Wood 1984). These studies have grouped the species in fewer genera than was previously the case (Kahl 1972e). We draw from the results of both sets of studies in recognizing two higher level divisions of the true storks, the wood storks and the openbills in the tribe Mycteriini, and the typical storks, giant storks and adjutants in the tribe Ciconiini. We have chosen to use tribes in subclassifying the storks, rather than the more traditional rank of subfamily, because the proper taxonomic rank of the typical stork group remains under discussion as does the appropriate positioning of the Shoebill and New World vultures. Sibley et al. (1988) classify the traditional storks as a subfamily, in which case tribe would be the next lower taxon. In the present monograph, we continue the more traditional approach of considering the storks to represent the family Ciconiidae. The stork tribe Mycteriini includes four wood storks (Mycteria) and two openbill storks (Anastomus). All locate food primarily by touch rather than by vision, and have bills especially, and even uniquely, suited for this behaviour. The wood storks have large, cylindrical, but slightly down-curved bills, which they place in the water and hold open for fish to swim into. They also use special movements, such as pumping their feet or opening and closing their wing, to increase the likelihood of a fish entering the bill. Mycteria have distinctive displays including aerial Flying

Classification

Around, a Gaping Display, and Display Preening (see subsequent chapters for descriptions of these and other behaviours). Courtship includes a gaping and snapping of the bill but not the Bill-clattering characteristic of other storks. The differences among the Mycteria storks are minor, consisting primarily of plumage and soft part coloration (Kahl 1972b). It is interesting to note that one of the wood storks, the Yellowbilled Stork, is called Mycteria ibis. Through the vagaries of nomenclatural history, the rules of priority result in the name 'ibis' being given not to an ibis, but to a stork (Mathews 1913). The openbills have bicurved bills that leave a central gap. Where they come together at the tip, the curved mandibles form a forceps with which the stork pincers its preferred and almost exclusive prey, freshwater snails. The two species are markedly different in colour, Anastomus lamelligerus being black and Anastomus oscitans being mainly white. The two possess an Advertising Sway behaviour that seems to be unique (Kahl 1972d); during copulation the male shakes his head rather than clattering his bill. In the tribe Ciconiini, we include 13 species in the genera Ciconia (typical storks), Ephippiorhynchus and Jabiru (giant storks), and Leptoptilos (adjutants). D.S. Wood (1984), arguing on the basis of his quantitative analysis of character distance, deviated from Kahl's (1972e) classification by

Figure 12. Asian Openbill Storks sunning (northern India).

11

including the genera Ephippiorhynchus and Jabiru in a tribe with the typical storks. This reflects the undoubted affinities between the genera Ciconia and Ephippiorhynchus. However, the Jabiru also shares characteristics with the adjutants, and is somewhat of an intermediate between the other two giant storks and the adjutants. In that the classification scheme should reflect that intermediacy, it seems to us that a more natural classification is achieved by grouping all four genera into one tribe, Ciconiini. The typical storks, members of the genus Ciconia^ have straight or slightly recurved bills; the head and neck are feathered, but the skin around the eye (lores) is bare. A display, the Head-shaking Crouch, performed by the male is found uniquely in the genus Ciconia (Kahl 1971f, 1972c). Bill-clattering and whistling vocalizations are characteristic, especially in courtship. We have concluded that the Black Stork is the most primitive and the White Stork the most advanced in the genus Ciconia (Kahl 1972c). Abdim's seems closely related to the Black Stork, both behaviourally and morphologically. The Woollvnecked Stork of Africa and Asia and the Maguari of South America share a number of characteristics, including a forked black tail. We here recognize the Storm's Stork (Ciconia stormi) as a separate species from the Woollynecked Stork and the Oriental White Stork (Ciconia boyciana) as a separate species from the White Stork.

12 Classification

Fossils of this group are known, including both extant and extinct species of Ciconia as early as the middle Miocene and as late as the late Pleistocene in Africa (CJ.O. Harrison 1980). In the giant stork genera Ephippiorhynchus and Jabiru, a Flap-dash Display is used by pairs on the feeding grounds. The Blacknecked and Saddlebill Storks are similar in many ways to the typical storks. The Jabiru shares an inflatable throat sac with the adjutant storks. Additionally, the Saddlebill Stork has a vascularized chest patch. Because of its intermediacy between the typical storks and the adjutants, unlike D.S. Wood (1984), we have retained it in its own genus. The adjutants (Leptoptilos) all feed to some degree as scavengers, have massive bills, and lack normal contourfeathering on the head and neck, both adaptations for eating large carrion. They are unique in giving a loud squealing or mooing vocalizations during courtship (Kahl 1966a, 1970, 1972a). The large inflatable throat sac is an obvious characteristic of these storks.

Relationships within the Ibises and Spoonbills Ibises and spoonbills are medium-sized birds that have been recognized as being related since at least the mid 1800s (Nitzsch 1840). They are almost always grouped at the family level (Sibley and Ahlquist 1972, 1990). The most widely used name for the group is the Threskiornithidae (Eisenmann et al. 1984). It has been argued that by priority the more appropriate name is Plataleidae (K.E. Campbell 1986, A.R. Phillips 1986, Olson et al. 1986). However,

Figure 13. A Woollynecked Stork shading its young at the nest (northern India).

following WJ. Bock (pers. comm. 1990) in his advocacy of retaining the well-used name, as well as by priority, we continue to use 'Threskiornithidae'. The notable differences in bill morphology have long resulted in the recognition of two subfamilies, the Threskiornithinae including the ibises, having a narrow downcurved bill with a flattened tip, and the Plataleinae including the spoonbills with a flattened bill. The two bill types correlate with the predominant non-visual feeding modes adopted by the two groups, Probing and Head Sweeping, respectively (Kushlan 1978a). The various species have a number of characteristics in common, especially a distinctive slit-like cranial morphology (called schizorhinal), which they share with waders (perhaps by convergence) and which is an adaptation to allow upper bill movement. They also have a distinctive musculature and sternum, 11 primary feathers ( l l t h minute), 12 tail feathers, long legs with lower half of tibia bare, loss of feathers on the head and even neck, distinctive ornamental plumes, and changeable soft-part colours. They lack powder-down patches, have a cupped middle toenail undoubtedly used for feather maintenance, and a grooved mandible that functions in removing secretions of the supraorbital salt gland. They have down on both feather tracks and non-feathered portions of the skin (apteria). An interesting aspect of the biology of ibises is the variability of chromosome structure among some species, a very unusual trait among birds (including storks) in which chromosomal uniformity is the rule (Capanna et al. 1982). In addition to the traits they share with storks, the ibises and spoonbills share characteristics with the waders, traditionally classified in the order Charadriiformes. Olson (1985b) included ibises and waders in the same order. However, a relatively close relationship is not supported by DNA evidence (Sibley and Ahlquist 1990). The ibises and spoonbills have been relatively little studied in the field because they are hard to observe and the various species are well scattered around the world. Extremely little is known about certain rare or inaccessible species. The ibis subfamily Threskiornithinae consists of 23 species distributed among 14 genera. It is likely that the present classification does not adequately represent relationships among the species. However, taxonomically useful anatomical trends among the species are not readily apparent. It is also an old group, dating back to the Miocene, about 25 million years ago. The earliest fossils are assignable to a still living genus, the widespread Plegadis (Plegadis peganus and Plegadis pharangitus—Olson 1981). Species representing two other living genera are known from the Pliocene of South Africa, Geronticus apelex and Threskiornis aethiopicus, the bones of the latter being indistinguishable from the modern form that occurs in the area today (Olson 1985a). Extinct species of present genera (Theristicus wetmorei and Eudocimus peruvianus) are known from the Pleistocene in South America (K.E. Campbell 1979). Especially intriguing are subfossils of flightless ibises, Apteribis glenos, from the islands of Maui and Molokai in the Hawaiian Islands and Jamaica (Xenicibis xympithecus) (Olson and Wetmore 1976, Olson and Steadman 1977, 1979, Olson and James 1991),

Classification

which indicate the occurrence of very recent extinctions of particularly vulnerable species of ibis. The Hawaiian forms were undoubtedly lost through the hand of man. Relationships among the ibis genera are not well understood. The glossy ibises (Plegadis], represented so anciently in the fossil record, today have a nearly worldwide distribution, the only ibis genus to do so. The Glossy Ibis itself (Plegadis falcinellus] is found on all inhabitable continents and may be a relatively recent invader of the New World. The Whitefaced Ibis is the South American and western North American form, whereas the Puna Ibis is a South American montane form. From available fossil and osteological evidence, it appears that the genus Eudocimus is closely related to the glossy ibises (Mayr and Short 1970, Olson 1981). The genus has traditionally been considered to include two forms, the American White Ibis (albus) ranging from North America to northern South America and the Scarlet Ibis (ruber) found in northern South America. Because their recent studies demonstrated considerable successful interbreeding of red and white birds in a natural zone of overlap, Ramo and Busto (1987) concluded that, by definition, the two forms must be considered the same species. This conclusion is supported by other evidence, not the least being that the presence of the carotenoid pigment canthaxanthin, which accounts for the colour of the red form, is unreliable as a taxonomic characteristic (Sibley and Ahlquist 1990). South America is a centre of ibis radiation. Most species there are at this time assigned to unique genera, reflecting the dearth of information available on extant and fossil South American forms, lack of taxonomic study of the subfamily as a whole, and the probable age of the group. Theristicus is here considered to include a single polytypic species. The genera Phimosus, Cercibis, Harpiprion, and Mesembrinibis are also represented by single species. Sibley and Ahlquist (1990:852) present the intriguing result of one experiment: that there may be a split within the family between the South American genera (Mesembrinibis, Theristicus, Cercibis, Eudocimus} and the Old World ibis genera and spoonbills. This deserves further study. The sacred ibises (Threskiornis] are widespread in the Old World. Once divided into four species, they were combined into two species and many subspecies by Holyoak (1970). More recent studies of their behaviour, structure, and chromosomes suggest that three species exist (Cramp 1977, De Boer and Van Brink 1982, Mees 1982, Lowe and Richards 1990). In this monograph we restore the division of this genus into four species, three of which are allospecies. De Boer and Van Brink (1982) suggested that the Australian White Ibis (Threskiornis molucca) may be the ancestral form from which the other two species were derived. Several species of ibis of Africa, Madagascar and the

Figure 14. A Strawnecked Ibis in breeding plumage (southeastern Australia). Figure 15. Black Storks at their nest; male nearest the camera (southwestern Poland).

13

14

Classification

spoonbills, and may be most closely related to each other (Kahl 1988a).

Species Sequence of the Storks, Ibises and Spoonbills Family CICONIIDAE—-the true storks Tribe MYCTERIINI—the wood storks and openbills Mycteria americana — American Wood Stork Mycteria cinerea — Milky Stork Mycteria ibis—Yellowbilled Stork Mycteria leucocephala—-Painted Stork Anastomus oscitans—Asian Openbill Stork Anastomus lamelligerus — African Openbill Stork Tribe CICONIINI—the typical storks, giant storks and adjutants Ciconia nigra — Black Stork Ciconia abdimii—Abdim's Stork Ciconia episcopus—Woolly necked Stork Ciconia stormi—Storm's Stork Ciconia maguari — Maguari Stork Ciconia ciconia — White Stork Ciconia boyciana—Oriental White Stork Ephippiorhynchus asiaticus — Blacknecked Stork Ephippiorhynchus senegalensis—Saddlebiil Stork Jabiru mycteria— Jabiru Stork Leptoptilos javanicus — Ltsstr Adjutant Stork Leptoptilos dubius —Greater Adjutant Stork Leptoptilos crumeniferus — Marabou Stork Figure 16. A Shoebill in a grassy marsh adjoining Lake Kyoga (north-central Uganda).

Family BALAENICIPITIDAE — the Shoebill Balaeniceps rex — Shoebill Family THRESKIORNITHIDAE — t h e ibises and spoonbills

Middle East are included in the genera Bostrychia, Lophotibis, and Geronticus. It is likely that these are fairly closely related to each other (Fry et al. 1985). Three genera of ibises are found only in Asia, Pseudibis, Thaumatibis, and Nipponia. Olson (1985b) suggested that Pseudibis is anatomically very similar to and possibly congeneric with Geronticus. Although the Giant Ibis has most recently been considered to be a Pseudibis (Holyoak 1970), Olson (1985b) has cautioned against removing this distinctive, large ibis from its specialized genus without anatomical study. We have followed this sensible advice and restore it to its separate genus. The spoonbill subfamily Plataleinae has a cosmopolitan, although somewhat patchy, distribution. Four of the Old World species are closely related, and largely replace each other geographically. We here identify the Royal Spoonbill (Platalea regia} as a separate species, based on our recent studies of its behaviour and morphology, which differ from the Eurasian species (Kahl 1988a). The specific distinctiveness between the Royal and Blackfaced spoonbills (Platalea minor) also needs scrutiny. The New World Roseate Spoonbill (Platalea ajaja) and the Australian Yellowbilled Spoonbill (Platalea flavipes) are more distantly related to the other

Subfamily THRESKIORNITHINAE —the ibises Eudocimus ruber—American White Ibis Phimosus infuscatus—Barefaced Ibis Plegadisfalcinellus—Glossy I bis Plegadis chihi—Whitefaced Ibis Plegadis ridgwqyi — Pun& Ibis Cercibis oxycerca — Sharptai/ed f bis Harpiprion caerutescens—Plumbeous Ibis Theristicus caudatus—BufFnecked Ibis Mesembrinibis cayennensis—Green Ibis Bostrychia kagedash — Hadada Ibis Bostrychia carunculata — Wattled Ibis Bostrychia olivacea—OVive Ibis Bostrychia rara — Spotbreasted Ibis Lophotibis cristata — Madagascar Crested Ibis Threskiornis aethiopicus—Sacred Ibis Threskiornis melanocephalus — Oriental White Ibis Threskiornis molucca—Australian White Ibis Threskiornis spinicollis — Strawnecked Ibis Geronticus eremita — Waldrapp Ibis Geronticus calvus — Bald Ibis Pseudibis papillosa—Black ibis Thaumatibis gigantea — Giant Ibis Nipponia nippon—Oriental Crested Ibis Subfamily PLATALEINAE—the spoonbills Platalea leucorodia — Eurasian SpoonbiJJ Platalea regia — Royal Spoonbill Platalea minor—Blackfaced Spoonbill Platalea alba — African Spoonbill Plataleaflavipes~ Yellowbilled Spoonbill Platalea ajaja — Roseate Spoonbill

Conservation

H

ow unfortunate it is that large water birds are particularly subject, either intentionally or otherwise, to continual attack from the human race. They are hunted, they are eaten, they are harassed, their feeding habitat is altered, their nesting sites are destroyed. On the other hand, they are also appreciated, for their size, their beauty or ugliness, or for the place many hold in the culture and history of a people. In our studies of the herons of the world, we were constantly amazed at the resilience of members of that family (Hancock and Kushlan 1984). In our work on the present volume, we found that for storks, ibises and spoonbills the story is a very different one. These birds have come under much the same pressures as have herons, but for reasons not quite clear, except as explicated for an individual species, they have responded poorly to the changes imposed upon them. A result is that many species of this diverse group of birds are at, or near extinction, over much of their former range (W.B. King 1981, Collar and Andrew 1988). In many areas, these bird populations are suffering from a combination of pressures including deliberate killing, deforestation, wetland drainage, use of intensive modern farming methods, and the accumulative effects of pesticides (J.F. Parnell et al. 1988). These problems seem universal, though the degree to which they apply varies according to the awareness and concern of the human inhabitants of the region. In fact, almost invariably for these large birds, it is the attitude of the local human population that decides whether a species survives in its midst. Solutions are by no means unclear, although they may be difficult to put into practice; J.F. Parnell et al. (1988) discuss these. Habitat improvement is required to reverse the impact of wetland destruction and alteration. In many cases improvement will require direct manipulative management of water, vegetation and breeding sites in compensation for loss of natural feeding and nesting sites, as well as monitoring and protection of existing functioning ecosystems (Kushlan 1979b, 1983, 1987, 1989a,b). More general environmental improvements are also required, especially regarding the long-term effects of chemical contaminants. It has been noted that 'People control is a foremost aspect of colonial waterbird management' (J.F. Parnell et al. 1988), and control of disturbance, including both passive disturbance and hunting, is needed, especially at colony sites. In some cases it is necessary to consider management to reduce natural interactions, such as predation and competition, for crucially endangered species. Multispecies fishery management is required in those areas where the needs of these birds are at odds with those of human fisheries. In some cases, the enhancement or re-establishment of a population may require reintroduction of birds, from wild or zoo stocks, and the artificial establishment of suitable nest or colony sites (e.g. Hirsch 1978a,b). Also critical are additional research on limiting factors and increased emphasis on education, especially of rural peoples. To be most effective, these management and conservation activities should be enacted on a regional basis. Luthin (1987), in discussing the conservation needs of storks, noted that a species' conservation concerns will often differ within its overall range. Also, because more than a single wading birds species may be dependent on regional resources, it is often within this multispecies context that management plans should be drafted. These plans, single-species or regional, are essential to define and guide conservation activity. In some parts of the world, mankind is belatedly attempting to halt the destruction of forest and wetland habitat in which the majority of storks, ibises and spoonbills live. But it is not clear if it is too late for some populations. Endangered means that a species or subspecies is in danger of extinction, and its survival is unlikely if the causal factors continue to operate (Collar and Stuart 1985). Populations that are decreasing may be

16

Conservation

Figure 17. A Milky Stork flying over the nesting colony (northwestern Java, Indonesia). considered vulnerable if further declines could make them endangered. Some species are vulnerable because of their rarity, which may or may not be due to human intervention. We consider five species and two subspecies to be endangered, and six species to be vulnerable. These species encompass over 20% of the world's species of storks, ibises and spoonbills. Amongst the storks, a number of species are endangered and many more are decreasing in numbers in some or all of the areas they inhabit (Luthin 1987, Collar and Andrew 1988). The Storm's Stork is a vulnerable species that has never been common and is seldom seen. Until recently we were not even sure of its status and so may never fully understand its biology before it becomes extinct. Other species are rare because of man, the Lesser Adjutant (vulnerable) and Greater Adjutant (endangered) Storks have all but disappeared over most of their range and are now in grave danger. The vulnerable Milky Stork remains secure only in the Indonesian archipelago. The Oriental White Stork and Black Stork, both vulnerable species, are increasingly confined to isolated breeding sites. Ibises and spoonbills have a number of species at risk (Manry 1986). Of the six species of spoonbill, the Blackfaced Spoonbill is the most endangered, having retreated as a breeding bird to a few islands off the North Korean coast

or perhaps to other unknown sites yet to be discovered. The endangered Giant Ibis may just be hanging on in isolated areas in Cambodia and perhaps Vietnam. The endangered Davison's race of Black Ibis now occurs only in Vietnam and possibly Indonesia. The endangered Waldrapp Ibis has now gone from Turkey, appears to be rapidly decreasing in Morocco, and seems to be on the verge of becoming extinct (Hirsch pers. comm.). The highly endangered Oriental Crested Ibis is now known to occur in the wild only in Qinling Shan in south Shaanxi, China, where the known population is in the low dozens. The continued existence of the dwarf Sao Tome Island race of the Olive Ibis has only recently been confirmed, but the situation is extremely precarious for this endangered race. The once widespread Madagascar Crested Ibis is vulnerable due to habitat changes that have been occurring in Madagascar. Everywhere pressures of human population have affected even the widespread species, and as suitable environments shrink so the pressures on all species increase. Regional decreases in numbers are documented in most species for which such data are available. It is likely that in the last 50 years that populations of most storks, ibises and spoonbills have been reduced by at least a half, and in many species by more. The extent of these reductions vary regionally. Bird populations in North America are probably the best

Conservation

17

known in the world. Trend in numbers and distribution suggest that considerable changes in ranges and populations have been underway for many decades. Perhaps the greatest decreases are in the Florida Everglades, where once thriving populations of American Wood Storks and American White Ibises have been decreasing to the point of local extinction (Frohring et al. 1988). The desperate position of the Ciconiiformes has developed with great rapidity. On the other hand, these species still maintain sizable populations elsewhere in North America and into South America. Within the United States the conservation and management needs are becoming well recognized (J.F. Parnell et al. 1988), but many countries have not the resources available in the USA to fight for the survival of decreasing species. Over much of South America, populations of all species remain high (or unknown). Even where historical decreases have occurred, there is evidence that populations may be returning. However in South America, apart from a few individuals and isolated concerned government or private bodies, conservation of habitat or species is not yet underway. In Europe the protection of species by concerned government and private bodies is holding back the deterioration, but species such as the White and Black Storks are falling in numbers annually. Eurasian Spoonbills are confined to a

few sites in western Europe. The former indifference, or active persecution, in western Europe has, except in Mediterranean regions, belatedly been replaced by a growing concern, manifesting itself into sometimes ineffective but now increasingly professional attempts to protect such species on their feeding and breeding grounds. Quite recently, however, it has become clear that in eastern Europe, pollution created by ill-conceived and inefficient industrialization is on a much larger scale than was previously realized. Some areas, it is thought, may never recover from the environmental damage, but all require attention. In Africa, the picture is probably the most favourable as most species remain widespread and even common (Collar and Stuart 1985). North of the Sahara, the Waldrapp Ibis is fast on its way to extinction. It is well to note that such local extermination is not new in the area. The ancient Egyptians mummified Sacred Ibises by the hundreds of thousands, perhaps over a million, undoubtedly reducing the populations that probably were finally extirpated by land use changes (Goodman 1988). Modern times have produced oil spills and the catastrophic effects of recent wars have yet to be evaluated. Historically the peoples of subSaharan Africa have shown tolerance to creatures around them, but increases in human population have begun to change the face of this continent, and the need to feed increasing numbers

Figure 18. A Bald Ibis (northeastern South Africa).

Figure 19. A female Blacknecked Stork foraging (northern Australia).

18 Conservation

of people has, together with ill-conceived development schemes often funded by western agencies, put pressures on all species of storks, ibises and spoonbills (Collar and Stuart 1985). Island forms in the Atlantic and Indian Oceans continue to be at great risk. In Asia the picture is very bad (Bain and Humphrey 1980, Hancock 1989). Here many wading bird species are very near extinction over much of their former range. Only in parts of India and in the, as yet, undeveloped areas of the Indonesian archipelago do some of the former abundant species survive in any numbers. The reasons for such a dramatic change, which has occurred since the end of the Second World War —a matter of about 45 years — are not hard to find. The elimination of endemic diseases and the resultant population explosion combined with a huge increase in the standard of living of many, though by no means all, its peoples have meant a rapid and urgent need to exploit all its natural resources (D.A. Scott and C.M. Poole 1989). Southern and eastern Asia comprises only 14% of the world's land surface, but support at the present time 56% of the world's human population. Furthermore this huge population is expanding at about the rate of 55 million per annum (D.A. Scott and C.M. Poole 1989). Also in Asia, not only has continuous warfare in many areas destroyed much of the formerly unexploited natural habitat, but the demand for food under such circumstances has meant the killing of large birds for food. Weapons are now readily available for shooting, and in many areas no feeding bird is safe from the gun, and no breeding bird safe from the trap. There has been a breakdown in the traditional religious beliefs in some nations, notably China, such that temple grounds and monastries, which traditionally provided protection for storks and ibis colonies, now seldom do so today. An exception which still enjoys protection in the temple confines is the Asian Openbilled Stork of Thailand. Today's awareness of the need for conservation measures has barely touched some nations or regions of Asia. The peoples of Pakistan, have a tradition of hunting which, using modern weapons, has served to nearly exterminate all the species of wading birds that bred in their land. War and poverty in the Philippines have brought with them heavy deterioration in the numbers of large birds. In Japan the recent stirring of conscience there has come too late in the day for most species. The subcontinent of India provides, perhaps, the clearest example of how religious and traditional tolerance has meant the survival of storks, ibises and spoonbills, and other resident and migratory species of large water birds. In areas where such ancient customs either do not pertain or have been lost beneath the struggle to survive, such species have

Figure 20. Jabiru Storks at the beginning of a copulation, with neck-bands a fiery red (northeastern Argentina). Figure 21. A female Saddlebill Stork at its nest with three nestlings about 45-50 days old (western Kenya).

Conservation

been exterminated. An example is the Black Ibis, which has its stronghold in Gujarat, a state composed mainly of tolerant, strict Hindu peasants and landowners. Here this species is quite tame, feeding around village ponds and nesting near to houses and other occupied buildings. But in states where hunting and trapping are still carried out, it is only in the nature reserves that the Black Ibis survives. Even so, in Gujarat, such tolerance has not saved the Blacknecked Stork. Drought conditions have dried out wetlands, and increasing demand for firewood has resulted in the destruction of trees on a vast scale, so that this bird has lost its traditional nesting sites, with the result that no breeding pairs remain there. Australia is the area of the world that retains its several species in greater abundance than perhaps any other. The Blacknecked Stork is at the southernmost limit of its range in Australia and New Guinea, having been almost totally wiped out in the rest of Asia apart from a few pockets of birds in India and perhaps Cambodia and Vietnam (Hancock 1989). Conservationists and conservation organizations in India, Africa and Asia are struggling with the problems they face. Private organizations such as Interwader in Asia are surveying the shrinking wetlands and recommending methods of protecting them. The ICBP is donating funds for work in many parts of the world. Brehm Fond has sponsored surveys and workshops on colonial bird conservation in Asia. In spite of all the measures being taken, there is an inevitability about the demise of some species. Given the need to preserve genetic stocks represented by these species and the hope that habitat restoration will allow reintroduction, it would seem likely that captive breeding remains the only hope in some cases. The Waldrapp Ibis thrives in captivity with some 500 birds, whereas in the wild the total population may well be below 100. The Oriental Crested Ibis numbers in China may be increased if a way can be found to breed them successfully in captivity. Many storks prosper under captive conditions and could be released in the wild if suitable protected areas can be found for them. The zoological parks of the world are attempting to breed these species, and such efforts should be encouraged ( L u t h i n e t a l . 1986, R.E.Johnson et al. 1987a,b). Of course, all proposed reintroductions require detailed consideration and study (Kushlan 1980, Seward 1987). The need for education is paramount to the success of any conservation measures. The recent discovery of the few colonies of Greater Adjutant Stork breeding in villages in Assam heightens the need in that area (Saikia and Bhattacharjee 1990) As a scavenger the species requires a food supply provided by man and his domestic animals, and does not appear to thrive in nature reserves where man is excluded. Conservationists, scientists, farmers and local landowners do not mix happily and readily together, and great care will be required if useful and productive understanding is to be reached in Assam, and indeed in many other areas where man, stork, ibis and spoonbill live in close proximity. Similarly, in South America, education efforts must be aimed at both the populace and government agencies.

19

Popular programmes, environmental education, and in country and out of country workshops for wildlife professionals may be used with success. Without such education, the best laid conservation scheme will come to naught.

Figure 22. White Storks copulating on a rooftop nest (southwestern Poland).

Courtship and Reproduction

I

n order to reproduce, storks, ibises and spoonbills must form pair-bonds, mate, build nests, lay, incubate and defend their eggs, and rear their young at least until the chicks are able to fly safely from the nest. Reproduction is a complicated affair, requiring in all these species the cooperation of both parents. A few species, such as the Black Stork, and the Hadada, Spotbreasted, and Black Ibises, nest alone in pairs. These usually are found in tall trees, especially in forests. Most species nest in colonies. Colonial species may nest on the ground, in reeds or in trees, depending on the species and the environment. Perhaps the most unusual are those that nest on cliffs, including the Waldrapp, Bald, Wattled and Buffnecked Ibises. Timing of nesting differs among species and locations. Both photoperiod and environmental conditions affecting food availability seem crucial in determining nesting timing. In the high latitudes, nesting is decidely seasonal and is associated with migrations out of the nesting area in the local winter. In the subtropics, nesting is often in the drying season. In the tropics, some species may nest nearly all year round. In all cases nesting occurs during the local period of high food availability. In times of low food availability, especially due to inappropriate rainfall or surface water conditions, nesting may be postponed or may be skipped in a given year. The Waldrapp and American White Ibises are examples (Smith 1970, Kushlan 1978b). The tight relationships of rainfall to nesting is particularly demonstrated in Australia, the Florida Everglades and Africa (Carrick 1962, Urban 1974b, Kushlan 1989a,b). Meeting and overcoming environmental and social constraints on reproduction are achieved through the use of appropriate behaviours, some of which have been ritualized into displays that allow one bird to communicate with another. THE PAIR-BOND The pair-bond is the key to successful nesting. Both members of the pair participate in building and protecting the nest, incubating the eggs, and feeding and caring for the young until they fledge from the nest site under their own power. It seems that in these species a single parent is insufficient to supply simultaneously the protection and food needed to raise young. Despite the importance to both individuals of maintaining a strong pair-bond to raise their young, it is becoming increasingly clear that promiscuous mating behaviour, called extrapair copulation, is extremely common in some species. Colonial nesting presents opportunities for promiscuity that are less available when, as in most species, birds nest in widely separated nesting sites within defended territories. The evolutionary advantage is clear: the promiscuous male increases its chance to sire additional offspring while the female is receiving a bold and clearly successful male who might father some of her young. But risks are also present, not the least of which is the potential loss of one's own nest or eggs if they are left unattended, as we have seen in the American White Ibis (Kushlan 1973b). An important question is whether extrapair copulations result in insemination, and this is a difficult question to study in these birds. Another question is what are the strategies the male uses to defend its mate against such intrusions, while also attending to its other duties such as supplying nest building materials? In the Eurasian Spoonbill, most extrapair copulations occur while the resident male is away gathering nesting material; and, perhaps as a response, males usually copulate with their mate just

Courtship and Reproduction 21

before leaving them to go on stick-collecting trips (Aguilera 1989, Aguilera and Alvarez 1989). Despite promiscuity, even if an intruder is caught in the act, the initial pair-bond usually persists, probably because the pair has little choice if they are to raise their young. Pair-bonds last for at least a nesting episode. If nesting is interrupted at an early stage, many species can nest again during the same season. Individuals of some species may repair in successive years, or even remain paired year-round. The White Stork is famous for maintaining pair-bonds over many years. It and similar species are said to 'mate for life'. In most cases, they re-mate as long as they both return to their previous site at the right time. If not, new mates may be taken. Several species of tropical ibises and several storks, such as the Saddlebill, Blacknecked and Jabiru, are solitary, non-migratory, and may remain together throughout the year. These pairs may not require extensive courtship at the start of the next breeding season. Pairs form during a courtship period in which both birds engage in a series of behaviours that help establish and then maintain the bond. The initial stages are crucial in that the birds have to overcome the tendency to exclude other birds nearby. That is, for much of the year, a bird tries to keep other birds at a comfortable distance, yet, when pairing, is faced with accepting the close proximity of another bird. One correlate of pair formation in most species is a seasonal change in the colour of the bill, face, legs or other exposed skin, called the soft parts. In the American White Ibis, the face turns a bright red and the bill black, and the females develop a red gular pouch under the chin. In the Sacred Ibis, the skin along the breast and under the wings turns bright scarlet and is highly visible when the bird flaps its wings. The Sacred Ibis also develops ornamental plumes at this time. The Oriental Crested Ibis has a unique type of cosmetic coloration. It uses its head to paint its pinkish back feathers grey with a secretion from its preen gland. These colour changes enhance the displays of the birds. The overall process is more similar between species of storks, ibises, and spoonbills than it is different. However, the details of the behaviours used in the process, called displays, differ among species. Species-specific courtship displays reduce the chances of mating with the wrong species. The occurrence of various displays and the different forms of display can serve as distinguishing characteristics between the species, just as morphological and plumage differences do. Most of the displays shown by storks, ibises and spoonbills during their courtship are 'derived' activities (Tinbergen 1952). These are behaviour patterns —usually borrowed from one or more other activities such as preening, fighting or fleeing™that have evolved as social signals. Displays have become specialized through a process known as 'ritualization' (Blest 1961), in which the elements of the display have become elaborated and stereotyped into the speciesspecific patterns that are recognized by the recipient bird. The various displays used by storks, ibises and spoonbills can be defined and categorized. Many of these behaviours have been previously described and named (Kahl 1966a, 1971f, 1972a-d, 1973b, 1983, 1988a). Displays are named

according to their component body movements, without assigning or assuming motivation. For purposes of enhancing clarity, increasing descriptiveness and reducing motivational inferences, we have renamed some of the terms currently found in the literature. These changes are noted in the accompanying table. So that the named ritualized behaviours will stand out and be easier to locate in the text, they have been capitalized. In general, using the same name for a behaviour in more than one species implies that the displays are homologous; that is, evolutionarily derived from a single display of a common ancestor. Certain displays show close similarities among the ibises and storks, and the herons (Hancock and Kushlan 1984). However, because of uncertainty as to the phylogenetic closeness of these groups, it is best to maintain a separate behavioural nomenclature, unless the similarities of the displays are so compelling as to lead us to imply their homology. In general, the process of courtship starts when birds begin to gather near a potential nesting site during the daytime. Depending on the species, these sites may be patches of trees, marsh reeds, or cliffs. In most cases the nesting sites are characterized by some degree of remoteness or inaccessibility, and many are located on islands. In these

Figure 23. American Wood Stork. Social interactions between birds in a treetop colony (southeastern USA).

22 Courtship and Reproduction

locations males behave in ways that serve to attract females and to advertise their readiness to mate. Most information available is on colonial species, and generalizations are often based on these species. It can be expected that solitary species may have slightly different approaches to mate attraction. In some non-colonial species, individuals remain on an activity area year-round and may return to the same nest-site in succeeding years. The Hadada Ibis appears to exhibit pair-bonding behaviour even within a flock, as pairs of birds separate out to preen mutually. In colonial species, the male birds stand in groups in a potential colony site and periodically fly about. The colony forms as displaying males and interested females gravitate toward each other, forming 'bachelor parties'. It is usual to find that a large colony is composed of smaller groups in which all the pairs are in the same stage of nesting, as a result of displaying birds having been attracted to each other. A male eventually selects a display perch within the selected colony site in a location where he is visible to interested females. In most species the eventual nest-site is located at or near the display perch, so it is possible that the quality of the site a male is able to secure and hold against competing males may be of considerable importance in formation of the pair. Mate choice is primarily a matter of the female selecting from among available, displaying males. The displays of the male also serve to intimidate nearby competing males. When the defensive content of a display is insufficient, fights can develop among males. The female approaching its chosen male also has to overcome his aggressive tendencies, and some of her displays at this time can be interpreted as appeasements. In the American White Ibis, the much smaller female can become quite bloodied during the process of gaining access to the male's defended display site. After mate selection is made, the pairing is reinforced by additional displays that often continue for several days. These eventually lead to copulation. Since the displays leading up to pair formation and copulation are defined differently for the two groups (storks, and spoonbills and ibises), it is worthwhile to describe these separately.

Figure 24. A Yellowbilled Stork feeding by tactolocation with bill held open and moved slowly about in water (central Kenya). Figure 25. A Painted Stork sunning in the 'delta-wing' posture (northern India).

COURTSHIP IN STORKS In all species, courtship begins with a male making himself conspicuous and available to an interested female. In many situations, the male just stands conspicuously. During Standing, a bird may try to look imposing. The Marabou blows up its gular air sac. One of the first active advertising displays used by male storks of the genus Mycteria in their bachelor parties or on their display sites is Display Preening, in which the bird repeatedly goes through the motions of running its bill down the primaries (the main flight feathers) of one wing then the other. It does this just as if preening, but the feather itself is not grasped, and perhaps not touched at all. Often occurring during episodes of Display Preening,

Courtship and Reproduction 23

Swaying Twig-Grasping display is executed by the male bending forward to one side and grasping a twig. Upon releasing the twig, the bird sways to the other side and repeats the process. A similar advertising behaviour found only in the openbills is the Advertising Sway. With bill gaping, the bird bends its head to its feet and lifts its wings slightly away from its body, while shifting its weight from foot to foot and swinging its bill from side to side. The Head-shaking Crouch, is given by storks of the genus Ciconia^ when a female approaches a displaying male. He inclines his body forward with head and bill low, wings slightly away from body and, while shifting from foot to foot, shakes his head in a 'no' motion, sometimes lowering the body to the nest in the process. A ritualized response to disturbance is the Arching Display, given by all stork species. In this 'anxiety' display, the bird, from an erect posture with feathers sleeked down, leans forward and looks at the offending object. In some species this may be accompanied by spread wings and vocalization. The generalized agonistic display among storks is the Forward Display. With the neck retracted, body horizontal, and feathers erect, the bird points its bill at its opponent and moves toward him. An antagonistic bird may also rattle its bill in a Forward Clattering Display. Although ritualized, these threat displays can lead to overt fighting should the intruder not choose to retreat. A hostile bird may pursue an opponent into the air using an Aerial Clattering Display. The pursuing bird flies after and above the opponent with its neck extended and bill pointed toward him. The attacking bird may clatter its bill but usually does not try to bite the opponent, who usually takes evasive action. An intriguing, apparently agonistic, display is called Mock Fighting. On or near the nest-site, a bird suddenly erects its neck feathers and makes lunging grabs into the air as if fighting or catching a flying insect. The bird frequently loses its balance and has to flap its wings to stay on the perch. The usual subordinate display of storks, is called the Upright Display. The submissive bird stands erect with head extended upwards, feathers compressed, and turns away from the threatening bird. At least two appeasement displays are used by the female stork in attempting to gain access to the male's courtship station, Balancing Posture and Gaping. Upon approaching the male, the female (of some species) assumes the Balancing Posture, holding her head low with her wings spread. This may be accompanied by bill Gaping, as if overheated, but the bird does not actually pant. As time passes without being driven away, she may close her wings but often continues to Gape with her head bowed. Soon after acceptance of a mate, a male Mycteria may engage in a Flying Around Display. He takes off from the nest and flies in a tight circle before returning to land next to his mate. Also during or soon after pair-formation, Bill Snaps become common. This behaviour, shown by both sexes in some species, involves a quick lift of the head to bring the bill to slightly above horizontal, followed by a snap of the

Figure 26. Asian Openbill Stork. The Up-Down Display at a nest with large young (south-central Thailand). bill. The display is used when levels of hostility are still high among the newly paired birds. A stork may infrequently give a bill snap when taking flight. During this display, the Pre-flight Snap, the bird bends forward and low, swinging its head between the legs. It then gives a single loud snap with its bill followed by a normal take off. Another display given before flight is the Erect Gape. The bird stands erect, with neck extended almost vertically and bill held slightly below the horizontal. Sometimes a low call is given. The Erect Gape may also be followed by a Bill Snap. The Flap-Dash display —probably derived from foraging

24 Courtship and Reproduction

A Nest-Covering Display has been observed in several species. The parent bird hides its eggs or young under partially opened wings.

Figure 27. A female Greater Adjutant Stork preening near her nest (northeastern India). behaviours —is performed on the ground or in shallow water by the Saddlebill, Blacknecked and Jabiru Storks. A bird, usually the male, flapping its wings, runs several metres away from its companion and then runs back again. Following pair-formation, mates greet each other with an Up-Down Display, which is performed by all storks but differs among species. In the openbills and wood storks, the bird lifts its head vertically, and, with its bill gaping, it may utter short hissing or grunting vocalizations, while lowering the bill toward the floor of the nest-platform. Depending on species, during the head-up part of the display, the bird may snap the bill once or twice or repeatedly, or it may not snap its bill at all. In the Blacknecked Stork, the bill is not raised, but the wings are spread and fluttered and the bill is clattered rapidly. Other species show varying degrees of head movements and clattering during the Up-Down. The most extreme is reached in the White and Oriental White Storks, in which the head is thrown all the way back toward the tail and the bill-clattering is loud and prolonged. This is, perhaps, one of the most widely known of all stork behaviours. Up-Down Displays continue throughout the nesting season. Copulation occurs at or near the nest-site with no precopulatory display. Characteristic of storks is for the male to clatter its bill or rattle it against that of the female during copulation. In storks, this behaviour is called Copulation Clattering.

COURTSHIP IN IBISES AND SPOONBILLS The courtship displays are more poorly known among ibises and spoonbills than in storks. Many species have never been studied at all. None the less, from the information available, a number of displays can be identified. Certainly, additional studies on these species are needed. As in storks, the males of colonial ibis and spoonbill species, and those that must pair each year, display for females. Upon choosing a display site, ibises and spoonbills tend to remain on it almost continuously. In fact, Display Standing may be considered the initial display in courtship. Display Standing can develop into Display Sleeping. In imitation of sleeping, the bird places its bill under its back feathers, although its eyes remain open. This behaviour appears to be quite contagious among neighbouring birds. It also occurs as a pause in active display, serving no doubt to maintain a bird's continued claim to a display- or nestsite. Species that remain paired throughout the year perform displays that serve to reinforce the pair bond, and perhaps to coordinate breeding readiness. Pairs display together with simple behaviours such as Standing, Head-Shaking, Stick-Shaking and allopreening. Ibises can be quite noisy when nesting, even those that are rather silent for most of the year. Calls differ among species from whistles to caws; they also may make noise with their wings when landing, taking off or flying around. Ibises and spoonbills use calls for contact, to communicate with mates, during copulation, and in agonistic situations. Some species, such as the Barefaced Ibis, perform duets (Luthin 1983a). The active solicitation behaviour of unmated males, in species so far studied, consists primarily of three displays: Display Preening (Front and Rear), Display Shaking, and Bill Popping. Display Preen (Front) and Display Preen (Rear) occur when the bird appears to use its bill-tip to preen its primary feathers (Front) or to preen over and behind the wings (Rear). The Display Shake is given by a bird standing erect and holding its wings loosely. It then shakes the wings alternately up and down. The Bill Popping display occurs as the bird snaps its gaped bill shut, usually without sound, although at times accompanied by a quiet pop. In African Spoonbills, this display is given as the bill is raised to the horizontal or slightly above. In the ibises so far studied, it is given with the head extended and bill downward. Ibises sometimes also grasp a twig and shake it briefly. Bill Popping is very reminiscent of the Snap Display of herons and also is reminiscent of the Bill Snap of storks. The solicitation displays may be given in any direction, not necessarily in the direction of a female, and sometimes toward neighbouring pairs. In some ibises, Bill Popping may be truncated into a more moderate form in which the neck is not extended. In ibises so far studied, the three

Courtship and Reproduction 25

displays are often used together but not in any standard sequence. In the African, Royal and Yellowbilled Spoonbills, these three displays are given in succession: Display Shake/Display Preen/Bill Popping. Another display, observed primarily in courting female spoonbills, is the Display Flight. The bird takes to the air, flying in a low circle or even a figure-of-eight while calling. It then lands near its point of origin, which is usually near an unmated male. This display is similar to the Flying Around of the storks. The initial phase of pair-formation involves considerable aggressiveness by the male toward the intruding female. As a result, many of the initial behaviour patterns of courting male ibises and spoonbills have components of aggression or fear. When a bird is alarmed, the Erect Posture is assumed. The bird stands upright, feathers sleeked, bill down, peering at the disturbing object. This behaviour is similar to the Arching display of storks and may precede, but does not necessarily lead to, escape behaviour. The basic aggressive behaviour of ibises and spoonbills is the Sparring Display, in which the bird assumes a horizontal posture and moves toward or lunges at an intruder using a bill thrust. The Sparring bill thrust may or may not make contact. An overt fight may follow. Two birds engaged in ritualized Sparring hold their feathers erect and bills gaped, Figure 28. Asian Openbill Storks. High water levels prevented breeding at Bharatpur, India.

while each reaches forward and snaps its bill towards the antagonist; it does not actually make contact. This behaviour is returned by the opponent, also without contact, and a stab-and-counterstab Sparring match commences, with birds alternating ritualized attacks. The aerial modification of Sparring is the Supplanting display, in which a bird flies at another bird, with its neck extended and bill gaped. The attacker slows its flight with its wings, and attempts to land on the opponent or, more likely, on the perch vacated by the displaced victim. One of the most common activities of some species of ibises is appearing to rub their head on their back. The Head Rub is ritualized into a mutual appeasement display and also an advertising display. The bird places the side of its head on its shoulders, bill pointing upwards. The bird may then turn it slightly to vertical or even to the horizontal on the other side, in the rubbing motion, or it may make only a quick touch of the nape to the upper back. The female ibis uses this display prior to attempting to enter the male's display area, and the male may use it with other attraction displays. The origin of this display is clearly the similar comfort movement. In some species it is reminiscent of aspects of the Up-Down display of storks. In its vertical form, it is similar to the Stretch Display of herons. Upon actually attempting to enter the male's display area, the female assumes a Bowing Display, in which she approaches the male keeping her body and head low, feathers sleeked back. She turns her head away from the

26 Courtship and Reproduction

male, exposing the side of her head to him. The male generally reponds with a Sparring display and even an overt attack, in response to which the female may flee but does not fight back. The male responds initially by aggressive actions and eventually by acceptance of the female at the nest-site. This behaviour is identical in function and similar in appearance to the stork's Balancing Posture, except that the wings are not opened. The aggressive phase of pair-formation ends as the male accepts a female. Bowing Displays continue, and when one of the pair returns to the nest, a Greeting display is given. With head feathers erect (if present) and bill gaped upwards, the bird calls and may flap its wings. In spoonbills the Greeting is given with the bill upward, whereas in ibises the head and bill are slowly brought downward to the vertical in a bowing motion, before the vocalization. Especially in spoonbills the downward component is recognizable as a distinct display, Head Shaking. Bending forward simultaneously with bills downward and closed, the birds shake their heads in a 'no' motion while lowering them to the nest. Some ibises also Head Shake. The birds' necks may overlap, the taller male usually being on the top. The Greeting displays appear similar to the Up-Down of storks. Stick Shaking may also occur. Bending forward with bills pointed down, the birds grasp nest material and shake their heads rapidly from side to side. Again, necks may cross and overlap, with the male on top. This behaviour is similar to Head Shaking, except for the involvement of twigs. Another mutual display of paired birds is the Head Quiver. Standing upright with head feathers erect, a bird twitches its head several times. Generally the display is performed by both sexes in succession or simultaneously. Following the acceptance of a mate, pair-formation is completed through copulation. Allopreening continues as do Head Shaking and other Greetings. During the Copulation Display, the male climbs on the back of the nearly horizontal female, and brings his cloaca into contact with hers, while flapping his wings for balance. In the process, the male may also perform a bill-shaking behaviour in which he grasps the female's head or bill or holds his bill alongside her neck while he shakes his head or nibbles at her feathers. A later defensive posture is the Nest-covering Display. A parent bird erects its feathers, orients toward the threat, and droops its wings down and over the nest. This display is clearly homologous to the behaviour of the same name in storks. Birds of a pair often engage in preening each other (allopreening). They especially nibble the feathers of the partners head and neck. Throughout nesting the birds commonly occupy themselves with preening, head-rubbing and shaking. Although these are functional maintenance behaviours, they probably also serve a display purpose, helping to maintain the pair-bond. Figure 29. A Eurasian Spoonbill with well grown young (Bharatpur, India). Figure 30. Young Asian Openbill Storks leave the nest early (Bharatpur, India),

Courtship and Reproduction 27

NESTING After courtship and pair-formation, nesting begins. Depending on the species and the habitat, nests may be made out of sticks, reeds or vines, often with green material added. Nestbuilding is generally the province of both sexes. The male is largely responsible for gathering the sticks. The female accepts the material. The sticks are then inserted by the female into the developing nest bowl by a behaviour called 'tremble-shoving', holding the stick by an end and wiggling it into the nest matrix. Behaviours during nest-building and maintenance may involve the real or feigned picking, grasping or presenting twigs to the female. At this time, appeasement and pair-formation displays become less frequent and less intense. The eggs are laid a day or more apart, and incubation usually begins with the first or second egg. The members of the pair alternate incubation duties. The larger male is often in attendance at the nest for much of the day. Young hatch asynchronously and are unable to stand from the first ('altricial young'). The bills of hatchlings are short, straight and rounded, even in ibises and spoonbills. The typical shapes begin to develop soon thereafter. Young are fed by both adults on regurgitated food. Storks regurgitate food onto the floor of the nest and their nestlings pick it up, whereas nestling ibises and spoonbills insert their entire head and bill down the parent's gullet to receive food. During the first few weeks after hatching, one adult remains at the nest at all times, as the parents alternate foraging trips. Storks also bring water to nestlings. When the young are further developed, both adults leave the nest to forage at the same time. Nestlings respond to disturbance with threat and alarm displays. Storks respond with a Nestling Bristle and may give Forward Displays. Ibises and spoonbills assume an Erect Posture. Nestlings solicit food from their parents using a Begging Display. Facing the parent while resting on its tarsi, the nestling raises its head and bill, bobs them up and down, flaps its wings, and gives a begging call. As the nestlings grow they may give the display while standing or— especially in the case of the smaller, more active ibises and spoonbills—while running or even flying after the parent. Larger young may catch their parent, and placing a wing over the parent's shoulder, pin him or her down while they force feed from the gullet. Large young spend considerable time flapping their wings. In some colonial species, it appears that young leave their nests to form groups, called creches. The young can fly long before they depart from the colony. They usually return to near their nests to be fed by the parents, which they appear to recognize by call and sight. Fledging periods vary widely, according to the size of the species, from less than a Figure 31. Storm's Stork. A unique photo of an adult standing on an active nest; only two nests of this species have ever been found (in Thailand and Sumatra). Photo by courtesy of the late Seub Nakhasathien. Figure 32.

An Asian Openbill Stork in Bharatpur, India.

28 Courtship and Reproduction

month in the smaller ibises to more than 3 months in the larger storks. Although for most species, parental care ends soon after fledging, the young of some species of ibises —those that appear to maintain long-term pair-bonds — continue to use the family's home range for a time, and are even fed there by the parents.

Changes to display names Original name

New name

Storks1

Forward Threat Aerial Clattering Threat Forward Clattering Threat Snap Display Anxiety Stretch Nestling Defence Display

Forward Display Aerial Clattering Forward Clattering Bill Snap Arching Display Nestling Bristle

Spoonbills and ibises2

Alarm Posture Appeasement Display Flight Intention

Erect Posture Bowing Display Pre-flight Tipping

1 2

Used in Kahl 1966a, 1972a-d, 1973b. UsedinKahl 1983, 1988a.

Figure 33. A captive Oriental White Stork at Vogel Park, Walsrode, Germany. Figure 34. African Spoonbills Sparring and nest building on a rocky island on Lake Turkana, Kenya.

Feeding Behaviour and Ecology

s

torks, ibises and spoonbills are among the more typical of the 'wading' birds, species that characteristically feed while walking about in shallow water. How these birds behave while feeding and what they can catch to eat are constrained by a general adaptive plan, millions of years old. That design includes long legs, long necks and long bills, all adaptations for feeding while walking through shallow water. Because of this body plan, most species of storks, ibises, and spoonbills are highly aquatic, characteristically occurring in and around shallow water. The wood storks and the openbill storks epitomize a near total dependency on water and are found only in very specific types of aquatic habitats. However, most species can also feed on land should compelling suitable food become available there. Some, such as Abdim's Stork and the Waldrapp, Bald and Hadada Ibises, are more terrestrial than aquatic. They feed mostly on insects in dry grasslands and farm fields. Yet they retain the primordial body design, modified by somewhat shorter legs, necks and bills. All species are both liberated by and constrained by their wading bird heritage. Indeed, it is instructive in understanding the ecology of these species to consider what foraging opportunities can be possible for large birds built according to these general specifications. The various behaviours used by storks, ibises, and spoonbills in their foraging activities can be delineated and defined (Kushlan 1978a). Recognizable similarities in most behaviours among the species allow use of the same terminology in both families. In this book we capitalize the named feeding behaviours in order to allow the reader to identify them in text. We also capitalize, as is the custom, courtship behaviours. However, unlike courtship behaviours, most feeding activities are not ritualized displays. The number of behaviours identified as being used in feeding is surprisingly extensive. No species is known to feed in all the ways so far documented. Some species do exhibit quite a variety. The Roseate Spoonbill, for example, has been documented as being able to use eight different feeding techniques. The most universal feeding behaviour is Standing. That is, a bird simply stands in one place and captures prey as it is encountered. Standing can be used on land or in the water, or even while mobile— African Openbill Storks Stand on the backs of moving hippos (Kahl 1971c). Typically, Standing foraging involves capturing prey that is on the ground or in the water. Two special types of Standing behaviour have been recognized when prey occur other than in the water. When Standing Flycatching, a stationary bird catches prey that is airborne. When Gleaning, a Standing bird catches prey that is on an emergent plant or some other above-water object. Another variation, which occurs while Standing, is Flipping. A bird turns over an object to catch what is underneath. In Foot Stirring, a bird vibrates its foot or leg, to stir up potential prey. No species, or even an individual, forages solely by Standing, because prey-capturing opportunities are soon depleted at most feeding sites. To move from place to place, birds intersperse Standing with more mobile behaviours. The most widespread behaviour is Walking Slowly, in which a bird walks or wades at slow speed usually continuing to seek prey. Faster passage is achieved by Walking Quickly and by Running. Walking Slowly can be recognized by the bird's slow rate of progress, as slow as a step or two per second. Running, on the other hand, can be recognized by its rapid rate of progress, such that the bird occasionally needs to use its wings for balance. Of course, these classifications are relative and not totally distinctive.

30 Feeding Behaviour and Ecology

The relative amount of Standing and Walking depends on the species and the situation. The Shoebill typically Stands in one place for long periods at a time, whereas ibises and spoonbills seem always on the move. The usual prey-capturing technique involves a Bill Grab in which the bill grasps a prey item, using the mandibles as pincers. Unlike herons, which strike from a recoiled neck (Hancock and Kushlan 1984), storks and ibises Grab prey without a darting stroke. The Shoebill is unusual in that its Grab is accompanied by a Body Lunge, in which the head and trunk are thrown at the prey. The bird often ends up partially submerged (Guillet 1979). Some food can be captured in a more leisurely manner by Pecking. A bird simply uses its bill to pick up an item from the water or substrate. Wood storks use a highly specialized prey capturing behaviour, the Bill-snap Reflex. Upon contact between a prey item and the open bill, the two mandibles close refiexively at a speed averaging 25 ms in the American Wood Stork (Kahl and Peacock 1963). Several special techniques have developed to allow birds to forage without using their eyes. The most widespread is Probing, in which a bird quickly and repeatedly moves its bill tip into and out of the water or substrate. This is the characteristic non-visual foraging method of ibises, which have innervated bill tips that register contact with a prey item. It is also used by many storks, including openbills, typical storks and giant storks. In ibises several types of Probing have been distinguished according to water depth and rapidity of the probe (Kushlan 1977a). One can recognize Shallow Probing (the bill is inserted fewer than 2 cm

into the substrate), Deep Probing, Standing Probing, Walking Probing and Multi-probing (Walking interrupted by Standing Probing). Groping is a similar behaviour in which a bird places and moves its open bill about in the water. It is the characteristic component of the feeding repertoire of the wood storks. Head Sweeping (Head Swinging of Kushlan 1978a; Slow Sweeping of Vestjens 1975b) is the typical tactile feeding behaviour of spoonbills but is also used by ibises. Vestjens (1975b) showed that the details of Head Sweeping differed in the two Australian spoonbills. It is usually accomplished while Walking Slowly. When performed while standing still and searching a site with short sweeps, it is called Intensive Sweeping. Forward Ploughing takes place when a bird walks with its bill under water and projecting forward, rather than swinging it sideways. Dragging is similar except that the bill is dragged through the water at the bird's side. Wings are also used during foraging. Feeding while actually in flight has not been recorded in storks, ibises or spoonbills, although it can be common in some species of herons. Feeding birds do take short repositioning flights, however. In Hopping, a bird flies up and alights nearby, even adjacent to its former feeding spot. In Wing Flicking, a bird quickly partially extends and retracts its wing; whereas in Openwing Feeding, a bird completely extends and retracts one wing at a time. These behaviours appear to serve to frighten prey, especially fishes, into movement. Several behaviours tend to be strung together in various ways to create a feeding sequence. The wood storks, for example, typically forage by alternating Standing with

Figure 35.

Figure 36.

Immature Painted Stork (Bharatpur, India).

An Oriental White Ibis at Bharatpur, India.

Feeding Behaviour and Ecology

31

Figure 37. African Openbill Storks feeding on snails in a lily pond (Tana River, Kenya). Walking Slowly, while Groping with their open bill and Foot Stirring. At the same time they Wing Flick or Openwing Feed on the same side as Foot Stirring. The mix of these actions increase the activity of potential prey, and therefore the chances of its swimming into the bird's Groping bill (Kahl 1964, 1972b). The Blacknecked and Saddlebill storks Walk Slowly or Run while Probing vertically in the water with each step. The American White Ibis is typical of a bird with less restricted sequencing (Kushlan 1977a). It will Walk Slowly while Probing; then, stopping at the edge of the water it may Peck at the mud surface or attempt to excavate a crayfish from its burrow by Standing Deep Probing. It then might move to deep water where it will Head Swing like a spoonbill. Once a bird captures a prey item, it must be 'handled'. Small prey can be consumed immediately. Ibises can swallow such items nearly instantaneously by closing their bill tips while slightly pushing their head forward. Large prey have to be softened or broken into smaller pieces. A large food item, dead or alive, can be handled by Biting (i.e. mandibulating the prey) or by Stabbing (i.e. repeatedly poking and piercing it with the bill). The Australian White Ibis holds mussels against a rock with its feet whilst stabbing them with its bilL Handling time goes up exponentially

as prey size increases, soon reaching impractical limits such that a bird might forgo eating a captured prey item in favour of resuming the search for a new one (Kushlan 1979c). Divergence among the several lineages of storks, ibises and spoonbills is reflected, indeed in large part influenced, by their feeding behaviours and accompanying morphological features. One major evolutionary and ecological division is among species that feed primarily visually, by seeing their prospective prey before catching it, and species that feed primarily tactilely, by feeling prospective prey with their bill (Kushlan 1978a). It has been shown that the American Wood Stork feeds just as effectively without use of its eyes as with them (Kahl 1964). The primarily visual foragers, including the typical storks and adjutants, generally Grab the prey items they see. The primarily tactile foragers, including the ibises, wood storks, openbills and spoonbills, typically use Probing, Groping and Head Sweeping. Despite these overall adaptive trends, most species have the flexibility to use both visual and tactile methods when the appropriate occasion presents itself. Given the correlations between feeding behaviour and morphology, much of the feeding story of each species can be told by examining their primary feeding apparatus, their bill. The divergence between ibises and spoonbills is essentially one of billshape. The ibises have thin downturned bills

32 Feeding Behaviour and Ecology

Figure 38.

A Hadada Ibis feeding in Kenya.

Figure 39. America.

A Buff-necked

Ibis in Argentina, South

whereas the spoonbills have straighter, flattened bills, each representing a basic morphological type for tactile aquatic foraging. Ibises that feed in terrestrial habitats, such as the Green Ibis, have shorter bills than those that feed in water, such as the American White Ibis. The storks have an even greater diversity of billshapes. Ranging from the enormous Grabbing bill of the Jabiru Stork, to the specialized bills of the openbill storks and the ShoebilL Openbill storks probe for snails, capturing and opening them in their bill-tips. The Shoebill feeds visually, Grabbing its very large prey. The wood storks have slightly down turned bills, reminiscent of the ibises. Another peculiar morphological correlation with feeding behaviour is the colour of the feet of the American Wood Stork. Its feet are pinkish, contrasting with its darkish legs. This colour possibly serves to enhance the effectiveness of Foot Stirring while feeding. Some species are solitary foragers. They often use simple feeding behaviours such as Standing or Walking Slowly, stalking prey in ways that are not enhanced by a crowd. The Shoebill is the archetypical example of a solitary species. Others are solitary at one time of year and more social at another. The Spotbreasted and Hadada Ibises are two examples of species that nest solitarily but feed in groups when not breeding. The great majority of species are capable of feeding in the company of other birds. Generally speaking most storks, ibises and spoonbills feed at least some of the time in social associations, called feeding aggregations, which form at sites of high food availability. Others show the higher level social organization typical of flocking. A correlation between nesting and feeding dispersion is too great to be a coincidence. Colonial nesting and communal roosting provide exceptional opportunities for feeding together (Krebs 1978). Birds nesting in the same place can take advantage of foraging opportunities found by other birds in a process called social facilitation. Either by following birds flying to the feeding grounds or by seeing other birds feeding, wading birds are able to identify promising feeding sites and join the aggregation of foraging birds assembled there. Many of the communally foraging species are white, a colour that has been demonstrated to attract wading birds to a feeding site (Kushlan 1977e). This raises an interesting question: once a bird finds a profitable foraging site, why should it advertise the place to other birds? At least two non-exclusive possibilities present themselves. The first relates to the character of the feeding sites used by most of these species. These sites, such as drying ponds and shallow wetlands, are ephemeral in that they are suitable for feeding only for short periods. Also in such cases the amount of food present is more than one bird can eat in the short time it will be available. Thus there may be no harm in inviting other birds to join the feast. The second possibility is that a foraging bird actually benefits from having more birds at its feeding site. One benefit may be in hiding from predators within a large aggregation of other birds. More directly, the prey present are disturbed by the feeding activity, which increases the probability of one bumping into a waiting bill.

Feeding Behaviour and Ecology

This is a form of mutualism, a form of interaction between individuals or species from which both benefit. A number of instances of commensalism (an interaction in which one animal benefits and the other is unaffected) have also been reported in storks, ibises and spoonbills. Generally a visually foraging species follows a tactilely foraging species, benefiting from the prey the latter stirs up (Kushlan 1978a). The subtle interactions among species, as noted above, may have lead to the social signalling function of the white plumage found in many species of storks, ibises and spoonbills (Kushlan 1978a). Feeding aggregations also include other aquatic birds, especially herons and cormorants, though aggregative foraging carries with it certain risks, especially that of losing a captured prey item to a robber. Small birds can fly over and snatch a prey item from the bill of an ibis or stork. Larger wading birds also steal from smaller wading birds (Kushian 1978c, Kushlan et al. 1985). Especially at risk are prey that take time for handling. Some species that feed in family groups may tend to feed in more permanently available feeding sites. Species that remain in family groups, represented by the Blacknecked and Saddlebill storks and several species of ibises, also use the same foraging area for large portions of the year. Members of the pair forage within sight or hearing of one another but only join aggregations if one happens to form in their home range. Most storks, ibises and spoonbills feed primarily during

33

daylight. However, some — especially among the tactile feeders —will forage at night, and even visual feeders can secure prey on bright moonlit nights. In general, the smaller species begin feeding at sunrise. Storks often do not leave the colony site for feeding grounds until later in the day. This is because they travel long distances, not by flapping flight, but by soaring high on thermals and then gliding a long way to the next thermal. Such species leave the colony or roost when the ground heats up in mid-morning, forming the thermals. In certain species, such as the American Wood Stork, some individuals may spend the night at the foraging ground, returning to the colony the next morning (Kahl 1964). Outside the nesting season, storks, ibises and spoonbills spend much of the middle of the day resting near the feeding ground. Storks, ibises and spoonbills are overwhelmingly carnivorous, each species eating various sorts of fish and amphibians, snails, crustaceans, insects and other aquatic invertebrates. The bulk of the diet of a population may consist of very few species of prey. The most specialized group is the openbill storks, whose peculiar bill allows them to extract aquatic snails from their shells (Kahl 1971c). Thus snails make up almost all of their diet. With their Probing bill, ibises are especially adept at capturing small invertebrates, such as prawns, crayfish and aquatic insect larvae that live on the bottom or on aquatic plants. Similarly, they can probe into loose soil or grass to extract terrestrial insects and their larva. Faster-swimming

Figure 40. White Storks feeding on drained carp ponds in Israel.

34 Feeding Behaviour and Ecology

prey, such as fish, are also caught but usually only consumed in numbers when they are available in high densities. With their Head Sweeping, spoonbills take prey from the water column or in submersed plants, generally catching fish, insects and prawns. The wood storks are restricted in diet by their tactile foraging techniques. Their food is primarily fish that are found in rather dense concentrations (Kahl 1964, Kushlan 1979b). Several groups depend heavily on scavenging for food for most of the year. The adjutants are specialized scavengers on large prey killed by predators, and the Sacred Ibis often scavenges at kills and dump sites. Scavengers revert largely to living food when feeding their nestlings. The typical storks have the most generalized diet, being able to select their prey visually. In addition to the expected fish and amphibians, they also consume small mammals, insects and reptiles, depending on what is available. Prey availability is, in fact, a key to understanding the foraging ecology of these species (Kahl 1964, Kushlan 1978a, 1981, 1986b). Many of the fundamental adaptations of storks, ibises and spoonbills are related to accommodating to changing patterns of prey availability. These include seasonal migration, intraregional population shifts, timing of the breeding season, colony site selection, foraging habitat use, and food consumption. Those species that have relatively specialized bill shapes and feeding behaviours often depend on very specific circumstances of prey availability, generally related to seasonal patterns of rainfall and water level fluctuations (Kushlan 1978a, 1981, 1989a). Aquatic ibises require shallow water; terrestrial ibises require ground conditions suitable for probing. Wood storks require high prey densities for efficient foraging, and their nesting success depends on the continuous availability for several months of shallow, ephemeral foraging sites. The less specialized typical storks can. forage in either wetlands or fields and depend on seasonal changes in food availability. The migratory patterns of many species are correlated with changing food supplies. The White Stork depends on frogs during the nesting season in Europe and insects —

especially plague locusts—in the winter in Africa. The survival of storks during the non-breeding season has been shown to depend on rainfall patterns and therefore food availability on the African wintering grounds. Fires can increase the prey available to upland foraging birds. Several species, such as the Marabou, Maguari, White and Woollynecked Storks and the Sacred and Bald Ibises, characteristically forage on recently burned land, even following the fires closely to catch insects and other prey exposed by the fire. Prey availability is especially critical for the tactile foragers (Kahl 1964, Kushlan 1979b,c). Their ultimate diet is the result of the interaction of their size, bill shape, feeding behaviour, sizes and types of prey available, and prey abundance. As a result the American White Ibis and American Wood Stork, feeding in the same area, take entirely different diets, owing to differences in where they can feed and what they can catch and handle. Food is required for the maintenance of adults and for the raising of young; the latter can as much as double the daily energy needs of a pair of adult wading birds. All the storks, ibises and spoonbills are relatively large birds and have substantial daily energy requirements for growth and maintenance (Kahl 1962, 1964, 1966a, Kushlan 1977b,c, 1978a). The daily energy needs of an American White Ibis has been calculated to be about 200 g of food, whereas the larger Marabou Stork requires 720 g. Nestlings increase this requirement. The rigours of delivering sufficient food to nestlings determines much about the seasonality of breeding and feeding habitat selection exercised by these birds. Nesting failure is not uncommon, generally brought about by changes in rainfall patterns, which affect food availability. Because the adults begin incubating their eggs as soon as the first is laid rather than waiting for the entire clutch, the young hatch at different times, i.e. asynchronously. As a result the older chick gets a head start on its siblings and may survive food shortages when the younger birds perish, thus permitting some production under difficult conditions.

Photographs. The photographs appearing in the above sections were kindly supplied by M. P. Kahl (Figures 1-27), J. A. Hancock (Figures 28-30, 32-40) and the late Seub Nakhasathien (Figure 31).

American Wood Stork Mycteria americana Linnaeus Mycteria americana Linnaeus, 1758, Syst. Nat., ed. 10, p. 140: Brazil Other names: Wood Ibis, Flinthead, Ironhead, Gourdhead, Gannet, Preacher, Spanish Buzzard, Colorado Turkey, Wood-Pelican (English); Gaban, Gaban heucito, Garzon, Coco, Cabeca seca, Cigiienon, Doroteo, Sargento, Ibis del Monte, Alcatraz, Sowewies, Negroscopes, Cabeza de piedra, Galletan, Cayama, Galan sin ventura, Faisan, Cigliena cabeza pelada, Tuyuyu, Fraile (Spanish); Passarao, Cabeca seca (Portuguese); Nengrekopoe, Amerikaanse Nimmerzat, Kaalkopooievaar (Dutch)

IDENTIFICATION The American Wood Stork is a large (85-113 cm tall) white-plumaged, wading bird with black in the wings and tail, a naked blackish head and neck, and a rather heavy, downcurved bill. In its adult plumage, the head and most of the neck are featherless and covered with scaly, almost plate-like skin, which forms a shield-shaped cap on the top of the head. The face and neck skin is blackish grey except the cap, which has a lighter bronze tinge to it, and the nape behind the cap, which is black. The bill is stout at its base and initially straight before tapering to a downcurved tip; it is noticeably thicker and shorter than the bills of the Old World species of wood stork. The bill is dusky grey with tan or reddish brown sides. Flight feathers, some coverts, and tail feathers are black with a greenish or purplish sheen. The legs are black, but the slightly webbed toes are pink. The iris is dark greyish brown. Males and females have similar plumages, but the male is larger (c. 3kg) than the female (2 kg); females also have a smaller head and thinner bill. A complete moult occurs before the nesting season. The breeding plumage features a pinkish orange tinge on the underlining of the wings and pure white plume-like undertail feathers (coverts), which extend beyond tail in flight. During courtship, the bill is shiny and appears polished or oiled. The feet and lower legs turn rose; this is probably due to vascularization, as the feet pale rapidly after death (pers. obs. M.P.K.). The legs often appear white, owing to the deposition of excreta for cooling when it is hot at the nest (Kahl 1963b). Nestlings are covered with a sparse grey down at hatching and weigh about 62 g. At about 10 days, the first-down is replaced by a thick woolly white down, excepting the forehead, lores, chin, and anterior throat, the skin of which are basically naked. By the sixth and seventh week the downy plumage turns smoky grey on the head and neck, darker on the top of head and paler below; there is a white V-shaped area on forehead at about time of fledging. The bill is pale yellow; the legs and feet are pale fleshcoloured; the iris is brown. Growth and development of nestings is described more fully in Kahl (1962). The juvenile plumage is much like the adult's but duller, and with feathering on the head and neck. The base colour is a dull, greyish white, with some wing coverts tipped with brown. The back of the head and upper neck are scantily feathered with coarse, smoky and buff hair-like feathers that are darkest and longest on the top of the head. The tail and flight feathers are brownish-black, with less sheen than in adults. As feathers are lost on the head, the skin darkens and begins to corrugate,

American Wood Stork

juveniles having less defined scaling on the naked skin. The bill is initially pale straw-yellow and contrasts strongly with the dark feathers of the head; the bill and legs darken gradually with age. The bird remains in juvenile plumage for the rest of the first year, with subadult plumage (feathering on neck, bill somewhat pale) emerging in the fall moult; some feathering remains on the neck into the third year. Nesting first occurs in full adult plumage in their fourth year (J.G. Ogden pers. comm. 1990). The adult American Wood Stork is generally silent, except during displays at the nest. In courtship, adults snap and rattle their bills, clatter their bills together when copulating, and give a rasping, hissing 'fizz' call during nest relief. Especially in the nesting season, their wings make a loud 'Woofing' sound during flight and landing. Large flying young give a flight call that is a trisyllabic goose-like sound. Nestlings utter a loud Begging Call that is a highpitched, rasping, nasal braying 'Nyah-nyah', which gets lower-pitched and hoarser with age. After 2 weeks, the call becomes a loud nasal 'Wonk' (J.A. Rodgers pers. comm. 1988). When distressed (e.g. cold, fearful, very hungry) nestlings give a short, repeated 'hiccoughing' call (pers. obs. M.P.K.). A large bird, the American Wood Stork flies with slow audible wing beats; the flapping rate averages 177/min (w = 44) (Kahl 1971b). Upon taking flight the neck is stretched out and down and legs left to dangle, until both can be fully extended for flight. In flapping flight, it alter-

37

nates 7-10 flaps with a short glide. It commonly uses thermals to rise to great heights, from which it glides until it picks up the next thermal. Thermal soaring and gliding are normally used for all long-distance travel, with low-level flapping flight used only for short journeys or when thermals are unavailable. Because thermals are dependent on the heating of the earth's surface by the sun, soaring/gliding flight is restricted to the warmer times of the day, generally between 09.00 and 16.00 h (Kahl 1964). Near the Corkscrew colony, soaring birds have been followed aloft in a small plane to 1525m (Browder 1984). Especially when descending from high altitudes, they may perform remarkable aerial acrobatics, diving steeply at high speeds and flipping over and over (pers. obs. M.P.K.). The rush of air through their wings sounds like a high-flying jet plane. Days with strong, gusty winds seem to stimulate these aerial high jinks. Distinguishable from most other wading birds by its height, stocky build, and downcurved bill, the American Wood Stork is larger than ibises and smaller than the Jabiru Stork which has all white wings and tail and massive upturned bill. The Maguari Stork has a similar flight pattern and size, but has a fully feathered white neck and head and a less conspicuous, forked, black tail and conspicuous white undertail-coverts. In flight, entirely black flight feathers are diagnostic in contrast to the black-tipped primaries of the smaller American White Ibis. In soaring flight, the American Wood Stork looks very similar to the American White Pelican (Pelecanus erythrorhynchos) but is identifiable by its extended neck and legs. In tropical America, it might possibly be confused at a distance in flight with a King Vulture (Sarcoramphuspapa); again the long legs and neck of the stork are diagnostic.

DISTRIBUTION AND POPULATION The nesting range includes the southern United States (primarily in Georgia and Florida; isolated breeding records have occurred further north and west (Hamel 1977, J.L. Dusi and R.T. Dusi 1968a), southward discontinuously through Cuba, Hispaniola, and both coasts of Mexico and Central America (although confirmation of breeding appears to be lacking for Guatemala and El Salvador). The species also occurs throughout the lowlands of much of South America. West of the Andes, it ranges to western Ecuador and Peru. East of the Andes, it extends from the Magdalena Valley, northern Colombia, south through Amazonas, to eastern Peru and Tarapaca in northern Chile, eastward through Bolivia to Cordoba, Santa Fe and northern Buenos Aires in Argentina, north through Uruguay, Brazil, and the Guianas to Venezuela (Palmer 1962:511512, Blake 1977:188). American Wood Storks frequently occur outside their usual breeding range, especially during post-breeding population movements. The extent of these vary from year to year. Basically, they can be divided into two classes: 'hardship movements', which are most extensive after nesting failure when ecological conditions in the breeding range may be poor; and 'dispersal', comprising mainly juveniles

38 American Wood Stork

from that season's reproduction. The usual northern dispersal limits extend from North Carolina through Tennessee to Arkansas. However, the actual limits of records reach northeast to New York, Massachusetts, Rhode Island, Maine, and southern New Brunswick, and north and northwest through southern Ontario, Michigan, Illinois, South Dakota, Montana, and Idaho. An extremely northern occurrence was recorded in northwestern British Columbia (in 1970). Most temperate populations undergo some migration seasonally. Northern-nesting birds from Georgia and northern Florida move south into central and southern Florida in the winter (Comer et al. 1987) and are present during the time of the initial nesting stages of local birds. In summer, storks that nested in southern Florida move northward to northern Florida and the coastal plain of Georgia and Alabama. Birds from eastern Mexico move north into Louisiana, Arkansas, and Texas in summer; birds from western Mexico move into southern California and southern Arizona. A similar movement probably occurs in southern South America with birds moving south to Chubut, Argentina, in summer and autumn (Hudson 1920, Yamasita and dePaula 1986). In general, they can be expected to avoid situations of extremely high water or drought, which make feeding difficult. The American Wood Stork is the most abundant and widespread of the three American storks, and the population status of this species is well-understood. During the late 1980s, between 150-1450 pairs have nested annually in southern Florida, and a total of 5000-6000 pairs throughout the entire southeastern United States (Kushlan and Frohring 1986, Ogden et al. 1987, J.C. Ogden, pers. comm. 1990). Several thousand storks are resident in Quintana Roo, Mexico, and colonies of 8000-10000 pairs occur in the Usumacinta Delta in Campeche, Mexico; between 1971 and 1979, several smaller colonies totalling about a thousand pairs were found on the west coast of Mexico (Knoder et al. 1980, Sprunt and Knoder 1980, Luthin 1987). In Quintana Roo, Mexico, 548 nests were counted by Cristina Ramo and Benjamin Busto (pers. comm. 1991). A colony of 3000 pairs was reported in Guanacaste Province, Costa Rica (M. Gochfeld persl comm. 1989); and smaller colonies have been found in Panama and Surinam (Spaans 1975b, Luthin 1984a). Large populations occur in the extensive freshwater marshes of South America, being common in the Pantanal of Brazil (Luthin 1987) and numbering nearly 6000 in the Llanos of Venezuela (Ramo and Busto 1984). An additional 3000 were censused on the Venezuelan coast. Large nesting colonies in southern Florida, particularly in the Everglades and the largest at Corkscrew, in the Big Cypress Swamp, once constituted the most abundant North American breeding segment, numbering about 1000012000 pairs in the 1930s (probably not 50000-75000 pairs as often reported) (Kushlan and Frohring 1986). There were still over 10000 pairs in Florida in 1960 (Kushlan and Frohring 1986, Ogden et al. 1987). However, changes in the hydrological conditions of the wetlands have resulted in a 75% decrease in southern Florida from 1967 to 1981-82. Population decreases have also occurred elsewhere. In

Belize, colonies known in the 1970s no longer are active, probably owing to disturbance (Luthin 1987). The species is considered to be threatened at the southern edge of its range, near Sao Paulo and Rio Grande do Sul, Brazil (Luthin 1987). Ogden et al. (1987) believe that the nesting population in the United States shifted northward into more but smaller colonies, expanded its range, and, perhaps, increased in size. However, previous censusing inaccuracies and omissions make such a shift, from south to north Florida very uncertain (J.A. Rodgers pers. comm. 1990). The Wood Stork is one of the few storks or ibises to have had population genetics studies completed. Stangel et al. (1990) found that there was considerable genetic similarity among birds nesting in different colony sites in Florida. ECOLOGY The habitat requirements of the American Wood Stork are quite restricted. It is primarily a bird of freshwater wetlands, including inland marshes, swamps (especially cypress swamps in the USA), lake-edge marshes and river bottoms, but also mangrove swamps and lagoons along the Neotropical coast. It feeds in shallow pools in inland and coastal wetlands. The wetlands used by American Wood Storks are characterized by seasonal fluctuation of surface water depth. In the rainy season, these wetlands flood, but in the dry season water levels fall, which increasingly restricts water to pools and ponds where fishes become highly concentrated. The American Wood Stork is highly gregarious, roosting and nesting in colonies and almost always feeding in small groups or in mixed aggregations with other species of wading birds. Interactions among birds in an aggregation are not common, but intraspecifically the storks use Forward Displays in defence of a feeding site or individual distance. At high intensities, the Wood Stork shakes its head from side to side during the display (Kahl 1972b). Such threats can progress to grabbing at an opponent and in bill duelling. On rare occasions, birds have been seen pirating food away from other species, such as the Great Egret (Egretta alba) (pers. obs. M.P.K.). The American Wood Stork feeds by tactile Groping (Kahl and Peacock 1963, Kahl 1964) in water usually less than 50cm deep. In trials on captives, birds temporarily deprived of frontal vision fed as efficiently as those with unaltered vision. It Stands or Walks slowly with its bill placed in the water and gaped 7-8 cm. It Foot Stirs by pumping the foot up and down on the substrate near the Groping bill (Rand 1956) and may Wing Flash on the same side as the pumping foot; it then repeats with the other foot and wing, sometimes taking a step or two and turning in a circle. While Walking, it will withdraw its bill from dense vegetation but keep it submerged in open water. The Foot stirring and Wing Flashing constitute a disturbance technique, and when potential prey touch the open bill, it closes within approximately 25 ms. When a fish is caught, the stork raises its head, bites at a large prey item, and then swallows. While most feeding is diurnal, the birds do also forage at night, and there is a fair bit of flight activity in and

American Wood Stork 39

out of breeding colonies on moonlit nights (pers. obs. M.P.K.). At various times many kinds of food have been reported, including fish, prawns, crayfish, frogs, young alligators, woodrats, young birds, crabs, snakes, turtles, and aquatic insects and their larvae. Early reports of vegetable matter were thought to be inadvertent inclusions, not eaten purposely; however, in the 1980s a number of dead nestlings and adults were found with impacted stomachs containing plant material (peat) (J.A. Rodgers pers. comm. 1988). Birds have even been reported eating cow and buffalo dung (Kahl 1964, J.A. Rodgers pers. comm. 1990), and during the drought of 1988 dead chicks had cow dung in their stomachs (M. Coulter pers. comm. 1990). Although a wide range of prey is taken, in any one location the bulk of the diet is limited to the few types and sizes of prey that its Groping technique can capture. In the Everglades, we found that storks ate only fish, with 72% of the biomass being composed of sunfish (Lepomis sp.), flagfish (Jordanella floridae), sailfin mollies (Poecilia latipinna) and marsh killifish (Fundulus confiuentus) (Ogden et al. 1976). Similarly, in Georgia, 96% of the biomass of food collected from nestlings consisted of sunfish, chubsuckers (Erymyzon sucetta), and redfin pickerel (Esox americanus Gmelin) (Coulter 1987b). The size of prey taken in Georgia was consistently larger than that taken in southern Florida (M. Coulter pers. comm. 1990). In Quintana Roo, Mexico, they ate only fishes, primarily cichlids (Ramo and Busto pers. comm. 1991). In general, the American Wood Stork's peculiar feeding technique results in its catching relatively large individuals of whatever fishes are the easiest to catch. In southern Florida, we were able to compare the prey taken to those available, and we found that the storks were able to capture selectively the species that dominated their diet, but were relatively poor at capturing other species, including the abundant but tiny mosquitofish (Gambusia qffinis). The energy requirements are well known (Kahl 1962, 1964). An adult requires approximately 450 kcal/day, which is supplied by about 520 g offish. A nestling requires about 16.5kg of food to fledge, so that an average stork family requires 200 kg of live food or 220 000 kcal per nesting season. A colony of 6000 nests requires over one million kilograms of fresh fish during the nesting season. The energy requirements of the more typical smaller colony are considerably less, of course. To forage effectively, the American Wood Stork requires extremely high densities offish, so in the heart of their range they nest during the dry season when falling water levels concentrate fish in relatively deeper pools within the marsh or swamp (Kahl 1963a, 1964, Kushlan 1976b). In the Everglades and Big Cypress areas of extreme south Florida, we found that they selected feeding locations where fish were being concentrated over sites with lesser densities of fish. The connections between falling water, feeding, and nesting are so precise that the timing of stork nesting is correlated with the speed at which water level falls in the dry season, and any appreciable rise of water level will result in higher nestling mortality or cause adult storks to abandon their

nesting attempt (Kahl 1964, Kushlan et al. 1985). As a result, nesting success depends on progressive drying for the entire 4-month nesting season. Similar relationships between nesting and rates of water-level decrease have been found elsewhere in Florida (E. S. Clark 1978a). And in that the American Wood Stork nests at similar stages of the local annual hydrological cycle in Central and South America (Thomas 1985), it is likely that nesting depends on falling water levels —and the concurrent increased food availability—throughout its range. To maintain suitable feeding opportunities, American Wood Storks may use many feeding sites during the course of the drying season, following falling water tables to a succession of suitable foraging sites (Ogden et al. 1978, Browder 1984, Kushlan et al. 1985). In southern Florida, we have followed Wood Storks that were travelling by soaring flight to feeding sites 140 km from their colony, although 70 km is the more usual limit; in Georgia the average distance flown was about 13km, and over 85% were within 20km (Bryan and Coulter 1987a,b). Storks tend to feed closer to the colony site when young are small, returning to the nest more frequently than later in the nesting cycle. Storks travel to nearer sites by flapping, and more distant sites by soaring and gliding. Thus, the nesting success of the American Wood Stork depends on finding and obtaining the appropriate quantity and quality of food, which in many cases is made available by fluctuating water levels (Kahl 1964, Kushlan et al. 1975, Browder 1978, 1984). The American Wood Stork is characteristically seen standing motionless, alone or with other waterbirds, in trees or on the ground and walks in an erect, 'dignified', sort of way (thereby the name Preacher). Its overall appearance may be summarized as stately but dull. During periods of high ambient temperature, storks actively excrete a dilute urine onto their legs, and the evaporation of this fluid acts to cool the blood in the legs, and— through circulation—in the body (Kahl 1963b). Such behaviour is seen most commonly during the breeding season, when birds are in the tops of trees for long periods. Parents bring water to their young in the nest, enabling them to cool themselves in the same manner. This cooling mechanism was first demonstrated in the American Wood Stork, but it has also been shown to occur in all other storks (as well as in New World vultures, anhingas and some cormorants, but not in the Shoebill). BREEDING The timing of the dry season differs in various parts of the American Wood Stork's range. In southern Florida, historically, they used to begin nesting mostly at the beginning of the dry season, in November-December, but more recently they often begin in January-March, or even, although often not successfully, as late as May. In 1988, nesting at Corkscrew Swamp extended into July because of unusually dry conditions. In most years, the rainy season begins in May or June and any remaining nesting is aborted as the adult storks are unable to find sufficiently concentrated food sup-

40 American Wood Stork

plies. In north Florida and Georgia, nesting occurs in the spring and early summer (Rodgers 1990) when water levels fall locally and the inhibiting effects of cold winter temperatures have passed. In Central and South America the birds nest during the season of decreasing water levels, beginning in February in the Yucatan, Mexico (Ramo and Busto in press). In Surinam, they nest from September to February (Spaans 1975b). They nest in wetlands, choosing isolated islands or clumps of trees, usually over water. In southern Florida, bald cypress (Taxodium distichwri) and red mangrove (Rhizophora mangle) are the principal species used. American Wood Storks nest in colonies, where they often choose nesting sites on the tops of trees, as high as 25-30 m in cypress trees and 10m in mangrove trees. In north Florida, they nest in low vegetation, often under the canopy (J.A. Rodgers pers. comm. 1988). Colony sites are traditionally reused year after year and are abandoned mainly in cases of disturbance or food failure. Counts of as many as 32-45 nests have been reported from single cypress trees (Phelps 1914, R.P. Allen pers. comm. 1959). The nest is a well constructed platform of sticks and vines, usually with fresh leaves and Spanish moss (Tillandsia usneoides) prominently included in the lining of the cup, perhaps for insulation (Rodgers et al. 1988). Nests vary from 45 to 90 cm in diameter and 10 to 15 cm deep. Courtship behaviour has been well studied (Kahl 1972b). Breeding is initiated as storks form daytime 'bachelor parties' at the future colony site. These clusters of unmated birds move about from tree to tree—with loud wingflapping and Bill-rattling fights—before settling down on a specific site. Such pre-pairing behaviour may last for weeks before nesting actually starts. Eventually, display sites are established by males and defended against both sexes by Aerial Clattering and Forward Clattering; bouts of Display Preening are frequent. If a mate is not obtained, males may abandon initial display sites in favour of new trees, especially when attracted by the mating activities of other pairs. Arching displays are given in response to disturbance, especially to potential predators below the nests. In most courtship displays American Wood Storks droop and spread their plume-like undertail-coverts, and bristle the feathers on their back. The principal advertising displays are: Display Preening, characterized by the stork's pulling the feathers of the nearly closed wing past the bill at a relatively high rate; Swaying Twig-Grasping; a subtle Flying Around display; and the Bill Snap. The female approaches a displaying male in a Balancing Posture, with wings spread open and bill Gaping. The Gaping continues after the wings have been folded in. The male often responds to the approach with a Bill Snap. Often females are driven off, but they generally return repeatedly until they are accepted by the male. Following pair formation, the Balancing Posture and Gaping cease after the female is allowed to enter the nest, and Bill Snaps are given by both sexes upon approach of the mate. In the Wood Stork's Up-Down display, the bird lifts its head and neck upwards, gapes the mandibles widely, and then lowers the head and gives a fairly loud 'fizzing' call, but few or no bill snaps. The Up-

Down may grade into Swaying Twig-Grasping, and Bill Snaps may occur between other displays. During Copulation Clattering, the male clatters his mandibles loudly and, at the same time, beats his bill roughly back and forth against the female's bill. Patterns of adult attendance at the colony site depend on the timing of thermal updrafts. Adults may stay in the colony until mid-morning, when they use thermals to soar and glide to the feeding grounds. They may return to the colony site later in the afternoon, or stay at distant feeding sites overnight before returning on the next morning's thermals. Nest-building is carried out by both sexes, and sticks are added to the nest throughout the breeding season. Stick gathering—which is done largely by the males during the early stages—is a highly social affair, with dozens of birds converging from various parts of the colony on a single tree to gather sticks and bicker with each other. At Georgia colonies, males did about twice as much stick gathering as females (M. Coulter pers. comm. 1990). At Corkscrew, in southern Florida, stick gathering activities peaked in the early morning hours (05.30-08.30 h), before thermals developed and the major feeding flights of the day occurred (pers. obs. M.P.K.) Eggs are laid at 1-2 day intervals, and are incubated by both parents from the first egg. The finely pitted shell is a dull or cream white, usually with considerable staining, and the eggs are ovate to elliptical ovate or elongate ovate (Bent 1926:60). Clutch size is 2-5, most typically 3, but can vary widely from year to year at any given location. In southern Florida, 85 clutches (pre-1940) averaged 3.48 eggs and 60 clutches (post-1970) averaged 3.33 eggs; in central and northern Florida, 81 clutches (pre-1940) averaged 3.05 eggs (J.C. Ogden, pers. comm. 1990). Rodgers (1990) found that 199 clutches collected in southern and central Florida averaged 3.28 eggs. In Yucatan, clutch size averaged 2.16 (C. Ramo and B. Busto pers. comm. 1991). Incubation takes 27-32 days, averaging 27.9 days in six clutches (pers. obs. M.P.K.). The average weight at hatching is 62 g. However, hatching is asynchronous and there is soon a substantial size difference among siblings, the smallest and youngest of which is the first to die if food resources prove inadequate. After the young hatch, one adult remains in attendance through the third or fourth week, shading the nestlings when necessary from the heat of the sun and protecting them from crows, which may attack the eggs or young, or other storks, which may attempt to take over the nest. Both parents feed the young by regurgitating onto the nest platform, where the food is picked by the nestlings. Small young are fed smaller quantities and more frequently than larger young; young 5-7 days old are fed nearly 15 times per day, whereas young 3-5 weeks old are fed about 4.5 times per day, and young 6-8 weeks old were fed about 2.5 times per day (n = 231 feedings at 4 nests) (pers. obs. M.P.K.). Parents also bring water in their gullet, which they drool over the nestlings, some of it going into their mouths. Nestlings Beg with a rhythmic nodding of the head, wings partially spread.

American Wood Stork 41

Nestling growth is rapid (Kahl 1962), although not so rapid as given by Ricklefs (1968), who based his calculations on too high an adult weight. By the third or fourth week, young have grown to half of adult size and can stand and exercise their wings (Kahl 1962). By 49-55 days (average = 51.2 days, n = 9), they can leave the nest to practice flying. Young leave the colony at about nine weeks, although sometimes returning to the nest site to roost and to be fed for several more weeks. At Corkscrew in 1959, young at least 90 days old, which had been observed throughout the nesting season, were still returning to their former nests to be fed by parents. Near large breeding colonies, young tend to gather in feeding flocks that contain few or no adults; young from smaller colonies seem to more often associate with adults at feeding areas (pers. obs. M.P.K.). Recent nesting success has ranged from 2.7 young per nest in Georgia to 0.8 in northern Florida, to 0.3 in Georgia and of course 0.0 in many instances (Coulter 1987a,b, Rodgers et al. 1987, M. Coulter pers. comm. 1990). The range everywhere is great and some northern Florida colonies can be as productive as the Georgia ones (J.A. Rodgers pers. comm. 1990). Recent nesting success in southern Florida has been minuscule: in the Everglades National Park, for example, 6 years of success and 23 years of failure between 1962-1990 (J.C. Ogden pers. comm. 1990). The greatest period of nest-mortality comes during the egg phase of the cycle; the second greatest period of nestmortality is when nestlings are between 0-14 days old. In north and central Florida colonies, between 1981 and 1985, 86% of nestling deaths were probably due to starvation, especially of the youngest nestling in a nest (J.A. Rodgers pers. comm. 1990). Other factors contributing to nesting mortality include intra- and interspecific attacks and weather disturbances. Strange, apparently unmated, American Wood Storks sometimes attack occupied nests, and the ensuing fights often result in egg or nestling mortality. The primary reason for nesting failure is rising water levels on the feeding ground, leading to chick starvation and colony abandonment. In the Everglades, this rise can be as little as a few centimetres (Kushlan et al. 1975). The phenomenon of water level related nesting failure has been consistently found in several portions of this species' range (Kahl 1964, E.S. Clark 1978a, Ayarzaguena Sanz et al. 1981, C. Ramo and B. Busto pers. comm. 1991). Predation by other birds (e.g. Fish Crows, Corvus ossifragus) or mammals (e.g. Raccoons, Procyon lotor) causes the loss of some eggs or nestlings, but this is often precipitated by other factors such as parental desertion or drying of the substrate below the nests (Rodgers et al. 1987, Ruckdeschel and Shoop 1987). At Georgia colonies, mammalian predation is often a major factor and reproductive success is often determined by the outcome of a 'race' to fledge the young before the swamp under the nests becomes dry, thus allowing predators access to the trees (M. Coulter pers. comm. 1990). Strong storms, especially when the nestlings are small, may result in heavy or even complete losses. When accompanied by severe cold, storms may kill some young directly. However, the primary effect seems to be that of dispersing

the food supply through the flooding of feeding sites, thereby causing parents to desert young they cannot feed (Kahl 1964). In recent decades, low productivity in southern Florida colonies has been due largely to man-caused ecological disturbances (see Conservation). TAXONOMY Formerly called 'Wood Ibis', this is a true stork and not to be confused with the ibises of the family Threskiornithidae. Behavioural and morphological similarities with the three Old World species of wood stork, formerly placed in the genus Ibis, have resulted in the combining of all the wood storks into a single genus, Mycteria (Kahl 1972e, 1979b). CONSERVATION Population decreases in southern Florida have resulted in the American Wood Stork being placed (in February 1984) on the List of Endangered Species in the United States. The decrease in the Everglades and Big Cypress regions is attributable, indirectly, to water management, which has altered the way in which the wetlands dry in the dry season and thereby effected the timing and success of nesting (Kushlan 1987). This has resulted in the beginning of the nesting season shifting from November—December, to February-April, and the storks cannot finish before the beginning of the summer rainy season (Kushlan et al. 1975). American Wood Storks nested successfully in the Everglades only twice between 1967 and 1987. In the Big Cypress Swamp of southern Florida, drainage and development have reduced the foraging area available to storks and altered the hydrology of the wetland that remains (Browder 1978); storks nested successfully there in only 6 years between 1963 and 1989 (J.C. Ogden pers. comm. 1990). Similar plans to drain or to manage wetlands for agriculture are proposed or in place for the llanos of Venezuela, coastal marshes of Mexico, and portions of the Brazilian Pantanal (Luthin 1987). The potential loss of an intermittently-used feeding site at the Savannah River Nuclear Plant, South Carolina, led to the successful development of artificial feeding ponds as a mitigation measure (Coulter 1986a-c, Coulter et al. 1987b). Elsewhere in its range, the American Wood Stork is often threatened by human disturbance at nesting sites. Nesting storks in Mexico and elsewhere in Latin America are hunted by local residents for food (Luthin 1987). Such hunting pressure is responsible for the loss of the known nesting sites in Belize (Luthin 1987). It is especially important to protect the nesting area in the Usumacinta Delta, Campeche, Mexico, which is the most productive population of American Wood Storks remaining in North America (Sprunt IV and Knoder 1980). It should be recognized, however, that the Wood Stork is still an abundant species over much of its natural range. In fact, the nesting population in the southeastern USA outside of southern Florida has expanded its range and, apparently, increased in recent years (Ogden et al. 1987). Although population estimates are very inexact, owing to errors in-

42

American Wood Stork

herent in aerial surveying (J.A. Rodgers pers. comm. 1990), it appears that the total US population has at least remained stable in the last decade. Expansion into northern Georgia and South Carolina began in 1980 and 1981, respectively. Although there was, in the past, some indication of excessive levels of pesticide contamination in the USA (Ohlendorf et al. 1978), there is no indication of any decrease in clutch size (Rodgers 1990—contrary to Ogden and Patty 1982). The conservation strategy for the American Wood Stork is two-fold: protection and habitat preservation and management. The protection of colony sites from disturbance, hunting and tree cutting is essential, as is prevention of loss and ecological changes in its wetland foraging habitat. In southern Florida, intensive management can create con-

ditions for successful nesting (Kushlan'1987). Loss of foraging habitat around the historically large Corkscrew Swamp colony site in Florida's Big Cypress Swamp has substantially reduced the area's carrying capacity for storks. It has been discovered that American Wood Storks will feed in artificial sites prepared for them (Coulter et al. 1987b), but this is an expensive undertaking. In Central and South America, conservation must centre on reducing wetland alteration in order to protect stork populations.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 284.

Milky Stork Mycteria cinerea (Raffles) Tantalus cinereus Raffles, 1822, Trans. Linn. Soc. London, 13, p. 327: Sumatra Other names: Malayan Wood Stork, White Malayan Tantalus, Southern Painted Stork (English); Javaanse Nimmerzat (Dutch); Milchstorch (German); Walengkada, Walangkada (Javanese); Kuntul kelabu (Sumatran); Bloewo, Burong upeh (Malay)

IDENTIFICATION Slightly smaller than the Painted Stork, standing 91-95 cm tall, the Milky Stork is an all-white bird, except for the flight feathers of the wing and tail, which are black, with a greenish gloss. Its downcurved bill, similarly shaped to that of other wood storks, is dull yellow and sometimes tipped with white. Its legs are dull red to flesh colour, its bare facial skin is greyish (this may be from faded skins) to dark maroon, blotched irregularly with black. The iris is dark brown to grey-brown. In appearance, the Milky Stork is altogether a rather dull version of its African relative, the Yellowbilled Stork. In courtship, the white plumage brightens, becoming suffused with a very pale cream-yellow tinge (hence 'milky' stork). The wing-coverts and back feathers have a lighter, nearly white terminal band. In some fresh skins the wing linings are washed with salmon-pink (H.C. Robinson and F.N. Chasen 1936:205), and breeding birds show a narrow pinkish band of bare skin along the underside of the wing (pers. obs. M.P.K.). The white undertail-coverts become elongated and fluffy. The bill turns a deep yellow, with a greyish tan cast on the basal third. The legs become a deep magenta. The facial skin turns dark wine-red, with a brighter red around the eye, and a pale grey or creamy smudge above each eye; soon after courtship the facial skin fades to a paler orange-red (pers. obs. M.P.K.). (Reports of grey, yellow, or brown soft-part coloration are probably taken from preserved skins, and these colours may not be as in life.) The sexes are similar, males slightly larger on average and with a relatively thinner, longer bill. At hatching, chicks are covered with white down, By 10-14 days the contour feathers start to appear, and by 5-6 weeks they are fully feathered. Nestlings have a dark brownish-grey bill (Hoogerwerf 1936c). Immature birds are greyish brown above with a white lower back, rump and tail-coverts, and black flight feathers; the wing-lining is mixed dark brown and white. The head and neck are darker brown. At about 10 weeks loss of the small feathers on the head starts and dark naked areas are visible on the face. After about 3 months, the bill colour changes to a warm yellow, with a greenish yellow tip. The iris remains dark brown, and the legs are flesh-coloured (Hoogerwerf 1936c). The sounds produced by the Milky Stork are not noticeably different from those of the other wood storks (Kahl 1972b). Generally silent, except at breeding colonies. A falsetto 'fizzing' vocalization, audible for about 75 m, is given during the Up-Down display, and bill-clattering and wing noise are also heard during other social displays. Nestlings make a frog-like croaking sound when Begging (Hoogerwerf 1936c). In flight, the flapping-rate averages 205/min (n = 21) (Kahl 1971b). Soaring on thermals occurs usually between 10.00 and 14.00 h (Hoogerwerf and Rengers Hora Siccama 1937). At breeding colonies and feeding areas, flight activities are contagious, with a take-off by one bird often followed by several other birds nearby (pers. obs. M.P.K.).

Milky Stork 45

In most of its range only the white egrets and the Asian Openbill Stork are likely to be confused with the Milky Stork. The egrets are smaller and entirely white, and the Asian Openbill is smaller and easily distinguished by its peculiar greyish bill. In the northern part of its range (Kampuchea, Vietnam), Milky Storks may sometimes occur with the more common and slightly larger Painted Stork, the latter, in adult plumage, is distinguished by the black and white breast-band and wing-coverts, pink inner secondaries, and generally brighter soft-part coloration. Immatures of the two species are very similar, the Painted Stork being slightly larger and somewhat darker in overall coloration. It is also said (B.F. King and E.G. Dickinson 1975:50-51) that immatures can be separated by the Milky Stork having a paler underwing-lining, which contrasts with the black flight feathers (whereas immature Painted Storks have the entire underwing blackish). DISTRIBUTION AND POPULATION The Milky Stork was formerly a widespread resident in Southeast Asia, including Kampuchea (=Cambodia), southern Vietnam, the lowlands of Malaysia, and several islands in Indonesia (H.C. Robinson and F.N. Chasen 1936, Kahl 1979b). An old specimen collected in Thailand in 1935 was recently located (Morioka and Yano 1990). Luthin (1987) and Verheugt (1987) have summarized its recent status. A few birds were still to be found —in mixed feeding flocks with Painted Storks, Lesser Adjutants and herons—at the Grand Lac, Kampuchea, in April 1968 (pers. obs. M.P.K.). No recent records exist from Thailand. Some

birds have apparently recolonized Vietnam since the war of 1963-75, but their breeding status is unknown. Until 1989, the species had not been found breeding in Malaysia since 1935 (Madoc 1936, Collar and Andrew 1988:14). However, storks in breeding plumage were seen at Kuala Gula in July 1984, and about 20 nests were located there in 1989; there are an estimated 100-150 Milky Storks in this area, and the population is thought to be increasing (K. Brouwerpers. comm. 1990). The Milky Stork nests on several islands in Indonesia. On the north coast of Java, the birds have bred on two island reserves. We saw breeding birds at Pulau Dua in 1968 and at Pulau Rambut in the 1970s (pers. obs. J.A.H.). The species nests in eastern Sumatra at Hutan Bakau Pantai Timor Reserve in Jambi Province. In Riau and South Sumatra, regular breeding is suspected but not confirmed in coastal lowlands (Silvius et al. 1986). Also in Sumatra, the species is regularly recorded in Lampung Province from the Way Kambas Game Preserve. On Sulawesi, Uttley (1987) noted adults in breeding plumage and immature birds at a mixed colony at the mouth of the Sulowath River, but no nests were found; there is apparently a small resident population of Milky Storks in Sulawesi (G.M.N. White and M.D. Bruce 1986:111). In addition, it is possible that the Milky Stork may yet be found in Kalimantan. The historic population centre was Indonesia, but never in the period since the Second World War has this species been numerous; and now it is rare everywhere. The Milky Stork is a wanderer and disperses after the breeding season, but the details of its movements are poorly known. It could wander from late September to April (Verheugt 1987). Probably it moves to traditional feeding areas annually after breeding, but no doubt it will shift when conditions alter, either due to changes in rainfall patterns or because of disturbance or harassment. It occurs seasonally in Bali, Sumbawa, and other areas of Java. Over 1000 individuals visit the northeast coast of West Java seasonally; it is likely that these are largely migrants from southeast Sumatra. In Sumatra non-breeding birds are sometimes found in the interior of the island and up to 900 m in altitude (Marie and Voous 1988:65). Because so much of the Milky Stork's range is inaccessible, population estimates are extremely difficult to obtain. It is possible that the entire population is as low as 5000 birds (Verheugt 1987). [The world total of '30QOO', given in Kahl (1987a) is a misprint and should instead have read '3000'.] We were fortunate to see this species in two island reserves in Java in the 1960s and 1970s, and at that time it was thought that the total breeding population was below sustainable levels. The bird continues to be rare, although greater searching effort has recently found isolated remnants. Several small subpopulations maintain a precious foothold in Sumatra, Indonesia. In Jambi Province, 74 active nests were found in July 1985 within the Hutan Bakau Pantai Timor Reserve. In September 1988, nesting Milky Storks were observed from aerial surveys at three colonies in South Sumatra Province (Danielsen et al. in press a); approximately 900 adults and at least 150 juveniles were

46 Milky Stork

counted in the colonies, and there were estimated to be about 1000 Milky Stork nests (MJ. Silvius pers. comm. 1989). The largest number of Milky Storks counted in South Sumatra was 1587 (Silvius et al. 1986), the remanant core of this dwindling population. In Lampung Province, 116 birds were seen at a single site in November 1985 (Milton and Marhadi 1985). Uttley (1986) found 33 adults and 4 immatures at a colony site in Sulawesi, although breeding was not confirmed. At Pulau Rambut, West Java, there were a minimum of 14 active nests in 1984, with a maximum count of 76 adults (Allport and Wilson 1986). It is estimated that between 356 and 408 birds are resident along the northern coast of West Java (Verheugt 1987). Approximately 100 birds are resident at Kuala Gula, northern Malaysia (Duncan Parish pers. comm. 1989), and some 20 pairs were found nesting there in November 1989 (K. Brouwer and M.G. Coulter 1990). ECOLOGY The Milky Stork is found primarily in lowlands, and is especially associated with coastal mangrove swamps. It also uses various aquatic habitats, including ponds, lakes, marshes, tidal mudflats and saline pools, and visits fish ponds and rice fields (B.F. King and E.G. Dickinson 1975:50). In intertidal areas, the level of the tide dictates the stork's feeding pattern. Except for the small population near the Grand Lac, Kampuchea, most Milky Storks seem to be associated with salt water and mangroves. The Milky Stork often feeds in aggregations with other wading birds, especially Lesser Adjutant Storks and Great White Egrets (Egretta alba). It forages by Walking Slowly and Groping with its bill slightly open, moving it from side to side, and also by Walking and Probing in the mud. It has been reported to Foot Stir, like the other wood storks (Hoogerwerf 1936c). The main method of feeding is by tactile location, but undoubtedly it visually locates mud-skippers which are abundant on the mudflats of Indonesia and Vietnam (Silvius 1988). On intertidal mudflats in western Malaysia, a bird was observed to Probe in the mud, to Grope in the water and to search visually for prey (Swennen and Marteijn 1987). We have seen this species on Pulau Rambut feeding in the shallow pools beneath the mangrove bushes in the company of egrets. Here food supplies were abundant, allowing this stork to consume sufficient food in a short period and enabling it to lounge around for prolonged periods of the day. At the Pulau Dua, Java, breeding colony in March 1968 there was considerable in-and-out flight activity on nights with a full moon, indicating that the birds not only forage, but also visit their nests, at night (pers. obs. M.P.K.). The main diet is small fish, primarily mud-skippers (Gobiidae) in Malaysia (Swennen and Marteijn 1987), but snakes and frogs have also been recorded (Hoogerwerf and Rengers Hora Siccama 1937). Hoogerwerf (1936c) observed parents feeding 20 cm fish, eels and mud-skippers to their young in the nest in western Java. Like other wood storks, the Milky Stork will spend time

soaring on thermals, and it uses this method of flight when moving from area to area. At high tide, birds are often seen resting in mangrove forests. They are said to be generally shy and vulnerable to disturbance (Verheugt 1987), although we found that birds at the nest could be approached easily. BREEDING In Indonesia, breeding takes place after the rains, during the dry season that usually lasts until October. The species probably depends on particular water levels for feeding; the start of the breeding season probably coincides with maximum fish stocks, following reproduction of fish during the rainy season. In Java, nests were often in black mangroves (Avicennia marina) 8-30 m high (Hoogerwerf 1936c, Verheugt 1987), and in Sumatra nests were located in red mangroves (Rhizophora apiculata) (Silvius 1988). On the north coast of Java, nests on Pulau Rambut were built on top of the highest trees, some 30 m above the ground and we found some of the nests in similar position on Pulau Dua when the species still nested there. A few large trees on one side of the island were used in preference to the lower mangroves and creeper-covered bushes occupied by the bulk of the other wading birds, although some Milky Storks nested as low as 4 m at Pulau Dua. Unfortunately, since the mid-1970s Pulau Dua has become joined to the mainland by accumulated silt, allowing access to humans and other predators, and since that time the storks have not nested at this site. Several South Sumatran colonies were built 2-15 m up in mangroves within 1—4 km of the coast; here the Milky Storks nested with Lesser Adjutant Storks, Oriental White Ibis, and several species of herons (Danielsen et al. in press a). Malaysian nests were 8-1 Om above the ground, mostly in dead trees (S.H. Yatim pers. comm. to K. Brouwer, 1990). The nests are constructed of sticks and lined with fresh vegetation. Hoogerwerf (1936a) described the nests as similar to those of the Grey Heron (Ardea cinerea) but slightly more robust and containing thicker branches. We have found the breeding and courtship of the Milky Stork to be similar to other members of the wood stork group, although certain threat displays, such as Aerial Clattering and the Forward display, have not yet been observed (Kahl 1972b). During many displays at the nest, performing birds retract the skin of their head and upper neck, exposing two or three times as much area of naked skin; the amplitude of this retraction is even greater than that seen in the Yellowbilled Stork. Between displays, when the retraction is relaxed, and the skin hangs in loose folds and wrinkles, like wattles (pers. obs. M.P.K.). The male advertises using Display Preening and the arriving female reponds with the Balancing Posture and Gaping, which may continue for several days during pairformation. The male uses a Flying Around display, just after the female occupies the nest site, and Up-Down displays are given in greeting. In the Milky Stork, the Up-Down includes four or five bill snaps in a series between hissingscreaming vocalizations. Swaying Twig-Grasping is also a

Milky Stork 47

typical display. In response to disturbances at the nest, the Arching display is given; the photograph in Hoogerwerf (1936c) shows it to be essentially identical to the homologous display in the other wood storks. Allopreening between pairs is frequently followed by body shaking. Head rubbing is another common comfort movement; the bare skin of the head may first be oiled by rubbing it on the preen gland and is then rubbed on the plumes of the back. This movement is carried out quite frequently by birds perched nearby an incubating partner. Nesting is colonial, and, in Java, Hoogerwerf (1936c) reported 5-8 nests in a single tree. Most colonies are rather small, with from 10-20 up to a few hundred nests. The eggs are chalky white and usually very dirty. At West Java colonies, 9 clutches contained 3 eggs each (Hellebrekers and Hoogerwerf 1967); sometimes the clutch is 4 (Hoogerwerf 1949). The incubation period is estimated at 27-30 days (Hoogerwerf 1936c). Parents feed by regurgitating food onto the floor of the nest, where the nestlings pick it up. Small young are fed more often than large young; up to 4 weeks, chicks may be fed twice per hour, whereas older chicks may only be fed once in the afternoon (Hoogerwerf 1936c). During the heat of the day, the young are often shielded by the open wings of the adult which stands over the nest. Adults sometimes bring water to cool nestlings and for them to drink (drooled from the parent's bill). Heat stress is also alleviated by birds excreting a dilute urine onto their legs, which turns them white. At Pulau Rambut, Java, in 1984, active nests held about 2 young each (Verheugt 1987). By 6-7 weeks the young are able to leave the nest and begin to fly, and by 8 weeks they are flying well but are still fed in the nest by the parents (Hoogerwerf 1936c). TAXONOMY The species was formerly and inappropriately called 'Wood Ibis', which led to unnecessary confusion with the true ibises (family Threskiornithidae). Because the morphological and behavioural similarities are so great, this species, formerly in the genus Ibis along the the other Old World wood storks, is now included with them and the American Wood Stork in the genus Mycteria. CONSERVATION This species was essentially eliminated from Vietnam by the widespread destruction of mangrove swamps during the

Southeast Asian War (1963-75). Extensive mangrove reforestation and protection by wardens may have resulted in some recolonization in recent years. Elsewhere it is seriously threatened by poaching, disturbances at nesting colonies, mangrove deforestation, tidal rice cultivation, fish-farming, and—in Indonesia—human resettlement (Luthin 1987, Verheugt 1987, Collar and Andrew 1988:14). Of the 26 sites at which the species is known to occur, 16 (62%) are reported to be under moderate to severe threat (D.A. Scott and C.M. Poole 1989). In addition, specimens appear from time to time in the zoo trade (Danielsen et al. in press b); some of these have been purchased as 'juvenile Painted Storks' and, on maturity, proved to be Milky Storks instead. At present, some 70 Milky Storks are kept by nine zoos outside of Indonesia (K. Brouwer pers. comm. 1990). Indonesia Interwader (the South East Asia Pacific Shorebirds Study Programme, with headquarters in Kuala Lumpur, Malaysia), the ICBP and other European-based conservation organizations participated in wide-ranging surveys in the mid 1980s, providing much of the status information we report here. A Milky Stork conservation programme has been proposed. It is especially critical to establish an integrated network of wetland preserves, as is being done in Vietnam. However, the loss of coastal wetland habitat continues, and in Indonesia is estimated to account for the loss of 100000 ha per year (Verheugt 1987). The near total destruction of the mangrove forest in Java is the most serious threat to the species there. Also, in many areas birds are netted throughout the year and eggs are taken during nesting. The Milky Stork is now an officially protected bird in Malaysia and Indonesia (Verheugt 1987), and as the core of the population is confined to areas near gazetted nature and wildlife reserves, it is hoped that the considerable devastation brought about in recent years by humans can be mitigated. Certainly without active conservation efforts, especially those sponsored by Interwader and supported by ICBP and other conservation organizations, this stork has little chance of survival. The species has been listed on Appendix I of CITES since 1987.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 285.

Yellowbilled Stork Mycteria ibis (Linnaeus) Tantalus ibis Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 241: Egypt Other names: Wood Ibis, African Wood Stork (English); Tantale ibis, L'ibis tantale (French); Afrikanischer Nimmersatt (German); Kamindo (Mandingo); Tsagtsagi, Tsagagi (Hausa); Okogliech (Luo); Gangala (Mozambique); Levlosyane (Sotho); Nepando (Kwangali); Geelbekooievaar (Afrikaans); Naedjah, Badja (Arabic)

IDENTIFICATION This medium-sized stork, standing 90-105 cm, appears all white, except for its black wings and tail. The back of its head and neck are a dirty white, whereas its back, wing coverts, breast, and belly are white, more-or-less suffused with pink on the wings. The tail, flight feathers, and greater primary coverts are black. The distinctive yellow bill is long and slightly decurved, as is the case for all wood storks. The head is feathered to the hindcrown, and the bare skin of the forehead, face and throat is orange to red; the legs are pale red. Courtship and breeding colours are much brighter. The bill becomes a rich golden-yellow, while the face turns a bright carmine-red, grading to orange posteriorly near the feathers. During courtship displays, the skin of the head is retracted, considerably enlarging the visible area of bare red skin (Kahl 1972b). The legs and feet become a brighter red. Much of the white plumage becomes suffused with pink and the wing-coverts are crimson, especially the greater underprimary- and underalula-coverts, each feather being tipped in white. When freshly moulted, the black feathers of the wings and tail are glossed with green and purple. The sexes are alike, but the male is slightly larger. At hatching, the nestling weighs about 60 g and is covered with a sparse grey down. By 10 days, the weight increases to about 500 g, and the grey first-down is superseded by a white, woolly second-down. The bill and bare facial skin start out mostly yellow and gradually change to black with age. By about 5 weeks of age the bill starts, from the tip, lightening to a pale yellow. Nestlings have dark brown irises and legs and feet that progress from a fleshy-pink to a cyano tic-blue (pers. obs. M.P.K.). Immature birds have the feathering on the head extending further forward than in adults. They are greyish brown, paler underneath, with a dull partially bare face of orange and a horn or dull yellowish bill. The legs and feet are a brown horn colour, and the primary, secondary and tail feathers are blackish brown. By about the age of fledging, they start to develop some salmon-pink coloration in the underwing linings (pers. obs. M.P.K.). After the first year, the body plumage is white washed with grey, the flight feathers of the wing and tail become black, and adult-coloured soft parts begin to appear, although the pink flush does not occur until later. Birds are probably not fully adult for 3 years. The Yellowbilled Stork is generally a silent bird, but during social displays a rather weak hissing scream is uttered. At breeding colonies a variety of bill-clattering and 'woofing' wing sounds are heard. Nestlings give a loud, repeated, and monotonous braying call when Begging adults for food. The Yellowbilled Stork flies with its neck and legs fully extended, using either flapping or soaring

50 Yellowbilled Stork

flight. The flapping rate averages 186/min (n = 20) (Kahl 1971b). For flights to distant feeding grounds, Yellowbilled Storks soar on thermals and glide, as do most storks, to cover long distances without wasting much energy. When descending from on high, either at the feeding areas or the breeding colony, birds often dive at high speeds, side-slip and flip; they appear to enjoy this aerial play. The Yellowbilled Stork can be confused with wintering White Storks. However, the decurved yellow bill, black tail and rosy pink cast of the adults are distinctive. In flight, at a distance, it might be mistaken for a Great White Pelican (Pelecanus onocrotalus) or an Egyptian Vulture (Neophron percnopterus), both of which have shorter necks and legs and white tails. DISTRIBUTION AND POPULATION The Yellowbilled Stork is widely distributed in suitable habitat in tropical Africa, south of the Sahara from Senegal and Somalia south to South Africa, where it is rare south of the Orange River, and also to Madagascar. It seems, generally, to avoid the heavily forested regions of central Africa. It is a straggler to southern Tunisia, Morocco, and Israel, and formerly to Egypt (F.J. Jackson 1938:80, Castan and Olier 1959, Paz 1987:34). The species is particularly widespread in East Africa, where it breeds in Kenya (3 sites), Tanzania (5 sites), and Uganda (common, but not recorded breeding) (Kasoma and Pomeroy 1987). In the Sudan it breeds at Malakol and probably other sites. In West Africa, it nests from Gambia to northern Nigeria, often inside walled cities (Bannerman 1953:180). It was found breeding in Zululand, South Africa, in 1949 and again in 1954, 1960 and 1961 (Clancey 1964a:47^8, Berruti 1980b). It has bred in northern Botswana (W. Fraser 1971) but is generally uncommon south of the well-watered parts of Zimbabwe and northern Bots-

wana. It occurs sparingly in the non-breeding season over much of South Africa, although Clancey (1964a:47) found it present year-round in the coastal waters of Natal. It is found quite widely in western Madagascar (Rand 1936:340); Langrand (1990) reported that there is no direct evidence of nesting there now, although in October near Lake Kinkony, young birds hardly able to fly were seen. The Yellowbilled Stork undertakes local movements in Kenya and migrates from north to south in Sudan with the rainy seasons, leaving the Sudd during the flooding period (Luthin 1984a,b). In the Bangweulu area of northeastern Zambia it leaves when the water is high—and fish are hard to catch —and returns in large numbers when the swamps begin to dry (Brelsford 1946). Although it migrates regularly to and from South Africa, its general movements are poorly known, and in some areas it remains year-round. It is probably best to describe the Yellowbilled Stork as a facultatively nomadic species, avoiding areas where water or rainfall conditions are inappropriate. The overall population may be considered abundant and stable. The species is not uncommon as far north as the Sudan, and in Kenya we found it to be the commonest species in a colony on the Tana river, numbering about 2000 birds. In the following year after the long rains it also nested there, some 200 birds being present. Larger numbers than this are found in eastern and central Africa in seasons when rainfall is adequate. A colony of at least 2000 pairs was found in western Tanzania in February 1962 (Stronach 1968). ECOLOGY This stork appears to use a somewhat wider range of habitats and have a broader diet than other wood storks. Its habitat use is particularly diverse, including large marshes, swamps, margins of lakes or rivers, rice fields, and even small pools. It feeds in tidal pools along the beach in East Africa (Brown et al. 1982:174). Most of its food is taken in water about 10-40 cm deep, using its partially open bill to Sweep and Grope. The Yellowbilled Stork feeds by tactile location, Standing, Walking Slowly, Groping, Foot Stirring, and Wing Flashing. It will often stand still to await the arrival of prey, snapping its partially open bill at its touch. Foot Stirring is often accompanied by the flashing open of the wing on the same side (Talent 1940), as with the Painted Stork and the American Wood Stork. Prey is quickly swallowed with a toss of the head. The species usually feeds singly or in small groups, but at times in the hundreds (F.J.Jackson 1938). Birds in foraging groups often threaten each other with Forward Displays, in which the neck is retracted, the black tail is cocked upward, the wings are slightly drooped, and the back and neck feathers are erected (J. Cooper 1969, Kahl 1972b). It will sometimes feed in company with other species, often herons, spoonbills, or Great White Pelicans (Pelecanus onocrotalus). On rare occasions, Yellowbilled Storks have been seen feeding in the wake of a moving crocodile or hippopotamus, apparently taking advantage of organisms stirred up by the

Yellowbilled Stork

larger animal (Pooley 1967a). The storks feed apparently for only a short time, obtaining their requirements often very quickly and then resting. Frogs, small fish, aquatic insects, worms and crustaceans, and possibly some small mammals and birds are eaten. A tilapia of around 150 g appears to be about the maximum size offish that can be swallowed (Bell-Cross 1974). Scavenging of food regurgitated by cormorants has been observed (Kasoma and Pomeroy 1987). The Yellowbilled Stork will roost with other species on the ground, and at night in large tree roosts. When resting on the ground, birds often sit on their tarsi. Sometimes, especially after a rain or following bathing, they stand with their wings fully extended to the side until dry (Kahl 1971g). BREEDING The breeding pattern of this stork varies throughout the African continent. Nesting areas are subjected to annual or semi-annual rainfall, and the Yellowbilled Stork will nest either at the peak of or at the end of the long rains, whenever food is most abundant because of local ecological conditions. Especially in extensive marshlands with little topographical relief, nesting is at the end of the rains, as declining water levels concentrate fish, similar to the situation with the American Wood Stork in southern Florida. However in other habitats, such as in the marshes near Lake Victoria in western Kenya, breeding occurs at the peak of the rains, as fish move out of the lake and adjacent rivers and into the flooded marshes (Kahl 1968). There, during the dry season, the fish are in deeper waters and unavailable to the storks; when fish move into shallower marshes to spawn, they and their fry are most available to all fisheating birds, who time their breeding cycle to this annual event. Cases of off-season breeding of Yellowbilled Storks have been reported from western Kenya (J. Parsons 1977), northern Botswana (Tree 1978) and eastern Kenya (Hancock 1984:41), when unseasonal rainfall caused local flooding and ideal feeding conditions. Breeding sites and feeding grounds are often far apart, and birds will travel quite long distances—usually by using thermals for soaring and gliding flight—to obtain food. Nesting is colonial, often in company with other waterbird species, but sometimes the Yellowbilled Stork is the sole or major occupant of a nesting site. Nests are built throughout a mixed colony, with perhaps up to 20 storks nesting close together in any one part of the colony. The nests may be built in small trees, usually over water, or in larger trees over dry land. As many as 50 nests have been counted in a single large tree (Bannerman 1953:180). Pair-formation begins when the male displays from its chosen position, using Display Preening which appears to expose the brightest parts of the wing to an approaching female. The interested female adopts a Balancing Posture, with her head low and forward, bill agape and wings spread. She will also use Gaping without the spread wings, a behaviour pattern unique to the wood storks. Aerial Clattering and Swaying Twig-Grasping are characteristic. In many

51

displays, the skin of the head is pulled back, allowing the bright-coloured bare skin to expand over the entire head. Up-Down displays are performed by the pair. Bill clattering occurs during copulation in a similar manner to other wood storks, and threat displays and anxiety movements are similar to those of the other members of this genus. In traditional sites, old nests—if any remain from a previous season—may be repaired, but usually new stick nests are constructed and lined with grass or green leaves. Nests take up to 10 days to build and are 80-100 cm in diameter and 20-30 cm thick. 2-4 eggs (usually 3) are laid; average clutch size (n = 45) was 2.5 (Brown et al. 1982:175). They are a dull white to greenish white, usually unmarked but rapidly become stained. J. Parsons (1977) found that there was a significant decrease in 'the width of Yellowbilled Storks' eggs from a Kenyan colony as the season progressed. Both parents share duties of incubation, brooding, guarding and feeding the young. Incubation takes about 30 days. Young hatch at 1- or 2-day intervals and grow rapidly. Their insatiable appetites make the German common name 'Nimmersatt' (never satisfied) particularly appropriate. One parent usually attends the young constantly until they are about 21 days of age, at which time both adults forage much of the time to fulfil the nestlings' food demands. At this age, young suddenly become quite agressive, and this may be in response to the need to defend themselves from attack, particularly from other Yellowbilled Storks that attempt to take over the nest when adults are absent. Feeding is by regurgitation on to the nest floor, and water is passed from adults (Stronach 1968) by dribbling it into the nestling's bill. Adults will shade the young with open wings, but brooding of young quickly ceases. The slower movements of these storks when nestlings reduces losses by falling from the tree, unlike heron species where fighting and rapid early movements away from the nest result in heavy losses. Generally, breeding success seems to be relatively high, with broods of 1-3/nest recorded in East Africa (Brown et al. 1982:175). Egg predation by Fish Eagles (Haliaeetus vocifer) was heavy at a colony near Kisumu, Kenya, in 1975. At this colony, at least 236 eggs were laid in 95 nests (average of 2.48 eggs/nest). An estimated 61% hatched and 38% were predated; only 31 young fledged, giving a success rate of only 0.33 young/nest (J. Parsons 1977). The young take their first flights at 50-55 days and soon leave the nest and become fully independent. Juveniles take at least 3 years to attain adult appearance and may not breed until later; firm data are lacking however. TAXONOMY In the past, the Yellowbilled Stork, along with the Painted and Milky storks, was classified in the genus Ibis. However, intensive studies have shown that they are closely similar to the American Wood Stork, and so all four species have been united in the genus Mycteria (Kahl 1971f, 1972e). In the older literature, storks of the genus Mycteria (including the former genus Ibis] were often called by the common name of'Wood Ibis'. This led to much confusion

52 Yellowbilled Stork

with the true ibises (family Threskiornithidae). It has long been recognized that these birds are, indeed, true storks (Garrod 1875a,b) and should not be called ibises. Names in current use reflect that fact and avoid much of the confusion of the past. However, readers consulting the older literature should be aware of these nomenclatural changes.

ditions. In East Africa, it is both abundant and stable but is subject to pressure from poaching and from reduction of habitat. Whilst increasing human populations in many parts of Africa constitute a long-term threat, it cannot be considered other than an abundant species with a comparatively large, stable population at present.

CONSERVATION

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 286.

This is a widespread species, which appears capable of responding to short-term natural changes in habitat con-

Painted Stork Mycteria leucocephala (Pennant) Tantalus leucocephalus Pennant, 1769, Indian ZooL, p. 11, pi. 10: Ceylon (Sri Lanka) Other names: Rosy Wood Ibis, Indian Wood Stork, Pelican-ibis (English); Indischen Nimmersatt, Buntstorch (German); Tantale indien (French); Janghil, Dhok (Hindi); Kankari (Mirshikars, Bihar); Katsarunga, Ram-jhankar, Sona-jangha (Bengali); Lamjang, Lungduk (Sind); Chitroda (Kutch); Yeru kald-konga, Yerri kali-konga (Telugu); Ghenga narai, Singa nareh (Tamil); Sangu-valai-narai, Changa vella nary (Tamil, Sri Lanka); Dae-tuduwa (Sinhalese); Hnet-kya (Burmese); Lai-tou-kuan, Tu-Tsae-kuan (Chinese)

IDENTIFICATION The Painted Stork is a medium-sized, conspicuously patterned, black and white stork, standing 93100 cm tall. The head is partially naked and orange; the bill is yellow and slightly decurved at the tip. The face and crown of the adult is featherless, and the skin is an orange-yellow to reddish orange colour. The nape, neck, back and breast are white, with a black-and-white band horizontally across the breast. The primaries, secondaries and tail are black. All the wing-coverts (except the upper greater secondary coverts, which are white) are black, glossed greenish, but feature prominent white terminal bands. These 'chequered' areas of the wings and breast are one of the distinguishing features of the species. The innermost secondaries and their coverts, close to the tail, are rose pink. The legs and feet are also pink to dull red, but they often look white from the excreta splashed over them, especially during the nesting season. The eyes are dark brown. The sexes are alike, but the male is slightly larger. In the breeding season, the plumage becomes neater and cleaner looking, with the black-and-white coverts and breast-band contrasting strongly with the surrounding feathers. The elongated inner secondaries and their coverts become a brighter pink, with white terminal bands. The bill colour deepens, and the face becomes reddish orange with a glaucous cast and the throat pinkish. By retraction of the skin, the extent of the bare head increases during some displays, but not as dramatically as in the Yellowbilled Stork of Africa. The legs turn a brighter reddish magenta, and the iris becomes a paler grey-brown (Kahl 1972b, Breeden and Breeden 1982). At hatching, the nestling is partially feathered with a pale greyish down. Partially grown chicks are covered in white down, accentuating their naked, jet black faces. The black bill is tipped with pale yellow, more on the upper than lower mandible (S. Ali and S.D. Ripley 1968:95). Immature birds have greyer plumage, with the head and neck pale brownish, arid the back darker with lighter buffy greater secondary coverts. They lack the band across the chest and the pink secondaries of adults. Head feathering of the young is lost gradually. The forehead and anterior of the crown become featherless by one year, and the entire head by two years of age. This feather loss is accompanied by an increase in lipoid secretion, probably to protect the exposed skin (Shah et al. 1977). Adults are generally silent away from the nest. During displays, the typical wood stork 'fizzing' call is given, but it is somewhat weaker in this species (Kahl 1972b). Also wing-woofing and various forms of bill-clattering are heard during displays, fights and copulation. Nestlings vocalize loudly when Begging to be fed; their voices are not noticeably different from the other wood storks. Year old juveniles can still vocalize loudly, but become practically voiceless, as adults, by 18 months (Shah and Desai 1972). In flight, the Painted Stork flies with neck extended and slightly downcurved. The flapping rate

Painted Stork 55

averages 190/min (n = 70) (Kahl 1971b). The Painted Stork readily soars high on thermals, and can be seen travelling long distances with other soaring species. In most of its range only the white egrets and the Asian Openbill Stork are likely to be confused with the Painted Stork. The egrets are smaller and all white, and the Openbill is smaller and is easily distinguished by its peculiar greyish bill. In parts of southeast Asia (e.g. Kampuchea, Vietnam), it may occur alongside the slightly smaller Milky Stork. Adults are easily distinguished by the more intricate black-and-white pattern on the wings and across the breast of the Painted Stork. It is also said (B.F. King and E.G. Dickinson 1975:50-51) that immatures can be separated by the Painted having the entire underwing blackish, whereas immature Milkys have a paler underwing-lining, which contrasts with the black flight feathers. DISTRIBUTION AND POPULATION A lowland species entirely dependent on water, the Painted Stork's range extends throughout the Indian subcontinent from Pakistan to Sri Lanka and eastward as far as south and eastern China, Thailand, Kampuchea and Vietnam. Although widespread in India, it is more localized elsewhere in its range. A small nesting colony remains in the Indus Delta of Pakistan, and a few birds appear erratically as far as Mainwali in the Punjab (T. Roberts pers. comm.). Numerous Indian colony sites are known from Gujarat, Saurashtra, Rajasthan, and elsewhere (Luthin 1984a). Colonies are located within the Colombo Zoo in Sri Lanka and the Delhi Zoo in India. There are no recent breeding records in Bangladesh (M.A.R. Khan 1987). In 1982-83, Painted Storks were observed in Burma. In Thailand, there is an old nesting record from Ban Yang (J.H. Riley 1938), and the only known recent nesting site is at the Thale Noi NonHunting Area, in southern peninsular Thailand, where only one pair remained in 1988 (P. Round et al. unpub. report). Four small colonies are known from Vietnam, in the coastal wetlands. Non-breeders occasionally wander southward in

the Malay Peninsula, as far as Kuala Lumpur (Gibson-Hill 1949:29). The species is non-migratory, moving nomadically after nesting to wet areas, dispersing in small parties to wherever food is available. The Painted Stork remains locally abundant in India and Sri Lanka, but elsewhere its numbers have been drastically reduced. One of the largest known colonies—the Keoladeo Ghana, near Bharatpur, Rajasthan, India—contains several thousand pairs in a good year (S. AH 1953:535, Kahl 1974). A colony in Baunagar, Gujarat, India, numbered 170 nests in 1983 (Luthin 1984a). A census in midwinter 1989, revealed a total of nearly 5000 birds in India and 850 in Sri Lanka (D.A. Scott and P.M. Rose 1989). The Pakistani population is small, and the species is now very rare in Bangladesh (M.A.R. Khan 1987), Burma, Thailand and Vietnam (Luthin 1987). It is unlikely that any substantial breeding occurs in any of these areas today. Earlier this century, it was said to be abundant in northwestern Kampuchea (Cambodia) (Delacour 1929); a few birds were seen there in April 1968, including some half-grown young, being held captive for human-consumption (pers. obs. M.P.K.), but nothing is known of its status now. In southern Vietnam, where it was common before the war, it now breeds only in four small colonies of 5-10 pairs each (Luthin 1984a); its present status there is uncertain. In China, it formerly occurred in summer in the southeast (La Touche 1931-34:441; Etchecopar and Hue 1978:71); it is almost certainly extirpated there now. ECOLOGY Painted Storks are found in freshwater marshes, ponds, flooded fields, sometimes river banks, and rarely on the coast and in salt pans. In the non-breeding season, they have been observed feeding in relatively dry agricultural fields (AJ. Urn pers. comm. 1990). It is a tactile feeder, searching shallow waters with its bill half open, Groping and making use of Foot Stirring and Wing Flashing to disturb its prey. Birds Walk Slowly, Groping as they move, and when food is plentiful they will feed in quite large parties. During Foot Stirring, the wing is often flicked quickly open on the same side in order to direct fish back toward the open bill. Only occasionally do feeding birds perform Forward Displays at neighbouring birds. The bilLis held forward, with the body near horizontal and the neck retracted; the feathers on the back, neck and breast are erected, undertail-coverts drooped and tail cocked. If the opponent does not retreat, its bill may be grabbed and brief grappling takes place. Their diet is almost entirely fish, although S. AH and S.D. Ripley (1968:93) reported that Painted Storks will also eat reptiles, frogs, crustaceans and insects. The length offish consumed by adults is usually 5-25 cm. In Delhi Desai et al. (1974) found that the fish taken were mainly carp and catfish during the storks' nesting season, which is also the time at which these fish breed and are most available. Stomach contents also showed that besides fish, these storks had consumed fragments of plant fibre, vegetable debris,

56 Painted Stork

small stones, aquatic insects and one small frog. It is estimated that an adult bird consumes as much as 660 kcal, or 600 g of fish, per bird per day to maintain body weight (Desai et al. 1974); a family of storks consumes approximately 240 kg of fish during a breeding season. In Delhi, breeding is in the last phase of the rainly season, when fish are available in large quantities (Desai et al. 1977). After feeding, Painted Storks frequently stand on the bank in a hunched posture (S. Ali and S.D. Ripley 1968:93). Standing birds often rest on one leg and clasp the tarsus of that leg with the other foot (Henry 1971:384). When resting on the ground away from the nest, they sit on their heels with their tarsi folded forward horizontally. BREEDING A colonial breeder, the Painted Stork often nests in mixed colonies of storks, ibises, spoonbills, cormorants and herons. In the larger colonies in India, such as at Bharatpur, up to several thousand pairs nest. Colony sites are traditional and are often used for many years if undisturbed (S. Ali and S.D. Ripley 1968:93). Breeding normally starts during the latter part of the rainy season, but its timing is dependent on water levels and food; if the rains fail, breeding will not start or will be abandoned for the year. Old nests may be repaired, if available, or new platforms are built (Desai et al. 1977), in trees or tall bushes, often standing in or overhanging water. Some colonies consist of nests of only Painted Storks, in a single large tree such as a peepul (Ficus religiosa); we know of several such colonies, in northern India, in the centre of villages. At the large colony near Bharatpur, India, nests are in many low, scattered babul trees (Acacia arabica). Nests have also been placed in trees such as kandi (Prosopis spicigera), keli-kadamb (Stephegyne parviflora), and Barringtonia racemosa (S. Ali and S.D. Ripley 1968:94). For 15 nests in Delhi, it took 4-8 days (average = 6.3 days) to build the nest. Sticks 30-60 cm long are used for construction, with green leafy material as the lining. Males do most of the stick gathering, making 5-6 trips per hour in the morning and afternoon (Desai et al. 1977). The nest itself is a rather flimsy stick platform some 55-80 cm in diameter, lined with a small amount of leaves, straw and water weed. Fresh vegetation or leafy twigs are added throughout the occupation of the nest. Groups of unmated males often settle only temporarily on nest sites, and if they do not acquire mates, they move on to another tree taking unmated females with them. Such 'bachelor parties' (Kahl 1972b) may contain a dozen or more males and an equal number of females, and they form a conspicuous feature of any colony of wood storks at the beginning of the breeding season. Painted Storks, like other wood storks, perform a number of ritualized displays at or near the nest (Kahl 197la). In order to form pair bonds, the male stork occupies a potential nest site while Display Preening, and is approached by an unmated female. Displays are often of a high intensity and are more frequent at the commencement of breeding. In Display Preening the stork pretends to strip down its wing feathers with its bill, in a rhythmic pattern. The female

reponds by landing near, assuming the Balancing Posture, with wings widely spread and bill open, in a display called Gaping. She may maintain this posture for a half of a minute or more, before closing her wings, although still Gaping. Males defend territories with Aerial Clattering, Forward Clattering and Bill Snaps. Such rituals are common to all the wood storks, as is the Arching display given usually at the approach of a possible predator below the nest site, and which is often copied by nearby birds and spreads across the colony. In the Bill Snap, given by both sexes during pair formation, the bill is lifted to the horizontal and a single audible snap is given. Once a female has been accepted at the nest, the male performs a Flying Around display, circling around and landing again next to the mate. Up-Down displays usually follow such flights, as they do whenever a member of a pair returns to the nest site. The birds swing their heads and necks upwards to near the vertical, with the bill open. They then lower the head, swinging it from side to side. The Up-Down of the Painted Stork is similar to the other wood storks except that its softer hissing vocalizations are interrupted by double or triple Bill Snaps as the head is lowered. After pair-formation is complete, returning birds carry out mutual Up-Down displays together at the nest. In the Painted Stork, the Swaying Twig-Grasping plays a particularly significant part in early courtship, being especially ritualized in this species. On the nest-platform the bird bends over and touches or grasps lightly at twigs at intervals of 1-2 s. Sometimes the neck is swung and the body pivoted from left to right and back again, grasping a twig after each motion. Multiple bill snaps are often given between each movement. Display Preening is carried out often by several birds at the same time even though out of sight of one another. This action resembles true preening except that the bill is not closed on the wing-feathers that appear to be treated. Copulation occurs at the nest. The female holds her wings open and the male hooks his feet above her shoulders. The male clatters his bill loudly and shakes his head vigorously, beating the female's bill back and forth with his own. Eggs are dull white, sometimes streaked with brown; 2-5 but most commonly 3 or 4 are laid (S. Ali and S.D. Ripley 1968:94). In 90 clutches over a 6 year period at Delhi, the clutch-size varied from 1 to 4, with an average of 2.49/clutch (Desai et al. 1977). The eggs are laid 2-3 days, occasionally 4 days, apart; incubation starts with-the first egg and is shared by both birds. At Delhi, for 37 eggs over a 4-year period, the time taken to hatch was 28-32 days, with an average of 30.2 days (Desai et al. 1977). At Bharatpur, 6 eggs in 3 different nests had a 27- to 29-day incubation period (Breeden and Breeden 1982). Parents regurgitate food onto the floor of the nest and, when they are strong enough, the chicks pick it up. Small nestings are frequently shaded from the hot sun by a parent in the 'delta-wing' posture, with wings partially opened to the wrist (Kahl 197Ig). Young are seldom, if ever, left unattended until they are about 3 weeks old (Desai et al. 1974). Predation by the ever present house crows (Corvus splendent) and Pariah Kites

Painted Stork 57

(Milvus migrant) account for heavy losses. Lowther (1949:140) reported heavy predation on nestlings by a Pallas's Fishing Eagle (Haliaeetus leucoryphus) in the Mainpuri District, India. Young birds begin to exercise their wings after 25-30 days, but remain unable to fly until about 6070 days. They are usually fully independent by 85 days, but sometimes they will return to the nest up to 115 days, feeding largely on their own but still soliciting food from their parents (Shah and Desai 1975b, Desai et al. 1977). Success in rearing young is dependent on food availability, and this in turn depends on fish stocks and the water level. Over six seasons in Delhi, Desai et al. (1977) found that 68% of the eggs hatched and 64% of the young fledged, giving an overall success rate of 44% or 1.1 chicks/nest. Maturity is slow, with the plumage changing over 2 years. By about 16 months of age, young loose the ability to vocalize, and the adult plumage is attained at 3 years. Three marked individuals first bred at 4 years old (Shah and Desai 1975b). TAXONOMY Formerly classified under the genus Ibis, the Painted Stork is now combined with the other wood storks in the genus Mycteria. The inaccurate common name of 'Wood Ibis' is, fortunately, rarely used today. Even though apparently not closely related, Painted Storks have hybridized with Lesser Adjutant Storks (Leptoptilos javanicus) on several occasions. A male Painted Stork and a female Lesser Adjutant Stork mated and reared young several times in the Colombo Zoo, Sri Lanka (W.G.O. Hill 1943, A.P. Gray 1958:5), and another pair (sexes undetermined) did the same in the Kuala Lumpur Zoo, Malaysia, in 1974 (P. Olney pers. comm. 1974).

CONSERVATION The Painted Stork is one of the most widespread and least threatened storks of Asia (Luthin 1987), but its future survival depends almost entirely on human behaviour and conservation of its wetland habitat. Human behaviour, in turn, depends very often on religious attitudes. In India and Sri Lanka, the human population lives very much in harmony with wildlife, and in most areas active support is given to birds in traditional colonies. Indeed villagers take pride in 'their' birds. Elsewhere throughout its range, such as Bangladesh, Burma and Thailand, this stork has suffered not only from drainage of swamps and marshlands and the felling of nest-trees, but also from increasingly active hunting. Young and eggs are taken from nests both for food, and for profit. A good price is obtained by hunters, often professionals, for young birds to be kept as pets or sent to zoos. Increases in the human population throughout Asia, and especially in Southeast Asia, has shown an upsurge in this activity. Unless these and other large wetland birds are given protection by laws that can be enforced, the Painted Stork will become extirpated over much of its range. All nesting colonies require protection from disturbance. Because captive populations exist in Asian zoos (R.E. Johnson et al. 1987a), these can be used for stocking other zoos and for restocking wild populations, once protection is afforded. If left undisturbed, the Painted Stork can become fairly tame and nest in close proximity to people, even in their villages.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 287.

Asian

Openbill Stork Anastomus oscitans (Boddaert) Ardea oscitans Boddaert, Table des planches enlumineez d'histoire naturelle, 1783, p. 55: Pondicherry Other names: Shell-Ibis, Shell-eater, White Openbill (English); Le bee ouvert (French); Weiss-KlafTschnabel (German); Naththai-kuththi-narai, Karu-narai (Tamil/Sri Lanka); Bellan-kokka, Beli-kava (Sinhalese); Karu-tsoke (Burmese); Nok Pak Hang (Thai)

IDENTIFICATION The Asian Openbill Stork is a small pale grey to whitish stork, standing about 68 cm tall, having scapulars, flight feathers, some coverts in the wing, and tail black. Together with the African species, it is distinguished by its peculiarly shaped bill that has a narrow gap between the two mandibles, formed by the lower being curved downwards, then upwards, meeting the nearly straight upper mandible again at the tip. Adult plumage is predominantly white, with black in the wings and tail. The bill is a dull greenishhorny colour, mottled and streaked with black and reddish; the average gap between the mandibles in 14 adult specimens was 5.87 mm (Kahl 1971c). The iris varies from dark grey to brown, and the lores are blackish to dull blue. The legs and feet are dull flesh colour. Sexes are alike, but the male is marginally larger and its bill is longer and heavier. Before the onset of breeding, probably through a moult, the greyish plumage becomes immaculate white and the black feathers take on an iridescent purplish green. The white changes back to grey soon after the eggs are laid, in a process not well understood, presumably through feather wear and/or soiling (Kahl 1970). [The Oriental Crested Ibis undergoes a similar colour change from white to grey, which is apparently caused by the bird rubbing a 'black substance' from the skin of the face onto the feathers (Uchida 1970).] There is a band of bright orange skin, about 2.0 cm wide and 15.0-17.5 cm long, on the underside of the forearm (radius/ulna), which shows when the wing is opened (pers. obs. M.P.K.). The bill is basically horn-coloured and blacker at the base, but becomes mixed with shiny patches of reddish brown. The dull flesh-coloured legs and feet flush a deeper pink at the start of the breeding season. Small chicks have sparse, pale greyish down, tinged with brown, over blackish skin. Their gular skin is a bright pinkish purple (pers. obs. M.P.K.). Larger nestlings are a dirty grey, with black flight feathers in the wing and tail. Immature birds are a darker grey than adults and the head and neck are browner; they have a blackish brown mantle, wings and tail. The legs and feet are pale horn colour (often streaked with white droppings). The short bill is dark and has little or no gap until the bird is approximately 4-6 months old (Legge 1880:1103). In captive birds the gap continued to increase over several years, even though the birds had no access to shelled molluscs on which to feed (Kahl 1971c, H.G. Klos pers. comm. 1970). During courtship displays, birds utter a series of hollow, slightly nasal honking sounds (Kahl 1972d). This call has also been described as a 'mournful "Hoo-hoo"' (Loke in J. Huxley 1962).

60 Asian Openbill Stork

Normal flight is buoyant, with several flaps followed by a short glide; the flapping rate is 208/min (n = 44) (Kahl 1971b). Between breeding colony and feeding grounds, Asian Openbills soar on thermals and glide for long distances. When returning to the colony, birds often dive almost vertically at dizzying speeds, banking, slide-slipping, and gyrating violently (Krishnan 1960, S. Ali and S.D. Ripley 1968:96). The grey plumage and gaped bill are quite distinctive. The Asian Openbill is much smaller than the Painted Stork, with which it often nests. Its range does not overlap with the darker African Openbill. At a distance, it might possibly be mistaken for the larger, whiter White Stork, which occurs within the range during migration. DISTRIBUTION AND POPULATION The Asian Openbill Stork historically was widespread throughout tropical Asia, from the Indian subcontinent (India, Sri Lanka, Pakistan, Nepalese terai) through Thailand to Vietnam. Populations now occur rather patchily in suitable protected habitat (see Luthin 1987). It no longer breeds in Pakistan and, indeed, is only a rare visitor there (T. Roberts pers. comm.). Indian colonies are scattered, whereas several colonies occur in protected sites in Sri Lanka, such as in Wilpattu, Yala and Kalametiya National Parks. The once extensive Bangladesh population persists only along the coast in the Sundarbans and in Ghittagong. In Thailand, one large colony and several smaller ones are known. In fact, the largest extant colony is at Wat Phai Lorn, near Pathum Thani, about 40 km north of Bangkok. Other recently colonized Thai sites are at Wat Ampuwararam, Wat Tarn En and Ban Tha Sadet (Suphanburi Province); the former was undoubtedly established by a break off from the Wat Phai Lorn colony. The Asian Openbill is now rare in Vietnam, being known from only two sites. Information from Kampuchea is lacking. Stragglers occur in the Malay Peninsula, south to southern Thailand or beyond. On the Indian subcontinent, the species moves in synch-

rony with the varying distribution of seasonal monsoons. Henry (1971:384) reported seeing a flock of c. 100 entering Sri Lanka, at the Jaffna Peninsula, as if arriving from India. We have found that the Asian Openbill does not return to its breeding sites in Bharatpur, India, when the monsoons fail. At various lighthouses along the coasts of India, birds dash themselves against the dome of the lantern, suggesting that movement is sometimes at night (S. Ali and S.D. Ripley 1968:96). It is well documented that in Thailand, the Asian Openbill has a well marked migration route (McClure 1974:93). It arrives at Wat Phai Lom in October and November and disperses in April and May after nesting, except for a few hundred birds that remain as residents in the wet season. Some birds travel only to northern Thailand and a few to Kampuchea. But many migrate to the deltas of the Brahmaputra and Ganges rivers in Bangladesh, 1500km away. They can be seen along this route in large numbers; and, outside the regular periods of migration, the species occurs in small numbers at almost any place along the well defined route. This amazing migratory pattern was discovered during the extensive ringing programme in 1968 to 1970, under the direction of Dr H. Elliott McClure. Although reduced in geographic extent and overall numbers, the Asian Openbill remains the most abundant Asian stork (Luthin 1987). Its largest colony at the Wat Phai Lom Buddhist temple in Thailand numbers between 8000 and 14000 nests in a relatively small area (4.7 ha). In 1980, about 30 000 birds were counted including young, and the other Thai populations totaled another 200-1000 individuals. During the period of December 1988 to February 1989, a total of 20 358 Openbills were counted in India and an additional 4172 in Sri Lanka (D.A. Scott and P.M. Rose 1989:56). In the 1950s, the species was common throughout Bangladesh, but is now reduced to about 30 in the Ghittagong Hills and 50-70 in the Sundarbans (M.A.R. Khan 1984, Luthin 1984a). These areas continue to serve as the off-season home for many of the Wat Phai Lom birds from Thailand. ECOLOGY The Asian Openbill Stork is found in marshes, jheels, flooded rice-paddies, mangrove swamps, and less commonly along river banks or on tidal mudflats. Loss of natural habitat in Thailand has meant that this species has shifted to feeding in rice fields and the edges of canals and rivers, where apple-snails (Pila) are still plentiful. Commonly foraging is in small to medium-sized flocks. Feeding is by tactile or visual methods (Kahl 1971c). Snails are secured by the forcep-like bill, while the bird Walks Slowly or Stands in shallow water. The Probing bill is jabbed downward, and is quickly closed on any prey. The snail is extracted from its shell in exactly the same way as by the African Openbill, using the tip of the thin lower mandible to extract the snail body, leaving the relatively intact shell behind (Kahl 1971c). The common supposition (J.S. Huxley 1960) that the unique bill is regularly employed in the manner of a 'nut-cracker' does not seem to apply,

Asian Openbill Stork

especially in the case of the favourite food, large Pila snails. However, small snails are sometimes crushed between the mandibles (Henry 1955, Mukherjee 1974). Jerdon (1864:766) reported that blinded captives removed a snail from its shell without crushing it, and he gave a graphic description of the extraction process: The bird secured a shell by its feet, and after sundry alterations of its position, succeeded in cutting off the operculum as cleanly as if it had been done by a razor, but so rapidly that I was unable to see the exact way in which it was accomplished. It then inserted the tip of its clumsy beak into the open mouth of the shell, and after working it about for a short time, pulled out the entire shell-fish almost to its utmost tip. Many individuals have the extreme tip of their lower mandible curved slightly to their right (N.F.R. Snyder and H.A. Snyder 1969). This asymmetry may help the bird sever the columellar muscle of the apple-snail. Such asymmetry is not usually found in the African Openbill (Kahl 1971c), and it would be interesting to learn if this is because the Asian birds feed more regularly on snails that coil in a given direction, as suggested by N.F.R. Snyder and H.A. Snyder (1969). Although foraging is usually done in water, in Thailand Asian Openbills feeding nestlings during the peak of the dry-season sometimes follow tractors in dry fields and obtain aestivating snails turned up by the blade (P. Round et al. unpub. report). E.C.S. Baker (1929:334) describes the main diet as being molluscs, chiefly those of the large Pila snails, but the Asian Openbill occasionally eats bivalve molluscs, crabs, frogs, insects and other small animals found in its accustomed feeding marshes (S. Ali and S.D. Ripley 1968, McClure and Kwanyuen 1973, Breeden and Breeden 1982). In the Sundarban area of northeastern India, Mukherjee (1974) collected 72 adult Openbills for stomach analyses. He found that 85% of the wet-season diet consisted of molluscs (mainly Pila snails), but that in the dry season more crustaceans, fish, reptiles, insects, etc., were taken. An adult Openbill, collected at Bharatpur, India, in September 1967, contained over 150 small snail bodies (Kahl 1971c). In the Indian region, during normal rainfall seasons, Asian Openbills arrive in an area at the commencement of the rains, coinciding with the snails' re-emergence from dryseason torpor. In Thailand, birds arrive and breeding starts at the beginning of the dry season (P. Round pers. comm. 1990). At Bharapur, India, we found that in 1987, when no rain fell, the Asian Openbills did not appear at all, whereas in 1988, exceptionally heavy rains brought the storks back and they commenced nesting at their traditional sites. However, this nesting failed because of excessively high water levels, owing to supplementary water pumped from a nearby canal and river system. This additional water raised the levels, apparently too high for the birds to forage, leading to abandonment of the traditional site and movement to a new site. The disruption of traditional feeding areas caused the storks to leave their nests unattended, to feed. Their eggs were

61

then robbed by House and Jungle Crows (Corvus splendens and macrorhynchos) as well as by Pariah Kites (Milvus migrans), which had prospered in the drought year and whose numbers were correspondingly high. Additionally these delays and disruptions to the normal breeding pattern meant that Painted Storks had started to nest before the Asian Openbills were fully established, leading to heavy fighting between the species, and the resultant desertion by many of the much smaller Asian Openbills. We saw large parties of these birds feeding in small ponds and flooded fields well away from the colony, many still retaining bright courtship colours, including red legs. Thus nesting failed in two successive years, one because of drought and one because of excess water, exacerbated by artificial flooding. Thus it seems that many of the ecological requirements of the Asian Openbill Stork revolve around having suitable water supplies on its feeding areas.

BREEDING The species is highly social, its colonies being exceptionally dense, with as many as 62 nests in a single tree in India (Hume and Oates 1890:225) and 156 in a single Sterculia foetida in Thailand (McClure and Kwanyuen 1973). In India, this stork often breeds in large mixed colonies with other storks, ibises, spoonbills, herons and cormorants. However, the largest colony, in Thailand, contains only Asian Openbills. In India and Sri Lanka, they are usually the first wading birds to begin nesting at the beginning of the monsoon. In Thailand, however, nesting begins at the end of the rainy season and progresses throughout the driest time of year. Nest-platforms are placed in the tops of trees, often over water. In India, they sometimes use large trees, such as peepul (Ficus religiosa) or neem (Melia azadirachta) (Hume and Oates 1890:224), or smaller trees, such as babul (Acacia arabica] (S. Ali 1953:533). In Thailand, we have seen up to 10 nests on a single bamboo shoot. They were precariously sited and the close proximity of nests can lead to interference from neighbours. This over-use of all available sites is due to the need for a safe nesting area, provided only on the temple grounds where the birds are safe from human persecution (see Conservation). Bamboo nesting sites are subject to wind and storm loss and, as a result, productivity is low. Courtship behaviour is described in detail in Kahl (1972d). Unpaired males choose display sites, while females move around observing their displays. In defence of its site, a bird will use Forward Clattering, very subtle Bill Snaps, and Arching Displays. Sexual displays include Swaying Twig-Grasping, Advertising Sway and the Up-Down. The latter greeting display is often given by both members of the pair, especially after the return of one to the nest. In this display, the bird stands erect, arching its neck forward and pointing its bill horizontally or slightly upward. It then moves its head slowly downward and utters a series of nasal honking sounds. The Up-Down often grades into Swaying Twig-Grasping, and is preceded and followed by Bill Snaps. During Copulation Clattering, the clattering sound comes

62 Asian Openbill Stork

from the male vigorously slapping his bill against that of the female. Pair-formation and behaviour during breeding are generally similar in the Asian and the African species. However, in hostile situations, the African birds erect the feathers of the crown and occiput, whereas the Asian birds are less prone to do so. Head shaking and quick rubs of the crown on the shoulders and upper back are also less common in the Asian birds. The Advertising Sway has not yet been reported for the African Openbill. The displays of both species are somewhat less ritualized than in some other storks (Kahl 197If), though the Up-Down greeting displays of the openbills are quite similar to those of the wood storks. Both sexes cooperate in the building of the nest, with the male collecting most of the sticks. Birds make up to 30 trips per day with material, and nests may take as long as 11 days to build (Mukhopadhyay 1980). Five nests measured 100125 cm in diameter (Mukhopadhyay 1980) and were built of small sticks and lined with leaves and other vegetation. Fresh foliage is brought to add to the nest even during incubation, possibly to provide moisture. Between 2 and 4 eggs are laid, rarely 5. They are dull white and are broad oval in shape, slightly compressed at the smaller end (E.C.S. Baker 1929:334). Both birds share incubation. Estimates of the incubation period vary from 24-25 days (S..A11 and S.D. Ripley 1968:97), 27-29 days (McGlure and Kwanyuen 1973), to 28-30 days (Mukhopadhyay 1980); we believe the longer periods (27-30 days) to be more accurate. Regurgitation of water over eggs was noted in India (Kahl 1970, Mukhopadhyay 1980) but it has not been reported in Thailand. On rare occasions, 3 adults have been observed tending a nest (B. Amget pers. comm., pers. obs. M.P.K.). Small chicks are shielded from the sun by the parents standing on the edge of the nest covering them with open and drooping wings (often not as erect as Painted Storks). Nestlings are showered with water by their parents during the hot part of the day (Breeden and Breeden 1982, B. Amget pers. comm.). Both parents feed the young by regurgitation into the nest, but some chicks will grab at the adult's bill and catch the food before it reaches the floor of the nest. Mukhopadhyay (1980) reported that chicks were fed 5-6 times per day. Predation on eggs and young in India is by monitor lizards (Varanus sp.). Jungle Crows (Corvus macrorhynchos] and several species of birds of prey. Breeding success at a colony in northeastern India was about 50% (Mukhopadhyay 1980). At the large Wat Phai Lorn colony, in Thailand, usually only 2 young survive per nest; in years without wind storms, the survival is higher. Predation is neglible there, but the nestlings are victims of parasitism by various ticks, flies and mites (McClure and Kwanyuen 1973). First flights are at about 35-40 days, and the young finally leave the nest at about 6-8 weeks (McGlure and Kwanyuen 1973, Mukhopadhyay 1980). At Bharatpur, India, fledged young are active in the nesting areas often before other species, including Painted Storks, have finished incubation. When really ready to leave the nest, the young flap their wings frequently, and when fully fledged, they gather in groups away from the nest.

TAXONOMY The Asian Openbill is very closely related to the African Openbill Stork, from which it mainly differs in colour. The two openbills form a natural group, here designated as a tribe, with the four wood storks of the genus Mycteria. Similarities in many of their courtship displays indicate this close relationship (Kahl 197If, 1972b, 1972d, 1972e). CONSERVATION Toleration by humans of nesting waterbirds in India and protection of temple sites in Thailand have served the species well. However this tolerance is not matched in Pakistan, where it now no longer breeds because of continual persecution, or in Bangladesh, where a tradition of hunting goes back to the British. Trapping remains a problem there. In Thailand, protection is given by Buddhist monks in their temple grounds at Wat Phai Lorn, but the large population of birds and the accumulated excreta have killed a number of the trees and are threatening the others. Inkapatanakul (1986) found that the heavy use of bamboo was due to the loss of other trees. It has even been suggested (Sitwell 1984) that the species of bamboo used by Asian Openbills at Wat Phai Lorn (Bambusa arundinaced) is due for a major die-off about 1993, which could be disastrous for the storks nesting there. Drainage of natural marshes in Thailand has led to the birds seeking food in cultivated fields, and the danger from pesticides has increased. The long flight path on migration, over what has become hostile territory, results in heavy losses particularly of first-year birds. Whilst Indian and Sri Lankan birds are afforded unmolested breeding areas, they are subject to drought in some years, artificial flooding in others, and a lack of alternative colony sites. Other breeding populations in Southeast Asia are poorly documented, but are certainly in trouble. Various evidence from the biology of the species demonstrates its dependence on particular hydrological conditions. With both drought and extreme artificial flooding causing nesting failure, water management in feeding sites becomes critical. The Asian Openbill Stork is in fact a species that could benefit from active management throughout its range. Temple sites are particularly suitable for active management. Remnant colonies could provide stock for repopulation, if productivity were increased and additional nesting and feeding sites provided. All colony sites need to be provided complete protection from both poaching and disturbance (Luthin 1987). Nesting Asian Openbills are in competition for trees with wood cutters; additional sites can be provided by protecting tree stands, and present sites can be enhanced through tree planting and construction of artificial nest sites (Inkapatanakul 1986). Formerly, river flooding washed excrement away annually; now flood control has prevented this and the birds' excrement accumulates. In restricted areas where nutrient accumulation is killing trees, some dilution of nutrients may be feasible. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 288.

African

Openbill Stork Anastomus lamelligerus Temminck Subspecies: Anastomus lamelligerus lamelligerus Temminck Anastomus lamelligerus Temminck, 1823, Planches Coloriees, livr. 40, pi. 236: Senegal Anastomus lamelligerus madagascariensis Milne-Edwards Anastomus madagascariensis Milne-Edwards, 1880, Compt. Rend. Acad. Sci., Paris, 91, p. 1037: Madagascar Other names: Claw-billed Stork (English); Bec-ouvert africain (French); Schwarz-Klaffschnabel (German); Masoklo (Luo); Etongorokofu (Kwangali); Oopbek-ooievaar (Afrikaans); Famakacora (Malagasy)

IDENTIFICATION The African Openbill Stork, a medium-sized stork, standing 55-60 cm, is an all black bird with a stout, distinctively shaped bill, which is open for part of its length. The peculiarly shaped bill, from which it gets its generic and common names, is a brownish horn colour but paler, often a dull yellowish tan, basally. The upper mandible is essentially straight, but the lower is curved upward, leaving a gap between the two, which meet again at the tip; the average gap in 16 adult specimens was 5.9 mm (Kahl 1971c). The lores and the skin around the eye are dull-bluish to blackish. The legs and feet are black and the eye is brown. The iris is dusky brown. The sexes are alike, but the male is on average larger, 1250 g compared with 1000 g for the female, and the male's bill is larger. The dark plumage is enlivened during the breeding season with shiny, sequin-like feathers of iridescent metallic brown, black, green and purple, most pronounced on the abdomen, owing to highly modified feather morphology (Stettenheim 1976). Those on the breast and abdomen are curly and jetblack, whereas those on the back and wing-coverts are flattened and green, brown or bronze (Mackworth-Praed and Grant 1973:67, pers. obs. M.P.K.). Newly hatched chicks weigh 35-40 g (n = 6) and are covered with a sparse, wispy grey down on the back, wings, and head. The skin of the head and neck are pink and that of the body 'almost purple' (Anthony and Sherry 1980). The bill is black, with the distal third cream-white, and becomes mostly black by about 25 days. The legs and feet of small nestlings are pinkish-orange, later fading to fleshypink and then dingy grey. The iris is dusky brown. At about 10 days a dense, sooty black down emerges; at this stage the naked gular pouch is a bright pink (pers. obs. M.P.K.). Immature birds are duller with brown feather-edgings below and on the back and upperwing-coverts, and the back of the neck speckled white. The bill is blacker, lacks the paler base and initially is shorter and lacks the gap of the adult (Anthony and Sherry 1980). Typical bill configuration develops slowly, possibly taking several months. The African Openbill is usually silent, except at the breeding colony. Courting adults utter loud, sonorous, rather raucous croaks or honks, 'Horrrh-horrrh', audible at up to 100 m. During copulation, a

African Openbill Stork

clattering sound is produced by the male beating his bill against that of the female. The Begging call of nestlings is similar to the Yellowbilled Stork, but higher-pitched and more nasal (Brown et al. 1982:176, pers. obs. M.P.K.). In flight the tail appears broad and rounded and the feet extend beyond it (Bannerman 1930:103). Often birds soar on thermals, especially during long-distance movements (Ruwet 1963). In flapping flight, a series of flaps alternates with a short glide; the flapping rate averages 201/min ( » = 18) (KahI1971b). Its gaped bill makes the African Openbill unique and easily distinguishable from other species. At distances too great to discern the bill, it could be mistaken for other dark waterbirds, such as the Glossy Ibis or Black Heron (Egretta ardesiaca). It is geographically isolated from the lighterplumaged Asian Openbill Stork. DISTRIBUTION AND POPULATION This wide-ranging stork is found in suitable habitat throughout the African tropics south of the Sahara. Most of the breeding seems to take place in the Southern Hemisphere. It mainly occurs below 1500 m elevation. In West Africa, there is a report of breeding only from Sierra Leone (Bannerman 1953:175). It has nested on occasions and in small numbers in Chad and Ethiopia (Salvan 1967, SchaufFele and Schiitz [sic] 1968). In East Africa, it is widespread through Kenya and Uganda, breeding erratically at Garsen, Kisumu, Rwenzori National Park, Busoga and Jinja. In Tanzania it breeds in large numbers at Utenge and Wembere and in smaller numbers elsewhere (Britton 1980:20). Breeding colonies have been found throughout Zambia in all months of the year. In southern Africa, this species is uncommon south of the Zambezi (Clancey 1964a) and breeds in Zimbabwe, northeastern Botswana, northern Namibia, and northern South Africa (Dutton 1972, Braine

65

1974, Newman and English 1975, Tree 1978, Anthony and Sherry 1980), In Madagascar there is a separate race, A.I. madagascariensis, distributed in the western parts of the island, including the highlands (Bonvallot and Randrianasolo 1975, Langrand 1990). The African Openbill wanders considerably within the wetter parts of the African tropics, probably moving back and forth across the equator. Its movements are not well known, but it becomes locally abundant when conditions are right. It is primarily a non-breeding visitor north of the equator and is more common in the east than in the west. It arrives in the dry season, presumably from breeding areas in the south. Large flocks gather at Lake Turkana, Kenya, each November-December, suggesting well-established movements from breeding sites in the Southern Hemisphere (Britton 1980:20). We have seen it in a mixed colony on the Tana river in January 1983, when it took advantage of exceptionally heavy short rains to breed, but in other years it was absent in the same area, when the short rains failed (Hancock 1984:41). The population is probably stable (Brown et al. 1982:176). The African Openbill is locally numerous yearround in East Africa, especially near breeding and migratory sites, and may be extending its range there; flocks up to 7000 have been reported (Kasoma and Pomeroy 1987). It is the most numerous stork for part of the year (Dec-May) in Sudan but is uncommon in the highlands of Kenya. In Tanzania, breeding colonies can number in the thousands. The largest reported colony there was of 5000 or more nests (Kahl 1968), but this may have been too high an estimate (Stronach 1968). O. Langrand (pers. comm. 1989) reports it to be common in Madagascar, between Mahajanga and Morombe, particularly near Bemamba Lake and Morondava where groups of 15-20 birds are commonly seen. Conversely it no longer is seen at Alaotra Lake. ECOLOGY Characteristically a floodplain species, the African Openbill Stork is found mostly in freshwater wetlands, and not usually in heavily forested areas. It requires shallow water, such as ponds, streams, rivers and temporarily flooded marshes; it will sometimes also forage on drier ground if appropriate food is present. In West Africa, it is occasionally seen on coastal mudflats (Bannerman 1953:175). Its main habitats in Madagascar are shallow lakes, rice paddies, and occasionally marine coasts and mangrove swamps (Langrand 1990). At other times it travels in large flocks and can be found in any habitat, including ploughed fields (Britton 1980:20). This species feeds either alone or in groups, or with Sacred and Hadada Ibises (FJ. Jackson 1938:74). The foraging strategy involves both tactile and visual feeding. Both day and night feeding have been reported (Clancey 1964a:46). Birds will probe deeply with bill open, often with head submerged, to capture snails in ponds and shallow lakes, which are often covered by floating vegetation. They may Stand in one place repeatedly Probing before moving to

66 African Openbill Stork

a new spot. We have seen up to 50 birds feeding close together in one such lily pond in East Africa (Hancock 1984) and riding on the backs of hippopotamuses to catch prey disturbed by these animals (Kahl 1971c). When feeding in groups, mutual harassment and attempted preyrobbing have been reported (Niven and Niven 1966a). Early authors suggested that the unique gap in the bill of this species was used to carry and then crush the shell of the snail (Chapin 1932a:472, F.J.Jackson 1938:74, J.S. Huxley 1960). We have found, however, that the snail is most often held in the tip of the bill, and the extraction accomplished by holding the shell against the substrate with the distal end of the upper mandible. The tip of the knife-thin lower mandible is then forced under the operculum, severing it from the body. The stork then shakes its head to release the body of the snail from the shell, which remains essentially intact. This process can take as little as 15 s (Kahl 1971c). Mussels are opened in a similar manner or by holding and squeezing until the animal opens its shell (Root 1963). In the latter case, when a mussel is caught, it is carried to the shore where piles of discarded shells can accumulate as the same site is used over and over again (Kasoma and Pomeroy 1987). Such specialized actions are aided by unique quadrate bones in the skull which have one narrow condyle and assist this form of feeding. Comparison between the bill shapes of openbills and other snail-eating species is discussed by J.S. Huxley (1960). Because openbills must spend some time manipulating their prey before swallowing it, other species, such as gulls, may sometimes attempt to steal it (Steyn 1988). The African Openbill Stork is a highly specialized species that depends almost entirely on molluscs. Heuglin (186973:1121) considered the overall diet to be broader, including fish, frogs, crabs, locusts, beetles, worms, mussels and snails. Since then, most other observers have mentioned only snails and freshwater mussels, including the genera Lanistes, Limicolaria, Achatina, Pila and Mutela (references in Kahl 1971c, Anthony and Sherry 1980). While some of the items of food described by Heuglin may be occasionally taken, it is clear that this stork feeds almost exclusively on mussels and snails, both from fresh water and dry land, with snails of the genus Pila predominating. It is the emergence of these snails, brought about by the onset of the rains, that triggers the arrival of these birds in their breeding area. Apparently the African species eats more mussels than does the Asian species (J.S. Huxley 1960, 1962), but they are not always successful in opening all the mussels that they catch (Root 1963). Many hours of observation have been spent without seeing these birds catch anything except molluscs (Bell-Gross 1974, pers. obs. M.P.K.). Captive-reared nestlings strongly favoured a diet of mollusc bodies, and refused other food (e.g. small fish) even when ravenously hungry (Kahl 1972f). A captive 15-dayold nestling weighing 148g consumed 143g of snail bodies (nearly 97% of body weight) in one 12 h period. At 37 days of age, a nestling weighing 720 g ate 243 g of snail bodies in 15-20 s and still begged for more (Kahl 1971c). The African Openbill is a very gregarious stork, in its breeding, foraging, flying and roosting. Often they arrive in

a large flock and then disperse to feed individually. Very heavy rains, with the resultant flooding and abundant supply of snails, often result in African Openbills suddenly appearing in an area and starting to breed. In years of exceptional flooding, breeding may span long periods (Anthony and Sherry 1980). It has been reported that this species is sometimes used as an animated feeding-perch by Carmine B.ee-eaters (Merops nubicus) (Cunningham-Van Someren 1970). BREEDING African Openbill Storks breed colonially, often alongside ibises, spoonbills, herons, darters, cormorants and other storks. They are opportunistic breeders, and nesting occurs only in years when local food supplies are plentiful; it does not occur regularly in the same place. Nesting is in trees or bushes, often over water, and sometimes as many as two dozen or more nests of this species are found in a single tree. Most populations of this species nest in the rains, at the height of flood-plains inundation. Commencement of breeding is probably related to the emergence from aestivation of snails (Pila). Males stand at the nest site and are approached by unmated females. Courtship and pair-formation are basically similar to the processes in other storks (Kahl 1972d). Males secure nest sites and perform a variety of displays. One prominent display in the courtship of the Asian Openbill is the Advertising Sway; this display has not been confirmed in the African species —contrary to the statement in Brown et al. (1982:177)—but probably does occur. Commonly given is the Up-Down display, which involves an arching of the neck and gaping of the bill; raucous calls are uttered but no bill snaps are given. The Swaying TwigGrasping display is often performed. The Bill Snap is frequently given early in pair-formation but less after pairing is completed. Forward Clattering is given by males defending sites from other males, and by females threatening other females. Disturbance near the nest by animals or humans elicits the Arching display, and this serves to warn other birds of danger. Copulation Clattering is accompanied by the male shaking his head vigorously and striking his bill against the bill of the female; he does not clatter his own bill, as most other species of storks do. Nest platforms are made of twigs and lined with grass, leaves, sedge, various water plants; the inner lining is often damp. Nests measure 45-75 cm in outside diameter and about 20cm across the inner cup (Rand 1936:341, Heiden 1973, Anthony and Sherry 1980). They are built either by repairing an old nest or building afresh with sticks and reeds, and lining with weed collected from the bottom of a pond. We have seen this carried out by birds bringing quite large dripping piles of weed, even after incubation has started, and the nest is kept wet throughout, sometimes by regurgitating water. Loud calls are given by both birds at the nest during these actions. Initial nest building or repair spans about a week before egg-laying (Anthony and Sherry 1980). Eggs are laid at 2-day intervals; 28 clutches varied from

African Openbill Stork 67

3-5 eggs, averaging 3.9 (Maclean 1985:68). They are dull white and become heavily stained. Both birds incubate, often with the partner standing nearby. Incubation has been estimated as 25-30 days (Brown et al. 1982:177). Egg predation has been reported by a Gymnogene or African Harrier Hawk (Polyboroides typus) from Zimbabwe (Anthony and Sherry 1980). Nestlings have shorter, straight bills, but possess a wide buccal cavity, which enables a nestling weighing 45 g to swallow large snails up to 5 g. Begging is done with open wings flapping and slightly nasal cries. Often the bill is pointed upwards exposing the pink gular skin of the chick. We found that captive nestlings usually refused food other than mollusc bodies even when ravenously hungry (Kahl 1971c). Parental feeding of young is accomplished by regurgitating food on to the floor of the nest. Parents often regurgitate water over the nestlings. Flight feathers begin to appear at about 25 days, and the body is fully covered with contour feathers by 40 days. The fledgling period is estimated to be 50-55 days (Brown et al. 1982:177, pers. obs. M.P.K.); that of 80 days, given by Anthony and Sherry (1980) and Maclean (1985:68), is almost certainly too long. Nesting success varies greatly with feeding conditions, from complete failure to 2-A young/ nest (Brown et al. 1982:177). TAXONOMY The two openbill species are closely related to each other, morphologically and behaviourally. The Up-Down and Copulation Clattering displays show close resemblance to homologous displays seen in the several species of wood storks (Mycteria). The most significant morphological differences between the openbills and wood storks involve the cranium and bill (Beddard 1901, Verheyen 1959, Kahl

1971c). Because of their similarities to each other, and differences from the other storks, we have combined the wood storks and openbills in the tribe Mycteriini (Kahl 1972e). The separation between the two subspecies is based on the Madagascar bird having more pronounced longitudinal ridges on the bills of adults and more white spots on the necks of juveniles (Bangs 1918, Chapin 1932a:470, Bannerman 1930:103); some authors report the Madagascar bird as larger, but the two specimens we measured in the Leiden Museum were smaller (see Appendix). Sharpe (1898:309) saw no reason to separate the Madagascar population. We tend to agree, but, pending further study, have retained it tentatively as a separate subspecies. CONSERVATION The African Openbill Stork is locally common and can form flocks numbering over 7000 in some seasons. In Africa, its population is considered to be stable, in part because its colonies are smaller and more widely spread than are the breeding sites of the Asian Openbill (R.E. Johnson et al. 1987a, Luthin 1987). The species may actually be expanding its range in eastern Africa. In Madagascar, it was common in the lowlands (Rand 1936), but in recent years the Madagascar population has been reduced. Its elimination from such places as Alaotra Lake reflects the increase in human population there, where hunting and poaching continue to take place.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 289.

Black Stork Ciconia nigra (Linnaeus) Ardea nigra (Linnaeus, 1758, Syst. Nat., ed. 10, p. 142: northern Europe ( = Sweden) Other names: Schwarzstorch (German); Cigogne noire (French); Zwarte ooievaar (Dutch); Svart stork (Swedish); Sorte Stork (Danish); Ciguena negra (Spanish); Bocian czarny (Polish); Ghyorny aist (Russian); Grootswartooievaar (Afrikaans); Endongondongo (Kwangali); Mokoroane (South Sotho); Unocofu (Xhosa); Gandahari, Gamdayaki, Gamyaki (Hausa); Surmal (Hindi); Hei-kuan, Wu-kuan, Hei-chuchi (Chinese); Hyun hak (Korean); Nabe-koh, Kuro-koh (Japanese)

IDENTIFICATION This medium-sized stork, standing 90-100 cm tall, is entirely glossy black save for the white of its lower breast and abdomen, which extends to the long undertail-coverts that form a frilly fringe to the black tail. The adult's head, neck and upper wings are glossed with metallic purple and bronze, and its back with an iridescent green. The tail-feathers are slightly duller and browner. Feathers of lower foreneck are elongated, forming a rather shaggy neck-ruff that can be erected in displays or when the ambient temperature is low. The bare skin around the eye, the bill and the legs are red; the iris is brown. The bill and legs are red. The sexes are similar in plumage, although the male is larger and has a longer, heavier, and sometimes more recurved bill (see photo in Siewert 1932b:33). Birds moult into fresh, bright plumage in the spring (Dementiev and Gladkov 1951:444), and their iridescence is stronger, especially on the head, neck and wing-coverts of the male (Siewert 1955:67, pers. obs. M.P.K.). During courtship, the red skin of the face, bill and legs deepens in colour to a rich scarlet. The area of red loral skin expands during displays and contracts when the bird is relaxed, so that the bill and orbital-skin are separated by black feathers in non-displaying birds (pers. obs. M.P.K.). After the breeding season, these soft parts take on a darker, less brilliant, or more brownish tint (Dementiev and Gladkov 1951:445, Cramp 1977:328). Some sources state that a complete annual moult lasts from May-June through winter (Naumann 1838:283, Cramp 1977:328). Nestlings are covered with a sparse down at first, which is superseded by a thicker down later; it is white with a greyish hue and quickly becomes soiled. Their bill is yellow, and the legs and feet are pink or flesh-coloured; later the bill is orange-yellow at the base, grading to pale olive distally, and the legs are pale olive. They develop full contour-feathering at about 2 months. The progression of the nestling's development is illustrated by photographs in O. Heinroth and M. Heinroth (1924-3 l:fig. 170) and described in more detail in Schroder and Burmeister (1974:6-7). After fledging the immature bird is browner, without the iridescence of the adult. The neck and wings are duller, flecked, especially on the lower neck, with lighter brown feather-tips, and some back feathers show a slight greenish tinge. The bill and legs are pale grey-green, as is the facial skin. The bill and legs become red in the spring of their second year, i.e. when about 10 months old (Cramp 1977:328). The adults have more developed vocal calls than White Storks —usually a whistling or prolonged piping or hissing—but their bill-clattering is much less developed. This could be because in the Black Storks' more secluded nesting sites, short-range communication between mates is more important than long-range threats against intruders. The bisyllabic calls have the sound of a sibilant inhale/exhale and are given at a rate of about 2 couplets of whistles per second (pers. obs. M.P.K.). Their vocalizations have been rendered as 'Cha-lie, cha lie, cha lie' (Siewert 1932a, 1955:75), or 'Huji-ji', 'Chi-chu',

70 Black Stork

Tleeeee, he fleece, hefleeeeehe'or 'Hhiio . . . hhiio' (Cramp 1977:327). A melodius flight call, *Fuo' is rarely heard (Bauer and Glutz von Blotzheim 1966:416), but birds migrating at night are heard calling (Collman and Croxall 1967). Various begging calls of the nestlings are described in Cramp (1977:327; further details are in Schroder and Burmeister 1974:26-27). Black Storks are agile fliers and able to manoeuvre nimbly between the branches in dense forest. The Black Stork is said to fly with faster wing-beats than the White Stork (Bannerman 1957:22). However, upon measurement this is found to be not the case; in normal, level flight, the average flapping rate of the Black Stork is 159/min (n = 22) versus 170/min for the White Stork (Kahl 1971b). Although it does soar and glide on long-distance migration, this species does not so readily soar on thermals as does the White Stork; it is said that this is because the Black Stork has narrower wings (Cramp 1977:324). On migration it usually moves in small groups and does not often travel with flocks of White Storks; it does often migrate with birds of prey (Porter and Willis 1968). Some migratory movement takes place at night (Dementiev and Gladkov 1951:440, Collman and Croxall

1967). Especially near the nest, aerial acrobatics have been reported and mild aerial hostile encounters with whistling vocalizations occur (Fincham 1971). The Black Stork can be mistaken at first sight for Abdim's Stork in Africa, but the latter species is much smaller, with white on the lower back and rump, and has greenish legs and bill. The Black Stork can be clearly distinguished from the Woollynecked Stork, with which it is frequently seen during migration in northern India, by its black neck, red bill and legs, and larger size. DISTRIBUTION AND POPULATION The Black Stork has the most extensive breeding range of any stork. It does or has occurred throughout much of the Palaearctic. It ranges from Europe—discontinuously from Scandinavia, Germany, and Portugal—eastward through the former USSR and Mongolia to the Maritime Territory, and northeastern China. It also breeds in southern Africa, from Zambia and Malawi to Cape Province. Within this overall range, it is thinly spread and widely dispersed and in many areas has been declining. It has been

Black Stork

extirpated from its former breeding grounds in southern Sweden, Denmark, Finland, the Netherlands, and, with the exception of a few odd pairs, from southern and western Germany, France, Greece, and Korea (Loiseau 1977, Luthin 1984a, 1987, Sonobe and Izawa 1987:43). Significant nesting still occurs in Portugal, Spain, eastern Germany, Hungary, Poland, Czechoslovakia, USSR, and possibly several other countries of eastern Europe. In the USSR, the species is found as far north as along the Baltic coast near Leningrad, Vologda and Kirov; in Siberia on the eastern slopes of the Urals it extends north to 61°N. It occurs patchily across much of the eastern USSR south of 60-63°N—in the mountainous and forested regions, but not in the steppe areas—to the Pacific Ocean (further details in Dementiev and Gladkov 1951:437, Vaurie 1965:85). It nests in the northern portion Mongolia, down to the Gobian Altai; and across much of northern China, westward to Sinkiang (Etchecopar and Hue 1978:74; Meyer de Schauensee 1984:138). In northeastern China, it may breed (formerly at least) south to about the vicinity of Beijing (T.H. Shaw 1936:124). It used to nest in southeastern Korea, near Andong, Kyongsang-Pukto (Austin 1948:46), but has not been reported to do so since 1966 (P. Won 1973:18); during the mid-1980s Black Storks were observed throughout the breeding season in northeastern Korea (Hamgyong-Pukto) and may be nesting there (Sonobe and Izawa 1987:44). Several Black Storks were reported in a colony of Black Kites (Milvus migrant) and Grey Herons (Ardea cinerea) near the Caspian coast in northern Iran during April and July, but it is not clear whether the storks were breeding (Passburg 1959). A nesting site is known from central Iran, in a wetland near Isfahan (Luthin 1984a). Tom Roberts (pers. comm.) considers it possible that one or two pairs might nest in the Tobakhar mountains of Baluchistan, northwestern Pakistan, and J.A.W. Anderson (pers. comm.) claims to have had 2 young newly fledged birds brought to him at Quetta, Pakistan, in 1969 by hunters who found a nest in that area. However, it is unlikely that they now remain, as hunting pressures on breeding and migratory birds of all species has intensified in recent years in Pakistan. There are several summer sightings from central Afghanistan but no definite evidence of breeding there (Paludan 1959:66). Migration Most Black Storks are migratory, but in a few cases populations are dispersive or sedentary. During its extensive migrations, the species occurs throughout much of the African and Asian tropics. The majority of eastern European birds migrate to Africa, whilst those from western Asia winter primarily in northern India, and those from further east apparently winter mainly in southern China. Some of, the Iberian population, as well as those in southern Africa, are essentially non-migratory, although they may wander widely in the non-breeding season. Although mainly migrating along defined routes, the Black Stork is apparently somewhat less dependent on soaring than the White Stork. Some passage birds have been reported from Italy, the Aegean Sea and various Mediterra-

71

nean islands, and they will cross wider stretches of water than White Storks normally do (Bauer and Glutz von Blotzheim 1966:421, Cramp 1977:324). Birds leave Europe, except from Iberia where they are mainly resident, from early August to October, with the greatest exodus in September. Nearly all European birds travel to Africa. Relatively few travel via Gibraltar and visit West Africa, because so few still breed in western Europe. Moreau (1967) reported a few birds in Nigeria and Chad, and some were seen in the Senegal Delta by Roux (1972). There are scattered reports from several other West African countries (Bouet 1955a). There is a small but regular spring passage northwards from Tunisia. The route around the eastern end of the Mediterranean is used by many more Black Storks. At the Bosphorus, a total of 6194 passed between 16 August and 25 October 1966, and over 7400 birds were counted between 8 September and 3 October 1973 (Porter and Willis 1968, Cramp 1977:325); the peaks came in late September. A total of 203 were seen in a 3-h period near the Bosphorus in March 1963 (Kumerloeve 1966b). Parties of up to 500 birds are reported from Israel in both spring and autumn; and, on their journey north, 3500 were counted 26 March 1981 above the Valley of the Moon near Eilat. Formerly a rare winter visitor to Israel, it has been increasing in recent years, possibly because of the increased number of artificial fish-ponds. As many as 290 Black Storks remained in the Hula and Bet She'an Valleys in the winter of 1982-83 (Paz 1987:34). A few birds winter as far north as Turkey (Vittery 1972). Birds heading for Africa apparently travel down Sinai and the Red Sea coast, for they are said to be rare in the Nile Valley (Cramp 1977:325). Some birds have been observed on the Red Sea, near Jiddah, and in small flocks near Aden, but no evidence is available to show the origin of these migrants. Wintering birds are recorded regularly in small numbers south in Africa to the Sudan (about 100 annually), Kenya and northern Tanzania, and these are most likely of European origin (Backhurst et al. 1973); birds in southern Tanzania may be of southern African origin (Britton 1980:21). Ringed birds from northeastern Europe have been recovered in Ethiopa and Uganda (Schiiz 1940b, Bauer and Glutz von Blotzheim 1966:422). There is one report of a bird ringed in Germany taken as far south as Malawi [formerly Nyasaland] (Belcher 1930:12); however, we feel this was an error, probably referring instead to the White Stork (see also C.W. Benson 1958). And we agree with Prigogine (1976) that reports of 'large flocks' of Black Storks in tropical Africa (e.g. V.G.L. Van Someren 1922, Verheyen 1953) almost certainly refer to the Abdim's Stork. Birds enter Pakistan on a broad front across northwest Baluchistan and via the northwestern Himalayas, and a flock of over 100 has been recorded in Gilgit in the autumn (TJ. Roberts pers. comm.). TJ. Roberts (1969) reported scattered sightings of birds in Sind and, in January 1969, saw 81 birds in the Punjab of Pakistan, and they have been noted in the Kurram valley. Certainly parties of Black Storks winter in northern India regularly. They have been seen at least as far south as 14-15°N in India (Madsen

72 Black Stork

1990), and once 2 birds were reported in Sri Lanka (Henry 1971:386, W.W.A. Phillips 1978:9). We saw a group of 7 birds, including 1 immature, in the Ranthambhor National Park in March of 1989, in the company of a pair of Woollynecked Storks. Wardens there reported seeing flocks of up to 20 birds on passage through this tiger reserve. Black Storks from the eastern USSR and China winter mostly in southern China (Cheng 1976:34). In Hong Kong, the Black Stork is an irregular and scarce winter visitor, with a few individuals arriving for short periods, but not every year (Chalmers 1986:41). It is an uncommon visitor further south, in Burma, northern Thailand and Laos (Delacour and Greenway 1940, Smythies 1953:521, B.F. King and E.G. Dickinson 1975:51, P. Round et al. unpub. report). In Korea, the Black Stork is a rare and local summer visitor, occasionally wintering in the south of the peninsula (Austin 1948:46, Gore and Won 1971:119). Generally this stork migrates in small flocks, but individual birds disperse nomadically and have been recorded from as far afield as Britain, Norway, Finland, Switzerland, Malta, Cyprus, Madeira, Taiwan and Japan. Old ringing records .show that from a brood of 4 ringed in Denmark, 2 were recovered southwestward in the Netherlands and northern France, and 2 southeastward in Hungary and Romania. 2 young from an East German nest were recovered in France and Hungary (Cramp 1977:324). Birds ringed in Latvia were recovered in Poland in September, in southern Asia Minor in November, and in Egypt in March of the following year. Birds ringed in Lithuania were recovered in Hungary and in the central Soviet Union. One ringed in the Kaliningrad, USSR, region was recovered in Peloponnesus (Greece) (G. Niethammer 1938). Spring migrants arrive back at their nesting areas between March and May, mostly in April. In many areas they return about 1-2 weeks later than the local White Storks (G. Niethammer 1938); but in southwestern Poland in the spring of 1969 Black Storks preceded the White Storks by a few days (pers. obs. M.P.K.). Populations and trends Populations are small; the Black Stork is nowhere common, and its numbers are diminishing everywhere. In the USSR, the Red Data Book describes its status as widely distributed but rare and numbers decreasing everywhere. The total population in the USSR is unknown, but no more than 1QO birds breed in the western Ukraine. In Estonia, in 1970, it was estimated that at most 80 pairs nested, compared to 150 recorded in 1962. In Latvia, in the 1950s, there were 260 nesting pairs. About 200 pairs are said to inhabit the Khabarovsk region of the extreme eastern USSR (Sonobe and Izawa 1987:43). Annually 2-3 pairs nest in Qianshan Nature Reserve, Liaoning Province, northeastern China (Sun 1991). The Black Stork apparently no longer occurs in southern Korea as a breeding bird (P. Won 1973:18); however, birds were recently observed during the spring and summer in northeastern Korea (Hamgyong-Pukto) and could nest there (Sonobe and Izawa 1987:44). A few pairs may still nest in western Germany, e.g. in the

Liineburger Heide (Miiller-Scheessel 1964, 1965, Makowski 1970), and 5 or possibly more pairs in Austria; in southern Greece there are under 20 pairs. The species has disappeared as a breeding bird from Denmark (Hans Skov pers. comm. 1990), where about 150 pairs bred in 1850; this number declined to about 70 pairs in 1900, 40 in 1910, 20 in 1920, 3-5 in 1930-38, 1 (irregularly) through the early 1950s. The last successful breeding was probably in 1953. Its disappearance in Denmark was mainly from destruction of old forests and too much human activity near the nests; also many ringed birds were shot during their migrations (outside of Denmark). In Yugoslavia, numbers have declined sharply, especially in the north of the country, but 50-80 pairs remained in the late 1980s along the Sava River in Croatia (M. Schneider 1988b). In Poland the situation is one of improvement with an increase of breeding since 1936, after 50 years of decline. In 1966, 480 occupied nests were found in Poland, and the total population estimate rose to some 500-530 pairs; by the mid-1980s the estimate for Poland was about 800 pairs. In eastern Germany, 18 pairs remained after 1963, and in Czechoslovakia there was an increase since 1940 of up to 100 pairs in 1960. There were 50-60 pairs in Hungary in 1941, and about 150 pairs in the mid-1980s. Two isolated populations remain elsewhere in Europe: one in northeastern Portugal, where some 25-30 pairs still breed, and 100-150 pairs in central and southern Spain (Cramp 1977:324, Luthin 1987). A few pairs breed in southern Africa, where the discovery of nesting is a comparatively recent occurrence (Clarke 1904). Siegfried (1967) reports 15 pairs breeding in Zimbabwe (formerly Southern Rhodesia), 9 in Cape Province and 4 in Natal, South Africa, 3 in Lesotho (formerly Basutoland), and 1 each in Mozambique, Botswana (formerly Bechuanaland) and Orange Free State, South Africa, a total of 34 breeding pairs. In a later estimate, Siegfried et al. (1976) state that there are 'probably no more than 100 breeding birds' in South Africa itself, and that this population is stable. Transvaal Province, South Africa, supports an additional 50-70 breeding pairs (Tarboton 1982); a few pairs also breed in Zambia (C.W. Benson et al. 1971:46), at least 2 pairs in Malawi (J.H. Ryder and B.A. Ryder 1978) and probably a few pairs in the Kuiseb River Canyon, western Namibia (Tilson and Kok 1980). ECOLOGY The Black Stork usually avoids human habitation and favours well-wooded areas with shallow lakes and ponds as well as marshlands, damp meadows, rivers, and streams. It is very dependent on water, being more an aquatic feeder and fish-eater than the White Stork, so much so that in southern Africa, at least, nesting is not carried out in drought years (Tarboton 1982). Although usually frequenting freshwater areas over most of its range, non-breeding Black Storks are often found in the estuaries of tidal rivers in South Africa (Siegfried 1967). It may feed on drier land in the winter, when it will take reptiles, mammals, and insects. Unlike most stork species, the Black Stork is often found

Black Stork

in hilly or mountainous regions, and has nested as high as 2000-2200 m in Soviet Central Asia (Dementiev and Gladkov 1951:441); it is also found up to at least 2000m in Lesotho, southern Africa (Brown et al. 1982:179) and at about 2500 m in the western Sudan (Lynes, in Bannerman 1957:20). Feeding is by deliberate stalking while Walking Slowly in shallow water, either singly or in small groups. Food location is mainly visual, and the bird Probes vertically and grabs prey in the tip of the bill with a quick forward lunge of the head. Breeding adults in Poland may fly up to 10km from the nest site to gather food (Cramp 1966). On rare occasions Black Storks have been observed foraging with both wings raised in an open canopy (England 1974, Breife 1982). This has been compared with the Black Heron (Egretta ardesiaca), which shades the water with open wings and darts its head forward to catch prey, but although the wings are bent and pushed forward of the normal flying position the Black Stork at no time forms the closed canopy characteristic of the Black Heron. Fish makes up the main food of this bird; species captured include loaches (Misgurnus and Cobitis], pike (Esox lucius), burbot (Lota lota), rudd (Scardinius erythrophthalmus], roach (Rutilus mtilm\ perch (Perca fluviatilis], eel (Anguilla anguilla) and sticklebacks (Gasterosteus sp.). They have been seen to catch and swallow fish up to 30 cm long (Meinertzhagen 1954:390). And they also sometimes eat insects, frogs, snakes, crustaceans, small mammals, lizards and nestling passerine birds (Cramp 1966, 1977:326, Bauer and Glutz von Blotzheim 1966:427, Dementiev and Gladkov 1951:444, Witherby et al. 1939:117, Schroder and Burmeister 1974:26). In Africa, the Black Stork is less dependent on insects than White or Abdim's Storks and does not commonly follow locust swarms or outbreaks of army worm (Spodoptera sp.). In Pakistan, TJ. Roberts (1969) reported them eating snails (Vivapara bengalensis and Lymnaea acuminata). This species is much shier and more solitary than the White Stork. Birds generally nest and feed in secluded places, rarely visited by man. Dementiev and Gladkov (1951:444) describe it as 'exceedingly circumspect', and Siewert (1955:48), who studied the species intensively at the nest in northeastern Poland, referred to it as a Kulturfluchter (a fleer from civilization). However, in the Transcaucasus they are known to feed in cultivated areas, and here they also nest close to human habitation, occasionally even in Azerbaijani hamlets (Dementiev and Gladkov 1951:441). While occasionally seen in flocks of up to 100, it is more usually observed singly or in small parties of 3 or 4 birds. Predation by eagles (Aquila sp.), especially during migration, has been reported (Ritzel 1980, Madsen 1990). BREEDING Nests are often used year after year and the pair-bond may be of long duration; in Europe it is rare for a given nest to be used for more than 10 years (Schroder and Burmeister 1974:16). Pairs often return to their traditional nest site together, and it may be that pairs migrate and pass the winter together.

73

Usually the Black Stork nests solitarily, and where it is numerous, nests are usually at least 1 km apart. In the central part of Transvaal Province, South Africa, the closest nests were 6-20 km apart; in Kruger National Park, eastern Transvaal, some nests were 2.2-4.7 km apart (Tarboton 1982). Nesting in Europe is usually in old, large trees (e.g. beech, oak, pine) in deep forests, but may be on a wooded river bank or, especially in Asia or Africa, on a cliff. Territory near the nest and surrounding area is defended against conspecifics. Contrary to other records, two occupied nests in the same tree have been reported in the Lenkoran Lowlands, Azerbaijan, USSR (Dementiev and Gladkov 1951:441). In Europe, nests are most often 4-25 m high in trees—a wide variety of both evergreen and deciduous; in many parts of Asia and Africa, nests may be in niches on cliffs, caves, large boulders, or under overhanging ledges. Usually nests are well away from human habitation. Some are high up in trees, which can be seen from long distances, whilst others are well hidden. The trees selected are usually very large and the nest is often on a large horizontal branch, about two-thirds of the way up the tree. Dementiev and Gladkov (1951:442) found that near Lenkoran, Azerbaijan, the storks nest on the outskirts of colonies with herons and cormorants, though this must be considered unusual. In Transvaal, South Africa, they sometimes nest in colonies of the Bald Ibis or near Cape Vultures (Gyps coprotheres), Black Eagles (Aquila verreauxi], Lanner Falcons (Falco biarmicus), or Peregrines (Falco peregrinus] (Tarboton 1982). In exceptional cases, nests have even been built on the ground (Bauer and Glutz von Blotzheim 1966:423). The nest is a large structure of sticks and twigs, and sometimes quite large branches. Old nests that are used again and again in traditional nesting sites are repaired and characteristically reinforced with earth and grass, and lined with moss, leaves, grass, paper, animal-fur and rags. Sometimes old raptor nests are taken over. In southern Africa, they occasionally nest atop Hamerkop (Scopus umbretta) nests, or those of the Black Eagle (Maclean 1985:66). Old Black Stork nests can be 1-2 m in diameter, but are considerably smaller if newly constructed (Dementiev and Gladkov 1951:442; Brown et al. 1982:179). In general, they are somewhat smaller and shallower than nests of the White Stork (Bauer and Glutz von Blotzheim 1966:423). Unless the pair arrives together, the male usually returns first to the nest site. Threat displays and fighting are less common than in the White Stork. Birds often spread their long, white undertail-coverts and press them and the black tail (not spread, but compressed into a c. 12 cm wide panel) downward to about the vertical during displays; in hostile encounters, the luxuriant feathers of the lower foreneck are fully erected, giving the bird a larger, fiercer appearance (Siewert 1932a, Bauer and Glutz von Blotzheim 1966:426, pers. obs. M.P.K.). A variation of the Up-Down display is given towards intruders at the nest, in which the bill is lifted to the vertical and clattered (Stoll 1934). The Up-Down given as a greeting between mates is similar but less vigorous and most often without the vertical bill-movements and clattering.

74 Black Stork

According to several observers, bill-clattering is only rarely recorded in Black Storks in Europe (O. Heinroth and M. Heinroth 1924-31:141, Stoll 1934, Siewert 1932b:35, 1955:70, Rosenberg 1941), but we have seen it during the Up—Down display between mates in South African birds (Kahl 1972c). Up-Downs are accompanied by a series of weak, melodious bisyllabic hissing whistles, synchronized with the tossing movements of the head and sinuous movements of the neck. Although the Up-Down display of the Black Stork is very different, some other mating displays are similar to those of the White Stork, including Head-Shaking Crouch, Copulation Clattering and Display Preening. Before copulation, the female often bends forward, with her body about horizontal, and presses up against the breast of the male; he sometimes hooks his bill over her back before stepping on (pers. obs. M.P.K.). After copulation, mutual Up-Down displays and allopreening are often performed. Before egg-laying the pair spend much time on the nest together, with bills resting on or buried in the luxuriant neck feathers. Nest-building is done by both sexes. This usually involves the repair of an old platform, from previous seasons, and the adding of soft material to the lining. As long as 2-3 weeks may elapse between the arrival of a pair at the nest-site and the laying of eggs (G. Niethammer 1938:303). Clutches vary between 2 and 5 eggs, rarely 6. For 82 clutches in Poland, the mean was 3.2 eggs/clutch (Mrugasiewicz in Cramp 1977:327); in southern Africa, 90 clutches averaged 2.8 eggs/clutch (Brown et al. 1982:179) and another 19 clutches averaged 3.4 eggs/clutch (Tarboton 1982). The eggs are smaller than in the White Stork, and the shell is white with a greyish hue; they become soiled as incubation progresses. Eggs are laid at 2-day intervals, and incubation, by both sexes, commences after the first or second egg is laid. The incubation period is variously given, as 30, 35-36, 32-38 or 35-46 days (Dementiev and Gladkov 1951:443; Bauer and Glutz von Blotzheim 1966:424, Cramp 1977:327); the lower figures are probably the more accurate. Hatching is asynchronous. At least one adult remains on the nest, to guard eggs or young, until the nestlings are 23 weeks old (Ruthke 1957, Cramp 1977:326,328). Both parents feed the young by regurgitation onto the floor of the nest. At Oka sanctuary, USSR, young were fed mainly on small (9-25 cm) fish and a few frogs; each parent fed 3-8 times daily and the daily consumption of each nestling was 400-500 g (Cramp 1977:326). Begging young utter monotonous braying calls similar to other stork species, and clatter their bills when aggressively excited (Stoll 1934). Nest predation is apparently uncommon; pine martens (Martes martes) occasionally take eggs or young (Ruthke 1957). Fledging takes about 60-71 days, and after that the young join the adults at their feeding grounds. Young continue to return to the nest for another 2 weeks, to be fed and to roost at night (Ruthke 1957, Bauer and Glutz von Blotzheim 1966:424). Nesting success varies from 31 to 92% from year to year, with about 3.4 young/nest in successful pairs. One pair in Czechoslovakia successfully reared 5 young (Stollmann 1961). Birds apparently stay together in family groups until migration (Dementiev and Gladkov 1951:443;

Cramp 1977:328). G. Niethammer (1938:302) considered that sexual maturity is reached at the age of 3 years, because a bird ringed as a nestling bred at that age. In northern South Africa, Tarboton (1982) found that pairs failed to breed in 38% of the years, usually when rainfall was below average. TAXONOMY For many years most classifications listed the Black Stork and the White Stork (including the Oriental White Stork) as the only members of the genus Ciconia. We, however, include the Abdim's, Woollynecked, Storm's and Maguari Storks in the genus Ciconia (Delacour and Mayr 1945, Kahl 1972e). Detailed study of the courtship behaviour of this group (Kahl 1972c) suggests that the Black and White Storks may be the least closely related species in the genus, with the others, formerly placed in separate genera, in intermediate positions between them. In spite of its extensive range, no subspecies of Ciconia nigra have been described. The birds in southern Africa are probably a comparatively recent offshoot from the Palaearctic population; they seem to be reasonably well isolated, reproductively, but have not yet developed any noticeable differences from the parent-group (Moreau 1966b:122, Snow 1978:30). CONSERVATION The Black Stork is a rare and threatened species throughout most of its range. It is undergoing a long-term population decline in western Europe, and habitat changes are proceeding at a rapid rate in much of eastern Europe and Asia. It has already been extirpated from Scandinavia and nearly so from France, West Germany and Greece (Luthin 1987). In eastern Denmark, for example, the population was estimated to be 150 pairs in 1850, but had decreased to half of that in 1900, and to 20 pairs in 1920 (E.V. Rasmussen 1979); the last stork nested there in 1953. Although Black Storks are seen each year, they no longer breed in Denmark. This has been the tale throughout western Europe. The main threat to this species is the destruction of forests, particularly large trees on which nesting traditionally has occurred. In Russia and other parts of eastern Europe, the rapid development of industry and farming has reduced areas of breeding. Heavy predation by man in southern Europe and tropical Asia, during migration, have reduced the population considerably. In addition, habitat changes, particularly the loss of wetlands and the use of pesticides, in wintering grounds in Africa will undoubtedly adversely affect the Black Stork as they have the White Stork. According to Sun (1991) nestlings in northeastern China are often stolen for sale to zoos. The Black Stork is a wilder, shier creature than the White Stork, but without full protection on its breeding grounds, and during its extensive dispersant and migratory journeys it will inevitably fall rapidly in numbers throughout its range. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 290.

Abdim's Stork Ciconia abdimii Lichtenstein Ciconia abdimii Lichtenstein, 1823, Verzeichniss Doubletten Zool. Mus. Berlin, p. 76: Dongola, Sudan Other names: Whitebellied Stork, Smaller Locustbird (English); Abdimstorch, Regenstorch (German); Cigogne d'Abdim (French); Abdim-ooievaar, Regenooievaar (Dutch); Kleinswartooievaar, Blouwangooievaar, Swart Sprinkaanvoel (Afrikaans); Sinbilah (Arabic); Shamuwa, Shamwona (Hausa); Eviyoyo (Kakamega); Omenena (Luo); Shuramurove (Shona); Mokoroane, Lekololoane, Roba-re-bese (South Sotho); Lekololwane, Mokotatsie (Tswana); Endongondongo (Kwangali)

IDENTIFICATION Abdim's Stork is a comparatively small member of the 'typical' stork group, standing 68-81 cm tall. It is predominantly blackish, with a white rump and belly. The adult is a predominantly black bird, with the dark head, neck, upper breast, and back glossed with purple and green; the tail and wings are also black. Its lower back, including the uppertail-coverts and underparts, from the lower breast to the belly are white. At rest, a narrow line of white breast feathers shows between the bend of the wing and the throat. The iris is greyish brown, and the red-tipped bill is otherwise a pale horny jade-green. The bare skin of the face is lead-blue, with a red patch over and in front of the eye and under the bill. The legs and feet are dull olive, with a reddish 'garter' at the tibio-tarsal joint and on the toes. The male is slightly larger than the female, and the male's bill is deeper and thicker at the base (Brown et al. 1982:180). At the commencement of breeding, most of the bare facial skin becomes bright French blue and the rest of the soft parts assume a richer colour. The lores, eye-ring, chin and inside the ears are vermilionred, the forehead a lighter flesh-colour, and the remainder of the naked face bluish-purple. Breeding birds have greenish grey legs, with pinkish red intertarsal joints and toes (Andersson 1872:280, A.L. Butler 1905, Kahl 1972c). Two captive young took 32-36 h from pipping to hatching. Newly hatched young weighed about 37 g; they are thinly covered with dark down, over pale blue-grey skin, and have black foreheads. By 10 days, nestlings are covered with soft light greyish down, the bill is black with a white tip, and the legs are pink. Half-grown nestlings are covered with whitish down, except for the front of the crown where the down is black; this head and neck pattern—as shown in the photo in Bigalke (1948) —betrays a close relationship to the Woollynecked Stork, which retains this pattern in the adult. By 4 weeks of age, the young are fully feathered and resemble the adults, although substantially duller (North 1940, Bigalke 1948, Farnell and Shannon 1987). Immature birds are patterned like adults but are much duller and browner, with less gloss, and less colourful soft parts. There is some down on the bare skin of the throat, and the bill is a dull reddish (Mackworth-Praed and Grant 1970:54). The bisyllabic whistles are similar to those uttered by the Black Stork. Calls given at a roost in Zimbabwe have been described (Brooke 1969) as 'solitary whispered whistles which carry some 25 yards'. 'For a stork its cry is most trivial —a high-pitched "peep . . . peep . . . peep . . .", worthy of a little more than a barn-door chicken' (Lynes 1925a:550). Flapping flight is used for short-distance travel, and soaring/gliding flight for longer distances, especially on migration. Our observations at Lake Shala, Ethiopia, demonstrated the pattern that is

Abdim's Stork 77

general for most storks. Abdim's Storks left the nesting island to fly, by flapping flight, to the mainland. On the way back, they were first able to gain altitude in the thermals over the land, and could soar and glide back to the island. However, if the wind was high and soaring not possible, they returned by flapping flight low over the water (Kahl 197Id). We have seen similar situations in several species of storks. Abdim's Storks engage in incredible acrobatics when descending from high altitudes (Mackworth-Praed and Grant 1962:81). In normal flapping flight, the flap-rate was 189/min(n = 28)(Kahll971b). Abdim's Stork could be confused with the more svelte Black Stork, but it is a considerably smaller and more 'dumpy' bird, shows a white rump and lower-back in flight, and differs in its soft-part colours. It is gregarious at most times, unlike Black Storks, which are seldom seen in flocks. The White Stork differs, of course, in being predominantly white. And the Woollynecked Stork is somewhat taller and longer necked, with a whitish head and neck. Other African storks are larger and distinctive in their own ways. Sympatric herons are thinner and differently coloured, have a different 'jizz' (Gestalt^ manner), and are found in more aquatic habitats. DISTRIBUTION AND POPULATION It nests in the northern semi-arid tropics from Senegal, northern Nigeria and Chad, eastward through Sudan, Ethiopia, northern Somalia, southward to western Kenya and Uganda; it also nests, in small numbers, in Yemen, southwestern Arabia (Dekeyser 1952, Meinertzhagen 1954:391, K.D. Smith 1955a, Montfort 1965, Moreau 1966b:247, Urban 1970, Kahl 1971d, 1979b, Gallagher 1986). In West Africa, its breeding is largely confined between 17° and 8°N (Bannerman 1953:172); in East Africa it breeds, in small numbers, south to the equator. Abdim's Stork is an intra-African, trans-equatorial migrant, breeding north of the equator and wintering mainly

to the south. Its migratory routes and times of movement are well recorded. Movements seem largely timed to follow the rains, and their associated insect abundance (Andersson 1872:280), back and forth from northern to southern hemisphere. This stork rarely spends much time in dry conditions (Moreau 1966b:246, Brown et al. 1982:180). Birds in a region concentrate in large numbers in preparation for their southward migration; in Sudan this occurs in August and September (Luthin 1984a). Birds from West Africa move eastward and then southward by SeptemberNovember; they take a more easterly route than in the spring and probably fly at great heights over the Congo forest (Chapin 1932a:457, Bannerman 1953:173). From eastern Africa, they fly south often in huge flocks, becoming a passage migrant in September-December in Uganda, western Kenya and western Tanzania. The vast majority of these birds spend the winter (Dec-Feb) further to the south (Britton 1980:20). Flocks arrive in South Africa in October, departing northwards towards the end of March; they are most frequent in the drier inland areas (i.e. Orange Free State, Transvaal), and also in Botswana and northern Namibia (Clancey 1965:229, McLachlan and Liversidge 1970:52). Up to 800 birds have been reported from Zambia in February and early March (C.W. Benson et al. 1971:46). They pass northward through Uganda, western Kenya and western Tanzania in late February to May. Abdim's Stork is said to migrate at night (G. Archer and E.M. Godman 1937:68), but we doubt if this is common. A migrant Abdim's Stork was captured in Zimbabwe carrying an arrowhead imbedded in its chest. The arrowhead, which was surgically removed, proved to be of a type originating in the eastern portion of West Africa. Later the bird was released in apparent good health (Condy 1966). Some birds, mostly subadults and immatures, wander irregularly eastward to Oman, eastern Arabia; this movement may be increasing owing to expansion of cultivation and irrigation (Gallagher 1986). It has been recorded from southern Spain as a vagrant (Heuglin 1869-73:1107). There appears to be a small, sedentary (i.e. non-migratory) population in eastern Eritrea, Ethiopia (K.D. Smith 1955a, Snow 1978:31); they breed during the January-March winter rainy season, when most other Abdim's Storks are in the southern hemisphere. The species is common to abundant throughout its range, at times gathering during migration in groups of thousands at appropriate feeding sites. Breeding colonies are generally small, a few dozen birds at the most, but the breeding range is extensive and total numbers are at least in the tens or hundreds of thousands. In January 1987, Donald A. Turner (pers. comm. 1990) observed over 40000 Abdim's Storks, in company with over 100000 White Storks, feeding on an army-worm infestation, approximately 50 km west of Arusha, Tanzania. Overall, its populations are considered to be stable (Brown et al. 1982:180). ECOLOGY This stork is a terrestrial species, seldom feeding in water, and most commonly found in grasslands, pastures and culti-

78 Abdim's Stork

vated fields. It is primarily insectivorous, and its migration patterns can be altered by the local appearance and distribution oflocust swarms, as storks move to take advantage of this food resource. According to Andersson (1872:280), 'The more plentiful the rain, the more abundant the birds, the cause being simply the greater abundance of food.' Burnt ground, where new grass grows after rain, also supports high densities of readily available insects. Abdim's Storks are also attracted to grass fires, where they patrol near the flames.

Foraging is mainly diurnal and food-location primarily visual. Abdim's Storks generally feed in loose aggregations. At fires and on burnt ground, they often are in the company of Marabou and White Storks, kites, rollers and other birds. At other times, small parties of this stork will Walk Slowly across dry fields with necks stretched forward and heads erect, looking for their prey. At locust swarms, flocks of thousands can be seen. Under these conditions, a stork Walks Quickly, picking up insects until satiated. As Condy (1965) observed, single birds or pairs feed in a more leisurely manner, whereas birds in larger groups seem stimulated to forage at a faster rate. Is this because of social stimulation or just better foraging conditions (e.g. more food)? In Zimbabwe, a 'fair-sized flock' of Abdim's Storks, along with 4 Pied Crows (Corvus albus), reportedly cleared a 4-acre (1.6ha) field that was heavily infested with army-worm caterpillars in 3 days (Hamling 1953). The Abdim's distinctive diet is made up primarily of insects, and it is best known for its exploitation of plagueinsect swarms. It has been known to take frogs, lizards, other small reptiles, small fish, water-rats, young birds, millipedes, scorpions, snails and crabs. Its insect prey includes beetles, grasshoppers, crickets and army worms (Spodoptera sp.), caterpillars, and especially locusts such as the red locust (Nomadacris septemfasciata) (Andersson 1872:280, Heuglin 1869-73:1107, Ghapin 1932a:458, Meinertzhagen 1954:391, Dean 1964, Brown et al. 1982:180). This species has been reported (Friedmann 1930:30, Curry-Lindahl 1961) foraging near large mammal carcasses; it was not determined whether they were actually eating carrion or just the insects attracted to it (most likely the latter). When not feeding, flocks often gather near pools or water-holes. Sometimes the birds sun themselves with wings extended widely (North 1940). Soaring is common at midday near feeding areas and may aid thermoregulation during hot weather (Brown et al. 1982:180). There appears to be little aggressive action between these birds, but occasionally they will use Forward Displays or an Upright Display, with bill held high and body feathers compressed and perhaps with the neck ruff intermittently erected for a few seconds (Kahl 1972c). Often Abdim's Storks associate in feeding groups with White and Woollynecked Storks. And sometimes they are followed, or even ridden upon, by Carmine Bee-eaters (Merops nubicus) (Reichenow 1900-01:345, Cunningham-Van Someren 1970). This species is often extremely tame around nests, yet much less so when feeding away from the nesting area (A.L. Butler 1905). At the colony at Lake Shala, Ethiopia, one could approach almost to within touching distance of birds

on nests before they flew off; even then they moved only a few metres (Kahl 197Id). Local people generally do not bother the Abdim's. However, they are reportedly eaten by the Ennedi Nomads, in Chad (Salvan 1967); being insectivorous, they are presumably rather tasty. BREEDING Abdim's Stork is a colonial breeder, unlike other 'typical' storks (genus Ciconia). Breeding groups are usually small, consisting of a few to a few dozen pairs; as many as 30 nests have been reported from one mimosa tree in the Sudan (Brehm 1854). They sometimes nest in mixed colonies, with Pink-backed Pelicans (Pelecanus rufescens), Marabou Storks, Cattle Egrets (Bubulcus ibis), and Sacred Ibises nesting alongside each other (Lynes 1925a:550, Kasoma and Pomeroy 1987). They have been found nesting as high as 2100m in northwestern Ethiopia (Cheesman and Sclater 1935). Breeding is during the rainy season north of the equator, coinciding the time when insect populations are at their most prolific. In Ethiopia we found that the actual initiation of nesting coincided with the first heavy rains of the season, which produced a rush of chironomid flies and probably the insect prey of the storks as well. Heavy rain a week and a half later was correlated with a marked increase in courtship and nest building activity (Kahl 197Id). Thus it seems to us that rains or the food supplies they produce are the proximal trigger of courtship and nesting. An exception to the 'spring' nesting season is in equatorial East Africa and coastal Eritrea, where heavy rainfall occurs at other times of year (K.D. Smith 1955a, FJ.Jackson 1938:73). Abdim's Storks build their nests in trees, such as baobab, acacia and palm, as well as on rooftops of African huts; sometimes on masses of rock, such as on the columnar basalt formations of Lake Shala, Ethiopia (Kahl 197Id). In northwestern Kenya, Blencowe (1962) found them nesting on cliff ledges. Particularly in West Africa, nests are often built in trees in villages or even on the roofs of huts, and tribesmen welcome their annual return as harbingers of the rainy season. At Lake Shala, Ethiopia, we found that the nests were close together, some actually touching, although most were separated by 30 cm or more. They are made of sticks and lined with softer vegetation, and are 1.0-1.5 m in diameter and 20-30 cm thick (Kahl 1971d, Brown et al. 1982:181). Nesting begins with the male establishing a territory either at an old nest or at a new site. The frequency of ritualistic displays is greater than in most other members of this genus, probably because it is the only member to nest colonially. During the selection of nest sites Aerial Clattering is very commonly used. The attacker flies after an opponent, attempting to maintain a position above and behind. As the attacker nears its opponent, it lifts its head slightly above the horizontal and clatters the bill loudly (up to 25 times). An unusual display by males before and during pairing is termed Mock Fighting (Kahl 1972c). This rather bizarre display seems sometimes to be triggered by an insect flying

Abdim's Stork 79

near the bird; at other times it appears to be spontaneous. It erects the neck, head and breast feathers and makes a series of wild lunges in the air, snapping and rattling the bill. It leans from side to side as though buffeted by wind, often losing its balance, flaps its wings to retain balance, and bites vigourously at nearby objects or nesting material. Sometimes this display is followed by the bird diving steeply from its perch and flying around erratically in a darting, bat-like flight, sometimes with the feet thrust forward in the manner of a bustard on take-off. In a letter to Margaret Nice, Konrad Lorenz describes this display precisely (Nice 1943:68) in captive Abdim's and Black Storks (it has not, to our knowledge, been reported in wild Black Storks): 'When [they] are let out into a larger enclosure . . . they begin a curious dance, evading by "hooks" and sudden ducking the non-existent attacks of an imaginary eagle, even thrusting at him vertically upward with their bills.' Cathy King (pers. comm. 1990) has observed a similar display in captive Maguari Storks. When an unmated female approaches a male at a potential nest site, the Head-Shaking Crouch is frequently performed. The male utters a series of bisyllabic whistles and inclines its body forward, with head lowered and bill down. The wings are held slightly away from the body and lifted. Back and neck feathers are erected and the tail cocked. As weight is shifted from foot to foot, the head is shaken from side to side, and then the bird lowers its body to lie flat on the nest. Sometimes the approach of the female after such a display will result in her being driven off. Only occasionally is this display performed by a newly mated female as her mate approaches the nest, and after pair-bonding is complete is it no longer carried out. The Up-Down display of this stork varies strikingly from the well-known display of the White Stork. Throughout the display, the wings are spread and held at an angle of 90°. The bill is first pointed downward, with the neck arched; a series of melodious whistles are given. Then the head is raised until the bill is pointing upward, and the mandibles are clattered rapidly a number of times. The tail is alternately lifted and depressed, with head movements. To our knowledge, the Nest-Covering display has not been described for this species. The very active displays of Abdim's Stork at the time of pair-formation are followed by the completion of the nest or addition to a used nest from the previous year. The male brings most of the material, which the female uses to build the structure. Completed clutches are 1-5 eggs, usually 2 or 3; 16 clutches averaged 2.2 eggs (Brown et al. 1982:181). They are laid at 2 or 3 day intervals. Eggs are chalky creamwhite and soon become soiled. Data are not available for incubation periods in the wild.

The incubation period of a captive pair was 30-31 days, and began with the laying of the second egg (Bigalke 1948); artificial incubation of 2 eggs from another captive pair took 28-30 days to hatching (Farnell and Shannon 1987). Both parents incubate and continue to accompany nest exchange with Up-Down displays. The only egg mortality noted at Lake Shala was caused by intraspecific fighting at the nests (Kahl 197 Id). Adults feed young by regurgitating onto the nest. Parents often shade nestlings from the hot sun, with outstretched wings. Predation is avoided mainly by one parent being present while the nestlings are small. Eggs and chicks are lost through disputes of neighbouring birds, and attempts to take over nests or nesting material. The chicks develop slowly, being able to sit by the sixth day, and standing at 2 weeks. They are fully feathered by 4 weeks of age (Farnell and Shannon 1987). The fledging period is probably 5060 days, and the entire breeding cycle, from arrival of adults to the flight of the young, takes about 90-100 days (Brown et al. 1982:181). According to Bannerman (1953:173) 3-4 young, rarely 5, are reared in a season. TAXONOMY Once placed in the monotypic genus Sphenorhynchus, and before that in Abdimia, the Abdim's Stork is now grouped with the other 'typical storks' in the genus Ciconia (Delacour and Mayr 1945, Meinertzhagen 1954:391, Kahl 1972e). It was named after Abdim Bey, the Governor of Dongola, Sudan, in 1823 (Kahl and Schiiz 1972). CONSERVATION This locally abundant stork is fortunate in that native Africans hold local individuals in high regard, realizing that their arrival signals the beginning of the rainy season (G. Archer and E.M. Goodman 1937:68). Of its significance to the local Africans, Lynes (1925a:550) states: 'If Abdim does not bring baby boys, at least it brings the precious rains'. It is also recognized as an aid to pest control, owing to its large intake of insects. Threats include habitat loss and pesticides. In the Sudan, it has been suggested that pesticides are adversely affecting young, a suggestion that needs additional study (Luthin 1984a). Efforts to control locust plagues may reduce the food available to these storks.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 291.

Woollynecked Stork

Ciconia episcopus (Boddaert) Subspecies: Ciconia episcopus episcopus (Boddaert) Ardea Episcopus Boddaert, 1783, Table des Planches Enlumineez d'Histoire Naturelle, p. 54: Coromandel Coast, SE India Ciconia episcopus microscelis Gray Ciconia microscelis G.R. Gray, 1848, Genera Birds, 3, p. 561, col. pi. 151: no locality given (Africa) Other names: Whitenecked Stork, Whiteheaded Stork, Bishop Stork, Parson-bird, Beef-steak bird [India] (English); Cigogne episcopate, Cigogne a cou laineux, Gigogne eveque (French); Wollhalsstorch (German); Witnekooievaar, Bisschopooievaar (Dutch); Wolnek-ooievaar (Afrikaans); isiThandamanzi (Zulu); Laglag (Hindi); Kali-tul (Gujarati); Manik-jor (Bengali); Vannati-narai, Pathiri-kokku (Tamil); Karim-kokku (Malayalam); Padili-kokka (Sinhalese); Ghi-gyin-sut (Burmese); Sandang-lawe (Javanese)

IDENTIFICATION The Woollynecked Stork, a medium sized stork, standing 75-92 cm tall, is mostly black, with a distinctive downy white neck, which gives it its name, and a black skull-cap. In the adult, the hindhead and neck are white, with a black 'parson's cap'. The lower belly and undertail-coverts are white and contrast strongly with the rest of the glossy black plumage. There is a conspicuous ruff of fluffy elongated (c. 150cm) black feathers, with paler tan or iridescent greyish purple tips, on the lower foreneck, which can be erected during displays (Harvey 1972, pers. obs. M.P.K.). The black portions of the plumage are tinged with glossy greenish blue and purple, brightest coppery purple on the breast, sides of breast, and lesser secondary wing-coverts. The deeply forked tail is black, but is often obscured by the protruding, white, stiffened undertail-coverts, giving the impression that they are the true tail (Priest 1933:95, FJ. Jackson 1938:71, Henry 1971). [It has sometimes been erroneously stated (e.g. Blanford 1898:371, E.C.S. Baker 1929:324, H.C. Robinson and F.N. Chasen 1936) that the upper tailcoverts—rather than the tail itself—are forked. This has led to some confusion.] The legs and feet are dull red, darker on the tarsal joints and toes, the naked skin of the forehead, orbital area, cheeks and throat is dull bluish or lead-grey; the iris is dark red or brownish scarlet (Chapin 1932a:458, S. Ali 1953:404, Henry 1971:381). The bill is black, showing red along the culmen, at the commisure, and often toward the tip. In some specimens, the bill is largely dark red, with only the basal third black. This is placed by some as a separate race, neglecta (J.H. Riley 1925:28, Chasen 1935), but we have not recognized it here. The African subspecies averages slightly smaller and has the edges of the black cap diffused, forming a rather jagged border of mixed black and white feathering, compared to the sharply delineated border between black and white on the Asian subspecies. The sexes are alike, with the male somewhat larger.

82 Woollynecked Stork

The breeding plumage is cleaner looking and glossy areas of black feathering are brighter. The iris is deep crimson or wine-red with posterior border of the sclerotic membrane bright yellow (E.C.S. Baker 1929:324, Bannerman 1930:98, Chapin 1932a:458, J.H. Riley 1938:36); this conspicuous yellow skin enlarges slightly during some courtship displays (pers. obs. M.P.K.). At close range and when the wings are opened, as in Up-Down display, a narrow band of brilliant neon orange-red unfeathered skin is visible along the underside of the forearm (radius/ulna), it is almost glowing, like a red-gold jewel (Legge 1880:1119, Meyer and Wiglesworth 1898:809, Bannerman 1930:98, Hachisuka 1931-32:349, pers. obs. M.P.K.). Small nestlings have pale greyish down over plumbeouscoloured skin, buffy down on the neck, and a black crown. Later the neck is thickly covered with a white down, and the sides of the abdomen and undertail-coverts are white, while the upper parts, lower neck and breast are a dark sooty grey down with minute grey tips. The head, face and throat are feathered forward to the base of the bill. The bill is brownish black, tipped with reddish orange or yellow, and the iris is dull brown or hazel to dull green. Legs are fleshy plum-

beous, speckled with dark brown (Whistler 1918b, Chapin 1932a:458, J.A. Scott 1975, Anthony 1977, pers. obs. M.P.K.). The immature bird's patterning is similar to the adult's, although the forehead is feathered and sometimes streaked with black and white (Clancey 1964a:45, Mackworth-Praed and Grant 1970:54, J.A. Scott 1975). The dark portions of the body and wings are dull brown, rather than black, and lack the iridescence of adults. The feathers of the neck are longer and more fluffy (Kuroda 1936:540, S. Ali and S.D. Ripley 1968:98). Bysyllabic whistling calls, very similar to those of the Abdim's and Black Storks, are given during Up-Down greetings at the nest (Kahl 1972c). It has been reported to utter a fierce hissing when attacked by a trained falcon (Dharmakumarsinhji 1955:91). Bill-clattering is heard, infrequently, during courtship displays (Kahl 1972c) and in threat to raptors near the nest (J.A. Scott 1975). We are sceptical of reports of 'harsh and raucous' calls (e.g. Mackworth-Praed and Grant 1962:80, Maclean 1985:67). When Begging, young are said to utter a call 'like a file being used on a metal pipe' (Brown et al. 1982:182).

Woollynecked Stork 83

The Woollynecked Stork has to run for a distance along the ground before taking off (Rabor 1977:27). In normal flapping flight, the flapping rate averages 160/min (n = 10) (Kahl 1971b). They often soar on thermals. 'The powers of flight of this stork are remarkable . . . [it performs] complicated evolutions, rolling and tumbling in the air and diving at a steep angle, when the wind passing through its quills makes a loud noise' (Bannerman 1953:171). Birds have been observed diving from the nest and flying away in a darting, bat-like flight (pers. obs. M.P.K.). Woollynecked Storks often associate with Black Storks, particularly during their extensive migrations in northern India and Africa, but the former is easily distinguished by its white neck. In limited areas of peninsular Thailand and Malaysia, and southern Sumatra, the Woollynecked Stork may occur with the very similar Storm's Stork; the former is distinguished by the more extensive white on neck, darker bill and facial skin; difficult to confuse with any other species.

DISTRIBUTION AND POPULATION This stork has a wide distribution in tropical Africa, Asia and associated islands. It ranges from Senegal and Ethiopia to Angola and eastern South Africa; then from Pakistan, India and Sri Lanka to Vietnam, Thailand, Philippines, the Sunda Islands, Celebes (but not Borneo). The episcopus subspecies is found in appropriate habitat throughout most of the Indian subcontinent, though Tom Roberts (pers. comm.) considers it now extinct in Pakistan. Formerly (Delacour and Jabouille, 1931:86) it was common in Kampuchea, southern Laos and Vietnam. It is still widespread in the southern part of Vietnam (Vo Quy, 1975) and several were seen in central Kampuchea, as recently as 1968 (pers. obs. M.P.K.). It occurs in the Malay Peninsula, at least as far south as southern Thailand, but there are no breeding records for Malaysia (Medway and Wells 1976:94). No recent breeding has been recorded from Thailand, Burma or Bangladesh (Bain and Humphrey 1980, M.A.R. Khan 1984, Luthin 1987). It may be quite widespread in portions of Sumatra and Sulawesi (Holmes 1977, Luthin 1987) and was, at one time, widespread in the Philippines (duPont 1971:23) but its present status there is not known. It also occurs in Java, Bali, Sumbawa, Flores and Lombok. The microscelis subspecies is confined to Africa where it is found sparingly from the Gambia River and the Sudan south to northeastern South Africa, Angola, and northern Namibia and Botswana; it is less common at the southern edge of its range. In East Africa, it is local and irregular (Short et al. 1990:73); we have found it primarily on the coast in Kenya, and it is not uncommon in coastal Tanzania, including Pemba Island, and in the western parts of Uganda. It seems certain that this species migrates, but its movements are not well documented or understood. When dispersing or migrating it will sometimes gather together in sizeable flocks. Movements of African and Asian birds are

restricted to their respective continents and adjacent islands, and there appears to be no interchange between the two populations (Bannerman 1953:171). The species is resident and locally migratory in India (S. Ali 1964:100). Here, as in West Africa, migrations are probably governed by the rains (Bannerman 1953:171). C.W. Benson et al. (1971:47) reported considerable movements indicative of migration in Zambia, where it is not known to breed; in the Luangwa Valley it occurs only during the rains from November to May. On the other hand, western African birds move irregularly back and forth across the equator, between the northern and southern savannas, and avoid the rains in the wetter areas. Southern African records suggest southward movements in the early dry season (Apr-Jun) and northward migration in February and March (Brown et al. 1982:182). The Woollynecked Stork is seldom common, and generally is rare in most areas within its range. Because of its rarity and inconspicuousness, we know little about population levels in most regions. In winter, we have seen as many as 30 pairs in less than three weeks while traveling in Rajasthan and Madhya Pradesh, India (pers. obs. J.A.H.). It is uncommon in Gujarat, and 8 storks and 2 nests have been reported from the Gir Forest in Saurashtra (Luthin 1984a). During January 1989, a total of 462 were censused in India and 21 in Sri Lanka (D.A. Scott and P.M. Rose 1989:56). Other known populations exist in southern Vietnam (where it may breed in the mangrove swamps), Sumatra and Sulawasi (Luthin 1987). The total Indonesian population has been estimated at 'less than 1000 individuals' (MJ. Silvius and W.J.M. Verheugt pers. comm. 1989). In Africa overall, the species is uncommon to rare, but population numbers are probably stable (Brown et al. 1982:181). Kai Curry-Lindahl (in Brown et al. 1982:182) has seen several hundred together in Zaire; and in Zimbabwe, flocks of up to 200 have been reported in the Gwaai Reserve (Smithers et al. 1957:28).

ECOLOGY The Woollynecked Stork is a solitary species, about which relatively little is known. It often occurs in wet grasslands or dry ground near water, often in or near open forest. Its overall distribution is patchy, as it occurs singly or in pairs, often in wooded areas where marshes occur. In India, it is 'partial to water-logged ground . . . in the process of desiccation leaving fish and frogs high and dry' (S. Ali 1964:100). It generally avoids deeply forested country in Africa, but it has been seen feeding in lowland rain forest clearings in the Congo (Curry-Lindahl in Brown et al. 1982:182). It is found in various habitats, including grasslands, where it is particularly attracted by fires in the dry season. In East Africa it can be found as high as 3000 m (Britton 1980:21). In India, we have generally seen this stork in secluded forest glades and often hidden by tall marshland reeds that surround small muddy pools. Observation proved extremely difficult, if for no other reason than such

84

Woollynecked Stork

habitat is the preferred resting place of the tiger, which itself discourages investigation. S. Ali and S.D. Ripley (1968:98) considered it to be uncommon in intertidal marine habitats in India, but this is contrary to our experience in Africa and C.M.N. White's and M.D. Bruce's (1986:110) in Wallacea. In Kenya, we have seen it mainly on the coast, and in Sulawesi it occurs on sea shores (where it eats sea snakes) and in lowland swamps and rice paddies. In Sumatra, where it is sympatric with Storm's Stork, this species is found mainly in open swamps, flooded grazing land, rice fields, grasslands and dry crops (Marie and Voous 1988:66). And in Thailand, where Storm's Stork has also bred once, the Woollynecked Stork inhabits open (logged) semi-evergreen rain forest with grassy clearings and small waterholes in the plains and foothills (P. Round et al. unpub. data). This species forages by Walking Slowly or Standing, visually searching for prey. It feeds mainly early in the morning or at dusk, usually singly or in pairs. It is sometimes seen in small flocks, especially outside the breeding season. Although we once saw 16 birds foraging together in a loose group in northern India in August. Much feeding behaviour is reminiscent of that of the closely related White and Black Storks. Woollynecked Storks have been seen catching winged termites, in India, by flying back and forth through a rising swarm and snapping up the insects (Ball, in S. Ali and S.D. Ripley 1968:99). They sometimes attend grass-fires, to search for injured insects and other small animals escaping from the flames. On the Kenya coast, they forage visually on the coral reefs, along the edge of the water as the tide recedes, and on mudflats in tidal creeks. The Woollynecked Stork feeds on animal matter, such as locusts, grasshoppers, other large insects and their larvae, water-beetles, molluscs, crabs, frogs and toads, lizards, worms, and some fish; they probably consume some vegetable matter, perhaps accidentally. They have been seen eating winged termites in India (E.C.S. Baker 1929:325) and southern Africa (P. Steyn pers. comm.), and polychaete worms on coral reefs at the Kenya coast (pers. obs. M.P.K.). In Sulawesi they are said to eat sea snakes, Hydrophidae. They have been recorded at carrion with vultures (Friedmann 1930:29), and often take burnt insects obtained from grass-fires. Oil-palm fibres have been found in stomachs, but their exact role in the birds' diet is not understood (Brown et al. 1982:182). This species is usually found singly, in pairs, or small parties. Whilst McLachlan and Liversidge (1970:51) stated that in southern Africa the Woollynecked Stork seldom settles on trees outside the breeding season, our experience in other parts of the range is that when disturbed it will frequently fly to the top of a tree, choosing whenever possible a bare branch. We have watched groups of 4 or 5 birds spend considerable time preening in tall dead tree branches. They commonly roost in trees at night. Several authors (e.g. A. Koenig 1910) have commented on how shy and difficult to approach these birds are; in Zimbabwe, J.A. Scott (1975) found them tame in the non-breeding season but 'extremely shy and secretive when breeding'. Predation by an immature Martial Eagle (Polemaetus

bellicosus) in Botswana was reported by P. Steyn (K. Brouwer pers. comm. 1990). BREEDING This species is usually a solitary nester; small, loose colonies of up to 4-5 nests are occasionally reported (Bouet 1955a). The same nest may be used annually, if disturbance has not taken place. Since birds are often seen in pairs even outside the breeding season, it is probable that they remain paired for more than one season, perhaps for life. The nest is placed at the extremity of a horizontal branch or near the top in a tall tree 10-30 m, sometimes up to 50 m, from the ground. In India, trees such as simul (Salmalia malabarica), neem (Azadirachta indica), or imli (Tamarindus indicus) are used; in Africa, Brachystegia glaucescens, B. speciformis, Ficus sycomorus, Acacia xanthophloea, Albizia adianthiflora or Adansonia digitata (baobab) (S. Ali and S.D. Ripley 1968:99; Hitchins 1974, J.A. Scott 1975, Anthony 1978b, pers. obs. M.P.K.). Pitman (193la) and J.A. Scott (1975) reported nests placed in trees overhanging a waterhole or pool in a secluded section of river. Though the nest is usually not built near other storks, if the tree is large enough other species such as vultures (in India, Gyps bengalensis) or egrets will often nest nearby. In India, the nesting season varies with local rainfall conditions (S. Ali 1953:404); most egg-laying seems to occur during the rains. In most of the African range, on the other hand, the species seems to be primarily a dry-season nester (see Appendix). Courtship displays are somewhat similar to the other typical storks (genus Ciconia), but in well-established pairs, which may well remain together for multiple seasons, such displays are usually minimal (Kahl 1972c). The Up-Down display is most similar to that of the Abdim's Stork, with the wings held open at about 90°, displaying the brilliant orange band of bare skin along the underside of the radius/ulna, and the tail and undertail-coverts fanned and pumped up and down. Rapid, wheezy bisyllabic whistles are given, and the bill is clattered as it is raised up to about the horizontal (Kahl 1972c). Mackworth-Praed and Grant (1973:65) report a 'remarkable tumbling display flight in the mating season', but Brown et al. (1982:182) concluded that more recent observations do not confirm these reports. However, we have seen birds in Uganda and India dive from a nest-tree and fly about 100 m in a characteristic darting display flight, similar to one that sometimes follows the Mock Fighting display in Abdim's Stork (Kahl 1972c and pers. obs.). J.A. Scott (1975:205) describes a display used in response to interspecific disturbances at the nest that sounds similar to the Nest-Covering display in other Ciconia storks (Kahl 1972c, C.E. King 1988). Both birds share in the nest-building, incubation, and the feeding of the young (Lowther 1949:117, J.A. Scott 1975). They build a stick nest 0.6-1.2m in diameter and about 30-60 cm deep; it has a central depression, sometimes deep enough to hide the sitting bird, and is lined with finer twigs, dry grass and leaves. During incubation, changeovers take

Woollynecked Stork

place at intervals of 2^4 h during the day. Green leaves, and sometimes grass, are brought every few days. Repairs and alterations to the nest are constantly carried out, especially during the early stages of incubation. When its mate approaches, the sitting bird will occasionally become agitated and will bill clatter. When the approaching bird lands at the nest, both birds stand, headbob, and clatter their bills and sometimes open their wings in an Up-Down display (Brown et al. 1982:182). This behaviour has not often been observed; however, most change-overs seem to occur without display (J.A. Scott 1975), as is often the case with stork species that form longterm pair-bonds. Such definite mating displays that have been seen consist of allopreening by the standing bird, directed to the top of the head and neck of the sitting bird. Following this, both birds stand with necks bent and heads and bills touching, circling the nest once with headbobbing movements repeated in unison three or four times. The original sitting bird then sits again, and its standing mate, with breast-feathers erect, preens both itself and its mate, going into a half crouch and sitting on its tarsus momentarily. Generally 3-4, rarely 5, eggs are laid. They are rough, white, broad to pointed ovals, and quickly become stained. Intervals between laying are usually 2 days but sometimes 3. The incubation period is 30-31 days (J.A. Scott 1975). Young hatch asynchronously, and the size difference between siblings may be considerable. They solicit food from the parents with a Begging display, in which the head is bobbed up and down and a 'filing' call is uttered (Brown et al. 1982:183). When approached by potential predators, such as House Crows (Corvus splendens) in India, nestlings threaten by pointing their bills at intruders and clattering their mandibles rapidly in a Nestling Bristle display (Kahl 1972c). Parents feed the young by regurgitating food onto the nest floor. A parent remains at or near the nest until the nestlings are about 30 days old. At this time they are well feathered and able to move about the nest. After 40 days, young move out onto the branches and exercise their wings. They fly after 55-65 days but remain in the vicinity of the nest for another 3 weeks. Broods of up to 4 may be reared, although breeding success is usually nearer to 1.5 young reared per nest (J.A. Scott 1975, Brown et al. 1982:183). In some areas, we have found nest observation of this species to be difficult due to extreme shyness, and this is confirmed by J.A. Scott (1975) who considers that behaviour at the nest may be adversely affected by human interference. However, in other places, e.g. several nests in India and Uganda, birds nest near villages or houses and are not bothered by observers nearby (pers. obs. M.P.K.).

85

ioural and morphological affinities with Abdim's Stork, and its deeply forked black tail is similar to that of the Maguari Stork of South America. Finsch (1904), J.H. Riley (1925:28) and others distinguished birds with more red on the bill and more naked skin on the throat as another, slightly smaller, subspecies, neglecta. The type-locality of neglecta is in question but, as described by Finsch (1904), its range covers Celebes (=Sulawesi), Java, Sumbawa, Lombok and the Philippines. Chasen (1935) gave its type location as Java, but C.M.N. White (1974) considered that he erred. E.C.S. Baker (1929:325) was unable to separate neglecta from the mainland populations; he stated that the amount of naked skin on the throat varies greatly according to age, with older birds having more extensive areas of bare skin. Whilst most birds on the Asian mainland are reported as having a black bill with red on the culmen and at the tips of both mandibles, this can not be taken as unalterable, and the bill may, like that of other species, vary according to season and maturity. Examination of museum skins has shown that, while specimens from the Sunda Islands do show more red in the bill than those from India, many African birds have nearly as much red (pers. obs. M.P.K.). The differences are in our opinion too few and inconsistent to recognize neglecta as a valid race. A further discussion of this problem is found in C.M.N. White and M.D. Bruce (1986:110-111). Previously, many ornithologists considered birds we now recognize as Ciconia stormi to be a subspecies of C. episcopus. The recent discovery of the Storm's Stork breeding in southern Thailand (Nakhasathien 1987) and southeastern Sumatra (F. Danielsen et al. pers. comm. 1990)—where it is sympatric with C. episcopus without interbreeding—has clearly shown to us that it is a separate species (see Storm's Stork). CONSERVATION

TAXONOMY

Once widespread, this species has suffered overall population reductions from habitat destruction and shooting (Henry 1971:381). Although it remains uncommon but stable in Africa, the widely dispersed populations of Asia require urgent conservation measures in the establishment of reserves. The Melaleuca forests in Vietnam, where the storks breed, are fully protected (Luthin 1987). The limited information on this species makes more a detail conservation programme difficult. Thus range-wide surveys are needed to determine population size and status in both Africa and Asia. These data could lead to the development of appropriately situated reserves. Like other solitary nesting storks, which are likely to pair for life, this species returns regularly to traditional nesting sites, which are usually in a large tree. Without protection of these trees, a number of which are well known, this species will undoubtedly suffer further decline.

This species was formerly placed in the monotypic genus Dissoura, because of relatively minor anatomical characters (Beddard 1896). It is now recombined with the other 'typical storks' in the genus Ciconia (Delacour and Mayr 1945, Kahl 1972e). The Woollynecked Stork shows close behav-

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on pages 292-293.

Storm's Stork Ciconia stormi (Blasius) Melanopelargus episcopus stormi Blasius, 1896, Mitteil. Geogr. Gesell. Naturhist. Mus. Liibeck, ser. II pts 10-11, p. 120: western Borneo, Pontianak Other names: Borneo wolhalsooievaar (Dutch); Nok kra su-um (Thai); Burung ginti (Pasir Panjang/Kalimantan)

IDENTIFICATION A largely black medium-sized stork, Storm's Stork stands 85 cm tall, with white on the hindneck, abdomen, and under tail. It is similar to, but slightly smaller than, the closely related Woollynecked Stork. The adult's colour and pattern are very similar to the Woollynecked Stork including a black cap, downy white sides and back of neck, dark body plumage, and white belly and undertail coverts. Unlike the Woollynecked, which has an entirely white neck, black feathering in the Storm's Stork projects up the front and sides of its white neck in two wedge-shaped patches. As a result, the only white on the head and neck is restricted to the lower cheeks and nape and to a very narrow wedge extending down the midline of the foreneck for the upper third of its length (Ogilvie-Grant 1903, Nakhasathien 1987). The black cap has a definitive edge where it meets the neck, as in the Asian race of the Woollynecked, but it is a bit more extensive, coming forward to the bill and further back on the nape (pers. obs. M.P.K.). The black tail is deeply forked contrasting with the longer white undertail-coverts. [Some confusion exists in the literature about which actual tail-feathers and undertail-coverts are black and which are white. D.R. Wells (pers. comm. 1990) and C. King (pers. comm. 1990) confirmed in museum specimens that, as in the Woollynecked, all the feathers of the deeply forked tail are black and all the undertail-coverts are white.] The bill is vermillion, often with some pinkish red-orange on the sides and a yellow tip. A substantial area of facial skin is a dull orange to orange-yellow or golden. Particularly below the eye, this naked, golden skin appears slightly swollen (C. King pers. comm.). On some but not all individuals, the culmen is slightly concave (i.e. recurved) and there is a small knob where the culmen meets the forehead (Ogilvie-Grant 1903, Medway and Wells 1964, 1976:95). The legs and feet in the adult are dull red, with darker grey joints, but often become covered in extreta turning them a paler colour. The iris is red. There is an excellent coloured plate of this species, accompanying the article by Ogilvie-Grant (1903:pl.V). A captive Storm's Stork at Walsrode, Germany, acquired as a juvenile in early 1988, had in May 1989 a stripe of bright orange, naked skin down the under side of the ulna, visible when the wing was opened (pers. obs. M.P.K.). In this it is similar to C. episcopus. The sexes are similar, although, as in other storks, the male is probably slightly larger. This needs to be confirmed. At a nest in Sumatra, one of the adults was slightly larger and more glossy on the breast, throat, and back; this was assumed to be the male (F. Danielsen et al. pers. comm. 1990). The bill and soft parts brighten during the breeding season. The bill becomes vermillion to orangered. The throat, cheeks and part of the facial skin orange grading to pinkish-orange near the neck feathers. A large area around the eyes is yellowish-golden. An adult in full breeding plumage is well illustrated accompanying the articles by Nakhasathien (1987) and Round et al. (1988). At 1-3 days old, young are covered in white down, with a black bill and black crown. In less than a month, they roughly double in size, and black feathers start emerging on their throats, wings and

88 Storm's Stork

clearings (Nash and Nash 1988). As with other stork species, soaring behaviour is apparently very 'contagious', for when one bird began to soar near the nest-site in Sumatra, several others came from different directions to join in, until there were 6 birds soaring together for 30-60 min (F. Danielsen et al. pers. comm. 1990). Storm's Stork is slightly smaller than the Woollynecked Stork, with which it might occur in Sumatra or southern Thailand. When viewed from below in flight, the neck of the Storm's appears wholly dark (Holmes 1977). Although there is probably some overlap where habitats merge, the two species are ecologically segregated, with Storm's Stork generally preferring denser, undisturbed forested areas and the Woollynecked occurring in more open country and more often in disturbed habitat (Nakhasathien 1987).

bodies. Their bill is a blackish-grey with a yellow-orange tip. The facial skin starts a pale yellow and changes to dark grey; the gular pouch is a brighter yellow, with small black spots. The iris of nestlings is brown. Their legs start as a light yellow, gradually changing to pink with darker joints. The areas covered by black feathering increase greatly after 30 days. By about 45 days, the nestlings resemble the adults in feather pattern, but they were smaller and had shorter bills; they were fully feathered by 60 days. Unlike the adult, the black feathering on the head extends below the eye to the base of the bill. The legs are dark greyish flesh initially, but the tarsus turned a dull reddish-orange at about 3 weeks. Captive nestlings in Thailand could fly by 90 days, but wild young in Sumatra left the nest at about 60 days (Nakhasathien 1987, Danielsen et al. pers. comm. 1990). Captive juveniles have browner and duller body plumage than adults. They show a slightly more densely feathered upper neck, and the pinkish orange area of facial skin is a less intense dull gold. Their bills are a lighter red than adults (C. King pers. comm. 1990). A bird we observed in captivity at Walsrode, Germany, on 10 May 1989, gave a short series of sibilant whistles, very similar to those heard from the Woollynecked and Abdim's storks; they were not loud and were just audible at 15 m distance (pers. comm. M.P.K.). Elsewhere the adult's voice is described as 'Karau' (Smythies 1981:30) and that of the nestlings as a 'loud harsh Krack, krack, krack' (Nakhasathien 1987). Begging calls of the young have also been characterized as a froglike 'Skhrekeh . . . keh . . . keh . .. keh, skhrekeh . . . keh . . . keh* (F. Danielsen et al. pers. comm. 1990). This species is often seen soaring high over rivers or forest

DISTRIBUTION AND POPULATION This species is resident in Borneo and southeastern Sumatra, and a rare resident or irregular visitor in peninsular Malaysia and Thailand (Collar and Andrew 1988:15). It has recently (Nakhasathien 1987) been discovered nesting in southern Thailand and in South Sumatra (F. Danielsen et al. pers. comm. 1990). The majority of recent sightings have come from Borneo, where it is uncommon but widespread (Smythies 1981:30; Luthin 1987). In recent years, observations of individuals or small groups have been reported from Central Kalimantan, East Kalimantan, eastern Sarawak, Brunei and eastern Sabah (MacKinnon 1983, Luthin 1987). In Sumatra and neighbouring islands, there are a number of sightings. A specimen was collected from eastern Sumatra, and another from North Pagai, West Sumatran Islands. There are several records in peninsular Malaysia, including Sungei Tekam Forest Reserve and Kuala Lompat (Medway and Wells 1976:95) and, more recently, in Taman Negara and on the Pahang-Johore border (G.W.H. Davison and D.R. Wells pers. comm. 1990). The discovery, in 1986, of a nest of Storm's Stork in southern Thailand extends the range of this species over 500 km further to the north (Nakhasathien 1987). Nothing is known about migration. Probably no real migration as such occurs but storks most likely wander in response to local conditions (e.g. drought, disturbance, changes in food supply). Transient birds have been reported almost 20 km from the nearest typical habitat in Sumatra (F. Danielsen et al. pers. comm. 1990). This is a very rare to uncommon species, many observations being of only single individuals or pairs. Never an abundant species, there has apparently been a decline in numbers and it is now considered rare everywhere (W.B. King 1978). It is distinctly local in the heart of its range in Borneo, where it is thought to be decreasing. In western Malaysia, recent records include a bird seen in the Sungei Tekam Forest Reserve, Pahang (February 1969) and 2 near Kuala Lompat, Pahang, in the Kerau Game Reserve (February 1970) (Medway and Wells 1976:95). Since then one or two birds have been reported each year (Wells pers. comm. 1990). An unusual concentration of 12 individuals was seen in Brunei (Holmes 1969 cited in Smythies

Storm's Stork

1981:30), and 6 (possibly as many as 12) were seen, in groups of 1-3, in East Kalimantan along the Telen River (Holmes and Burton 1987; D. Holmes pers. comm. 1990). Up to 7 individuals have been seen near the Sembilang River, in South Sumatra (F. Danielsen pers. comm. 1990). The recent northward extension of its known range into southern Thailand is based on a single known nest. As it was found in an area of forest which is now flooded by dam construction, the small population may well have perished even at the point of its discovery (Nakhasathien 1987, P. Round et al. unpub. report). A recent estimate of the total population in Indonesia (i.e. Kalimantan and Sumatra) stated that they 'may well number less than 300' (MJ. Silvius and W.J.M. Verheugt pers. comm. 1989), although M. Silvius (pers. comm. 1990) now considers that estimate 'rather too high'. In total numbers this is probably one of the least numerous of all the storks in the world. ECOLOGY Storm's Stork is a solitary species of dense primary forest and the swamps, rivers, and small streams running through it. Most recent sightings in Borneo were along rivers adjacent to dense forest. In Borneo, Smythies (1981:30) reported it frequenting open, grassy, freshwater swamps subject to periodical flooding, river banks in remote forest, and even paddy-fields. In Sumatra it frequents dense primary and logged riverine and swampy lowland forest (Marie and Voous 1988:66) and has been seen feeding in small forest ponds (M. Silvius pers. comm. 1990). In the Malay Peninsula it frequents rivers and damp clearings in lowland forest (Medway and Wells 1976:95). The nesting habitat in southern Thailand was thick lowland forest with very large trees, such as Hopea ferrea, Cynometra bijuga, Dipterocarpus spp., Mesua ferrea and Ficus spp. Trees exceeding 10 cm (diameter at breast height) had a density of roughly 500800 trees per hectare and there was an 80-90% canopy cover (Nakhasathien 1987). The undergrowth was mainly rattans and other palms, bamboos, shrubs and climbers, and this correlates with similar lowland forest regions in Borneo. In general this species seems to be less tolerant of disturbed habitat than is the Woollynecked Stork (MJ. Silvius and W.J.M. Verheugt pers. comm. 1989). Little is known of feeding behaviour. Birds forage along the mudbanks of main rivers in central Kalimantan (Nash and Nash 1988). Its local name in Thailand ('nok kra suum') refers to the birds' way of fishing, by quiet stalking, along the bank of a stream in dense forest (Nakhasathien 1987). Adults from the Sumatra nest foraged mainly in primary swamp forest 2-3 km from the nest (F. Danielsen et al. pers. comm. 1990). They were most active away from the nest in the early mornings. D. Holmes (pers. comm. 1990) has seen them feeding in Lampung, Sumatra, in a small wetland within a dryland secondary forest. Similarly, little is known of the diet of Storm's Stork. Food items brought to the young by parents at the Sumatra nest were mostly fish, 5-7 cm long and weighing an estimated 10-30 g each, also some frogs, and worms 10-15 cm long (F. Danielsen et al. pers. comm. 1990). It is reported that it eats

89

earthworms in Borneo (Smythies 1981:30). Other food is probably similar to that of the Woollynecked Stork, but more data are needed. BREEDING The discovery of a nest in southern Thailand, by Seub Nakhasathien on 27 September 1986, was the first nest of this species known to science. The nest was 19 m up in the fork of a 27m tall dipterocarp tree (Dipterocarpus baudii) on the bank of the Khlong Mon, a tributary of the Khlong Phra Saeng, at 69m above sea level. This is in Khlong Saeng Wildlife Sanctuary, Surat Thani Province, Thailand, at 9°05'N 98°30'E. The nest was a flat platform 15cm deep, with an external diameter of 50 cm. It was constructed of dry sticks 15-60 cm in length and lined with dry leaves and some down. A second, unoccupied, platform was found some 200 m away in the top of another 30 m tall dipterocarp. A local man reported that before leaving the area in 1985, he had seen 4 birds feeding along a nearby tributary of the Khlong Mon (Nakhasathien 1987). On 26 April 1989 a half-built nest, the second known, was found in the Sembilang area of South Sumatra. This site was on the bend of a river 8-12m wide, about 20km upstream from the sea, near the boundary between mangrove and freshwater swamp forest; within 5 km of the nest, 82% of the area was covered with primary swamp forest and 17% by mangrove forest, as determined from satellite photos. The nest was 8.3 m above the river on the second lateral branch of a 18m high Rhizophora mucronata tree. The nestplatform, when complete, was 30-40 cm in diameter and 10-15 cm in height. It was constructed mainly of branches of Rhizophora mucronata 15-25 cm long and 0.5-1.0 cm thick, and lined with leaves ofXylocarpus granatum and draped with epiphytes (F. Danielsen et al. pers. comm. 1990). [It is not known whether the birds nested again, at the same site, in 1990; the area could not be checked because of budget restraints (F. Danielsen pers. comm. 1990).] Practically nothing is known of courtship behaviour. It is probably similar to that of the closely related Woollynecked Stork. At the Sumatra nest, very little in the way of ritualized displays was seen; when one bird returned to the nest to feed the young, the other adult usually flew away (F. Danielsen et al. pers. comm. 1990). If crows, monkeys, or raptors came near the nest, the adult spreads its wings to cover the nestlings, in what sounds much like the NestCovering display reported in the Maguari Stork (Kahl 1972c). A bird we observed in captivity at Walsrode, Germany, on 10 May 1989 (pers. obs. M.P.K.), gave a low-intensity form of the Up-Down display. Initially it raised its bill to about horizontal, then it clattered rapidly but softly 15-20 times, while raising its bill further to about 20° above the horizontal. The entire performance was reminiscent of that seen in the Woollynecked Stork (Kahl 1972c). Very little is known about behaviour at the nest. When the Thai nest was discovered on 27 September, the adults were apparently incubating. Two chicks hatched about 21-24 October, and when 1-3 days old were seen to crouch down

90 Storm's Stork

and remain motionless when the loud calls of a Crested Serpent Eagle (Spilornis cheela) were heard. By 22 November the nest had been destroyed and the young stolen by a local villager, who dislodged the nest and chicks with a long bamboo pole. The stolen young from the nest were located and delivered, for captive rearing, to the Nature and Wildlife Educational Centre at nearby Surat Thani. Here they were reared on freshwater fish, cut into small pieces. The newly constructed dam having now completely flooded this area of lowland forest, any remaining Storm's Storks must be considered lost to the area, and possibly to Thailand. In the 1989 Sumatra nest, an unhatched egg was found. When taken from the nest, it was 'olivebrown with black and greyish black areas'; however, after cleaning, it was 'basically white in colour' (F. Danielsen et al. pers. comm. 1990). This egg, apparently the only one of this species known, is preserved at the Zoological Museum, Bogor, Indonesia. When humans approached, the adult at the Sumatra nest crouched low on the platform, until only a small mass of black feathers, 'with only the eyes free to follow the intruder', was visible from below (F. Danielsen et al. pers. comm. 1990). Adults brought food to the young about 5 times/day during their first month and about 2 times/day during their second month. Food was regurgitated onto the nest-floor and picked up by the nestlings. Mostly small fish, plus some frogs and worms (still alive when presented) were fed to the young (F. Danielsen et al. pers. comm. 1990). While the young were in the nest, both adults returned to sleep in or near the nest each evening. TAXONOMY Previously the Storm's Stork has frequently been considered a race of Ciconia episcopus (Kahl 1972e, 1979b). It is clear now that both species occur, or have occurred, within the same areas. Holmes (1977) and Silvius and Verheugt (1986) gave evidence for the occurrence of both Storm's and Woollynecked Storks in southeastern Sumatra, and Storm's has now been found nesting there (F. Danielson et al. pers. comm. 1990). In Thailand, the discovery of nesting Storm's Storks (Nakhasathien 1987) establishes its presence within the former range of the Woollynecked Stork there. Unlike the putative race C. episcopus neglecta, C. stormi is quite distinct from C. episcopus. It has a striking orange and yellow facial skin coloration, which is apparent from the nestling stage onward, and a different feather-pattern on the sides of its neck. The habitat preferences of the two populations are different, with Storm's keeping to more heavily forested areas. On the basis of available information, especially the plumage and soft part differences, ecological differences, and its wide zone of sympatry with C.e. episcopus, we have concluded that Ciconia stormi should be considered a separate species. CONSERVATION This lowland forest bird is clearly at risk throughout its range, due to forest destruction and hunting. Although it is

listed as a protected species in Indonesia, Sarawak, and Thailand, such designation has brought no real protection or conservation action. The saga of the discovery of the first Storm's Stork nest ever to be found (Nakhasathien 1987), its destruction by local hunters, its flooding by dam construction, and the likelihood of this species' complete extirpation from Thailand at the very moment of its discovery, presents vividly the situation as it exists in Asia today. There appears to be little or no appreciation of the needs of conservation in many areas. Nakhasathien (1987) stated that the impact of the Chiew Larn Dam upon lowland bird communities was never addressed. In what was supposed to be an environmental impact assessment before the dam was approved, it was stated (Anon. 1980 cited in Nakhasathien 1987) that 'in regards to birds . . . the adverse effects of the impoundment are likely to be minor, because most birds are mobile and able to flee away in the event offloading [our emphasis]. It would seem difficult to find a statement that more clearly indicates the need to explain the effects of such developments to government bodies unfamiliar with conservation issues. W.B. King (1981) stated that 'the extensive destruction of forests in Borneo is believed a major factor in the decline.' Such widespread lowland forest destruction occurs throughout Southeast Asia. The only hope for the survival of species like Storm's Stork is for areas of complete protection to be established, not only against the destruction of the trees for commercial use or for dam construction but also against local hunters. Smythies (1981:30) stated that in Borneo, Storm's Stork is 'probably decreasing because it is shot for food.' In many cases the hunters are not original forest dwellers by tradition but have been attracted to such areas either for exploitation of the land or water, under their own volition or by government edict. Although direct persecution kills some birds, habitat destruction is by far the greater danger. Range-wide surveys are needed to locate the species and assess its breeding and feeding requirements, followed by its protection on large tracts of land. [According to F. Danielsen (pers. comm. 1990), the best time to locate this shy species is during early mornings when it is most active.] Clearly, with so little known about its biology and ecology, much basic study is required. The most critical long-term threats appear to be the Indonesian Governmentencouraged movement of people into the storks' forest habitat, and unsuitable forestry practices in swamp forest where regeneration is poor. Capture, mainly of nestlings, for supply to the international zoo trade has apparently increased in recent years. Storm's Storks have recently appeared in zoos in Kuala Lumpur, Malaysia and Walsrode, Germany. The price of the species in the bird markets of Singapore has doubled (from S$300 to S$600) in the past year (F. Danielsen et al. pers. comm. 1990). Such trade, if not restrained, could put fatal pressure on such a rare and localized species. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 294.

Maguari Stork Ciconia maguari (Gmelin) Ardea Maguari Gmelin, 1789, Syst. Nat., 1(2), p. 623; based on 'Maguari' of Marcgrave, 1648, Hist. Rerum Nat. Brasiliae, p. 204: northeastern Brazil Other names: American Stork (English); Cigiiena comun, Cigiiena Americana, Maguari tabuyuyi, Magoray, Yiulo, Gulo, Cabeza de Heuso, Sa pyta, Baguari Pillo, Gaban Peonio, Tuyango, Malvinero (Spanish); Cauaua, Cegonha, Joao-grande (Portuguese); Erie, Reddie foetoe (Dutch); Pillo, Pillu (Araucano); Tuyuyu (Guarani)

IDENTIFICATION The Maguari is a large stork, standing 97-120 cm tall. It has a straight bill and is largely white, with black flight feathers in the wing and a forked black tail. The glossy black of the adult's wing involves most of the feathers, including scapulars, greater coverts and flight feathers. The forked black tail—created by the outer tail feathers being longer than the inner—is shorter than the stiff white undertail-coverts (well illustrated in Chubb 1916:156), which may function aerodynamically. Nonetheless, the black tail can be seen through the under-coverts in overhead flight (Wetmore 1926:62). The bill is mainly bluish grey bordered with red, shading to dark maroon and blackish in its distal third. It is long, relatively thin and, except for its thicker base, markedly heron-like. In fact, in its original scientific descriptions, the bird was confused with a heron (Molina 1782). The head and neck feathers are long semi-plumes, which are erectile and function in display. The throat and pebblytextured lores and orbital-skin are orange-red, turning redder during courtship. The legs are purplish red, and the iris pale lemon-yellow to cream-white (Wetmore 1926:63, Sick 1985, B.T. Thomas 1988). The sexes are similar, although males are slightly larger and have a more recurved bill (Kahl 1971e). During courtship the plumage is bright from a fresh moult, the base of the bill is greenish grey whereas the tip is violet, with a diffuse zone of whitish violet between them (S. Narosky pers. comm. 1990). At hatching, nestlings weigh 76-90 g, and their sparse down is 'snow-white' (B.T. Thomas pers. comm. 1990). The bill is dull grey, the gape pink, the gular area whitish yellow, the legs and feet pink, and the rest of the skin bluish grey. By 8-10 days the body down is greyer, the head-down being curled and blackish; the nestling's bill, legs and feet become shiny black. The unfeathered throat is orange, and there is also a pale yellow stripe extending up the ventrum. This iris is dark brown. Succeeding the whitish down present at hatching, the young have two coats of very dark feathers, which together last for much of the nestling period (c. 10-75 days). By 3 weeks it may fade to olive-brown (Kahl 1971e, B.T. Thomas 1979b, 1984). By about 8 weeks, the juvenile plumage begins to appear. Development of this white plumage continues during the remainder of the first 3 months. By fledging, the gular patch has turned to pale scarlet. By 3 months, the legs and feet start to turn pinkish, the bill becomes bi-coloured, and the plumage resembles that of adults. The skin around the eye remains black for about a year, before turning red, and the dark brown iris does not begin to turn to yellowish until the second year. In fact, the dark iris is the best field mark for distinguishing juvenile Maguari Storks from adults (B.T. Thomas 1984). Nestlings of no other stork species have dark coloured down. Black nestling plumage may function to make young in terrestrial nests less visible to predators (Fernandez 1920, Kahl 1971e, B.T. Thomas

Maguari Stork

93

the Andes in Chile, where it has occurred as far south as Magallanes Province (A.W.Johnson 1965:150). The species is at least partially migratory, although little is known of its movements. Birds leave the Venezuelan Llanos when the marshes dry and the aquatic food disappears (B.T. Thomas 1985). It also occurs only seasonally in Bolivia and northern Argentina. Occurrences further south and in Chile may be related to drought dispersal (A.W. Johnson 1965:150). In Surinam, it is a rare visitor to coastal areas from March to May (Spaans 1976a). B.T. Thomas (1986) observed that several months after leaving the nest, the young left the nesting area, and none of the 128 ringed birds were seen again in the area until they returned to breed at 3-4 years of age. Apparently widespread in typical habitat over much of its range, this species is found nowhere:in large numbers (B.T. Thomas 1987). Although quantification is lacking, it appears to be most numerous in the Pantanal and Chaco of Argentina, Brazil and Paraguay (Short 1975, Nores and Yzurieta 1980). Brooks (1991) found it to be relatively common in the Paraguay Chaco, at a density of one stork per 1.16 km2. Populations appear larger in Amazonas and Rio Grande do Sul, Brazil (Sick 1985). Surveys in the Llanos of Venezuela found 82 individuals in 1983 (Luthin 1984a); however, studies there have shown that it has decreased markedly within the past decade (B.T. Thomas 1986, 1987). 1984, C.E. King 1988), or it might aid thermoregulation in cold weather by absorbing solar radiation (Cowles et al. 1967). Adults give wheezy, hissing, bisyllabic whistles during the Up-Down display; these are hoarser, lowerpitched and slower than those given by other species of Ciconia (Kahl 1971e, 1972c). During some social displays, especially hostile ones, they loudly clatter their bills (B.T. Thomas 1986, C.E. King 1988). Young beg with a 'Ehehe-ehehe' call. A variety of other sounds made by nestlings is described by B.T. Thomas (1984). The Maguari flies with neck and legs outstretched with strong wing beats alternating with gliding. It soars regularly in Buenos Aires Province, Argentina (Gibson 1880, Weller 1967) and in the Venezuelan Llanos (B.T. Thomas pers. comm. 1990). The flapping rate in normal flight averages 181/min( B = 32) (Kahl 1971b). The Maguari Stork is distinguishable from herons by its white colour, large grey and maroon bill, and black wings, and in flight by its outstretched neck. The two other large sympatric storks have featherless black heads; the Wood Stork is smaller and the Jabiru is larger than the Maguari.

DISTRIBUTION AND POPULATION The Maguari is found in South America, primarily in lowland areas east of the Andes, from eastern Colombia, through inland Venezuela, and the Guianas (sparsely), Brazil (mainly south of the Amazon), eastern Bolivia, southward through Paraguay, Uruguay, and Argentina to Chubut Province. It is rare, and probably non-breeding, west of

ECOLOGY Preferring shallow marshes, mudflats, tropical wet savannas, the Maguari Stork is a bird of open country and generally stays away from forested regions (Wetmore 1926:62). Its usual foraging site is very shallowly flooded wet meadow, in water less than 12 cm deep and more rarely in water as much as 30cm deep. It feeds solitarily or in pairs, especially during the nesting season, but will feed in larger aggregations and with other species of wading birds, especially outside the nesting season, Hudson (1920:114) reported that in the evening they gather in large numbers around water but in the daytime disperse over the feeding grounds. Primarily a lowland species, it occurs occasionally in the foothills of the Andes (e.g. at Cochabamba, Bolivia— Remsenetal. 1985). The Maguari Stork feeds visually by Walking Slowly, usually in shallow (2-30 cm) water, with its neck extended and partially open bill near the water, Pecking and Grabbing prey items. It may search carefully in dry fields and has been observed to lift up pieces of cow-dung to inspect underneath (B.T. Thomas 1986). It also will Wing Flash and Run short distances after prey. B.T. Thomas (1986) described observations of 'aerial pawing', a behaviour that would appear to be Foot Stirring in the air rather than in the water. Maguari Storks will sometimes Grope in the water or Probe in mud. When they catch a large prey item, they bite it a few times before swallowing. They have been observed foraging at grass-fires in Paraguay (Kahl 1972c). The diet is quite broad, including earthworms, insects, insect larvae, freshwater crabs, frogs, tadpoles, snakes, fish

94 Maguari Stork

and eels, small mammals, and eggs and young of marsh birds (L. Fraser 1843, Pelzeln 1868-71:304, Gibson 1919, Fernandez 1920, Zotta 1940, Kahl 197le, B.T. Thomas 1984, 1985). A large rail was found in the gullet of a bird shot in Patagonia (Bent 1926:70), and a Maguari was once observed eating cow-dung in Venezuela (B.T. Thomas 1985). In the Llanos of Venezuela, of 344 food items eaten by adults or brought to the young, 44% of the diet was fish and eels, 32% frogs and tadpoles, 12% rats, and 12% other invertebrates and vertebrates (B.T. Thomas 1985). Food brought to young differs in different years, presumably depending on availability. BREEDING In Venezuela, storks return to their breeding areas in flocks and gather in groups on feeding and loafing grounds. Most of the time in these parties is spent standing or preening. Some pairs reform in successive years on the same nest-site or in the same area (B.T. Thomas 1986). Sites may be reused, but most nests are rebuilt each year, as they do not usually last between seasons. Nesting generally starts early in the rainy season, so that young fledge at the end of the rains (B.T. Thomas 1987). However, it differs in timing throughout the storks range, and to some extent from year to year. Egg-laying occurred between May and November in different years in the Llanos of Venezuela (B.T. Thomas 1984), depending on variations in the beginning of the rainy season. Within an area in a given year, egg laying is fairly synchronized (B.T. Thomas 1985, 1986). In Argentina, nesting occurs in August and September (S. Narosky pers. comm. 1990). In the northern part of its range (e.g. Venezuela), the Maguari nests in small trees and bushes, usually less than 6 m high; nests in dead palms tend to fail because the trees break off or young fall out (B.T. Thomas 1984). In other areas (e.g. eastern Brazil, Argentina), it nests on the ground in marshes (Hagmann 1907, Kahl 1971e). Birds may nest singly, semi-colonially in well-dispersed groups, near colonies of other species, or in colonies of 5-15 nests; in colonial situations, nests may be within 50 cm of each other but are separated vertically in the tree or bush (B.T. Thomas 1984, 1986). In Venezuela, 77% of nesting was colonial, and colonial nesters were more successful than solitary nesters (B.T.Thomas 1984, 1985). Pair-formation and the reunion of pairs from previous years, which appears likely, may take place in assemblies on the ground away from the nest-site, as birds display to each other by walking about stiff-legged and giving Up-Down displays (B.T. Thomas 1986). This appears to be in contrast to pair-formation in other species of storks, which takes place in trees at or near the nest-site; however, in the Maguari such aberrant pair-formation may reflect an adaptation to ground-nesting, which these birds do over much of their range. Thomas' observations were made in the Llanos of Venezuela, where the storks are tree-nesters and primarily colonial; nothing is known of pair-formation in the remainder

of the range, where they nest mainly on the ground and mostly solitarily or semi-colonially. The elongated feathers on the lower foreneck are strongly erected in many displays. A variety of ritualized breeding displays were described by Kahl (1972c) and C.E. King (1988). Many of the displays are similar to those seen in other species of 'typical storks'. The Head-Shaking Crouch, a display which appears characteristic of and confined to storks of the genus Ciconia, has not been observed in the Maguari, although it may be expected to occur. The Up-Down displays observed in Argentina involved the displaying bird throwing its head to near the vertical, clattering the bill loudly and slowly 6-10 times; this was preceded and/or followed by 4-6 wheezy, two-part whistles (Kahl 1972c). B.T. Thomas (pers. comm. 1990) described the vocalization as a throaty hiss. During Up-Downs observed in captives, C.E. King (pers. comm. 1990) recorded that birds stood erect with the body nearly vertical, the tail lowered and the wings held close to the body. The head was thrown back, but not so far as in the Up-Downs of White and Oriental White Storks. Males tended to throw the head further back than females. The bird then bent forward, brushing the bill through the erected neck feathers, and the display terminated with the bill near or touching the ground. The wheezy vocalizations were given during the 'back' portion of the display and before the commencement of the bow. Up-Downs observed in Venezuela were rarely associated with bill-clattering (B.T. Thomas pers. comm. 1990). And C.E. King (1988, pers. comm. 1990) did not observe clattering in over 562 Up-Downs given by 10 captive birds from Paraguay and Argentina. It would appear that clattering during the Up-Down is rare, perhaps absent in some birds, in the Maguari Stork. This is similar to the situation with the Black Stork, in Eurasia and southern Africa. After nests were constructed, the storks continued to give Up-Down displays, which lasted an average of 9.25 s (n= 7) (B.T. Thomas 1986). Bill-clattering in Venezuela was observed to be associated with instances of aggression, threat and annoyance (B.T. Thomas pers. comm. 1990). It was given when humans or other birds, such as the Great White Egret (Egretta alba), Barefaced Ibis or Carib Grackle (Quiscalus lugubis). Copulation occurs on the nest, often after presentation of grass by the male. The Copulation Clattering display includes the members of the pair snapping their bills together or the male biting in the neck feathers of the female; 13 successful copulations were measured as lasting an average of 8.8s (B.T. Thomas 1986). Both B.T. Thomas (1986) and C.E. King (1988) observed courting birds carrying grass or other material in their bills, and this may have display function. In nest-building, males bring most of the sticks and females do most of the construction, although both do participate in both jobs. Nesting material, especially the lining, is added throughout the breeding season (B.T. Thomas 1986). The nests of the Maguari are unusual among storks in the extensive use of grass or reeds. Marsh nests may be made entirely of reeds (Cyperus giganteus), grasses and other aqua-

Maguari Stork 95

tic plants [Polygonaceae, Solinaceae (S. Narosky pers. comm. 1990)]. They look very much like crane nests. These nests are large conical structures, which can be up to 2.5 m wide and 1 m tall, built up from the floor of the marsh rising to 0.5 m above the water (Hagmann 1907, Fernandez 1920, Kahl 1971e). S. Narosky (pers. comm. 1990) found nests 80 X 150 cm in external diameter. We found similar ground nests in tree islands. Arboreal nests, too, may be made entirely of grass but most are made of sticks (<1 m long X <2cm diameter), covered with wet grass that is poked into the nest structure and hardens as it dries. The eggs are nearly smooth, dull white, and oval to subelliptical; the yolk is flame scarlet in colour. The clutch size is 2-4 eggs, averaging 3.2 in 49 Venezuelan nests (1974— 76); smaller clutches may occur in dry years. Eggs are laid on alternate days and incubation begins with the second or third egg. Egg-laying is highly synchronized in closely adjacent nests. Both adults participate in incubation, which takes 29-32 days. Young may hatch as much as a week apart and it is common to find weight differences of 5001400 g between siblings (B.T. Thomas 1984, 1986). Young 6-7 days old emit a guttural call. They give UpDown greeting displays—with rapid, but largely inaudible, bill-clattering—to returning adults and then drop to their tarsi and commence the Begging display (Kahl 1972c). Both adults feed the young, with undigested food regurgitated onto the nest floor. The size and quantity of food brought to nestlings increases with their age. An 18-day old nestling was given an eel 60 cm long and weighing 326 g; it took several hours to swallow (B.T. Thomas 1984). Parents are occasionally observed giving water to nestlings at midday, and they often shaded eggs or nestlings, with their backs to the sun and wings partially spread (B.T. Thomas 1986). One parent usually remains with the young at all times until they are about 6 weeks old. After the age of 25-35 days, but well before they can fly, young in terrestrial nests often wander off the nest, into the adjacent marsh (Kahl 1972c); however, young in elevated nests are 'nest-bound' until they can fly (B.T. Thomas 1984). The nest and its contents are defended by both parents, who give Arching displays, Aerial Clattering and Nest-Covering displays in response to potential predators near the nest. At solitary nests, adults perform a Lead-Away display, in which they approach a human intruder and clatter their bill, then walk away slowly, leading them from the nest (B.T. Thomas 1984, 1985). Feathered young also defend their nest, emitting a grunting or bleating cry and rattling their bills (Centro Editor 1983b, S. Narosky pers. comm. 1990). Young remain in the nest for 60-72 days and weigh about 4 kg when they leave the nest. Upon fledging, young form groups with other young at feeding areas. Juveniles apparently can recognize their parents individually, and they emerge from these groups to be fed by them, either in the marsh or back at the natal nest, for 5-6 weeks after fledging. Juveniles also return to their nest to roost at night for up to 6 weeks after fledging. There is little sibling rivalry, and success from 55 nests over 9 years in Venezuela was 52% (i.e. 89 young fledged from 171 eggs laid). Loss of eggs, presumably from snake predation, was higher than loss of

nestlings from all causes combined. Most nestling losses were caused by large young falling out of nests that contained 3^ chicks. No brood-reduction through nestling starvation was observed in any of the 123 nests studied. Within 3.5-4.5 months after fledging, the young had left the area (B.T. Thomas 1984), and, although 128 young were ringed over four seasons, none were seen again in the nesting area until they returned to breed at 3-4 years of age (B.T. Thomas 1986). Males apparently become sexually mature at 3 years of age and females at 4 years (B.T. Thomas 1987). TAXONOMY This species shows a number of similiarities to the Oriental White Stork of northeastern Asia, with which it shares some similar aspects of morphology and some behaviours, and to the Woollynecked Stork, with which it shares its distinctively forked tail. The fossil stork Ciconia maltha from the Pleistocene of North America may provide a past link between the ancestors of the modern-day Maguari and Oriental White Storks (H. Howard 1942; Brodkorb 1963). Like the Black Stork, the Maguari apparently clatters only rarely during the Up-Down display and may not do so in all areas of its range. As pointed out by C.E. King (1988), these are the only two members (among those that have been studied) of the genus Ciconia that habitually nest in heavily vegetated sites; in such situations it might be advantageous to possess displays that are less conspicuous to potential predators that might not otherwise find the nest over a long range. There has been much confusion regarding the scientific name of this species. It was formerly called Euxenura galeata, after Molina (1782). However, this description was of a composite nature (Hellmayr and Conover 1948, Kahl 1979b) and cannot be used. Following the suggestion of Delacour and Mayr (1945) and Kahl (1972e) we have merged the monotypic genus Euxenura into Ciconia, with which it shows a close relationship. This decision was further supported by the work of D.S. Wood (1983, 1984) and B.T. Thomas (1988). CONSERVATION Status throughout the range is poorly known, but in the Venezuelan Llanos, where the population was studied for 13 years, it has decreased markedly within the past decade (Luthin 1984a, B.T. Thomas 1986, 1987). Aerial censusing of nesting areas could add greatly to our knowledge of its overall status. Recently, the first chicks hatched in captivity in 30 years were produced at Disney World, Florida (Healy 1991). The Maguari is considered a gamebird in Amazonia (Sick 1985) and elsewhere. Hunting (including the taking of young for food), human disturbance and loss of habitat to agricultural activities are primary threats, and possibly the causes of population decreases seen in recent years (B.T. Thomas 1986, 1987). Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 295.

White Stork Ciconia ciconia (Linnaeus) Subspecies: Ciconia ciconia ciconia (Linnaeus) Ardea Ciconia Linnaeus, 1758, Syst. Nat., ed. 10, p. 142: Europe, Asia, Africa: restricted to Sweden by Linnaeus, 1861, Fauna Svecica, ed. 2, p. 58 Ciconia ciconia asiatica Severtsov Ciconia alba asiatica Severtsov, 1873, Izvestiia Imp. Obshchestva Liubitelei Estest. Antrop. Etnogr., Moscow, 8, pt 2 (1872), p. 113: Turkistan Other names: European Stork (English); WeiBstorch, WeiBen Storch, Adebar (German); Gigogne blanche (French); Ooievaar (Dutch); Ciguena comun (Spanish); Cegonha branca (Portuguese); Ciconia bianca (Italian); Hvid stork (Danish); Vit stork (Swedish); Bely Aist (Russian); Capa bfly (Czechoslovak); Bocian bialy (Polish); Roda bijela (Yugoslav); Barza alba (Rumanian); Feher golya (Hungarian); Pelargos (Greek); Bajbar, Badjah, Belaridj, Haji, El-haz laqlaq, Laklaka, Najeh, Mehab (Arabic); Leklek, Bu-laqlaq (Turkish); Lag-lag, Haji lag-lag (Hindi); Mashowori (Shona); Witooievaar, Grootsprinkaanvoel [='Great Locust Bird'] (Afrikaans); Lima (Hausa)

IDENTIFICATION Standing 100-125 cm tall, this large, mostly white, stork has black flight feathers in the wings, a straight red bill, and long red legs. The adult plumage is all white or near-white, except for the primaries, secondaries, greater primary and secondary coverts, alula, and scapulars, which are black glossed with green or purple. There is sometimes a small, variable amount of white or silvery grey on the outer vanes of the secondaries and/or inner primaries. The feathers of the lower foreneck and chest are elongated, forming a ruff that can be erected. The bill, legs and feet are red to orange-red; the legs may be whitish, owing to coating with uric acid from excrement. The iris is brown, and the bare skin surrounding the eye and on the lores is black. The naked gular skin is black, grading to orange-red posteriorly and along the border adjacent to the feathered neck. The sexes are similar except that the male is usually larger and with a more massive bill, which is sometimes subtly upcurved on the ventral border of the lower mandible (Schierer 1960a). In pairs where morphological differences are obscure, the sexes can often be distinguished on the basis of behavioural differences in ritualized displays (Schiiz 1942b, Kahl 1972c). The subspecies asiatica is on average larger than ciconia and its bill is redder, upcurved, longer and heavier. There are no dramatic differences between the non-breeding and breeding plumage, except that in the latter the whites of the feathers are cleaner and the areas of bare skin are more brightly coloured. The white plumage, especially the neck-ruff, becomes soiled as the breeding season progresses. The first-down of hatchlings is short, sparse, and off-white; the second-down (at 1 week) is dense, woolly, and white. Weight at hatching is 70-77 g (O. Heinroth 1922:186, Dementiev and Gladkov 1951:450, Schropel 1984). By the third week, black scapulars and flight-feathers, begin to appear. At hatching the bill'is black with a brown tip, and legs are pink, later becoming grey-black.

98 White Stork

tered White Storks at 3600 m above the ground. Birds occasionally clatter during flight, especially in the vicinity of the nest-site (pers. obs. M.P.K.). Within their breeding range, White Storks might possibly be confused with one of the white egrets, which have no black in the wings. In the winter range, they could be confused with the Yellowbilled Stork, which has a decurved yellow bill, naked red face, and a black tail. When soaring high, they might be confused with the Great White Pelican (Pelecanus onocrotalus) or the Egyptian Vulture (Neophron percnopterus)) which show similar wing-patterns, but the extended long neck and legs of the stork set it apart. DISTRIBUTION AND POPULATION

Juveniles are similar to adults but the black wing-coverts and scapulars are tinged with brown, and the primaries and secondaries are less glossy. The bill is, at first, shorter than the adult's and changes gradually from blackish to brownish or pale red with a blackish tip. The legs are a duller brownish red. Measurements of developing captive young are given by L. Gangloffet al. (1989). The only true vocalization by adults is a hissing, given primarily at the nest in conjunction with some forms of the Up-Down and other displays. Loud and prolonged clattering of the bill produces a rapid machine-gun-like rattle, with a marked crescendo and diminuendo. The tongue-bone expands the gular pouch, which acts as a resonator for the sound during clattering. Nestlings give a Begging call reminiscent of the mewing of a cat; they also clatter their soft bills, but without much sound. White Storks fly with neck and legs fully extended. They often soar high on thermals of rising warm air, especially during migration. Between thermals they glide in loose scattered formation, not usually in rows or Vs as some other soaring birds. During flapping flight, slow, steady wingflaps are interspersed with short glides. The flapping rate averages 170/min (27 recordings) (Kahl 1971b); the flight speed has been timed at 40-47 km/h (6 recordings) (Pennycuick 1972). Migrants have been reported at 4300 m in the Himalayas (Meinertzhagen 1955) and at 3000m in the southern Sudan (J. Vincent 1948). They have also been seen at 3300 m in the western Sudan (J. Vincent 1948) and at 1600 m near the Dead Sea, where one flock was mistaken for enemy planes (apparently during World War I) and fired on for ten minutes, one stork being killed (Meinertzhagen 1954:389). M.I.M. Turner (1964), while flying a small plane over the Serengeti Plains, Tanzania, encoun-

The overall distribution of this species includes much of Europe, the Middle East, west-central Asia, northwestern and extreme southern Africa. Vagrants have occurred in Iceland, Britain, Ireland, Norway, Finland, Malta, Azores, Madeira, Canary Islands, Cape Verde Is., St Helena I., and Sri Lanka. Ciconia ciconia ciconia is found in Europe in southern and eastern Portugal, western and central Spain and eastern France; in The Netherlands, Denmark, eastward to the Gulf of Finland and in the Pskov/Leningrad regions of the USSR, southward to the Sea of Azov, Turkey, northern Greece, eastern Yugoslavia, sporadically occurring in northern Italy. Formerly it bred in southern Sweden, Switzerland, western France, Belgium and southern Greece. In northwestern Africa it is found in northwestern Tunisia, northern Algeria, northern and northwestern Morocco. In the Middle East it is found in Turkey and Azerbaijan, USSR, southward to western Iran, and, increasingly, to Israel. A few pairs breed in the extreme southern tip (Bredasdorp, Mossel Bay and Oudtshoorn regions) of Cape Province, South Africa. The reports of breeding in Zambia, Zimbabwe and Zululand, South Africa, reported in MackworthPraed and Grant (1962:79) were in error (Clancey 1964b). It may have bred in Kenya in 1982 (Dittami and Haas 1982). A few birds winter in the southern parts of the breeding range; most migrate to tropical Africa or, in lesser numbers, to southwestern Asia. Ciconia ciconia asiatica is found in Turkistan, USSR, Uzbek SSR, Tadzhik SSR, Kirgiz SSR, and southern Kazakh SSR, from Amu Dar'ya to Issyk KuF and Lake Balkhash and extreme western Sinkiang, China (Kashgar). The summer range covers approximately between 39^3°N and 59-79°E (Creutz 1988:17). It migrates through Afghanistan and winters mainly in the northern part of the Indian subcontinent, mixing with birds of the nominate race from the Middle East. As many as 360 White Storks (race not determined) were seen in extreme southern India in January 1981 (J.M. Johnson 1984). Two White Storks with red bills seen west of Bangkok, Thailand, in January 1964, may have been from this population (P. Round et al. unpub. report). The White Stork is a long-range, diurnal migrant, which is very much dependent on soaring/gliding flight, assisted by rising thermals of warm air, for covering long distances with relatively little expenditure of energy. Movements take

White Stork 99

place mainly during the warmer times of the day, and birds can transit 100-200 km per day. Probably owing to this reliance on thermals, most European birds pass around both ends of the Mediterranean; only a few cross in the middle (see map in Schulz 1988:16). When water must be crossed, storks generally do so at the narrowest point. The main departure from the European breeding grounds is in August; as a rule juveniles leave before adults. A majority of birds from eastern Europe cross the Bosphorus, go around the eastern end of the Mediterranean, and enter Africa across the southern end of the Gulf of Suez. Whereas most of those from extreme western and southwestern Europe cross at the Straits of Gibraltar. The migration-divide (German: Zugscheide) in Europe is discussed in detail in Schuz(1963a). Autumn migration at the Bosphorus peaks between midAugust and early-September. It is estimated that approximately 370000 adult and immature White Storks pass through this area each autumn (Goriup and Schulz 1990). Flocks of up to 11 000 storks and total numbers of over 207 000 were observed between 23 July and 6 October 1966; 80% of the movement took place between 09.00 and 13.00 h (Porter and Willis 1968). Up to 315000 were counted at the Bosphorus in the autumn of 1972 (Kasparek and Kilic 1989a). At Burgas Bay, northwest of the Bosphorus in Bulgaria, 226956 were counted between 10 August and 30 October 1981; during the 5 years 1979-83, peak migration dates fell between 21 and 31 August; flocks averaged 577, but one flock contained 21 000 birds (Michev and Profirov 1989). On the western route, 27414 birds were counted crossing the Straits of Gibraltar between 20 July and 4 September 1985 (Lazaro and Fernandez-Cruz 1989). Birds using the eastern route winter mainly in the eastern half of Africa, many as far south as Cape Province, South Africa. In extreme cases, the round-trip journey from European breeding grounds to wintering areas in southern Africa may be over 22 000 km. It was estimated that a 25-year-old bird from East Germany may have covered over 500 000 km on migration in its lifetime (Creutz 1979b). Birds using the western route winter mainly in the northern tropics (12-17°N) of West Africa (Moreau 1972:240; ringing recoveries summarized in Cramp 1977:330, and Kanyamibwa et al. 1990). The return journey in spring is more rapid, with most birds arriving back on European breeding grounds in March-April. In certain years adults return late to the breeding sites and their nesting success is seriously reduced; the cause of these 'disturbance years' (German: Storungsjahre) is not known but may relate to unfavourable conditions on the wintering grounds or during migration. Wintering birds in Africa move around to a large extent in response to changing abundance of food, such as locust swarms (Schiiz 1959a,b). When food is abundant, the birds may gather in very large numbers: for example, 100000 White Storks were estimated in a 25 km2 area of northern Tanzania in January 1987, feeding on an army-worm infestation (Schulz 1988:17, 364, D.A. Turner pers. comm. 1990). There is some evidence that young birds often migrate further south than adults (Schiiz 1959a), and non-

breeding immatures often remain in the winter range during the northern summer. One juvenile, ringed as a nestling in extreme southern South Africa, was recovered 4 months later near the Tanzania/Zambia border, approximately 3100 km to the north (McLachlan 1963). Extensive ringing in Europe over many decades has furnished us with valuable information on movements, longevity and life-history. However, recent reports indicate that ringing may result in increased mortality because the rings can injure the bird's legs (Schulz 1987, Lohmer 1989). It is also possible that some birds are shot, out of curiosity, specifically because they are wearing a ring (D. Jonkers pers. comm. 1989). Counts of breeding birds in Europe are extensive and often of long duration. International Stork Censuses have been conducted in 1934, 1958, 1974, 1984 (Schiiz 1936a, 1979b, Schiiz and Szijj 1960a, Boettcher-Streim and Schiiz 1989, Rejman and Folk 1991). The trend is generally downward in northwestern Europe, with the last regular breeding in Belgium in 1895, Switzerland 1949, and Sweden 1954. Populations appear to be declining at a slower rate in eastern Europe (they may even be increasing and extending their range to the north and east in the Baltic states and European USSR). In The Netherlands numbers dropped from about 500 breeding pairs in 1910 to 209 in 1929, 85 in 1950, and 5 in 1985 (Jonkers 1989). In Alsace, France, the population declined from 177 pairs in 1947 and 145 pairs in 1960, to less than 20 pairs in 1986 (Schierer 1986, Kanyamibwa et al. 1990). In western Germany the number of breeding pairs dropped from about 2500 in 1958 to only about 500 in 1987 (Schulz 1988). And in Denmark numbers went from about 4000 breeding pairs in 1890, 1000 in 1930, 222 in 1952, 111 in 1964, 54 in 1971, 19 in 1984, to 12 in 1989 (Schiiz 1986c, H. Skov pers. comm. 1990). Numbers have increased in some areas of eastern Europe. In Estonia, the first nest was found in 1841; there were approximately 320 pairs in 1939 and 1954, and by 1974 there were 1060 pairs. Similarly, in the Pskov Region of the RSFSR, numbers climbed from 400 pairs in 1958 to 1218 pairs by 1974 (Veroman 1976). Population figures through the mid-1980s are summarized in Rheinwald et al. (1989). These recent estimates are (breeding pairs): Spain 6753, Portugal 2004, France 19, The Netherlands 2, Denmark 14, western Germany 649, eastern Germany 2722, Austria 318, Poland 30500, Czechoslovakia 1670, Bulgaria 5422, Rumania 3700, Hungary 4693, Yugoslavia 4500, Greece 1500, USSR 37086, Turkey 9000, Iran 2394, northwest Africa 15830. The total from these surveys is approximately 128000 breeding pairs. Some areas of the USSR and Middle East were not included, because of lack of data. About 19000 of these pairs belong to the western European population (birds nesting west of the Zugscheide), migrating through Gibraltar to West Africa. Between 1974 and 1984 this western population declined by 20% on average, considerably more in the north than the south. The eastern population, which migrates via the Bosphorus, declined by about 12%, more in the south than the north (Rheinwald 1989b).

100 White Stork

At the fringes of breeding range, there are about 6 pairs in the South African breeding population (Siegfried et al. 1976) and an estimated 600+ pairs in the C. ciconia asiatica population (Schiiz and Veromann 1982). Wintering numbers are more poorly known. Estimates of numbers wintering in Africa are at least 450 000 individuals in the eastern population and 90000-100000 in the western (Schulz 1988). A total of 310 000 White Storks were counted passing through Israel in the spring of 1984 (Paz 1987:33), with some remaining throughout the year (pers. obs. J.A.H.). Between 20000 and 24000 birds winter in the Kirov Bays area of the southwestern Caspian sea, about 140 km south of Baku (Goriup and Schulz 1990). According to a recent survey .{Goriup and Schulz 1990), the White Stork population level today is under half of what it was 40 years ago and is still falling. ECOLOGY The White Stork favours open country. In breeding areas, it is most often found in wet pastures, moist meadows, shallow lakes and marshes; it is fond of fields during ploughing or mowing. Feeding areas are usually adjoining or close to the nesting site. In the winter range, it is more often found in drier savannas and open grasslands, sometimes gathering at lakes, ponds or rivers. Food locating is mainly visual. A bird Walks Slowly or rapidly forward, with measured strides, holding its head forward of the body and the bill pointed down. Sometimes the White Stork locates aquatic prey by tactolocation, in a manner similar to the wood storks (Mycteria) (Lohrl 1957). Foraging is done singly, in small flocks, or in large groups if prey is abundant; at locust swarms up to 2000 have been reported in Uganda (FJ. Jackson 1938:69), 5386 in South Africa (Milstein 1966), and tens of thousands in northern Tanzania in January (Moreau and Sclater 1938). It is normally a diurnal feeder but is also active at night, by moonlight, during the breeding season (Darazsi 1958, Golodushko quoted in Pinowska and Pinowski 1989, Bauer and Glutz von Blotzheim 1966:405). Foraging efficiency of juvenile storks in Poland was found to be significantly lower than that of adults (Pinowska and Pinowski 1989). The White Stork's food is small animals, the diet varying considerably with feeding conditions. Prey includes insects and their larvae (especially beetles, grasshoppers, locusts), reptiles (lizards, snakes), small mammals (especially voles, mice, moles, shrews, young rats), and earthworms (especially important in early spring and during flooding). Less commonly it eats the young and eggs of surface-nesting birds, frogs (an important food item in some areas, not in others), toads, tadpoles, fish, molluscs and crustaceans. Indigestible remains of food are later regurgitated as pellets, measuring about 50 X 34 mm and weighing about 16 g (Haverschmidt 1949:47). Grasshoppers and locusts seem to be the most important food items on the African wintering grounds. Two storks examined had 1000 and 1600 locusts in their stomachs (Mell 1951). A first-year bird in western Kenya had eaten so many locusts that it was unable to fly and was killed by a

'wild cat' (Schiiz and Bohringer 1950). There is some evidence that storks may store excess fat before migrating. Milstein (1966) found birds that had been feeding on abundant locusts to be very fat in February in South Africa. Adults in Europe seem to gain weight late in the breeding season: in the month of June, 14 males weighed an average of 3.341 kg, 14 females 3.15kg; in July, 12 males averaged 3.97 kg and 12 females 3.521 kg (G. Niethammer 1938:291). Birds are often very gregarious on migration and in the winter quarters. They sometimes gather in large numbers, hundreds or even thousands, at abundant food sources such as locust swarms and grass-fires. The stork rests, often on one leg, with the bill buried in the ruff of feathers on the lower foreneck; both of these postures probably help to minimize heat loss at low ambient temperatures (pers. obs. M.P.K.). They are able to swim although rarely do so (Zimmermann 1955). Habitat losses in European breeding areas are probably very important in causing population declines. However, an analysis of long-term population and breeding trends in northwestern Europe suggests that 'critical mortality occurs in the winter in Africa' (Lack 1966:228). Changes in African feeding conditions, owing to factors such as droughts and locust-control, may be of major importance (Dallinga and Schoenmakers 1989). On migration, particularly in Europe, collision with wires and electrocution are important causes of mortality (K. Brouwer and C.E. King pers. comm. 1990). Hailstorms sometimes cause mass mortality of wintering birds in South Africa (J. Wittmann 1932, Badenhorst 1956, Kuhk, 1956). The heat and adverse winds (Arabic: khamsins] of desert crossings also take their toll of migrants (Schiiz 1935b, Bannerman 1957:11-12, C.A. Reed and T.E. Lovejoy 1969:152). Particularly on migration, there is evidence that storks are sometimes attacked by birds of prey (Schiiz 1965a, 1975, Breach 1970, Harwin 1970), and many are also killed by humans (Hartert 1912-3, Giraudoux and Schiiz 1978, Schulz 1988). A total of 484 ringing recoveries from southern Africa shows mortality rates in birds which died there to be 59% in the first year, 47% in the second, 36% in the third, and 20% in the fourth to tenth years, after which few recoveries were made (Oatley 1989). Other analyses indicate a first-year mortality between 55% and 74% (Schiiz 1955a).

BREEDING Usually the White Stork nests in the lowlands, but it does nest at up to 2000 m in Morocco (Heim de Balsac and Mayaud 1962:74) and Armenia, USSR (Dementiev and Gladkov 1951:446). Especially in Europe, the White Stork often breeds in close association with man, on rooftops, towers, chimneys, telephone-poles, high-wire towers, walls, hay-stacks and special poles erected for the purpose. In Armenia in 1988, 53% of nests were on poles and other telegraph wire supports, 29% on traditional buildings and 18% on modern structures (Adamian 1991). It also sometimes nests on trees, atop cactus (e.g. in Spain), or on cliff

White Stork 101

ledges, and, very rarely, on the ground (Thienemann 1911). Nests are built singly or in small, loose colonies; in Spain as many as 9 nests have been found on one rooftop (Chozas, 1984), and colonies may contain up to 30 pairs (Goriup and Schulz 1990). Some nests have been in use annually for hundreds of years; one known to exist in 1549 was still occupied in 1930 (Goriup and Schulz 1990). The bulky nest is constructed of branches and sticks, lined with twigs, grass, paper and rags; it is reused in successive seasons and added to each year and throughout the nesting season. Nest platforms are approximately 0.75-1.7m (exceptionally to 2.25 m) in diameter and 0.5-2.0 m (exceptionally to 2.8 m) high; the size increases with age. One nest, taken down from a cathedral weighed 800kg (Madon 1935). European nests are known to have been in use regularly for over 100 years (Naumann 1838). Ringing recoveries in central Europe show that about 80% of first-breeders settle on a nest site within 50 km of where they were hatched (Hornberger 1943a, Schiiz 1949c). Nesting is sometimes in mixed colonies with other species, such as with Cattle Egrets (Bubulcus ibis] in Spain. Pair-bonds are monogamous; rarely extra females attend a nest (Thienemann 1911, Assfalg and Schiiz 1988). Partners apparently do not migrate or winter together but often re-mate in successive years, probably owing to mutual attachment to the same nest-site. Males usually arrive back at the nest-site first. Sexual maturity is usually attained at 3-5 years; one male bird was still breeding successfully at 25 years (Creutz 1979b). Nesting (and sometimes roosting) birds greet newcomers with loud bill-clattering and Up-Down displays. Strange storks that approach an occupied nest often trigger a violent attack by its owner. If a newly-arrived female is quickly accepted by a male at a nest, it may signify that she is a returning mate from a previous season. Unwelcome strangers are threatened with a version of the Up-Down display in which the wings are held away from the sides and, especially in the male, pumped rhythmically up and down. Thus, the Up-Down display, in its various forms and contexts, can function as an advertising display, a threat or a greeting between mates (Camanni et al. 1989). When used as a greeting between mates, the Up-Down probably functions to suppress aggression (pers. obs. M.P.K.). The throat-sac is extended by the tongue-bone and acts as a resonator during clattering (Siewert 1932b:136). When a strange female attempts to approach an unmated male on a nest, he often displays by giving the HeadShaking Crouch, orientated toward the female. In this display, the male drops to the nest, as if incubating, fully erects the feathers of the neck-ruff, and shakes his head vigorously with a side-to-side motion. If, eventually, the female is accepted by the male, mutual Up-Down displays follow and soon thereafter copulations. A successful copulation lasts about 20 s (Schiiz 1934), and there may be as many as 107 day (Schiiz 1942b). Copulations have been witnessed, in wild pairs, with the female standing, sitting on her tarsi or lying flat on the nest (pers. obs. M.P.K.). After a pair is formed, mutual allopreening, especially in the feathers of the head and neck, is common between mates. For further

details on ritualized displays observed during courtship, the reader is referred to Schiiz (1942b) and Kahl (1972c). Nest-building is by both sexes; males bring most of the nest-material (Nagy 1955). Although old nests are usually reused, a new nest can be constructed in as little as 8 days (Goriup and Schulz 1990). A clutch contains 3-5 eggs, usually 4, the extremes being 1-7. Eggs are chalky white; they are laid at intervals of 1-4 days, usually 2 days. Of 80 clutches in The Netherlands, 5% had 2 eggs, 25% 3, 48% 4, 16% 5, 6% 6; the average was 3.93 eggs per clutch (Haverschmidt 1949:39). Of 837 clutches in southwestern Poland, the figures were: 1% 2, 48% 3, 44% 4, 8% 5; the average was 3.58 eggs per clutch (Mrugasiewicz 1972). The incubation period usually lasts 31-34 days. In Spain, 97 eggs took between 24 and 33 days to hatch, and the average was 30.4 days (Chozas 1986a). Incubation is by both sexes, female probably at night; it starts with the first or second eggs, and hatching is asynchronous. Young billclatter to parents from day one (Siewert 1932b:155). Both parents feed nestlings by regurgitation onto floor of nest. Nestlings are fed about once per hour when 5-6 days old and about once every 2 h when 15-16 days old (Schiiz 1943c). Water is regurgitated to the young on hot days (Schiiz 1943c, Haverschmidt 1949:50, Dementiev and Gladkov 1951:450). Parents often forage and feed their young on clear nights (Schiiz and Schiiz 1932b). Young are guarded constantly by one adult during their first 2-3 weeks. The fledging period is 54-70 days; young become independent and leave the area within another 7-20 days (G. Niethammer 1938, Dementiev and Gladkov 1951, Bauer and Glutz von Blotzheim 1966, Cramp 1977). Additional details on the care and development of nestlings may be found in Schiiz (1943c). The elapsed time from the completion of the clutch until first flight of the young averaged 91-92 days, based on 540 records (A. Mrugasiewicz pers. comm. 1969). There is considerable mortality (mainly of eggs, but also of some nestlings and even of adults) during violent fights between owners and strange storks that often attack and attempt to take over occupied nests (Schiiz 1936b). In East Prussia during 1931, 2660 cases of fighting were recorded at 8835 nests, with 77 adult birds left dead (Schiiz 1944). These fights are strongest between adult storks of the same sex (Schiiz 1944). Nesting success is higher in years when birds return earlier to their breeding sites from the winter quarters. In nests with several young, the smallest is reared successfully only in years when food is plentiful; when all young in a nest die, it is often the result of infestation with intestinal trematodes from frogs in the diet (Szidat 1935, Schiiz 1936b; additional discussion of parasites is given in Szidat 1940, 1943). When environmental conditions are optimal, as many as 6 young may be fledged from a single nest (Schneider 1988 cited in Goriup and Schulz 1990). In Europe overall, the number of young reared varied from 0.8 to 3.4 and averaged about 2 (Bauer and Glutz von Blotzheim 1966:404) There are several long-term studies of nesting success. In Germany (Oldenburg) during 1928-63, the brood size reared varied from 0.8 to 2.84, with a mean of

102 White Stork

1.94 (Tantzen in Lack 1966:216, Schiiz 1981a:86); in The Netherlands during 1934-45, from 1.3 to 2.6, mean 1.96 (Haverschmidt 1949:52) and during 1950-75, from 1.1 to 2.9, mean 1.7 (Jonkers 1975:265); in Denmark during 195271, from 1.3 to 2.9, mean 1.9 (Dybbro quoted in Cramp 1977:335). Breeding success appears to be highest when the parents are 10-16 years old (Meybohm and Dahms 1975:60). TAXONOMY The Oriental White Stork, in the Far East, has often been treated as a subspecies (C. ciconia boyciana) of the European bird. However, recent studies (e.g. Archibald and Schmitt pers. comm. 1990, G.E. King 1988) have shown the morphological and behavioural differences to be so great that we feel it should be considered a full species (Ciconia boyciana). Existing data suggest that the storks of the Central Asian population are considerably larger (see Appendix) than the European population (Vaurie 1965:84); some authors (e.g. Meinertzhagen 1930:432; J. Niethammer 1967b) doubt whether the differences are sufficient for distinction. Tentatively, we do recognize it as a subspecies (asiatica); however, additional measurements and other information on the bird's biology are desirable. The genus Ciconia as now constituted includes storks that have, at times in the recent past, also been classified under the genera of Sphenorhynchus (Abdim's), Dissoura (Woollynecked and Storm's), and Euxenura (Maguari) (Delacour and Mayr 1945, Kahl 1972Q. CONSERVATION The biology of the White Stork is one of the best known of all birds (Zink 1967, Rheinwald et al. 1989:470), primarily because of the species' close association with man, and its large size, popularity and abundance in Europe. In spite of this extensive scientific knowledge about the bird, we have not been able to prevent a continued decline in numbers over much of its range. Essentially it has been extirpated from Sweden, Denmark, The Netherlands, France, Switzerland, Belgium; and, if present trends continue, it may be gone from western Germany by the end of this century (C. Luthin pers. comm. 1989). Reasons for the decline in Europe are not firmly established (see discussions in Dallinga and Schoenmakers 1987, 1989, Goriup and Schulz 1990), and it is probable that several independent causes are involved. The following factors have been cited by various authors: destruction or alteration of feeding areas in breeding range; reduced food supplies in winter range (owing to droughts and insecticides); chemical pollution, in both summer and winter ranges; electrocution and collision with overhead wires; shooting (mainly on migration). It would seem most likely that it is loss of habitat—owing to development, industrialization, and intensive agriculture—in western Europe that has caused much of the decline (C. Luthin pers. comm. 1989). Otherwise why would there be such a distinct difference in numbers of storks at the borders of eastern and western Europe? It remains to be seen what effects the recent political changes in Europe will have on stork popu-

lations. Storks will undoubtedly suffer if future economic growth in eastern Europe is not tempered with ecological sense. Whatever the reasons for the decline, it would appear to be largely caused by man. However, long-term climatic changes may also be involved, especially in the protracted decrease of populations wintering in the West African Sahel, where there has been a generally declining trend in rainfall during 1943-85 (Kanyamibwa et al. 1990). Lack (1966) analysed data from German stork populations and concluded that the reasons for the decline should be looked for in Africa. Goriup and Schulz (1990) summarized it best when they said that 'western birds suffer most since they find no respite in either western Africa or the western European breeding grounds, whereas the eastern birds have the opportunity to compensate for poor breeding success by better survival in eastern and southern Africa during the winter'. In many areas of Europe and the Middle East, the White Stork is revered as a symbol of good luck and prosperity and is protected from direct persecution by popular custom. People sometimes say of an unlucky man: 'he must have killed a stork', and in ancient times the penalty for killing one was the loss of a hand (Gary 1973:38). Unfortunately the killing of a stork carries no such stigma in many parts of the world where the birds go during their non-breeding seasons, and many are killed by humans (Schulz 1988). However, direct killing by humans is of slight significance in the overall mortality of White Storks (Goriup and Schulz 1990). Attempts are being made to re-establish artificially the White Stork as a breeding bird in some areas of Europe, primarily Switzerland, France, The Netherlands and Italy, where it has disappeared (see Schierer 1972, Bloesch 1980, 1989a, Vos 1989, Camanni and Tallone 1990b, P. Arnold 1990, Blackwell 1990). Perhaps the most successful results have been achieved by Max Bloesch, of Switzerland, who re-introduced young birds taken from nests in Algeria, and —as last reported (1985) —had 108 nesting pairs of freeflying birds (Bloesch 1989a). Most of these birds remain the entire year in the immediate area and are artificially fed, so they cannot be equated with a truly 'wild population' that migrates to Africa each winter (cf Rheinwald, 1989a:436). Since the offspring of these birds sometimes do migrate away in the winter, it remains to be seen if a truly wild, naturally behaving population will eventually become reestablished. This is, of course, the hope. Such captive reintroduction programmes are controversial with many biologists. Some doubt whether 'project' storks can ever become truly independent of care by man. There are concerns about the wisdom of introducing foreign gene pools (e.g. Algerian storks into Switzerland). Since they often winter in the area and occupy nest-sites early in the spring, 'project' storks may out-compete wild migrants for nesting areas in regions where these are in short supply. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on pages 296-297.

Oriental

White Stork Ciconia boyciana Swinhoe Ciconia boyciana Swinhoe, 1873, Proc. Zool. Soc. London, p. 513: Yokohama, Japan Other names: Eastern White Stork, Japanese White Stork, Japanese Stork, Chinese White Stork, Korean White Stork, Blackbilled White Stork, Blackballed Stork, Far Eastern Stork (English); Schwarzschnabelstorch (German); Kitaisky Bely Aist, Dalnevostochny Bely Aist (Russian); Kono-tori, K6 (Japanese); Pai-kuan, Bai-guan (Chinese)

IDENTIFICATION Standing 110-150 cm tall, the Oriental White Stork is a very large white bird with black flight feathers in the wing. The bill is black and the legs and feet dark red. Adult plumage, including tail, is white or off-white; the primaries, secondaries and greater wingcoverts are black with a slight purplish or greenish sheen. Several of the inner primaries and the secondaries have the upper side of their outer vanes silvery grey to white (more pronounced than in the White Stork). At the base of the foreneck, there is a loose ruff of longer feathers, often soiled in appearance, that are sometimes erected during displays. The bill is black, tinged purplish or deep red at the base, slightly upcurved and, sometimes, with a slight gap between the mandibles in the middle (Vaurie 1965:85). The bare skin around the eyes is red and has a pebbly texture. Although there is some overlap in size, males are usually noticeably larger than females and have a relatively more massive bill. In the breeding season, there are no major changes in appearance, except that the newly moulted plumage is cleaner and the soft parts become more brightly coloured. The naked skin around the eye, lores, and on the throat-pouch becomes a deeper, richer red. The legs and feet are dark red. The iris is pale yellow-grey to pale blue. At 1 day of age, the nestling is covered with dirty white down and weighs 82-88 g. The bare skin near the eyes is a light pinkish orange and the iris is dark brown. The bill at first is ivory to greyish white with a yellow hue and later becomes a dirty greenish and yellow-orange in colour. The legs and feet are yellowish pink at first and later become light orange. By 16 days the nestling is covered with pure white down, and the bare parts of the head are a bright reddish orange. By 62 days the bill has become a dark grey with a yellowish brown dorsal edge, and the iris is a light yellowish grey. Further details on the growth and development of the nestlings are given in Winter (1982, 1991). C.E. King (pers. comm.) found that at fledging, captive young had plumage similar to that of adults, but they had brown on some feathers; their legs, feet, bill and bare areas of the face did not achieve the full colour of adults. Juveniles are described as being similar to adults, except that black feathers show a brownish cast and there are brown patches on the scapulars (Dementiev and Gladkov 1951:455). A loud and prolonged clattering of the bill is heard during many displays at the nest; clattering is also heard, occasionally, at feeding areas or in flight. By lowering the hyoid apparatus into the base of the gular region, a sound chamber is created that gives the clattering a reverberative component (C.E. King

Oriental White Stork 105

Stork, which occur in the extreme southern portion of the Oriental White Stork's winter range. DISTRIBUTION AND POPULATION

Shaded red area indicates former range.

1988:53). A 'hissing* vocalization is a component of agonistic displays by adults in captivity (C. King pers. comm. 1990, K. Archibald pers. comm. 1990). A *Kaa, kaa' or £Yia, yia' Begging call is heard from young, starting several hours before they hatch. A few days later a hoarse 'Yeee' call is given, growing louder and lowerpitched with age (Winter 1982, Fei et al. 1991b). C.E. King (1988:105) described the Begging call of captive nestlings as 'a bisyllabic vocalization that resembled a cat's "meow" in cadence1. In migration, Oriental White Storks fly in a scattered formation, unlike the V-formations of cranes (Hemmingsen 195la). In China, birds within the same flock soaring to gain height in thermals circle in both clockwise and counterclockwise directions. Migrants progress with both flapping and gliding flight. Flocks of autumn migrants often contained several hundred birds (M.D. Williams et al. 1991). On the wintering grounds, they sometimes clatter their bills when gliding (Wang 1991). This species might be confused with the white cranes (Grus japonensis or G. leucogeranus), with which it sometimes migrates; however, the cranes have less black in the open wings, and they usually call loudly in flight and form Vformations, whereas the storks do neither (Hemmingsen and Guildal 1968:188). In southwestern parts of the winter range, the Oriental White Stork may mingle, occasionally, with the White Stork. The Oriental species is considerably larger and has a black (as opposed to a red) bill. It is not easily confused with the Milky Stork or Asian Openbill

The Oriental White Stork breeds in the Amur and Ussuri River regions of the Soviet Far East (westward to Blagoveshchensk and southward to Lake Khanka) and extreme northeastern China (Heilongjiang, Jilin Provinces), and formerly bred in Korea and Japan. It winters mainly in the Changjiang (=Yangtze) valley, China, a few wintering south to Fukien Province and rarely to Hong Kong, occasionally to Taiwan, southern Ryukyu Is. It has wandered northward to southern Yakutsk and Sakhalin Is., USSR, and southwestward to Bangladesh, northeastern India and southern Tibet (Vaurie, 1965:84—85, Fisher et al. 1969:183, fitchecopar and Hue 1978:73, Neufeldt and Wunderlich 1982). Soviet birds depart from their nesting area in the Middle Amur River region between late August and midSeptember; other migrants, from further north, move through the area in early-October (Winter 1982:82). In Heilongjiang Province, China, spring migrants have been observed to appear in the breeding area in late-March and autumn migrants had departed by late-October (Fei et al. 199la). Further south, in Liaoning Province, spring migrants pass rather quickly in March and autumn migrants pass from mid-September to mid-November; the autumn migration has more birds and is more protracted (Sun 1991). Most of the migrants seen passing Beidaihe, in eastern China, did so between 11.00 and 15.30 h. The autumn stork migrations there exhibited a wavelike pattern, probably related to weather conditions, with 71% of the 1986 migration (1945 out of 2729 birds) passing on only 4 days: 29 October; 3, 6, and 14 November (M.D. Williams et al. 1991). Most Oriental White Storks apparently winter in the extensive wetlands near the middle and lower Changjiang River in south-central China, with a few wandering south as far as Hong Kong (Chalmers 1986:43). They start appearing in mid-October, and the majority of birds have arrived by December; they leave for the north in early March (Luthin 1987, Wang 1991). Occasionally individuals may winter as far north as Korea (Austin 1948:45, P. Won 1973:17, Sonobe and Izawa 1987:42), southern Liaoning Province, China (Sun 1991), or Maritime Territory, USSR (L. Taczanowski 1891-93:974, Dementiev and Gladkov 1951:454). Some birds winter at the mouth of the Ussuri River, USSR, fishing through the ice (Wilder and Hubbard 1938:422). In eastern China, at Beidaihe, most spring birds are seen passing during the latter half of March. Many more storks are seen passing south in the autumn than are seen going north in the spring (M.D. Williams et al. 1991). This could reflect some losses during the winter, but probably means that the main spring migration occurs elsewhere. In the days when Japan held a breeding population, those birds were mainly residents or short-distance migrants,

106 Oriental White Stork

spending the entire year in Japan (Jahn 1942, Haverschmidt 1949:70, Schiiz 1963b, Neufeldt and Wunderlich 1982). Populations are small. Once common in Japan, the species is now extinct there as a breeding bird. The major decline in the Japanese population took place between 1868 and 1895. Illegal shooting and, later, poisoning by mercury and other agricultural chemicals were the main causes for the decline. The last successful breeding in Japan took place in 1959 (W.B. King 1981). Only 7 birds still remained in the wild in Japan in 1966 (Sakamoto 1966), and only one in 1970 (Kuroda 1970); it now occurs only as a rare migrant or winter visitor. The species was still 'locally common' in Korea through the 1940s (Austin 1948:46). However, it had disappeared as a breeding bird in South Korea by 1971, when the male of the last remaining pair was shot (P.O. Won 1971, Luthin 1987). By some unknown date prior to 1989, it also ceased breeding in North Korea (K. Sonobe pers. comm. 1989). In 1978, the USSR population, in the Far East, was estimated at approximately 660 breeding pairs and was thought to be declining (Flint 1978). On the USSR side of the Middle Amur River in the Bureya-Arkhara depression, Winter (1982:77) found 31 pairs nesting in a 162 km2 area in 1975, a density of about 1 pair in every 5.2 km 2 . Most of the known nesting birds are on the USSR side of the border (Neufeldt and Wunderlich 1982); little is known about the breeding populations in Chinese territory. Martin Williams (pers. comm. 1989, M.D. Williams et al. 1991) reported a total of 2729 Oriental White Storks migrating past Beidaihe, Hebei Province, in eastern China during the autumn (11 Oct-16 Nov) of 1986. This is more than double the previous estimates of total world population and may represent most of the birds alive at that time. The current estimate for world population is about 3000 birds (Luthin 1987). A total of 1367 Oriental White Storks was counted on the wintering grounds, mostly in the Changjiang River area of China, during January 1989 (D.A. Scott and P.M. Rose 1989:80) and 121 in Hong Kong in 1991 (Chan 1991). ECOLOGY The favoured habitats of this species are river valleys, wet meadows and marshes with scattered clumps of trees. In the Soviet Far East, they have been observed breeding primarily in river lowlands on islands of forest, in areas with scattered moss-sedge-cottongrass bogs (Winter 1982). During times of former breeding in Japan and Korea, they frequented cultivated fields, especially rice-paddies, with nearby wooded hillsides (W.B. King 1981). Wintering birds in China are found at lakes, marshes, and, sometimes, sandy beaches and intertidal areas (Shi 1991). In the wet spring of 1975, adults in the Soviet Far East foraged within 100-600 m of their nests; later in the summer, when conditions were drier, they foraged 3-4 km away. In the drier spring of 1976, foraging was done 3-7 km from the nests (Winter 1982). Foraging birds Walk Slowly in shallow or fairly deep

water, often up to their belly feathers, and make swift downward and forward pecking movements (Winter 1982). Food finding is mostly visual (Fei et al. 199la). Sometimes, especially in water where visibility may be limited, feeding is apparently by tactolocation. Birds plunged the partially open bill into the water, as they walked, much in the manner typical of the Blacknecked and Saddlebill Storks. Up to 17 plunges per min have been counted, when storks caught an average of 1.22 ± 1.72 S.D. prey items per 5 min observation period (n = 9). Handling-time varied from 3-5 s for a carp to 2-3 min for catfish (Coulter 1990c, Coulter et al. 1991). Shi (1991) found that wintering birds foraged mainly in the early morning (07.00-09.00 h) and late afternoon (15.00-17.00 h) and rested at midday. One bird, watched for nearly 10 h, caught 486 food items in 742 attempts, a 65.5% success rate. The food is primarily fish, especially loaches (Misgurnus fossilis and sleepers (Perccottus glehni), frogs, aquatic beetles and other insects, also small rodents, snails, arachnids, snakes, crustaceans, clams, worms and young songbirds (Austin 1948:46, Austin and Kuroda 1953:333, Neufeldt and Wunderlich 1982, Winter 1982, Wang 1991, Shi 1991, Winter 1991). According to S.W. Winter (pers. comm. 1990) the Oriental White Stork is a specialist, with fish making up 70-90% of the total number of food items and 80-90% by volume. He has recently compiled an analysis (Winter 1991) of 2600 food items eaten by storks in the Amur region, USSR. In the Soviet Far East, vegetable matter (e.g. leaves, moss, sticks, seeds) made up a large percentage (70-90%) of the volume in stomachs of some adults and nestlings (Winter 1982:94). However, most of this material is probably ingested accidentally during the capture of animal prey, and, being less digestible, it remains in the stomach for a long time (S.W. Winter pers. comm. 1990). Wintering birds in China have been observed purposefully eating plant material, sand, and small stones (Shi 1991). A Chinese bird was seen catching a catfish weighing about 750 g (Fei et al. 199 la). Estimates of daily food-intake of wild, wintering birds in China ranged from 1.5-3.Okg; c. 70% (by weight) of the food taken was fish (Shi 1991). Apparently the Oriental White Stork is more dependent on wetlands and more tolerant of cold than is the White Stork (Hemmingsen 195la). It migrates south in the autumn later than the European species and winters further north; its larger size may be an adaptation to cold weather (M. Williams pers. comm. 1990). Wintering birds, in southcentral China, roost in flocks in open shallow water (Wang 1991). BREEDING In the Soviet Far East, nests are placed atop lone trees or at the borders of woodlands; they have not been closer than 4.55.5km to human settlements (Winter 1982). On the other hand, in Korea, nests were sometimes built near houses or villages (Austin, 1948:45). Nests are most often placed in the top of a large tree (e.g. birch, oak, ginko, ash, pine, elm)

Oriental White Stork 107

at a height of 4-17m; 50 nests in the Soviet Far East averaged 8 m above the ground (Winter 1982). There are a few known instances of Oriental White Storks building their nests on man-made structures, such as buildings, telephonepoles and artificial nest-platforms (Sakamoto 1966). The nest-platform is constructed of dead branches 20150cm long and lined with down, dried grass, sedge, and moss; it is 1.0-2.5 m in diameter and about 0.5-2.0 m high (Austin 1948:45-46, Austin and Kuroda 1953:333, Winter 1982). Storks often use the same nest-site year after year, if not disturbed. In the Soviet Far East the foundations of stork nests are often used as nesting sites by smaller birds, such as Tree Sparrows (Passer montanus) and White-cheeked Starlings (Sturnus cineraceus) (Winter 1982). Adults continue to bring material for the nest even after the young have become large (Winter 1982, Fei et al. 1991a). Courtship behaviour has not been fully described in wild birds. Most observations have been made on captives (see C.E. King 1988, Archibald and Schmitt 1991). Most of the described displays seem to be similar to those of the closely related and well-studied White Stork. Certain aspects of the Up-Down and Threat Up-Down in the Oriental White Stork differ in having more side-to-side head and neck movements, and the head is thrown further back. So far, no wing-pumping movements have been observed in the Oriental White Stork however, they might occur in wild birds when other storks are flying overhead. It has been reported that the Oriental White Stork's clattering is of shorter duration than the White Stork's (Shulpin 1936 cited in Dementiev and Gladkov 1951:455, Grote 1954, Hornberger 1967:78), conversely it has also been reported that it is longer (K.M. Schneider 1952, Kahl 1972c, C.E. King 1988:57). Some captive Oriental White Storks apparently clatter more rapidly than White Storks (C.E. King 1988:60), whereas others clatter more slowly (Kahl 1972c). [In both the above cases C.E. King's (1988) observations were by far the greatest in number.] The duration and intensity of displays may vary widely, according to context and environmental stimuli. Obviously more study is needed, especially of birds in the wild. In captives, mutual allopreening, synchronized UpDown displays (C.E. King 1990) and cooperative nestbuilding (K. Archibald pers. comm. 1990) are indicative of pair-bonding. Wild birds copulate on the nest both before and during egg-laying and after the young have fledged. In wild birds in China, it has been reported that the female initiates copulation by putting her head under the male's wing (Fei et al. 199la). In captive birds, mates sometimes circled each other on the nest prior to copulation (C.E. King 1988:96). The clutch-size is 2-5 eggs. Eggs are milky white to dirty white and oval to elliptical. In the Soviet Far East, 20 full clutches in 1975 averaged 3.6 eggs, and in 1976, following an exceptionally dry winter, 11 clutches averaged 2.73 eggs (Winter 1982). In captivity, eggs are usually laid at 2-day intervals (C.E. King pers. comm. 1990). Fei et al. (1991a) have reported that eggs at a nest in China were laid at intervals of 4-5 days; however, they gave no details of how these intervals were determined. There is sometimes as

much as a 5 to 7-day difference in age between the chicks in one nest (Dymin and Pankin 1975). At a Chinese nest, the birds began incubation 2 days after the third egg (of 4) was laid (Fei et al. 1991a). Both sexes share incubation duties and change places on the nest 4-6 times per day (Winter 1982). In one pair (captive), the incubation duties were shared about equally by male and female (C.E. King pers. comm. 1990), but in another pair (wild) the female incubated more and also at night (Fei et al. 199la). Incubation by wild birds, lasts about 30-34 days (Austin and Kuroda 1953:333, Fei et al. 1991a). In the Amur region (USSR), egg shells were observed to be thrown out by the parents after hatching, and were found on the ground under the nest (S.W. Winter pers. comm. 1990). Parental feedings occur about 4-8 times per day, and the male does most of the feeding when the young are less than a month old. The bulk of the food given to young in Soviet nests is fish 80-140 mm long (Winter 1982). Parents reingest any uneaten food (Fei et al. 199la). Young remain in the nest for 53-60 (Austin and Kuroda, 1953:333), 55-63 (Fei et al. 1991a) or 65-70 days (Dymin and Pankin 1975, Winter 1982). After fledging, Soviet young remain within 100-500 m of their nests for about a week. Then they gather into flocks of 8-40 birds at wetland areas in the general vicinity, before leaving on their migration in late August (Winter 1982). Nestlings perform the Up-Down display from 1 day old onward; the loudness of the clattering increases with age, as the nestling's bill grows and hardens. Adults were heard to bill-clatter in response to humans near the nest, and one female with young performed a Nest-Covering display and clattered briefly upon sighting the observer near the nesttree (Winter 1982). At Soviet nests, Winter (1982) reported that at 12 nests in 1975, 73.3% of the eggs hatched and 81.8% of the young fledged. At 11 nests in 1976, 76.7% of the eggs hatched and 91.3% of those young survived for at least 1 week (when observations ceased). Causes of egg/nestling mortality were infertility, eggs or small nestlings kicked out of nest by a parent, large nestlings falling from nest and destruction of nest by a storm. From 38 nests near the Ussuri River during 1974 and 1975, 62 young fledged, for an average of 1.6 young per nest (Shibaev et al. 1976). Nests are reportedly sometimes destroyed by Asiatic black bears (Selenarctos thibetanus) (Przhevalsky 1870:196-197, Dementiev and Gladkov 19^1:454, Grote 1954). TAXONOMY Whether the Oriental White Stork is a race of its western cousin, Ciconia ciconia, or whether it is a valid species has long been debated (Vaurie 1965:85, Kahl 1972c, 1972e, W.B. King 1981, Neufeldt and Wunderlich 1982, Creutz 1988:18). The two populations are widely allopatric; their breeding ranges are separated by 50° of longitude, c. 4000 km, and so no tests of reproductive isolation are possible in the wild. Morphological and behavioural differences between the two are considerable (C.E. King 1988, Archibald and Schmitt 1991).

108 Oriental White Stork

Because the differences are so great, we have decided here to treat the two as separate species, although they are certainly so closely related that they should be regarded as a superspecies (Amadon 1966). Morphologically, C. boyciana is much larger than C. ciconia in body size, culmen and tarsus length. The bill in C. boyciana is black in adults and greenish with yellow-orange hues in nestlings, whereas the bill in C. ciconia is red in adults and black in nestlings. The skin surrounding the eye and lores in adult C. boyciana is red, whereas in adult C. ciconia it is black. The iris in adult C. boyciana is pale yellowgrey to pale blue, whereas in adult C. ciconia it is brown. There is considerably more silvery grey or white in the secondaries of C. boyciana. Behaviourally, in natural breeding areas C. boyciana is more solitary (i.e. nests are usually 200-300 m or more apart) and is usually less tolerant of close association with man than is C. ciconia. Also, in captivity, C. boyciana is much more nervous and temperamental, needs more solitude and shows more intraspecific aggression than does C. ciconia. Noticeable differences have been described in the Up-Down and Threat Up-Down displays (C.E. King 1988, Archibald andSchmitt 1991). CONSERVATION In the Middle Ages this stork was common in Japan until 1879 and even nested within the city of Tokyo, on top of Buddhist temples (Austin and Kuroda 1953:332, Yamashina 1962b). They decreased drastically during the Meiji Era (1868-1912), mainly because of illegal hunting. Many nesting trees of the small remaining population were cut down during the Second World War. And the few remaining birds were finished off by agricultural chemicals during the 1950s and 1960s. The last successful breeding in Japan took place in 1959, when 3 young fledged from 2 nests (Ornithological Society of Japan 1975:35). Several breeding attempts during the 1960s produced no young (Fisher et al. 1969:183), and the species has now disappeared entirely as a breeding bird in Japan. Although considered a 'good luck omen' by Korean people (Austin 1948:45), the species has now disappeared as a breeding bird from that country (Gore and Won 1971:119). For the Soviet Far East, Winter (1982) lists the following factors as important in the decline of the species: forest burning and cutting of large trees used for nesting; drainage of wetlands and disturbance of hydrological conditions by

construction of power stations upstream; poisoning of food supplies by chemical pesticides; poaching; and weather disturbances, such as droughts and strong winds. S.M. Smirenskii reports (pers. comm. to G.E. King 1989) that recent droughts and shortage of nesting trees has resulted in low productivity in storks in the Amur region. In China, the species is protected by law (M. Williams pers. comm. 1990). However, hunting and trapping (for food and for zoos) as well as the conversion of wetlands to agriculture are threatening the birds (Luthin 1987). Apparently many Oriental White Storks, especially nestlings, are captured for sale to zoos. Some Chinese zoos sell them to foreign zoos. At one nest in Heilongjiang Province, China, only 4 nestlings 'escaped into the wild' during an 18-year period, most of the others having been poached by humans (M. Coulter pers. comm. 1990). Nests themselves, as well as the trees they are in, are sometimes taken for fuel (Fei et al. 199la). Storks wintering in south-central China are killed by poisons used in hunting wild ducks; they are also threatened by overfishing—which reduces their prey—industrial/ agricultural pollution of feeding areas, and other activities of man (Wang 1991). Their long bones are sometimes used as chop-sticks (Dementiev and Gladkov 1951: 454). This species is currently listed in Appendix 1 of the CITES Convention, which prohibits international trade. It is totally protected by law in the USSR, Japan and South Korea (W.B. King 1981); however, that has not prevented its extinction in two of these three countries! Eventually, captive breeding may be important in maintaining this species, especially if natural populations continue to decrease. Over 100 Oriental White Storks are currently (1990) maintained in zoo collections in several countries. Although this species does not adapt as well to captivity as does the White Stork, there has already been some success in captive breeding (C.E. King pers. comm. 1990). Reintroduction programs are highly desirable in Japan and Korea, if and when it is determined that ecological conditions there are suitable for the survival of the birds. While agreeing that captive breeding is desirable, we strongly recommend that the taking of young from natural nests to supply zoos be strictly limited to a few institutions with facilities and interest in breeding programmes.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 298.

Blacknecked Stork Ephippiorhynchus asiaticus (Latham) Mycteria asiatica Latham, 1790, Index Ornith., 2, p. 670: India Other names: Jabiru, Policemanbird, Green-necked Stork [Australia] (English); Asien-Grossstorch (German); Banaras, Loharjang, Loha sarang (Hindi); Lohar jangh, Ram salik (Bengali); Telia-hereng (Assamese); Periya-narai (Tamil); Pedda nalla konga (Telugu); Ali-kokka (Sinhalese): Hnet-kalah (Burmese)

IDENTIFICATION The Blacknecked Stork, one of the tallest, most stately and sedate of the storks, stands 129-150 cm tall. A slim black and white bird with a long, thin, slightly upturned black bill and very long bright coral-red legs. Its large size is well described by its Sinhalese name which means 'man-heron'. The head, neck, scapulars, greater and median wing-coverts, and tail are glossy black, showing purple, green and blue iridescence according to the angle of the light. The back, flight feathers, lesser wing-coverts and underparts are pure white. When spread, the white wings have a diagnostic black band, running longitudinally down their centre, both top and bottom. The large bill is a shiny black. The sexes are easily separated by eye colour: the male's is brown, the female's bright yellow and contrasts sharply with the dark feathers of the head; furthermore, males are noticeably taller. [Some published accounts, such as Andersson's (1872:282), have the eye-colour of the sexes reversed.] The most noticeable change from the regular adult plumage to the breeding plumage is that the legs become brighter. The gloss on the black portion of the plumage is heightened during the courtship period. And there is a 'skull-cap' glossed crimson-purple or bronze-purple (Iredale 1956:112, Henry 1971:382). The small, downy young are snow-white at first, the head and neck changing to dusky grey later. Their bills are straight, not upturned as in adults (Gates 1883:264, Lucas and Le Souef 1911:127, Lowther 1949:pl. 58, S. Ali and S.D. Ripley 1968:104). Immature birds are dull brown with whitish underparts, lower back, and rump; they have flight feathers tipped blackish brown and basally white and dull blackish legs. The white parts including the belly are dusky brown, turning white with age. Juveniles are patterned like adults, but the glossy black areas of adults have a brownish cast in young birds. We have seen fully adult plumed birds with legs still dark brown; such birds are probably subadults. Immatures of both sexes have brown eyes (Dharmakumarsinhji 1955:93). Vocalizations include bill-clattering loudly during Up-Down and other displays (Kahl 1973b). The species is said to make 'low grunts' at the nest (Beehler et al. 1986:60) and to utter 'dull booms' (Pizzey 1980:62), but we find these statements doubtful. 'In flight it is a noble sight with its neck extended, its long, bright red legs projecting behind the tail, and the huge white wings marked with a broad black band formed by the greater and median coverts; this pattern is repeated on the underwing' (Henry 1971:382). The flap rate was measured at 167/min (n = 20) by Kahl (1971b). The Blacknecked Stork looks 'rather skeletal' in flight (Pizzey 1980:62).

Blacknecked Stork

This species is unlikely to be confused with any other bird in its range. During the winter months, in India, it could be mistaken for a Black Stork, but the latter species is smaller, all dark above, and has a red bill. In Australia, there is a slim possibility that it might be confused, at a great distance, with a Magpie Goose (Anseranas semtpalmata), although the latter is much smaller and flies with a faster wing-beat.

DISTRIBUTION AND POPULATION This species formerly ranged throughout the entire Indian subcontinent, from Sri Lanka southeastward through Burma, Thailand to the northern Malay Peninsula, Kampuchea, southern Laos and Vietnam. It also occurs discontinuously in southern New Guinea (Irian Jaya and Papua); islands in the Torres Strait; and northern, northwestern and eastern Australia. Occasionally it wanders south as far as southern Victoria, Australia (Bell 1963, Beruldsen 1980;155, Blakers et al. 1984:62). Accidental (once: 1898) in western Java (Kuroda 1936:541). The birds appear to be sedentary, and we know of no evidence of regular long-distance migration. In Australia, it wanders far from its breeding grounds but there is nothing to suggest regular seasonal movements (Blakers et al. 1984:62). Solitary nesting habits limited the species' numbers in suitable habitat, and historically it was nowhere abundant. However, within the last two decades, numbers on the Indian subcontinent have fallen dramatically. In Pakistan it is very rare and no longer resident, being only an occasional straggler to this country (TJ. Roberts pers. comm. 1988). In India, it remains widespread but thinly spread: the states of Rajasthan, Uttar Pradesh and Assam seem to be the present stronghold, and the greatest declines have come in

111

Bihar and West Bengal; it was never common in the south (Rahmani 1989a). In Gujarat we have seen non-nesting pairs of birds in areas where traditional nest trees have been destroyed, and we suspect that failure to breed may be because of lack of alternative suitable sites (pers. obs. J.A.H.). In the Keoladeo National Park, Rajasthan, Kahl (1970:457) found 5 nesting pairs in 1967, but we have recently seen only 2 pairs and these are dependent on water supplies that are subject to poor monsoon periods. During a mid-winter census (Dec 88-Feb 89), 120 were counted in India and 4 in Sri Lanka (D.A. Scott and P.M. Rose 1989:56) [It is apparently an error that 63% of those were listed for the state of Tamil Nadu (D.A. Scott and P.M. Rose 1989:43), for Rahmani (1989a:104) states that there has been only one record for the entire state in recent years.] In Bangladesh, the species has disappeared (M.A.R. Khan 1987:230), though it is still an occasional visitor to Nepal where it sometimes nests in small numbers. In Sri Lanka, a relict population of 6-7 pairs persists in the Yala National Park (Rahmani 1989a:105). Elsewhere, scattered birds have been reported from Burma and Thailand, but there are no reports of recent nesting (P. Round pers. comm. 1989). Four birds were still present in the Hat Chao Mai National Park, near Trang, southern Thailand, in December 1980 (P. Round et al. unpub. report), where it was once found breeding (on cliffs) in 1917 (Medway and Wells 1976:96). It is believed that a few pairs remain in the Mekong Delta in Vietnam in the Melaleuca forests (Le Dien Due pers. comm. 1987) and possibly in Cambodia (V. Quy pers. comm.). It is thought that the species has been in decline for several decades due to increased cultivation, especially for rice (Luthin, pers. comm. 1988), and we confirm from our own observations over four decades that this is so. Only in southeastern Irian Jaya (Indonesian New Guinea), where an estimated 650 birds were counted during a dry-season concentration (Bishop in Silvius and Verheugt 1990), and in Australia does this species seem to still prosper. In Australia, it is widespread and common in the north. It breeds in the Kimberleys, Cape York and the Atherton regions and in the swampland areas of the Top End. We have seen it in some numbers on the flood plains of the South Alligator River. Further south it occurs in swamps associated with large river systems. In winter, there is some movement to the Pilbara and Murray-Darling regions. The only population estimates from Australia are 3743 birds in New South Wales in 1965 (Blakers et al. 1984:62). ECOLOGY The species appears to require freshwater swamps, rivers, lakes with shallow water, coastal areas and river estuaries together with suitable large trees, or at least one large tree, some distance away. Most pairs used the same territory continuously, but where such areas dry out intermittently, birds may wander to other areas that continue to provide food. A lone escaped bird that survived in the Everglades (Florida, USA), for 4 years, followed a similar pattern,

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Blacknecked Stork

remaining in one local area for most of the year but retreating to deeper water some 30 km away in the dry season (pers. obs. J.A.K.). This tall species typically forages in shallow water, Walking Slowly, with measured strides, at about 1 m/sec, and Probing in the water and submerged vegetation. The bill is held open about 75-100 mm at the tip and raised about 200 mm above the water between probes (Kahl 1973b, pers. obs.). Food location seems to be mainly tactile but sometimes also visual. Occasionally individual birds Run a short distance in pursuit of active prey, and we have seen this quite frequently in Australia. The escaped bird observed in the Florida Everglades fed by Standing and Walking, and infrequently captured fish (pers. obs. J.A.K.). A bird watched feeding for 2 h at Kakadu National Park, northern Australia, stalked forward at about 1 step/s with the bill held open and jabbing vertically into the water as deep as the junction of the mandibles. The bird caught many small fish, bit them several times in the tip of the bill, and then swallowed by raising the bill to the horizontal and tossing the head back. Food location seemed to be non-visual, as the bird jabbed in the water at regular intervals while wading, without appearing to spot any prey. It crossed and re-crossed an area about 20 X 50 m in size within a large marsh. In the 2 h period the bird was seen to capture only 10-15 small (25-75 mm) fish, although it made an estimated 2000-3000 probes in the water. Another bird, also in Kakadu Park, was seen and heard foraging almost constantly for over 4 h (02.0006.15h) on a clear, starry and moonless night. It Probed deeply and forcefully into the water, making a loud splash with each plunge, at intervals of just under 1 s, as it waded silently back and forth in a small billabong just 10 m from a tent (pers. obs. M.P.K.). Pairs forage independently but often within sight of each other; if one bird takes wing, the other will usually follow within a few seconds. We have seen small parties feed quite closely together, usually consisting of a pair of adults and juveniles. Despite this communality in family groups, the birds are normally highly territorial during the nesting season. In one instance, two birds rose into the air, chasing each other, the dominant bird grasping the wing of the other in midair in a spectacular aggressive display. In the Keoladeo National Park, India, we have seen a bird land amongst a party of feeding egrets and drive them from the area (pers. obs. J.A.H.). At the same place, a pair, accompanied by a large, but still Begging, young bird, threatened and attacked a flock of White Pelicans (Pelecanus onocrotalus) at a feeding area (Breeden and Breeden 1982). Drinking is remarkably frequent, and Kahl (1970, 1973b) reports one bird drinking 21 times in succession by placing the open bill in water and scooping upward and forward to above the horizontal to swallow. Kahl (1973b) described a Flap-Dash display, usually seen at feeding areas. One bird, usually the male, stands erect then runs with long strides and flapping wings. It dashes away from its mate, then dashes back and stands with wings widely spread. This is probably a social display, making use of the flashing white wing pattern, in pair-

formation and maintenance of the pair-bond. The FlapDash has been reported only from the Blacknecked, Saddlebill and Jabiru Storks, also the only species with large areas of white in the wing. Prey consists mainly offish, but also frogs, reptiles, crustaceans and large insects as well as other small animals (Rand and Gilliard 1967:48). Breeden and Breeden (1982) saw a Blacknecked Stork catch, kill and fly off with a Pond Heron (Ardeola grayii) at Bharatpur, India. They have also been observed catching and swallowing whole live Coots (Fulica atra) (Panday in S. AH and S.D. Ripley 1968). Blacknecked Storks are usually solitary or in pairs, but in Australia, they have been found in parties of up to 18 (Blakers et al. 1984:62). The flock of 50-60 reported in India (Whistler 1918a) was probably an error (A. Rahmani pers. comm. 1990). Earlier this century, it was said that Cambodians prize their blood as a medicine (Delacour 1929, Delacourandjabouille 1931:88). BREEDING Most nests in India are 6-25 m above ground and have been found in trees such as kadamb (Acanthocephalus kadamba), peepul (Ficus religiosd), simul (Bombax malabarica) and sheeshum (Delbergia sissoo); in Australia, nesting is in trees such as Eucalyptus spp., Nauclea orientalis and Melaleuca spp. Construction of a new nest may take more than a month (Hume and Oates 1890:266). It is built by both birds and often reused annually. This species prefers nesting in a high, isolated tree, which presumably allows easy access for its large wing-span. One nest was built on cliffs, on Telibun Island off the west coast of peninsular Thailand, near Trang (H.C. Robinson and F.N. Chasen 1936:208, Madoc 1950:57). In various parts of its range, egg-laying seems to be mainly as the rains cease (P. Slater 1959). In northern India, this is usually in October; if the rains end earlier than normal, the birds may lay in September (Hume and Oates 1890:267). Pairing is long-term, thought to be for life, so that when building or rebuilding commences there is less demonstrative behaviour than in stork species that pair anew each year. The greeting ceremony, which is not often seen, consists of Up-Down displays (Kahl 1970, 1973b) in which the mates face each other with wings fully open and fluttering rapidly in a horizontal arc. Loud clattering occurs 810 times/s for 5-10 s. Less commonly the Up-Down may be performed away from the nest, at the feeding grounds (Hume and Oates 1890:266, Barrett 1954:64). The Erect Gape can occur either at or near the nest, shortly before flight; it is similar to the homologue in the Marabou Stork, but no sound is uttered. A single snap or a short series of bill rattles can take place during the Arching display, which occurs when the birds are disturbed at the nest. Clattering by the male does occur during copulation, which starts for the most part without obvious preliminaries, except that sometimes the female presses up against the standing male. During Copulation Clattering, the female's wings are partially or wholly open, and the male flaps his

Blacknecked Stork

wings slowly for balance. Copulations average 24 s in duration (range = 15-32, n = 4). Both sexes share in building the nest. The nest is a massive structure—usually 1-2 m in diameter and 1.01.5 m deep, with a considerable cavity—built high in a large tree, made of sticks and lined with straw, leaves, rushes, grass, sometimes mud and often rags. The clutch size is 3-4, rarely 5. The eggs are broad ovals, compressed at one end, and dull white with the inner membrane dull green. Eggs become soiled and stained after a few days in the nest and become a dingy yellowish white (Hume and Gates 1890:268, S. Ali and S.D. Ripley 1968). Parents at nests in India were seen regurgitating water over the contents of nests presumed to contain eggs (Kahl 1970). Both sexes share in incubation, but the incubation period is unknown. Both parents feed the young by regurgitation onto the nest floor (S. Ali and S.D. Ripley 1968). Frith (1976:86) mentions that parents give water to the young on hot days. However, he does not give a reference for this information, and it may have come from a mis-reading of Kahl (1973b), which is not very clear on this point. Kahl (1973b) did not observe the young of the Blacknecked Stork but, instead, recorded Saddlebill parents watering young. The fledgling period is given as 100-115 days (Frith 1976:86). After they can fly, young still solicit food from their parents away from the nest (Breeden and Breeden 1982). TAXONOMY The Blacknecked Stork was formerly placed in the monotypic genus Xenorhynchus\ however, it is so obviously closely related to the Saddlebill Stork (Ephippiorhynchus senegalensis) that it makes good sense to combine the two into one genus (Kahl 1970, 1972e). Plumage differences from E. senegalensis are minor. In E. asiaticus the lesser and median under wingcoverts are white and the uppertail-coverts are white, whereas in E. senegalensis most of the lesser, all of the median and the greater underwing-coverts are black (only a narrow band of underwing-coverts on the leading edge of the forearm are white) and the uppertail-coverts are black (Sharpe 1898:311, 313; pers. obs. M.P.K.). Other morphological differences involve mainly the coloration of the bill and legs. Two subspecies have been described and were previously

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recognized—E. asiaticus asiaticus (Latham) and E. australis (Shaw) —but having studied them both in the field and in museum trays, we cannot separate the populations. Supposedly the New Guinea/Australian birds have a greener gloss on the neck (Iredale 1956:112). However, birds we have observed in both India and Australia appeared the same, and the amount of blue or green iridescence on the neck varied with the quality of the ambient light. Amadon and Woolfenden (1952) and Legge (1880:1116) also found the Australian birds difficult to separate from those on the Asian mainland. Therefore, we do not recognize any subspecific division.

CONSERVATION Habitat destruction, in the form of cutting of traditional nesting trees and draining of wetland feeding areas, is an inevitable consequence of the growing human populations throughout Asia. Blacknecked Storks are solitary and highly territorial—at least during the breeding season—and successful reproduction means little to the overall population if offspring have no place to go. Trapping for zoos, both foreign and Indian, is given as another likely cause of some of the drastic decline in India in recent years (Rahmani 1989a). Overfishing competes for food supplies, and pesticide poisoning is a possible threat in highly agricultural areas. Shooting is probably not a serious concern, because the flesh of the Blacknecked Stork is too fish-tasting to eat (A. Rahmani pers. comm. 1990). For the conservation of this species in the Indian subcontinent, Rahmani (1989a) lists the following six steps: (1) a survey to identify important wetland breeding sites; (2) protection of critical habitat; (3) legal protection of the birds themselves; (4) ecological research; (5) captive breeding; and (6) publicity, to gain public support. The Government of India has recently placed the Blacknecked Stork in Schedule 1 of the Indian Wildlife (Protection) Act, and its trapping has been banned.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 299.

Saddlebill Stork Ephippiorhynchus senegalensis (Shaw) Mycteria senegalenis Shaw, 1800, Trans Linn. Soc. London, 5, p. 35, pi. 3: no locality (=Senegal) Other names: African Jabiru (English); Jabiru de Senegal (French); Sattelstorch (German); Saalbekooievaar (Afrikaans); Abu Mijeh, Abu Milieh (Arabic); Hello (Mandingo); Baba da Jika, Zuguntumi (Hausa); Kandjendje (Kwangali); Hukumihlanga, Kokwasabi (Tsonga); Kololoane (Chuana); Nyamantando (Sotho)

IDENTIFICATION The Saddlebill Stork is one of the tallest of the storks, standing at 135-150 cm. A slim, sedate, and stately black and white bird, it has a long, thin, slightly up-turned, multi-coloured bill and very long legs. The adult's head and neck are black, and the underparts white. The white wings have a diagnostic black longitudinal band, above and below. The black head, neck, wing scapulars and tail- are glossy, showing purple or green according to the angle of the light. The tail- and uppertail-coverts are black. The large, slightly recurved bill is alternately red, black, red; the distal half (c. 15 cm) is deep red, the middle third (c. 10cm) is black, and the basal fifth (c. 6cm) is covered by a deep red fleshy lappet, which extends back to the eye. Above this red lappet is a bright yellow lappet, forming the 'saddle'. The legs are blackish to greyish green, with pink tibio-tarsal joints and toes. Males are taller and have a dark brown iris, whereas females have a pale yellow iris [erroneously reversed in Andersson (1872:282)]. Breeding plumage is not known to differ noticeably from regular adult plumage, except that the iridescence on the black regions of the head and neck is more pronounced. One or both members of the pair often have two small yellow, sometimes red, wattles that hang down 15-35 mm under the base of the bill, like 'stirrups' under the 'saddle' (Kahl 1973b; Brown et al. 1982:185). A small (45-50 mm diameter), round or oval-shaped patch of bare, dark red skin is sometimes exposed in the middle of the breast (Chapin 1932a:462); it is often hidden by feathers. These colourful areas of bare skin may indicate fully mature birds in breeding condition. Details of the development of nestlings are minimal. The downy young are fluffy and white, the head and neck later changing to a darker, smoky grey. The bill is horn-coloured and 'not particularly long' (Mackworth-Praed and Grant 1962); by 21 days the bill is still only 88mm long (Pitman 1965). At 45 days, nestlings are still largely downy, with dark feathers showing on the wings (Brown et al. 1982:186). The plumage of large nestlings is mainly a 'lustreless greyish black; the head and neck are covered with grey down; the bill is blackish, with only a slight indication of red at the tip; the "saddle" at the base of [the bill] is blackish and edged with a fringe of whitish down' (A.L. Butler 1905). Other, captive reared, young still did not show red on the bill at 6 months of age (F. Junor, pers. comm. 1965). Soon after fledging, young still have the head and neck covered with thick and woolly, snuff-coloured down (Niven and Niven 1966b). Juveniles show little or no red in the bill or legs; areas that are black in adults are dark grey; areas that are white in adults are pale grey (C.W. Benson and C.R.S. Pitman 1958). By the second year, the bill is dull red and black, and the 'saddle' is a dull yellow (Brown et al. 1982:185). Vocalizations, if any, are unknown in adults, although they do bill-clatter during some displays, as does the closely related Blacknecked Stork of Asia. Begging calls of nestlings are less loud than those of the colonial storks (Kahl 1973b).

116 Saddlebill Stork

(Kasoma and Pomeroy 1987). In the Kafue flats, Zambia, Dowsett and de Vos (1965) counted an average of 27 Saddlebills in an area of 58 square miles (=151 km 2 ), 1 bird/ 5.6 km2; further northeast in the Bangweulu Swamps, Zambia, G.W. Howard and D.R. Aspinwall (1984) estimated a density of 1 bird/5.8 km2. In South Africa, where the species occurs regularly only in Zululand and the eastern Transvaal, Siegfried et al. (1976) estimate a total population of 'fewer than 50 birds*. ECOLOGY

An impressive bird on the wing, with large areas of white in the flight feathers and a span of over 2.5 m, it often soars on thermals until nearly out of sight from the ground. The flap rate was measured as 150/min (n = 13) (Kahl 1971b). It flies with the neck and feet fully extended, and, because of its extreme length, 'it flies in a curiously bent position with head and feet lower than the body' (Mackworth-Praed and Grant 1962:83). The tallest African stork, the Saddlebill is slimmer and more graceful than the Marabou. Juveniles, in grey plumage, might be mistaken for juvenile Yellowbilled Storks but are much taller. Saddlebill juveniles, if not in the company of their parents, might possibly be misidentified as young Marabous (Turner in Pitman 1965). DISTRIBUTION AND POPULATION This species occurs throughout tropical Africa, from Senegal to Sudan and Ethiopia, south to northern Namibia, northern Botswana and northern South Africa (Transvaal, Zululand). It occurs sporadically further south but is quite rare south of the Orange River. The birds appear to be sedentary, and we know of no evidence of regular long-distance migration. They are to some extent nomadic, especially during drought conditions or when the large rivers are in flood (Chapin 1932a:463, Bannerman 1953:177). The Saddlebill Stork is widespread but never very numerous. Its territoriality and solitary nesting limits numbers in suitable habitat, and historically it was nowhere abdundant. There are 6 recorded breeding sites in Kenya, 5 in Uganda, and 11 in Tanzania (Kasoma and Pomeroy 1987). The record of 100 birds at Lake Turkana, Kenya (Kasoma and Pomeroy 1987) is apparently an error. Along the 52km length of the Kazinga Channel, in western Uganda, only 1 pair of Saddlebills was found over an 8-month survey

The Saddlebill Stork is found in aquatic habitats in open or semi-arid country, generally avoiding deep forest. It favours shallow freshwater marshes, rivers, lake shores and floodplains, with suitable large trees, often some distance away, for nesting. Most pairs use the same territory continuously, except during drought, when the birds may wander nomadically. Birds of this tall species typically forage in shallow water, Walking Slowly and Probing in the water and submerged vegetation. They usually take 0.7-1.0 steps/s and each stride covers 0.8-L2 m. The bill is held open about 5070 mm at the tip and raised about 200 mm above the water between probes (Kahl 1973b). There is some evidence that large fish may be impaled with one of the mandibles, rather than just grasped (Bell-Cross 1974). Most prey finding appears to be tactile. However, occasionally individual birds Run a short distance, often with wings open for balance, in pursuit of active prey. Foot Stirring has been reported (Brown et al. 1982:185). Birds have been seen washing frogs in water before eating them (Curry-Lindahl 1961). Pairs forage independently but usually within sight of each other. Small parties sometimes feed quite close together and usually consist of an adult pair and their offspring. Despite the communality in family groups, Saddlebills are usually highly territorial (Brown et al. 1982:186); however, this territoriality may become weaker outside the breeding season (G.W. Howard and D.R. Aspinwall 1984). Most foraging is diurnal, although a bird has been reported feeding at night at an artificially lighted waterhole at Mulika Lodge, Meru National Park, Kenya (Lassus 1973). Kahl (1973b) described a Flap-Dash display, usually given at feeding areas. The male (usually) stands erect, runs away from its mate with long strides and flapping wings, turns and runs back to the mate, and stands with wings widely spread. This is almost certainly a social displaymaking use of the flashing white wing pattern —and probably functions in maintenance of the pair-bond, or even in pair-formation. Prey consists mainly offish, but also frogs, reptiles, crustaceans, some molluscs and other small animals. Where they are common, lungfish (Protopterus aethiopicus) are favorite food (F J.Jackson 1938:76, Kahl 1973b). Adults commonly swallow fish 15-30 cm in length and weighing up to 500 g, occasionally 1300 g. They have also been reported eating large water beetles (Chapin 1932a:463), other insects

Saddlebill Stork

and carrion (Heuglin 1869-73:1113). Prey is often washed before swallowing head-first; and birds commonly drink after eating (Brown et al. 1982:186). Drinking is frequent, with the bird placing the open bill pointed forward in the water and scooping upward and forward to above the horizontal to swallow. Snipping off catfish spines before swallowing the fish has been seen (A. Morris 1979). Scavenging on fish regurgitated by cormorants has been reported (Kasoma and Pomeroy 1987). This species usually occurs solitarily or in pairs. Small groups seen are often parents and several young (C.W. Benson and C.R.S. Pitman 1958). The largest recent aggregation that we know of is a group of 12 birds seen in Zambia (G.W. Howard and D.R. Aspinwall 1984), although Bannerman (1953:177) reported 'flocks of twenty to thirty may be observed at times on the sandbanks' and 'one record of thirty seen on the Upper Niger'. Curry-Lindahl (1961) reported a pair of Saddlebills that stayed near a herd of hippopotamuses and moved to the centre of the herd when humans approached. Saddlebill Storks roost on trees. Normally they are rather shy but they can become tame in areas such as national parks where they become accustomed to vehicles (Brown et al. 1982:185).

BREEDING At the northern and southern ends of the range, Saddlebills normally begin nesting late in the wet season or at the start of the dry season; in equatorial regions they may nest during the rains. In both cases, the young usually fledge at the height of the dry season, when food is concentrated and relatively easy to obtain. The nest is usually built in the top of a large tree (e.g. Euphorbia ingens, Euphorbia dawei, Acacia giraffae, Kigelia pinnata, Adansonia digitata). Especially in Zambia, they often build nests in trees (e.g. Albizzia harveyi) growing out of termite-mounds (C.W. Benson and G.R.S. Pitman 1958). Unless in a well-protected area such as in an inaccessible swamp, the nest is often as high as 20-30 m from the ground (Brown et al. 1982:186). They have been reported as sometimes nesting on cliffs (Mackworth-Praed and Grant 1962:83, Clancey 1964a:46), but this needs confirmation (Pitman 1965). Saddlebill Storks are reported to occasionally use old nests of other large birds, such as Tawny Eagles (Aquila rapax). Sometimes other large birds, such as Secretary Birds (Saggitarius serpentarius), usurp unoccupied Saddlebill nests as well (PFIAO nest-record cards, University of Gape Town). A SaddlebilPs nest in western Kenya was surrounded, on two consecutive seasons, by many nests of Blackheaded Weavers (Ploceus cucullatus) that arrived after the storks began to nest (Kahl 1973b). These smaller birds probably gain considerable protection from predators by establishing their colony near a Saddlebill's nest (Moreau 1942). Pairing is long-term, thought possibly to be for life, so that when nesting starts the birds are less demonstrative than stork species that mate with different partners each

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year. Sometimes many days pass during the breeding season without any obvious social displays taking place. This is the only species of stork for which the Up-Down display has not been adequately described in nature. It was not seen at a Kenyan nest in 146 h of observation over two seasons (Kahl 1973b). Professor H. Mendelssohn (pers. comm. 1972) observed the Up-Down in a pair of captive Saddlebills at the Tel-Aviv Zoo. In this pair, the period of display was prolonged, because the birds broke their eggs and had to re-lay. They gave Up-Downs, as a greeting when one bird was on the nest and its mate approached: The approaching partner came with quick and very long steps, head and neck stretched forward. Arriving at the nest, he (or she) spread his wings fully and began a quick bill-clattering, lowering the head and sometimes lifting it again to [the] horizontal position and even higher. Then the wings were closed, the head lowered and the bird generally pecked at sticks in the nest. The partner on the nest displayed similarly, but less intensively, beginning after its mate. The wings were less widely spread, sometimes not at all. Other social displays are detailed and illustrated in Kahl (1973b). In the Arching display, which often occurs when the birds are disturbed, the stork stands erect, leans forward and compresses its plumage; a loud snap or a short series of bill-rattles may be given. The Erect Gape occurs on or near the nest, usually before flight; it is similar to a homologous display in the Leptoptilos storks, but no sound is uttered by the Saddlebill. Copulation has not been described in the Saddlebill. We once observed a Forward Display by a captive immature Saddlebill as it threatened several Yellowbilled Storks in its feeding-pond. The Saddlebill stood erect, spread its wings, and clattered its bill slowly, as it approached the smaller storks (Kahl 1973b). A similar display was given, as a threat, to humans by a captive adult in the Tel-Aviv Zoo (H. Mendelssohn pers. comm. 1972). The bird approached with widely spread wings and lunged with quick billclattering. Frank Junor (pers. comm. 1965) described how captive-reared juveniles, once they could fly, 'would often fly low over the head of a human, lower their heads in flight and snap with a most audible clack above the head of the stranger'. This species does not necessarily breed every year in a given locality (Kemp 1974:15, Brown et al. 1982:186). It is sometimes said to be shy and nervous at the nest (Pitman 1965), but a pair near Kisumu, Kenya, nested in two successive years in a tree within 20 m of a busy, tarmac highway (Kahl 1973a). Both adults cooperate in building the nest, but at a Kenya nest the male brought most of the sticks. The platform is a massive structure made of sticks and lined with reeds, sedges, or mud. It is built or repaired by both birds and commonly reused year after year. Nests may be up to 2 m in diameter and 50 cm deep (occasionally larger), with a central depression deep enough to completely hide a sitting bird (Pitman 1965, Brown et al. 1982:186). The eggs are oval, dull white, slightly glossed and coarse-

118 Saddlebill Stork

textured (Pitman 1965). The clutch size is 1-5, usually 2-3; the mean of 54 clutches was 2.8 eggs (Brown et al. 1982:186). According to Pitman (1965), the clutch size north of the Equator is usually 3 eggs, whereas in Zambia and Zimbabwe clutches of 4 are more frequent. Both sexes incubate, in shifts that vary from 1 h 25 min to 5 h 50 min. The incubation period has not been accurately recorded, but has been estimated at 30-35 days (Brown et al. 1982:186). Old birds often sit in the nest on their tarsi (Heuglin 1869-73:1113). Parents often bring water back to the nest and regurgitate up to 1-2 litres onto the nest and eggs(Kahl!973b). Both parents feed the nestlings by regurgitation onto the floor of the nest. Young are also given water by their parents, who stand upright, hold their bills high, and regurgitate it over them (Kahl 1973b). Young perform Begging displays, either standing or on their tarsi, with rhythmic 'up-down' nods of the head, slightly opened wings and rather weak braying vocalizations; they sometimes billclatter to the parent. Unattended nestlings threaten intruders with a Nestling Bristle display, which includes erection of head, neck, and upper back feathers, and clattering of the bill (Kahl 1973b). Chicks are brooded closely for the first 10 days. The fledgling period is not accurately known but is probably at least 100 days (Kahl 1973b, Brown et al. 1982:186); this is considerably longer than Pitman's (1965) estimate. Fledged young still give Begging displays and are fed by the parents after they leave the nest (Niven and Niven 1966b). At what age juveniles become fully independent and separate from their parents is not known; however, it is probably several months after they leave the nest. Family groups are often seen together long after the breeding season, and, according to Pitman (1965), 'they are still together when the red colour is appearing on the bill [of the juveniles] and when the yellow "saddle" begins to show'.

Nesting success has not been recorded. 1-2 young normally fledge from a clutch of 3, apparently more in southern Africa (Pitman 1965, Brown et al. 1982:186). Sample counts at Lochinwar and Blue Lagoon, in Zambia, during AugustNovember at the end of the breeding season showed 'no more than 5% immature birds' (Dowsett and de Vos 1965). TAXONOMY The Saddlebill Stork (genus Ephippiorhynchus) is so closely similar to the Blacknecked Stork (formerly classified in the monotypic genus Xenorhynchus) that they have been combined (Dekeyser 1962, Kahl 1972e). Plumage differences from E. asiaticus are minor. In E. senegalensis most of the lesser, and all of the median, and all of the greater underwing-coverts are black (only a narrow band of underwing-coverts on the leading edge of the forearm are white) and the uppertail-coverts are black, whereas in E. asiaticus the lesser and median underwing-coverts are white and the uppertail-coverts are white (Sharpe 1898:311, 313; pers. obs. M.P.K.). Other morphological differences involve mainly the coloration of the bill and legs. CONSERVATION The population is probably stable but could easily become threatened (Brown et al. 1982:185). The taking of nestlings for sale to zoos could make an impact on a bird that nests solitarily and is nowhere abundant, such as the Saddlebill. In 1989, the Dallas Zoo (USA) paid US$6000-6500 per bird (P. Osten pers. comm. 1990). Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 300.

Jabiru Stork Jabiru mycteria (Lichtenstein) Ciconia mycteria Lichtenstein, 1819, Abhandl. K. Akad. Wissen. Berlin, Phys. Kl. (1816-17), p. 163; (based on 'Jabiru' of Marcgrave, 1648, Hist. Rerum Nat. Brasiliae, p. 200): Brazil. Other names: Garzon soldado, Garzon bianco, Cigiiena de cuello pelado, Galan sin ventura, Juan grande, Baguari (Spanish); Blaasman (Dutch); Doih (Warrau); Tararamu (Macusi); Mora-coyasehre (Arawak); Tuyuyu (Guaram); Tuiuiu-corol, Jaburu-moleque, Jaburu (Portuguese)

IDENTIFICATION The Jabiru Stork is a robust, all white-plumaged stork, standing 120-150 cm tall and having a black head and black, massive, slightly upturned bill. The common name 'Jabiru' apparently derives from the South American Indian word 'Yabini', meaning 'blown out with the wind', and referring to the loose, brilliant red skin of the lower neck, which becomes inflated during threat, anger or courtship. This is the only stork with all-white feathers, including wings and tail. The head and neck are featherless, except for a silvery grey tuft of feathers on back of the crown. The upper neck is dull black skin that abuts a band of loose pink skin, forming an inflatable throat sac around the lower third of the neck; it inflates and turns fiery scarlet during anger, excitement or physical exertion (P.L. Sclater and W.H. Hudson 1889:106, Kahl 1971e, 1973b). A small oval-shaped patch of bare skin also occurs on the chest; although normally covered with feathers, it becomes exposed when the bird stands erect, as before take-off or during displays. The flight feathers (both primaries and secondaries) are white, making it the only all-white species of stork. The legs and feet are black, and the iris is brown in both sexes. The sexes have similar plumage, although the females are slightly smaller and have thinner and more upturned bills (Kahl 1971e, B.T. Thomas 1985). Individual birds can be distinguished by variations in the configuration of the black/red junction on the bare neck (Sick 1985, B.T. Thomas pers. comm. 1990). Some birds even have small spots of red within the black or black within the red. Such differences, which seem not to change as the bird ages, may be helpful to researchers studying birds that cannot be individually marked. Plumage, which is often soiled at other times of the year, moults into immaculate white in the breeding season. The bare skin of the neck is smooth and silky to the touch (B.T. Thomas 1985). The red band on the lower neck brightens and, because of increased sexual excitement, is a rich scarlet much of the time. Hatchlings are covered—except for the naked lores and orbital skin—with a greyish white down, with localized yellowish tinges, which is especially thick on the head and uppersides, and more scantily covered on sides and undersides. They have straight, rather than recurved, bills (Lloyd 1895, Bent 1926:69). Juvenile plumage is mottled brownish grey, the feathers being light grey with darker tips. A sparse, dusky brown, hair-like down is retained on the head and neck, being longest on the occiput where it forms a blackish crest. The undersides are mostly white, and the white wing feathers tipped with grey or buffy brown. The remainder of white plumage develops gradually over the first 2 years, beginning at the undersides and above the tail (Nutting 1882, Hagmann 1907, Bent 1926:69). Generally this species is voiceless, but it bill-clatters loudly in displays and in alarm. A coughing sound is occasionally given in Up—Down greeting displays and during copulation.

Jabiru Stork 121

Having the second-longest wingspan of any New World bird-'nearly eight feet' ( = 2.4m) (Hudson 1920:117)-the Jabiru flies gracefully, with legs and neck extended, the latter appearing disproportionate because of a bulge formed by the loose skin on the neck (Roosevelt 1914:91). This bulge gives the appearance of flying with the neck partially retracted. Taking off requires a running start. The bird alternates a few slow flaps with gliding, and frequently uses thermals for soaring (Chubb 1916:159, Wetmore 1926:60, Sick 1985). Dickey and van Rossem (1938:87) commented that 'There is a smoothness and flow to the wing stroke which is astonishing in so large a bird*. The flapping rate is 179/min (n = 13) (Kahl 1971b). One of the world's largest storks, the Jabiru is considerably larger than the American Wood Stork and has a massive upturned, rather than downturned, bill. Its size and black head and bill distinguish it from the Maguari Stork. It is a stately bird, that stands tall in the marsh, often alone or in pairs. In flight, it is similar to an American White Pelican (Pelecanus erythrorhynchos), but is larger, with extended neck and legs and all white wings. DISTRIBUTION AND POPULATION The Jabiru Stork breeds from southern Mexico (Campeche, Tabasco, possibly Quintana Roo), locally through the lowlands of Central America (Belize, Guatemala, Nicaragua, Costa Rica) and most of tropical South America east of the Andes. Its overall South American range is from Colombia (lower Magdalena), Venezuela and the Guianas, southward through eastern Peru (rare on the coast), eastern Bolivia, Brazil, Paraguay and Uruguay to northern Argentina (Corrientes, Santa Fe Provinces; accidental to Buenos Aires)

(Meyer de Schauensee 1970, Blake 1977:189-190, Koepcke 1963, pers. obs. J.A.K.). Populations are concentrated in the vast wetlands of the Colombian and Venezuelan Llanos, the Brazilian Pantanal (Mato Grosso) and the ParaguayanArgentine Chaco. Vagrants have been recorded north to Texas and Oklahoma (Haucke and Kiel 1973, Oberholser and Kincaid 1974:123, K.A. Arnold 1978). In South America, the Jabiru Stork is generally resident and non-migratory (B.T. Thomas 1985), Reportedly, it leaves Belize between June and November (Luthin 1984a). It is said to leave the main rivers in the Amazon region of eastern Peru during the high-water season and to move to smaller streams and higher country (Koepcke 1963:403). Formerly, it was found in Honduras only in December and January (Monroe 1968); recent reports are lacking. It does wander, being recorded rarely as far north as Texas and Oklahoma, as far west as coastal Peru, and as far south as Buenos Aires, Argentina (Olrog 1963, Hughes 1970, Haucke and Kiel 1973, K.A. Arnold 1978, C.T. Clark 1982). Populations in Central America have historically been small and highly localized. In recent surveys, only a few nests have been documented in most areas of the region, and the entire Central American population is estimated at between 150-250 individuals. A maximum of 83 adults and 6 juveniles were counted along the Gulf coast of Mexico in May 1978 (Knoder et al. 1980). Populations in northern South America are larger, but nevertheless relatively low. Only about 200 individuals were found on surveys in the Llanos of Venezuela in 1983 (Ramo and Busto 1984), although their dispersed nesting makes surveys difficult. Coastal populations are generally smaller; a maximum of 100 birds was seen along the Surinam coast (Spaans 1976a). The largest populations are found in the Pantanal and eastern Chaco in south-central South America. Brooks (1991) counted one bird per 4.38 km2 in the Paraguayan Chaco. It is said to be very rare in Uruguay (Cuello and Gerzenstein 1962:35). ECOLOGY Jabiru Storks favour extensive shallow marshes, wet meadows, ponds, rivers and pastures, both inland and along the coast. They are birds of open country but prefer wooded areas nearby for roosting and nesting. This species feeds by Standing or by Walking Slowly, with the neck extended forward and the bill pointed downward, while non-visually Probing every few steps. It will also Stab at food items visually, Probe quickly, or Run and Hop short distances after prey. When a prey item is captured, the birds frequently carry it ashore, bite it several times before swallowing. Buried eels are sought by Probing into the mud. In Venezuela, 43 individuals located about half of their prey by feel and half visually (B.T. Thomas 1985). Jabirus usually feed singly or in pairs, but when prey becomes concentrated, such as by falling water levels, they join loose, sometimes large, flocks, and mixed-species aggregations. They also are attracted to fires. Large numbers

122 Jabiru Stork

were observed gathered during a mouse plague in Argentina. 'Armies of these majestic white birds were seen stalking over the grass on all sides, or at the close of day winging their flight to the distant watercourses in a continuous flock' (Hudson 1951:58-59). Naumburg (1930:89) observed 200300 together at ponds in the Pantanal of the Mato Grosso, Brazil. Jabirus are resident in large territories, which they defend (B.T. Thomas 1981). Males are highly aggressive in captivity both to conspecifics and to others, including humans (Shannon 1987). In Venezuela, a Jabiru aerially chased an American Wood Stork that had caught a 30cm eel; however, the owner retained and managed to swallow the prey (B.T. Thomas 1985). The food is primarily fish and eels (e.g. Symbranchus marmoratus in Venezuela), but they can take many kinds of prey including insects, snails (Ampullaria guianensis and A. papyracca, in Guyana), mussels, crabs, frogs, small mammals and snakes (Schomburgk 1922-23:121, Pelzeln 1868-71:305, Chubb 1916:159, Meyer de Schauensee and Phelps 1978:22, Ogden and Thomas 1985b, B.T. Thomas 1985). BREEDING The Jabiru Stork nests singly on the tops of tall broadleaved trees (e.g. Bombax munguba, Calycophyllum spruceanum, Ceiba pentandra, Eriodendron anfractuosum, Enterolobium contortisiliquum), and often in palms (e.g. Copernicia tectorum, Butia yatay). In coastal Mexico, nests are in tall red mangroves (Rhizophora mangle). Nests are sometimes near mixed colonies of other species but usually not near other Jabirus, which are driven away from the nesting area. B.T. Thomas (1981) found 5 active nests within a 250 ha area in Venezuela. The large, stick nest is placed in or near the top of a large tree, often one that towers over its neighbours and affords a view of the surrounding country. Many authors have commented on the Jabiru choosing the largest tree around for nesting (Roosevelt 1914:90, Bent 1926:67-68, Naumburg 1930:90). We have observed a typical situation in the species' northernmost nesting site in the Usumacinta Delta, Mexico. The nest was situated in a tall mangrove tree adjacent to mangrove swamp and open marshes. The tree projected well above the surrounding canopy with a commanding view of the countryside. On rare occasions (Schomburgk, in Bent 1926:67) the birds may nest atop large rocks. Nests themselves are flat, massive structures, up to 2 m across and 1 m deep, made of branches up to 5 cm thick and 1.5-2.0 m long, intermeshed with smaller twigs and lined with dry grass. Nests are often reused in successive years; however, nests in palms are not reused if the palm dies. A new nest in Venezuela took nearly 2 months to complete (Hagmann 1907, Kahl 1971e, B.T. Thomas 1981). The nesting season is variable across the range (see Appendix) and seems largely determined by seasonal rainfall patterns. In both Venezuela and Argentina, birds nest from the latter part of the rainy season into the dry season (Kahl 1971e, B.T. Thomas 1981). In Surinam, they nest in the long dry season (Spaans 1975b). Especially during the nesting season, Jabirus may exhibit

intra- and interspecific territoriality near the nest and at feeding areas. B.T. Thomas (1985:928) recorded that Jabirus twice chased Maguari Stork fledglings that flew within 100m of Jabiru nest sites. A captive male has been observed, threatening a tapir by throwing his head back until the bill was vertical and snapping the bill, and males were very aggressive toward their keepers in the breeding season (Shannon 1987:245). It is likely that solitary pairs remain mated in successive seasons and use the same nesting site, so courtship display is not extensive (Kahl 1971e, 1973b). On the feeding grounds, pairs engage in Flap-Dash displays, which may function in pair-bond maintenance. In captive birds (Shannon 1987), pair-formation is initiated by the female, who overcomes the aggressive territoriality of the male by using a display in which she rushes up to the male with her head down while coughing and bill clattering. The first greeting displays also appear to be initiated by the female, with the male responding. After pairs have formed, courtship displays are those typical of storks (Kahl 1973b). The Up-Down display in greeting involves the birds facing each other, with bodies erect, wings closed, necks upward and bill nearly horizontal. The birds then bill-clatter loudly and rapidly (8-10 times/ s), while slowly swaying their heads and necks from side to side and slightly raising and lowering the bill across the horizontal plane. The Flap-Dash is a spectacular social display, usually seen away from the nest at feeding areas. A member of a pair, usually the male, suddenly stands erect, runs away from its mate with open and flapping wings, then turns and dashes back to the mate, and stands for a moment with wings fully spread (Kahl 1973b). This display has been reported only in Jabiru, and in the Blacknecked and Saddlebill storks (genus Ephippiorhynchus), the only three species of stork with white flight feathers in the wing. In wild Jabiru Storks this display was usually seen performed by pairs, but Shannon (1987) saw it performed in captivity by single birds as well. Arching displays, with loud bill snaps, occur in response to disturbance near the nest. Copulation takes place on the nest, usually begins without preliminary display and lasts 13-19 s; Copulation Clattering may involve gasping coughs from one of the birds (Kahl 1973b, B.T. Thomas 1981). During early nest-building, the sticks are mainly gathered by the male and positioned by both birds (Kahl 1971e, Shannon 1987). Both sexes incubate and feed the young. When disturbed at the nest, adults give loud claps of their bill. (Lloyd 1895, Schomburgk 1922-23:121). The eggs are a dirty white, laid in clutches of 2-4, usually 4 but sometimes 5 (Lloyd 1895; Hagmann 1906; Kahl 1971e). 11 nests studied by B.T. Thomas (1981) in Venezuela contained the following: single chicks (2 nests), 2 chicks (5 nests), 3 chicks (2 nests), 4 chicks (1 nest), or 5 chicks (1 nest). The fledgling period was estimated at 10-11 weeks, but this is based upon observations at a single nest and may be something of an underestimate (B.T. Thomas 1981, 1985).

Jabiru Stork

TAXONOMY The Jabiru Stork shares many characteristics with the Saddlebill and Blacknecked Storks (genus Ephippiorhynchus] on the one hand, and with the Lesser and Greater Adjutant and Marabou Storks (genus Leptoptilos) on the other (Dekeyser 1962, Kahl 1971e). The Jabiru appears closer behaviourally to the former and morphologically to the latter. We retain its monotypic genus to demonstrate this intermediate taxonomic position (Kahl 1972e). CONSERVATION Habitat destruction and human disturbance is recognized as a primary threat to the Jabiru throughout its range (Luthin 1987). Wetland drainage, especially for agricultural conversion, tree cutting and hunting are the principal threatening human activities.

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The Jabiru has become a symbol of conservation and wetland protection in Central America, particularly in Belize and Costa Rica. It is considered to be threatened in Central America and endangered in Nicaragua (Luthin 1984a). Specific needs include population ecology studies and the creation of Jabiru reserves to protect wetland habitats important to many species (Luthin 1984a). In Surinam, the species is 'uncommon nowadays and reduced by relentless shooting' (Haverschmidt 1968:28). It is, in fact, considered to be a gamebird in the Amazon, and larger nestlings are particularly sought (Sick 1985).

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 301.

Lesser

Adjutant Stork Leptoptilos javanicus (Horsfield) Ciconia Javanica Horsfield, 1821, Trans. Linn. Soc. London, 13, p. 188: Java Other names: Hair-crested Adjutant, Smaller Adjutant, Javan Adjutant, Hair-crested Stork, Marabou Crane (English); Marabout d'Asie (French); Javaanse Maraboe (Dutch); Malayischer Kropfstorch, Stirnplatten-Marabu, Sunda-Marabu (German); Chinjara, Chandana, Chandiari, Bang gor, Chota garur (Hindi); Madan-chur, Madantak, Modun-tiki (Bengali); Tokla moora [='bald head'], Bor tokola (Assamese); Dodal kongo, Dodal gatti gadu (Telugu); Meva kokku, Mana (Tamil); Mana kokka (Sinhala); Vayalnaiicken (Malayalam); Don-mi-gwet (Burmese); Tontsap (Arakan); Bangau, Bango (Javanese); Burong botak [='with bald head'], Burong babi (Malay); Burong Kambing, Burong Gajah (Sumatran); Hoang (Hainan Chinese)

IDENTIFICATION The Lesser Adjutant Stork is a massive, heavy-bodied stork, standing 110—120 cm tall, if erect, but often shorter because it stands hunched with the head drawn in on the shoulders. It has a dark grey back, white underparts, and sparsely feathered head and neck. It is generally dignified and sluggish in demeanour. This stork is glossy metallic blue-black above, white below, with sparse hairlike feathers on the nearly naked, reddish yellow head and neck. The bill is dull yellow, with the tip whitish and the base tinged red to pinkish. The bare crown or 'frontal plate' varies from greenish brown to light yellowish tan. The featherless face and neck are a mixture of pale pink and bright tumeric-yellow, tinged with brick-red. The iris is pale slaty grey to pale bluish grey. The legs and feet are greenish brown to almost black but are usually difficult to see because they are covered with white excreta. The sexes have identical plumages. However, the males are noticeably taller and heavier, with a more massive bill. This is a distinction that can be recognized in wild birds. At the start of breeding, the back remains a dark slaty blue-black, and small, oval-shaped copper spots about 2.0 cm long develop near the tips of the median secondary coverts (these spots are not on the greater coverts, as sometimes stated in the literature). Narrow white edging appears on the scapulars and 4-5 of the innermost greater secondary coverts. The other greater secondary coverts remain dark. Nestlings are covered with white down, except for the head and face, which are largely naked. The juvenile is similar to the adult, except that the upperparts are less glossy, and they lack the copper spots on the coverts. The head and neck are more densely feathered than in the adult. During the Up-Down display, given at the nest, birds utter a series of high-pitched squeaks and cowlike 'Moos'. Vocalizations heard during these displays were hoarser and more rasping than those given by Greater Adjutant Storks. Their quality was reminiscent of some of the vocalizations of young Marabou Storks at about the time of fledging (Kahl 1970, 1972a). M.C. Thompson (1966) reported calls resembling the 'deep growl of a dog' from non-breeding birds in Borneo. A loud 'Woofing' wingnoise can be heard near the nest.

126 Lesser Adjutant Stork

Unlike other storks that fly with the neck outstretched, birds in the genus Leptoptilos fly with the head drawn back on their shoulders (Glenister 1951, Henry 1971, Kahl 1972a). The Lesser Adjutant commonly soars when thermal air-currents are available; soaring/gliding flight is a favoured mode of transport between nesting/roosting areas and foraging grounds. The flapping rate has been timed at 153/min(w = 8)(Kahll971b). The Lesser Adjutant can be most easily confused with the Greater Adjutant. It is distinguished by its smaller size; somewhat thinner and slightly recurved bill; dark slateblack (as opposed to paler bluish grey) back and upperside to wings; pale light-tan forehead; more dense down and hairlike feathering on the neck and back of head, most pronounced in a small crest on the occiput; much smaller (rarely even noticeable) gular pouch, which is actually only a band of expandable, bare, reddish skin around the base of the neck and is not pendant, feathers that form a white ruff at the dorsal base of the neck with large black spots; white (not sooty grey) undertail-coverts; tibia black up to the feathers. These differences are illustrated here, and in colour in plate VIII of Kahl (1971f). The adult Lesser Adjutant resembles the Juvenile Greater Adjutant in many aspects of morphology. DISTRIBUTION AND POPULATION The Lesser Adjutant Stork occurs in tropical Asia and adjacent islands. It was once widespread from India (mainly eastern and southern) and Sri Lanka to southern China, Vietnam, Thailand, Malaysia, Sumatra, Java and Borneo. As far as is known, it is only an occasional visitor to Nepal and Bali (M.D. Bruce 1982) and apparently no longer breeds in Burma or Thailand. This is mainly a non-migratory species. However, it can be nomadic, especially in response to rains. It may be locally migratory, although this needs confirmation. The Borneo population is reported to increase in the winter, with up to 200-300 birds seen there in January (Smythies 1981:31). There are few censuses available. Formerly this species

was less abundant than the Greater Adjutant (E.C.S. Baker 1929:330); today, however, the Lesser is the more common species in almost all areas of their common range. The population in India is now more thinly scattered, with recent reports of nesting only in Madhya Pradesh, Assam and perhaps the Indian Sundarbans (Luthin 1987, P. Saikia and P.C. Bhattacharjee pers. comm. 1990). During a survey in Assam, India, during 1987-89, it was estimated that there were about 400 Lesser Adjutants. The survey found 40 nests in Orang, 11 in Manas, 20 in Kaziranga, 11 in Lowkhowa reserves, and 53 nests outside protected areas (Saikia and Bhattacharjee 1989a,b). Small breeding colonies in Yala National Park, Sri Lanka, number about 100 pairs. A single colony is known from Vietnam, and no information is available from Laos and Kampuchea (Luthin 1987). A population of about 200 birds is estimated to occur on the west coast of peninsular Malaysia (D. Wells pers. comm. 1989). In Sumatra, its main stronghold is in South Province. As many as 620 Lesser Adjutants were counted there during 1984—89. The entire Indonesian population (Sumatra, Java, Kalimantan, Bali) is believed to be no more than 2000 (M.J. Silvius and WJ.M. Verheugt pers. comm. 1990). Elsewhere no data exist on population numbers. ECOLOGY The details of the Lesser Adjutant Stork's ecology are little known. It inhabits flood-plains, swamps, pools in forests, marshes, drying river beds, lakes, and rice-paddies. It sometimes moves to agricultural fields and grasslands. Particularly in Southeast Asia and the Greater Sunda Islands, it is a bird of mangroves and coastal mudflats. The Lesser Adjutant is apparently less of a scavenger than the Greater Adjutant or Marabou Storks. It feeds alone or in small flocks and shows considerable alertness and activity when hunting for food. Most food location appears to be visual, with the bird Walking Slowly on dry ground or shallow water and Grabbing prey in the tip of its bill. We have seen up to 6 birds perched on top of wooden telephone poles lined across a marsh in Assam, apparently scanning the marsh from their high vantage point for potential prey (J.A.H. pers. observ.) Food consumed includes fish, frogs, reptiles, crustaceans, locusts, rodents, small mammals, lizards, insects and some carrion (Kahl 1970). We have seen birds bring whole vertebrates, such as frogs, fish and rats, back to the nest. The Lesser Adjutant is in fact known to 'eat any living thing, including chickens, not too big to swallow' (E.C.S. Baker 1929:330). Because of its preference for natural prey, the Lesser Adjutant is less attracted to the vicinity of human habitation than is the Greater Adjutant. While resting, it often sits on its intertarsal joints, with the tarsi stretched out in front. BREEDING In India and Burma, nests are placed from 12-30m up in tall forest trees, such as satian (Aistronia scholaris) or simul

Lesser Adjutant Stork

(Salmalia malabarica) (Kahl 1970). In Southeast Asia, it breeds in coastal mangroves on the shoreward edge of mangrove forests or along the banks of tidal rivers (H.C. Robinson and F.N. Ghasen, 1936:210). Some tree-nests in southern Burma were as high as 46m (Hopwood 1921, E.C.S. Baker 1935:448). The Lesser Adjutant Stork usually nests near waterbodies, although the water may dry before breeding is complete (P. Saikia and P.G. Bhattacharjee pers. comm. 1990). It is a semi-colonial or colonial breeder. It generally breeds in small colonies, sometimes with as many as 12-20 nests in a single large tree (E.C.S. Baker 1935:448). In the past, at least, it bred in mixed colonies with the Greater Adjutant (Smythies 1953:523). The nesting season of the Lesser Adjutant in Assam, India, can begin as early as July (P. Saikia and P.C. Bhattacharjee pers. comm. 1990). In southern Burma, where they formerly nested in large numbers in the company of the Greater Adjutant, the Lesser Adjutant was said to begin nesting somewhat later than its larger cousin (Blanford 1898:375). However, such was not the case in Assam, India, where we observed nests of both species in NovemberDecember 1967. Here 36 nests of the Lesser Adjutant contained young 35-75 days old, whereas a week later a single nest of the Greater Adjutant was found with young about 30-35 days old (Kahl 1972a). The nest is a large but rather flimsy stick structure 60150 cm in diameter and 10-120 cm deep, lined with smaller twigs and green leaves. Old nests from previous years are often used again by birds who add sticks to the old foundations in the new season. The general courtship behaviour of the Lesser Adjutant appears to be very similar to that of the Marabou Stork in Africa, with the exception of a few specific displays. The Balancing Posture is given by females during pairformation. This display in the Lesser Adjutant involves lifting the head in a scooping motion and contains a billclattering component lacking in both the Greater Adjutant and Marabou (Kahl 1972a). During the Up-Down display, given as a greeting between mates at the nest, both species of Adjutant clatter while the bill is held above the horizontal, whereas the Marabou clatters with the bill below the horizontal. We have observed few threat displays in this species. The Bill Snap, Pre-flight Snap and Erect Gape are similar to those of the Greater Adjutant and Marabou. In the Arching display, given in the presence of intruders, the Lesser Adjutant extends its neck and gives a hoarse wail. The Arching displays that we saw, were not as ritualized as in the Marabou (Kahl 1972a). Eggs are white, but become quickly soiled during incubation. A full clutch contains 2-4 eggs. The eggs of this species are quite similar to those of the Greater Adjutant, and in spite of the considerable size-difference of the two species, their eggs differ only slightly (E.C.S. Baker 1935:449). The incubation period is 28-30 days (P. Saikia pers. comm. 1990). Both adults feed the nestlings; young are also given water in the nest by their parents. Some observers have reported'

127

the Lesser Adjutant to be more shy than the Greater; however, this was not found to be the case at the nest (Kahl 1-970). Young often gave the Up-Down display to their parents before the Begging display (Kahl 1972a). In an odd situation, at the Dehiwela Zoo, Columbo, Sri Lanka, a female Lesser Adjutant hybridized with a male Painted Stork (Mycteria leucocephald). Both parents were freeflying and ranged over neighbouring paddyfields at will. The 'odd couple' hatched 4 broods between 1940 and 1942, and a number of viable offspring were produced. One could still be seen walking freely about the zoo as late as December 1967 (pers. obs. M.P.K.). The hybrid offspring in adulthood most resembled the Lesser Adjutant Stork in plumage-coloration and bill-size, and the Painted Stork in stance and bill shape (W.C.O. Hill 1943, Henry 1955, A.P. Gray 1958, pers. obs. M.P.K.). The same species combination (sexes unknown) also hybridized in 1974 at the Kuala Lumpur Zoo, Malaysia, and produced young (P. Olney pers. comm. 1974). TAXONOMY Of the three species in the genus Leptoptilos, the Lesser Adjutant is the least specialized in its morphology and behaviour. In some morphological characters, such as billshape and external structure of the gular pouch, the Lesser Adjutant shows more affinities with the Jabiru than do the other two Leptoptilos species (Kahl 1971e). CONSERVATION Numbers of this stork appear to be much reduced over most of its range. In Assam, India, habitat loss, both from the cutting of nest-trees and the draining of feeding areas, is a threat to the Lesser Adjutant. At least one ethnic group kills the birds so that the bill and skull can be used for medicinal purposes (Saikia and Bhattacharjee 1989a,b). This species is poached in Bangladesh and is sold in the market in Bangkok, Thailand (M.A.R. Khan 1987, Luthin 1987). Although there are about 200 birds in peninsular Malaysia, there is very little breeding there owing to the loss of suitable large mangrove trees and the robbing of young by humans. Birds are still widely but thinly distributed in Sabah, Borneo, especially on the remote northeastern and eastern coasts (D. Wells pers. comm. 1989). However, they are said to be decreasing rapidly there, probably owing to shooting (Gore 1968). As far as we can determine, the large numbers that once bred in southern Burma no longer exist. The healthiest population may be in southeastern Sumatra, but the birds there are at risk because of swamp-forest clearance, direct persecution and transmigration of people into the area (M. Silvius and W. Verheugt pers. comm. 1990). A few breeding sites are protected (in Assam, the Sundarbans, and Vietnam), but even there human disturbance and poaching continue. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 302.

Greater

Adjutant Stork Leptoptilos dubius (Gmelin) Ardea dubia Gmelin, 1789, Syst. Nat., 1, p. 246; based on 'Gigantic Crane' of Latham, 1785, General Synop. Birds, 3, p. 45: India Other names: Adjutant Stork, Larger Adjutant, Pouched Adjutant (English); Indische Marabu, Argala, ArgalaMarabu (German); Hargila [=bone-swallower], Garur, Peda dhauk (Hindi); Dusta (Dakhani), Dhenk (Mirshikars, Bihar); Hargila, Chaniari Dhauk (Bengali); Hargila, Bortukla (Assamese); Peenigala konga (Telugu); Don-zat (Burmese)

IDENTIFICATION The Greater Adjutant Stork is the most massive and ugliest of the Asian storks, standing 120-150 cm to the top of the head when erect. It often appears shorter, owing to its hunched stance. Its general colour is slaty-black, grey and white, with a nearly featherless head and neck, a massive wedge-shaped bill and a naked ruddy pinkish gular pouch, hanging from the front of the neck. It was named 'Adjutant' because it walks with the deliberate, measured gait of a military adjutant. The adult's upperparts are slate grey, with a pale band along the wing formed by the pale grey greater secondary coverts. The underparts are white. The head, neck and gular pouch are almost naked, with a few scattered dark brown hairlike feathers on the nape, neck, and sides of head. Dorsally there is a ruff of white feathers around the base of the neck, surrounding a red or orange air sac, c. 15 cm in diameter, that is only visible when inflated. The iris is white to yellowish white. The bill varies from pale yellowish to greenish, more reddish near the base. The bare skin of the head is a dull reddish brown, turning to brick-red on the hindneck and blackish on the forecrown. The gular pouch and neck are mostly yellow, with more pink, spotted with black, on the pouch. The legs and feet are actually a horny brown to dark grey, but often appear pale greyish white or chalky white because they are coated with uric acid from excreta. The male and female have the same plumage and soft-part colours. Sexual dimorphism in size is slight. However, males are somewhat larger than females, with heavier bills; the female's bill is a bit more decurved. At the start of breeding, the upperparts become a pale bluish grey, and the paler wing band turns a silvery grey. The undertail-coverts are white at their bases and dark smoky grey at their tips; they become luxuriant and fluffy at breeding time. The face and forehead become encrusted with what appears to be dried blood. The gular sac becomes a bright saffron-yellow (see colour plate VIII in Kahl 1971f). Nestlings are covered with a dense white down, except for the bare head and face. They are similar in appearance to nestlings of the Lesser Adjutant and the Marabou, except that Lesser Adjutant nestlings have a slightly thinner bill. Juveniles are similar to adults but have darker uppersides, without the pale band along the top of the wing, and with denser pale-brownish and grey feathering on the head and neck. Like adults, they also have smoky grey undertail-coverts.

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This is generally a silent bird, except around the nest. Bill-clattering is common during various displays. During displays at the nest a high-pitched squealing, slurring to a lower-pitched cow-like 'Mooing', is uttered; these calls are quite similar to those given by the Marabou Stork of Africa (Hume and Gates 1890:262, Kahl 1966a, 1972a). When the wings of adults are viewed from above in flight, there is a conspicuous pale grey band formed by the silvery grey greater secondary coverts. Despite heavy flapping on take-off, the bird flies strongly with noisy rhythmic flaps when once airborne and circles gracefully in thermals. 'On the wing .. . they are very majestic and a flight of these birds sailing round in great circles is very imposing' (E.C.S. Baker 1929:329). Unlike other storks, birds in the genus Leptoptilos fly with their necks partially retracted back onto their shoulders, similar to the herons. The flapping rate has been timed as 139/min (a = 8) (Kahl 1971b). The only other species within its range with which the Greater Adjutant could be confused is the Lesser Adjutant. The Greater is distinguished by its larger size, paler grey upperparts, pale grey band along the upperside of wing (which is conspicuous both at rest and in flight), ruff of unspotted white feathers around the dorsal base of neck, more massive bill (often with a slightly decurved culmen), dark (as opposed to pale-tan) forehead; dark sooty grey (not white) undertail-coverts, more sparsely feathered head and neck, c. 50 mm of red skin visible at the upper end of unfeathered tibia; and much larger gular pouch (20-35 cm long when inflated). All these differences are shown in plate VIII of Kahl (1971*). DISTRIBUTION AND POPULATION The Greater Adjutant is sparsely distributed in tropical Asia. Formerly it bred from northeastern India, Burma to Thailand, Kampuchea and southern Vietnam; it has now been extirpated or is uncommon in most of its former haunts (Deignan 1963:10, Wildash 1968:46, B.F. King and E.G. Dickinson 1975:52, S. Ali and S. D. Ripley 1968:106). Recent breeding is known only in Assam, India (Kahl

1970, Saikia and Bhattacharjee 1990b, A. Rahmani pers. comm. 1990). However, the species may still be nesting in the swamp forest of Minh Hai Province, Vietnam, although it has not been sighted in recent years (Luthin 1987). Earlier accounts of the range extending to the Greater Sunda Islands (Sumatra, Java, Borneo) (E.C.S. Baker 1929:328, J.L. Peters 1931:130, S. Ali and S. D. Ripley 1968:106) are almost certainly in error, being instead the Lesser Adjutant (Kuroda 1936:542). According to David Wells (pers. comm. 1989) there are 'no acceptable records [of Greater Adjutants] south of Thailand/ The Greater Adjutant is largely resident but also nomadic, depending on local conditions. During the period of their former abundance, Greater Adjutants came to India mainly during the rains and then returned to Burma to breed during the dry season. In years when desert locust swarms were heavy, the storks sometimes appeared in large numbers in areas of India (e.g. Rajasthan) where they usually were not common (Singh and Singh, 1960). Birds wandered irregularly, primarily during the rains, to Pakistan, Nepal, north-central India, and south (rarely) to Tamil Nadu ( — Madras Province). In the last few years, few, if any, Greater Adjutants have been reported outside Assam (none at Bharatpur since 1987; Bholo pers. comm. 1990), and this may suggest that nearly all the original migratory stock which formerly bred in Burma have been extirpated, and that the isolated populations in Assam are mainly sedentary as reported by P. Saikia and P.C. Bhattacharjee (pers. comm. 1990). Up through the 1930s very large numbers ('literally in hundreds of thousands', according to E.C.S. Baker 1929:328) gathered to breed in Burma, and it was thought that perhaps all of the Indian population of Greater Adjutants went there each year at the end of the rains (Hume and Oates 1890:261). The immense breeding colonies once reported near Shwaygheen, in the Pegu District of southern Burma (c. 150km northeast of Rangoon) are apparently gone today. Similarly the colony site in the Sunderbans of Bangladesh has been eliminated (M.A.R. Khan 1987). The species was formerly 'rather common' in Chiang Rai Province, northern Thailand (Deignan 1945:46), but it is much reduced in that country now (Lekagul 1968). Even as far back as the 1920s it was apparently much less common in Indochina than was the Lesser Adjutant (Delacour and Jabouillc 1931:89). Between 1987 and 1989 the largest number of Greater Adjutants seen in Assam, India, on a single day was 56 in Tezpur and 83 in Guwahati, and the total population there was estimated at 300 birds (Saikia and Bhattacharjee 1989a,b). In April-May 1989 Rahmani (1989c) counted 80 Greater Adjutants at 10 locations in Assam. More recently, in October 1989-January 1990, a special search was launched, and 75 active nests were located in Lower Assam (Saikia and Bhattacharjee 1990b). ECOLOGY Greater Adjutant Storks occur in various wetlands, such as jheels, marshes, paddyfields and lakes, particularly 'where

Greater Adjutant Stork

the water is drying and concentrating the fish life in shallow puddles' (S. Ali and S.D. Ripley 1968:106). Their use of wetland habitats is primarily during the breeding season; during the non-breeding period they often feed on human refuse or at abattoirs. Formerly the species was very common around human habitations, where it fed on carrion. During the 19th Century, 'large numbers of [Greater] Adjutants haunted the city of Calcutta . . . for the offal and refuse, and rendered valuable service as scavengers' (S. Ali and S.D. Ripley 1968:106). 'At that time . . . Adjutants could be seen on the highest points of almost every house in Calcutta' (E.C.S. Baker 1929:329). The usual feeding behaviour is for the bird to Walk looking for food, or alight at a food source and Stand to eat what is available. Food finding is primarily visual. It has been suggested (Jerdon 1864:730) that the inflated gular sac rriay help to supply extra oxygen 'during protracted acts of deglutition'. We doubt this. We feel that the gular sac functions primarily during social displays and, possibly, in thermoregulation. When feeding on swarms of the desert locust in Rajasthan, India, a stork was reported running a few steps and 'flapping its wings briskly to beat the bushes . . . on which large concentrations of the hoppers and fledglings of the desert locust were noted to be resting in the morning. Disturbed by the heavy and noisy flaps of the wings the insects jumped down from their roosting places, when they were immediately attacked with the wedge-shaped bill, picked up, and devoured' (Singh and Singh 1960). The food includes carrion, fish, frogs, reptiles (Vipera russelli, Uromastix hardwickii), crustaceans, locusts, and even maimed ducks (Panday 1974). It is likely that the diet changes during the breeding season to include many whole vertebrates for the essential calcium requirements of the fast-growing young. Outside the nesting season, this is a species well known for its appalling eating habits. According to E.C.S. Baker (1929:329) 'their tastes are omnivorous and there are few things an Adjutant will not swallow'. According to Akhtar (1947) a captive Greater Adjutant 'swallowed a shoe wellshod with iron; on another occasion it swallowed a goodsized fowl right down, with its wings and feathers.' In Assam, India, one bird was seen swallowing pieces of vertebral column from a dead buffalo that were over 30 cm long (Rao and Murlidharan 1989). A local Indian name, 'hargila', means 'bone-swallower'. The birds are fairly gregarious in foraging, associating with other Adjutants and with vultures and kites at animal carcasses and refuse dumps. When using wetland sites, they feed until the food supply is exhausted; then the group moves on to another site (P. Saikia and P.C. Bhattacharjee pers. comm. 1990). We have seen aggressive encounters of captive birds at feeding sites elicit Forward Displays and Upright Displays (Kahl 1972a). But even adult Adjutants are often dominated by the more aggressive vultures at feeding sites. The Greater Adjutant is often found in and around urban centres, where there are sources of food, and so seems not disturbed by close association with man.

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BREEDING The Greater Adjutant Stork nests, as well as roosts, in tall trees, often near villages and not necessarily in wetlands. In Burma it has nested on limestone pinnacles. A nest we saw in Assam, India, was placed in the crown of a satian tree (Aistronia scholaris) about 27-30 m from the ground; this nest was in the tallest tree within 500m (Kahl 1970). In late 1989, 75 nests were found concentrated in 6 colonies in Assam, India. They were placed at heights of 12-23 m in a variety of trees, such as kadam (Anthocephalus cadamba), simul (Bombax ceiba) and others. Nest trees were surrounded and protected by dense stands of bamboo (Saikia and Bhattacharjee 1990b). The nest itself is a large platform of coarse sticks, 1-2 m in diameter and up to 1 m deep. Pairs may breed singly, semi-colonially or colonially, sometimes with other birds, including the Lesser Adjutant Stork. At recent Assamese sites, some pairs nested singly, and the greatest number of nests in a single colony was 31 (Saikia and Bhattacharjee 1990b). Eggs are layed from September to January in Assam. In some areas, this species has been said to begin breeding about a month earlier than the Lesser Adjutant (Hume and Oates 1890:260). However, that was not the case with a single nest found at Kaziranga Wildlife Sanctuary, Assam, India, in late 1967. Nor was it the case in Assam in 1990, where some nests were still being built in November, after most Lesser Adjutants had fully grown young (pers. obs. J.A.H.). Most courtship displays described for the Greater Adjutant (Kahl 1972a) are similar to those shown by the Marabou Stork of Africa, which has been studied in more detail (Kahl 1966a). Adults are quite noisy at the nest, with many displays incorporating a loud clattering of the bill, plus a variety of squealing and 'Mooing' calls and loud wing noise. The Bill Snap is used most commonly when birds were being approached by another adult: the bird holds its body either erect or horizontal, with neck retracted, and gives a single, audible snap with the bill. It also gives a pop in the Preflight Snap. The Erect Gape is given without vocalizations. The Arching display is given in response to disturbance, accompanied by a repeated high-pitched wail. We did not see this display in the fully ritualized form, with neck puffed out and wings fully spread, as occurs in the Marabou Stork. The Balancing Posture is given, mainly by courting females, in submission; Swaying Twig-Grasping is given mainly by unmated males, as they are approached by a female. The Up-Down display, given as a 'greeting' between mates at the nest, terminates with the bill being clattered upward in this species, instead of the downward clattering seen in the Marabou. The clutch size is 2 or 3 eggs, rarely 4; at 75 Assamese nests in 1989, the average was 2.9 eggs (Saikia and Bhattacharjee 1990b). The shells are white, usually becoming badly soiled as incubation advances. Both sexes share in nest-building, incubation (E.C.S. Baker 1935:448) and feeding of the young. The incubation period is 28-30 days (P. Saikia pers. comm. 1990). At 75 Assamese nests in 1989-90,

132 Greater Adjutant Stork

c. 75% of the eggs laid produced fledged young, giving an average of 2.2 per nest (Saikia and Bhattacharjee 1990b). Young often give Up-Down displays to their parents, before the Begging display. They gape widely, throw their head to vertical, and then give several nasal squawks, before starting the repetitious movements of Begging. TAXONOMY In many aspects of behaviour and morphology, the Greater Adjutant closely resembles the Marabou Stork of Africa. Some authors (e.g. Meinertzhagen 1951) have even suggested that the two species are closely enough related to be considered conspecific. However, studies of both species at the nest (Kahl 1966a, 1972a) have shown that one of their principal courtship displays, the Up-Down, differs in significant ways, and this difference would probably be sufficient to provide reproductive isolation if the populations were sympatric. Therefore, we feel it is biologically sound to consider L. dubius and L. crumeniferus as valid, separate species. However, they do appear to be closely enough related to form a superspecies (Amadon 1966). CONSERVATION This is, perhaps, the most endangered of all the Asian storks. In the early years of the 20th Century, the Greater Adjutant was a very common bird in Indian cities during the non-breeding season, and they gathered in immense numbers to breed in southern Burma (Smythies 1953). Today the bird is uncommon to rare in many of its former haunts, and the Burmese breeding grounds are said to be deserted (C. Luthin pers. comm. 1989). Investigations to discover currently active breeding sites and measures to protect them are urgently needed. If the decrease of the past 50 years continues at the same rate, the species might cease to exist in another few years. It is puzzling that the Marabou Stork seems to be increasing in Africa, whereas the apparently ecologically similar Greater Adjutant is decreasing in Asia. In India the species has rarely been persecuted directly, for, being a carrioneater, it is considered by Hindus to be unclean and therefore inedible. It is believed that many of the trees in the traditional Burmese nesting areas have been cut and the stork is probably eliminated from that country (J. Sayer pers.

comm. 1989). The Greater Adjutant colony site in the Sunderbans, Bangladesh, has been lost because of logging operations going back into the 19th Century (M.A.R. Khan 1987); and the stork has been eliminated from elsewhere in Bangladesh by hunting, except for one recent sighting in December 1988 (A. Rahmani pers. comm. 1990). The discovery, in late 1989, of 75 breeding pairs of Greater Adjutants in Assam, India, is encouraging (Saikia and Bhattacharjee 1990b). A major problem is that most of the nest-trees are on private land, and the owners often cut them down for their own use. Wetlands have been drained for agriculture and remaining sites have been filling in, due to the enormous growth of water hyacinths and grasses. Industrialization is also encroaching on wetlands. Other possible reasons for decline in India are pesticide poisoning and increased competition from larger populations of vultures (A. Rahmani pers. comm. 1990). Furthermore, adults and nestlings are sometimes killed for food by members of a nomadic (non-Hindu) tribe that visits the nesting areas during the winter months, when the birds are breeding (Saikia and Bhattacharjee 1990b). The problems of the Greater Adjutant Stork are further complicated by the fact that the local human population is completely unaware of the status and need for protection of this species. None the less, the Brahmaputra Valley of Assam is the only site where conservation measures for this species would have a chance of success. P. Saikia and P.C. Bhattacharjee (1990b, pers. comm. 1990), who assessed nesting locations, population status, and habitat use, provided a conservation strategy. This includes increasing the conservation awareness of villagers, protection of nest-sites, planting of roosting and nesting trees, and providing financial help to villages where nests are located. Known nesttrees could be purchased as 'national treasures' and strict laws against any harm to the birds should be enforced. We strongly urge that the Indian Government take immediate steps to ensure the safety of this rare species within their borders.

Note'. Body measurements, egg measurements and nesting season date for this species can be found in the Appendix on page 303.

Marabou Stork Leptoptilos crumeniferus (Lesson) Ciconia crumenifera Lesson, 1831, Traite Ornith., livr. 8, p. 585: Senegal Other names: Marabout d'Afrique (French); Afrika-Marabu (German); Abu Sen (Arabic); Bambu (Somali); Jimmu (Mandingo); Borintunke (Hausa); Mmakaitsimeletsa (North Sotho); Svorenyama (Shona); Qandlopfu (Tsonga)

IDENTIFICATION This massive extremely ugly stork stands about 120cm tall, when erect rather than in its more usual hunched posture, and has an impressive wing-span of up to about 2.9 m. It is basically slaty black, grey and white, with a nearly featherless head and neck and a massive wedge-shaped bill. A distinctive feature is its gular pouch of nearly naked ruddy pinkish to pale magenta skin. When inflated, the air pouch hangs down 25-35 cm from the front of the neck. To some eyes, the Marabou seems one of the ugliest birds in the world. It has been referred to as a 'gaunt-looking object at the best of times', 'the world's ugliest bird', 'entirely lacking any charm', 'stiffjointed and dried-up as though it had lived for a thousand years', or 'a dirty old man with a skin disease of the scalp'. However, biologists who know the Marabou think otherwise and seldom speak of it in such derogatory terms; we tend to notice the more endearing qualities in the bird. Referring to the lovely white undertail-coverts, grown in the breeding season and formerly used in high fashion, F J.Jackson (1938:79) stated: 'It is almost inconceivable that such delicate and beautiful plumes, when adorning a fan or some finery dear to a woman's heart, are the product of this huge, gaunt and repulsive looking scavenger, with a bald, scabby head and a body alive with vermin.' The upperparts are dark slate-grey, the underparts white. The flight feathers are glossed with green, whereas the back and mantle are glossed with blue. Head, neck and gular pouch are nearly naked, with a few scattered dark reddish brown, hair-like feathers on the nape, neck, and sides of head. Dorsally, at the base of the neck, there is a ruff of white feathers, surrounding a deep crimson or orange-red air-sac c. 10-15 cm in diameter and 5.0-7.5 cm high; this structure varies greatly among individuals and is visible only when inflated and exposed by separation of the feathers. The iris is sepia-brown to greyish brown. The bill is primarily horn-coloured to greenish yellow, mottled with black especially near the base. The forehead, lores and bare skin of the head are mainly orange-red with numerous small black pigmented spots, these increasing with age and most prevalent around the ear-opening and at the base of the bill. The gular pouch and neck are pinkish or magenta, spotted with black especially on the pouch. The legs and feet are nearly black but usually heavily coated with white excrement. There is a narrow band of smooth reddish skin at the extreme upper end of the naked tibia, just below the beginning of the feathers, visible only when the bird bends over or raises its wing. When birds are together, the sexes are usually distinguishable by size. Males are larger than females, with longer and heavier bills. Over 50 adults were measured in Uganda, and Pomeroy (1977b) found that 'wherever the sum of wing-length and tail-length exceeded 100 cm, the bird was a male. At the start of the breeding season upperparts become a paler bluish-grey, with a ladder-like band of white marks along the upper surface of the wing, formed by the white edges of the greater secondary coverts (Kahl 1987b). Luxuriant, elongated and fluffy undertail-coverts (the 'Marabou down' of the former millinery trade) are pure white. The face and forehead become blackish red, encrusted with

Marabou Stork

what appears to be dried blood. Colours of the bare skin on the head, neck and gular pouch intensify, with the back of the neck becoming a bright pale blue. Iris colour, especially in some females, turns a lighter brownish grey for several days just prior to egg-laying (Kahl 1966a). The pale bluish grey of the back is due to a 'waxy bloom' of keratin on the surface of the feathers (Pomeroy 1977b, Jacob and Pomeroy 1979); this gradually wears oft'and the adults revert to the darker slate-grey backs, usually long before the young have fledged. Hatchlings have wrinkled, pinkish skin and are sparsely feathered with a pale grey down; newly hatched young weigh about 70 g. Older nestlings of 8-35 days are covered with a denser white down, except for the bare head, foreneck and face; they are similar in appearance to nestlings of the Greater and Lesser Adjutant Storks. Blackish contour feathers appear on the back and wings at 35-50 days. At 5165 days, nestlings become predominantly black on the dorsal side, and after 66 days they are fully feathered and change little in appearance thereafter (Kahl 1966b). Juveniles are similar to adults. However, their uppersides are dark slaty-black, with a moderate bluish gloss mixed with brownish; they are also without the white hatchmarks (secondary coverts) along the top of the wing. Denser pale brownish or black filamentous feathers, mixed with pale grey woolly feathers, grow on the head and neck. The softparts are less brightly coloured, and the gular pouch is less developed (or less commonly fully inflated). Bill-length increases slowly with age, requiring 1-2 years to reach full length (D. Pomeroy pers. comm. 1989). For a more complete description of the development of plumages, and their sequence, the reader is referred to Pomeroy (1977b). The Marabou is generally a silent bird, except around the nest and when roosting. During social displays birds clatter

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their bills loudly and utter a variety of high-pitched squeals and yelps, slurring to a lower-pitched, grunting cowlike 'Moo'; these vocalizations are quite similar to those given by the Greater Adjutant Stork of Asia (Kahl 1966a, 1972a). Their wings make loud 'Woofing5 sounds, especially during landing and take-off during social encounters with other Marabous. Heavy on take-off, the Marabous flies strongly with noisy rhythmic flaps once airborne. Groups soar on fixed wings in thermals, sometimes climbing to such high altitudes that they are nearly invisible from the ground. A group was observed, from a light aircraft, soaring at 3000-4000 m above Uganda with their gular pouches fully inflated (Akester et al. 1973). Colin Pennycuick (1972) provided an interesting description of soaring behaviour in Marabous, as observed from a motor-glider over the Serengeti Plains, Tanzania. Unlike other members of the family, storks in the genus Leptoptilos fly with their necks partially retracted back onto their shoulders, similar to the herons, except that the bill and legs are angled slightly below the horizontal. We timed the flapping rate as 143/min (n = 8) (Kahl 1971b). On the ground or at close quarters, the Marabou cannot be mistaken for any other bird within its range. When soaring on high, it can be distinguished from vultures and the Great White Pelican (Pelecanus onocrotalus) by the long legs trailing out behind, and from the rare Shoebill by the thinner bill.

DISTRIBUTION AND POPULATION The Marabou Stork is resident in tropical Africa. It occurs from Senegal across to Ethiopia and Somalia, south to northern South Africa (northeastern Transvaal, Zululand), northern Botswana and northeastern Namibia. The Marabou undertakes short, intratropical migrations according to rainfall conditions. It often leaves areas during the rains, to appear again the following dry season (Chapin 1932a:465, 467, Pomeroy 1973). In Uganda, migrant Marabous seem to move mainly in a north-south direction (Pomeroy 1977b). They also wander erratically to areas of temporarily abundant prey. The species is a rare vagrant south of its breeding range, and is an accidental visitor in Israel (3 records; Paz 1987:34). This species is common to abundant in most parts of its range. Apparently the Marabou has increased in recent years throughout its range, because of its close association with humans. The total Uganda population was estimated at 4000-5000 in 1970-72 (Pomeroy 1973) and had increased significantly by 1975-76 (Pomeroy 1977c). The total Kenyan population has been estimated at 1000-2000 (Pomeroy 1986). The age-structure of the population in Uganda (based on plumages of 4566 birds) is heavily dominated by adults: 13% juveniles, 15% subadults 3^ years old, and 71% adults more than 4 years old. The maximum calculated age exceeds 25years (Pomeroy 1977b). The estimated annual mortality rate of adults is 8% per year (D. Pomeroy pers.

136 Marabou Stork

comm. 1990). The species has lived 31 years in captivity (Flower 1938). ECOLOGY The Marabou is found mainly in open country, often in the vicinity of large lakes or rivers. It does not occur in the densely forested regions of Africa, except where forests are bisected by large rivers. It seems to prefer arid or semi-arid areas, but often within flying range of an aquatic habitat. Often it is associated with vultures and other scavengers in 'big game country5, where it feeds on carrion provided by the remains of kills left by large predators. It is common, sometimes abundant, around human habitations, garbage dumps, abattoirs and fish-camps, where refuse is available. It also uses natural wetlands, especially during the nesting season, and favours drying pools where fish are concentrated. Feeding behaviour is surprisingly reserved. When feeding at carrion with vultures and eagles, the Marabou is usually a rather shy bird. It hangs around the outskirts of the group, dancing in occasionally to steal a morsel from the smaller, but more aggressive, scavengers. In such situations Marabous play more the part of'pirate', allowing the vultures— with their stronger, hooked bills—to tear off pieces of meat from a carcass and then attempting to steal it away (Houston 1975). At rubbish dumps and fishing villages, Marabous lose all fear of humans and can be seen within a meter or so of people working there, waiting patiently for a scrap to be tossed their way. Especially during the breeding season, but occasionally at other times as well, Marabous forage for living prey. When termites are swarming, the storks often stand at their mounds and consume large quantities of these fat-bodied insects as they emerge on their mating flights. Aquatic prey is often hunted using Groping, the tactolocation technique characteristic of the wood storks, Mycteria. The partially open bill is immersed vertically in shallow water and moved slowly about as the bird wades forward. When a prey item is contacted, the bill snaps rapidly shut, and the prey is quickly swallowed with a backward toss of the head. At other times, when visibility is good, such prey is located visually and Grabbed in the tip of the bill. Marabous often feed together. In feeding groups of Marabous, dominant individuals usually stalk about with their gular pouches fully inflated, and subordinant birds give way to them. Large numbers appear, as if from nowhere, at infestations of locusts or army worm caterpillars and at grass-fires, where they consume insects and small vertebrates flushed or injured by the flames. Here they often feed in the company of migrant White Storks and Abdim's Storks. One Marabou collected while feeding on a swarm of desert locusts (Schistocerca gregaria) had a total of 993 locusts in its alimentary tract (K.D. Smith and G.B. Popov 1953). The food of the Marabou is almost anything organic. According to Brown et al. (1982:188), it will eat Virtually any animal matter from termites to dead elephant5. Over

200 Marabous have been recorded at one elephant carcass in Kruger National Park, South Africa (Kemp 1974:15). Carrion is an important staple in the diet, especially among non-breeding Marabous. 'Wild5 carrion, such as the remains of lion kills, has been largely replaced in many areas by 'man-made5 carrion from garbage, fish scraps and abattoir refuse around settlements. And today in many urban areas, vegetable matter forms a large part of the Marabou's diet (D. Pomeroy pers. comm. 1990). Whole vertebrates (primarily fish, frogs and rodents) are favoured by breeding adults during the period that their nestlings are growing most rapidly, probably owing to the increased need for calcium by the young. Hand-reared nestlings did not survive when the supply of whole vertebrate food failed (Kahl 1966b). Marabous relish the eggs and young of crocodiles, which they often consume as they are emerging from their shells (Pitman 1957). Birds also sometimes fall prey to Marabous. Wildlife film-makers Simon Trevor and Des Bartlett have photographed them catching and eating nestlings of Quelea quelea in Kenya's Tsavo National Park (see also Disney and Marshall 1956, Leuthold and Leuthold 1972, and Kemp 1974) and adult doves in Namibia5s Etosha National Park. Several authors tell of predation on larger birds, such as young and adult flamingos in East Africa (Brown et al. 1982:188), cormorant nestlings in South Africa (Milstein 1975) and Great White Pelican chicks in Nigeria (Dunger 1965). With its massive bill and elastic throat, a Marabou is able to swallow pieces of meat weighing as much as 600 g. FJ. Jackson (1938:77) reported one 'struggling with the entire windpipe of a bull buffalo that was halfway down its throat5. A worker at the Kitale abattoir (Kenya), reported that a Marabou stole and swallowed a bloody butcher-knife that was laid aside for a few moments; days later the knife, digested spotlessly clean of all blood and residue, was found nearby in the grass, where it had been regurgitated as 'indigestible5 by the stork (Kahl 1988b). It is estimated that an adult Marabou, weighing 5-8 kg, requires about 0.7-1.Okg of food per day to survive (Kahl 1966b, Pomeroy 1973). Often all the birds in an area gather in the late afternoon at traditional communal roosts, which may be used for many years if not disturbed. One such roost, near Nairobi, Kenya, far from any known breeding colony, contained up to 1000 individuals (D. Pomeroy pers. comm. 1990). The next morning they often remain at the roost for several hours after sunrise, until thermal air-currents develop on which they can soar, effortlessly, to distant feeding grounds. Marabous commonly sit for long periods on their tarsi and often sun, either sitting or standing, with wings fully spread. Opening the wings to the sun in this manner is a very contagious behaviour pattern, and many birds in a group may do so simultaneously. The inflatable gular pouch is primarily used as a signal of dominance in social displays. However, it and other subcutaneous air-sacs may also serve other, secondary physiological functions, such as thermoregulation. Marabous are sometimes kept as pets in African villages, and they become quite tame. However, their massive bill is

Marabou Stork 137

a formidable weapon, and they have been reported to have killed children who were trying to catch them (Mackay 1950). In the Sudan, they are occasionally used as 'animated feeding perches' by the Carmine Bee-eater (Merops nubicus) (Cunningham-Van Someren 1970). BREEDING Marabou Storks nest in colonies, often spread over a rather wide area, of a few pairs to a several hundred pairs, most commonly 20-60 pairs. They often build in association with Pink-backed Pelicans (Pelecanus rufescens) or, less commonly, with other waterbirds. Sites are frequently traditional and are used for many years barring disturbance. Consistent requirements for a successful breeding site seem to be safety from disturbance, plus reasonable proximity (<50-60 km) to a good food-source (ideally both carrion and live vertebrate prey) for the young. The largest colonies in eastern Africa were near to extensive swamps (Pomeroy 1978a). Nests are usually placed 3-40 m up in trees; they occasionally nest on cliff-ledges, as is the case in northeastern Uganda, southern Sudan and southern Tanzania. Tree-nests are often in euphorbia (Euphorbia dawei], mvule (Chlorophora excelsa), baobab (Adansonia digitata) or acacia. Such trees are sometimes in the midst of villages, especially in West Africa (Bannerman 1930:107) or in larger cities, as in Kampala, Uganda (Kasoma and Pomeroy 1987). The nest platform is a rough structure of coarse sticks, lined with smaller twigs and green leaves. Foundations from previous years are often used again but usually require considerable renovation, owing to weathering and stickpirating by neighbours. A finished platform is approximately 1 m in diameter and 20-30 cm thick. Occasional sticks are added to the structure throughout the nesting season, until the young have fledged. Because nestling growth is so slow, the nesting cycle is protracted: from egg-laying to fledging takes about 165 days (Kahl 1966b). In the northern and southern tropics, breeding usually begins in the dry season and young fledge during the next wet season. In the equatorial zone, where there are two wet/dry cycles per year, seasonality is less clear-cut (Pomeroy 1978c); breeding cannot be completed in one short dry season, and young may fledge in the next dry season. Pair-formation and ritualized displays in the Marabou were described in detail in Kahl (1966a). Males establish themselves on nest-sites and wait to be approached by courting females. Their large gular pouches are usually fully inflated at this time, indicating a high level of hostility and dominance in the males. All trespassers are treated with hostility at first; other males generally fight back, whereas females react passively and with submissive displays, such as the Balancing Posture. Eventually a persistent female is accepted by the male. After pairing the female also inflates her gular pouch, and both members maintain them fully inflated much of the time, at least up until the time of egg-laying. This seems to signal a 'feeling of dominance', at the nest-site, in relation

to all other Marabous in the vicinity. When the pair is firmly established, the male then periodically leaves her alone on the nest-site while he flies out to collect sticks. A mutual Up-Down is performed by the pair whenever one member returns from an absence. Frequent copulations follow, interspersed with nest-building and mutual display activities. During the early stages of nest-construction, the male collects most of the material and delivers it to the female on the nest, who does most of the building. Later in the breeding cycle, a majority of the stick-collecting is done by the female. Sticks continue to be brought to the nest by both parents until the young are ready to fledge. Clutch size ranges from 1-4, rarely 5, and usually 2-3. The mean of 108 clutches was 2.4 eggs. Eggs are laid at intervals of 1-3 days (Brown et al. 1982:190). The shells are dull chalky white when first laid but are soon stained. Both sexes share in incubation, feeding and guarding of the young. Incubation begins at or soon after the laying of the first egg. Both sexes incubate for about 84% of the daylight hours: males 48% of the time, females 36% of the time. Males also incubate more than females at night. The incubation period is 29-31 days (average = 30.3 days). The hatching process takes about 24 h. Eggs usually hatch at intervals of 2-3 days; extremes observed are 1-7 days (Kahl 1966b, Pomeroy 1977b). Parents take turns brooding the young much of the time for first 10 days. Thereafter, brooding is reduced, except during periods of rain or cold. Nestlings are shaded by a standing parent, with wings either open or closed, when the sun is hot. Young are almost never left alone on the nest until after 25-30 days of age. Even then a parent is present at or near the nest most of the time. Almost full-grown nestlings are sometimes left alone while both their parents forage. They are usually able to defend the nest against intruding Marabous that sometimes attempt to enter it. Nestlings solicit for food by dropping to their tarsi, with partially open wings, cocked tail and up/down nodding of head in the Begging display. Begging calls of newly hatched young are a high-pitched chitter. This call gradually deepens, as the young grow older, until it is a hoarse, nasal braying. Both parents feed the young, by regurgitation onto the floor of the nest. We found that males fed more (55%) than did females (45%) in 386 recorded feedings. Most feedings occurred between lO.OOh and 15.00h, coinciding with the daily thermal development that permitted the parent to fly to and from distant feeding grounds. We estimated food consumption of young, from captives. It increases from <100g/day at age 1-10 days, to 500600 g/day at age 60+ days. Nestlings receive a large proportion of whole-vertebrate food, such as fish, probably because of their high calcium requirements during rapid growth. Of 421 feedings observed at the Kitale colony (Kenya), 29% consisted of whole-vertebrate prey (Kahl 1966b). Larger nestlings are also given water, by regurgitation, by their parents. This watering may be related to the thermoregulatory habit of excreting a dilute urine onto the bare

138 Marabou Stork

skin of their legs for evaporative cooling (Kahl 1963b). Without a supplementary water supply, from the parents, this behaviour might lead to dehydration of the young. Nestlings can stand and flap their wings at 17 days old, hover over nest at 65 days, and make short flights within the nest-tree at 95 days. The first recorded flight out of the tree was at 98 days, although more commonly first flights were at 110-115 days. Nestling growth is very slow, and young remain dependent on their parents until an age of 130 days or more. Such a protracted growth pattern is perhaps an adaptation to reduce the nestlings' daily food demands to a level the parents can meet (Kahl 1966b, Pomeroy 1977b). Marabous are probably 6 or 7 years old when they first breed (Pomeroy 1977b). Some adult Marabous may not attempt to breed in any given year and among those that do breed, success is fairly low. Depending on conditions, 5295% of eggs hatch. In nests where 3 eggs hatch, it is rare for the last-hatched chick to survive, although it occasionally does (Pomeroy 1979). Only 36% of eggs in the Kitale colony (Kenya) resulted in fledged young (Kahl 1966b). About 0.9 young per nest fledged in Uganda colonies (Brown et al. 1982:190). In recent years (1984-89), the Marabou has increased within the city of Kampala, Uganda, and breeding success has risen to 1.18-1.75 young/nest (D. Pomeroy pers. comm. 1990). Pomeroy (1978a) found that nesting success was greatest in the medium sized colonies (i.e. 40-60 nests). The reasons for this are unknown. It may be that medium-sized colonies had a larger proportion of mature, experienced adults and, therefore, had a higher success rate. According to the calcu-

lations of Pomeroy (1978b), only about one egg in three [gives] rise to fledglings .. . less than a quarter of these survive to join the breeding population. Hence any extra recruit to the breeding population would require 12 extra eggs [to be laid].

TAXONOMY The relationships of the Marabou to the Greater Adjutant Stork have been a matter of some discussion, owing to their similarity. However, we have found marked differences in their behaviour, sufficient to continue to recognize them as separate species. We discuss this further in our account of the Greater Adjutant Stork.

CONSERVATION At the present time, the Marabou Stork appears to be doing very well indeed in most parts of its range. It is one of those unique species that appears to benefit greatly from its association with man and his urban environment. It is interesting and puzzling that this species is increasing in Africa, when its close relative, the Greater Adjutant Stork, is decreasing in Asia.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 304.

Shoebill Balaeniceps rex Gould Balaeniceps rex Gould, 1850, Athenaeum, no. 1207, p. 1315: Upper White Nile (=Sudan) Other names: Whaleheaded Stork, Boat-bill, Bog-bird [Sudan], Lesser lechwe-eater [Zambia] (English); Schuhschnabel, Konig Walkopf [='King Whalehead'] (German); Bec-en-sabot du Nil (French); Becco a scarpa [Italian]; Schoenbekooievaar (Dutch); Abu markub [='Father of the shoe'] (Arabic); Adungwek or Adumwueg (Dinka); Bany (Nuer); Okomo warjok (Shilluk); Bulue (Luganda); Mkumabimo or Nkubabimu (Bakenya/L. Kyoga, Ugandan); Ekurududu (Teso/Ugandan); Chifumpafumpa (Tabwa/Zambia); Fumpa-fumpa (Balamba/Katanga, Zaire); Motuta or Mututa (Lualaba/Katanga, Zaire)

IDENTIFICATION Slow moving, deliberate, morose, often statue-like, the Shoebill is a large grey bird, standing 110140 cm tall. It is immediately recognizable by its huge bulbous bill, reminiscent of a Dutch wooden clog. The adult's head is dark grey, with a slightly darker blue-grey crown, while its back and mantle are ashy grey with a slight greenish gloss. The lower neck feathers are lanceolate, with black shafts. The primary wing feathers are dark grey, although black near the tips. The secondaries have a slight greenish gloss. The underparts are paler grey. The enormous, prominently hooked bill is yellowish with dark irregular patches. It is a formidable tool and weapon, with cutting edges like a turtle's jaws (FJ. Jackson 1938:65). A dark keel on the upper mandible terminates in the hook, which covers the tip of the lower mandible when the bill is closed. The eye is a pallid yellowish or greyish white, and the legs blackish to horn-coloured. The toes are extremely long (middle toe = 16.8-18.5 cm long; W. Fischer 1970:13) and completely divided (i.e. not partially webbed, as in the true storks); the claw of the hind toe is larger than those on the fore-toes (Reichenow 1900-01:357). A small crest or tuft on the back of the head can be erected during alarm or excitement (Owen 1958, M. Burton and C.W. Benson 1961, C.W.Benson 1961). The male is noticeably larger than the female and has a longer bill (see Abb. 14 and 15 in W. Fischer 1970:20, and Figs. 4 and 5 in L. Buxton et al. 1978). The breeding plumage is not known to differ noticeably from the regular adult plumage. The eye may become more yellow at sexual maturity. Newly-hatched young are covered with a silvery grey, silky down. The bill, though not yet bulbous, is markedly hooked and the gape is wide; the head is large relative to the body. Exaggerated swelling of the bill becomes noticeable at 23 days and is well-developed by 43 days (L. Buxton et al. 1978). Later, 'nestlings are covered with grey down of a fulvous tinge' (A.L. Butler 1905). The iris colour of immatures is said to change from milky blue-grey, to pearl-grey (in the second year), to bluish white (at about 5 years), to pale yellow (perhaps at sexual maturity) (W. Fischer 1970:10). A chronology of the development of the nestling is given by L. Buxton et al. (1978): Days 1-35, entirely or mainly downy, unable to stand, requires constant brooding or shading; days 35-75, developing contour feathers, stands unsteadily or sits on tarsi, requires little brooding by day; days 7595, largely feathered, left alone by parents, seldom shaded, stands steadily and exercises wings, begins to leave nest on foot, soon can fly. When fully feathered, juveniles are a darker grey than the adults with a slight brownish tinge. The bill develops its bulbous shape but is not quite as exaggerated as the adult birds. In one Zambian

Shoebill

141

sharp and needle-like, especially from below, (pers. obs. M.P.K.).

juvenile, it was a pinkish colour with ill-defined blackish markings (L. Buxton et al. 1978). Plumage parts that are slate-grey in adults are grey-brown in juveniles (Reichenow 1900-01:357, Schouteden 1912). Captives assume adult plumage at 3^ years (Moller in Collar and Stuart 1985:66). During displays at the nest, the bill is clattered and a high pitched whine or 'Moo' is uttered (W. Fischer 1970:45, 58, L. Buxton et al. 1978). A soaring bird uttered a 'rather loud, hoarse croak' as it dived at a soaring Marabou Stork nearby (pers. obs. M.P.K.). Also reported are a hoarse but feeble cry (Dekeyser 1962), a raucous and guttural cry (Dechambre 1936) and an 'occasional loud cry while soaring' (Mackworth-Praed and Grant 1970:51). Both adults and young clatter their bills (Gilliard 1958). Recordings of the voice are available (Boswall and North 1967). Begging calls of nestlings resemble human hiccups (L. Buxton et al. 1978). Near-vertical take-offs and landings are possible, because of the large, broad wings (Guillet 1979). When disturbed, this stork is able to rise more calmly than any of the larger herons, without noise, and flap slowly away (FJ. Jackson 1938:65). The neck is retracted in flight (Schouteden 1912, Bouet 1955a). It does not readily take flight and is unlikely to travel more than 400-500 m when flushed (Owen 1958). It soars on thermals (Berg 1940). On 8 December 1963, a single Shoebill was watched soaring high up with several Marabou Storks just soutwest of Kampala, Uganda. The wings were held perfectly flat (i.e. no dihedral) and the primaries were widely slotted; the extremely long toes were conspicuous in flight. The flap rate has been timed once; at 150 flaps/min (pers obs. M.P.K.) it is slower than all but the largest of storks (Kahl 1971b). The Shoebill is unmistakeable if seen well. In flight or when soaring, it might be mistaken for a Marabou Stork or possibly for a Goliath Heron, but the Shoebill has a shorter, blunt bill, whereas the heron's and Marabou's bills look

DISTRIBUTION AND POPULATION The Shoebill is widely but locally distributed in freshwater swamps of central tropical Africa, from southern Sudan (north to about 9°N) through parts of eastern Zaire, Rwanda, Uganda and western Tanzania to northeastern Zambia. The distribution of this species generally coincides with that of papyrus (Dekeyser 1962), and it also largely coincides with that of the lungfish (Protopterus aethiopicus) (Greenwood 1958:14). The Shoebill is most numerous in the White Nile Sudd and adjacent areas of the southern Sudan; significant populations also occur in the wetlands of Uganda and western Tanzania and the Bangweulu Swamp of northeastern Zambia. In Uganda, it is widespread—but with a fragmented distribution in suitable habitat—eastward to Teso and Busoga, and south to the Kagera river. Records of its occurrence in the extreme west of Kenya (A. Koenig 1910, Britton 1978, 1980) are apparently erroneous (I.S.C. Parker 1984a). It has been found in Tanzania at several sites, all in the west [Moyowosi Swamp, Lake Mujungwizi, Lake Mujunju, Kagera Swamp, Nzilindagaza, Malagarasi Swamp, and Lake Rukwa (rarely)]. In Zambia, we have isolated records from the Itawa Swamp, near Ndola, and the Kasanka National Park; but in the west of the Northern Province and in the Luapula Province, it is more widespread, in particular in the Bangweulu Swamps. It is also reported uncommonly from Fungwa Swamp, around Mweru Marsh, Lake Kako, and near the mouth of the Luapula River. In Rwanda, it occurs in the Akagera depression. In Zaire, it is found in the western Rift Valley (Lake Edward) and mainly in the southeast, in Katanga Province. Isolated birds are reported or rumoured from the Central African Republic (Blancou 1939), northern Cameroon (Lisle 1956), southwestern Ethiopia (Duckworth 1974), Malawi (K. Brouwer pers. comm. 1989); and strays have been reported as far southwest as the Okavango Delta, Botswana (N.J.C. Mathews 1979, pers. comm. 1989). Long ago it was rumoured on the headwaters of the Upper Congo River, in Angola (H. Johnston 1902). These and other distribution records are discussed in Collar and Stuart (1985:59-63). Drawings of the Shoebill dating from approximately 3500 BC, have been found in Egyptian tombs (A. Koenig 1910:525; Schouteden 1912). Fossil relatives of the Shoebill have been found from the Oligocene of northern Egypt (D.T. Rasmussen et al. 1987) and the late Miocene of Tunisia (Rich 1972). The Shoebill appears to be non-migratory, remaining in or near its home range throughout the year (Kasoma and Pomeroy 1987), so long as foraging conditions are appropriate. Sudanese birds make limited seasonal movements, in response to changes in food availability, habitat availability and disturbance (Guillet 1978b, and pers. comm. 1990). It is generally thought that this is a rare species, its

142 Shoebill

population being limited mainly by the distribution of its habitat. It may be that isolated pockets of birds or of individual pairs exist in suitable areas throughout its range. Even in areas where it is relatively numerous, estimates of numbers can vary considerably. As I.S.C. Parker (1984b) stated, an impression of scarcity probably derives from the general inaccessibility of the species' habitat. In the few areas where it can be searched from a boat or from a vehicle, the habitat is usually a thin ribbon along the water's edge. In such circumstances the bird's scarcity is not surprising. Where the habitat is extensive, in the shallows between dry land and papyrus banks, it is difficult to search from either boats or wheeled vehicles. In order to find it 'its home must be entered; cruising to and fro outside ... would never reveal... that it may be ... just beyond the barrier of papyrus'(F.J.Jackson 1938:66). In the Jonglei area of the southern Sudan, 3000 were counted in 1979 along the White Nile system (A. Guillet in Collar and Stuart 1985:63), and Jonathan Kingdon (in Collar and Stuart 1985:63) estimates that the whole area of the Sudd may contain as many as 10000 birds. In Rwanda, numbers have fallen from 30-50 in the early 1970s to between 15-25 (Vande Weghe 1981), and in Uganda the population has been estimated 400-600 birds (Collar and Stuart 1985:64). However, Kasoma and Pomeroy (1987) suggest that this estimate may be too low, since the birds are difficult to see, and Moller (1978) found 26 birds in 20 km2 of swamp in Uganda. I.S.C. Parker (1984b) estimated about 0.64 birds/km2 in a swamp in western Tanzania, surveyed from the air. In a portion of the Bangweulu Swamps of northeastern Zambia, G.W. Howard and D.R. Aspinwall (1984) estimated that 232 (c. 0.145 birds/km2) Shoebills were present in October 1983. A reasonable estimate of the total number of Shoebills would seem to be between 11000-15000 birds over its whole range (Anon. 1985b, A. Guillet pers. comm. 1989); this is nearly 10 times greater than previous estimates (Guillet 1978a, Brown et al. 1982:191). ECOLOGY Shoebills are characteristically birds of extensive, dense freshwater marshes, especially flood-plain interspersed with undisturbed papyrus (Cyperuspapyrus) and reedbeds (Phragmites, Typha, etc.) having floating vegetation on which to stand; they generally avoid areas in which the plants are taller than the bird's back or are too dense to penetrate easily. They seem to prefer areas of poorly oxygenated water, where fish and other aquatic organisms are forced to the surface to breathe (Guillet 1987:222). Because of this, Vande Weghe (1981) found that, in the Akagera National Park and probably elsewhere, the Shoebill avoids pure papyrus swamps, preferring mixed swamps of Cyperus papyrus (papyrus), Miscanthidium and floating vegetation consisting of ferns and grasses with clumps of Cladium and Typha. It is also found in marshes dominated by Pistia stratiotes, Typha (cattail), Miscanthidium., Vossia cuspidata, Cyperus latifolius and

Echinochloa pyramidalis. When conditions are favourable, birds have been seen on flooded grassland and open floodplain (Collar and Stuart 1985:64). In the southern Sudan, the Shoebill has recently expanded its feeding habitat to include rice fields and other flooded plantations (A. Guillet pers. comm. 1990). This species especially prefers open parts of the swamp (Owen 1958), possibly due to the ease of taking flight under such conditions, and also because in dense papyrus swamps fish are harder to see and capture. Shoebills feed in shallow or deep water by Standing and waiting or Walking Slowly, usually on floating vegetation or along channels (L. Buxton et al. 1978, Guillet 1979). These feeding methods are the 'lurking and stalking' (Lauern und Pirschen) described by Moller (1982a) and by Guillet (1979). 'Instead of searching for his prey, [the Shoebill] waits patiently for it to come to him' (A.L. Butler 1905). Ambush is their speciality. Standing in one place, motionless, may last for 20-30 min (L. Buxton et al. 1978, Brown et al. 1982:192), but at other times Standing and Walking tend to blend together. Walking tends to be used more commonly in sites where uninterrupted movement is possible, such as along channels. On floating platforms of vegetation, feeding is restricted to Standing and waiting (Guillet 1979); it uses its extremely long toes to support its weight on floating plants—much in the manner of a giant jacana—often sinking slowly up to the tarsal joint or even deeper. It does not habitually wade in deep water, nor stalk prey in a crouched heronlike posture. Hopping is observed only as a startle reaction and it does not often run, although we once saw a bird run a few steps after something, with wildly flapping wings (pers. obs. M.P.K.). Normal progress is with a very slow stride, each step taking on average 5 s, and carrying the bird forward about 60cm (Moller 1982a). Occasionally it may briefly adopt a crouched posture during pursuit of a prey item. Owing to these preferred feeding behaviours, channels, lagoons and waterways that carry comparatively slowmoving water are used for feeding. Especially favoured feeding places are along trails made by large animals, which open up areas that would otherwise be too thick for Shoebills to utilize. Fishing is by visual location, rather than by touch, and hearing may also play a role (Guillet 1979). The bird feeds by holding the bill pointed vertically downward, to facilitate binocular vision. Fish, such as lungfish, are captured as they come to the surface to breathe, particularly in stagnant (i.e. deoxygenated) water. The final attack is performed extremely rapidly and, apparently uniquely, by 'collapsing' on the prey, throwing the wings forward and plunging down with the head and bill (Guillet 1979, Moller 1980, 1982a). The bird sometimes becomes entangled in vegetation, taking some time to recover. The tip of the bill absorbs the major force of the bird's relatively violent strike (Guillet et al. 1985), and the prey is secured by the sharp hook on the upper mandible. Vegetation, which is often taken into the bill along with the prey, is separated (Guillet 1979), and the prey is often decapitated by the extremely sharp mandibles before swallowing. From 1 to 10 min may elapse between

Shoebill

capture and swallowing of prey; turtles and snakes may take even longer (Moller 1982a). After the prey has been swallowed, water is often scooped up in the bill for drinking. On rare occasions 'collapsing' may be done from on the wing (Guillet 1979). The Shoebill's unique, boat-like bill is highly adapted to the bird's specialized mode of feeding (Guillet et al. 1985, Guillet 1987). Thanks to the anatomy of the bill and related skull structures, the Shoebill is able to catch unwieldy prey amid dense aquatic vegetation. A unique anatomical locking mechanism in the articulation of the jaw (Bock 1964:18) probably assists the bird in holding onto heavy, hard, or slippery prey. There is no evidence to confirm the rumour that Shoebills excavate aestivating lungfish from dried mud (Brown et al. 1982:192). Shoebills tend to forage singly, even if otherwise paired (Moller 1982a), seldom closer than 20 m to each other. We once saw 9 birds, scattered over a large marsh, southeast of Lira, Uganda (pers. obs. M.P.K.). Loose aggregations occur, particularly where water is drying out and large numbers offish have become concentrated (Guillet 1979). There is no evidence that Shoebills ever fish communally, in the manner of some pelicans, as suggested by P. A. Cottam (1957). Moller (1982a) reported that, in Uganda, most feeding takes place during peak periods between 06.00 and 08.00 h in the morning, and between 18.00 and 20.00 h in the evening. Reports of night feeding are often discounted (Brown et al. 1982:191), although Guillet (1979) reported that they can feed by the light of fires set by fishermen, and their very large eyes would suggest some noctural, or at least crepuscular, abilities (FJ. Jackson 1938:66). W. Fischer (1970:50) refers to the Shoebill as a Dammerungsvogel ( = twilight-bird). A bird in the marshes bordering Lake Edward, in western Uganda, was followed on all of 3 consecutive days (Moller 1982a); it moved approximately 0.5-1.5 km during foraging activities each day but apparently did not move from its resting place at night. On the other hand, C.R.S. Pitman (in M. Burton and C.W. Benson 1961), who had considerable experience with the species, reported that they feed 'mostly by night' and that they become active in the evening. In 551 h of observation in Uganda, Moller (1982a) recorded 111 strikes, 60 of which (54%) were successful. The shortest time between successful strikes was 23 min, and birds usually changed locations after a strike (Moller 1982a). This may indicate that the disturbance of a strike makes another successful one unlikely at the same place soon after. It seems that birds are fortunate to make about 2 kills per day. The diet of Shoebills is primarily composed of fish, especially lungfish (Protopterus aethiopicus), bichirs (Polypterus senegalus), catfish (Clarias spp.) and tilapia (Tilapia spp.). They also eat frogs, watersnakes, monitor lizards (Varanus niloticus), turtles and young crocodiles (Crocodilus niloticus). Rodents, snails and young waterfowl are taken on occasion. It has been reported that Shoebills will take newborn lechwe (Kobus leche) calves, but such reports (M. Burton and C.W. Benson 1961) are probably without foundation. (L. Buxton et al. 1978, Brown et al. 1982:192), especially since lechwe

143

calves are large and Shoebills swallow their prey whole (A. Guillet pers. comm. 1990). In any case, fish provide the bulk of the diet, and in many areas these include impressively large lungfish. In Uganda, Moller (1982a) observed that fish 15-50 cm long were eaten; a lungfish 1 m long was caught but then released. In areas frequented by hippopotamuses, the cloudy water may enhance the accessibility of large fish (Moller 1980). Ideal feeding sites have high densities of fish but are low in dissolved oxygen, causing the fish to rise for air. The diet does vary geographically. Moller (1982a) reported that lungfish and catfish are the main prey taken in Uganda, whilst L. Buxton et al. (1978) found that catfish (Clarias mossambicus) and water snakes were the main diet fed to young in the Bangweulu Swamps, Zambia. Zambian birds caught catfish weighing up to about 500 g and snakes 50-60 cm long (L. Buxton et al. 1978:205). A captive juvenile consumed approximately 0.91 kg of fish and meat per day (G. Steinbacher 1937, Owen 1958). The Shoebill's usual stance is with the bill resting on the breast (Mackworth-Praed and Grant 1970:51). It habitually stands motionless and statuelike for many minutes at a time, apparently waiting patiently for a lungfish or other prey to surface; it was characterized by FJ. Jackson (1938:65) as 'sombre-looking and listless .. . often looking more like a grey stump or post than a bird'. Often it stands on one leg while scratching with the other (Reichenow 1900-01:358, Guillet pers. comm. 1990). It does perch in trees, although none exist in much of its habitat (A.L. Butler 1905, M. Burton and G.W. Benson 1961, Guillet 1984). Shoebills soar on hot afternoons. At times, at least, this may have an advertising and territorial function (L. Buxton etal. 1978, Moller 1980). At least in some areas neither nesting nor fishing birds can tolerate even a low level of human disturbance (Guillet 1978a). They will desert a breeding area if disturbed before egg-laying (A. Guillet, pers. comm.). When the surrounding vegetation had not been burned and heavy cover remained intact, birds at established nests tolerated native fisherman as close as 20 m (Guillet 1984). Other observers have also commented on the lameness of the birds after the nest has been established (L. Buxton et al. 1978). Captive birds in the Tierpark Berlin periodically shed some of the covering on their bill (W. Fischer 1970:80-81); it is not known whether birds in the wild also do this. Also in the Berlin Tierpark, we have seen a captive Shoebill sunbathe, with its back to the sun and its wings spread with the tips touching the ground; when a cool wind started, the bird closed its wings (pers. obs. M.P.K.). BREEDING In Sudan, Shoebills nest solitarily, usually with less than 3 nests/km2. An area around the nest is often cleared by the adult birds of all tall vegetation for an area of about 3 m in diameter (Guillet 1984). In Uganda, a breeding pair may defend a 2-4 km2 area from intruders (Moller 1978). In the northern and southern extremes of its range, nesting begins after the rains end, just as water levels begin to

144 Shoebill

drop. At both Bangweulu, Zambia, and the White Nile region, Sudan, eggs are laid at the end of the rains, and young fledge late in the dry season or early in the next rains (Brown and Britton 1980:38). The very long breeding cycle of the Shoebill forces it to lay eggs in a wet season in some equatorial areas that have two rainy seasons per year. This is the case in Uganda, where Shoebills nest in March-June during the main rains (FJ. Jackson 1938, Kasoma and Pomeroy 1987). Breeding at this time would produce young fledging in the main dry season. Both birds build the nest, performing Up-Down greeting displays often in association with collection and presentation of nesting material (L. Buxton et al. 1978). Up-Down displays accompanied by bill-clattering are the commonest displays seen at the nest (Guillet 1984). Both birds stand erect with bills pressed close against the neck. They bow, point the bill downward and shake it repeatedly from side to side, accompanied by bill-clattering. The bill-clattering is rather slow (4-5/s) and has a deep, hollow quality, clattering sounds of male and female are individually distinguishable (Brown et al. 1982:191), probably owing to the relative sizes of their bills. Sometimes, at the beginning of an UpDown, the bird gives a hoarse, rasping squeal, then the bill is lifted to about 45° above the horizontal, so that the back of the head and crest rest on the upper back; then it brings the head down while clattering the bill (W. Fischer 1970:45, 58, pers. obs. M.P.K.). Vocalizations are also described as a high pitched whine or 'Moo' (L. Buxton et al. 1978). The Up-Down shows some apparent similarities to the display of the Marabou Stork (Kahl 1965, 1966a). Up-Downs are also given as a greeting to familiar people by captive birds (A.L. Butler 1905, G. Steinbacher 1937, Poulsen 1950, W. Fischer 1970). Other displays described by W. Fischer (1970) also resemble stork displays more than they do those of herons or pelicans (other suggested relatives of the Shoebill —see Taxonomy). In most areas pairs seem to defend all-purpose territories, 2.5-3.8 km 2 in size, with the nest fairly centrally located (Moller in Collar and Stuart 1985:66). Adults on the nest defend it with displays similar to the Forward Display of the storks, and also with bill-clattering and a display termed Bill Hook Exposure, in which the bill is raised, opened and pointed at the intruder. If ritualized threats do not intimidate an intruder, a bird 'jumps into the air and falls feet first onto its rival; the bill then follows, with the whole body weight behind it' (Guillet 1984:234). Nest-building is by both sexes and, before the eggs are laid, involves the birds 'sewing' new grass into the structure by jumping up and down and working the material in with their toes. After the eggs are laid, new additions of grass are just dropped on the nest, without any 'sewing' movements (Guillet 1984). The description of the behaviour of the Shoebill when collecting or adjusting nest material sounds somewhat similar to the Swaying Twig-Grasping display in Marabou Storks (Kahl 1966a, L. Buxton et al. 1978:218). The nest is usually built on a floating platform, in extensive marshes, and surrounded by high emergent vegetation and sometimes deep water (Guillet 1978a), sometimes it is

in an open, grassy glade within stands of dense papyrus (C.W. Benson 1961). Nest-mounds are large and flattish, up to 3 m across, built of floating vegetation, which may be partially submerged. The actual nest, often constructed of grass stems and measuring c. 1.0—1.7 m across, is placed on top of this mound. In Sudan, floating Shoebill nests will often support the weight of a man (Guillet 1984), but such was not the case at a floating nest in Zambia (L. Buxton et al. 1978). The report (Brown et al. 1982:192) that nests become water-logged and sink during the nesting season is incorrect (A. Guillet pers. comm. 1990). In some areas of the Sudan, nests are placed on termite mounds projecting about 1.5m above water level (C.W. Benson 1961, Critchley and Grimsdell 1970, Guillet 1978a). Nests are often added to and reused in successive years (Guillet 1984). The white eggs have a thick chalky coating and become much stained and dull during incubation. The normal clutch size is 2 eggs; 1- and 3-egg clutches do also occur (average = 1.7, n — 7; L. Buxton et al. 1978). Chicks can hatch with as many as 5 days between them, so that the first egg probably is incubated immediately upon being laid. Eggs (and later young) are regularly wetted with regurgitated water on hot days, up to four times daily. Eggs are turned either by the bill tip or by pressing the grass around the eggs with the toes. L. Buxton et al. (1978) considered this behaviour to be intentional rather than accidental; we have not seen egg-turning with the toes in any of the true storks. Both parents incubate. Nest-reliefs at a Zambian nest usually came in mid-morning and again in the afternoon, with the female doing about 75% of the incubation, including the night hours (L. Buxton et al. 1978). The duration of incubation is not known, but it is probably about 30 days. Nestlings are brooded, shaded, guarded and fed by both parents; however, at a Zambian nest the female was the more attentive (L. Buxton et al. 1978). The chicks give a hiccuping Begging Call, and peck at the legs of the adult and the hook on their bill-tip, thereby stimulating regurgitation. Adults hold the food, which is well champed up into soft pulp, between the mandibles from which the chick can obtain it; this mode of feeding is continued until whole fish can be swallowed by the chick, unaided, at about 30 days. Any food dropped onto the floor of the nest is later re-eaten by the adult. This method of feeding young differs from true storks, who regurgitate directly onto the floor of the nest. During the last "6 weeks of its time in the nest, a single Zambian young was fed an average of 6 fish and/or snakes per day (L. Buxton etal. 1978). Small chicks are continually shaded by the attending parent and on hot days regularly doused with water; some of the regurgitated water is drunk by the chicks. Shading continues until feathers are fully developed at about 60 days (L. Buxton et al. 1978). The pre-fledging period lasts about 95 days, at which time young first leave the nest on foot. They can fly at about 105 days and fly well by 112 days, but are still fed by the parents for a further period—possibly a month or more. Thus, from egg-laying to first flight takes 135-140 days, according to Guillet (1978a:42) and L. Bux-

Shoebill

ton et al. (1978); however, the latter authors stated that 'in all, from the start of nest-building to independence of the young, the nesting cycle would occupy 6 to 6i months' (180195 days). Most nests have been reported to contain 1 or 2 young, occasionally as many as 3 (A.L. Butler 1905, L. Buxtonetal. 1978). TAXONOMY As discussed in the chapter on classification, there has been historical debate on the affinities of the Shoebill. Powderdown patches on the lower back suggest possible affinities with the herons (Bartlett 1861), tongue morphology is similar to the Hamerkop (Scopus umbretta), and the syrinx is heron-like (Beddard 1888). The structure of the skull has led some authors to conclude that the Shoebill is a heron (Boehm 1930) .and others to suggest a relationship to the pelicans (P.A. Cottam 1957, Saiff 1978). Even in the last century, von Heuglin (1869-73:1096) called it 'a Marabou . . . with a pelican-like bill'. However, we feel that the structure of the bill—and related features of the skull—are so closely tied to the bird's specialized foraging techniques that they are probably not a reliable indication of ancestral relationships. Some recent studies, both behavioural and morphological, have converged on the conclusion that the species is more closely related to storks than to any other group (A. Feduccia 1977, L. Buxton et al. 1978). However, based on recent biochemical analyses, Sibley et al. (1988) have classified it as a subfamily of the Pelecanidae (with the true storks sharing an adjacent family with the New World vultures). The question as to whether the Shoebill excretes a dilute urine onto its legs in reaction to hyperthermia, as do all true storks (Kahl 1963b), is still unsettled. Heat-stress trials (Kahl 1967a) resulted in no urohydrosis by a captive Shoebill. However, L. Buxton et al. (1978) present some evidence [not sure proof, as suggested in Brown et al. (1982:190)] that Shoebills may at times excrete on their legs as a cooling mechanism. Watering of nestlings by parents may be related to this behaviour, since other similar birds (e.g. herons) are not known to do it. Because urohydrosis is probably rather complicated, from a physiological and genetic standpoint, it may be valuable in assessing taxonomic relationships. Clearly more work needs to be done on this problem. The Shoebill is distinctive in many aspects of its biology, and is thus, we feel, most appropriately placed in its own family, the Balaenicipitidae (L. Buxton et al. 1978, Kahl 1979a). The fact that we have included it in this book, while excluding the Hamerkop, indicates that we believe the Shoebill to be closer to the storks (Kahl 1967b, 1979c); however, the jury remains out as to how close. CONSERVATION The highly specialized ecology, patchy distribution, small populations sizes and extreme dependence on marshland that is subject to disturbance and drainage renders the Shoebill a highly sensitive species. The ICBP rates its position as of'special concern' (Collar and Stuart 1985:59). It has been included in Appendix II of the CITES Convention (K. Brouwerpers. comm. 1990).

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The main area of concern for this species is undoubtedly the southern Sudan. Major dangers are large-scale drainage (especially for the Jonglei Canal, irrigation and rice farming), cattle grazing and burning of trails, and increased fishing activity. All of these factors threaten Shoebills, especially when water levels drop late in the breeding season and nests are particularly vulnerable to cattle, fire and humans. In some areas, boundaries between Sudanese tribal holdings create a sort of 'no-man's land', in which Shoebills can sometimes nest undisturbed (Guillet 1978a). A major concern in Sudan was the planned construction of the Jonglei Canal; however, civil war has halted work on this project, at least temporarily. There is a demand for the Shoebill by zoos, because of its large size and distinctiveness. Brown et al. (1982:193) stated that 500 young were reportedly taken in 1978, but all of these birds were said to have perished in the Khartoum Zoo (K. Brouwer pers. comm. 1990). By late 1990, Shoebills were being offered for sale in the USA for US$10 000-20 000 each, making them the most expensive of all the available birds in the zoo trade (Bill Toone, San Diego Wild Animal Park, pers. comm. 1990). Captive breeding has apparently been reported only once, in 1964 in Elizabethville ( = Lubumbashi), Zaire (Jarvis 1966); since then, about 100 birds have been kept in zoos in the past 20 years, with no reproduction known (Collar and Stuart 1985:70, Guillet 1987, R.E.Johnson et al. 1987a). Not only does the taking of birds for sale reduce wild populations, but it teaches local people that such birds are extremely profitable, and encourages hunting and consequent disturbance. Many tribal customs forbad the hunting of Shoebills, which were feared and protected by myths and legends (M. Burton and C.W. Benson 1961). For example, people around Lake Salisbury, Uganda, believe that when in a canoe, you must never mention the name of the Shoebill, otherwise there will be a storm and the canoe will overturn (L. Tennant pers. comm. 1964). The policy of buying Shoebills from local people is helping to destroy these protective beliefs. Other people, not bound by such customs, are also penetrating these areas (Guillet 1978a). We feel that only zoos with serious breeding programmes for the species should be encouraged to purchase Shoebills. The Shoebill is protected by law in Sudan, the Central African Republic, Uganda, Rwanda, Zaire and Zambia (Collar and Stuart 1985:67). It is included in Class A of the African Convention of Nature and Natural Resources (K. Brouwer pers. comm. 1990). However, the construction of large water projects will potentially upset the ecological balance in several crucially important areas. Should the three-phase project envisaged for the eventual re-routing of the White Nile (the Jonglei Canal) be undertaken, the flooding pattern in the Sudd region of the Sudan would be completely altered, destroying the habitat. In Uganda, swamp drainage for rice and sugar cane plantations threatens the Shoebill. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 305.

American

White Ibis Eudocimus ruber (Linnaeus) Scolopax rubra Linnaeus, 1758, syst. Nat., ed. 10, p. 145; based chiefly on 'The Red Curlew' of Catesby, 1732, Nat. Hist. Carolina, pt. 5, p. 84, pi. 84: America ( = Bahamas) Subspecies: Eudocimus ruber ruber (Linnaeus) Eudocimus ruber albus (Linnaeus) Scolopax alba Linnaeus, 1758, Syst. Nat., ed. 10, p. 145; based on 'The White Curlew' of Catesby, 1732, Nat. Hist. Carolina, pt. 5., p. 82, pi. 82: America (=Carolina) Other names: Scarlet Ibis, White Curlew, Spanish Curlew, Red Curlew, Lesser Ibis, Currie Currie, Brown Ibis [juvenile], Brown Curlew [juvenile] (English); Corocoro, Corocoro Rojo, Corocoro Blanco, Corocoro Colorado, Coco, Coco Blanco, Curri-curri, Ibis Blanco, Ibis Escarlatea, Coclito Blanco, Coclito Colorado, Coclito Rojo, Coclito Mojoso [juvenile] (Spanish); Flamant, Ibis Flamant, Courly Rouge de Bresil, Ibis Rouge, Ibis Blanc (French); Rode Ibissen, Rode Ibis, Witte Ibis, Scharlaken Ibissen, Korikori, Flamingo (Dutch); Guara (Portuguese).

IDENTIFICATION The American White Ibis, more appropriately called the Scarlet Ibis in South America, is easily identified as a medium-sized ibis (56-68 cm long) typically with pure white or red plumage. Over most of its range, it is often the most abundant wading bird and, because of its numbers, flocking habits and obvious plumage, one of the more spectacular. The basic plumage of the adult is a solid colour, an immaculate white, a deep scarlet-red or an orange-pink. The wing-tips are glossy blue-black, so dark as to appear black in the wild. The facial skin, bill and legs are pink-red, and the iris is blue; darker in scarlet birds. The species lacks specialized plumes such as are found on many other species of ibis. The reddish plumage is variable among individuals and populations, in that the colour depends on a combination of season, food and genetics. In northern South America, individuals of various shades of pink or orange typically occur among the darker red birds. Birds are duller and more variably coloured in the inland marshes of Venezuela than on the coast (Luthin 1983a, P.O. Frederick pers. comm. 1990). Some mostly white birds have pink mottling. Males and females have identical plumages, but the male is substantially larger (Kushlan 1977d). Perhaps this difference is most apparent in bill length: the male's bill is 1.24 times that of the female in both subspecies (Kushlan 1977d, Luthin 1983a). The differences are such that they can be used to identify the sexes in the field, especially when they are together. During courtship, the bare skin of the legs, face and upper bill turns bright red, primarily through increased vascularization (Frederick 1987c). The bill usually darkens during courtship, but this feature is variable in ways that are not entirely

148 American White Ibis

documented as yet. In general, the distal portion of the bill darkens to a glossy black, and this darkening extends towards the head to a variable degree. In North America usually about two-thirds of the bill darkens, whereas in some individuals the colouring is more extensive. In northern South America, the entire bill is usually coloured in both white and red birds, but many have only partial darkening (see Ramo and Busto, no date). In southeastern South America, the female's bill remains brownish with a blackish tip during the breeding season (H. Sick 1985, pers. comm. 1990). The pattern of variability in this seasonal feature requires much additional study before its geographic trend can be understood. Also during the height of pair-formation, courting birds develop a bright red, naked throat-pouch, which can extend up to 3 cm. Both sexes develop some expansion of the bare throat area, but the extent of development is variable among individuals and populations (Beebe 1914, Kushlan 1973b, Rudegeair 1975a).Jn North America, the female usually (Kushlan 1973a, Rudegeair 1975a) but not always (Beebe 1914, C.R. Schmidt 1968) seems to have a more developed sac. It has been reported that in South America both sexes develop a sac (Luthin 1990, Wieringen and Brouwer 1990). However, the primary source of this information, Risdon (1969), described wattles on both sexes, rather than a gular sac, and that author admitted to being unable to tell the sexes apart. Sick (1985, translated by Belton 1990, pers.

comm.) speaks of seeing a small compact sac on either side of the throat rather than one sac. The photography of Ramo and Busto (no date) clearly show development is more extensive in the female Scarlet Ibis. Additional data are needed regarding geographic, sexual and individual differences in throat-sac development. Colour changes are very fleeting, and the intense colours fade soon after courtship is over. The intense skin colour pales almost immediately. Bills begin to fade after a few weeks, although a dusky wash can remain for months. During nesting, white plumage becomes soiled and discoloured and such birds can appear to have a dark or buffy wash. In the llanos, this wash may be yellowish (Ramo and Busto no date:81). A complete post-nuptial moult begins as nesting winds down and is well underway by the time nesting is completed. This moult begins on the neck and back, spreading to the wing scapulars, coverts, tail and finally the primaries. In the red form, necks may turn a pale pink as a result of this moult (M.v. Wieringen and K. Brouwer pers. comm. 1988), making the birds look scruffier and duller than in breeding plumage, and accentuating individual differences. An incomplete moult occurs prior to nesting (Beebe 1914). The chick appears grey. The head, back and wings of newly hatched young are covered with a grey to dark black down, more blackish on the head and more brownish on the wings. The ventrum is initially bare, becoming whitefeathered. Some individuals have a patch of white feathers on the head. This distinctive tuft may persist throughout nesting and provides a measure of individuality (De Santo et al. 1990). The skin is initially pink but turns grey within the first 2 days of hatching; the iris is brown; the feet and legs are pink-grey but turn grey during the nestling period (Beebe 1914, ffrench and Haverschmidt 1970, Antas 1979, M.v. Wieringen and K. Brouwer pers. comm. 1988, De Santo et al. 1990). The short, comparatively straight bill of the hatchling is pink with a dark tip, but becomes ringed by a variable number of black bands (usually 3), starting to develop at c. 6 days after hatching (M.v. Wieringen and K. Brouwer pers. comm. 1988). The coverage and pattern of the rings vary among chicks. By 6 weeks, the entire bill is grey and then turns pinkish over the next few weeks (Beebe 1914, De Santo et al. 1990). Juvenile plumage, which differs from that of the adult, begins to develop as the primaries emerge on the fourth or fifth day (De Santo et al. 1990). Other wing and body feather tracks emerge within 2 weeks, and the plumage is generally complete by the end of the third to fourth week except for retention of down on the head and neck through the fifth to sixth week. The head, back, upper breast, wings and tail are slate grey and the undersides white; the face and bill are pink, the legs grey and the iris slate grey. Juveniles fledge at a smaller than adult size and continue to grow after fledging, taking up to 2 years to achieve adult weight (Kushlan 1977f). Beginning as early as 4.5 months old in captivity and certainly by 8-9 months old in the wild, the juvenile feathers, now faded to a sandy grey, begin to be replaced with white or red feathers resulting in a mottled appearance until moult is completed. Most of the plumage

American White Ibis 149

has changed by the end of the second year, but brown mottling remains on the head and neck into the third year. First nesting takes place when young are 3 years old. Vocalizations are variable, but are generally characterized as honks. The primary flight and disturbance call is a coarse 'Hunk-hunk-hunk'. Nestlings beg with a distinctively shrill 'Zziu'. They often vocalize while feeding, the flock giving off a murmuring, muffled 'Huu-huu-huu'. During courtship 'Hunking5 and squealing calls are given. The American White Ibis is an accomplished aerialist, the birds flying in excess of 40 km/h, alternating rapid wing beats with gliding, typically in a variable V-formation (Pennycuick and De Santo 1989, pers. obs. J.A.K.). They soar to great heights. This behaviour is especially spectacular over a colony, even before the initiation of nesting. The birds also soar when gaining height to glide to feeding sites. Upon disturbance, a foraging aggregation will take to flight all at once and may perform manoeuvres before returning to the feeding site or landing on a roost nearby to await passage of the disturbance. Individual birds may engage in spectacular aerobatics. Especially characteristic is a free fall into a roost. Young and adults tend to flock and feed separately; typically first- and second-year birds are found together. DISTRIBUTION AND POPULATION Overall, the range of this species is quite extensive, but the breeding range is more patchy. It nests over much of the coastal plain of southeastern North America through Central America, the Greater Antilles, in northern South America, and in southeastern Brazil. The subspecies albus occurs in the northern portion of the range through Central America and perhaps into western South America. Although the line of demarcation between races is not clear, the subspecies ruber is found in the remaining part of the South American range. In North America, the American White Ibis has been slowly expanding and consolidating its range for the past several decades and now nests as far north as Virginia, south through Florida, to Louisiana (Sprunt 1944, Frohring and Beck 1978, Shields and Parnell 1983). It also nests in Baja California, both coasts of Mexico, Belize, both coasts of Nicaragua (Chinandega, Leon, Managua, Chontales, Caribbean Region of Zelaya), the Pacific but not the Caribbean coast of Costa Rica (Guanacaste, Puntarenas including Golfo de Nicoya and Golfo Dulce), and Panama (Panama Bay, Herrera, Los Santos, Chiriqui) (Slud 1964, Luthin 1984a, D.A. Scott and M. Carbonell 1986, Stiles pers. comm. 1988, Tejera 1989a,b). In the Greater Antilles, it nests in Cuba, Hispaniola and reportedly in Jamaica (Garrido and Garcia 1975, D.A. Scott and M. Carbonell 1986). In South America, nesting occurs in northwestern Colombia, interior and coastal Venezuela, and along the Atlantic coast through Guyana, Surinam, French Guiana and northeastern Brazil in Amapa to Maranhao, and in southeastern Brazil (Sick 1985, D.A. Scott and M. Carbonell 1986, Marcondes-Machado and Monteiro 1989, 1990, Teixeira et al. 1990, Antas et al. 1990). It previously nested in Trinidad, along the northeast Brazilian coast to Ceara and along

the southeast Brazilian coast (from Rio de Janeiro to Santa Catarina). The relative abundance of white and red birds varies over the overall range. Northern breeding populations are composed primarily of white birds, but not entirely so as scarlet birds have regularly been reported from the range of white birds since their description from there by Catesby two centuries ago. Fully scarlet birds continue to nest in the wild in Florida (pers. obs. J.A.K., P. Frohring pers. comm. 1990). The origin of these birds must remain uncertain owing to previous introductions, but the fact that we have observed them regularly for two decades is of some importance. In South America, nesting ibises are mostly red birds, with far fewer white birds from northern Colombia through Venezuela to Surinam. Red birds also are typically found down the Pacific coast to Ecuador and northeastern Peru (F.P. Penard and A.P. Penard 1908, Ramo and Busto 1982, 1984, 1988, Luthin 1984a, Serna 1984, Hilty and Brown 1986). In the llanos, about 4-6% (depending on where they are counted) of the birds are white (Ramo and Busto 1987). This highly nomadic species undertakes seasonal migrations, intra-regional population shifts and wide ranging dispersals (Palmer 1962, Kushlan 1976d, 1979c, Kushlan et al. 1985). In migration, North American birds move into Florida and Mexico, and perhaps into Cuba and Central America. This migration mostly goes around the Gulf of Mexico, with post-migration arrival dates at the northern Gulf coast in first 2 weeks of March (Stevenson 1957). Population shifts on a similar scale also occur. In Florida, the population shifts between seasonal interior wetlands and the coast (Kushlan 1976d, 1979c). A similar shift occurs in Venezuela, from the higher llanos to middle and lower llanos and to the coast (in Dec-May) where birds are also found year-round (Luthin 1983a, M.v. Wieringen and K. Brouwer pers. comm. 1988, McNeil et al. 1990, Kushlan et al. 1985). Coastal nesting birds also migrate, as evidenced by their seasonal occurrence in Trinidad (Hislop and James 1990). It has long been thought that these birds nest at the mouth of the Orinoco in Venezuela (fFrench 1967). Scarlet birds breeding in Surinam shift along the coast after the nesting (Held 1990). Shifts also take place in Brazil with birds sometimes moving up the Amazon into Para (Sick 1985) and to the coast in Sao Paulo and Parana (Guiz pers. comm. 1990). Wide-ranging dispersals take white and scarlet birds outside of their usual nesting and migratory ranges, and in fact they have been reported as vagrants or rare visitors over much of North, Central and South America. The extent of dispersal varies among years, probably related to water and feeding conditions. In North America, the species occurs occasionally as far as New York, Quebec, Pennsylvania, Illinois, Missouri, Kentucky, Wyoming, South Dakota, Colorado and California (e.g. Dresser 1866, Lincoln 1923, Palmer 1962, D.H. Davis 1979, Ernst 1974, Irmscher and Irmscher 1979, J.C. Miller 1982a,b,). White birds also occur rarely along the Caribbean slope of Nicaragua and Costa Rica (G. Stiles pers. comm. 1988), in the Bahamas, Jamaica, Cayman Islands, Puerto Rico, Trinidad, Aruba, Curasao, and along the Pacific coast as far south as Tumbes

150 American White Ibis

in northwestern Peru (Brudenell-Bruce 1975, Schulenberg and Parker 1981, Raffaele 1983, Bradley 1985). Red-plumaged birds are naturally accidental in Jamaica, Grenada, Cuba and elsewhere in the West Indies, and also (despite views to the contrary) in North America (Audubon 1840, Brewster 1883, Sell 1918, R.T. Dusi 1965, Simpson 1966, Cypert 1971, Garrido and Garcia 1975, Maehr and Hintermister 1982). A record from Honduras is considered to be doubtful (Monroe 1968). Scarlet birds also have been seen east of the Andes in Ecuador (Stevenson 1972). Population sizes of white and scarlet ibises are among the best known of any ibis species, because they nest in large colonies and are historically the most numerous ciconiiform wading birds in most of the areas in which they occur (with the possible exception in recent years of the immigrant Cattle Egret, Bubulcus ibis). Along eastern North America, over 64000 were counted nesting in about 60 coastal colonies in 1975 (Custer et al. 1980). Over 87000 nested in Louisiana in 1967 (Portnoy 1981). Colonies of over 10000-20000 birds have recently been reported from coastal North America (Osborn and Custer 1978, Frederick 1986a, Bildstein et al. 1990b). However, the largest of these colonies have been abandoned in the past few years (W. Post 1990, K.L. Bildstein pers. comm. 1990). In southern Florida, historical populations numbered near 100000; reports of historic numbers in the millions have no credence (Frohring et al. 1988). Until very recently, populations have remained relatively high there; in the 1970s populations numbered over 50000, including over 35000 in one colony (Kushlan 1973a, Kushlan and White 1977). Nesting numbers have been decreasing rapidly in the Everglades in recent years. In Mexico, numerous small colonies of 20-600 nests occur along the Pacific coast and smaller numbers of larger colonies, of more than 2000 nests, have been counted on surveys on the Gulf of Mexico coast (Knoder et al. 1980). More recently, 1750 pairs were counted nesting at Bahia de Santa Maria in Sinaloa, 1250 pairs in Tabasco and 1000-1500 pairs in Quintana Roo (D.A. Scott and M. Carbonell 1986). In Central America, colonies number in the hundreds to thousands. In Belize, numbers may be increasing; 400-500 pairs recently nested in Belize District (D.A. Scott and M. Carbonell 1986, D.S. Wood and R.C. Leberman 1987). Colonies are found on the Pacific coast of Costa Rica and in Panama; about 1000 pairs nest at Palo Verde and 600 near Canas in Guanacaste, Costa Rica (Delgado in Luthin 1984a, D.A. Scott and M. Carbonell 1986, Stiles pers. comm. 1988). In northern South America, the scarlet subspecies of the American White Ibis remain abundant and recent surveys have well-documented its nesting status (Spaans 1975a, 1982b, Ramo and Busto 1984, 1988, Luthin 1984a, Dujardin and Tostain 1985, D.A. Scott and M. Carbonell 1986, C.S. Luthin pers. comm. 1990, C. Ramo pers. comm. 1990, Ramsamujh 1990, Held 1990, Dujardin 1990, Hislop and James 1990, Teixeira et al. 1990, Antas et al. 1990). Numbers in Colombia are poorly known, but birds nest

along the coast and in the western llanos of that country. The population numbers in the hundreds of thousands in northeastern South America. The total in Venezuela is over 128000, including a colony of over 52000 birds; in the Venezuelan llanos, over 95% of the nesting population surveyed in 1983 were found in 22 colonies (Ramo and Busto 1988). About 300 pairs are known to have nested in Guyana in mid 1970s (Ramsamujh 1990). Nesting populations in Surinam numbered over 13000 pairs in the early 1970s, decreasing over the decade to 2000 nesting pairs, then increasing to nearly 35000 pairs in 1986 (Spaans 1982b, K. Brouwer pers. comm. 1988, Held 1990). In French Guiana, populations totalled about 3000 pairs in 1988 (Dujardin 1990). In 1991, 1700 pairs were counted in two colonies (Dujardin 1991). In Trinidad, ibises have been counted at their roost-site for many years (Hislop and James 1990). Birds using the roost increased from 1500 in 1969 to 10000 in 1987, but no nesting has taken place in Trinidad for 30 years. In Brazil, the species' status is becoming better understood (Luthin 1984a, Sick 1985, Bokermann and Guix 1987, 1990, Teixeira et al. 1990, Antas et al. 1990), although censuses of nesting colonies are few. In the northeast, one colony included approximately 5000 ibis nests. The species was once reported to be the most common wading bird of the Amazon region but over the last century has become scarce there (Sick 1985). In 1981-82, 7500 adults were counted at the mount of the Amazon. During the nonbreeding season, up to 9000 have been counted by R.G. Morrison (pers. comm.) in the northeast of Brazil. In the southeast of Brazil, these ibises were thought to have been extirpated, but birds have been observed there repeatedly since the early 1980s, particularly in Rio de Janeiro and Sao Paulo states. A small nesting colony was recently discovered at the junction of the Oncas and Quilombo Rivers (Marcondes-Machado and Monteiro 1990). Throughout the range of this species, because of its propensity to change colony sites, nesting numbers fluctuate from year to year and decade to decade in any one area, and such fluctuations may or may not reflect the underlying population sizes (ffrench and Haverschmidt 1970, Kushlan 1979b, Spaans and de Jong 1982, Dujardin and Tostain 1985, M.v. Wieringen and K. Brouwer pers. comm. 1988, W. Post 1990). For example, along the eastern North American coast, nesting numbers decreased 40% within 1 year, between 1975 and 1976 (Custer et al. 1980). Similarly, at two colony sites in North Carolina, nesting numbers decreased from 12 000 pairs at each site in the wet year of 1984 to 2000 pairs at each in the following very dry year (Bildstein et al. 1990a). Population fluctuations along the Suriname coast have been similarly extensive and impermanent. Some population fluctuations have resulted in local longterm decreases, however. This is especially the case in southern Florida, the Atlantic Coast of North America, French Guiana and southeast Brazil. The nomadic tendency of this species also suggests a low degree of isolation among populations. This has been shown to be the case genetically (Stangal et al. 1991).

American White Ibis

ECOLOGY The American White Ibis is a bird of interior and coastal marshes and swamps. Historically it probably reached its peak abundances in the great grass and sedge wetlands of the eastern USA and Venezuela, but throughout most of its range it occurs in coastal areas. In southeastern Brazil its historic range coincided with the extent of mangrove swamps. However in North America, it extends well into temperate areas, where it uses salt marsh and nearby freshwater swamps. This ibis actually can use a wide variety of feeding habitats, usually in shallow water (frrench and Haverschmidt 1970, Kushlan 1979c, Guster and Osborn 1978, Desenne and Shimotake 1990). Inland, it is most typically found in shallow seasonal marshes, swamps and river-edge hardwood forests, and almost as often in wet pasture and shallow ephemeral ponds. Along the coast its typical feeding habitat is marsh and mangrove swamp, where feeding is dependent on tides (Desenne and Shimotake 1990). Evidence has accumulated that the American White Ibis suffers reduced nesting success when restricted to foraging in salty habitats. So they often choose freshwater feeding habitats, even if the nesting colony itself is situated in a marine or estuarine environment. Several studies have found smaller clutch sizes on the coast than inland (Kushlan 1977d, Rudegeair 1975b). More recent studies have shown that young are incapable of enduring the high salt content of marine food (J.W.Johnston and K.L. Bildstein, 1990). Throughout its range, the species can take advantage of particularly good feeding opportunities that happen to become available. It should not be unexpected to see this species feeding in such places as open marine lagoons, on exposed tidal flats, along the tide line on beaches or even in the surf, in gardens, residential yards, ball fields and golf courses, in agricultural fields, following tractors, in garbage dumps, or in the accumulating manure under chicken coops. It feeds heavily in rice fields at the beginning of the wet season in the llanos (Wieringen and Brouwer 1990). Ibis populations move about to take advantage of available foraging opportunities, establishing a succession of roosts as conditions change from year to year, or even as a season progresses. Feeding locations are usually within 10 km of a colony site, although we have followed flights for over 40 km in southern Florida, as has Bildstein in coastal Carolina (Bateman 1970, Kushlan 1976d, 1979c, Custer and Osborn 1978, K.L. Bildstein pers. comm. 1989). This species can feed effectively whether water levels rise or fall as long as foraging sites of appropriate water depth are available (Kushlan 1979c). In general it appears that water depth is more important than direction of water-level change in determining foraging patterns of ibises in the Florida Everglades. A flocking species, American White Ibises may feed in large aggregations and fly in cohesive flocks between feeding and roosting sites (Kushlan 1978a, Wrege 1980, Petit and Bildstein 1987). Juveniles often flock separately, taking 2 months to begin flock flying; they are less successful at

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foraging than are adults and may feed on the periphery of adult flocks (Bildstein 1983, 1984, Petit and Bildstein 1986). Feeding aggregations form by local enhancement—the attraction of birds to other birds already feeding at a site, which is one way birds are able to locate suitable feeding locations (Kushlan 1976b, 1978a, 1981). Birds foraging within a flock have an advantage in being able to spend less time on surveillance (Petit and Bildstein 1987). The American White Ibises feed primarily by non-visual Probing in shallow water but also use such techniques as Head Swinging, Groping and Pecking when appropriate (Kushlan 1977c). They hunt crabs visually. Their usual behaviour is to Probe deeply in the water while Walking Slowly. They may Stand to concentrate their Probing under a plant or within a crayfish or crab burrow. On dry or mostly dry land they generally feed visually, picking up prey items from the surface of the soil (Pecking). Details of the foraging techniques, such as rates of probing, capture, walking and turning, vary depending on the nature and availability of the food supply (Henderson 1981). The close association with other wading birds can result in intense interactions among individuals. Ibises defend their individual distances aggressively and are subject to having their captured prey stolen by nearby birds (Kushlan 1978c, Kilham 1984). Ibises can be aggressive and can successfully steal prey from other species (P.C. Frederick pers. comm. 1990). Conversely, they are used as 'beaters' by other species that walk after them as they forage (Courser and Dinsmore 1975, Kushlan 1978b, Kilham 1980, 1984). Desenne and Shimotake (1990) found them feeding commensally with capuchin monkeys (Cebus sp.). The diet has been well documented. The American White Ibis is a specialist on aquatic invertebrates, with the bulk of its diet in most instances consisting of crayfish, small crabs, and aquatic insects and their larvae. Their feeding methods make them selective feeders on these prey in shallow freshwater habitats (Kushlan 1979b,c). In coastal marshes, they seek out fiddler crabs (Uca spp.), the females of which appear to be more vulnerable to predation (Bildstein et al. 1989). All types of aquatic and terrestrial insects are taken, along with a wide variety of other prey types (Nesbitt et al. 1974, Kushlan and Kushlan 1975, Haverschmidt 1968, Sick 1985, Kushlan 1979c, M.v. Wieringen and K. Brouwer pers. comm. 1988, 1990, Tejera Nunez 1990, Desenne and Shimotake 1990). Among their other prey, they eat worms, snails, bivalves, frogs and small fishes, when they can catch them, and almost any other small item they can grab and handle. Adults require 165kcal/day; young require 8620 kcal of food during the nestling period (Kushlan 1977b,c). The larger male requires more energy than does the female, and the male's size is detrimental to meeting his energy requirements because he obtains no compensating increase in foraging effectiveness (Bildstein 1987). Because of its large population sizes, the American White Ibis can contribute substantially to energy flow through the avian community. The southern Florida population of 62000 birds required 930 000 000 kcal in 1 year (Kushlan 1977b,c). In a temperate saltmarsh, the species accounted

152 American White Ibis

for 17% of the avian biomass and 10% of the energy flow through the bird community of the wetland (Christy et al. 1981; Bildstein et al. 1982a, K.L. Bildstein pers. comm. 1989). BREEDING In the American White Ibis breeding is highly communal, colonies often being both large and densely packed with nesting birds. This species usually nests in trees and bushes but also nests in such places as cactus, grasses and sedges, or on the ground. Colonies are situated especially on true islands, on islands of trees within marshes, in mangrove swamps and in gallery forests. Except on true islands, the presence of water under the nest sites seems to be crucial to colony formation. Colony sites usually develop at roost sites through the formation of daytime bachelor parties of males that remain at the roost and on the nearby ground to begin their display activity. Throughout its range, the species is highly vagile in its use of nesting colony sites. Populations move around frequently, and it is more typical for a site to be used only once or just a few times than for it to be used regularly over many years. This is clearly a rather nomadic species. In general, nesting appears to take place when local conditions are suitable for effective foraging, primarily the availability of sufficiently shallow wetlands or the shallow flooding of higher habitat (Kushlan 1976d, 1979b). In the Florida Everglades, this is usually during drying conditions, although ibises there can feed effectively on both falling and rising water levels (Kushlan 1979c). Along the coast of North America and in the llanos, nesting is on rising water levels at the onset of the wet season (M.v. Wieringen and K. Brouwer pers. comm. 1988, K.L. Bildstein pers. comm. 1990, C.S. Luthin pers. comm. 1990). Nesting along the coast of South America is more variable; and, although they appear to be less influenced by climatic conditions, nesting is mostly in the wet season (M.v. Wieringen and K.L. Brouwer pers. comm. 1988, C.S. Luthin pers. comm. 1990). Because of their dependence on water conditions, the actual timing of nesting varies throughout the species' wide range, and also in any area from one year to the next. In South Florida, American White Ibises may nest any time from the more usual spring nesting period (Mar-Apr), through the summer, and even into the autumn (Rudegeair 1975a, Kushlan 1976d, Allen-Grimes 1982). Nesting may be a bit later in coastal colonies, lasting into the summer. Timing in coastal Carolinas is similar, April-May (K.L. Bildstein pers. comm. 1990). In the llanos, they begin to nest in June and July (M.v. Wieringen and K. Brouwer pers. comm. 1988, Luthin 1990). In Trinidad, egg-laying varies from April to September (ffrench and Haverschmidt 1970). Nests with eggs have been recorded in Surinam from April to August, and in French Guiana in mid-April to June (Haverschmidt 1968, Dujardin 1990). Less exact information is available from Brazil; in the northeast, active colonies were found from March to September (Antas et al. 1990). Courtship behaviour has been well documented, and

there appears to be no substantive differences between the colour forms (Palmer 1962, Kushlan 1973b, 1977d, Rudegeair 1975b, Spil et al. 1985, Ramo and Busto 1985, C.S. Luthin pers. comm. 1990). Display Sleeping, Display Shaking, Display Preening and Supplanting Flights are common displays seen in bachelor parties. Display sites are tightly defended by males with Forward display and short Supplanting Flights. Display Preening, Head Rubbing and Bill Popping are the primary Displays of the male. Head Rubbing is quickly initiated and may include a complete roll of the bill from left to right or only a quick touch of the back of the head in a pumping motion, rather reminiscent of the Stretch display of herons. Full Bill Popping includes twig grasping and nibbling, and may be accompanied by complete extension of the neck until the head is down between the legs. In South American birds a rocking motion has been seen (M.v. Wieringen and K. Brouwer pers. comm. 1990). A female approaches displaying males very cautiously in the Bowing display, especially showing the side of her face to the male. This posture presents the fire red face and gular pouch, highlighting the eye (Kushlan 1977d). The male gives Forward display and may actually attack, sometimes going so far as to grab the female's head and shaking it like a twig; this often appears to be redirected Twig-grasping. M.v. Wieringen and K. Brouwer (pers. comm. 1988) did not see such head-grasping in South America. Before pairbonding is completed, the female can become quite bloodied in her attempts to remain in the male's display area. During courtship, the pair uses Greeting displays and mutual Stick Shaking, typically with necks crossed. Both also spend considerable time Display Preening and Standing touching each other. Greeting displays may include twig-grasping and hunking vocalizations by the males and squealing vocalizations by the females, the latter ceasing later in courtship. Nests are usually near but not necessarily on the display site; actual nest-site selection falls to the female, who builds the nest with sticks brought by the male. She shoves the twig into the nest-platform and positions it by moving it up and down in a tremble shoving motion. Nests are typically closely grouped, even almost touching each other. Nestplatforms often contain fresh leaves, replenished by the male during incubation. Copulation begins with the male placing his neck over that of the female as they engage in mutual twig-grasping. The male then mounts, achieving and maintaining his position by treading his feet and flapping his wings. Copulations occur throughout the day. The first egg is laid 5-6 days after the beginning of copulations, and additional eggs are laid at 1- to 2-day intervals, the latter being usual (P.C. Frederick pers. comm. 1990). Extrapair copulations are common in the later stages of pair-formation and even into incubation. The male initiates these—the female, although capable of resisting, seldom does so—and it is estimated that over 6% of successful copulations are outside of a mated pair (Frederick 1987a). Clutch size is 2-5, averaging 2-3 depending on location:

American White Ibis

clutch sizes averaged 1.97 in coastal Louisiana, 2.07 in inland northern Florida, 2.20 on the Florida coast, 2.23 in coastal North Carolina, 2.43 in Venezuela, 2.45 in the Everglades, 2.49 in interior central Florida (Kushlan 1977c, Rudegeair 1975b, Allen-Grimes 1982, Shields 1985, Hammatt 1981, B. Busto and C. Ramo pers. comm. 1988). Eggs are smooth and not glossy, splotched with brown over a base colour ranging from cream to blue-green. Shape varies from ovate to elongate, some surprisingly elongate. Nest-attendance patterns of the parents depend on the stage of nesting and distance to the feeding sites (Kushlan 1976c, M.v. Wieringen and K. Brouwer pers. comm. 1988). The male tends to remain at the nest throughout the nest building period, as departures put the nest and its contents at considerable risk (Kushlan 1973b, Frederick 1986a). During the 21-day incubation period and during early hatching, the pair alternates nest attendance. In the Everglades, the males were observed to remain at the nest during the day early in nesting (Kushlan 1973b), but this probably is more related to food location and the number of foraging trips. Observations in other locations and later in nesting indicate that the parents alternate more frequently (Wieringen and Brouwer 1990), and this is probably the more typical situation. It is clear that the larger size of the male is effective in defending against nest predation or nest material robbing, in that lone females are susceptible to being driven from the nest site (Frederick 1986a). Contests between males seem to be settled by size difference (P.C. Frederick pers. comm. 1990). Predation by Black-crowned Night-Herons (Nyeticorax nyeticorax), crows (Corvidae), and other birds is a particular threat to unguarded young (Kale 1965, Shields and Parnell 1986). Crows may steal eggs and unintentionally drop them in other nests (Shields 1987). The guardian defends the nest or young with Forward display and Bill Thrusts. Renesting after disturbance is possible, but its frequency is not known. Eggs in a clutch hatch 1-2 days apart, and the hatchlings are incapable of even holding their head up for the first day or so. To feed the hatchling, the adult grasps the young's bill stimulating it to raise its head and then regurgitates into its mouth. After a week, the young initiate feeding with a Begging display by touching the parents bill while calling and vigorously flapping their wings. Males may pirate food from females and young, and considerable piracy occurs among juveniles (Frederick 1987a, De Santo et al. 1990). The feet of the young develop rapidly and the birds can move from their nest as early as 2 weeks, forming creches (Kushlan 1977f, K. Brouwer and M. van Wieringen 1990a). At this time, both adults are away from the nest foraging at the same time, returning to feed the young up to 9 times/ day. Young leave the colony when about 50 days old, at which time the bill is under-developed relative to adult size whereas the legs are nearly fully grown, an adaptation for walking foraging (Kushlan 1977f, De Santo et al. 1990). Nesting success is often poor, usually averaging near one or so young per nest and about a 50% mortality (Kushlan 1977c, Girard and Taylor 1979, Allen-Grimes 1982, Frederick 1987a). Colonies suffer large mortality due to high tides,

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predation or lack of food. In southern Florida, colonies fledged about 1.03 young per nest. In Louisiana, 64% of all nests were lost during incubation by high tides (Hammatt 1981). A similar situation in South Carolina led to repeated flooding at a coastal colony, and subsequently its total elimination by a hurricane (A. Dinep pers. comm. 1990, K.L. Bildstein pers. comm. 1990). In this site, production of fledglings decreased from 39% in a wet year to 3% in a dry year (Bildstein et al. 1990a). Over its range, rainfall and water-level conditions are usually the primary determinant of nesting success in this species. TAXONOMY The Eudocimus ibises (Parkes 1951) appear to be most closely related to the Plegadis ibises (Mayr and Short 1970). However, the Plegadis ibises have a distinctive conformation of the tarsometatarsus and so are readily distinguishable (Olson 1981). Eudocimus is the more primitive group, with the fossil Plegadis paganus being somewhat intermediate. Examination of additional fossil material would be desirable to resolve the question of derivation of the Plegadis ibises from Eudocimus stock. The relationship between extant North and South American populations of the genus Eudocimus has long been a matter of interesting discussion. Ridgway (1884) long ago proposed that they were the same species. However, subsequent interpretations gave special credence to one of the earliest visits to a colony in the llanos by Zahl (1950), who found no mixed-colour pairs, although his visit was very short. None the less until recently, this weak evidence suggesting that the two forms do not interbreed has influenced interpretations of their taxonomic relationships. Recent studies, especially by Cristina Ramo, Benjamin Busto, Charlie Luthin, Eduardo Aguilera Prieto, Koen Brouwer, and Marcel van Wieringen, have provided additional evidence. Ramo and Busto (1982) formally proposed that the white and scarlet ibises be considered to subspecies. As first revisors, they chose the name Eudocimus ruber for the species. The primary differences of possible taxonomic importance between North and South American populations are in plumage colour, size and some details of the soft-part coloration. The fundamental taxonomic question is: how different are these forms and are these differences sufficient to inhibit interbreeding if they were in geographic contact? The most obvious distinction, of course, is in plumage coloration of the adult. (The juveniles are identical.) The red coloration of the birds is due to the deposition of specific carotenoid compounds secondarily derived from dietary pigments (Fox 1962, Fox and Hopkins 1966). This implies a genetic basis. However, the actual colour expressed in an individual (its phenotype) is the result of a combination of dietary influences and genetics. As a result, plumage colour is variable within and between populations. In the llanos, for example, birds range from deep scarlet, to orange, to white. The genetic component is probably a simple difference in a biochemical pathway that could be under the control of a

154 American White Ibis

single enzyme system, and perhaps a single gene. Salmoncoloured birds result from the mating of white and red adults, the faded colour apparently resulting from the incomplete dominance of the dominant colour gene. If this indeed is the case, back-crossing of a genetically salmoncoloured bird (heterozygote) to a homozygous white or red bird would produce either another pink bird or a pure white or red bird like the parent. Thus, we would expect to see all three forms in the population, which is the case in South America. That the genetics is slightly more complicated is suggested by the occurrence of slightly mottled white and pink birds. South American birds, both white and red, are smaller than those from North America. However, such a size cline is to be expected in a bird which ranges from the temperate to tropical climates. Wieringen and Brouwer (1990) provide strong evidence for the existance of such a cline in their comparison of the sizes of birds from Florida, Venezuela and Surinam. There remains more differences in size between the sexes than between the two colour forms, however. It has been suggested that the North and South American forms differ in soft-part colours in that the entire bill of the scarlet ibis turns black during the height of courtship in the South American birds whereas it only partially turns black in North American birds. However, there is much variation and these colours hold for only a very short period at the height of courtship. Some South American birds do not have a fully black bill (see photos in Ramo and Busto no date) and some North American birds do (pers. obs. J.A.K.). H. Sick (1985, pers. comm. 1990) stated that, in the southern South American populations, the bill of the male turns shiny black but the female's bill stays brownish with a blackish tip only; this also occurs in northern South America. The presence of the throat-sac is another character of interest. A case has been made that in North America the female ibis has a throat-sac during breeding, whereas both sexes have a sac in South America (Wieringen and Brouwer 1990). However, both sexes show some enlargement of the throat during courtship in both populations (Beebe 1914, C.R. Schmidt 1968, pers. obs. J.A.K.). Several observations do suggest that more males may have obviously enlarged sacs in South America, although the citation generally used to support this argument (Risdon 1969) notes the occurrence of small wattles, not fully developed throat-sacs (he also admitted that he could not tell the sexes apart). All observations indicate that the courtship behaviour of the two forms are virtually identical (Spil et al. 1985, C.S. Luthin pers. comm. 1990). Some minor differences have been reported under zoo conditions, such as a scarlet ibis rocking with its head down (M.v. Wieringen and K. Brouwer pers. comm. 1988). But, of course, birds used to each other, especially in captivity, could be expected to exhibit somewhat artificial behaviour. If one were to draw a conclusion from the above discussion alone, the weight of the evidence provides no reason why these two clearly related forms should not be considered the same species. However, this is inferential and definitive evidence can only be obtained from observations

of interbreeding. Fortunately such evidence exists, and it was such data that compelled Ramo and Busto (1982) to propose that the two forms be considered the same species. The two forms are entirely interfertile; this despite any of the potential differences in size and soft parts discussed above. They have long been known to interbreed in zoos (A.P. Gray 1958), although such evidence is scarcely more than suggestive as even distantly related animals interbreed in captivity. However, in this case, an introduction experiment demonstrated that interbreeding can occur in the wild. Eggs from scarlet birds were introduced into Florida (Bundy 1965, Zahl, 1967). These, and their presumed descendants, interbred freely with the resident white ibises. Birds of scarlet plumage remain in the South Florida population, and we have been observing their interbreeding with white birds for nearly 20 years. If they had derived from the introductions, they would persist after several generations of outbreeding with white birds in this way. The next question is whether hybridization could occur in the wild, should white and scarlet birds nest together naturally. This indeed takes place in northern South America, where Ramo and Busto (1982, 1985, 1988) and G.S. Luthin (pers. comm. 1990) studied such pairs in the wild. They observed no differences in nesting or courtship behaviours in wild birds and saw considerable interbreeding between white and scarlet forms. There is no evidence that the offspring of these pairings would be in any way disadvantaged. Thus, we conclude that colour alone is not a characteristic that distinguishes between two species. Based on this basic information, two hypotheses have recently been proposed to describe taxonomically the relationship between the two forms. Ramo and Busto (1982) proposed that these two forms be considered the same species, but that the scarlet and the white birds be considered to represent two subspecies. Wieringen and Brouwer (1990) proposed that the North American form and the South American form be considered separate species and that the South American form be considered dimorphic. In considering these competing proposals, it seems first that no taxonomic characters are known that separate either white and scarlet birds or North and South American birds into populations that are incapable of interbreeding. This supports the suggestion of Ramo and Busto that white and red birds and North and South American birds are conspecific. It is not appropriate to define subspecies on the basis of plumage colour alone, which Ramo and Busto appear to have done, when plumages do not sort out geographically. This supports the suggestion of Wieringen and Brouwer that the South American form, at least, is dimorphic. It is even possible that the North American form also is dimorphic, considering how regularly scarlet birds have historically been encountered in the normal range of the North American birds. To our knowledge, consideration has never been given to such a possibility. The clearest concordance of the evidence with taxonomy is to recognize a single species and two subspecies, the latter being identified (for now) by a combination of character-

American White Ibis

istics. The South American subspecies ruber is smaller, usually having an all or mostly dark bill during courtship, and males more often showing a gular sac. This subspecies includes both white and red birds. The North American albus is larger, usually having a part of the bill dark-coloured during courtship, and the males seldom showing as full a development of the gular sac as females. It should be noted that none of these characteristics is definitively understood. We suggest that this species is an example of the allopatric divergence of the two stocks followed by secondary contact in northwestern South America. Such a situation would be identical to that believed to have occurred in two other forms in the New World, the Great Blue Heron/Great White Heron (Ardea herodias) and the Green-backed/Striated Heron (Butorides striatus), both of which are thought to represent subspecies in secondary contact distinguished by plumage differences (Hancock and Kushlan 1984). The Great Blue/Great White situation is also similar in that the southern Florida race is dimorphic. Our hypothesis of polytypic conspecificity should in no way be taken to imply that the question of the relationship between the predominantly white North American birds and the predominantly red South American birds has been resolved. Like all taxonomic proposals, this is a hypothesis to be further investigated. It is our hope that detailed quantitative studies of morphological and biochemical variation and genetic control of plumage colour will be undertaken to determine what, if any, are the appropriate specific and subspecific limits of populations of scarlet and white ibises. Among problems to be addressed are documentation of the sizes and other characteristics of birds nesting in Central America and the birds in western South America. Further information is also needed on the size and other characteristics of red birds in northern populations, and of the southern South American forms. Although such studies would test our hypothesis of conspecificity, more importantly the scarlet-white ibis system could provide a mechanism by which fundamental issues regarding gene flow, polymorphism, gene expression and speciation could be studied. CONSERVATION The white and scarlet ibises are well recognized birds throughout their range, because of their impressive population sizes and, in South America, because of their remarkable coloration, which Beebe and Beebe (1910) called jets of flame. The sight of flocks of white or scarlet ibises flying to roost is one of the more memorable wildlife experiences and may be one of the great spectacles of nature. Such roosting flocks were prime causes of the establishment of the great Everglades National Park in Florida in the 1930s and for the fame of the Caroni Swamp in Trinidad. F.P. Penard and A.P. Penard (1908) captured the scene of a scarlet ibis roost by describing one 'as mangroves . . . splattered with blood'. The impression made by white ibis flocks is scarcely less, paling only in comparison with the scarlet form. These birds are increasingly taken to be symbols of conservation activities in the areas in which they occur. Populations remain high over most of the species' range,

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over 128000 birds nesting in the llanos alone (Ramo and Busto 1984). However, local populations have decreased in several areas, such as in the Florida Everglades and along French Guiana (Spaans and de Jong 1982, pers. obs. J.A.K.). The scarlet ibis has been placed on the endangered species list in Brazil (Sick 1972, 1985) and little information is available on the status of the disjunct populations in Sao Paulo and Parana, which have reappeared and are in need of conservation action, especially as they may be threatened by future pollution of their newly discovered colony site (Marcondes-Machado and Monteiro 1989, 1990). Over its range, white and scarlet ibis adults have traditionally been hunted, and eggs and young collected for food (F.P. Penard and A.P. Penard 1908, Bent 1926, Wetmore and Swales 1931, ffrench and Haverschmidt 1970, Herklots 1961, T. Barbour 1943, Ramsamujh 1990). Its appealing taste may derive from the predominantly invertebrate and crustacean diet. Disturbance and killing from hunting and other causes continues to be one of the primary causes of reduction in numbers of breeding birds in various areas (Haverschmidt 1968, Luthin 1984a). Hunting was allowed in French Guiana until 1984 and hunting in Guyana continues year round (Ramsamujh 1990, Dujardin 1990). Restrictions on hunting have been in effect in Surinam since 1984 but enforcement is lax, as is the case throughout much of the range (Held 1990). In Trinidad, where the scarlet ibis is a national symbol, hunting has persisted even within its sanctuaries (Hislop and James 1990). Conservation of these species in the neotropics will require education as well as enactment and enforcement of protective laws. Recently a commercial threat has emerged in French Guiana. Ben de Jong and Jean-Luc Dujardin found that an extensive local industry had developed that produced artificial flowers from red ibis feathers (Betlem and de Jong 1983, Dujardin 1990). Despite efforts to outlaw the taking of ibises and to find alternative sources of feathers, ibises continue to be killed (Dujardin 1990). With the concentration of a large portion of the species' population in a few colonies, protection and management of colony sites become crucial. Over the wide range of the species, nesting sites are threatened by lumbering, wetland reclamation, and water management. Protection is made difficult by the inaccessibility of many colony sites, and the species' propensity to move colony sites frequently. In developed countries, such as in the USA, potential nesting sites are lost through alteration of the site or nearby wetlands. W. Post (1990) documented the recent extinction of the largest White Ibis colony along the Atlantic Coast. This colony was stable from 1974 to 1984, when over 18 000 pairs nested, but then declined until it was abandoned by the ibises in 1987. Causes included susceptibility to predation, drought and floods that affected ibises more than the herons. Thus complete protection and management of remaining colony sites is critical in North America. Colony site protection in Central and South America is even more difficult because of the political and economic situation. However, long-term sites must still be protected

156 American White Ibis

in sanctuaries and reserves. One of the most famous sites (although no longer a colony site) is in the Caroni Swamp of Trinidad, where many people have observed it coming into its night roost and a tourist industry has developed surrounding this nightly event. Critical too, of course, is the protection and management of wetland feeding habitat. Population decreases in the Everglades are a response to alteration of the natural hydrological regime of the marsh by water management practices (Kushlan 1986a, 1987). This species has the advantage of being able to shift easily and quickly among foraging areas to avoid unfavourable conditions or take advantage of newly created foraging opportunities. Wetlands will need to be managed to maintain suitable foraging conditions for the White Ibis and other wading birds. The recently elucidated story of the Scarlet Ibis in the Garoni Swamp of Trinidad is especially instructive in this respect (ffrench 1967, 1978, 1985, ffrench and Haverschmidt 1970, Betlem 1984, Bildstein 1990a). It was not clear why

breeding ceased after this time until Bildstein, drawing upon his studies of the White Ibis in coastal environments, suggested that the ultimate cause was the drainage of freshwater marshes that served as feeding grounds during nesting. This conservation story holds a message for management of this species elsewhere, the importance of protecting the feeding habitat. The best hope is to preserve large tracts of coastal and inland wetland habitat. The species serves as both an indicator of environmental health and as a symbol of the need to undertake such habitat conservation. Several groups are now working on conservation plans for the species, and these must be encouraged and implemented.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 306.

Barefaced Ibis Phimosus infuscatus (Lichtenstein) Subspecies: Phimosus infuscatus infuscatus (Lichtenstein) Ibis infuscata Lichtenstein, 1823, Verzeichniss Doubletten Zool. Mus. Berlin, p. 75, based on 'Afeytodo', no. 365, of Azara, 1805, Apuntamientos Hist. Nat. Paxaros Paraguay Rio Plata, 3, p. 201: Paraguay Phimosus infuscatus berlepschi Hellmayr Phimosus berlepschi Hellmayr, 1903, Verh. Zool.-Bot. Gessell, Wien, 53, p. 247: Orinoco River region Phimosus infuscatus nudifrons (Spix) Ibis nudifrons Spix, 1825, Avium Species Novae Itinere Brasiliam, 2, p. 69, pi. 86, 'Ad litora lacuum fl. St. Francisci: Rio Sao Francisco, Bahia, Brazil Other names: Whispering Ibis (English); Coclito, Coclito Negro, Cocli, Ibis Negro, Zamurita, Cuervillo de Cara Pelada, Cuercillo de Cara Afeitada, Cuervillo Pico Marfil, Cuervillo Cara Roja, Cuervo de Frente Pelada, Curri-curri, Corocoro Negro, Tara, Corocoro Carirrojo, Yervetacho (Spanish); Tapicuru-decara-pelada, Macarico-de-bico-branco, Magarico-de-cara-pelada, Macarico-preto, Tapicuru, Frango d'agua, Chapeu-velho (Portuguese); Zwarten Clans-Ibis (Dutch)

IDENTIFICATION The Barefaced Ibis is a small, dark ibis with green-glossed plumage and a reddish face. It is 46-56 cm long weighing about 578 g (n = 8, range 500-600 g; Belton 1984, C.S. Luthin pers. comm. 1990). The basic plumage colour of the adult is black but glossed with a bronzy green or blue, more so above and on the tail and wings than below. This coloration gives the bird one of its Spanish names, Corocoro Negro (B.T. Thomas pers. comm. 1990). The flight feathers and underwing are bluish green. The underparts tend to a greenish brown. The forehead, face and chin (gular area or pouch) are bare, thus providing the most common English name, Barefaced Ibis. These and the bill are reddish, ranging from flesh to reddish brown in coloration during the non-breeding season. The long decurved bill is slightly serrated, the same colour as the face. The forehead may be covered with papillae or corrugations. Feathering around the face is glossy violet. The leg and feet colours are very variable, reported as yellowish to whitish pink, reddish, grey and tan to brown. The iris is light brown (F.P. Penard and A.P. Penard 1908), although Belton (1984) found the iris to consist of brown, white and red concentric circles. Additional information is needed to understand geographic, seasonal and developmental differences in soft-part colouring. The three named races are described based on slight differences in plumage and soft-part colours (Blake 1977). All have bare pink to reddish faces. The subspecies berlepschi has a relatively dark reddish brown bill tipped in blackish and a deep red face. The nudifrons subspecies has a paler clay-coloured bill, brick red face, and a green gloss on its hindneck feathers. Its forehead and face are rough and ridged. The infuscatus subspecies is reported to be smaller and shorter-billed than nudifrons and to have a pale clay- to cream-coloured bill, though Belton (1984) reported this as a greyish red. Its forehead and gular sac are yellowish pink, and strong purple reflections gloss the feathers of the hindneck. During the breeding season the soft parts attain their brightest colours, in some specimens appearing bright red (Luthin 1983a). The throat area, also called the gular patch or pouch, becomes red and is prominently displayed during courtship.

Barefaced Ibis

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typical quietness of this species outside the nesting season is quite odd within the loud-voiced ibis family. The species is more vocal during nesting. Luthin (1983a) characterized the courtship vocalizations as falling into four types, which he called multiple hooting, duetting, longhooting and pair-calling. Multiple hooting is the most commonly heard call, especially early on in nesting. This is a series of several hoarse hoots, with the first sound being louder than those following. A duet of these calls may be performed by members of a pair, usually at the nest site or future nest site. The duet is often preceded by a 'Whoop' sound, the long-hoot. Pair-calling is a series of 9-30 rapid soft hoots, by one of a pair. It is used early in nesting as a contact call. This dark ibis is distinctive from others in its range in its bare, pink face and reddish bill. It can be confused with the all-dark Glossy and Green Ibises in poor light. The Green Ibis is larger with a dark bill, greenish gloss and short legs. The Glossy Ibis and Whitefaced Ibis have a chestnut sheen. The Sharptailed Ibis is larger and distinctively shaped, with its stocky build and long tail. DISTRIBUTION AND POPULATION

The sexes are considered to be similar, but a review of specimens in the US National Museum suggests that males tend to be larger then females, at least in weight and in bill and tarsus length. With one exception, of 8 specimens examined, the bills of birds labelled Temale' were smaller than those of 2 identified as 'male' (pers. obs. J.A.K.). Belton (1984) recorded that a male weighed 575 g, a female 550 g, and an unsexed (live) bird 600 g. The males also have a thicker bill and tarsus than the females (Luthin 1983a), and are 10% larger in bill and tarsus (C.S. Luthin pers. comm. 1990). These differences are not noticeable in the field. Nestlings are black with a pale grey face and fleshy soft parts. After several days, banding develops on the bill, which, from the differing reports, may be variable. The bill is reported to be dark with a pink tip or several pink bands (Luthin 1983a), and to be pinkish with a dark tip (Belton 1984, B.T. Thomas pers. comm. 1990). The facial skin above the bill is grey (or even bluish—Luthin 1983a) with a white band, the latter being very striking within the first few days but fading to grey by 1 week (B.T. Thomas pers. comm. 1990, regarding berlepschi). The bill loses its banding by fledging, although the light patch in front of the eyes is retained. The plumage lacks the metallic sheen of the adults, and the birds have noticeably shorter bills (Dugand 1947). The Barefaced Ibis is characteristically a silent bird outside the nesting period, having a weak, gobbling call rendered 'Ghew-ghew-ghew' or £Cu-cu-cu-cu' (Sick 1985, Hilty and Brown 1986). Hudson (1920:124) described the voice as little sigh-like puffs of sound, hence the name he gave it, the Whispering Ibis. As Hudson noted many years ago, the

The Barefaced Ibis occurs in the tropics and parts of the subtropics of South America east of the Andes, from Colombia to Argentina, excluding the Amazon River itself. This hiatus creates disjunct range-segments on either side of the Amazon. The overall range north of the Amazon is from northeastern Colombia, and Venezuela (eastern, central, coastal), Guyana, Surinam, and northwestern Brazil (Rio Branco). South of the Amazon, the range is from central Brazil, to Bolivia, Paraguay, Uruguay, and northern Argentina south to Buenos Aires and Cordoba. Occupying the species' range north of the Amazon, berlepschi is the more northern subspecies. Nudifrons occurs in higher lands of Brazil from central Mato Grosso, Roraima, Maranhao, east to Piaui and Bahia, southward to Sao Paulo. P.i. infuscatus occurs from Rio Grande do Sul (Belton 1984) to the Chaco of eastern Bolivia (Beni, Santa Cruz), through Uruguay into northern Argentina to Tucuman, Cordoba, Santa Fe, and northern Buenos Aires (Olrog 1963, D.A. Scott and M. Carbonell 1986). Delimitation of the ranges of the two southern subspecies is worthy of further clarification. They have been thought to be contiguous, but Belton (1984) suggests that the northeastern littoral of Rio Grande do Sul may be a hiatus separating the southerly infuscatus from the more northerly nudifrons. Populations undertake seasonal migrations (B.T. Thomas pers. comm. 1990). Movements in the llanos relate to the rainy season. Barefaced Ibises are most common there in the breeding months and are less frequently seen in the dry season, January to June (B.T. Thomas 1979b, E. Aguilera pers. comm.). They arrive in the llanos soon after the rains begin and initiate courtship immediately. Belton (1984) observed, similarly, that there may be some movement out of Rio Grande do Sul in the cold season.

160 Barefaced Ibis

Populations can be rather substantial locally. It is one of the most numerous species in the pantanal of Mato Grosso, but is not found commonly in some other locations in the region (Dugand 1947, Sick 1985). Similarly, it was the most locally abundant ibis in the Paraguayan Chaco (Brooks 1991). Ramo and Busto (in press a) found them nesting in 7 mixed colonies in Guanare, in the Venezuelan llanos. Nesting numbers per colony are low: in 1980 numbers were 22, 50, 6, 15, 100, 5 and 20 in the colonies censused. C.S. Luthin (pers. comm. 1990) found nesting groups of similar sizes. ECOLOGY The Barefaced Ibis is a bird of open marshes, savanna, wet meadows and fields. We studied its feeding habits in the llanos, where it forages on moist ground, shallow water, marshes and puddles, in rice fields, along streams and ditches, and on the shore of lagoons and ponds (pers. obs. J.A.K.). Along the coast, it feeds in tidal flats. It also uses damp and drier sites, such as newly mowed fields and flooded pasture (Sick 1985, Belton 1984). Although primarily a bird of the lowlands, it has been observed at 2600 m near Bogota, Colombia (Fjeldsa and Krabbe 1990). It feeds by Walking Slowly on mud, moist land, or in shallow water, while Probing with its bill into the soil or shallow water. Owing to its relatively long bill, it can Probe rather deeply in the water or soft mud but also Pecks rapidly at the surface. We have observed them to feed very much like the American White Ibis, by actively walking along while Probing every 1-3 steps (pers. obs. J.A.K.). Probing sites are typically at the edge of puddles and muddy spots. The Barefaced Ibis typically feeds gregariously, often in small flocks of a few birds but occasionally in flocks of up to several hundred (Naumburg 1930). Larger single-species flocks, as are not uncommon in Scarlet or Whitefaced Ibises, have seldom been reported. However, one of us has observed them feeding commonly among other wading birds in rather large mixed aggregations (pers. obs. J.A.K.). In Mato Grosso they commonly form mixed flocks with Sharptailed, Glossy and Whitefaced Ibises (Naumburg 1930, pers. obs. J.A.H.). In the llanos, we have seen them feeding with Scarlet and Sharptailed Ibises especially, and also with cattle (pers. obs. J.A.K.). In such cases they are brought into interaction with other species. The Scarlet Ibis, for example, steals prey from their Barefaced feeding associates (Kushlan et al. 1985). The Barefaced Ibis has been documented to eat soil invertebrates, such as worms (probably annelids) and beetle larvae, as well as small crabs, fish, molluscs, including freshwater clams, and aquatic insects (Olivares 1969:80, Belton 1984, R. Cintra pers. comm., G. Morales pers. comm. 1990). Sick (1985) has found seeds and leaves among stomach contents. BREEDING Barefaced Ibises nest in dense bushes, in colony sites that are reused in successive seasons. They generally nest in small groups within larger mixed-species colonies contain-

ing cormorants, spoonbills, night herons, Maguari Storks and other ibises. They also nest in small single-species colonies of a few to 10 nests (Belton 1984, C.S. Luthin pers. comm. 1990). Luthin (1983a) considers the species to nest in loose aggregates, somewhat intermediate between colonial and solitary nesters. They nest in the rainy season, with some variability being attributable to annual differences in the rainfall timing. Typically, they begin nesting in the llanos in the rainy season from May to late October, and in Brazil in October and November (B.T. Thomas 1979b, Luthin 1983a, Belton 1984, E. Aguilera pers. comm.).. Nesting appears to be prolonged and continuous through the season, as large young can be present simultaneous with nest-building in the same colony (Luthin 1983a, Belton 1984). They appear to respond quickly to onset of the rainy season, commencing nesting within 2 weeks of the beginning of the rains (C.S. Luthin pers. comm. 1990). Courtship may be interrupted by windy conditions. Pair-formation was studied by Luthin (1983a) in the llanos, and most information on the process is derived from this important study. The males gather in display groups in variable locations with the composition of the display party changing frequently. Fighting among males seems to be minimal. Erect Postures and Display Shakes occur as other males approach too closely. There are occasional Sparring and Supplanting displays, but these are not common. The birds display more often in the mornings and late afternoons, departing the display grounds in midday except during the peak of the courtship period. Birds arriving at the display area, land with a characteristic 'whoosh' of their wings. Luthin reported seeing both sexes displaying. The most common behaviour is Display Preening, directed to all parts of the body, most often to the back with the wings held open and one of them drooped. The feathers on the neck and back may be raised a bit, and the red skin of the gular patch is often flared. During this display, the bird may also perform a Head Rub, touching, holding or rubbing the top of its head to its back. Both members of a pair may Display Preen together, or one may lead and the other follow. Luthin reported that Display Preening by the male is more vigorous and exaggerated, and the female seldom droops her wings. Allopreening between the members of the pair appears to be rare. The female will use a Stick Shaking display when she lands next to the displaying male. This is followed by a Stick Shake by the male. It appears that this is primarily an appeasement display. A pair may engage in duetting, which becomes more synchronized as the pair-bond develops. This is often followed by Stick Shaking. Pairing birds often stand erect, side by side, with the males standing taller. An Erect Posture is assumed in response to disturbance. Display and nest-sites are defended with a Sparring display, accompanied by a high intensity multiple-hoot and extension of the gular area. At this stage, fights may occur between displaying males. To displace another bird, an attacking male will use Supplanting display, in which it flies towards or drops down on the other bird with feet and wings

Barefaced Ibis

extended. A male encroached upon by another may raise and shake the feathers of his back and neck. Luthin (1983a) called this a 'Ruffle display' but it may be considered a variation of the Display Shake. The nest-site is selected after initial pair-formation. The new pair moves from the highly visible display perch to a more suitable nest-site within the canopy, where pairformation displays and site defence are continued. At this point, defence may be quite aggressive, including Sparring displays, with the body tipped towards the foe, wings spread and feathers raised. In this posture, a bird will walk towards the intruder and may even chase it away, following it from his nest site. Females give a variation of the Sparring display directed towards intruding birds, while remaining in sitting position on the nest. This posture, which shows the red chin patch, is held for as long as the intruder is present. Luthin observed that this intense territorial behaviour keeps closely synchronized birds from nesting near each other. After nesting gets underway, a pair allows establishment of new nests nearby, so neighbouring nests are usually in different stages of nesting. Early copulation takes place at the display site, later copulations at the nest site. Copulation is preceded by Stick Shaking and Bill Shaking, which in this species is more appropriately called Bill Swinging. The male comes up to the female, and may put his head over hers, Stick Shakes, and then begins to slowly swing his head back and forth bumping the female's bill. Upon placing a foot on the female's back, the male continues this Bill Shaking (Swinging) by taking the female's bill in his. During copulation both sexes raise their feathers and flare their gular sacs. Copulation may be followed by tail wagging or ruffling the body feathers. Promiscuous matings are common, perhaps a reason why nesting birds so strongly resist the establishment of neighbouring nests until after incubation begins. C.S. Luthin (pers. comm. 1990) found one such event occurring when young were in the nest. These matings appear to be forced. The male gathers material for the nest, which is built by the female. Early twigs are used for Stick Shaking rather than nest-construction. Later, both birds simultaneously Stick Shake as the twig is handed from the male to the female. Nest-construction may continue into incubation. The nest is placed rather inconspicuously deeply within the branches of thick, often thorny, shrubs and trees usually over water (C.S. Luthin 1983a, pers. comm. 1990). The nest is a rather small, flat platform of sticks, 30-35 cm wide (Meyer de Schauensee and Phelps 1978:23). Eggs are blue-green and unmarked (B.T. Thomas pers. comm. 1990). Nests with 2-6 eggs have been observed in the llanos (Luthin 1983, Ramo and Busto in press), 3-4 being most common; the average is 3.8/nest in the Venezuelan llanos (Luthin 1983a). These eggs averaged 47.3 X 34.5 mm

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in size. Luthin (1983a) suspects that egg dumping may occur, as suggested by the appearance of additional eggs rather late in incubation. Both parents incubate, over a period of 21-23 days, and attend the nest until the young are about 1 week old. In addition to the threat of intruding ibises, eggs are at risk to predators. For example, eggs are eaten by the boa (Constrictor constrictor), the tegu lizard and predatory birds (B.T. Thomas pers. comm. 1990, Luthin 1983a). Luthin observed that adults did not interfere with the lizard eating their eggs. The threat of predation and lack of nest defence undoubtedly accounts for the secluded positioning of the nest. Exchanges at the nest occur up to 6 times per day and are accompanied by a Greeting display in which the incubating bird gives a long-hoot while the bill is pointed upward and the head back, followed by Stick Shaking. When the relieving bird arrives, both duet and display together. Males may also continue to supply twigs. Later in nesting, exchanges include less display. Young hatch asynchronously and are altricial. After several days all chicks appear to be the same size, so brood reduction may not be common. Young are silent, except for the light buzzing call given at feedings by older nestlings (C.S. Luthin pers. comm. 1990). Like the eggs, nestling are at risk to predation, and the young will leave their nest in response to disturbance. They depart the nest permanently in 22-27 days and fledge at 27-30 days (Luthin 1983a).

TAXONOMY Based on his observations of behaviour, C.S. Luthin (pers. comm. 1990) concluded that the Barefaced Ibis appears to have no close relatives. Resolution of the question of its relationships to other ibises will require additional study. Similarly, additional study is needed of the named subspecies. The highly variable soft-part colours, especially, need to be more carefully evaluated, as do the range and degree of differentiation between the two southern subspecies.

CONSERVATION The Barefaced Ibis is widespread, locally quite common and under no specific threat. However, a lack of detailed information on its habitat needs, food, and migratory and dispersal strategies would hamper the development of conservation plans, should these become needed locally.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 307.

Glossy Ibis Plegadis falcinellus (Linnaeus) Tantalus Falcinellus Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 241; based on 'Numenius rostro arcuarto' of W.H. Kramer, 1756, Elenchus Veg. Animal. Austriam Inferiorem Observatorum, p. 350, and 'Le Gourly verd' of Brisson, 1760, Ornith., 5, p. 326, fig. 2 (immature): Austria Italia ( = Italy) Other names: Black Curlew (English); Corocoro Gastano, Coco, Coco Prieto, Coco Oscuro, Ciguena, Morito, Coclito Moro, Coclito Moreno (Spanish); Ibis Falcinelle, Pecheur, Ibis Noir, Pecheur (French); Sichler, Brauner Sichler (German); Zwarte Ibis (Dutch); Mignattaio (Italian); Tambala Dahakatti, Kokka (Sinhalese); Karuppu-Kottan (Tamil); Voromainty, Renikato (Malagasy); Karavaika (Russian)

INTRODUCTION The widespread Glossy Ibis is a small dark ibis, standing 56-63 cm tall and weighing 500-800 g. It is characterized by having a striking bronze green tinge to its otherwise dark plumage. During the non-breeding season, the adult's head and neck are sooty brown with thin white streaking originating from white-margined feathers. A shiny chestnut gloss is particularly obvious on the feathers of the back, wings, tail and front of the head. The flight feathers are greenish black. The exposed skin of the face is blackish purple, the iris is dark grey to brown, and the bill dark brown to blackish. The legs are dark brown. The sexes are similar in plumage coloration, but the male is larger in all body measurements. The male's total length averages about 63.5 cm, contrasted with 55.3 cm for females (Dementiev and Gladkov 1951). The differences are so noticeable that they can be used as field marks. A partial pre-nuptial moult occurs, and, during the courtship season, the plumage increases in radiance. The sheen becomes a rich chestnut on the neck, mantle, shoulders and underparts, and purple and green on the head, upperparts, tail and wings. The lores remain dark brown, but the face turns pale blue at courtship, fading to dark purple thereafter. A distinctive white to pale blue line develops on the skin bordering the feathering of the forehead and side of the face but does not extend behind the eye or under the chin. A complete post-nuptial moult occurs prior to winter. The nestlings are sooty black. They have a red patch of skin on the crown, which fades over the first week, as well as an accompanying patch of white feathers on the head and throat, the white differing in extent among individuals. The bill is pink with a variable amount of black banding. The lores, edge of mouth, legs and feet are pink; the iris is greyish brown. The juvenile plumage is dark dusky green, but not glossed to the extent of the adult and without chestnut overtones. The head and neck are greyish brown, mottled with white on the face and streaked with white on head and neck; the small white patches above, on the crown, fray off rapidly. The undersides are brownish, with a faint greenish bronze tinge. The bill, initially pink, gradually darkens, starting at the tip. Legs and feet become a dark olive-grey, the iris a darker brown. With moult, the plumage becomes more like the adult's, acquiring a full sheen over the first 2 years. The Glossy Ibis has a call similar to the American White Ibis, a sheep-like 'Huu-huu-huu-huu'. An adult uses a guttural 'Graa-graa-graa' when approaching the nest and engages in mutual cooing sounds during courtship. The flight is rapid wing-beats interspersed with gliding in small tight flocks or in elongated V flights. Birds often fly long distances to and from feeding grounds. They may be seen in acrobatic manoeuvres,

164 Glossy Ibis

rising high and then falling out of the air with their legs dangling. The Glossy Ibis looks black from a distance and can be confused with similar dark birds under such conditions. It might at a distance be confused with a curlew, but the Glossy Ibis is darker, larger, carries itself more upright, and has longer appendages. Adults are not safely distinguished from the Whitefaced Ibis at a distance in the field, even during the nesting season when both birds have white bands on their forehead. However, the Glossy Ibis is larger; its facial skin is bluish white rather than reddish; its legs are dark rather than reddish; and the white, facial line is the result of skin pigmentation rather than feathering as in the Whitefaced Ibis, nor does it extend behind the eye as in the latter species. Juveniles and wintering birds of the two species may not be distinguishable in the wild at all, although juvenile Glossy Ibis have darker upperparts than juvenile Whitefaced Ibis. The Glossy Ibis might be confused in the field with several dark South American ibises. However, it is larger and slimmer than the crested Green Ibis and has darker, longer legs than the Barefaced Ibis. Juvenile American White Ibises have lighter undersides. In many cases, close study is required to distinguish these dark ibises, which may feed together. In Asia, the Black and Giant Ibises are bigger, and the former differs in having white wing patches. In Africa, the species can be distinguished from the Hadada by the latter's duller plumage, jerky flight and distinctive call. It can be told from the other Bostrychia ibises by its lack of a crest and by its glossy sheen. The dark Waldrapp Ibis has a bald head and a neck ruff.

DISTRIBUTION AND POPULATION The Glossy Ibis is the most widespread species of stork or ibis, occurring in parts of Eurasia, Africa, Australia, and North, Central and South America, and on various islands (Keve 1967). In the New World, the breeding range encompasses eastern North America, from New Brunswick, Canada, and Maine through Florida and Louisiana, sporadically inland as far as Arkansas, USA. It has also been recorded as nesting in the Greater Antilles (Cuba, Hispaniola, Puerto Rico), Central America (Palo Verde in Costa Rica), and northern South America in Venezuela (Aragua, Zulia, Apure) (Meyer de Schauensee and Phelps 1978). In Eurasia, it breeds particularly in southeastern Europe through Russia, in northeastern Italy, Austria, Yugoslavia (Sasko Lake), Albania, Greece, Bulgaria, Romania, Moldavia, the Ukraine, southern Russia (Kuban and Terek Rivers), Armenia, Turkamen, Kazakh, Turkey, Israel, Iraq, Iran, Pakistan (Sind), India (Punjab, Uttah Predesh, Orissa, Madhya Predesh, Manipur, Assam), Bangladesh, Burma, Vietnam (Minh Hai), Indonesia (West Java, Kalimantan, Irian Jaya, probably Sulawesi), the Philippines (Mindanao, Luzon), and northern and eastern Australia. In Africa, it nests locally in Tunisia, Morocco, Mali, Kenya, Tanzania, Zambia, Zimbabwe, Botswana, South Africa, Namibia, Angola and Madagascar. The species is highly nomadic and shifts its nesting sites quickly. As a result, it has been documented as breeding in only a small portion of its overall range, sometimes for only a few years in any specific location. It also nests in relatively

Glossy Ibis

inaccessible areas such that ephemeral colony sites can be overlooked easily. In any one year, it undoubtedly nests in more places and over a wider area than is documented. The nesting range of the Glossy Ibis has changed markedly in the present century. In North America, the breeding range has greatly expanded. Although known from 1817, it was first found breeding on that continent in the late 1800s. The known nesting population in Florida grew to 1200 pairs over the next quarter-century, after which it declined while simultaneously expanding its range elsewhere in the USA: it nested in Louisiana in the 1930s, in Virginia and New Jersey by the mid 1950s, New York and Arkansas by the 1960s, Maine in 1972, and New Brunswick, Canada, in 1986 (Semple 1932, Sprunt 1933, Bock and Terborgh 1957, Hailman 1959, P.W. Post 1962, Hanebrink and Cochran 1966, Finch 1973, McAlpine et al. 1989). Populations apparently began expanding and increasing in northern South America by the mid 1960s (Gochfeld 1973). Expansion also may have occurred in Africa and the Middle East, with nesting first reported in South Africa in mid 1950s and Israel in 1969 (Paz 1987). It has recently been reported to nest in Pulua Dua, West Java (Milton and Marhardi 1985). In other areas, the range has contracted markedly, especially in western Europe and North Africa. It no longer nests in southern Spain, southern France, Sicily, Austria, Morocco, northern Algeria or Egypt (Cramp 1977, Pineau and Giraud-Audine 1977). Previously it was also reported to nest in Sri Lanka and northwestern Mongolia. The species disperses widely outside its more usual range, occurring as a straggler in many additional places. It has, in fact, been seen in most temperate to tropical areas at some time. After the breeding season, it wanders as far afield as Iceland, Bermuda, Azores, Madeira, Cape Verde Is., Sao Tome, Faroe Is., Great Britain, Scandinavia, Germany, Poland east through Tibet, China (Chekiang, Fukien, Kwangton including Hainan), Sri Lanka, Hong Kong, Cambodia, Borneo, Moluccas, New Guinea, New Zealand and Tasmania. In the New World, it wanders to Quebec, Newfoundland, Nova Scotia, the Bahamas (Great Inagua, Cay Lobos), the Cayman Is., Puerto Rico, Jamaica, the Virgin Is., Lesser Antilles (Guadeloupe), eastern Mexico, through Central America (Honduras, Costa Rica, Panama) and northern South America (coastal Colombia and Venezuela). Dispersive and nomadic wanderings turn into a true migration in autumn. In North America, Glossy Ibises from the eastern USA winter in Florida and the Gulf Coast and also move through Florida to the West Indies (Bahamas, Greater Antilles, northern Lesser Antilles), Central America and northern Venezuela. Eurasian birds winter along the Mediterranean in southern Europe and the Middle East through western USSR to Iraq, and Iran through to India, and also in northern Africa from Morocco through the Nile Valley and the Sudan. They also cross the Sahara and follow the Nile valley into central and east Africa. Birds from eastern Europe move either to East Africa or to India. In Asia, ibises winter through India to Southeast Asia, the East Indies, and Australia. It is likely Australian birds winter in large numbers in Irian Jay a (M J. Silvius and

165

W.J.M. Verheugt pers. comm. 1990). There may be migratory movements in Africa as well (Tarboton 1977a). The return north is rather protracted in the spring and many begin to nest after the other birds in mixed colonies. The Glossy Ibis is never abundant though widespread, as individual populations are small and changeable, and in most areas is considered to be locally rare. In Australia, Glossy Ibises wander nomadically and do not use regular nesting sites (K.W. Lowe 1983). Colony sites and groups of colonies tend to number in the hundreds to low thousands of nesting pairs (Luthin 1984a). However, over its entire range the population is considerable. Available information reflects changing local abundances. In Greece, the population has decreased from 1840 breeding pairs before 1970 to 50-71 pairs in 1985-86 (Crivelli et al. 1988). Colonies of over 12 000 birds were found in the Volga Delta in the 1930s, but few remain. Flocks of over 1000 birds are still seen in the USSR. Our observation in northern India suggests that, throughout the Asian region, the Glossy Ibis is now very sparsely distributed, and is absent from many apparently suitable areas, probably due to the lack of well-protected breeding sites. It occurs on Sulawesi in fairly large numbers and in Irian Jaya in flocks of up to several thousand (C.M.N. White and M.D. Bruce 1986, Uttley 1987, MJ. Silvius and W.J.M. Verheugt pers. comm. 1990). Even where it occurs more abundantly, year-to-year variation in breeding numbers may be substantial. Comprehensive censuses along the North American east coast found 12300 breeding Glossy Ibis in 1975 (Custer et al. 1980). The following year, 25% fewer birds were found, demonstrating the rather typical variability in nesting effort and distribution from one year to the next. The present status of the species and nesting populations are unknown in many areas, especially in China, the Philippines, Madagascar, the USSR, West Africa and northern South America. More information is needed in these areas. ECOLOGY Glossy Ibises are marsh birds, using fresh and saltwater marshes throughout their range. They particularly favour lake and river-edge marshes, such as are found in river deltas, and their breeding distribution often follows watercourses. Along the coast, they frequent saltmarshes and estuaries. They are particularly found in irrigated farm fields, especially rice paddies, throughout the range, and they are also attracted to muddy substrate exposed by falling water levels in wetlands and reservoirs. Population shifts have been in response to drainage of river flood-plain, which destroys nesting and feeding habitat, and to development of reservoirs, which provides new nesting sites. The core of the Old World populations is in the USSR, where the species occupies the deltas of rivers emptying into the Black, Azov, Caspian and Aral seas, and islands in reservoirs in that area. The Glossy Ibis feeds in very shallow water, generally in small flocks, often well dispersed. It feeds by Walking Slowly while Probing rapidly into the water and soil, usually

166 Glossy Ibis

in mud away from plant cover or in short vegetation. Few other observations of foraging behaviour have been made. The Glossy Ibis often feeds in mixed-species aggregations of wading birds. When feeding in mixed-species flocks, it is often followed by commensals such as egrets, which appear to benefit by catching prey items frightened into flight by the probing ibis flock (Erwin 1983). In Florida, we have seen them feeding alone in mixed flocks, especially with Snowy Egrets (Egretta thula) and American White Ibises. However, we have also seen them feed alone, especially males. In northern Australia, they feed with Royal Spoonbills and on the fringes of areas where Blacknecked Storks feed. In the Bharatpur Reserve in northern India, they are found only in small numbers and feed solitarily in the shallow marsh edges of pools. The food is primarily invertebrate, including insects and insect larvae (dragonflies, crickets, water beetles, scarabs, weevils, soldierflies, leafminer beetles and flies), worms, snails, spiders, and crayfish and other crustaceans. Less frequently, fishes, frogs, tadpoles and snakes are consumed in numbers, when these become available. The diet varies seasonally, depending upon what is accessible. Tadpoles are eaten when abundant, and also locusts when swarming. BREEDING The Glossy Ibis is primarily a marsh bird, usually nesting in dense reeds near its preferred feeding areas. Secluded lakes, thickets of reeds, river oxbows and islands of vegetation are favoured nesting sites. Reed nesting is probably the primitive nesting habit, but the species also uses a wide range of nest sites including trees, bushes or the ground. Nesting is usually in mixed species colonies with other waterbirds, but Glossy Ibises have also been reported to nest singly (Le Souef 1917). In Florida, we have found them nesting intermingled with the American White Ibis and Cattle Egrets (Bubulcus ibis). In Kenya, we found their nests to be scattered throughout a mixed colony consisting mainly of egrets. In Australia, 1000 Glossy Ibis have nested with 40000 Strawnecked Ibises, 1000 Oriental White Ibises, Yellowbilled Spoonbills, Royal Spoonbills and other wading birds (K.W. Lowe 1983). They also nest with cormorants and pelicans. The nesting season is generally in the spring, May to August in eastern Europe and eastern North America, and earlier further south. In southern Florida, we have found them nesting as early as March, but even here they are more usually associated with summer nesting colonies. Such variability in nesting timing is not unusual. In Africa, on the Tana River, Kenya, we counted 50 pairs nesting in the winter rains and at the Garsen Colony Site, we found 60 pairs nesting in June of the same year. Courtship behaviour is virtually undescribed (Baynard 1913a,b). After pairing, a Greeting display continues to be given throughout nesting. It involves considerable mutual bowing, rubbing of bills together, and guttural cooing vocalizations. Prior to nest-relief, the incoming bird gives a call some distance from the nest and is answered by the mate. Later in nesting the incubating bird may leave at the call,

just prior to the arrival of the relieving bird without interaction. The nest is built by both birds and both collect material. However the male brings most of the nesting material. Ibisbuilt nests are relatively bulky for their size, averaging 3040 cm wide, and 4-27 cm thick. They are made of reeds or twigs and lined with green leaves or grass, which are continually added throughout nesting. Within a colony site, the birds choose to nest in the denser available vegetation and to be rather dispersed throughout the colony, as noted above. Most typically, they place their nests low to the ground, especially on marsh vegetation or even on the ground. In Kenya, we found the nests built in heavy-foliaged trees and tall bushes, near and beneath the nests of African Spoonbills and African Openbill Storks. In Australia, they have been reported nesting on the trunks and branches of knocked-down lignum trees (K.W. Lowe 1983). They also reuse nests built by egrets, and these are positioned higher than the nests that the ibis build for themselves (Burger and Miller 1977). The eggs are deep, solid blue, fading to a lighter blue during incubation. The clutch size is 2-6, averaging about 3, and egg dumping may occur (Jozefik 1960b). Eggs average 53 X 36mm in size and weight 31-39 g (K.W. Lowe 1983). Egg-laying within a colony may be spread over 5 or more weeks (K.W. Lowe 1983), and there is evidence of renesting within a breeding season. Incubation, by both sexes, takes 21 days, the larger male taking the daytime shift. Chicks hatch asynchronously, and a substantial brood reduction occurs, with an average of 1 or fewer chicks fledging per nest in most studies, the reduction being attributed to starvation and predation (L.M. Miller and J. Burger 1978, K.W. Lowe 1983). Both adults care for young, one parent being present during the first week. We found nest-relief to be done furtively, with the relieving bird creeping along branches well hidden until the change-over occurred. The young are fed by regurgitation and are not as aggressive as American White Ibis chicks. Young beg by touching the parent's bill, which inspires regurgitation: as the chick grows, it will place its entire head into the parent's gullet. Chicks are able to move from the nest in 2 weeks to form creches in nearby branches or on the ground, but continue to be fed at the nest site. They are surprisingly docile and are with difficulty described in any other way than Affectionate'. They fledge in 25-28 days. Depending on the colony, they may stay in the colony for another 3 weeks or fly to the feeding grounds with the parents soon after fledging. Young fledge at less than adult size, which is not attained for 6 months. TAXONOMY The Glossy Ibis is closely related to the Whitefaced Ibis. The two species can interbreed in captivity and are sometimes considered to be conspecific (Parkes 1955, Palmer 1962). However, the two forms nest sympatrically in Louisiana and Alabama, USA (Duncan and Johnson 1977), and so appear to be specifically distinctive. The unusual situ-

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ation of having such closely related forms nesting together in North America is best understood by noting the lack of subspecific differentiation between Old World and New World populations. We believe that the Glossy Ibis is a natural invader of North America, perhaps within the historic period. The Glossy Ibis is often divided into two subspecies with P. f. peregrinus being the form found in Australia, Indonesia, and Madagascar. It was described as having smaller wings and tarsus, but this distinction has not held up to closer scrutiny (Amadon and Woolfenden 1952, Vaurie 1965). Although we can reject this subspecies, the apparent uniformity of the Glossy Ibis over such a wide range deserves careful study.

nikov et al. 1975). The situation in the southern USSR is typical. Populations there decreased due to development and then stabilized due to the creation of reservoirs and protection breeding sites (Oleinikov et al. 1975). In the Volga Delta, the high populations of 1930s have more recently declined to a single pair by the 1960s (Cramp 1977). The actual status of this species is very poorly known owing to its tendency to shift nesting sites so frequently and quickly. Apparent population decreases in one area are sometimes found to correspond to increases in other areas. Perhaps the most important contribution to the conservation of this species is to conduct additional studies of its biology, movements and population structure, and to protect the reedmarshes on which it depends.

CONSERVATION In the Old World, nesting and feeding sites are threatened by drainage and development of wetland habitats, and nesting sites are enhanced by reservoir development (Olei-

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 308.

Whitefaced Ibis Plegadis chihi (Vieillot) Numenius chihi Vieillot, 1817, Nouv. Diet. Hist. Nat., nouv. ed., 8, p. 303; based on 'Cuello jaspeado', no. 364, of Azara, 1805, Apuntamientos Hist. Nat. Paxaros Paraguay Rio Plata, 3, p. 197: Paraguay and campos of Buenos Aires, Argentina Other names: Whitefaced Glossy Ibis, Black Curlew, Bronze Ibis, Black Curlew (English); Atotola, Morito Cariblanco Atotola, Acolote, Cuervo, Gallareta, Cuervo de Pantano, Cuervo de Canada, Cuervillo de Canada, Cuervo de Pantano Comun (Spanish); Carauna, Tapicuru, Magarico-preto (Portuguese)

INTRODUCTION The Whitefaced Ibis is a small (46-56 cm, 450-525 g), dark ibis, with glossy purple, green and bronze reflections, and, in the breeding season, white feathers around the bill. This is a striking bird, which, because of the brilliance of its metallic sheen, is often called 'a work of art'. Outside the breeding season, the adult's base colour is a dark maroon-brown. The underparts are deep brown with vinaceous overcast and minor dull dark green reflections on the breast (Belknap 1957). The head and throat are streaked in white. The legs and face are reddish, and the iris reddish brown. The sexes are similar in plumage but the male is larger, heavier and has a longer bill. Bill length can be used to tell the sexes apart in the field, as overlap in size between them is minimal (Belknap 1957). There is considerable variation among individuals, however, and the South American forms may be smaller than their North American counterparts. The full plumage brilliance is achieved in the nesting season following a pre-nuptial moult. This occurs in April and May in North America and August and September in South America (Belknap 1957, S. Narosky pers. comm. 1990). The head, neck, upper back, wing-coverts and undersides assume a deep, dark, chestnut-maroon coloration. The dark uppersides reflect iridescent purple, violet and green. The dark flight feathers are metallic green and bronze, whereas the wings have purplish overtones. The bare skin of the face becomes reddish or purplish. A line of white feathers, variable among individuals, separates the forehead and the face, extending completely around the eye. The chin is also white, and this colour sometimes extends down the throat. The iris is reddish brown. The bill, legs and toes are blackish grey. Complete post-nuptial moult occurs in late summer, July and August in North America. Plumages and coloration of young have been well documented (Belknap 1957, S. Narosky pers. comm. 1990). Hatchlings are covered with a sparse brownish black down, darker and denser on the uppersides than elsewhere (Belknap 1957). The legs and bill are pink orange. The short, straight bill has a single central band, which, like the tip, is lead coloured. The bare crown of the head is pink with cinnamon and white down. The literature suggests to us that there may be more white on the crest of North American nestlings than on South American birds. The legs are black with bluish grey nails having pinkish tips. The iris is a very dark greyish brown. The juvenile plumage begins to develop within 2 weeks. It is basically a shiny dark green, initially retaining traces of down, especially on the head. By the third week, the bare area of the top of the head develops a light orange colour. The head and neck acquire white mottling, and so are lighter than in the adult. The body colour lacks the rich chestnut hue. Upperparts are glossed with greenish highlights;

170 Whitefaced Ibis

Glossy Ibis and Whitefaced Ibis. In the breeding season, the two species are distinguished by the difference in plumage sheen, bare-skin colour, and the character of the eye-line. However the latter is not a reliable feature under most field conditions, because the Glossy Ibis has a similar white line of skin during that time of year. The white feathering of the Whitefaced Ibis does extend all the way round the eye, which the white skin patch of the Glossy Ibis does not. The otherwise dark juvenile American White Ibis has white undersides. The glossy sheen and facial coloration will also distinguish this ibis from other dark ibises in its South American range. The Whitefaced Ibis is often seen, and can be confused, with the Barefaced Ibis, which is dark, less glossy, and has a pale face and bill (S. Narosky pers. comm. 1990). In flight the two are easily told apart, as the legs of the Whitefaced Ibis extend well beyond the tail whereas those of the Barefaced Ibis cannot be seen. The Puna Ibis is distinguished by its black undersides (Sick 1985).

underparts are lighter, with faint purplish or greenish sheen, depending on the light. The iris is brown, and the legs and bill are black. The bill is a whitish pink, with black rings at the base, middle and tip. Newly fledged young are smaller than adults and have noticeably smaller and less downcurved bills. The bill stripes may be retained into the first winter, during which the bill darkens. The post-juvenile moult begins in late summer and fall, the birds developing a brownish plumage with purple-chestnut sheen and the streaked brown and white neck of the wintering adult. The Whitefaced Ibis flies strongly, quickly and gracefully, alternating bouts of quick flapping with gliding. It flies with its head and neck completely extended, appearing black from a distance. However, at closer range or when the sun hits the birds, the bronze and green sheen become obvious. They fly in small to large flocks, often in ever changing' V or linear formations. Flocks can number in the hundreds, and dusk roosting flights can be spectacular. These birds also soar, especially on migration, and can engage in spectacular aerial manoeuvres, in which they descend rapidly in a twisting path, falling nearly to the ground, before rising again. When in flight or on the ground, the usual call is a loud nasal (Ka-onk, ka-onk, ka-onk', which Hudson (1920) likened to human laughter. In courtship, both sexes give repeated 'Geek' and other calls. In the wild, it is difficult or impossible to distinguish between juveniles and some wintering individuals of the

DISTRIBUTION AND POPULATION The Whitefaced Ibis has an exceptional breeding distribution with two entirely disjunct nesting populations, one centred in the Great Basin of western North America and the other in the pampas of central and southern South America. The reasons for this odd distribution are unclear (R.A.Ryder 1967). The northern population is concentrated in the Great Basin states of the United States and Canada, including the eastern Oregon, southern parts of Idaho, Wyoming, Montana, North Dakota, Alberta and Minnesota, east to Alabama, west to central California (San Joaquin Valley), and south to New Mexico, Texas, Louisiana, and Colima, Zacatecas, Veracruz and Tabasco, Mexico (R.A. Ryder 1967, Steffen et al. 1978, Skaar 1975, Salt and Salt 1976, Lokemoen 1979, Portnoy 1981, Ivey and Severson 1984, Findholt and Berner 1988, Stahlecker 1989, D.M. Taylor et al. 1989). The southern population nests in eastern Bolivia, Paraguay (Tebicuari), Uruguay, Brazil (Mato Grosso, Sao Paulo, Rio de Janeiro, Santa Catarina, Rio Grande do Sul), central and southern Chile, and south to Argentina (Los Yngleses, Ajo, Buenos Aires, Rio Negro). A disjunct population occurs in coastal Peru. Even within this larger range, breeding populations are quite localized. The Whitefaced Ibis is, in fact, transient over a large part of its overall breeding range. The small Chilian population has apparently not nested in the last decades (Schlatter pers. comm. 1990). The northern population is migratory. They disperse after nesting to various feeding locations, some of which they remain at until fall (Kotter 1970, D.M. Taylor et al. 1989). Wintering birds also are found widely from southern California, Colorado, Arizona, coastal Louisiana, Texas, Baja California and Mexico. Most of the wintering population appears to be focused in Mexico, including Jalisco, Michoacan, Nayarit, Guanajuato, Queretaro and the Federal District. Wintering birds also have been reported from further south in Central America, in Guatemala, El

Whitefaced Ibis 171

Salvador and Honduras, but considering the difficulty of identifying the species, additional information is needed on this point (Monroe 1968:58). According to S. Narosky (pers. comm. 1990), the southern population in Argentina does not migrate; this conclusion is contrary to that of Hudson (1920). Juvenile birds in South America do disperse after nesting, however. Banded young have been recovered as far as 1800 km from their natal colony, and have moved as far as 1400 km within a few months (Olrog 1975, Sick 1985). Individuals also wander, especially prior to nesting in response to dry conditions. Birds have been found as far north as southern British Columbia, Alberta, Saskatchewan and Manitoba, as far east as Ohio, New York and Maryland, as far south as Florida and Costa Rica, and as far west as Hawaii. In South America, stragglers are reported as high as 4300 m in the mountains of Bolivia (Fjeldsa and Krabbe 1990) and as far south as the Straits of Magellan. We consider that the previously accepted records of Whitefaced Ibis nesting in Venezuela and Colombia (e.g. American Ornithologists' Union 1983) refer in fact to the Glossy Ibis (Gochfeld 1973). The single nesting record from Florida appears to be valid (Brewster 1886, Parkes 1955), but we conclude that it was an isolated incident. These locations are therefore not included as part of overall range of the species. The Whitefaced Ibis is a common species and can be abundant locally. However, nesting colony sites are widely dispersed within its overall range. In the mid 1960s, North American nesting populations decreased precipitously due to pesticide contamination, especially from feeding in agricultural fields. At that time, R.A. Ryder (1967) estimated 10000 breeding pairs remained in the USA. The Texas nesting population decreased from 9200 pairs to 2100 pairs, from 1969 to 1974 (K.A. King et al. 1980). However, populations remained high in most of the species' southern range. They are abundant on the pampas of Argentina (Hudson 1920); Weller (1967) reported 15000 nests from San Antonio, Argentina, and Belton (1984) reported 12000 nests in Rio Grande do Sul, Brazil. In the marshes of Rio Saladillo, Cordoba, Argentina, 20 000 wintering Whitefaced Ibis were reported (D.A. Scott and M. Carbonell 1986:20). This species has recently been increasing in North America, reclaiming and surpassing its historic breeding range, north to Oregon and Canada and east to Alabama. By the mid 1970s over 6000 pairs of nesting Plegadis ibises (almost certainly this species) were counted in Louisiana, and another 5300 pairs occurred in Texas (Portnoy 1981, K.A. King et al. 1980). In 1985, 5000 breeding pairs were studied at a single colony in Nevada (Bray and Klebenow 1988). They are known to nest as far south as the Tampico lagoons (100 pairs) and the Tabasco Lagoons (500 birds counted) in Mexico (D.A. Scott and M. Carbonell 1986). ECOLOGY A marshland bird, the Whitefaced Ibis is found in fresh and saltwater marshes and wet grassy areas, along ponds, rivers

and reservoirs, and in open grassland. They feed in water less than 20 cm deep, in damp meadows, and on open dry grasslands. In recent years, they have accommodated to flooded pastures and irrigated fields, especially rice fields. In the Great Basin, they feed often in agricultural fields, mostly in puddled water in recently tilled and irrigated alfalfa fields (Bray and Klebenow 1988). They can travel considerable distances to feeding areas, over 30 km from colony sites in Argentina (Weller 1967). They are, however, quite dependent on rainy conditions to provide foraging habitat, and nesting success depends on weather, failing in drought years and succeeding in wet years (R.A. Ryder 1967). Whitefaced Ibises feed actively by Walking, slowly to quickly, while Probing rapidly, usually at each step. Periodically they stop and Stand while Probing at the base of aquatic plants. They also Head Swing while standing still in open water, vibrating their mandibles rapidly (Belknap 1957). The species is highly gregarious, probably more so than the Glossy Ibis. Flocks can exceed 1000 birds (Belton 1984). Flocking, feeding birds are often reported to engage in Leapfrog-feeding, in which the rear birds of the flock hop to the head of the moving flock. Birds in the flock are often subjected to piracy by other birds, such as grackles (Quiscalus spp.). Birds fly to and from feeding areas and roosts in flocks, and they roost together in trees or on the ground in marshes. They feed on small fish, frogs (Hyla), crayfish, snails, bivalves, earthworms and aquatic insects such as whirligig beetles, beetle larvae, midge fly larvae and pupae and dragonfly nymphs (Belknap 1957, Centro Editor de America Latina 1983a). In Nevada and Utah, they feed primarily on earthworms but also on dragonfly larvae, horsefly larvae, beetles caterpillars, and other insects, snails and spiders (R.A. Ryder 1967, Bray and Klebenow 1988). In Argentina they feed with gulls on flies and fly larvae associated with livestock carrion (Weller 1967). BREEDING Whitefaced Ibises are highly conspecifically gregarious, nesting in large numbers in compact colonies, usually in marshes. This species preferentially nests in herbaceous marsh vegetation, although it will nest on short bushes, on dry land and on spoil islands. The most common nesting habitat throughout its range is tule marsh (Scirpus spp.), as well as common cane (Phragmites) inland and cordgrass (Spartina spp.) along the coast. In North America, it is likely that the use of high ground on spoil islands may increase as natural marshes become less suitable. This species changes its nesting locations frequently, causing considerable year-to-year shifts in breeding areas. In part, such shifts are due to changing availability of nesting and feeding habitat in flooded shallow marshes and wet meadows. It is likely that birds return to previously used nesting sites, but if conditions are unfavourable begin to search for more appropriate conditions, becoming attracted to waterbird colony sites elsewhere. This sugges-

172 Whitefaced Ibis

tion is supported by the fact that colonies are unsynchronized, suggesting independent arrival of breeders. Typically, the Whitefaced Ibis nests in the spring, April to May in North America, and November to December in South America (Belton 1984, S. Narosky pers. comm. 1990). However, they can delay breeding, for example to mid June in the USA, if conditions are adverse (Belknap 1957), or may even skip years in nesting. Because nesting is often not highly synchronized, large chicks and eggs may be present at the same time. Early courtship has not been recorded in detail. The male displays from several potential nest-sites, especially from abandoned nest-platforms. He may also attempt to displace other birds from their platforms using Bill Thrusts and Supplanting Flights (Belknap 1957). At the display site, the male uses a ritualized bill probing, and gives a 'Hunk-hunkhunk' call, which the female answers. At a potential nestsite both birds preen, perhaps Display Preening. The actual nest-site selection is by the female (Belknap 1957). The nest is built on top of grasses and sedges, which are bent over to support the structure. The birds tend to nest relatively low, within 1 m of the water, even when in shrubs. Often, however, the nests end up floating on the water surface or on the ground. In Texas, 86% of nests were found to be on the ground, the rest in the branches of low shrubs (Burger and Miller 1977). After pairing, the male and female spend considerable time preening, perhaps Display Preening, and Allopreening. Initially the female gathers reeds for the nest, which the male guards. However, after construction or renovation begins in earnest, the male gathers while the female constructs the nest. Copulation occurs at the nest. Nest exchange is accompanied by mutual 'Geeeek, geeeek5 calls (Belknap 1957). Usually, although not always, the nest is relatively large: 40 cm across, solid and flat, but with a well-developed cup. Nests are usually made of reeds or other herbaceous vegetation, sometimes with sticks included. The cup is lined with shredded bits of plant material such as reed leaves. Whitefaced Ibises also frequently take over previously built nests of their own or other species, adding to and reconstructing them to meet their own needs. Burger and Miller (1977) found them to be more tolerant of other Whitefaced Ibises near their nest than they were of other species. None the less, they are extremely defensive of their nest and an area of up to 1m around it. The bird defends with Bill Thrusts and Supplanting Attacks, occasionally landing on an opponent's back, pulling its feathers or striking blows with its bill, while giving 'Gheeee' calls. Eggs are laid at 1-2 day intervals and hatch asynchronously after 20-22 days' incubation. The clutch size is 2-5, averaging 2.6 in Texas, 2.7 in Utah, 3.0 in Louisiana (Belknap 1957, Kotter 1970, K.A. King et al. 1980). The eggs are deep blue to greenish blue lacking any mottling. Utah eggs averaged 52.0 X 36.7 mm and 37.6 g. The young call with a vibrating 'Cheeeeeeeeu' and are fed by regurgitation (Belknap 1957). Parents arriving to feed young give a 'Goick, goick' call. Eggs and young often suffer

heavy losses, especially from heat and predation (Belknap 1957). In Utah, 66% of eggs hatched and 53% fledged. After fledging, young fly with adults to foraging areas, returning at dusk to roost. The young birds can fly in their fifth week and are independent in 8 weeks. First-year mortality is estimated to be 54%. TAXONOMY The Whitefaced Ibis is closely related to the Glossy and Puna Ibises. Palmer (1962) considered the two North American forms to be conspecific. Nesting areas overlap slightly in southern United States, and the American Ornithologist's Union (1983) considers the two forms to be separate species, but part of a superspecies. The specific name has previously been given as guarauna and as mexicanus, but these properly refer to other species (Parkes 1955). CONSERVATION The general decline and contraction of breeding colonies in the 1960s was caused by pesticide contamination. High pesticide levels in various populations have been shown to affect clutch size, eggshell thickness and other aspects of reproductive physiology (Flickinger and Meeker 1972, Capen 1977a, 1978, K.A. King et al. 1978, Capen and Leiker 1979, K.A. King et al. 1980, Steele 1984). Population declines have occured in Nevada, Utah and Texas (R.A. Ryder 1967). In Texas, the population decreased by 77% from 1969 to 1974 because of pesticides (K.A. King et al. 1980). Despite discontinuation of the use of aldrin in rice fields in the USA, pesticides are continuing to be accumulated by Whitefaced Ibises wintering in Mexico (R.A. Ryder 1985, Hennyetal. 1985). In many areas, ibises are used for food, and this species is no exception. An open hunting season existed for Whitefaced Ibis in California until 1915 (Bent 1926). They remain hunted in Mexico and in areas of South America. Chilean populations have apparently been persecuted in the past, wherever colonies were accessible to poachers (Schlatter pers. comm. 1990). Such hunting would be particularly troublesome in areas that are otherwise marginal, such as Chile. In recent years, the species has expanded its range in North America, with nesting records occurring over all of its former range in the USA in places where it had been absent for decades, (e.g. Minnesota, Idaho, New Mexico, and California) and even over an expanded area. In Oregon, numbers have increased from 600 to 2595 pairs during 1980-1987 alone (Ivey et al. 1988). This expansion coincides with a change in pesticide use as well as creation of aquatic habitats by dams and irrigation. Additional information on the status of various populations is needed. The most critical concerns are a reduction, throughout the Whitefaced Ibis' range, in the use of persistent pesticides that adversely effect them and the protection of aquatic habitats, especially marshes. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 309.

Puna Ibis Plegadis ridgwayi (Allen) Falcinellus Ridgwayi J.A. Allen, 1876, Bull. Mus. Comp. ZooL, 3, p. 355: Lake Titicaca, Peru Other names: Andean Glossy Ibis (English); Cuervo de Pantano de la Puna, Cuervo de Pantano del Norte, Cuervillo de la Puna, Cuervillo de Bolivia (Spanish); Make-make, Yanahuiku (South American Indian)

IDENTIFICATION The Puna Ibis is a medium-sized (56-61 cm tall) dark ibis, looking black from a distance, with a dark red downcurved bill. The dark purplish brown base colour of the adult's body is highlighted by deep glossy purple and green reflections above on the lesser wing-coverts, becoming almost black below. The head and neck are dull chestnut. The wings and tail are glossy green, the tail a darker green. The iris is red; the legs and feet are black. In winter, overall coloration is duller, and the head and neck become darker brown with fine white streaking. During the breeding season, the head and neck turn deep chestnut. The species is sexually dimorphic, in that males are considerably larger than females. In 10 males, the culmen averaged 11.7 cm (10.812.7 cm), whereas in 7 females it averaged 9.2 cm (8.6-9.5 cm) (Blake 1977). Chicks are dark brown with banded bills. The juvenile is similar to the non-breeding adult, except that it has more conspicuous white streaks on the neck, extending onto the belly, and lacks much of the adult's sheen. Its iris is brown and its bill lighter and more brownish. The Puna Ibis flies in flocks, sometimes large ones, forming Vs or wavy lines, usually low over the terrain. It is usually silent, but gives 'Wut' and 'Cwurk' calls when disturbed (Fjeldsa and Krabbe 1990). Juveniles give a whispering call. The Puna Ibis is a glossy ibis, although larger than the Whitefaced Ibis. It is also duller and stockier than the Whitefaced, with shorter legs and lacks the band of white feathers around the bill. It is darker and longer-legged than the co-occurring Buffnecked, and does not have the stocky, broad-winged and short-tailed appearance of that species (J. Fjeldsa pers. comm. 1990). DISTRIBUTION AND POPULATION The Puna Ibis is endemic to the Andean Mountains of western South America above 3500 m. It occurs in north-central Peru (Ancash and Junin) southward to the Titicaca basin of Peru and Bolivia, to northern Chile (Arica, Tarapaca). It also occurs very locally in Cochabamba, Bolivia and in northwestern Argentina (Jujuy, Salta) (Blake 1977, Fjeldsa and Krabbe 1990). It is a wanderer and is seen regularly as high as 4800 m above sea level in Peru (J. Fjeldsa pers. comm. 1990). It winters, from May to September, beyond its breeding range as far as valleys at the foot of the Andes, along the Peruvian coast (D.L. Pearson and M.A. Plenge 1974, Hughes 1984, Fjeldsa and Krabbe 1990). It is accidental in La Libertad, north Peru, but it is likely that reports from Central America and Patagonia are misidentifications (J. Fjeldsa pers. comm. 1990). Little is known about population sizes but, overall, they may be considered common. The species can be seen at most Peruvian puna lakes, though usually only in small numbers. However, it can be locally

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for Plegadis ibises, this species feeds socially, sometimes in flocks containing large numbers of birds. BREEDING The Puna Ibis nests in tall reeds in highland marshes. In the enormous reed-beds of Lake Junin, it locates its nesting colonies in places that are quite inaccessible to humans (J. Fjeldsa 1983, pers. comm. 1990). The birds especially use very tall Scirpus reed beds, isolated by trenches of deep water. In other areas, such as on the Puno—Cuzco border, they nest in Juncus-lintd ponds. They may nest in mixed colonies with other waterbirds. The nesting season is somewhat variable geographically (Fjeldsa 1983, pers. comm. 1990, Fjeldsa and Krabbe 1990). At Lake Junin, Peru, nesting is in April to July, with some clutches also in January and February. Nesting is in January to March in Ayacucho, and in November to January and February in the southern parts of the range. Nothing is known about the species' courtship behaviour.

abundant, and in the Lake Junin area of Peru, the population has been estimated to be about 8000 birds (Dourojeanni et al. 1968). However, it may nest only in a dozen or so of the puna lakes of the Andes (J. Fjeldsa pers. comm. 1990).

ECOLOGY This species is characteristic of the high-altitude puna, the temperate zone of the Andes. It may even be found in the higher arctic zones, but it seldom occurs in the subtropical zones except on the Peruvian shores (J. Fjeldsa pers. comm. 1990). Puna Ibises frequent large marshes and damp pastureland, where they forage on muddy flats, along creeks and in short grass (Fjeldsa and Krabbe 1990). They also feed terrestrially, in grasslands quite far from water. They appear to be rather opportunistic, congregating, for example, in places having large numbers of brightly coloured lepidoptera larvae (J. Fjeldsa pers. comm. 1990). The typical feeding behaviour is Probing deeply in the water or mud to extract prey, usually along the edges of denser vegetation (J. Fjeldsa pers. comm. 1990). As is usual

CONSERVATION The biology of the Puna Ibis is almost entirely unknown, and its status throughout the Andes is not well understood. It historically inhabited a large portion of the puna zone, but it is now considered vulnerable in Chile (Glade 1988). Fortunately, a reasonable recovery of the local population has occurred in Lauca National Park (Schlatter pers. comm. 1990). Much of the marshy vegetation of the puna zone, which supports the ibis, is under threat (Fjeldsa 1985a). A combination of human overpopulation, harvesting of tules and cattle grazing have opened much of the marsh. J. Fjeldsa (pers. comm. 1990) reports that some clearing of tules, as has occurred along Lake Titicaca, may be advantageous because it creates feeding habitat for the ibis. However, it becomes harmful when safe nesting areas are eliminated. In many parts of the high Andes, grazing by cattle has completely eliminated all the tall marsh vegetation. It has been suggested that the rarity of ibises in the Bolivian highlands is due to egg-collecting traditions of the Ayumara Indians (Fjeldsa and Krabbe 1990). Lake Junin, one of the primary nesting sites for the Puna Ibis, remains one of the most productive waterbird habitats in the Andes. Proper conservation and management of the lakes and wetlands of the puna zone are essential to the preservation of this endemic ibis. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 309.

Sharptailed Ibis Cercibis oxycerca (Spix) Ibis oxycercus Spix, 1825, Avium Species Novae Itinere Brasiliam, 2, p. 69, pi. 87: in Provincia Para Other names: Longtailed Ibis (English); Tarotaro (Spanish); Trombeteiro, Tara (Portuguese); Groote GlansIbissen (Dutch)

IDENTIFICATION The Sharptailed Ibis is a dark, medium-sized ibis, c. 75-86 cm long, having a conspicuously long tail that has suggested its common English name. The adult's plumage is black overall, usually showing a green glossing. The upper back, wings and tail have a purplish gloss. A fuzzy occipital crest is often noticeable, running along the back of its head and upper neck. The bill, legs, toes, and bare face are orangish red. The throat is yellow-orange, and a feathered strip of grey extends from the lower bill below the eye. The iris is greyish red, but Betsy Thomas (pers. comm. 1990) has observed a bird with scarlet coloured irides; this colour may be associated with breeding. Additional information on soft-part colours, especially during courtship, is needed. The sexes appear to be similar. Although the sizes of museum specimens labelled as to sex overlap, in the wild we have observed what appears to be a size difference between members of a pair, suggestive of sexual differences in size (pers. obs. J.A.K.). Hatchlings appear not to have been carefully described. Juveniles are similar to adults but lack a metallic sheen to their feathers. This is a noisy bird, which has attracted our attention from a distance by its calls, given while perched in trees. Its usual vocalization is a single or doubled 'Cuk' or 'Tur-cuck' that can blend with a bugled 'Tur-dee' call, which gives this ibis its Spanish common name, Tarotaro. The call is similar to but less nasal and raucous than that of the Buffnecked Ibis (Sick 1985, W. Belton pers. comm. 1990). The flight call is a 'Cuk-cuk-cuk-cuk'. In flight the Sharptailed Ibis is relatively slow, and flying appears somewhat laboured. When a bird flies up, it usually goes but a short distance, usually to a nearby tree. This is a quite distinctive ibis, rather larger and stockier than sympatric species. When at rest, its long tail projects well beyond the tips of its folded wings, and, when in flight, beyond the trailing legs and feet. It is larger than the other dark ibises, such as the superficially similar Glossy Ibis, and can be further identified by its reddish face and soft parts. DISTRIBUTION AND POPULATION The Sharptailed Ibis is a lowland bird of the Amazon and Orinoco River basins. It occurs from east of the Andes in Colombia through inland Venezuela, following the Orinoco and Apure rivers to the coast, and again in a disjunct distribution, along the inland portion of the northern Amazon drainage from southwestern Guyana, to northwestern Brazil (Rio Negro), western Brazil (Rio Guapore), and south to northwestern Mato Grosso. The species is very locally distributed within its overall range. Although probably not substantially migratory, it does undertake seasonal population shifts. Both E. Aguilera (pers. comm. 1990) and

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1990) has observed a pair feeding on dry ground by tossing leaves aside before probing in the soft sand underneath. Sharptailed Ibises do not flock, and seldom are seen alone. Rather they typically feed in pairs or in very small groups, usually including 2 adult birds (parents) and fledged young. These young may Beg food from the adults. Little is known about the diet. They appear to eat invertebrates. More information is needed on this point.

BREEDING The Sharptailed Ibis nests solitarily. The nests are placed in trees in gallery forests along streams and in isolated coppices within the grassland. Birds appear to remain paired year-round, and courtship is probably rather perfunctory. In any case, we know of no information on courtship behaviour. Mutual preening occurs between members of a pair (Ogden and Thomas 1985b). In the llanos, E. Aguilera (pers. comm. 1990) found nesting to begin in August and September and to continue through February. This is in the drying season, suggesting a dependence on exposed higher ground for effective foraging.

B.T. Thomas (1979a) found it to be absent from the interior llanos during the dry season (December, January). It can be fairly common and obvious locally, but nothing is known about population sizes. More detailed information on range and population is much to be desired. ECOLOGY This is an ibis of llanos savannas and river-edges of Amazonia, below 300-500 m. During studies in the Venezuelan llanos, we found it to be more terrestrial than the flocking ibises, feeding in relatively short grass and open areas in relatively higher and drier (but still moist) ground (pers. obs. J.A.K.). It generally feeds in water less than 3 cm deep, but it also frequents pools and muddy spots along the edges of ponds and streams in open areas, in short grass, and in shallow water (Kushlan et al. 1985). Food finding seems to be primarily tactile. It usually forages by Walking rather quickly while slowly Probing deeply into the soft substrate. B.T. Thomas (pers. comm.

TAXONOMY The Sharptailed Ibis is generally similar to the other dark ibises, but its taxonomic relationships remain to be discovered through careful study.

CONSERVATION Basic to even considering the conservation of this species is our limited knowledge. Little is known about its basic biology and ecology, and such information is sorely needed. Because it is a solitary bird, its status may decline without anyone's knowledge (C.S. Luthin pers. comm. 1990).

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 310.

Plumbeous Ibis Harpiprion caerulescens (Vieillot) Ibis caerulescens Vieillot, 1817, Nouv. Diet. Hist. Nat., nouv. ed., 16, p. 18; based on 'Curucau aplomado', no 363, of Azara, 1805, Apuntamientos Hist. Nat. Paxaros Paraguay Rio Plata, 3, p. 195: Paraguay Other names: Blue Ibis (English); Curicaca, Curucau Moroti, Bandurria, Bandurria Mora, Vanduria (Spanish); Macarico-real, Curicaca, Guricaca-cinza (Portuguese)

IDENTIFICATION The Plumbeous Ibis is a relatively large grey ibis (71-77 cm long), identifiable especially because of a shaggy crest along the back of its head and neck and a distinctive white bar on its forehead. The basic plumage colour of the adult is grey to bluish-grey, lighter below than above because the dorsal feathers are each broadly mottled with greyish brown. The head is almost entirely feathered, with a very noticeable white band of feathers on the forehead and a less noticeable smaller band at the lower side of the bill. The distinctive nuchal crest, extending a full 10 cm down the back of the neck, is composed of long, grey plumes each having a cream-coloured centre, which in some lights gives the neck a pinkish tinge. These are arranged so that they appear furrowed (Sick 1985, W. Belton pers. comm. 1990). The flight and tail feathers are dark brown to black, glossed with a greenish bronze that can look silvery in the appropriate light. The strong, downcurved, distinctly serrated bill is black. The exposed skin of the lores and chin are grey to black. The legs are a pinkish orange, which brightens in the courtship period (F. Olmos pers. comm. 1990). The iris colour is orangish and has been variously reported as brownish yellow, orangeyellow and orange (Naumburg 1930, Blake 1977, Belton 1984). Wetmore (1926:63) noted that the iris was somewhat lighter on its inner margin. He also noticed in one specimen that its lower eyelid was coloured lilac. Based on our examination of museum specimens the species appears to be sexually dimorphic, with males larger than feinales on average (pers. obs. J.A.K.). Nestlings have a light grey down, reddish skin, a white forehead and a black bill (Cintra 1986). Young have a less noticeable crest, and the white forehead band seems broader (F. Olmos pers. comm. 1990). The Plumbeous Ibis flies with strong, steady flapping with head and neck extended, frequently gliding. The calls are well known. Hudson (1920:123) likened its voice to a dozen strokes on the brass strings of a banjo so big that it could be heard a mile and a half away. He called it the voice of the desolate marshes. Belton (1984:433, Sick in prep, as transliterated by W. Belton pers. comm. 1990) have described its vocalizations. The basic call is a rapid, high, loud Ti-pi-pi-pi'. The flight call is a powerful, distinctive, trumpeting, which has been rendered as 'Kree-kree-kree', 'K-k-kuh-kuh-kee-kee-kee-keekeh-kuh-kuh', and 'Ki-ki-ki, Ghew-ghew-ghew'. These distinctive calls are an indication of the need for long-range communication, particularly between mates (Remsen 1986). The Plumbeous Ibis is rather distinctive among sympatric ibises owing to its thick neck and solid coloration. The Buffhecked Ibis is found in many of the same areas but differs in coloration (F. Olmos pers. comm. 1990, W. Belton pers. comm. 1990).

Plumbeous Ibis 181

Usually the foraging method is Walking slowly, Probing in the mud and shallow water. Birds frequently Probe rather deeply, completely submerging their bills in the process. This is a solitary species, found singly or in pairs and more rarely in small groups of 4-6 birds. It would seem that the larger groups include members of a mated pair, seasonally accompanied by fledged juveniles (F. Olmos pers. comm. 1990). It also may feed in company with Whitefaced Ibises (Sick 1985:179). Brooks (1991) reported a flock of more than 40 birds, but it does not form large conspecific flocks as do some ibis species. It is known to feed on snails (Pomacea), crabs, aquatic insects, frogs, fish, and eels (Sick 1985:179, Cintra 1986, R.W. Storer 1989). Further details of its diet are needed.

DISTRIBUTION AND POPULATION The Plumbeous Ibis is a bird of the great marshy grasslands or pantanal of south-central South America. It occurs from northern and eastern Bolivia east to Beni, Brazil, in central Mato Grosso and in Rio Grande do Sul, Paraguay, Uruguay and Argentina, including Tucuman, Formosa, Santiago del Estero and northern Santa Fe. It is casual south in Argentina to Cordoba and Buenos Aires Province. All indications are that individuals appear to remain in a specific area year round (F. Olmos pers. comm. 1990). However, they are occasionally found outside the normal range and earlier reports spoke of some seasonality to movements, so additional detailed information is desirable on any short-range seasonal or dispersive shifts that might occur. This species is locally well known, but numerical information on population sizes are lacking. It is fairly common in some areas, such as the northern pantanal of Pecone, Mato Grosso, Brazil (F. Olmos pers. comm. 1990). But it has, overall, a rather patchy distribution, occurring regularly in some locations but more rarely in others (Wetmore 1926:63, Belton 1984:433). ECOLOGY The Plumbeous Ibis is a bird of wetlands, especially the seasonal marshes of the chaco and pantanal. It feeds in ponds, marshes, swamps, flooded pasture and other moist ground, especially drying pools of water. Although usually occurring in wet areas, it may also be encountered on drier grasslands and is commonly found in association with humans. It roosts in trees near water and is often seen resting in such locations.

BREEDING The Plumbeous Ibis nests solitarily within sparsely wooded areas (Dubs 1988), As it is very likely that these birds remain paired year round, the nest is probably placed within the usual home range of the parents. In Mato Grosso, nesting begins in March and April and continues at least until mid-October (Dubs 1988, Olmos 1990, Cintra 1986). This timing corresponds with the drying season, with fledging occurring towards the end of the dry period. Both members of the pair construct the nest, which is made of twigs and lined with leaves and grass (Dubs 1988). Nests are about 50 cm wide and 25 cm high (Cintra 1986). They are placed on horizontal limbs of the large trees; nests in the pantanal of Mato Grosso are reported to be up to 12 m above ground (Cintra 1986, Dubs 1988). Clutch size is 2-3 in the pantanal. The eggs are pale grey. Incubation is 28 days, with young hatching in early September (Cintra 1986, Dubs 1988). The young are heavily guarded by one of the parents during the first 10 days. When approached by humans, the ibis screams loudly, lowers its body, bristles its feathers and jabs at the intruder with its bill. The young also defend themselves with a forward lunge of the bill (Cintra 1986). These behaviours suggest that the risk of predation is high. Olmos (1990) described predation on nest contents by monkeys and a hawk. Nests in the pantanal have 2 young that fledge at 4043 days (Cintra 1986, Dubs 1988, F. Olmos pers. comm. 1990). They initially feed in marshes near the nest and then move away to feed with the family, returning to the nest-tree to sleep for several weeks (Cintra 1986).

TAXONOMY The Plumbeous Ibis has long been considered to be a very distinctive ibis, having been variously placed in several genera over the years, including Harpiprion, Molydophanes, Geronticus and, more recently, Theristicus, the latter suggesting a relationship to the BufFnecked Ibises. Short (1975:196) retained the genus Harpiprion, considering this species to have no close relatives. Until additional research

182 Plumbeous Ibis

is conducted into its taxonomic relationships, it seems best to retain the species in its traditional monotypic genus. CONSERVATION As is the case for some of the other South American ibises, so little is known about this species that neither its biology nor its status is sufficiently understood. Such information is essential if its conservation is to be assured. Fabio Olmos (pers. comm. 1990) reports that the species is not persecuted in the pantanal of Mato Grosso and is often associated with humans. As such, it is vulnerable to the burning and logging of nesting trees, which are found on

the higher ground also used for farming. Human association may also place them at risk from pesticides, especially in heavily cultivated regions such as the Taquari Basin. Although the pantanal of Mato Grosso remains relatively remote, future threats, such as hydroelectric dams, mining and expansion of agriculture, will also adversely affect habitat protection (D.A. Scott and M. Carbonell 1986:91).

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 310.

Buffnecked Ibis Theristicus caudatus (Boddaert) Subspecies: Theristicus caudatus caudatus (Boddaert) Scolopax caudatus Boddaert, 1783, Table Planches Enlum, p. 57; based on 'Courley a col blanc', de Cayenne of Daubenton, 1765-81, Planches Enlum, pi. 976: Cayenne, French Guiana Theristicus caudatus hyperorius Todd Theristicus caudatus hyperorius Todd, 1948, Proc. Biol. Soc. Washington, 61, p. 50: Buena Vista, Bolivia Theristicus caudatus melanopis (Gmelin) Tantalus melanopis Gmelin, 1789, Syst. Nat., 1, p. 653; based on 'Black-faced Ibis5 of Latham, 1785m General Synop. Birds, 3, p. 108, pi. 79: 'In insula novi annf = New Year's Island, near Staten Island, ex Latham Theristicus caudatus branickii Berlepsch and Stolzmann Theristicus branickii Berlepsch and Stolzmann, 1894, Ibis, p. 404: Maraynioc, Pariayacu, Junin, Peru Other names: Whitenecked Ibis, Blackfaced Ibis, Whitethroated Ibis, Andean Ibis, Grey Ibis (English); Bandurria, Bandurria Amarilla, Bandurria Comun, Bandurria del Norte, Bandurria del Sud, Bandurria de Verano, Bandurria de la Palma, Vandurria de Invierno, Mandurria, Tautaco, Curacau, Tutachi, Carcel, Code, Corococco, Gangola, Garza Pelata (Spanish); Curicaca, Despertador (Portuguese); Le Petit Courly d'Amerique (French); Witkin-Glans-Ibissen (Dutch)

IDENTIFICATION The Buffnecked Ibis is a relatively large light-coloured ibis, 71-81 cm long, standing c. 43 cm tall. It is distinguished by its distinctively buffy white neck and head and, in flight, by a large white patch contrasting with dark wing feathers. The basic colour pattern of the adult is rather complex and also variable among tropical, temperate and montane populations. The uppersides are basically grey highlighted by a slight greenish gloss and light grey to white wing-coverts, which show as the wing-patch noted above. The cream to buff colour of the head and neck, which is set off from the body coloration, turns a deeper yellow during courtship, fading to a whiter shade afterwards, probably due to wear. The top of the head is cinnamon rufous, as often are feathers at the base of the neck. The undersides are paler or darker than the base colour, depending on the population; the rump and undertail are black. A distinctive pectoral band of grey feathers crosses the chest in light-bellied forms. The grey flight feathers have a greenish gloss. The two northern subspecies, caudatus and hyperorius, have dark undersides, whereas the Andean and southern subspecies, branickii and melanopis, have light undersides with pale colour on the breast reaching further down on the belly although interrupted by the grey pectoral bar (Fjeldsa and Krabbe 1990, J. Fjeldsa pers. comm. 1990). It is not certain that the trends of variation in the species are appropriately captured within the definitions of the four subspecies, as there is much intergradation among the typical forms, and examples exist of birds outside the defined range of their plumage type (Friedmann and Smith 1950). Much of the variation is probably clinal, that is, associated with climatic differences from tropical environments to cooler and high-altitude environments, which tend to be

Buffhecked Ibis

occupied by smaller and lighter plumage birds. Essentially, the undersides become paler going from north to south and from low to high altitudes. The northern tropical subspecies, caudatus, has the darkest, blackish, underparts. Its neck is whitish buff with a rufous patch below. The wing-feathers have some white on them, especially on the coverts and parts of the primary feathers. The southern tropical subspecies, hyperorius^ is very similar to caudatus, and may not be well separable. It has lighter but still relatively dark undersides, grading from brownish to grey-brown. Its dorsal colour is also a bit browner than caudatus. The grey feathers have a lighter grey edging, making them more conspicuous (Todd 1948). Its tail has a green sheen. These birds generally have larger bills than caudatus (C.S. Luthin pers. comm. 1990). The southern temperate melanopis, has a richly buff foreneck and breast and a rather restricted throat feathering, sometimes limited to the chin alone, giving it the fullest gular pouch of the recognized subspecies. The wing-coverts are pale grey to amost silvery white. The Andean subspecies, branickii, is somewhat smaller than the other subspecies but has the longest wings (C.S. Luthin pers. comm. 1990). The base of the neck is tinged with rufous. It has the palest breast and undersides, which are pale buff to creamy white. Its wing-coverts are a rather dark grey barred with paler feather-tips showing as a greysilver (J. Fjeldsa pers. comm. 1990, C.S. Luthin pers. comm. 1990). Overall, its dorsal coloration is more consistently grey than in the other forms. In all subspecies, the skin around the eye and on the sides of the throat or entire throat is black, and may be somewhat

185

warty. The dark face has inspired one of the species' common English names, the Blackfaced Ibis. The iris is brownish red, often described as being red to crimson. The legs and feet are pink, orange-red or purple-red. The chin retains a thin line of feathering, less in melanopis. Along with the plumage, the soft parts brighten during the breeding season. The legs turn redder at the start of courtship. They have also been reported as black tinged with crimson (P.S. Humphrey et al. 1970:104). Both males and females possess an expanded gular sac during this time. The sexes are similar in plumage, but males are larger in body measurements and weight. In Chilean populations, males averaged 1758 g, and females averaged 1598 g (Gantz in R. Schlatter pers. comm. 1990). At hatching, the young have little feathering, but a grey down soon develops. J. Fjeldsa (pers. comm. 1990) reported that a young bird he collected in Patagonia was uniformly grey, but Blaauw (1917) reported that the head and breast of nestlings are yellow-white. The legs are initially light pink, the throat blue-black and the bill blue-white with black at the tip. The legs then turn grey, with a purple tinge. The iris is brown; the facial skin is black, with a white patch on either side at the base of the bill. Juveniles are similar to adults, but their buff colour is duller and their abdomens are paler than in the adult. The head and neck are streaked brownish. The wing-coverts have buff edges, giving a scalloped effect (Fjeldsa and Krabbe 1990). The head turns more rufous with age. Soft parts are duller than in the adult. The legs are pink and the iris is dark brown (B.T. Thomas pers. comm. 1990, Schlatter pers. comm. 1990). An immature bird in the American Museum of Natural History had a greenish bill and brownflesh feet (P.S. Humphrey et al. 1970:104). In feeding and flying, the Buffnecked Ibis is quite vociferous, frequently giving its distinctive two-note call. On the ground, this call may be given with the head thrown back; in flight, only the first syllable may be called. It is a musical, bell- or trumpet-like call that has been variously rendered because it sounds different depending on distance and terrain. In English, it may be rendered as 'Tur-tac*, accenting the second syllable. It is better described as being reminiscent of a blacksmith's hammer on an anvil. Hudson (1920) described the call of melanopis as short, abrupt notes of extraordinary power resembling the blows of a powerful hammer on a metal plate. In 1860, Darwin described the call as being like the neighing of the wild llama (the guanaco) (Darwin 1962:166). The Buffnecked ibis flies with great dexterity on quick, strong wing-beats. In flight, the legs do not extend beyond the tail, and the light wing patches flash conspicuously. The birds often fly low over to the ground but sometimes soar to great height during their daily wanderings (J. Fjeldsa pers. comm. 1990), and they characteristically undertake complex aerial manoeuvres. Their acrobatics going into roost have been much remarked upon. Hudson (1920) described the performance as frenzied, the birds simultaneously dashing downward with 'a violence wonderful to see'. Individuals and flocks, where they occur, often wheel and zigzag while calling loudly. The birds rise and fall, sometimes

186 Buffnecked Ibis

together and sometimes scattered, while calling with their 'loud percussive screams' (Hudson 1920). The species should not be easily confused with any other ibis, owing to its distinctive call, neck colouring and, in some races, the bicoloured undersides. The Plumbeous Ibis is monocoloured and has a thicker neck. The Buffnecked Ibis might look like a goose from a distance, but not up close. DISTRIBUTION AND POPULATION The Buffnecked Ibis occurs over much of South America, from Colombia southward. Hiatuses in distribution occur, such as the highlands of southern Colombia, southern Venezuela and Brazil, parts of the Ecuadorian and Peruvian Andes, the coastal desert of Chile, inland northern Chile, and western Argentina (Zapata and Martinez 1972, Blake 1977, Fjeldsa 1985a, Fjeldsa and Krabbe 1990, J. Fjeldsa pers. comm. 1990, Luthin pers. comm. 1990). It is accidental in Panama to the north and the Falklands to the southeast. Theristicus caudatus caudatus is the northern lowland form, ranging from Colombia south to Valle del Cauca and Meta and Venezuela south to Apure and Bolivar, to the Guianas, and into northern Brazil. It is this race that is casual in Panama (Pacora and San Jose) (Blake 1977:194). Theristicus caudatus hyperorius occurs in central and southern Brazil, eastern Bolivia, northern Argentina (Tucuman, Chaco, Santa Fe, Entre Rios), Paraguay, southeastern Brazil and Uruguay, with an isolated small population above 2000 m in the Cordoba Mountains near Cordoba, Argentina (Arballo 1990, J. Fjeldsa pers. comm. 1990). Theristicus caudatus branickii is the northern highland subspecies, occurring very locally in the mountains of Ecuador (Antisana, Cotopaxi), Peru (Ancash, Junin, Huancavelica, Ayacucho, Cuzco, Puno), northwestern Bolivia (La Paz, Oruro), and extreme northern Chile (Arica), rarely moving down to the lowland valleys of Chile, Peru and Bolivia (Cochabamba) (Fjeldsa and Krabbe 1990). Theristicus caudatus melanopis is the Patagonian subspecies (Blake 1977, Fjeldsa and Krabbe 1990, J. Fjeldsa pers. comm. 1990). To the north, it occurs in relict populations along the Andes in Peru (Lambayeque, La Libertad, Lima, Tacna). It is absent from the Atacama Desert, occurring again south of Antofagasta, then down the coast to Tierra del Fuego, including the offshore islands. In the northernmost part of this range, it occurs in the high mountains, whereas further south it uses the central valley of the Andes and the coastal zone, and even further south it occurs across the whole of Chile (A.W. Johnson 1965). In the southern Argentine, the range extends northward through the Patagonian grasslands to Neuquen and Rio Negro. Many populations appear to occur in the same area yearround. However, the southern populations are migratory, and other birds may shift seasonally, especially in the cool or dry seasons. The southern melanopis populations move northward to Chile and Argentina as far as Tucuman, Cordoba and Buenos Aires Province, the latter destination being the main wintering ground for Patagonian birds

(MacDonagh 1942, Nores and Yzurieta 1980, J. Fjeldsa pers. comm. 1990). They may be more common there in dry years (Weller 1967). This subspecies is also the one that is accidental in the Falklands. The movement of southern populations to the north in the winter is the origin of several of southern South American common names, such as Bandurria de Verano. Although the northern populations of the llanos remain year-round, some movements apparently take place, as they are less abundant in the dry season (E. Aguilera pers. comm. 1990). Similarly, Andean birds may shift to lower or wetter locations, including into the pampas of Argentina, during the winter dry months (Fjeldsa and Krabbe 1990, C.S. Luthin pers. comm. 1990). Little is known about population sizes, undoubtedly because of the birds' wide dispersal. Overall, however, where they occur, most populations appear to be relatively high. The species seems particularly common in the grasslands of the llanos and Patagonia. Montane populations of branickii have always been considered to be rare, perhaps because of their highly localized distribution, their remoteness and their wary habits (Fjeldsa 1988, Fjeldsa and Krabbe 1990, C.S. Luthin pers. comm. 1990). It appears that this species is fairly common in the southern Andes but much rarer further north. More information is needed on the status of this species in the Andes. ECOLOGY The Buffnecked Ibis is a bird of the damp to dry grasslands and meadows of South America (Vuilleumier 1985, Nores and Yzurieta 1980, Fjeldsa 1988, Fjeldsa and Krabbe 1990, pers. obs. J.A.K. and J.A.H., F. Olmos pers. comm. 1990, R. Schlatter pers. comm. 1990). It occurs from sea level up to about 3000 m, though a specimen in the US National Museum was collected at 4200 m, and it occurs in the Puna Zone at 5000m in the Andes (Fjeldsa and Krabbe 1990, pers. obs. J.A.K.). This species feeds primarily in relatively dry, short grass, such as on relatively dry savannas, agricultural fields, pasture land, and scrub lands in Patagonia and other parts of southern South America, and is the most terrestrial of the Neotropical ibises (Nores and Yzurieta 1980, Kushlan et al. 1985, pers. obs. J.A.H. and J.A.K.). It uses very dry ground during the dry season and after fires, but also makes use of a variety of damp habitats when available, including edges of lakes, ponds and rivers, damp meadows, and openings in forests. Montane populations occur in boggy glades fringed with forests of the valdivian (rain forest) zone and in grasslands of the high altitude Puna and paramo zones. J. Fjeldsa (pers. comm. 1990) points out that the species is highly localized in the montane grasslands. Along the coast, it feeds in shore meadows. F. Olmos (pers. comm. 1990) did not see them feeding in or around ponds or lagoons. The birds feed alone, in pairs or in variably sized, but generally small, loose flocks of up to 50 individuals, which travel together between feeding sites. They are often reported in small flocks of up to 6-8 birds in the llanos. We have seen them feed there primarily in pairs and in the

Buflfnecked Ibis

scrublands of Patagonia we found them feeding singly and in pairs (pers. obs. J.A.K. and J.A.H.). In the pantanal, F. Olmos (pers. comm.) found them feeding in pairs and in groups of up to 4. The montane forms are almost always solitary or in pairs. It is likely that they primarily remain in family groups. Brooks (1991) estimated their territories to average 0.65-1.15 km2 in the Paraguayan Ghaco. Feeding is by Probing in soft soil and Pecking at the surface of hard soil. (Sick 1985, R. Schlatter pers. comm. 1990, J. Fjeldsa pers. comm. 1990). In Probing, the birds can drill deeply into loose soil. On hard ground, they probe into bunch grass tussocks. The species consumes a variety of prey found in and on the soil (Darwin 1962:166, Hudson 1920, MacDonagh 1942, P.S. Humphrey et al. 1970:103, San Martin 1959, Sick 1985, Robert Storer pers. comm. 1990, R. Schlatter pers. comm. 1990). The diet is primarily soil invertebrates, including grasshoppers, crickets, beetles, caterpillars, insect larvae, spiders, centipedes, scorpions, earthworms and also some vertebrates, including frogs, toads, tadpoles, salamanders, lizards, snakes and rats. It swallows all kinds of items, such as stones, seeds, grass pieces and bones, perhaps as a grit (R. Schlatter pers. comm. 1990). On Tierra del Fuego a rodent tooth and shell fragments of a goose egg have been reported from a large chick (P.S. Humphrey et al. 1970:103). Gantz found that they consume 590 insect larvae per day (R. Schlatter pers. comm. 1990). Their flights to roost have been often remarked upon. They are heard before being seen and repeatedly take the same flight paths at night to their roosts in trees. These are often the largest trees around. Sick (1985) reported that they roosted in parana pines (Araucaria angustifolia) and in the Amazon estuary in kapoks (Bombax munguba). We have seen them roost in palms (pers. obs. J.A.K.). BREEDING Buffnecked Ibises nest solitarily or in loose colonies of 10-30 pairs situated in a wide variety of locations (A.W. Johnson 1965, Sick 1985, P.S. Humphrey et al. 1970:102). They are sometimes found in mixed colonies of night herons, cormorants and other wading birds (J. Fjeldsa pers. comm. 1990), but more usually alone. In the high Andes and on rocky islands, they nest on the ground and on cliffs. At lower forested elevations, they nest in trees or rocks scattered on the savanna or in woods. They have also been reported as nesting in marshes, on abandoned dredges, on poles, in reeds in a lake-edge marsh and on the ground along a stream. The nesting season varies but usually corresponds to rainy periods or, in the mountains and in subantarctic areas, to warm periods (C.S. Luthin pers. comm. 1990). In the llanos, nests may be found year-round though they are most frequent in March and October, at the end of the drying season and in the wet season (Hilty and Brown 1986:72, C.S. Luthin pers. comm. 1990, E. Aguilera pers. comm. 1990). B.T. Thomas (pers. comm. 1990) observed birds in the llanos copulating in November. The southern populations nest in September-December,

187

including those in extreme southern Chile and Argentina (A.W.Johnson 1965, P.S. Humphrey et al. 1970:103, Nores and Yzurieta 1980:27, Fjeldsa and Krabbe 1990). In Peru, they nest in June. In Tierra del Fuego, they arrive at the time of the first break up of winter ice, nesting in November to February. Almost no information is available on courtship and nesting behaviour. B.T. Thomas (pers. comm. 1990) reported observing allopreening and copulation involving a pair of birds that were otherwise occupied by feeding in a dry field. The male mounted the female, they snapped their bills together, then the male nibbled at the female's neck feathers. Afterwards, the female walked away shaking her feathers, and both resumed feeding. Clearly, much more needs to be known about courtship in this species. The nest is a large, bulky, loose structure of sticks, with grass or rush stems intermeshed. Nests are usually placed high in trees, 3 to 10m above the ground. The nesting site may be reused in subsequent years, if it is not disturbed (Vigil 1973:50). The usual clutch is 2-3 eggs, which are white to greenish white with some light brown speckling. The incubation period is 28 days, and in captivity the young are capable of fledging at 8 weeks (Blaauw 1917). TAXONOMY The Buffnecked Ibis is a highly variable species with respect to size and plumage coloration. As a result the relationship among the several populations has been much debated, with several different species and subspecies having been proposed at various times (Salvador! 1900, Todd 1948). Although some populations have distinctly different coloration, their ranges and intergradations are not well understood. It is likely that most variation is clinal, and it is unclear how or whether the species might best be subdivided taxonomically. CONSERVATION The Buffnecked Ibis has long been known, perhaps first being noted by Cook in Tierra del Fuego, who reported a species of curlew as big as a heron (Crawshaw 1907). The species is widespread and abundant over a large part of its range, especially in the humid pastures and grasslands of the llanos in the north and pantanal in the south. Overall, it has adapted well to agricultural practices in the grasslands, making heavy use of pastures and fields; as a result populations have increased in southern Chile in the past 40 years (R. Schlatter pers. comm. 1990). However, montane populations from Ecuador to Bolivia are highly restricted. Populations in coastal Peru and northern Chile have decreased, and the bird can only be seen in a few places (Hughes 1970, Koepcke 1970, Gantz in R. Schlatter pers. comm. 1990, J. Fjeldsa pers. comm. 1990). Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 311.

Green Ibis Mesembrinibis cayennensis (Gmelin) Tantalus cayennensis Gmelin, 1789, Syst. Nat., 1, p. 652; based on Courly verd, de Cayenne of Daubenton, 1765-81, Planches Enlum., p. 820: Cayenne, French Guiana Other names: Cayenne Ibis, River Ibis, Bush Ibis, Curry-curry, Bush Curi-curi, Bronze Ibis (English); Corocoro Negro, Corocoro de Monte, Cocoquam, Zanurita, Garza Negra, Cocli Negro, Bandurria Habladora, Caracolero Verde-negro, Cuervillo Grande (Spanish); Korokoro (Dutch); Chapeu-velho, Corocoro, Tapicuru, Carauna (Portuguese)

IDENTIFICATION The Green Ibis is a relatively small ibis (45-60 cm long), somewhat stocky, short-legged and longbilled, with a glossy dark plumage. The adult's base colour is black, which is how it appears in poor light. However, its entire back is highlighted by a bronzy green sheen. The forehead is greyish. The green plume-like feathers on the back of the neck are a distinctively iridescent emerald, but the bushy nuchal crest formed by these feathers is not terribly conspicuous. The tail and flight feathers are deep blue-black. The ventral plumage is duller than the dorsal. The bare face and orbital areas are grey, the bill is olive, and the legs and feet are pea green (S. Olson pers. comm. 1990). The iris is pale brown to black (Luthin 1983a, S. Olson pers. comm. 1990). Storrs Olson (pers. comm. 1990) found a bird in breeding condition to have intensified soft-part colour. The legs and feet were a sparkling metallic green, whereas the throat skin was a violet indigo. When the gular pouch is distended during courtship, this shows its bright blue cheek-line. The sexes are alike. Museum specimens labelled as to sex overlap in size, although the largest birds tend to be males. 2 females collected by S. Olson (pers. comm. 1990) weighed 700 and 740 g whereas 2 males weighed 745 and 800 g. A bird whose sex was not determined weighed 890 g. Bill-length overlaps similarly. Additional information is needed to determine whether dimorphism occurs in this species. Nestlings are dark grey, with a distinctive circumorbital white line (C.A. Wood 1923). By fledging, the juveniles are very much like adults. Luthin (1983a) reported that they have adult soft-part colours, a green sheen on the feathers, and even a slight nuchal crest. The white eye-line is often apparent in young for some time after fledging (B.T. Thomas pers. comm. 1990), although it eventually fades with age. The Green Ibis is a noisy bird, well known to lockl inhabitants by its dawn and dusk calling, which gives it and other ibises their most common Spanish name, Corocoro. At night, in the morning and other times at roost, it is especially noisy. When perched in trees, it often gives a long, low, throaty, mammal-like growl on an even pitch, repeated for minutes on end (S. Olson pers. comm. 1991). The primary call of this species is a growling, rattling, rolling 'Kro-kro-kro -kro' or 'Clu-clu-clu-clu'. When feeding it is quieter, but clucks a throaty, watery 'Koke' or 'Uk' (Slud 1964). Green Ibises often fly from feeding site to feeding site in small groups in a jerky, heavy, stiff-winged flight. Its flight call has been variously described, as a rattling 'Klaw-klaw-klaw', an accented 'Wok-wok-wok-wok', and a series of 3-6 loud 'Ga-lups' followed by 4^9 'Brups' (the latter described by Luthin 1983a). The Green Ibis is generally recognizable by its size and coloration. It is stockier, shorter and has a thicker bill than similar species. The Glossy and Whitefaced Ibises have maroon iridescence rather

190 Green Ibis

locally common, and overall population levels are probably as high as could be expected for such a bird, ECOLOGY

than green and relatively longer legs. The Barefaced Ibis is smaller with pink rather than green soft parts. The Plumbeous Ibis is grey.

DISTRIBUTION AND POPULATION The Green Ibis is a forest bird, primarily of the wooded basins of the Amazon and Orinoco rivers of South America, a habitat preference reflected in several of its older common names, such as River Ibis and Bush Ibis. Its range extends northward into Central America. In the north it ranges from the Caribbean coast of Costa Rica to Limon, through Panama, northern Colombia to the Venezuelan interior (Orinoco and Apure Rivers) as well as coastal Falcon, the Guianas, west again to east of the Andes in Brazil, Bolivia east to Beni, Paraguay, and northern Argentina in Misiones (Olrog 1963:88, D.A. Scott and M. Carbonell 1986). Seasonal population shifts seem to occur in some areas. In the pantanal, individual Green Ibises are reported to occur in May-November, and in the llanos from November to August (B.T. Thomas 1979a, R. Cintra pers. comm.). Individuals are frequently found outside the nesting areas. Dispersing birds are found as far northward as Sarapique, Costa Rica and Rio Plantano, Honduras (American Ornithologists* Union 1983) and stragglers in Rio Grande do Sul in January (Sick 1985). Nothing is known about population sizes and the birds are well dispersed in suitable habitat. However, they are

The Green Ibis is a bird of the water's edge and shallow water. However, it is found locally in a variety of habitats, including wooded swamps, forest pools, small marshes, heavily timbered gallery forests along rivers, on forested streams having muddy banks and along roadside ditches. It also occurs in secondary growth, such as abandoned coffee plantations (Haverschmidt 1968) and, as we have seen, in more open pasture as well (pers. obs. J.A.K.). It feeds in muddy areas particularly, on the edge of river banks and forested pools, where its dark coloration renders it well camouflaged. Generally, this is a rather solitary bird. In a study of its feeding habits in the llanos, one of us found it feeding solitarily, in pairs, or in small loose flocks of up to 6 or 8 birds (pers. obs. J.A.K.). Gonzalo Morales (pers. comm. 1990) observed groups of up to 30 feeding in the Venezuelan llanos. Betsy Thomas (pers. comm. 1990) has also seen flocks in the same region, reporting one of 26 birds. When it feeds near other species of ibis—though this is infrequent— it is usually near the edge of the group (pers. obs. J.A.K.). Green Ibises appear to use single feeding areas and roost sites for much of the year. They seem to establish territories that they appear to claim by calling. Luthin (1983a) found only solitary birds or pairs during the nesting season; but, after nesting, the larger groups of 2-6 typically seen are probably families. Often the individuals of a probable pair are well separated when actually feeding, but they and the larger family group roost together at night. Upon disturbance, the Green Ibis turns its head jerkily while nodding and making its clucking sound (Slud 1964). It then walks or flies to nearby trees where it calls in its distinctive manner, often hidden from view. Green Ibises forage for long periods throughout the day, and spend the remainder perched in nearby trees. They feed by walking relatively quickly, Picking at the surrace and Probing rather deeply in soft mud (pers. obs. J.A.K.). They also feed while rapidly opening and closing their wings (Ogden and Thomas 1985a). A muddy feeding spot is characteristically chosen, and birds are often seen with bills partially mud-covered. We have observed them feeding slowly by a sequence of Standing-Probing-Walking-Probing in a thorough and deliberate manner, completely working over a patch of mud, a small pool or a stream edge, spending many minutes or an hour in a single area (pers. obs. J.A.K.). This deliberate foraging tactic has been well known for many years (R.H. Schomburgk 1848:503). Luthin (1983a) reported numerous probes around soft earthworm holes in the forest. Green Ibises are documented to eat aquatic insects (Orthoptera, Homoptera, water bugs and water beetles), worms, molluscs, including snails, small fishes and frogs (Sick 1985, Ogden and Thomas 1985a,b, G. Morales pers. comm. 1990, pers. obs. J.A.K.). Sick (1985) has reported two instances of its eating plants. He found large quantities

Green Ibis 191

of fibrous vegetable matter in two specimens taken at Linhares, Esprito Santo, Brazil (H. Sick pers. comm. 1990). Undoubtedly, the overall diet is more varied than what has so far been recorded. BREEDING Green Ibises nest solitarily and inconspicuously high in trees and thickets, usually in gallery forests along rivers. They appear to nest within their long-held activity areas. Birds in breeding condition have been collected in Panama and Colombia in February-April (S. Olson pers. comm. 1990, US National Museum specimens). They have been reported as nesting in April in Colombia, September in Surinam, and June-August in Venezuela (Haverschmidt 1968, B.T. Thomas 1979a, 1990, Hilty and Brown 1986:73). In the llanos, courtship and nesting occurs during the rainy season, typically beginning 2 months after the onset of the rains (Luthin 1983a). Its nesting season is shorter and more consistent than that of the Barefaced Ibis, which nests in the same area. Luthin (1983a) was able to study some of the displays of this species, providing most of what is known and described below. It is likely that birds remain paired year-round, so some courtship displays may be truncated, and the courtship period is rather extended. As the breeding season approaches, display sites are selected at the top of tall trees, probably within the pre-existing home range. During courtship, Green Ibises alight together on a display tree, standing no more than 2 m apart. Courtship is initiated by one bird performing a Head Shaking display, and the second bird responding with its own Head Shaking. The pairs then may Duet, the call consisting of a soft, deep 'Brrr*. They hold their necks near vertical and elevate their head feathers. At the end of the duet, both birds shake their heads sideways and bump bills. One or both birds may flare their gular pouch, showing the bright blue line along the cheek. After a pause some birds then begin Stick Shaking or bouts of allopreening directed to the head and neck. Occasionally, Display Preening occurs, in which the birds raise the feathers on their necks and back and slightly droop their wings. Display Preening and Stick Shaking are less common in the Green Ibis than in other, similar species (Luthin 1983a). Copulation displays are, similarly, less developed than some of the other Neotropical ibises. The male Green Ibis stands close to the female, sweeps his bill back and forth in front of her sometimes with his head over hers. The male tries to grab the female's bill while mounting her, and may keep hold during copulation. The pair constructs a rather flimsy, flat structure made loosely of sticks, lined with finer material, according to C.A. Wood (1923), although Luthin (1983a) failed to find such finer material in the nests he examined. The birds are secretive in their nesting, entering and leaving the nest-site quietly. Luthin (1983a) found that

activity at the nest was limited and vocalizations consisted only of a quiet gurgle during exchanges at the nest. Eggs are a dark olive green, without markings or gloss. 2 eggs from Surinam measured 54.9 X 38.8 and 55 X 37.9 mm and weighed 46 and 43 g (Haverschmidt 1968). Clutch size, when recorded, has been 2-4. Hatching is asynchronous. Luthin (1983a) found that only 2 young survive to fledging. Hatchlings have distinct facial markings that disappear in 2 or more weeks (Luthin 1983a). Like the adults, young also are quiet at the nest, begging with a light buzzing call (C.S. Luthin pers. comm. 1990). After 2-3 weeks, the adults return to the nest-site only to feed the young, which continue to remain relatively inactive. Young in the nest crouch down when approached by humans (C.S. Luthin pers. comm. 1990). The secretive behaviours of the young and adults at the nest suggest that predation risks may be high. Luthin (1983a) relays a report of capuchin monkeys (Cebus nigrivittatus) eating young. Other predators are undoubtedly a threat also. The young fledge at 23-27 days. They remain near the nest for a few days and then move with the adults to their feeding sites, thereby forming the family groups so apparent at the end of the rainy season and thereafter.

TAXONOMY Like most ibises, the taxonomy of the Green Ibis has not been thoroughly studied. Based on behavioural information, Luthin (1983a) has suggested that Mesembrinibis might be related to the solitary Old World ibises of the genus Bostrychia, such as the Spotbreasted, Olive and Hadada ibises. Both groups are secretive, solitary, forest-dwelling species, loudly calling at dawn and dusk, having dark plumage and nuchal crests, and building loosely constructed nests high in trees. Behavioural patterns held in common include mutual Head Shaking while bumping bills and elevating the feathers, bill grasping during copulation, and Mutual Preening. However, these behaviours are not unique, and additional behavioural and morphological study is called for.

CONSERVATION Little is known about the biology, detailed distribution and population sizes of this species. Such information is needed to understand its ecological requirements and its conservation needs. However, the Green Ibis appears to be widely dispersed within its range, and is, overall, probably reasonably common.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 312.

Hadada Ibis Bostrychia hagedash (Latham) Subspecies: Bostrychia hagedash hagedash (Latham) Tantalus hagedash Latham, 1790, Index Ornith., p. 709: Cape of Good Hope Bostrychia hagedash brevirostris (Reichenow) Theristicus brevirostris Reichenow, 1907, Ornith. Monatsber., 15, p. 147: Alen, Southern Cameroon (=Equatorial Guinea) Bostrychia hagedash nilotica Neumann Bostrychia hagedash nilotica Neumann, 1909, Ornis, 13, p. 193: Kimo, NW of Addis Ababa, Ethiopia Other names: Hadada, Hadedah (English); in-Kankane, 1-Han, Ing'ang'ane (South Africa); L'ibis Hagedash (French)

IDENTIFICATION The Hadada Ibis is a short-legged, medium-sized (610mm long), dull brown bird, with a somewhat greyer head and neck. Close up, the basically brown coloration of the mantle and back of the adult has an iridescent green to bronze cast. The head is greyer than the body, with a prominent huffish white stripe running across the cheek. The neck is a brownish or smoky grey. The colour of the primaries, secondaries and major coverts vary somewhat according to the light, from greenish brown to pale red and purple, forming coloured patches on the wing. The tail is dark blue, with the rump and tail coverts green. The bill is dark horn with a strong red marking at the base of the upper mandible. The featherless lores and base of the bill are black, and eye is dark brown ringed by a prominent white ring. The legs and feet are dark horn-brown. During courtship, the red marking on the bill brightens to a crimson, and the bill itself darkens. The legs and feet turn darker, and a pale reddish flush appears on the toes. The nestlings are pink with brown down, black bill, and pink legs and feet. Immature birds are similar to adults but are greyer and have less iridescence on the back and wings. The best-known characteristic of this ibis is its jerky irregular flight and its loud echoing call, 'Ha-dada', from which it gets its common English name. This call is given by both male and female and is particularly heard when flying to and fro between the roost and feeding ground. It is also heard quite often during the day, as birds move between feeding grounds. It feeds silently and allows quite close approach where not persecuted. The main differences between the three subspecies are size and some aspects of coloration (Amadon 1953). The hagedash subspecies is relatively short-billed and paler than the others. The brevirostris subspecies is a darker brown colour with its glossy wing patch much greener and more purple, according to the light. The bill of nilotica is noticeably longer, and some specimens are strikingly larger. (Comparative measurements are: hagedash—bill 11.7-15.3, wing 33.4-37.Ocm; brevirostris—bill 12.616.3, wing 33.0-38.3; nilotica—H\\ 15.2-17.4, wing 33.5-39.2—Brown et al. 1982.) This species is easily confused with others in the genus Bostrychia, as noted in those accounts. Voice differences are important in identification. The Hadada has a very noisy, harsh call with three distinct syllables. It has been compared to the wail of a child in distress and the hideous mocking laughter of a witch (F J.Jackson 1938). The three part call contrasts with the two syllable call of the Olive Ibis. The

194 Hadadalbis

Hadada's call also contrasts with the more raucous call of the Spotbreasted Ibis. The other species of Bostrychia also have a very prominent nuchal crest, which is minimal in the Hadada Ibis. DISTRIBUTION AND POPULATION This ibis is a widespread species throughout the African continent south of the Sahara desert (Mackworth-Praed and Grant 1962, Snow 1978, Brown et al. 1982:195-196); hagedash is found in Zimbabwe, southern Mozambique, and eastern South Africa. North of the Zambezi Valley, hagedash is replaced by brevirostris, which is found throughout central and east Africa, northwards through Kenya and Somalia and westward to Gambia and Sierra Leone, and along the Congo River. The nilotica subspecies occurs from Uganda and western Tanzania to Zaire, Sudan and Ethiopia (Pitman 1928, 193 Ib). Near Mount Kenya, it is a lowland species and is replaced at higher altitudes by the shyer, rarer Olive Ibis. In southern Africa, the Hadada has in recent decades expanded its range from the southeastern coastal belt to the two higher inland plateaus, as well as westward along the coast (Skead 1966). This expansion has been in response to the construction of dams and reservoirs (Snow 1978). Little is known about the population sizes of this species, despite (or because of) its wide range. Bannerman (1953) found it to be a common bird in certain areas of west Africa but rare in some coastal provinces, particularly in Nigeria. A colony in Cape Province, South Africa, was recorded as numbering 70 birds (Raseroka 1975a). ECOLOGY The Hadada Ibis inhabits damp ground along rivers and streams, especially preferring open woodland, savanna and

marshlands near trees. It is attracted to lakes and manmade irrigation, avoiding the desert regions of north and southwest Africa as well as thick forest areas. It is found close to farms, cultivated farmland and lawns. On Mount Kenya and other mountain areas, it is not found above 1800m. Hadada Ibises are, in the main, sedentary birds feeding in pairs, small groups or in some cases parties of up to 50 birds, sometimes in association with other birds, especially Cattle Egrets (Bubuleus ibis). We have also seen single birds quietly feeding in shallow pools in the company of Goliath Herons (Ardea goliath), Wood Sandpipers (Tringa glareola) and Greenshanks (Tringa nebularia). Flocks vary in size during the day, birds breaking away periodically to feed elsewhere. In a tree-lined marshy pool in the Masai Mara in Kenya, we have watched Hadada Ibis arrive in small parties, noisily flying in at dawn, but then settling to feed quietly throughout the day, until they move off to feed around village huts on their way to their roost. Between feeding sites, they fly haphazardly with jerky flaps. Preferring open ground, they feed on both dry and damp areas of short grass and cultivated fields and even gardens, usually by Probing in soft soil. On hotel lawns along the shoreline of Lake Naivasha in Kenya, we have seen them Walking Slowly in small parties in company with Sacred Ibis, Probing the grass and flower beds and vegetable patches. They particularly favour the base of trees where the soil is exposed. The availability of soft soil may be critical to the species. Ossowski (1952) suggested that heavy mortality during severe droughts was due to the hardness of the soil. The birds may also be surface feeders under some conditions (Raseroka 1975b), but this needs additional study. Hadada Ibises are attracted to wattle plantations in Natal, South Africa, and are thought thus to act as effective pest controllers (Ossowski 1952). This work also underlined the value of the Hadada in keeping golf courses free of the larvae of chaffer beetles (Ossowski 1952). The Probing extracts a variety of invertebrates, including earthworms, beetles, Lepidoptera, fly larvae, millipedes, water bugs and crustaceans. Raseroka (1975b) found that 96% of the animals ingested were insects, and 60% of the diet was made up of animals obtainable from the soil surface. They have not been recorded as taking large prey. BellCross (1974), in his observations offish eating birds, never found this species to eat fish. Inedible items, such as stones and glass, are often found to be taken by Hadada Ibises (Skead 1951, Ossowski 1952). Skead (1951) concluded that they were attracted by bright objects on the ground. As evening approaches, birds straggle to roosts, some pairs roosting apart from the main flock. At the roosts, the birds will call loudly to each other and will be answered by birds at other nearby roosts (Manry in Brown et al. 1982: 195-196, pers. obs. J.A.H.). These large nocturnal roosts are of 100 or more birds, usually away from other species. They are often used year-round, and year after year.

Hadadalbis 195

BREEDING It is likely that some birds maintain a pair-bond through the year, as we have seen interactions outside the nesting season that suggest this. Commonly, a female, with wings half open, approaches a male and touches her bill to his. After this brief mutual bill touching the female will resume feeding elsewhere amongst the party. When disturbed, the two birds will separate from the other birds and fly up to the branch of a tree standing close together. Hadada Ibises nest solitarily in trees or bushes, often over or near water (Raseroka 1975a). In South Africa, nests have been seen in trees some distance from water, and even on a telegraph pole (Line 1941, Raseroka 1975a, Uys and Broekheysen 1966). Time of nesting varies geographically. Raseroka (1975a) showed that nesting ranged from February in Zambia to October-November in southern Africa (also observed by Ossowski 1952). In Eastern Cape Province, eggs were laid from the beginning of November to mid-January. Nesting is recorded in April in Uganda (Pitman 1931b). As the breeding season approaches, paired birds will separate from the feeding parties, which gradually break up as the nesting season commences. It is likely that established pairs have a lower intensity of courtship interaction than do those nesting for the first time. We saw one such incident at the commencement of the rains in Kenya, and it was noticeable that red parts of the upper mandible were brighter on the interacting pair than others of the feeding party. In a party of 8 feeding birds, one bird picked up single sticks, grass and straw at intervals, and paraded through the group and then away from it. It then crouched amongst the other birds, vibrating its half-open bill. One of the birds then came from the group and stroked the quivering bill of the displaying bird with its own closed bill in a 'crossed sword' action. The first bird then touched the prominent white stripe on the second bird and the interplay ceased. We saw this behaviour by the two birds about eight times in over 15 min. Skead (1951) noted that the billing behaviour of pairs included rattling of bills, up and down, and side to side, while nodding the head. This took place at roosts, but more intensively at nesting sites, where sticks were offered by each bird to the other, followed by neck intertwining, mutual preening, head shaking and jibbering of the lower mandible. W. Wennrich (1983) noted that this behaviour preceded copulation and was followed by mutual head raising and lowering in unison. Stick presentation continues into the nesting period, and during incubation the arriving bird will offer a stick to the sitting bird, which will then leave the nest without further display between the pair. Large sticks form the base of the nest and these are lined with small sticks and then with grasses and lichen. Nests can be completed within 14 days but can take up to a month or longer, with periods of suspended activity (Raseroka 1975a, Brown et al. 1982:195-196). Sticks are both removed and added as building goes on. Nests in Combretum caffrum trees averaged 17 m apart and varied from 1.2 to 7.0 m high

above the water or ground (Raseroka 1975a). Nests are used for a number of years, and Skead (1951) saw a nest in a kaffir plum tree used for 6 consecutive nesting seasons, although there is no certainty that the same pair used the nest each year. Some birds will take over partly built nests. 2-3 eggs are laid, but sometimes 4, on alternate days. Eggs are large, often heavier than those of a domestic fowl, but highly variable even within a single clutch. They are a buff colour and variably spotted and flecked with light brown. Often the flecks are overlaid with sepia pigment. Size varies from 56.9 X 41.5 to 64.0 X 41.0mm, with an exceptional egg measured at 67.2 X 42.5 (Skead 1951). Pitman (1928) noted an average egg size of about 60 X 42 mm. Incubation takes between 25 and 28 days and is shared by both parents (Raseroka 1975a). Chicks hatch at long intervals, some taking up to 48 h to leave the egg, and thus the developmental stage of nestlings vary. Young remain in the nest for up to 33 days. They are blind for the first 3 days and have pink skin covered in brown down. Their bill is black and straight. They utter a purring sound when food begging, and are fed by regurgitation, which continues after the young have left the nest. Before fledging, young are left at the nest, but are brooded at night (Raseroka 1975a). Each parent takes young to feeding areas, coming together as a family regularly and roosting together in the vicinity of the nest. TAXONOMY With the merging of the genus Hagedashia into Bostrychia, the relationships of the Bostrychia ibises seems well settled. The Hadada Ibis is polytypic, the several subspecies differing somewhat in size and colour. In fact, another race, B. h. erlangeri was once recognized, which is now included in brevirostris: Related to this diversity, an interesting aspect of the Hadada Ibis is that the structure of its chromosomes (its karyotype) differs from that of other ibises, which is surprisingly variable anyway (see Classification). Capanna et al. (1982) have noted that karyotypic variability is found in taxa that are characterized by strict territoriality, small individual populations and gregariousness, traits that encourage inbreeding. These results suggest the possibility the Hadada Ibises have a higher rate of inbreeding than might otherwise be expected, leading to interesting questions of breeding biology and conservation. CONSERVATION This species has acclimatized well to human habitation (Wolstenholme 1961), and in fact has been a part of human culture for centuries (Schiiz 1970a). It is not subject, over the majority of its range, to persecution. The main danger to its population stability comes from extended droughts. Conversely where additional water tanks and ponds are constructed, this species will move in to colonize the newly irrigated areas where trees are available. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendi" dix — on page 313.

Wattled Ibis Bostrychia carunculata (Riippell) Ibis carunculata Riippell, 1837, Neue Wirbelthiere Fauna Abyssinien, Vogel, p. 49, pi. 19: Taranta Mts, Ethiopia

IDENTIFICATION A medium-sized, dark brown, crested bird, the Wattled Ibis is distinguished by the combination of its dark colour, its crest and its white wing-patch. It has a relatively short bill, a relatively long tail, and a small wattle hanging from its throat. The basic colour of the adult is dark brown, darker above than below, with a greenish gloss on the back. The crown is dusky, and a crest of the same colour extends behind the neck. The wing-feathers are black and glossy, the primaries having a bluish gloss and the secondaries, a greenish gloss. The upper wing-coverts are also black, except for the white wing-patch. The brown face is feathered with a dusky red bill. The red eye is surrounded by a white ring. The wattle is a thin, dusky red pendant, up to 20 mm long, which hangs from the lower throat. The legs and feet are also dusky red. The bill-length has been given as 11.0-12.7 cm (Friedmann 1930) and as 8.3 cm (de Naurois 1973), the difference suggesting that there may be a sexual dimorphism in the species. Friedmann also gives the wing-length as 32.5-38.0 cm and the tarsus as 6.3-6.6 cm; de Naurois gives the wing length as 32.538.0 cm. During courtship the soft-part colorations of the bill, wattle, legs and feet brighten. Nestlings are covered in black-brown down. They have bare skin around the eyes; their straight bill is grey with a black tip (Dorst and Roux 1972). In flight, like the other Bostrychia ibises, the Wattled Ibis is very noisy. Flying to and from roost, often in flocks, it gives a cry described as 'Kowrr-kowrr-kowrr'. This is louder than the call of the Hadada and, in a chorus, can be heard up to 5 miles away. Besides the raucous calls, roosting birds utter a series of grunts, groans, and squeaks (Urban 1978). The Wattled Ibis may be confused with other Bostrychia ibises. It is larger than the uncrested Hadada and is recognized in flight by its white wing patch. At closer range, the red pendant shows clearly. Its crest is less well developed than those of the Spotbreasted and Olive ibises but more developed than the uncrested Hadada Ibis. DISTRIBUTION AND POPULATION This species is resident all over the Ethiopian plateau of northeastern Africa. It occurs from about 1500 m to the high moorlands of about 4100 m elevation. The population has not been censused, but it has been common within its restricted range, especially near Addis Ababa and Bale. Flocks of over 100 birds were commonly reported. ECOLOGY Difficulty of access to the Ethiopian plateau region has meant that few observations have been made of the ecology of this species. The birds appear to be sedentary, often roosting near the breeding sites. They roost in trees singly, in pairs or in small groups, and sometimes in larger groups. They also roost on rock cliffs some distance from water (Brown et al. 1982:196-197). After dawn they fly to the feeding

198 Wattled Ibis

trees or even on the ledges of buildings (Fry et al. 1985). Schiiz (1970a) found a nest-building pair near Lake Awasa at 1700 m, which he considered was probably near the lower limit of nesting. The nest is made of branches and large sticks, lined with softer plant material. It is c. 20-50 cm across. On cliff edges, the nests may be close enough to touch or may be several metres apart. The breeding birds often choose to nest on eastern slopes, probably so as to obtain maximum early morning sun, because night-time temperatures at the altitudes at which they nest can be very cold (Dorst and Roux 1972). Males claim and defend territories, and females remain at the nest site. Laying dates are well dispersed, March-May, July and December, which include the small rains, the big rains and the dry season. The eggs were described by de Naurois (1973) and by Brown et al. (1982: 196-197) as dirty white, rough-shelled and marked with brown. On the other hand, Dorst and Roux (1972) described them as being olive green with brown markings. The eggs are 59-61 X 3940cm and weigh c. 50 g. Upon fledging, the young feed away from the main colony. grounds, where they forage singly or in flocks of up to 100 birds. The Wattled Ibis feeds in open grasslands, marshes, alpine woodlands, among crops or in forest glades down to 1500m altitude (Brown et al. 1982: 196-197). Their foraging behaviour is Walking while Probing the ground regularly. They will accompany herds of domestic animals, turning over dung heaps for beetles. If disturbed, they will rise as a flock, calling loudly. Their food consists of small prey including worms, insect larva, other insects, molluscs, frogs, small reptiles and mammals. BREEDING Nests are usually built in small or large colonies on cliff edges at over 2400 m altitude. Occasionally they nest at lower altitudes, singly or in small groups of 2 or 3 pairs, in

TAXONOMY The Wattled Ibis is the only montane, non-forest species of Bostrychia. Its habit of nesting on rocky cliffs and ledges is similar to the Geronticus ibises, and in this it may show considerable affinity to the latter genus (Fry et al. 1985). CONSERVATION Although previously a common species, within its very restricted range, long periods of recent warfare and drought raise concerns for this ibis. Little is known about its present status. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 314.

Olive Ibis Bostrychia olivacea (Du Bus de Gisignies) Ibis olivacea Du Bus de Gisignies, 1838, Bull. Acad. Roy. Sci. Lettres Beaux-Arts Beligique, 4 (1837), p. 105 and pi: 'Cote de Guinee' (=upper Guinea, as determined by Chapin, 1923, Amer. Mus. Novit., no. 84, pp. 1-3) Subspecies: Bostrychia olivacea olivacea (Du Bus de Gisignies) Bostrychia olivacea cupreipennis (Reichenow) Theristicus cupreipennis Reichenow, 1903, Ornith. Monatsber., 11, p. 134 Bostrychia olivacea rothschildi (Bannerman) Lampribis rothschildi Bannerman, 1919, Bull. Brit. Ornith. Club. 40, p. 6: Roca Infante D. Henrique, Prince's or Principe Is. Bostrychia olivacea bocagei (Ghapin) Lampribis bocagei Chapin, 1923, Amer. Mus. Novit., no. 84, p. 5, figs 2A, 3A: Rio de Sao Tome, Sao Tome Bostrychia olivacea akleyorum (Chapman) Oreoibis akleyorum Chapman, 1912, Bull. Amer. Mus. Nat. Hist., 31, p. 235, pis 23-24: south slope of Mt Kenya, altitude 9000 ft Other names: African Green Ibis, Green Ibis, Kenya Ibis, Dwarf Ibis and Dwarf Olive Ibis (bocagei) (English); Gallinhola and Gallinho do Matto (bocagei) (Portuguese); L'ibis Olivatre, Ibis Olive a Crete (French)

IDENTIFICATION The Olive Ibis is a small to medium-sized, dusky brown ibis with a prominent crest and red face and bill. The back is a dusky brown with a green iridescent tinge. It is very dark brown below, which also is tinged with green. The very obvious head-crest is dark brown with a purplish sheen to the long feathers that extend well back from the top of the head. Secondaries and wing-coverts show green, bronze and pinkish lustre; the tail is dark blue. The bare face and bill are coral red, and the red eye is set off by blue-black lores. The legs and feet are described as yellowish green to pinkish brown and also as dull dark red (Brown et al. 1982). It seems certain that the differing descriptions of the soft parts refer to birds in different stages of breeding coloration. These colour changes need to be better documented. The subspecies differ in details of coloration and geographical range. The subspecies olivacea has faint pale striping on the head and neck. There is more green on the neck and body of cupreipennis. The crest of rothschildi is more purple. The subspecies bocagei is duller plumaged and very much smaller than olivacea. The subspecies akleyorum is larger than olivacea, and its body is noticeably greener. Other than noted above, there is a considerable overlap in size among the races. In view of the small number of specimens, patterns of size variation cannot be considered definitively known. The very limited data for rothschildi suggest that males are larger than females, but this needs to be confirmed. In other ways there is no indication that sexes differ.

Olive Ibis 201

(7) olivacea (lowland forest); @ cupreipennis (lowland forest); @ akleyorum (mountain forest); (4) bocagei (Sao Tome Island only).

The nestlings have dark brown down. Immature birds lack the long crest and are duller coloured than the adults. The voice of the Olive Ibis is a strong resonant double squawk, rendered *Gar-wa, gar-wa*, and also a goose-like call heard at a long distance (Brown et al. 1982). When annoyed it will give call 'Gar' at an intruder (Meinertzhagen 1937). This ibis has a straggling flight and travels high over the canopy at dawn and dusk calling loudly as it flies. The Olive Ibis looks very similar to the closely related Hadada and Spotbreasted ibises. Although the Spotbreasted Ibis has its neck and underparts well spotted and also has a green spot in front of the eye and a green streak crossing the face in line with the bill, A. Brosset (pers. comm. 1990) has stressed to us that the Olive and Spotbreasted Ibises are practically impossible to distinguish by plumage and size in the field. Similarly, many observers have confused the Olive Ibis with the more familiar Hadada, especially when the Olive Ibis is in deep or mottled shade, where little of its iridescent colouring is visible. It appears to fly more heavily than the Hadada Ibis and appears to have a thicker neck (Meinertzhagen 1937). In many cases, voice is the best character for separating these species. The call of the Spotbreasted Ibis lacks the honking quality of the Olive Ibis; and, as Chapin (1932a) pointed out, the Olive Ibis does not stress the second note as does the Spotbreasted Ibis. The Hadada has three distinct notes instead of the two of the Olive Ibis. (Tapes of calls are listed by or available from Chappuis 1980, The British Library of Wildlife Sounds, and Keith and Gunn 1971.)

DISTRIBUTION AND POPULATION This widespread sedentary species has seldom been reported in the wild, especially in recent years, so very little is known about the details of its distribution and nothing about its population levels. It may be more common than we can document because its habitat is so inaccessible and because it is so difficult to identify with certainty in the field. The subspecies olivacea is found in West Africa in Sierra Leone and Liberia; cupreipennis is also found in West Africa, in Cameroon, Gabon, Congo and Zaire. The subspecies akleyorum is found in the mountains of Kenya and Tanzania, particularly Mount Kenya and the Aberdares, the Usambara Hills, and probably Mount Meru (F J. Jackson 1938, Snow 1978). Collected only from the Gulf of Guinea Island of Principe (Prince's), rothschildi is now thought to be extinct. The other island subspecies, bocagei, is found only on the nearby island of Sao Tome. In 1988, John Burlison and Peter Jones, along with Ana-Maria de Sa Almeida and Claudio Menezes do Espirito Santo, visited southwestern Sao Tome and received credible accounts of the survival of this race (World Birdwatch 1988). In June, 1989, Jan Haft and Jacob Fahr reported seeing birds of this species along the lo Grade River, about 4 kilometres north of Ribeira Peixe (World Birdwatch 1989). The reports of these birds, the first since a specimen was collected in 1928, are one of the more exciting discoveries in ibis conservation of the past decade. ECOLOGY The Olive Ibis is a solitary species of deep forest, but has a rather intriguing habitat distribution over its range (Snow 1978). It is widespread in the lowland forests of West Africa but is absent from the Cameroon highland forest and from the mountains forming the eastern edge of the Congo basin. Yet it is a montane bird in Kenya and Tanzania, where it occurs upwards from about 1800 metres. In West Africa it occurs with the Spotbreasted Ibis, but in East Africa it appears to be replaced at low altitude by the Hadada Ibis (Snow 1978). Certainly akleyorum is more often seen than the lowland races to the west. We found akleyorum near streams in the montane forests of Kenya. Vincent C. Fayad has reported seeing it more often on the Sirimon route on Mount Kenya than on the Naro Moru route. In September 1988, he encountered 10-12 birds at an altitude of 3000 m near the Sirimon road head (V.C. Fayad pers. comm. 1989). I.S.C. Parker (pers. comm. 1990) also reports that this species is found in some numbers in the Hegenia abyssinica forest of some 100-200 acres, surrounded by moorland, between 2700 and 4000 m in the Aberdare Mountains. We have searched, unsuccessfully, for olivacea in the forest of West Africa, but have only succeeded in attracting very large numbers of leeches which fall from the trees in large numbers! However the fast running streams strewn with boulders, which we found there, mirrored the similar East African streams where we have observed akleyorum. The other western subspecies, cupreipennis^ occupies a similar lowland forest habitat.

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Olive Ibis

On Sao Tome, Burlison and Jones (pers. comm. 1990) reported that the endemic bocagei occurs in the remote, forested regions at high altitude. They reported that local people had never seen it in tall forest trees grown on longabandoned oil palm plantations but only in the 'abo', or untouched primary forest. In June, 1989, 5 individuals were observed on lava gravel along the Rio Grande River (World Birdwatch 1989). In its forest habitat, the Olive Ibis feeds along the mountain streams and riverlets, although Bannerman (1953) encountered it in the vicinity of larger rivers such as St. Paul's River in Liberia and the Gamma River in Gabon. I.S.C. Parker (pers. comm. 1990) suggests that the Hegenia forests of Kenya contain a number of birds, and indicates that they prefer to forage on the forest floor where mature trees have little or no undergrowth. The species is described by Bannerman as having regular habits, feeding in marsh, swamps and backwaters and returning to high trees, such as a silk-cotton, to spend the night. It prefers to roost in such very large trees, with dead tops (Brown et al. 1982). Bannerman (1953) stated that when disturbed it takes refuge in thickly leaved trees. It travels high above the canopy at dusk and dawn calling loudly. It is silent throughout the day when feeding, and so is difficult to locate. Nothing is reported regarding feeding techniques of the Olive Ibis. It appears to feed in pairs or small groups (W.L. Sclater and R.E. Moreau 1933a). Its food consists of insects such as beetles, larvae, grubs, snails and worms. A male specimen from East Africa contained beetles, and a female, green leaves, beetles and segments of Myriopods (FJ. Jackson 1938). It is reported by Brown et al. (1982) to eat snakes and some vegetable matter from the forest floor. BREEDING The earliest, and one of the few, reports of breeding is the account of Mr and Mrs Carl Akeley who found a nest of the East African race in 1910 on the southern slopes of Mount Kenya in dense forest at about 3000 m elevation (Chapin 1923, F.J. Jackson 1938). It was seen in a bamboo forest near their hunting camp. The nest was loosely constructed of dead branches on the limb of a small tree about 8 m from the ground. The nest contained 3 young at the time, and a fragment of an eggshell which was pea-green and more or less stained with cinnamon rufous and irregular blotches of chestnut. It was much less heavily marked than the egg of the Hadada Ibis (F.M. Chapman 1903). No further nests were discovered until 1959, when A. Dyer found one at the headwaters of the Pesi River in the Aberdares (Kenya) in a mixed Juniperus-Podocarpus forest (I.S.C. Parker 1982). This section of the Aberdares is relatively dry forest, being in the range's rain shadow, and the Pesi is a very small stream at this point, no more than 2 m across. This nest was in the fork of a slender branch over a pool about 2 m below a small waterfall. The structure was flimsily made, of large twigs between 30 and 40 cm across. Both birds were observed building, which is accomplished silently.

During a visit in 1980, we discussed the matter with ornithologists in Kenya, remarking on the sparsity of sightings of this species. The next year, Mr I.S.C. Parker was able to find a nest again in the Aberdares (I.S.C. Parker 1982). The nest was observed in July and September at a small stream near the Thiririka river, on the southeastern slopes of the Aberdares in mixed Podocarpus and Ocotea forest. This nest was also over water, precariously placed about 4 m high on a very slender branch 2 cm thick and partially supported by several even smaller branchlets and a vine. It was about 30-40 cm in diameter, and made of large twigs such that it could be seen through from the ground. In late July it was empty (although what Parker thought to be a leaf when viewed through the nest bottom could have been an egg), but by the first week in September contained 2 halfgrown nestlings (I.S.C. Parker 1982). When visited by Ian Parker and Peter Davey later in the month, the nest had largely fallen down. Since that date, no further observations have been made due to heavy mists and rainfall at that time of year (P. Davey pers. comm.). No other nesting records are known, and there remains no record of nesting from West Africa. Almost nothing is known of the nesting behaviour of the species. The sounds of nearby running water may help mask any noise made by the birds during nesting. TAXONOMY The Bostrychia are very similar, and since few specimens are available, much about their taxonomic relationships remains to be understood. It was once suggested that the Hadada Ibis and the Olive Ibis were the same species (Akeley in F.M. Chapman 1912). As it is so much smaller, Bannerman (1930) thought bocagei distinctive enough to be considered a different species. The subspecies rothschildi is considered now to be extinct and few specimens are available. The presently recognised Bostrychia species were once separated into several genera (Hagedashia, Lampribis and Bostrychia), and appear to be related to the Madagascar Crested Ibis (Lophotibis) and the Geronticusibises (Amadon 1953, Appert 1966, Fry etal. 1985). Clearly, the taxonomy and relationships of the ibises in this group need additional study. CONSERVATION This species inhabits deep forest areas that are rarely visited, and as a result its status is almost completely unknown. It is thought that one race is already extinct, and another has only been seen once by biologists in recent years. Present dangers appear to include predation by forest birds and mammals, especially baboons (Papio anubis, P. cynocephalus], as well as the cutting of forest trees. On Sao Tome and Principe Islands, deforestation is a major problem, especially since it appears that the birds are found only in virgin forest. Dr P. Jones (pers. comm. 1990) reported that the Sao Tome natives hunted the Olive Ibis in the last century, when they were a 'favourite quarry'. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 315.

Spotbreasted Ibis Bostrychia rara (Rothschild, Hartert, and Kleinschmidt) Lampribis rara Rothschild, Hartert, and Kleinschmidt, 1897. Novit. Zool., 4, p. 377: Denkera, Ghana Other names: Crested Ibis, Crested Wood Ibis (English); Kala (Congolese); L'Ibis Vermicule (French)

IDENTIFICATION The Spotbreasted Ibis is a small (59 cm long) short-legged crested ibis confined to deep forests. It is a dark-plumaged bird recognized by its crest and characteristically spotted neck and underparts. The adult is brown with a long, loose crest, darker than the base colour and glossy green. The neck, breast and underparts are reddish brown; the feathers are edged in dark brown with a central cream spot. The mantle is also brown, edged in buff but with a glossy green sheen. The rump and tail are blue green. The wings are a blue-black, the underwing black. The scapulars and inner secondaries sport an iridescent bronze green. The bill varies between red and dark red, whereas the naked face and lores are black, with a green to turquoise spot in front of the eye and another behind it. A green streak crosses the face in line with the bill. The eye is dark brown; legs and feet are brown with a pinkish flush. Typical measurements are: wing 27.0-29.0, tail 11.2-12.0, bill 11.5-13.0, tarsus 5.6-6.5 cm (Brown et al. 1982). The species is sexually dimorphic, the bill of the female being smaller than that of the male. Brosset and Erard (1976) noted that they observed the female to be less well marked on the head: the turquoise spots and the facial streak were smaller, and the bill was not as bright red as the male. This singular observation of sexual differences in soft parts may not be definitive as the female's courtship colours may have started to fade before the male's. In any case, it would be expected that any difference in markings would become inconsequential as nesting progressed. Nestlings have red-pink skin, lightly purple-blue, and are covered with a sparse dark brown down. The face is blackish blue and the chin pink. White down replaces the brown down after about 6 days, when the sheaths of contour feathers begin to appear. The base of both mandibles are red. Immature birds are similar to adults though with shorter bills, less pronounced markings and no metallic sheen. This ibis is, like other Bostrychia, extremely noisy, especially at dawn and dusk when leaving or returning to roost, and even at night (Brosset and Erard 1976). It is quiet during the day, while feeding and at the nest. Brown et al. (1982) described its raucous call as 'K-hah, k-hah'. The second note is accented, as was pointed out by Chapin (1932a) and Bannerman (1953) but missed by Brown et al. (1982). When flying, the call may be given with alrpost every wing-beat (Bannerman 1953). The call differs from those of the other Bostrychia ibises. Ash et al. (in press) were able to identify this species in a new area of Uganda by its call, which lacks the honking, goose-like quality of the Olive Ibis. A tape of the Spotbreasted Ibis by C. Chappuis (pers. comm. 1990) presents one bird with a soft voice and a second one much harsher, with both birds accenting the second, or a third, syllable. This species clatters its mandibles during nest relief and when threatened. The Spotbreasted Ibis is distinguished among ibises by the speckling on its neck, breast and undersides. The sympatric Olive Ibis is larger, lacks spots on the face and has purple on its crest. The Hadada Ibis, which lacks a crest and is more likely to be found in the open, is also larger. All these marks might be apparent close by and in good light, but the Spotbreasted Ibis's wooded habitat and crepuscular flying habits make it difficult to distinguish in the field from either the Olive or Hadada ibis. A. Brosset (pers. comm. 1990) considers them to be almost impossible to distinguish in the wild

Spotbreasted Ibis 205

ent in an area year-round, and use traditional roosts for sleeping (Brosset and Erard 1976).

except by voice, and J.S. Ash (pers. comm. 1990) certainly confirms this with respect to distinguishing the Spotbreasted from the Olive Ibis in Uganda.

DISTRIBUTION AND POPULATION This species is found in West Africa from Liberia through Ghana, and from Cameroon and Gabon through Zaire to western Uganda. The presence of the species in Uganda was recently discovered by Ash et al. (in press), where they found it on the banks of the Kirimia River. The Spotbreasted Ibis is seldom seen and is considered to be rare, although widespread. It is most often reported from northeastern Gabon, but much remains to be understood about the details of its population status or distribution.

ECOLOGY The Spotbreasted Ibis occurs in lowland Guinean and Congo forests, where it is seldom seen, and so it is not well understood (Bannerman 1953, de Naurois 1973). In Uganda, Ash et al. (in press) found it along the river bank associated with Afromomum swamps. Bannerman (1953) stated that it is a bird of small forest streams and wooded swamps and avoids the larger river banks and forest clearings. It is primarily, although not entirely, a bird of virgin primitive forest areas subject to periodic flooding. This ibis feeds in pairs or alone by forest streams and in wooded swamps and pools. It Probes in mud and other soft substrate, consuming aquatic snails, worms, beetles and grubs (Bannerman 1953, Bouet 1955a). The birds are pres-

BREEDING This species nests alone in the thick forests in which it is found year-round. It apparently always nests near or over water within the forest. Nesting takes place when water is rising or is high and appears to correspond with peak rainfall periods. In Gabon, these are September-December and March-May (Brosset 1974). Brosset and Erard (1976) found that the two periods of maximum nesting activity coincide with the two rainy seasons, with some minimal activity also in the short dry season of January to February but not in the longer dry period from June to August. They nest in September and November in Zaire, also the time of the big rains (Lippens and Willie 1972)* However, nesting can probably occur at any time of year under appropriate conditions. Brosset and Erard (1976) suggested that initial pairbonding may take place in the traditional roosting sites. As it is also likely that members of the pair remain together year-round, courtship would be expected to be perfunctory. On arrival at dusk, the birds circle around, changing positions frequently, perching on the main branches of trees. 5-8 birds perch side by side, bowing by lowering the head and bill with crest feathers erect. Pairs touch bills and mutually preen. The nest is a small, crude platform of twigs very similar to those reported for other Bostrychia, but smaller than that of the Hadada Ibis and bulkier than that of the Olive Ibis (F.M. Chapman 1912:24, Brosset 1974, I.S.C. Parker 1982). It is some 30 cm across and between 5 and 15 cm thick; the lining is of dead leaves, fibre, roots or fresh bits of epiphytes. One nest was made entirely of liana shoots. The first nest to be found was brought to P. Herroelen (1955:187). It had been built at a height of about 3 m from the ground near a stream in secondary forest. It was made of short, dry branches and lined with fibre and roots. Brosset and Erard (1976) observed a nest that was 80 cm above the level of the water in the centre of the mouth of a stream that was about 4 m deep during the rainy season (0.8 m in the dry season). This nest was on the main horizontal stem of one of the lateral branches of a leaning shrub located at the edge of an area of dense flooded vegetation. Another nest was in a shrub, 4.5 m above water at the edge of the Ivindo River, and a further one was 6 m above the water in a marsh on the Libumba, a tributary of the Ivindo. Other empty nests were found at heights from 1.5 to 3.0m in areas subject to flooding but which were dry during Brosset's and Erard's (1976) visit. Although the nest is not hidden, it is effectively screened by thick foliage, and the building, laying, incubating and nest-relief are all carried out in a secretive, quiet manner. The normal clutch size appears to be 2. The egg is pale green, spotted with small irregular rusty markings, lightly glossed and fairly large—55 X 35 mm. Both sexes incubate, incubation taking about 20 days. It appears that the female broods the eggs at night, the male replacing her in the

206 Spotbreasted Ibis

morning and early afternoon. During incubation, except for occasional head or wing tosses to ward off insects, the bird is immobile with its neck folded back and its bill placed under the wing. It takes advantage of the need to turn the eggs for stretching and shaking. If threatened, the sitting bird will stretch out its neck and clatter its bill strongly. On three occasions, monkeys (Cercopithicus neglectus) passed near the nest being observed. Twice the bird flattened itself against the nest, remaining immobile. The third time, the bird, having just risen to turn the egg, adopted an upright position with plumage tight, neck extended and bill pointing upwards. It remained in this position for up to 2 min without moving. The nest observed was robbed of its young 9 days after hatching, but it is not known whether the predator was animal or human. A. Devez (in Brosset and Erard 1976) described sexual differences in nest-relief after hatching, when the mate comes to feed the young and replace the sitting bird. A female arriving will raise her plumes, lower her bill and make clattering noises, inserting her bill under the sitting male to encourage him to rise. Bowing the body slowly causes the male to rise and bill clatter with slightly ruffled feathers. The young are then fed by the female. Should the male not then leave the nest, the female extends her neck, raises both crest and plumage and clatters loudly with her bill until he departs. When a female is sitting and is relieved by the male, he will pass his bill across her neck and they will face in the same direction, bill clattering simultaneously. Sometimes this ceremony varies, with the male passing his bill through the neck feathers of the female. Feeding of nestlings is by regurgitation. Young chicks are stimulated by the adult's tapping of the chick's chin and clacking of its bill. The young bird then taps the adult's bill starting at the tip and moving upward to the base then inserting it into the open bill of the adult, whose crest is often raised. As the chick grows, the initial stimulation of

the adult is no longer needed. Chicks Beg food by making hissing noises described as shrill but only faintly audible. Indeed the whole ceremony, including feeding and nestrelief, is done quietly and with minimum noise and movement. It is apparent that all activity at the nest is extremely discreet, and creates the minimum of attention (Brosset and Erard 1976).

TAXONOMY Little is known about the biology, and hence the taxonomy, of this species. Its relationships with other species, especially in the genus Bostrychia deserve additional study.

CONSERVATION The status of Spotbreasted Ibis is really unknown. It appears to be more abundant than does the Olive Ibis, which however is overall more widely distributed. Hunters are known to collect young from nests, and it is likely that native hunters, as well as predators such as monkeys, present the major threats to the species, but data are insufficient to establish whether it is under severe threat other than from man. Any encouragement to local hunters to capture young for collections at museums or zoos should be resisted to avoid the destruction of this rare species. The forest of the Spotbreasted Ibis is rapidly shrinking, so they will eventually become vulnerable through loss of their habitat.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 316.

Madagascar Crested Ibis Lophotibis cristata (Boddaert) Subspecies: Lophotibis cristata cristata (Boddaert) Tantalus cristatus Boddaert, 1783, Table Planches Enlum., p. 51; based on 'Courly hupe, de Madagascar' of Daubenton, 1765-81, Planches Enlum., pi 841: Madagascar Lophotibis cristata urschi Lavauden Lophotibis cristata urschi Lavauden, 1929, Alauda, 1, p. 233: Ankarafantsika and 100km SE of Majunga, western Madagascar Other names: Crested Wood Ibis, White-winged Ibis (English); Akohoala, Akohovohitra, Akoholahialu, Lampirara (Malagasy); Coq des Bois, Ibis Huppe de Madagascar (French); Madagaskar-Mahnenibis (German)

IDENTIFICATION The shy Madagascar Crested Ibis is a medium sized (50 cm long) rather handsome white-winged ibis, with a long neck, long bill and long crest. It is the largest bird in the Madagascar forest. The basic body colour of the adult is a heavy rufous brown. The crest on the nape and back of the relatively long neck has a metallic sheen, the colour of which differs between the subspecies. The chin, throat and neck are brown. The wing is mainly white. The bare face is bright red, including around the eye. The rather long bill is horn colour. The iris is dark, and the tarsus and foot are red. The sexes are alike. Breeding colour changes have not been well described, but it is likely that the head, feet and crest colours brighten during courtship, as is the case in most ibises. The crest of the nominate subspecies consists of long feathering of dark metallic green and white. The subspecies urschi has a deeper reddish brown body colour, and its throat, neck and crest are a metallic green mixed with orange yellow and less mixed with white (Langrand 1990). Individuals having intermediate characteristics are known. Nestlings are initially all white turning brownish. The bill is grey, darker toward the tip. The bare skin and feet are pale reddish yellow. Juveniles have a paler body colour than the adult, speckled brown on the white wings, and white on the tail. The head is very pale brown. The grey bill is shorter and paler, darkening from the base. The skin about the eye is grey, the eye itself is brown (Rand 1936). The feathers on the back of the neck are long and blackish blue, but they lack the metallic sheen. The legs are a paler red. The voice is usually heard at dusk or when birds are flying to roost, but also when alarmed. It is described as a monotonous, subdued, toneless 'Goo, goo, goo' by Langrand (1990) and as 'Grooh' by Appert (1966). A call was described as a loud, far-carrying, slightly raucous 'Work, work, work' by G.S. Keith et al. (1974). Rand (1936) stated that the bird can sometimes be heard calling during the day,

Madagascar Crested Ibis 209

plantations. The subspecies urschi lives in dry deciduous forest habitat, from sea level up to 2000 m above sea level. This shy bird feeds alone and, more usually, in pairs. The birds forage on the forest floor, by Walking through the undergrowth. They stop to Probe amongst dead leaves and forest litter, in the moss and around tree trunks. They also Peck at the surface picking up insects and millipedes from among the dry leaves (Gill in G.S. Keith et al. 1974). They particularly feed along wooded streams at low water or in debris covering the dry stream bed. Generally, the diet of the Madagascar Crested Ibis consists of invertebrates and small vertebrates (M.E. Nicoll and O. Langrand unpub. report). Of 13 stomachs of cristata examined, 10 contained various insects, including beetles, one centipede, one spider and three earthworms. In one stomach of urschi,, Rand (1936) found two 100 mm long lizards, one 250 mm long snake and various insects. The Madagascar Crested Ibis roosts on strong branches, and flies to the top of large trees when disturbed. It prefers shaded woodland areas. and described the sound as a loud, rather creaky *Ack, ack'. Appert heard a bird at the nest give a quiet *Kr'. The brown plumage makes this bird inconspicuous on the ground, especially in its forest habitat. However, the white wings flash strikingly when the ibis is in flight. Rand (1936) saw 2 or 3 birds flying to a tree top with their wings flashing conspicuously in the brilliant moonlight. When flushed, it often flies to a perch in a nearby tree. Or it may walk quietly further into the woods. In flight, it cannot be confused with any other species in its restricted range, as its wide, white wings contrast with its dark body. It is the largest living terrestrial bird in Madagascar, similar in size to the distinctive Helmeted Guineafowl (Numida meleagris). DISTRIBUTION AND POPULATION The Crested Ibis is endemic to Madagascar, where previously it was distributed throughout the island, except in the dry country of the far southwest. The subspecies cristata is found in the east and north. The subspecies urschi occurs in the west and south. It is present throughout the year, at least in some areas (Appert 1966). It appears to be an entirely sedentary bird. No estimate is available of population size. Rand (1936) found the species to be common, but G.S. Keith et ai. (1974) reported it to be 'distinctly uncommon' in 1972. The western subspecies remains widespread and common (Appert 1966, Turner in Collar and Stuart 1985). ECOLOGY A forest ibis, this species is found in every type of original forest on the islands and is not necessarily restricted to wet habitats. In the east cristata is found in humid forest, in the wet undergrowth of the dense rain forests by small forest streams. It has also acclimatized to slightly degraded forest, being found in thick and shady areas in vanilla and oil palm

BREEDING This species is entirely dependent on tall trees for nesting in the forest. The breeding period is from September to January according to Langrand (1990), and certainly includes November since a female ready to lay was taken at Tabiky on 2 November 1929. Appert (1966) observed nesting in January and in November. A juvenile was collected at Marotony on 2 January 1931 (Rand 1936). Unfortunately little is known of the courtship behaviour. Appert (1966) saw one bird of a pair display on the ground. The bird ruffled its head, neck and chest feathers, and held its wings out pointing to the ground. It took up one or more dry leaves and held them for a relatively long time, and also played with them and dropped them to the ground. The distinctive crest is presumably used in courtship displays, but it is kept lowered most of the time, except when disturbed or alarmed. Pairs nest solitarily. The nest is made of long sticks, 12 cm in diameter. It is a large structure, up to 85 cm in diameter, the cup being about 30 cm wide and 8 cm deep. There is a lining of leaves and fine sticks. The nest is constructed in the shade high up in the fork or on several branches of a tree, as many as 7-15 m up in rain forest trees (Appert 1966). Appert (1966) found a nest in a tamarind tree (Tamarindus indicus) which was in leaf at a time when other trees in the area were bare. The eggs are a matt white; they measure, on average, 41 X 58mm. The usual clutch is 2-3, and both parents share incubation (Langrand 1990). The incubating bird sits tightly on the nest when approached from below. Young in the nest gave a Teh' call when encountering a person there; they also raise their wings and bristle their feathers. They remain in the nest for 20-30 days. TAXONOMY The genus Lophotibis^ whilst superficially resembling Bostrychia (= Lampribis, Hagedashia), continues to be treated by us as a separate genus pending additional study.

210 Madagascar Crested Ibis

The fact that distinctive differences exist in populations on the same island is somewhat surprising. However, the two forms occupy different habitats, and both apparently are entirely sedentary. The forms are separated by the wide central savanna, which stretches down the centre of Madagascar.

food or captured, for payment of a small fee (W.B. King 1981). It is also threatened by forest habitat destruction, including forest fires, as are much of the native fauna and flora of Madagascar. The protection of forest habitat is essential to the continued existence of this and other forest species on the island.

CONSERVATION The Madagascar Crested Ibis was the only bird endemic to Madagascar to be afforded legal protection, as of 1974 (G.S. Keith et al. 1974). However, it is still widely shot, hunted for

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 316.

Sacred Ibis Threskiornis aethiopicus (Latham) Subspecies: Threskiornis aethiopicus aethiopicus (Latham) Tantalus aethiopicus Latham 1790, Index Ornith., p. 706: 'Aethiopia' = PEgypt (Bruce, 1790, Travels Source Nile, 5, p. 172, pi. 35) Threskiornis aethiopicus bernieri (Bonaparte) Ibis bernieri Bonaparte, 1855, Consp. Gen. Avium, 2, p. 151: Madagascar Threskiornis aethiopicus abbotti (Ridgway) Ibis abbotti Ridgway, 1893, Proc. US Natl. Mus., 16, p. 599: Aldabra Island Other names: White Ibis, Black-necked Ibis, Sticklebill (English); Heilige Ibis (Dutch); Heiliger Ibis (German); Ibis Sacre (French); Ibis Sagrado (Spanish); Helig Ibis (Swiss); Ibis Sacro (Italian); CBHineHHbiH Hbnc: "Ibis" (Russian)

IDENTIFICATION The Sacred Ibis is a medium-sized, white ibis, 65-75 cm long, with black highlights, including its head and neck, wing-tips and ornamental plumes along its back. The adult's head and neck are bare and covered with a black scaly skin. The iris is brown and the lower eyelid pale pink. The downcurved bill is black, and rather heavy looking for its size. Grey grooves are quite apparent along either side of the mandibles. The distinctive plumes originate from the scapulars and form a loose lace-like net across the back, closed wing and tail. When new, the blacktipped primaries and secondaries are glossed with purple-blue and sometimes with iridescent bluegreen. A dull grey bare spot at the sides of the breast extends as a strip across the leading edge of the underwing. The bare skin of the legs and feet also are black. The island subspecies are smaller than birds on the African mainland. The subspecies bernieri has a light blue eye and less black on the wing-tips. The subspecies abbotti has a bright blue eye, black wingstripe and reduced feather pigmentation on the wing tips. The sexes are alike in appearance, but males are larger than females. Brown et al. (1982) reported that the bill-length of males is 162-183 mm, whereas that of females is 135-157 mm. Urban (1974a) reported that the differences were not sufficient to tell the sexes apart in the field except when both birds of a pair were together or when a displaying group formed. In courtship, the black scapular plumes develop a metallic bluish sheen. The bare skin underneath the wings and the similar patch next to the breast become a vivid red. The black skin on the head and neck becomes glossy, and the black legs flush red or dark red. A red ring becomes apparent in the iris. Urban (1974a) reported that the plumage of birds nesting at Lake Shala became tinged with yellow or brown, and underparts generally become stained as breeding proceeds. There are some differences between subspecies in courtship plumage and colour. The ornamental plumes are more developed in the subspecies aethiopicus but are less so in the subspecies abbotti and bernieri^ in which the black plumes are also less intense during nesting. The African subspecies also has a neck-sac, apparently absent in the other two races, which is especially obvious when half-open bills are pulsated during hot weather such as when the bird is on the nest.

Sacred I bis 213

heavy bill. In flight it can be distinguished from herons and egrets by its quicker wing-beat, and from storks because its legs are hidden beneath its tail.

Nestlings have short, straight bills which are pale white or pink, as are the feet. They are covered with a white down, but the head, neck and back are black. At 21 days the dorsal and lateral surfaces are covered with a brownish black down (Urban 1974a). Wing-feathers are black as they develop, and legs become greyish and distinctively swollen. Juveniles are whitish with black speckling or streaking, especially on the head and neck. Their wing-tips lack metallic gloss, and the scapulars tend to be greenish brown. The iris is brown with a narrow, outer, white ring. The feet and legs are dull grey to blackish and these darken as the birds get older. The bill takes 3 months to develop fully. Loss of neck feathering and acquisition of full adult plumage is not complete until the third year (FJ. Jackson 1938). The Sacred Ibis is generally a silent bird but sometimes utters a harsh groan or croak, especially when in flight. In antagonistic encounters, both sexes call and squeal. Urban described these calls as 'Whoot-whoot' and 'Pyuk-pyuk' and also a wheezing 'Hnhhh-hnhh'. The female gives a 'Whaank* call in soliciting copulation. Adults call to young with a 'Turrooh' (Cramp 1977). Like all members of the Threskiornis genus, these birds have strong, powerful wing-beats, which they alternate with gliding, achieving 38 km/h (Urban 1974b). The head and neck are fully stretched, and they fly in loose V-formations or in line astern. They usually fly low over water but can soar very high over land. R.A. Clark and A. Clark (1979) report them travelling up to 30 km away from colonies when feeding. The ontogeny of landing behaviour has been studied by G. Riippell (1977). The Sacred Ibis is rather easily distinguished from other ibises in its range by its white plumage, black head and

DISTRIBUTION AND POPULATION The Sacred Ibis is a widespread resident in Africa south of the Sahara desert, from Senegal in the west to Ethiopia in the east, and southward to the Cape. Outside Africa, it breeds only in Iraq, especially in southern areas of Amara to Fao, although definite nesting records are few (Allouse 1953 in Cramp 1977). It was formerly found in Egypt, from which country the type specimen and English name came, but it had almost completely disappeared from there by 1850 (Meinertzhagen in Cramp 1977). The subspecies bernieri is only found on Madagascar, especially on the west of the island between Port-Berge and Moromoe. The subspecies abbotti is endemic to the islands of Aldabra, including West and Middle islands and lie Polymnie. African birds move around seasonally. There is a definite movement of birds towards the equator in Africa, with northern birds moving northwards and birds south of the equator moving southwards at the commencement of breeding, but movements vary depending upon seasonal rainfall. Ringing records from South Africa show that birds ringed in that country were recovered mostly in Zambia, with fewer in Botswana and Namibia. The furthest travelled 1336km (Dowsett 1969, R.A. Clark and A. Clark 1979). Birds in Iraq are present year-round but probably also wander to Kuwait and Iran. After breeding birds disperse, moving in the dry season especially to coastal areas and permanent bodies of water, travelling considerable distances to traditional winter feeding grounds. Numbers in these areas fluctuate seasonally and annually. Neither of the island subspecies show signs of either dispersal or migration. The species occurs rarely in southern Arabia. Vagrants have appeared in the USSR on the Caspian and Black sea coasts, and 7 birds were seen in 1944 south of Baku, and it is thought that they may breed in Khuzestan (Dementiev and Gladkov 1951, Cramp 1977). It is possible, however, that the birds seen in these areas were Oriental White Ibises. Bururlin (1935 in Dementiev and Gladkov 1951) considered them to be 'Indian ibises' (Oriental White Ibis), but Dementiev disagreed, having seen them at close quarters. That Eurasian Spoonbills migrate annually between the Caspian Sea and Bharatpur in India raises the possibility that Oriental White Ibises might do the same. Few population estimates exist, but the species is common over much of the area in which it occurs. On the Witwatersrand in South Africa, numbers fluctuate between 4900 in summer and 2500 in winter (Dowsett 1969, Tarboton 1977b). Elsewhere, feeding flocks vary from several hundred birds to groups of 10 or 12, or even single birds. They roost in the hundreds on islets in rivers or floodland, on trees near water and sometimes even in villages. Their dispersal in small parties make assessments of numbers difficult over their large African range. Bernieri is uncommon

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(Langrand 1990). Abbotti numbered 150-200 in 1967-68 (W.B. King 1981).

In Witwatersrand, South Africa, only 3 of 28 roosts were used year-round (Tarboton 1977a).

ECOLOGY

BREEDING

This species is found in many habitats. It occurs in all types of freshwater habitat and often feeds on dry land in grasslands and cultivated fields. Particularly in the dry season, it frequents coastal mudflats. On Lake Turkana in northern Kenya, we have seen birds feed on the rocky islands and shores around the lake. Individuals wander some distance into the desert areas where they feed on insects. At Lake Naivasha, Kenya, they feed on hotel lawns, and along the muddy shores of the lake. We have seen individual birds feeding alongside Brown-necked Ravens (Corvus corax edithae) scavenging at camps and villages. In South Africa, they feed in sewage, dung heaps and refuse, as well as cultivated fields, flooded grasslands and marshes (R.A. Clark and A. Clark 1979). Abbotti feeds in coral pools and along lagoon shores (W.B. King 1981). They respond quickly to rainfall or flooding of potential feeding habitat, moving into such areas immediately after water rises (R.A. Clark and A. Clark 1979). Such flooded feeding areas seem to be critical for the initiation and successful completion of nesting. The Sacred Ibis has simple methods of feeding. It will Peck at the surface for insects on dry land, and Probe or Deep Probe for crustaceans and aquatic insects in mud and soft soil. Deep Probes can result in the bill being buried up to the eye. They will Probe into carrion. Occasionally they will run forward with wings half open to catch a flying insect or moving reptile. They will also follow locust or grasshopper swarms. Although they do sometimes feed alone, Sacred Ibises more typically feed in groups. In shallow pools, they are often found alongside African Spoonbills and Yellowbilled Storks. They follow Little Egrets (Egretta garzetta) feeding on prey that the egrets disturb (F.T. Morris 1978). The diet consists of a wide variety of prey, especially insects. It includes grasshoppers, locusts, crickets, water beetles, worms, molluscs, crustaceans, fish, amphibians and lizards. The birds consume the eggs of waterfowl, Great White Pelicans (Pelecanus onocrotalus) and crocodiles (Crocodylus niloticus) (especially after the shells have been broken by vultures), and the young of Cape Cormorants (Phalacrocorax capensis) (Bolster 1931, Urban 1974a). Carrion and offal figure in the diet near human habitation. R.A. Clark (1979c) reported that South African birds in Pretoria consume mainly human, animal and vegetable refuse, and insects. In summer, the diet is 50% animal food and 34% vegetable food; in winter, 47% vegetable food and 34% animal food (by wet weight consumed). Urban (1974a) found regurgitated food from a 1-month-old chick contained 22 dung beetles, 2 lepidopteran larvae, 2 coleopteran adults, 1 bead, 1 small piece of bone, 1 seed, bits from a millipede, several small ants, and a small piece of egg shell. Birds use communal roosts that are located in trees or on islands. They arrive in the evening singly and in flocks (S.M. Evans et al. 1981). These roost sites shift frequently.

This species nests colonially in many different types of sites. We have seen them nesting in trees and bushes, on the ground and among rocks on islands. The species is surprisingly resourceful in this respect. Sacred Ibises will nest in single-species colonies or in mixed colonies of storks, herons, spoonbills and cormorants, but usually they choose to nest in discrete groups, somewhat separate from other species. Colonies range from a few birds to as many as 2000 pairs at Wembere, Tanzania (Britton 1980) and on the Witwatersrand in South Africa 2180 pairs nested in six colonies in 1974 (Tarboton 1977a). The breeding season varies geographically (E.K. Urban 1974a, pers. comm. 1988). Breeding is generally associated with wet-season inundation, and often occurs at the height of the local rainy season. Birds do even nest at the edge of the desert in the wet season. They can nest in a single site more than once in a year. We have seen up to 100 pairs nesting in a mixed colony on the Tana River, Kenya, in June during the long rains and 10 pairs in the same colony during the short rains in January (Hancock 1984). In some years nesting occurs at unusual times in response to unusual flooding. In Madagascar, bernieri lays eggs in November (Rand 1936). Often nests are placed very close together, but they are seldom touching. Nests are made of large sticks and lined with softer material, mainly vegetation. They usually are about 10cm across and 20cm deep, but sometimes much larger, depending on the availability of nesting material. When this is in short supply there is much stealing of sticks and other material from nearby nests of other species in the colony, as well as from each other. On Lake Turkana, where sticks are scarce and the African Spoonbill nests before the Sacred Ibis, there is considerable fighting between ibises and spoonbills as well as with Little Egrets whose young have hatched before the Sacred Ibis begins to nest. The courtship behaviour of the Sacred Ibis has been recorded by Urban (1974a) at a colony on Lake Shala in Ethiopia. Nesting activity begins at the colony site as birds fly in a Display Flight; these are male birds in small parties. For the first few days they settle for short periods and display by standing in groups with tail and ornamental plumes spread and wings held down. The throat-sac of the African subspecies is expanded. At this stage, the male birds carry out a Sparring display, standing in a horizontal position with head and neck extended and performing an openbill thrust at nearby opponents. As they strike the bills, they make a wheezing squeaking call. The Sparring display is performed by the displaying bird adopting a horizontal stance with its head and neck stretched towards the opponent, and with partially open wings revealing the bright red skin. It then lunges with open bill. Other displays at this time are the modified Sparring, in which 2 or more males stand side by side facing the same

Sacred Ibis 215

way and open their wings and point their heads skyward. The Head Rub display is performed by the bird smoothly moving its neck up and back and then, with bill pointed upwards, flicking its neck backwards so that the crown touches one side of the mantle. A Bill Popping display is performed from a low horizontal position when the neck is extended forward and down but the mandibles are not snapped together. Males chase each other in pursuit flights, and by supplanting attacks force opponents from their perch by flying directly at them. After 1 or 2 days the females arrive and constant chasing and fighting occurs. If the female remains with a male and no chasing occurs, pair-bonding usually commences. The two birds face each other, in a Bowing display, with downward-stretched necks and heads and intertwining of necks and bills. The Head Rub and, infrequently, the Bill Popping displays follow. Pairs then stand, with bodies touching, on the nest territory which, when they have established it, is used for copulation. The male stands over the crouching female and grasps and shakes her bill, moving his partially opened mandibles up and down it. After this, both birds stand and preen, probably a Display Preen. The male collects nesting material and the female remains on site to build the nest. On the male's arrival with material, the female crouches and both birds point their bills upwards and intertwine them. When she has nearly finished building, the female will utter a few sharp calls (Brown etal. 1982). Kim Lowe (pers. comm. 1990), in his study of South African birds, considers that the behaviour of the Sacred Ibis at the nest is considerably less aggressive and relatively more silent than that of the Australian White Ibis. On the other hand, Penny (1974) described a colony on Aldabra of abbotti where fighting often occurs and the birds Tence with their bills and kick each other like game cocks'. Once the nest is completed, eggs are laid. Brown et al. (1982) reported 2-5 eggs are laid, occasionally 5 in Ethiopia and East Africa. The mean (of 535 clutches) in South Africa was 2.43; in southwestern Cape 2.4 (291 clutches); in Transvaal 2.0 (271 clutches). On Aldabra, Penny (1974) reported 2 eggs were laid, rarely 1 or 3. The eggs are oval or slightly rounded, rough and dull white with a faint blue tinge. A few dark red spots are sometimes visible on African eggs, but Penny (1974) stated that, on Aldabra, eggs were all-white. According to Urban (1974a) incubation takes 28-29 days. A replacement clutch after egg loss may be laid but this is not proven (Cramp 1977). Both sexes incubate, changing over at least once every 24 h. Incubation commences after the clutch is complete and hatching is synchronous (Urban 1974a, but see Stronach 1968). The young are fed by regurgitation by both parents, one of which remains at the nest for the first 710 days. Young leave the nest between 14 and 21 days after hatching and assemble in groups nearby. Adults continue to feed them once a day until they fledge at 35-40 days. They leave the colony after 35-48 days. Birds do not acquire full adult plumage for up to 3 years. It is not known when they first commence breeding.

Nesting success is extremely variable. Manry (1978b) reported 85% success from 54 nests on Schaapen Island, South Africa. J. Parsons (1977) reported 43% from 353 nests at Kisumu, Kenya. Only 18 fledged from 420 chicks at Kisumu in 1977. Factors affecting success include low rainfall affecting food supply, heavy rain storms causing young to fall from the nest, and predation by African Fish Eagles (Haliaeetus vodfer) (J. Parsons 1977) and Kelp Gulls (Lams dominicanus) (Manry 1978b). Parsons reported Fish Eagles eating 7 eggs per visit, and Manry reported the eating of eggs, young and fully grown chicks by Kelp Gulls. On Aldabra, disturbance by man caused desertions (C.W. Benson and MJ. Penny 1971) and, in Africa, low-flying aircraft have had the same effect. TAXONOMY We consider Threskiornis aethiopicus to be one of three species forming a superspecies that includes T. melanocephalus and T. moluccus. We thus follow Snow (1978) and K.W. Lowe and G.C. Richards (1991) but not Holyoak (1970). These three species are widely distributed in the Old World, and there are difficulties in deciding the specific limits in this group. Snow (1978) pointed out that two of the forms are themselves dimorphic, indicating that the clearest delimitation of relationships would be by the recognition of three species. K.W. Lowe and G.C. Richards (1991) found no evidence of intergradation among the species in areas of overlap. The Sacred Ibis has two isolated island populations, bernieri on Madagascar and abbotti on Aldabra. K.W. Lowe and G.C. Richards (1991) detected little difference between these two forms. Holyoak (1970) stated that abbotti has little or no black in the wing-tips and has duller display plumes than bernieri. He also described the eye of the abbotti as light blue, and bernieri as nearly white, though pale blue is the more common description of the eye of the latter. In any case, they both differ from the nominate race, which has a brown to brownish red eye. Pending more details and in view of the wide geographical separation of these sedentary forms we prefer to treat them as two races. Interestingly, aethiopicus has once been recorded from Aldabra, where it was distinguished from the local subspecies abbotti by its black wing tips and brownish-red eye (Penny 1974). CONSERVATION Despite its adaptability, this species is dependent on safe nesting sites and on appropriate feeding areas. The African subspecies migrates or disperses widely in search of the latter and, except where pollution is present, finds ample areas in which to feed. Nesting sites are, however, subject to disturbance, drainage and predation by man. The two islands subspecies are also under greater pressure, mainly from predation and disturbance by man. The small population still resident on Madagascar also needs careful protection as eggs and young are reported as being collected by local people (Langrand 1990). Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 317.

Oriental White Ibis Threskiornis melanocephalus (Latham) Tantalus melanocephalus Latham, 1907, Index Ornith., p. 709: India Other names: White Ibis, Oriental Sacred Ibis, Oriental Ibis, Black-headed Ibis, Indian Black-necked Ibis, Oriental Black-necked Ibis (English); Witte Ibis, Zwartkop Ibis (Dutch); Didhar (Hindi); Hei-tou-lu (Chinese); Dahakatti-Kokka, Thalkaththi-Chondam (Sri Lanka); Ibis a Tete Noire (French); Kuro Toki (Japanese)

IDENTIFICATION The Oriental White Ibis is a medium-sized (65-75 cm long), white ibis with a prominent black head and neck, and a long downcurved bill. The base colour of the adult is a pure white with bare orange patches under each wing and along the flank. Like the closely related Sacred Ibis, the Oriental White Ibis has a featherless head and neck that is covered by black skin. Its long bill is also black. Tom Roberts (pers. comm. 1990) reports a few scattered blue spots on the head in winter. The eye is reddish brown. The legs and feet are dull grey to black. The sexes are alike in plumage. Sizes differences between the sexes are yet to be documented. With the onset of the courtship season, a slate grey colour develops on the scapular plumes, and the secondaries become elongated and frayed. These, and straw-like feathers that develop on the neck, are the courtship plumes. The bare patches along the flank and under the wings turn a bright blood red. The bare skin on the head and neck develops a bluish tinge. The feet and legs turn glossy black, flushing red or dark red for a short period. The tail and axillaries are a pale yellowish brown. Nestlings are thinly covered by white down, but the head, neck and back are black. The bill is black and quite short. At 21 days the dorsal and lateral surfaces are covered in blackish brown feathers. The legs at that stage are grey and become distinctively swollen. In the juvenile, only the grey face and around the eye are bare. The rest of the head and neck are white-feathered with brownish or blackish markings. The iris is brown and the bill shorter than in the adult. Juveniles lack ornamental plumes. Full adult plumage takes up to 3 years to acquire. The Oriental White Ibis usually is a silent bird. In the breeding season, however, it gives a remarkably loud booming call (S. Ali and S.D. Ripley 1968). It also makes a series of grunts and mumbles at the nest. It flies with rapid wing-beats in strong, steady flight with occasional glides, often in loose V-formation or in lines. In flight it is faster than egrets or storks. This species is distinguished from other ibises in its range by its white plumage and black head and neck. It differs from other members of its group in several ways (Holyoak 1970, K.W. Lowe and G.C. Richards 1991). Compared to the Sacred Ibis of Africa, its bill is less robust, it lacks a neck-sac, it lacks black on its wing-tips, its display plumes are light grey rather than black, it has straw-like neck plumes, and it has a yellowish tail. It is distinguished from the Australian White Ibis by its more robust bill

218 Oriental White Ibis

and lack of black on wing-tips and display plumes. Its white base-plumage distinguishes it from the Strawnecked Ibis. It is distinguished from the Oriental Crested Ibis by its black neck and head and dark legs and feet. DISTRIBUTION AND POPULATION The Oriental White Ibis was a wide-ranging species found across continental Asia from India and Sri Lanka, north and east to Nepal, Burma, Thailand, and China northeast to Heilongjiang, south to Nam Phan (Cochinchina) in Vietnam and southern Cambodia, through the Malay Peninsula in Malaysia to Java and probably Sumatra in Indonesia. The species' distribution is somewhat patchy within its overall range. Perhaps it is most widely distributed in India, where it breeds throughout the subcontinent. Andrew Robertson (pers. comm. 1990) found it in no fewer than 24 localities from Corbett National Park, across Rajasthan, at Sultanpur Jheel near Delhi, Bharatpur, and in Gujarat in the Gulf of Kutch, the Little Rann of Kutch, and southward through Maharastra and Karnataka down to the tip of India. It is common in the protected national parks and lagoons of Sri Lanka. On the east coast of India it is found at various jheels and tanks and along the coast at Point Calimere. It is also found in Assam, India, and in the Sundarbans, but it is rarer elsewhere in Bangladesh. There are no breeding records for Burma. In Vietnam, Vo Quy (in D.A. Scott 1989) reported 7 large colonies in mangrove and Melaleuca forest. While 1 colony has been abandoned, at least 2 or 3 new ones have become established more recently. Isolated populations exist in China. We have seen them nesting in small numbers in the Zhalong Nature Reserve in the northeast of Heilongjiang. Although its overall range is a broad one (Etchecopar and Hue 1978), we have found no

trace of the Oriental White Ibis in many of the regions we have travelled over and consider it as probably no longer extant over much of the illustrated range in China. T.-H. Cheng (pers. comm. 1990) supports our impression. The species is a resident of the Malay peninsula. Nesting is not yet reported for Sumatra, although it may occur in the Banyuasin Musi River delta (Silvius 1985). In East Java, ibises nest at Ujung Pangkah near the Brantas (MJ. Silvius and W.MJ. Vergheugt pers. comm. 1990). On Pulau Dua and Pulau Rambat we found nesting birds in 1970 (Hancock 1984). Since that time, two colonies only remain on Pulau Rambat, and the ibises do not nest on Pulau Dua (Allport and Wilson 1986). Birds are still found at Muara Bubos and Clanok and from Sutolilo (D.A. Scott 1989). The Oriental White Ibis is somewhat migratory or dispersive over the most northern parts of its range. The northernmost population remains in Heilongjiang until October before migrating south (G. Archibald pers. comm. 1990). Birds that migrate from Manchuria traditionally wintered in southern and eastern China and in Japan. Today it is only a rare visitor to Japan, from Honshu southward. It is found in Hong Kong in small numbers each winter, and also in the Malay Peninsula. There seems to be very limited evidence elsewhere that regular migration takes place. Most birds do leave their breeding grounds and disperse to feed along the coasts and rivers, sometimes a long way away. They wander to any suitable feeding areas, deserting them when drought or disturbance affects foraging. Occasionally birds wander to the Philippines (as well as to Japan as noted above) (Itoh 1986, duPont 1971). Nonbreeding birds wander to the Nepalese terai, and a few still visit the lower Sind valley in Pakistan. The species is occasional in summer in Hong Kong. It is a vagrant in Sulawesi and Kalimantan, Borneo, where birds come into contact with the Australian White Ibis (C.M.N. White and M.D.Bruce 1986). Population estimates are few, but the numbers and range have become much more restricted in recent years. There were recently 25 nests in the northernmost colony site in Heilongjiang, China (G. Archibald pers. comm. 1990). Little is known about colonies elsewhere in China. Probably the largest concentration of this species today occurs in Vietnam at the southern tip of the Mekong River valley. Vo Quo (in D.A. Scott 1989) reports 800-1000 birds in the Bee Lieu coastal marshes and 500-600 breeding at Cal Nuoc at the southern tip of the Mekong. There are 2 or 3 new colonies at Nam Can, and there are 7 large colonies in the mangroves and Melaleuca forests in this area. The largest population in India is undoubtedly at Bharatpur, where D.A. Scott (1989) reported 389 birds in summer and 50 birds in winter at the Keoladeo Ghana Reserve in 1984. However at this reserve in 1983, the annual census showed no fewer than 1435 birds (Hancock 1984). Up to 100 were counted at the Ranganthiloo Wildlife Sanctuary, south of Mysore, where we saw them in 1988. Elsewhere colonies of up to 50 or 60 birds are found in many areas of India and Sri Lanka, though at Kaliveti Tank in Tamal Nadu, India, up to 230 birds were counted, and at Kullar

Oriental White Ibis 219

Sandai and Vembakottai in the same state 160 were reported (D.A. Scott 1989). This contrasts sharply with Pakistan, where the largest recorded concentration was 100 birds in Sind in 1965, now reduced to only a few birds in the 1980s (Tom Roberts pers. comm. 1990). The species is scarce in Burma, though a few winter near Rangoon. In the southern central plains of Thailand, 90 birds roosted at Wat Tarn En and 70 birds were reported on passage at Ban Lung Jorm (D.A. Scott 1989). It is rare in Formosa and Japan. Similarly, it is now very rarely seen in Malaysia (P. Round pers. comm. 1990). In Indonesia only very small colonies have been reported. Although breeding colonies have not been found, large flocks occur in southern Sumatra, where the population may be 2000 individuals (MJ. Silvius and WJ.M. Verheugt pers. comm. 1990). Silvius et al. (1985) found 700 birds feeding in Sumatra Selaan. The species occurred widely in Java in the early part of the century but only recently were breeding sites rediscovered, at Ujung Pangkah near Brantas, East Java (MJ. Silvius and WJ.M. Verheugt pers. comm. 1990). Also on Java, we reported 20-30 breeding pairs on Pulau Dua in 1970 (Hancock 1984). Elsewhere along the east coast, only small numbers were found. ECOLOGY This wide-ranging, adaptable species is both an aquatic and terrestrial feeder, both inland and along coastlines. It will feed quite often in dry areas and will even scavenge in open drains as well as in city streets where it is not molested. It is primarily a bird of the lowlands. In Sri Lanka, it is found in coastal lagoons, in swamps and, in the dry zone, in irrigation reservoirs. Usually the Oriental White Ibis feeds in flocks or small groups, but sometimes it feeds singly. It has simple foraging methods, Walking Slowly, Probing or Pecking at the surface. It will turn over vegetation and adopt tactile feeding methods in shallow water, often in the company or other species. The ibises use their long, curved bills to Deep Probe on coastal mud flats. They shift territories as the tide rises. Fish, frogs, molluscs, insects, worms and small reptiles make up the bulk of the diet, but some vegetable matter is also consumed. Stomachs of small downy nestlings contained pulpy olive-brown and green vegetable matter and elytra of tiny blackish beetles about 5 mm long (S. AH and S.D. Ripley 1968). BREEDING Nesting occurs in well-protected areas, often in traditional colonies mixed with herons, storks and cormorants, but sometimes separately. Nests are typically built in mediumsized trees such as Acacia, Prosopis, Barringtonia and others, that are standing in or near water. Nests are also found on the top of partially submerged shrubs such as Zizyphus and Capparis horrida (S. Ali and S.D. Ripley 1968). Where they were left unmolested, we have seen nests in tall trees on the edge of human habitation. Amongst other species, ibises nest in small conspecific

groups, though the ibises' nests are usually above those of egrets and cormorants. On islands in the Gulf of Kutch, they sometimes nest on or very close to the ground often with Indian Darters (Anhinga rufa melanogaster]. In Indonesia, they nest alongside Milky Storks. Certainly here, as at nearly all Indian Sites where we have seen Oriental White Ibises nesting, they have nested in trees already occupied by other species. The availability of nest sites in mixed colonies is to some extent dictated by other species within the chosen site. As an example, desertion of a traditional site by Asian Openbill Storks at Bharatpur in 1989, due to artifically raised water levels, left the trees clear for other species. Nesting takes place after the rains and varies according to the monsoons. The species nests in December to April in Sri Lanka. Eggs have been found in April-August in West Java. The nesting biology of this species has been studied by Hoogerwerf (1935), but insufficient information is available to determine the differences in the courtship and nesting behaviour of any of the species of Threskiornis. One thing that is fairly certain, however, is that the Oriental and Australian White Ibises do not have the throat-sac of the Sacred Ibis. Oriental White Ibises commence breeding activity with Display Flights, landing for short periods while they display by standing in groups with tail spread, wings held down and ornamental plumes spread. The blood-red stripe under the wing is prominent in Display Flights. Males carry out Sparring displays standing in a horizontal position with head and neck extended and feinting with an open bill thrust at opponents. No bill snapping occurs. Circular pursuit flights are common at this stage. We have seen, in this species as in the Sacred Ibis, a modified Sparring display in which two birds stand side by side facing in the same direction and point their bills upwards and, nearly touching each other, call with a harsh grunt. Male birds advertise for females by performing Bill Popping and Head Rubbing displays, and these are carried out also by females as they approach the proposed nest sites. Bill Popping- is the less common of the two. The bird keeps low, crouching to extend its neck forward and down in a sudden quick movement. In the Head Rub, the neck is raised up and back with the bill pointing skyward and then the head touches the mantle on one side. When a female approaches a displaying male, either she is chased away or performs a Bowing display. Both birds Bow with neck and head extended, necks may be inter-twined, bills are crossed. Birds will then Display Preen, but not allopreen. We have found the Oriental White Ibis to be less aggressive than the Sacred Ibis and to have less elaborate displays. In fact, we have seen considerable variability in the intensity and duration of courtship displays. There is less fighting and less intense displays in small colonies, where only a few pairs are present, than in larger ones. Our observations of courtship in a year following highly successful breeding suggested that breeding activities were particularly strong then, whereas in a year after a nesting failure, displays were not so extended (pers. obs. J.A.H.). Once nesting pairs are established, the remainder of the competing group will depart and re-form at another site,

220 Oriental White Ibis

although they may return to attempt to dislodge the first group of paired birds. Copulation takes place at the nest-site after the territory has been successfully established. The male straddles the crouching female and grasps her bill, shakes it, and then moves his partially opened mandibles up and down her bill. Preening follows. Copulation is carried out usually before any nest-building is started. K. Lowe (pers. comm. 1990) noted frequent promiscuity between unpaired birds as well as between males already paired, which copulate with other females visiting the nest-site. Stick gathering is carried out by the male and the nest built mainly by the female. S. Ali and S.D. Ripley (1968) stated that both birds actually take part in nest building. However, competition for sticks is usually such that one bird remains at the nest, although it does steal sticks from neighbouring nests whilst remaining on its own platform. Nests are often close together, in some cases with only a few centimetres separating them. Nests built close together inevitably lead to aggressive action around them. Once the nest is completed, one bird stays there. Nests consist of a small platform of sticks c. 25-30 cm across. 2-4 eggs are laid. They are long, oval-shaped and chalky white with a faint bluish tinge, and sparingly marked with small spots and blotches of light or dark brown, more densely at the broad end. The average size of 150 eggs measured was 63.5 X 43.1 mm (E.C.S. Baker 1929). Incubation takes about 23-25 days. Young are fed by regurgitation and both parents feed the young. The chick will grasp the adult's bill to open it and then thrust its bill into the adult's opened gullet. During the heat of the day chicks are not usually fed, and often both parents will stand on the nest to shield the chicks from the hot sun. At that time the adult's bill is half open and the throat of the bird pulsates continually. There is constant raiding of the nest by House and Jungle Crows. Young ibises are attacked by a variety of birds of prey in India. Fledglings leave the nest to join other young from surrounding nests and sit on branches close to one

another. At this time they will still food Beg when adults pass over head. Seldom do more than 2 chicks survive per nest. TAXONOMY The Oriental White Ibis has variously been considered a separate species or a race of the Sacred Ibis (Holyoak 1970, J. Steinbacher 1979, Van Tets 1978). However, K.W. Lowe and G.C. Richards (1991) showed that morphological variations in museum specimens were sufficient to separate T. melanocephalus from both T. aethiopicus and T. moluccus. They found no intergradation of characters between moluccus and melanocephalus, despite the close contact of the forms in the Sulawesi/Moluccan island area. C.M.N. White and M.D. Bruce (1986) consider their distribution to be allopatric. K.W. Lowe and G.C. Richards were not able to document detailed courtship displays, although suggesting that the behaviour of the forms is strikingly different. We consider that these forms have been isolated for a long period and that it is best to treat them as separate species (K.W. Lowe and G.C. Richard 1991, D. Holyoak pers. comm. 1990). Certainly there is room for considerably more study before the taxonomy is fully resolved. CONSERVATION Threats to this species include loss of habitat due to drainage, disturbance by human predation and pesticides. The establishment of protected breeding areas in reserves, particularly in India and Vietnam, has helped the species to maintain its numbers in these areas, but in most other regions it must be regarded as a threatened species.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 318.

Australian

White Ibis Threskiornis molucca (Cuvier) Ibis molucca Guvier, 1829, Regne Animal, nouv. ed., 1, p. 520: Moluccas Subspecies: Threskiornis molucca molucca (Guvier) Threskiornis molucca pygmaeus Mayr Threskiornis aethiopicus pygmaeus Mayr, 1931, Amer. Mus. Novit., no. 486, p. 6: Rennell Island, Soloman Is. Other names: Australian Ibis, White Ibis, Black-necked Ibis, Sticklebill (English)

IDENTIFICATION The Australian White Ibis is a medium sized, white ibis with a black bill, bare black neck and obvious black plumes covering the lower back. This Australian bird looks very much like its relative, the Sacred Ibis of Africa. The adult's white base colour is set off by the dark, green-black wing-tips and black, lace-like scapular plumes that extend over the lower back and over the yellow tail. The bare face is black to blueblack; the bill is black. The iris is brown. A pink band traverses the back of the head. The neck is covered by bare, black skin and lacks a neck-sac. The upper part of legs are pink, lower portions dark grey to black. Patches of bare reddish skin occur at the sides and under the wings. Elongated feathers form a tuft at the base of the foreneck. The sexes have similar plumages, but males are larger than females. Some comparative measurements for molucca are, for the male, culmen 16.8-19.0, wing 36.8-38.1, tarsus 9.1-10.0, tail 12.1-12.5 cm; and for the female, culmen 13.9-15.1, wing 34.8-35.5, tarsus 8.3-8.5, tail 11.5-11.8 cm. The size of the molucca varies along a cline, increasing from New Guinea through southern Australia (K.W. Lowe and G.C. Richards 1991). The subspecies are distinguished primarily by size, the island subspecies (pygmaeus) being considered a dwarf form. In comparison to the data presented above on the Australian form, the body-part sizes of a pygmaeus specimen are culmen 12.3, wing 30.4, tarsus 7.4 and tail 11.0 cm. During courtship, the bare patch under the wing and flank turn bright red, and the other bare parts become glossy black. Ornamental plumes reach their yearly maximum extent, and the tuft of feathers at the base of the foreneck reaches maximum elongation. Nestlings are at first covered in white down except for the head and neck, which have black down. The white feathers are replaced after 3 weeks with brownish black feathers. At this time the legs are grey and swollen. The juvenile is whitish but has black feathers on the head and lacks neck and wing ornamental plumes. The bill is shorter and straighter than is the adult's. Like the other Threskiornis species, this is generally a silent bird, except during courtship and at the

Australian White Ibis 223

nest. The voice, as described by Pizzey (1980), consists of harsh barks and shouts. The normal call is an 'Urk', whereas calls associated with breeding are 'Koaha*, 'Tawtaw' and 'Taw-aw' (Frith 1976). Young birds utter a squeaking, hissing sound. The Australian White Ibis is often found in company with the similar Strawnecked Ibis but is easily distinguished by its white body. Its small neck-tufts contrast with the full, straw-coloured neck plumes of the Strawnecked Ibis. It is distinguished from the remaining Threskiornis species by a combination of characters including its dark wing-tips, dark ornamental plumes, a neck-tuft, no neck-sac, yellow tail and coloured tubercules on the head. In comparison with the Sacred Ibis of Africa, it has a more delicate bill, lacks a neck-sac and has a neck-tuft. Unlike the Oriental White Ibis, it has black wing tips and display plumes. DISTRIBUTION AND POPULATION This species is widely distributed on the Australian mainland north to southern New Guinea, the Moluccas and the Solomon Islands. In Australia, this ibis is particularly found in the east and north. In the southeast it is a common breeder along the coast and inland but not further than 3CFS. There are some colonies in the northeast, and we have seen colonies on the South Alligator river east of Darwin. Only since 1952 have birds moved into the southwest region, and breeding there was first reported in 1979. In New Guinea, the bird is resident in Irian Java and Papua New Guinea. It also inhabits islands of the Moluccas (Seram, Geser, and great Kai). The subspecies pygmaeus is found on Rennell and Bellona in the Solomon Islands. Both migratory and nomadic, this species responds to seasonal and localized dry periods by undertaking population shifts (Carrick 1962). Bekle (1982) found it to be somewhat unpredictable in its wanderings in the Perth area. Banded birds from Victoria and New South Wales moved northward along the east coast and to the Cape York Peninsula. Birds cross the Torres Strait to winter in Papua New Guinea (Purchase 1976). Draffan et al. (1983) reported migratory flocks of 20-500 birds crossing into southern New

Guinea. Thus the resident population in New Guinea is joined by migrants from Australia in winter. The species is accidental in New Zealand and Tasmania (W.R.B. Oliver 1955, R.H. Green 1959a). In Tasmania there were major eruptions in 1957-58 and again in 1975, but breeding has not been recorded. It is also vagrant in the Lesser Sundas (Tinimbar and Babar), Moluccas (Kai, Geser), Sulawesi, and Kilimantan on the Borneo coast, where it may or may not coincide with the Oriental White Ibis (K.W. Lowe and G.C. Richards 1991, MJ. Silvius and W.J.M. Verheugt pers. comm. 1990). Population sizes are in places fairly well known, at least in Australia. The major colonies are found in Victoria and New South Wales. In 1979-80, numbers in 16 colonies ranged from 150 to as many as 15000 per colony, with a total of 33000 birds (Cowling and Lowe 1981), and on Barren Box Swamp in 1967 there were up to 1000 nests. However, colonies in the north are smaller. There were an estimated 184 nests at Kapalga and up to 1000 nests on the Fitzroy and the Ord Rivers. We visited a colony site on the South Alligator, which was also occupied by a mixed breeding colony of egrets and cormorants. There we saw up to 50 nests of ibises, whilst, across the river from the main colony, 2 more colonies of up to 10 nests were found with no other species present. The Australian White Ibis is not uncommon at Seram and has been considered to be scarce or uncommon in western Irian Jaya. However, a recent survey in New Guinea found large mixed colonies of nesting birds in southeastern Irian Jaya, in the Pulau Kimaam and Wasur reserves (Bishop 1984). Over 15000 Australian White Ibis were recorded at Pulau Kimaam and over 30 000 at Wasur. This exciting discovery alters the picture completely as to the status of this species, as well as the status of other storks and ibises in New Guinea. ECOLOGY The Australian White Ibis much prefers aquatic or wet situations and tends to avoid arid areas (there are few records from the centre and central south of Australia except during exceptional rains). It feeds along the shallow edges of lakes and inland lagoons, on mudflats, in swamps, irrigated pasture land, mangroves, creeks, marshes, and at dams, reservoirs and rivers. In times of drought, birds feed along the coast. In Irian Jaya, birds occupy the rich grasslands that lie within the Wasur reserve and extend eastward to join the extensive Bulla Plains of the Tonda Wildlife Management area (Bensbach, Papua New Guinea) (Bishop 1984). In northern Australia we watched birds feeding along the mudbanks of the heavily tidal rivers on the vast expanse of exposed mud, and on the grassy flood plains. Human habitats are not avoided. Birds commonly visit garden lawns, playing fields and sewage dumps. They scavenge around garbage tips and poultry yards. Like others of the genus they will also feed on burnt savanna. Australian White Ibises feed in conspecific flocks or in the company of other wading birds. We have seen them with Royal Spoonbills, Pied Herons (Egretta picata) and Black-

224 Australian White Ibis

necked Storks. Whitefaced Herons (Ardea novaehollandiae) follow Australian White Ibises, capturing prey disturbed by their probing (W.E. Davis pers. comm. 1987). The Australian White Ibis feeds by Probing, although it also has been reported to Head Swing. A bird will wade steadily through the water Probing near its feet (Vestjens 1973a). In doing so, it will often penetrate 25cm or more into the water, deeply submerging its head and neck. This behaviour contrasts with that of the Strawnecked Ibis, which prefers shallower water (Carrick 1959). Australian White Ibises also use 'anvils' for opening mussels. On finding a mussel, the bird takes it out of the water, holds it with the feet on thick roots of living trees, pebbles or a flat rock and hammers it with the bill until the shell breaks (Vestjens 1973a). The diet is varied, consisting of earthworms, freshwater crayfish, spiders (adults and egg-sacs) crickets, mole crickets, grasshoppers, shieldbugs, moths and caterpillars, bees, ground beetles, water beetles (adults), dung beetles, click beetles (larvae), weevils, freshwater mussels, water snails, fish, frogs, and lizards (Carrick 1959). In eastern Australia, Carrick found that the White Ibis fed largely on freshwater crayfish and other aquatic foods, and made limited use of drier habitats. He also found that this ibis favoured larger prey than the Strawnecked Ibis, with most insects being consistently about 5 cm long; a large proportion of the diet involved Probing for burrowing types of prey such as crayfish, crabs and earthworms. Cowling (1974) found that captive birds preferred to eat live locusts rather than dead ones, but preferred sluggish ones available in areas sprayed with insecticides. Among grasshoppers and locusts (Acrididae), Carrick (1959) found that the birds preferred the spur-throated locust (Austracris guttulosa), which is abundant around the margins of marshes after feeding, and that in many cases dry-land species were consumed by Strawnecked Ibises but not by the Australian White Ibis. BREEDING K.W. Lowe (1984) stated that, in Australia the White Ibis exhibits two broad responses to its environment. Some populations accommodate their breeding to an unpredictable environment and other populations use complex breeding tactics dependent on a highly predictable set of environmental conditions. Overall, the species breeds during the wet season in locations where recent rains have produced the flooding that sustains breeding. Nesting is in trees, bushes or reeds. In Victoria, birds nest in lignum (Muehlenbeckia cunninghamii), reeds (Phragmites spp), rushes (Juncaceae) or eucalyptus trees (Eucalyptus). In the north and in the south of the State, nest sites are usually in reeds, rushes or paperbarks (Melaleuca ericifolia). Rarely do Australian White Ibises nest on the ground, though they do so occasionally on islands on the perimeter of wetlands. They have been recorded nesting in trees on dry land well away from wetlands (Cowling and Lowe 1981). This species often nests in close proximity to the Strawnecked Ibis in Australia. In New Guinea, it nests alongside

herons, egrets, cormorants and storks. Here we have seen that it has a tendency to nest in groups of its own species, and sometimes in separate colonies at some distance from the mixed colonies. Courtship behaviour has not been well documented, but some information is provided by Frith (1976) and K.W. Lowe (1984). Males and females gather in 'courting groups'. Courting groups disperse and reform elsewhere. The nest is started when groups have dispersed, leaving a pair that has mated to establish it. Within a group, synchrony of breeding is established, but sometimes other groups will take over occupied nest-sites, evicting the pairs and destroying the nests, which may contain eggs or young. Males display by vocalizing, erecting feathers, spreading their wings and snapping their bills. Raising their wings displays the scarlet skin underneath and alongside the breast. The male chooses a potential nest-site and tramples a small area. The male uses Bowing displays, and when a female approaches, the male grasps a twig in its bill. Both birds will grab the stick together, after which they Display Preen. Fighting is common. When one male was seen to interrupt the display of another, they fought by Sparring with their bills. Fights occurred between paired and unpaired males over the possession of a nest-site, or when an unpaired male or female attempted to intervene between a female and her male 'escort'. Females that fight for access to a male are usually individuals that have bred with the same male previously. These fights occur during courtship bouts and on nest-sites after a pair had become established. Fights between females have resulted in eggs being destroyed by the interloper. The male and female share the task of nest building. A nest consists of a substantial, compact, shallow cup of reeds or sticks lined with vegetation. In dense colonies, nests become trampled into a continuous platform (Pizzey 1980). The Greeting displays at the nest consist of deep bowing. Promiscuity is common both before and after pair-bonding, and some birds make several attempts to breed during the season. Between 1 and 4 eggs are laid, most pairs having 3 eggs, which are dull white and sometimes become heavily stained. Eggs are laid at night on alternate days. The mean size of eggs is 67.8 X 43.8mm (n = 30) (K.W. Lowe 1984). Both sexes incubate, incubation being 20-23 days. Hatching is asynchronous, over a 2-day period. Feeding of nestings is by regurgitation. Young remain in the nest for approximately 3 weeks, and then stay together in groups for another 2 weeks, perching together on trees while still being fed by the parents. The mortality rate of chicks depends largely on availability of food. However, colonies are often subject to flooding and are deserted. Conversely, in areas where sites dry out, nests are also deserted through lack of food availability or because of land predators. Predation does not seem to be as important a factor as it is for the Sacred Ibis in Africa. We have, however, seen birds of prey circling nests in the South Alligator colony site, and eggs and young are certainly taken by Whistling Kites (Haliastur sphenurus), which

Australian White Ibis 225

also harry adult birds taking food to the nest (Pizzey 1980). Lowe (1984) found the number of young fledged to be between 0.50 and 0.60/clutch. TAXONOMY The forms of the Australian White Ibis have in recent years usually been considered to be representatives of the Sacred Ibis (J. Steinbacher 1979). However, more recent considerations have concluded that the Sacred group is best classified as three species, which we follow (Snow 1978, Mees 1982, K.W. Lowe and G.G. Richards 1991). The aethiopicus and molucca species resemble each other superficially in plumage colour. However, it is likely that molucca and melanocephalus are more closely related, sharing such features as the tuft of feathers on the breast and juvenile plumage (Snow 1978). K.W. Lowe and G.C. Richards (1991) show that there is no blurring of characteristics between molucca and melanocephalus, despite the closeness of their ranges. Three subspecies have been recognized in recent years within the molucca group, molucca, pygmaeus and strictipennis. The subspecies molucca represented the birds in the Moluccas, and strictipennis the birds in Australia. The birds in New Guinea were considered to be intermediate (J. Steinbacher 1979). Holyoak (1970) had distinguished between the two based on the colour of the shafts of the secondary wing-feathers, but Lowe and Richards show this character to be variable among birds and probably among

ages. Mees (1982) and K.W. Lowe and G.C. Richards (1991) concluded that molucca and strictipennis are identical. However there is a geographical variation in size, decreasing from north to south. The name molucca has priority for the species name. The name moluccus is often used, but Sibley and Monroe (1990) show that the original name, molucca, is correct. The subspecies pygmaeus is similar to molucca but is a dwarf form, being considerably smaller than the nominate race. Thus we consider that the Australian White Ibis is a polytypic species.

CONSERVATION This species flourishes in suitable habitat throughout its range and is to our knowledge, in no difficulty. Recent discoveries in New Guinea are especially encouraging. Reserves in Australia, at least, are well protected, and the species is not subjected yet to the pressures of human populations, either there or in Irian Jaya. Furthermore, it appears to accommodate well to human habitations.

Note: Body measurements, egg measurements and nesting ne season data for this species can be found in the Appendlix on page 318.

Strawnecked Ibis Threskiornis spinicollis (Jameson) Ibis spinicollis Jameson, 1835, Edinburgh New Philos. Journ. 19, p. 213: Murray River, New South Wales Other names: Dryweather Bird, Farmer's Friend, Letterbird, Black Ibis, Black-and-white Ibis (English)

IDENTIFICATION A medium-sized ibis, 65-70 cm long, the Strawnecked Ibis is a predominantly black and white bird, with straw-like display feathers at the base of its neck. The adult is in many ways similar to the more widespread Australian White Ibis, with which it is often found. The back and wings are black with a glossy green sheen. Part of the neck, the undersides and tail are white, which provides a conspicuous contrast when the bird is in flight. Just below the head, at its side and base, the neck turns predominantly white, merging with the stiff straw-coloured plumes protruding from the lower neck. A stripe of white feathers crosses the undersides of the wings. The unfeathered head, back of the neck, and throat are black, as is the long, curved bill. The feet and lower legs are black, but the upper legs are paler, flushed pink to red. A patch of bare black skin occurs on the flanks. Males are larger than females. McKilligan (1975) found the length of the culmen (straight line from tip to base) averaged 165mm in males contrasted with 137mm in females. This difference was statistically significant and could be used to determine the sex of birds at close range in the field. During courtship and breeding, the legs turn pink to dark red, the green sheen brightens and the straw-coloured plumes are at their maximum length. Immature birds have feathered heads, and overall are duller coloured than adults and lack the sheen on their black feathering. The chin and the back of the neck are black, the legs and feet dusky. The head feathering is lost and neck plumes develop in the third or fourth year (Le Souef 1917). The Strawnecked Ibis is generally a silent bird. Its voice has been described as consisting of grunts, hoarse coughs, and drawn-out croaks (Pizzey 1980, Simpson and Dat 1984). In flight this bird is a study of black and white contrast. Flocks will wheel and turn in loose formation often descending from great heights, circling widely toward the ground in a series of gliding movements. The Strawnecked Ibis is distinguished from its two sympatric ibises, the all-dark Glossy Ibis and the Australian White Ibis, by its colour pattern. It is less confined to wetlands than are these species.

DISTRIBUTION AND POPULATION The Strawnecked Ibis has long been considered to be an Australian endemic. It nests at least occasionally in all states and territories of the mainland, although it is usually absent from the arid interior (McKilligan 1975). The population appears to be centred in Victoria, with fewer nesting elsewhere in the continent. The nesting populations in western Australia and in southeastern Australia are separated by deserts. Recently the Strawnecked Ibis has been found breeding in southeast Irian Jaya, which considerably extends the breeding range of the species (Bishop 1984). Prior to this discovery, the species was

228 Strawnecked Ibis

Creek, New South Wales (McKilligan 1975, Cowling and Lowe 1981). In earlier years, even larger colonies have been reported (Waterman et al. 1971). The largest colonies in the southwest are a few hundred birds. The newly discovered nesting population in Irian Jaya included about 350 birds. The Strawnecked Ibis remains abundant throughout its range in Australia, with no indication of population declines. In Victoria, Cowling and Lowe (1981) did not detect a change in numbers of nesting ibises over 25 years. However the breeding range appears to have expanded towards the south-central and southeast of that State. The species was rare in southwestern Australia, until 1982.

known to be found in New Guinea nearly year-round, but in varying numbers. It was reported over 100 years ago by D'AIberis at Attack Island, Fly Delta, where in December large flocks were seen flying at a great elevation in a northwest direction. It was found throughout the year in the Bensbach River area of the southern trans-Fly region. Its abundance there varied with rainfall, peak numbers being reached at the beginning of the rainy season, when several thousand were present, the numbers falling to hundreds at the height of the dry season. It was also seen fairly regularly by Hoogerwerf at Kurik in the nearby south of Irian Java, but seldom in the rainy season. Clearly, much of the movement of birds into and around New Guinea is migratory. It has in fact long been known to be a migrant to Irian Jaya, and flocks of 50-100 birds are seen in the dry season over the Torres Strait (MJ. Silvius and W.J.M. Verheugt pers. comm. 1989). Flocks have been found drowned at sea (Pescott 1968). Such a migratory dispersal appears to be the rule, especially northward from the densely packed breeding areas in southeastern and western Australia.^ Birds nesting in New South Wales migrate to the east coast and the northeastern Northern Territories, whereas birds nesting along the west coast migrate north to Darwin and Arnhem land. We have seen flocks arriving in April, apparently from great distances and heralding the onset of the hot, dry season there. The arrival of the Dryweather Bird, landing among the Magpie Geese, Anseranus semipalmata, with their young is the harbinger of the season to local residents. Immature birds disperse widely and rapidly after the nesting season. These birds wander widely, especially in response to changing water conditions, occurring at times nearly anywhere in Australia. The species is a vagrant in Tasmania and in Indonesia, beyond New Guinea. Of the three species of ibis found in Australia, this is the most numerous. In 1979-80, 76500 adult Strawnecked Ibises were estimated to have bred in the 26 largest colonies in Victoria (Cowling and Lowe 1981). Other documented nesting concentrations are 150000 birds in most years at Bool Lagoon in South Australia, 200 000 at Reedy Lake, near Kerang, Victoria, 100000 on the Lachlan River, near Booligal, New South Wales, and 15000 on the Yanko

ECOLOGY The Strawnecked Ibis is more likely to be found in dry areas than the other ibises in Australia, often moving into dryland territory. It seems particularly attracted to insect plagues in dry areas, although there is no evidence that it can control outbreaks. It also follows grassland fires (M.J. Silvius and W.J.M. Verheugt pers. comm. 1989). The unpredictable climate of Australia means that the Strawnecked Ibis is often compelled to assemble in wandering nomadic flocks to use these sporadic food sources. Among its typical foraging habitats are grasslands, including city parks and road edges, swamps, and cultivated land after the crop has been harvested, but birds have not been seen in long, dense grass or in woodland with thick undergrowth (McKilligan 1975). They also avoid marine habitats. Use of dry habitats becomes especially prevalent during the non-breeding dry season, in the North. During this period the ibis chooses sites of moderate to average moisture content rather than deep water or croplands. Unlike the Australian White Ibis, the Strawnecked Ibis does not usually visit garbage dumps. Feeding in wetland areas is by deep or shallow Probing along the edges of marshes and in wet grasslands. Here the birds are more likely to keep some distance from one another, but when they move to terrestrial feeding areas they move in closer formations. When grasshopper and other swarms of big insects are found, the ibises feed by sight in large close parties moving rapidly forward. They feed more by visual techniques than do Australian White Ibises. They also have been reported to feed in trees, by probing into cracks in the bark (Haffenden 1981). The food of this species has been well studied by Carrick (1959), Cowling (1974), and McKilligan (1975). Their diet predominantly includes aquatic and terrestrial invertebrates but is rather varied. Among the more common prey are caterpillars, crickets, locusts, beetles and their larvae. Compared to the Australian White Ibis, they take more small insects and spiders and fewer crayfish. In the dry season, beetles, spiders and small lizards particularly are taken. As noted above, the Strawnecked Ibis eats plague locusts; Cowling (1974) found it prefers live to dead locusts, but is nevertheless attracted to those rendered sluggish by pesticides. While nesting in suitable freshwater wetlands in the rainy

Strawnecked Ibis 229

season, Strawnecked Ibises forage on flooded areas where food is temporarily available. There they feed on dormant aquatic prey and terrestrial animals drowned by rising water levels (Carrick 1962). Continued flooding increases plant growth, which supplements the supply of herbivorous insects. In most seasons, a bird will spend much of its time foraging, occupying more than 75% of its time budget (McKilligan 1975). Feeding activity tends to cease in midday. Travel time is minimized by choosing roost sites, often in woods or tall eucalyptus trees, near feeding and drinking sites. BREEDING During its nesting season, the Strawnecked Ibis moves to areas that flood seasonally and there takes advantage of temporally abundant food supplies (Carrick 1962, Waterman et al. 1971, Gosper et al. 1983). Because of the dependence of breeding on the rainy season, it nests most reliably in areas where water supplies are dependably seasonal. McKilligan (1975) stated that breeding in southwestern Australia is well defined from August to December. In South Australia and Victoria much the same pattern emerges. Occasional breeding takes place in the Northern Territories from January to March, though this is on a very small scale. In New South Wales and southwestern Queensland, the breeding season is not well defined (Carrick 1962). We did not find evidence of breeding in colonies containing Australian White Ibis on the Adelaide and Alligator Rivers, N.T., even though rains are reliable in this area. Nesting is an annual event in southeastern and southwestern Australia (Victoria, New South Wales, Western Australia) but may not occur each year in the Northern Territory or in Queensland, where nesting occurs in isolated breeding colonies. Although usually absent from the arid interior (McKilligan 1975), the species has been known to nest there after heavy rains (Waterman et al. 1971). Strawnecked Ibises begin to nest several months after the beginning of the wet season (Carrick 1962, McKilligan 1975), unlike the Australian White Ibis which initiates nesting soon after the rainy season begins. In some cases, nesting is begun too late, the flood subsides and nesting is abandoned. Regional differences in the timing of nesting over Australia are explainable by variability of the timing of the rainy season, and the situation is similar to that described for Australian waterfowl (Braithwaite and Frith 1969). In permanently flooded swamps, temperature may also be a factor in determining nesting season (McKilligan 1975). Nests are built in low bushes, or on well-protected reed beds on islands. Sometimes, however, the Strawnecked Ibis will nest on dry ground where there are no trees, bushes or reeds (Carrick 1962). Birds have been reported as nesting in flood debris and large lignum clumps (Waterman et al. 1971). They will roost in, but apparently not nest in, trees. Breeding is in large colonies, although sometimes only a few pairs are to be found in mixed-species colonies of

Australian White Ibis and herons. Sometimes they will take over the breeding sites of the earlier-nesting Australian White Ibis after the latter has raised its young. Nothing has been recorded of the display of this species, nor of its behaviour at the nest. The nest is built of sticks, reeds and rushes. It is a shallow cup, usually sited in a bush over water. When nests are in reeds, the surrounding vegetation is trampled down, and the areas becomes a single platform of vegetation. Clutches vary from 1 to 4. Lowe found that in one year the majority of the nests contained 2 eggs, with a mean of 2.29 eggs per nest, whereas the following year the mean clutch was 2.42. Eggs are dull white but soon become stained brown (Pizzey 1980). Incubation takes 3i weeks, and fledging occurs 4 weeks after hatching (Carrick 1962). Both parents feed and brood the young (Carrick 1962). Feeding is by regurgitation and continues up to 2 weeks after the young have left the nest. Predation is not great, but food availability determines success of the number of young raised, and when its supply ceases young are readily abandoned. Lowe found that in one year 112 clutches produced 52 fledged young. TAXONOMY Although long considered to represent a distinct genus (Carphibis), the Strawnecked Ibis appears to be fairly closely related to the Sacred Ibis group (Holyoak 1970), one of which—the Australian White Ibis—is also found in Australia. It differs primarily in its dark plumage, ornamental neck plumes, and lack of ornamental display plumes derived from the secondary feathers. It also occupies a slightly different ecological niche, being better adapted to feed in dry conditions and nesting on a schedule slightly delayed behind that of the more aquatic Australian White Ibis. A subspecies fozroi was described by Matthews based on its being a smaller size than other populations. However, the principal correlate of size differences in this species is sexual, not geographic (Amadon and Woolfenden 1952). CONSERVATION Historically, the Strawnecked Ibis played an important role in the lives of aborigines, who used it as part of their natural diet. European occupation and habitat changes have benefited the Strawnecked Ibis through the clearing of forest and woodland, and the drainage of swamps for pasture and farm land (McKilligan 1975). The bird is welcomed in dry farming areas where it is recognized as a destroyer of insect hordes. There seems to be no threat to the Strawnecked Ibis in its dry-season refuge areas. However, the Strawnecked Ibis does depend on freshwater swamps for nesting. Further wetland drainage and flood prevention will restrict the available nesting areas. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 319.

Waldrapp Ibis Geronticus eremita (Linnaeus) Upupa Eremita Linnaeus, 1758, Syst. Nat., ed. 10, p. 118: Switzerland Other names: Hermit Ibis, Bald Ibis, Bare-faced Ibis, Red-cheeked Ibis, Northern Bald Ibis, Crested Ibis, Redfaced Ibis (English); Heremietibis (Dutch); Ibis chauve (French); Ibis calvo (Spanish); Waldrapp or Waldrappe is used in German and Swiss as well as English

IDENTIFICATION The Waldrapp is a dark, relatively large ibis, 70-80 cm long, with a bare red head set off by an untidy ruff of feathers. Bannerman and Bannerman (1968) describing the striking plumage called the Waldrapp a grotesque, yet handsome bird. The base plumage colour of the adult is a dark metallic green, which is tinged with a purple iridescence. Its upper wings are glossed with copper and violet, forming a wing-patch that seems more maroon in specimens we have seen. Elongated purple-glossed black feathers form a loose ruff or collar emanating from the back and sides of the neck. The neck tuft, which is used in displays, gives this bird its unkempt, dishevelled look. The head is featherless, and the face and throat are red, except for the black crown topped by a central streak of orange-red, which varies with age. The iris is grey, and the curved bill, along with the legs and feet, are a dull red. The sexes have similar plumage, but the male is slightly larger, a difference that is discernible among groups of wild birds. Sample measurements are: wing, 40.3-42.0 cm (male) and 39.0-40.8 cm (female); bill, 13.3-14.6 cm (male) and 11.5-13.1 cm (female). Siegfried (1972) found less sexual difference in bill length in Moroccan birds than in eastern birds. In both sexes, the red of the soft parts brightens during courtship. Nestlings are covered with a grey-brown down, darker above than below and have a pink and black bill. The juvenile is similar to the adult, but duller with very little iridescence and no copper coloration on the wing. Unlike the adult, the head of a young bird is feathered (grey with white edges). The ruff feathers are also underdeveloped. This permits recognition of young in the field without difficulty. The head remains feathered through the second year. The eye is yellow-grey to grey, turning red only in the bird's third year. Generally, this is a silent bird when foraging, but noisy at the roost and at breeding sites. It uses bill clappering at the nest, and its calls are either guttural or high pitched. The nearest description is that of a domestic turkey. Such noises are made when bowing to Greet a mate, by an unpaired male at a nest site, and at the time of nest-relief. Young give a call described by some as 'Lib, lib' when food Begging (Brown et al. 1982). The Waldrapp's flight is like that of a gull, taking a few wing-beats followed by a glide (Bannerman and Bannerman 1968). Birds fly to and from roosts in V-formation, using shallow wing-beats interspersed with glides on slightly bowed wings. In flight, the short legs are hidden beneath the tail. The Waldrapp Ibis is a rather distinctive bird within its much restricted range. It is larger and heavier than the Glossy Ibis, having shorter legs and neck. It is also distinguishable by its head and bill colour.

232 Waldrapp Ibis

DISTRIBUTION AND POPULATION Historically, the Waldrapp Ibis nested in the mountains of central Europe, across northern Africa and into the Middle East. But this range is now much reduced. Nesting is now confined to Morocco, irregularly in Boghari in Algeria, and in Birecik, Turkey. The Waldrapp occurred in central Europe from the 12th to the 17th centuries, ranging from southern Germany and Austria in the valleys of the upper Rhine and Danube Rivers, and in the Alps of Switzerland, Italy and Germany, and perhaps in Hungary and Greece (Rothschild et al. 1897, DesFayes 1987). In Africa and the Middle East, it occurred in two distinct populations. The first nested in Morocco and sporadically in northern Algeria; the second nested in southeastern Turkey and Syria (Geroudet 1965a, Kumerloeve 1978a, 1984, Cramp 1977, DesFayes 1987). Evidence is lacking for reported nesting elsewhere, including Yemen, Iraq and Ethiopia (K.D. Smith 1970). It remains possible that it bred in ancient times in the Red Sea Hills of Egypt and Sudan, where it was depicted as early as 2430-2410 B.C. and was part of the symbolism of ancient Egypt (Meinertzhagen 1930, Kumerloeve 1983). Migratory patterns differ between the eastern and western populations. Certainly, the population that previously nested in central Europe would have migrated south in the winter, and the eastern-nesting birds are still migratory. These birds, nesting in Turkey, winter in Ethiopia, especially along the Red Sea (Bourne 1959). The Waldrapp Ibises nesting in the west may only experience a post-breeding dispersal rather than a true migration (K.D. Smith 1970). The birds nesting in Morocco disperse

from their colonies in June and July, and return in February and March. Their wintering grounds are poorly known, but many individuals have been reported remaining in Morocco and in nearby Mauritania. It is estimated that 70% of the population remained in Morocco for the winter, using three main areas including the Sous and Massa Estuaries (Hirsch 1979b, Thevenot et al. 1982). Even in the middle Atlas Mountains, birds appear to remain all winter (Snow 1952, Hirsch 1982b). Most immature birds do not return to the colony site during the nesting season, and, overall, the Waldrapp Ibis appears to be rather nomadic. It has been reported casually in Iraq, Israel, Saudi Arabia, the Nile Valley of Egypt and Sudan, Yemen, Somalia, Mali, Tunisia and Spain, and even as far as the Cape Verde Islands and the Azores (Hartert and Ogilvie Grant 1905, Salvadori 1899, Valverde 1959, Bannerman and Bannerman 1968). The source of some of these nomadic birds is not known for certain, leading to speculation that undescribed colonies exist in Turkey or elsewhere (K.D. Smith 1970), though the possibility seems to be increasingly remote (Hirsch 1979a). The world's population of this species is small, and continues to decrease, and as a result the Waldrapp is severely endangered (Pegoraro 1989, Department of the Interior 1990). In 1982, the known breeding population was about 550 birds in 13 colonies, 12 of which were in Morocco, and an additional 408 birds in 33 zoos (Michelmore and Oliver 1982). By 1989, there were about 750 birds in captivity. The several nesting populations have their independent numerical trends. The Moroccan subpopulation, nesting in many small colonies and some isolated nests, has been decreasing gradually for decades. At the beginning of the century, the Waldrapp Ibis was an abundant species in Morocco, with a large number of colonies being described by many authors. Meade-Waldo (1903) described a colony on the cliffs of Sale, with each bay occupied by many birds, so thick that the nests touched. By 1920, however, this colony had been extirpated (Jourdain 1922, Bede 1926). The population stood at 1500 birds in 1940 (P. Jackson 1978). In 1965, Geroudet (1965a,b) reported that the species was in "alarming decline'. By 1977, only 13 colonies were extant (Hirsch 1976b). In 1982, the total population there was estimated to be 350-400 birds (367 were counted), including 93 pairs in 12 colonies (Michelmore and Oliver 1982). In 1982, Hirsch (1982b) found 5 colonies in the High Atlas and adjacent plains in eastern Morocco, containing 26 pairs and a total of 108 birds. Two colonies on the coast, outside the proposed Massa National Park, contained 10 pairs, with a total of 76 birds. 5 colonies inside the proposed park contained 57 pairs and a total of 183 birds. By 1984, this had increased to 220 birds. In May, 1989, only 43 pairs were counted in the coastal colonies (Pegoraro 1989). However, in 1990, 78 pairs fledged 72 chicks, and the Morocco population of about 300 birds is now considered reasonably stable (U. Hirsch pers. comm. 1990). The subpopulation nesting in nearby Algeria has always been erratic, but continued to breed there periodically into the 1980s. A small colony was discovered in 1974 in the

Waldrapplbis 233

Djebel Amour near El Bayadh in the central Saharan Atlas Mountains (Ledant et al. 1981). It consisted of 8-12 pairs each year, and never more than 14 (Hirsch 1981b,d, 1982b). In 1982, 14 nests fledged 10 young (R.E. Johnson pers. comm. 1990). By 1984, only 7 birds were seen. By 1990, about 5-12 birds arrived annually but did not breed because of drought. (U. Hirsch pers. comm. 1990). A second colony at Kasar el Boukhari had only up to 12 pairs and appeared to depend on rainfall, which, when absent, resulted in this nesting site being left unoccupied. Sporadic breeding may have occurred up to 1953 at Kasar, where recently used nests were discovered in that year (Kumerloeve 1958). The colony site is now considered to be abandoned (Ledant et al. 1981). The colony in Birecik, Turkey, was for many years the stronghold of the eastern population segment. Some 3000 pairs nested there in 1890 (Kumerloeve 1962). Weigold (1912-13) estimated 1000 birds there in 1912. This site supported 1300 birds in 1953 (including immature birds that were up to one-third of the total), 200 pairs in 1961, and 5 pairs in 1984 (Kumerloeve 1962, 1965, 1984, P.Jackson 1978, Hirsch 1982b, Luthin 1984a). By 1989, only 3 individuals returned to the Birecik colony, and 2 of these birds were killed in a storm. So the colony now appears doomed (Baris 1989a, Pegoraro 1989, H. Peters 1990). These population decreases have been occurring for a long time, with each population trend appearing to be rather independent. It seems worthwhile to consider in more detail the historic and recent status in segments of its historic range, other than those where it has nested most recently. Any nesting that had occurred in Egypt ended millennia ago. European nesting sites were last occupied in the 17th century. In Syria, the last breeding was in 1916 or earlier. In Iraq, there have been no records since the 1920s (Allouse 1953, Cramp 1977), and there is no proof that the species ever bred there, contrary to Bannerman and Bannerman (1968). In Israel, there are no breeding records. A few records of non-breeding birds have occurred since 1935. Up to 16 birds were seen at Elat and one in 1980 near the Dead Sea (Hirsch 1980a, Lambert and Grimmett 1983). Attempts to release captive birds into the wild have been unsuccessful. We witnessed these birds from the Tel Aviv University flying free around the area, perching on the balconies on nearby apartment buildings. In North Yemen, birds have been seen in marshes near Ta'izz. 14 birds, including 2 juveniles, were seen in autumn 1985. In 1986, 9 birds were seen roosting with Cattle Egrets (Bubulcus ibis) in January, and in January of 1987 at least 4 birds were seen again at Ta'izz. There is speculation as to whether these birds might come from Turkey, or from some undiscovered breeding site (Hirsch 1977a, Brooks et al. 1987, Bezzel and Wartmann 1990). In Mali, single birds have been seen on the northern edge of the Niger inundation zone, but certainly not in the numbers suggested by K.D. Smith (1970). In Mauritania, however, a flock of up to 50 birds was seen in February 1951 at Bir Magrein in the northwest of the country, and about 20

birds were seen in the south near Avatane Wells in the Djouf (Dekeyser 1954, Valverde 1957, K.D. Smith 1970). ECOLOGY The Waldrapp Ibis is a terrestrial ibis, but both present and past distributional records suggest that it is not confined to such habitats (Meinertzhagen 1940, H.B. Peters 1928, Hirsch 1981b). Typically it feeds in mountain meadows, stubble fields, short grass, rocky semi-arid ground, on damp ground in lower areas of high plateaus. The colony site at Birecik is on and near the banks of the Euphrates River, and those in North Africa are along streams or on the coast, so they also feed in wetlands, on coastal sandy strips, on river beds and in sandy banks on the seashore. They may also be found feeding amongst cattle and village huts. Ruthke (1977) observed them feeding at a rubbish dump. They avoid tilled fields and pastures, especially if the grass exceeds c. 25 cm. Despite its extensive use of terrestrial habitats, the Waldrapp Ibis appears to be dependent on rainfall, which probably determines the timing and extent of the nesting season and its nomadic movements (K.D. Smith 1970). In winter, birds in the Ethiopian highlands feed on high moors, wet meadows, and by fast-flowing mountain streams and lake margins, up to 3500 m above sea level (Ash and Howell 1977). Birds feed singly or in loose parties, by Pecking and by Probing deeply in the ground, beneath stones, or under plants and clumps of grass. Characteristically they feed while Walking rather rapidly and will Run after insects. The prey is lifted by the tip of the bill and quickly swallowed. Hirsch (1979b) stated that these birds seldom drink, obtaining most of their liquid intake from their food. In fact we have not found any observation of how they catch fish in water. They feed all day long. The diet is broad, reflecting their ability to exploit whatever local prey abundances might occur (Meinertzhagen 1954, Siegfried 1966a, K.D. Smith 1970, Pala 1971, Parslow 1973, Porter 1973, Rencurel 1974). Invertebrates eaten include locusts, crickets, grasshoppers, ants, beetles, spiders, caterpillars, earwigs, scorpions, centipedes, wood lice, earthworms and snails. Vertebrates include fish, amphibians, lizards and snakes, small rodents, and small birds either live or dead, including nestlings. Vegetation eaten includes berries, shoots, duckweed and rhizomes of aquatic plants. Favourite prey for birds in Moroccan colonies are snails picked from shrubs, and in Birecik they eat larvae of beetles and other insects. Aharoni saw one decapitate a young horned viper (Cerates cerates). One Turkish bird contained 95% locusts (Parslow 1973), and one in winter from South Yemen nothing but black beetles (Meinertzhagen 1954). Hirsch and Schenker (1977) reported that the majority of the food of birds at Birecik consisted of mole crickets (Gryllotalpa gryllotalpa), and various beetles and their larvae. Roosts used away from the breeding area can be in trees, on cliff ledges or sometimes in fields. On landing at the roost, birds raise their plumage, croak and peck at each

234 Waldrapp Ibis

other. They interrupt feeding to sunbathe with well extended, drooping wings. BREEDING The Waldrapp Ibis nests in large or small colonies or alone. The Moroccan and Syrian birds tend to nest in many small colonies whereas the Turkish birds formerly nested in large assemblages. Nests are placed on the ground, among boulders on cliffs, on mountain ledges or in holes in rocky cliff faces. These colony sites may be 100 m above the ground. Nest-sites are usually near a river or next to the sea. Historically, Waldrapps nested on castles in Europe and have used artificial platforms erected on their behalf in the Turkey colony site (Robin 1973). Nesting occurs in March and April. However, in dry years, most colonies on the plains are abandoned (Robin 1973). The behaviour of the Waldrapp Ibis at its breeding sites has been intensively studied, and the results have been supplemented with extremely detailed accounts of behaviour under captive conditions (Mallet 1977, Hirsch 1979b, W.L.R. Oliver et al. 1979, Thaler et al. 1981, Michelmore and Oliver 1982, Sahin 1982a,b, 1983a,b,c, Pegoraro 1985). The first birds to occupy breeding sites are males, who acquire and defend a territory with fierce fighting. The Forward Threat involves thrusting or touching with the mantle held erect, the head retracted, and the bill raised and gaping. Birds may beat their wings and grab and pull each other backwards and forwards. Males stand at the potential nest site calling. Activity increases with the arrival of the females, who fly in circles to which the males respond with a guttural 'Hoop hoop' call, while lowering their bill to touch the breast, after which it is raised sharply. This is probably the Bill Popping display. The approach of another bird intensifies this action. The male crouches, passing its bill under its flanks, simulating the insertion of nest material, whilst swaying and making a rumbling sound. If the approaching bird is a female, she will remain inactive until approached but will then lower her head, soliciting allopreening of the neck, back and head feathers. During this action the male continues to utter his 'Hoop, hoop' call. Following allopreening, billing and entwining of necks takes place; the male moving around the female. Such activities may take place for up to a week before pair-formation is completed. Once paired, the arrival at the nest site of one of the birds initiates a mutual Greeting display. Facing each other, heads are first raised and then lowered so that the open bills are nearly touching the ground. This is repeated two or three times; on each occasion the crest and breast feathers are raised and each bird utters a 'Jum' call as the bill is lowered. At other times a slower more deliberate deep bowing is followed by an upright posture, without a call. Such ceremonies are formal and quite deliberate and are followed by self-preening. They seldom occur away from the nest site. Male birds, after selecting a favoured site for nesting, will clean away all remnants of previous nesting material and,

after the pair-bond has been formed, the male will collect new material, arriving at the nest to present sticks to the waiting female. Males will also steal sticks from nearby nests. Both birds assist in building the nest. Nest-building takes about a week to complete. The nest is a loosely constructed structure of twigs and sticks. The birds place a softer lining in the nest-cup, consisting of leaves, grass and paper or other waste materials. Promiscuity is reported in both captive and wild birds, although, in observations of captive birds, approaches by males to other females are usually rejected. Intrapair copulations occur throughout the nesting period, long after insemination could occur (Sahin 1983a). This behaviour appears to be a means of communication and reinforcement of pair bonding in this species. Such behaviour especially occurs in the event of breeding failure at the first attempt, and sexual activity increases amongst unpaired or failed pairs towards the end of the first nesting period (W.L.R. Oliver et al. 1979). Clutches are 3-A eggs. However, often only 2 are laid and only 1 may be laid during droughts (Aharoni 1911, 1929). Rencurel (1974) found that nests constructed in holes usually contained 4 eggs whilst those on ledges only 3 or even 2. The eggs are oval and rough pitted, their bluish white colour becoming stained with brown during incubation. Eggs are 63 X 44 mm, and the average weight is 68 g (Schonwetter 1967). Double clutching can occur (Michelmore and Oliver 1982, Sahin 1983a). Eggs are laid 1 or 2 days apart (Hirsch 1979b, W.L.R. Oliver et al. 1979). Time to hatching averages 28 days. Both birds share incubation; and throughout incubation, the incoming bird often presents nesting material and may have to push its mate off the nest. Hatching success is low. Both birds feed young by regurgitation, in captivity more feedings are made by the male. Non-parental birds also feed young (Sahin 1982a). Eggs hatch 2-3 days apart, and sibling competition is severe, even in captivity where food is readily available. Overall nesting success is low, owing to hatching failure, low food supplies, disturbance and, perhaps, pesticides. In Morocco, 20% of the nests reared no young, 60% 1 young, 15% 2 young and only 5% 3 young (Cramp 1977). In dry years, half the pairs reared no young and most of the rest only one. Chick stealing by neighbouring pairs occurs sometimes. Nestlings fledge after 43-47 days, and the young leave the nesting site to travel to the feeding grounds with their parents. Soon after the young join their parents in the feeding areas, the colony site will be abandoned for the year and nearby roosts occupied at night. The young eventually separate into their own flocks (Ruthke 1977). This ibis does not breed until its fifth year even though it has acquired full adult plumage after 3 years. Older birds appear to be more successful in rearing nestlings than do younger birds. TAXONOMY Although considered to be monotypic, the eastern population of the Waldrapp Ibis may have shorter bills than

Waldrapplbis 235

western birds, leading Siegfried (1972) to suggest that they were isolated. CONSERVATION The Waldrapp Ibis is extremely endangered, owing to longterm population declines, which have accelerated in recent years. In 1990, it was proposed for inclusion in the Endangered Species List by the USA (Department of the Interior 1990). This relict species has attracted an enormous amount of attention from scientists and conservationists. In spite of these considerable efforts, it is likely that the bird may become extinct in the wild very shortly. Although all of the reasons for the population decrease and localized extinctions are not understood, several causes are identifiable. Over the past several thousands of years, climatic change has probably led to a gradual range contraction. The species occurs in Pleistocene fossils in Europe, and the climate there prior to the 16th century was different from that of today. Similarly, the climate of Egypt was more conducive to the species 5000 years before present than it is now (K.D. Smith 1970). Over the shorter term, rainfall variations have coincided with fluctuations in the colonies in Asia Minor and Syria (Safriel 1980). However, failure of the population to respond to more recent climatic amelioration suggests that factors other than climate are now holding numbers down. Waldrapp Ibises were killed and eaten in Europe and Africa, according to early accounts going back to the 1500s (Jourdain 1934, Hirsch 1977a). Extirpation of the population nesting in Syria is particularly ascribed to hunting. Syrians had long believed that eating the flesh of this bird transferred sacred powers from the bird to the eater, and so the species was eliminated from Syria early in this century. Conversely, the Turkish colony, known to have existed since 1879, has been protected by the goodwill of the local human population, who traditionally identify them with birds released from the Ark by Noah. The decline of this colony began with local human population increases and changed agricultural practices. The nearby town has greatly expanded, its population nearly doubling with the immigration of Kurds and Syrian Arabs from about 1958. Thus the colony was confronted by people with little history of toleration for the colonial birds nesting in their midst, and disturbance was severe. Stone-throwing by children caused panic and desertion. At about the same time pesticides against malaria and locusts came into use, and it is strongly suspected that this contributed to the rapid population decrease at about 1959-60, a period when over 600 birds were found dead in the area (Parslow 1973, Hirsch 1976b, 1980a). This highly endangered species has attracted considerable attention from a wide range of observers and conservationists who have initiated attempts to protect breeding areas and the dwindling numbers of breeding birds. The

small Turkish town of Birecik has been the location of much of this activity. Hans Kumerloeve, followed by Udo Hirsch, made valiant efforts to document and save the remaining birds at this colony (Kumerloeve 1962, 1967, 1984, Hirsch 1976b, 1978b). Studies, publications, appeals to international organizations, lectures and the assistance of local politicians created some support. Locally a Waldrapp Ibis day was initiated. In 1973, the World Wildlife Fund supplied Hirsch with funds to increase his attempts to support the colony. Wooden platforms were built; and, with difficulty, birds were encouraged to use these safer nesting sites. Nets were secured below the platforms to save young who fell out of the nests due to disturbance and an attempt was made to hand-rear some of these. In 1983, the returning adults abandoned the traditional colony site to nest near the aviary holding captive nesting birds. This placed the wild birds in a safer situation. In Morocco, disturbance by local people, tourists, and egg and zoo collectors has similarly reduced the colonies, and more protection is vital. Efforts are underway to protect the wetlands critical to the population, at the Souss-Massa Estuary (Salathe 1990). Pesticide use has been and may continue to be one of the most severe threats to the species, which appears to be extremely susceptible (Parslow 1973, Department of the Interior 1990). Dry-land pesticides may have decimated the Turkish population, and residual poisons in fish stocks in both fresh and marine habitats may well have further affected already weakened birds on their wintering grounds. In zoos, breeding of captive birds is meeting with considerable success. The Basel Zoo pioneered this work, and strong success has been attained in Jersey, Tel Aviv, Innsbruck, Nuremberg and Wuppertal. We have observed the captive birds at the Jersey Wildlife Preservation trust and the Tel Aviv University Zoo, where release programmes are in effect. At Alpenzoo Innsbruck-Tyrol 150 young have been produced since 1963 (Pegoraro 1989). Because as many as 750 Waldrapp Ibises are in captivity (all derived from Moroccan birds), the reintroduction of birds back to their natural habitat is, perhaps, the species' best hope for the future (Pegoraro 1989, Akcakaya 1990, K. Brouwer and M.C. Coulter 1990). Attempts to release captive birds are being made in Israel and Spain. Additional studies in preparation for expanded release programmes are now needed to assess hazards and the benefits. In June 1991, a meeting was held to begin developing a conservation strategy for the Waldrapp Ibis, especially to manage a viable captive population for reintroduction programmes (K. Brouwer 1991).

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 319.

Bald Ibis Geronticus calvus (Boddaert) Tantalus calvus Boddaert, 1783, Table Planches Enlum., p. 52; based on 'Gourly a tete nue, du Gap de bonne-Esperance' of Daubenton, 1765-81, Planches Enlum., pi. 867: Gape of Good Hope Other names: Southern Bald Ibis, Bare-headed Ibis, Wild Turkey (English); Glattnackenrapp (German); WildeKalkoen (Afrikaans); Umowanele (Zulu)

IDENTIFICATION The Bald Ibis is a strikingly iridescent dark ibis of relatively large size, about 81 cm long. It is brightly coloured with a whitish head, red bill and fluffy neck-ruff. The adult's base plumage colour is dark green, highlighted by iridescent green, bronze and violet overtones and iridescent copper patches on its shoulders. The neck-ruff is formed by long, fluffy, hackleshaped feathers of a metallic blue-green hue. The ruff surrounds the whitish-tan skin of the bare head and neck. The crown of head is bright red and domed posteriorly. The relatively short bill as well as the feet and legs are dull red. The eyes are red to an orange red. The bare skin of the crown turns bright red during the courtship period. The sexes are alike, although the males are larger than the females, with longer bills and tarsi. Nestlings are covered with a dark grey down, darker on the crown. The soft parts are also dark: the bill is black with a grey tip; the legs and feet are blackish grey. The eye is dark blue. Immature birds are dark iridescent green and lack the copper wing-patches. The head and neck are feathered light grey. The crown lacks the red colour of the adult, and the bill is flesh-coloured with a greyish base. At 1 year, the plumage is similar to the adult, but the copper wing markings are less bright. There is some white on the face at the base of the bill. The crown becomes pinkish orange. At 2 years, the crown becomes wrinkly red and less pronounced in colour than the adult. The bare face is white with some red spots, and the neck feathers begin to form the ruff. This is a relatively quiet bird, having a feeble gobbling sound (leading to an old Afrikaans common name of Wilde-Kalkoen or wild turkey—Elliot 1877). It is, however, noisy at the nest. In flight it will utter a high-pitched call similar to a gull or a feeble edition of an African Fish Eagle (Haliaeetus vocifer). It is described as a high-pitched 'Keeuw-klaup, klaup' by J. Vincent and G. Symons (1948). It also sometimes calls when evading a predator, or when a flock gathers. At the nest it has a wide variety of calls. A piping call is uttered with the bill pointing downwards or resting on the back. The tail is flicked with each note. Similar pipings and gruntings are giver! when fighting and during courtship. Bill clappering occurs before and after copulation. The young utter a shrill Begging call at the approach of the parents. The Bald Ibis has powerful wing-beats in flight and is very crow-like in action (J. Vincent and G. Symons 1948). It flies by gliding and soaring, using air currents and flapping flight to gain altitude. It descends in a long glide, with its bill pointed forward. Within its range, the red cap, pale neck and wing-patch of the Bald Ibis are distinctive among ibises. Contrasted with the Hadada Ibis, it has a shorter bill, appearing straighter, and in flight its wings are more pointed.

238 Bald Ibis

depending on rainfall conditions (Siegfried 1971). In Natal, one survey noted a large decline at 1 colony but increases at 2 others (K.H. Cooper and K.Z. Edwards 1969). There are undoubtedly single breeding pairs that have been overlooked. ECOLOGY

DISTRIBUTION AND POPULATION The Bald Ibis occurs in a restricted area of the high grasslands of South Africa, Natal (Transvaal, Orange Free State), Lesotho, and Swaziland (Manry 1985a, Clancey 1964a, Siegfried 1971). It is no longer found in the Cape Province, having disappeared during the present century. Generally, this is a sedentary species, often remaining near its nesting areas year round. However, some seasonal movements do take place, as some birds move out of their nesting area, but this shift is generally not more than a few kilometres. Historically, some birds moved closer to the coast in winter (Elliot 1877). It has wandered as far as Cape Town, where a single bird was collected at Milnerton in 1906 (L.E. Taylor 1909), and also to Tigerhoek near Caledon and Little Namaqualand, at the mouth of the Orange River (Siegfried 1966a, 1971, Brooke in press). The species is rare. Until recently the population was considered to be about 2000 birds, including non-breeders (Siegfried 1966b, 1971, W.B. King 1981). Recent surveys, however, estimate the population to be much higher. The total breeding population is now estimated at 5000-8000 birds (Manry 1985a,b, Collar and Stuart 1985). The population appears to be stable, although continuing to be threatened (Jaarsveld 1979, 1980, Bonde 1981, Manry 1985a, Luthin pers. comm. 1990), The population is centred primarily in the Orange Free State where 64 colony sites were known by 1980 and 2400 birds, including non-breeders, were counted in one season (Manry 1985a, Luthin pers. comm. 1982). In Transva'ai in 1982, 492 pairs were counted at 44 colony sites, 2 other sites not being censused. In addition, 1294 birds were counted at non-breeding roosts (Allan 1982). In Natal, 50 colonies with about 400 breeding pairs were known in 1982. In Lesotho, Jacot-Guillarmond (1936) reported that the population was decreasing in the lowlands, but it is still common in the mountains. A small population in 2 breeding sites remain in the western mountains of Swaziland (Manry 1985a). At least 4 nesting locations are known in Lesotho. Surveys to determine the population status of this species are rather difficult, as it nests solitarily in some places. The breeding season is staggered, and nesting can be sporadic

This is a dry-land ibis, preferring short grasslands and cultivated pastures, resorting to water only for drinking or bathing. In western Natal, 41% of the ibis* feeding habitat use was in short grassland, 40% was in pasture, about 1 % was on sport fields and lawns, and the rest was in active and abandoned cropland and ploughed fields (Manry 1982). Habitat use changes seasonally, grassland being used exclusively in January, which is early in the wet season. The birds then move to rye pasture in mid-year where they remain until December. They avoid medium to long grass and other vegetated areas. So, as the grasses grow too tall for foraging, the ibises move to heavily grazed pasture and rye fields. In the same way, they are attracted to native and artificial grasslands and rye grass pastures. They are particularly attracted to burnt grasslands in the first week after burning and then return again to forage in the new growth following the rains (Manry 1984a). Arthropods are particularly abundant in winter-burnt grasslands, and the ibises feed on insects disturbed or killed by burning. Their return to the site occurs after the insects have recolonized the regenerating grass. Natural lightning fires, control burning and accidental fires provide much of the short-grass habitat on which this species depends. Its dependence on fire and grazing is longstanding—the former range of the Bald Ibis coincided with that of the black wildebeest (Connochaetes gnou), suggesting they have similar environmental requirements (Siegfried 1971). Manry (1983) pointed out that cultivation has been practised in the species' range in Natal as early as 1600 to 1700 B.C. He proposed that the Bald Ibis was associated with this patchily distributed foraging habitat and has continued feeding in such locations until modern times. Today, grasslands are burned in winter to create firebreaks, and again in the spring to enhance fresh grass growth; ibises benefit from both of these customs (Manry 1985b). The birds feed by walking while Pecking and Probing the soil. On burnt ground, they Walk Quickly picking up living and dead insects. Although wary, they forage around cattle. They also turn over dung and take advantage of plagues of locusts and grasshoppers. They also feed quite close to human habitation, as the local people are benign towards them. The Bald Ibis is extremely gregarious, feeding and roosting in large flocks. The birds move rapidly over the ground, well spaced out. Feeding flocks fly to a foraging area from roosts. They often stay in feeding area for up to a week, before moving to another food source nearby. The diet is poorly known, but Bald Ibises are clearly both hunters and scavengers. Arthropods (especially caterpillars, beetles, grasshoppers), earthworms, snails, frogs, small

Bald Ibis 239

dead mammals and birds form the bulk of their food (Stark and Sclater 1906, J. Vincent and G. Symons 1948, Milstein 1973, 1974, Manry 1984a). They readily take plague insects. The Bald Ibis by no means disdains carrion and acts as an efficient scavenger (Stark and Sclater 1906). Milstein (1973, 1974) reported on Bald Ibises' habit of collecting buttons, which he suggests they may mistake for beetles. But he also found, at the nest, a hollow porcelain tube, a beer-can pull-tab, a toothpaste cap and two metal rims of vaccine bottle tops. In one small colony, he found 156 small buttons, items not readily available in the foraging habitat. At dusk the ibises assemble in their roosts, on cliffs or in trees, in groups of 40-50 with other ibises and herons. Birds fly to foraging areas from their roosts, usually located in or near their breeding areas amongst rock cliffs. As many as 40-150 birds will share their roost with Cattle Egrets, Sacred Ibises, and cormorants (Pocock and Uys 1967). Many of the roosts have been in use for a very long time; Pocock and Uys (1967) suggested that this might be up to 100 years. Bald Ibises will sunbathe with wings wide open and touching the ground. BREEDING This ibis is a ground-nester, especially on cliffs along deep river gorges. It often nests by waterfalls, even in the spray zone or under the falls. In other colonies, however, no water is present (Milstein and Siegfried 1970). Birds may nest alone, but the Bald Ibis is more commonly a highly gregarious species nesting in colonies of 40 pairs or more. Numbers vary at traditional sites from one year to the next. Although sedentary for the most part, some intercolony movement has been documented (Milstein and Wolff 1973). Claiming of potential nest-sites and pair-bonding occur as early as June, as the birds spend increasing time at the colony sites and less time off foraging. Males occupy the nest-site first, defending them by bill jabbing and sparring. Courtship behaviour has been difficult to observe. At the display site, initial interactions seem perfunctory with copulations appearing to be somewhat randomly directed at different partners. A male and female will behave as a pair, and then separate after a few days. A pair will move from ledge to ledge, performing courtship displays. Mutual preening takes place, particularly directed at the distinctive neck feathers. Pairs grab each other's bills, wagging their heads strongly. Twig presentation can be made by either one of the pair, and this is accompanied by raising and lowering of the sideways-turned head. A second bird will often grab the twig and wiggle it. After being apart for some time, the male will peck at the female's bill, body, neck or wings with his closed bill. The females responds by extending her neck vertically with the bill resting against her neck, concealing her face and bill within raised neck plumes. Sometimes the female will rest the extended bill against her neck or on the ground. To copulate, a male will bite a female's foot to induce

crouching, and then mount to copulate. During the brief copulation, the male grasps the female's bill and shakes it vigorously. Often the two birds will then stand erect with bills raised, snapping their mandibles together. Upon return of the mate during incubation or brooding, a bird on the nest will flick its tail and make piping noises, bending its neck so that the nape of its head touches its back. Intruders receive snapping bills pointed at them. Upon successfully repelling the invader, the bird on the nest will rub faces with its mate. They squirm, twist their necks and utter squealing noises. The male collects nesting material for the female to use in building the structure. The female presses her breast down to form a hollow of the cavity (Brown et al. 1982, Manry unpub. report). The nest itself is placed on a cliff ledge or in holes in the cliff face. There is no confirmation of treenesting, though the possibility is discussed by Pocock and Uys (1967). The nest is quite large, about 50 cm in diameter and 15 cm high. Eggs are laid between August and October, in the dry winter season. Egg-laying can be carried out over quite an extended period. The clutch is 1-3 pale blue-brown spotted eggs, averaging 1.9/nest in the Transvaal (Milstein and Wolff 1973). Eggs are laid at 2-day intervals. They measure 63 X 43 mm and weigh about 68 g (Milstein and Wolff 1973, Milstein and Siegfried 1970). In Transvaal, 1.8 young hatched per nest and in Natal 0.83, although this varies according to the season (Milstein and Wolff 1973, Manry unpub. report). Both adults feed the young by regurgitation, which develops into a tussle with older chicks, who Beg by head bobbing while giving shrill cries. To provide food from its gullet, the adult rests its bill on the floor of the nest. Older chicks tap the bill of their parent after its arrival at the nest and then insert their bills into the adult's opened mouth. The young fly after about 55 days, in mid-October to December (Manry 1985b), but depend on their parents for food for up to 2 months. Nesting success depends on available food supplies. In droughts, many birds do not breed, and, when food shortages occur, the young starve (Manry 1985c). Ibises are also lost to predation, including by Jackal Buzzards (Buteo rufofucus), Cape Eagle Owls (Bubo capensis), Pied Crows (Corvus albus) and man (Milstein 1973). Attacks by Black Eagles (Aquila verreauxi) have been witnessed, and one pair of these birds, which occupied the nearby cliffs, caused the Bald Ibises to abandon their nests (Allan 1982). Overall, however, breeding success is generally good. In Transvaal, South Africa, in 1983 about 400 pairs raised 277 young. TAXONOMY The genus Geronticus contains two species widely separated in Africa. Both have similar plumage and nest on cliffs. The southern species is sedentary, whereas the more northern species is migratory and somewhat nomadic. The relationship of the Geronticus to other ibises is obscure (Holyoak 1970), but Fry et al. (1985) suggested that Geronticus and Bostrychia may be related.

240 Bald Ibis

CONSERVATION The Bald Ibis has long played a role in human history (Kumerloeve 1978a, J. Vincent and G. Symons 1948). Historically, young were taken to be used as medicine by the Zulu. It certainly was once more widespread in southern Africa than it is at present (Siegfried 1971, Milstein and Wolff 1973). It is now confined to a limited range and maninduced foraging habitat. Hunting has always been a factor. The Boer farmers shot this ibis during the South African wars to supplement their meat supplies. Settlers also hunted it for both food and feathers. Although the species is officially protected throughout its range and colony sites are under protection in Transvaal (Milstein and Wolff 1973), local hunting pressure remains, as people take young from the nest for food and for other reasons. Such hunting may be the largest shortterm threat to the species. Other disturbance, as well as loss of nesting sites, also threatens the bird. However, the main long-term reason for the species' decline during the present century was the spread of karoo vegetation following severe overgrazing of the grasslands (Siegfried 1966a,b, 1971, Brooke in press). The encroachment of brush and karoo vegetation in the Cape Province and afforestation by conifers in Natal and Transvaal cer-

tainly make the grassveld unsuitable (Siegfried 1971, Manry 1985a). This shy bird is now entirely dependent on farming communities, who own the sheep and cattle pastures in which they feed and the cliffs on which they nest. In the Transvaal, cash incentives and warnings of prosecution were successful in re-establishing one colony (Allan 1983a). The habitat requirements of the species are very exacting. For example, it is found only in areas where the grasslands are burned annually, and not where the fire cycle is longer (Manry 1985c). Some parts of its former range may be made more suitable through restoration of its grassland environment, and irrigation schemes may be increasing habitat quality and consistency. A small captive population in the Pretoria Zoo may become the core of a reintroduction programme (Brooke in press). A determined programme of education, habitat improvement, and cooperation of land owners can save this bird from further declines.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 320.

Black Ibis Pseudibis papillosa (Temminck) Subspecies: Pseudibis papillosa papillosa (Temminck) Ibis papillosa Temminck, 1824. Planches Color., livr. 51, p. 304 and text: India and Ceylon (=Sri Lanka) Pseudibis papillosa davisoni (Hume) Geronticus Davisoni, Hume, 1875, Stray Feathers, 3, p. 300: Pakchan Estuary, Tennasserim, India Other names: White-shouldered Ibis, White-necked Ibis, Red-naped Ibis, Burma Ibis (davisoni), Tenasserim Black Ibis (English); Schwarzen-Ibisse (German); Ibis Noir Oriental (French); Baza, Kala Baza, Karan Kul (Hindi); Kulo Dochara (Bengali); Kala Akohi Bog (Assam); Nella Kankanam (Telugu)

IDENTIFICATION The Black Ibis is a dark ibis, about 68 cm long, distinguished by the white wing-patch that is visible while the bird is in flight. The common name belies the adult base colour of this ibis, which has a dark brown body, neck, head and wings. The dark, glossy blue secondary feathers of the wing are sometimes flecked with white. A large, broad patch of white runs along the leading edge of the wing on the lesser wing-coverts to well below the carpal joint. Although prominent in flight, this feature shows only as a thin white line when the wings are closed. The short legs are hidden beneath the steel-blue tail in flight. The tibia has feathering halfway to the tarsus joint. The head, somewhat square shaped, is black and .distinctively capped with a triangular patch of warty red skin, absent in davisoni. The eye is orange-red to bright red. The curved bill and the broad, stubby feet are horn coloured. The iris, legs and feet turn bright red at the commencement of breeding. The male is slightly larger than the female. The subspecies davisoni and papillosa arc distinguished by the former lacking the red cap. Young birds are dark brown all over, with paler breasts. They lack the iridescent tinge on their feathers. The pale horn-coloured bill is shorter than that of the adult. The Black Ibis is generally a silent bird when feeding but is noisy at the evening roost. There, its call is loud and trumpet-like. It is sounded particularly when the breeding season approaches, as the male birds advertise at nest sites prior to pair-formation. This species can be confused at a distance with the Glossy Ibis, but the Black Ibis is larger and shorter legged and is clearly distinguished by the brilliant red head (in papillosa) and white wing-patch. In the Mekong Delta, it is said to join feeding associations with the Giant Ibis, but the latter is substantially larger. In all comparisons, the white wing-patches distinguish this species in flight. DISTRIBUTION AND POPULATION The Black Ibis was formerly widespread in the Indian subcontinent through Southeast Asia, in Pakistan, Nepal, India, Burma, western China, Thailand, Laos, Cambodia, Vietnam and the Malaysian Peninsula. The western subspecies is papillosa, formerly occurring in Pakistan and Nepal, south to India (Mysore and east to Bengal and Assam), and Bangladesh.

Black Ibis

The blue ? indicates the race davidsonii.

The eastern subspecies is davisoni, formerly found throughout Southeast Asia, from Burma, to northwestern, central and peninsular Thailand, Malaysia, Cambodia, Laos, Vietnam (to southern Annam and Cochinchina), and north into Yunnan, China. It has been reported from Borneo for many years, but its breeding status has not been documented. The species is now rather patchily distributed throughout its remaining range. In Pakistan, where it was once widespread, occasional sightings are made during the rainy season only. On the Indian sub-continent, it is found in small populations (S. Ali and S.D. Ripley 1968, T. Roberts pers. comm. 1990). It is less rare in Rajasthan southward through India, and in Gujarat it is still quite common (D.A. Scott 1989). In Bengal and Assam, it is locally common. In the Nepalese terai, it is found in some numbers along the river banks. There is a doubtful record ofpapillosa from the Arakan, which probablyvrefers to davisoni. It is not recorded from Sri Lanka or the Andaman Islands. The subspecies davisoni is increasingly rare and has been brought to near extinction (Bain and Humphrey 1980). In Thailand there have been no records since 1937, except for one unconfirmed report from the Thale Noi wetlands (W.B. King 1981), nor are there recent reports from Burma or Malaysia. The last record from China was from Yunnan in 1889. Vo Qoy and Le Due (pers. comm. 1990) consider that the Black Ibis is still to be found in parts of Cambodia and Laos, and also in southern Vietnam near the Cambodian border (Wildash 1968, D.A. Scott 1989). Two birds were seen in Dong Nai, Vietnam in 1991 (Eames 1991). Since 1979, it has been reported, without total certainty, several times from Indonesian Borneo, most recently from the Kalimantan wetlands in 1983 and from the Barito Swamps in 1989 (MJ. Silvius and WJ.M. Verheugt pers. comm. 1990). Silvius and Verheugt report that a small number are believed to survive along interior rivers in Kalimantan, Indonesia.

243

The species is generally non-migratory in India and elsewhere. We have seen it in Gujarat even during droughts. However there is some evidence for population movements in response to monsoons, particularly dispersal of young birds. Birds are found in Pakistan only occasionally and during the rainy season. They also occur casually elsewhere outside their normal range, such as in Burma and Borneo. Population sizes are not known. Thailand was once the centre of the subspecies davisoni population and it was once common there (Bain and Humphrey 1980). In Gujarat, India, V.C. Soni (pers. comm. 1990) of Saurastra University found 10 roosts totalling 325 birds in the Rajkot area and a further 8 roosts totaling 270 individuals. We have seen flocks of 20-30 birds in many parts of this state. S. Faisa (pers. comm. 1990) found 30 birds in Tamil Nadu, where it is generally rare, though we have seen them in small numbers in nearby regions. Other recent censuses include 200 in winter at Chandpata Lake, Madhya Pradesh, and 45 in winter at the Karnataka Tanks, Anantapur and Bangalore, India. The Asian Midwinter Waterfowl Survey in 1988 recorded 220 in India (Rajasthan, Uttar Pradesh, Madhya Pradesh) and 62 in Nepal (Royal Chitwan National Park) (Van der Ven 1988). The population size of davisoni is impossible to ascertain, if it survives at all. ECOLOGY The Black Ibis occupies a similar ecological niche in Asia as does the Hadada Ibis in Africa. It prefers to feed near water, but uses a wide range of aquatic and terrestrial foraging sites, especially in shallow water, around the edge of marshes, in other wetlands, jheels or man-made tanks, coastal estuaries, river edges, paddy-fields, grasslands, cultivated fields. Where it is not persecuted, it will feed close to human habitation, in urban areas foraging amongst open sewage, water-well spills and flooded fields (V.L. Dharmakumarsnhji pers. comm. 1990, pers. obs. J.A.H.). Soni (1988) identified several specific feeding sites, including crop fields (usually after harvest), grassland, sewage, garbage, litter, carrion, manure heaps, shallow muddy shorelines, estuaries and marshlands. Characteristics of behaviour varies, depending on the history of disturbance. In India, the Black Ibis is not a shy bird; it feeds almost under the feet of villagers, and when disturbed it often flies round in a short circle to return to the same area from which it took off. In Southeast Asia, however, where it is rare and widely persecuted, it is extremely shy (Wildash 1968). The Black Ibis is a tactile feeder, like all ibises, and adapts the techniques used according to food taken. Its feeding behaviours include Standing, Shallow and Deep Probing, Pecking, Groping and Head Swinging. Soni (1988) described the Black Ibis as using Groping and Head Swinging along muddy shores. It will Stand quietly while Probing deeply or shallowly, or Pecking. In small parties, the birds Walk Slowly in the same direction across a field keeping well spaced out and seldom interacting. The birds remain separate from other species, except when gathered at carrion and sewage, where they are often in the company of Cattle Egrets (Bubulcus ibis), House

244 Black Ibis

Crows (Corvus spendens) and Jungle crows (Corvus macrorhynchos). During pair-bonding, females will occasionally beg food from males on the foraging grounds (pers. obs. J.A.H.). Soni (1988) considered the simple repertoire of feeding behaviour to be something of a constraint. The species is nevertheless, rather adaptable. It can change diet according to availability of food supply, as demonstrated during the recent drought of several years duration in Gujarat. When groundnuts ripen in Gujarat, huge flocks of Demoiselle (Anthropoides virgo) and Common (Grus grus) Cranes feed extensively on this food, and they are joined by Black Ibis. With the lack of fresh water and failure of the crop, all the cranes left the area, but the ibises remained, changing their diet by exploiting the carcasses of cattle killed in the drought. They fed on the carrion and on large quantities of associated insect larvae. This food source, along with domestic sewage, provided a major portion of the Black Ibises' intake throughout the period. The species' omnivorous diet includes earthworms, millipedes, centipedes, larvae and pupae of house flies, rat-tailed maggots, dung beetles, locusts, grasshoppers, thrips, scarab beetles, cocktail beetles, white grubs, collembola, frog tadpoles, garbage beetles, small crustaceans, lizards, small slow-moving mammals, and even birds. It also appears to consume groundnuts, and leaves of cultivated and wild plants (Soni 1988, pers. obs. J.A.H.). As dusk falls, the scattered birds collect to roost communally usually in large trees. Where there are no trees, we have seen them use small islands or the centre of large fields, where they will stand close together in the open and fly off silently if disturbed. BREEDING In Gujarat, Black Ibis have two distinct breeding seasons, one in October and November and a second more regular one in March and April. These seasons precede the periods of coastal monsoons. Elsewhere, the timing of the nesting season varies geographically depending on the timing of the monsoons. The Black Ibis nests singly, usually high in a tree such as a peepul or banyan, nearly always occupying an old nest of a bird of prey, a vulture or a crow. Often the tree is very close to human habitation. We have seen a nest used annually in the grounds of the Palace of Jasdan, and in the tall palm trees in the Palace grounds in Porbandar. Conversely we have seen several nests quite close together, with at least 2 nests in one large tree in the riverine forest belt along the rivers in the Nepalese terai. Nests are often used annually in spite of nest failures due to harassment by House and Jungle Crows. Courtship behaviour has not been well studied. Prior to pair-formation, the male issues a trumpet-like call when advertising from its nest site or on the wing. Copulation is on the nest or in a nearby tree, but not on the ground. Both birds collect new material for the nest and this activity continues well after incubation commences. Two to four eggs are laid. They are pale bluish green, usually but not always spotted and flecked with red. The average size of 59 eggs measured was 63.0 X 43.8 mm

(Baker in S. Ali and S.D. Ripley 1968). Incubation lasts 33 days (Kumar and Soni 1984). Young are fed at the nest by regurgitation, and fledged young will still beg for food after leaving the nest. TAXONOMY The two races of the Black Ibis were once placed in separate genera (Inocotis papillosus and Graptocephalus davisoni). Elliot (1877) and other authors recognized their similarity but considered them to be seperate species, Pseudibis papillosa and P. davisoni, and even today some authors give specific status to each. Holyoak (1970) pointed out how similar they really were, and despite their apparently allopatric distribution, proposed that they be considered a single species, which we and most others have followed. CONSERVATION The eastern subspecies of the Black Ibis (davisoni) is highly endangered. In fact, this formally widespread species is almost entirely dependent on human tolerance. Its adaptable food choice means that, without persecution, it can survive under both wet and dry conditions in a wide range of wet and dry habitat, including near habitation. However without secure nest-sites, it has no chance of long-term survival. Hunting in Pakistan, often for sport, and shooting in Southeast Asia for food have played a large part in its demise from those areas. In other areas, warfare has reduced both populations and natural habitats. It is a sad fact that former havens provided by religious sites are now less available throughout the species' range. The tolerance of the Hindu population has until now played a major part in the Black Ibis' well-being in Gujarat, although even here continuing drought and the felling of trees present a growing threat to its survival. A combination of the rapidly increasing human population in all these regions, necessitating more intensive cultivation, draining of wetlands, use of pesticides, and nearly continuous warfare has virtually wiped out this species, as it has many other storks, ibises and spoonbills that were formally present in very large numbers. There is a need for increased surveys over the range of this species (Holmes and Burton 1987), and for identification and protection of nesting and feeding sites from human intrusion. M.J. Silvius and WJ.M. Verheugt (pers. comm. 1990) suggest that in Indonesia, surveys should be conducted in the Mahakam Lakes region and upper Mahakam River. The species must also be protected from hunting throughout its range, as it is now in Thailand (W.B. King 1978). The very recent discovery of two birds in Vietnam, in the Nam Bai Cat Tien National Park, is a matter of great importance. However, it is also a matter for concern in that serious conflicts exist between park officials and fishermen living in the park, which led to a shoot-out and deaths (Anon 199Id). It is hoped that relocation of residents will provide an opportunity to maintain these wetlands. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 320.

Giant Ibis Thaumatibis gigantea (Oustalet) Ibis gigantea Oustalet, 1877, Bull. Soc. Philomath. Paris, Ser. 7, 1, p. 25: Mekong River, Cambodia Other name: Mekong Ibis (English)

IDENTIFICATION The Giant Ibis is the largest member of the ibis family, up to 103 cm tall. It is a dark bird, with a featherless black head and neck. The base plumage colour of the adult is brownish. The lower part of the neck is covered with greenish brown feathers edged with grey. The back is dark brown with a green gloss. The scapulars are dark brown, the primaries brownish black, and the secondaries and wing-coverts dark green. The wings extend to the end of the tail. The head is large and nearly square. The head and upper neck are bare, covered with black skin with prominent black bars across the head and back of the neck. The bill is brown to corn brown, or pale brown (Delacour and Jabouille 1931, B.F. King and E.G. Dickinson 1975, pers. obs. J.A.H.). The legs and feet are red to violet, and have also been reported to be straw brown (Delacour and Jabouille 1931, Wildash 1968, B.F. King and E.G. Dickinson 1975, Lekagul and Cronin 1974). For an ibis, this species is remarkably large. Example measurements are wing 52.3-57.0, tail 30.0, tarsus 11.0, culmen 20.8-23.4 cm (Delacour and Jabouille 1931). The sexes are alike, but females are, on average, slightly smaller than males. The differences in the description of soft-part colours suggest that these change seasonally. Little is known about the plumage of juveniles. One juvenile measured was smaller than an adult (wing 53 cm and culmen 18 cm) (Delacour and Jabouille 1931). Elliot (1877) described one specimen he considered to be a juvenile, but this description did not differ from that of an adult. It seems likely that juveniles lack the iridescent gloss of the adult. Its size alone will distinguish this species from other ibises. It has a heavy body, long neck and long bill, though the tarsus is relatively short. It has been seen with the Black Ibis, which is smaller and has a white wing-patch. DISTRIBUTION AND POPULATION This bird formerly nested in southeast Thailand, central and northern Cambodia, southern Laos, and southern Vietnam (King in Collar and Andrew 1988). Its historic nesting range was centred in Cambodia, extending into the Mekong Valley wetlands of eastern Thailand and the Saravane district of Laos. It nested at Phu-Rieng near Saigon in 1925. Fisher et al. (1969) suggested that it could be called the 'Mekong Ibis', to emphasize that its range centres around this great river. In fact, the species has never been proven to breed outside of the Southeast Asian mainland drained by this river. The species is certainly extinct in Thailand (P. Round pers. comm. 1989). Fisher et al. (1969) believed that its recent breeding grounds are scattered over a stretch of lowlands from the Mekong River on the Laos border area, 325 km to the southwest into the valley of the River Sen for 50-65 km north of Kompong Thorn. The latter region is where Delacour found the species numerous in 1927-28.

Giant Ibis 247

E.G. Dickinson 1975). One was collected from a paddy-field in Trang in the Malay Peninsula in 1896. It is reported to occupy the same feeding habitats as the smaller Black Ibis, and to feed with it. The birds appear to feed in pairs and small groups (Wildash 1968, Milton in Fisher et al. 1969). In times of drought they are said to aggregate at permanent water holes (Bain and Humphrey 1980). Virtually nothing is known of their behaviour. It is reported to be a shy bird. BREEDING Again, virtually nothing is known of the nesting habits of this rare species.

Vo Quy (pers. comm. 1989) considers that the birds remain only in the Mekong Delta of southern Vietnam and along the border of Cambodia. He suggests that they may be found in the mangrove forests of Nam Can, the Melaleuca forests of U Munh, and in Dong Thap Muoi. Indeed, C.S. Luthin (in Collar and Andrew 1988) reported small numbers in southern Vietnam at Dong Thao Muoi, an inland delta of the Mekong. Giant Ibises have not been seen in this area, however, since 1980 (D.A. Scott 1989). Birds have been collected as vagrants outside their usual area. These were found in peninsular Thailand and Malaysia on the opposite side of the Gulf of Thailand (Fisher et al. 1969). Little is known about population sizes. The Giant Ibis was numerous in 1928 and 1929 in the valley of the River Sen, north of Kompong Thorn (Deiacour and Jabouille 1931). Wildash (1968) found it quite common in places in 1968. Population estimates range from 100 birds (Bain and Humphrey 1980) to statements that the species is extinct. ECOLOGY The Giant Ibis is a lowland bird, feeding around lakes, in swamps and marshes, on open plains, in open forests and clearings, and in ponds in deeper forests (B.F. King and

TAXONOMY Although for many years this distinctive, stork-sized ibis was assigned to its own genus, some recent treatments merge it into the genus Pseudibis, implying a close relation to the Black Ibis (Pseudibis papillosa), with which it occurs (J. Steinbacher 1979). Olson (1984) stated that the appropriate anatomical study has not been done and cautioned against relegating the species to Pseudibis without such a study. This is a very distinctive ibis, and it is quite logical that this distinctiveness should be recognized taxonomically until morphological, behavioural and biochemical studies have been completed that demonstrate otherwise. Therefore, we have kept it in the genus Thaumatibis. CONSERVATION The Giant Ibis is in as much trouble as any of the endangered ibises. It is not clear how many survive and where. As the species account suggests, almost nothing is known of its basic biology. This is a striking, unique ibis—owing to its size, which is remarkable for a member of the family —and it deserves to be saved from extinction. Unfortunately, no conservation measures can be proposed because no study has been made of the species in the wild. This must be remedied: the present distribution must be determined and secure reserves set up to accommodate the species. Much of this species' habitat has been destroyed by draining for rice and other crops and by the ravages of war, which has plagued this region for decades. Add to this the ever-increasing human population in parts of its range, and it is clear that only protection in nature reserves can give this vulnerable species a slim chance of survival. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 247.

Oriental

Crested Ibis Nipponia nippon (Temminck) Ibis nippon Temminck, 1835, Planches Color., livr. 93, pi. 551: Japan Other names: Japanese Crested Ibis, Japanese Ibis, Rosy Ibis, Chinese Ibis, Asian Crested Ibis (English); Toki (Japanese)

IDENTIFICATION The Oriental Crested Ibis is a striking bird, with its rosy-tinged white plumage, dark bill and pronounced crest. It is a medium-sized ibis, c. 56 cm long, appearing to have long wings and a rounded tail in flight. The non-breeding plumage is basically white. The crest feathers are sometimes tinged with pink, and the back and wings are also tinted a pale shade of pink. We have seen museum specimens from Japan in which the remaining coloration is orange. In flight, the underwings show quite strong pink colouring as does the tail. The black, downcurved bill has a red tip. The unfeathered portion of the face is a bright red. The iris is red, and the eye has a yellow ring encircling it (G. Archibald and S. Lantis 1979). The legs and feet are reddish brown. Prior to the breeding season a gradual colour change takes place, as the head, back and neck take on a grey coloration. This change is cosmetic, rather than structural, and originates in a black secretion from an area of skin under the feathers surrounding the ibis' naked face. The bird applies this tint to its feathers by daubing (Uchida 1970, 1974). The black substance adheres to the proximal feather barbules creating a grey or blackish tinge, which becomes more marked each time the daubing takes place. Firstly the crest, then the head and neck, and finally the back become dark grey. This coloration remains throughout the breeding season until a post-breeding moult returns the feathers to white. Immature birds have cheeks covered with downy feathers, and the rest of the face is a bare and orange yellow in colour. The general immature plumage is a dusky cream, glossed with rose. The primaries are a blackish brown, and the legs and feet light brown. The irides are a light yellowish brown. The Oriental Crested Ibis is generally silent. Its infrequent call is a 'Gak', given before taking flight and while flying. It appears to be used as a low-intensity alarm call and for localization. A 'Gak-gakgak' is given in response to disturbance. A growl-like call is given in agonistic situations. This is the only all-white crested ibis, and should not be confused with any other in its extraordinarily limited range.

DISTRIBUTION AND POPULATION Originally the core of the breeding range of the Oriental Crested Ibis was in eastern Asia. It occurred from the Ussuri River in eastern USSR and in eastern Manchuria, as far north as Heilongjiang through central China (south to Anhwei and Chekiang west to Shensi and Kansu), Korea, and east to Japan (Dementiev and Gladkov 1951:433-437, Yamashina 1977a,b, Cheng 1976).

250 Oriental Crested Ibis

In recent years the bird has been known in the wild only from Shaanxi Province, China, the Demilitarized Zone of Korea, and Sado Island, Japan (G. Archibald and S. Lantis 1979). At least the northern populations of China, Japan, and Manchuria appear to have been migratory, moving in autumn into southern Japan, the Yangtze River basin of China, and the islands of Taiwan and Hainan. It is thought that the recent sightings in Korea were of birds wintering there. Southern populations in China and Japan were more sedentary, remaining year-round except for localized movements. In China, Yinzeng found that after fledging in July the ibises moved to the lowlands 20 km from their nesting area (W.W. Brehm-Fonds 1984). Populations of the Oriental Crested Ibis were reasonably large until the mid-1800s. They subsequently decreased due to shooting, habitat loss, and pesticides. It is presently one of the world's most endangered birds. The changing population in Japan has reflected the level of its protection. Before the Meiji Era, the species was protected by the ruling classes and often seen in suburbs of Edo (now Tokyo). However, from the Meiji Restoration in 1886 and the introduction of firearms, down to 1892 when hunting restrictions were established for the first time, the ibis and other large birds were hunted (Takashima 1957a). The decrease was precipitous, occurring in the three decades after hunting was permitted (G. Archibald and S. Lantis 1979). The species apparently survived only in the Niigata and Ishikawa Prefectures, on the Noto Peninsula and on Sado Island, where it was rediscovered by Dr S. Uchida in 1933 (Takashima 1957a). In 1936, 27 individuals were known to exist, and for many decades this population was the only known to survive. They last bred in the wild in 1973. In 1981 the remaining 5 wild birds were taken into captivity, joining another bird, to form the nucleiis of a captive breeding population. By 1984 the captive population had fallen to 3. The population in the USSR has never been well documented. It has not been observed there since 1983. Although the present status there is unknown, on the basis of a report by Roslyakov (1977), it is suspected that some may survive (G. Archibald and S. Lantis 1979). Oriental Crested Ibis populations in Korea are well documented, but the bird apparently never nested there (Gore and Won 1971). L. Taczanowski (1891-93) reported 50 north of Seoul in 1888. There were no more reports until 1965 (Fisher et ah 1969:184-185). Individuals were dis-

covered there in the winter of 1977-78 by George Archibald (Thiede 1982, G. Archibald and S. Lantis 1979). No confirmed reports exist after 1981 (W.W. Brehm-Fonds 1984). In China, the populations apparently decreased gradually over the past century (Vaurie 1972). Oriental Crested Ibises were unreported from China for over two decades, from the late 1950s until 1972, when several were caught for the Beijing Zoo. One captive ibis remained there as of 1984. A government-sponsored survey in 1974 found no additional wild birds. After a 3-year search of eleven provinces (20000km 2 ), a breeding group was finally located in the Qinling (Tsingling) Mountains of central China in 1981 (Dr Liu Yinzeng in Luthin 1984a). As a result of the discoveries in Japan and China, in 1984 the known world population numbered 21 birds, including 1 captive in China and 3 in Japan. By 1989, it had grown to 49, including 3 captives in China. ECOLOGY The remaining ibises in China occur in forest patches in the high valleys of the Qinling Mountains, 1200m above sea level. The birds have been reported roosting in pairs throughout the year. Such pairs choose trees near other pairs for winter night roosting. Oriental Crested Ibises feed by Probing in shallow open water, especially in rice-paddies, freshwater ponds, lakes and streams in forested areas. The presence of shallow water may be crucial; in China, Dr Liu Yinzeng of the Institute of Zoology in Beijing found that high water levels in streams apparently decreased food availability during nesting (W.W. Brehm-Fonds 1983). During the non-breeding season they feed in groups and frequently shift feeding sites among rice paddies. Juveniles disperse in winter and forage solitarily in lowland aquatic habitats. An upright posture is assumed in alarm. In Japan the ibises appear to be rather tame, especially the juveniles. They feed on small fishes and molluscs. In China, they feed on aquatic vertebrates and aquatic and terrestrial soil invertebrates. The diet includes fish (carp, catfish, weatherfish), frogs, newts, river crabs, crayfish, water-scavenger beetles, diving beetles, beetle larvae, crickets, snails, mussels and earthworms (Yinzeng in Luthin 1984a, Yamashina 1978). The diets change seasonally, and intake appears to be reduced in winter. BREEDING Oriental Crested Ibises returned to their nesting area in October in China and January in Japan. The remnant birds in China nest only in forest patches in remote high mountain valleys. These (and similarly high mountain birds in Japan) nest solitarily. However, lowland populations in central China and Japan apparently were colonial. The birds appear to be monogamous, and, as noted above, paired outside of the nesting season. Mutual preening is common within such pairs. It appears to be solicited by an ibis nibbling the bill of its associate before lowering its head to be preened (G. Archibald and S. Lantis 1979). A rejection of the invitation to preen is indicated by a Head Bobbing display.

Oriental Crested Ibis 251

During the breeding season, ibises preen in such a way as to distribute black pigment over their feathers. As described by Uchida (1970), daubing behaviour includes rubbing the head and neck against the shoulder feathers for as many as 30 min per bout. This occurs after bathing, taking 5 or 6 daubing sessions to distribute the pigment. It seems that the complete repertoire of nesting behaviour is not yet known. Twig presentation with Head Bobbing seems to be an important aspect of pair-formation and pairmaintenance and may occur several months prior to nesting. When no potential mate is receptive or present, a bird may engage in Twig Carrying for many days. This behaviour often precedes a Mounting display in which the male mounts the receptive female and shakes his tail without actual copulation (G. Archibald and S. Lantis 1979). Such pseudocopulation occurs frequently, many times per day especially when other ibises approach the pair. Bill touching and allopreening are common during courtship. The nest, constructed of twigs, is 50-70 cm in diameter. It is placed in a pine or oak tree, rather far out on a branch (G. Archibald and S. Lantis 1979). The nest-cup is lined with soft material, including leaves and mosses. The clutch is 2-4 brown-spotted blue-grey eggs, each weighing 65-75 g and 63-68 mm long. In 1983, one of the captive Chinese pairs built several nests, but never laid any eggs. Incubation begins in March in China and (formerly) mid-April in Japan. Each pair defends its nesting territory. Both sexes incubate for 28-30 days. When departing the nest during incubation, the adults cover the eggs with the nest-lining material. Hatching in China is in mid-May. At hatching young are light grey, with a featherless head and orange-red legs. In China, nestlings are fed by regurgitation and fledge in June. Juveniles remain with the parents near the nesting area until July in China and until early September in Japan. Known birds have had limited nesting success. In China, Zing found in 1983 that crows destroyed clutches, but that the ibises can renest. One of the captive birds in Japan died from egg binding. Encouragingly, in 1989, the 6 pairs of wild birds laid and hatched 7 eggs. Survival of hatchlings depends on continued food availability (W.W. Brehm-Fohds 1983). The adults and young appear to remain together for some time. Birds from the Qinling Range move to the Hanjang River, 20km away during the dry summer months, but the adults return with their young to the breeding grounds in winter. TAXONOMY The interesting cosmetic coloration of this species caused considerable confusion among early taxonomists. Pere David obtained specimens of the grey form, naming them Ibis sinensis. D.G. Elliot (1877), following Oustalet, believed them to be young of the year. Yasuo Uchida definitively explained this unique type of cosmetic coloration. Perhaps owing to its coloration and the close relationships between many plant and animal species of China and the southeastern USA, the Oriental Crested Ibis is thought to be most closely related to the American White Ibis (Eudoci-

mus). Of course, detailed morphological study is required to address this supposition. CONSERVATION Both known recent nesting locations of the Oriental Crested Ibis are in preserves set aside for its protection. The conservation of this species is a matter of special concern, considering its precarious status. Individuals, conservation organizations, and the governments of Japan and China have been actively involved in the efforts. The participants include the Institute of Zoology in Beijing, the Toki Center in Japan, the W.W. Brehm Fund for International Bird Conservation at Vogelpark Walstrode, Germany, the Worldwide Fund for Nature, the International Crane Foundation, and the Storks, Ibises, and Spoonbills Specialists Group of the I.C.B.P. The Oriental Crested Ibis has for many years been a wellknown bird in Japan. The pale rose-coloured feathers were used by Samurai for their ornamental arrows and for feather brooms in the tea ceremony (Takashima 1957a). The decrease in this species was caused by hunting, loss of habitat by deforestation, pesticides, and modern rice farming practices that remove water after crop harvest. After their drastic reduction and rediscovery, they were made a Special National Monument in 1952. The Toki Center was founded in Japan to protect the Crested Ibis. A programme initiated to attract ibises to uncontaminated sites in Japan failed to influence the widely-ranging birds to avoid other feeding sites. In 1966, a captive breeding programme was established with 6 young ibises. One of these birds survived through 1984. Additional birds were brought into captivity in the early 1980s (G. Archibald 1981). More recently, eggs laid in Japan appear to be infertile, accounting for the lack of hatching there for over a decade. Because of poor success of the captives, attention was turned for a time to providing feeding sites in rice-paddies continuously stocked with fish (G. Archibald and S. Lantis 1979) . . . to no avail. In China, the nesting area is protected, and a research station has been established nearby. Observational studies were conducted and food provided to the birds. In 1989, the first Oriental Crested Ibis was hatched in captivity in the Bejing Zoo, and another was hatched in 1990 (Specialist Group on Storks, Ibises and Spoonbills 1989, 1990). In 1984, a Crested Ibis Workshop met in Beijing to discuss a management plan for the species. It proposed a three-point conservation plan for continued protection, expanded research and captive propagation. This is being undertaken at the Beijing Zoo in cooperation with the W.W. Brehm Fund and Vogelpark Walsrode, Germany. The government of China issued commemorative postage stamps in honour of the ibis, and funds generated from the sale of firstday covers were to be used for the conservation effort (W.W. Brehm-Fonds 1985). Despite these efforts, the conservation situation with respect to this species could hardly be more severe. Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 321.

Eurasian

Spoonbill Platalea leucorodia Linnaeus Subspecies: Platalea leucorodia leucorodia Linnaeus Platalea leucorodia Linnaeus, 1758, Syst. Nat., ed. 10, p. 139; based on 'The Spoonbill' of Albin, 1734, Nat. Hist. Birds, 2, p. 61, pi. 66: Europe: restricted to Sweden by Linnaeus, 1761, Fauna Svecica, ed. 2, p. 57 Platalea leucorodia balsaci Naurois and Roux Platalea leucorodia balsaci Naurois and Roux, 1974, Oiseau, 44, p. 77: Zira Island, Bane d'Arguin, Mauritania Platalea leucorodia archeri Neumann Platalea leucorodia archeri Neumann, 1928, Journ. Ornith., 76, p. 783: Dahlak Island, Red Sea Other names: White Spoonbill, Black-legged Spoonbill, Common Spoonbill, Indian Spoonbill, Japanese Spoonbill (English); Spatule blanche (French); Loffler (German); Lepelaar (Dutch); Espatula (Spanish); Spatola (Italian); Skedstork (Swedish); Kolpitsa, Kolpik, Lopaten ['shovel'] (Russian); Abu Malaqah ['father of the spoon'] (Arabic); Herasagi ['spatula heron'] (Japanese); Ta-pyi-lu, Ta-shaotsui (Chinese); Chamach baza, Chamcha (Hindi); Ghinta, Khunte bak (Bengali); Handi-alawa (Sinhala)

IDENTIFICATION The Eurasian Spoonbill is a medium-sized wading bird, standing about 60-85 cm tall, with a wing-span of c. 115-145 cm. It is white, rather stout and has a long neck and legs and a black spatulate bill. The adult's all-white plumage can become dingy with soiling. The black bill has irregular horn-grey and black barring on the upper side and an individually variable amount of yellow near the tip. The legs and feet are black. There is a narrow band of bare black skin at the base of the upper mandible and on the lores to the eye. The naked chin and throat are mainly pale sulphur-yellow. The iris is dark red or carmine. The plumages of the sexes are the same, but they are dimorphic in size. Males have slightly larger and heavier bodies, longer legs and longer bills, differences that are evident within wild flocks. During the breeding season, the plumage is immaculately white and the crest becomes particularly obvious, especially during displays. The crest consists of long narrow feathers (up to 12.0-15.Ocm in males, 10.5-13.5 cm in females). A crescent-shaped gorget or collar, with a variable amount of yellowish buff to cinnamon, develops on the lower foreneck; sometimes the crest is lightly tinged with the same colour. The bare skin of the lower throat becomes a richer yellow, often with a reddish band bordering the feathers (Cramp 1977:357). The two African subspecies differ from the nominate race as follows. The smaller balsaci has a completely black bill, or black with only traces of yellow; the yellow-buff on the chest is faint or absent in breeding season. The subspecies archeri is similar to balsaci^ but smaller still.

254

Eurasian Spoonbill

At hatching, the bill, legs and feet are orange, becoming pink-flesh after a few days. The bare skin around the eye is blue; the naked chin, throat and base of upper mandible are pink; the iris is grey. A nestling's first-down is sparse and white with silky tips; the second-down is short and dense, cream-white. Bills are not spatulate in young nestlings. Rather they gradually thicken and become bulbous at the tip and slightly decurved. Juveniles are similar to non-breeding adults, except that the bill is pinkish orange, becoming dark during the first winter. Full bill-length is not attained until 3^6 months. Legs are pinkish flesh coloured, later changing to slate. The iris is grey-brown. Some outer primaries and primary coverts have horn-black tips, and primaries, secondaries and some other feathers have dark shafts, although this is variable. Dark primary-tips and feather-shafts gradually decrease as birds mature, though some individuals, in otherwise full breeding plumage, retain them (Cramp 1977:357). This is a generally silent bird, except during social displays at the nest, when a deep grunting or groaning 'Huhhuh-huh' is given. These soft vocalizations can be heard over short distances only. During Sparring and other fights,

various snapping sounds are produced by the bill's rapid closure and by bills striking against each other. This is not true bill-clattering, as occurs in most storks. Small nestlings Beg with high-pitched 'Chittering' calls; in older nestlings these calls become a lower-pitched 'Tschibb-tschibbtschirr-tschirr' (Bauer and Glutz von Blotzheim 1966:430). Like all spoonbills, this species flies with neck and legs extended. A series of wing-beats alternates with a short glide. Flocks often fly evenly spaced in single file or in diagonal wavy ribbons. They sometimes soar to considerable heights on thermals. The flapping rate has been recorded as 186 ± 1.8/min (Palmer 1962:533). The Eurasian Spoonbill is distinguished from the white egrets by its novel bill, slightly heavier body and neck, and characteristic feeding behaviour. In flight it can be separated from egrets by the extended neck and faster wing-beats. In the southern parts of its range, it might be mistaken, at a distance, for a white ibis, but the straight, spatulate bill of the spoonbill is unmistakable. In Palearctic regions, it could be mistaken, in flight at long range, for a swan (Cygnus), but the spoonbill flaps and glides, and its long legs extend well beyond the tail. In northern Africa, it is distinguished from

Eurasian Spoonbill 255

the African Spoonbill by the latter's bare red face, red legs, red on the bill and pale blue iris. DISTRIBUTION AND POPULATION This spoonbill breeds from Europe to northeastern China, India, Sri Lanka, and the eastern and western coasts of northern Africa. It winters south to Mali, Sudan, Persian Gulf, India, Sri Lanka, southeastern China, and southern Japan. Platalea leucorodia leucorodia formerly bred rather widely throughout Europe. Until the 17th century it nested in southeastern England. It now breeds more locally in Europe: southern Spain, Portugal, northwestern Netherlands, Denmark (sporadically, north to 57°04'N; Kortegaard 1973), eastern Austria, Hungary, Greece, the Balkans and southern USSR. Recently, tentative breeding attempts have occurred again in France, northern Germany and Czechoslovakia. The range extends discontinuously across Asia: from Turkey and southwestern Siberia, eastward to western Mongolia, southeastern Siberia and northeastern China, and southward to India and Sri Lanka. It is a regular visitor to the UK, and casual or accidental in Scandinavia, Madeira, Canary Islands, Cape Verde Islands, Faeroes, Azores, and also Greenland (Cramp 1977:353). Platalea leucorodia balsaci is resident on islands off the coast of Mauritania, West Africa, primarily the Bane d'Arguin, at about 20°N. It is probably non-migratory. Platalea leucorodia archeri is resident on the coasts and islands of the Red Sea, the Gulf of Aden and Socotra. It is said to be a 'very common breeding resident' along the northwestern coast of Somalia (Ash and Miskell 1983). Several small colonies have recently been established northward to Nabq, southern Sinai and Tiran Island at the mouth of the Gulf of Aqaba (Paz 1987:36). This population is probably non-migratory. The wide-ranging leucorodia subspecies is migratory and also undergoes marked disperals. European populations winter mainly in Africa, regularly south to Mali and Sudan, rarely to Nigeria, Kenya and Uganda. A few remain around the Mediterranean bas,in, occasionally as far north as The Netherlands and England in mild winters. Eastern European populations migrate via Greece to the Nile Valley in Sudan and via Italy to Tunisia (Miiller 1984). Western Asian populations winter from Israel, Iraq, Persian Gulf, north coast of the Arabian Sea, and India southward to Sri Lanka. Eastern Asian populations winter mainly in southeastern China, rarely to southern Japan and Taiwan. It is mainly the temperate Palaearctic populations that are migratory, whereas the more southerly populations are primarily resident. The migration routes and timing are for the most part well known. Migratory movements are usually diurnal, in small parties or large flocks. In the late summer there is a post-breeding dispersal, mainly of juveniles, which generally occurs over short distances, although sometimes it is further, e.g. Yugoslavia to Scotland, Hungary to the Camargue, France, and The Netherlands to the Azores. Birds depart European colonies on regular migration in

August-September and return in March-April. Colourringed birds from The Netherlands are known to follow the Atlantic coast of western Europe and have been found, in winter, along the west coast of Africa in Senegal and Mauritania. It is believed that many Dutch and Spanish birds winter among the larger population of P.L balsaci in and just south of the Bane d'Arguin, Mauritania (Roux 1962, 1973, Poorter 1982); this has been confirmed by ringing recoveries (Trotignon and Trotignon 1981). Ringed birds from eastern Europe have been recovered in winter in southeastern Europe, Italy, Turkey, Tunisia, Libya, Egypt, Sudan and Niger. The virtual absence of recoveries from the northwestern Mediterranean, eastern Morocco and coastal Algeria suggest there is little contact between eastern and western European birds on their wintering grounds. Birds ringed at the large colony at Lake Manyas, Turkey, have been found in Israel, Egypt, Sudan, Iraq and Pakistan (Schiiz 1957b). They more often utilize marine habitats—such as deltas, estuaries, lagoons and shallow inlets —as their winter quarters. During the last century, vast flocks of migrant Eurasian Spoonbills would arrive in India from the north during October. More recently, 5 spoonbills, ringed as nestlings in the Caspian and Black Sea regions of the USSR, were later recovered in India (Sapetin 1968, McClure 1974:94). Wetten (1986) estimated the world population of Eurasian Spoonbills to be 31 000-34500 individuals. The species is abundant nowhere in Europe; in most countries where the bird breeds there are only a few hundred pairs in the best of years. Population trends in Europe have been generally downward throughout this century, with only some very recent cause for encouragement (G.A. Brouwer 1964, Bauer and Glutz von Blotzheim 1966:432-435, Cramp 1977:353354). The history in The Netherlands is illustrative. From 300 pairs around 1900, the population rose with protection to 450 pairs in 1925, and to about 500 pairs in the 1950s. It decreased to below 200 pairs in 1964. In 1989, owing to increased environmental protection, the population had rebounded to 423 pairs nesting in 8 colonies (K. Brouwer pers. comm. 1990). The first nesting of the Eurasian Spoonbill in France occurred in 1981 (Marion and Marion 1982). In Spain during 1990, there were 850-900 nesting pairs in 2 colonies (E. Aguilera pers. comm. 1990). Tbe breeding population in Austria and Hungary has remained almost constant since 1950 (Miiller 1984). In contrast, it may have disappeared completely as a breeding bird from Romania (Wetten 1986). The West African population (P.L balsaci) in the Bane d'Arguin, Mauritania, may have the densest concentration known for the species, with an estimated 1430 breeding pairs during the years 1959-65 (de Naurois 1969:106) and a total population of 5000-6000 birds in 1974 (Poorter 1982). Between 8600 and 10000 spoonbills were counted in the Bane d'Arguin area during December 1979 and January 1980 (Trotignon and Trotignon 1981); an unknown portion of these were winter visitors from Europe. The Red Sea population (P.L archeri) has the smallest numbers, with an estimated 200-500 birds in eastern coastal Sudan (Luthin 1984a).

256 Eurasian Spoonbill

During censuses in mid-winter 1989, 5559 Eurasian Spoonbills were counted in India and Sri Lanka, the greatest numbers being 1751 birds in Gujarat and 1065 in Rajasthan (D.A. Scott and P.M. Rose 1989). It is not known how many of these were migrants from further north and how many were birds resident in the Indian region. ECOLOGY The habitat of the Eurasian Spoonbill is rather broad, including many types of shallow-water areas with mud, clay or fine-sand bottoms. Birds usually avoid waters with a rocky bottom, thick vegetation or swift currents. They are usually found in coastal lowlands or alluvial river valleys, in fresh, brackish or salt water. They feed in estuarine areas, especially on the wintering grounds. However, European birds are relying increasingly on estuaries for feeding during nesting (J. van Wetten in Luthin 1984a). The birds feed both singly and gregariously in aggregations of up to 100 birds, generally in water less than 30 cm deep. Their foraging behaviour is characteristic of spoonbills. Food detection is primarily non-visual. A bird wades vigorously forward, Walking and Sweeping its partially open bill from side to side in the water with a semi-circular scything motion. The bill is gaped about 5 cm, and the tip often touches the substrate. When active prey such as fish are being hunted, birds may hold their bills to one side and run rapidly, Dragging them through the water (Poorter 1969). When less active prey is sought, birds usually walk more slowly and search with wider sweeps of the bill (Wetten and Wintermans 1986:13). They also use an Intensive Sweeping, called terugzoeken (trying to relocate) by Poorter (1969), when they remain searching in one place. Spoonbills exhibit a Probing behaviour (wroeten—rooting; Poorter 1969), using a stabbing motion. S. Ali and S.D. Ripley (1968:117) state that 'where food is plentiful a compact, eager, jostling herd will advance almost at a run, working methodically back and forth over a particularly rewarding patch5. Foraging is most active during mornings and evenings, and like most spoonbills, this species often also feeds at night (Aguilera 1990b). In coastal areas feeding is primarily at low tide, regardless of the time of day (van Oordt in Bannerman 1957:30). The diet consists of small fish (up to 10-15 cm), aquatic insects and their larvae, crustaceans, frogs and tadpoles, molluscs, worms, leeches, reptiles, and some plant material (possibly ingested accidentally). At freshwater feeding areas in The Netherlands, spoonbills captured 1.5-2.0 prey items (mostly small fish) per minute, i.e. 540-960 prey items during a 6-8 h feeding day (Wetten and Wintermans 1986:44). The amount of food eaten has been recorded: 2 captive spoonbills together ate about 1 kg of freshwater fish per day (O. Koenig 1952); an adult weighing 1.9kg ate 300 g/day, and a juvenile weighing 0.9 kg ate 140 g/day (I. Sterbetz 1984). Although the usual prey is small, there is a report of a bird in The Netherlands that choked while attempting to swallow a large eel (R.P. Allen 1942:101). During the breeding season, feeding areas are usually within 35-40 km of the nesting colony (Wetten and Winter-

mans 1986:47). They often walk ashore after feeding and rest on one leg. Spoonbills commonly sleep with their bill and face rotated to the rear and tucked beneath the feathers of the back. They roost either on the ground (if secure) or in trees. They can swim well, and do so when necessary. BREEDING The Eurasian Spoonbill usually nests in colonies of a few to a few hundred pairs. They nest in dense stands of emergent aquatic vegetation, especially reeds and trees, such as willows (Salix sp.), oaks (Quercus sp.) and poplars (Populus sp.) (Beetham 1910, Marion and Marion 1982, Miiller 1983, Aguilera and Alvarez 1989). Breeding colonies often face pools or channels. Almost invariably the nest site is within 10-15 km (often much less) of feeding areas. Nests are usually at low elevations, but an exceptional colony was at nearly 2000 meters on Lake Sevan, Armenia (Dementiev and Gladkov 1951:417, Cramp 1977:352). Sometimes Eurasian Spoonbills nest in or near mixed colonies of herons, ibises and other waterbirds, but they usually remain somewhat segregated from other species. A colony site is often used in successive years. In most areas birds are intolerant of disturbance, especially early in the breeding cycle; however, they often nest in trees in the midst of villages in India (E.C.S. Baker 1935:433). Nests are usually placed 1-2 m apart, but sometimes so closely together that they touch. They are built in trees (up to 2-5 m, sometimes higher), reedbeds or on the ground, as is the case in The Netherlands and Mauritania. The nest itself is a large pile of reeds, sticks or twigs, lined with grass and leaves, 50-100 cm in diameter and 30-60 cm deep. Pair-formation displays, described by Poorter (in Cramp 1977:355-356), appear to be similar to many of those seen in the African and Royal Spoonbills (Kahl 1983, 1988a). During the Greeting display, the brightly coloured throatskin is exposed to the mate when the head is raised. The luxuriant crest is raised and fanned laterally into a 'warbonnet5 during many behaviours. Sparring, Nest-Covering displays and Begging displays also occur. Mutual allopreening by mates is common, especially on the head and neck, during quiet moments on the nest. Copulations are frequent in the early breeding season: as many as 11 attempts in 1 h have been observed (Dragesco 1961b). The average interval between the first observed copulation and the day on which the first egg is laid is 6.7 days (n = 9) (Aguilera and Alvarez 1989). Attempted extrapair copulations by paired males with neighbouring females, while their mates are absent, are common (Aguilera 1989). During copulation, we observed that the male rapidly bites and slaps the bill of the female, making a soft clattering or slapping sound (pers. obs. M.P.K.). In 18 copulations timed by Aguilera and Alvarez (1989) the duration was 9.7 ± 2.3 s. The same authors mentioned, but did not describe, a pre-copulatory display by the male, termed 'Rubbing back5. Each pair defends its own nest and the immediate area nearby, in which only close neighbours are tolerated. Birds from adjacent nests often cooperate to drive away strange

Eurasian Spoonbill

adults. Both sexes assist in the construction of the nest, with males collecting most of the nest material at Spanish nests (Aguilera and Alvarez 1989). Eggs are slightly elongated ovals, chalky white with small reddish brown spots and lines. It was found in Romania that larger clutches have smaller eggs (Vespremeanu 1968a). The clutch size is usually 3-4, rarely 6 and exceptionally 7 (de Naurois 1959). In Macedonia, 54 clutches were 3 eggs (56% of clutches), 4 (37%), 5 (6%), and 6 (1%). In Romania, 415 clutches were 2 eggs (23%), 3 (54%), and 4 (23%), with a mean of 3.0 (Vespremeanu 1968a). In southwestern Spain a fifth egg was laid in only 3% of 937 clutches (Aguilera 1990a). Eggs are laid at 2-3 day intervals (Cramp 1977:356). The incubation period is given by Holstein (1929) as 21 days, but it is probably nearer to 24—25 days (Dementiev and Gladkov 1951:419, Vespremeanu 1968a). Incubation is by both sexes and usually begins with the laying of the second egg; hatching is asynchronous (Aguilera 1990a). Pairs within a colony may be relatively unsynchronized, as all stages of nesting may occur simultaneously (Vespremeanu 1968a). At a colony in southwestern Spain, males generally attended the nests in the daytime and foraged at night, whereas females tended the nests mostly at night (Aguilera 1990b, Aguilera and Alvarez 1989). Eggs and small nestlings are often shaded by their parents on hot days. At feeding a nestling inserts its bill into the buccal cavity and upper throat of the adult to receive food that is regurgitated for it. Unfledged young wander about the colony from about 3 weeks of age, often visiting other occupied nests; they return to their own nest for feeding up until about 6-8 weeks. The fledging period is 45-54 days (Vespremeanu 1968a, Cramp 1977:356). Both parents tend the young until well after fledging, females were observed to feed the chicks more than males at Spanish nests (Aguilera and Alvarez, 1989). The bond between parents and young may continue until about 10 weeks, with the parents gradually providing fewer meals and the young foraging more on their own. Most nestling mortality occurs in the first 10-15 days. Breeding success was recorded by Vespremeanu (1968a) in Romania in 1958-63: of 1244 eggs laid, 95% hatched, and of 1188 young, 70.3% fledged. Nestling mortality varied from 11.3 to 100.0%, depending on the season. At the Goto Donana National Park, Spain, 4 is the maximum known number of young fledged from any spoonbill nest (Aguilera 1990a). In broods of 5, the death of the youngest chick is sometimes attributable to attacks by their own parents, rather than by siblings (Aguilera 1990a). High water levels during the nesting season can lead to very low success (Vespremeanu 1968a). At some European colonies, food availability may be a limiting factor, because adults have to fly longer distances (often as much as 30 km) to feed as the season progresses (J. van Wetten in Luthin 1984a). Avian predators, such as Jackdaws (Corvus monedula), Magpies (Pica pica), and Black Kites (Milvus migrant), prey on eggs and young in Spain (Aguilera and Alvarez 1989). At other sites mammalian predators account for much of the egg and nestling mortality, except on offshore islands that are out of reach of terrestrial predators.

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TAXONOMY Some authors have considered the Royal Spoonbill, Platalea regia to be a subspecies of the Eurasian Spoonbill (i.e. Platalea leucorodia regia). However, based on field studies of the various populations on their breeding grounds (Kahl 1988a and pers. obs.), we conclude that the two are distinct enough to warrant specific status. There have been suggestions that the eastern and western populations could be considered subspecifically distinct. We agree with earlier statements (Seebohm 1890:230, Meinertzhagen 1954:394, Vaurie 1965:78) that the size differences used to separate the eastern and western populations in Eurasia are not sufficient to warrant the recognition of the far-eastern race of P.I. major. We do, however, recognize three other subspecies, based on differences in size and breeding coloration: P. 1. leucorodia, P.I. balsaci and P.I. archeri. It seems that P. leucorodia, P. regia and P. minor are sufficiently closely related to comprise a superspecies. But P. alba is more distinctly different from the other three and not so closely similar as indicated by some authors (e.g. Snow 1978:42).

CONSERVATION The breeding range of the Eurasian Spoonbill, especially in western Europe, has been drastically reduced owing to drainage, alteration or pollution of feeding areas and, mainly in earlier years, to human exploitation of eggs and nestlings for food (G.A. Brouwer 1964). An important consideration over the past century has been the disappearance of reed swamps, the favourite nesting habitat, due to agriculture and hydropower development (Vespremeanu 1968a). The decrease of the Dutch population from about 500 pairs in the 1950s to 150-200 pairs in the early 1970s, probably owes to pollution of feeding areas by chlorinated hydrocarbons (Rooth and Jonkers 1972). Dutch birds recovered to about 400 pairs in the late 1980s, and by 1989 there were 423 pairs in 8 Dutch colonies (Osieck and deVries 1987; K. Brouwer pers. comm. 1989). Populations in Greece continue to decrease owing to degradation of habitat, agricultural practices, overfishing and disturbance (Crivelli et al. 1988). The species is now seriously endangered in most areas of its European range (Cramp 1977:353). During their migrations, the western European populations are dependent on a succession of estuaries along the coasts of Belgium, France, Portugal, Spain and Morocco. Eastern European birds depend on sites in Italy and Greece. It is important that these migratory and staging areas are protected, as well as feeding and nesting habitats in the summer and winter ranges.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on pages 322-323.

/v

Royal Spoonbill Platalea regia Gould Platalea regia Gould, 1837, Proc. Zool. Soc., pt. v, p. 106: east coast of New South Wales Other names: Blackbilled Spoonbill, Blackfaced Spoonbill (English)

IDENTIFICATION The Royal Spoonbill is a medium-sized, rather stout, white waterbird, standing 74-85 cm tall, with a wing-span of about 120 cm. It has a long neck and legs and a black, flattened bill, spatulate at the tip. Although the plumage of the adult is entirely white, it is sometimes soiled. The bill is mostly black, with irregular bluish grey and black barring on the upper side; it lacks the yellow tip seen in Eurasian Spoonbills. The skin of the face and unfeathered throat is black. The legs and feet are shiny black. The iris is a deep, dark crimson. The species is sexually dimorphic. Males have slightly larger bodies, longer legs, and longer bills (Vestjens 1975b, Kahl 1988a). During breeding the plumage is immaculately white, and the nuchal crest becomes prominent. It is more luxuriant than in any other species of spoonbill, composed of up to 100 long, narrow plumes (individual feathers measured up to 19.4 cm long). The crest is especially conspicuous when erected and fanned laterally during displays. Some adults begin losing their crest feathers even before the young have left the nest. Also during breeding, a horizontal collar of pale yellowish buff appears on the lower foreneck; this is variable and less conspicuous than in the Eurasian Spoonbill. Some individuals have a faint hint of the same buff colour on their crest. A naked, narrow band of bare skin under the wings is salmon-pink to saffron coloured (Kahl 1988a). Oval spots, about 20 X 10 mm, of deep yellow-ochre develop on the skin above (and sometimes, to a lesser extent, below) the eyes, contrasting strongly with the black facial skin. Also a triangular patch of dull red skin appears at the top of the black forehead, just below the white feathers of the crown. These yellow and red facial markings are variable between individuals. According to K.W. Lowe (pers. comm. 1990) some Royal Spoonbills can be seen at all months of the year with these breeding colours. (In preserved specimens, the yellow eye-spots and red forehead-triangles of living Royal Spoonbills both appear as dull yellowish tan areas. This has resulted in misleading descriptions by those unfamiliar with the living birds.) The bill, legs and feet are pinkish orange at hatching; the bare skin around the eye is bluish. The firstdown is sparse and white; the second-down is more dense and off-white. Bills are not spatulate in nestlings but gradually thicken and become bulbous at the tip, with a slight decurvation (Kahl 1987c). Juveniles are similar to non-breeding adults, except that the bill is shorter and dusky pinkish, becoming longer and darker during the first year. The facial skin is dusky and lacks any coloured areas, becoming blackish during the first year. The legs change gradually to slate-grey. In certain individuals, some outer primaries and primary coverts have horn-black tips, and primaries, secondaries and some other feathers have dark shafts. Dark primary-tips and feather-shafts gradually fade or disappear as the birds mature. However, some apparent adults retain dusky primary-tips. The Royal Spoonbill is generally silent, except during social displays at the nest, when a low growling, grunting, or honking *Huh-huh-huh-hutf is given (Kahl 1988a). These vocalizations are soft and audible only over short distances. During fights, some rattling sounds are produced by snapping of the bill. True clattering, as made by storks, is not heard, contrary to some reports in the literature. Small

260 Royal Spoonbill

nestlings perform the Begging display with high-pitched 'Chittering* calls; in older nestlings these calls become lower-pitched (Kahl 1988a).Vestjens (1975a) found that the shape of the trachea changes with age, and this may be related to changes in voice. Flight is with the neck and legs extended, a series of wingbeats alternating with a short glide. Flocks often fly in single file or in diagonal wavy ribbons, with each bird evenly spaced behind and to one side of the one in front. The flapping rate averages 230/min (n = 44) (pers. obs. M.P.K.). The Royal Spoonbill is distinguished from the white egrets by a longer, heavier and spatulate bill, slightly heavier body and thicker neck, characteristic feeding behaviour, and, in flight, by an extended neck and faster wingbeats. It might be mistaken at a distance for an Australian White Ibis, but the straight, spatulate bill is diagnostic. It is distinguished from the slightly larger Yeilowbilled Spoonbill by the black face, bill and legs. The Royal Spoonbill usually has a whiter, 'cleaner' plumage than either the Australian White Ibis or the Yeilowbilled Spoonbill. In the extreme northwestern part of the Royal Spoonbill's range (e.g. Borneo), it may rarely overlap with wintering Blackfaced Spoonbills. In non-breeding plumage, it might not be possible to distinguish this bird from the Blackfaced Spoonbill, as discussed in that species' account. DISTRIBUTION AND POPULATION The Royal Spoonbill is primarily Australian, found in the well-watered parts of northern, eastern and southeastern Australia. In Indonesia, it is possibly resident in the Mollucas, Lesser Sunda Islands, Sulawesi and Irian Jaya; it has occurred in Java, where it formerly bred (Hoogerwerf 1952, Silvius and Verheugt 1990). In southern Papua New Guinea, it is commonest in the dry season and probably only a non-breeding visitor from Australia; a small, isolated population exists on Rennell Island, Solomon Islands (Coates 1985:90, Beehler et al. 1986:61). Prior to 1950, it was a vagrant in New Zealand, but more recently it has bred in small numbers at Okarito and Otago, South Island (Holdaway 1980, H.A. Robertson and B.E. Preece 1980). It

wanders to Tasmania (Blakers et al. 1984:66) and Borneo (Smythies 1981:33). This species is sedentary or nomadic, moving mainly in response to periodic droughts. However, it is reported to migrate across the Torres Straits, between Australia and New Guinea, in large flocks (Coates 1985:90). Five recoveries, 1-36 months after ringing, show movements of between 277 and 1472km within Australia (Blakers et al. 1984:66, K.W. Lowe, Australian Bird and Bat Banding Schemes pers. comm.). Few estimates of population numbers and trends are available. However, this is a common to locally abundant species in the well-watered parts of northern and southeastern Australia. In 1988, a dry-season survey by the Asian Wetland Bureau found over 250 birds in southeastern Irian Jaya (Silvius and Verheugt 1990). Except for these New Guinea birds, possibly, populations outside Australia are not large. ECOLOGY The Royal Spoonbill inhabits large shallow waters, inland and coastal marshes, lake shores, mud flats and mangroves. It does frequent farm dams, but is less often found on such small waters than is the Yeilowbilled Spoonbill. In tidal areas most feeding occurs at low tide. Royal Spoonbills are often found in small flocks. They habitually feed in water less than 40 cm deep. They rest on the shore or in trees, with bills rotated about 150° and tucked into the feathers of the upper back. Feeding occurs both day and night. As do other spoonbills, the Royal Spoonbill feeds by touch. The partly open spatulate bill is swept from side to side in the water. Although the birds feed mostly by Slow Sweeping, they also do a rapid, short-stroked Intensive Search, after detecting a potential prey item. Vestjens (I975b), who described the feeding behaviour in detail, found that this species also used Dragging, Probing and Grabbing foraging methods. All these techniques involve tactolocation of food, except for Grabbing, in which prey is visually located. It is apparently the food items that touch the widest portion of the 'spoon' that are detected and grasped. Food is then transferred from the distal portion of the bill to the throat with a quick, backward toss of the head and then swallowed. They are specialized and rather efficient foragers. During 'Slow Sweeping' foraging birds captured an average of 6.4 items per minute in the freshwater Lake Cowal, NSW (Vestjens 1975b), and 3.5/min in the marine tidal area of Westernport Bay (K.W. Lowe 1982). Spoonbill predation of intertidal shrimps has been shown by R.K. Howard and K.W. Lowe (1984) to be sufficient to influence their population levels. The results of this study demonstrate how a non-visual predator can be selective in the prey it catches. Spoonbills were particularly selective for adult female shrimps rather than small shrimps. The diet includes small fish, aquatic insects, crustaceans, small freshwater snails, tadpoles, and, occasionally, spiders and crickets (Vestjens 1975b). Prey caught at night is different to that caught during the day (K.W. Lowe 1982).

Royal Spoonbill

When feeding in the same habitat, the Royal Spoonbill takes more fish and fewer crustaceans and insects than does the Yellowbilled Spoonbill; this is due to differences between the two species in bill-structure and speed of movement. In the stomachs of 20 Royal Spoonbills at Lake Cowal, NSW, Australia, the ratio of crustaceans and insects to fish was 4:3 (Vestjens, 1975b). At Westernport Bay, Victoria, Australia, the stomach contents of 10 birds was 73% crustaceans (wet-weight), especially the striped prawn (Macrobrachium intermedium), and 23% fish, especially (K.W. Lowe 1982) the Bridled Goby (Arenigobius bifrenatus). BREEDING The Royal Spoonbill nests singly or in small colonies (up to 100 pairs), often with ibises, darters or the smaller cormorants. The nest-site is usually within 5 km of feeding areas, sometimes as near as 500 m (Vestjens 1975b). The nest is a shallow platform of sticks in trees or bushes; it is usually over water, occasionally up to 20m high, although usually considerably lower. At Lake Cowal, nests were placed in medium-sized trees of river cooba (Acacia stenophylla), river red gum (Eucalyptus camaldulensis) or emergent lignum bushes (Muehlenbeckia cunninghamii). Nests are sometimes constructed of reeds in reed-beds (Beruldsen 1980:158). In the small colonies at the periphery of the range in New Zealand, nests were built in the tops of very tall trees (at Okarito) or on the ground (at Blenheim) (Holdaway 1980). The timing of nesting is somewhat variable: AugustJanuary in southern Australia and February-May in northern Australia. On South Island, New Zealand, nesting is in November-December, and in western Java was formerly March-September. Unmated birds of both sexes congregate in 'bachelor parties', where many forms of ritualized displays are seen. Males then establish themselves on potential nest-sites, and females attempt to approach them. However, the females are repeatedly driven away, with Supplanting or overt attacks. Eventually a female is accepted by a male, and the pair proceeds to build a nest. Mutual allopreening is common between mated birds, especially of the feathers of the head and upper neck. Whenever one member of the pair returns to the nest after an absence, both birds perform the Greeting to each other. Other commonly performed ritualized displays during courtship are Bowing displays (shown by females), Sparring, Head Shaking, Stick Shaking, Display Sleeping, Display Shake, and Display Preen (shown by both sexes). Copulations and egg-laying follow closely after pair-formation (Kahl 1988a). Both adults build the nest, with the male gathering most of the material. The clutch size varies from 2-5 eggs, usually 3-A. The eggs are tapered oval in shape; they are dull white, spotted with yellow-brown or red-brown. Incubation is performed by both sexes for about 25 days; hatching is asynchronous at 1- to 3-day intervals. Parents may desert the nest if disturbed during the early stages of the breeding cycle, especially during nest-construction before the eggs are laid (pers. obs. M.P.K.).

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Both parents brood, guard and feed the nestlings. At feeding a nestling inserts its bill into the throat of its parent, who then regurgitates food from the stomach for the young to eat. Only one young is able to feed from the parent's throat at a time. During the 1984-85 season at Lake Cowal, a total of 110 nestling feedings averaged 6.9s in duration (range = 4.7-12.5s); a brood was given between 8 and 28 individual portions during a session, and most of the feedings took place within 10-20 min of the parent returning from a foraging trip (pers. obs. M.P.K.). Young are capable of their first flights at 4-5 weeks, but they continue to return to the nest for feedings by the parents until about 2 months. Family groups remain together for several weeks more. Young have been observed performing the Begging display on the wing, while flying after an adult (Hoogerwerf 1952). Egg and nestling mortality are most commonly caused by inclement weather or predation by Australian Ravens (Corvus coronoides), lace monitors (Varanus varius) and, possibly, various snakes (Vestjens 1977a). TAXONOMY According to some earlier classifications (e.g. J. Steinbacher 1979), the Royal Spoonbill was considered a subspecies of the Eurasian Spoonbill, Platalea leucorodia regia. However, owing to significant morphological differences, especially in soft-part coloration, we prefer to treat it as a full species, Platalea regia (Amadon and Woolfenden 1952, Gyldenstolpe 1955a,b). If the Royal Spoonbill were conspecific with any other population, it would be with the Blackfaced Spoonbill (P. minor) of northeastern Asia, with which it shares many morphological traits. As can be seen in the tables of body measurements (see Appendix), there is considerable overlap in size between the two populations, and the soft-part coloration is strikingly similar (see Ogilvie-Grant, 1889:48, col. pi. 1). The extent and shape of the bare skin on the throat, which has been suggested as a means to distinguish between the Royal and Blackfaced Spoonbills, is quite variable, individually and with age. It is, therefore, unreliable as a taxonomic character. Pending further study, we chose to retain Platalea minor as a full species; however, it is certainly very close, taxonomically, to Platalea regia. In our opinion, the three spoonbills, Platalea regia, P. minor and P. leucorodia, are closely enough related to comprise a single superspecies. CONSERVATION There appear to be only a few Royal Spoonbill breeding colonies of significant size. However, in general, the species seems to be doing well in Australia, the stronghold of its range, and expanding slowly in outlying areas such as New Zealand. Status in the northern part of the range is less well known. The species has apparently disappeared from Java, where it nested in small numbers in the 1950s (Hoogerwerf 1952). Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 324.

Blackfaced

Spoonbill Platalea minor Temminck and Schlegel Platalea minor Temminck and Schlegel, 1849, in Siebold's Fauna Japonica, Aves, p. 120, pi. 76: Japan Other names: Lesser Spoonbill; Swinhoe's Blackfaced Spoonbill (English); Petite Spatule du Japon (French); Kurotsura-herasagi (Japanese); Hsiao-pyi-lu, Hsiao-shao-tsui (Chinese)

IDENTIFICATION The Blackfaced Spoonbill is a medium-sized white waterbird, c. 60-83 cm tall. As the common name suggests, it has a black face, and the long bill is flattened and ends in a broad spoon. The adult plumage is entirely white. The bare black skin on the face extends to the throat and 5-15 mm behind the eye. The bare area in back of the eye and on the throat is quite variable, and it is difficult to judge in the field (pers. obs. M.P.K.). The bill is slate-coloured and, as described by Sharpe (1898:51), 'transversely barred with black, the bars broken and disconnected on the spatule'. The legs are black. The iris is crimson to blood red. Sharpe (1898:51) also reported 'a patch and scattered spots of orange-ochre over the spatule of the bill. Peter Kennerley (pers. comm. 1990) stated that this coloration on the bill was 'never seen on adults in Hong Kong, even in nuptial plumage'. We wonder if Sharpe may have confused this species with the Eurasian Spoonbill. If the Blackfaced Spoonbill does, indeed, show orange or yellow near the end of the bill, it apparently does not occur in many individuals, nor to as great an extent as in the Eurasian. C. Poole (pers. comm. 1990) tells us that birds 'can show pale areas on the spatule, which I would best describe as fleshed coloured'; this description may refer to subadult birds. The matter of bill coloration is one that requires additional observation. Females are similar to males but slightly smaller. The plumage and soft-part colours change during breeding, but the details remain somewhat uncertain. A yellowish half-moon marking develops on the face, but its size, shape and position can vary. H.G. Won (1966) described the marking as being on the lores. Referring to the facial spot, others place the 'bright yellow-ochre patch before the eye, extending [across] the under-lid, and a thin line [across] the upper lid' (Sharpe 1898:51) or 'a yellow spot in front of and under the eye' (La Touche 1931-34:432). The yellow eye-spots of living spoonbills appear as dull yellowish tan in preserved specimens. Breeding adults develop a long nuchal crest. According to Peter Kennerley (pers. comm. 1989) the crest plumes of birds in Hong Kong take on a distinctive golden-yellow colour in the spring, just before they begin migration. They also show a bright yellow bar across the lower foreneck or upper breast (not on the throat, as described by H.G. Won 1966). In some cases, this develops into a complete golden collar around the lower neck. In full breeding plumage the yellow collar of the Blackfaced Spoonbill is a richer colour than the similar collar of the Eurasian Spoonbill (P. Kennerley pers. comm. 1990). Colour photos in Chung (1986) show that by late in the breeding season, after the young have hatched, the yellow marking on the lower foreneck has become faint, and the crest shows only a wash of pale buff. The hatchling is little known, except that 'chicks bills are shorter than adults' and are round' (Chung

264 Blackfaced Spoonbill

DISTRIBUTION AND POPULATION

1986) [from a translation by Kyoko Archibald]. The juvenile is white; it 'differs from the adults in having a more yellowish bill and blackish shaft-stripes to the quills, the outer primaries blackish at the ends of the outer webs' (Sharpe 1898:51). These dark primary tips are obvious in the field, making it easy to separate juveniles from adults in flying groups. Vocalizations of the Blackfaced Spoonbill have not been described. Birds we observed briefly in Hong Kong appeared to have a flight typical of other spoonbills. The flapping rate which was timed on a single occasion was 231/min (pers. obs. M.P.K.). This species is similar to the Eurasian Spoonbill but noticeably smaller, with more extensive bare black skin on the face and throat, and usually, perhaps always, lacking the prominent yellow mark on the 'spoon' of the bill. In its summer plumage this species shows more yellow or buff on the crest than does the Eurasian Spoonbill (Gore and Won 1971:123). It is possible that the Blackfaced and Royal Spoonbills may occur together in Borneo (Mann 1989). For the reasons discussed below (see Taxonomy), separation in the field would be difficult. In breeding plumage (unlikely in Borneo) the Blackfaced Spoonbill would have more yellow below the eyes, whereas the Royal Spoonbill would have more yellow above the eyes. The Blackfaced Spoonbill may also lack the dull red triangle at the top of the bill.

The breeding distribution of the Blackfaced Spoonbill is poorly known. It is confirmed to nest only in Korea, on a few small islands (Sogam-do, Tegam-do, Tok-to; formerly also on Ai, Kamsang) along the western coast of North Korea (Chung 1986, Sonobe and Izawa 1987:36-37). Prior to the Korean War (1950) this species was said to be common and was found breeding on islands (e.g. Wido, Ito) along the western coast of South Korea (Austin 1948:49, P. Won 1973:26). There are unconfirmed reports of breeding in central Manchuria in the vicinity of Harbin (Vaurie 1965:78), northwestern Jilin Province, Inner Mongolia, and, possibly (formerly at least), in eastern China south to Fukien. As far as is known, most of the population winters in southern China (Kwangtung Province, Hainan Island), Hong Kong (Mai Po marshes), northern Vietnam (Hong River delta) and southwestern Taiwan (Tsen-wen Hsi, Chiayi Hsien). It has been recorded as far north as Korea in December (H.G. Won 1966, Gore and Won 1971:123). [The Eurasian Spoonbill was said 'to be numerous in the mangroves' of coastal Vietnam in winter (Delacour and Jabouille 1925), but this may have referred to this species instead.] A few winter stragglers are found in Thailand (P. Round pers. comm. 1989), Japan (Takara 1979, Sonobe and Izawa 1987:36), and the Philippines (duPont 1971:23). A record from Brunei was assigned to this species, but it was apparently a subadult bird and could conceivably have been a Royal Spoonbill from the south (Mann 1989, pers. comm. 1990). There may be other, as yet undiscovered, small flocks wintering along the south China coast; probably few winter inland in China (D.A. Scott pers. comm. 1990). Wintering birds are present in Hong Kong mainly between late October and early May (Chalmers 1986:45); they are found in Taiwan from September through May (Lucia Liu Severinghaus pers. comm. 1990). According to H.G. Won (1966) summering birds arrive in North Korea in March (although this must be exceptionally early) and leave again in October/November. Migrating birds must pass through China, but little is known about their movements between the breeding areas in North Korea and the wintering grounds. Some do cross South Korea, where up to 46 were recorded on South Kanghwa Island on 5 September 1989; 20 were still present into the second week of October 1989. About two-thirds of these birds were immatures (C. Poole pers. comm. 1990). An undetermined number of Blackfaced Spoonbills were identified in flocks of Eurasian Spoonbills during December 1985 and January 1986 at Poyang Lake, Jiangxi Province, China (Kennerley 1987), and single birds were seen there also in December 1989 (P. Kennerley pers. comm. 1990). The known population of the Blackfaced Spoonbill is small, and it is currently endangered. Information on the breeding population is scarce. In June 1965, a total of 36 adults were counted at a breeding colony off the coast of North Korea (Choi 1966); in the mid-1980s about 30 were

Blackfaced Spoonbill 265

counted in that area (Chung 1986, Sonobe and Izawa 1987:37). More information is available from the wintering grounds. Prior to 1982 the highest winter counts in Hong Kong were about 20 birds; however, since then, there has been a slow but steady increase (Chalmers, 1986:45). In most recent years about 40 birds have wintered in Hong Kong. A high of 47 was recorded on 28 January 1989 (P. Kennerley pers. comm. 1990). Of the 40 Blackfaced Spoonbills we observed in Hong Kong on 31 January 1989, approximately 30-40% were immature birds (pers. obs. M.P.K.). Between September 1988 and March 1989 a group of 130 Blackfaced Spoonbills wintered in southwestern Taiwan (D.A. Scott and P.M. Rose 1989:80, Lucia Liu Severinghaus pers. comm. 1990). Approximately 145 wintered there in 1989-90 (Lucia Liu Severinghaus pers. comm. 1990), and about 100 were counted in the area on 12 March 1990 (C. Poole pers. comm. 1990). In the winter of 1987-88 a total of 62 were counted, by Dr Le Dien Due, in the Hong River delta area of Vietnam. A high count of 27 birds was recorded there on 27-28 March 1990. Over 50% of birds seen in Vietnam during 1988 and 1989 were immatures (D.A. Scott pers. comm. 1990). A few Blackfaced Spoonbills have recently been reported during the winter in South Korea (Coulter and Brouwer 1991). 3-6 birds have wintered at Songsanp'o on Cheju Island, and 4 were reported in 1987 in the Nakdong Estuary. If the Hong Kong, Taiwan and Vietnam populations represent most of the wintering Blackfaced Spoonbills, the total world population is, perhaps, about 285 birds (Kennerley 1990). However, only c. 30 adults are known to summer in North Korea. So it remains to be discovered where the others go. Perhaps there are as yet undiscovered breeding areas in the eastern part of China. The most likely area would seem to be the offshore islands in Korea Bay, to the east of Dalian, China, and adjacent to the North Korean border. ECOLOGY The ecology of the Blackfaced Spoonbill is little known. Like other spoonbills, it frequents shallow waters (ptchecopar and Hue 1978:80). The known population occurs largely in coastal areas, inhabiting tidal flats, saltmarshes, estuaries and tidal creeks; it sometimes uses inland lakes. It appears to favour mangrove areas during the winter. The feeding behaviour is described as similar to other spoonbills. The bird places the partly open bill in water, Swinging it from side to side (Etchecopar and Hue 1978:80, Chung 1986). According to H.G. Won (1966), 'In the seasons when they were not breeding, they search [for] food in the shallow waters along the seashore at dawn or late evening (frequently in creeks and streams of tidewater)'. It is 'met with in small parties, fishing in the shallows' (La Touche 1931-34:433). Foraging birds seen on the wintering grounds in Vietnam were mainly solitary (D.A. Scott pers. comm. 1990).

Blackfaced Spoonbills eat small fish, shrimps, tiny crabs, snails and insect larvae (H.G. Won 1966). An immature female collected in southern South Korea in December 1959 had 'many small fresh-water shrimp' in its stomach (Fennell and King 1964). Delacour and Jabouille (1931, cited in R.P. Allen 1942:101) reported that on the wintering grounds, in what then was French Indo-China, they fed on insects, crustaceans and vegetation. The general behaviour (comfort movements, resting, flight) that we observed in wintering birds in Hong Kong did not differ noticeably from that of the other spoonbills (pers. obs. M.P.K.). On the wintering grounds in Vietnam, birds gather in groups only at high-tide to roost (D.A. Scott pers. comm. 1990). They 'often stand on one foot with their necks on the backs when roosting' (H.G. Won 1966). BREEDING The nesting biology of this spoonbill is poorly known. Nesting is in June and July in Korea. All the presently known breeding sites are on small, rocky offshore islands. From the descriptions and photographs in H.G. Won (1966) and Chung (1986), we know that nests are built on rocky cliffledges and are constructed of dried sticks and grass. The courtship behaviour is not known. Nests are described as 20-30 cm in diameter (Choi 1966). The clutch size is reported as 4-6 eggs (Chung 1986). The eggs are 'white and of prolonged ellipsoid shape with some dotted markings' (H.G. Won 1966). TAXONOMY The Blackfaced Spoonbill has always been regarded as a valid full species, because it is plainly distinct from the sympatric Eurasian Spoonbill (Platalea leucorodid). However, we believe that the question of the relatedness of this form revolves not around the Eurasian Spoonbill but around the Royal Spoonbill (Platalea regia) of Australia. It is possible that the two forms are sufficiently closely related to be considered races of one species (Kahl 1988a). If so, the Blackfaced Spoonbill would be Platalea regia minor. The possible affinities between Platalea minor and Platalea regia were pointed out a hundred years ago by Seebohm (1890:231). It is commonly assumed that the Blackfaced Spoonbill, sometimes called the Lesser Spoonbill, is a very small bird. However, it is small only in relation to the Eurasian Spoonbill. There is in fact considerable overlap in size (see Appendix) between Blackfaced and Royal Spoonbills. As shown in the excellent colour-plate in Ogilvie-Grant (1889:48, pl.l)t the soft-part coloration on the faces of the Blackfaced and Royal Spoonbills is strikingly similar. It appears that the Blackfaced Spoonbill usually has more yellow under the eye, whereas the Royal Spoonbill usually has more yellow over the eye. The Blackfaced Spoonbill also appears to lack the red triangle at the top of the forehead. The yellow eye-spots and red forehead triangles of living I This plate is apparently not in all copies of The Ibis.

266 Blackfaced Spoonbill

spoonbills fade quickly after death, and both appear as dull yellowish tan patches in preserved specimens. It is sometimes claimed that the Blackfaced and Royal Spoonbills can be separated by the extent and shape of the bare black skin under the throat. Of course, this is not a mark that can be used easily in the field. From our admittedly limited experience with museum specimens, we feel that this character is probably much too variable to be of any use in separating the two populations (pers. obs. M.P.K.). However, as with most aspects of the biology of this spoonbill, further study on this point is desirable. A bit of trivia regarding this species, to show the genetic relatedness of spoonbills and ibises, is that a female Platalea minor hybridized successfully with a male Threskiornis melanocephalus in the Berlin Zoo. The hybrid male offspring had the head feathering of the mother spoonbill and the bill of the father ibis. The hybrid later paired with a female Platalea ajaja and produced a number of young (O. Heinroth 1905, A.P. Gray 1958:5). CONSERVATION Considering the small total population, this bird must be regarded as one of the more seriously endangered in the world. According to H.G. Won (1966) the 'breeding grounds on Ai island and other islands off the west coast of North Korea have long been proclaimed [a] sea-bird's sanctuary and guard boats patrol the territorial waters . . . During the breeding period fishing boats and other ships are not allowed to approach the breeding islands. Moreover, landing on the islands and collection of eggs there are strictly prohibited.' According to Chung (1986) and Sonobe and Izawa (1987:37) these areas are still protected by the North Korean government. This is an extremely encouraging situation. A sizeable portion of the population winters at the Mai Po marshes, Hong Kong, where it currently receives strict protection. The Mai Po area is well guarded because it is not only a nature reserve but is also adjacent to the Hong Kong/China international border. Steps should be taken to

assure the integrity of this area—and the ecological health of the adjacent Deep Bay feeding areas—after 1997, when Hong Kong merges with the People's Republic of China. The main wintering grounds in Taiwan are said to be under a threat of reclamation for industrial development or an airport (Lucia Liu Severinghaus pers. comm. 1990). And the Hong River estuary, Vietnam, has been greatly reduced in the last few years to make shrimp-ponds and ricepaddies. There remain only 6000 ha of mudflats, mangroves and islets in the main delta area. Spoonbills are sometimes taken for food in nets or snares in Vietnam. There are plans to establish a nature reserve in the Hong River estuary (D.A. Scott pers. comm. 1990), a conservation measure to be applauded. Any wetland that regularly holds more than 1 % of the world population of any species of waterfowl is considered to be of international importance under the Ramsar Convention. In the case of the Blackfaced Spoonbill, therefore, any wetland holding 3 or more individuals is internationally important (P. Kennerley pers. comm. 1990). At present, however, not a single wetland regularly visited by Blackfaced Spoonbills has been proposed for inclusion under the Ramsar Convention. What losses may occur during migration across China, and in other wintering areas along the South China Sea, is unknown. However, direct persecution of migrating or wintering spoonbills is nearly certain in this densely populated part of the world. It is important that surveys be conducted to find additional nesting, migratory and wintering sites, especially in China. The protection of all wintering and staging areas in China, Korea, Taiwan and Vietnam would be highly desirable, as would the protection of additional breeding sites in China, should they be found.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 325.

African

Spoonbill Platalea alba Scopoli Platalea alba Scopoli, 1786, Deliciae Florae Fauna Insubricae, 2, p. 92; based on 'La Spatule blanche de L'Isle de Luc.on' of Sonnerat, 1776, Voyage Nouvelle Guinee, p. 89, pi. 51: Luzon, Philippines (error), Gape of Good Hope Other names: Red-legged Spoonbill (English); Spatule d'Afrique (French); Lepelaar (Afrikaans); Apamo (Luo); Da-dosa (Mandingo), iNkenkane, isiXulamasele (Zulu); Sotrotsoina (Antakara, Sakalava/ Madagascar)

IDENTIFICATION Standing 75-90 cm tall, the African Spoonbill is a medium-sized, rather stocky, bird with all-white plumage, a grey spoon-shaped bill and red face. In the adult the predominantly grey bill is edged on its upper surface with red; the lower surface is grey and black, sometimes with yellow edges and spots. In addition to the face, the forehead, lores, upper throat, legs and feet are red. The iris is pearly white to pale blue or pale grey. When seen together, the sexes are distinguishable by their relative sizes. They look alike, but the male is slightly larger and has a proportionately longer, heavier and slightly more decurved bill (Kahl 1983). Changes at the start of breeding are subtle. The soft parts become a brighter red, the plumage a more immaculate white. A fluffy and luxuriant crest develops, with loose plumes, rather than the lanceolate ones of the Eurasian and Royal spoonbills. The crest becomes slightly tinged with pale cream colour at the start of the breeding season, which may be from oil-gland secretions. The iris becomes a bright pale blue, and the bill is paler bluish grey with brighter red edges (Kahl 1983). At hatching the skin is pinkish and covered with white down; the bill is pink to orange and the legs are pink. A description of nestlings at various stages of developement is given by Whitelaw (1968). The spatulate tip on the bill begins to develop when the chick is about 11 days old. Immature birds are generally similar to adults, but can be distinguished by the following differences: the feathers extend a bit further forward on the face and forehead; the h^ad is streaked with blackish brown, and the crest is small or non-existent; the tips of the primaries and some under wing-coverts are sooty black; the bill is yellowish to horn-coloured; and the legs and feet are sooty grey to black. The African Spoonbill is usually silent. Rather soft, nasal 'Kor, kor, kor' notes, having a crow- or gull-like quality, are heard in flight, especially as a flock is taking off or preparing to land, and at breeding colonies during displays (FJ. Jackson 1938:90). Also rather loud wing-noise occurs during landing, take-off and in flight. It is especially noticeable around the breeding colony (Kahl 1983). Faint rattling sounds, audible at close range during copulation, are produced by the male beating his bill back and forth against the bill of the female; however, the soft, leathery bills of spoonbills are incapable of producing the loud clattering sounds given by some species of storks. Aggressive interactions at the nest elicit a hoarse 'Kwaark' (Whitelaw 1968). 'Yip-yip' calls are given during copulation. The bird flies with its head and neck extended and legs trailing. When in a flock, spoonbills usually

African Spoonbill

269

al. 1982:208). The one known breeding colony in southern Sudan had 50-100 pairs in the mid-1980s (G. Nikolaus in Luthin 1984a), and the Garsen colony, in eastern Kenya, had over 100 pairs in January 1983 (Coverdale et al. 1983). This species has apparently increased and extended its breeding range southwestward in Cape Province, South Africa, since the 1950s (PJ. Wilson 1957, Blaker 1967, Neame 1968b, Siegfried 1968), possibly because of the increase in farm-ponds in the area.

fly in single file or diagonal line formation. Their flapping rate averages 228/min (n = 25) (pers. obs. M.P.K.). In the northern extreme of its range this species may, occasionally, coexist with the rarer Eurasian Spoonbill. However, the African Spoonbill is slightly larger, has a bare red face, grey (rather than black) bill, and red (rather than black) legs. Although this species can be confused with other wading birds at a distance, its heavier stature and outstretched neck in flight distinguish it from the white herons. The straight, spoon-shaped bill separates the spoonbill from the Sacred Ibis. DISTRIBUTION AND POPULATION This is the characteristic spoonbill of Africa, occurring from southern Mauritania, Senegal, central Sudan and Ethiopia (from about 17°N) south to Cape Province, South Africa, and in Madagascar. Migratory patterns are poorly known. It seems likely that they are mostly nomadic wanderings, in response to local rainfall irregularities, rather than true seasonal migrations. None the less, numbers fluctuate seasonally: African Spoonbills are more common in winter than summer in parts of southern Africa (Brown et al. 1982:208). Birds nesting in Sudan are known to migrate north to Khartoum, Rahad and Kassala in the late dry season (G. Nikolaus in Luthin 1984a). One bird ringed in Transvaal, South Africa, was later recovered several hundred kilometres north in Zambia; other ringed birds have been recovered as far as 680 km from their colony of origin (McLachlan and Liversidge 1970:57). A young spoonbill ringed at the Nata Delta, in northern Botswana, was later recovered in the Orange Free State, South Africa (Tree 1978). No estimates of the African Spoonbill's population size are available. It is widespread but with a patchy distribution, and varies from uncommon to common (Brown et

ECOLOGY The African Spoonbill occurs in many aquatic habitats, including shallow lakes, rivers, marshes, artificial ponds, and along the coast in estuaries and coastal lagoons. It feeds by Walking Slowly through shallow water, Sweeping the bill from side to side. Prey is captured in the 'spoon' of the bill and swallowed with a backward toss of the head. Occasionally it dashes about frantically, in pursuit of fish or other aquatic prey that has been visually detected. In Zululand, Natal, South Africa, spoonbills have been observed feeding behind a wading hippopotamus in shallow water (Pooley 1967a). They also feed in association with herons (J. Reynolds 1965, Connor 1979). In Zululand, Pooley (1968) observed a spoonbill following a Great White Egret (Egretta alba), 'with his bill almost between the heron's legs, apparently feeding off small insects and crustaceans disturbed by the heron as it stalked through the water... the heron flew off several times, but each time the spoonbill followed, until the heron eventually turned and pecked the spoonbill to drive it away5. The diet consists primarily of small fish, aquatic invertebrates and, occasionally, locusts (McLachlan and Liversidge 1970:57). Three stomachs collected by Rand (.1936:336) in Madagascar contained 'twenty small fish and six small crayfish, one large water beetle and many small crayfish, one large water beetle'. The African Spoonbill is gregarious when feeding and is usually found in small flocks. Usually, it is a rather shy bird, but sometimes it becomes quite tame at nesting areas after the eggs are laid. Often the spoonbill will rest on the shore of feeding areas or in nearby trees, standing on one leg with its head rotated to the rear and bill nestled in the feathers of the upper back. BREEDING This species is gregarious in its nesting behaviour. It breeds in colonies of up to 250 nests, often in association with, but somewhat segregated from, other waterbirds, such as ibises, herons, storks, cormorants and darters (Brown et al. 1982:208, Kahl 1983). African Spoonbills usually nest over shallow water, in trees (e.g. Acacia spp.), bushes and reedbeds (Phragmites australis). Whitelaw (1968) found them nesting on an island of reeds in a sewage lagoon. They sometimes nest on the ground on rocky islets (Bannerman 1930:125, pers. obs. J.A.H.) or on rocky ledges (Quickelberge 1972). Breeding seasons are variable throughout the species'

270 African Spoonbill

extensive range in Africa. African Spoonbills nest mainly in the dry season from West Africa to the Sudan, in the rainy season in East and Central Africa, and mixed in both wet and dry seasons in South Africa (Brown et al. 1982:209, Maclean 1985:76). Nesting is also variable from year to year, being suspended in a site when the rains do not occur (FJ. Jackson 1938:90). Courtship displays of the African Spoonbill have been described by Kahl (1983). During pair formation, unmated males establish themselves at potential nest-sites, and unmated females approach them. Males initially attack and drive away any other spoonbill that comes near. Through persistent and repeated approaches, accompanied by Display Flights and Bowing displays, a female is eventually accepted at the nest-site by a male. During Display Flights, the woofing flight-noise is particularly loud and rasping. A pair is established when mutual Greetings and other pairing displays are performed, and copulations follow. The Greeting consists of raising and opening the bill and fully erecting the crest, while uttering a series of loud calls; sometimes the wings are spread or flapped slowly. Soon after pairing, birds engage in several displays that appear to help cement the pair-bond. The series Display Shake/ Display Preen/Bill Popping occurs in a fixed sequence. In the Head Quiver, the erect head feathers shake slightly. Birds also stand side by side on the nest and engage in mutual Head Shaking, in which the bill is pointed vertically downward and the head is shaken in a 'no' motion. In Display Sleeping, birds spend long periods with their bills tucked under the feathers of their back, as if sleeping, but keep their eyes open. During the Copulation display, one bird (probably the male) gives short 'Yip, yip, yip' calls. Copulations last about 10 s and may occur as often as every 5-10 min. Pairs cooperate to defend the nest-site against intrusion by other spoonbills with Sparring and, sometimes, with overt attack. In the Sparring display, the bird erects its crest, gapes its bill and makes biting movements at the opponent. The opponent may respond with a similar action. Allopreening by mated birds is frequent, most often performed by the female. They commonly preen each other at the same time, and most often direct their attentions to the feathers of the head and neck. Both sexes build the nest, the male flying off to collect most of the material; addition of material to the nest continues throughout the breeding season. A flat platform is constructed with small sticks, twigs, or reeds and is sometimes lined with dry grass. Five nests in South Africa averaged 48 X 37 cm and were 6-11 cm deep (Whitelaw 1968). The clutch size is 2-4 eggs. Mean clutches were 2.8 (47 clutches in East Africa; Brown et al. 1982:209), and 2.6 (303 clutches in South Africa; Maclean, 1985:76). Eggs are dull

white to pale buff, blotched with various amounts of reddish brown or dark brown. Incubation, by both sexes, begins after the second or third egg is laid. Females are reputed to incubate more by day, males more by night. In exposed sites, incubating birds sit facing into the wind. The incubation period varies from 25 to 29 days; the average is 26 days (Brown et al. 1982:209). Nestlings are constantly guarded by one or the other of their parents until about 16 days old. Feeding, by both parents, is by regurgitation, the nestling placing its head well inside the throat of the adult. At a South African colony of 12 nests, parental feedings were nearly twice as common in the late afternoon (15.00-17.30 h) than they were in the morning or at midday. By 21 days of age, young begin to wander away from the nest on foot or in short flights, and by about 35 days they are able to fly out of the colony; however, they are still fed by their parents until at least 46 days of age (Whitelaw 1968). Breeding success is generally less than one young reared per pair (Brown et al. 1982:209). In a South African colony, 59.5% of the eggs hatched, and 49.1% of those young survived to fledging, giving an overall survival rate of 34% and 0.82 young per nest (Whitelaw 1968). Nestlings are sometimes taken by birds of prey: Brown (1970:73) stated 'I have seen [an African Fish Eagle, Haliaeetus vocifer] visit a Spoonbill colony repeatedly till it had practically wiped out every brood.' TAXONOMY We agree with Snow (1978:42) that the African Spoonbill is closely allied with Platalea leucorodia, Platalea regia and Platalea minor, and these may be considered to form a 'speciesgroup'. However, Platalea alba is distinct enough from the other three, not to be included with them in a defined superspecies. CONSERVATION As is the case for many African species, this spoonbill faces no apparent major problems. Although it may be affected adversely in some areas by the drainage of wetlands, the expansion of cultivation and ponds may be having positive effects in other areas. Eggs or nestlings are sometimes taken by humans for food. African Spoonbills have been bred in captivity (Wylie 1982).

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 326.

Yellowbilled

Spoonbill Platalea flavipes Gould

Platalea flavipes Gould, 1837, Proc. Zool. Soc., pt.v, p. 106: New South Wales, Australia Other names: Yellowlegged Spoonbill (English)

IDENTIFICATION The Yellowbilled Spoonbill, standing 76-92 cm tall, with a wing-span of about 140 cm, is a rather stout, whitish waterbird with a long neck and legs, and a flattened yellow bill. The adult's entire plumage is a very pale off-white. The bill and legs are dull straw-yellow to grey, and the bare skin of the face is pale yellow. The iris is a pale greyish yellow. The male has a noticeably larger body than the female, as well as a longer bill and legs. Of 5 adult males and 9 adult females examined in museums, the average culmen length of females was 85.8% that of males (18.64 vs. 21.72 cm) and the average tarsus length of females was 90.4% that of males (11.88 vs. 13.14cm) (Kahl 1988a). At the approach of the breeding season, adults moult into a cleaner-looking, very pale yellowish white plumage. A ruff of stiffened, spiky, pale straw-coloured feathers develops on the lower foreneck. More than 100 feathers comprise a typical ruff and the longest ones are 6.5-7.5 cm in length. Several long, filamentous black or dark grey inner-secondaries contrast with the white feathers of the lower back. Also during breeding, the bare skin of the face becomes a pale cyanotic bluish yellow, with areas of pinkish blue. A thin line, 2-4 mm thick, of black skin surrounds the bare face at the junction with the feathered head and neck. The iris ranges from a pale greyish yellow to a pale blue, and the eye is surrounded by a narrow ring of pink skin. A crescent-shaped spot of dark red skin, variable in shape but usually about 9 X 25 mm in size, appears on the base of the bill in front of each eye. The legs become a brighter greyish green-yellow (for additional details and coloured photographs see Kahl 1988a). Hatchlings have pinkish skin and are covered with a sparse white to light grey down; their thick, blunt bill is pinkish yellow. Juveniles appear similar to non-breeding adults, except that they have sooty tips on some primaries, shorter apricot-coloured bills, a darker brown iris, and duskier grey to olivegrey legs. The dark tips to the primaries are sometimes retained by birds that otherwise appear to be fully adult; these may be older subadults. The Yellowbilled Spoonbill is usually silent, except at the nest where a variety of wheezing, coughing and hissing vocalizations are given during displays (Kahl 1988a). The voice was also described as a 'feeble reedy grunt' (Pizzey 1980:64). This species flies with alternate shallow flaps and glides, with the neck and legs extended. The flapping rate averages 218/min (n = 23) (pers. obs. M.P.K.). This species is easily distinguished from the white herons by its stockier build, spatulate bill, and outstretched neck in flight. It often occurs in the same habitat with the slightly smaller Royal Spoonbill, from which it is distinguished by the pale yellow, as opposed to black, bill, face, and legs.

Yellowbilled Spoonbill

DISTRIBUTION AND POPULATION This species occurs primarily on the Australian mainland. Its main breeding area is in southeastern Australia (Blakers et al. 1984:67). It has occurred as a vagrant on Lord Howe Island, King Island, Kangaroo Island, Tasmania (all Australian territory), and New Zealand (Rangaunu Bay, Kaitaia, on North Island; Billing 1977). The Yellowbilled Spoonbill is a nomadic bird, wandering according to rainfall and local conditions of flooding. Little is known of specific population numbers. This spoonbill may be considered common but seldom abundant. However, the species seems to be holding its own in most areas. A maximum of 250 birds was recorded at Lake Cowal, NSW, at a time when the number of Royal Spoonbills was 100, and Strawnecked Ibises 20000 (Vestjens 1975b).

ECOLOGY The Yellowbilled Spoonbill is found more or less wherever there is water. It uses various types of shallow water, including farm-ponds, borders of lakes, swamps and rivers. It is seldom found in tidal areas but does occasionally frequent saltpans and saline swamps. It is found on small, isolated waters more often than is the Royal Spoonbill. The species is most often seen feeding in loosely associated pairs, and sometimes in small groups. It may also feed in mixed-species aggregations, with herons, ibises and other spoonbills. It seems to be intimidated by the somewhat more aggressive Royal Spoonbill. Foraging usually takes place in water less than 40 cm deep. The feeding areas at Lake Cowal were often within 300-500 m of the nesting site (Vestjens 1975b, pers. obs. M.P.K.). As is the case for the other spoonbills, foraging is mainly tactile. The partly open bill is placed in shallow water and, in Head Sweeping, the bill is moved slowly or rapidly from side to side. When a food item is contacted by the 'spoon' portion of the bill, it is grasped, the head raised and the item swallowed with a quick backward toss of the head. Birds also feed in a more directed way by Intensive Search, in which the bill is moved rapidly from side to side in short

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strokes (Vestjens 1975b). It also uses Probing to explore under plants and Grabbing to pick up a visually located prey item. Their Feeding takes place both in the day and at night, although we found foraging activity to be low during the middle of very hot days (pers. obs. M.P.K.). Two Yellowbilled Spoonbills, watched continuously for 8 h by Vestjens (1975b), fed for about 7 h and waded 12 km while feeding. The Yellowbilled Spoonbill has a longer bill, with a narrower spoon than the Royal Spoonbill (Vestjens 1975b). Such differences in bill-shape and slower feeding movements result in different types of prey being taken by the two spoonbills in the same habitat. The diet is mainly aquatic insects (Notonectidae, Corixidae), but also crustaceans, small fish (e.g. the introduced Gambusia affinis) and freshwater snails. Vestjens (1975b) examined a total of 28 Yellowbilled Spoonbill stomachs. He found they ate four times as many crustaceans and insects as fish, and in doing so consumed far fewer fish than did Royal Spoonbills feeding in the same habitat. At Lake Cowal, Vestjens (1975b) observed that Yellowbilled Spoonbills captured about 1.3 items per minute of foraging during the non-breeding season (April-August) and about 7.1 items/min in the same habitat during the breeding months (September-March). A total of 792 items were found in one bird's stomach. Often this spoonbill rests on the shore or in dead trees, with the neck retracted or with the head rotated and the bill buried in the feathers of the upper back. It has the reputation of being shy, distrustful and difficult to approach. Some adult birds at the nest allow a close approach, whereas others are extremely wary.

BREEDING Yellowbilled Spoonbills usually nest singly, although sometimes within 30-75 m of another pair. Colonial nesting seems to be exceptional; however, AJ. North (1913-14:17) reported 40-50 nests of this species, plus many cormorant nests, in one large red gum tree. Other waterbirds, such as Little Pied Cormorants (Phalacrocorax melanoleucos), often build their nests near a spoonbill's. The nest-platform is constructed of sturdy sticks, usually over water, on horizontal limbs or in forks of trees or, less commonly, in bushes or rushes. They usually nest higher than the Royal Spoonbill. Of the 28 Yellowbilled Spoonbill nests found during the 1984-85 breeding season at Lake Cowal, 27 were in river red gum (Eucalyptus camaldulensis) and 1 in a river cooba tree (Acacia stenophylla}\ all the trees were standing in shallow water. These 28 nests varied from 2.5 to 10.0m (average = 5.34 m) above the surface of the water (pers. obs. M.P.K.). Yellowbilled Spoonbills occasionally take over old nests of other large birds, such as the Wedgetailed Eagle (Aquila audax), for use as their own (pers. obs. M.P.K.). The pair-formation process is poorly known, because birds often arrive at the nest site already paired. It is possible that the pair-bond may last for more than a single season (Kahl 1988a). Because pair-formation apparently

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occurs elsewhere and nests are widely spaced, there is less social interaction, and this species is less demonstrative at the nest than other, more colonial, spoonbills. During the Greeting, shown between mates at the nest, the feathers of the head, neck, neck-ruff and scapulars are strongly erected. In this display, each bird emits a series of 4—10 explosive coughing hisses (Kahl 1988a). Birds often perform mutual Head Shaking and Stick Shaking displays at the nest with their mates, especially early in the breeding season. As noted above, this species is generally rather shy around the nest. When approached, the bird on the nest adopts the Erect Posture and gives 'hoarse coughing hisses' in a series, 0.3-1.0 s long and 1-15 s apart (Kahl 1988a). The clutch-size is 2—4, perhaps rarely 5. The eggs are symmetrical, elongate oval in shape and dull, matt white. Incubation is by both parents. The exact incubation period is unknown but probably lasts about 24—26 days. Nestlings use the Begging display, with up and down movements of the head and vocalizations, to solicit food from their parents. In very small young, the call is a weak chittering, high-pitched trill; as they grow older, the call becomes deeper and lower-pitched, beginning to resemble some calls given by adults (Kahl 1988a). Young are fed by regurgitation by both parents. During the 1984-85 season at Lake Cowal, a total of 23 nestling feedings averaged 5.4 s in duration (range = 4.1-7.2s). A brood was usually fed between 11 and 14 times by a parent, and most of the feedings took place within 10-30 min after the parent returned from a foraging trip (pers. obs. M.P.K.). Nestlings leave the nest at about 4 weeks and roam over nearby bushes and trees; they are able to fly at about 7 weeks (Frith 1976:91). If forced to leave their nest-tree before they are able to fly, nestlings are able to swim strongly. The birds appear to suffer predation at the nest site. On

23 October 1984 an Australian Raven (Corvus coronoides) was seen taking a 1-week-old nestling from a Yellowbilled Spoonbill nest at Lake Cowal (pers. obs. M.P.K.). And, in the same area on 19 October 1984, a 2 m carpet python (Morelia spilotes variegata) was observed ascending to a Yellowbilled Spoonbill nest containing eggs. Vigilance and aggression by the attending parents were apparently successful in this case, for the snake was gone the next morning and the adults continued with normal incubation (Kahl 1988a). TAXONOMY The Yellowbilled Spoonbill has sometimes been placed in a monotypic genus, Platibis. However, we agree with Amadon and Woolfenden (1952:6) that all the spoonbills are similar enough to be contained within a single genus, Platalea. Certain morphological features, such as the spiky neck-ruff, show some affinities with the Roseate Spoonbill, Platalea ajaja. And the blackish, filamentous inner secondary plumes that develop during the breeding season are reminiscent of the ibises of the genus Threskiornis. CONSERVATION This species seems to be doing well, and no specific conservation problems are apparent. Some shooting and nestrobbing, for zoo collections, occurs sporadically at places such as Lake Cowal, NSW, but overall losses do not appear to be serious.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 327.

Roseate

Spoonbill Platalea ajaja Linnaeus Platalea ajaja Linnaeus, 1758, Syst. Nat., ed. 10, p. 140; based chiefly on 'Ajaia Brasiliensibus' of Marcgrave, 1648, Hist. Rerum Nat. Brasiliae, p. 204: 'in America australi' = Rio Sao Francisco, eastern Brazil, ex Marcgrave, designated by Berlepsch, 1908, Novit. Zool., 15, p. 301 Other names: Pink Spoonbill, Rosy Spoonbill, American Spoonbill, Pink Curlew, Flame Bird, Pink, Banjo-bill, Flamingo (English); Sevilla [Cuba], Chocolatera [Yucatan] Cuchareta, Cucharon [= large spoon], Cucharera, Chuchara, Espatula rosada, Flamenco, Flamenco espatula, Garza paleta, Garza colorada, Garza cuchara, Garza rosada, Ganso cucharon, Pato rosado, Penitente, Pico de cuchara rosada, Planeta, Plateada, Sidra (Spanish); Spatule, Spatule rouge (French/Creole); Rode Lepelaar (Dutch/Netherlands Antilles); Lepelbek (Dutch/Surinam); Colherado, Colhereiro (Portugese/ Brazil); Ayaya (Brazilian Indian)

IDENTIFICATION The Roseate Spoonbill is a stunningly pink wading bird, with the distinctive spatulate bill characteristic of spoonbills. It is a medium-sized, rather stocky, bird standing 75-80 cm tall, with a wing-span to 120133 cm and weighing about 1.2-1.75 kg. It is the only spoonbill with a brilliantly coloured plumage. The adult's wings, abdomen and tail-coverts are bright pink to carmine. The neck, upper back and upper breast are white. The tail is a surprising tawny-buff to orange. The naked head is greenish, whilst the legs are magenta-ruby, blackish at the joints and toes. The sexes are similar in plumage but dimorphic in size. The male is somewhat larger in body and bill-length. These differences are apparent when a pair is together. During the breeding season all the colours brighten. The lesser wing-coverts and upper and lower tail-coverts become a brilliant carmine, whereas the wings and abdomen become a brighter pink. A patch of stiff, recurved feathers is present on the upper breast and lower foreneck; these are coloured magenta-ruby to carmine. A yellow wash develops on the side, adjacent to the bend of the wing. The soft-part coloration also intensifies. The bill is greyish green to greyish tan with mottled areas of green, black and yellow. The bare skin of the head assumes a pale green to golden-buff colour. A black area of skin develops around the ear-openings, at the edge of the facial feathering, and across the back of the neck (Palmer 1962:534). The black area on the head is individually variable in size and shape (A. Sprunt IV pers. comm. 1990). The iris becomes cherry-red to scarlet. During courtship, some individuals develop bright saffron-orange gular-sacs and orange rings around their eyes. We have seen this in captive birds (from Surinam) at Vogelpark, Walsrode, Germany, as they were courting and preparing to nest. This coloration was first reported by Audubon (1840) from the southern USA and subsequently from South America by other first-hand observers (W.E.D. Scott and R.B. Sharpe 1912, Hudson 1920:125, Wetmore 1926:66). There seems to be a dearth of first-hand reports of this colour change in North American birds, and some accounts may have been secondarily derived (Baird et al. 1884:103, Oberholser and Kincaid 1974:133). However, A. Sprunt IV (pers.

Roseate Spoonbill 277

comm. 1990) states that it does occur in Florida birds, that it is of short duration and indicates 'the highest of high' colours. Additional information is desirable on the gular-sac and colour changes of various populations. At hatching, the nestling weighs about 50 g and is covered with a sparse, short, white down. The skin, including the bill and feet, is a rich salmon-pink (see photo in R.P. Allen 1962; Bent 1926:16, R.P. Allen 1942:119, D.H. White et al. 1982). The skin of a newly hatched spoonbill is not blackish as stated in Palmer (1962:534). At about 2 days, the legs and feet begin to darken, and by fledging they are a dark grey-brown. By about 7 days, the downy coat becomes longer, thicker and woollier. Except for the developing flight-feathers, the body remains largely covered in down until about 21 days (D.H. White et al. 1982). The bill remains basically flesh-coloured until fledging. As in all spoonbills, the bill is initially tubular. The spoon on the end begins to develop at about 9 days. Near fledging (39 days), it is only 67% the length of the adults (D.H. White etal. 1982). Large nestlings are faintly pink and have almost fully feathered faces and heads (W.B. Robertson pers. comm. 1990). The plumage of juveniles is mostly white, with a slight and variable suffusion of pink on the tail and under the wings. The head and face remain more fully feathered than in adults. The iris is dark brown to black. There are dusky tips to the wing-coverts and primaries. The bill is a pale dirty yellow to dull green, and the legs and feet are dark

grey-brown. As birds mature, the amount of pink in the plumage increases. And the head loses feathers, until the full adult plumage is reached at about 3 years. [There is a fuller discussion of plumaged succession in R.P. Allen (1942:119121,123); however, adults may lose the bright carmine in their wings and tail during the non-breeding season, and this would alter Allen's sequence.] 'When disturbed or alarmed, [it utters] a low 'Huh-huhhuh-huh, repeated rapidly, without change in pitch or volume' (Palmer 1962:535). Like the American White Ibis, it makes a soft grunting 'Uh-uh-uh' note while feeding contentedly. Low clucking calls are heard during change-over at the nest and in threat displays. In giving all these calls, the head is raised and the bill opened. We have heard a 'guttural croak' —more guttural and more voiced than in other species of spoonbills —during courtship displays in captive birds, which may be the calls that R.P. Allen (1942:86) referred to as 'low, cackling and clucking sounds.' Soft billrattling is sometimes heard during courtship activities and displays. Probably referring of a colony at feeding time, Chapman (in Bent 1926:17) described the nestling's Begging call as 'a chorus of tremulous, trilling whistles.' We have found the whistling, cheeping calls of nestlings to be quite distinctive, but reminiscent of ibises (pers. obs. J.A.K.). The Roseate Spoonbill has a strong direct flight, often in flocks that arrange themselves in clusters, diagonal lines or wedge-shaped formations. It flies with its neck outstretched, flaps alternating with short glides. Large, but still partly dependent young, follow adults in flight while Begging for food (R.P. Allen 1962). The flapping rate has, apparently, not been recorded. A variation of the normal flight pattern was described by Ogden (1976): 'birds made a short upglide, with their backs arched and wings stiff and angled down, before resuming normal flight.' Bright pink adults could be confused with flamingos and the scarlet phase of the American White Ibis. The Caribbean Flamingo (Phoenicopterus ruber ruber) is taller and thinner. The American White Ibis is smaller. The spoonbill is easily distinguished from both by its characteristic bill shape, but many reports of the flamingos outside their normal range actually refer to Roseate Spoonbills. Pale juveniles might be mistaken for an egret or the white phase of the American White Ibis. Again, the bill shape is diagnostic. DISTRIBUTION AND POPULATION The Roseate Spoonbill's overall breeding range can be defined by the Neotropics. It nests in the southern USA (mainly near the coast in Texas, southwestern Louisiana, southern Florida), the Bahamas (Great Inagua), Cuba, Isle of Pines, Hispaniola and adjacent islands, Mexico (outside the central plateau region), Central America (although breeding records are lacking for some countries), northern and eastern Colombia, coastal and inland Venezuela and the Guianas, southern Brazil (infrequently reported from the main part of the Amazon basin), Ecuador, Peru (mostly east of the Andes), eastern Bolivia, Chile (no recent nest-

278 Roseate Spoonbill

ing), Paraguay, Uruguay, Argentina (more common in the north, south sparingly to Buenos Aires Province) (Bond 1961:41-12, Narosky 1978, Vigil 1973, Short 1975, Blake 1977:203, J. Steinbacher 1979, American Ornithologists' Union 1983:58, R.B. Smith and D.R. Breininger 1988). The species is a casual visitor or accidental over a much greater range and well into the temperate areas of North and South America, including various parts of the northern and western USA, Bimini and elsewhere in the Bahamas, Jamaica, Puerto Rico, the Lesser Antilles, Trinidad, Aruba, Bonaire, coastal Ecuador, southern Patagonia and the Falkland Islands (Baird et al. 1884, Bent 1926:22-23, Bond 1961:41-42, Herklots 1961:41, Voous 1983:60, J. Steinbacher 1979, American Ornithologists' Union 1983:58, Raffaele 1983, Luthin 1984a). The species undertakes dispersals and migrations in various parts of its range. Fledglings as well as adults from Florida disperse rapidly northward from their breeding sites after leaving the nest, with juveniles going as far as 400 km (Bent 1926:21, W.B. Robertson et al. 1983). Unmarked juveniles have been found even further northward (e.g. to Tennessee and South Carolina coast) (W.B. Robertson pers. comm. 1990). Individuals from Texas disperse northward as well (Oberholser and Kincaid 1974:132; Sutton 1967:45^46). In Brazil, they spread out widely after nesting is over (Belton 1984). There are not enough recoveries of marked spoonbills beyond the first year to demonstrate conclusively the seasonal patterns of movement. R.P. Allen (1942 and in Palmer 1962:536) believed that migratory exchange took place between Florida and Cuba, but his view was based on very limited information. Birds move out of Louisiana and much of Texas in winter, returning in February-April (Oberholser and Kincaid 1974:132). However, spoonbills also winter in both Texas and Florida. Although numbers have increased recently in both Florida and Texas/ Louisiana, there seems to be little interchange between the two populations (W.B. Robertson et al. 1983). Clearly, more marking is necessary before the pattern of movement is clear. Spoonbills appear to disperse within South America as well. Although present in Rio Grande do Sul, Brazil, throughout the year, they are scarcer in winter, when they are more restricted to the coast (Belton 1984). The Roseate Spoonbill appears to be a common bird throughout the core of its range in South America, being particularly abundant in the wet savannas of Brazil, the Chaco and Mato Grosso (R.P. Allen 1942:30). This species has always been less abundant at the periphery of its range, in the southern USA and northern Argentina. The current population trend is an overall increase in the USA. The trend was generally downward from 18501920; thereafter, recovery was fairly rapid on the Texas coast and slower in southern Florida. In the summer of 1941, R.P. Allen (1942:29-30) estimated that the total US population was 5698 birds (513 in Florida, 150 in Louisiana, 5035 in Texas), 35% of which were breeding adults. By 1976, it was estimated that there were over 6400 nesting birds in this area (2400 in Texas, 2600 in Louisiana, 1400 in

Florida) (Kushlan and White 1977, Blacklock et al. 1978, Custer et al. 1980, Portnoy 1981). Complete censuses have not been carried out elsewhere, but some numbers are available. These include 3230 pairs on the Gulf coast of Mexico in April-May 1971 (Sprunt and Knoder 1980), 600 pairs on Isla Rajaros and Isla San Pablo in Costa Rica, and 10 pairs on the Surinam coast (Luthin 1984a). Today few are seen in Belize, and the colony there is apparently no longer extant (Luthin 1984a, D.S. Wood and R.C. Leberman 1987). A 1964 nesting colony in Honduras, near Cauquira, numbered c. 300 pairs (Monroe 1968:58). The species is uncommon in Panama, although it was reported as breeding there until 1930 (Ridgely 1976:50). Nearly 1000 birds were counted in aerial surveys in Venezuela (Luthin 1984a), but they were not breeding at the time of the surveys. This species is rare on Hispaniola (Wetmore and Swales 1931, de Dod 1978), but it is common in Cuba, nesting on the Isle of Pines and along the mangrove coast of the main island (Garrido and Garcia 1975:31). It apparently does not nest on the other Antilles. ECOLOGY Roseate Spoonbills feed in various shallow, relatively open, aquatic sites. They utilize heavily saline, salt, brackish and freshwater areas. Over most of the range, these include pools, lagoons, marshes, flooded pastures, stock-ponds and rice-paddies. They also forage in tidal ponds, mangrove pools and streams, mudflats, and estuaries. Feeding areas may be as far as 20-35 km from the roosting or breeding site (J.C. Lewis, 1983, pers. obs. J.A.K.). Although Roseate Spoonbills are usually found in the lowlands, a wandering bird was seen at about 4400 m in the Peruvian Andes (Stott 1959). Feeding sites are usually rather shallow, with maximum foraging depths in Florida Bay being about 20cm (Powell 1987). Although they will feed singly, Roseate Spoonbills are very gregarious, foraging in small or large flocks with other spoonbills, herons, ibises or storks. They often join feeding flocks, and we have attracted them into a feeding site using decoys (Kushlan 1977e). They appear to feed cooperatively with other spoonbills, and other species sometimes use them as 'beaters', gaining a foraging advantage by following them around (M.L. Morrison et al. 1978, J.K. Russell 1978). As with the other spoonbills, food detection and capture are primarily by feeling rather than sight. They typically feed by Head Sweeping, with the bill placed in the water between vertical and nearly horizontally forward. Bob Allen (1947:162) described it graphically: 'The broad, spatulateshaped bill is swept from side to side in wide semi-circles, the mandibles slightly parted, the tips digging gingerly into the surface film of the soft bottom, beneath waters that are nearly always opaque. Delicate, sensitive to the small wrigglings and the darting, skittish movements offish or prawn a quarter of an inch in length or of insects of even lesser dimensions, this keen, responsive instrument must serve as both eyes and hands.' The entire head and neck, or just the bill, is scythed

Roseate Spoonbill

laterally until a prey item is encountered. Then the bill is clamped shut, raised and the food swallowed with a backward jerk of the head. Spoonbills may manipulate large or awkward prey by shaking, biting or beating them for many minutes. They mostly wade in shallow water, Walking Slowly, and moving along with the flock. However, we have seen them Running after prey that they see near the surface. When feeding on active prey, such as killifishes, birds 'will sometimes run from side to side and flap their wings, leaping ahead in order to intercept a school of minnows', but when feeding on slower prey, such as prawns, the pace is more leisurely (R.P. Allen 1942:93). They will Grab at prey located visually, and they also use Probing and Intensive Sweeping to obtain prey in localized areas. We have seen birds feeding in water past their chests, with the entire head and neck immersed (see also photo in Heinrichs 1986), but more often they forage in shallower water. Feeding can take place at any time of the day or night. In tidal areas, feeding activity peaks at low tide (R.P. Allen 1942:93, J.C. Lewis 1983:2). In many areas the birds feed frequently, or even largely, at night (R.P. Allen 1947, Friedmann and Smith 1950, Powell 1987). Many years ago John James Audubon (1840, and in Bent, 1926:19) stated that spoonbills 'are as noctural as the night heron'. Although this is a bit exaggerated, we have seen them feeding actively at night many times, in Florida Bay on the falling tide. R.P. Allen (1957:16-17) suggested that at night many of their prey items are more active and easier to catch. However, in Florida Bay, spoonbills fly out of their colonies and roosts in the morning to feed in mangroves and freshwater marshes during the day, retiring to the roost at night (pers. obs. J.A.K., W.B. Robertson pers. comm. 1990, P. Bjork pers. comm. 1990). Especially at the end of the nesting season, we have seen juveniles feeding together away from the adults. The diet consists almost entirely of small aquatic organisms, such as small fish (e.g. Cyprinodon variegatus, Gambusia qffinis, Fundulus sp.), crustaceans (shrimps, prawns, crayfish, crabs), insects (mainly Coleoptera and Odonata), occasionally molluscs (snails, including periwinkles, slugs) (Hallinan 1924, Bent 1926:19, Palmer 1962:540-541, Haverschmidt 1968:30). Plant material may also be ingested; this seems usually to be done accidentally, along with animal prey, but it is sometimes taken intentionally (R.P. Russell 1982). Three full stomachs obtained in Florida during February-March, contained fish (81%), water beetles (11%), shrimp (5%), plant material (3%), and traces of gastropods and aquatic bugs. One of those stomachs contained a total of 246 small fish, plus other miscellaneous items (C. Cottam and P. Knappen 1939). R.P. Allen (1942:102) gives figures for stomach contents (by volume) as: fish 62%, Crustacea 20%, insects 14%, vegetation 3%, molluscs a trace. At least for the US population, immature sheepshead minnows (Cyprinodon variegatus) are 'probably the most important single species in the diet of the Spoonbill' (R.P. Allen 1942:95). More recent analyses of nestlings' stomach contents in Florida Bay show about 85% fish and 15% palaemonid-shrimp (R. Bjork pers. comm. 1990).

279

Most prey eaten is small (usually <30 mm), but captives were able to swallow pieces offish as large as one's fist (R.P. Allen 1942:101). There are occasional reports of predation on spoonbills, mainly the young, by birds of prey such as Peregrine Falcons (Falco peregrinus) (E.S. Clark 1981) and Bald Eagles (Haliaeetus leucocephalus) (Robertson pers. comm. 1975). Raccoons (Procyon lotor) and Great-tailed Crackles (Quiscalus mexicanus) often eat the eggs (R.P. Allen 1942:59-60). In fact, we find it likely that the distribution of nesting sites in Florida Bay reflect their inaccessibility to raccoons. We have seen eagles and vultures taking eggs and young from the nest, from which the parents were absent. Hurricanes (W.B. Robertson and H.B. Muller 1961), droughts, and winter storms cause some mortality as well. Man is the chief enemy (see Conservation). Ectoparasitic infestations sometimes result in a high mortality of nestlings and desertion by the parents (R.P. Allen 1942:88). In Texas, birds have been observed leaving the water where they have been feeding to defecate (McDaniel et al. 1963); this may help to reduce the cycling of parasitic infections.

BREEDING The Roseate Spoonbill usually nests on islands away from ground predators, in colonies of up to 500 or more pairs. (R.P. Allen 1942:89, W.B. Robertson et al. 1983). In Brazil, it nests on Ilha do Governado, at the gates of the city of Rio de Janeiro (Sick 1985). In most areas, such as the Venezuelan llanos, it typically nests in the company of various herons, ibises, cormorants and storks. Spoonbills are sometimes dominated by herons at the nest-site (R.P. Allen 1942:89). The nesting season varies geographically, as would be expected for such a widespread species. Depending on the region, nests are built in bushes, low trees —such as mangroves, Rhizophora mangle, and Avicennia germinans in Florida—and sometimes in taller trees (up to 20-25 m high). Spoonbills also nest in rushes or, rarely, on the ground in secure sites (J.W. Harrison 1978). In Texas, where larger trees are not available, spoonbills use low vegetation on dredged-material islands (R.P. Allen 1942:50, D.H. White et al. 1982, J.C. Lewis 1983). In general, the preferred vegetation height for nesting is greater at mainland than at island sites, possibly because of decreased wind and increased need for protection from terrestrial predators on the mainland (J.C. Lewis 1983). R.P. Allen (1942:50) felt that a suitable feeding area near the nest is essential for justfledged young to use for several weeks before they join adults at more distant feeding grounds. The nest is often described as 'frail'. We have found, however, that it is a rather sturdy platform of sticks and twigs, often lined with finer twigs, grasses, leaves, bark or moss. At one Texas colony nests were on average 55 cm (range 43-71 cm) wide and 10-13 cm deep (D.H. White et al. 1982). R.P. Allen (1942:84) found nests in Texas 'more substantial in their construction' than nests in Florida. Birds arrive at breeding areas in flocks, presumably

280 Roseate Spoonbill

unpaired. Bob Allen (1942:73-74) described 'up flights' and 'sky gazing' as typical behaviours seen in the pre-pairing stage. During 'up flights', flocks suddenly take wing en masse, circle around, and then land again. In 'sky gazing' a group of standing adult spoonbills react to another flying over by extending their necks to full length, pointing their bills skyward and appearing to gaze upward for a few seconds. These two behaviour patterns seem not to have been reported in other species of spoonbills. However, the 'up flights' might possibly represent a group version of the Display Flight, described for lone African, Royal and Yellowbilled Spoonbills (Kahl 1983, 1988a). Pairing birds also engage in the Display Preen. This display of the Roseate Spoonbill, as witnessed in courting captive birds at Vogelpark, Walsrode, Germany, was quite different from the homologous displays of the other spoonbills so far studied (details in Kahl 1983, 1988a). In the Roseate, the head is shaken in a very loose-jointed way, with the tip of the bill describing a 'figure-eight'. Then there is a quick, loose-jointed preen of the upper wing-coverts, the primaries or the neck-ruff, which is erected. During courtship, some soft bill-rattling is heard, especially during the Display Preen. Potential mates bite repeatedly at each other's bills when excited. Displaying and courting birds often fly and land with the tongue-bone extended, causing the throat-skin (or gular-sac) to be conspicuously exposed (pers.obs. M.P.K.). Copulations, once begun, are frequent. In one pair the first successful copulation was observed 14 days after pairformation (R.P. Allen 1942:89). In one period of 2 h 40 min, a pair copulated three times and the male brought 11 twigs to the nest (Palmer 1962:538). An apparently successful copulation lasted 20-25 s (R.P. Allen 1942:84). Other breeding displays described by R.P. Allen (1942) and Palmer (1962:538) appear to be generally similar to those observed in other species of spoonbills (Kahl 1983, 1988a). However, further detailed studies are needed to fully describe the display repertoire of this species. Both sexes cooperate to build the nest, with the male collecting most of the material and the female doing most of the building. They may use the nests of earlier-nesting ibises (Weller 1967). There is apparently an increase in nestbuilding activity just before the eggs hatch (R.P. Allen 1942:88). Eggs vary from ovate to elliptic-ovate or even elongateovate (Bent 1926:16). They are white, or nearly white, with no gloss, and with spots and small blotches of various shades of brown. The clutch size is 1-5, usually 2 or 3. R.P. Allen (1942:34) gives the average clutch size as 2.7. In a Texas colony, 87 nests had an average of 2.64 eggs per clutch on 27 May, when egg-laying was complete (Palmer 1962:538). In 154 marked nests over a 3 year period in Texas, clutch size varied significantly from year to year but averaged 3.0 (D.W. White et al. 1982). One female laid her first egg 6 days after the first observed copulation (R.P. Allen 1942:89). Eggs are usually laid every other day. Incubation, by both sexes, begins the day after the first egg is laid and lasts from 22 to 24 days, probably most often

22 days (R.P. Allen 1942:88, Palmer 1962:540, D.H. White etal. 1982). R.P. Allen (1942:34) stated that breeding adults are quite nervous and 'will desert a nest that is disturbed'. Care must be taken, as with other spoonbill species, in approaching the nests for observation or photography, especially before eggs have been laid. However, comparisons between intensively studied and relatively undisturbed colonies (D.H. White et al. 1982) show that careful observations can be done without causing significant mortality. Both parents feed their young by regurgitation. Young put their bills in the side of the adult's bill and down into the throat to obtain the meal. Parental feedings last about 10s (F.M. Chapman 1914, and in Bent 1926:17). Young leave the nest at about 4—5 weeks (R. Bjork pers. comm. 1990). They remain nearby and are fed by parents until at least 8 weeks. Nesting success varies widely, depending on environmental conditions. In some years success is good, whereas in others most nests may be lost. R.P. Allen (1942:34) reports that In some seasons a colony appears to hatch and rear all eggs and young; in other seasons I have recorded nest mortality as high as 98.5%. The average nest mortality for Florida Bay for the five seasons from 1936-37 to 1940-41, inclusive, is estimated at about 35%. In Texas the average nest mortality over a six-year period has been about 22% . . . the average number of young produced annually per pair is about 1.8 in Florida Bay; 2.1 in Texas. At a recent Texas colony, D.H. White et al. (1982) found that among 154 nests in 3 seasons, 73% of the eggs laid were hatched and 50% of the eggs laid were fledged, giving an average production of 1.5 young per nest. Roseate Spoonbills probably breed, at the earliest, in their third year (R.P. Allen 1942:33). Although Allen stated that 'only adults in full breeding plumage pair', Dunstan (1976) observed successful breeding by adult spoonbills that lacked the full breeding plumage. Pairing is probably for a single breeding season only (R.P. Allen 1942:34). TAXONOMY The Roseate Spoonbill has sometimes been placed in a monotypic genus, Ajaia, because of its distinctive plumage and its simplified trachea (R.P. Allen 1942:126, Vestjens 1975a). However, we agree with Mayr and Short (1970) and J. Steinbacher (1979) that it should be included within the genus Platalea. Certain morphological features, such as the spiky neck-ruff and lack of nuchal crest, show an affinity with the Yellowbilled Spoonbill (Amadon and Woolfenden 1952). It may be more closely related to this species than to spoonbills of the Eurasian/Royal/Blackfaced group. CONSERVATION The large and widespread populations of Roseate Spoonbills throughout tropical America have apparently

Roseate Spoonbill

remained more or less stable. Based on its history, this species appears to be rather resilient in recovering and maintaining its population levels. It also may be more resistant to the effects of pesticides than some other species of waterbirds (Oberholser and Kincaid 1974:133, D.H. White etal. 1982). In North America and Cuba, however, the species was at one point brought to the verge of extinction (Sprunt 1939, R.P. Allen 1942, T. Barbour 1943:19, Oberholser and Kincaid 1974:133). At the low point, between 1890 and 1919, there may have been only 20-25 breeding pairs left in the USA. 'If a reservoir of supply had not existed in countries to the south, it would probably have become extinct in the United States before 1900' (R.P. Allen 1942:14). The drastic and widespread decline of the US population came during the plume-hunting period in the late 19th century. Spoonbills were occasionally killed so that their wings could be made into fans. However, because the feather colour fades rapidly, the Roseate Spoonbill was not, itself, hunted on a large scale for the millinery trade. Their decline was probably brought about by the wanton disturbance of breeding areas, also containing egrets that were hunted on a large scale for their plumes. According to R.P. Allen (1942:3) 'the shooting of egrets in colonies that these birds shared with the "Pink Curlews" would have had an effect on both species, even though no Spoonbills were actually killed. Death of the egrets would, in my opinion, coincide with desertion of the area by the Spoonbills.' During the current century, populations in the southern USA have shown a dramatic recovery, first on the Texas/ Louisiana coast and later, and to a lesser extent, in southern Florida. Without a doubt much of that recovery was due to increasingly stringent wildlife legislation and vigilant protection of breeding areas in sanctuaries, primarily those administered by the National Audubon Society. In Texas and Louisiana, recovery occurred between 1920 and the 1970s. The Roseate Spoonbill began to reappear as a breeder on the Texas coast in the 1920s, and by 1976, it was estimated that there were over 2400 nesting birds in Texas and 2600 in southwestern Louisiana (Blacklock et al. 1978, Portnoy 1981). Recovery in Florida came more slowly, possibly because Florida Bay, the spoonbill's main nesting ground there, was still too remote for adequate law enforcement. Spoonbills were still being exploited, mainly for food, through the 1930s (Powell et al. 1989). Partial recovery did eventually occur in Florida, and Robert Porter Allen (1962) could

281

write: 'From a single tiny colony of 15 pairs in 1939, the Florida nesting population increased to a peak of eight colonies and more than 400 breeding birds in 1955-56'. From 1955 to 1978 the nesting population in Florida Bay roughly doubled every 10years (Powell et al. 1989). By 1978-79 there were approximately 1400 breeding pairs in southern Florida (W.B. Robertson et al. 1983). They are now breeding, in small numbers, as far north as Tampa Bay (west coast), where they had not nested since 1912 (Dunstan 1976), and on Merritt Island (east coast) (W.B. Robertson pers. comm. 1990). They also are expanding their range from west to east in Louisiana (Portnoy 1981). Surveys during 1984-87 suggested that the Florida Bay breeding population dropped back from the 1978-79 high of 1254 pairs to approximately 450-590 pairs (Powell et al. 1989). At the same time, populations of other fish-eating birds using Florida Bay have also suffered reproduction and population decreases. These include the Osprey (Pandion haliaetus} (Kushlan and Bass 1983), the Brown Pelican (Pelecanus occidentalis] (Kushlan and Frohring 1985), and the Great Blue Heron (Ardea herodias] (Powell 1983). These simultaneous declines are probably due to a common cause, the reduction of food availability (Kushlan and Frohring 1985). The source of these problems may lie in disturbance to the ecology of south Florida. Reduced bird populations stem from decreased production of young in years past. This may be due to inadequate food resources in Florida Bay, which in turn may be caused by man-made changes in the freshwater run-off from the Everglades. Additionally, commercial and residential development in the Florida Keys has claimed wetlands that were once used by foraging spoonbills (Powell etal. 1989). The very high numbers (1254 pairs) in Florida Bay during 1978-79 may just be an anomaly, reflecting the unusually high recruitment of new birds from a very successful season 3 years before; except for that one year, the Florida Bay population has been at 550-750 pairs for the past 15-20 years (A. Sprunt IV, pers. comm. 1990). According to W.B. Robertson (pers. comm. 1990), there has been a considerably better nesting success in Florida Bay during the past 2-3 years.

Note: Body measurements, egg measurements and nesting season data for this species can be found in the Appendix on page 328.

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Appendix

American Wood Stork (Mycteria americana) BODY M E A S U R E M E N T S

Sex

Male Male, average Male Male, average Male Male, average Male Male, average Male Male, average Male Male, average Male Male Male Fern. Fern., average Fern. Fern., average Fern. Fem. Fem. Fem. Fem. Fem. Fem. Both Both, average Both Both, average

[n]

Wing (mm)

[3] 464-475

[3]

[6] [6]

[2] [2]

[4] [4] [2] [9] [?] [?] [1] [1] [1]

470 453-492 476 455-479 467 — — — — 452-484 468 — — 493

[6] 435-455 446 [6] [?] 422-464 [?] 443 [1] [1] [1] [1] [1] [1] [1] [?]

[?] [?]

[?]

460 — — — — —

— 443-488 — — —

EGG M E A S U R E M E N T S

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

227-240 233 215-235 227 221-244 233

196-205 200 194-213 203

186-204 195 155-166 158 — — — — — — 155-160 158 — — 155 143-150 148 132-155 146 — — — — — — — — — — —

1664-1790 1727 —

— — —

— — 205-230 — 215-239 227 — 225 235 190-205 196 192-208 202 205 — —

— 197-205 — 197-214 208 — — 208 175-192 183 183-202 193 — — —

— 205 200 — — — — —

— 185 180 — 175-213 — — —

— — — — — — — — — — — — — — — — — — — — — — — — 1650 1550-1650 1650

Weight (kg) — — — _ — 3.05-3.30 3.18 _

2.70 _ _ 3.09 2.50 — — — — — — 2.05 2.78 2.40 2.15 — 2.80 — — _ —

Reference Extremes Average Extremes Average Average Average Extremes Average Calculated Average

(1) (1) (2) (2) (5) (5) (6) (6) (10) (7) (13) (13) (1) (10) (12) (2) (2) (13) (13) (5) (7) (8) (9) (10) (10) (11) (3) (3) (4) (13)

No.

Length (mm)

Width (mm)

Weight (g)

Reference

35 35 40 40 20 — 64 64 64 —

62.5-72.3 67.7 60.8-73.0 67.9 68.1 — 62.0-73.0 68.0 — 67.8

43.8-48.8 46.7 34.3-54.9 46.0 46.5 — 43.0-51.0 46.0 — 46.0

— — — — 83 — — 80 —

(1) (1) (2) (2) (3) (4) (5) (5) (5) (6)

References: (1) Pers. obs. M.P.K. (2) Bent 1926:60. (3) Palmer 1962:513. (4) O. Heinroth 1922. (5) Schonwetter 1967. (6) Oberholser and Kincaid 1974:125.

EGG-LAYING Place S Florida (formerly) S Florida (currently) Central/N Florida Georgia, South Carolina SE Texas Mexico (Tabasco, Campeche) Honduras Costa Rica Venezuela (Llanos) Surinam (coastal) Brazil (Matto Grosso) Argentina (E central)

Longevity in wild = 11 years, 8 months (pers. obs. M.P.K.; from USFWS Banding Office). Longevity in captivity = 30-1- years (M. Davis pers. comm., 16 July 59). Body temperature = 40.65°C (n = 66) (Kahl 1963b). Breathing rate = 11-12/min (Kahl 1963b). References: (1) Pers. obs. M.P.K. (2) Blake 1977:187. (3) Eaton 1910:244. (4) Forbush and May 1955:41. (5) Palmer 1962:510. (6) Hartman 1955. (7) Hartman 1961:41. (8) Haverschmidt 1968:26. (9) O. Heinroth 1922. (10) B.T. Thomas 1985:923. (11) Tashian 1953. (12) Sharpe 1898:322. (13) Oberholser and Kincaid 1974:124125.

284

Months Nov-Dec (a few Jan+) Jan-Mar (a few Nov-Dec) Feb-May Apr-Jun May-Jul Jan-Mar Jan-Feb Dec-Jan Sep-Dec Sep-Oct Jul-Sep Oct-Nov

Milky Stork (Mycteria cinered) BODY MEASUREMENTS

Sex

[n]

Fern. Fern. Both Both, average p p p

[1] [1] [?] [?] IU [1] [1]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

435 450

201 195 194-238 220 203 270 275

200 190 188-225 188 188 223 225

145 153 156 160 170 165 165

— — — — — —

— — — — — —

(1) (5) (3) (1) (2) (4) (5)

4.4.4

494

470 438 495 500

References: (1) Kuroda 1936:539. (2) Pers. obs. M.P.K. (Malaysian Game Dept./25 Jan 68). (3) H.C. Robinson and F.N. Chasen 1936:206. (4) Sharpe 1898:326. (5) Delacour and Jabouille 1931:93.

EGG M E A S U R E M E N T S

Extremes Average Calculated Extremes Average Extremes Average Extremes Average

No..

Length (mm)

Width (mm)

Weight (g)

Reference

44 44 44 24 24 3 3 3 3

59.0-74.5 66.3 — 62.0-68.2 64.8 62.7-64.1 63.3 66.0-70.0 68.3

43.0-48.0 45.9 — 43.3-47.9 45.4 43.9-45.7 45.1 45.0-46.0 45.7

— 77 — — — — — —

(1) (1) (2) (3) (3) (3) (3) (4) (4)

References: (1) Hoogerwerf 1949b:19. (2) Schonwetter 1967. (3) Hellebrekers and Hoogerwerf 1967:13. (4) Pers. obs. M.P.K. (Mus. Zool., Bogor/8 Mar 68).

EGG-LAYING

Place

Months

WJava SE Sumatra W Malaysia

Mar—Aug Jun-Aug Aug

285

Yellowbilled Stork (Mycteria ibis) BODY MEASUREMENTS

Sex

W

Fern. Fern. Fern. Both Both Both Both Both Both Both Both Both ?

[1] [1] [1] [?] [?] [?] [?] [?] [?] [?] [?] [?] [1]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

425 — —

200 — — 205-242 205-260 208-229 205-242 205-230 — — 203-243 203-243 245

200 — — 197-229 190-230 200-221 197-229 190-230 — — 195-229 195-218 225

175 — — 168-183 160-190 175 168-183 160-180 — — 168-190 170-190 183

— — 1500-1650 — — — 1500 — 1500-1650 1500-1650 — —

1.19 2.33 — — — — — — — — — —

(10) (11) (11) (1) (2) (3) (4) (5) (6) (7) (8) (9) (12)

455-513 450-515 450-475 455-513 450-500 490-526 — 455-517 455-517 500

Longevity in captivity = 19-1- years (K. Brouwer et al., pers. comm., 1990). References: (1) Brown et al. 1982:173. (2) Schouteden 1954. (3) Heuglin 1869-73:1130. (4) Bannerman 1930:108. (5) Reichenow 1900-01:334. (6) Clancey 1964a:47. (7) Cramp 1977:322. (8) Maclean 1985:70. (9) McLachlan and Liversidge 1970:50. (10) Andersson 1872:296. (11) Britton 1970b. (12) Sharpe 1898:328.

EGG MEASUREMENTS

Extremes Average Calculated Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Single

No.

Length (mm)

Width (mm)

17 17 17 — — 15 15 3 3 36 36 51 51 —

59.0-72.0 66.0 — 59.0-66.0 61.3 62.1-71.1 66.7 65.0-68.0 66.0 59.0-72.3 66.0 59.0-72.3 66.2 —

42.0-46.0 44.0 — 40.0-45.0 42.0 42.3-47.6 45.3 44.0-46.0 45.0 42.0-46.1 44.0 42.0-47.6 44.4 —

EGG-LAYING Weight (g)

Reference

— 71 — — — — — 77 — — — — 62

(1) (1) (1) (2) (2) (3) (3) (4) (4) (5) (5) (6) (6) (7)

Place West Africa: Gambia to N Nigeria Gambia, Senegal, Guinea-Bissau Ghana N, central Nigeria Chad S Sudan S Somalia W Kenya, Uganda NW Kenya [L. Turkana] E Kenya NW Tanzania S Zaire Zambia Malawi NE Botswana NE South Africa (Natal)

References: (1) Schonwetter 1967. (2) Reichenow 1900-01:334. (3) W. Fraser 1971. (4) Mundy and Cook 1974. (5) Brown et al. 1982:175. (6) Maclean 1985:71. (7) Heinroth 1922.

286

Months Nov-Feb Nov-Feb Nov-Jan Aug-Jan Jun-Sep Oct-Nov Oct Mar-Jun Aug-Sep Jan, May-Aug Jan-Jul Jul-Aug Feb-Sep Jul-Sep Mar, Aug-Sep May-Oct

Painted Stork (Mycteria leucocephald) EGG MEASUREMENTS

BODY MEASUREMENTS

Sex

[n]

Male Male Fern. Fern. Fern. Fern. Both Both Both Both Both Both Both Both, average Both Both, average Both, average

[1] [1] [1] [1] [1] [1] [?] [2] [?] [?] [?] [3] [6] [6] [10] [10] [?]

Wing (mm)

Culmen (mm)

500 513 488 525 488 505 490-510 -

— 215 252-278 —

490-500 450-510 490-523 505 495

— — 215-278 — 230-256 246 — — 279

263 275 225

Tarsus (mm)

(mm)

Tail

203 — 240 194 230 210 240-250 — — — 210-250 — 205-242 225 — — 243

156 — 150 163 155 — 150-172 — — — 150-172 — — — — — 152

Wing-Span

Weight

(mm)

(kg)

— — — — — —

— — — — — —

1500-1600 — — — — —

2.02-3.5 2.0-3.0 — — — — 3.03-3.37 3.18 —

Reference Extremes Average Calculated Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Average

(7) (9) (9) (7) (7) (10) (2) (3) (4) (8) (10) (10) (5) (5) (6) (6) (1)

No.

Length (mm)

Width (mm)

60 60 60 60 60 27 27 50 50 3 3 2 50 —

65.0-80.0 70.0 — 66.0-75.0 69.6 64.5-73.8 69.3 65.3-80.2 65.9 67.6-68.2 67.9 — 69.5 71.0

43.2-51.6 48.0 — 41.5-48.0 43.9 43.8-49.5 47.0 43.2-51.6 45.0 47.3-48.5 47.7 — 49.0 48.0

Weight (g) — 89 66-85 74 — — — — — — 82-98 — —

Reference

(i) (i) (i)

(2) (2) (3) (3) (4) (4) (8) (8) (5) (6) (7)

References: (1) Schonwetter, 1967. (2) Desai etal. 1977. (3) Hume and Gates 1890:223. (4) E.C.S. Baker 1935:451. (5) O. Heinroth 1922. (6) S. AH and S.D. Ripley 1968:94. (7) Henry 1971:385. (8) Pers.obs.M.P.K. (N India/lOSep67).

Longevity in captivity = 28+ years (K. Brouwer et al. pers. comm. 1990). References: (1) T.H. Shaw 1936:120. (2) E.G.S. Baker 1929:332. (3) S. AH and S.D. Ripley 1968:95. (4) O. Heinroth 1922. (5) Abdulali 1968. (6) Desai et al. 1974. (7) Legge 1880:1100. (8) Tom Roberts pers. comm. (9) Sharpe 1898: 325. (10) Delacour and Jabouille 1931:92.

EGG-LAYING Months

Place Pakistan N India S India Sri Lanka Burma Thailand (Ban Yang, formerly) Kampuchea (Grand Lac)

287

Jun-Nov Jul-Oct Nov-Mar Mar-Apr Aug-Nov Jul Jan-Feb

Asian Openbill Stork (Anastomus oscitans) BODY MEASUREMENTS Wing (mm)

Sex

[n]

Male Fern. Fern. Fern. Both Both Both, average

[1] 388 [1] 400 [1] 385 [1] 370 [?] 400-425 [?] 392-408 [14] -

Culmen (mm)

Tarsus (mm)

Tail (mm)

153 160 — 158 158-163 153-162 160

158 150 — 134 150-158 140-150 —

160 155 — — 150-188 183-217 —

Wing-Span (mm)

Weight (kg)

Reference

— —

— — — — — —

(4) (4) (5) (6) (1) (2) (3)

1190 1475-1488 — —

Longevity in captivity = 7+ years (K. Brouwer et al. pers. comm. 1990). References: (1) Legge 1880:1103. (2) E.C.S. Baker 1929:333. (3) Kahl 1971c:27. (4) Sharpe, 1898:307. (5) Delacour and Jabouille, 1931:94. (6) Pers. obs. M.P.K. (northern India/13 Sep 67).

EGG MEASUREMENTS

Extremes Average Average Extremes Average Extremes Average Extremes Average Calculated Extremes Single

No.

Length (mm)

Width (mm)

Weight (g)

Reference

80 80 40 100 100 57 57 30 30 30 2 —

48.3-64.0 57.8 55.0 48.3-63.1 57.9 50.0-63.0 56.0 51.0-62.0 58.0 — 55.8-59.7 59.0

36.4-43.4 41.1 37.3 36.4-45.2 41.2 37.0-45.5 40.0 36.4-44.2 41.5 — 40.0-41.2 41.0

— — — — — — — — 55 — —

(1) (1) (2) (3) (3) (2) (2) (4) (4) (4) (6) (5)

References: (1) E.C.S. Baker 1929:334. (2) Hume and Gates 1890:225. (3) E.C.S. Baker 1935:454. (4) Schonwetter 1967. (5) Herbert 1926. (6) Pers. obs. M.P.K. (N India/10 Sep 67).

EGG-LAYING Months

Place N India, Burma S India Sri Lanka Thailand

Jun-Sep Nov—Mar Dec-Apr Nov-Feb 288

African Openbill Stork (Anastomus lamelligerus] BODY M E A S U R E M E N T S

Sex

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

172 194 — — — — _

160-180 — 154-171 153-194 — — 175

— 160 — — — — — 140-160 — 119-146 133-165 — — 160

— — — — — _ — 180-210 — 185-196 165-206 — — 175

— — — — — — — 1400 — — 1400 — — —

— — 1.25 1.0 1.14 1.26 1.17 — — — — — — —

(6) (11) (7) (7) (8) (12) (12) (1) (2) (3) W (5) (10) (9)

153 140

133 130

— —

— —

— —

(11) (11)

A. I. lamelligerus

— Male, average [16] 432 Male [1] — Male [1] — Fern. [1] — Fern. [1] — Fem. [1] — Fern. [1] Both [?] 400-420 Both [?] 384-436 Both [?] 369-400 Both [30] 370-433 Both [?] 400-440 Both [?] 390-435 393 ? [1] A. I. madagascariensis

Male Fem.

[1] [1]

400 360

EGG M E A S U R E M E N T S

No.

Extremes Average Extremes Average Extremes

39 39 14 14 13

Average Extremes Average Extremes Average Extremes Extremes Average Extremes Average Calculated

13 5 5 12

EGG-LAYING Months

Chad Ethiopia (L. Tana) Kenya, Uganda Tanzania Zambia Zimbabwe NE Botswana N Namibia N South Africa Madagascar

Sep Aug-Sep Recorded all months Jan-Apr/Jul-Aug Recorded all months Sep-Jun Jan-Feb Jan-Mar Feb-Mar Sep-Dec

289

4 15 10 10 10

Width (mm)

Weight (g)

51.0-61.4 55.4 51.0-59.5 54.7 53.5-55.5 [sic] 55.7 53.4-59.8 56.2 52.0-60.5 56.4 55.3-61.4 56.0-58.0 53.3 51.0-59.5 54.4 —

36.7-43.2 40.0 39.5-43.2 40.5 36.7-41.5

35-50 — — c. 50 —

(2) (2) (3)

39.9 38.2-40.3 39.3 36.5-40.5 39.0 40.8-41.3 40.0-42.0 39.9 39.5-43.2 40.9 —

— — — — — — — — — — 50

(3) (4) (4) (5) (5) (6) (7) (8) (9) (9) (9)

Reference

(i) (i)

References: (1) Maclean 1985:68. (2) Brown etal. 1982:177. (3) Anthony and Sherry 1980. (4) Benson and Pitman 1958. (5) Braine 1974. (6) Bouet 1955a. (7) F.J.Jackson 1938:75. (8) Heiden 1974. (9) Schonwetter 1967.

References: (1) Reichenow 1900-01:336. (2) Macworth-Praed and Grant 1973:67. (3) Heuglin 1869-73: 1120. (4) Bannerman 1930:102. (5) FJ. Jackson 1938:73. (6) Kahl 1971c. (7) Brown et al. 1982:176. (8) Maclean 1985:68. (9) Sharp 1898: 309. (10) Clancey 1964a:45. (11) Pers. obs. M.P.K. (Leiden Mus./26 Apr 89). (12) FitzPatrick Inst., Univ. of Cape Town (unpubl. data).

Place

12 —

Length (mm)

Black Stork ( Ciconia nigra) BODY M E A S U R E M E N T S

Wing (mm)

Sex

[n]

Male Male Male Male Fern. Fern. Both Both Both Both Both Both Both Both Both Both Both Both Both Both Both Both, average Both ? p p

[1] 592 [1] 560 [1] 538 [1] 525 [1] 559 [1] [?] 520-580 [?] 520-600 [?] 520-550 [9] 520-600 [?] 530-570 [?] 516-580 [?] 540-615 [?] 525-590 [?] 530-570 [7] 520-575 [4] 540-585 [?] 520-600 [9] 520-600 — [?] [8] 520-580 539 [8] [?] 520-600 550 [1] — [1] — [1]

Culmen (mm) 188 205 193 175 185 — — 170-195 160-190 180-195 — — 170-195 180-195 175-194 193-198 160-190 160-190 — 150-190 — 160-195 185 — —

Tarsus (mm) 213 195 205 190 205 — — 190-205 180-200 185-195 — — 185-200 185-195 182-207 198-204 180-(280?) 180-200 — 175-200 — 180-200 — — —

Tail (mm) 260 250 200 213 237 — 230-250 250 190-240 230-250 — — — 230-250 220-245 245-264 190-240 — — — 190-250 — —

EGG M E A S U R E M E N T S

Wing-Span (mm) — 1900 — — — -

1440-1550 — 1440-1550 — —

1550 — —

Weight (kg)

Reference

3.2 — — — — — — 3.0 — — 3.0 — — — — — — c. 3.0 3.0 — — c. 3.0 3.0 3.0 2.59 2.25

(15) (16) (17) (21) (21) (16) (1) (2) (3) (4) (5) (6) (7) (8) (9) (10) (10) (11) (12) (13) (14) (14) (19) (18) (20) (20)

Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Average Average Extremes Extremes Average Extremes Average Calculated

No.

Length (mm)

Width (mm)

Weight (g)

Reference

11 11 84 84 47 47 21 21 6 6 3 3 3 3 3 3 — 50 — 20 20 80 80 80

61.2-67.1 64.0 60.3-74.3 65.4 60.0-74.0 69.0 63.0-74.0 69.4 64.5-69.5 66.7 70.0-74.0 71.3 68.5-69.5 68.8 68.0-73.0 70.0 — 65.5 60.0-74.3 62.8-70.8 67.0 (60.0)-72.0 65.4 —

46.1-49.0 47.0 45.2-51.7 48.8 45.0-56.0 48.7 45.0-56.0 48.8 45.0-48.3 47.3 48.5-50.0 49.2 48.5-49.0 48.8 47.8-48.0 47.9 — 49.0 44.0-54.7 47.3-51.8 49.6 44.0-53.0 48.8 —

— — — — c. 86 — — — — — — — — — — 83 — 86 — — — — 86

(1) (1) (2) (2) (3) (3) (4) (4) (5) (5) (6) (6) (7) (7) (8) (8) (9) (10) (11) (12) (12) (13) (13) (13)

References: (1) Dementiev and Gladkov 1951:442. (2) Hartert 1912-21:1216. (3) Maclean 1985:66. (4) McLachlan and Liversidge 1970:53. (5) Priest 1933. (6) P.R. Barnes 1950. (7) Ellenberger 1951. (8) Howells 1968. (9) O. Heinroth 1922. (10) Etchecopar and Hue 1978:73. (11) Schroder and Burmeister 1974:31. (12) Adam Mrugasiewicz pers. comm. (6 April 1969). (13) Schonwetter 1967.

Longevity in wild = 18 years (Bauer and Glutz von Blotzheim 1966:424). Longevity in captivity = 30 years (Dresser 1871-81:316). Longevity in captivity = 314- years (K. Brouwer et al. pers. comm. 1990). References: (1) Clancey 1964a:43. (2) Sokolowski 1968:146. (3) Reichenow 1900-01:347. (4) McLachlan and Liversidge 1970:52. (5) O. Heinroth and M. Heinroth 1924-31:139. (6) Mackworth-Praed and Grant 1973:64. (7) Meinertzhagen 1954:390. (8) Kozlova and Tugarinov 1947:263. (9) Hartert 1912-21:1216. (10) Bannerman 1930:97. (11) Brown et al. 1982:178. (12) G. Niethammer 1938:299. (13) Cramp 1977:323. (14) Dementiev and Gladkov 1951:445. (15) S. Ali and S.D. Ripley 1968:104. (16) LaTouche 1931-34:440. (17) Taczanowski 189193:975. (18) Paz 1987:33. (19) Schroder and Burmeister 1974:5-6. (20) FitzPatrick Inst., Univ. of Cape Town (unpubl. data). (21) Sharpe, 1898:305.

EGG-LAYING Place Europe USSR, Mongolia, NE China, SE Korea Zambia Malawi Zimbabwe Mozambique Botswana N South Africa, Lesotho S South Africa (Cape Prov.)

290

Months Apr-Jun Apr-May May-Jun Jul Jan-Sep (mainly May/Jun) Sep-Oct Jun May-Sep (mainly Jun/Jul) Aug-Sep

Abdim's Stork (Ciconia abdimii) BODY M E A S U R E M E N T S

Wing (mm)

Sex

[n]

Male Fern. Fern. Both Both Both p p

438 [1] 435 [1] — [1] [22] 400-475 [?] 413-425 [?] 430-460 — [1] — [1] — [1]

9

Culmen (mm)

Tarsus (mm)

Tail (mm)

110 105

120 118 — 117-136 115-119 110-130 — — —

180 175 — 167-205 175 170-200 — — —

— 103-127 108-113 100-120 — — —

EGG MEASUREMENTS

Wing-Span (mm) — — — — — — — —

Weight (kg)

Reference

— 1.08 — — — 0.98 0.94 0.93

(5) (5) (4) (1) (2) (3) (4) (4) (4)

Extremes Average Extremes Average Calculated Extremes Extremes Average Extremes Average Extremes Average Extremes Average Singles Singles Singles Singles Singles

Longevity in captivity = 16 years 5+ months (Flower 1925:1399). Longevity in captivity = 2 1 + years (K. Brouwer et al. pers. comm. 1990). References: (1) Bannerman 1930:100. (2) Heuglin 1869-73:1105. (3) Reichenow 1900-01:344. (4) FitzPatrick Inst. Univ. of Cape Town (unpubl. data). (5) Sharpe 1898:293.

No.

Length (mm)

Width (mm)

41 41 23 23 23 4 37 37 11 11 — — — — 1 1 1 1 1

56.2-64.1 59.6 55.0-63.5 58.3 — 55.0-61.0 54.6-64.8 60.4 56.2-62.0 58.3 51.0-56.0 52.1 50.0-58.0 59.6 55.5 58.0 60.8 60.3 —

40.1-45.8 43.3 39.7-47.1 42.5 — 40.0-47.0 40.8-45.3 43.1 40.5-44.0 42.0 39.1-41.1 (37.5?) 40.0-42.0 43.3 41.0 41.5 40.0 39.6 —

Weight (g)

Reference

— — — 58 — — — — — — — — — — — — — 60

(1) (1) (2) (2) (2) (3) (4) (4) (11) (11) (5) (5) (6) (7) (8) (8) (9) (9) (10)

References: (1) Serle 1943 in Brown et al. 1982:181. (2) Schonwetter 1967. (3) Reichenow 1900-01:344. (4) Kahl 1971d. (5) Heuglin 1869-73:1107. (6) Bouet 1955a. (7) Bannerman 1953:173. (8) Bannerman 1930:101. (9) Bigalke 1948. (10) Salvan 1967. (11) Duignan et al. 1988 (K. Brouwer pers. comm. 1990).

EGG-LAYING Months

Place W Africa to Ethiopia, Somalia, N Kenya Equatorial Kenya and Uganda E Eritrea (winter rainfall area) SW Arabia, Yemen

291

Feb-Jul (mainly Apr-May) Jan-May

Jan-Mar Mar-May

Woollynecked Stork (Ciconia episcopus} BODY MEASUREMENTS

Sex

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

149-185 162 — — 143-165 137-157 148 155-171 163 — —

161-178 170 — — — 140-163 157 —

175-205 192 _ _ _ _ _ _ _

_ _ _ _ _ _ _

_ _ _ _ _ __ _

— — —

— 146 137 145 — 146-147 147 — — 148-152 140-157 145 140-171 150 — 125-169 153 137-157 146 — 145-168 141-145 165-176 145-168 [131?] 153

— — — 160 — 153-166 158 — — — 147-160 154 — — — — — — — 150-175 152-180 157-179 152-158 152-190 148 155

_ — —

_ — —

__ — 1.93 2.5

182 193 169 _ 193-195 194 — _ _ — _ _ — _

_ — _ _ — _ _ — _

_ — _ _ — _ __ — _

_ — _ _

_ — _ _

__ — _ _

175-205 199-225 183 158-166 169-190 193 173

1750-1850 -

2.05-2.79 -

Sex

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

[1] [1] [1] [1] [1] [?] [?] [16] [?] [?] [1]

500 463 ~ ~ 454-491 413-438 440-485 450-490 440-470 440

163 _ 143 _ _ _ 117-131 135-168 _ 130-140 140

175 _ 155 _ _

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

180 _ 180 _ _ _ 175-179 180-214 _ 170-180 177

_ _ _ _ _ -

— 1.59 — 1.85 1.59 — — — — — —

(24) (19) (24) (23) (23) (14) (15) (16) (17) (18) (20)

C. e. microscelis

C. e. episcopus

Male M 471-507 Male, average W 488 Male [7] 471-523 Male, average [7] 493 — Male [8] Male [7] 446-500 Male, average [7] 469 Male [8] 440-500 Male, average [8] 478 Male 515 [1] Male 519 [1] ~\/f i Male — Male [1] Male 465 [1] Male 464 [1] Male 497 [1] Male 525 [1] Fern. [3] 460-480 Fern., average [3] 472 Fern. [4] 450-516 Fern., average W 485 — Fern. [2] Fern. [5] 442-462 Fern., average [5] 450 Fem. [10] 435-480 Fern., average [10] 458 Fem. 490 [1] Both [9] 440-490 Both, average [9] 472 Both [5] 475-490 Both average 485 [5] Both [?] 450-515 Both [?] 444-497 Both [?] 470-508 — Both [?] Both P] 444-525 ? 470 [1] P 450 [1]

Reference (1) (1) (2)

(2) (3) (21) (21) (25) (25) (9) (9) /(10) 1 r\\ (10) (13) (13) (22) (22) (1) (1) (2) (2) (3) (21) (21) (25) (25) (22) (25) (25) (4) (4) (5) (6) (7) (8) (22) (11) (12)

Male Fem. Fem. Fem. Fem. Both Both Both Both Both p

_ _ 129-144 140-171 _ 145-150 152

Longevity in captivity = 30+ years (K. Brouwer et al. pers. comm. 1990). References: (1) Riley 1925:29. (2) Abdulali 1968. (3) Medway and Wells 1976:94. (4) Meyer and Wiglesworth 1898:807. (5) Legge 1880:1119. (6) E.C.S. Baker 1929:324. (7) Hackisuka 1931-32:348. (8) Kuroda 1936:540. (9) S. Ali and S.D. Ripley 1968:99. (10) J.H. Riley 1938:36. (11) duPont 1971 :23. (12) Rabor 1977 :26. (13) Rensch 1931. (14) Mackworth-Praed and Grant 1970:53. (15) Heuglin 1869-73:1109. (16) Bannerman1930:98. (17) Clancey 1964a:45. (18) Reichenow 1900-01:348. (19) Britton 1970a. (20) McLachlan and Liversidge 1970:51. (21) Pers. obs. M.P.K. (Zool. Mus., Bogor, Indonesia/8 Mar 68). (22) Delacour and Jabouille 1931: 85-86. (23) FitzPatrick Inst., Univ. of Cape Town (unpubl. data). (24) Sharpe 1898:296. (25) Pers. obs.M.P.K. (British Mus./28 Aug 70).

292

Woollynecked Stork continued EGG M E A S U R E M E N T S

Extremes Average Extremes Average Extremes Average Extremes Extremes Extremes Average Extremes Average Extremes Average Singles Singles Extremes Average Calculated Extremes Average Calculated

No.

Length (mm)

Width (mm)

3 3 100 100 50 50 — — 3 3 3 3 7 7 1 1 4 4 — 120 120 120

58.8-62.3 60.8 57.0-72.3 62.9 57.5-66.5 62.5 60.0-61.0 59.0-64.0 59.3-61.2 60.4 60.0-61.3 60.7 59.3-67.0 62.0 67.0 65.7 60.0-67.0 63.5 — 57.0-72.0 63.0 —

45.0-47.5 46.2 43.6-49.3 47.4 43.8-48.0 45.8 41.0-42.0 47.0-48.0 44.7-45.4 45.0 41.2-42.3 41.9 43.0-45.4 44.4 43.0 43.2 41.2-43.2 42.4 — 43.6-49.3 47.3 —

EGG-LAYING

Weight (g)

— — — — — — — 70-70 70 — — 53-70 59 55 55 — — 59 — — 77

Place

Reference

Months

C. e. episcopus

N India S India Sri Lanka Burma Malaysia (Tanjong Badak) Philippines Indonesia (Flores)

(1) (1) (2) (2) (3) (3) W (5) (6) (6) (7) (7) (9) (9) (8) (8) (10) (10) (10) (11) (11) (11)

May-Sep Nov-Mar Jan-Apr Feb-Mar/May-Jul/Sep Mar-Apr Jan Feb-Mar/Jun/Aug/Nov

C. e. microscelis

Liberia Sierra Leone N Nigeria Chad Sudan (Upper Nile) Ethiopia Uganda Zambia Zimbabwe N South Africa (Zululand, Natal)

References: (1) Inglis 1898. (2) E.C.S. Baker 1935:443. (3) Hume and Gates 1890:269. (4) Bannerman 1953:172. (5) Bouet 1955a. (6) Anthony 1978b. (7) Pitman 1931a. (8) Anthony 1977. (9) Maclean 1985:67. (10) Brown et al. 1982:182. (11) Schonwetter 1967.

293

Sep-Oct Jan-Feb Jan Jul-Sep Get J«l Nov-Feb Aug-Sep Aug-Nov Sep-Nov

Storm's Stork (Ciconia stormi) BODY M E A S U R E M E N T S

Wing (mm)

Sex

W

Male Male Male Fern. Fern. Fern. Fern. Both Both, average p P ?

405 [1] 398 [1] — [1] 375 [1] 390 [1] 404 [1] 420 [1] [5] 380-470 402 [5] — [1] 395 [1] 395 [1]

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

163 — 137 143 — — 147 120-135 131 168 — —

163 — 136 138 138 — 150 — — — 150 150

155 136 _

_

_

133 143 143 180 _ _ _ _

-

-

_ _ _ _

_ _ _ _

—

—

—

Reference (1) (2) (3) (1) (1) (2) W (6) (6) (4) (5) (5)

References: (1) Ogilvie-Grant 1903. (2) Mayr 1938. (3) Pers. obs. M.P.K. (Zool. Mus., Bogor, Indonesia/8 Mar 68). (4) Medway and Wells 1976:95. (5) Medway and Wells 1964:156. (6) Pers.obs. M.P.K. (British Mus./ 28Aug70).

EGG M E A S U R E M E N T S

No. Single

Length (mm)

Width (mm)

60.2

41.9

Weight (g)

(1)

Reference: (1) F. Danielsen et al. pers. comm. 30 Apr 1990.

EGG-LAYING

Place

Months

Sep Apr-May

S Thailand SE Sumatra

294

Reference

Maguari Stork (Ciconia maguari) BODY M E A S U R E M E N T S

Wing (mm)

Sex

[n]

Male Male Male, average Male, average Male Male Male Male Fern. Fern. Fern., average Fern., agerage Fern. Fern. 6oth Both, average ? P

[8] [6] 531-614 562 [6] — [9] — [1] — [1] — [1] — [1] — [6] [5] 535-566 553 [5] — [5] — [1] — [1] [2] 535-603 569 [2] 590 [1] 540 [1]

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

225-255 203-245 231 241 235 230 235 230 205-227 205-229 219 213 200 205 207-227 217 230 —

265-275 — 266 — 260 260 260 270 — — — — 240 235 — — 280 —

__ _ 200 _ — — — — — — — — — — 196-208 202 250 —

_ _ _ — — — — — — — — — — —

_ — — 4.2 4.0 4.2 4.25 4.35 — — — 3.8 3.4 3.5 — — — —

Reference

(i)

(2) (2) (6) (1) (1) (1) (1) (1) (2) (2) (6) (1) (1) (3) (3) (4) (5)

Longevity in captivity = 20+ years (K. Brouwer et al. pers. comm. 1990). References: (1) B.T. Thomas 1985:923. (2) Blake 1977:188. (3) Goodalletal. 1951:101. (4) Zotta and Fonseca 1937. (5) Chubb in Haverschmidt 1968:27. (6) C.E. King 1988:162.

EGG-LAYING

EGG MEASUREMENTS

Extremes Average Extremes Average Calculated Single Single Single Single Single Single

No.

Length (mm)

Width (mm)

Weight (g)

54 54 18 18 18 1 1 1 1 1 1

69.1-85.0 75.2 70.0-84.0 75.1 — 77.4 76.3 75.6 74.0 70.4 71.5

46.0-57.0 52.6 46.0-57.0 53.4 — 56.2 56.2 51.9 52.0 50.0 51.5

100-140 116 — — 118 — — — — — —

Place Reference

Venezuela (Llanos) E Brazil (Isla Mexiana) NE Argentina (Corrientes Prov.) E-central Argentina (Buenos Aires Prov.)

(i) (i)

(2) (2) (2) (3) (3) (4) (5) (6) (6)

References: (1) B.T. Thomas 1984. (2) Schonwetter 1967. (3) Kahl 1971e. (4) Gibson 1880. (5) Gibson 1919. (6) Haedo-Rossi 1969. 295

Months May-Nov (mainly Jul-Sep) Aug-Sep Jun-Aug Jul-Oct

White Stork (Ciconia ciconid) BODY MEASUREMENTS

Sex

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

530-630 600-620 — 540-610 580 556-598 576 — — — 530-590 560-580 —

150-190 180-200 — 169-206 — 157-198 179 — — — 140-170 150-175 —

195-240 — — — — 191-230 214 — — — — — —

215-240 240-260 — — — 221-268 247 — — — — — —

— — — — — — — — — — — — —

— — 2.9-4.4 — — — — 3.571 3.97 4.3 — — 2.7-3.9

(1) (2) (3) (4) (4) (22) (22) (3) (5) (14) (1) (2) (3)

543-582 558 — — — 585-605 542-580 550-600 540-610 — 530-630 — 565 — —

155-180 164 — — — — 158-191 175-200 169-206 — — — 176 —

184-211 197 — — — — 213-225 190-220 — — — —

218-256 237 — — — — 218-251 210-240 — — — — 227 — —

— — — — 1550-1650 — — — — up to 2000 up to 2200 — — — 'almost 2.5m'

Sex

&

C. c. ciconia ~^ Male Male Male Male Male, average Male Male, average Male, average Male, average Male Fern. Fern. Fern. 1 trem., average Fern Fern., average Fern., average Fern. Both Both Both Both Both Both Both Both Both, average Both, average Both, average

[9] [?] [41] [9] [9] [18] [18] [41] [12] [1] [?] [?] [27] ro7i U'J [15] [15] [12] [1] [?]

[6] [6] [?] [9] [?] [?] [?]

[12]

[?] [?]

—

220 — —

0 OO^ o.azO

— —

3.52 3.5 — 3.5-4 — — — — 2.3-4.4 2.6-4.5 — 3.5-4 —

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

— 213-247 234 211-234 220 -

-

-

-

-

-

-

-

Reference

C. c. asiatica

Male Male, average Male Male, average Male Fern. Fern., average Both Both Both Both Both Both, average

/Q\ W)

a

[14] 580-630 184-235 [14] 610 [18] 581-615 188-223 [18] 596 204 [1] 653 220 [9] 548-596 178-196 [9] 577 187 [?] 590-615 195-215 [?] 600-620 198-222 [ 9 ] 550-640 [14] 580-630 184-235 [16] 188-210 [9] 589 -

(4) (4) (22) (22) (17) (22) (22) (15) (15) (16) (13) (18) (16)

Longevity, oldest ringed bird = 29 years 3 months (Rydzewski 1973). Longevity, free-flying semi-captive = 33 years 8 months (Bloesch 1986). c Longevity in captivity = 33 years 3 months (Flower 1938). d Longevity in captivity = 35-1- years (K. Brouwer et al. pers. comm. 1990). References: (1) Witherby et al. 1939:115. (2) Hartert 1912-21:1214. (3) Bauer and Glutz von Blotzheim 1966:390. (4) Vaurie 1965:84. (5) Hornberger 1967:7. (6) Cramp 1977:328. (7) Dementiev and Gladkov 1951:451. (8) Cramp 1977:335. (9) Reichenow 1900-01:346. (10) O. Heinroth and M. Heinroth 1924-31:133. (11) O. Heinroth 1922:185-186. (12) Mell 1951:24. (13) J. Niethammer 1967. (14) Milstein 1966 .(15) Hartert 1912-21:1215. (16) Dementiev and Gladkov 1951:453. (17) Mendelssohn 1975a. (18) G. Niethammer 1972b:208. (19) Schierer 1972. (20) Grzimeketal. 1968:208. (21) Creutz 1988:8. (22) S.W7. Winter pers. comm. 1990.

(22) (22) (5) (11) (6) (7) (8) (9) (13) (19) (20) (21) (8) (10) (12)

b

296

White Stork continued EGG MEASUREMENTS

Extremes Average Calculated Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Extremes Extremes Average Extremes Average Single

EGG-LAYING

No.

Length (mm)

Width (mm)

Weight (g)

Reference

150 150 — 103 103 27 27 20 20 80 80 16 16 120 120 140 13 — 37 37 1

65.0-81.5 73.0 — 65.6-81.5 73.2 68.6-77.6 77.3 65.6-81.5 70.7 61.6-78.6 71.3 71.0-78.2 73.8 65.5-81.5 73.2 64.0-79.0 — — 66.6-88.6 73.2 —

46.5-56.0 51.8 — 46.5-55.7 51.8 49.0-55.9 52.5 46.0-55.7 50.7 47.0-54.5 51.5 47.9-57.0 53.8 46.5-55.7 [58.8?] 48.0-53.2 — — 48.5-57.1 52.3 —

— 110 — — 95.5-129.3 111.5 — — — — — — — — — 95.5-129.3 120 — — 118

(1) (1) (1) (2) (2) (3) (3) (3) (3) (3) (3) W (4) (5) (5) (6) (7) (8) (10) (10) (9)

Place Spain, Portugal N Europe W and central Asia NW Africa S South Africa

References: (1) Schonwetter 1967. (2) Witherby et al. 1939:114. (3) Bauer and Glutz von Blotzheim 1966:403. (4) Dementiev and Gladkov 1951:449. (5) E.C.S. Baker 1929:322. (6) Haverschmidt 1949:40. (7) G. Niethammer 1938:297. (8) O. Heinroth and M. Heinroth 1924-31:133. (9) Mell, 1951:25. (10) Adam Mrugasiewicz pers. comm. 6 Apr 69.

297

Months Feb-May Apr-Jun Apr-Jun Feb-Apr Sep-Nov

Oriental White Stork (Ciconia boy eland) BODY M E A S U R E M E N T S

Sex

M

Male, average Male Male, average Male Male, average Male Male, average Male Male, average Male Male, average Male, average Male, average Male Male, average Male Male Fern., average Fern. Fern., average Fern. Fern., average Fern. Fern., average Fern. Fern., average Fern. Fern., average Fern., average Fern., average Fern. Fern., average Both Both

[5] [11] [11] [8] [8] [14] [14] [2] [2] [2]

[?]

?

[9] [31] [16] [16] [1] [1] [7] [7] [7] [7] [7] [20] [20] [4] [4] [3] [3] [26] [8] [15] [15] [?] [8]

EGG MEASUREMENTS

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

616-670 654 580-678 645 592-680 640 — — 650-690 670 — — — — 665 590 — 598-655 615 540-658 615 576-635 606 — — 615-705 653 — — — — 620-670 620-670

266 235-279 257 223-276 243 207-282 256 — — — 260 — 255 226-276 248 — 227 239 222-288 243 210-245 224 203-248 228 — — 231-288 255 232 — 210-245 224 195-222 200-260

301 256-300 277 280-315 296 250-310 282 — — 290-305 298

245-307 271 250-285 264 — — — — 238-260 249

— _ -

— — 4.5-5.5 5.22 — — 4.13-5.5 4.82 — _

— 285 — — 265 241 271 241-266 253 245-278 265 215-282 255 — — 245-265 252 254 — — — — —

— — — — 245 223 — 216-284 242 220-265 252 — — — — 230-256 241 — — — — — —

_ — _ _ _

5.1 _

— — — — _ — _ — — —

— — — — _ — _ 3.7-5.0 4.35 — — 2.89-4.5 3.75 —

_ —

_ —

— — — —

4.1 — — — —

Reference

No.

Extremes Average Calculated Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Average

(1) (2) (2) (3) (3) (4) (4) (4) (4) (6) (6) (8) (11) (12) (12) (9) (10) (1) (2) (2) (3) (3) (4) (4) (4) (4) (6) (6) (H) (H) (12) (12) (5) (7)

10 10 10 24 24 4 4 4 4 4 4 10 10

— — —

Length (mm)

Width (mm)

68-77 75.0 — 72-80.3 76.7 76-79 77.5 74-79 77.3 78.1-81 79.6 72.7-80.2 78.9 77-80 75.2 77.0

53-58 54.5 — 51.5-58.3 56.3 58-60 59 56-58 56.9 57-62.3 58.4 52.1-55.6 54.6 54-56 58.1 58.0

Weight (g) __ 124 — — — 147.5 — — — — — — — — —

Reference (1) (1) (1) (2) (2) (3) (3) (4) (4) (5) (5) (8) (8) (6) (7) (9)

References: (1) Schonwetter 1967. (2) Winter 1982:78. (3) Fei et al. 1991. (4) Leonovich and Nikolyevsky 1976. (5) Pankin and Neufeldt 1976. (6) Austin and Kuroda 1953:333. (7) E.C.S. Baker 1929:323. (8) S.V. Winter pers. comm. 1990. (9) Taczanowski 1891-93:974.

EGG-LAYING Place Korea (formerly) Japan (formerly) SE USSR/NE China

Longevity in captivity = 31 years (Schtiz 1965b). Longevity in captivity = 48+ years (C.E. King and K. Brouwer et al. pers. comm. 1990). Body temperature of large nestlings; 6 readings = 40.5-41.0°C (Fei et al. 1991). References: (1) K. Archibald and B. Schmitt 1991. (2) S.W. Winter pers. comm. 1990. (3) K. Murata, unpubl. data (via K. Archibald pers. comm.). (4) K. Archibald pers. comm. 1990. (5) S. Ali and S.D. Ripley 1968:102. (6) Taczanowski 1891-93:974. (7) Hartert 1912-21:1215. (8) C.E. King pers. comm. 1990. (9) LaTouche 193134:438. (10) T.H. Shaw 1936:121. (11) C.E. King 1988:162. (12) Murata et al. 1988.

298

Months Mar Mar-May Apr-May

Blacknecked Stork (Ephippiorhynchus asiaticus] BODY M E A S U R E M E N T S

Sex

[n]

Male Fern. Both Both, average Both p ?

[1] [1]

Wing (mm)

563 538 [4] 488-625 566 [4] 565-645 573 —

r?i [i] [i]

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

300 288 288-300 297 298-324 320 —

300 275 295-338 313 300-333 323 —

213 213 235-250 242 257-281 200 —

— — 2125 — — c. 2000

— — — _ — —

(3) (3) (1) (1) (2) (4) (5)

Longevity in captivity = 34+ years (K. Brouwer et al. pers. comm. 1990). References: (1) Legge 1880:1116. (2) E.C.S. Baker 1929:326. (3) Sharpe 1898:312. (4) Rand and Gilliard 1967:48. (5) Pizzey 1980:62.

EGG MEASUREMENTS

Extremes Average Calculated Extremes Average Calculated Extremes Average Extremes Average Extremes

No.

Length (mm)

Width (mm)

50 50 50 11 11 11 45 45 30 30 p

67.0-76.0 72.0 — 71.0-76.0 74.5 — 66.3-78.3 72.8 67.9-74.9 69.5 70.3-71.9

51.0-57.0 53.7 — 52.0-55.0 53.8 — 49.5-57.5 53.0 51.0-55.2 53.2 52.4-54.0

Weight (g)

Reference

— 115

(i) (i) (i)

— — 120 — — — — —

(2) (2) (2) (3) (3) (4) (4) (5)

References: (1) Schonwetter 1967 (India). (2) Schonwetter 1967 (Australia). (3) Hume and Gates 1890:268. (4) E.C.S. Baker 1929:327. (5) Medway and Wells 1976:96.

EGG-LAYING

Months

Place N India Burma (S Pegu) Sri Lanka N Australia/SE Papua New Guinea SE Australia (coastal NSW)

299

Sep-Dec Dec-Jan Early months of year Mar-Jun Aug-Nov

Saddlebill Stork (Ephippiorhynchus senegalensis) BODY M E A S U R E M E N T S

Sex

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

Male Fern. Fern. Fern. Both Both Both Both Both Both

[1] [1] [1] [1] [?] [?] [?] [10] [4] [?]

— , — 675 600-670 590-625 630-700 600-670 620-670 650-700

— — 315 273-334 — — 273-334 282-334 320-360

— — 375 311-365 — — 311-365 293-353 330-360

— — 270 250-288 —

— — — up to 2700 — — — — —

7.3 5.0 6.0 — — — — — — —

(3) (3) (7) (8) (1) (2) (3) (4) (5) (6)

— 260-288 257-272 270-300

Longevity in captivity = 19+ years (Flower 1925), 36 years (Schiiz et al. 1955). References: (1) Brown etal. 1982:185. (2) Mackworth-Praed and Grant 1973:68. (3) F.J.Jackson 1938:76. (4) Bannerman 1930:104. (5) McLachlan and Liversidge 1970:50. (6) Reichenow 1900-01:342. (7) O. Heinroth 1922:185. (8) Sharpe 1898:314.

EGG MEASUREMENTS

Extremes Average Extremes Average Calculated Extremes Average Extremes Average Single Single Single Single

No.

Length (mm)

Width (mm)

12 12 6 6 6 9 9 — — 1 1 1 1

79.0-86.3 82.5 75.6-81.3 80.0 — 75.6-81.3 79.1 76.0-80.5 77.0 78.0 78.5 77.3 76.1

55.3-62.0 58.5 56.7-58.0 57.0 — 56.0-58.0 56.7 56.0-57.5 57.0 57.0 56.2 56.0 56.2

EGG-LAYING Weight (g) — — — 146 — — — — — — — —

Place

Reference

Senegal (Boghe) N Ivory Coast Chad (L. Agau) Ethiopia S Sudan N Uganda W Uganda Kenya, Tanzania Zambia Zimbabwe Malawi N South Africa (Zululand, E Transvaal)

(1) (1)

(2) (2) (2) (3) (3) (4) (5) (6) (7) (7) (7)

References: (1) Pitman 1965. (2) Schonwetter 1967. (3) Maclean 1985:69. (4) Chapin 1932a:463. (5) Bouet 1955a. (6) Mackworth-Praed and Grant 1973:69. (7) McLachlan and Liversidge 1970:50. Two eggs (in the British Museum collection), which actually belong to the Secretarybird (Sagittarius serpentarius), are within the size-range of the Saddlebill and were formerly misidentified as such; they may have been included in some of the above measurements (Pitman, 1965).

300

Months Jan-Feb Sep-Oct Dec—Jan Oct-Nov Sep-Apr Nov-Jan May-Jul Dec-Jun, Sep-Oct (mainly Mar-Apr) Jan-Aug, Nov-Dec (mainly Mar-Apr) Jan-Apr, Jul (mainly Feb-Mar) Jan Mar-Jul

Jabiru Stork (Jabiru mycterid) BODY M E A S U R E M E N T S

Sex

[n]

Wing (mm)

Male [10] 625-715 Male, average [10] 660 Male [4] 686-730 Male, average [4] 707 Male [?] 620-686 Male, average [?] 647 Male [1] Male [1] Male [1] Fern. [8] 585-665 Fern., average [8] 620 Fern. [2] Fern., average [2] 648 Fern. [?] 610-622 Fern, average [?] 614 Fern. [1] Fern. [1] p [1] 650 p [1] 630 p [1] p [1] -

Culmen (mm)

Tarsus (mm)

295-343 324 318-345 329 280-354 315 310 300 330 284-328 299 267-292 280 248-320 295 — 305 350 — — 300

285-330 308 — — 292-338 314 310 320 320 — — — — 286-305 297 — 295 385 — — —

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

— — — 205-245

— — — -

— 6.3-8.1 7.03 — — 6.7 7.7 — — — 4.3-5.9 5.07 — — — — — — 5.47 —

(1) (1) (2) (2) (8) (8) (3) (3) (3) (1) (1) (2) (2) (8) (8) (8) (3) (4) (5) (6) (7)

236 — — — — — — — 198-241 213 — 250 — — -

— — — — — — — 2325 — — — 2350

Longevity in captivity = 16 years (Flower 1925). Rectal temperature = 39.1-40.5°C (Benedict and Fox 1927). References: (1) Blake 1977:189. (2) Shannon 1987. (3) B.T. Thomas 1985:923. (4) Zotta and Fonseca 1937. (5) Chubb in Haverschmidt 1968:28. (6) Benedict and Fox 1927. (7) Bent 1926:71. (8) Oberholser and Kincaid 1974:123-124.

EGG M E A S U R E M E N T S

Extremes Average Average

No.

Length (mm)

Width (mm)

8 8 —

71.5-75.3 73.4 73.2

55.0-60.5 58.2 58.2

EGG-LAYING

Place

Weight (g)

Reference

— —

(1) (1) (2)

SE Mexico, Belize Venezuela (Llanos) Colombia (Rio San Jorge) Guyana, Surinam E Brazil (Isla Mexiana) SW Brazil (Mato Grosso) NE Argentina (Corrientes, Chaco)

References: (1) Hagmann 1906. (2) Oberholser and Kincaid 1974:123. The egg-measurements given in F.P. Penard and A.P. Penard (1908), and repeated in Schonwetter (1967), Haverschmidt (1968:28) and Kahl (1971e), are much too large and are probably an error, referring instead to the flamingo (Phoenicopterus ruber ruber) (Spaans 1975c).

Months Dec-Jan Aug-Nov Oct-Nov Aug-Oct Jul-Aug Sep-Nov Aug-Oct

The breeding dates given (May/June) for Surinam by F.P. Penard and A.P. Penard (1908) and repeated by Haverschmidt (1955b:32, 1968:28) and Kahl (1971e), are probably an error (F. Haverschmidt pers. comm. 27 Jun 71, A.L. Spaans pers. comm. 5 Jul 1971, Spaans 1976b).

301

Lesser Adjutant Stork (Leptoptilosjavanicus) BODY

Wing (mm)

Sex

W

Male Male Male Fern., average Fern. Both Both, average

[1] — [1] 575 [1] — [?] [11 — [?] 580-660 — [?]

MEASUREMENTS

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

— 258 — 263 260-305 —

— 225

— 233 — 258 230-253 —

— — — — — —

4.09 5.23 — 4.5 — — 5.71

(1) (1) (5) (2) (5) (3) (4)

— 245 228-268 —

Longevity in captivity = 30+ years (K. Brouwer et al. pers. comm. 1990). Rectal temperature = 38.8-40.2°C (Benedict and Fox 1927). References: (1) J.H. Riley 1938. (2) O. Heinroth 1922. (3) E.C.S. Baker 1929:330. (4) Benedict and Fox 1927. (5)Sharpe 1898:318.

EGG M E A S U R E M E N T S

No.

Extremes Average Extremes Average Average Extremes Average Extremes Average Average Extremes Average Extremes Average

50 50 2 — 3 7 7 9 9 — 66 66 — —

Length (mm)

Width (mm)

58.8-86.2 76.4 70.5-71.5 73 73 61.9-78.0 70.23 70.6-80.3 75 — 60.0-86.0 75.7 — 72.3

49.0-62.0 55.3 51.8-52.8 53.5 55 49.9-53.0 51.23 52.2-57.3 54.2 — 50.2-58.8 54.8 — 53.1

EGG-LAYING

Weight (g) — — — — — — — — 113 — 127 [calc.] 120-130 —

Place

Reference

NE India, Burma S India, Sri Lanka Sri Lanka Malaysia (Johore) Malaysia (Selangor) Malay Peninsula Sumatra (Belitung) EJava WJava Samarahan Delta, Borneo

(i) (i)

(2) (3) W (5) (5)

(6) (6) (7)

(8) (8) (9) (9)

References: (1) E.C.S. Baker 1935:449. (2) Hume and Gates 1890:265. (3) Henry 1971. (4) Hoogerwerf and Rengers Hora Siccama 1937. (5) Hoogerwerf 1949b: 18-19. (6) Hellebrekers and Hoogerwerf 1967a. (7) O. Heinroth 1922. (8) Schonwetter 1967. (9) P. Saikia (pers. comm.).

302

Months

Jul-Jan (mainly Jul-Aug) Feb-May Sep (sometimes) Feb-Apr Mar-Jun

Jan-Apr May Feb-Mar May-Aug May

Greater Adjutant Stork (Leptoptilos dubius] BODY MEASUREMENTS

Wing (mm)

Sex

W

Both p

[?] 800-820 800 [1]

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

320-345 300

320-330 325

310-335 313

-

Weight (kg) —

Longevity in captivity = 43+ years (K. Brouwer et al. pers. comm. 1990). References: (1) E.C.S. Baker 1929:328. (2) Sharpe 1898:317.

EGG M E A S U R E M E N T S

Extremes Average Extremes Extremes Average Calculated Average

No.

Length (mm)

Width (mm)

Weight (g)

50 50 — 60 60 60 —

70.1-82.8 77.3 69-81.5 71-83 78.5 — 78.7

51.5-64.7 57.5 50.5-63.8 53-64.7 57.5 — 59.0

— — — — 142 160

Reference

(i) (i)

(2) (3) (3) (3) (4)

References: (1) E.C.S. Baker 1929:329. (2) Hume and Gates 1890:264. (3) Schonwetter 1967. (4) P. Saikia (pers. comm.).

EGG-LAYING Place

Months

Assam, India/Burma

Sep-Jan

303

Reference (1) (2)

Marabou Stork (Leptoptilos crumeniferus] BODY M E A S U R E M E N T S

Sex

Male Male Male Male Male, average Male, average Male, average Male, average Male, average Male, average Male Fern. Fern. Fern. Fern. Fern., average Fern., average Fern., average Fern., average Fern., average Fern., average Fern. Fern. Fern. Both Both Both Both Both, average P

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

[?] 705-794

279-346

266-319

278-358

— _ 750-765 745 — — — 745 — — 631-709 — — 620 678 — — — 679 — — — — 643-656 650-700 620-720 650-700 673 725

— — — — — — — 311 — — 241-314 — — — — — — — 278 — — — — 225-281 — 226-280 240-280 244 270

266-319 — 290-320 — 293 — — 293 — — — 245-283 — 205 — 266 — — 266 — — — — 225-248 — 215-270 250-270 242 275

— — — 323 — — — 323 — — 240-302 — — — 282 — — — 282 — — — — 244-256 — 250-295 250-300 276 295

[?] [37] [2] PI [?] [26] [8] [37] [9] [1] [?] [?] [22] [2] [?] [?] [14] [6] [22] [7] [1] [1] [1] [?] [?] [10] [?] [10] [1]

EGG M E A S U R E M E N T S

Wing-Span (mm)

Weight (kg)

a

2870 (max.) — — — 2630 — — — — 2630 — — — — — 2470 — — — — 2470 — — — — — — 2540 — —

Reference Extremes Average Calculated Average Extremes Extremes Extremes Average Extremes Average Extremes Average Extremes Extremes Average Single Extremes Average Calculated

(1) — 5.6-8.9 5.2-6.0 7.1 — 7.0 7.3 7.06 — 7.22 4.0-6.8 — 4.0-6.8 4.3-4.9 5.7 — 5.7 5.8 5.66 — 6.12 6.7 4.82 — 7.4 — — — —

(2) (3) (5) (1) (2) (4) (4) (3) (3) (11) (1) (2) (3) (5) (1) (2) (4) (4) (3) (3) (5) (11) (11) (6) (7) (8) (9) (8) (10)

No.

Length (mm)

Width (mm)

22 22 — — — — 18 18 40 40 18 18 2 3 6 1 14 14 14

71-84.5 79.1 — — 80-84.5 77-84.5 72.5-86.2 79.2 71-86.2 79.1 72.5-86.2 79.2 81.5-84 71-76.5 80 73 73-84 79.5 —

50-62 55.6 — — 54-57.5 54-60 52.3-58.5 56.3 50-62 55.9 52.3-58.5 56.3 54-57.5 50-53.5 55 61 52-62 56.0 —

Weight

(g)

Reference

— 138 140 140-145 _ — — — c. 138 — c. 138 — — _ — — — 138

(1) (1) (1) (2) (3) (4) (5) (5) (6) (6) (7) (7) (8) (8) (9) (10) (11) (11) (11)

References: (1) Brown et al. 1982:190. (2) O. Heinroth 1922. (3) Reichenow 1900-01:341. (4) Chapin 1932a:468. (5) W. Fraser 1971. (6) Maclean 1985:70. (7) MacLachlan and Liversidge 1970:47. (8) M.E.W. North 1943. (9) Pomeroy 1977b. (10) FJ. Jackson 1938:80. (11) Schonwetter 1967.

EGG-LAYING Place

a

Reputed spans of 4 m or over can be discounted (Brown et al. 1982:187). Longevity in captivity = 41+ years (K. Brouwer et al. pers. comm. 1990). Resting body temperature = 39.2-41°C (Bamford and Maloiy 1980). Mean resting heart-rate = 90.6 ± 1.8 beats/min (Bamford and Maloiy 1980). References: (1) Brown et al. 1982:187. (2) Maclean 1985:69. (3) Pomeroy 1977b. (4) Pomeroy 1973. (5) K.M. Schneider 1952. (6) Heuglin 1869-73:1115. (7) F.J.Jackson 1938:77. (8) MacLachlan and Liversidge 1970:46. (9) Reichenow 1900-01:338. (10) Sharpe 1898:320. (11) FitzPatrick Inst., Univ. of Cape Town (unpubl. data).

N South Africa (Transvaal) NE Swaziland Zimbabwe N Botswana S Tanzania, N Zambia, Malawi N Tanzania, SE Kenya NE, W Kenya Uganda, E Zaire S Sudan S Ethiopia, Somalia N Nigeria S Chad Gambia

304

Months May Jul May-Sep Jul-Sep May-Sep Jul-Sep Sep-Jan Oct-Jan Sep-Jan Oct-Nov Aug-Mar Nov-Jan Dec-Jan

Shoebill (Balaeniceps rex) BODY M E A S U R E M E N T S

Sex

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Male Male Male Fern. Fern., average Fern. Fern. Fern. Both Both Both Both Both, average Both, average ? p p

[1] [1] [1] [3] [3] [1] [1] [1] [?] [?] [4] [?] [?] [?] [1] [1] [1]

780 705 710 — — 655 667 700 588-700 655-685 660-700 655-780 707 700 650 721 730

191 213 238 — — 208

245 246 255 — — 217 — — 221-244 — 254-255 — — 250 250 — —

258 281 284

— — 188-200 — 228-235 — — 230 203 — 245

Wing-Span (mm)

Weight (kg)

— — — — — — 2605 — — — — — — — — 2620

— 6.70 5.4-5.9 5.57 4.36 — — — — — __

— — 276 _ — 244-260 — 279-305 — — 280 250 — 250

— — — — —

Reference (7) (8) (4) (1) (1) (4) (8) (9) (2) (3) (4) (5) (5) (6) (10) (8) (4)

Longevity in captivity = 36 years (Flower 1938:220, Bourliere 1946). References: (1) H.G. Klos 1985. (2) Heuglin 1869-73:1096. (3) Mackworth-Praed and Grant 1973:62. (4) W. Fischer 1970:13. (5) Maclean 1985:63. (6) Reichenow 1900-01:357. (7) Bannerman 1930:87. (8) Chapin 1932a:449. (9) FJ. Jackson 1938:63. (10) Sharpe 1898:287.

EGG

Extremes Average Calculated Extremes

MEASUREMENTS

No.

Length (mm)

Width (mm)

10 10 10 3

80.1-90.0 85.7 — 81.0-82.0

56.9-61.3 59.4 — 60.0-61.0

EGG-LAYING

Weight (g)

Reference

— 164 —

(1) (1) (1) (2)

Place S Sudan Uganda (L. Kyoga, Rwenzama Swamp) SE Zaire (L. Kabamba) NE Zambia (Bangweulu)

References: (1) Schonwetter 1967. (2) F.J.Jackson 1938:66.

305

Months Sep-Feb (mostly Nov-Jan) Mar-Jun Aug-Sep Apr-Jul

American White Ibis (Eudocimus ruber) BODY M E A S U R E M E N T S

Sex

[n]

Male Fern. Male Fern. Male, average

[6] [6] [5] [5] [6]

(ruber}

Fern., average Male, average (albus) Fern., average ? (ruber) ? (albus) Male Male, average Male Fern. Fern., average Male Fern.

[6] [5]

Wing (mm)

Culmen (mm)

Tarsus (mm)

266-289 156-169 — 251-269 123-131 279-285 145.4-163.0 90.3-102.5 260-268 118.5-130.0 83.8-88.0 — 282.3 163.2 260.6 282.6

265 [5] — 254 — 279 [7] 205-295 — 277.3 310 [1] [9] 263-278 P] 271.5 [2] 258-263 [2] 258-263

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

94.97 84-95 107.1-119.8 93.3-104.8 95.6

— — — —

— — — —

(1) (1) (1) (1) (1)

127.9 155.1

— 95.9

90.2 121.1

— —

— —

(1) (1)

125.6 110 162 136-164 153.3 153 119-130 124 157-168 120

85.5 — 89 — — — — — — —

98.6 114 114 — — — — — — —

— — — — _

— — — — _

— — —

— 0.772-0.797 — 0.935 —

(1) (2) (3) (4) (4) (4) (4) (4) (5) (5)

References: (1) Blake 1977:199. (2) D.G. Elliot 1877. (3) D.G. Elliot 1877. (4) Palmer 1962:524. (5) Palmer 1962:530.

EGG-LAYING

EGG M E A S U R E M E N T S

No.

Length (mm)

Width (mm)

20

57.53

38.65

Weight (g)

Place Reference S Florida Carolinas Venezuela (Llanos) Trinidad Surinam French Guiana Brazil

(1)

Reference: (1) Palmer 1962:528.

Months Mar-Apr, or later up to Sep Apr-May or later Jun/Jul onwards Apr-Sep Apr-Aug Apr-Jun Mar-Sep

Nesting is highly variable, depending on water conditions; on the South American coast it is mostly during the wet season. Elsewhere nesting is either during rising or falling water levels.

306

Barefaced Ibis (Phimosus infuscatus) BODY MEASUREMENTS

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

[8] 270-292 [8] 281.3 [8] 252-275 [8] 265.2

114-127 120.2 106-121 108.8

-

122-129 125.7 110-124 114.1

-

-

(1) (1) (1) (1)

Male [10] 273-299 Male, average [9] 285 Fern. [8] 251-295 Fern., average [8] 264.6

104-136 127.5 101-112 108.9

-

H9_134 124.2 105-130 117.7

_ -

_ -

-

-

(1) (1) (1) (1)

104-124 113.6

-

116-131 121.7

-

-

(1) (1)

Sex

[n]

Wing (mm)

P. i. berlepschi

Male Male, average Fern. Fern. , average P. i. nudifrons

P. i. infuscatus

Male [10] 270-289 Male, average [10] 282.1 Reference: (1) Blake 1977:198.

EGG MEASUREMENTS

No. P. i. nudifrons P. L berlepschi

Length (mm)

Width (mm)

Weight (g)

Reference

45.0 48.4

33 32

28 27.5

(1) (1)

References: (1) Schonwetter 1967.

EGG-LAYING

Months

Place

Venezuela (Llanos) Brazil

May-Oct Oct-Nov

Nesting is prolonged and continuous throughout the rainy season.

307

Glossy Ibis (Plegadis falcinellus] EGG M E A S U R E M E N T S

BODY M E A S U R E M E N T S

Sex

[n]

Wing (mm)

Male [8] 275-294 288.3 Male, average [8] Fern. [10] 252-275 Fern., average [10] 262.2 INDIA Both EUROPE Male Fern. AUSTRALIA Male Fern. Fern., average Fern., average Male, average Male EUROPE Male, average Fern. Fern., average ? Fern.

—

248-298

Culmen (mm)

Tarsus (mm)

Tail (mm)

127-129 131 98-114 104.2

99.2-112.0 103.1 78-88 81,8

98-108 102.3 90-100 93.6

99-144 — —

85-100 _

— —

290-306 264-277

—

275-294 126.3-140.1 96.8-107.7 97.8-112.7 67.5-86.3 79.4 267 106.3 259.8 — — — — 286 280-306 126-141 101-113

—

[5] 265-275

[5]

[5] [5] [7]

297 [7] 267-281 273 [7] 257-304 — 250-275 [?]

[7]

132 106-114 110 114-136 100-135

107 82-90 86 73-111 —

Wing-Span (mm)

Weight (kg)

94-106 — —

93-107 96-106 98.8 — — 96-111 106 90-99 94 — —

Reference

No.

Length (mm)

Width (mm)

Weight (g)

Reference

(1) (1) (1) (1)

133 13 — 100 — 13

45-59 47.5-57.6 49.3-56.1 52.18 54 48.9-54.7

33.40 31.5-38.2 33.8-37.6 36.9 35 35.5-37.3

38 — — — — —

(2) (2) (3)

(2)

-

0.750

(2) (2)

-

0.485-0.970

(3) (3) (4) (4) (5)

(i)

(4) (5)

References: (1) Brown etal. 1982:194. (2) Marchant and Higgins 1990:1075. (3) Baker in S. Ali and S.D. Ripley 1968:115. (4) Mackworth-Praed and Grant 1973: Series 1, Vol. 78. (5) McLachlan and Liversidge 1970:P56.

(3)

EGG-LAYING Months

Place India Southeast Asia, Burma (formerly) Australia (NSW Balvanald)a Australia (Cowal)a Algeria, Morocco, Egypt Mali Kenya Tanzania Zambia Namibia Zimbabwe South Africa (SW Cape) South Africa (Transvaal) NEUSA Florida, USA West Indies Venezuela Trinidad & Tobago Europe

(5) (5) (5) (6) (7)

References: (1) Blake 1977:201. (2) S. Ali and S.D. Ripley 1968:115. (3) Marchant and Higgins 1990:1077. (4) Amadonand Woolfenden 1952 in Marchant and Higgins 1990:1077. (5) Cramp 1977:343. (6) Palmer 1962:517. (7) Witherby et al. 1939 in Palmer 1962:517.

May-Jul (but sometimes in rains) Jan—Feb May—Jun Oct-Nov Oct-Feb Jun (previously) Mar, Jul-Sep May-Aug, Jan-Mar (short rains) Apr Feb Aug Jan Sep-Nov Sep-Jan, Oct-Dec May-Aug Mar-Jun Feb-May Feb-May May-Aug

Nesting highly variable and sporadic throughout the extensive range. a Nest times depend on rainfall and nesting periods of other ibises, egrets and spoonbills.

308

Sex

Wing [n] (mm)

Male [11] 261-280 Male, average [11] 269.6 Fern. [8] 238-249 Fern., average [8] 243.5

Whitefaced Ibis (Plegadis chihi]

Puna Ibis (Plegadis ridgwqyi)

BODY M E A S U R E M E N T S

BODY M E A S U R E M E N T S

Culmen (mm)

Tarsus (mm)

118-143 133.9 100-111 103.6

91-98 94.6 63-73 70

Tail (mm)

Wing-Span (mm)

Weight (kg)

-

-

92-100 93.7 83-94 87.8

Reference

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Male [10] 292-302 Male, average [10] 298.4 Fern. [7] 266-287 Fern., average 276.8

108-127 117.4 86.95 92

80-88 84 59-68 63.5

117-125 120.2 106-112 109

-

-

Sex

(1) (1) (1) (1)

Reference: (1) Blake 1977:200

[n]

Reference: (1) Blake 1977:202.

EGG M E A S U R E M E N T S

Average Extremes

EGG MEASUREMENTS

No.

Length (mm)

Width (mm)

Weight (g)

-

50.8 42-55

35.8 32.4-39.4

36 -

Reference (1) (1)

Average Extremes

Reference: (1) Schonwetter 1967.

No.

Length (mm)

Width (mm)

Weight (g)

-

52 49.0-55.4

35.4 33.2-37.3

35 -

Reference: (1) Schonwetter 1967.

EGG-LAYING Place USA South America

EGG-LAYING Place

Months

Peru Ayasuchu South Bolivia

Apr-May, Jun (occasionally) Nov-Dec

309

Months Apr-Jul, Jan-Feb Jan-Mar Nov-Jan/Feb

Reference (1) (1)

Referei (1) (1) (1) (1)

Sharptailed Ibis

Wing (mm)

Sex

[n]

Male Male, average Fern. Fern., average

[9] 390-415 [9] 399.7 [4] 376-413 [4] 394.2

Plumbeous Ibis

(Cercibis oxycercd)

(Harpiprion caerulescens]

BODY MEASUREMENTS

BODY MEASUREMENTS

Culmen (mm)

Tarsus (mm)

Tail (mm)

156-168 161.6 144-167 152.7

-

250-301 272.6 256-272 262.2

Wing-Span (mm)

Weight (kg)

Reference (1) (1) (1) (1)

Reference: (1) Blake 1977:195.

Wing (mm)

Sex

[n]

Male Male, average Fern. Fern., average

[10] 397-450 [10] 413.4 [10] 360-406 [10] 384.8

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight "(kg)

Reference

145-167 153.8 126-147 134.6

— -

165-190 179.5 165-192 176

-

— -

(1) (1) (1) (1)

Reference: (1) Blake 1977:192.

EGG M E A S U R E M E N T S

Average 1? t-*~a.*-^t* Extremes

EGG M E A S U R E M E N T S

No.

Length (mm)

Width (mm)

Weight (g)

— —

65.9 C^A, 1/—OD.J ££ t% o4.

44 4 O 7 44. A.A. Q 4-z./— o

70

Reference (1) (1)

Average

No.

Length (mm)

Width (mm)

Weight (g)

Reference

-

68.6

44.9

74

(1)

Reference: (1) Schonwetter 1967.

Reference: (1) Schonwetter 1967.

EGG-LAYING

EGG-LAYING Place Llanos (S America)

Place

Months

Mato Grosso

Aug/Sep-Feb

Months Mar/Apr-Oct

Corresponding with commencement of the dry season.

Nesting is in the dry season.

310

Buffnecked Ibis (Theristicus caudatus] BODY MEASUREMENTS

Sex

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

131-160 149 131-150 138.8

-

193-220 204.8 180-203 190.9

-

-

(1) (1) (1) (1)

[7] 395-418 [7] 406.3 [6] 384-410 [6] 395.1

143-172 156.8 140-156 146.8

-

196-224 209.3 194-211 202.8

-

-

(1) (1) (1) (1)

[7] 391-407 [7] 400.7 [6] 375-399 [6] 388.8

113-134 126.5 109-130 120

-

187-203 195.5 178-199 186

-

-

(1) (1) (1) (1)

[6] 374-398 383 [6] [7] 351-381 368.4 [7]

120-151 138.5 113-132 122.6

-

169-191 179.8 158-180 172.3

-

-

(1) (1) (1) (1)

[n]

Wing (mm)

Caudatus

Male [10] 385-429 Male, average [10] 402.7 Fern. [10] 378-405 Fern., average [10] 389.8 Hyperorius

Male Male, average Fern. Fern., average Branickii

Male Male, average Fern. Fern., average Melanopis

Male Male, average Fern. Fern., average

Reference: (1) Blake 1977:194.

EGG M E A S U R E M E N T S

Average

EGG-LAYING

No.

Length (mm)

Width (mm)

Weight (g)

Reference

-

66.9 63.6-68.6

44.3 44.0-44.6

69

(i)

References: (1) Schonwetter 1967. (2) Kreugerin Schonwetter 1967.

(2)

Place South Chile Argentina Tierra del Fuego Peru Llanos (Paraguay, Brazil, Venezuela)

311

Months Sep-Dec Nov-Feb Nov-Feb Jun Mar-Oct (but found all the year round depending on climate and altitude)

Green Ibis (Mesembrinibis cayennensis) BODY MEASUREMENTS

Wing (mm)

Culmen (mm)

Sex

W

Male Male, average Fern.

[51 263-300 104.5-118.5 112.6 280 [51 103-113 [41 280-289 108.5 283

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

61.8-63.5 62.4 56.8-62.2 59.5

127-155 138 131-148 142

— — —

— — —

(1) (1) (1) (1)

Reference: (1) Blake 1977:196.

EGG M E A S U R E M E N T S

Average Extremes

No.

Length (mm)

Width (mm)

Weight (g)

Reference

-

54.6 53-57

38.6 37.1-39.5

44

(i) (i)

Reference: (1) Schonwetter 1967.

EGG-LAYING

Place

Months

Panama Colombia Surinam Venezuela

Mar-Apr Mar-Apr Sep Jun-Aug

312

Hadada Ibis (Bostrychia hagedash} BODY MEASUREMENTS Wing (mm)

Sex

[n]

B. h. hagedash Both Average

[17] 334^370 - 353

B. h. brevirostris Both

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

117-153 134

63-73 68

137-170 154

-

1.262

Reference

(1) (1)

-

330-383

126-163

-

335-392

152-174

-

-

-

-

(1)

B. h. brevirostris Both [5] 335-365

130-141

67-73

145-153

-

-

(2)

B. h. nilotica Both sexes alike

(1)

References: (1) Brown et al. 1982:195. (2) Gates 1930:117.

EGG M E A S U R E M E N T S

EGG-LAYING

No.

Length (mm)

Width (mm)

Weight (g)

172 — — 90

54-66 62 51 57.0-65.1

38-48 42 45 38.5-46.8

55-62 — — —

Place

Reference

Guinea-Bissau Gambia Niger Nigeria Ethiopia Kenya, Uganda Tanzania Zambia Zaire Zimbabwe Mozambique South Africa

(i)

(2) (2) (3)

References: (1) Brown et al. 1982:196. (2) Mackworth-Praed and Grant 1973:75. (3) McLachlan and Liversidge 1970:56.

Months May-Oct (rains) Jan-Mar (dry) Jun Jun-Jul (rains) Sep Mar-Jun (peak long rains; but can be year round) Apr-Jul, Oct Jan, Mar-Apr, Jul-Dec Jan-Dec Oct-Dec Dec Jan-Dec

This species has an extended breeding season throughout the year but peak laying occurs during and after the main rains.

313

Wattled Ibis (Bostrychia carcunculatd) BODY M E A S U R E M E N T S

Wing (mm)

Sex

[n]

Both

[7] 358-380

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

110-127

63-66

150-180

-

-

(1,2)

References: (1) Moltoni and Ruscone 1942. (2) Friedman 1930 in Brown et al. 1982:196.

EGG M E A S U R E M E N T S

No.

Length (mm)

Width (mm)

Weight (g)

Reference

-

59-61

40-59

50

(1)

Reference: (1) Brown et al. 1982:197.

EGG-LAYING

Place Ethiopia

Months Mar-May, Jul, Dec

314

Olive Ibis (Bostrychia olivacea) BODY M E A S U R E M E N T S

Sex

Wing Culmen Tarsus (mm) (mm) (mm)

W

Tail (mm)

Wing-Span (mm)

Weight (kg)

EGG M E A S U R E M E N T S

Reference

B. o. olivacea — Male Male — — Ad. Ad. —

333 330 330 334

96 c. 100 95 96

67 72 73 68

146 150 150 147

— — — —

— — — —

(1) (1) (2) (2)

B. o. akeleyorum Male —

358

108

71

160

-

-

Fem. Male

362 355

102 115.5

64 73

165 150

-

-

( 1 ) (Type) Mt. Kenya 9000 ft (Akeley) ( 1 ) Mt. Kenya 9000 ft (Akeley) ( 1 ) Mt. Kenya 6500 ft (van Someren)

— —

B. o. rothschildi Male Fem. —

328 313

95 c. 90

66 67

110

B. o. cupreipennis Male (?) Imm. — Ad. Ad. Fem. -

310 280 317 309 335

85 76 86.5 85 94

57

130

62 59 70

125 127 133

255 256 233 255 250

75 76 75 68 75

53 55 50 50 —

103 101 88 96 —

B. o. bocagei Ad. Male Male Fem. p -

B. o. akeleyorum B. o. akeleyorum

(Type) Guinea Coast (Type of splendida) Liberia Bannerman collection Bannerman collection

—

— -

Length (mm)

Width (mm)

Weight (g)

Reference

— —

56-58 57.1

40-41 40.2

50 50

(1) (2)

No data available on other races. References: (1) Brown et al. 1982:198. (2) Schonwetter 1967.

EGG-LAYING Place Kenya

(1) (Type) Prince's Is. (L. Fea) (1) Prince's Is. (Dohrn and Keulemans) (1) (1) (1) (1) (1)

No.

Months Jun-Aug

This species is little known, with no breeding records outside Kenya.

(Type) Bipindi (Zenker) Efulen (Bates) Gamma R. (Du Chaithu) Near Bolobo (Schouteden) Avakubi (Lang and Chapin)

(1) (Type) Sao Tome ( 1 ) Sao Tome (F. Newton) ( 1 ) Sao Tome (F. Newton) ( 1 ) Sao Tome (F. Newton) (3)

References: (1) Chapin 1923. (2) de Naurois 1973. (3) Specimen in British Museum of Natural History. Measurements taken by Chapin were with the wing straightened and this is probably the case in other reports, but slight differences occur, with some authors using the same skins. There are now estimated to be 9 skins only of B. o. bocagei:, in spite of this small number it is to be hoped that should this race be re-discovered the temptation to collect further specimens will be resisted.

315

Spotbreasted Ibis

Madagascar Crested Ibis

(Bostrychia rara)

(Lophotibis cristatd)

BODY M E A S U R E M E N T S

Wing (mm)

Sex

W

Both Male Fern.

[10] 270-290 [3] 265-291 [3] 253-275

BODY M E A S U R E M E N T S

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

115-130 122-131 122-131

56-65 57-63 57-63

112-120 102-120 102-120

-

Weight (kg) —

Reference

Sex

W

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight •(kg)

Reference

(1) (2) (2)

Both

-

355

134

64

184

-

-

(1)

Reference: (1) Elliott 1877:500.

References: (1) Brown et al. 1982:198. (2) Gates 1930:118. EGG M E A S U R E M E N T S

EGG M E A S U R E M E N T S

No.

Average

Length (mm)

Width (mm)

55

35

Weight (g)

No. Reference

Length (mm)

Width (mm)

55

41

Weight (g)

(1)

Reference: (1) Langrand 1990:117.

(1)

Reference: (1) Brosset and Erard 1976. EGG-LAYING EGG-LAYING

Place Gabon Throughout West Africa

Months Jan-Jun, Sep-Dec Mar-Jun, Sep-Dec

Nest building carried out when flood waters are high. Laying coincides with rainfall peaks, but some may occur if water remains high in dry season.

316

Reference

Place

Months

Madagascar

Sep-Jan

Sacred Ibis (Threskiornis aethiopicus) BODY MEASUREMENTS

Sex

Male Male, average Fern. Fern., average Male

M

Wing (mm)

380-385 383 [2-5] [3] 361-362 [3-81 362 396 [1] [2]

EGG-LAYING

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

162-183 170 135-157 146 167

100-116 108 94-103 98 105

129-140 134 129-135 131 140

1120-1240 — — — —

1.53

(1,2)

—

— •

— —

Place Aldabra (Seychelles) Senegal Gambia Guinea-Bissau Nigeria Mali Chad Sudan Ethiopia W Kenya Central Kenya Uganda Burundi Tanzania Zambia Malawi Zimbabwe Namibia South Africa

(1,2) (1,2) (1,2)

(3)

References: (1) Brown et al. 1982:200. (2) Cramp 1977:351. (3) Gates 1930:118.

EGG MEASUREMENTS

T. a. aethiopicus

No.

Length (mm)

Width (mm)

Weight (g)

Reference

_ 42 — 60

56-70 58-68 64 60.2-70.5

38-48 39-46 42 39.7-51.2

62 62 — —

(1) (2) (3) (4)

—

59.5

41.3

55

(5)

T. a. bernieri

Average

References: (1) Brown et al. 1982:201. (2) Schonwetter 1967 in Cramp 1977:350. (3) Mackworth-Praed and Grant 1973:73. (4) McLachlan and Liversidge 1970:54. (5) Schonwetter 1967.

317

Months Nov-Dec Nov-Dec Jan, Jul, Nov-Dec Mar-Sep Jan-Feb, May-Jun, Dec Jul-Feb Jun Mar, Jun-Aug Mar-Jun, Aug-Sep, Jan-Jun Apr May-Jun, Aug, Nov Feb, May, Nov Jul Jan-Feb, May, Oct Jan-Sep Jan-Sep Apr Jul-Aug Jan-Mar, May, Aug-Dec

Oriental White Ibis

Australian White Ibis

( Threskiornis melanocephalus]

( Threskiornis moluccd)

BODY M E A S U R E M E N T S

BODY M E A S U R E M E N T S

Sex

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Both Male

-

343-370 381

139-170 185

99-118 119

133-145 129

-

-

Reference: (1) S. Ali and S.D. Ripley 1968:1 12.

EGG MEASUREMENTS

Average

No.

Length (mm)

Width (mm)

Weight (g)

Reference

150 11 11

63.5 62.31 59.4-65.1

43.1 41.43 39.2-44.2

5.073

(1) (2) (2)

Reference (1) (1)

Sex

[n]

Male Male, average T?A<~% rem. Fern., average Male Male, average Fern. Fern., average Male Male average Fern. Fern., average

[3] [3] r/ll [4J [4] — —

Wing (mm)

Culmen (mm)

355-398 183.5-197.1 94.9-108.1 380.6 188.6 102.7 "2Ei£ Q"7O 149.1— 1/iO 1 15o.4 1 £Q A QO A O£ A 5D3—672 9^.4—95.4 363.7 153.7 94.1 380-397 186-190 94-107 389 188 103 348-373 131-157 81-90 361 148 85.8 173-210 190 5 140-168 — 154 —

Wing-Span (mm)

Weight (kg)

120-134 126.3 1 OO 150 1 QA \2.i— 125 — -

1100-1250 — — -

1.7-2.5 — 1.4-1.9 — -

—

—

—

EGG MEASUREMENTS

No.

EGG-LAYING

N India S India, Sri Lanka

Tail (mm)

Reference (1) (1) (1) (1) (2) (2) (2) (2) (3) (3) (3) (3)

References: (1), (2), (3) in Marchant and Higgins 1990:1086. (1) SAM, MV. (2) Healesville Fauna Reserve Victoria (ABBBS). (2) Vic. (K.W. Lowe).

References: (1) Baker in S. Ali and S.D. Ripley 1968:111. (2) Hellebrekers and Hoogerwerf 1967: 14.

Place

Tarsus (mm)

-

Months

-

Jun/Jul-Oct Nov-Feb/Mar

Length (mm)

Width (mm)

Weight (g)

60 71 59-75 59 78 60-70

41-47 39-49 39-49 40-46

668 -

-

Reference (1) (1) (1) (1)

References: (1) Marchant and Higgins 1990:1084.

EGG-LAYING Place S Victoria (Australia) S Australia NSW (Australia)

Months Jun-Nov, May-Mar (rarely) Sep-Dec, Jan Sep-Apr

This species does not always breed annually and variations of nesting times occur according to rainy season.

318

Sex

[n]

Male Fern.

-

Wing (mm)

Strawnecked Ibis

Waldrapp Ibis

( Threskiornis spinicollis]

(Geronticus eremita)

BODY MEASUREMENTS

E&ODY M E A S U R E M E N T S

Culmen (mm)

364-415 158.2-179.8 351-375 133.4-142.2

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

Sex

[n]

83.6-102 78.4-83.2

130-152 126-142

-

1.4-1.57 1.15-1.32

(1) (1)

Male Fern.

-

Reference: (1) Marchant and Higgins 1990.

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

403-420 390-408

133-147 115-131

68-72 68-72

196-220 196-220

-

-

(1) (1)

-

-

-

-

(1) (1)

Western population (Morocco) Male 132-146 Fern. 128-145 Northeastern population (Turkey) Male 126-132 Fern 121 127

EGG MEASUREMENTS

Average Extremes

Wing (mm)

No.

Length (mm)

Width (mm)

Weight (g)

14 -

61.4-69.1 64.8 57.5-70.0

42.4-47.0 44 38.6-48.0

69 -

(1)

(\\ \l)

Reference: (1) Brown et al. 1982.

Reference (1) (2) (2)

EGG M E A S U R E M E N T S

f

"D f^»-^v-.^^r- • (1) / 1 \ Marchant A/f ^*~^\->^n + and ~nA tTi/QQfV 1lUyo. flO*-? ((t) )\ Qr«J-»orMA7*»t*-«»»r /- r 4v-,r- 1 References. rliggms lyyu. ocnonwetter 1Q£7 iyt>/.

No. 46 13

EGG-LAYING

Place S, SE Australia W Australia N Australia

Months

Length (mm)

Width (mm)

Weight (g)

61-69 63

42-46 44

68 54-67

Reference (1,2) (3) (4)

References- (1) Brown et al 1982*204 (2) Schonwetter 1967 in Cramp 1977:347. (3) Wackernagel 1964 in Cramp 1977:347. (4) Mackworth-Praed and Grant 1973:74.

Sep-Feb Sep-Feb Sporadic not annually

This species is largely dependent on rainfalls but uses permanent water also. It often nests in company with Australian White Ibis.

EGG-LAYING Place Morocco Algeria

319

Months Mar-May Only after successful rains

Bald Ibis

Wing (mm)

Sex

[n]

Both

[7] 369-403

Black Ibis

( Geronticus calvus)

(Pseudibis papillosa)

BODY M E A S U R E M E N T S

BODY M E A S U R E M E N T S

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

140-152

66-73

175-200

-

-

(1)

Note: females are slightly larger with a slightly longer bill and tarsus. Reference: (1) Brown et al. 1982.

Wing (mm)

Sex

[n]

P. p. papillosa Both

— 365-400

P. p. davisoni Both

-

Culmen (mm) •

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Reference

138-158

75-85

165-194

—

—

(1)

197

83

229

-

-

419

References: (1) Baker (in Ali and Ripley 1968). (2) D.G. Elliott 1877:49. EGG MEASUREMENTS

No.

Length (mm)

Width (mm)

Weight (g)

Reference

24

51-71 57-71

38-49 38-49

68 -

(1) (2)

EGG M E A S U R E M E N T S

No.

Length (mm)

Width (mm)

Weight (g)

Reference

References: (1) Brown et al. 1982:206. (2) McLachlan and Liversidge 1970:55.

P. p. papillosa Average Average Extremes

-

63.0 63.0 56-70

43.8 43.8 38-50

64 -

(1) (2) (2)

EGG-LAYING

P. p. davisoni Average Extremes

-

63.7 61.0-68.2

45.2 43.9-46.7

71 -

(2) (2)

Place South Africa

References: (1) Baker in S. Ali and S.D. Ripley 1968:133. (2) Schonwetter 1967

Months Aug-Oct

EGG-LAYING

Place

Months

P. p. papillosa N India Mar and Oct Gujarat, India Apr-May, Dec-Jan S India Apr-May, Dec-Jan Nepal Mar and Oct In all areas two distinct nesting periods occur, but these are extremely variable due to rains. P. p. davisoni Burma (formerly)

320

Feb-Mar

(2)

Wing (mm)

Sex

[n]

Male Fern.

- 546 546

Giant Ibis

Oriental Crested Ibis

(Thaumatibis giganted)

(Nipponia nippori)

BODY M E A S U R E M E N T S

BODY M E A S U R E M E N T S

Culmen (mm)

Tarsus (mm)

Tail (mm)

236 241

112 108

267 279

Wing-Span (mm)

Weight (kg)

Reference

Sex

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

(1) (2)

Both

—

419

165

89

165

Wing-Span (mm)

Weight (kg)

(1)

Reference: (1) D.G. Elliott 1877.

References: (1) Robinson and Kloss 1911. (2) D.G. Elliott 1877.

EGG MEASUREMENTS EGG MEASUREMENTS

No.

Length (mm)

Width (mm)

Weight (g)

Reference Average Extremes

Not recorded

No.

Length (mm)

Width (mm)

Weight (g)

-

50.5-66.0

57.3

35.6 31.5-40.5

39(?)

Reference

(1) (2)

References: (1) Schonwetter 1967. (2) Swinhoe and Nehrkorn in Schonwetter 1967.

EGG-LAYING EGG-LAYING

Place

Months

Not recorded

Place China Japan (formerly)

321

Reference

Months Mar mid-Apr

Eurasian Spoonbill (Platalea leucorodid) BODY M E A S U R E M E N T S

Sex

Culmen (mm)

Tarsus (mm)

Tail (mm)

[14] 367-409 389 [14] [3] 382-401 394 [3] [13] 386-412 394 [13] [8] 350-388 373 [8] [5] 375-395 — [1]

194-222 208 211-224 216 195-231 213 175-223 206 205-230 —

148-150 149 144-153 149 140-163 149 113-150 141 140-150 —

— — 121-126 124 108-126 117 — — 110-120 —

— — 394 410 355-388 370 364-369 367 360-377 370 350-373 359 350-370 — — — — 365 _ 350-395 360-397 350-410 _

— — 238 230 176-223 191 187-194 191 168-191 182 168-183 177 175-190 — — — — 183 — 180-228 183-223 180-238 — — — 173-225 199 203 — —

— — 158 145 133-148 139 134-139 137 123-141 131 115-140 129 — — — — — 125 — 130-165 120-145 130-165 140-150 — — 128-163 143 143 — -

— — 132 — — — 110-113 112 108-118 113 — — — -

[n]

Wing (mm)

P. I. leucorodia

Male Male, average Male Male, average Male Male, average Male Male, average Male Male •Male \,ri Male Male Male Male Fern. Fern., average Fern. Fern., average Fern. Fern., average Fern. Fern., average Fern. Fern. Fern. Fern. Fern. Fern. Both Both Both Both Both Both Both, average Both Both, average Both, average Unsexed Unsexed

[1] [1] [1] [1]

[22] [22]

[2] [2]

[10] [10] [10] [10]

[8] [1] [1] [1] [1] [1] [?] [?] [?] [?] [?]

[12] 370-410 [12] 387.5 — [?] — [?] 373 [?] — [1] — [1]

— — 108-122 — 110-132 — — — _ —

Wing-Span (mm)

'

— — — — —

— — — — — — — — 1258 1400 1450 — — 1150-1300 — — — — _ —

118 _

_

—

—

Weight (kg)

Reference

— — — — — — — — — 1.96

(1) (1) (2) (2) (3) (3) (16) (16) (12) (8)

/-vr 2 .DJ

2.08 1.656 — — — — — — — — — — — — — — 1.94 — — — — — — — — — — — 1.77 1.785

Culmen (mm)

Tarsus (mm)

Tail (mm)

[9] 364-390 [9] 372 [1] 382 [8] 340-362 [8] 352.5

157-200 184 205 151-180 167

123-144 130 140 115-129 120

132 -

(1) (1) (11) (1) (1)

[?] 325-360 [1] 325 [1] 308

145-170 153 150

102-118 103 113

-

(3) (13) (16)

Sex

[n]

P. I balsaci Male Male, average Male Fern. Fern., average

Wing (mm)

Wing-Span (mm)

'Weight (kg)

Reference

P. I. archeri

Both Unsexed Unsexed

/n\

U)

Longevity in wild = 28 years 2 months (Rydzewski 1973). Longevity in captivity = 27 years (Schenker 1978). References: ( 1 ) de Naurois and Roux 1974. (2) T.H. Shaw 1936:128-129. (3) Cramp 1977:352, 357. (4) E.C.S. Baker 1929:312. (5) Dementievand Gladkov 195 1:421. (6) Hartert 1912-21 :1217-1218. (7) Lippens 1954:26. (8) Vasvari 1955. (9) S. Ali and S.D. Ripley 1968:117. (10) La Touche 1931-34:431. (11) de Naurois 1959. (12) Witherbyetal. 1939: 121.(13)Pers.obs.M.P.K. (Leiden Mus./26 Apr 89). (14) Kobayashi 1963:67. (15) Sharpe 1898:46-47. (16) Ogilvie-Grant 1889.

(2) (3) (10) (13) (1) (1) (2) (2) (3) (3) (16) (16) (12) (5) (5) (5) (2) (13) (3) (4) (6) (14) (7) (5) (5) (15) (15) (15) (9) (9)

322

Eurasian Spoonbill continued EGG MEASUREMENTS

No.

Length (mm)

Width (mm)

250 250 250 105 105 17 17 40 40 100 100 100 100 160 160

56-74.5 67.2 — 47.9-77.1 61.2 65.1-73.0 69.5 61.1-72.1 65.6 62.2-78.3 68.2 58.2-74.6 67.2 58.5-76.4 67.4

40.0-49.5 45.5 36.0-51.7 42.4 43.2-49.0 45.0 41.0-47.7 44.2 41.5-49.5 46 41-49.2 45.4 41.2-49.0 46.8

P. L balsaci Average

12

67

45

P. L archeri Extremes Average

15 15

57-65 60

40-45 43.2

P. L leucorodia Extremes Average Calculated Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average

EGG-LAYING Weight (g) — 76 — -

Place

Reference

(i) (i) (i) (2) (2) (3) (3) (4) (4) (5) (5) (7) (7) (8)

(8) (9)

-

(6) (6)

References: (1) Schonwetter 1967. (2) Vespremeanu 1968a. (3) Dementiev and Gladkov 1951:418. (4) E.C.S. Baker 1929:312. (5) Hellebrekers 1967. (6) Archer and Godman 1937:82. (7) Witherbyetal. 1939:119. (8) Bauer and Glutz von Blotzheim 1966:438. (9) Etchecopar and Hue 1967:63.

323

Months

P. I. leucorodia Spain, Portugal N, central Europe N India, Pakistan S India Sri Lanka Algeria (formerly: 19th century)

Feb-Apr Apr-Jun Jul-Oct Nov—Jan Dec-Apr May

P. L balsaci Mauritania

Apr-Jul

P. I. archeri Red Sea, Somalia (coastal)

Apr-May, Jun-Aug

Royal Spoonbill (Platalea regid) EGG MEASUREMENTS

BODY MEASUREMENTS

Sex

W

Male M Male, average [4] Male [3] Male, average [3] Male [3] Male, average [3] Male [3] Male, average [3] Male [6] Male, average [2] Male [1] Male [1] Male [1] Male [1] Fern. [5] Fern., average [5] Fem. [9] Fern., average [9] Fem. M Fem., average [4] Fem. [4] Fem. [1] Fem. [1] Unsexed [?] Unsexed [?] Unsexed [?] Unsexed [?] Unsexed [1] Unsexed [1] Unsexed [1] Unsexed [1] Unsexed [1] Unsexed [1] Unsexed [1]

Wing (mm)

Culmen (mm)

Tarsus (mm)

355-370 361 353-374 362 — — — — 347-385 — — 370 390 — 325-358 339 — — — — 350-380 — 375 375 355 375 375 360 370 — — — — —

165-198 187 180-196 190 183-196 188 176-187 180 205-219 183 — 168 189 188 163-175 166 136-189 164 164-172 167 158-187 190 164 200 220 212 200 195 187 — —

120-143 133 120-143 134 121-137 129 — — 116-127 130 — 121 130 135 113-125 120 — — 115-128 123 100-120 128 126 138 130 137 138 128 126 — —

— — —

— — —

Tail (mm)

Wing-Span (mm)

Weight (kg)

— — — — — — — — — — — — — — — — — — — — — — 115 110 137 115 — — — — — _ —

— — _

— — _

— — — — — 1200a — _

— — — — 1.65-2.07 — — _

_ — — — — — —

_ — — — — — — 1.4-1.8 — — — — — _

— — — — — _ — — -

— — 1.70 1.68 1.29 1.726 1.80

Reference

(i) (i)

Extremes Average Calculated Extremes Average Average Average Average Average Extremes Ova) Average Java)

(2) (2) (4) (4) (3) (3)

(15)

(4) (4)

(8) (8)

(10)

No.

Length (mm)

Width (mm)

16 16 16 18 18 — — 8 16 11

59.5-69.3 65.7 — 58.5-69.6 63.6 65 64.3 65.5 68 60.5-64.5

41.7-44.5 43.2 — 40.1-44.9 43.3 44 42.8 42.7 45 42.8-44.5

11

62.5

43.5

Weight (g)

66

Reference (1) (1)

(1)

(7) (7) (2) (4) (5) (6) (3) (3)

References: (1) Schonwetter 1967. (2) Beruldsen 1980:158. (3) Hoogerwerf 1952. (4) Lucas and LeSouef 1911:126. (5) AJ. North 1913-14:13. (6) Vestjens 1977b. (7) Hellebrekers and Hoogerwerf 1967:14.

(1) (1) (3) (3) (9) (9)

(15) (10)

EGG-LAYING

(9) (") (12) (13) (14)

Place S Australia N Australia New Zealand (South Island) W Java (formerly)

(8) (8) (5) (5) (5) (6) (7)

a

From specimen label. References: (1) Ogilvie-Grant 1889. (2) Pers. obs. M.P.K. (British Mus./lO Apr 89). (3) Vestjens 1976:56. (4) Pers. obs. M.P.K. (S. Austr. Mus./lO Apr 85). (5) B.P. Hall 1974:46. (6) Lavery 1971:52. (7) Serventy and Whittell 1962:125. (8) Pers. obs. M.P.K. (Leiden Mus./26 Apr 89). (9) Pers. obs. M.P.K. (CSIRO, Canberra/4 Mar 85). (10) Pers. obs. M.P.K. (W. Austr. Mus./24 Apr 85). (11) Lucas and LeSouef 1911:126. (12) Rand and Gilliard 1967:49. (13) Gould 1865:288. (14) Sharpe 1898:48. (15) K.W. Lowe (pers. comm., 4 Mar 90).

324

Months Aug-Jan Feb-May Nov-Dec Mar-Sep

Blackfaced Spoonbill (Platalea minor) BODY M E A S U R E M E N T S

Sex

[n]

Wing (mm)

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

Male Male, average Male Male Fern. Fern. Fern. Fern, (imm.) Both Both Both Both Both Both Both Unsexed Unsexed Unsexed Unsexed Unsexed Unsexed Unsexed

[2] [2] [1] [1] [1] [1] [1] [1] [?] [?] [?] [?] [?] [?] [4] [1] [1] [1] [1] [1] [1] [1]

338-360 249 365 362 376 350 350 — 356-371 338-376 350-370 334-358 — 338-371 338-369 350 — 367 366 361 367 361

173-180 176 180 180 188 180 183 _

118-125 121 119 125 135 125 118 — 119-121 — — 106-115 108-134 119-134 108-133 125 — 126 125 132 131 120

_ _ 105 _ _ _ 100 102-107 105-116 _ 103-122 _ 100-117 105-113 _ _ _ — _ _ 101

_ _ _ _ _

_ _ _ _ _

-

1.338

_

_

_ _ _ _ — _ _ -

_ _ _ _ — _ _ -

183-185 — 190-193 151-196 — 145-193 145-163 193 155 192 178 190 191 178

Reference (1) (1) (2) (10) (10) (1) (2) (13) (3) W (5) (6) (7) (8) (9) (1) (1) (10) (10) (10) (11) (12)

References: (1) Ogilvie-Grant 1889. (2) Sharpe 1898:51. (3) Hackisuka 1931-32:347. (4) Hackisuka and Udagawa 1951:102. (5) Hartert 1912-21:1220. (6) H.G. Won 1966. (7) B.F. King and E.G. Dickinson 1975:54. (8) Kobayashi 1956b:67. (9) La Touche 1931-34:433. (10) Pers. obs. M.P.K. (British Mus./lO Apr 89). (11) Pers. obs. M.P.K. (Leiden Mus./26 Apr 89). (12) duPont 1971:23. (13) Fennell and King 1964:241.

EGG M E A S U R E M E N T S

No.

Extremes Average Calculated Extremes Average

10 10 10

—

10

Length (mm)

Width (mm)

58.2-66.0 62.5 — 63-67 63.7

40.4-48.0 44.6 — 41-42 42.6

EGG-LAYING

Weight (g)

Reference

— 68 — —

(1) (1) (1) (2) (3)

References: (1) Schonwetter 1967. (2) Won 1966. (3) Etchecopar and Hue 1978:80.

325

Place

Months

Korea

Jun-Jul

African Spoonbill (Platalea alba) BODY M E A S U R E M E N T S

Sex

[n]

Male Male Fern. Fern. Fern. Fern. Both Both Both Both, average Unsexed

[?] [1] [?] [?] [1] [1] [?] [?] [15] [15] [1]

Wing (mm) 378-405 405 363-380 363-380 370 370-415 365-414 384 -

Culmen (mm)

Tarsus (mm)

Tail (mm)

226 — 160-184 175 — 170-226 — 172-230 193 —

166 — 130-139 135 — 130-160 — 131-157 144 —

137 — 130-136 128 — 135-137 — 105-152 124 —

Wing-Span (mm)

Weight (kg)

Reference

— _ — — —

— — — — 1.79 — — — — 1.59

(1) (2) (1) (2) (5) (4) (1) (3) (4) (4) (6)

Longevity in wild = 11 years 5 months (McLachlan and Liversidge 1978:57). References: (1) Bouet 1955a:132. (2) Bannerman 1930:124. (3) Clancey 1964a:51. (4) Brown et al. 1982:208. (5) Sharpe 1898:50. (6) FJ.Jackson 1938:88.

EGG-LAYING

EGG M E A S U R E M E N T S

Extremes Average Calculated Extremes Average Extremes Average Extremes Average Extremes Average Extremes Average Extremes

No.

Length (mm)

Width (mm)

23 23 23 205 205 15 15 33 33 72 72 4 10 —

59.1-72.2 65.4 — 57-82 68 65.2-73.9 69.1 61.0-71.4 66.4 54.6-72.4 66.2 67-73 66.8 65-70

40.6-48.0 44.0 — 40-48 45 40.5-47.2 44.5 39.7-47.8 44.2 35.5-47.5 44.5 44-46 43.4 42-44

Place

Weight (g)

Reference

— 69 — 69 — — — — — — — — —

(1) (1) (1) (2) (2) (3) (3) (4) (4) (5) (5) (6) (6) (7)

Senegal Guinea-Bissau Sierra Leone Niger Mali Chad Nigeria Sudan Ethiopia Kenya Uganda Burundi Tanzania Zambia Malawi Zimbabwe Botswana Namibia South Africa (Cape Prov.) South Africa (Transvaal, O.F.S.) South Africa (Natal) Madagascar

References: (1) Schonwetter 1967. (2) Brown et al. 1982:209. (3) Broekhuysen 1957. (4) McLachlan and Liversidge 1970:57. (5) Whitelaw 1968. (6) FJ. Jackson 1938:90. (7) Bouet 1955a:132.

326

Months Oct-Mar Jan-Feb, May, Nov-Dec Sep Oct Jan-Apr, Aug-Oct Jan-Apr Jan-Feb Feb-Mar, Sep Jul-Aug, Oct Dec-Jul, Sep-Oct (peak in May) Feb-Jun Jun-Oct Feb— Mar, May Feb-Jul Sep Jan-Sep Mar-Jun Feb-Aug, Oct Aug—Nov Mar-Sep Feb-Nov Feb

Yellowbilled Spoonbill (Platalea flavipes) BODY MEASUREMENTS Wing (mm)

Sex

W

Male Male, average Male Male, average Male, average Male, average Male Male Male Fern. Fern., average Fern. Fern., average Fern., average Fern. Fern. Fern. Both, average Unsexed Unsexed

[5] — [5] [3] 410-420 415 [3] 400 [?] 413 [?] — [1] — [1] _ [1] [8] 186-231 205 [8] — [4] — M 370 [?] 395 [1] — [1] — [1] 413 [?] — [1] 363 [1]

Culmen (mm)

Tarsus (mm)

Tail (mm)

Wing-Span (mm)

Weight (kg)

225-238 231 217-236 227 213 225 — 222 222 — — 179-191 185 — — 195 205 225 — 181

— 133-140 136 133 138 — 130 134 — — 113-122 118 — — 119 115 138 — 119

— — — 175 170 175 — — — — — — — 172 — — 170 — 138

— • — — — — — 1375a — — — — — — — 1340a — — 1450 —

— 1.75-2.0a 1.92a — — — 1.82a — — — — — — 1.5a 1.7a — — 1.8 —

Reference

(i) (i)

(2) (2) (3) (4) (2) (6) (6) (1) (1) (2) (2) (4) (2) (6) (6) (5) (7) (8)

a

From specimen labels. References: (1) Vestjens 1975a. (2) Pers. obs. M.P.K. (CSIRO, Canberra/4 Mar 85). (3) AJ. North 191314:14. (4) Sharpe 1898:52. (5) Lucas and Le Souef 1911:127. (6) Pers. obs. M.P.K. (S. Austr. Mus./lO Apr 85). (7) Serventy and Whittell 1962:126. (8) Gould 1865:289.

EGG MEASUREMENTS

Extremes Average Calculated Extremes Average Average Average Average Average Average

No.

Length (mm)

Width (mm)

27 27 27 13 13 12 8 — — —

65.0-78.2 69.6 — 65.5-77.0 70.1 69 68 70 68 60[?]

44.0-49.0 46.0 — 43.3-47.5 45.2 47 45 47 45 47

EGG-LAYING Weight (g)

— 81 — — — — — — —

Place

Reference

S Australia N Australia

(i) (i) (i)

(2) (2) (3) (4) (5) (6) (7)

References: (1) Schonwetter 1967 (2) AJ. North 1913-14:18. (3) Vestjens 1977b:56. (4) Serventy and Whittell 1962:126. (5) Lucas and Le Souef 1911:127. (6) Beruldsen 1980:159. (7) Frith 1976:91.

327

Months Aug-Jan Mar-Jun

Roseate Spoonbill (Platalea ajaja) BODY M E A S U R E M E N T S

Culmen (mm)

Tarsus (mm)

Tail (mm)

Male [13] 345-375 Male, average [13] 356 Male [?] -

156-173 166 —

107-112 109 —

93-107 99 —

Male [?] Male [?] 341-369 Male, average [?] 351 Male [1] 350 Male [1] 345 Fern. [11] 325-365 Fern., average [11] 343 Fern. [?] Fern. [?] 322-352 Fern, average •[?] 335 Fern. [1] 345 Fern. [1] 333 Both [?] Both [?] 360^370 Both, average [?]

— 154-179 168 — 163 145-177 157 — 151-162 156 — — 150-180 163-172 —

— 107-124 114 — 105 97-117 104 — 97-107 104 — — 107-123 — —

— 94-103 98 — 93 87-108 97 — 85-100 94 — — — 97-105 —

Sex

[n]-

Wing (mm)

Wing-Span (mm)

Weight (kg)

Reference

— 'to about 1300' — — — — — — — — — — — — — — 1275

— 'up to 1.6'

(1) (1) (2)

1.2-1.8 — — — — — — 1.4-1.7 — — — — — — 1.6

(3) (8) (8) (4) (5) (1) (1) (3) (8) (8) (4) (5) (6) (7) (8)

Longevity in captivity = 10 years 5 months (Terres 1980:554). References: (1) Blake 1977:202. (2) Palmer 1962:534. (3) Haverschmidt 1968:30. (4) ffrench 1973:78. (5) Sharpe 1898:54. (6) J.C. Lewis 1983:1. (7) Goodalletal. 1951:108. (8) Oberholser and Kincaid 1974:132-133.

EGG MEASUREMENTS

Extremes Average Calculated Extremes Average Extremes Average Average Average Average

No.

Length (mm)

Width (mm)

50 50 50 70 70 40 40 20 — —

59.5-71.0 65.2 — 56.9-71.2 64.4 60.2-71.5 65.0 65.0 64.3 65.0

41.2-46.5 43.8 — 41.3-47.9 43.7 41.0-47.0 43.9 44.2 43.3 43.7

EGG-LAYING

Weight (g)

Reference

— 68 52.4-77.0 62.1 — — — — —

(1) (1) (1) (2) (2) (3) (3) (4) (5) (6)

Place E Texas, SW Louisiana Florida Bay Peninsular Florida Mexico (Gulf coast) Honduras Cuba (Camaguey Prov.) Surinam Argentina (Buenos Aires Prov.)

References: (1) Schonwetter 1967. (2) D.H. White et al. 1982. (3) Bent 1926:16. (4) Palmer 1962:538. (5) Sprunt 1954:49. (6) Oberholser and Kincaid 1974:133. 328

Months Apr-Jun Nov-Dec Mar-Apr Feb Feb-Mar Sep-Nov Aug-Sep Oct-Dec

Bibliography The following bibliography is based extensively on a list of references kindly supplied by Malcolm Coulter and the ICBP Storks, Ibises and Spoonbills Group.

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Index Page numbers in bold type refer to the main entries for each species Abdim's Stork (Ciconia abdimii) 9, 11, 29, 70, 71, 73, 74, 75-9, 82, 84, 85, 88, 102, 137, 291 Adjutant Stork see Greater Adjutant Stork; Lesser Adjutant Stork African Openbill Stork (Anastomus lamelligerus) 11, 29, 31, 60,61,62,63-7, 166,289 African Spoonbill (Platalea alba) 24, 25, 28, 166, 214, 255, 256, 267-70, 277, 280, 326 American White Ibis (Eudocimus ruber) 13, 17, 20, 21, 22, 31, 32, 34, 37, 146-56, 161, 163, 164, 166, 170, 251, 306,318 American Wood Stork (Mycteria americana) 17, 21, 30, 31, 32, 33, 34, 35-42, 47, 50, 51, 93, 121, 284 Anastomus lamelligerus 11, 29, 31, 60, 61, 62, 63-7, 166, 289 A. oscitans 11, 18, 23, 25, 26, 27, 45, 55, 59-62, 65, 66, 67, 105,219,288 Asian Openbill Stork (Anastomus oscitans) 11, 18, 23, 25, 26, 27, 45, 55, 59-62, 65, 66, 67, 105, 219, 288 Australian White Ibis (Threskiornis molucca) 6, 13, 215, 217, 218, 219, 221-5, 227, 228, 229, 300 Balaeniceps rex 7, 9, 10, 14, 30, 32, 135, 139-45, 305 Bald Ibis (Geronticus calvus) 17, 20, 29, 34, 73, 236-40, 321 Barefaced Ibis (Phimosus infuscatus) 24, 94, 157-61, 164, 170,307 Black Ibis (Pseudibispapillosa) 16, 19, 20, 164, 241-4, 245, 247, 320 Black Stork (Ciconia nigra) 11,13, 16, 17, 20, 68-74, 75, 77, 79, 82, 83, 84, 94, 95, 111, 290 Blackfaced Spoonbill (Platalea minor) 14, 16, 257, 261, 262-6, 325 Blacknecked Stork (Ephippiorhychus asiaticus) 12, 17, 19, 21, 24, 31, 33, 106, 109-13, 115, 118, 122, 123, 166, 224, 299 Bostrychia carunculata 20, 196-8, 314 B. hagedash 20, 22, 29, 32, 65, 191, 192-5, 197, 201, 202, 203,205,237,243,313 B. h. brevirostris 193, 194, 195 B. h. erlangeri 195 B. h. hagedash 193, 194 B. h. nilotica 193, 194 B. olivacea 16, 191, 197, 199-202, 203, 205, 206, 315 B. o. akleyorum 199,201 B.o.bocagei 199,201,202 B. o. cupreipennis 199, 201, 202 B. o. olivacea 199,201 B. o. rothschildi 199, 210, 202 B. rara 20, 32, 191, 197, 201, 203-6, 316

Buffnecked Ibis (Theristicus caudatus) 181,183-7,311

20, 32, 173, 177, 179,

Cercibis oxycerca 159, 176-8, 310 Ciconia abdimii 9, 11, 29, 70, 71, 73, 74, 75-9, 82, 84, 85, 88, 102, 137,291 C. boyciana 11, 16, 24, 74, 94, 95, 102, 103-8, 298 C. ciconia 11, 17, 19, 21, 24, 33, 34, 50, 60, 69, 70, 71, 72, 73, 74, 77, 78, 79, 94, 96-102, 103, 105, 106, 107-8, 137, 296-7 C. c. asiatica 97, 98, 100, 102 C. c. boyciana 102 C. c. ciconia 97, 98 C. episcopus 11, 12, 34, 70, 72, 74, 75, 77, 78, 80-5, 87, 88, 89, 90, 95, 102, 292-3 C. e. episcopus 90 C. e. neglecta 90 C. maguari 2, 11, 34, 37, 74, 79, 85, 89, 91-5, 102, 121, 122, 123, 161,295 C. nigra 11,13, 16, 17, 20, 68-74, 75, 77, 79, 82, 83, 84, 94,95, 111,290 C. stormi 1, 11, 16, 27, 74, 83, 84, 85, 86-90, 102, 294 Crested Ibis see Madagascar Crested Ibis; Oriental Crested Ibis Ephippiorhynchus asiaticus 12, 17, 19, 21, 24, 31, 33, 106, 109-13, 115, 118, 122, 123, 166, 224, 299 E: a. asiaticus 114 E. senegalensis 12, 18, 21, 24, 31, 33, 106, 112, 113, 114-18, 122,300 Eudocimus ruber 13, 17, 20, 21, 22, 31, 32, 34, 37, 146-56, 161, 163, 164, 166, 170, 251, 306, 318 E. r. albus 149 E. r. ruber 149 Eurasian Spoonbill (Platalea leucorodia) 3, 17, 20, 26, 213, 252-7, 259, 261, 263, 264, 265, 267, 269, 322-3 Geronticus calvus 17, 20, 29, 34, 73, 236-40, 321 G. eremita 1, 16, 17, 19, 20, 29, 230-5, 319 Giant Ibis (Thaumatibisgigantea) 14, 16, 164, 245-7, 321 Glossy Ibis (Plegadisfalcinellus) 2, 13, 65, 159, 162-7, 170, 171, 172, 177, 189, 227, 231, 241, 308 Greater Adjutant Stork (Leptoptilos dubius) 16, 19, 24, 123, 126, 127,128-32, 135, 138,303 Green Ibis (Mesembrinibis cayennensis) 32, 159, 164,

188-91,312

Hadada Ibis (Bostrychia hagedash) 20, 22, 29, 32, 65, 191, 192-5, 197, 201, 202, 203, 205, 237, 243, 313

384

Index

Harpiprion caerulescens 179-82, 310 Jabiru mycteria 12, 18, 21, 24, 32, 37, 93, 112, 119-23, 301 Jabiru Stork (Jabiru mycteria) 12, 18, 21, 24, 32, 37, 93, 112,119-23,301 Leptoptilos crumeniferus 8, 22, 34, 78, 112, 116, 125, 126, 127, 129, 130, 131, 132, 133-8, 141, 144, 304 L. c. cristata 209 L. c. urschi 207, 209 L. dubius 16, 19, 24, 123, 126, 127, 128-32, 135, 138, 303 L.javanicus 16, 45, 46, 57, 123, 124-7, 129, 130, 131, 135, 302 Lesser Adjutant Stork (Leptoptilos javanicus) 16, 45, 46, 57, 123, 124-7, 129, 130, 131, 135, 302 Lophotibis cristata 16, 202, 207-10, 316 Madagascar Crested Ibis (Lophotibis cristata) 16, 202, 207-10,316 Maguari Stork (Ciconia maguari) 2, 11, 34, 37, 74, 79, 85, 89, 91-5, 102, 121, 122, 123, 161, 295 Marabou Stork (Leptoptilos crumeniferus) 8, 22, 34, 78, 112, 116, 125, 126, 127, 129, 130, 131, 132, 133-8, 141, 144, 304 Mesembrinibis cayennensis 32, 159, 164, 188-91, 312 Milky Stork (Mycteria cinerea) 16, 43-7, 51, 55, 105, 219, 285 Mycteria americana 17, 21, 30, 31, 32, 33, 34, 35-42, 47, 50, 51,93, 121,284 M. cinerea 16, 43-7, 51, 55, 105, 219, 285 M. ibis 11, 22, 25, 46, 48-52, 53, 65, 98, 116, 117, 214, 286 M. leucocephala 22, 30, 43, 45, 50, 51, 53-7, 60, 61, 62, 127,287 Nipponia nippon

16, 19, 21, 59, 218, 248-51, 321

Olive Ibis (Bostrychia olivacea) 16, 191, 197, 199-202, 203, 205,206,315 Openbill Stork see African Openbill Stork; Asian Openbill Stork Oriental Crested Ibis (Nipponia nippon) 16, 19, 21, 59, 218, 248-51,321 Oriental White Ibis (Threskiornis melanocephalus) 6, 28, 30, 46, 166,213,216-20,223,318 Oriental White Stork (Ciconia boyciana) 11, 16, 24, 74, 94, 95, 102, 103-8, 298 Painted Stork (Mycteria leucocephala) 22, 30, 43, 45, 50, 51, 53-7,60,61,62, 127,287 Phimosus infuscatus 24, 94, 157-61, 164, 170, 307 P. i. berlepschi 157, 158 P. i. infuscatus 157, 158, 159 P. i. nudifrons 157, 158 Platalea ajaja 14, 29, 274, 275-81, 328 P. alba 24, 25, 28, 166, 214, 255, 256, 267-70, 277, 280, 326 P.flavipes 8, 14, 43, 166, 260, 261, 271-4, 280, 327

P. leucorodia 3, 17, 20, 26, 213, 252-7, 259, 261, 263, 264, 265, 267, 269, 322-3 P. L archeri 253, 255, 257 P. L balsaci 253, 255, 257 P. L leucorodia 255, 257 P. L regia 257 P. minor 14, 16, 257, 261, 262-6, 325 P. regia 7, 14, 25, 166, 223, 256, 257, 258-61, 264, 265, 266,267,271,273,280,324 Plegadis chihi 13, 159, 161, 164, 166, 168-72, 173, 189, 309 P.falcinellus 2, 13, 65, 159, 162-7, 170, 171, 172, 177, 189, 227,231,241,308 P. f. peregrinus 16 7 P. ridgwayi 13, 172, 173-5, 309 Plumbeous Ibis (Harpiprion caerulescens) 179-82, 310 Pseudibispapillosa 16, 19, 20, 164, 241-4, 245, 247, 320 P.p.davisoni 241,243,244 P. p. papillosa 241,243 Puna Ibis (Plegadis ridgwayi) 13, 172, 173-5, 309 Roseate Spoonbill (Platalea ajaja) 14, 29, 274, 275-81, 328 Royal Spoonbill (Platalea regia) 7, 14, 25, 166, 223, 256, 257, 258-61, 264, 265, 266, 267, 271, 273, 280, 324 Sacred Ibis (Threskiornis aethiopicus) 5, 6, 17, 21, 34, 65, 78, 194, 211-15, 217, 219, 221, 223, 224, 229, 239, 269, 317 Saddlebill Stork (Ephippiorhynchus senegalensis) 12,18,21, 24, 31, 33, 106, 112, 113, 114-18, 122, 300 Sharptailed Ibis (Cercibis oxycerca) 159, 176-8, 310 Shoebill (Balaeniceps rex) 7, 9, 10, 14, 30, 32, 135, 139-45, 305 Spotbreasted Ibis (Bostrychia rara) 20, 32, 191, 197, 201, 203-6,316 Storm's Stork (Ciconia stormi) 1,11,16, 27, 74, 83, 84, 85, 86-90, 102, 294 Strawnecked Ibis (Threskiornis spinicollis) 4, 13, 166, 218, 223,224,226-9,319 Thaumatibis gigantea 14, 16, 164, 245-7, 321 Theristicus caudatus 20, 32, 173, 177, 179, 181, 183-7, 311 T. c. branickii 183, 185, 186 T. c. caudatus 183, 185, 186 T. c. hyperorius 183, 185, 186 T. c. melanopsis 183, 185, 186 Threskiornis aethiopicus 5, 6, 17, 21, 34, 65, 78, 194, 211-15, 217, 219, 221, 223, 224, 229, 239, 269, 317 T.a.abbotti 211,213,214,215 T. a. aethiopicus 5,211,215 T.a.bernieri 5,211,213,214,215 T. melanocephalus 6, 28, 30, 46, 166, 213, 216-20, 223, 318 T. molucca 6, 13, 215, 217, 218, 219, 221-5, 227, 228, 229, 300 T. m. molucca 225 T. m. pygmaeus 221, 223, 225 T. m. strictipennis 225 T. spinicollis 4, 13, 166, 218, 223, 224, 226-9, 319

Index

T. s. faroi 229

Whitefaced Ibis (Plegadis chihi)

385

13, 159, 161, 164, 166,

168-72, 173, 189,309 Waldrapp Ibis (Geronticus eremita) 1, 16, 17, 19, 20, 29, 230-5, 319 Wattled Ibis (Bostrychia carcunculata) 20, 196-8, 314 White Ibis see American White Ibis; Australian White Ibis; Oriental White Ibis White Stork (Ciconia ciconia) 11,17, 19, 21, 24, 33, 34, 50, 60, 69, 70, 71, 72, 73, 74, 77, 78, 79, 94, 96-102, 103, 105, 106, 107-8, 137, 296-7 see also Oriental White Stork

Wood Stork, American 17, 21, 30, 31, 32, 33, 34, 35-42, 47, 50, 51, 93, 121, 284 Woollynecked Stork (Ciconia episcopus) 11,12, 34, 70, 72, 74, 75, 77, 78, 80-5, 87, 88, 89, 90, 95, 102, 292-3 Yellowbilled Spoonbill (Plataleaflavipes) 8, 14, 43, 166, 260, 261, 271-4, 280, 327 Yellowbilled Stork (Mycteria ibis) 11, 22, 25, 46, 48-52, 53, 65, 98, 116, 117, 214, 286