ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI CHRISTCHURCH, NEW ZEALAND 2-9 DECEMBER 1990 f VOLUME I Vi. t Museum of Comparative Zoology Library Harvard University ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI VOLUME I CHRISTCHURCH, NEW ZEALAND 2-9 DECEMBER 1990 20TH INTERNATIONAL ORNITHOLOGICAL CONGRESS LOGO The distinctive logo for the Congress was designed early in 1986 by Karen Pinnel, a Wellington Polytechnic design student. As part of her third year course at the Design School she was asked to design a simple graphic using a native bird from New Zea¬ land. The final graphic chosen by the New Zealand Organising Committee for its sim¬ plicity, style and eye-catching appeal is based on the endemic Yellow-eyed Penguin, but the slightly more golden crest colour is representative of the distinctive crested penguin species concentrated in the New Zealand region. The Organising Commit¬ tee wishes to record its thanks to Karen for her donation of the copyright for the use of the design to the New Zealand Ornithological Congress Trust Board. With penguins only being found in the southern hemisphere this logo is representative of the ‘South¬ ern Perspective’ theme for the 20th International Ornithological Congress. New Zealand 1990 [QFBCIAl PROJECT 1 NEW ZEALAND 1990’ Coordinated and promoted by the New Zealand 1990 Commission, activities under this theme are designed to commemorate a milestone year in the nation’s history. Some of these formal milestones are 1000 years of known habitation of New Zealand; 150 years of the founding of modern government, and the cities of Auckland and Wellington; 100 years of a one person one vote electoral system. Throughout 1990 the country is formally recognising and celebrating the achievements of past and present with a wide range of local and international functions and expositions. Par¬ ticipating events will encourage New Zealanders to highlight and celebrate their natu¬ ral advantages; increase awareness, understanding and appreciation of all cultures in their society; promote harmony, goodwill and tolerance; increase opportunities for sharing and partnership; and encourage thinking about the future. The 20th International Ornithological Congress is an integral part of New Zealand 1990. The New Zealand Ornithological Congress Trust Board is pleased to acknowl¬ edge the support and participation of the 1990 Commission in the Congress. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI CHRISTCHURCH, NEW ZEALAND 2-9 DECEMBER 1990 VOLUME I EDITORIAL SUBCOMMITTEE: BEN D. BELL (CONVENER), R.O. COSSEE, J.E.C. FLUX, B.D. HEATHER, R.A. HITCHMOUGH, C.J.R. ROBERTSON, M.J. WILLIAMS NEW ZEALAND ORNITHOLOGICAL CONGRESS TRUST BOARD National Library of New Zealand Cataloguing-in-Publication data Congressus International^ Ornithologici (20th : 1990 : Christchurch, N.Z.) Acta XX Congressus International^ Ornithologici, Christchurch, New Zealand, 2-9 December 1990. Wellington, N.Z. : New Zealand Ornithological Congress Trust Board, 1990-1991. 4 v. + 1 supplement ISBN 0 - 9597975 - 1 - 3 (Vol I) ISBN 0 - 9597975 - 2 - 1 (Vol II) ISBN 0 - 9597975 - 3 - X (Vol III) ISBN 0 - 9597975 - 4 - 8 (Vol IV) ISBN 0 - 9597975 - 0 - 5 (supplement) ISBN 0 - 9597975 - 5 - 6 (set) 1. Ornithology - Congresses. 2. Birds - Congresses. I. New Zealand Ornithological Congress Trust Board. II. Title. III. Title: Acta Twentieth Congressus International^ Ornithologici : Christchurch, New Zealand, 2-9 December 1990. 598 Reference to material in this volume should be cited thus: Author(s), 1991. Title . Acta XX Congressus International^ Ornithologici: pages. ISBN 0 - 9597975 - 5 - 6 (five-volume set) ISBN 0 - 9597975 - 1 - 3 (Vol. I) MCZ LIBRARY Copyright © New Zealand Ornithological Congress Trust Board OCT 04 m 1991 Published by New Zealand Ornithological Congress Trust Board UNIV P O Box 12397, Wellington, New Zealand / Typeset, printed and bound in New Zealand by Hutcheson, Bowman & Stewart Ltd Wellington The New Zealand Ornithological Congress Trust Board acknowledges support for the publication of this volume from the Science & Research Division, New Zealand De¬ partment of Conservation; Victoria University of Wellington; the New Zealand 1990 Commission; and the New Zealand Lottery Board. PREFACE These Acta XX Congressus Internationalis Ornithologici provide a full and representa¬ tive record of the activities of the 20th Congress held in Christchurch, New Zealand, 2-9 December 1990. Following the tradition of some recent International Ornithological Congresses, the Acta include all symposia papers - as well as plenary papers and business reports. In a departure from tradition, we have included as a Supplement to the Acta, the Programme and Abstracts distributed at the Congress so as to more fully report the scientific content of the Congress. Most of the symposium papers presented are included in these Proceedings. An abstract is presented if a complete manuscript was not received by the due date. We have sought to minimise the time lag between the Congress and the publication of its Proceedings. Authors of published papers were asked to follow strict editorial guidelines. Conveners of the 48 symposia agreed to take on the task of editing and refereeing papers in their symposia. The post-Congress editing of these Acta has thus been substantially reduced. To streamline production, most text was optically char¬ acter read directly from the manuscript for typesetting. Proofs were not returned to authors for checking but were proof-read by the editors. Our policy, advised to contributors when they were sent editorial guidelines, has been to reproduce papers in the form in which they were received, subject only to stand¬ ardisation of typestyle and format during the typesetting process. In general, texts and figures are reproduced in the form the authors submitted them, although some tables have required resetting. If authors did not conform to the recommended style their papers were not changed, except where inconsistency within the paper was evident. We thank conveners and authors for responding to our request to adhere to editorial guidelines and deadlines, and we particularly appreciate the assistance of Prof. Dr. Peter Berthold, Chairman of the Scientific Programme Committee, for negotiating with many conveners and authors on our behalf. Ben D. Bell Secretary-General Convener, Editorial Subcommittee * i I / ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 7 TABLE OF CONTENTS VOLUME I PREFACE 5 INTERNATIONAL ORNITHOLOGICAL CONGRESSES 1884-1990 10 ACTA OF PREVIOUS CONGRESSES 1 1 COMMITTEES OF 20TH CONGRESS Officers 12 Permanent Executive Committee 12 Scientific Programme Committee 12 New Zealand Organising Committee 13 New Zealand Sub-committees 13 MEMBERS OF 20TH CONGRESS 1 5 PROFESSOR HELMUT SICK 1910-1991 61 SIR CHARLES ALEXANDER FLEMING 1 91 6-1987 62 REPORT OF THE SECRETARY-GENERAL 65 REPORTS OF THE I.O.C. STANDING COMMITTEES Standing Committee on Ornithological Nomenclature 84 Standing Committee for the Coordination of Seabird Research 87 Standing Committee on Applied Ornithology 91 XXI CONGRESSUS INTERNATIONALIS ORNITHOLOGICUS AND INTERNATIONAL ORNITHOLOGICAL COMMITTEE 1990-1994 Officers 95 Executive Committee 95 Past Presidents 95 Past Secretaries-General 95 Senior Members 96 National Representatives 96 STATUTES & BY-LAWS OF THE INTERNATIONAL ORNITHOLOGICAL COMMITTEE 99 PATRON S MESSAGE H.R.H. THE PRINCE PHILIP 107 8 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI PLENARY LECTURES PRESIDENTIAL ADDRESS Phylogeny and Classification of Birds from DNA Comparisons C.G. SIBLEY 109 An Ornithological Glimpse into New Zealand’s Prehuman Past I.A.E. ATKINSON & P.R. MILLENER 127 Recent Avifaunal Changes and the History of Ornithology in New Zealand BEN D. BELL 193 Communal Breeding along the Changing Face of Theory J.L. CRAIG 231 Applied Ornithology: Putting Theory and Practice together E.H. BUCHER 247 Respiration of Avian Embryos at High Altitude C. CAREY 263 Ecological and Physiological Constraints on Reproduction in Albatrosses J.P. CROXALL 279 SYMPOSIUM PAPERS SYMPOSIUM 1 Biogeography and Speciation in Neotropical Birds 303 SYMPOSIUM 2 Origins and Evolution of the Australasian Avifauna 357 SYMPOSIUM 3 Ornithogeography of the Pacific Region 417 SYMPOSIUM 4 Systematics and Biogeography of Afrotropical Birds 447 SYMPOSIUM 5 Patterns and Processes of Population Differentiation in Birds 491 SYMPOSIUM 6 The Methodology of Reconstructing the Past 551 SYMPOSIUM 7 Modern Biochemical Approaches to Avian Systematics 589 AUTHOR INDEX 637 VOLUME II • SYMPOSIUM 8 Ecology and Social Behaviour of Parrots and Parakeets 651 SYMPOSIUM 9 Bird Flight 699 SYMPOSIUM 10 New Aspects of Avian Migration Systems 749 SYMPOSIUM 11 Ecological and Evolutionary Consequences of Body Size 789 SYMPOSIUM 12 Ecological and Behavioural Adaptations of Southern Hemisphere Waterfowl 839 SYMPOSIUM 13 The Avian Feeding System 887 SYMPOSIUM 14 Parent-offspring Relationships 927 SYMPOSIUM 15 Brood Parasitism 999 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 9 SYMPOSIUM 16 Filial and Sexual Imprinting 1049 SYMPOSIUM 17 Nocturnality in Birds 1089 SYMPOSIUM 18 Social Organisation of Nectar-feeding Birds 1137 SYMPOSIUM 19 Social Behaviour in the Non-breeding Season 1193 SYMPOSIUM 20 Acquisition and Functions of Avian Vocalisations 1241 AUTHOR INDEX 1287 VOLUME III SYMPOSIUM 21 Co-operative Breeding - A Second Phase 1299 SYMPOSIUM 22 Mating and Mate Choice 1343 SYMPOSIUM 23 Feeding Ecology of Antarctic and Sub-antarctic Seabirds 1375 SYMPOSIUM 24 Mechanisms of Interspecific Competition 1415 SYMPOSIUM 25 The Generality of Community Concepts in Avian Ecology 1459 SYMPOSIUM 26 Long-term Population Studies of Birds 1497 SYMPOSIUM 27 Food Limitation in Breeding Terrestrial Bird Populations 1557 SYMPOSIUM 28 Bird-piant Interactions 1603 SYMPOSIUM 29 Recruitment in Long-lived Birds 1637 SYMPOSIUM 30 Avian Brood Reduction 1701 SYMPOSIUM 31 Adaptations to Extreme Environments 1753 SYMPOSIUM 32 Sensory Basis of Orientation 1801 SYMPOSIUM 33 Physiology of Diving Birds 1851 SYMPOSIUM 34 Pain and Stress in Birds 1901 AUTHOR INDEX 1941 VOLUME IV SYMPOSIUM 35 Avian Energetics 1955 SYMPOSIUM 36 The Avian Pineal 2003 SYMPOSIUM 37 Endocrinology of Avian Breeding Systems 2053 SYMPOSIUM 38 Integrative Aspects of Osmoregulation in Birds 2103 SYMPOSIUM 39 Avian Nutritional Ecology 2147 SYMPOSIUM 40 Habitat Loss: Effects on Shorebird Populations 2195 SYMPOSIUM 41 Seabirds as Monitors of Changing Marine Environments 2237 SYMPOSIUM 42 Bird Conservation at a Landscape Scale 2281 SYMPOSIUM 43 Disease Ecology and the Conservation of Avian Species 2321 SYMPOSIUM 44 Superabundance in Gulls: Causes, Problems and Solutions 2359 SYMPOSIUM 45 Contributions of Captive Breeding to the Conservation of Endangered Species 2399 SYMPOSIUM 46 Genetic Aspects of Population Structure 2431 SYMPOSIUM 47 Birds as Indicators of Global Change 2471 SYMPOSIUM 48 Integrating New Zealand Conservation 2511 AUTHOR INDEX 2563 10 ACTA XX CONGRESSUS INTERN ATIONALIS ORNITHOLOGICI INTERNATIONAL ORNITHOLOGICAL CONGRESSES 1884 - 1990 No. City Year President Secretary-General 1 Vienna 1884 II Budapest 1891 III Paris 1900 IV London 1905 V Berlin 1910 VI Copenhagen 1926 VII Amsterdam 1930 VIII Oxford 1934 IX Rouen 1938 X Uppsala 1950 XI Basel 1954 XII Helsinki 1958 XIII Ithaca 1962 XIV Oxford 1966 XV Den Haag 1970 XVI Canberra 1974 XVII Berlin 1978 XVIII Moscow 1982 XIX Ottawa 1986 XX Christchurch 1990 Dr G.F.R. Radde Prof. Victor Fatio, Otto Herman Dr Emile Oustalet R. Bowdler Sharpe Prof. Dr Anton Reichenow Dr E.J.O. Hartert Prof. Dr A.J.E. Lonnberg Prof. Dr. Erwin Stresemann Prof. Alessandro Ghigi Dr Alexander Wetmore Sir Landsborough Thomson Prof. J. Berlioz Prof. Ernst Mayr Dr David Lack Prof. Dr Nikolaas Tinbergen 1966-1969 Prof. Dr Finn Salomonsen 1969-1970 Prof. Jean Dorst Prof. D.S. Farner Prof. Dr Lars von Haartman Prof. Dr Klaus Immelmann Prof. Charles G. Sibley Dr Gustav von Hayek Jean de Claybrooke Dr E.J.O. Hartert J. Lewis Bonhote Herman Schalow E. Lehn Schioler Prof. Dr L.F. de Beaufort Rev. F.C.R. Jourdain Jean Delacour Prof. Dr Sven Horstadius Prof. Dr Adolf Portmann Dr Lars von Haartman Dr Charles G. Sibley Prof. Dr N. Tinbergen Prof. Dr Karel H. Voous Dr H.J. Frith Rolf Nohring Prof. Dr V. Ilyichev Dr Henri Ouellet Dr Ben D. Bell ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 11 Acta of Previous Congresses I. Sitzungs-Protokolle des ersten Internationalen Ornithologen-Congresses, der vom 7. bis 11. April 1884 in Wien abgehalten wurde. Wien, Verlag des Ornithologischen Vereines in Wien, 1884. vi + [90] pp. Mitteilungen des Ornithologischen Vereins Wien, Band viii-x, 1884-86. II. Bericht . . . Zweiter International Ornithologischer Congress, Budapest, 1892. (Blasius) [n.p.; 1891] 58 pp. III. Ill6 Congres Ornithologique International, Paris, 26-30 juin 1900. Compte rendu des seances publie par E. Oustalet . . . et J. de Claybrooke . . . Masson et Cie, Paris, xii + 503 pp. 1901 . [= Ornis, vol. 1 1] IV. Proceedings of the IVth International Ornithological Congress, London, June 1905. Edited by R.B. Sharpe, E.J.O. Hartert, J.L. Bonhote. Dulau & Co., London. 696 pp. 1907. [= Ornis, vol. 14] V. Verhandlurigen des V. Internationaler Ornithologen-Kongresses, Berlin, 30. Mai bis 4. Juni 1910. Herausgegeben von Herman Schalow . . . Deutsche Ornithologische Gesellschaft, Berlin, x + 1186 pp. 1911. VI. Verhandlungen des VI. Internationalen Ornithologen-Kongresses in Kopenhagen, 1926. Herausgegeben von Dr. F. Steinbacher. Berlin, vi + 641 pp. 1929. VII. Proceedings of the Vllth International Ornithological Congress at Amsterdam. Amsterdam, vii + 527 pp. 1931. VIII. Proceedings of the VII Ith International Ornithological Congress, Oxford, July 1934. Edited by F.C.R. Jourdain. Oxford University Press, Oxford, x + 761 pp. 1938. IX. IX6 Congres Ornithologique International, Rouen, 9 au 13 mai 1938. Compte rendu publie par Jean Delacour . . . Rouen. 543 pp. 1938. X. Proceedings of the Xth International Ornithological Congress, Uppsala, June 1950. Edited by Sven Horstadius. Almqvist & Wiksells, Uppsala. 662 pp. 1951. XI. Acta XI Congressus Internationalis Ornithologici, Basel, 29. V.- 5. VI. 1954. Herausgegeben von Adolf Portmann und Ernst Sutter. Birkhauser Verlag, Basel und Stuttgart. 680 pp. 1955. XII. Proceedings of the XI Ith International Ornithological Congress, Helsinki, 5. - 12. VI. 1958. Edited by G. Bergmann, K.O. Donner, L. v. Haartmann. Tilgmannin Kirjapaino, Helsinki. 2 vols. 820 pp. 1960. XIII. Proceedings of the XI I Ith International Ornithological Congress, Ithaca, 17-24 June 1962. Edited by Charles G. Sibley, Joseph J Hickey and Margaret B. Hickey. Published by the American Ornithologists’ Union. 2 vols. xvi + 1246 pp. 1963. XIV. Proceedings of the XIVth International Ornithological Congress, Oxford, 24-30 July 1966. Edited by D.W. Snow. Blackwell Scientific Publications, Oxford and Edin¬ burgh. xxiv + 405 pp. 1967. XV. Proceedings of the XVth International Ornithological Congress, The Hague, 30 August - 5 September 1970. Edited by K.H. Voous. E.J. Brill, Leiden, viii + 745 pp. 1972. XVI. Proceedings of the XVIth International Ornithological Congress, Canberra, 12- 17 August 1974. Edited by H.J. Frith and J.H. Calaby, Australian Academy of Science, Canberra, xviii + 765 pp. 1976. XVII. Acta XVII Congressus Internationalis Ornithologici, Berlin, 5-11. VI. 1978. Herausgegeben von Rolf Nohring. Verlag der Deutschen Ornithologen-Gesellschaft, Berlin. Vol I, pp. 1-747; vol. II, pp. 756-1463. 1980. XVIII. Acta XVIII Congressus Internationalis Ornithologici, Moscow, August 16-24, 1982. Edited by V.D. Ilyichev and V.M. Gavrilov. “Nauka”, Moscow. Vol. I, pp. 1-576; vol. II, pp. 577-1335. 1985. XIX. Acta XIX Congressus Internationalis Ornithologici, Ottawa, 22-29. VI. 1986. Edited by Henri Ouellet. Published for National Museum of Natural Sciences by Uni¬ versity of Ottawa Press, Ottawa. Vol. I, pp. 1-1404; vol II, pp. 1405-2815. 1988. 12 ACTA XX CONGRESSUS INTERN ATIONALIS ORNITHOLOGICI XX CONGRESSUS INTERN ATIONALIS ORNITHOLOGICUS Patron H.R.H. The Prince Philip Honorary President Dr N.K. Kuroda Japan Honorary Vice-Presidents Professor W. Hsu China Professor H. Sick Brazil President Professor Charles G. Sibley USA Vice-President Professor J.K. Pinowski Poland Secretary-General Dr Ben D. Bell New Zealand Permanent Secretary, International Ornithological Committee Professor Walter J. Bock USA COMMITTEES Permanent Executive Committee 1986-1990 Chairman: Prof C. G. Sibley USA Vice-Chairman: Prof Dr J.K. Pinowski Poland Dr B.D. Bell New Zealand Prof W.J. Bock USA Dr C. Erard France Dr S. Haftorn Norway Dr E.N. Kurochkin USSR I.C.R. Rowley Australia Professor Dr K. Immelmann Prof Dr P. Berthold Germany Dr E.H. Bucher Argentina Dr B.K. Follett United Kingdom Prof J.R. King USA Dr H. Ouellet Canada Germany until his death in 1987 Scientific Programme Committee 1986-1990 Chairman: Prof Dr P. Berthold Germany Vice-Chairman: Dr M.J. Williams New Zealand Dr B. D. Bell New Zealand Dr E.H. Bucher Argentina Dr L.S. Davis New Zealand Dr H. Masatomi Japan Dr C.M. Perrins United Kingdom Prof C. G. Sibley USA Prof W. J. Bock USA Dr J.L. Craig New Zealand Dr S. Haftorn Norway Dr H. Ouellet Canada I.C.R. Rowley Australia Dr L. Tomialojc Poland ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 1 New Zealand Organising Committee Ben D. Bell (Chairman)* M.N. Clout C.J.R. Robertson* S.M. Usher* Brian D. Bell* L. J. Fairbairn H.A. Robertson* M. J. Williams* B. Brown A. Hudson P.M. Sagar B. G. Wybourne P.C. Bull* R.G. Powlesland* R. B. Thomson Sir Charles Fleming served on the Committee until his death in 1987. * NZ Executive Committee and NZ Ornithological Congress Trust Board members Subcommittees of New Zealand Organising Committee Business Management Subcommittee C.J.R. Robertson (Convener) C. Cheer L. Fairbairn A. Hudson D.E. Hurley S.M. Usher Christchurch Local Organising Subcommittee R.B. Thomson (Convener) P. Bell C.N. Challies I.G. McLean E.B. Spurr C.F.J. O’Donnell J. West P.M. Sagar K-J. Wilson Ben D. Bell (Convener) B.D. Heather Editorial Subcommittee R.O. Cossee J.E.C. Flux R.A. Hitchmough C.J.R. Robertson M.J. Williams Excursions Subcommittee Brian D. Bell (Convener) C.J.R. Robertson H.A. Robertson Film Review Subcommittee K. Westerskov (Convener) L.S. Davis N. Harraway Grant Review Subcommittee Ben D. Bell (Convener) Brian D. Bell C.J.R. Robertson M.J. Williams IOC-ICBP Liaison Subcommittee M.N. Clout (Convener) Brian D. Bell R. Hay P.J. Moors C.J.R. Robertson Publications Subcommittee R.G. Powlesland (Convener) R.O. Cossee R. Pickard C.J.R. Robertson Publicity & Circulars H. A. Robertson (Convener) A. Ballance J. Cockrem J.E.C. Flux B.D. Heather M.J. Meads Subcommittee R.E. Brockie J.R. Hay R.G. Powlesland Professional Conference Organisers Conference Makers Limited P.O. Box 9126, Newmarket Auckland 14 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI - ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 15 MEMBERS OF THE 20th INTERNATIONAL ORNITHOLOGICAL CONGRESS Aasgaard, Mr Arne, Norwegian Broadcasting Corporation, Radio Norway, 0340 Oslo 3, Norway Able, Dr Kenneth P., State University of New York at Albany, 1400 Washington Av¬ enue, Albany, N.Y. 12222, U.S.A. Able, Mrs Mary A., 1400 Washington Avenue, Albany, N.Y. 12222, U.S.A. Abs, Dr Michael, Ruhr-Universitat, LS for General Zoology & Neurobiology, Box 10 2148, Bochum, D-04360, Germany Academy Press, P.O. Box 8900, Symonds Street, Auckland, New Zealand Adam, Ms Elizabeth M., 20 South Duncan Avenue, Fayetteville, AR 72701, U.S.A. Adams, Ms Lynn, 24 Jordan Avenue, Ashburton, New Zealand Alatalo, Dr Rauno, University of Jyvaskyla, Department of Biology, Yliopistonkatu 9, Jyvaskyla 40100, Finland Alexander, Mr David, 49 Coopers Road, Shirley, Christchurch, New Zealand Allen, Mr David, MAF Fisheries, Ministry of Agriculture & Fisheries, P.O. Box 3437, Auckland, New Zealand Allen, Ms Jan, 12 Oku Street, Island Bay, Wellington, New Zealand Alves, Miss Vania S., Universidade Federal do Rio de Janeiro , Instituto de Biologia - Zoologia, Cidade Universitaria - llha do Fundao, Rio de Janeiro, 21 941 , Brazil Amundsen, Mr Trond, University of Oslo, Department of Biology, Division of Zoology, Box 1050, N-0316 Oslo 3, Norway Anderson, Mr Andy, Box 1, Tarras, Otago, New Zealand Anderson, Mr Alf-lnge T., Helmershusv, Box Number 107-1, Kristianstad 29194, Swe¬ den Anderson, Dr John G.T., College of the Atlantic, 105 Eden Street, Bar Harbor, ME 04609, U.S.A. Anderson, Ms Karen B., College of the Atlantic, 105 Eden Street, Bar Harbor, ME 04609, U.S.A. Anderson, Mr Peter J., Department of Conservation New Zealand, P.O. Box 842, Whangarei, New Zealand Anderson, Ms Hayley, Earthwatch, P.O. Box C360, Clarence Street, Sydney, NSW 2000, Australia Anderson, Ms Betty, Alaska Biological Research, P.O. Box 84338, Fairbanks, 99208, Alaska, U.S.A. Anderssen, Ms Jorid, C /- Dr G W Gabrielsen, Norwegian Institute for Nature Re¬ search, Haakon Vll’s 6T 5A, Tromso 9000, Norway Andersson, Mr Niis A., Abisko Scientific Research Station, P.O. Box 52, S-980 24 Abisko, Sweden Andors, Dr Allison V., American Museum of Natural History, Department of Ornithol¬ ogy, Central Park West at 79th Street, New York, NY 10024-5192, U.S.A. Andrew, Mr John J., Department of Conservation New Zealand, Private Bag, Christchurch, New Zealand Angelstam, Dr Per, Grimso Wildlife Research Station, Riddarhyttan, S-77031, Swe¬ den Ankney, Dr Dave, University of Western Ontario, Department of Zoology, London, Ontario, N6A 5B7, Canada 16 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Antram, Mr Frank, Traffic Oceania, P.O. Box R594, Royal Exchange, Sydney, N.S.W. 2000, Australia Arctander, Dr Peter, University of Copenhagen, Institute of Population Biology, Universitetsparken 15, DK-2100 Copenhagen, Denmark Armstrong, Dr Doug, University of Sydney, School of Biological Sciences A08, Syd¬ ney, NSW 2006, Australia Armstrong, Mr Barry, 60 Frankleigh Street, Spreydon, Christchurch, New Zealand Arnold, Mr Todd W., University of Western Ontario, Department of Zoology, London, Ontario, N6A 5B7, Canada Arrowood, Dr Patricia C., New Mexico State University, Department of Biology, Box 30001 Department 3AF, Las Cruces, New Mexico 8800, U.S.A. Arvidsson, Mr Bjorn, University of Gothenburg, Department of Zoology, P.O. Box 25059, S-400 31 Gothenburg, Sweden Ash, Mrs J.W., Godshill Wood, Nr. Fordingbridge, Hants. SP62LR, United Kingdom Ash, Dr John S., Godshill Wood, Nr. Fordingbridge, Hants. SP62LR, United Kingdom Atkinson, Dr Ian, N.Z. Department of Scientific & Industrial Research, C/- DSIR Land Resources, Private Bag, Lower Hutt, New Zealand Aubin, Mr Thierry, Laboratoire D’Ethologie Experimentale, Saint Lucien, 28210 Nogent-Le-Roi, France Aulen, Dr Gustaf, Swedish Ornithological Society , Box 142 19, S-104 40, Stockholm, Sweden Austen, Ms Madeline, P.O. Box 103, Campbellville, Ontario, LOP 1B0, Canada Avise, Prof John C., University of Georgia, Department of Genetics, Athens, Georgia, 30602, U.S.A. Baha El Din, Ms Mindy, ICBP Project Co-ordinator, Executive Business Service, Cairo Marriott Hotel, P.O. Box 33, Zamalek, Cairo, Egypt Baines, Mr Alun, 41 Marion Street, Macandrew Bay, Dunedin, New Zealand Bairlein, Prof. Franz, University of Koeln, Zoology Institute, Weyertal 119, Koeln 41, D-5000, Germany Baker, Mr David, O.S.N.Z., 103 Campbell Road, Auckland 6, New Zealand Baker, Prof. Allan J., Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6, Canada Baker-Gabb, Dr David, Department Conservation and Environment, 123 Brown Street, Heidelberg, Victoria 3084, Australia Ball, Dr Gregory F., Boston College, Department of Psychology, 140 Commonwealth Avenue, Chestnut Hill, MA 02167, U.S.A. Balmford, Mrs Rosemary, 459 The Boulevard, East Ivanhoe, 3079, Australia Bankovics, Dr Attila, Hungarian Natural History Museum, Baross u. 13, Budapest, H- 1088, Hungary Bannasch, Dr Rudolf, Akademie der Wissenschaften der DDR, Forschungsstelle fur Wirbeltierforschung, Am Tierpark 125, Berlin, D - (0) 1136, Germany Baptista, Dr Luis, California Academy of Sciences, Department of Ornithology, Golden Gate Park, San Francisco, 94118, U.S.A. Barber, Mr Lindsay, 49 Oakley Crescent, Christchurch, New Zealand Barfknecht, Dr Ralf, Wolfskaul 2, 5000 Koln 80, Germany Barker, Mr David A., Department of Conservation New Zealand, P.O. Box 86, Hokitika, New Zealand Barkla, Mr John, Department of Conservation New Zealand, Private Bag, Wanganui, New Zealand Barlow, Mrs Maida, 38 Filleul Street, Invercargill, New Zealand ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 17 Barlow, Dr Jon C., Royal Ontario Museum, Department of Ornithology, 100 Queen’s Park, Toronto, Ontario M5S 2C6, Canada Barnes, Miss Mercia, O.S.N.Z., Milwood Building, Collingwood Street, Hamilton, New Zealand Barrett, Mr Geoffrey W., University of New England, Department of Zoology, Armidale, NSW 2351, Australia Barrowclough, Dr George F., American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, U.S.A. Bartle, Mr John A. (Sandy), National Museum of New Zealand, P.O. Box 467, Wel¬ lington, New Zealand Bartlett, Mr Ken, 242c Main Road, Redcliffs, Christchurch 8, New Zealand Barton, Mrs Edith, 29 Ridge Road, Fairhaven, Victoria 3221, Australia Bassett, Dr Terence H., 31 6-5th Street South, Lethbridge, TIJ 2B5, Alberta, Canada Bateson, Prof Paul P G, University of Cambridge, Sub-Department of Animal Behav¬ iour, High Street, Madingley, Cambridge CB3 8AA, United Kingdom Batten, Dr Leo, Nature Conservancy Council, Northminster House, Peterborough, Cambridge, PEI 1UA, United Kingdom Battley, Mr Phil, 163 College Street, Palmerston North, New Zealand Baumann, Ms Sabine, Universitat Kiel, Institut fur Haustierkunde, Olshausenstrasse 40, D-2300 Kiel, Germany Baverstock, Dr Peter, University of New England Northern Rivers, Military Road, East Lismore, NSW 2480, Australia Beason, Dr Robert, State University of New York, Biology Department, Geneseo, NY 14454, U.S.A. Beattie, Ms Jane, 242c Main Road, Redcliffs, Christchurch, New Zealand Bech, Dr Claus, University of Trondheim, Department of Zoology, Dragvoll, Trondheim, N-7055, Norway Bednarz, Dr James C., Hawk Mountain Sanctuary Association, Route 2, Kempton 19529, U.S.A. Beggs, Ms Jacqueline, Cnr Milton & Halifax Streets, Private Bag, Nelson, New Zea¬ land Beichle, Mr Ulf, Institut fur Haustierkunde, Olshausenstrasse 40, D-2300, Kiel, Ger¬ many Beigel-Heuwinkel, Mrs Ursula, Universitat Munster, Erlanger Str. 37, D-4000 Dusseldorf 13, Germany Beissinger, Dr Steven R., Yale University, Forestry & Environmental Studies, New Haven, CT 06511, U.S.A. Beletsky, Dr Les, University of Washington, Department of Zoology, NJ15, Seattle, WA, 98195, U.S.A. Belinsky, Ms Ann, Tel Aviv University, Department of Zoology, Ramat Aviv, Tel Aviv 09778, Israel Bell, Mr Brian D., Ornithological Society of New Zealand, P.O. Box 12397, Welling¬ ton, New Zealand Bell, Dr Ben D., Victoria University, School of Biological Sciences, P.O. Box 600, Wellington, New Zealand Bell, Mrs Gillian F., 66 Seatoun Heights Road, Seatoun, Wellington 3, New Zealand Bell, Mr Martin, D.O.C. National Wildlife Centre, R.D.1, Masterton, New Zealand Bell, Mr Paul, 9 Ferry Street, Seatoun, Wellington, New Zealand Bell, Mr David, 9 Ferry Street, Seatoun, Wellington, New Zealand Bell, Mr Michael, 9 Ferry Street, Seatoun, Wellington, New Zealand 18 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Bell, Mrs Patricia, 70 Woodbury Street, Russley, Christchurch, New Zealand Bellingham, Mr Mark, Forest and Bird Society, Box 728, Wellington, New Zealand Belton, Mr William, Rocky Hollow, Great Cacapon, WV, 25422, U.S.A. Belton, Mrs Julia, Rocky Hollow, Great Cacapon, WV, 25422, U.S.A. Benavides, Dr Nancy, ICBP - Ecuador, Calle Hullgria 275, Vaucouver, Quito, Box 9068 5-7, Ecuador Bendell, Prof. James, University of Toronto, Faculty of Forestry, 33 Willcocks Street, Toronto, Ontario M5S 3B3, Canada Bendell, Mrs Yvonne, 469 Apple Lane, Mississauga, Ontario, L5J 2T2, Canada Benito Espinal, Dr Edouard, AGETL-IGEROC, BP 795, Pointe a Pite, 97173, Guadeloupe Bennett, Mr Andy, Department of Zoology, South Parks Road, Oxford, 0X1 3PS, United Kingdom Bennun, Dr Leon A., National Museums of Kenya, Box 21149, Nairobi, Kenya Berg, Mr Ake, University of Agricultural Sciences, Department of Wildlife Ecology, Box 7002, Uppsala, 750 07, Sweden Berger, Mr Rudolf H., Hockegasse 55-57/St. 1, A-1180, Vienna, Austria Berger, Mrs Margaretha, Hockegasse 55-57/St. 1, A-1180, Vienna, Austria Bering Andersen, Mrs Else, Zoological Museum, Universitetsparken 15, Copenhagen, DK-2100, Denmark Berkhoudt, Dr Herman, Zoological Laboratory, Box 9516, 2300 RA Leiden, Nether¬ lands Berthold, Prof. Peter, Max-Planck-lnst., Vogelwarte, Schloss Moeggingen, Radolfzell, 7760, Germany Bertrand, Dr Gerard, The Massachusetts Audubon Society, 297 Sagamore Street, Hamilton, 01982, U.S.A. Best, Mr Hugh, Department of Conservation New Zealand, P.O. Box 10-420, Welling¬ ton, New Zealand Beudels-Jamar de Bolsee, Mrs Roseline C., Institut Royal des Sciences Naturelles de Belgique, 29, Rue Vautier, Bruselles, 1040, Belgium Bevanger, Dr Kjetil, Norwegian Institute for Nature Research, Tungasletta 2, Trondheim N-7004, Norway Bezzel, Dr Einhard, Institut fur Vogelkunde, Gsteigstrasse 43, Garmisch- Partenkirchen D-8100, Germany Biber, Dr Olivier, Swiss Institute of Ornithology, CH-6204 Sempach, Switzerland Biebach, Dr Herbert, Max-Planck-lnstitut fur Verhaltensphysiologie, Vogelwarte, Andechs, D-8138, Germany Bierregaard, Dr Rob, Smithsonian Institution, World Wildlife Fund, NHB 106, Wash¬ ington, DC 20560, U.S.A. Binsbergen, Mr Andri, Ministry of Agriculture, Nature Management & Fisheries, Prinses Irenelaan 323, Rijswijk, 2285 GA, Netherlands Bird, Prof. David M., McGill University, Raptor Research Centre, 21,111 Lakeshore Road, Ste. Anne de Bellevue H9X ICO, Quebec, Canada Birks, Ms Sharon, Cornell University, Mudd Hall, Ithaca, NY 14853, U.S.A. Birt-Friesen, Ms Vicki, Memorial University of Newfoundland, Department of Psychology, St Johns, Newfoundland, A1B 3X9, Canada Bishop, Mr K. David, Victor Emanuel Nature Tours Inc., “Semioptera”, Lot 15, Kerns Road, Kincumber, New South Wales 2251, Australia Black, Mrs Mary, 43 Wairarapa Terrace, Fendalton, Christchurch, New Zealand Blanchard, Dr Kathleen A., Quebec-Labrador Foundation, Atlantic Center for the En¬ vironment, 39 South Main Street, Ipswich, MA 01938, U.S.A. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 19 Blokpoel, Dr Hans, Canadian Wildlife Service, 49 Camelot Drive, Nepean, KIA 0H3, Canada Bloxham, Mr Arnold, Arnold’s Agencies, P.O. Box 5302, Palmerston North, New Zea¬ land Boag, Dr Peter T., Queen’s University, Department of Biology, Kingston, Ontario, K7L 3N6, Canada Boag, Dr David, University of Alberta, Department of Zoology, Edmonton, T6G 2E9, Canada Bock, Mrs Katharine L., Columbia University, C /- Walter J. Bock, Department of Bio¬ logical Sciences, New York, NY 10027, U.S.A. Bock, Prof. Walter J., Columbia University, Department of Biological Sciences, New York, NY 10027, U.S.A. Boere, Dr Gerard C., Ministry of Agriculture, Nature Management & Fisheries, C /- National Forest Service, P.O. Box 20020, Utrecht 3502LA, Netherlands Boersma, Dr P. Dee, University of Washington, Institute for Enviromental Studies, FM- 12, Seattle, WA, U.S.A. Bogel, Mr Ralf, Nationalpark-Administration, Doktorberg 6, 8240 Berchtesgaden, Germany Boles, Mr Walter E., Australian Museum, 6-8 College Street, Sydney, NSW 2000, Australia Bolton, Mr Mark, University of Glasgow, Zoology Unit, Glasgow G12 8QQ, United Kingdom Bonniface, Ms Adrienne, 25 Rountree Street, llam, Christchurch, New Zealand Booth, Mrs Louise, 128 Glandovey Road, Fendalton, Christchurch, New Zealand Bosque, Dr Carlos, Universidad Simon Bolivar, Dept. Biologia De Organismos, Apt. 89000, 1080 Caracas, Venezuela Boyd, Ms Shaarina, Department of Conservation New Zealand, Private Bag 8, Auck¬ land, New Zealand Bradfield, Mr Philip, Department of Conservation, Kokako Cottage, Mapara South Road, R D 3, Te Kuiti, New Zealand Bradley, Dr J. Stuart, Murdoch University, South Street, Murdoch, Perth, 6150, West¬ ern Australia, Australia Bradley, Mrs Diana, 53 Osterley Road, Isleworth, TW7 4PW, United Kingdom Bradley, Dr Richard A., Ohio State University, 1465 Mt Vernon Avenue, Marion, OH 44302, U.S.A. Bradley, Dr David W., California State University, Academic Computing Services, 1250 Bellflower Boulevard, Long Beach, CA 90840, U.S.A. Bradshaw, Mr Harry, Oxford University, Animal Behaviour Research Group, Depart¬ ment of Zoology, South Parks Road, Oxford OX1 3PS, United Kingdom Braithwaite, Ms Victoria, Oxford University, Animal Behaviour Research Group, De¬ partment of Zoology, Southparks Road, Oxford, OX1 3PS, United Kingdom Braun, Dr Eldon J., University of Arizona, Tucson, Arizona, U.S.A. Braun, Mrs Shirley, C /- Dr Eldon J. Braun, University of Arizona, Tucson, Arizona, U.S.A. Braun, Dr Michael, Smithsonian Institution, Laboratory of Molecular Systematics, Museum Support Center, Washington, DC 20560, U.S.A. Brauning, Mr Daniel, The Academy of Natural Sciences, 19th and The Parkway, Phila¬ delphia, PA 19103, U.S.A. Breitwisch, Dr Randall, University of Dayton, Department of Biology, 300 College Park, Dayton, Ohio 45469, U.S.A. 20 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Brejaart, Ms Ria, 10 College Road, Lyttelton, New Zealand Brieschke, Ms Heike, Alexander Koenig Zoological Institute & Zoological Museum, Adenauerallee 150-164, Bonn, 5300, Germany Briggs, Ms Sue V., CSIRO, National Parks & Wildlife Service, P.O. Box 84, Lyneham, ACT 2602, Australia Brindley, Miss Emma, Nottingham University, Animal Behaviour Research Group, University Park, Nottingham, NG7 2RD, United Kingdom Brisbin, Mrs Brenda, 1220 Evans Road, Aiken, SC 29801, U.S.A. Brisbin Jr., Dr I. Lehr, Savannah River Ecology Laboratory, P.O. Drawer E, Aiken, SC 29801, U.S.A. Briskie, Mr James V., Queen’s University, Department of Biology, Kingston, Ontario, K7L 3N6, Canada Brom, Dr Tim G., University of Amsterdam, Institute of Taxonomic Zoology, P.O. Box 4766, Amsterdam 1009 AT, Netherlands Broni, Mr Stephen C., Department of Conservation New Zealand, P.O: Box 5244, Dunedin, New Zealand Brooke, Dr Michael, Cambridge University, Department of Zoology, Downing Street, Cambridge, CB2 3EJ, United Kingdom Brooke-White, Ms Julia, 129 Aro Street, Aro Valley, Wellington, New Zealand Brouwer, Mrs Sara, 37A Mountain View Road, Mount Albert, Auckland, New Zealand Brown, Dr Richard, Kansas State University, Department of Anatomy & Physiology, College of Vet. Med., Manhattan, Kansas 66506, U.S.A. Brown, Mr George, 28 John Street, Stokes Valley, New Zealand Bruce, Mr Murray, Biocon Research Group, 8 Spurwood Road, Turramurra, N.S.W. 2074, Australia Brugger, Dr Kristen E., U.S. Department of Agriculture, Denver Wildlife Research Centre, 2820 East University Avenue, Gainesville, FL 32601, U.S.A. Brumley, Ms Cathie, Department of Conservation, C /- E Kennedy, Private Bag, Christchurch, New Zealand Brummermann, Dr Marjarethe, Boeselager Strasse 6, W-4600, Dortmund, Germany Bruno, Dr Massa, Intituto de Zoologia, Via Archirafi 18, 90123 Palermo, Italy Bryant, Prof David M., University of Stirling, Biological Sciences, Stirling, FK9 4LA, United Kingdom Bucher, Dr Terry, University of California, Department of Biology, 405 Hilgard Avenue, Los Angeles, CA 90024, U.S.A. Bucher, Dr Enrique, Colorado State University, Department of Biology, U.S.A. Bucher, Mrs Elizabeth, C /- Colorado State University, Department of Biology, U.S.A. Buck, Dr Hugh, Pitman-Moore International Inc, 3 Pinggir Ridley, 55000, Kuala Lumpur, Malaysia Budgey, Ms Helen, The Open University, Box 49, Walton Hall, Milton Keynes, MK7 6AA, United Kingdom Buhler, Dr Paul, Institute fur Zoologie, University Hohenheim, D-7000, Stuttgart 70, Germany Buhlmann, Dr Jost, Else-Zublinstrasse 1, Zurich, CH-8047, Switzerland Buitron, Prof. Deborah, North Dakota State University, Fargo, ND 58105, U.S.A. Bull, Dr Peter, Ornithological Society of New Zealand, 131a Waterloo Road, Lower Hutt, New Zealand Bullock, Mrs Irene K.J., Department of Conservation New Zealand, P.O. Box 10-420, Wellington, New Zealand Burbidge, Dr Allan H., W Australia Dept of Conservation & Land Management, Box Number 51, Wanneroo, W.A. 6065, Australia ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 21 Burger, Dr Ivan H., Waltham Centre For Pet Nutrition, Freeby Lane, Waltham-On-The- Wolds, Melton Mowbray, Leicestershire, LE14 4RT, United Kingdom Burger, Dr Joanna, Rutgers University, Department of Biology, Piscataway, NJ 08855, U.S.A. Burgess, Dr Elizabeth, University of Wisconsin, 2015 Linden Dr. West, Madison, Wis¬ consin, 53706, U.S.A. Burke, Dr Terry, University of Leicester, Department of Zoology, University Road, Leicester, LEI 7RH, United Kingdom Burley, Dr Nancy, University of Illinois, Department of Ecol., Ethol., & Evol., 606 E. Healey Street, Champaign IL, U.S.A. Butz, Mr Oliver, Interconvention, C /- Mr John Dittami, Institute F. Zoologie, Althanstrasse 14, Vienna A-1090, Austria Buurma, Mr Luit S., RNLAF, Head Section Ornithology, P.O. Box 20703, 2500 ES The Hague, Netherlands Byrd, Dr Mitchell A., College of William and Mary, Department of Biology, Williamsburg, Virginia, U.S.A. Cade, Prof. Tom J., Boise State University & The Peregrine Fund Inc., 5666 West Flying Hawk Lane, Boise, ID 83709, U.S.A. Cade, Mrs Renetta M., 6484 Hollilynn Drive, Boise, ID 83709, U.S.A. Cairns, Dr David, Department of Fisheries & Oceans, Science Branch, Box 5030, Moncton, NB EIC 9B6, Canada Caithness, Mr Tom, 32 Mawson Street, Lower Hutt, New Zealand Cameron, Ms Heather, University of Canterbury, Department of Zoology, Christchurch, New Zealand Cameron, Miss Margaret A., RAOU, 4 Connor Street, East Geelong, Vic 3219, Aus¬ tralia Capparella, Dr Angelo P., Illinois State University, Department of Biological Sciences, Normal, Illinois, 61761, U.S.A. Carey, Dr Cynthia, University of Colorado, Department of EPO Biology, Box 334, Boulder, 80309, U.S.A. Carlson, Dr Allan, Department of Wildlife Ecology, Box 7002, 75007 Uppsala, Swe¬ den Carpenter, Prof. F. Lynn, University of California, Department of Eco. Evo. Biology, Irvine, California 92717, U.S.A. Carter, Mr Clide, P.O. Box 71793, Ndola, Zambia Carter, Mrs Loretta E., P.O. Box 71793, Ndola, Zambia Cash, Mr William (Bill) F., Department of Conservation New Zealand, P.O. Box 161, Picton, New Zealand Casotti, Mr Giovanni, Murdoch University, School of Veterinary Studies, South Street, Murdoch, Perth, Western Australia 6150, Australia Cassidy, Ms Alice, University of British Columbia, Department of Zoology, 6270 Uni¬ versity Blvd., Vancouver, B.C., V6T 2A9, Canada Cates, Ms Iris, 378 East King Street Extension, Hillsborough, North Carolina, 27228, U.S.A. Catterall, Dr Carla, A.E.S. Griffith University, Nathan, Brisbane, 4111, Australia Challies, Dr Chris, 22A Highfield Place, Avonhead, Christchurch, New Zealand Chambers, Mrs Alison, 649 River Road, Hamilton, New Zealand Chambers, Dr Geoffrey K., Victoria University, School of Biological Sciences, P.O. Box 600, Wellington, New Zealand Chandola-Saklani, Dr Asha, Garhwal University, Box 45, 246174 Srinagar Garhwal, India 22 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Cheng, Dr Kimberly M., University of British Columbia, Department of Animal Science, Suite 227 - 2357 Main Mall, Vancouver B.C., V6T 2A2, Canada Chorlton, Mr Roger J., South Makara Road, Makara, Wellington 5, New Zealand Christian, Dr Pete, CSIRO, Division of Entomology, P.O. Box 1700, Canberra, 2601, Australia Christidis, Dr Les, Museum of Victoria, Department of Ornithology, 71 Victoria Cres¬ cent, Abbotsford, Melbourne, 3067, Australia Church, Jane P., P.O. Box 39, Sonoita, 85637, U.S.A. Clark, Mr Robert, 135 Peterborough Street, Christchurch, New Zealand Clarke, Dr Mike, McGill University, Department of Biology, 1205 Avenue Docteur Penfield, Montreal PQ, H3A 1B1, Canada Clayton, Dr Nicky, E.G.I, Department of Zoology, South Parks Road, Oxford, OX1 3PS, United Kingdom Clements, Dr James F., 2001 North Soto Street, Los Angeles, California, 90032, U.S.A. Clout, Dr Mick, Department of Conservation New Zealand, P.O. Box 10-420, Welling¬ ton, New Zealand Cockrem, Dr John, Massey University, Department of Physiology & Anatomy, Palmerston North, New Zealand Cogswell, Dr Howard L., California State University Hayward, 1548 East Avenue, Hayward, CA 94541, U.S.A. Cogswell, Mrs Bessie W., 1548 East Avenue, Hayward, CA 94541, U.S.A. Cohn, Dr Jean W., 4787 Beaumont Drive, La Mesa, CA 92041 , U.S.A. Colbourne, Mr Rogan, Department of Conservation New Zealand, 15 Seavista Drive, Pukerua Bay, Wellington, New Zealand Collias, Prof. Nicholas, University of California, 405 Hilgard Avenue, Westwood, Los Angeles, California, 90024-1606, U.S.A. Collias, Dr Elsie, University of California, 405 Hilgard Avenue, Westwood, Los Ange¬ les, California, 90024-1606, U.S.A. Collier, Dr Kevin J., Department of Conservation New Zealand, P.O. Box 10-420, New Zealand Collins, Dr Charles T., California State University, 1250 Bellflower Blvd., Long Beach, CA, 90840, U.S.A. Collins, Mrs Patricia H., 6001 Fairbrook Street, Long Beach, CA, 90815, U.S.A. Collins, Prof. Brian G., Curtin University, School of Biology, GPO Box U 1987, Bent¬ ley, Perth 6102, Australia Conant, Prof Sheila, University of Hawaii, 2450 Campus Road, Honolulu, Hawaii, HI96822, U.S.A. Congdon, Mr Brad, Griffith University, Australian Environmental Studies, Nathan, Bris¬ bane, 4111, Australia Constable, Mr Jeff, 51 Aotea Terrace, Huntsbury, Christchurch, New Zealand Cooke, Prof. Fred, Queen’s University, Kingston, Ontario, K7L 3N6, Canada Cooke, Mrs Sylvia, C/- Prof. Fred Cooke, Queen’s University, Kingston, Ontario, K7L 3N6, Canada Cooper, Mr Alan, Victoria University & , University of California - Berkeley , C /- S.B.S, P.O. Box 600, Wellington, New Zealand Cossee, Mr Roderick, Department of Conservation New Zealand, P.O. Box 10-420, Wellington, New Zealand Costa, Dr Daniel, University of California, Long Marine Laboratory, 100 Shaffer Road, Santa Cruz, CA 95060, U.S.A. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 23 Cowie, Dr Richard, Cardiff University, School of Biology, Cardiff, Wales, CF1 3TL, United Kingdom Cowie, Mr Jack, Leslie Street, Waiau 8275, New Zealand Cowling, Mr Sid, R.A.O.U., Box 22, Glenhuntly, Vic 3163, Australia Cowling, Mr Ben, Caulfield Grammar School, Box 22, Glenhuntly, Vic 3163, Australia Craig, Dr John, University of Auckland, Zoology Department, Private Bag, Auckland, New Zealand Crick, Dr Humphrey, British Trust for Ornithology, Beech Grove Station Road, Tring, Hertfordshire, HP23 5NR, United Kingdom Crocker, Mr Tony, 14 Thames Street, St Albans, Christchurch, New Zealand Crockett, Mrs Ruth, O.S.N.Z., 21 McMillan Avenue, Kamo, Whangarei 0101, New Zealand Crockett, Mr David, O.S.N.Z., 21 McMillan Avenue, Kamo, Whangarei 0101, New Zealand Cross, Mrs Milli, 47/2 Sale Street, Howick, Auckland, New Zealand Crossland, Mr Andrew, O.S.N.Z., 46 Frensham Crescent, Woolston, Christchurch, New Zealand Crouchley, Mr Dave, Department of Conservation New Zealand, P.O. Box 29, Te Anau, New Zealand Crowe, Prof. Timothy M., University of Cape Town, Fitzpatrick Institute, Rondebosch, 7700 Cape Town, South Africa Crowell, Prof. Kenneth, St. Lawrence University, Canton, New York, 13617, U.S.A. Crowell, Mrs Mamie R., RD4, Box 97, Canton, New York, 13617, U.S.A. Crowell, Mr Tom L., RD4, Box 97, Canton, New York, 13617, U.S.A. Croxall, Dr John, British Antarctic Survey, Madingley Road, Cambridge, CB3 0ET, United Kingdom Crozer-Jones, Dr Betsy, Tri State Bird Rescue & Research, 110 Possum Hollow Road, Newark, DE 19711, U.S.A. Culik, Dr Boris, Institut fur Meereskunde, Duesternbrooker Weg 20, 2300 Kiel 1 , Ger¬ many Cullen, Prof. Mike, Monash University, Clayton, Melbourne 3168, Australia Cunningham, Mr Duncan, Department of Conservation New Zealand, P.O. Box IQ- 420, Wellington, New Zealand Curl, Mr David, Monash University, Dept, of Ecology & Evolutionary Biology, Clayton, Victoria 3168, Australia Curry, Dr Robert L., Archbold Biological Station, P.O. Box 2057, Lake Placid, Florida, 33852, U.S.A. Custer, Dr Thomas W., U.S. Fish and Wildlife Service, P.O. Box 2506, Victoria, Texas, 77902, U.S.A. Cuthbert, Dr Francesca J., University of Minnesota, Department of Fisheries and Wildlife, 1980 Folwell Avenue, St Paul, MN 55108, U.S.A. Dahlgren, Dr Jens, University of Lund, Department of Ecology, Helgona vagen 5, 22362 Lund, Sweden Dahlsten, Prof. Donald L., University of California, Division of Biological Control, Berkeley, CA 94720, U.S.A. Danchin, Dr Etienne, E.N.S., 46 Rue d’Ulm, 75230, Paris CEDEX 05, France Danks, Mr. Alan, Department Conservation and Land Management, Two Peoples Bay Nature Reserve, Via Albany, W.A. 6330, Australia Dann, Mr Peter, Penguin Reserve Committee of Management, Box 403, Cowes, VIC 3992, Australia 24 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Date, Dr Liz, University of New England, Department of Ecosystem Management, Armidale, NSW 2351, Australia Daugherty, Dr Charles H., Victoria University of Wellington, P.O. Box 600, Welling¬ ton, New Zealand Davenport, Mr John, 214 Maungatapu Road, Tauranga, New Zealand Davey, Mr Chris, CSIRO, P.O. Box 84, Lyneham, ACT 2602, Australia Davidson, Mr Peter, Civil Aviation Authority (Australia), 23 Huelin Circuit, Flynn, Can¬ berra, ACT2615, Australia Davis, Mr Ken, 2618 NE 86th, Seattle, WA 98115, U.S.A. Davis, Dr Lloyd S., University of Otago, Department of Zoology, P.O. Box 56, Dunedin, New Zealand Davis, Ms Alison, Department of Conservation New Zealand, P.O. Box 10-420, Wel¬ lington, New Zealand Davis, Miss Marjory, 211 Hills Road, Shirley, Christchurch, New Zealand Dawnay, Mr Andrew, The Wildfowl & Wetlands Trust, Mill Road, Arundel, Sussex, BN18 9BP, United Kingdom Dawson, Prof Terence J., University of New South Wales, School of Biological Sci¬ ence, P.O. Box 1, Kensington, N.S.W. 2033, Australia De Hamel, Mr Richard J. B., 49 Hartley Avenue, Papanui, Christchurch, New Zealand De Rebeira, Mr Perry, 12 Glenwood Avenue, Glen Forrest, Perth, WA 6071, Australia Degnan, Ms Sandie, University of Queensland, Department of Zoology, St Lucia, Bris¬ bane 4072, Australia Dekker, Dr R.W.R.J., Universiteit van Amsterdam, Zoologisch Museum, Postbus 4766, 1009AT Amsterdam, Netherlands Delahaye-Brown, Mrs Anne-Marie, Kansas State University, Department of Anatomy & Physiology, Manhattan, Kansas 66506, U.S.A. Dennett, Dr Xenia, 100 Mountain View Parade, Rosanna, Victoria, 3084, Australia Derrickson, Dr Scott, Smithsonian Institution, National Zoological Park, Conservation & Research Centre, Front Royal, Va 22630, U.S.A. Desrochers, Mr Andre, University of Cambridge, 864 King’s College, Cambridge, CB2 1ST, United Kingdom Dhindsa, Dr Manjit S., Punjab Agricultural University, Department of Zoology, 141 004 Ludhiana, India Dhondt, Dr Andre, University of Antwerp UIA, Department of Biology, B-2610 Wilrijk, Belgium Dickson, Ms D. Lynne, Canadian Wildlife Service, Room 210, 4999-98 Ave, Edmon¬ ton, Alberta, T6B 2X3, Canada Dickson, Mr H.Loney, Canadian Wildlife Service, Rm 1104999-89 Avenue, Edmonton, Alberta T6B 2X3, Canada Diehl, Miss Barbara, Wysockiego 22 m82, P.O. Box 163, Warsaw, 03388, Poland Dilks, Peter J., Department of Conservation New Zealand, 23 Chrystal Street, Christchurch, New Zealand Disney, Mr John, Australian Museum, 12 Yallambee Road, Berowra, Sydney, N.S.W. 2081 , Australia Dittami, Mr John, Institut F. Zoologie, Althanstr. 14, Vienna, A-1090, Austria Divers, Mrs Margaret, R.D.2, Winton, Southland, New Zealand Dobler, Mr Gerold, Edelweissweg 13, Weingarten, D-7987, Germany Dodunski, Miss Krystyna, 151 Clarence Street, Riccarton, Christchurch, New Zealand Donaghey, Dr Richard, 1 Wembley Street, Burnie, Tasmania 7320, Australia Douglas, Mr Murray, Department of Conservation New Zealand, 44 Apuka Street, Brooklyn, Wellington, New Zealand ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 25 Douthwaite, Mr Bob, Natural Resources Institute, Central Avenue, Chatham Maritime, Chatham, ME4 4TB, United Kingdom Dowding, Dr John, T.V.N.Z., Natural History Unit, Box 474, Dunedin, New Zealand Dumbell, Dr Grant, Ducks Unlimited (NZ) Inc., P.O. Box 44-176, Lower Hutt, New Zealand Dunkerley, Dr Gillian, 69 Batemans Road, Gladesville, NSW 2111, Australia Dunnet, Prof. George M., University of Aberdeen, Tillydrone Avenue, Aberdeen, AB9 2TN, United Kingdom Dunnet, Mrs Margaret, C /- George M. Dunnet, University of Aberdeen, Tillydrone Avenue, Aberdeen AB9 2TN, United Kingdom Dunning, Dr John, University of Georgia, Department of Zoology, Athens, GA 30602, U.S.A. Dyck, Dr Jan, Copenhagen University, Insitute of Population Biology, Universitetsparken 15, Copenhagen, DK2100, Denmark Dyer, Ms Pamela K., University of Queensland, Department of Geographical Sci¬ ences, St Lucia, 4072, Australia Dyer, Dr Antoinette, Davidson College, Department of Psychology, P.O. Box 1719, Davidson, North Carolina 28036, U.S.A. d’Auzon, Mr Jean-Louis, Association Pour la Sauvegarde de la Nature Neo- Caledonienne, P.O. Box 1772, Noumea, New Caledonia Eadie, Dr John, University of Toronto, 1265 Military Trail, Scarborough, MIC 1A4, Ontario, Canada Ebihara, Dr Shizufumi, Nagoya University, Department of Animal Physiology, Faculty of Agriculture, Chikusa, Furo-cho, Nagoya, Japan Eckert, Mr H. John, Box 143, Langhorne Creek, SA 5255, Australia Eckert, Mrs Shirley Y., Box 143, Langhorne Creek, SA 5255, Australia Edwards, Mr Scott, University of California, Berkeley, Museum of Vertebrate Zoology, Berkeley, 94720, U.S.A. Eens, Mr Marcel, University of Antwerp, U.I.A., Universiteitsplein 1, B-2610 Wilrijk, Belgium Egan, Ms Judy, 40 Martin Road, Paraparaumu, Kapiti Coast, New Zealand Eguchi, Dr Kazuhiro, Kyushu University, Department of Biology, Faculty of Science, Fukuoka, 812, Japan Eichorst, Mr Bruce, University of North Dakota, Department of Biology, Box 8238, Grand Forks, North Dakota 58202, U.S.A. Eichorst, Mrs Jean, University of North Dakota, Department of Geography, Box 8274, Grand Forks, North Dakota 58202, U.S.A. Eller, Ms Gillian, 12 Rochdale Avenue, Glendowie, Auckland, New Zealand Elliott, Dr Graeme, 549 Rocks Road, Nelson, New Zealand Ellis, Mr Brian, 33 Bleakhouse Road, Howick, Auckland 1705, New Zealand Ellison, Mr Walter G., University of Connecticut, RFD 1 Box 748, Woodstock, VT 05091, U.S.A. Elzanowski, Dr Andreas, Max-Planck-lnstitut fur Biochemie, Martinsried 8033, Ger¬ many Elzer, Mrs Eleonore, Box 226, Philipsburg, Quebec, J0J IN0, Canada Emmerton, Dr Jacky, Purdue University, Department of Psychology, West Lafayette, IN 47907, U.S.A. Empson, Ms Raewyn A., Department of Conservation New Zealand, P.O. Box 5086, Wellington, New Zealand Enoksson, Dr Bodil, Uppsala University, Department of Zoology, Box 561, S-751 22 Uppsala, Sweden 26 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Erikstad, Dr Kjell E., Norwegian Institute for Nature Research, C /- Tromso Museum, University of Tromso, Tromso, Norway Erskine, Mrs Janet M., Box 1327, Sackville, NB EOA 3C0, Canada Erskine, Dr A.J. (Tony), Canadian Wildlife Service, Box 1327, Sackville, NB EOA 3C0, Canada Erz, Prof Wolfgang, Federal Research Centre for Nature Conservation, Konstantinstrasse 110, Bonn, D(W)-5300, Germany Escalante, Ms Patricia, American Museum of Natural History, Department of Ornithol¬ ogy, Central Park West at 79th Street, New York, NY, 10024-5192, U.S.A. Evans, Prof Peter R., University of Durham, Department of Biological Sciences, Sci¬ ence Laboratories, South Road, Durham DH1 3LE, United Kingdom Evans, Ms Janet, 87 St. Dunstans Bay, Winnipeg, R3T 3H6, Canada Evans, Dr Roger, University of Manitoba, Winnipeg, R3T 2N2, Canada Ezaki, Dr Yasuo, Natural History Museum , Hyogo Pref Board of Education, Shimoyamate-dori 5-10-1, Chuo-ku, Kobe 650, Japan Faizi, Mr S., National Commission for Wildlife Conservation and Development, P.O. Box 61681, Riyadh, Saudi Arabia Falls, Prof. J. Bruce, University of Toronto, Department of Zoology, 25 Harbord Street, Toronto, Ontario M5S 1A1, Canada Falls, Mrs Ann, University of Toronto, Department of Zoology, 25 Harbord Street, Toronto, Ontario M5S 1A1, Canada Fanshawe, Mr John, International Council For Bird Preservation, 32 Cambridge Road, Girton, Cambridge, CB3 0PJ, United Kingdom Feare, Dr Chris, ADAS Central Science Laboratories, Tangley Place, Worplesdon, Surrey, GU3 3LQ, United Kingdom Feduccia, Prof. Alan, University of North Carolina, 704 Wellington Drive, Chapel Hill, North Carolina, 27514, U.S.A. Feduccia, Mrs Olivia, C/- Prof. A. Feduccia, University of Carolina, 704 Wellington Drive, Chapel Hill, North Carolina, 27514, U.S.A. Feinsinger, Dr Peter, University of Florida, Department of Zoology, Gainesville, FL 32611, U.S.A. Ferdinand, Dr Lorenz, Danish Ornithological Society, Frederiksborgvej 525, DK-4000 Roskilde, Denmark Ferolla, Prof Maria Ignez, Universidade Federal De Minas Gerais, Av. Antonio Carlos, 6627, Pampulha, Belo Horizonte, Brazil Ficken, Dr Millicent, University of Wisconsin, Department of Biological Sciences, Milwaukee, Wl 53201, U.S.A. Filewood, Mr Win, University of N.S.W., 26 Trelawney Street, Eastwood, N.S.W. 2122, Australia Fitzgerald, Dr Mike, DSIR Land Resources, 66 Bloomfield Terrace, Lower Hutt, New Zealand Fitzpatrick, Dr John W., Archbold Biological Station, Box 2057, Lake Placid, FL 33852, U.S.A. Fitzpatrick, Miss Mary, 4 Molesworth Place, Hoon Hay, Christchurch, New Zealand Fjeldsa, Dr Jon, Zoological Museum, Universitetsparken 15, Copenhagen, DK2100, Denmark Flade, Mr Martin, Technische Universitat Berlin, Seelingstr. 32, Berlin - West 19, D- 1000, Germany Fleischer, Dr Rob, University of North Dakota, Department of Biology, Grand Forks, ND 58202, U.S.A. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGIC! 27 Fleming, Dr Michael, Conservation Commission of N.T., 51 Larapinta Drive, Alice Springs, 0870, Australia Fleming, Mr Brian, 27 Clark Road, Ivanhoe, Victoria 3079, Australia Fleming, Mrs Anthea, Bird Observers Club of Australia, 27 Clark Road, Ivanhoe, Vic¬ toria 3079, Australia Flint, Dr Elizabeth, Johnston Atoll National Wildlife Refuge, P.O. Box 396, APO San Francisco, California 96305, U.S.A. Flux, Ian, 230 Hill Road, Belmont, Lower Hutt, New Zealand Forbes, Dr Scott, Simon Fraser University, #315-11675 7th Avenue, Richmond V7E 4X4, Canada Forbes, Mrs Margaret, C/- Dr Scott Forbes, Simon Fraser University, #315-11675 7th Avenue, V7E 4X4, Canada Ford, Dr Hugh A., University of New England, Department of Zoology, Armidale, NSW 2351 , Australia Fordham, Dr Robin, Massey University, Palmerston North, New Zealand Forslund, Mr Par, Uppsala University, Department of Zoology, Box 561, Uppsala, S- 75122, Sweden Fotso, Mr Roger C., The University of Yaounde, Faculty of Sciences, P.O. Box 812, Yaounde, Cameroon Fountain, Mr Kevin, 39 Amyes Road, Hornby, Christchurch, New Zealand Fox, Mr Eric K., Otorohanga Zoological Society, P.O. Box 222, Otorohanga, New Zealand Fox, Dr Nick, National Avian Research Centre, (Abu Dhabi), UK Facility, Penllynin, College Road, Carmarthen SA33 5EH, Wales, United Kingdom Francis, Dr Charles M., Queen’s University, Biology Department, Kingston, Ontario, K7L 3N6, Canada Frank, Ms Michelle, 19 Grove Road, Kelburn, Wellington, New Zealand Franke, Ms Irma, Museo de Historia Natural, Box 14-0434, Lima 14, Peru Frantzen, Mr Bjorn, Greenpeace Norway, Postboks 30, N-2966, Slidre, Norway Freed, Dr Lenny, University of Hawaii, Department of Zoology, 2538 The Mall, Hono¬ lulu, Hawaii 96822, U.S.A. Freeman, Mr Alastair, 2/527 Cashel Street, Linwood, Christchurch, New Zealand Freeman, Ms Amanda, 2/527 Cashel Street, Linwood, Christchurch, New Zealand French, Dr Kristine, Macquarie University, Nth Ryde, Sydney, NSW 2109, Australia Friend, Dr Milton, United States Department of the Interior, Fish & Wildlife Service, 6006 Schroeder Road, Madison, Wisconsin 53711, U.S.A. Frund, Mr Jean-Louis, Les Productions Jean-Louis Frund Inc., 4100, Grande coulee, St Edouard, Quebec J0K 2H0, Canada Fujimaki, Prof. Yuzo, University of Obihiro, Inada, Obihiro, 080, Japan Fullagar, Dr Peter J., CSIRO Division of Wildlife & Ecology, P.O. Box 84, Lyneham, Canberra 2602, Australia Fuller, Dr Robert, British Trust for Ornithology, Beech Grove, Tring, Hertfordshire, HP23 5NR, United Kingdom Furness, Dr Robert, University of Glasgow, Department of Zoology, Glasgow, G12 8QQ, United Kingdom Gabrielsen, Dr Geir W., Norwegian Institute for Nature Research, Haakon VH’s 6T 5A, Tromso, 9000, Norway Galbraith, Dr Colin A., Nature Conservancy Council, Northminster House, Peterbor¬ ough, Cambridge, PEI 1UA, United Kingdom Galbraith, Mrs Maria, 10 The Parslins, Deeping St. James, Peterborough, Lines PE6 8NR, United Kingdom 28 ACTA XX CONGRESSUS INTERN ATIONALIS ORNITHOLOGICI Ganter, Miss Barbara, Bundesstrasse 26, Hattstedt, 2251, Germany Gardarsson, Prof. Arnthor, University of Iceland, Institute of Biology, Grensasvegur 12, IS-108 Reykjavik, Iceland Garnett, Dr Stephen, SEARCH, Garden of St Erth, Blackwood, 3458, Australia Garrett, Mr Kimball L., Natural History Museum of L.A. County, Section of Ornithol¬ ogy, 900 Exposition Boulevard, Los Angeles, CA 90007, U.S.A. Garrick, Mr Andy, Department of Conservation, P.O. Box 1146, Rotorua, New Zea¬ land Garrido Calleja, Mr Orlando H., Museo Nacional de Historia Natural, Industria- Capitolio Nacional, La Habana, 2, Cuba Gaston, Dr Anthony, Canadian Wildlife Service, 100 Gamelin Boulevard, Hull, Que¬ bec KIA OH3, Canada Gauthier, Dr Gilles, Universite Laval, Department of Biology, Ste Foy QC, G1K 7P4, Canada Gauthreaux, Prof. Sidney A., Clemson University, Department of Biological Sciences, Clemson, SC 29634-1903, U.S.A. Gehlbach, Dr Frederick R., Baylor University, Waco, TX 76798, U.S.A. Gehlbach, Mrs Nancy Y., 13 Sugar Creek, Waco, TX 76712, U.S.A. Gelter, Dr Hans P., University of Uppsala, Department of Genetics, Box 7003, Uppsala 75007, Sweden Gemmill, Ms Daphne, Earthwatch, 215 10th Street, SE, Washington, DC, 20003, U.S.A. Gerstberger, Dr Rudiger, Max-Planck-lnstitut for Physiologie, W.G. Kerckhoff-lnstitut, Parkstr. 1, D6350 Bad Nauheim, Germany Gesche, Dr Melita, 48 N.Helderberg Pkwy, Slingerlands, NY 12159, U.S.A. Gessaman, Dr James, Utah State University, UMC 5305, Logan, Utah 84322, U.S.A. Getty, Dr Thomas, Michigan State University, Kellogg Biological Station, Hickory Corvers, Ml 49060, U.S.A. Geven, Miss Helen, 117C Waimairi Road, llam, Christchurch, New Zealand Gibb, Dr John A., 3 Wairere Road, Belmont, Lower Hutt, New Zealand Gichuki, Mrs Cecilia, National Museums of Kenya, Department of Ornithology, Box 40658, Nairobi, Kenya Gichuki, Mr Nathan N., National Museums of Kenya, Ornithology Department, Box 40658, Nairobi, Kenya Gidall, Miss Kim, 291 Hills Road, Shirley, Christchurch, New Zealand Giese, Ms Melissa, Griffith University, 116 Dixon Street, Sunnybank, Brisbane 4109, Australia Giezentanner, Mr William, Massachusetts Audubon Society, 278 Eliot Street, Natick, MA, 01760, U.S.A. Gilbert, Dr Betty, RT. 1 Box 29, Dorset, Vermont 05251, U.S.A. Gill, Dr Brian, Auckland Museum, Private Bag, Auckland, New Zealand Gillett, Prof Dorothy K, 5521 Opihi Street, Honolulu, HI 96821, U.S.A. Gnam, Ms Rosemarie, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024, U.S.A. Gochfeld, Dr Michael, Robert Wood Johnson Medical School, 675 Hoes Lane, Piscataway, NJ 08854, U.S.A. Goerner, Mr Martin, Institut fur Landscheiftsforschung und Naturschutz, Steiger 17, Jena, 6900, Germany Goldsmith, Dr Arthur, University of Bristol, Department of Zoology, Woodland Road, Bristol BS8 IV6, United Kingdom ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 29 Gooch, Mr Stephen, Sheffield University, School of Biological Sciences, Western Bank, Sheffield, SIO 2TN, United Kingdom Goodey, Mr Wayne, Monash University, Dept of Ecology & Evolutionary Biology, Wellington Road, Clayton, Victoria 3168, Australia Goodman, Mr Steve M., Field Museum of Natural History, Roosevelt Road, at Lake Shore Drive, Chicago, IL 60605, U.S.A. Goodwillie, Ms Carol D., University of the South Pacific, Box 9874 Nadi Airport, Nadi, Fiji Gordon, Mr David, Royal Society for the Protection of Birds, The Lodge, Sandy, Bed¬ fordshire, SGI 9 2DL, United Kingdom Goriup, Mr Paul, Nature Conservation Bureau, 122 Derwent Road, Thatcham, Berk¬ shire, RG13 4UP, United Kingdom Gosler, Dr Andrew G., Edward Grey Institute, Department of Zoology, South Parks Road, Oxford, 0X1 3PS, United Kingdom Goslow Jr., Dr George E., Brown University, Box G-BMC, Section of Population Bi¬ ology, Providence, Rhode Island 02912, U.S.A. Gowaty, Dr Patricia A., Clemson University, Department of Biological Sciences, Long Hall, Clemson, SC 29634-1903, U.S.A. Graham, Enfys, 34 Rawhiti Road, Pukerua Bay, Wellington, New Zealand Grant, Prof. Peter R., Princeton University, Biology Department, Princeton, 08544, U.S.A. Grant, Dr B. Rosemary, Princeton University, Biology Department, Princeton, 08544, U.S.A. Grant, Mr Andrew D., Department of Conservation New Zealand, Private Bag, Christchurch, New Zealand Gray, Dr Russell, University of Otago, Department of Psychology, P.O. Box 56, Dunedin, New Zealand Green, Dr Ronda J., Griffith University, Australian School of Env. Studies, Kessels Road, Nathan, Brisbane 4111, Australia Green, Dr Rhys, The Royal Society for the Protection of Birds, ‘The Lodge’, Sandy, Bedfordshire, SGI 9 2DL, United Kingdom Greene, Mr Terry, 13 Reydon Place, Howick, Auckland, New Zealand Gregory, Mr Brian J., Welsh Ornithological Society, Monmouth School, Monmouth, Gwent, NP5 3XP, United Kingdom Gretton, Mr Adam, I.C.B.P., 32 Cambridge Road, Girton, Cambridge, CB3 0PJ, United Kingdom Griffin, Miss Andree, “Wirrawilla” CMB 16, Paluma, Queensland 4816, Australia Griffin, Ms Janine, 133 Summervale Drive, Christchurch, New Zealand Griffiths, Geraldine, College of St. Paul & St. Mary, 22 Elm Grove, Watford, Herts., United Kingdom Grove, Christine, 15 Ravenna Street, Avonhead, Christchurch, New Zealand Grubb Jnr., Prof. Thomas C., Ohio State University, Department of Zoology, 1735 Neil Avenue, Columbus, Ohio 43210, U.S.A. Guilford, Dr Tim, Oxford University, Department of Zoology, South Parks Road, Ox¬ ford, OX1 3PS, United Kingdom Gwinner, Prof. Eberhard, Max Planck Institut fur Verhaltensphysiologie, Vogelwarte, D-8138 Andechs, Germany Habmann, Mrs Gertrud, Akademiestr. 23, D-8000 Munchen 40, Germany Hadden, Mr Don, 288 Yaldhurst Road, Avonhead, Christchurch, New Zealand Hafner, Dr Heinz, Tour Du Valat, Station Biologique, Le Sambuc, Arles, 13200, France 30 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Haig, Dr Susan, Smithsonian Institution, Department of Aquaculture, Fisheries & Wild¬ life, Clemson University, SC 29634, U.S.A. Haila, Dr Yrjo, University of Helsinki, Department of Zoology, P. Rautatiekatu 13, 00100 Helsinki, Finland Hailman, Prof. Jack, University of Wisconsin, 3205 Tally Ho Lane, Madison, Wl 53705, U.S.A. Hailman, Mrs Elizabeth, University of Wisconsin, 3205 Tally Ho Lane, Madison, Wl 53705, U.S.A. Hake, Mr Mikael, University of Gothenburg, Department of Zoology, P.O. Box 250 59, S400 31 Gothenburg, Sweden Hall-Jones, Dr John, Ornithological Society of New Zealand, 74 Park Street, Invercargill, New Zealand Hall-Jones, Mrs Pamela, 74 Park Street, Invercargill, New Zealand Halliburton, Mr Gavin, 6 Barlow Street, llam, Christchurch, New Zealand Halliburton, Mrs Beth, 6 Barlow Street, llam, Christchurch, New Zealand Hanotte, Mr Olivier, University de Mons Hainaut, Faculte de Medecine, Avenue du Champ de Mars 24, 7000 Mons, Belgium Hanowski, Ms Jo-Ann, Natural Resources Research Institute, 5013 Miller Trunk High¬ way, Duluth, MN 55811, U.S.A. Hanski, Mr llpo, University of Helsinki, Department of Zoology, P Rautatiekatu 13, Helsinki 00100, Finland Hanssen, Ms Guri, Sverdrupsgt 12, N-0559, Oslo 5, Norway Harrison, Mr Malcolm, Department of Conservation New Zealand, P.O. Box 10-420, Wellington, New Zealand Harrison, Mrs Kathleen C., 50 Athol Terrace, llam, Christchurch, New Zealand Hartley, Mr Ian, Leicester University, Zoology Department, University Road, Leices¬ ter, LEI 7RH, United Kingdom Harty, Mr Tom, R D 2, Drury, South Auckland, New Zealand Harty, Mrs Hazel, R D 2, Drury, South Auckland, New Zealand Haslett, Mrs Kay, 47 Ranui Road, Remuera, Auckland, New Zealand Hausberger, Dr Martine, Laboratoire d’Ethologie, Universite de Rennes, 1 Avenue du General Leclerc, Rennes, Cedex 35062, France Hawkins, Mrs Jenny, 772 Atawhai Drive, Nelson, New Zealand Hay, Dr Rod, Department of Conservation, P.O. Box 10-420, Wellington, New Zea¬ land Hay, Mrs Nan, 14 Holmwood Road, Christchurch, New Zealand Healy, Ms Sue, University of Oxford, EGI, Department of Zoology, South Parks Road, Oxford OX1 3PS, United Kingdom Heather, Barrie, 10 Jocelyn Crescent, Silverstream, New Zealand Hebert, Mr Percy N., University of Manitoba, Zoology Department, Winnipeg, Mani¬ toba R3T 2N2, Canada Hegelbach, Dr Johann, Zoologisches Museum Zurich, Winterthurer str. 190, Zurich, CH-8057, Switzerland Heinsohn, Mr Robert, Australian National University, Department of Zoology, GPO Box 4, Canberra, ACT 2601, Australia Hejl, Dr Sallie, United States Forest Service, Forestry Sciences Laboratory, P.O. Box 8089, Missoula, MT, 59807, U.S.A. Helbig, Dr Andreas J., University of Frankfurt, Zoological Institute, Siesmayerstr. 70, Frankfurt, D-600, Germany Heldmaier, Prof. Gerhard, Phillips University, Department of Biology, Box 1929, ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 31 Marburg, D-3550, Germany Henderson, Dr Ian, University of Leicester, Department of Zoology, University Road, Leicester, LEI 7RH, United Kingdom Henderson, Mr John R., Ithaca College, Ithaca, N.Y. 14850, U.S.A. Henley, Mr Jim C., Ornithological Society of New Zealand, 6/456 Aberdeen Road, Gisborne, New Zealand Herring, Ms Charlene, 2 Clifton Bay, Sumner, Christchurch, New Zealand Heuwinkel, Dr Hubert, Natural History Museum of Benrath, Benrather Schlossallee 102, D-4000 Dusseldorf 13, Germany Hewlett, Mrs. Maureen G., Lake Head University, C /- Prof. Ryder, Oliver Road # 955, Thunder Bay, P7B 5EI, Canada Hidasi, Prof Jose, Fundacao Museo de Ornitologia, Avenida Para 381,395, CEP 75110, Goiania, Brazil Higgins, Mr Peter J., Royal Australasian Ornithologists Union, 21 Gladstone Street, Moonee Ponds, Victoria 3039, Australia Higuchi, Dr Hiroyoshi, Wild Bird Society of Japan, Research Centre, Higashi 2-24-5, Shibuya-ku, Tokyo, 150, Japan Hilgerloh, Dr Gudrun, Nationalparkverwaltung, Okosystemforschung, Virchowstrasse 1, D 2940 Wilhelmshaven, Germany Hill, Dr Gary, 162 Opanuku Road, Henderson, Auckland, New Zealand Hills, Ms. Sarah-Jane, 35 de Bretagne Street, St. Lambert, Quebec, J4S 1A3, Canada Hillstrom, Mr Lars, Department of Zoology, Box 561, 75328 Uppsala, Sweden Hino, Dr Teruaki, Hokkaido University, Institute of Applied Zoology, N9 W9, Kita-ku, 060 Sapporo, Japan Hirt, Mr Fritz, Schweizer Vogelschutz SVS/ASPO, Oberdorf 43, CH 8164 Bachs, Switzerland Hitchmough, Mr Rod, Victoria University of Wellington, School of Biological Sciences, Box 600, Kelburn, Wellington 2, New Zealand Hocken, Mr Tony, “East Riding”, White Rocks Road, 6-D RD, Oamaru, New Zealand Hogstad, Prof. Olav, University of Trondheim, Department of Zoology, N-7055 Dragvoll, Trondheim, Norway Hogstad, Mrs Anne-M., University of Trondheim, C /- Prof Hogstad, Department of Zoology, N-7055 Dragvoll, Trondheim, Norway Hogstedt, Prof Goran, Museum of Zoology, Bergen, N5007, Norway Hoi, Dr Herbert, University of Vienna, Institute of Zoology, Althanstr. 17, Vienna 1090, Austria Hoi-Leitner, Ms Maria, Althanstrasse 14, Vienna, 1090, Austria Holmes, Mrs Deborah, Dartmouth College, Department of Biological Sciences, Hano¬ ver, NH, 03755, U.S.A. Holmes, Prof. Richard T., Dartmouth College, Department of Biological Sciences, Hanover, NH, 03755, U.S.A. Holt, Mr Steven, The Academy of Natural Sciences, 19th & The Parkway, Philadel¬ phia, PA 19103, U.S.A. Holyoak, Dr David T., College of St. Paul & St. Mary, 22 Elm Grove, Watford, Herts., United Kingdom Homberger, Dr Dominique G., Louisiana State University, Department of Zoology & Physiology, Baton Rouge, LA 70803, U.S.A. Hornabrook, Miss Judith S., 9B Pitarua Street, Thorndon, Wellington, New Zealand Horne, Ms Jennifer F.M., National Museums of Kenya, P.O. Box 40658, Nairobi, Kenya 32 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Horrigan, Mr Bill, 57 Piddington Street, Watson, ACT, 2602, Australia Horton, Dr Philippa, South Australian Museum, North Terrace, Adelaide 5000, Aus¬ tralia Houston, Dr David C., University of Glasgow, Department of Zoology, Glasgow, G12 8QQ, United Kingdom Houston, Mr Dave, Department of Conservation New Zealand, P.O. Box 1130, Dunedin, New Zealand Houston, Dr David, British Ornithologists’ Union, British Museum (Natural History), Department of Ornithology, Tring, Herts. HP23 6AP, United Kingdom Howard, Mr Richard P., I.C.B.P. British Section Chairman, Hogg House, Piggy Lane, Lower Basildon, Reading RG8 9NH, United Kingdom Howard, Mrs Sheila, Child-Beale Wildlife Trust, Hogg House, Piggy Lane, Lower Basildon, Reading RG8 9NH, United Kingdom Howden, Mr Peter, “Helmsdale”, No 8 R D, Ashburton, New Zealand Howe, Prof. Robert W., University of Wisconsin - Green Bay, Green Bay, Wl 5431 1 - 7001, U.S.A. Howell, Dr Thomas R., University of California Los Angeles, P.O. Box 950, Gualala, CA 95445, U.S.A. Howell, Mrs Eleanor, P.O. Box 950, Gualala, California 95445, U.S.A. Howland, Prof. Howard C., Cornell University, Section of Neurobiology & Behavior, W- 201 Mudd Hall, Ithaca, NY, U.S.A. Howland, Mrs Monica, Cornell University, 103 Stimson Hall, Ithaca, NY 14853, U.S.A. Hsu, Prof. Weishu, Peking Natural History Museum, 126 Tien Chiao Street, Beijing, 100050, China Hudde, Ms Barbara, Konrad Lorenz Institut fur Vergleichende Verhaltensforschung, Savoyenstrasse 1A, A-1160 Wien, Austria Hughes, Dr Maryanne, University of British Columbia, Department of Zoology, 6270 University Boulevard, Vancouver, BC V6T 2A9, Canada Hulscher, Dr Jan, University of Groningen, Zoological Laboratory, Kerklaan 30, Haren, 9750-AA, Netherlands Hulsman, Dr Kees, Griffith University, Nathan, 4111 Brisbane, Australia Hultsch, Dr Henrike, Freie Universitat, Institut f. Verhaltensbiologie, Haderslebener Strasse 9, D-1000 Berlin 41, Germany Hummel, Prof. Dietrich J., Trinchenberg 4, D-3302 Cremlingen, Germany Hunt, Dr Peggy W., 555 Bluebird Canyon Drive, Laguna Beach, California 92651, U.S.A. Hunt, Mr Gavin, 26 Carruthers Street, llam, Christchurch, New Zealand Hunt Jr., Prof. George L., University of California, Dept. Ecology & Evolutionary Bi¬ ology, Irvine, California 92717, U.S.A. Hunter, Dr Mac, University of Maine, Wildlife Department, Orono, 04469 Maine, U.S.A. Hunter, Ms Fiona M., University of Sheffield, Western Bank, Sheffield, SIO 2TN, United Kingdom Huntington, Prof. Charles E., Bowdoin College, RFD 2, Box 357, S. Harpswell, Maine, 04079, U.S.A. Huntington, Mrs Louise S., RFD 2, Box 357, S. Harpswell, Maine, 04079, U.S.A. Hurrell, Ms Mei, 1 Helmore’s Lane, Fendalton, Christchurch, New Zealand Hussain, Mr S. A., Bombay Natural History Society, Hornbill House, S. B. Road, 400023 Bombay, India Hussell, Dr David J.T., Ontario Ministry of Natural Resources, P.O. Box 5000, Maple, Ontario, L6A 1S9, Canada Hutchinson, Mr Wayne, Department of Conservation New Zealand, Private Bag, Wanganui, New Zealand ACTA XX CONGRESSUS INTER NATION ALIS ORNITHOLOGICI 33 Hvenegaard, Mr Glen, University of Alberta, Department of Forest Science, 855 Gen¬ eral Services Building, Edmonton, Alberta T6G 2H1, Canada Ickes, Dr Roy A., Washington and Jefferson College, Washington, PA, 15301, U.S.A. Illingworth-Cooper, Mrs Janet, 2a Oxford Street, Gloucester, GLI 3EQ, United King¬ dom Imber, Dr Michael J., Department of Conservation New Zealand, P.O. Box 10-420, Wellington, New. Zealand Ingolfsson, Dr Agnar, University of Iceland, Institute of Biology, Grensasvegur 12, 108 Reykjavic, Iceland Innes, Mr John, Forest Research Institute, Private Bag 3020, Rotorua, New Zealand Irons, Mr David B., U.S. Fish & Wildlife Service, 1011 E. Tudor Road, Anchorage, Alaska 99503, U.S.A. Ishida, Dr Ken, The University Of Tokyo, University Forest In Chichibu, 1-49 Hinoda- Machi 1-Chome, Chichibu, Saitama 368, Japan Ishii, Prof. Susumu, Waseda University, Biology Department, 1-6-1 Nishi-Waseda, Shinjuku-ku, Tokyo, 169, Japan Ishikawa, Mr Nobuo, Laboratory of N. Japan’s Ornithology, 244-3, 3-Jo, 2-Ku, Shunko- cho, 070 Asahikawa, Japan Ivison, Tessa, 133 Waratah Street, Katoomba, Sydney, NSW 2780, Australia Jackson, Dr Sue, University of Capetown, P.F.I.A.O., Private Bag, Rondebosch, Cape Town 7700, South Africa Jacquier, Mr Andre G. H., Association Pour la Sauvegarde de la Nature Neo- Caledonienne, P.O. Box 1772, Noumea, New Caledonia Jacquier, Mrs Therese M. E., Association Pour la Sauvegarde de la Nature Neo- Caledonienne, P.O. Box 1772, Noumea, New Caledonia Jahnel, Mr Mathias, Johann Wolfgang Goethe Universitat, Zoologisches Institut, Siesmayerstr. 70, Frankfurt, D-6000, Germany Jaksic, Prof. Fabian M., Catholic University of Chile, Box 114-D, Santiago, Chile James, Prof. Douglas A., University of Arkansas, Department of Zoology, Fayetteville, AR 72701, U.S.A. James, Dr Helen, Smithsonian Institution, Bird Division, NHB E611, Washington, DC, 20560, U.S.A. James, Dr Frances, Florida State University, Tallahassee, Florida 32306, U.S.A. James, Mr Roger, 27 Orbell Street, Dalmore, Dunedin, New Zealand Jamieson, Dr Ian, University of Otago, Zoology Department, P.O. Box 56, Dunedin, Otago, New Zealand Jamous, Dr Rene, International Wildlife Preservation Fund, Chateau De Sauvage, Rambouillet, 78120, France Jane, Mr Dave, Department of Conservation New Zealand, Private Bag, Wanganui, New Zealand Jansen, Mr Paul, Department of Conservation New Zealand, P.O. Box 1 146, Rotorua, New Zealand Jarvinen, Mr Jaakko, Partiotie 26 B, SF-00370 Helsinki, Finland Jeal, Mrs Doreen, 5 Farndale Place, Avonhead, Christchurch, New Zealand Jenkins, Mrs Pauline, 234 Howick Road, Blenheim, New Zealand Jenkins, Mr Peter, New Zealand Jenner, Ms Peggie, 26 Page Street, Morrinsville, New Zealand Jenni, Dr Lukas, Swiss Ornithological Institute, Sempach, 6204, Switzerland Jenni-Eiermann, Dr Susi, Swiss Ornithological Institute, Sempach, 6204, Switzerland Johnson, Prof. Ned K., University of California, Museum of Vertebrate Zoology, De¬ partment of Integrative Biology, Life Sciences Building, Berkeley CA 94720, U.S.A. 34 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Johnson, Dr Stephen R., LGL Limited, 9768 2nd Street, Sidney, British Columbia, V8L 3Y8, Canada Johnson, Ms Helen, C/- Post Office, Oxford, North Canterbury, 8253, New Zealand Johnsson, Miss Kristina, University of Agricultural Sciences, Department of Wildlife Ecology, Box 7002, Uppsala, 750 07, Sweden Joiris, Prof Claude, Free University of Brussels (V.U.B.), Pleinlaan 2, Brussels, B1050, Belgium Jolly, Mr Jim, Box 29035, Ngaio, Wellington 6004, New Zealand Jones, Ms Frances, 2142 Marne Street, Victoria B.C., V8S 4J9, Canada Jones, Prof. Davrd R., University of British Columbia, Department of Zoology, 6270 University Boulevard, Vancouver, V6T 2A9, Canada Jones, Dr Darryl N., Griffith University, Div. of Australian Environmental Studies, Nathan, Queensland 4111, Australia Jordhoy, Dr Per, Norwegian Institute for Nature Research, Tungasletta 2, N7004 Trondheim, Norway Joseph, Mr Leo, University of Queensland, Department of Zoology, St Lucia, Brisbane 4067, Australia Kaiser, Mr Andreas, Max-Planck-lnstitut fur Verhaltensphysiologie, Vogelwarte Radolfzell, Schloss Moggingen, Moggingen, Radolfzell, 7760, Germany Kalaber, Mr Ladislau, Society Ornithological of Romania, (4225 Reghin), Sor M. Eminescu 26, Mures, Reghin 4225, Romania Kalas, Dr John A., Norwegian Institute for Nature Research, Tungasletta 2, N7004 Trondheim, Norway Kalchreuter, Dr Heribert, European Wildlife Research Institute, Glashutte, Bonndorf, 7823, Germany Kalisch, Dr Hans-Joachim, Dorfstrasse 22, Allerbuttel, 3178, Germany Kampp, Dr Kaj, Zoological Museum, Universitetsparken 15, Kobenhavn, 2100, Den¬ mark Kanyamibwa, Mr Samuel, RWANDA, CNRS - Route de Mende, B.P. 5051, Montpellier, 34090, France Karasov, Dr William, University of Wisconsin-Madison, Department of Wildlife Ecol¬ ogy, 1630 Linden Drive, 226 Russell Labs, Madison, Wl 53706, U.S.A. Katz, Ms Mary, American Museum of Natural History, 79th Street & Central Park West, New York, NY, 10024, U.S.A. Keall, Ms Sue, 56 Middleton Road, Johnsonville, Wellington, New Zealand Kear, Dr Janet, The Wildfowl & Wetlands Trust, Martin Mere, Burscough, Lancashire, L40 OTA, United Kingdom Keast, Prof Allen, Queen’s University , Kingston, Ontario, K7L 3N6, Canada Keith, Mr G. Stuart, American Museum of Natural History, Central Park West, New York, NY 10024, U.S.A. Keith, Mrs Sallyann, American Museum of Natural History, C/- G Stuart Keith, Cen¬ tral Park West, New York, NY 10024, U.S.A. Keith, Mr J. Anthony, Canadian Wildlife Service, 563 Fairview Avenue, Rockcliffe Park, Ontario, KIM 0X4, Canada Kelynack, Mr Lindsay, 6 Truro Road, Camborne, Plimmerton, Wellington, New Zea¬ land Kennedy, Mr Martyn, University ot Otago, Department of Zoology, P.O. Box 56, Dunedin, Otago, New Zealand Kennedy, Mr Euan S., Department of Conservation New Zealand, Private Bag, Christchurch, New Zealand ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 35 Kerle, Dr Anne, 51 Larapinta Drive, Alice Springs, 0870, Australia Kerr, Ms Liz, Andrew Isles Bookshop, P.O. Box 358, Prahran 3181, Australia Ketterson, Dr Ellen D., Indiana University , Department of Biology, Bloomington, IN 47405, U.S.A. Kettle, Mr Ron, 75 Dupont Road, London, SW20 8EH, United Kingdom Kikkawa, Prof. Jiro, University of Queensland, St Lucia, Brisbane 4072, Australia Kikkawa, Mrs Naoko, C/- Prof. Jiro Kikkawa, University of Queensland, St Lucia, Bris¬ bane, Australia Kikuchi, Mr Motoshi, Waseda University, Biology Department, 1-6-1 Nishi-Waseda, Shinjyuku-ku, Tokyo, 169, Japan King, Prof. James R., Washington State University, Department of Zoology, Pullman, WA 99164 - 4220, U.S.A. King, Mr Warren B., P.O. Box 77, Ripton, VT 05766, U.S.A. King, Mr Ben, American Museum of Natural History, 79th Street at Central Park West, New York, NY 10024, U.S.A. Klein, Ms Nedra K., University of Michigan, Museum of Zoology, Ann Arbor, Ml 48109, U.S.A. Klempt, Mr Martin, Institute for Small Animal Research, Doernberg Strasse 25/27, D3100 Celle, Germany Klomp, Mr Nicholas, University of Glasgow, Glasgow, G12 8QQ, Scotland, United Kingdom Knight, Dr Richard, Colorado State University, Department of Fishery and Wildlife, 233 Wagar Hall, Ft. Collins, CO 80523, U.S.A. Knox, Dr Alan, Buckinghamshire County Museum, Tring Road, Walton, Buckingham¬ shire, HP22 5JP, United Kingdom Komdeur, Mr Jan, I.C.B.P., Fregate Island, Box 330, Victoria, Mahe, Seychelles Komdeur, Mrs Mariette, ICBP, 7 Newnham Croft Street, Newnham, Cambridge, CB3 9HR, United Kingdom Korf, Dr Horst-Werner, Department of Anatomy and Cytobiology, Aulweg 123, D 6300 Giessen, Germany Korpimaki, Dr Erkki M. T., University of Turku, Laboratory of Ecological Zoology, Department of Biology, Turku, 20500, Finland Koskimies, Mr Pertti, University of Helsinki, Department of Zoology, Division of Ecol¬ ogy, P. Rautatiekatu 13, SF-00100 Helsinki, Finland Kruijt, Prof. Jaap, University of Groningen, Zoological Laboratory, Postbox 14, Haren 9750AA, Netherlands Kruijt, Ms Tineke, C /- Prof. Jaap Kruijt, University of Groningen, Postbox 14, Haren 9750AA, Netherlands Kubat, Miss Susanne, Institut fur Vergleichende Verhaltensforschung, Savoyenstrasse 1 A, Vienna A-1 160, Austria Kubokawa, Dr Kaoru, Waseda University, Department of Biology, 1-6-1 Nishiwaseda, Shinjuku, Tokyo 169, Japan Kuitunen, Dr Markku T., University ofJyvaskyla, Department of Biology, Yliopistonkatu 9, SF-40100 Jyvaskyla, Finland Kuitunen, Ms Pirjo Y., University of Jyvaskyla, Yliopistonkatu 9, SF-40100 Jyvaskyla, Finland Kurochkin, Dr Evgeny N., Paleontological Institute of the U.S.S.R., Academy of Sci¬ ence, Profsouzhaya 123, 117647 Moscow, U.S.S.R. Kuroda, Dr Nagahisa, Yamashina Institute for Ornithology, 1 15 Konoyama, Abiko-si, Chiba 270-1 1 , Japan 36 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Kwon, Prof. Ki Chung, Dong-A University, Department of Biology, 840 Hadan Dong, Sahagu, Pusan, 604-714, Korea(S) Lambeck, Dr Robert H.D., Delta Instituut voor Hydrobiologisch Onderzoek, Vierstraat 28, Yerseke, 4401 EA, Netherlands Landfried, Dr Steven E., Steven Landfried Consultants, Route One, Evansville, Wl 53536, U.S.A. Landmann, Dr Armin, University of Innsbruck, Institute of Zoology, Technikerstr. 25, Innsbruck, A-6020, Austria Landre, Ms Elissa, Massachusetts Audubon Society, 278 Eliot Street, Natick, MA, 01760, U.S.A. Langham, Dr Nigel, D.S.I.R., Ecology Division, Goddard Lane, Havelock North, New Zealand Langlands, Mr Peter, 90A Kaiwara Street, Hoon Hay, Christchurch, New Zealand Langmore, Miss Naomi, Australian National University, Department of Zoology, G.P.O. Box 4, Canberra, ACT 2601, Australia Langslow, Dr Derek, Nature Conservancy Council, Northminster House, Peterbor¬ ough, PEI 1UA, United Kingdom Lanyon, Dr Scott, Field Museum Of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, IL 60605, U.S.A. Laughlin, Ms Sarah B., Vermont Institute of Natural Science, P.O. Box 86, Woodstock, VT, 05091, U.S.A. Le Maho, Dr Yvon, Laboratoire d’Etude des Regulations Physiologiques C.N.R.S., 23 Rue Becquerel, 67087 Strasbourg, France LeCroy, Ms Mary, American Museum of Natural History, Central Park W. at 79th Street, New York, NY 10024, U.S.A. Lechte, Ms Ruth E., World YWCA: Energy & Environment, Box 9874 Nadi Airport, Nadi, Fiji Lee, Dr William, D.S.I.R., Private Bag, North Dunedin, Dunedin, New Zealand Lehikoinen, Dr Esa, University of Turku, Department of Biology, SF 20500, Turku, Finland Leisler, Dr Bernd, Max-Planek-lnstitute, Verhaltensphysiologie, Vogelwarte, Obstberg, Radolfzell, D7760, Germany Lemon, Prof. Robert, McGill University, 1205 Avenue Docteur Penfield, Montreal, H3A 1 B1 , Canada Leonzio, Dr Claudio, Dipartimento Biologia Ambientale, Via Delle Cerchia 3, 53100 Sienna, Italy Lepson, Mr Jaan K., University of Hawaii, P.O. Box 1 1292, Hilo, HI 96721 , U.S.A. Levey, Dr Doug, University of Florida, Department of Biology, 223 Bartram Hall, Gainesville, FL 32611, U.S.A. Lewis, Mr Milton, Australian National University, Zoology Department, P.O.Box 4, Canberra 2601, Australia Lill, Dr Alan, Monash University, Botany & Zoology Department, Wellington Road, Clayton, Victoria 3168, Australia Lind, Miss Pamela B., Para, Bovey Tracey, Newton Abbot, Devon, TQ13 9JT, United Kingdom Lindberg, Dr Peter, University of Goteborg, Department of Zoology, P.O. Box 6400, Goteborg, S400 31, Sweden Linden, Dr Mats, Uppsala University, Department of Zoology, Box 561, Uppsala, S- 751 22, Sweden Linden, Prof. Harto H. H., Finnish Game & Fisheries Research Institute, Turunlinnantie 8, Helsinki 00930, Finland ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 37 Lister, Ms Heather, Griffith University, Australian Environmental Studies Division, Nathan, Brisbane 4111, Queensland, Australia Liversidge, Dr Richard, McGregor Museum, 92 Central Road, Kimberley 8307, South Africa Liversidge, Mrs Vivienne, C/- Dr R. Liversidge, 92 Central Road, Kimberley 8307, South Africa Lloyd, Mr Brian, Department of Conservation New Zealand, P.O. Box 10-420, Welling¬ ton, New Zealand Lock, Mr Bill, Wallaceville Research Centre, P.O. Box 40063, Upper Hutt, New Zea¬ land Loehrl, Mrs Hildegard, Bei den Eichen 5, D-7271 Egenhausen, Germany Loehrl, Dr Hans, Bei den Eichen 5, D-7271 Egenhausen, Germany Loetscher, Mrs Naomi P., 2064 Cardinal Drive, Danville, KY, 40422, U.S.A. Loetscher Jnr., Dr Frederick W., 2064 Cardinal Drive, Danville, KY, 40422, U.S.A. Loman, Mr Jon, Lund University, Department of Ecology, Helgona vagen 5, 223 62 Lund, Sweden Lopez-Calleja, Dr M. Victoria, Universidad De Chile, Las Palmeras 3425, 653 San¬ tiago, Chile Lotz, Ms Aileen, 6950 SW 71 Ct. , Miami, FL 33143, U.S.A. Louette, Dr Michel, Koninklijk Museum voor Midden-Afrika, Tervuren, 1982, Belgium Lovegrove, Mr Tim, University of Auckland, Zoology Department, Private Bag, Auck¬ land 1 , New Zealand Lovei, Dr Gabor L., DSIR Fruit and Trees, C /- E. Vincze, Private Bag, Palmerston North, New Zealand Lovis, Mrs J. Valerie, Flat 1, 87A Totara Street, Fendalton, Christchurch, New Zea¬ land Lovvorn, Dr James, University of Wyoming, Department of Zoology & Physiology, Laramie, WY 82071, U.S.A. Loyn, Mr Richard H., Department of Conservation Forests and Lands, 123 Brown Street, Heidelberg, Victoria 3084, Australia Lubcke, Dr Ursula, Deutsche Ornithologen Gesellschaft, Oevelgonne 66, Hamburg 52, D2000, Germany Lyall, Mr John, Department of Conservation New Zealand, Private Bag, Hokitika, New Zealand Lyon, Mr Bruce, Princeton University, Department of Biology E.E.B., Princeton N.J., 08544-1003, U.S.A. Macbeth, Mrs Marion, 31 Hewitts Road, Merivale, Christchurch, New Zealand MacMillen, Dr Richard E., University of California, Irvine, Dept of Ecology & Evolution¬ ary Biology, School of Biological Sciences, Irvine, California 92717, U.S.A. Madas, Ms Katalin, Hungarian Ornithological and Nature Protection Society, Timaru. 19, H7621 Pecs, Hungary Maddock, Prof. Max, University of Newcastle & Shortland Wetlands Centre, Newcas¬ tle, NSW 2308, Australia Magrath, Dr Robert D., University of Oxford, Edward Grey Institute, Department of Zoology, South Parks Road, Oxford OX1 3PS, United Kingdom Magrath, Mr Michael, 29 Gingana Street, Aranda, ACT 2614, Australia Major, Dr Richard, The Australian Museum, 6-8 College Street, Sydney, NSW 2000, Australia Mating, Dr Peter, Flat 1, 203B Clyde Road, Fendalton, Christchurch, New Zealand Maloney, Mr Richard, University of Canterbury, 61 Main Road, Redcliffs, Christchurch, New Zealand 38 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Markwell, Mr Tim, 62 Elizabeth Street, Riccarton, Christchurch, New Zealand Marshall, Miss Ulrike, Johann Wolfgang Goethe-University, AG Stoffwechselphysiol, Department of Zoology, Siesmayerstr. 70, Frankfurt, D-6000, Germany Martens, Prof. Jochen, Institut fur Zoologie, Saarstr. 21, Mainz, D-6500, Germany Martin, Dr Graham, University of Birmingham, School of Continuing Studies, Edgbaston, Birmingham, B15 2TT, United Kingdom Martin, Mrs. Marie-Anne, 26 Macaulay Road, Rugby, CV22 6HE, United Kingdom Martin, Dr Jean-Louis, American Museum of Natural History, Department of Ornithol¬ ogy, Central Park West at 79th Street, New York, NY 10024-5192, U.S.A. Martin, Dr Thomas, University of Arkansas, Co-operative Fish & Wildlife Unit, Depart¬ ment of Zoology, Fayetteville, AR 72701, U.S.A. Martin, Ms Nancy L., Vermont Institute of Natural Science, PO Box 86, Woodstock, VT 05091, U.S.A. Martin, Dr Kathy, University of Toronto, Division of Life Sciences, 1265 Military Trail, Scarborough, Ontario MIC 1A4, Canada Martin, Ms Pamela, University of Guelph, Department of Environmental Biology, Guelph, Ontario, NIG 2W1, Canada Mathiasson, Dr Sven H, Natural History Museum, Slottsskogen, Goteborg, 40235, Sweden Mathiasson, Mrs Gunnel, Natural History Museum, Slottsskogen, Goteborg, 40235, Sweden Mathiasson, Mr Jesper, Natural History Museum, Slottsskogen, Goteborg, 40235, Sweden Matson, Ms Shirley, Maruia Society Incorporated, Wild Places Shop 46, Shades Ar¬ cade, Christchurch, New Zealand Mauersberger, Dr Gottfried, Museum fur Naturkunde, Invalidenstrasse 43, Berlin, DDR-1040, Germany Maurer, Dr Brian A., Brigham Young University, Department of Zoology, 574 WTDB, Provo, Utah 84602, U.S.A. Maxwell, Ms Jane, 50 Cheviot Road, Lowry Bay, Eastbourne, Wellington, New Zea¬ land May, Prof Robert, Oxford University, Department of Zoology, South Parks Road, Ox¬ ford, OX1 3PS, United Kingdom May, Ms Judith, Oxford University Press, Walton Street, Oxford, OX2 6DP, United Kingdom May, Ms Margaret L., Toronto Historical Board, C/- Dr J.C. Barlow, Royal Ontario Museum, 100 Queen’s Park, Toronto ONT M53 2C6, Canada M ay h ill, Mr Keith, Ornithological Society of New Zealand, 18 Ruamoana Place, Omokoroa, Tauranga R.D.2, New Zealand Mayhill, Mrs Pauline, 18 Ruamoana Place, Omokoroa, Tauranga R.D.2, New Zealand McAllan, Mr Ian A.W., 46 Yeramba Street, Turramurra, NSW, 2074 Australia McCabe, Prof. Robert A., University of Wisconsin-Madison, College of Agricultural & Life Sciences, 226 Russell Laboratories, 1630 Linden Drive, Madison Wl 53706, U.S.A. McCabe, Mrs, 4501 Keating Terrace, Madison, Wl 53711, U.S.A. McCallion, Mrs Jean, 24A Soleares Avenue, McCormacks Bay, Christchurch, New Zealand McCarthy, Ms Mary, 64 Waimairi Road, llam, Christchurch, New Zealand McCoy, Mr John, 9 Nash Road, Halswell, Christchurch, New Zealand McDonald, Dr Mary V., University of Central Arkansas, Department of Biology, Conway, AR 72032, U.S.A. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 39 McGill, Dr Patricia, Chicago Zoological Society, 3300 Golf Road, Brookfield Zoo, Brookfield, Illinois 60513, U.S.A. Mcllroy, Dr John, CSIRO, Division of Wildlife & Ecology, P.O. Box 84, Lyneham, Can¬ berra, 2602, Australia McIntosh, Mr Angus, University of Otago, Department of Zoology, Great King Street, Dunedin, New Zealand McKean, Mr John L., 1055 East Evelyn Street, Building A, Apt. 1, Sunnyvale, Califor¬ nia 94086, U.S.A. McKee, Dr Joseph, Nuclear Sciences Group - DSIR, P.O. Box 31-418, Lower Hutt, New Zealand McKenzie, Dr Keith, Centrepoint Community, P.O. Box 35, Albany, New Zealand McKilligan, Mr Neil, University College of Southern Queensland, Darling Heights P.O., Toowoomba, 4350, Australia McKilligan, Mrs Helen, University College of Southern Queensland, Darling Heights P.O., Toowoomba, 4350, Australia McKinlay, Mr Bruce, Department of Conservation New Zealand, P.O. Box 5244, Dunedin, New Zealand McKinney, Prof. Frank, University Of Minnesota, Bell Museum of Natural History, 10 Church Street S.E., Minneapolis, MN 55455, U.S.A. McKinney, Mrs Meryl, 4393 Arden View Court, Arden Hills, MN, 55112, U.S.A. McKnight, Ms Margo, University of South Florida, 1814 Marvy Avenue, Tampa, Florida 33612, U.S.A. McLachlan, Mr Andrew, 28 Parker Street, Hornby, Christchurch, New Zealand McLean, Mr Leckie, CSIRO, C /- S. Briggs, P.O. Box 84, Lyneham, ACT 2602, Australia McLean, Dr Ian G., University of Canterbury, Department of Zoology, Private Bag, Christchurch 1, New Zealand McNab, Prof. Brian K., University of Florida, Department of Zoology, Gainesville, FL 32611, U.S.A. McNeil, Prof. Raymond, Universite de Montreal, Departement de Sciences Biologiques, C.P. 6128, Succ. “A”, Montreal, Quebec H3C 3J7, Canada McNicholl, Dr Martin K., I.C.B.P. - Canada, 218 First Avenue, Toronto, MYM 1XY, Canada McPherson, Mr Les, 1/57 Perth Street, Richmond, Christchurch, New Zealand McQueen, Ms Shirley, University of Otago, Medwyn, Reservoir Road, Sawyers Bay, Otago, New Zealand Meathrel, Ms Catherine, Murdoch University, South Street, Murdoch, Perth, Western Australia 6150, Australia Medway, Mr David G., Ornithological Society of New Zealand, 25a Norman Street, New Plymouth, New Zealand Mees, Mr G.F., Rijksmuseum Van Natuurlijke Historie, Postbus 9517, 2300 RA, Leiden, Netherlands Mees-Balchin, Mrs V.J., Grotiuslaan 32, 2353 BW Leiderdorp, Netherlands Mehlum, Dr Fridtjof, Norwegian Polar Research Institute, P.O. Box 158, 1330 Oslo Lufthavn, Norway Melville, Mr David, WWF Hong Kong, GPO Box 12721, Hong Kong Mench, Dr Joy, University of Maryland, Department of Poultry Science. College Park, MD 20742, U.S.A. Mendelssohn, Prof. Heinrich, Tel-Aviv University, Department of Zoology, P.O. Box 39040, Tel-Aviv, 69978, Israel Mendelssohn, Mrs Tamar, Tel-Aviv University, Department of Zoology, Tel-Aviv, 69978, Israel 40 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Merritt, Maj Ron, United States Air Force, Bird Aircraft Strike Hazard, Tyndall Air Force Base, Florida 32403, U.S.A Merton, Mr Donald V., Department of Conservation New Zealand, P.O. Box 10-420, Wellington, New Zealand Metz, Ms Karen J., Carleton University, Department of Biology, Ottawa, Ontario, K1S 5B6, Canada Middleton, Dr Alex L.A., University of Guelph, Department of Zoology, Guelph, NIG 2W1 , Canada Milder, Mr Alvin S., 134 Greenfield Avenue, Los Angeles, CA 90049, U.S.A. Milder, Mrs Sharon L., L.A. County Museum of Natural History, 134 Greenfield Av¬ enue, Los Angeles, CA 90049, U.S.A. Miles, Prof John, University of Otago, Box 17, Lake Hawea, 9190, New Zealand Miller, Dr Edward H., University of Victoria , Royal B.C. Museum, 675 Belleville Street, Victoria B.C., V8V 1X4, Canada Mills, Mr Robert G., University of Otago, Department of Zoology, Box 56, Dunedin, New Zealand Mills, Dr Jim, Department of Conservation New Zealand, Science & Research Divi¬ sion, P.O. Box 10-420, Wellington, New Zealand Mills, Miss Suzanne, P.O. Box 382, Christchurch, New Zealand Milone, Prof. Mario, University of Naples, Department of Zoology, Via Mezzocannone, 80134 Napoli, Italy Minot, Dr Edward O., Massey University, Biology Department, Palmerston North, New Zealand Miskelly, Dr Colin, University of Canterbury, Zoology Department, 1/31 A Lansbury Avenue, Strowan, Christchurch 5, New Zealand Mitchell, Ms Christine, Louisiana State University, U.S. Fish & Wildlife Service, P.O. Box 16028, Baton Rouge, Louisiana 70893, U.S.A. Mock, Dr Douglas W., University of Oklahoma, Norman, OK 73019, U.S.A. Moermond, Prof Timothy C., University Of Wisconsin, Birge Hall, 430 Lincoln Drive, Madison, Wl 53706, U.S.A. Molina, Ms Kathy C., Natural History Museum of L.A. County, 900 Exposition Boul¬ evard, Los Angeles, CA 90007, U.S.A. Moller, Dr Anders P., Uppsala University, Zoology Department, P.O. Box 561, Uppsala S-751 22, Sweden Monaghan, Dr Pat, University of Glasgow, Glasgow G12 8QQ, United Kingdom Mongeon, Ms Marie-Therese, C /- Les Productions Jean-Louis Frund Inc, 4100, Grande coulee, St-Edouard,, Quebec J0K 2H0, Canada Monk, Dr James, Bulletin of BOC, The Glebe Cottage, Goring, Reading, Berkshire RG8 9AP, United Kingdom Monk, Mrs Diana, The Glebe Cottage, Goring, Reading RG8 9AP, United Kingdom Monroe, Mrs Rose S., P.O. Box 23447, Anchorage, KY, 40223, U.S.A. Monroe, Mr Mark S., P.O. Box 23447, Anchorage, KY, 40223, U.S.A. Monroe Jnr., Dr Burt L., University of Louisville, Department of Biology, P.O. Box 23447, Anchorage, KY 40223, U.S.A. Montevecchi, Prof. Bill, Memorial University of Newfoundland, Ocean Sciences Cen¬ tre, St. John’s, A1B 3X9, Canada Montgomerie, Dr Robert, Queen’s University, Department of Biology, Kingston K7L 3N6, Canada Montgomery, Mrs Mary, Society for the Protection of Birds, 309 Melville Avenue, Westmount, Quebec, H3Z 2J7, Canada Moon, Dr Geoff, 19 Waima Crescent, Titirangi, Auckland 7, New Zealand ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 41 Moore, Mrs Amberley, British Ornithologist’s Club, Honorary Secretary, 1 Uppingham Road, Oakham, Rutland, LEI 5 6JB, United Kingdom Moore, Prof Frank, University of Southern Mississippi, Department of Biological Sci¬ ences, Box 5018, Hattiesburg, MS 39406, U.S.A. Moore, Mr Stuart, Department of Conservation New Zealand, Private Bag, Christchurch, New Zealand Moore, Mr Geoffrey J., James Cook University North Queensland, Zoology Depart¬ ment, Townsville, Old 4811, Australia Moorhouse, Mr Ron, Victoria University, Box 600, Kelburn, Wellington, New Zealand Moors, Dr Philip, Royal Australasian Ornithologists Union, 21 Gladstone Street, Moonee Ponds, 3039 Melbourne, Australia Morel, Dr Gerard J., O.R.S.T.O.M., Route de Sallenelles, Breville-Les-Monts 14860, France Morel, Dr Marie-Yvonne, Route de Sallenelles, Breville-Les-Monts, 14860, France Moreno, Dr Juan, Museo Nacional de Ciencias Naturales, J Gutierrez Abascal 2, Madrid 28006, Spain Morin, Ms Marie P., University of Hawaii, Department of Zoology, P.O. Box 3543, Kailua-Kona, HI, 96745, U.S.A. Morioka, Dr Hiroyuki, National Science Museum, Hyakunin-cho 3-23-1, Shinjuku-ku, Tokyo 169, Japan Morris, Dr Ralph D., Brock University, Biological Sciences, St Catharines, Ontario L2S 3A1 , Canada Morris, Mr Chris, R.A.O.U., 32 Peate Avenue, Glen Iris, Melbourne, Vic 3146, Aus¬ tralia Moser, Dr Mike, International Waterfowl and Wetlands Research Bureau, I.W.R.B., Slimbridge, Gloucester, GL2 7BX, United Kingdom Movalli, Dr Paola, University of Milan, Department of Pharmacology, Via Balzaretti, Milano 20124, Italy Mulder, Mr Raoul A., Australian National University, Department of Zoology, G.P.O. Box 4, Canberra, ACT 2601, Australia Muller, Mrs Mary N., Loveday’s Mill, Painswick, Gloucestershire, GL6 6SH, United Kingdom Muller, Mr Charles A., Loveday’s Mill, Painswick, Gloucestershire, GL6 6SH, United Kingdom Mumme, Dr Ronald L., Allegheny College, Department of Biology, Meadville, PA 16335, U.S.A. Mundkur, Mr Taej, Saurashtra University, Department of Biosciences, Rajkot, 360 005, India Munn, Mr Allan, Department of Conservation New Zealand, Chatham Islands, New Zealand Munro, Ms Ursula, University of New England, Department of Zoology, Armidale, 2351 , Australia Munro, Mr Allan, 14 Onehuka Road, Lower Hutt, New Zealand Murie, Dr Jan, University of Alberta, Department of Zoology, Edmonton, Alberta, T6G 2E9, Canada Murphy, Dr Miles E.B., 132 Windermere Road, Hamilton 4007, Brisbane, Queensland, Australia Murphy, Dr Mary, Washington State University, Department of Zoology, Pullman, WA 99164 - 4220, U.S.A. Murphy, Dr Elaine, T.V.N.Z., Natural History Unit, Box 474, Dunedin, New Zealand 42 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Murray, Mr Durno, 17 Ashmore Avenue, Pymble, NSW 2073, Australia Murray, Mrs Barbara, 17 Ashmore Avenue, Pymble, NSW 2073, Australia Murray, Mr David P., Department of Conservation New Zealand, Private Bag, Twizel, New Zealand Murray, Ms Sarah, New Zealand Nagata, Mr Hisashi, Kyushu University, Department of Biology, Faculty of Science, Hakozaki 6-10-1, Higashi-ku, Fukuoka, 812, Japan Nagy, Prof. Ken A., University of California, 405 Hilgard Avenue, Los Angeles, CA 90024-1606, U.S.A. Nagy, Mrs Patricia, University of California, 405 Hilgard Avenue, Los Angeles, CA 90024-1606, U.S.A. Nakajima, Mr Michio, 1383-4 Sezakimati, Sovkasi, 340, Japan Nakamura, Mr Masahiko, Osaka City University, Laboratory of Animal Sociology, Faculty of Science, Sugimoto, Sumiyoshi-ku, Osaka 558, Japan Nakamura, Dr Kazuo, National Agricultural Research Centre, Bird Control Lab., Kannondai, Tsukuba City 305, Japan Nakamura, Prof. Tsukasa, Yamanashi University, Department of Biology, Kofu 400, Japan Nee, Dr Sean P., AFRC Unit of Ecology & Behaviour, Department of Zoology, South Parks Road, Oxford, OX1 3PS, United Kingdom Nee, Mr Mark, 1 1 Shortland Street, Napier, New Zealand Neergaard, Mr Raimo, University of Gothenburg, Department of Zoology, Section of Animal Ecology, Box 25059, S-40031 Gothenburg, Sweden Neill, Miss Molly, 15A Wairere Grove, Paraparaumu, New Zealand Nelson, Dr Joseph B., Aberdeen University, Balkirk, Glenlochar, Castle Douglas, DG72LU, Scotland, United Kingdom Nelson, Mr Dean, Department of Conservation New Zealand, P.O. Box 1130, Dunedin, New Zealand Nettleship, Dr David N., Canadian Wildlife Service, Bedford Institute of Oceanogra¬ phy, P.O. Box 1006, Dartmouth, N.S. B2Y 4A2, Canada Neve de Mevergnies, Mr Gabriel, Unite Ecologie & Biogeog, , Universite Catholique de Louvain, Place Croix du Sud 5, B-1348 Louvain-La-Neuve, Belgium Newton, Dr Ian, Institute of Terrestrial Ecology, Monks Wood Experimental Station, Abbots Ripton, Huntingdon, PEI 7 2LS, United Kingdom Newton, Mrs Halina, Institute of Terrestrial Ecology, Monks Wood Experimental Sta¬ tion, Abbots Ripton, Huntingdon, PEI 7 2LS, United Kingdom Niemi, Prof. Gerald J., Natural Resources Research Centre, 5013 Miller Trunk High¬ way, Duluth, MN 55811, U.S.A. Nilsson, Mr Ron, New Zealand Nol, Dr Erica, Trent University, Biology Department, Peterborough, K9J 7B8, Canada Nolan, Prof Val, Indiana University, Department of Biology, Bloomington, IN 47405, U.S.A. Noon, Dr Barry R., U.S. Forest Service, Redwood Sciences Laboratory, 1700 Bayview Drive, Areata, California 95521, U.S.A. Noon, Mrs Paige T., 1871 Golf Course Road, Areata, California 95521, U.S.A. Nores, Dr Manuel, Centro de Zoologia Aplicada, Box 122, 5000 Cordoba, Argentina Noske, Dr Richard, Northern Territory University, Dripstone Road, Ellengowan Drive, Casuarina, Darwin 0811, Australia Ntiamoa-Baidu, Dr Yaa, University of Ghana, Zoology Department, P.O. Box 67, Legon, Accra, Ghana Nuechterlein, Prof. Gary, North Dakota State University, Fargo, ND 58105, U.S.A. ACTA XX CONGRESSUS INTERNATION ALIS ORNITHOLOGICI 43 Nugroho, Dr E., Indonesian Swiftlet Lovers, Jalan Indrapura 4 Candi Baru, Semarang, 50132, Indonesia Nye, Mr Greg, Unit 3/11 Newton Street, P.O. Box 234, Coorparoo, 4151, Australia Nygard, Mr Torgeir, Norwegian Institute for Nature Research, Tungasletta 2, Trondheim 7004, Norway O’Brien, Mr Rory, 3 Valentine Avenue, Kew, Victoria, Australia O’Donnell, Dr Colin, Department of Conservation New Zealand, Private Bag, Christchurch, New Zealand Oakley, Miss Margaret, 4 Warbler Grove, Waikanae, New Zealand Oates, Mr Kerry E., Ornithological Society of New Zealand, 12 Jackson Terrace, Porirua, Wellington, New Zealand Oba, Dr Teruyo, Natural History Museum & Institute, 955-Z AOBA-CHO, Chiba 280, Japan Obst, Prof. Bryan S., University of California, Department of Biology, Los Angeles, CA 90024, U.S.A. Ohori, Mr Satoshi, Waseda University, Environmental Research Laboratory, 2-579-15, Mikajima, Tokorozawa, 359 Saitama, Japan Oka, Miss Nariko, Yamashina Institute for Ornithology, Khonoyama, Abiko City, Chiba Prefecture 270-1 1 , Japan Olstad, Ms Ronnaug, C/- Dr Kjetil Bevanger, Norwegian Institute for Nature Research, Tungasletta 2, Trondheim N-7004, Norway Ormerod, Dr Steve, University of Wales, C /- N.R.A., 19 Penyfai Lane, LLanelli, Dyfed SA15 4EL, United Kingdom Orchard, Ms Suzanne M., 50 French Street, Masterton, New Zealand Oring, Prof. Lewis, University of North Dakota, Department of Biology, Grand Forks, ND 58202, U.S.A. Osborne, Mrs Yvonne, Osborne Editions International, P.O. Box 1768, Auckland, New Zealand Osieck, Dr Eduard R., Netherlands Society For The Protection Of Birds, Driebergseweg 16-C, 3708 JB, Zeist, Netherlands Ouellet, Dr Henri, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa KIP 6P4, Canada Owen, Dr Myrfyn, The Wildfowl and Wetlands Trust, Slimbridge, Gloucester, GL2 7BT, United Kingdom Owen, Mrs L.M.V., C/- Dr Myrfyn Owen, The Wildfowl & Wetlands Trust, Slimbridge, Gloucester GL2 7BT, United Kingdom Owen, Mr Keith, Department of Conservation New Zealand, 24 Sloane Avenue, Tihiotonga, Rotorua, New Zealand Owens, Mr Ian, University of Leicester, Zoology Department, University Road, Leices¬ ter, LEI 7RH, United Kingdom Paasivirta, Mr Olli, Ministry of the Environment, Ratakatu 3, Helsinki, 00120, Finland Pain, Dr Deborah, Station Biologique De La Tour Du Valat, Le Sambuc, Arles, 13200, France Palmer, Mr John, Arnold Books, 11 New Regent Street, Christchurch, New Zealand Parkin, Dr David T., University of Nottingham, Department of Genetics, School of Medicine, Queens Medical Centre, Nottingham NG7 2UH, United Kingdom Parrish, Mr Richard, Department of Conservation New Zealand, 143 Church Street, Onerahi, Whangarei, New Zealand Part, Dr Tomas, Uppsala University, Department of Zoology, Box 561, S-75122 Uppsala, Sweden 44 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Paterson, Mr Adrian, University of Otago, Department of Zoology, P.O. Box 56, Dunedin, New Zealand Paton, Dr David, University of Adelaide, Department of Zoology, GPO Box 498, Ad¬ elaide, SA 5001, Australia Peal, Mr Ronald, British Ornithologists’ Club, 2 Chestnut Lane, Sevenoaks, Kent, TN13 3AR, United Kingdom Peal, Mrs Betty, 2 Chestnut Lane, Sevenoaks, Kent, TN13 3AR, United Kingdom Pearson, Dr David, Arizona State University, Department of Zoology, Tempe, AZ 85287, U.S.A. Peckover, Mr Bill, 14 Balanda Street, Jindalee, Queensland 4074, Australia Peckover, Mrs Joan, 14 Balanda Street, Jindalee, Queensland 4074, Australia Pemberton, Ms Ruth, 68B Mortimer Terrace, Brooklyn, Wellington, New Zealand Pepperberg, Prof. Irene M., University of Arizona, Dept of Ecology & Evolutionary Biology, Biological Sciences West, Tucson, Arizona 85721, U.S.A. Perez-Rivera, Prof Raul A., University of Puerto Rico, Humacao Campus, Department of Biology, C.U.H. Station, Humacao 00661, Puerto Rico Perrins, Dr Christopher, Edward Grey Institute, Department of Zoology, South Parks Road, Oxford OX1 3PS, United Kingdom Perrins, Mrs Mary, Edward Grey Institute, C/O Dr C.M. Perrins, Department of Zool¬ ogy, South Parks Road, Oxford OX1 3PS, United Kingdom Pessino, Ms Catherine, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, U.S.A. Petch, Mrs Sheila, 90 Balrudry Street, Avonhead, Christchurch, New Zealand Peters, Dr Margarete, Senckenberg-Anlage 25, Frankfurt a.M., D-6000, Germany Peters, Prof. D. Stefan, Forschungsinstitut Senckenberg, Senckenberg-Anlage 25, Frankfurt a.M. D-6000, Germany Peters, Ms Michele A., Wildlife Rescue Inc., 10432 Casador del Oso N.E., Albuquerque, NM 87111-3771, U.S.A. Petersen, Mr Peter C., Wm. Butterworth Memorial Trust, 235 McClellan Blvd., Dav¬ enport, IA 52803, U.S.A. Peterson, Dr Roger T., P.O. Box 825, Old Lyme, Connecticut 06371 , U.S.A. Peterson, Mrs Virginia M, P.O. Box 825, Old Lyme, Connecticut 06371, U.S.A. Petrie, Dr Marion, The Open University, Department of Biology, Walton Hall, Milton Keynes, MK76AA, United Kingdom Phillipson, Mr Stephen, P.O. Box 8, Arthur’s Pass, New Zealand Piatt, Dr John, Alaska Fish and Wildlife Research, 1011 E. Tudor Road, Anchorage, AK 99503, U.S.A. Pickwell, Mr Warwick R., 29 Drem Street, Toogoolawah, 4313, Australia Pienkowski, Dr Mike, Nature Conservancy Council, Northminster House, Peterbor¬ ough, PEI 1UA, United Kingdom Pierce, Dr Ray, Department of Conservation New Zealand, P.O. Box 482, Whangarei, New Zealand Piersma, Mr Theunis, Netherlands Institute for Sea Research & Zoological Laboratory, University of Groningen, P.O. Box 59, 1790 AB Den Burg, Texel, Netherlands Pietz, Dr Pamela, Northern Prairie Wildlife Research Center, Route 1, Box 96c, Jamestown, ND 58402, U.S.A. Pinowski, Prof. Jan K., Institute of Ecology PAS, Department of Vertebrate Ecology, Dziekanow Lesny, 05-092 Tomianki, Poland Pinxten, Ms Rianne, University of Antwerp, U.I.A., Department of Biology, Universiteitsplein 1, B-2610 Wilrijk, Belgium ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 45 Place, Prof. Allen R., University of Maryland , System Center of Marine Biotechnology, 600 East Lombard Street, Baltimore, MD 21202, U.S.A. Plenge, Mr Manuel A., Apartado 18-0839, Lima 18, Peru Pohjanen, Ms Elisabet, Blodstensvagen 17, 75244 Uppsala, Sweden Ponganis, Dr Katherine, Scripps Institution of Oceanography, University of California, 13326 Landfair Road, San Diego, CA 92130, U.S.A. Poonswad, Ms Pilai, Mahidol University, Department of Microbiology, Faculty of Sci¬ ence, Rama 6 Road, 10400 Bangkok, Thailand Potter, Dr Murray, 11 Upham Terrace, Palmerston North, New Zealand Powell, Ms Abby N., University of Minnesota, Department of Fisheries and Wildlife, 1980 Folwell Avenue, St Paul, MN 55108, U.S.A. Power, Dr Dennis M., Santa Barbara Museum of Natural History, 2559 Puesta del Sol, Santa Barbara, CA 93105, U.S.A. Power, Mrs Leslie, C /- Dr. Dennis Power, Santa Barbara Museum of Natural History, 2559 Puesta del Sol, Santa Barbara, CA 93105, U.S.A. Powlesland, Dr Ralph, Department of Conservation, Conservation Sciences Centre, P.O. Box 10420, Wellington, New Zealand Powlesland, Mrs Mary, 64 Roseneath Terrace, Roseneath, Wellington, 6001, New Zealand Poysa, Dr Hannu, Finnish Game & Fisheries Research Institute, Evo Game Research Station, Evo SF-16970, Finland Pratt, Dr Thane K., U.S. Fish & Wildlife Service, Hawaii Research Station, P.O. Box 44, Hawaii National Park, HI 96718, U.S.A. Prawiradilaga, Miss Dewi, Centre for Research & Dev. in Biology, P.O. Box 110, 16122 Bogor, Indonesia Price, Ms lola, Dept Fisheries & Oceans, 563 Fairview Avenue, Ottawa, KIM 0X4, Canada Priddel, Dr David, New South Wales National Parks & Wildlife Service, Box 1967, Hurstville, Sydney, NSW 2220, Australia Pringle, Mrs Gail, 14a Jacaranda Place, Queenspark, Christchurch, New Zealand Prinzinger, Prof. Roland, University of Frankfurt, Zoological Institute, Siesmayerstrasse 70, 6000 Frankfurt, Germany Prove, Prof. Ekkehard, Universitat Bielefeld, Fakultat Biologie, Lehrstuhl fur Verhaltensphysiologie, P.O. Box 8640, D-4800 Bielefeld, Germany Prum, Dr Richard O., American Museum of Natural History, 237 W 18th Street, Appt 5RW, New York, NY 10011, U.S.A. Pulham, Ms. Gwenda, O.S.N.Z. New Zealand, Unit2, 1 Parkhill Road, Birkenhead, Auckland 10, New Zealand Quayle, Mrs Gail, 6 Tresillian Avenue, Atawhai, Nelson, New Zealand Quinn, Dr Tom W., University of California, Department of Biochemistry, Berkeley, CA 94618, U.S.A. Quinn, Mr Patrick J., Ornithological Society of New Zealand, 41 Aorangi Crescent, P.O. Box 5, Lake Tekapo, New Zealand Quinn, Ms Megan E., Ornithological Society of New Zealand, P.O. Box 5, 41 Aorangi Crescent, Lake Tekapo, New Zealand Rackham, Mr Allan, Boffa Miskell Partners, P.O. Box 110, Christchurch, New Zealand Radcliffe, Dr Joan, 28 Newbridge Place, llam, Christchurch, New Zealand Ralph, Dr C.John, U.S. Forest Service, 1700 Bayview Drive, Areata, CA 95521, U.S.A. Ramenofsky, Dr Marilyn, University of Washington, Department of Zoology NJ-15, Seattle, WA 98195, U.S.A. 46 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Ramos-Olmos, Dr Mario, World Wildlife Fund & Conservation Foundation, 1250 24th Street NW, Suite 500, Washington, DC 20037, U.S.A. Randi, Dr Ettore, Instituto Nazionale di Biologia della Selvaggina, Via Ca Fornacetta 9, 1-40064 Ozzano Emilia (BO), Italy Rands, Dr Michael, International Council For Bird Preservation, 32 Cambridge Road, Girton, Cambridge, CB3 0PJ, United Kingdom Rands, Dr Gillian F., Trends in Ecology & Evolution, 68 Hills Road, Cambridge, CB2 1 LA, United Kingdom Rappoport, Ms Ann, U.S. Fish & Wildlife Service, 1011 E. Tudor Road, Anchorage, Alaska 99503, U.S.A. Rasch, Ms Gretchen, Department of Conservation New Zealand, P.O. Box 10-420, Wellington, New Zealand Rasmussen, Ms Pamela C., University of Kansas Museum of Natural History, Dyche Hall, University of Kansas, Lawrence, Kansas 66045-2454, U.S.A. Ratcliffe, Dr Laurene, Queen’s University, Department of Biology, Kingston, Ontario K7L 3N6, Canada Rayner, Dr Jeremy, University of Bristol, Department of Zoology, Woodland Road, Bristol BS8 IU8, United Kingdom Recher, Dr Harry F., University of New England, Armidale, NSW 2351, Australia Reed, Miss Christine, Department of Conservation New Zealand, Private Bag, Twizel, New Zealand Reese, Mr Peter, 4A Gibbs Place, Halswell, Christchurch, New Zealand Reid, Mr Ian, O.S.N.Z., 629 Grey Street, Claudelands, Hamilton, New Zealand Reilly, Ms Pauline, RAOU, 19 Lialeeta Avenue, Box 67, Fairhaven, Victoria 3221, Australia Reinertsen, Dr Randi, University of Trondheim, Department of Zoology, N-7055 Trondheim, Norway Reitan, Mr Ole, Norwegian Institute for Nature Research, Tungasletta 2, N-7004 Trondheim, Norway Rhymer, Dr Judith A., Smithsonian Institution, Laboratory of Molecular Systematics MSC, Washington, DC 20560, U.S.A. Ribeiro, Miss Anna, Universidade Federal Do Rio De Janeiro, CCS Dept Zoologia Inst. Biologia, llha Do Fundao, P.O. Box 21 941, Rio de Janeiro, Brazil Richards, Ms Mary, 24 Tarikaka Street, Ngaio, Wellington, New Zealand Richardson, Dr Ken, Murdoch University, Veterinary Studies, South Street, Murdoch, Perth, Western Australia 6150, Australia Richardson, Dr W. John, LGL Ltd., Environmental Research Associates, P.O. Box 280, King City, Ontario LOG 1K0, Canada Richardson, Mrs Dorothy, C /- LGL Ltd, P.O.Box 280, King City, Ontario, LOG 1K0, Canada Richford, Dr Andrew, Academic Press/Poyser Limited, 24-28 Oval Road, London, NW1 7DX, United Kingdom Ricklefs, Prof. Robert E., University of Pennsylvania, Department of Biology, Phila¬ delphia, PA 19104-6018, U.S.A. Rinke, Dr Dieter, Brehm Fund South Seas Expedition, Private Bag 52, Nuku’alofa, Kingdom of Tonga Rising, Prof. James D., University of Toronto, Department of Zoology, Toronto, On¬ tario, M5S 1A1, Canada Risley, Mr Christopher, C /- Dr Erica Nol, Trent University, Peterborough, K9J 7B8, Canada ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 47 Ristau, Dr Carolyn, Rockefeller University, 1230 York Avenue, New York, NY 10021, U.S.A. Rivera, Ms Valli M., Calle Espana D-3, Urb. Oasis Garden, 00657 Guaynabo, Puerto Rico Robbins, Mr Mark, Academy of Natural Sciences, 19th & The Parkway, Philadelphia, PA 19103, U.S.A. Roberts, Dr Julie, University of New England, Armidale, NSW 2351, Australia Roberts, Mr Bernie, 25B Sumnervale Drive, Sumner, Christchurch, New Zealand Robertson, Mr Christopher J.R., N.Z. Department of Conservation, P.O. Box 12397, Wellington, New Zealand Robertson, Mrs Gillian B.H., P.O. Box 12397, Wellington, New Zealand Robertson, Dr Raleigh J, Queens University, Department of Biology, Kingston, On¬ tario, K7L 3N6, Canada Robertson, Mr Peter, University of East Anglia, University Plain, Norwich, NR4 7TJ, United Kingdom Robertson, Dr David G., 11 Shaftsbury Street, Eden Hills, SA 5050, Australia Robertson, Mrs Minnie, 11 Shaftsbury Street, Eden Hills, SA 5050, Australia Robertson, Dr Hugh A., Department of Conservation New Zealand, P.O. Box 5086, Wellington, New Zealand Robinson, Ms Ann, Griffith University, Nathan, Brisbane, OLD 4111, Australia Rogers, Mr Danny, Royal Australasian Ornithologist’s Union, 21 Gladstone Street, Moonee Ponds, Melbourne, VIC 3039, Australia Rogers, Mr Alan, RAOU, 26 Westleigh Drive, Westleigh, NSW 2120, Australia Root, Dr Terry, University of Michigan, 430 East University, Ann Arbor, Ml 48109, U.S.A. Roper-Lindsay, Dr Judith, Forestry Road, Ashley, RD2, Rangiora, New Zealand Rosner, Mr Hans-Ulrich, Hohle Gasse 5, D2250 Husum, Germany Ross, Dr Graham, Australian Biological Resources Study, Australian National Botanic Gardens, P.O. Box 1383, Canberra, ACT 2601, Australia Rothwell, Mrs Nan, 3 Marine Terrace, Bayswater, Auckland, New Zealand Rounce, Mr John, 2/146 Victoria Avenue, Palmerston North, New Zealand Rov, Mr Nils, Norwegian Institute for Nature Research, Tungasletta 2, N-7004 Trondheim, Norway Rowe, Mrs Stella, O.S.N.Z., 1 Corrin Street, Hamilton, New Zealand Rowe, Mr John, O.S.N.Z., 1 Corrin Street, Hamilton, New Zealand Rowell, Ms Alison, Civil Aviation Authority, 14 Wales Street, Belconnen, Canberra, ACT 261 7, Australia Rowley, Mr Ian, CSIRO, Division of Wildlife & Ecology, Midland, Box LMB 4, Perth, WA 6056, Australia Ruff, Mr Robert, University of Wisconsin, Department Chair, 226 Russell Labs, Madi¬ son, Wl 53706, U.S.A. Rusch, Dr Donald H., University of Wisconsin, 21 1 Russell Labs, 1630 Linden Drive, Department of Wildlife Ecology, Madison, Wl 53620, U.S.A. Russell, Dr Eleanor M., CSIRO, Division of Wildlife & Ecology, Midland, Box LMB 4, Perth, WA 6056, Australia Russell, Ms Gail L., Armata Environmental Consultants, 227 River Street, 2710 Deniliquin, Australia Rutschke, Prof. Erich, Centre for Wildfowl Research GDR, Allee nach Sanssouci, Villa Liegnitz, Potsdam, 1570, Germany Ryan, Mr Peter, University of Capetown, Fitzpatrick Institute, Rondebosch 7700, South Africa 48 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Ryder, Prof. John P., Lakehead University, Oliver Road # 955, Thunder Bay, P7B 5EI, Canada Ryder, Prof. Ronald A., Colorado State University, Dept, of Fisheries & Wildlife Biol¬ ogy, Ft. Collins, CO 80523, U.S.A. Ryder, Mrs Audrey, 748 Eastdale Drive, Fort Collins, Colorado 80524, U.S.A. Sadleir, Dr Richard, Department of Conservation New Zealand, P.O. Box 10-420, Wellington, New Zealand Safina, Dr Carl, National Audubon Society, 306 South Bay Avenue, Islip, New York, NY 11751, U.S.A. Sagar, Mr Paul, 38A Yardley Street, Christchurch 4, New Zealand Sageder, Dr. Gabi, Institut f. Zoologie, Althenstr. 14, Vienna A-1090, Austria Saitou, Dr Takashi, University of Tsukuba, Institute of Biological Sciences, 2-3 Samon-cho, Shinjuku, Tokyo, 160, Japan Salama, Mr Mohamed, National Commission for Wildlife Conservation and Develop¬ ment, P.O. Box 61681, Riyadh, Saudi Arabia Salpeter, Mrs Rosalia, 47 La Boheme Avenue, Caringbah, Sydney, NSW 2229, Aus¬ tralia Sanaiotti, Ms Tania, Depto de Ecologia, Inpa, C.P. 478 Manaus, AM,, Brazil Sandberg, Mr Roland C.E., University of Lund, Department of Animal Ecology, Ecol¬ ogy Building, Helgonav 5, Lund, S-223 62, Sweden Sandley, Ms Sarah, Oxford University Press, P.O. Box 11149, Ellerslie, Auckland, New Zealand Saris, Mr Frank J. A., Sovon, P.O. Box 81, 6573 ZH Beek-Ubbergen, Netherlands Satterthwaite, Mr Charles, 228 Main Road, Monck’s Bay, Christchurch, New Zealand Saunders, Dr Denis A., CSIRO, Division of Wildlife & Ecology, Midland, Box LMB 4, Perth, WA 6056, Australia Saunders, Mrs V., CSIRO, Division of Wildlife & Ecology, C /- Dr Denis Saunders, Midland, Box LMB 4, Perth, WA 6056, Australia Saunders, Dr Alan, Department of Conservation New Zealand, 154 Campbell Street, Karori, Wellington, New Zealand Saunders, Mr Alec, New Zealand Saurola, Mr Pertti, University of Helsinki, Zoological Museum, P. Rautatiekatu. 13, SF- 00100 Helsinki, Finland Saurola, Mrs Helmi-lrene, Kylatie 20 A 15, SF-00320 Helsinki, Finland Schelbert, Mr Bruno, Baudepartement Aargau, Neumattstrasse 4, Widen 8967, Swit¬ zerland Schels, Dr Herbert F., FAO, Kornweg 5, Mainleus 8653, Germany Schels, Mrs Christa, Kornweg 5, Mainleus 8653, Germany Schenk, Mr Gary, C/- Mrs Betty Schreiber, Natural History Museum, 900 Exposition Boulevard, Los Angeles, CA 90007, U.S.A. Schleucher, Miss Elke, JWG University AG Stoffwechsel Physiologie, Siesmayerstrasse 70, Frankfurt M., D-6000, Germany Schmid, Dr Herbert, Max-Planck-lnstitut, W.G. Kerckhoff-lnstitut, Parkstrasse 1, D- 6350 Bad Nauheim, Germany Schmidt-Koenig, Prof. Klaus, Verhaltensphysiologie, Beim Kupferhammer 8, Tubingen, D-7400, Germany Schmokel, Ms Bianca, Seelingstrasse 32, D-1000 Berlin 19, Germany Schodde, Dr Richard, CSIRO, Division of Wildlife & Ecology, Box 84, Gungahlin, Lyneham, Canberra, ACT 2602, Australia Schodde, Mrs Rosemary, 30 Bamford Street, Hughes, Canberra, ACT 2605, Australia ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 49 Schreiber, Mrs Betty A., Natural History Museum, 900 Exposition Boulevard, Los Angeles, CA 90007, U.S.A. Schuchmann, Dr Karl L., A. Koenig Zoological Institute & Zoological Museum, Adenauerallee 150-164, Bonn 1, D-5300, Germany Schuckard, Mr Rob, Te Towaka, R D 3, Rai Valley, New Zealand Schulz, Dr Holger, National Wildlife Research Centre, P.O. Box 1086, Taif, Saudi Ara¬ bia Schuster, Dr Astrid, Nationalparkverwaltung Berchtesgaden, Doktorberg 6, D-8240 Berchtesgaden, Germany Scofield, Mr Paul, University of Auckland, 8 Crocus Place, Remuera, Auckland 5, New Zealand Seay, Mr David, 9520 Poole Street, La Jolla, California 92037, U.S.A. Seddon, Mrs Betty H., 11 Grey Street, Cambridge, Waikato, New Zealand Seto, Ms Atsu, The Ornithological Society of Japan, 2-3-6, Aogakidai, Nara, 631, Japan Seto, Ms Kyoko, 7-2-204, Tsukiwaka-cho, Ashiya, 659, Japan Seubert, Dr John, U.S. Department of Agriculture, 1800 Zinnia Street, Golden, Colo¬ rado 80401 , U.S.A. Seubert, Mrs Jean, C /- Dr J Seubert, U.S. Department of Agriculture, 1800 Zinnia Street, Golden, Colorado 80401, U.S.A. Seutin, Mr Gilles, Queen’s University, Department of Biology, Kingston, Ontario K7L 3N6, Canada Severinghaus, Dr Sheldon, The Asia Foundation, Box 3223, San Francisco, 94119, U.S.A. Severinghaus, Dr Lucia Liu, Academia Sinica, Institute of Zoology, Nankang, Taipei, 1 1 529, Taiwan Shannon, Dr. Geoff, 13 Banksia Street, Bunbury, WA 6230, Australia Shannon, Mr George, The Lower House, Stoney Lane, Tardebigge, Bromsgrove Worcs. B60 1LY, United Kingdom Sharrock, Dr Tim, British Birds, Fountains, Park Lane, Blunham, Bedford, MK44 3NJ, United Kingdom Shepard, Ms Margaret B., Cornell University, Section of Ecology & Systematics, Ithaca, NY 14853, U.S.A. Shepherd, Mr Michael, University of Nottingham, Department of Zoology, University Park, Nottingham, NG 72D, United Kingdom Sherley, Dr Greg, Department of Conservation New Zealand, Science & Research Division, 1 1 1 Taylor Terrace, Tawa, Wellington, New Zealand Sherry, Dr Thomas W., Tulane University, Department of Biology, 2000 Percival Stern Hall, New Orleans, LA 70118, U.S.A. Shobrak, Mr Mohamed, National Commission for Wildlife Conservation and Develop¬ ment, P.O. Box 61681, Riyadh, Saudi Arabia Short, Prof. Lester, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, U.S.A. Short, Mr Jeffrey J., USAFR, 259 Eutaw Forest Drive, Waldorf, MD 20603, U.S.A. Sibley, Prof. Charles G., San Francisco State University, 95 Seafirth Road, Tiburon, CA 94920, U.S.A. Sibley, Mrs Frances, 95 Seafirth Road, Tiburon, CA 94920, U.S.A. Silveira, Mr Charles E., 4 Payne Street, Gladstone Park, Melbourne, VIC 3043, Aus¬ tralia Silverin, Dr Bengt, University of Gothenburg, Department of Zoology, Box 25059, Gothenburg, S-40031, Sweden 50 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Simasko, Dr Joel, P.O. Box 3543, Kailua-Kona, HI, 76745, U.S.A. Simpson, Dr Marie M., 13 Warrington Road, Remuera, Auckland, New Zealand Sirgouant, Ms Serge, Societe Caledonienne d’Ornithologie, P.O. Box 4338, Noumea, New Caledonia Sjoberg, Dr Kjell, Swedish University of Agricultural Sciences, Department of Wildlife Ecology, S-901 83 Umea, Sweden Slack, Mr Roy, C /- Post Office, Pauatahanui, Wellington, New Zealand Sladen, Mr Fred W., Winchester Tours Ltd., P.O. Box 706, New London, New Hamp¬ shire 03257, U.S.A. Sladen, Mrs Phyllis M., P. O. Box 706, New London, New Hampshire 03257, U.S.A. Sladen, Prof. William J.L., Johns Hopkins University, C /- Airlie Center, Airlie, VA 22186, U.S.A. Slagsvold, Dr Tore, University of Oslo, Zoological Museum, Sarsgate 1 , N-0562 Oslo 5, Norway Slater, Prof. Peter, University of St Andrews, Bute Building, St Andrews, Fife, KYI 6 9TS, United Kingdom Slater, Dr Fred, University of Wales College of Cardiff, Llysdinam Field Centre, Newbridge-On-Wye, Llandrindod Wells, Powys LD1 6NB, United Kingdom Slotta-Bachmayr, Mr Leopold, Universitat Salzburg, Rettenpacherstrasse 5, Salzburg 5020, Austria Small, Ms Delle, 23 Maarama Crescent, Brooklyn, Wellington, New Zealand Smith, Mrs Susan, Everglades National Park, P.O. Box 1341, Homestead, Florida 33090, U.S.A. Smith, Mr P. William, P.O. Box 1341, Homestead, Florida 33090, U.S.A. Smith, Dr Susan M., Mount Holyoke College, Department of Biology, South Hadley, Mass, 01075, U.S.A. Smith, Mrs Meredith, Flat 30, 64 Cunliffe Street, Johnsonville, Wellington 4, New Zea¬ land Smith, Dr Geoffrey C., NSW National Parks & Wildlife Service, P.O. Box 1967, Hurtsville, Sydney, NSW 2220, Australia Smith, Dr Kimberly G., University of Arkansas, Department of Zoology, Fayetteville, AR 72701, U.S.A. Smith, Ms Peggy J., University of Arkansas, Department of Zoology, Fayetteville, AR 72701, U.S.A. Smith, Dr Henrik G., Lund University, Department of Ecology, Animal Ecology, Helgonavagen 5, 22362 Lund, Sweden Smith, Dr Thomas B., University of California, Department of Integrative Biology, Berkeley, CA 94720, U.S.A. Smith, Ms Sue, 43 Dudley Street, Shirley, Christchurch, New Zealand Smith, Jamie, University of British Columbia, Department of Zoology, 6270 University Boulevard, Vancouver, British Columbia, Canada Soler, Prof. Manuel, Universidad de Granada, Faultad de Ciencias, Fuente Nueva S/ N, Granada, 18001, Spain Somadikarta, Dr S., University of Indonesia, Jalan Salak 12, Bogor 16151, Indonesia Sonnenburg, Mr Holger, Turnerstrasse 3A, 4500 Osnabruck, Germany Sorenson, Dr Michael, Smithsonian Institution, National Zoological Park, Conserva¬ tion & Research Centre, Front Royal, Virginia 22630, U.S.A. Sorenson, Dr Lisa Guminski, Smithsonian Institution, National Zoological Park, Con¬ servation & Research Centre, Front Royal, Virginia 22630, U.S.A. Spaans, Dr Arie L., Research Institute for Nature Management, Kemperbergerweg 67, 6816 RB Arnhem, Netherlands ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 51 Spaans-Scheen, Mrs Marianne J., Research Institute for Nature Management, C /- Dr Arie L. Spaans, Kemperbergerweg 67, 6816 RB Arnhem, Netherlands Spencer, Mr Michael D., 1321 Sherbrooke Street West D-31 , Montreal, Quebec, H3G 1J4, Canada Sperling, Ms Jette, C /- Dr Claus Bech, University of Trondheim, Dragvoll, Trondheim N-7055, Norway Spidso, Mr Tor K., Department of Nature Conservation, Box 39, As-NIH, N-1432, Norway Spina, Dr Fernando, Istituto Nazionale di Biologia della Selvaggina, Via Ca’ Fornacetta 9, 1-40064 Ozzano Emilia (Bo), Italy Springer, Ms Paige, 275 Gravatt Drive, Berkeley, CA 94705, U.S.A. Spurr, Dr Eric, Forest Research Institute, P.O. Box 31-011, Christchurch, New Zea¬ land St. Clair, Ms Colleen C., University of Canterbury, Department of Zoology, Christchurch 1, New Zealand Stahl, Mr Jean-Claude, Victoria University, 48a McKinley Crescent, Brooklyn, Welling¬ ton, New Zealand Stangel, Dr Peter, Savannah River Ecology Laboratory, Drawer E, Aiken, South Caro¬ lina, U.S.A. Stark, Mr David, 2 Harland Road, Castletown, Thurso, KW14 8UB, United Kingdom Stattersfield, Ms Alison, 32 Cambridge Road, Girton, Cambridge, United Kingdom Steidal, Mr Gunnar, University of Jena, Section Biology, Branch Ecology, Neugasse 23, Jena, 6900, Germany Sternberg, Mr Helmut, Im Schapenkamp 11, Braunschweig, D-3300, Germany Stettenheim, Dr Peter, Biographies of North American Birds, Meriden Road, P.O. Box #64-255, Lebanon, NH 03766, U.S.A. Stettenheim, Mrs Sandy, Meriden Road, P.O. Box #64-255, Lebanon, NH 03766, U.S.A. Stevens, Dr Susan, 12 Anderson Street, Westmead, 2145, Australia Stewart, Mrs Helen, Ornithological Society of New Zealand, 1005 Port Road, Whangamata, New Zealand Stewart, Dr Anne, Department of Conservation New Zealand, Private Bag 8, Auck¬ land, New Zealand Stewart, Ms Annie, 6/101 Carlton Mill Road, Merivale, Christchurch, New Zealand Stewart-Cox, Ms Belinda, Long Mead, Brixton Deverill, Warminster, Wiltshire BA12 7EJ, United Kingdom Stickney, Mrs Eleanor, Yale University, Peabody Museum, 700 Leetes Island Road, Guilford, CT 06437, U.S.A. Stickney, Jnr., Mr. Albert, Yale University, Peabody Museum, 700 Leetes Island Road, Guilford, CT 06437, U.S.A. Stinson, Mr Derek, Division of Fish & Wildlife, Commonwealth of Northern Mariana Islands, Saipan, MP 96950, U.S.A. Stinson, Ms Colleen, Northern Marianas College, P.O. Box 1250, Saipan, MP 96950, U.S.A. Stokes, Ms Sarah Jayne, University of Canterbury, 14 Patterson Tee, Halswell, Christchurch, New Zealand Stokes, Mr Adrian, University of New England, Department of Ecosystem Manage¬ ment, Armidale, NSW 2351 , Australia Stone, Dr Charles, National Park Service, Tuscon, Arizona, U.S.A. Storch, Ms Use, University of Munich, Inst, of Wildlife Research & Management, Amalienstrasse 52, 8000 Munich 40, Germany 52 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Stracy, Mr Don, 14 Fitzwilliam Terrace, Tawa, Wellington, New Zealand Strahl, Dr Stuart, Wildlife Conservation International, 185th Street and Southern Boul¬ evard, Bronx, NY 10460, U.S.A. Straw, Mr Phil, New South Wales National Parks & Wildlife Service, 43 Bridge Street, Hurstville, Sydney, NSW 2220, Australia Stuart-Jones, Mrs Elisabeth, 36 Tuawera Terrace, Sumner, Christchurch, New Zea¬ land Styche, Mr Andrew, 24 Salek Street, Kilbirnie, Wellington, New Zealand Suhonen, Mr Jukka I., University of Jyvaskyla, Konnevesi Research Station, SF-44300 Konnevesi, Finland Sullivan, Dr Kim, Utah State University, Biology Department, Logan, Utah 84322- 5305, U.S.A. Summers-Smith, Dr Denis, Merlewood, The Avenue, Guisborough, TS14 8EE, United Kingdom Summers-Smith, Mrs Margaret, Merlewood, The Avenue, Guisborough, TS14 8EE, United Kingdom Suter, Dr Werner, Schweizerische Vogelwarte, CH-6204 Sempach, Switzerland Suter, Mrs Dorothy, Schweizerische Vogelwarte, C/- Dr Werner Suter, CPI-6204 Sempach, Switzerland Sutherland, Dr Bill, University of East Anglia, School of Biological Sciences, Norwich, NR4 7TJ, United Kingdom Suwelo, Dr Ismu, Forestry Training & Education Centre, Indonesian Swiftlet Lovers, Jalan Indrapura 4 Candi Baru, Semarang, 50132, Indonesia Sveinbjarnardottir, Ms Gudrun, University of Iceland, C /- Prof. Gardarsson, Inst, of Biology, Grensasvegur 12, IS- 1 08 Reykjavik, Iceland Svensson, Dr Soren, University of Lund, Department of Ecology, Ecology Building, S- 22362 Lund, Sweden Swennen, Dr Kees, Netherlands Institute for Sea Research, Box 59, Den Burg, Texel, 1790 AB, Netherlands Swift, Ms Nina, Department of Conservation New Zealand, P.O. Box 10-420, Welling¬ ton, New Zealand Szijj, Prof Laszlo J., California State Polytechnic University, 3801 West Temple Boul¬ evard, Pomona, CA 91768, U.S.A. Taborsky, Dr Michael, Konrad Lorenz Institut fur Vergleichende Verhaltensforschung, Savoyenstrasse 1A, A-1160 Wien, Austria Tallman, Dr Erika, Northern State University, P.O. Box 740, Aberdeen, 57401, U.S.A. Tallman, Dr Dan, Northern State University, P.O. Box 740, Aberdeen, 57401, U.S.A. Tarburton, Dr Michael, Keilor Adventist High School, Lot ITaylors Road, St Albans, Victoria 3021, Australia Tarr, Ms Cheryl, University of North Dakota, 807 Chestnut Street, Grand Forks, ND 58201, U.S.A. Tasker, Mr Mark L., Nature Conservancy Council, 17 Rubislaw Terrace, Aberdeen, AB1 1XE, United Kingdom Taylor, Mrs Mona, “Strathearn”, Wairuna R.D., Clinton, Sth Otago, New Zealand Taylor, Mr Rowley, DSIR - Land Resources, Cnr Milton & Halifax Streets, Private Bag, Nelson, New Zealand Taylor, Mr Graeme, Department of Conservation New Zealand, Threatened Species Unit, P.O. Box 10-420, Wellington, New Zealand Teixeira, Prof Dante Martins, Museu Nacional, Quinta da Boa Vista, Sao Cristovao, Rio de Janeiro (RJ), 20942, Brazil ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 53 Temple, Prof. Stanley A., University of Wisconsin, 1630 Linden Drive # 209, Madison, Wl 53705, U.S.A. Temple, Ms Anita J., The Nature Conservancy, Dunlap Hollow Road, Mazomanie, Wl 53560, U.S.A. Templeton, Mr Ronald, 30 Carcoola Road, St Ives, Sydney, NSW 2075, Australia Tennyson, Mr Alan, 222A Karori Road, Karori, Wellington, New Zealand Thaler, Prof. Ellen, Alpenzoo Innsbruck-Tirol, A-6020 Innsbruck, Weiherburg, Austria Thiede, Dr Walther, Ornithologische Mitteilungen, An der Ronne 184, Koeln, D5000, Germany Thiede, Dr Ulrike, University of Duesseldorf, An der Ronne 184, Koeln, D5000, Ger¬ many Thiollay, Dr Jean Marc, C.N.R.S., Laboratoire d’Ecologie, E.N.S., 46 Rue d’Ulm, Paris, Cedex, 75230, France Thomas, Dr David H., University of Wales at Cardiff, Cardiff, CF1 3TL, Wales, United Kingdom Thomas, Mr Tony, 20 Edward Street, Prebbleton, Canterbury, New Zealand Thompson, Mr Jeremy J., University of Queensland, Department of Zoology, St Lucia, Brisbane 4067, Australia Thompson, Dr Charles F., Illinois State University, Department of Biological Sciences, Normal, Illinois 61761, U.S.A. Thompson, Mrs Karen M., C/- Dr Charles F. Thompson, Illinois State University, De¬ partment of Biological Sciences, Normal, Illinois 61761, U.S.A. Thompson, Mr Chris, The City University of New York, Biology Department, City Col¬ lege, Convent Avenue at 138th Street, New York 10031, U.S.A. Thompson, Rosemary, 213 llam Road, Fendalton, Christchurch, New Zealand Thoms, Mr Michael, Unit 3, 13 Forbes Street, Beckenham, Christchurch, New Zealand Thomson, Mr Philip J., Department of Conservation, Private Bag 3072, Hamilton, New Zealand Thomson, Mr Bob, Chairman - Christchurch Organising Committee, 34 Whitewash Head Road, Sumner, Christchurch, New Zealand Thoresen, Dr Asa C., Andrews University, 2229 Wilderness Drive, Berrien Springs, Michigan 49103, U.S.A. Thornhill, Prof. Randy, University of New Mexico, Department of Biology, Albuquerque, New Mexico 87131, U.S.A. Tidemann, Dr Sonia, Conservation Commission of the Northern Territory, P.O. Box 496, Palmerston, Darwin, 0831, Northern Territory, Australia Tiebout III, Dr Harry M, University of Florida, Department of Zoology, Gainesville, Florida 32611, U.S.A. Tisdall, Mrs Christine, 15 Knox Street, Dunedin, New Zealand Todal, Ms Malfrid, Norwegian Institute for Nature Research, Tungasletta 2, N-7004 Trondheim, Norway Todt, Prof Dietmar, Freie Universitat, Institut f. Verhaltensbiolgie, Haderslebener Strasse 9, D-1000 Berlin 41, Germany Tomialojc, Prof. Ludwik, Wroclaw University, Museum of Natural History, Sienkiewicza 21, 50-335 Wroclaw, Poland Torok, Mr Janos, EOTVOS University, Dept Systematic Zoology & Ecology, H - 1088 Budapest, Puskin U 3, Hungary Torr, Mr Nick, Department of Conservation New Zealand, P.O. Box 29, Te Anau, New Zealand Trapnell, Ms Helge, C /- Dr David B. Wingate, Dept. Agriculture, Fisheries & Parks, Box HM 834, Hamilton HM CX, Bermuda 54 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Traylor, Mr Melvin A. Jr., Field Museum of Natural History, Roosevelt Road & Lake Shore Drive, Chicago, IL 60605, U.S.A. Traylor, Mrs Marjorie S., Field Museum of Natural History, Roosevelt Road & Lake Shore Drive, Chicago, IL 60605, U.S.A. Triggs, Dr Sue, Department of Conservation New Zealand, P.O. Box 10-420, Welling¬ ton, New Zealand Trouvilliez, Dr Jacques, Office National de la Chasse et de la Faune Sauvage, 5 Rue de St Thibault, Auffargis, 78610, France Tsuji, Mr Atsuo, Meijo University, Faculty of Science & Technology, Shiogamaguchi, Tempaku-ku, 468 Nagoya, Japan Tunnicliffe, Mr Geoffrey, Canterbury Museum, Rolleston Avenue, Christchurch 1, New Zealand Turbott, Mr Graham, 23 Cathedral Place, Parnell, Auckland, New Zealand Turner, Dr Angela, Glasgow University, Department of Zoology, Glasgow, G12 8QQ, United Kingdom Turner, Mr John V.N., The Wildfowl & Wetlands Trust, Martin Mere, Burscough, Lan¬ cashire, L40 OTA, United Kingdom Tyler, Dr Stephanie J, Royal Society for the Protection of Birds, RSPB Wales Office, Bryn Aderyn, Newtown, Powys, SY16 2AB, United Kingdom ten Cate, Dr Carel, University of Groningen, Zoological Laboratory, Postbus 14, Haren 9750 AA, Netherlands Udvardy, Dr Miklos, California State University Sacramento, 6000 Jay Street, Sacramento, California 95819, U.S.A. Udvardy, Mrs Maud E., C/- Dr. Miklos Udvardy, California State University Sacramento, 6000 Jay Street, Sacramento, CA 95819, U.S.A. Ueda, Dr Keisuke, Rikkyo University, Laboratory of Biology, Ikebukuro, Tokyo 171, Japan Ulfstrand, Mrs Astrid, C /- Prof. S.UIfstrand, Zoology Department, Uppsala University, S-751 22 Uppsala, Sweden Ulfstrand, Prof. Staffan, Uppsala University, Department of Zoology, S-751 22, Uppsala, Sweden Urban, Prof. Emil K., Augusta College, Department of Biology, Augusta, Georgia, 30910, U.S.A. Usher, Mrs Sue, Royal Society of New Zealand, 4 Halswell Street, Thorndon, Welling¬ ton, New Zealand Vader, Dr Wim, Tromso Museum, University of Tromso, Tromso, N-9000, Norway Van Bocxstaele, Mr Roland, Royal Zoological Society of Antwerp, Koningin Astridplein 26, Antwerp, B2018, Belgium Van Den Elzen, Dr Renate, Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 150-164, Bonn D-5300, Germany Van Der Lande, Dr Virginia, University of Nottingham, Department of Zoology, Univer¬ sity Park, Nottingham NG7 2RD, United Kingdom Van Esbroeck, Mr Jacques, Societe d’etudes Ornitholgiques “AVES”, Rue de la Cambre 16/2, Brussels, 1200, Belgium Van Horne, Dr Beatrice, Colorado State University, Department of Biology, Fort Collins, CO 80523, U.S.A. Van Meter, Dr Robert G., 1673 East Imperial Highway, Los Angeles, California 90059, U.S.A. Van Noordwijk, Dr Arie J., Zoologisches Institut, Rheinsprung 9, CH 4051 Basel, Switzerland ACTA XX CONGRESSUS INTERN ATIONALIS ORNITHOLOGICI 55 Varty, Dr Nigel, I.C.B.P., 32 Cambridge Road, Girton, Cambridge, CB3 OPJ, United Kingdom Vaz-Ferreira, Prof. Raul, Facultad de Plumanidades y Ciencias, Department of Ver¬ tebrate Zoology, Isabelino Bosch 2482, Montevideo, Uruguay Veen, Dr Jan, Research Institute for Nature Management, Postbus 9201, 6800 HB Arnhem, Netherlands Veit, Dr Richard, University of Washington, Department of Zoology, Seattle, WA 98195, U.S. A. Veit, Mrs Barbara, St. Margaret’s School, 128 Rockview Drive, Irvine, CA 92715, U.S. A. Veitch, Mr Dick, Department of Conservation New Zealand, Private Bag 8, Newton, Auckland, New Zealand Velarde, Dr Enriqueta, National University of Mexico, Instituto de Biologia, Apartado Postal 70-153, Mexico DF 04510, Mexico Veltman, Dr Clare, Massey University, Botany & Zoology Department, Palmerston North, New Zealand Vermeer, Dr Kees, Canadian Wildlife Service, C/O Institute of Ocean Sciences, P.O. Box 6000, Sidney, BC V8L 4B2, Canada Verstrael, Dr Theo J., Netherlands Central Bureau of Statistics, P.O. Box 959, 2270 AZ Voorburg, Netherlands Vickery, Dr Juliet, University of East Anglia, School of Biological Sciences, Norwich, County Norfolk, NR2 7TJ, United Kingdom Vickery, Mr Peter D., University of Maine, P.O. Box 127, Richmond, Maine 04357, U.S. A. Vielliard, Prof. Jacques M.E., UNICAMP, Department of Zoology, Box 6109, Barao Geraldo, Campinas 13081, Brazil Vigorita, Dr Vittorio, Wildlife Service, Agriculture & Forestry, IV Novembre 5, Milano 20124, Italy Violani, Dr Carlo G., Departimento di Biologia Animale, Universita, Piazza Botta 9, 27100 Pavia, Italy Virkkala, Dr Raimo, University of Helsinki, Department of Zoology, P Rautatiekatu 13, SF-00100 Helsinki, Finland Von Meyer, Mr Andreas, Cas. 711, Puerto Montt, Chile Vuilleumier, Prof Francois, American Museum of Natural History, Department of Or¬ nithology, Central Park West at 79th Street, New York, NY 10024-5192, U.S. A. van Balen, Mr Bas, ICBP, P.O. Box 47, Bogor, Indonesia Wada, Prof. Masaru, Tokyo Medical & Dental University, Department of General Edu¬ cation, Z-8-30 Kohnodai, Ichikawa-shi, 272 Chiba, Japan Wada, Mrs Kaori, Tokyo Medical & Dental University, Department of General Educa¬ tion, Z-8-30 Kohnodai, Ichikawa-shi, 272 Chiba, Japan Waghorn, Dr Elspeth J., Department of Conservation New Zealand, 58 Tory Street, Wellington, New Zealand Wagner, Mr Steven J., Clemson University, Department of Biological Sciences, Clemson, SC 29634-1903, U.S. A. Waide, Dr Robert B., Center for Energy & Environment Research, GPO Box 3682, 00936 San Juan, Puerto Rico Wakabayashi, Prof. Shuichi, Nikon University, Biology Department, School of Den¬ tistry, 1-8-13 Kanda Surugadai, Tokyo, 101, Japan Waller, Mr Malcolm, O.S.N.Z., South Head, R.D.1 Helensville, New Zealand Wallschlager, Dr Dieter, Humboldt-Universitat, Department of Animal Behaviour, Invalidenstrasse 43, Berlin, DDR 1040, Germany 56 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Walter, Mr John, 9 Hill Street, Pittsworth, Queensland, 4356, Australia Walter, Mrs Ruth, 9 Hill Street, Pittsworth, Queensland, 4356, Australia Walters, Dr Jeffrey R., North Carolina State University, Department of Zoology, Raleigh, NC 27695-7617, U.S.A. Warburton, Dr Bruce, Forest Research Institute, P.O. Box 31-011, Christchurch, New Zealand Ward, Mr Dave, DSIR - Sirtrack, Private Bag, Havelock North, New Zealand Warham, Dr John, University of Canterbury, 14 Konini Street, Christchurch 4, New Zealand Warncke, Dr Gunther, Institut fur Neurophysiologie, Universitat zu Koln, Robert-Koch- StraBe 39, 5000 Koln 41, Germany Warncke, Mrs Maria, Cl- Institut fur Neurophysiologie, Universitat zu Koln, Robert- Koch-Strasse 39, 5000 Koln 41, Germany Wartmann, Dr Beat A., Sonnenberg 33, Oberengstringen, CH-8102, Switzerland Watt, Dr Doris J., St. Mary’s College, Department of Biology, Notre Dame, Indiana 46556, U.S.A. Wattel, Dr Jan, Zoologisch Museum Amsterdam, Postbus 4766, 1009 AT Amsterdam, Netherlands Way, Mr Ritchie, 87 New Windsor Road, Avondale, Auckland 7, New Zealand Way, Mrs Rosemary, 87 New Windsor Road, Avondale, Auckland 7, New Zealand Weatherhead, Dr Patrick, Carleton University, Department of Biology, Ottawa, On¬ tario, K1S 5B6, Canada Weathers, Dr Wesley W., University of California, Department of Avian Sciences, Davis, CA 95616, U.S.A. Weathers, Mrs Deb, University of California, Department of Avian Sciences, Davis, CA 95616, U.S.A. Webb, Prof. Randy, University of Wyoming, Department of Zoology & Physiology, P.O. Box 3166, Laramie, Wyoming 82071, U.S.A. Webster, Dr J. Dan, Hanover College, P.O. Box 292, Hanover, IN 47243, U.S.A. Webster, Mrs Juanita R., P.O. Box 292, Hanover, IN 47243, U.S.A. Webster, Mr Rick, 227 River Street, Deniliquin 2710, Australia Webster, Dr Marcus D., St. John’s University, Collegeville, MN 56321, U.S.A. Wechsler, Mr Doug, VIREO/Visual Resources for Ornithology, The Academy of Natu- ral Sciences of Philadelphia, 19th & The Parkway, Philadelphia 19103, U.S.A. Weimerskirch, Dr Henri, Centre National de la Recherche Scientifique, CEBAS, Villiers-en-Bois, Beauvior-Sur-Niort, F-79360, France Weinberger, Dr Marc, 10 Wilputte Place, New Rochelle, New York 10804, U.S.A. Weise, Prof Charles M., University of Wisconsin-Milwaukee, Department of Biologi¬ cal Sciences, Milwaukee, Wl 53201, U.S.A. Weislogel, Ms Winifred S., Apartment 803, 1016 South Wayne Street, Arlington, Vir¬ ginia 22204, U.S.A. Weller, Prof. Milton W., Texas A & M University, Department of Wildlife & Fisheries, 210 Nagle Hall, College Station, TX 77843, U.S.A. Weller, Mrs Doris L., 2709 Red Hill Drive, College Station, TX 77845, U.S.A. Wendel, Mrs Linda, Keilufell 16, 111 Reykjavik, Iceland Wenzel, Prof. Bernice M., University of California, Department of Physiology, School of Medicine, Los Angeles, CA 90024-1751, U.S.A. Werner, Miss Sabine, Rettenpacherstr. 5, Salzburg 5020, Austria West, Ms Jill, 59 Strickland Street, Christchurch 2, New Zealand Westerbeke, Mr Paul, 7 Ruskin Road, Newlands, Wellington, New Zealand ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 57 Westerskov, Dr Kaj E., University of Otago, 7 Wells Street, Brighton, Otago, New Zealand Westin, Dr Jan, University of Goteborg, Faculty of Science, Fysikgarden 1 , Goteborg, Sweden Wettenhall, Dr Norman, Royal Australasian Ornithologist’s Union, 14 Lascelles Av¬ enue, Toorak, Melbourne 3142, Australia Wheeldon, Mr Rob, Tiercel Cottage, RD 2 Eketahuna, Wairarapa, New Zealand Wheeley, Mrs Anne R., College of William and Mary, Department of Biology, Williamsburg, Virginia, U.S.A. White, Prof Clayton “M”, Brigham Young University, Provo, 84602, U.S.A. Whitehead, Mr Michael, Monash University, Wellington Road, Clayton, Victoria 3168, Australia Whiteley, Ms Pam, CSIRO Australian Animal Health Lab., P.O. Bag 24, Geelong, Vic¬ toria 3222, Australia Widen, Dr Per, Grimso Wildlife Research Station, S-77031 Riddarhyttan, Sweden Wienecke, Miss Barbara, Murdoch University, South Street, Murdoch, Perth, WA 6150, Australia Wieneke, Ms Jo, 22 Bishop Street, Belgian Gardens, Queensland, 4810, Australia Wiens, Prof. John A., Colorado State University, Department of Biology, Fort Collins, CO 80523, U.S.A. Wilds, Ms Claudia P., American Birding Association, 3331 North Street North West, Washington, DC 20007, U.S.A. Wilkes, Mr Ross, 33 Elector Street, Seatoun, Wellington, New Zealand Wilkinson, Dr Roger, North of England Zoological Society, Chester Zoo, Upton-By- Chester, Chester, CH2 1LH, United Kingdom Wilkinson, Mrs Lynn, North of England Zoological Society, Chester Zoo, Upton-By- Chester, Chester, CH2 1LH, United Kingdom Willard, Dr David, Field Museum of Natural History, Roosevelt Road at Lakeshore Drive, Chicago, IL 60605, U.S.A. Williams, Dr Tony D., British Antarctic Survey, High Cross, Madingley Road, Cam¬ bridge, CB3 OET, United Kingdom Williams, Dr Joseph B., University of Cape Town, Percy Fitzpatrick Institute, Rondebosch 7700, South Africa Williams, Dr Murray, Department of Conservation New Zealand, P.O. Box 10-420, Wellington, New Zealand Willis, Prof. Edwin O., University Estadual Paulista Department of Zoology, CP 178, 13500-Rio Claro, Sao Paulo, Brazil Willson, Dr Mary F., Forestry Sciences Laboratory, P.O. Box 20909, Juneau, AK 99802, U.S.A. Willson, Dr Rory, Institut fur Meereskunde, Duesternbrooker Weg 20, 2300 Kiel 1, Germany Wilson, Mr Ian D., Puketi Road, R.D.1, Okaihau, Northland, New Zealand Wilson, Ms Kerry-Jayne, Lincoln University, Entomology Department, Canterbury, New Zealand Wilson, Dr Peter, DSIR Land Resources, Private Bag, Nelson, New Zealand Wiltschko, Dr Roswitha, FB Biologie der Universitat, Siesmayerstrasse 70, D6000 Frankfurt, Germany Wiltschko, Prof. Wolfgang, FB Biologie der Universitat, Zoologie, Siesmayerstrasse 70, D6000 Frankfurt, Germany Winchester, Mrs Yvonne, 2/21 Wells Road, Bucklands Beach, Auckland, New Zealand 58 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Winding, Dr Norbert, Salzburg University, Institute of Zoology, Hellbrunnerstrasse 34, Salzburg, A-5020, Austria Wingate, Dr David B., Department of Agriculture, Fisheries & Parks, Box HM 834, Hamilton, HM CX, Bermuda Wingfield, Prof John C., University of Washington, Department of Zoology NJ-15, Seattle, WA 98195, U.S.A. Winkelman, Dr Johanna E., Research Institute for Nature Management, C /- Dr. E. Osieck, Driebergseweg 16-C, Zeist, 3708 JB, Netherlands Winkler, Dr Hans, Institut fur Vergleichende Verhaltensforschung, Savoyenstrasse 1A, Wien, A-1 1 60, Austria Winkler, Dr Raffael, Natural History Museum Basel, Augustinergasse 2, Basel, 4001, Switzerland Winkler, Ms Amanda, Reiterstrasse 19, Basel, 4054, Switzerland Witt, Dr Klaus, Hortensienstr. 25, D-1000 Berlin 45, Germany Witt, Dr Jutta, Hortensienstr. 25, D-1000 Berlin, 45, Germany Wobeser, Dr Gary A., University of Saskatchewan, Western College of Veterinary Medicine, Saskatoon, Saskatchewan, S7H 3C5, Canada Wobeser, Mrs Amy G., 27 Simpson Crescent, Saskatoon, Saskatchewan, S7H 3C5, Canada Woehler, Dr Eric J., University of California, Dept of Ecology & Evolutionary Biology, Irvine, California, 92717, U.S.A. Wong, Miss Cathy, 76 Picton Avenue, Riccarton, Christchurch, New Zealand Woodbury, Dr C. Jeffery, American Museum of Natural History, Department of Orni¬ thology, 79th Street and Central Park West, New York, NY 10024, U.S.A. Woodward, Mr Gavin, Victoria University, School of Architecture, P.O. Box 600, Wel¬ lington, New Zealand Woolfenden, Prof. Glen E., University of South Florida, Biology Department, Tampa, FL 33620, U.S.A. Woolfenden, Mrs Jan, Archbold Biological Station, Lake Placid, FL 33852, U.S.A. Wooller, Dr Ron, Murdoch University, Biological Science Department, Murdoch, West¬ ern Australia 6150, Australia Worthy, Mr Trevor, 43 The Ridgeway, Nelson, New Zealand Woudberg, Mrs Lesley, 335 Wigram Road, Halswell, Christchurch, New Zealand Wright, Dr Jonathan, University of Oxford, AFRC Unit, Department of Zoology, South Parks Road, Oxford OX1 3PS, United Kingdom Wright, Dr Patricia, Duke University, Durham, NC, U.S.A. Wright, Mr Alan, “Albatross Retreat”, 676 Portobello Road, Broad Bay, Otago Penin¬ sula 9004, New Zealand Wright, Neil, 92c Riccarton Road, Riccarton, Christchurch, New Zealand Wuerdinger, Prof Irene, Universitat Hildesheim, Marienburger Platz 22, Hildesheim, D 32, Germany Wynne, Mr Graham, The Royal Society for the Protection of Birds, The Lodge, Sandy, Bedfordshire, SG19 2DL, United Kingdom Yamagishi, Prof Satoshi, Osaka City University, 3-3-138 Sugimoto, Sumiyoshi-ku, Osaka 558, Japan Yamaguchi, Mr Nobuyuki, Waseda University, Dept of Biology, School of Education, Nishiwaseda 1-6-1, Shinjuku-ku, Tokyo 169, Japan Yearbury, Mr Peter, 72 Conway Street, Somerfield, Christchurch 2, New Zealand Yearbury, Mrs Joce, 72 Conway Street, Somerfield, Christchurch 2, New Zealand Yom-Tov, Prof Yoram, Tel Aviv University, Department of Zoology, Tel Aviv 69978, Israel ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 59 Yuan, Mr Guoying, Xinjiang Environmental Institute of Protection, Beijing Road 38, Urumqi, China Zack, Dr Steve, Yale University, P.O. Box 5 OML, New Haven, CT 06511, U.S.A. Zahavi, Dr Amotz, Tel Aviv University, Institute Nature Conservation Research, Tel Aviv, Israel Zahavi, Mrs Avishag, C /- Amotz Zahavi, Tel Aviv University, Institute Nature Conser¬ vation Research, Tel Aviv, Israel Zann, Dr Richard, La Trobe University, Department of Zoology, Bundoora 3083, Aus¬ tralia Zink, Dr Robert M., Louisiana State University, 119 Foster Hall, Baton Rouge, Loui¬ siana 70803, U.S.A. Zusi, Dr Richard L., Smithsonian Institution, National Museum of Natural History, Washington, DC 20560, U.S.A. Zweers, Dr Gart, University of Leiden, Zoological Laboratory, Box Number 9516, 2300 RA Leiden, Netherlands 60 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI . . / ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 61 PROFESSOR HELMUT SICK 1910-1991 HONORARY VICE PRESIDENT 20TH INTERNATIONAL ORNITHOLOGICAL CONGRESS It was my pleasure and privilege to appoint my old friend, Helmut Sick, as one of the Honorary Vice-Presidents of the 20th International Ornithological Congress. On Au¬ gust 24, 1990, he wrote to tell me that he would be unable to participate in the Con¬ gress because he was seriously ill with malaria; he wrote: “You will be astonished - Malaria! I got my first malaria in 1941.” He died on March 5, 1991. Helmut Sick was born in Leipzig, Germany, in 1910 and studied science at German universities. He obtained the Ph.D. degree in ornithology in 1937 at the University of Berlin under the direction of Professor Erwin Stresemann. He then joined the staff of the Zoological Museum in Berlin and went to Brazil on an ornithological expedition just before the outbreak of World War II. He was not permitted to leave during the war and elected to remain in Brazil after the war. In 1946 he joined the Central Brazilian Foun¬ dation which was formed to explore the unknown interior of Central Brazil. Many of his publications on the birds of Brazil record results of this work over a period of seven years. He described his experiences in a popular book entitled “Tukani” (published in 1957 in German; 1959 in English) after his charming pet toucan. For the rest of his life he studied Brazilian birds and was a major influence in science and conservation in his adopted country. Helmut was the Brazilian delegate to the 1 1th IOC (1954) and was elected to the International Ornithological Committee at the 12th IOC in 1958 as the member from Brazil. I met him at the 1954 Congress in Basel, visited him in Rio de Janeiro in 1956, and we corresponded over the years. He was a productive and respected ornithologist, an accomplished artist, and a kind and valued friend. Charles Sibley 62 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI CHARLES ALEXANDER FLEMING K.B.E., D.SC., F.R.S., F.R.S.N.Z. 1916-1987 Sir Charles Fleming was an eminent and valued member of the New Zealand Organ¬ ising Committee for the Congress until his death in September 1987. The following tribute by Dr Peter C. Bull, Minutes Secretary to the Committee, is reproduced from the minutes of the 6th meeting of the Committee held in Wellington on 30 October 1987. New Zealand lost one of its most distinguished scientists with the sudden death of Sir Charles Fleming at his home in Wellington on 11 September 1987 at the age of 71. Members of the New Zealand Organising Committee have particular reason to mourn his passing. Not only was he a personal friend of many of us, but he was also a highly respected and influential member of the Committee. As a distinguished scientist on the world scene, his presence and influence at the 19th International Ornithological Congress in Canada last year greatly helped New Zealand’s successful bid to host the 20th Congress. Sir Charles was a former president of both the Royal Society of New Zealand and of the Ornithological Society (the two organisations sponsoring the 20th Congress) and his experience and sound judgement have been of great value to the Organising Committee. He will be sadly missed. Sir Charles was born in Auckland and educated at Kings College and Auckland Uni¬ versity where, after majoring in both geology and zoology, he gained his masterate with a thesis on prions (whalebirds). He joined the New Zealand Geological Survey in 1940 as an assistant geologist and remained in that organisation for the rest of his working life except for a period of war service as coastwatcher at the Auckland Is¬ lands. He returned to the Survey after the war and eventually became Chief Palae¬ ontologist, a post that conveniently linked his geological and zoological interests. Declining further promotion, he concentrated his energies on research, the affairs of the Royal Society, and his varied cultural interests (music, art, languages and history). His many outstanding research contributions in geology were matched by others in zoology - notably in ornithology, but also in biogeography, conservation, and the tax¬ onomy and songs of cicadas. At the time of his death he had some 500 publications to his credit, including books, major research papers and numerous shorter articles. Apart from research, he took a very active part in the affairs of the Royal Society of New Zealand and was its president from 1962 to 1966. He also served a term (1968- 69) as president of the Australian and New Zealand Association for the Advancement of Science. The quality of his work earned him many honours, both at home and abroad. He was a Fellow of the Royal Society (one of very few New Zealanders to hold this distinc¬ tion), a Foreign Member of the American Philosophical Society (the only New Zealand resident to be so honoured), a Corresponding Fellow of the American Ornithologists’ Union, and a Fellow of the Royal Society of New Zealand, In 1977 he was made a Knight Commander of the Most Excellent Order of the British Empire (KBE) for serv¬ ices to science. His ornithological contributions began with his classic study of the birds of the Chatham Islands (1939), closely followed by other major studies on the prions (1941) ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 63 and New Zealand flycatchers (1950). In studying the life history of the Silvereye (1943) he was one of the first New Zealanders to use coloured leg bands (home-made in those days) to identify individual birds in the field. Another pioneering effort, under¬ taken jointly with the late Dr K. Wodzicki, involved a census of the Gannet population of New Zealand (1952) by counting occupied nests shown on aerial photographs and checking the results by ground visits to some of the gannetries. This resulted in the first full census of any New Zealand seabird. Sir Charles had less opportunity for sus¬ tained ornithological research in later life, but he nevertheless kept up with current advances by extensive reading, and he watched and photographed birds whenever the opportunity offered. He enjoyed such activities immensely and his observations and experiences were often the subject of articles and lectures. Thus, in addition to his main ornithological papers, he has published several shorter ones plus many ar¬ ticles and short notes which together record a life-time’s observations and thoughts on a wide range of ornithological topics. The Ornithological Society of New Zealand has particular reason to be grateful to Sir Charles. He was one of its founding fathers, a very active regional organiser in its early days, its president in 1948-49, and a faith¬ ful attender and contributor at meetings of the Wellington Branch over a period of some 40 years. As a palaeontologist, Sir Charles was naturally interested in the geographical affini¬ ties of elements of the New Zealand fauna and flora, and in their appearance and disappearance throughout geological time, and he published extensively on these topics. He was particularly interested in the array of ancient forms (including several kinds of birds) preserved from extinction by New Zealand’s long isolation, and he became very critical of the human mismanagement that had caused the recent extinc¬ tion of some of these species and threatened the survival of others. Indeed, the proper conservation of New Zealand’s native plants and animals, based on sound scientific principles, became a major concern for Sir Charles during the latter years of his life, and he fought long and hard to improve matters. Future generations will have much for which to thank him in this regard. Those of us who were privileged to know Charles as a friend will remember him for his ready wit and agile mind, for his infectious enthusiasm in his appreciation and understanding of beautiful things both natural and man-made, and for his helpfulness and unobtrusive generosity. No less characteristic was his determination to use his intellectual abilities, possessions and reputation in the service of others, particularly in the promotion of good science and the conservation of natural resources for the physical and aesthetic enjoyment of future generations. The contributions of Sir Charles to the scientific and cultural life of New Zealand were indeed exceptional, but they were not his alone. Lady Fleming, a naturalist and historian in her own right, made these achievements possible through her dedication as Charles’ adviser, sec¬ retary, field companion and competent manager of home and family. We extend our sympathy to Lady Fleming and her family, and also our grateful thanks for all she and Sir Charles have given us. Peter Bull US INTERNATIONALIS ORNITHOLOGICI ' ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 65 XX CONGRESSUS INTERNATIONALE ORNITHOLOGICUS REPORT OF THE SECRETARY-GENERAL The 20th International Ornithological Congress was held in Christchurch, New Zea¬ land, over 2-9 December 1990. Following an Opening Ceremony and Civic Reception in the Christchurch Town Hall on the evening of 2 December, the major activities of the Congress took place on the campus of the University of Canterbury. Patron His Royal Highness, The Prince Philip, Duke of Edinburgh, agreed to be Patron of the 20th International Ornithological Congress. A message from Prince Philip to the Con¬ gress was read at the Opening Ceremony and is reproduced later in this volume. Major sponsors The two major sponsors of the Congress were the Ornithological Society of NZ (OSNZ) and the Royal Society of NZ. History As part of an international enquiry regarding possible venues for the 20th Congress, the President of the 19th Congress, the late Prof. Dr. Klaus Immelmann, wrote to New Zealand in September 1983. The present Secretary-General referred the matter to the Council of the OSNZ at its meeting in October 1983. In 1984 the Council established an investigative Congress Committee, comprising Ben D. Bell (convener), Brian D. Bell, Peter C. Bull and Sir Charles Fleming (adviser), to study the issue. In July 1985 the Congress Committee circulated a questionnaire to 85 NZ ornithologists seeking their views on a NZ Congress. Respondents strongly supported New Zealand plac¬ ing an invitation for the 20th Congress in 1990, the favoured venue being Christchurch. In November 1985 the OSNZ Council authorised the committee to pre¬ pare a formal invitation to President Immelmann and the International Ornithological Committee (IOC). The OSNZ investigative committee was expanded to include B. Brown (ex officio President OSNZ), R.G. Powlesland, C.J.R. Robertson and H.A. Robertson. The NZ National Committee for the International Union of Biological Sci¬ ences (IUBS) and the Council of the Royal Society of NZ added their support for the invitation proposal. Both the Royal Society of NZ and the OSNZ agreed to act as major sponsors. A formal invitation prepared by the OSNZ Congress Committee was sent to President Immelmann and the Permanent Executive Committee in May 1986. A team of NZ ornithologists, together with their NZ agents, Conference Makers Lim¬ ited of Auckland, attended the 19th Congress in Ottawa in June 1986 and New Zea¬ land’s invitation to host the 20th Congress in 1990 was approved by a large major¬ ity of members at the IOC meeting on 27 June 1986. Local organisation The NZ Organising Committee (NZOC) for the 20th Congress was formed in August 1986, drawing most of its membership from the OSNZ Congress Committee, which preceeded it. The NZOC handled planning of all aspects of the Congress within New Zealand. Eight members served on an Executive Committee which held regular meet¬ ings in Wellington. In 1987 the NZ Ornithological Congress Trust Board, comprising all eight members of the Executive, was formed as a charitable trust to handle finan¬ cial and contractual aspects of Congress organisation. The NZOC held 13 meetings, 66 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI its Executive 65 meetings and the NZ Ornithological Congress Trust Board 6 meet¬ ings before the Congress. The NZOC delegated further organisational work to nine key subcommittees : Business Management, Christchurch Local Organising, Editorial, Excursions, Film Review, Grants Review, IOC-ICBP Liaison, Publications, and Pub¬ licity & Circulars. Air New Zealand was appointed official carrier to the Congress. Conference Makers Limited of Auckland were appointed professional conference organisers and success¬ fully handled a wide range of consultancy and organisational activities, including reg¬ istrations, bookings, accommodation and tours. “World of Birds” initiative In late 1990 four major international events involving birds and natural history were coordinated in New Zealand under the theme “The World of Birds - a Southern Per¬ spective”. These were the 20th International Ornithological Congress in Christchurch, the 20th World Conference of the International Council for Bird Preservation (ICBP) in Flamilton, the Pacific Festival of International Nature Films in Dunedin and the BirdPex ’90 Stamp Exhibition in Christchurch. The coordinated registration, vehicle, tour and accommodation service, for all the conferences, was organised to provide a centralised service for all participants and encouraged participation in more than one event. Membership Members of the Congress comprised 1303 persons from 63 countries as follows: Full members 877 Student members 131 Accompanying persons 135 Non-attending members 6 Staff volunteers 154 Opening and Closing Ceremonies of the Congress The Opening Ceremony of the Congress took place in the Christchurch Town Flail on Sunday evening, 2 December. Following a traditional Maori challenge, members were welcomed by the Secretary-General who introduced the speakers of the opening ad¬ dresses: Mr Tipene O’Regan, Chairman of the Ngai Tahu Trust Board; the President of the Ornithological Society of NZ, Brian D. Bell; the Mayor of Christchurch, Ms Vicki Buck; the President of the Royal Society of NZ, Professor John Dodd; the President of the 20th International Ornithological Congress, Professor Charles G. Sibley; the NZ Minister of Conservation, the Hon. Denis Marshall MP, who also delivered the mes¬ sage from the Congress Patron, H.R.H. The Prince Philip; and Sir Edmund Hillary, who formally opened the Congress. A Reception and Dinner hosted by the Mayor and City of Christchurch, was held in the Christchurch Town Hall after the Opening Ceremony. Over 1000 members and guests attended, which helped to establish a friendly and cordial atmosphere for the rest of the Congress week. The Closing Ceremony was held in the display pavilion at the University of Canter¬ bury on Saturday, 8 December, after the last oral presentation. The President, Pro¬ fessor Charles Sibley, announced Vienna, Austria, as the venue for the 21st Interna¬ tional Ornithological Congress in 1994, with Christopher M. Perrins (United Kingdom) ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 67 as President. Professor Sibley also announced the names of new officers of the 21st Congress and the names of 65 ornithologists, from 35 countries, elected to the Inter¬ national Ornithological Committee. The Secretary-General paid a tribute to the late Sir Charles Fleming (New Zealand) in recognition of his service to ornithology and the NZ Organising Committee. The President and Secretary-General expressed their warm¬ est thanks to all those who had contributed much to make the Congress such a memorable and successful event. The 20th International Ornithological Congress was then formally closed by the President. Exhibits A special exhibit on New Zealand birds was provided at the Congress venue by the Canterbury Museum, allowing visiting ornithologists to examine a range of skins of New Zealand birds. Commercial and institutional exhibits were sited in the Pavilion near the poster paper exhibits. As part of the “World of Birds” initiative the Christchurch Philatelic Society held an international stamp exhibition (BirdPex ’90) in the Student Union building at Canterbury University during the Congress. Further details of exhibits are given in the Supplement (Programme and Abstracts). Receptions and Conversaziones Oxford University Press, in conjunction with the Royal Australasian Ornithologists Union (RAOU), held a reception to launch “The Handbook of Australian, New Zealand and Antarctic Birds” on Wednesday evening. The Secretary-General introduced the Minister of Conservation, the Hon. Denis Marshall MP, who launched the first two volumes of this major new work on the avifauna of the region. Other receptions in¬ cluded those hosted by the President and Mrs Sibley at the Chateau Blanc (Wednes¬ day); by the RAOU to invest Brian D. Bell (NZ) as a Fellow of the RAOU (Friday); by the NZ Organising Committee for members of the PEC and assisted delegates (Fri¬ day). Conversaziones held during the Congress were for the Frank Chapman Memo¬ rial Fund (Tuesday) and the British Ornithologists Union (Wednesday). Local Activities Early-morning and day excursions were well attended during the Congress. On Thurs¬ day, 6 December, a Field Day and High Country Fair was held at the Mount Hutt Sta¬ tion Resort, near Methven. Most members went on bird excursions before arriving at Mount Hutt later in the day. The Methven Lions Club and residents of the Mount Hutt district delighted members with displays and the informal entertainment of a high- country fair. The day ended with a relaxing social and barbecue featuring some of the best of NZ country fare and country-style dancing. The final local activity of the Con¬ gress, the closing banquet, was held on Saturday evening, 8 December, and was attended by over 700 members and guests. Pre- and Post-Congress Tours Under the “World of Birds - a Southern Perspective” theme an integrated programme of pre-Congress and post-Congress group tours was offered to Congress members in and around New Zealand. Fifteen main tour routes were used to give members a chance to view a wide range of New Zealand birds, habitats and landscapes. These included visits to major national reserves, such as Little Barrier, Stephens and Cod¬ fish Islands. As alternatives to conducted group tours, self-drive tours, using cars or motor camper vans, were offered. 68 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI A major event was to have been a subantarctic island cruise on board the brand new M.V. Frontier Spirit from 9-22 December 1990. Unfortunately, due to damage sus¬ tained by the ship in a cyclone in Fiji during November, the Frontier Spirit cruise was cancelled. Fortunately passage on an alternative subantarctic cruise with the M.V. World Discoverer was arranged by 77 Frontier Spirit passengers. Scientific Programme The President hosted a meeting of members of the international Scientific Programme Committee (SPC) at Tiburon, Marin County, California, in November 1987. All mem¬ bers of the SPC were able to attend the meeting, chaired by Dr Peter Berthold (Ger¬ many). The SPC reviewed over 300 proposals for symposia and other presentations, then allocated the programme of plenary speakers, symposia topics and conveners. In consultation with Dr Berthold, review and allocation of contributed papers (oral and poster), round table discussions and special interest groups was carried out in New Zealand at a meeting of local SPC members convened by Dr Murray Williams (Vice- Chairman). The resulting scientific programme of the 20th International Ornithological Congress followed the broad format of its predecessors. Six major ornithological themes were adopted for the plenary and symposia sessions: Systematics, Evolution and Ornithogeography; General Biology; Behaviour; Ecology; Physiology; and Applied Ornithology. The scientific programme included over 800 contributions including 7 plenary papers; 240 papers in the 48 symposia; 276 oral contributed papers; 233 contributed poster papers; 31 round-table discussions; and 10 special interest groups. Further details are given in the Supplement (Programme and Abstracts). Film Programme A programme of recent ornithological and natural history films was screened through the Congress week in the Ngaio Marsh Theatre at the University of Canterbury. Un¬ der the “World of Birds - a Southern Perspective” theme, Congress organisers and Television New Zealand (TVNZ) cooperated in the selection of film material. A total of 39 films for the Congress programme was selected from bird films submitted to TVNZ for the Pacific Festival of International Nature Films in Dunedin. Further details are given in the Supplement (Programme and Abstracts). Report of the Permanent Executive Committee The Permanent Executive Committee held five meetings in Christchurch on 2, 3, 4, 5 and 7 December. All members, with the exception of B.K. Follett (UK), S. Haftorn (Norway) and C. Erard (France), were able to attend. The topics discussed were re¬ ported on at the meetings of the International Ornithological Committee on 4 and 7 December. On 8 December an informal meeting of the old (1986-1990) and new (1990-1994) Executive Committees was held. Report of the International Ornithological Committee The International Ornithological Committee (IOC) met on 4 and 7 December. Both meetings were chaired by the President, Professor Charles Sibley. Fifty members were present on 4 December and 49 members on 7 December. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 69 At the first meeting reports were received from the President, Secretary-General, Chairman of the Scientific Programme Committee and Permanent Secretary of the IOC. The President outlined issues discussed by the Executive Committee at its three meetings over 2-4 December. Dr John Dittami and Dr Hans Winkler gave a slide presentation on the proposed venue and facilities that Austria was offering for the 21st Congress in August 1994. On the recommendation of the Executive, the IOC voted to accept Vienna as the venue for the 21st Congress (no other formal invitations having been received). The Permanent Secretary introduced proposed amendments to the Statutes & By- Laws for consideration at the second IOC meeting. The Secretary-General tabled reports from the three IOC Standing Committees: Ornithological Nomenclature (Pro¬ fessor W.J. Bock); Coordination of Seabird Research (Dr David Nettleship); Applied Ornithology (Professor Dr V.D. Ilychiev and Dr R.W. Peterson). Dr Nettleship and Professor Bock addressed their Standing Committee reports. All three reports appear later in this volume. Professor Burt L. Monroe Jr. tabled a report from P. William Smith, Convener of Round Table Discussion No. 1 - Standardisation of English Bird Names. That discus¬ sion on 3 December had up to 60 participants and had voted to recommend to the IOC that it appoint Professor Monroe as convener of a working group on Standardi¬ sation of English Bird Names, with power to co-opt members to represent the broadest possible range of opinion and geography. The IOC voted to approve the establishment of such a working group which should report through the Permanent Secretary to the 21st Congress in 1994. It was felt such a group should not be limited to only a few people but should be widely representative. The IOC also voted to establish an inter¬ national group to standardise French bird names under the Chair of Dr Henri Ouellet. Dr Ouellet outlined progress of the working group to date. A Resolutions Committee was appointed to consider resolutions presented at the second IOC meeting. Committee members were W.J. Bock (USA), E. Bucher (Argen¬ tina) and J. King (USA). A Nominations Committee established by the Executive to receive nominations for members of the IOC was endorsed. The members were P. Berthold (Germany), H. Ouellet (Canada), C.M. Perrins (UK), L. Short (USA) and L. Tomialojc (Poland). At the second meeting Dr Hans Blokpoel, at the invitation of the President, reported on the first term of the Standing Committee on Applied Ornithology. The officers, Executive Committee and new members of the IOC were then elected. IOC approved the following executive positions for the next Congress: Karel H. Voous (Netherlands) as Honorary President, Christopher M. Perrins (UK) as President, Svein Haftorn (Norway) as Vice-President, and Walter J. Bock as Secretary of IOC. The elected membership of the Executive Committee was raised from eight to ten. Accordingly, three members were re-elected to the Executive Committee (EC): Peter Berthold (Germany), Enrique H. Bucher (Argentina) and E.N. Kurochkin (USSR); and seven new members were elected: Asha Chandola-Saklani (India), Hiroyuki Morioka (Japan), Cynthia Carey (USA), Henri Ouellet (Canada), Richard Liversidge (South 70 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Africa), Jacques Blondel (France) and Murray J. Williams (New Zealand). Other mem¬ bers of the EC ex officio are the President (Chairman), Vice-President, Secretary- General, Secretary, immediate Past President and immediate Past Secretary-General. Sixty two new members of the International Ornithological Committee were elected and three were re-elected: Argentina Manuel Nores Australia Jiro Kikkawa, Phillip Moors, Pat Rich Austria John Dittami, Ellen Thaler, Hans Winkler Belgium Andre Dhondt, Michel Louette Brazil Paulo Antas Canada Jon C. Barlow, Fred Cooke, Raymond McNeil, David N. Nettleship Chile Fabian Jaksic Colombia F. Gary Stiles Cuba Orlando H. Garrido-Calleja Czechoslovakia Karel Hudec (re-elected), Aladar Randiik, Andrev Stollman Eire Oscar Merne Finland Olavi Hilden Germany Roland Prinzinger, Klaus Schmidt-Koenig, Rosi Wiltschko Hungary Lajos Sasvari Iceland Arnthor Gardarsson (re-elected) India Manjit S. Dhindsa, Zafar Futehally, S.AIi Hussain Israel Yoram Yom-Tov Italy Fernando Spina, Carlo Violani Japan Juzo Fujimaki, Hiroyoshi Higuchi Kenya Nathan Gichuki Netherlands Arie L.Spaans, Gart Zweers New Zealand John L. Craig, Murray J. Williams Norway Tore Slagsvold P. Republic of China Zheng-jie Zhao, Guang-mei Zheng Poland Tomasz Wesolowski Republic of China Lucia Liu Severinghaus Rumania Laszlo Kalaber South Africa Timothy M. Crowe, Alan Kemp, W. Roy Siegfried (re-elected) Sweden Staffan Ulfstrand Switzerland Lucas Jenni Uganda Derek Pomeroy USSR Alexander V. Andreev, R. L. Potapov, Yuri V. Shibaev United Kingdom John P. Croxail, George M. Dunnet, Peter R. Evans USA John W. Fitzpatrick, Richard T. Holmes, Dominique G. Homberger, Ellen D. Ketterson, Burt L. Monroe Jr., John C. Wingfield Zimbabwe Peter J. Mundy Changes to the Statutes and By-Laws (recommended by the Executive) were adopted by a vote of IOC. General changes included dropping the term “Permanent” from ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 71 “Permanent Executive Committee” and “Permanent Secretary” and the removal of sexist language. Most changes were to better accommodate the new position of Sec¬ retary of the IOC, with formal division of the duties and responsibilities of the Secre¬ tary-General and the Secretary, (as detailed under the “Statutes and By-Laws” sec¬ tion of this volume). A further change in the By-Laws is the new requirement for ad¬ vance notice to be given by nations intending to offer invitations to host a future Con¬ gress. The Secretary-General informed the IOC of his recent discussions with the Director of the International Council for Bird Preservation (ICBP). The relationships between the IOC and ICBP were discussed, particularly in relation to the IOC Committee on Applied Ornithology and the scientific programmes of future Congresses. Resolutions The following Resolutions were adopted by the IOC at its second meeting on 7 De¬ cember after review by the Resolutions Committee and the Executive: Resolution 1. Submitted by the Standing Committee on Ornithological Nomenclature: The International Ornithological Committee at its meetings during the XXth Inter¬ national Ornithological Congress, Christchurch, New Zealand, 2-9 December 1990 congratulates and supports the International Commission on Zoological Nomencla¬ ture in its efforts to increase continuity of zoological nomenclature by the conser¬ vation and stabilisation of established names, and directs its Standing Committee on Ornithological Nomenclature to assist the International Commission on Zoologi¬ cal Nomenclature in these efforts. The International Ornithological Committee rec¬ ognises the pioneering actions of the Standing Committee on Ornithological No¬ menclature in developing a list of available family-group names of birds and urges this committee to undertake similar projects on genus-group and species-group names of birds. Resolution 2. Submitted by T. Crowe: The members of the International Ornithological Congress protest in the strongest possible terms the decision by the executive of the British Museum (Natural His¬ tory) [BM(NH)] to abolish the research-oriented posts in the museum’s Sub-depart¬ ment of Ornithology. The bird collection at the BM(NH) is one of the most compre¬ hensive resources of its kind internationally, and the BM(NH) scientists utilising this resource have, in the past, made major contributions to the areas of systematic bi¬ ology and biogeography. A scientifically revitalised Sub-department of Ornithology could do the same in the future. It is unrealistic to expect that scientists visiting from other institutions can fill the gap created by the abolition of these research posts, especially if they are required to pay “bench fees” to use the collection. Therefore, we request that the BM(NH) executive and Board of Trustees consider reinstating research posts and rescinding the implementation of bench fees (at least for bona fide scientists) in its Sub-department of Ornithology. Resolution 3. Submitted by the Israeli ornithologists: We, the amateur and professional ornithologists participating at the 1990 XXth In¬ ternational Ornithological Congress, are deeply concerned about the possible im- 72 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI pact on millions of migrating birds by the immense Voice of America relay station projected for construction in the Aravah Valley. We fear the following effects of this station on birds: 1. Collision with the huge 140-200 meter high towers, meteorological antennas and mazes of cables. 2. Overheating of approaching and roosting birds by high-intensity short wave ra¬ diation emitted by 16 500kw transmitters. 3. The as yet unknown effects of electromagnetic radiation on orientation, naviga¬ tion and behaviour of migrating birds. We strongly urge the governments of Israel and the United States of America to stop all activities connected with the implementation of this project until thorough, independent environmental impact assessments are studied and completed, and the relevant recommendations are adopted. Resolution 4. Submitted by the Resolutions Committee: A. WHEREAS the participants of the XXth International Ornithological Congress have enjoyed a stimulating programme, rich in professional and social variety, in a com¬ fortable and congenial setting, AND WHEREAS the efficient and hospitable ar¬ rangements and their execution bespeak thorough, thoughtful, and wise planning, THEREFORE BE IT RESOLVED that the International Ornithological Committee, on behalf of the participants of the XXth International Ornithological Congress, con¬ vened in Christchurch, New Zealand, extends its deepest thanks to the New Zea¬ land Organising Committee, and their associates and sponsors, for their part in making the XXth International Ornithological Congress a rich and memorable ex¬ perience. B. WHEREAS the participants of the XXth International Ornithological Congress have experienced a rewarding scientific programme of exceptional variety, depth and quality, AND WHEREAS the development and management of such an excellent programme, including the timely publication of its Proceedings can result only from experienced insights, skillful negotiations, and long effort, THEREFORE BE IT RESOLVED that the International Ornithological Committee, on behalf of the par¬ ticipants of the XXth International Ornithological Congress, extends its special thanks to the Scientific Programme Committee and its advisors for their role in ensuring the scientific success of the XXth International Ornithological Congress. C. WHEREAS the contributions made by all those who took part in the development and management of the XXth International Ornithological Congress were indispen¬ sable, but WHEREAS the guidance and coordination of their efforts required the skills and labour of special leaders, THEREFORE BE IT RESOLVED that the In¬ ternational Ornithological Committee, on behalf of the participants of the XXth In¬ ternational Ornithological Congress, extends its heartfelt thanks to Ben D. Bell, Murray Williams, Chris Robertson and Peter Berthold for fulfilling so well their spe¬ cial roles in making the XXth International Ornithological Congress a first class, world-class event. Funding of the Congress Unlike its immediate predecessors, the NZ organising committee was not supported by a major government grant to fund necessary Congress planning and organisation. However, various government organisations provided important services and assist¬ ance, most notably the Department of Conservation’s Science and Research Division OPENING CEREMONY - 1 . Maori challenge. 2. Of¬ ficial party arrives. 3. Vicki Buck (Mayor, Christchurch) & President. 4. Prof. Charles Sibley (President). 5. Tipene O’Regan (Chair. NgaiTahu Trust Board). 6. L to R. Sir Edmund & Lady Hillary; Hon. D. Marshall; Prof. J. Dodd (Pres. Royal Soc. of NZ); Brian Bell (Pres. Ornithological Soc. of NZ). 7. Hon. Denis Marshall (Minister of Conservation). 8. Christchurch Town Hall. SCIENTIFIC PROGRAMME 1. Lecture audience. 2. Cynthia Carey (Plenary). 3. Ian Atkinson (Plenary). 4. Enrique Bucher (L) and staff preparing for Plenary. 5. Murray Williams (Vice-Chair. Scientific Programme Committee). 6. Poster Paper Pavilion. BETWEEN THE LECTURES % k HL oft 2f® rj\ t*. ■ ^ 1 R;{J ..... f JSm ffi \ w t >3® i» A ™ * fe;I My ij w| / 4 jjfjg i lu 4% ■ y < dL aJBBJ • ' \ Vx ' i^K AM f J _ ■[.. <4 £1 SVlaBr"" EXCURSIONS 1. Lake Ellesmere. 2. Heathcote-Avon estuary; 3. Field excursion group. 4. High Country Cook Out. 5 & 6. Mount Hutt High Country Fair. 7. Whitewash Head excursion. 8. Wind storm or the Upper Rakaia River. PFvqft ** / 1 -M m i vn . v ;.aB |||||. .v 1 ' '' [pW» H ■ ■TVS** ; >V 'I 1 ' tjSfrilM ; A. 1 •‘Hi ‘^1*? '. **1 farorw/tujno STAFF - 1. Conforonco Managomont & Registration. 2. Rodorick Cossoo, Sue Koall, Paul Wostorboko of ‘Communications Centro’. 3. Airlines, Cruise & Tours Desk. 4. Congress Shop in tho Pavilion 5. Two of tho 148 Volunteer Staff. 6. Brian Boll (Chairman, Tours Organising Committee). 7. Tho Accommodation Dosk. V" < 1 1 r . 0 1 1 mmm II n i mm CLOSING CEREMONY - 1. President & Lesley Fairbairn (Conference Makers Limited). 2 & 3. Members of NZ and Local Organising Committees. 4 & 6. Congress members & Volunteer staff. 5. Bob Thomson (Chair. Local Organising Committee). 7. President & Christopher Robertson (Congress Business Manager). 8. L to R. N Kuroda (Hon. President); C Sibley (President); J Pinowski (Vice- Pros.); Ben Boll (Secrotary-General); W Bock (Pe¬ rmanent Secretary IOC). Inset. L to R. Gillian Boll, Gillian Robertson, oFrancos Sibloy, Christopher Perrins, Peter Borthold. A m'g *2 9 HP I Jm I : - ■ \r* -m, h * ^^^H ^ >. ^ Ht * - ** ^ ^ % ii ft , r h~ *& fflff A ^ T j TP * wit yj 4 j m *a r ? e \ r & | 1 UlIV W (VI 'B W .wfei !r.v NJpte; r ^r^Tljr in^HH V’ T 4t| !/ ■€%£ ,«>- j Jr j I Kfli • 1 / i mlltiigg*:- *&,s 1 I; * # yJBJ , v * t* J"Jl -y^ • V jflT. * a 'i 1 k ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 81 in Wellington and the Ecology Division of the Department of Scientific and Industrial Research (now DSIR Land Resources). The Congress was an official project of the NZ 1990 Commission who gave generous financial support. The Science and Re¬ search Distribution Committee of the NZ Lottery Board provided a major grant towards the publication of the Proceedings. Support for the Secretary-General and for editing the Acta was provided by Victoria University of Wellington. Other sources of revenue included proceeds from membership fees, rental of display spaces during the Con¬ gress, commissions from tours, sales and accommodation, grants or loans from the OSNZ, Royal Society of NZ, Air New Zealand and the Todd Foundation. A full list of sponsors is given in the Supplement p. 28. Acknowledgements The organisation of such a large Congress has been a major team effort and many persons, both overseas and within New Zealand, have contributed in numerous ways to its success. The New Zealand Organising Committee expresses its thanks to His Royal Highness, The Prince Philip, Duke of Edinburgh, for agreeing to act as the Patron of the Con¬ gress. I have had a close and constructive working relationship with the President through¬ out the planning of this Congress. I thank Professor Sibley for his constant support and wise counsel. I am also appreciative of the work carried out by other overseas colleagues on behalf of the Congress, particularly by Peter Berthold (Germany) and members of the Scientific Programme Committee; and by Walter Bock (USA), Perma¬ nent Secretary of the IOC. I am indebted to former President, the late Klaus Immelmann (Germany) and former Secretary-General, Henri Ouellet (Canada), for assistance during the early days of Congress bidding and organisation. To a substantial degree, the Congress owes its success to the unstinting support of numerous people in New Zealand, most of whom worked in a voluntary capacity and well beyond the call of duty. In particular, I thank our army of staff and volunteers - including members of the Ornithological Society of NZ, students and tour guides - who worked in many parts of the country on our behalf. The particular responsibility of successfully planning and organising the Congress rested on the shoulders of the various organising committees in New Zealand and I thank all of them for their hard work. The eight member Executive Committee in Wellington were the core management team for the Congress and I wish to record my thanks to them for their considerable individual and collective efforts. Brian Bell, convener of the Excursions Subcommit¬ tee, organised an excellent tours programme using his wide knowledge of New Zea¬ land and its birds; Peter Bull, Minutes Secretary, skillfully and meticulously recorded the minutes of our many meetings, providing the Congress archives with an exemplary record; Ralph Powlesland, convener of the Publications Subcommittee, developed and maintained our international mailing database, and played a key role in the pro¬ duction of the two circulars and Programme and Abstracts volume; Chris Robertson, convener of the Business Management Subcommittee, took on an enormous area of responsibility, handling a wide range of business activities, including financial man¬ agement, liaison with Conference Makers Limited, and the complex negotiations 82 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI regarding the Southern Ocean Cruise; Hugh Robertson, convener of the Circulars and Publicity Subcommittee, was responsible for Congress circulars and publicity, promot¬ ing the Congress in New Zealand and overseas; Murray Williams, Vice-Chairman of the Scientific Programme Committee, competently and amicably handled scientific programme matters, including liaison with SPC Chairman, Peter Berthold, with sym¬ posia conveners, and other contributors; and Sue Usher, Royal Society of NZ repre¬ sentative, offered wise counsel on a wide range of business and planning issues, and represented at executive level our senior sponsor, the Royal Society of NZ. I also extend my thanks to other members of the NZ Organising Committee: the late Sir Charles Fleming, for support and advice; Beth Brown, representing the OSNZ; Mick Clout, representing the 20th ICBP World Conference organising committee; Leslie Fairbairn, representing Conference Makers Limited; Audrey Hudson, our ac¬ counting advisor; Paul Sagar and Bob Thomson, successive conveners of the Christchurch Local Organising Subcommittee; and Brian Wybourne, representing the Council of the Royal Society of NZ and the academic community in Christchurch. Central planning by the Wellington Executive was complemented by much work in Auckland by our professional consultants, Conference Makers Limited, and in Christchurch by the Local Organising Subcommittee. The experience, sound advice and professionalism provided by Leslie Fairbairn and her colleagues at Conference Makers Limited was an essential ingredient in the smooth running and success of the Congress, and I warmly thank them for carrying out their role as professional confrence organisers so well. The organisational demands of such a large Congress were immense and theirs was a job well done. Another substantial organisational burden fell on the Christchurch Local Organising Subcommittee, particularly in the latter part of 1990. I thank all members of that committee for their hard and often demanding work. Theirs was a critical role and I express thanks to Patricia Bell, Chris Challies, Ian McLean, Colin O’Donnell, Paul Sagar, Eric Spurr, Jill West, Kerry-Jane Wilson for their contributions. I especially thank their convener - Bob Thomson - for his excellent leadership and support. I thank Ian Atkinson, Bill Lock, Ralph and Mary Powlesland for assisting the Editorial Subcommittee in the large proof-reading exercise necessary for the preparation of these Proceedings; and Sue Keall for taking on the task of Minutes Secretary to the NZ Executive Committee during 1991. My thanks also go to many others who assisted in Congress organisation in a vari¬ ety of ways. These include: Dr Richard Sadleir, Director of Science and Research, Department of Conservation; Professor John Wells, School of Biological Sciences, Victoria University of Wellington; Dr Malcolm Crawley, Director of the former Ecology Division, Department of Scientific and Industrial Research; the invited speakers at the Opening Ceremony - Mr Tipene O’Regan, Ms Vicki Buck, Professor John Dodd, the Hon. Denis Marshall and Sir Edmund Hillary. Finally, I thank our many sponsors for their support and confidence in our organisational and planning endeavours. Ben D. Bell Secretary-General ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 83 REPORTS OF THE STANDING COMMITTEES OF THE INTERNATIONAL ORNITHOLOGICAL COMMITTEE The reports of the three Standing Committees were submitted to the Secretary-General in 1990 prior to the 20th International Ornithological Congress. 84 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI REPORT OF THE STANDING COMMITTEE ON ORNITHOLOGICAL NOMENCLATURE At the 18th International Ornithological Congress (Moscow, 1982), the Standing Com¬ mittee on Ornithological Nomenclature (SCON) passed a resolution to begin work on a historical analysis of avian family-group names. This project was essential because of the changes in the rules of zoological nomenclature in the 1961 Code of Zoologi¬ cal Nomenclature which threatened continuity of several well-established avian family- group names. This project was well under way by the 19th Congress (Ottawa, 1986) at which time the SCON passed several resolutions which were presented to the In¬ ternational Ornithological Committee (IOC) and were passed at the first meeting of that body (the latter action was not mentioned in the report of the IOC in the Proceed¬ ings of the 19th Congress). This project on the history and status of avian family-group names is basically completed and will be submitted for publication as soon as correc¬ tions and suggestions are received from members of the SCON and other interested ornithologists and the final proof reading is completed. Copies of the penultimate draft of this monograph will be available to interested ornithologists at the 20th Congress at the display of materials on zoological nomenclature. It is hoped that the “History and Nomenclature of Avian Family-Group Names” will be in press during 1991 . After it is published an application will be submitted to the ICZN proposing that this publication be accepted as the base line for avian family-group names. Only the names included in this list with the authors and dates of publication as given will be available for zoological nomenclature. Names published prior to its publication but overlooked will be treated as unavailable for zoological nomenclature. An information table on zoological nomenclature with materials provided by the Sec¬ retariat of the International Commission on Zoological Nomenclature will be set up at the Congress. With the support of a number of ornithologists, Walter Bock was elected as a Commissioner of the International Commission on Zoological Nomenclature in 1988. This provides an important direct link between the Standing Committee on Or¬ nithological Nomenclature and the International Commission on Zoological Nomencla¬ ture (ICZN). The SCON functions as a specialist committee on avian nomenclature for the ICZN. The ICZN met during the ICSEB, Maryland, July 1990; Walter Bock, as a Commis¬ sioner, took part in this meeting and reported on some of the more significant deci¬ sions. Perhaps the most important result of this meeting was the decision by the ICZN to develop several methods to insure continuity of nomenclature. These methods will include the reintroduction of a stronger and automatic regulation to conserve estab¬ lished names versus forgotten senior synonyms (a return to a form of the 50 year rule), and the development of lists of known available names with dates of prec¬ edence, etc. The list of avian family-group names will be among the first, if not the first, of these lists for zoological names. Once such a list is established and approved by the ICZN, names on the list and those published in the future will be the only ones available for that group of animals. Any names not on the list cannot be used with¬ out application to and approval by the ICZN. These actions by the ICZN and the pro¬ posed amendments to the Code are most positive steps by the ICZN toward continuity of nomenclature. The system of using endings “i/ii” for male personal names will be rationalised. Gender agreement of generic and specific names will be simplified. The ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 85 stem for family group names formed from Greek words ending in “is” or “es” will be simplified; the proposed rule will be that such names would be treated simply for any family-group name in which the grammatically correct form is not in common usage, e.g., Drepanididae, type genus Drepanis. The ICNZ also considered briefly the question of extending the Code to names above the family-group, and still expressed the strong opinion that these names should be excluded from the Code. Problems still exist for many names in cladistic classifica¬ tions in which the authors are unclear as to whether the groups belong to the family- level or to the order-level. A new project was proposed for the SCON to begin work on a list of available generic- group names for birds. Such a list has been started by Murrey Bruce. Walter Bock will discuss with M. Bruce, methods for continuing this list under the auspices of the SCON. A report will be presented at the 1994 meeting of the committee. Burt Monroe moved and David Holyoak seconded a motion that the IOC adopt a reso¬ lution supporting the ICZN in its actions to insure the stability and continuity of zoo¬ logical nomenclature and to direct the SCON to assist the ICZN in these efforts. RESOLUTION: “The International Ornithological Committee at its meetings during the XXth International Ornithological Congress, Christchurch, New Zealand, 2-9 Decem¬ ber 1990 congratulates and supports the International Commission on Zoological Nomenclature in its efforts to increase continuity of zoological nomenclature by the conservation and stabilization of established names, and directs its Standing Commit¬ tee on Ornithological Nomenclature to assist the International Commission on Zoologi¬ cal Nomenclature in these efforts. The International Ornithological Committee recog¬ nizes the pioneering actions of the Standing Committee on Ornithological Nomencla¬ ture in developing a list of available family-group names of birds and urges this com¬ mittee to undertake similar projects on genus-group and species-group names of birds”. Burt Monroe moved and Richard Schodde seconded a motion giving a vote of thanks for the outstanding efforts of Walter Bock in compiling the list of avian family-group names and writing a history of these names and of zoological nomenclature during the period in which the new regulations covering family-group names were developed. At its meetings during the 20th International Ornithological Congress, the SCON con¬ sidered a number of existing and pending applications on avian names before the ICZN and discussed several new cases. a) A major application is one to be prepared by Walter Bock on avian family-group names based on the analysis of these names as mentioned above. Basically this application will request that the ICZN accept the list of the valid names and their synonyms for the currently recognized family-level taxa of Recent (non-fossil) birds and the decisions contained within it, including the date of publication and author of the valid names and their synonyms, names to be conditionally con¬ served and suppressed, and names which are not available for purposes of zoo¬ logical nomenclature or which are objectively invalid. This list will serve as the base for all future nomenclatural actions on avian family-group names for living birds. The date of this base line will be 1 January 1 991 . 86 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI b) Other applications cover nomenclatural problems in the family-group names Phororhacidae and Phororhacos , Hydrobatidae and Threskiornithidae, and in the genera Macronectes, Catharacta, Carpophaga and Serresius, Rallus, Cacatua, Geospiza, Creadion, Calyptorhynchus, Dendrocygna, Eurystomus, Calidres, and Corvus. c) The SCON urges the ICZN and its Secretariat to speed the process by which ap¬ plications are processed and published, and final action is taken by the ICZN. Fur¬ ther the SCON urges the ICZN and its Secretariat to increase its use of the SCON as a specialist advisory committee and to submit all applications dealing with birds to the SCON for its consideration. Further ornithologists are strongly urged to interact with the SCON to discuss possible nomenclatural matters and in the de¬ velopment of applications to submit to the ICZN. W. Bock was asked to raise these concerns at the meeting of the ICZN in September 1991. Members of the SCON appointed by the President Charles G. Sibley for the period 1986-1990 were: Walter J. Bock (Chairman), Pierre J. Devillers, Christian Erard, David Holyoak, Ernst Mayr, Gerlof F. Mees, Burt Monroe, Hiroyuki Morioka, Henri Ouellet, Richard Schodde, L.S. Stepanyan, Karel H. Voous, George Watson, David Wells and Hans Wolters. Respectfully submitted for the Standing Committee on Ornithological Nomenclature of the International Ornithological Committee. Walter J. Bock Chairman, SCON Members of the Committee on Ornithological Nomenclature for 1990-94 are: Walter J. Bock (USA), Chairman* Murray D. Bruce (Australia)* David Holyoak (United Kingdom)* Ernst Mayr (USA) Gerlof F. Mees (Netherlands)* Burt L. Monroe Jr (USA)* Hiroyuki Morioka (Japan)* Henri Ouellet (Canada)* D. Stefan Peters (Germany)* Richard Schodde (Australia)* L. S. Stepanyan (USSR) Karel H. Voous (Netherlands) David Wells (Malaysia) Hans E. Wolters (Germany) * Members attending the meetings of the SCON at the XX International Ornithologi¬ cal Congress. ACTA XX CONGRESSUS INTERN ATIONALIS ORNITHOLOGICI 87 REPORT OF THE STANDING COMMITTEE FOR THE COORDINATION OF SEABIRD RESEARCH The Standing Committee for the Coordination of Seabird Research (SCCSR) of the International Ornithological Committee (IOC) is an international group of marine bird biologists appointed by the President of the IOC. The SCCSR was established in 1966 with the principal aim of providing a mechanism for enhanced information exchange and integration of research on seabirds worldwide. That liaison function has tradition¬ ally been achieved through organised discussions and presentations at each con¬ gress, culminating with the presentation of a special interest symposium as part of the formal IOC Scientific Programme. One major outcome of the open meeting at the 19th Congress in Ottawa in June 1986 and discussions that followed, was the clear desire by all participants (committee members and others) to see more emphasis placed on action-orientated activities to be pursued between congresses. With that as a goal, the development of working groups was initiated to undertake specific review exercises (based on consensus of research needs) including the formulation of special symposia and workshops. Final decisions resulted in the establishment of four working groups - “Seabirds-at-sea”, “Bibliographies”, “Nomenclature”, and “Technological innovations” - and a task force to solicit recommendations on possible symposia titles for the SCCSR’s official con¬ tribution to the 20th Congress. The following is a brief summary of major activities of the working groups and task force, and the overall accomplishments of the standing committee during the review period, 1986-1990. IOC-SCCSR Special Symposium After considerable input and discussions by committee members and non-members between 1986-1988, a consensus was reached to undertake a review of the useful¬ ness of seabirds as bio-indicators of changing environments. The topic was consid¬ ered to be of global interest with more than 70% of respondents to the call for pos¬ sible symposia titles suggesting the subject area. A decision was taken to adopt a symposium proposal submitted by Dr R.W. Furness entitled “Seabirds as monitors of changing marine environments” with Dr Furness and Dr D.N. Nettleship as conven¬ ers. The emphasis is on seabird and fishery interactions with the intent of assessing the potential use of marine birds (direct and indirect) as indicators of fish stock sta¬ tus in a variety of marine ecosystems. Technological Innovations Workshop Efforts by members of the Technological innovations and Seabirds-at-sea working groups resulted in the formulation of a workshop to review recent technological ad¬ vancements in examining activity budgets of seabirds. Dr G.W. Gabrielsen, of the Norwegian Polar Research Institute/Norwegian Institute of Nature Research, accepted the committee’s invitation to serve as chairman of the workshop. Between 1988-1990, Dr Gabrielsen and his co-chairman, Dr K.L. Kooyman, assembled an impressive group of scientists from eight countries that will meet at the 20th Congress to iden¬ tify and display new measuring devices, and discuss parameters of measurement, the need for standardization of methodologies, and the interpretation of physiological function. 88 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Other Special Interest Congress-related Initiatives There have been a number of actions taken by the standing committee to enhance information exchange and the standardization and integration of approaches taken in various seabird research areas. Calls for further review on aspects of survey/census techniques and the structure and function of single-species and multi-species moni¬ toring systems are high, and advancements in cooperative approaches to problem solving have been made. For example, there is now good liaison between eastern and western Atlantic and eastern Pacific, including northern regions of Norway (Svalbard), Denmark (Greenland), Canada and USA (Alaska). Two special interest groups have formed since 1986 and will meet at the 20th Congress to further discuss and review cooperative monitoring programs (“Cooperative seabird studies in the North Atlantic”) and the need for computerized seabird colony databases (“Computerized colony reg¬ isters - their design and use in seabird research, management and conservation”). The agenda for the SCCSR Open Meeting is robust, comprising numerous topics for discussion put forth by the seabird research community at large. Those discussions should culminate in the development of resolutions and action plans that will facilitate a more effective seabird research effort worldwide. Bibliographies Two major bibliographic undertakings were discussed in 1986: (1) petrel bibliography - review of progress in a comprehensive collection of citations on the biology of the Procellaridae (shearwaters and petrels) and Hydrobatidae (storm-petrels) [compiler: Dr J. Warham], and (2) initiation of the compilation of materials on the pelagic distri¬ bution and ecology of seabirds [principal compiler: Dr R.G.B. Brown]. The petrel bib¬ liography now contains about 6,000 citations, with an expected total of about 8,000 entries. Dr Warham will be making a formal report at the SCCSR open meeting in December 1990, and may also give a demonstration of how this computerized bibli¬ ography is searched. The pelagic distribution and ecology bibliography has been ad¬ vanced to an incomplete rough-working stage, comprising about 2,000 entries con¬ centrated on work in the northern hemisphere; its present status is “inactive”, but work is expected to resume early in 1991. Nomenclature Discussions on current issues related to seabird nomenclature and systematics by members of the SCCSR Nomenclature working group usually take place either at specialist Round Table Discussions or in association with the Standing Committee on Ornithological Nomenclature. No progress report has been received from the SCCSR working group chairman, Dr P. Devillers, although an oral report is expected to be presented at the 20th Congress SCCSR open meeting followed by a written submis¬ sion. Discussions of proposals for action have been on-going throughout the review period involving committee members and a broad cross-section of other seabird scientists from many nations. Questions of work priorities predominated, focussing on the need to bring certain information from seabird organisations, regional groups, and individual researchers together to permit unified action on specific problems and issues. Initia¬ tives proposed by the standing committee since 1986, several of which have already been actioned or are under careful consideration, include: Seabird conservation - there is a need for a careful collaboration between IOC and ICBP for the development of a global strategy plan for the conservation of seabirds. (Status: under discussion). ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 89 Long-term population studies - committee members consider long-term population studies of seabird populations to be essential to a better understanding of population and community dynamics and ecological requirements of seabirds. Equally important is the provision of mechanisms to facilitate publication of these extensive life-historical researches. (Status: under discussion). Journal publications - there is a strong consensus among committee members of the need for an international journal of marine ornithology. The long-term objective is for the establishment of one first-class global journal rather than an overabundance of secondary regional seabird journals and bulletins. The committee feels that a major effort is required to discuss the roles of the various seabird groups and their coordi¬ nation. The magnitude of the task to achieve that goal is formidable. (Status: under discussion). Population surveys and monitoring - the committee recommends the development of a global view on population surveys and monitoring with international cooperation. The approach under consideration is for the establishment of a matrix of regions worldwide, each region with a subcommittee responsible for the development of a list of work priorities and specific recommendations for the placement and initiation of re¬ gional survey/monitoring systems. The SCCSR role would be one of coordination and the preparation of a preliminary proposal. (Status: under discussion). Seabird colony registeries - the development of regional, national, and international computerized databases is considered important to seabird research and manage¬ ment. Access to seabird colony data is a problem that will benefit from being tackled globally, with standardization of methods and procedures derived from groups already operating colony databases. The first international meeting on seabird colony databases was organised and chaired by the SCCSR chairman as a special paper session at the joint meeting of the Colonial Waterbird Society and the Pacific Seabird Group, Washington, D.C., USA, 12-16 October 1988. Participants, representing six colony databases from four countries [Britain (2), Canada (1), Norway (1), USA (2)], made detailed presentations in the program entitled “Computerized colony registries: their design and use in waterbird research, management and conservation” as a first step in the coordination of systems development. A second meeting is being held at the 20th Congress (see above). (Status: on going). Commercial fisheries - review existing knowledge of the impact of competition for food with fisheries (which is at best fragmented and imperfect) in an attempt to identify information gaps and approaches that might be taken to correct them. The overall objective is to develop an agenda for cooperative international research efforts. (Sta¬ tus: under discussion). Seabirds as bio-indicators - several investigators and research groups are address¬ ing questions relating to the use of bird populations as indicators of environmental change. Several committee members are key coordinators of research activity within this subject area: measurement of certain parameters in population status and repro¬ ductive performance. The standing committee serves an important communication and liaison function. (Status: on going). Population differentiation and quantitative characters - the committee feels that there is a strong need to identify the extent to which natural populations of seabirds mix with 90 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI one another and to determine the structure of populations in genetic terms. A recom¬ mendation is before the committee to set up a working group of interested parties to discuss and develop ideas of how best to tackle these questions. (Status: under dis¬ cussion). Climate change and seabirds - require the formulation of a workshop to assess the implications of climate change on populations of seabirds. (Status: in progress). Bibliographies - develop an agenda for the preparation of additional reviews on cer¬ tain species, families, and specific subjects. (Status: in progress). The central task of the standing committee continues to be the difficulty in maintain¬ ing long-term communication and cooperation between members and their working groups, and seabird researchers at large. The exchange of information function of the committee has been largely successful, as has its role in the identification of subjects for special interest symposia and workshops on seabirds. However, it is felt that the committee’s influence and effectiveness can only be enlarged by an increase in the number of active participating members, additions that will provide a broader repre¬ sentation of seabird researchers throughout the world. These and other issues will be reviewed and discussed at our New Zealand meetings as we plan our activities for the next four years. Members of the Standing Committee for the Coordination of Seabird Research for the period 1986-1990 (20th Congress) are as follows: P.H. Becker (Germany) Brian D. Bell (New Zealand) W.R.P. Bourne (United Kingdom) R.G.B. Brown (Canada) P.A. Buckley (USA) J. Cooper (South Africa) J.C. Coulson (United Kingdom) J.P. Croxall (United Kingdom) P. Devillers (Belgium) A.W. Diamond (Canada) D.C. Duffy (USA) P.G.H. Evans (United Kingdom) R.W. Furness (United Kingdom) A.N. Golovkin (USSR) G.L. Hunt (USA) W. Hsu (China) J.R. Jehl (USA) C. Jouanin (France) N. Kuroda (Japan) J. L. Mougin (France) D. N. Nettleship (Canada) C.J.R. Robertson (New Zealand) R.P. Schlatter (Chile) W.R. Siegfried (South Africa) K. Vermeer (Canada) J. Warham (New Zealand) G. Watson (USA) V. Zubakin (USSR) The members of the IOC-SCCSR for the period 1986-1990 were appointed by Presi¬ dent Charles G. Sibley in 1986 with three additions made later by the committee chair¬ man to offset the loss of members (deaths and resignations). Respectfully submitted for the Standing Committee for the Coordination of Seabird Research. David N. Nettleship Chairman, SCCSR 1 0 November 1 990 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 91 REPORT OF THE STANDING COMMITTEE ON APPLIED ORNITHOLOGY The Standing Committee on Applied Ornithology was set up at the 19th International Ornithological Congress in Ottawa in 1986 at the suggestion of the President, the late Professor Klaus Immelmann. No precise terms of reference were given to the Com¬ mittee, which was therefore left to decide for itself what the scope of its activities should be in principle and what it should try to achieve in practice. A first short, informal meeting of the available members of the Committee was held in Ottawa. Those present included Professor Valery Ilyichev, R. Dowsett, Professor V.E. Flint, Dr M. Luniak, Dr N. Nankinov, Dr E. Rutschke, and Dr J. Temple Lang. The structure, officers, method of operation and a variety of possible activities for the Com¬ mittee were discussed. It was agreed that the Committee must not duplicate the work of the existing international bodies concerned with bird conservation and applied or¬ nithology, but should work as closely as possible with them when appropriate. The Committee would have to work primarily through scientific cooperation and exchange of information. The possibility of the Committee arranging an international conference on applied ornithology was raised. After this informal meeting in Ottawa the members of the Committee were formally appointed by Professor Sibley, the President of the 20th International Ornithological Congress. Professor Ilyichev and Dr Russell Peterson were appointed Co-Chairmen, and Dr Vladimir Yacoby and Dr John Temple Lang were appointed Co-Secretaries. The members of the Committee were asked to say what activities they thought the Committee should undertake. A very large number of suggestions were made, and it became clear that it was necessary for the Committee to choose some areas on which to concentrate, bearing in mind that the Committee has no funds of its own and that its members are widely spread throughout the world. The first formal meeting of the Committee was on 19 May 1987 in Kecskemet, Hun¬ gary. Present were Professor Ilyichev, Dr Peterson, Dr Bankovics, Dr Cooch, Profes¬ sor Nicolai, Dr Yacoby and Dr Temple Lang. Most of the meeting was devoted to dis¬ cussing possible topics for discussion at the 20th Congress, the conference of the International Council for Bird Preservation in 1990, or a suggested conference to be arranged by the Committee in the USSR. It was agreed to make a number of recom¬ mendations to the Scientific Programme Committee of the 20th Congress, including a plenary session on bird conservation; symposia on collisions and conflicts between birds and aircraft, powerlines, vehicles and lighthouses; transmission by birds of dis¬ eases to man; and population explosions of wild bird species. In addition, it was agreed to support other symposia topics to be suggested by other scientists on con¬ trol of damage by birds to agriculture, and captive breeding of wild bird species. The meeting also agreed that in general the Committee should promote discussion, research and exchange of information on practical, conservation-oriented subjects. The Committee would need to set up working groups on particular areas of study, in particular on collisions involving birds, transmission by birds of diseases, and popu¬ lation explosions of wild bird species. 92 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI This discussion was on the basis of a paper summarising the ideas and suggestions received by Dr Temple Lang about the priorities which the Committee might adopt for its activities. These suggestions were concerned primarily with harm done by birds to man, harm done by man to birds, and also with education, research techniques, ur¬ banisation problems concerning birds, and human use of birds. The second formal meeting of the Committee was held in Cambridge, England, on 29 April 1989. Present were Dr Peterson, Professor Ilyichev, Dr Alvarez Lopez, Dr Blokpoel, Dr Imboden, Dr Yacoby and Dr Temple Lang. Various topics which could be described as applied ornithology were included in the scientific programme of the 20th Congress. The recommendations of the Committee had usefully influenced the programme of the Congress. It was agreed to propose a Round Table Discussion at the 20th Congress on the future role and work of the Committee. The possibility that the Committee might arrange or sponsor a conference on applied ornithology was again discussed. It was agreed that the future work of the Committee should concern itself with applied ornithology, understood as optimising bird-man relations. This covers both reducing damage by birds to man (broadly, economic ornithology) and damage by man to birds (conservation). It is not possible wholly to separate these, since for example collisions between birds and aircraft both damage planes and kill birds. The aim should be to minimise damage to man with the minimum killing of birds. It was agreed that the Committee should try to avoid overlapping with the work of the International Council for Bird Preservation. It was pointed out that the 19th Congress had set up the Com¬ mittee to interest scientists in conservation, and to raise the standards of conserva¬ tion research. It is characteristic of applied ornithology that its conclusions need to be acted on by non-ornithologists, such as airport managers, agricultural advisers etc. who do not attend the International Ornithological Congresses (so that special meet¬ ings may be needed) and who may neglect or harm birds unless informed by applied ornithologists. The Committee therefore needs to make information available to these non-ornithologist users. The Committee could gather information worldwide, arrange exchange of information, and use the conclusions of scientific ornithology for practi¬ cal purposes. It was agreed that the Committee should continue after the 20th Con¬ gress. As in the case of other IOC Standing Committees, the members of the Standing Com¬ mittee on Applied Ornithology were appointed by the President of the 20th Congress until the close of that Congress. During the period of 1986 to 1990 the members of the Standing Committee on Applied Ornithology were: Co-Chairmen: Professor Dr Valery D. Ilyichev, Dr Russell Peterson Co-Secretaries: Dr John Temple Lang, Dr Vladimir E. Yacoby Dr Humberto Alvarez-Lopez Dr George W. Archibald Dr Attila Bankovics ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 93 Dr Miroslava Beklova Dr A. Bold Dr Hans Blokpoel Dr Donald Bruning Dr F. Graham Cooch Dr Stephen J. Davies Dr Robert Dowsett Professor Dr V.E. Flint Dr Luc Hoffmann Professor Dr W. Hsu Dr S.A. Hussain Dr Noritaka Ichida Dr Christoph Imboden Dr Maciej Luniak Dr Phil Moors Dr Dmitri Nankinov Professor Dr Jurgen Nicolai Professor Dr Erich Rutschke This report is respectfully submitted on behalf of the Standing Committee on Applied Ornithology of the International Ornithological Congress. Valery D. Ilyichev, Russell W. Peterson Co-Chairmen, SCAO 94 ACTA XX CONK — * . ' 7 . ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 95 XXI CONGRESSUS INTERNATIONALIS ORNITHOLOGICUS INTERNATIONAL ORNITHOLOGICAL COMMITTEE 1990-1994 Honorary President Karel H. Voous Netherlands President Christopher M. Perrins United Kingdom Vice-President Svein Haftorn Norway Secretary-General John Dittami Austria Secretary Walter J. Bock USA EXECUTIVE COMMITTEE Chairman: Christopher M. Perrins United Kingdom Vice-Chairman: Svein Haftorn Norway Peter Berthold Germany Walter J. Bock USA Cynthia Carey USA John Dittami Austria Richard Liversidge South Africa Henri Ouellet Canada Murray J. Williams New Zealand Ben D. Bell New Zealand Jacques Blondel France Enrique H. Bucher Argentina Asha Chandola-Saklani India Evgeny N. Kurochkin USSR Hiroyuki Morioka Japan Charles G. Sibley USA PAST PRESIDENTS Charles G. Sibley (1986-1990) USA Lars von Haartman (1978-1982) Finland Jean Dorst (1970-1974) France Ernst Mayr (1958-1962) USA PAST SECRETARIES-GENERAL Ben D. Bell (1986-90) New Zealand Henri Ouellet (1 982-1 986) Canada Valery Ilyichev (1978-1982) USSR Karel H. Voous (1966-1970) Netherlands Charles G. Sibley (1958-1962) USA Lars von Haartman (1954-1958) Finland 96 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI INTERNATIONAL ORNITHOLOGICAL COMMITTEE (* Newly elected members; ** Re-elected members; + Permanent members) SENIOR MEMBERS Jurgen Aschoff Germany Biswamoy Biswas India Tso-Hsin Cheng P.R. China Kai Curry-Lindahl Sweden (died 5/12/90) Camille Ferry France Paul Geroudet Switzerland Lars von Haartman+ Finland Holger Holgersen Norway Christian Jouanin France Richard Liversidge South Africa Bernt Loppenthin Denmark Gerlof F. Mees Netherlands Tsukasa Nakamura Japan Jurgen Nicolai Germany Floriano Papi Italy S. Dilion Ripley USA Eric Rutschke Germany Ernst Schiiz Germany (died 8/3/91) Flelmut Sick Brazil (died 5/3/91) Ernst Sutter Switzerland George D. Vasiliu Rumania John Warham New Zealand Kurt Bauer Austria Andre Brosset France G.R.Cunningham-van-Someren Kenya J. Bruce Falls Canada Zafar Futehally* India Friedrich Goethe Germany Svein Flaftorn Norway Thomas R. Flowell USA J. Allen Keast Canada Hans Lohrl Germany Ernst Mayr+ USA Wilhelm Meise Germany E. Max Nicholson United Kingdom Hans Oeme Germany Roger Tory Peterson USA Ian C.R. Rowley Australia Alfred Schifferli Switzerland Charles G. Sibley+ USA David W. Snow United Kingdom Melvin A. Traylor USA Karel H. Voous+ Netherlands Gerhardt Zink Germany NATIONAL REPRESENTATIVES ARGENTINA Enrique H. Bucher Manuel Nores* AUSTRALIA Sidney John Cowling Jiro Kikkawa* Phillip Moors* Pat V. Rich* Richard Schodde AUSTRIA John Dittami*+ Herbert Schifter Ellen Thaler* Hans Winkler* BELGIUM Pierre Devillers Andre A. Dhondt* Michel Louette* BRAZIL Paulo de Tarso Z. Antas* Roberto B. Cavalcanti BULGARIA Dimiter Nankinov CANADA Jon C. Barlow* David A. Boag Fred Cooke* Anthony J. Erskine Raymond McNeil* David N. Nettleship* Henri Ouellet+ W. John Richardson CHILE Fabian M. Jaksic* ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 97 COLOMBIA F. Gary Stiles* CUBA Orlando H. Garrido-Calleja* Hiram Jose Gonzalez-Alonso CZECHOSLOVAKIA Karel Hudec** Aladar Randik* Andrey Stollmann* DENMARK Jan Dyck Jon Fjeldsa ECUADOR Fernando I. Ortiz-Crespo EIRE Oscar Merne* FINLAND Olavi Hilden* Pertii Lauri Saurola FRANCE Jacques Blondel Jean Dorst+ Christian Erard Bernard Frochot Jean-Marc Thiollay GERMANY Peter Berthold Eberhard Gwinner Jochen Martens Roland Prinzinger* Klaus Schmidt-Koenig* Roswitha Wiltschko* Wolfgang Wiltschko HUNGARY Attila Bankovics Lajos Sasvari* ICELAND Arnthor Gardarsson** INDIA Asha Chandola-Saklani Manjit S. Dhindsa* S. Ali Hussain* INDONESIA Soekarja Somadikarta ISRAEL Yoram Yom-Tov* Amotz Zahavi ITALY S. Frugis Fernando Spina* Carlo Violani* JAPAN Yuzo Fujimaki* Hiroyoshi Higuchi* Hiroyuki Morioka Masashi Yoshii KENYA Nathan N. Gichuki* MALAYSIA David R. Wells MEXICO Mario A. Ramos-Olmos NETHERLANDS R.H. Drent Arie L. Spaans* Gart Zweers* NEW ZEALAND Ben D. Bell+ John L. Craig* Murray J. Williams* NORWAY Tore Slagsvold* PEOPLE S REPUBLIC OF CHINA Wei-shu Hsu Zheng-jie Zhao* Guang-mei Zheng * 98 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI PERU Manuel A. Plenge POLAND Jan Krystyn Pinowski Ludwik Tomialojc Tomasz Wesolowski* REPUBLIC OF CHINA Lucia Liu Severinghaus* REPUBLIC OF KOREA (S. KOREA) Pyong-Oh Won REPUBLIC OF SOUTH AFRICA Timothy M. Crowe* Alan Kemp* W. Roy Siegfried** RUMANIA Laszlo Kalaber* SPAIN Carlos M. Herrera SWEDEN Soren Svensson Staffan Ulfstrand* SWITZERLAND Bruno Bruderer Urs Glutz von Blotzheim Lucas Jenni* THAILAND Pilai Poonswad UGANDA Derek E. Pomeroy* UNION OF SOVIET SOCIALIST REPUBLICS Alexander V. Andreev* Victor R. Dolnik Valery E. Flint Vladimir M. Galushin Valery Ilyichev* Evgeny N. Kurochkin Roald L. Potapov* Yuri V. Shibaev* UNITED KINGDOM Philip J.K. Burton John P. Croxall* George M. Dunnet* Peter R. Evans* Brian K. Follett Janet Kear Ian Newton Peter J. Olney Christopher M. Perrins+ UNITED STATES OF AMERICA Russell P. Baida Luis F. Baptista Walter J. Bock+ Cynthia Carey John W. Fitzpatrick* Frank B. Gill Richard T. Holmes* Dominique G. Homberger* Frances C. James Ned K. Johnson Ellen D. Ketterson* James R. King (died 7/4/91) Burt L. Monroe Jr.* Lester L. Short John C. Wingfield* Glen E. Woolfenden ZIMBABWE Peter J. Mundy* ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 99 THE STATUTES AND BY-LAWS OF THE INTERNATIONAL ORNITHOLOGICAL COMMITTEE The Statutes and By-Laws of the International Ornithological Committee were origi¬ nally prepared by Donald S. Farner, President of the XVII International Ornithologi¬ cal Congress and adopted at that Congress in Berlin, 1978. They replaced the Reglement des Congres Ornithologiques Internationaux adopted at the IX Interna¬ tional Ornithological Congress in Rouen, 1938, and all amendments passed thereaf¬ ter. STATUTES Article I Objectives and Purposes The International Ornithological Committee (IOC) (1) promotes international collabo¬ ration and cooperation in ornithology and (2) as it deems desirable and useful, encour¬ ages international collaboration and cooperation between ornithology and other bio¬ logical sciences. • To effect these objectives and purposes the IOC sponsors and promotes International Ornithological Congresses; establishes and sponsors commissions and committees as it deems appropriate and desirable; establishes or sponsors other international ornithological activities as it deems appropriate; and functions as the Section of Or¬ nithology of the International Union of Biological Sciences. Article II Membership and Functions 1. Size. The size of the membership of the IOC is determined by the Committee, but may not exceed the number specified in the By-Laws (Art. I). 2. Representation. The members shall be representative of the international distribution of ornithologists, and the number of members from each country shall be proportional to its ornithologi¬ cal activity. 3. Election. New members are elected by the IOC at a regular meeting at the International Orni¬ thological Congress from a list of nominations prepared and presented by the Execu¬ tive Committee (EC). Proposals for this list can be made by any member of the IOC; they should be in writing with adequate documentation and submitted to the President and the Secretary at least six months prior to the next congress. Election to the IOC requires a simple majority of the members present and voting. 4. Term. The term of membership is indefinite unless the member resigns voluntarily or is 100 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI absent from regular meetings of the IOC at two consecutive congresses, which con¬ stitutes automatic resignation. Resigned members may be re-elected. 5. Meetings. The IOC meets at least twice during each International Ornithological Congress. The quorum for transaction of business at a regular meeting consists of the members present at the meeting. A member must be in attendance at a meeting in order to cast his/her vote. 6. Duties. The duties of the IOC are: (a) to select the site of the next congress; (b) to elect new members of the IOC; (c) to elect the President, Vice-President, Secretary and any Honorary Officers of the next congress; (d) to elect members to the EC; and (e) to take actions appropriate and necessary to carry out its stated objectives and functions (Art. I). 7. Special meetings. The President, under extraordinary circumstances, may call a special meeting of the IOC, and is obligated to do so on receipt of a petition signed by one-quarter of the members. The date set for a special meeting must permit reasonable time for consid¬ eration of the agenda and for travel arrangements. A quorum for a special meeting is one-third of the members of the IOC. Failure to attend a special meeting shall not count toward automatic resignation (Art 11:4). 8. Presiding officer. The President presides at the meetings of the IOC. 9. Communications. Actions of the IOC are communicated to the congress and published either in the pro¬ ceedings of the congress or in some other publication, as approved by the EC. Article III Officers A. The President 1 . Election. The President is elected by a simple majority of the members present and voting at a regular meeting of the IOC at an International Ornithological Con¬ gress and is not eligible for election to the same office in two successive con¬ gresses. 2. Term. The President holds office from the conclusion of the congress at which elected until the conclusion of the following congress. 3. Duties. The President of the IOC also serves as chair of its EC, as President of the International Ornithological Congress, and (or designates a representative) as Chair of the Section of Ornithology of the International Union of Biological Sciences. The President presides at meetings of the IOC, of its EC and of the International Ornithological Congress, and appoints committees and commis¬ sions (with the exception of the EC) of the IOC and of the congress. After ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 101 consultation with the host organisation of the forthcoming congress, the Presi¬ dent shall appoint the Secretary-General. The President appoints the Secretary of the Section of Ornithology of the IUBS. 4. Membership in the IOC. Past Presidents are permanent members of the IOC. The immediate Past President serves ex officio as a member of the EC. B. The Vice-President 1 . Election. The Vice-President is elected, following the election of the President, by a simple majority vote of the members present and voting at a regular meet¬ ing of the IOC at an International Ornithological Congress and is not eligible for election to the same office in two successive congresses. 2. Term. The Vice-President holds office from the conclusion of the congress at which elected until the conclusion of the following congress. 3. Duties. The Vice-President of the IOC also serves as Vice-Chair of the EC. 4. Succession. The Vice-President shall serve as president of the IOC in case of the inability of the elected President in office to continue until the completion of the normal term of the President in office. C. The Secretary-General 1. Appointment. The Secretary-General is appointed by the President (Art. 1 1 1 :A,3) after consultation with the host organisation of the forthcoming congress. 2. Term. The Secretary-General serves until the Secretary-General of the follow¬ ing International Ornithological Congress is designated. 3. Duties. The Secretary-General serves as Secretary-General and Treasurer of the congress, having all local and financial responsibilities for the preparation and running of the congress, including publication of the congress proceedings. The Secretary-General may nominate, for Presidential appointment, persons to serve in definite capacities such as treasurer and editor or on various local com¬ mittees for the congress. The Secretary-General serves ex officio as a voting member of the EC. 4. Membership in the IOC. Past Secretaries-General are permanent members of the IOC. The immediate Past Secretary-General serves ex officio as a member of the EC. D. The Secretary 1 . Election. The Secretary is elected, following the election of the President and the Vice-President, by a simple majority of the members present and voting at a regular meeting of the IOC at an International Ornithological Congress, and is eligible for re-election. 2. Term. The Secretary holds office from the conclusion of the congress at which elected until the conclusion of the following congress. 102 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 3. Duties. The Secretary shall keep the records of the IOC and its EC, prepare the agenda of meetings of the IOC and EC, serve as parliamentarian at these meet¬ ings, record and distribute within two months following the congress the minutes of the IOC and the EC meetings, prepare the published communications of these meetings, deal with communications of the IOC and EC as directed by the Presi¬ dent, and assist the President and Secretary-General in preparation of the con¬ gress. The Secretary is responsible for communicating with and assisting orni¬ thologists of potential host countries in the preparation of invitations for future congresses. The Secretary is a member of the EC. 4. Membership in the IOC. Past Secretaries are permanent members of the IOC. E. Honorary Officers The IOC, or the President with the consent of the EC, may appoint honorary officers, such as Honorary Presidents and Honorary Vice-Presidents of the congress, to rec¬ ognise the contributions of ornithologists and other persons to the cause of interna¬ tional ornithology. Honorary Presidents and Honorary Vice-Presidents are members ex officio of the IOC. Article IV The Executive Committee 1 . Membership a. The President (Art. III:A,3), Vice-President (Art. 1 1 1 :B,3) , the Secretary-General (Art. 1 1 1 :C,3) , the Secretary (Art. 1 1 1 : D ,3) , the immediate Past President (Art. 1 1 1 :A,4) , and the immediate Past Secretary-General (Art. 1 1 1 :C,4) of the IOC un¬ til the end of the following congress. b. An even number of elected members, as specified by the By-Laws (Art. III). No more than one of these members may be from a single country. These members shall be elected with proper attention to an adequate international distribution in the EC. 2. Election. Nomination and election of members of the EC shall follow election of the President, Vice-President, Secretary and any Honorary Officers. Nomination shall be proposed by the existing EC. Any member of the IOC present at the meeting may make addi¬ tional nominations; if seconded, these are added to the nominations proposed by the EC. Election of members of the EC is by simple majority vote of members of the IOC present and voting. Elected members are eligible for re-election as an elected mem¬ ber of the EC for one additional term. 3. Term. The EC shall serve from the conclusion of the congress at which it is elected to the conclusion of the following congress. 4. Duties. a. During the inter-congress period, the EC acts on the behalf of the IOC. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 103 b. During the inter-congress period, the EC has general responsibility for the sci¬ entific policy of the IOC including the programme of the congress, as specified in the By-Laws (Art. IV:4). c. At meetings of the IOC at an International Ornithological Congress, the EC pro¬ vides: (1) Nominations for the offices of President, Vice-President, Secretary, and Honorary Officers and for the elected members of the EC; (2) A recommendation concerning the host country and organisation for the en¬ suing congress after due consideration of all invitations; (3) Nominations for new members of the IOC with due consideration of Art. 11:2; (4) Recommendations for re-election of members considered to have resigned because of absence from two consecutive meetings, as specified in Art. 11:4; (5) Advice and counsel concerning any other matters deemed to be of interest within the purview or among the responsibilities of the IOC. Article V Amendment of the Statutes 1. Proposal of amendment. Proposals to amend the statutes require the signatures of at least five members of the IOC from at least three countries, and must be transmitted to the President and Sec¬ retary at least twelve months before the next International Ornithological Congress. The Secretary will distribute the proposed amendments to all members of the IOC at least four months prior to the congress. At the meeting of the IOC at the congress the EC will present its recommendation on each proposed amendment. 2. Adoption. Adoption of an amendment by the IOC requires a two-thirds majority vote of the mem¬ bers present and voting. Adopted amendments become effective at the close of the congress. Article VI Enabling Clause Adoption of these statutes requires a two-thirds majority vote of the members of the existing IOC present and voting at a regular meeting of the International Ornithological Congress at which they are presented, having been distributed to the members prior to that meeting. Adoption of these statutes shall replace the Reglement des Congres Ornithologiques Internationaux adopted in Rouen in 1938 and all amendments passed thereafter. If adopted, these Statutes become effective immediately. 104 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI BY-LAWS Article I The Size of the International Ornithological Committee (IOC) The membership of the Committee shall not be more than 140. Members over 65 years of age (Senior Members), Past Presidents, Past Secretaries-General and Past Secretaries are not counted in this limit. Members of the IOC must be residents of the country that they represent. Senior Members are permanent members of the IOC and are not subjected to the requirements of Art. 11.4 of the Statutes. Article II Meetings of the International Ornithological Committee 1 . Sufficiently prior to the regular meeting of the IOC at an International Ornithologi¬ cal Congress, the Secretary prepares the agenda of the meeting after consultation with the President and the Secretary-General. The Secretary-General then distributes to all members the agenda of the meeting. 2. Members are requested to inform the President and the Secretary of their intention to attend the meeting and/or resign from the Committee. 3. An agenda and information on matters to be covered shall be sent to members with the notice of any special meeting called by the President. Article III Membership of the Executive Committee (EC) In addition to the officers specified in Art. IV of the Statutes, the IOC elects ten mem¬ bers of the EC in accordance with Art. I V: 1 , b of the Statutes. Article IV , The International Ornithological Congresses 1. The frequency of congresses. Congresses will be held at four-year intervals unless, for compelling reasons, the IOC, or the EC acting on its behalf, deems otherwise. 2. The site and time of congresses. After consultation with the EC and the host organisation, and due consideration of the interests and convenience of the members, the site in the host country and time of the congress are fixed by the Secretary-General. 3. Membership of congresses. Membership in an International Ornithological Congress shall be open to all ornitholo¬ gists and students of avian biology without distinction as to country of origin upon payment of the stated congress fee, if any. Membership and attendance at a congress shall be in accordance with the general policies of the International Union of Biological Sciences. Any limitation on the number of active members of the congress may be ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 105 made by the Secretary-General only after consultation with and agreement by the EC. Such limitation must be clearly stated in congress announcements. In the case of limi¬ tation in the number of active members, provision must be made for associate mem¬ bers which may not be limited in number. Members of the IOC may not be denied a place as an active member upon payment of the congress fee. 4. The scientific programme of the congress. After consultation with the EC and the host organisation, the President appoints the Scientific Programme Committee. This Committee consists of three or more members from the host country and members from at least three other countries. The President, Secretary-General and Secretary are members ex officio of the Scientific Programme Committee. This committee is responsible to the EC for the scientific programme of the congress. 5. The organisation of the congress. The general organisation of, and the arrangements for, the congress are the respon¬ sibilities of the Secretary-General. 6. The proceedings of the congress. The Secretary-General is responsible for the publication of the proceedings of the congress, and serves as editor of the proceedings or appoints an editor after obtaining concurrence from the President. 7. Finances of the congress. The Secretary-General is the treasurer and principal officer of the congress and as such is responsible for all financial matters of the congress. In consultation with the President, the Secretary-General develops the budget and fixes congress fees. After all fiscal obligations have been absolved, any surplus funds, including any from the proceedings, are made available for inter-congress activities, including arrangements for the ensuing congress. 8. Hosting of future congresses. (a) Invitations from countries to host an International Ornithological Congress im¬ mediately following the next scheduled congress should be sent to the Presi¬ dent, Secretary-General and Secretary no later than six months before the next scheduled congress. (b) On request, the Secretary-General and the Secretary shall provide representa¬ tives of intending host countries with a list of information required in their in¬ vitation document and with general guidelines for submitting such an invitation. Article V Amendment of the By-Laws 1. Proposal of amendment. Proposals to amend the By-Laws require the signature of at least three members from at least three countries and must be transmitted to the President, Secretary-General and Secretary at least twelve months in advance of the next International Ornithologi¬ cal Congress. At least four months prior to the congress, the Secretary shall distrib¬ ute the proposed amendments to the members of the International Ornithological 106 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Committee. At the meeting of the IOC at the congress, the EC will present its recom¬ mendation on each proposed amendment. 2. Adoption. Adoption of the proposed amendments to the By-Laws by the IOC requires a simple majority vote of the members present and voting. Adopted amendments become ef¬ fective at the close of the congress. 3. Conflict with the Statutes. No amendment of the By-Laws can have the effect of modification of the Statutes. Article VI Enabling Clause Adoption of these By-Laws requires prior adoption of the proposed Statutes and re¬ quires a simple majority vote of the members of the existing Committee present and voting at a regular meeting of the IOC at the congress at which they are presented, having been distributed to the members prior to that meeting. Adoption of these By- Laws shall replace any existing By-Laws and regulations (formal and informal) of the IOC and of the International Ornithological Congresses. If adopted these By-Laws become effective immediately. / ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 107 BUCKINGHAM PALACE. As the human population of the world continues to increase, so the populations of most other species are made to contract and give up their habitats. The endemic birds of New Zealand have also been made to suffer in other ways. The introduction of exotic animals, most of which have either displaced native species or become predators, has caused the extinction of a number of endemic bird species and given New Zealand a disproportionately high share of the world's rare and endangered birds. This in turn has led to the added threats of poaching and the illegal trade in endangered species. The situation would undoubtedly have been far worse but for the heroic endeavours of voluntary and professional ornithologists and conservationists in New Zealand. Their struggle is at a critical stage and I hope that the presence of the 20th International Ornithological Congress in New Zealand will bring them much needed recognition and encouragement. Of all the wild animals of this world, birds are the most frequently seen by the human population, and any changes in the numbers of resident and migratory birds is quickly noticed and reported. Birds have therefore become the 'indicator' species for wild populations as a whole. This has given ornithology a special significance in the field of nature conservation and I know that the deliberations of this Congress will be followed with great interest by the biological research community and by the whole conservation movement. I have no doubt that the organisers have arranged an interesting and varied program, which will be of special interest to those unfamiliar with New Zealand’s unique natural history. I send you all my very best wishes for a happy and productive meeting and I hope that you will return home with greater confidence in the knowledge that you are part of a worldwide community of dedicated enthusiasts. 1990 108 ACTA XX CONGRESSUS IN' NALIS OR * / . ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 109 PRESIDENTIAL ADDRESS PHYLOGENY AND CLASSIFICATION OF BIRDS FROM DNA COMPARISONS CHARLES G. SIBLEY ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 111 PHYLOGENY AND CLASSIFICATION OF BIRDS FROM DNA COMPARISONS CHARLES G. SIBLEY Tiburon Center for Environmental Studies, San Francisco State University, Box 855, Tiburon, California 94920, USA ABSTRACT. DNA-DNA hybridization was used to compare the single-copy nuclear DNAs of ca. 1700 species of birds, representing 167 of the 171 families in Wetmore’s (1960) classification. The DNA-DNA hybridization technique provides a measure of the percentage of base-pair mismatch that has accu¬ mulated in the two lineages since the two species that are being compared last shared a common ancestor. The DNA hybridization distance measures were used to reconstruct the phylogeny of the higher categories and a new classification was derived from the phylogeny. Examples of the most in¬ teresting and/or controversial results are included herein, including the ratites, barbets and toucans, sandgrouse, pelecaniforms, tubenoses, lyrebirds and bowerbirds, vireos, starlings and mockingbirds, and nectar-feeding passerines. This paper is a synopsis of research performed at Yale University (1974-1986) with the collaboration of Dr. Jon E. Ahlquist and the assistance of many persons in the field and laboratory. A classification based on the DNA-DNA hybridization data has been published in The Auk (Sibley et al. 105: 409-423,1988.) Sibley & Ahlquist (1990) present details of the technique and the complete data. Keywords: Phylogeny, classification, DNA-DNA hybridization. INTRODUCTION The avian lineage branched from its reptilian ancestor in the Jurassic Period, about 150 million years ago. The ca. 9700 living species of birds may be less than 10% of the species that have existed since the origin of the group. There are few fossils of Mesozoic birds and most may not have left living descendants. Cenozoic fossils are more abundant, but bird bones are fragile and the record is sparse compared with that for other groups of vertebrates. Until the advent of molecular methods some 35 years ago, avian systematists had to rely mainly on comparative anatomy for evidence of the phylogenetic relationships among living lineages of birds. Closely related species are usually easy to recognize, but as the phylogenetic distances among taxa increase the interpretation of morphological characters becomes more difficult. For example, we can see that ducks, geese, and swans are closely related to one another, but which group is most closely related to them? This question applies to most of the higher categories of birds and during the 19th century it led to many proposals con¬ cerning the relationships among the major groups of birds. The classification of Hans Gadow (1892,1893) became the principal basis for that of Alexander Wetmore (1930, 1960), and Erwin Stresemann (1934) presented a system based mainly on those of Gadow and Maximilian Furbringer (1888). These two classifications have been widely used for the past 60 years. The history of avian classification and evidence of the phylogeny of birds have been reviewed by Sibley & Ahlquist (1990). This paper is a synopsis of some of our proposals concerning the phylogeny and classification of birds, based on studies using the technique of DNA-DNA hybridization. Some new data are presented and some corrections are noted. 112 ACTA XX CONGRESSUS INTERNATION ALIS ORNITHOLOGICI METHODS DNA-DNA hybridization measures the degrees of genetic similarity between species. “Hybrid” double-stranded DNA molecules are formed from the single strands of the DNAs of two species. The hybrid molecules are then dissociated (“melted”) in a ther¬ mal gradient under controlled conditions such that a measure of the difference be¬ tween the two nucleotide sequences may be calculated. The experimental conditions are set so that only homologous sequences can form double-stranded structures. The melting temperature of a DNA duplex molecule is a function of the number of correctly base-paired nucleotides, thus it is a measure of the degree of genetic similarity be¬ tween the two single strands forming the duplex. Data are expressed as melting curves and as distances between the midpoints of the melting curves. Dendrograms that do, and do not, assume equal rates of genomic evolution along all branches may be constructed to represent the branching pattern of the phylogeny indicated by the distance values. The technique, data analysis, and other aspects of the procedures are described by Sibley & Ahlquist (1983, 1986, 1987, 1990). The principal steps in the DNA-DNA hybridization technique follow: 1. Extract and purify DNA from cell nuclei = remove proteins, RNAs, etc. 2. Shear long-chain DNA strands into fragments ca. 400-600 bases in length. 3. Remove most of the copies of repeated sequences from selected species to pro¬ duce “single-copy DNA.” 4. “Label” the single-copy DNA with a radioactive isotope to produce a “tracer” DNA of one species = Species A. 5. Combine the single-stranded tracer DNA of Species A with the single-stranded “driver” DNA of the same species (A + A), and with the single-stranded driver DNAs of other species (A + B, A + C, A + D, etc.). Each combination is placed in a separate vial. 6. Incubate the vials in a waterbath at 60°C for 120 hours to permit the formation of double-stranded hybrid molecules composed of one strand of the tracer (A) and one strand of the driver (B, C, D, etc.) to produce the hybrids: A x A, A x B, A x C, A x D, etc. 7. Place the DNA-DNA hybrids on hydroxyapatite (HAP) columns. Double-stranded DNA binds to HAP; single-stranded DNA does no? bind to HAP. 8. Place the columns in a heated waterbath and raise the temperature in 2.5°C in¬ crements from 55 to 95°C. At each temperature, wash off (elute) the single- stranded DNA resulting from the “melting” of the hydrogen bonds between base pairs. Collect each eluted sample in a separate vial and assay the radioactivity in each vial. This is a measure of the percentage of hybrid molecules that melted at each temperature. 9. The melting temperature of a DNA-DNA hybrid is proportional to the degree of genetic similarity between the two single strands forming the hybrid molecule. Use the amount of radioactivity in each sample to construct melting curves and to calculate genetic distance values. Construct “trees” from the genetic distance values. In the following brief accounts of the major groups of birds, it is assumed that read¬ ers are familiar with the geographic distributions of most groups and with the English names of groups and/or species. English and scientific names follow Sibley & Monroe (1990). ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 113 In the classification of Sibley et al. (1988) the boundaries of categories are based on the DNA hybridization distance values (AT50H). For example, Orders are groups that differ from one another by an average AT50H value between 20 and 22; Families dif¬ fer by A9-1 1 , etc. We use the following categories and ranges of AT50FI values: Class (31-33), Subclass (29-31), Infraclass (27-29), Parvclass (24.5-27), Superorder (22- 24.5), Order (20-22), Suborder (18-20), Infraorder (15.5-18), Parvorder (13-15.5), Superfamily (1 1-13), Family (9-11), Subfamily (7-9), and Tribe (4.5-7). RATITES AND TINAMOUS The living ratites (Ostrich, rheas, Emu, cassowaries, kiwis) form a monophyletic group (Struthioniformes) with the tinamous (Tinamiformes) as their closest living relatives. The Emu and cassowaries (Casuariidae) are closely related to one another, and more closely related to the kiwis (Apterygidae) than to the Ostrich (Struthionidae) and rheas (Rheidae). The branches leading to the Ostrich, rheas, and the Emu-cassowary-kiwi cluster occurred close together, probably in the late Cretaceous. The present distri¬ bution of the ratites is a result of the breakup of Gondwanaland and the drift of the southern continents to their present positions. The ancestor of the kiwis and moas may have reached New Zealand via stepping-stone islands across the northern Tasman Sea. GALLINACEOUS BIRDS AND WATERFOWL The guans (Cracidae) and megapodes (Megapodiidae) are placed in a separate order (Craciformes) from the Galliformes. The New World quail (parvorder Odontophorida) are not closely related to the pheasants, Old World quail, grouse, turkeys, and guineafowl (parvorder Phasianida). The waterfowl (Anseriformes) seem to be the closest living relatives of the gallinaceous birds, but the divergence was probably in the late Cretaceous or early Tertiary. Sibley et al. (1988) included the parvclass Galloanserae (Craciformes, Galliformes, Anseriformes) with the ratites and tinamous (parvclass Ratitae) in the infraclass Eoaves. Sibley & Ahlquist (1990: 255, 288) and Sibley & Monroe (1990: 5) moved the Galloanserae to the beginning of the infraclass Neoaves. BUTTONQUAILS The Turnicidae are distant from all other living groups. The Australian Plains-wanderer Pedionomus is not closely related to Turnix, but is a charadriiform related to the seedsnipe, as proposed by Olson & Steadman (1981) and supported by our DNA hybridization evidence. Turnix species begin to breed at less than one year of age, at least in captivity, and may have evolved at an exceptionally rapid rate. This may account, in part, for the large genetic distance between the buttonquails and other groups. Sibley et al. (1988) placed the “Family Turnicidae” in the “Infraclass?” and in “Order Turniciformes, inc. sedis”. Sibley & Ahlquist (1990: 255, 257) and Sibley & Monroe 114 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI (1990: 42) substituted the parvclass Turnicae for “Infraclass?” and deleted the “inc. sedis” after Order Turniciformes. The revised classification for this section follows. Class Aves Subclass Neornithes . Infraclass Eoaves Parvclass Ratitae Infraclass Neoaves Parvclass Galloanserae Parvclass Turnicae Order Turniciformes WOODPECKERS, HONEYGUIDES, BARBETS, AND TOUCANS These groups (Piciformes) produced some intriguing results. As expected, the wood¬ peckers and honeyguides (infraorder Picides) are closest relatives, but the toucans proved to be more closely related to the New World barbets than the New World barbets are to the Old World barbets. Thus, the toucans are specialized New World barbets, as indicated in the following classification: Infraorder Ramphastides Superfamily Megalaimoidea: Asian Barbets Superfamily Lybioidea: African Barbets Superfamily Ramphastoidea Family Ramphastidae Subfamily Capitoninae: New World Barbets Subfamily Ramphastinae: Toucans JACAMARS AND PUFFBIRDS The Galbuliformes often have been viewed as close relatives of the Piciformes be¬ cause they share several morphological characters. The DNA evidence indicates that these two groups are related, but that the divergence between them was ancient, hence we placed them in separate, but adjacent, parvclasses. The jacamars and puffbirds seem to be more closely related to the Coraciae (hornbills, hoopoes, trogons, rollers, motmots, etc.) than to the woodpeckers, honeyguides, barbets, and toucans. HORNBILLS, HOOPOES, TROGONS, ROLLERS, MOTMOTS, TODIES, KINGFISHERS, AND BEE-EATERS We assigned these groups to four orders: Bucerotiformes (hornbills); Upupiformes (hoopoes); Trogoniformes (trogons), and Coraciiformes (rollers, motmots, todies, king¬ fishers, bee-eaters) in the superorder Bucerotimorphae. They are morphologically diverse, but the DNA evidence indicates that they are more closely related to one another than any one of the orders is to another group. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 115 The kingfishers are morphologically similar, but the DNA comparisons revealed sub¬ stantial genealogical diversity. We divided them into two parvorders (Alcedinida; Cerylida) and the Cerylida into two superfamilies, each with one family. The family name Halcyonidae has priority over Dacelonidae (W. Bock, pers. comm.), thus our classification of the kingfishers is revised as follows: Suborder Alcedini Parvorder Alcedinida Family Alcedinidae: Alcedinid Kingfishers Parvorder Cerylida Superfamily Halcyonoidea (replaces Dacelonoidea) Family Flalcyonidae: Halcyonid Kingfishers (replaces Dacelonidae) Superfamily Ceryloidea Family Cerylidae: Cerylid Kingfishers MOUSEBIRDS OR COLIES The Mousebirds of Africa possess several unique characters and all attempts to ally them closely with another group have failed. The DNA evidence also shows that the Coliiformes have no close living relatives; they are the survivors of an ancient line¬ age. Their closest relatives are probably the groups that include the hornbills, rollers, kingfishers, and cuckoos. CUCKOOS, HOATZIN, AND TURACOS The cuckoos and turacos often have been associated in classifications but we found no convincing evidence of a close relationship between them. The cuckoos (Cuculiformes) proved to be remarkably diverse and we found it necessary to divide them into two infraorders, five parvorders, and six families — far different from the traditional assignment of all species to the Cuculidae. The cuckoos are another an¬ cient lineage in which morphological conservatism has obscured their genealogical diversity. Their next nearest living relatives are uncertain and we cannot identify a single group with confidence. The owls and nightjars may be the nearest living rela¬ tives of the cuckoos, but the divergence must have been so long ago that the idea of “close relatives” becomes irrelevant. It seems likely that the living cuckoos are about equally distant from several of the other non-passerine groups. The Floatzin Opisthocomus hoazin occurs in northern South America and has been a taxonomic puzzle since its discovery over 200 years ago. It was usually assigned to the Galliformes, perhaps because it somewhat resembles a chachalaca. The Floatzin feeds mainly on plants and has a large, muscular crop and other adaptations related to its diet. The DNA evidence is clear; the Floatzin is a highly specialized cuckoo, most closely related to the Guira Cuckoo, the anis Crotophaga, and the roadrunners, as indicated in the following arrangement of the infraorder Crotophagides. Parvorder Opisthocomida: Floatzin Parvorder Crotophagida: Anis and Guira Cuckoo Parvorder Neomorphida: Roadrunners, Ground Cuckoos 116 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI PARROTS Parrots are parrots. They seem to have no close living relatives and there is no doubt about what is and what is not, a parrot. They are the descendants of an ancient lin¬ eage that has maintained the characteristic structures of beak, feet, and plumage. Our limited data do not provide the basis for subdivisions of the Psittacidae, but it seems likely that such subunits exist, possibly in relation to major geographic regions of the world. SWIFTS AND HUMMINGBIRDS The swifts and hummingbirds have been associated in most classifications of the past 150 years. Our DNA comparisons, and morphological evidence, indicate that they are one another’s closest living relatives, but the divergence was ancient, possibly in the late Cretaceous. OWLS, NIGHTJARS, AND ALLIES (STRIGIFORMES) The owls and nightjars usually have been thought to be related and the DNA compari¬ sons agree, but the divergence was a long time ago. The owls and diurnal raptors (Falconiformes) are not closely related. The next nearest relatives of the Strigiformes may be the swifts and hummingbirds, but the evidence is not conclusive. The “nightjars and allies” include the owlet-nightjars, frogmouths, Oilbird, potoos, eared-nightjars, nighthawks, and whip-poor-wills. We recognize two suborders in the Strigiformes for the nightjars and, allies: Aegotheli for the owlet-nightjars ( Aegotheles ) and Caprimulgi for the others. Thus, contrary to most classifications, the DNA com¬ parisons indicate that the owlet-nightjars and frogmouths are not close relatives, al¬ though both occur in Australasia. The frogmouths are separated as the infraorder Podargides from the remainder of the Caprimulgi (infraorder Caprimulgides). The South American Oilbird ( Steatornis ) is most closely related to the Neotropical potoos ( Nyctibius ). The eared-nightjars ( Eurostopodus ) are distinct from the typical nightjars and we place them in the Eurostopodoidea as the sister taxon of the Caprimulgoidea, which includes the typical nightjars, nighthawks, and whip-poor-wills. Thus, like the parrots and kingfishers, the nightjars and allies have retained a simi¬ lar external morphology while diverging substantially at the genomic level. Their plum¬ age coloration is obviously correlated with their crepuscular and nocturnal habits, but it also obscures their diversity. PIGEONS AND DOVES Pigeons and doves occur on all continents and many islands. They share a common morphology and the group is certainly monophyletic. Their nearest relatives have been uncertain but the sandgrouse and parrots have often been suggested. The DNA comparisons support the monophyly of the Columbiformes, but show that the sandgrouse are related to the Charadriiformes and that the parrots are no closer to ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 117 the pigeons than to other groups. Thus, the Columbiformes seem to have no close living relatives. CRANES, BUSTARDS, SUNGREBES, TRUMPETERS, RAILS, AND ALLIES The Gruiformes are morphologically diverse and it would not have been surprising if the DNA comparisons had revealed at least equal degrees of genomic diversity, but the monophyly of most of the traditional members of the order is reflected in the data, although the differences among the subgroups are substantial. Two genera often in¬ cluded in the Gruiformes were misplaced there. The buttonquails (Turnicidae) are not members of this order; we have assigned them to the Turniciformes, noted above. The Plains-wanderer ( Pedionomus ) of Australia was transferred to the Charadriiformes by Olson & Steadman (1981) and the DNA data agree. The buttonquails and the Plains-wanderer were usually included as the only members of the Turnicidae in the traditional Gruiformes. The DNA evidence indicates that the Limpkin ( Aramus ) and the Neotropical Sungrebe ( Heliornis ) are closest relatives. The DNAs of the African and Asian finfoots were not available. SHOREBIRDS: SANDGROUSE, SANDPIPERS, PLOVERS, GULLS, ETC. In our classification the suborder Charadrii includes the traditional charadriiforms in the infraorder Charadriides and the sandgrouse in the infraorder Pteroclides. The clas¬ sification of the subgroups in the Charadriides was modified by the DNA evidence, but is congruent with other sources of evidence. We divide the Charadriides into the parvorders Scolopacida (seedsnipe, Plains-wanderer, snipe, sandpipers, phalaropes, painted-snipe, jacanas) and Charadriida (thick-knees, oystercatchers, avocets, stilts, plovers, Crab Plover, pratincoles, skuas, skimmers, gulls, terns, auks). The sandgrouse (Pteroclidae) have been the subject of a long debate; are they pi¬ geons or plovers? Sandgrouse occur in arid regions in Africa, southern Europe, and parts of Asia. The pigeons, plovers, and galliforms most often have been proposed as their closest relatives. The relationship to the shorebirds seems clear, but the di¬ vergence was ancient and the morphological characters provide the basis for argu¬ ments in favor of both pigeons and plovers as closest relatives. Sibley & Ahlquist (1990: 463-470) reviewed the history of the debate and the evidence for and against each hypothesis. Another debate involving the sandgrouse concerned the early report that the adults transport water to their nestlings by wetting the breast feathers from which the young birds suck the water (Meade-Waldo 1896). This was dismissed by several authors, but confirmed by Cade & Maclean (1967) who observed and filmed adult male sandgrouse transporting water in their specialized ventral feathers. HAWKS, EAGLES, OLD WORLD VULTURES, FALCONS, AND ALLIES The diurnal birds of prey provide ample material for debate and conjecture. Are the owls closely related to the hawks? Are the falcons closely related to the hawks? What are the relationships of the Secretary-bird, the Osprey, and the New World vultures? We concluded that (1) the owls are not closely related to the hawks; (2) the 118 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Secretary-bird and the Osprey are members of the parvorder Accipitrida, which in¬ cludes the hawks, eagles, and Old World vultures; (3) the falcons are members of the parvorder Falconida, the sister group of the Accipitrida; and (4) the New World vul¬ tures are most closely related to the storks, as suggested by Garrod (1873) and sup¬ ported by Ligon (1967). In effect, the New World vultures are carrion-eating storks. GREBES Most classifications have viewed grebes and loons as closest relatives, although they differ in many morphological characters and their similarities have been ascribed to convergence by some authors. The problem has been to identify the nearest relatives of each if they are not considered to be one another’s closest relatives. The DNA comparisons show that the grebes have no close living relatives and that the grebe lineage branched early from a common ancestry with several other groups in the or¬ der Ciconiiformes, suborder Ciconii, infraorder Ciconiides. The Ciconiides includes the grebes, tropicbirds, boobies, cormorants, herons, flamingos, New World vultures, storks, ibises, Shoebill, pelicans, frigatebirds, penguins, loons, and tubenoses. Within this assemblage the loons are most closely related to the penguins and tubenoses, as noted below under Albatrosses, Petrels, Penguins, Loons, and Frigatebirds. Thus, the loons are no closer to the grebes than are the members of several other groups of waterbirds. THE TOTIPALMATE SWIMMERS: TRADITIONAL ORDER PELECANIFORMES The traditional Pelecaniformes includes the pelicans, boobies, gannets, cormorants, anhingas, frigatebirds, and tropicbirds. These groups share several morphological characters, including the totipalmate foot, intraorbital salt gland, and lack of a brood patch. All but the tropicbirds have an obvious gular pouch, although that of the frigatebirds is of uncertain homology. The monophyly of the “Pelecaniformes” has seemed to be beyond doubt, but the DNA comparisons indicate (1) that the boobies, gannets, anhingas, and cormorants are closely related to one another; (2) that the tropicbirds are distant from the other pelecaniforms; (3) that the frigatebirds are most closely related to the tubenoses, penguins, and loons, and (4) that the pelicans are closest to the Shoebill Balaeniceps rex. Many ornithologists will reject at least some of these suggestions, but there is congruent evidence. Several studies have con¬ cluded that the tropicbirds and frigatebirds are the most distant from one another and from the pelicans, boobies, gannets, anhingas, and cormorants. An alliance between frigatebirds and tubenoses has been proposed before, and the Shoebill-pelican rela¬ tionship was suggested by Cottam (1957) from an anatomical study. This may be the most controversial question we have raised, but the polyphyly of the traditional order Pelecaniformes should be considered as an alternative hypothesis to be tested by independent studies of molecules and morphology. HERONS, FLAMINGOS, IBISES, STORKS, AND ALLIES The long-legged, long-necked wading birds usually have been placed in the order Ciconiiformes. In our classification they are members of the parvorder Ciconiida (in ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 119 the Ciconiiformes) which also includes the pelicans, New World vultures, frigatebirds, penguins, loons, and tubenoses (Procellariidae). The other members of the Ciconiida are the herons, Hamerkop {Scopus), flamingos, ibises, spoonbills, Shoebill, and storks. Most of these groups will elicit no surprise, but we have been accustomed to finding the tubenoses and penguins together at the beginning of the series of non¬ passerine groups where, in our classification, now reside the woodpeckers, barbets, rollers, and their allies. This suggests that, except for the anseriforms, the oldest volant non-passerines were terrestrial birds, not waterbirds. We concluded (Sibley & Ahlquist 1990: 527) that “the herons, Hamerkop, flamingos, ibises, Shoebill, pelicans, New World vultures, and storks are more closely related to one another than any one of them is to another group.” However, we noted the com¬ plications introduced by different ages at first breeding and the correlated differences in average genomic rates of evolution. We are not satisfied that our data are without error and additional experiments should be devised to test our conclusions. The final word, if ever uttered, is certain to be instructive. ALBATROSSES, PETRELS, PENGUINS, LOONS, AND FRIGATEBIRDS A relationship between the tubenoses (our Procellariidae = traditional Procellariiformes) and the penguins has long been accepted. The loons have some¬ times been placed with, or near, the tubenoses, and the frigatebirds share several morphological characters with petrels. These groups also differ and the large alba¬ trosses are among the taxa with the greatest ages at first breeding. Members of some populations of the Wandering Albatross may not breed until 15 years old, but most albatrosses begin to breed between 6 and 12 years of age. The effect of delayed maturity is to cause a slower rate of accession and drift of neutral alleles which con¬ stitute a substantial percentage of the genome and therefore have a major effect on the average rate of genomic evolution. This subject is discussed by Sibley & Ahlquist (1990: 165-183). We concluded that the tubenoses, penguins, loons, and probably the frigatebirds are more closely related to one another than any one of them is to another group. We also noted that the relationships of the frigatebirds require further study and that the dif¬ ferent average rates of genomic evolution among members of these groups compli¬ cate the interpretation of the data. The major groups in our classification of the Order Ciconiiformes are as follows: Order Ciconiiformes Suborder Charadrii: sandgrouse and traditional Charadriiformes. Suborder Ciconii Infraorder Falconides: Osprey, hawks, Secretary-bird, Old World vultures, falcons. Infraorder Ciconiides Parvorder Podicipedida: grebes. Parvorder Phaethontida: tropicbirds. Parvorder Sulida: boobies, gannets, anhingas, cormorants. Parvorder Ciconiida 120 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Superfamily Ardeoidea: herons, bitterns, egrets. Superfamily Scopoidea: Hamerkop or Hammerhead. Superfamily Phoenicopteroidea: flamingos. Superfamily Threskiornithoidea: ibises, spoonbills. Superfamily Pelecanoidea: Shoebill, pelicans. Superfamily Cic'onioidea: New World vultures, storks. Superfamily Procellarioidea: frigatebirds, penguins, loons, petrels, albatrosses. ORDER PASSERIFORMES: THE PASSERINE BIRDS This order includes 5712 (59%) of the 9672 species recognized by Sibley & Monroe (1990). Most species begin to breed at the age of one or two years and we have de¬ tected little or no effect on relative rates of genomic evolution of different ages at first breeding. The suboscine passerines (suborder Tyranni) are characterized by syringeal and other characters and the DNA data delineate essentially the same groups defined by morphology. The oscine passerines (suborder Passeri) were treated by Wetmore (1960) and other systematists as a linear series of families with little or no structure in the form of superfamilies and subfamilies. In most cases, superficial similarities were used to classify the oscines and the classifications concealed the genealogical diversity, adap¬ tive radiations, and other aspects of the evolution of the group. The DNA hybridiza¬ tion comparisons revealed a different picture of passerine phylogeny and provided the basis for a new classification. Perhaps the most interesting aspect of our phylogeny and classification is the recognition of the Australo-Papuan endemic radiation and its effects in other parts of the world. OLD WORLD SUBOSCINES New Zealand Wrens (Acanthisittidae). The acanthisittids seem to be the survivors of an ancient lineage with no close living relatives. We include them in the Tyranni be¬ cause they are not oscines. We place them in the infraorder Acanthisittides, but it is possible that they should be assigned to a third suborder. Pittas (Pittidae) and Broadbills (Eurylaimidae). The pittas and broadbills are placed in the infraorder Eurylaimides. We lacked DNAs of the philepittids of Madagascar. NEW WORLD SUBOSCINES The DNA comparisons show that the New World suboscines (infraorder Tyrannides) are more closely related to one another than any one of them is to any of the Old World suboscines. This agrees with some other studies and disagrees with the Wetmore (1960) classification. Tyrant Flycatchers and allies (Tyrannidae). In this group we include the typical tyrants (Tyranninae), mionectine flycatchers (Pipromorphinae), tityras and becards (Tityrinae), ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 121 cotingas (Cotinginae), and manakins (Piprinae). The Pipromorphinae has been substituted for “Mionectinae” and “Corythopinae” for a group of tyrants delineated by the DNA comparisons. Lanyon (1988) rejected our separation of the pipromorphine genera because syringeal characters are not congruent with the boundaries we set. There are other controversies concerning the details within the Tyrannida which are discussed by Sibley & Ahlquist (1990: 590-597). We conclude that the Tyrannidae, as defined above, is a monophyletic cluster representing an adaptive radiation composed primarily of insectivores and frugivores. The Neotropical antbirds were shown to be composed of two groups (typical antbirds and ground antbirds) on the basis of sternal notches (Heimerdinger & Ames 1967) and syringeal characters (Ames 1971). The DNA data delineated the same clusters and also revealed that the typical antbirds (Thamnophilidae) may be separated in the parvorder Thamnophilida, distinct from the Furnariida which includes the ovenbirds, woodcreepers, ground antbirds, gnateaters, and tapaculos. The New World suboscines are the descendants of an adaptive radiation that oc¬ curred while South America was isolated from other continents during the Tertiary. OSCINES OR SONGBIRDS: SUBORDER PASSERI (PASSERES) The complex syringeal musculature and other characters define the Passeri. In our classification the Passeri contains 4561 species in 870 genera, thus by far the larg¬ est suborder of living birds. The DNA hybridization data made it possible to subdivide this group into two parvorders, each composed of three superfamilies, and to recog¬ nize 35 (or 36) families. Other classifications have usually recognized more families (36-91), but several of our subfamilies are equivalent in content to the families of other classifications. Our classification of the oscines follows. Tribes are not indicated. Changes from Sibley et al. (1988) are indicated by an asterisk*. Parvorder Corvida Superfamily Menuroidea Family Climacteridae: Australo-Papuan tree-creepers. Family Menuridae Subfamily Menurinae: lyrebirds. Subfamily Atrichornithinae: scrub-birds. Family Ptilonorhynchidae: bowerbirds. Superfamily Meliphagoidea Family Maluridae Subfamily Malurinae: fairywrens, emuwrens. Subfamily Amytornithinae: grasswrens. Family Meliphagidae: honeyeaters. Family Pardalotidae Subfamily Pardalotinae: pardalotes. Subfamily Dasyornithinae: bristlebirds. Subfamily Acanthizinae: scrubwrens, thornbills, whitefaces. Superfamily Corvoidea ‘Family Petroicidae: Australo-Papuan robins. (Bock 1990: 629 notes that Petroicidae, not Eopsaltriidae, is the correct name for this group.) Family Irenidae: fairy-bluebirds, leafbirds. 122 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Family Orthonychidae: Logrunner, Chowchilla. Family Pomatostomidae: Australo-Papuan babblers. Family Laniidae: true shrikes = Lanius, Corvinella, Eurocephalus. Family Vireonidae: vireos, greenlets, peppershrikes, shrike-vireos. Family Corvidae Subfamily Cinclosomatinae: quail-thrushes, whipbirds. Subfamily Corcoracinae: White-winged Chough, Apostlebird. Subfamily Pachycephalinae: sittellas, Mohoua (incl. Finschia), shrike-tits, Oreoica, Rhagologus, whistlers, shrike-thrushes. Subfamily Corvinae: crows, jays, magpies, birds-of-paradise, Melampitta, currawongs, wood-swallows, Bornean Bristlehead, Peltops, orioles, cuckoo-shrikes. Subfamily Dicrurinae: fantails, drongos, monarchs, magpie-larks. Subfamily Aegithininae: ioras. Subfamily Malaconotinae: bushshrikes, helmetshrikes, Batis, Platysteira, vangas. Family Callaeatidae inc. sedis: New Zealand wattlebirds. ‘Family Picathartidae inc. sedis: Picathartes, Chaetops. (Sibley & Ahlquist 1990:625). Parvorder Passerida Superfamily Muscicapoidea Family Bombycillidae: waxwings, silky flycatchers, Palm Chat. Family Cinclidae: dippers. Family Muscicapidae Subfamily Turdinae: thrushes. Subfamily Muscicapinae: Old World flycatchers, chats ( Erithacus , Saxicola, et al.). Family Sturnidae: starlings, mynas, mockingbirds, thrashers, American catbirds. Superfamily Sylvioidea Family Sittidae Subfamily Sittinae: nuthatches. ‘Subfamily Tichodrominae: Wallcreeper. (Not Tichodromadinae). / Family Certhiidae Subfamily Certhiinae: tree-creepers, Spotted Creeper (Salpornis). Subfamily Troglodytinae: wrens. Subfamily Polioptilinae: gnatcatchers, gnatwrens, Verdin. Family Paridae Subfamily Remizinae: penduline-tits. Subfamily Parinae: titmice, chickadees. Family Aegithalidae: long-tailed tits, bushtits. Family Hirundinidae Subfamily Pseudochelidoninae: river-martins. Subfamily Hirundininae: swallows, martins. Family Regulidae: kinglets, goldcrests. Family Pycnonotidae: bulbuls, greenbuls. Family Hypocoliidae: Grey Hypocolius {inc. sedis). Family Cisticol idae : African warblers ( Cisticola , Prinia, Apalis, Camaroptera, et al.). Family Zosteropidae: white-eyes, silvereyes, Cleptornis). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 123 Family Sylviidae *Subfamily Acrocephalinae: leaf warblers. (Phylloscopinae of Sibley et al. 1 988). Subfamily Megalurinae: grass warblers, incl. “ Bowdleria ” = Megalurus. Subfamily Garrulacinae: Garrulax. Subfamily Sylviinae: babblers, Wrentit, Sylvia. Superfamily Passeroidea Family Alaudidae: larks. Family Nectariniidae Subfamily Promeropinae: African sugarbirds. Subfamily Nectariniinae: flowerpeckers, sunbirds, spiderhunters. Family Melanocharitidae: Melanocharis berrypeckers, longbills. Family Paramythiidae: Crested Berrypecker, Tit Berrypecker. Family Passeridae Subfamily Passerinae: sparrows, Petronia, snowfinches. Subfamily Motacillinae: wagtails, pipits. Subfamily Prunellinae: accentors, Dunnock. Subfamily Ploceinae: weaverbirds. Subfamily Estrildinae: waxbills, indigobirds, whydahs. Family Fringillidae Subfamily Peucedraminae: Olive Warbler. Subfamily Fringillinae: chaffinches, goldfinches, crossbills, Hawaiian honeycreepers. Subfamily Emberizinae: buntings, longspurs, towhees, wood warblers, tanagers (incl. Neotropical honeycreepers, Swallow-Tanager, Plushcap, tanager-finches), cardinals, troupials, American blackbirds, et al. DISCUSSION Most of the Corvoidea occur only or mainly in Australia and/or New Guinea. The ex¬ ceptions are the Irenidae, Laniidae, Vireonidae, crows, jays, magpies, orioles, cuckooshrikes, drongos, monarchs, ioras, bushshrikes, helmetshrikes, Batis, Platysteira, and the vangas. Most species of wood-swallows are endemic to Australia and New Guinea, but some species occur in southern and southeastern Asia, and on Southwest Pacific islands. Mohoua and Finschia are endemic to New Zealand; whis¬ tlers and honeyeaters occur on many South Pacific islands. The bowerbirds and the birds-of-paradise usually have been treated as closely related groups, sometimes placed in the same family or subfamily. The DNA hybridization comparisons show that the bowerbirds are closest to the lyrebirds and scrub-birds (Menuroidea); the birds-of-paradise are closest to the currawongs, orioles, and corvines (Corvoidea). The division of the Passeri into the Corvida and Passerida is correlated with the mor¬ phology of the head of the humerus. In the Corvida the tricipital fossa is single and pneumatic, that is, there is an opening from the fossa into the hollow shaft of the humerus that forms a connection from the air sac system. About 90% of the species of the Passerida have two fossae and there is no opening into the shaft of the hu¬ merus. A few species (waxwings, for example) are intermediate between the two 124 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI extremes. These conditions are randomly distributed in relation to the Wetmore (1960) classification. This subject is discussed by Sibley & Ahlquist (1990: 571, 626, 629, 633, 639). The African genera Picathastes (rockfowl) and Chaetops (rock-jumpers) have corvoid humeral fossae and the DNA hybridization evidence suggests that they may be clos¬ est relatives. Sibley & Ahlquist (1990: 625-627) placed the Picathartidae in the limbo of inc. sedis because our comparisons were incomplete. The Corvoidea includes a disproportionate number of species that are co-operative (or communal) breeders. Russell (1989) noted that 68 co-operative breeding species occur in Australia, of which 58 are passerines. All belong to the old endemic families of the corvoid early Australian radiation. Thus, of 258 species of old endemics, at least 22% are co-operative breeders, compared with a world-wide incidence of ca. 3%. This correlation is only with the Sibley et al. (1988) classification; it does not hold with other classifications. Russell (1989) suggested that the relationship between Australian endemics and co-operative breeding evolved as a response to climatic and other environmental influences that favored this pattern of reproduction. The old endemics of Australia also tend to lay smaller clutches and to live longer than passerines in other parts of the world. A comparison between the groups in our classification and the families of Wetmore’s (1960) classification reveals many differences, including the treatment of the nectarivorous oscines. In Wetmore’s classification, the honeyeaters, sunbirds, flowerpeckers, and white-eyes were listed in sequence. Wetmore viewed the se¬ quence of groups as being on a scale of increasing specialization from the larks and swallows at the beginning of the series of oscine families to the New World nine- primaried groups at the end. By placing the nectarivorous families in sequence he implied a closer relationship among them. The DNA hybridization evidence has shown that the meliphagids are corvoids, the nectariniids are passeroids, and the zosteropids are sylvioids. The Dicaeidae of Wetmore (and others) included the pardalotes of Aus¬ tralia and the flowerpeckers. The DNA hybridization comparisons show that the pardalotes are corvoids and the flowerpeckers are nectariniids, hence passeroids. The African sugarbirds ( Promerops ) have been assigned to several groups, including the Meliphagidae, Nectariniidae, Sturnidae, and Turdidae. The DNA comparisons show that the sugarbirds are specialized sunbirds and we place them in the Promeropinae of the Nectariniidae. The vireonids usually have been placed near the nine-primaried wood warblers (Emberizinae: Parulini) because they are small, greenish or yellowish insectivores with a tendency to have a reduced 10th primary. The DNA evidence allies them with the Australian endemics in the Corvoidea. The ancestral vireonid may have arrived in South America via Antarctica when Antarctica had a temperate climate. The starlings have been allied with the corvines in many classifications. Wetmore placed them between the New Zealand wattlebirds and the honeyeaters. The DNA comparisons showed that the starlings are most closely related to the New World mockingbirds and thrashers (“Mimidae”) and we have included these groups as tribes (Sturnini, Mimini) in the family Sturnidae. This conclusion has been one of the most ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 125 difficult for many ornithologists to accept, but it is based on some of the best data in our study and it is supported by serological and morphological evidence. In addition, starlings and muscicapids are the only oscines that lack the enzyme sucrase (Martinez del Rio 1990). The several types of “creepers” often have been placed together. Wetmore included the Northern creepers ( Certhia ) and Australo-Papuan treecreepers ( Climacteris , Cormobates) in the Certhiidae, but the DNA data show that the Climacteridae is a menuroid group and Certhia is a sylvioid genus most closely related to the wrens, gnatcatchers, gnatwrens, and Verdin. in some classifications the muscicapine flycatchers, thrushes, and sylviine warblers have been placed in the same family or in adjacent families. Other classifications have included the thrushes, babblers, mockingbirds, wrens, dippers, and accentors as sub¬ families in the same family. Hartert (1910) included the muscicapine flycatchers, monarchs, sylviine warblers, babblers, and thrushes in his family Muscicapidae. This arrangement, usually called the “Primitive Insect Eaters,” has been adopted by sev¬ eral subsequent authors. Hartert’s Muscicapidae was a polyphyletic assemblage that included members of both of our parvorders and most of the superfamilies in the Passeri. Some classifications have included members of all six of our oscine superfamilies in the “Muscicapidae” when the Australo-Papuan treecreepers (Climacteridae) have been included in the Certhiidae. “The convergently similar members of the Australo-Papuan Corvida and the Afro- Eurasian Passerida have presented the most difficult problems in the classification of the muscicapoid and sylvioid Passeri. The association of superficially similar but ge¬ netically unrelated ecotypes in such polyphyletic taxa as the “Muscicapidae” of many classifications has obscured the zoogeographic and phylogenetic histories of many oscine taxa. The result is a classical example of the difficulties encountered in using morphological characters to determine the boundaries of monophyletic clusters of convergently similar species.” (Sibley & Ahlquist 1990: 634). Other examples are discussed by Sibley & Ahlquist (1990) and indicated in the clas¬ sification of Sibley et al. (1988). ACKNOWLEDGMENTS Jon Ahlquist shared equally in the research at Yale University that produced the data and publications on which this synopsis is based. Burt Monroe contributed his exper¬ tise to the classification (Sibley et al. 1988) and brought our book on the distribution and taxonomy of birds of the world (1990) to completion. The more than 300 persons who helped us in so many ways are listed in the Acknowledgments in Sibley & Ahlquist (1990) and Sibley & Monroe (1990). We thank them again. LITERATURE CITED AMES, P.L. 1971. The morphology of the syrinx in passerine birds. Bulletin of the Yale University Peabody Museum of Natural History 37: 1-194. 126 ACTA XX CONGRESSUS INTERN ATIONALIS ORNITHOLOGICI BOCK, W.J. 1990. A special review: Peter’s “Check-list of Birds of the World” and a history of avian checklists. Auk 107: 629-648. CADE, T.J., MacLEAN, G.L. 1967. Transport of water by adult sandgrouse to their young. Condor 69: 323-343. COTTAM, P.A. 1957. The pelecaniform characters of the skeleton of the Shoebill Stork, Balaeniceps rex. Bulletin of the British Museum (Natural History) Zoology 5: 49-72. FURBRINGER, M. 1888. Untersuchungen zur Morphologie und Systematik der Vogel. Vols. 1, 2. Amsterdam, Von Holkema. GADOW, H. 1892. On the classification of birds. Proceedings of the Zoological Society of London 1892: 229-256. GADOW, H. 1893. Vogel. II. Systematischer Theil. In Bronn’s Klassen und Ordnungen des Thier- Reichs. Vol. 6(4), 303 pp. Leipzig, C. F. Winter. GARROD, A.H. 1873. On certain muscles of the thigh of birds and on their value in classification. Part I. Proceedings of the Zoological Society of London 1873: 626-644. HARTERT, E. 1910. Die Vogel der Palaarktischen Fauna. Berlin, R. Friedlander. HEIMERDINGER, M.A., AMES, P.L. 1967. Variation in the sternal notches of suboscine passerine birds. Postilla 105: 1-44. LANYON, W.E. 1988. A phylogeny of the thirty-two genera in the Elaenia assemblage of tyrant flycatch¬ ers. American Museum Novitates No. 2914: 1-57. LIGON, J.D. 1967. Relationships of the cathartid vultures. Occasional Papers, University of Michigan Museum of Zoology 651: 1-26. MARTINEZ DEL RIO, C. 1990. Dietary and phylogenetic correlates of intestinal sucrase and maltase activity in birds. Physiological Zoology (in press). MEADE-WALDO, E.G.B. 1896. Sand grouse breeding in captivity. Zoologist, 3rd series, 20: 298-299. OLSON, S.L., STEADMAN, D.W. 1981. The relationships of the Pedionomidae (Aves: Charadriiformes). Smithsonian Contributions in Zoology No. 337. RUSSELL, E.M. 1989. Co-operative breeding — a Gondwanan perspective. Emu 89: 61-62. SIBLEY, C.G., AHLQUIST, J.E. 1983. Phylogeny and classification of birds based on the data of DNA- DNA hybridization. Current Ornithology 1: 245-292. New York, Plenum Publ. Corp. SIBLEY, C.G., AHLQUIST, J.E. 1986. Reconstructing bird phylogeny by comparing DNA’s. Scientific American 254 (2): 82-92. SIBLEY, C.G., AHLQUIST, J.E. 1987. Avian phylogeny reconstructed from comparisons of the genetic material, DNA. Pp. 95-121 in Patterson, C. (Ed.). Molecules and morphology in evolution: conflict or compromise? London, Cambridge University Press. SIBLEY, C.G., AHLQUIST, J.E. 1990. Phylogeny and classification of birds. A study in molecular evo¬ lution. 976 pages. New Haven, Yale University Press. SIBLEY, C.G., MONROE, B.L., JR. 1990. Distribution and taxonomy of birds of the world. 1111 pages. New Haven, Yale University Press. SIBLEY, C.G., AHLQUIST, J.E., MONROE, B.L., JR. 1988. A classification of the living birds of the world based on DNA-DNA hybridization studies. Auk 105: 409-423. STRESEMANN, E. 1934. Aves. Handbuch der Zoologie, vol. 7, part 2. Kukenthal, W. & Krumbach, T. (Eds). Berlin, Walter de Gruyter. WETMORE, A. 1930. A systematic classification for the birds of the world. Proceedings of the United States National Museum 76(24): 1-8. WETMORE, A. 1960. A classification for the birds of the world. Smithsonian Miscellaneous Collections 139(1 1 ) : 1 -37. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 127 PLENARY LECTURE AN ORNITHOLOGICAL GLIMPSE INTO NEW ZEALAND’S PRE-HUMAN PAST I. A. E. ATKINSON and P. R. MILLENER ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 129 AN ORNITHOLOGICAL GLIMPSE INTO NEW ZEALAND’S PRE-HUMAN PAST I. A. E. ATKINSON1 and P. R. MILLENER2 1 DSIR Land Resources, Private Bag, Lower Hutt, New Zealand 2 National Museum of New Zealand, P O Box 467, Wellington, New Zealand ABSTRACT. Data from subfossil birds, and current and past vegetation, are used to reconstruct four lowland and coastal forest-bird systems of pre-human New Zealand. These systems had relatively sim¬ ple trophic structures: 67 bird species used the forests, distributed between 10 feeding guilds; 33% of the species were flightless and 21% were nocturnal or semi-nocturnal. Feeding level, feeding mode, feeding site and foods eaten, differentiated the guilds and facilitated coexistence of species within guilds. Differences in guild composition between habitats were largely a consequence of biogeographic history. The ground herbivore, arboreal herbivore, and subsurface-feeding group of the ground insec- tivore guilds are considered to have no counterparts elsewhere. Some 40% or more of the bird spe¬ cies present originally in these systems are extinct. There are limited opportunities to partly restore some systems which, if taken, may increase chances of survival for remaining members of the avifauna. Keywords: trophic structure, guilds, resource partitioning, forest birds, feeding behaviour, species co¬ existence, ecological restoration, coastal forest, competition, extinctions, flightlessness, lowland for¬ est, moas, nocturnal activity, subfossil faunas, weights. “The past of systems influences their present behaviour and represents an often unexplained source of variance in the relationship between current en¬ vironment and current process”. Peter S. White 1 990 INTRODUCTION When humans first stepped ashore in New Zealand a thousand or more years ago, they encountered one of the most remarkable plant-animal communities in the world. The largely forested landscape was dominated by birds. The absence of mammals, apart from a few species of bat, would not have been surprising to these Polynesian people given the nature of the islands from which they sailed. What was new was the large size and abundance of flightless, herbivorous birds called moa providing a plen¬ tiful supply of food. To an ornithologist, moas would not have been the only unusual feature of this pris¬ tine terrestrial community. Other species, some of them flying birds, were also her¬ bivorous, an uncommon behaviour among birds, and these coexisted with various other frugivorous, nectivorous, insectivorous, omnivorous and raptorial birds. Flightlessness or reduced flight capacity was found in birds of all sizes and few spe¬ cies were specialist feeders. Most groups showed little radiation although rails, acanthisittid wrens, and moas were notable exceptions. 130 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI What kinds of food resources were available to these birds and how was food parti¬ tioned between species? To what extent had birds in New Zealand adopted the eco¬ logical role of mammals in plant-animal systems of the continents? Partial answers to these questions are now possible as a result of both new finds of subfossil birds and recent studies of living birds. The most detailed information relates to low-altitude environments and, since forest covered more than 80% of the land area during the few thousand years immediately preceding human arrival (McGlone 1989), this study is focused on lowland and coastal forest. Reconstructions of four examples of these forests are used as the basis for a preliminary analysis of avian trophic structure in such habitats as it might have been in pre-human times. CHARACTERISTICS OF LOWLAND AND COASTAL FOREST IN PRE-HUMAN NEW ZEALAND The greater part of this forest consisted of a mixture of tall long-lived southern coni¬ fers and shorter-lived hardwoods of variable height. Key characteristics of this com¬ munity are summarised in Table 1. TABLE 1 - Characteristics of New Zealand lowland conifer/hardwood forest. Characteristic Degree of development References High level of endemism in both plants and animals Angiosperms (85%); gymnosperms (100%); ferns (45%); landbirds (34%); reptiles (100%); amphibians (100%); butterflies and moths (90%) Godley 1975; Brownsey & Smith-Dodsworth 1989; C.J.R. Robertson 1985; Daugherty et al. 1990 Low species diversity in most plant and animal groups c. 36 significant tree species forming canopy; seldom more then 5-10 important at any one site; forest birds (c. 67 spp); reptiles (c. 22 spp ); amphibians (7 spp.); invertebrates (unknown but beetles very diverse) I.A.E. Atkinson (unpub); D.R. Towns (pers. comm.); Worthy 1987 Plant species predominantly evergreen Leaves rather dark green, of medium to small size, most frequently glabrous, entire with glossy upper surface Canopy of long-lived small-leaved conifers without cones mixed with shorter lived hardwoods frequently overtopped by the conifers Life span of conifers commonly 500-1000 years, sometimes >1000 years, usually 25-40 m tall. Cones replaced by fleshy arils supporting the ‘seeds’. Life span of hardwoods commonly 100- 450 years although some species live longer. / Tree ferns and ground ferns Often prominent in the understorey and on the ground, particularly in wetter environments Epiphytes and lianes Prominent at all levels in the forest, particularly in wetter environments Smaller areas of lowland forest dominated either by southern beeches ( Nothofagus spp.), which are of major importance in the montane zone, or kauri Agathis australis, which occurs in a mosaic pattern with southern conifer/hardwood forest in northern North Island are excluded. Both these forests occur with soils of low fertility. New Zealand forests have low species diversity. Many important tree genera of Aus¬ tralia, such as Eucalyptus and Acacia, are not indigenous to New Zealand. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 131 FOOD SOURCES FOR BIRDS WITHIN LOWLAND CONIFER/HARDWOOD FOREST Woody plants In addition to leaves and buds, up to 20% of the species present in any one site may produce nectar. There are very few species with ornithophilous flowers. Unlike the southern beech forests, production of honeydew excreted by bark inhabiting scale in¬ sects is only of minor importance in conifer/hardwood forest. Manna and lerp (Paton 1980) have not been investigated as food sources for birds in New Zealand. Up to 60% of the species at a site may produce fleshy fruit. Fruit are commonly red in colour although significant proportions are black or white (Lee et al. 1991). Few species have large fruit with none exceeding 20 mm diameter or 40 mm length; most fruit are less than 10 mm in diameter (Lee et al. 1991). The availability of nectar and fleshy fruit varies greatly from place to place depending on species composition and natural annual fluctuations in production. Herbaceous plants Ferns, sedges, lilies and dicotyledonous herbs are all common in conifer/hardwood forests but most are perennial and few species appear to be of high nutrient value. Grasses, including some species of tussock grass, are less common; they may be a useful source of nutrients. Mosses, liverworts, lichens and fungi These groups of plants are all abundant in conifer/hardwood forest; both fungi and lichens are potential food sources for vertebrates. Invertebrates The invertebrate life of lowland forest was once characterised by an abundance and variety of large (> 20 mm) flightless insects occupying habitats from the forest floor up into the canopy. These included very large nocturnal wingless crickets called wetas in the Stenopelmatidae (giant wetas, tree wetas and ground wetas) and Rhaphidophoridae (cave wetas). More than half the known insect fauna of New Zea¬ land are beetles and this group was also very common in forest, particularly the flight¬ less ground beetles and large flightless weevils. Another formerly abundant food source that may have been eaten by birds are veined slugs of the family Arthorocophoridae. These live in vegetation, decaying logs and leaf mould and may reach 150 mm in length (Burton 1962, 1963). A key source of food for many insectivorous birds is the invertebrate life in the leaf and branch litter of the forest floor. This fauna was formerly characterised by high densities of amphipods, litter-feeding caterpillars of moths, and large (up to 20 mm width) pill millipedes (Watt 1975). Where lime was not limiting there were populations of very large (up to 100 mm diam.) carnivorous molluscs which were specialised to capture and eat large subsoil earthworms, members of an extensive annelid worm fauna that occurs in leaf mould (36 spp), topsoil (48 spp), as well as in the subsoil (Lee 1959). Many invertebrates generally associated with the forest floor are also available to arboreal insectivores. Perched leaf litters among epiphytes and in tree cavities pro¬ vide important habitat for many invertebrates such as caterpillars, ants, amphipods, 132 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI ostracods, isopods and earthworms that are otherwise associated with the forest floor (Moeed & Meads 1983). The numbers and variety of all these groups of animals have been greatly reduced since the advent of mammalian predators in New Zealand particularly rats and mice. Vertebrates Evidence from cave deposits and rat-free islands indicates that skinks and geckos were originally very abundant in lowland and coastal forests (Whitaker 1973, Daugherty et al. 1990). Nocturnal and diurnal skinks were largely restricted to the ground but some geckos foraged arboreally. Tuatara ( Sphenodon spp.) were also widespread on the mainland and may have been eaten by some bird predators. Subfossil evidence and island distribution of living species of leiopelmatid frogs sug¬ gest that these also were a potential food, particularly since they lack a free-living tadpole stage. Three species of small nocturnal bats were originally widespread in lowland forests and roosted in caves or hollow trees (Daniel & Williams 1984, Hill & Daniel 1985). Birds themselves provided prey for raptors. METHODS USED IN HABITAT RECONSTRUCTIONS Habitat reconstructions were attempted for two examples each of lowland and coastal forest. These were warm temperate humid conifer/hardwood and cool temperate dry conifer/hardwood forests, and warm temperate and cool temperate coastal forests. Classification of thermal regimes follows Meurk (1984). Major plant species likely to have been present were derived from knowledge of surviving forest remnants as well as identifications of charcoal and fossil wood, in each habitat a list of the birds for¬ merly present, both extinct and extant, was derived primarily from subfossil bone deposits accumulated in caves, sand dunes or alkaline swamps. The list includes birds considered to have made some major use of these forests even if breeding in other habitats. Additional records were derived from current distribution data (Bull et ai. 1985) but several species now present in particular habitats were excluded be¬ cause of evidence that they may not have been present 1000 years ago. A significant number of New Zealand birds have only established since human modification of the landscape began. The trophic position of each species was assessed from feeding habits considered in the light of body weight which is correlated with many life history parameters (West¬ ern 1979, Peters 1983). Data on feeding habits were derived from published studies and the authors’ observations. Feeding habits of extinct birds were inferred where possible from bill structure together with the feeding behaviour of related extant birds where such existed. The observed feeding patterns of some extant species may have changed in response to altered competitive relationships following species extinctions and establishment of some introduced species. For this reason the analysis is restricted to major differ¬ ences in trophic behaviour between species. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 133 TABLE 2 - Classification of avian feeding guilds in New Zealand forest. Food source Feeding level Guild1 Foliage, twigs ground ground herbivores arboreal arboreal herbivores Fruit, buds, foliage ground arboreal | frugivore/herbivores Nectar, honeydew, fruit arboreal nectivores Invertebrates aquatic ground/subsurface arboreal aquatic insectivores ground insectivores arboreal insectivores Vertebrates all levels major predators (of vertebrates) Carrion ground/arboreal carrion feeders ’Guild classification modified from Lein (1972) and Terborgh et a). (1990). TABLE 3 - Methods of foraging by New Zealand forest-inhabiting birds. Foraging method Description Grazing Browsing/plucking Crushing Husking Grinding Lapping Gleaning1 Probing/prising1 Stripping Hovering’ Snatching’ Pouncing’ Hawking’ Flushing Raking Gauging Digging Foliage cut or pulled at or near ground level. Plants parts, including fruit, cut or pulled from vegetation, often woody. Use of bill to crush seeds before ingestion. Use of bill to husk seeds from enclosing structures or extract seeds from seed capsules. Use of mandible to grind fibrous foods against maxilla and extract sap or other juices. Use of tongue to drink nectar, honeydew or sap exudate, or pick up pollen. Picking up stationary food items by a standing or hopping bird. Use of bill to penetrate or lift the substrate to locate concealed food. Use of bill to peel or strip bark to locate concealed food. Picking up food while the bird hovers. Food plucked from the substrate by the bird as it flies past. Food taken from the substrate by a bird flying from a perch. Bird sallies into the air to catch flying prey. Disturbance of prey by bobbing action with whole body, rapid opening and closing of wings, spreading of tail, or vibration of foot. Use of feet or bill to scatter loose material such as leaf litter and expose concealed food. Use of bill to cut linear incisions in bark to reach cambium and release sap. Use of bill to excavate holes in decaying or live wood, or in soil, to expose concealed food. ’After Holmes & Recher (1986). Body weights of extant birds were taken from publications or unpublished data. The weights of extinct birds were estimated either from allometric relationships between femur dimensions and body weight or, for a very few species, assessed by compari¬ son with similar-sized living birds (Appendix 1). Trophic relationships were described by means of feeding guilds (Table 2), i.e. sets of birds deriving their subsistence from common pools of resources and thus co-ex¬ isting in the same habitat (Terborgh & Robinson 1986). Although the guild concept is useful for comparisons, the diversity of food sources used by some birds precluded any simple analysis of avian trophic structure. Where a species used different food sources between the breeding and non-breeding season, both sources were used to determine guild position. 134 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI TABLE 4 - Stratification of feeding levels used by New Zealand forest inhabiting birds. Feeding level Description of level Height zone (m) Canopy The uppermost storey of tree (or shrub) crowns, regardless of height, unshaded by other crowns. 5-40+ Upper understorey Plant and foliage shaded by canopy foliage. >2 (Subcanopy) (In taller forest only : the uppermost part of the upper understorey where such distinction is useful). >10 Lower understorey Plant crowns and foliage shaded by canopy foliage. 0.5-2 Ground storey Plants with growing points < 0.5 m above ground. variable, not necessarily always < 0.5 m Ground The ground surface including the litter. 0.0 Subsurface Feeding levels below the ground surface. variable Aquatic Feeding levels on or below a water surface. variable Unless indicated, nomenclature for bird species follows Turbott (1990) although subspecific epithets are excluded from the tables. Only major foods taken are tabu¬ lated. Foraging methods are categorised in Table 3. Classification of feeding levels (Table 4) extends the system used by Atkinson (1966a). In the guild tables, feeding levels are listed in decreasing frequency of use and emphasise feeding rather than other activities. The four habitat reconstructions of the tables correspond with those described under l-IV in the text. Where two weights are given, male weights precede those of female. I. WARM-TEMPERATE HUMID CONIFER/HARDWOOD FOREST ON KARST TERRAIN Site description This region of karst landscape on limestone is south-west of Hamilton, North Island. Its western boundary lies 8 km or more inland from the west coast and to the east it is bounded by the Waitomo Caves; northern and southern limits are at Raglan and Awakino respectively (Figure 1). In these Oligocene limestones (Kear 1960), karst features are prominent with numerous broken ridges and steep bluffs separated by wide intricately gullied basins with sink-holes, underground streams and caves. Al¬ though Jurassic siltstones and Pliocene Pleistocene andesites are present in the gen¬ eral area, this habitat reconstruction relates to limestone and will be referred to as the “Waitomo” region. This kind of landscape varies between 100 and 400 m altitude with a mean annual temperature of 12.5 to 15°C and annual rainfall from 2000 to 2500 mm. Soils are weakly or moderately leached central yellow-brown loams that are not particularly fertile as they have been formed from a subaerially deposited layer of andesitic vol¬ canic ash, Mairoa Ash (Orbell 1974) overlying the limestone to depths of 1 m or more (D. Hicks, pers comm). Soil pH is 5.8 to 6.0 and both base saturation and available phosphorus are low (New Zealand Soil Bureau 1954). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 135 Nature and age of the subfossil material. The subfossil bird bones recovered are from 37 caves that together formed one of two major study areas described and analysed by Millener (1981a). These originate mainly from pitfall deposits but include some from underground stream deposits derived from larger catchments. Pitfalls may introduce a bias of unknown size towards flightless birds. The caves themselves are much older than the deposits they contain: radiocar¬ bon dates from samples of avian bone collagen collected from 10 caves span an age range of 1075 ± 75 yr BP to 24,800 ± 500 yr BP. This age range extends into Otiran time and raises the question of whether the cave- deposited bones are derived only from animals living in forest that grew in a humid warm-temperate climate? A botanical survey of parts of Tawarau Forest, immediately west of Waitomo revealed a number of “cool-climate” plant species associated with matai/kamahi1 forest and Olearia virgata scrub, species that are not present in the widespread rimu/tawa forest now present in the generally mild climate of the region (Ogle & Druce 1987). Although some caves may have been available for trapping of cool-climate animals during Otiran times, many features of the deposits demonstrate that the animals rep¬ resented are those of humid forest, frequently forest associated with a warm-temper¬ ate climate. These features are: (i) The abundance of bones of moa species such as Pachyornis mappini, Euryapteryx curtus and E. geranoides, all recognised as lowland species (Atkinson & Greenwood 1989, Worthy 1990). 1 Scientific names of plants are given in the tables. 136 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI (ii) The common and relatively abundant bones of Anomalopteryx didiformis, a low¬ land moa which is associated with deposits of Holocene rather than Otiran age (Worthy 1990), and which was recovered from 31 of the 37 caves examined. (iii) The abundance of birds.such as Weka, New Zealand Pigeon, Kaka, Red-crowned parakeet, Tui and Saddleback, all commonly associated with lowland forest. (iv) The widespread occurrence of shells of many landsnail species obligately de¬ pendent on humid forest environments. (v) The absence or rarity of birds such as Harrier Circus approximans, Pipit Anthus n. novaeseelandiae and New Zealand quail Coturnix n. novaezelandiae which would be expected in more open habitats with cool-climate vegetation. A humid lowland forest thus dominated the area throughout the period when faunal remains were accumulating in the caves. This fauna could not have been living in scrub communities dominated by Olearia virgata, manuka Leptospermum scoparium or other species, but could have been derived in part from matai/kamahi forest of the kind described by Ogle and Druce (1987). The dominance of lowland forest animal species in the cave deposits also suggests that most of these deposits are Holocene in age. An alternative explanation is that the area remained largely under humid forest throughout the last 25 000 years, even though pollen evidence suggests that much of the Waikato lowlands to the north-west of the Waitomo study area may have been unforested between c. 18000 and 14000 yr BP (Newnham et al. 1989). In listing the major plant and bird species for this reconstruction, we have included only those we consider to have been present in the Waitomo district throughout the period c.6000 to 1000 yr BP, whether or not they may have been present during ear¬ lier periods. Vegetation Rimu/tawa forest as described by Nicholls (1980) and Ogle and Druce (1987) is prob¬ ably similar to the plant cover of the 6000-1000 yr BP period. This forest type covered much of the area prior to the arrival of Europeans last century. Plant species likely to have contributed the greatest proportion of biomass to the community are listed in Table 5. The structure of the forest (Figure 2) is characterised by a three-layered canopy in which the crowns of spaced conifers, particularly rimu, and of northern rata emerge clear of the main canopy to reach heights of 40 m or more. Northern rata is more prominent on rocky outcrops above bluffs. The main canopy at 2025 m height is com¬ posed of tawa, although kamahi is of major importance on some steeper sites sub¬ ject to disturbance. Hinau, mangeao and sometimes black maire form a second dis¬ continuous canopy layer between the discontinuous emergent layer and the semi- continuous main canopy. The understorey is relatively open and is unlikely to have impeded large birds except where supplejack was abundant. Tree ferns and ground ferns are both prominent in the understorey. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 137 TABLE 5 - Major plant species of humid conifer/hardwood forest in the Waitomo region. Scientific name Common name Abundance1 CANOPY AND SUBCANOPY: Southern conifers Dacrydium cupressinum rimu c Prumnopitys ferruginea miro Hardwood trees c Beilschmiedia tawa tawa a Elaeocarpus dentatus hinau a Knightia excelsa rewar ewa c Laurelia novae-zealandiae pukatea c Litsea calicaris mangeao a Metrosideros robusta northern rata c Nestegis cunninghamii black maire c Quintinia serrata tawheowheo c Weinmannia racemosa kamahi Woody lianes a Metrosideros diffusa climbing rata a M. fulgens climbing rata a M. perforata climbing rata a Passifiora tetrandra NZ passion vine c Rubus dssoides bush lawyer a UPPER AND LOWER UNDERSTOREY: Hardwood trees and shrubs Alseuosmia macrophylla toropapa a Carpodetus serratus putaputaweta c Coprosma grandifolia raurekau c Geniostoma ruprestre hangehange a Hedycarya arborea pigeonwood a Melicytus ramiflorus mahoe a Myrsine australis mapou a Oiearia rani heketara c Pseudopanax arboreus fivefinger a P. crassifolius lance wood c P. laetus c Rhabdothamnus solandri waiuatua a Streblus heterophyllus Juveniles of canopy and trees and shrubs small-leaved milk tree Tree ferns c Cyathea dealbata ponga a C. medullaris mamaku a Dicksonia squarrosa wheki Woody lianes a Freycinetia baueriana kiekie c Griselinia lucida puka c Metrosideros diffusa climbing rata a M. perforata climbing rata a Rubus dssoides bush lawyer Epiphytes a Astelia solandri (lily) astelia a Collospermum hastatum (lily) collospermum a GROUND STOREY: Shrubs Coprosma rhamnoides a Gaultheria antipoda snowberry c 138 ACTA XX CONGRESSUS INTERN ATIONALIS ORNITHOLOGICI TABLE 5 - (Continued) Scientific name Common name Abundance1 Herbs, including sedges Astelia fragrans astelia a Gahnia pauciflora cutty grass a Peperomia urvilleana a Uncinia uncinata hook grass a Ferns Adiantum cunninghamii c Asplenium bulbiferum a A. oblongifolium a Leptopteris hymenophylloides a Phymatosorus diversifolius a 'a = abundant species, present in 80% or more of stands c = common species, present in 20-80% of stands FIGURE 2 - Structure of pre-human conifer/hardwood forest in the Waitomo region. The site includes a young stand of southern conifers (rimu) on an alluvial terrace and rata grow¬ ing on a limestone bluff. Scientific names of plants are given in Table 5. Ground herbivores (Table 6, Figure 3) Moas dominated this feeding guild with seven species represented. Bones of two or more species are frequently mixed together in pitfall deposits suggesting that all seven species were present throughout the 6000 to 1000 y BP time period. Different moa species could, however, have used the forest at different times of the year according to food availability. The bones of Anomalopteryrx didiformis are more than five times more frequent as those of the next most common species, Dinornis struthoides (Millener 1981a). This may not necessarily reflect their relative abundance as they are unlikely to have had similar foraging behaviour. If, for example, A. didiformis alone regularly included moist fern-covered hollows within its foraging range then its chances of falling into deep fis¬ sures or cave shafts would be increased. The weights of species within this feeding guild ranged from 0.45 to c. 170 kg with large gaps in adult weights between some species (Table 6, Fig. 3). These gaps are to some extent more apparent than real because the weights of immature birds are ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 139 not included. Under natural conditions a more continuous distribution would have ex¬ isted. FIGURE 3 - Moas in the ground herbivore guild of the Waitomo region. From left to right: Dinornis giganteus, D. novaezealandiae, D. struthoides, Anomalopteryx didiformis, Pachyornis mappini, Euryapteryx curtus, E. geranoides. Estimated weights (kg) are shown for each species. How did the members of this guild of comparatively large herbivores co-exist? Stud¬ ies of mammalian herbivores in Africa have shown that the ability of these mammals to utilize food of poorer nutritional quality, i.e. foods high in structural carbohydrates such as cellulose, hemicellulose and lignin, is correlated with body size (Bell 1982). Larger body sizes are associated with adaptations for using low-quality herbage whereas smaller herbivores are more restricted to foods of high nutritional value. As a result, smaller herbivores generally tend to be more specialised in feeding behav¬ iour (Owen Smith 1982). The eight-fold difference in body weights between moa species of the Waitomo for¬ est (Table 6) suggests that species differed substantially in the nutritional quality of the foods eaten. The three Dinornis species were broad-spectrum feeders, probably exploiting a variety of food plants daily, and were able to use herbage of poor nutri¬ tional quality (Burrows et al. 1981). This is consistent with the heavy weights (total¬ ling up to 5 kg for D. giganteus) and large sizes of their gizzard stones (Gregg 1972, Burrows et al. 1981). Since both the ostrich and the rhea have hindgut modifications allowing significant fermentative digestion of fibre (McLelland 1979) it is probable that Dinornis spp., which apparently ate an even more fibrous diet, had similar specializations. Although it has been suggested that D. giganteus was largely restricted to forest margins and other more open habitats (Anderson 1989, Worthy 1990), in the Waitomo region bones of at least 22 individuals of this largest moa occur in 8 (22%) of the caves examined (Millener 1981a). This indicates that it was a forest animal in this region. Its rarity is predictable from its great size and probable diet overlap with the other two species of Dinornis (Table 6) which had similar bill shapes and musculature. If the horizontal foraging patterns of these three species were broadly similar, the chances of any one species falling into a cave would vary according to its relative abundance; recoveries of bones of the three Dinornis species from caves in the Waitomo region have been in the approximate ratio of 3:2:1, the smallest species, D. struthoides, being the most frequent (Millener 1981a). TABLE 6 - Foods and feeding behaviour of birds in the ground herbivore guild. 140 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI c I o « ] i E T3 8 8 II jn -5.0 © CL l| => Q. B "8 .52 T9 13 o E © £ o c © £ © T3 .C JS © © S .§ illf I* S-8* .2, I e ■g-e 3 © © > S I £ o © g> o £ - 2 § °- CL © E CD 8 1 it .. CD ^ c £ w II CD ll- i|| is i ^ 0 .. |S| Q TJ ® It? m © $ * ® ■DC® 5 8s i § S- o ~ - g Ms §2 .g I £ . ? S' I |8 t © .3 g 3 L. ©mg ^ -J JC 6 i* & I o (0 — VO 602 ■Q 8 CO £ I ■e 3 O k CD O) U5 C\j cvi Ill a CD T5 c 13 ■'t c osi CD I i * 5 < Q ia co ■S J8 ID •S *o c ^5 Q> N $ 2 O c .= x» 8 >» II <3 2 oi CD I III T> 5 3 2 CD I £ -8 1? CD 3 CO 8 8 8 3 a is O) CD a c ro fi § |tr a “ .£ IB CD co 6 10 s 6>‘ U9 “ cvi cvi d 1 $ g 0 o> U) !$ — 6 d cvi cvi d 1 d d 2 •» ar 2 ■» ST & c , = II »ss i | I a 8- « <0 C O) cd a m. o) 8, ■8 x Q a £ CD II U. a i ra I c 8 b z II E E 8 a ® CL ■c o $ 8 ■g o £ 5 CO co 'c L- o c b S o> c CD c _o £ CO ro ■o © u= •43 c © T5 8 .8 8 c ‘«0 © ifl £ C o o S» J3 <0 a 1 l E .8 ■e s a - o w « 4— * © £ 8 m >» S 5k 15 £ c © *-* c 8 XI £ ro N N CD ® © £ "8 ts £ £ 2 nJ 3 a c S .O) o> o I a CD rS £ c o '•8 CL E 8 8 © a 3 15 c 8 x> 2 ro JC § £ Q. 2 I E a TJ C n3 c a ® co I 8- o a £ E 3 C C ® > CD <0 • 2 Jr o) a «= > co- a -8 1 6 2 F (A C. <0 ji € Q 51 E 3 S 141 Extinct species; brackets are used to indicate speculative inferences relating to such species. 142 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI How then were food resources partitioned between these three species? Atkinson and Greenwood (1989) gave measurements indicating that there was a differential of be¬ tween 0.4 and 0.7 m between each pair of most similar-sized species in the heights they could stretch for food. Thus in times of food shortage, such as dry summers or cold winters, it could be expected that feeding by the most common species, D. struthoides, would deplete the understorey of foods suitable for Dinornis spp. Survival and thus coexistence of the two larger Dinornis spp. would then be possible only as a result of their height advantage. The two species of Euryapteryx, judged by their U-shaped bills (Atkinson and Green¬ wood 1989), rather weak bill musculature (Cracraft 1980) and small stone sizes and total weights of gizzard stones (Worthy 1989), probably fed on plant parts low in fi¬ bre such as leaves, buds and fruit. Presumably the much smaller E. curtus was more specialised in feeding than E. geranoides although the height difference would have made some foods available only to the taller species. The foods eaten by Pachyornis mappini are unknown although bill structure and cra¬ nial musculature (Cracraft 1980), suggest an animal adapted to cut fibrous leaves and stems. Fewer individuals are recorded from the Waitomo region (42) than of E. curtus (56) and D. struthoides (61), but P. mappini has been recovered from 12 (32%) of the caves studied (Millener 1981a). The diet of Anomalopteryx didiformis is also unknown. Its formerly widespread distri¬ bution in a variety of forest habitats, including low fertility beech forests, suggests that it was specialized to feed on the most nutritious parts of the plants present. The stud¬ ies of Emu by Davies (1978), Dawson and Herd (1983) and Herd and Dawson (1984) may provide a model for understanding A. didiformis. Weighing 28-48 kg, the Emu is comparable in size to this moa and is a generalized omnivore with high rates of pas¬ sage and no large chambers in the gut for fermentative digestion. They forage widely for a wide range of high-quality foods such as insects, seeds, fruits, shoots and green herbage. Although emus avoid plant material high in lignins they can digest fibre rich in hemicellulose in the distal segment of the small intestine (Herd & Dawson 1984). One of the more selective feeders among the ground herbivores present at Waitomo is likely to have been the Takahe Porphyrio mantelli. In their present alpine grassland habitat Takahe favour the basal meristems of tussock-forming grasses (Table 6), tis¬ sue low in fibre and rich in mineral nutrients, as well as fern rhizomes (Mills & Mark 1977). The gizzard is small (J.A. Mills, pers. comm.) and there are apparently no spe¬ cial hindgut modifications for microbial fermentation (Morton 1978). In lowland forest, suitable food plants such as the tussock grass Cortaderia fulvida occur along streambanks and a range of other plants including rhizomatous ferns grow within the forest itself The recovery of Takahe bones from 14 (38%) of the caves studied, in association with bones of many obligate forest birds, suggests that Takahe were feed¬ ing in the forest even if not breeding there. Although Mills et al. (1984, 1988) identify Takahe as a relict species of Pleistocene grasslands, the evidence both from the Waitomo region and elsewhere points to Takahe as having been widespread in low¬ land forest although not necessarily very abundant (Millener & Tempter 1981, Beauchamp & Worthy 1988, Caughley 1989). The great depth of the Takahe’s bill in relation to its width contrasts greatly with the proportions of a moa’s bill and suggests that co-existence of Takahe with moas was achieved through differing modes of feeding as well as foods eaten. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 143 c/) p '5 cr> 0 o > o CD CO c q3 v_ o > O) 13 ~o c 03 CD O > !o 0 JZ 0 O > D) 0 0 i_ O > !q i _ 0 0 0 O _Q l_ 0 0 _£Z C/3 u !5 o ZJ o > 0 _£Z 0 -Q 03 C T3 0 0 T3 C 0 0 "O o o LL HI _i m < i- I 1 £ 5 o < S3 O) lZ O) 8S o o o 0 0 88 d o §0 8 o o — o o Is Is ij $ih ill! (/) I'll ii«® = e U) $ -c -6 9- c 5 £ ~ © _ © | 2 $8 s 1 is 3 if £ re 0 Q. a 5 Q. >* 3 P S a. o I O V) £ __ £ 12 'S ffl ® •§ 8 § 73 £ 8 (0 o> If Is (/) 1 £ 2 o> jO *” i o ffl >* = £ o> g .S 6 g Ja (5 1 £ g S _Q 8 e i o _ 5f |S U) T3 is 49 8 si 8 CD o <0 £ o > O >- Q. S re 8. g CL 3 ti S 3 "o o fc C c ® 3 g * 2 O £ o> £ a o g 1 cn © (A 1 cn 2 re £> re 8 Z . c >% 3 o £ a o © 8 §■ sl 2 'o 4? 1 (/) ^ II Is 1 S re o»N c z c 5 Q X to 8 <0 9 © re 3 § 5 5 8.'? o §•« 5 $ a o o o o 0 Tf o o 6 6 o> cn 38 0S90 o CVi CO 0 0 S 5 o o d CM ^ d o cn o) 38 § o C\J CO 0 o 88 o o o o CM »— o o 6> cn 38 059 0 o CM X' CO 0 0 8 5 o o o cm o o >> © T> g i- O as CL © T> • gas .£ W Q 8 o > ■f3 o 8 c £ o > cn 2 LL .© re c L. .O £ 5 ffi s ^ E J2 5 F = flighted; F.less = flightless; (F) = weak or reduced flight powers; D = diurnal; N = nocturnal; N,D = both activities observed. I = Waitomo conifer/hardwood forest; II = Canterbury conifer/hardwood forest; III = Northern Capes coastal forest; IV = Chatham I. coastal forest. 144 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI The smallest member of the ground herbivore guild at Waitomo was Gallinula hodgeni which is only present rarely in bone deposits, and which probably fed mostly in riparian vegetation within or adjacent to the forest. The New Zealand Coot Fulica prisca although present at Waitomo, is principally a water-bird and has, therefore, not been included in the ground herbivore guild. Arboreal herbivores, frugivore/herbivores and frugivore/nectivores (Table 7, Figure 4) Strict arboreal herbivores are limited to two species of parakeet both of which eat principally buds, flowers and seeds. However the smaller Yellow-crowned parakeet Cyanoramphus auriceps includes a significant proportion of insects in its diet, particu¬ larly during the breeding season (R.H. Taylor, pers. comm.) and the Red-crowned Parakeet C. novaezelandiae forages on the ground and in forest openings or margins. FIGURE 4 - Aboreal herbivores and frugivore/herbivores in the Waitomo region. From left to right: Red-crowned Parakeet, Kakapo, Yellow-crowned Parakeet, N.Z. Pigeon, Kokako. Figures show weights in g except Kakapo (kg); male weights precede female weights (see Table 7). The Pigeon Hemiphaga novaeseelandiae, Kokako Callaeas cinerea and Kakapo Strigops habroptilus, although mainly leaf-eating, are here classified as frugivore/ herbivores, because their successful breeding appears dependent on adequate sup¬ plies of fleshy fruit. Kokako may sometimes breed earlier than Pigeons, thus allow¬ ing the two species to use fruit that ripen at different times, but there is overlap in their breeding seasons. Pigeons do not take insects but the significance of small insects fed by Kokako to nestlings as a further mechanism for ecological separation of the two species is unknown. The reduced flight powers of Kokako may have made them more vulnerable than Pi¬ geons to entrapment in caves but Millener (1981a) recorded nearly four times as many Kokako as Pigeons in the Waitomo cave deposits. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 145 3 03 CD O > o 0 c 0 0 "O !5 o 3 o > 0 _c 0 03 c TD 0 0 T3 C 0 0 "O O O LL I 00 LU _l CD < 1 1 * ■5 9 O < is 00 o 88 o o oo o 88 o o -o ^ S Q ^ 12 q! 8 oiS 08 0) 3 o £ © e £ « o « 3 S o c* o> 5 2 ® >•- o» 8^ e * J.s S r.-g ® gt< > S' gT £“-8 0 0 o o 6 88 o o 88 o o 8 CM o o o "D © 5 2 >- 12 © ® §1 >» Q. 8 8 £2 §> « 8 5 J a ^ 2 3 2 cO® -go—'’- if fl M|S *1^8 •E 3 I CM- 1 si? « s « « t= <2 a © > z § S’ •s g S ii 3: E E *> © ® P Q> 2 2 tj 1-81 fl oo 8 § a z 1= u. 0 O C\J O o o 6 6 0 o cm g> o o 0 o cm cn o o 0 o £8 o d r-- o2 |a| 1 1 <3 o> t o>< 2 XI CO f. sa 1 ”5 ssl 8 8 •o L z -6 £ © 2l 9 Q 9 LL LL LL t3 .C O 5 3 C -55 © © Sis 8 E x"2 6 S | a . |i £ =S © © CD 1 8 «o 5 £ o a. c £ 0 146 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI The flightless nocturnal Kakapo is not only the world’s heaviest parrot but the only lek breeding member of the Psittacidae (Merton et al. 1984). Its feeding habits are unique. The tip of the mandible rolls and crushes plant material against a series of serrated transverse ridges and grooves on the ventral surface of the maxilla. This allows sap and other juices to be extracted from leaves that are eaten while the fibre is rejected as pellets (Gray 1977, Merton 1985). The gizzard is relatively small and the extrusion of fibre during feeding means that no hindgut modifications are required for fibre di¬ gestion. Kakapo climb readily and feed arboreally as well as on the ground with other ground herbivores. Like the Takahe they feed very selectively but, unlike the gallinule, Kakapo eat an extremely wide range of species (Table 9). Kakapo breed later than Pigeons and Kokako and their apparent dependence on fruit (or possibly grass seed) during the nestling period in late summer, rather than insects, suggests that they did not compete with these species for food during the breeding season. Judged by the numbers of individuals recovered, Kakapo together with Anomalopteryx didiformis were the two most common flightless herbivores in the Waitomo forest. Fruit are also a crucial component of the diet of Kaka Nestor meridionalis but this parrot eats a large variety of high-protein foods not used by Pigeon, Kokako or other parrots in lowland forest (Table 7). Nectivores (Table 8) The New Zealand avifauna has no birds dependent mainly on nectar for much of the year, unlike that of Australia where nectar is more generally available. Nevertheless, because the three species listed in Table 8 are all Meliphagid honeyeaters with well developed brush tongues (McCann 1964) and all use nectar as a major food source, it is convenient to group them as a nectivore guild. The largest of the three species, the Tui Prosthemadera novaeseelandiae is the most dependent on nectar (Gravatt 1971). It is also the most aggressive, driving other honeyeaters and other birds away from trees where it feeds. A social hierarchy has been demonstrated by Craig (1984, 1985) in which Tuis dominate over Bellbirds Anthornis melanura which in turn dominate Stitchbirds Notiomystis cincta. Intraspecific dominance also occurs. The three species differ in their use of certain flowers as well as in the seasonal pro¬ portions of fruit and insects taken, especially in winter (Gravatt 1970, 1971, Table 8). Bellbirds are more insectivorous during autumn and winter while Tuis and Stitchbirds rely more on fruit at this time. Kaka when feeding on nectar or honeydew, also be¬ come part of the nectivore guild (Table 7). Aquatic insectivores (Table 9) This guild of three ducks was closely related to the riparian system within the forest and its associated habitats. Blue duck Hymenolaimus malacorhynchos are specialized to feed in turbulent or fast-flowing water and probably made little use of the forest it¬ self beyond the stream banks. Finsch’s Duck Euryanas finschi may have foraged extensively on land as suggested by Worthy & Mildenhall (1989) but the particular use it made of its very short bill is unknown. This species is the commonest guild mem¬ ber at Waitomo, perhaps because it may have nested in caves (McCulloch 1975). Brown Teal Anas aucklandica can use vegetated areas beyond stream courses where its nocturnal feeding may have allowed it to take foods unavailable to other members of the guild. TABLE 9 - Foods and feeding behaviour of birds in the aquatic insectivore guild. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 147 TABLE 10 - Foods and feeding behaviour of birds in the ground insectivore guild. 148 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI T i 1 CD C I o x: ® E « 1 * ■5 < S ■D > C II o = CM CO CM CO E E o £ * 0 9 8 1 II c o _ ® 2 E ?I | . s o © e “ i-8 ® bs.? a © a5 *- U) c\i c\i E sl So® it!. 3-8 J= u> o» SI £ 2 3 a o CM b T- in CM CM E E E s 8 f sz* fill 1 JS 0 3-8^5t 8 1 JS 0 3 o i* c 8 0 jj 8. § JS c 2 8 O'¬ 'S? 8 o f'- co o b 8 |1 1 JS 0 3 5 V) cn c 1 Q. T3 O '£ 8 0 <0 & £ o b I 0 vf o fl CD 1 fl R 5 2 5 S- fi I £ |S > £ c 5 wh II _c >< c .9 CD CL • I c? tr mt& S h. o o 2§ cn 2 pis §> 3 % 8 J & s N .55 0 J2 I ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI ■o CD 3 c c o O I o LU _J CQ < O) 05 c I o (0 * £ © E *c 8 * o < s> f CD « 0) b o T3 > C •* I S 5 £ 00 § o oo s o T> > ® a c 1 £ o 8 ff - k-* C 05 (0 1 1 © ;o® o . a: I 8^ £|S © § ® > o S’ _ 05 > § .E ro _© (D 2 8 3 2 05 8 c o O) c © «3 ■o © to ©* I? i « * o o ^ _o 03 CL O'- a 8. 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T" £ 3 00 = C. = ^ ^ o X o> in © © o o a> m o o o o m -o CO 0 © d Gerygone igata F/D Beetles, moths, caterpillars, spiders taken from leaves and terminal Upper and lower Grey Warbler shoots by gleaning, hovering and hawking (Gibb 1961, Gravatt understorey, 1971, Moeed & Fitzgerald 1982, Gill 1983) canopy TABLE 11 - (Continued) ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 1 53 co I, 0 11 8 « £ c oc £ CD I O 0) £ % E « TO 8 5 5 O < Is £ CD <0 0 I « 1 2 « S E « (0 £ y- O o o o o oo 8 o oo 8 o oo 8 o co 8 o co o d 7 > 0 >i © Q. 8 o O) O 8 ^B c e 1 © CL 2 •8 1 $ 8-8 0 U. c c 3 CD i 3 § >. © CD X) c 3 2 CD O o o o ■o c ra >* >; s 2 B Q w to "U T3 | % § § I C 2 2 J2 3 O) O) Q LL a CD 1 1 £ ra cb o © -e i to © -g 8« i.§3 -2> c £ "5 © tt « u. ■8 its © © IN i Ez 3 TJ II o kf © i CD 2 £ TJ 8 I © 8.1 - s g> 8 “= ^ T3 3 8 ■D ?o I 2 © © JL 8 io" "D W O © O II « € £ §.= ■D I 8 ll & CT'S TJ II •,3 II x B § ii 8 154 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI >> ~o o jD CD O LO A if) 0 O > * o 0 < f ) 0 0 0 O JD 0 i_ 0 CD i_ 0 3 CD 0 v_ O > H— > o 0 co c 0 0 O JD 0 0 if) ~o o 3 O > 0 JZ 0 JD CD 0 ~o 0 ^0 "0 0 0 if) ~ © * > s 2 5a £ ® Q to CM to h. o 8 CM o g i~" O) 88 o o 6 6 o o o .175 005 0 0.200 .085; .072 o o « c to to § 8 i-’ <5> o CM o CM o 88 £ _ .I o o 6 6 i o o CL o >, © k- o 2 T5 C 3 ,s c 2 O 3 CD ©‘ © it S' If .1 8g -g x> E ” § 5 o> g> ii <5 0.0 * ©5 w "S -s a8s W CD CO O) c ^ (/) “O 5 8 © is-SdE al* | s| 2-3 = -g | « ® 1 18 I i 2 2 CD* S« ■Q-i £ o O- 5 ft 2 O (0 CD c o o c 8 © 3 ■« a s* o 2 8 -8 | 8 § §> © jo "c3 T3 2 i ' CD <£ ||8 1 (3 . ® a i 2 do 0 is h © 8 £ o E •o I »| IP 111 * 5 dlfi iali o 5 o 3 o S .c © © -6 © © in © 2 o c ■6 a 2 o 2 is! | §£ ill T3 £ O 0 “8 8. | * I a’ i2'i £ S I’P 8 w ® 8 ® T3 3 - = § 8.13 © co V m 1/3 JS .© 5) o >» © B 2 ^ -8 S c o 3 CD CD C 3 O CL o> CO o> >N ra % S o X B C © CD 2 © © © € o s. !i 5 c N 2 Si CL 8-8 8 " si 3 © g- | "-<3 1 I? & CL 3 c 2 - 3 CD .E >» «- 2 si § | -o 8^-8 § 8 § § 2 CD (3 © 'O' ■§ "g1 |'£ © _® a © © £ m in c o £ TB n 3: 2 CD c k 3 |i5 11 o © I? 3(55 E a © .© £ TABLE 12 - (Continued) ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 155 o> s «— o CO £ % E « TJ s £ > O (0 £ 05 O * 2 Si o > 05 O O) S in a o m 8 o § o 6 o o o 6 o 6 7 > Ss £ Ss £ 2 C 3 2 05 aT k_ O) 2 <0 . o _C 1/5 k_ 0 T> c 3 "O -D c C *u c >; £ >. Q. 1 UL 3 3 2 2 05 05 3 2 O 8| (0 3 o c s sfl JS 8|* ® - o c >5§o 05 ■- C i_ .c 1- X) ® « - rt XI ° 313 I* w 2 2 M re £ ° g i1 -E 2 re •s o re - •£ SJ ^ « ® u 2 X re x re y» o I I'S- K 8 cl 8 w. O o £ k— u_ 0 ^ = f! 05-6 05 J3 £ (5.?® 156 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Larger arboreal insectivores: > 50 g wt (Table 12, Figure 6) This part of the insectivore guild contains both generalists and specialists. The larg¬ est of the generalists is the Long-tailed Cuckoo Eudynamys taitensis, a summer mi¬ grant that eats a wide range of small animals from all levels in the forest and which possibly also feeds at nigh't. The extinct Piopio Turnagra capensis included inverte¬ brates as an important part of its diet but also ate fruit, seeds and foliage, and could perhaps be classified as an omnivore. Both the Saddleback Philesturnus carunculatus and the extinct Huia Heteralocha acutirostris are specialized to take invertebrates by probing into living or dead wood. The much greater size and strength of the Huia, together with its marked sexual di¬ morphism in bill shape, allowed the Huia to extract invertebrates from a wider range of woody substrates too difficult for Saddlebacks to use. Three nocturnal insectivores are also included in this guild; two raptors and an Owlet- nightjar. The largest of the raptors, the extinct Laughing Owl Sceloglaux albifacies probably foraged in forest openings and along forest edges and is only rarely present in the Waitomo cave deposits. The much smaller Morepork Ninox novaeseelandiae, which hunts both in forest and forest edges, is equally rare in these deposits. The extinct Owlet-nightjar Megaegotheles novaezealandiae is much better represented in the Waitomo region than either of the nocturnal raptors. Its diet is unknown but large flightless insects and nocturnal moths would have been potential foods. When feeding on wood-boring insects the Kaka parrot would be functioning as a member of this guild as would Kokako when taking foliage insects (Table 7). It should not be forgotten that some members of this guild could at times be in competition with bats for food. Major predators of vertebrates (Table 13) At Waitomo this guild consisted of four species, three of them diurnal raptors. The largest of the raptors was the extinct Circus eylesi whose phylogenetic relationship is with the harriers Circus spp. but whose morphology, particularly wing shape, sug¬ gests that it behaved more like a goshawk (Holdaway 1989, pers. comm). It probably preyed on medium-sized birds by rapid pursuit through the forest from a perch rather than hovering or soaring to find prey. Thus moa chicks are likely to have been part of its diet. The much smaller falcon Falco novaeseelandiae is little more than a tenth of the weight of C. eylesi and takes much smaller prey from both perching and soar¬ ing positions. The harrier C. approximans specializes in stationary or slow-moving prey and carrion. Food is sighted while soaring, possibly assisted by hearing for live prey. The feeding habits of the North Island Aptornis Aptornis otidiformis are something of an enigma. The position of the eye orbits suggests it had binocular vision. R.L. Zusi (pers. comm.) has found evidence of very heavy musculature in the upper neck re¬ gion behind the skull which he considers gave the bird considerable levering or wedg¬ ing power. The bill is large and broad, angled at the edges and both flattened and slightly downcurved at the tip. Gizzard stones have not been recorded with bones of Aptornis. Thus this bird may have fed by tearing open rotten logs or digging for inver¬ tebrates, Tuatara, or petrels in shallow burrows (Holdaway 1989). 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O) w O t 00 1 ^ W5 00 ^ e ® O & m 8.73 Q LL Q ^o 8 (D *r C .© iS E E © * x 5 C Q. © 9- 9 © re o o in CD ■o § - £ « - fi CL - o s§& c cr 2 2 -fi 05 8 2 -fi ® 8 re « — s °. $ auu»o i canopy undersl ground canopy subcan ground 5 a>; § O £ 8 v, ground above i ground marine 3 ^ § 2 2 fi O) (D 1 >; 5 2 2 2 O) (D ground ground, underst canopy near co CO *s © N 23 88 •So S>.<5 4.i 8| JO o © • © CD CO C fi! «= o Jfl °- © CD to c *s . fi «D © ‘ C LjJ o LU CL *•««« UJ Q | * * ENDEMIC LANDBIRDS . - . 9 -80 -70 -60 50 -40 -30 -20 -10 — i . . i ■ . "r . . -i — 1 — i - 1 - 1 1 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 2000 < LU CC < Q Z 65% — Brooker et al. 1989 White-fronted Bee-eaters -> 9-1 2% <— Wrege and Emlen 1987 White Crowned Sparrows 34-38% — Sherman and Morton 1988 Indigo Buntings 27-42% — Westneat 1987 Eastern Kingbirds 30-53% — McKitrick 1990 Cliff Swallows 6% 24-43% Brown and Brown 1988 Swallows (Denmark) — 17% Moller 1987 Pied and Collared Flycatchers 24% — Alatalo et al. 1 984 Tree Swallows 24-32% — Morrill and Robertson 1990 Eastern Bluebirds 9% — Gowaty and Karlin 1984 Even though it is not possible to separate familiarity and kinship in already published studies their results are highly supportive for familiarity being an important independ¬ ent variable in addition to kinship. Wrege & Emlen (1987), Curry (1988), Clarke (1989), Davies (1990) and many other studies (see Stacey & Koenig 1990) found in¬ dividuals helping familiar but unrelated individuals. The more familiar a “helping” in¬ dividual is with the adults controlling a nest, the more likely it will encounter the nest and hence a begging stimulus. The consequence will be to feed. Curry (1988) and Curry and Grant (1990) even argue for the importance of familiarity but because they did not separate it from kinship they were left with no alternative but to preferentially reject the stimulus-response model. Future studies should acknowledge the likely importance of both kinship and famili¬ arity in determining associations among individuals. Combining observation of behav¬ iour with molecular tests of parentage to determine kinship, and measuring familiar¬ ity by frequency of close association will allow more realistic tests of existing models and their interdependence (Figure 1). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 241 Kin mating and incest avoidance Kin groupings so commonly found in communally breeding birds provide ideal situa¬ tions for evaluating ideas of inbreeding avoidance. In spite of the theoretical notion that inbreeding is but one extreme of a continuum of mating with outbreeding the other extreme (e.g. Shields 1982), most workers treat inbreeding in isolation. Inbreeding can be potentially disadvantageous in a normally outbreeding population but it is often forgotten that outbreeding can be equally problematic in typically inbred populations (Craig & Jamieson 1988). Inclusive Fitness model Stimulus-Response Help Non-kin Kin Non-kin Kin univariate model multivariate model FIGURE 1 - The relationship between kinship and familiarity in expression of helping be¬ haviour. a) Inclusive fitness model that subsumes familiarity as the recognition mechanism for kinship. This model has current acceptance, b) Stimulus-response model that assumes that the degree of association individuals have will determine familiarity and hence the like¬ lihood of encountering a begging stimulus. Kinship may or may not add additional explana¬ tory power. Few workers have attempted to consider the full range of variables. Whether individu¬ als mate with relatives is affected by a number of issues including incest avoidance, dominance, reproductive competition and mate choice. Reproductive competition by intra-sexual dominance can either be by fathers or replacement males dominating and preventing helpers (sons) from copulating with their mate (the helpers mother) or by a group of outsiders displacing a smaller number of existing male residents (e.g. Acorn Woodpeckers - Koenig & Pitelka 1979, Hannon et al. 1985). Dominance may be so extreme that individuals are psychologically castrated (Reyer et al. 1986) and hence appear incapable of responding sexually or aggressively in the short term. In¬ ter-sexual dominance may explain why a female will not copulate with her son but would rarely apply to fathers avoiding mating with daughters. In addition some fe¬ males may not choose to mate with young males because of reproductive inexperi¬ ence. A number of bird studies have shown that females with young males have lower 242 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI reproductive success than those with older males and that females prefer older part¬ ners (see Bateson 1983, Walters 1990). However, Walters ( 1990) suggests that even though young males make poor reproductive partners in Red-cockaded Woodpeck¬ ers the movement of the female away from her sons relates to incest avoidance. Many recent reports of communal breeders still suggest the likelihood of incest avoid¬ ance (Brooker et al. 1990, Zahavi 1990, Koenig & Stacey 1990, Ligon & Ligon 1990, Rabenold 1990, Woolfenden & Fitzpatrick 1990) and some even assign attributes such as reduced hatchability of eggs to inbreeding depression (Ligon & Ligon 1990). With the exception of the work on Acorn Woodpeckers (Shields 1987, Craig & Jamieson 1988, Koenig & Stacey 1990) none have tried to systematically test alter¬ natives. Some workers argue that dispersal of one sex further than the other is a form of in¬ cest avoidance (e.g. Zahavi 1990, Rabenold 1990, Walters 1990). A contrasting view is that reproductive competition (both inter- and intra-sexual) will ensure dispersal and that the consequence of this may be a reduced probability of incestuous mating. As Ligon & Ligon (1990) found dispersal can still allow incestuous mating to be common. Realistic evaluation of incest avoidance should contrast all explanations of observed mating patterns and where possible attempt estimates of the frequency that close relatives of equal status are available at the same time and the same place for mat¬ ing. This can then be compared with observed levels. The observation that few or no incestuous matings are seen is poor evidence for incest avoidance behaviours. TABLE 2 - Incestuous mating among communally breeding birds. Species % Incestuous N mating Source Pukeko >70% 107 Craig & Jamieson 1988 Splendid Fairy Wren 27% 270 Rowley et al. 1 986 9% Brooker et al. 1990 Galapagos Mockingbirds 7% 156 Curry & Grant 1990 Groove-billed Anis 1.6% 127 Koford et al. 1990 Some communal breeding studies report relatively high levels of incestuous mating (Table 2). Many studies suggest that levels of inbreeding between 1-3 % are of little consequence among typically outbreeding animals (e.g. Soule 1983, Shields 1982). Where levels of inbreeding regularly exceed 3%, inbreeding may be considered part of the mating system and hence it is unlikely that deleterious effects of inbreeding will be detectable. Rowley et al. (1986) reported high levels of incestuous mating but no significant dif¬ ference in reproductive success of inbred versus outbred pairings. However, the over¬ whelming scientific belief in the deleterious effects of inbreeding led Rowley to con¬ clude that his results were anomalous. DNA fingerprinting has shown that at least 65% of young do not belong to the male in the territory. The results allowed the re¬ searchers to return Splendid Fairy-Wrens to theoretical normality by suggesting they may have an inbreeding avoidance mechanism in promiscuity! Recalculated incest levels are still well above most species at around 9% but were conveniently not cal¬ culated. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 243 These results of high levels of extra pair copulations greatly reduce the level of relatedness between helpers and the young they feed. As Rowley & Russell (1990) state, this will markedly lessen indirect fitness gains. Many individuals, thought to be feeding relatives are provisioning non-relatives. Follow up of future relationships will allow an interesting test of whether kinship or familiarity better explains helper asso¬ ciations and mating patterns. SUMMARY AND CONCLUSIONS Science is influenced by changing fashions in theory as well as local and regional biases. For example, studies of communally breeding birds have increased greatly since the advent of theoretical ideas such as kin selection and inclusive fitness (see Brown 1987). Prior to 1970 most studies were largely on aspects of natural history. Acceptance of the communal breeding phenomenon appears to have been far more rapid among Southern Flemisphere ornithologists. The high frequency of pair breed¬ ing birds that hold breeding territories only seasonally in the Northern Flemisphere led to skepticism toward the phenomenon among researchers from that part of the world. Similar biases due to theory and our own encounters with nature greatly influence the questions we ask and the conclusions we reach. Even though there has been in¬ creased questioning of adaptive or selectionist reasoning since the late 1970s (e.g. Gould 1978, Gould & Lewontin 1979) this debate is only just impacting on studies of communal breeding (e.g. Jamieson 1986, 1989, Jamieson & Craig 1987, 1990, Emlen et al. 1990, Ligon & Stacey 1989, Mumme 1991). Initial responses to the plea to in¬ clude realistic evaluation of many alternatives including nonselectionist models have been confusing (e.g. Curry 1988, Clarke 1989, Jamieson & Craig 1991). A major prob¬ lem has been the subsuming of important variables such as familiarity as a mecha¬ nism for the theoretically important kin recognition. This assumption has meant that attempts to formulate models from stimulus-response ideas have largely produced straw alternatives that have been easily dismissed. Other examples of investigation where workers appear driven by conventional inter¬ pretation of theory include the idea of incest avoidance. In all examples, workers are urged to expand the range of questions asked to include the four questions suggested by Tinbergen (1963): causation, development, function and evolution. There is a need to increase our efforts at identifying more of the likely variables that influence the behaviour of communally breeding birds and how these variables interact. Future studies need to be multifactorial and be aimed more at describing, understanding and predicting the birds’ behaviours rather than simply supporting the conventional read¬ ings of theory . ACKNOWLEDGEMENTS I am grateful to my colleagues on the Scientific Committee, especially Lloyd Davis, Murray Williams and Ben Bell, for nominating me for this address. My work and this paper have greatly benefited from discussion with, and helpful criticism from Grant Dumbell, Ian Jamieson, Anne Stewart, Chris Craig, Russell Gray & Sandra Anderson. My research is supported by the University Grants Committee, Auckland University Research Committee and the NZ Lotteries Board. 244 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI LITERATURE CITED ALATALO, R.V., GUSTAFSSON, L., LUNDBERG, A. 1984. High frequency of cuckoldry in Pied and Collared Flycatchers. Oikos 42: 41-47. AXELROD, R., HAMILTON, W.D. 1981 . The evolution of cooperation. Science 21 1 : 1390-1396. BARLOW, G.W. 1980. 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BUCHER . . / ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 249 APPLIED ORNITHOLOGY: PUTTING THEORY AND PRACTICE TOGETHER E. H. BUCHER Centro de Zoologia Aplicada, Universidad de Cordoba, Casilla de Correos 122, Cordoba 5000, Argentina Man as a biological species has opted for manipulating the environment instead of passively adapting to its vagaries and restrictions. As a result of that, we have enough energy, resources, and time available to allow ourselves to get involved in many ac¬ tivities besides our basic biological duties, including science. During this century, scientific research has become a widespread, full-time professional activity, which in turn has contributed to increase our ability to manipulate the external world still fur¬ ther. Consequently, scientists are expected to be able to solve nearly all kind of prac¬ tical problems, including those caused to the environment precisely by our ever in¬ creasing technological capabilities. Having played a crucial role in alerting the world on the risks of forest destruction and biodiversity loss, ornithology is certainly well involved in the applied-versus-pure re¬ search conflict. Ornithologists are permanently requested to be relevant to society, and the IOC parallel meetings organized by the International Council for Bird Pres¬ ervation have been alerting us for several years on the magnitude and difficulties of the problem of protecting birds from extinction. The decision of including this topic in an IOC meeting is a clear indication of the ornithologists’ growing concern regarding their duties and responsibilities in society. Indeed, the science of ornithology has an outstanding record of contributions to the solution of applied problems. Many of the basic concepts firmly established in man¬ agement today were first coined within ornithology, some of them after longstanding, vigorous theoretical arguments. Emphasis in applied ornithology has shifted through¬ out the years from gamebird hunting and birds as agricultural pests to species con¬ servation, and the preservation of biodiversity. Well established principles originally derived from studies of birds include: a) The existence of density-dependent mechanisms of population regulation, and their counter-intuitive connotations for management. This may preclude achiev¬ ing increased harvests via predator control in game species, and make massive killing an unsuccessful method of pest control (Murton 1971). b) The existence of a critical minimal size below which a population is susceptible to extinction due to behavioural, genetic, and stochastic factors. c) The importance of behavioural aspects in bird management, including captive propagation and reintroduction of bird species to the wild. d) The existence of a complex correlation between habitat structure at different di¬ mensions and scales, and species survival, which has important implications for the design of reserves. e) The existence of areas of high diversity (“hot spots”) which deserve special con¬ servation efforts. f) The importance of some bird species for pollinating flowers and dispersing seeds. 250 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI There are numerous examples of successful applied research projects in ornithology throughout the world, too many to enumerate here. Several of them have been car¬ ried out by New Zealand ornithologists. New Zealand’s conservation research in¬ cludes fine examples of well designed and well implemented projects that have saved local species from extinction by means of intelligent and innovative manipulations (Napper 1989, Bell 1991). A growing concern Despite the outstanding record of contributions made by ornithology and other envi¬ ronmental sciences to management, there is a growing feeling among the public, politicians, decision makers, and funding agencies that not enough effort and exper¬ tise is being dedicated by academic sectors to the solution of environmental problems at this time, when the future and fate of our planet is at risk. In fact, we all face, to a varying degree, a moral and practical dilemma between pur¬ suing our own “pure” interests in science or concentrating on the solution of problems, some of them of enormous magnitude, such as the effects of global changes in cli¬ mate or the massive extinction of species. These conflicting feelings are currently being debated in nearly all academic circles dedicated to environmental sciences. Discussions usually include points such as whether we can afford to continue to be involved in “hobby projects” (as called by the utilitarians) that might lead to the dis¬ covery of another subtle ecological process in the tropics, instead of concentrating all our efforts on crucial problems such as saving the tropical rainforests from total de¬ struction. Unfortunately, long debates often lead only to more debates without prac¬ tical results. Even more frequently, empiricists, theoreticians, and “applied” scientists seem to have become increasingly isolated and entrenched in their own intellectual positions allowing little room for effective collaboration. Another aspect of the same moral dilemma has to do with a growing availability of funds for applied research that has led some scientists to proclaim interest in applied problems as a way of obtaining extra funding, without a real commitment to solving them. As a result, it is not unusual that funds targeted for the solution of specific prob¬ lems may in practice be used to carry out research that is irrelevant to the final goal. Moreover, in many cases priorities set for applied research programs often reflect the personal interests and perceptions of the members of the board rather than the real needs. Although reasons for such a strategy in a competitive world of constantly changing priorities are evident, it cannot be justified on both ethical and practical grounds. I am convinced that unproper targeting of funds intended for applied re¬ search is greatly reducing the effectiveness of the available resources throughout the world. Indeed, the whole issue of applied versus pure research is a moral one. Clearly, if we are studying birds that live in the real world, we must be prepared to help to manage that world as efficiently as possible. It is also very clear that management decisions need to be taken within short time, and that pressures and vested interest are always present. We can then expect that decisions will be taken anyhow, and without the best possible advice, unless scientists endeavour to find better alternatives and are deter¬ mined to advocate in favour of them in an effective and articulate way (Williams 1990). Despite its obvious relevance, I do not intend to insist here on the moral aspect of the applied versus pure research dilemma, for two reasons. In the first place, I feel that ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 251 awareness and moral issues in the environmental sciences were critical a couple of decades ago but have already been widely discussed. Today, the academic commu¬ nity is already well aware of the implications. Second, I firmly believe that other cru¬ cial aspects of the problem, particularly those related with the implementation of ap¬ plied science, are equally critical and deserve consideration. In other words, I do not feel that we will solve our growing environmental crisis by simply demanding every ornithologist to become permanently involved in practical problems, particularly when by doing so they may be forced to divert from the fundamental basic questions in which they have deep interest and expertise. On the contrary, priority should be given to find practical ways of making useful contributions to problem-solving without los¬ ing the motivation and the productivity that each one of us has when pursuing the fundamental questions and the inquisitive awe which first led us to ornithology. Furthermore, I would like to discuss alternatives that may enable us to make the most from the intellectual polymorphism currently in the scientific community. We need to promote a positive feedback between the search for general principles in science and the efforts aimed at solving specific problems. In other words, I argue that the funda¬ mental challenge we face today is for us to perceive the need for an improvement in our traditional approach to applied problems, and endeavour to implement them as fast as possible. In order to analyze that possibility, I would like to discuss the following points: a) What are the problems that interfere in the interaction between pure and applied research? b) Are there better alternatives for implementing applied research in ornithology and related environmental sciences?. Problems in developing applied research From my own experience, there are some identifiable factors in the present academic world that conspire against a better interaction between the academic and manage¬ ment sectors. Some of these problems are deeply rooted in human behaviour. The following deserve special consideration: Universality versus practicality: In academic circles applied research tends to be considered both less attractive and less prestigious. Such feelings are rooted in their perception of science as a discipline aiming at the discovery of basic laws and prin¬ ciples that govern the universe. In Einstein’s words, “science is a search for those highly universal laws. ..from which a picture of the world can be obtained by pure de¬ duction” (Popper 1965). Accordingly, theory-minded ornithologists are sometimes reluctant to become involved in what they consider economically useful but intellec¬ tually trivial investigations. As a result, many theories are never tested in practice due to the lack of interaction between “theoreticians” and “empiricists” (see Roughgarden et al. 1989). Furthermore, according to Slobodkin (1988) “Ecology in the absence of practical questions is in danger of deluding itself by a vain hunt of generality, answer¬ ing only easy questions that it poses for itself, and becoming irrelevant to anyone outside its own academic village.” On the other side, ornithologists involved in applied problems may lack exposure to the present-day status of the art in theoretical ecology. Consequently, applied scien¬ tists may miss the opportunity to gain access to a wider conceptual framework that 252 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI may provide them with new techniques and perspectives of great practical value. Lack of interaction between theory and practice may also make applied scientists unable to take advantage of management practices (such as predator control, prescribed burning, etc.) as large scale experiments where theories can be tested. Consequently, practical questions in environmental management in the absence of sound ecology are likely to receive misleading and even dangerous answers. Increasing specialization: a growing tendency to specialize does not favour selection for a holistic approach to science. In any specialty, the need of keeping pace with a discipline tends to preclude investments in widening field experience and the aware¬ ness of developments in other areas, not to mention seeking ways for making the specialist’s work useful. Ecology was once considered a truly “holistic” science, but to a large extent is losing its conceptual unity. Although specialization is absolutely essential in science, given the exponential in¬ crease in the body of knowledge and a growing competition between scientists, it should not mean a drastic (and debilitating) narrowing of a scientists’ interests. Un¬ fortunately, this is not usually the case. As a consequence, scientists capable of keep¬ ing a holistic approach when dealing with applied problems are increasingly rare. In fact, there are growing pressures to specialize, in order to be successful, but at the same time the questions we ask increasingly require broad, synthetic answers. Fashions: research fashions change periodically, often without having completely solved the initial challenge. So do the search for suitable species and field conditions that may favour testing the presently discussed hypothesis. Although fashions have proved to be very important in promoting an accelerated de¬ velopment of specific areas of science, they can also have the negative effect of dis¬ couraging at least momentarily other areas that are equally important or necessary. Examples include the “shading out” of population ecology during the community ecol¬ ogy era, and the lack of support for projects of sustainable management during the “reserve-oriented” times in conservation. Preference for fashionable themes and disdain for research relevant to the local re¬ ality is sometimes an unwanted result of postgraduate training of students from un¬ derdeveloped countries in centers of high academic excellence in the developed world. The problem may be designated as “type 2 brain drainage”, type 1 being the actual emigration from the country. Territoriality: Although man is a relatively gregarious species, cooperation is not al¬ ways easy, particularly within the academic environment. Communication and collabo¬ ration is difficult between individuals, departments, institutions, and disciplines. The widespread “lonely wolf” attitude corresponds well with the average scientist’s incli¬ nation for independence and personal control of his or her environment. Lack of communication and interaction between disciplines is also widespread. The following quotation exemplifies a common situation: “A quantum leap in progress could be made if evolutionary and population biologists mellowed in their widespread animosity for ecosystem science” (Schlesinger 1989). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 253 Isolation is commonplace between government agencies and research centers, de¬ spite the obvious need for collaboration between both. Researchers at government agencies can very rarely carry on in-depth research on the large variety of problems they have to face. At the same time, universities tend to lose contact with reality un¬ less they are connected with those applied biologists on the “front line.” Full-time and full-life careers in the university that do not expose scientists to the “real world,” may lack the formative input and motivation that even short periods in government or pri¬ vate agencies may provide. Lack of institutional expertise in inter-disciplinary research: Institutional programs aimed at the solution of applied problems are usually managed in exactly the same way as any other scientific program, ignoring their unique needs for the implementa¬ tion and coordination of inter-disciplinary research. As a result, a substantial propor¬ tion of the budget may go to scientifically sound but irrelevant research, key compo¬ nents of the problem may remain undetected, no effective interaction between man¬ agers and extensionists develops, and findings remain unnoticed by the potential users. Short-term funding also poses another problem that may seriously impair applied pro¬ grams which generally require more than one fiscal year to attain meaningful results. Misconceptions about applied science: Several pervasive “myths” about applied science tend either to make people reluctant to become involved in applied work or lower the success of applied research programs. The more noticeable misconceptions include: 1) Applied research is always intellectually unchallenging and trivial. 2) Any good scientific study contributes to better management. 3) Comprehensive surveys and descriptions are the necessary first step. 4) Each new problem is unique. There are few background principles, information, or even comparable past cases. 5) Management, implementation, evaluation, monitoring, and extension is for some¬ one else, somewhere else. Such misunderstandings are largely the consequence of approaching applied prob¬ lems with the same point of view and methodologies used in basic research. The alternatives I believe that the goal of solving applied problems in environmental sciences implies not only commitment, but also the need for the application of adequate concepts, pro¬ cedures, and techniques. Although we scientists are well aware of the importance of sound methodologies and experimental design in our own research, we tend to dis¬ regard the need for appropriate techniques in implementing applied projects, feeling that this is something that anybody can do just by inspiration, which usually is not the case. At the same time, the need for more efficient methodologies should not lead us to forget that techniques alone are not enough. The best of techniques, unless guided by a clear vision of the fundamental issues and by a concept that gives them form, can turn solutions into larger problems. Improving our approach to problem solving requires at least a) finding creative ways of facilitating interaction between theoreticians, empiricists, and managers, and b) providing them with the intellectual and logistic framework necessary for interaction. 254 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Preliminary model j FIGURE 1 - Basic steps in the development of a “strategic” management program. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 255 One possible way of achieving that goal is to replace the traditional, discipline-ori¬ ented style of research by a goal oriented approach, in which all research is subor¬ dinated to the management goal in a systematic, interdisciplinary way. Such a “strategic” approach is not new in other areas of science and technology, in¬ cluding business, industry, and military science, usually associated with operational research. Moreover, Holling (1971) has already demonstrated that many sophisticated techniques developed in systems sciences (like optimization methods and decision theory) can be usefully incorporated in environmental impact assessment. However, it has not been widely adopted in ornithology or in the ecological sciences in general. Reasons for that include the problems and constraints already discussed, as well as the fact that these ideas are usually presented in a complex mathematical language, which is less accessible to non mathematic-minded naturalists. In simple terms, a strategic approach consists essentially of the following steps (Figure 1): a) Definition of the management goal. b) Elaboration of an initial model based on available information. c) Detection of research priorities and needs in terms of the management goal, and elaboration of the respective research projects. d) Re-elaboration of the original model based on the information obtained from the first round of research projects, iteration of the whole process if necessary, and elaboration of management recommendations. e) Monitoring and readaptation if required. As such, a strategic approach does not rely on a once-forever set of recommenda¬ tions of predictions, but rather provides a flexible, interactive approach to manage¬ ment needs. And we must accept that uncertainty is an inevitable component of the behaviour of all complex systems (Holling 1978, Ehrlich 1989). A strategic approach to applied science has several advantages over the traditional basic approach. First, the goal determines the priorities. A strategic approach ensures that all the relevant components and driving forces of the problem are considered in an interactive way, instead of allowing useless competition for the available resources among disciplines. This also avoids the selection of only those aspects that are intel¬ lectually fascinating as well as technically feasible, and the rejection of those aspects that are less fascinating or less feasible but equally important in terms of the final goal. In the second place, by proceeding in an interactive way, the program can be permanently evaluated, improved, and redirected if necessary. A constant process of re-elaboration of the original model can provide decision makers with the best avail¬ able advice whenever necessary, instead of postponing any recommendation until the final report. This is particularly important given that management problems generally require assessment and corrective action long before comprehensive models can be constructed. On critical issues, where only quantitative assessments are needed by policy makers, the strategic approach may prove to be extremely useful even in its early stages of development. Finally, a goal-oriented project is easier to be evaluated, improved, and redirected if necessary. By promoting only goal oriented research it also helps to maximize the benefit to cost ratio of the investment. Under this basic structure, the process of research can be expanded and refined by using all available methodologies. Each subset of the problem can be reanalyzed 256 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI following the same procedure, until the single-project level of detail is reached. Math¬ ematical modelling and other techniques can be included to better understand spe¬ cific sectors or interactions. At this point, an ideal stage is set for facilitating interac¬ tion between theoreticians, empiricists, and managers regarding specific sub-prob¬ lems within the project. * Implementation of a strategic approach to research requires the interaction of special¬ ists under a coordination capable of keeping the whole process without deviating from its original course. In its basic form, the whole project can be conducted through a series of workshops (Holling 1978), although other mechanisms may be equally prac¬ tical, providing that the basic steps and goals are maintained. In practice, however, implementation of a strategic approach to research is not without difficulties, and several problems can be expected at the researcher, institution, and funding agency level. Coordination is critical. The role of the coordinator is a complex one, that requires not only scientific but also managerial skills. Lack of clear leadership or clear operational rules (authorship, etc.) may prove fatal to any project. Unfortunately, training for such a role is not usually included in university programs in natural or environmental sci¬ ences. Other potential problems include the lack of specialists, lack of motivated peo¬ ple, and personal conflicts (territorialism, “prima donnas”, politics, etc.). Goal-oriented research has two basic forms of implementation at the institution level. One is to concentrate all researchers in one institution devoted to solving a specific problem or a group of related problems (the “institutional” approach). The other pos¬ sibility is to set up temporary teams integrated by scientists of different institutions to deal with specific problems (the “horizontal” approach). Expected problems of the institutional approach include a lack of flexibility, ageing and overgrowth. Researchers may sometimes feel frustrated by the need to deal in a superficial way with a succession of problems without having the chance of deepening their skills and knowledge in one specific subject, and therefore become reluctant to keep apace with new needs and challenges. Moreover, in some cases problems appear and disappear at a much faster scale than originally expected, creating the need for sudden shifts in priorities. Finally, it is also possible that an institution will become dependent on the continued existence of the problem for its own justification and survival, which obvi¬ ously conflicts with its ultimate goal. The “horizontal” approach is well established in the industry and business sector, but is uncommon in the academic arena, where strong individualism is widespread. Even when research teams are implemented, there is usually a confusion between fully interactive, goal oriented, “interdisciplinary” teams, and “multidisciplinary” groups where members work on different aspects of the problem but in an unrelated way. Although difficult to implement, a strategic approach to research is by no means im¬ possible, and certainly is one of the few practical alternatives potentially capable of providing the adequate framework for effective interaction between ornithology and other environmental sciences, and critical present applied research needs. Moreover, it may well help us to take the lead in stimulating interaction with other disciplines of crucial relevance to environmental issues. In the words of Ehrlich (1989), “Here, as ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 257 in virtually all aspects of the application of the theory of population biology for the benefit of the humanity, we must either gain the cooperation of social scientists or invade their turf. The former is infinitely preferable, for obvious reasons.” t t t j r~ Loss of habitat I POPULATION DECLINE I High mortality T FIGURE 2 - Main factors influencing the present decline of the Blue-fronted Amazon in Argentina (see Beissinger & Bucher in press, Bucher in press). Finally, isolated scientists can also benefit from adopting a comprehensive, goal ori¬ ented approach in their applied research projects. By keeping a wide interest and awareness on the whole system within which their own problem is included, they can gain a much better perspective of how their own subject is related to other interact¬ ing factors. In summary, by following a strategic approach to applied problems, research can be conducted in a more productive way, better questions can be asked, and more inter¬ action elicited between disciplines. The use of a strategic approach does not dismiss the value of other kinds of research approaches on the subject. Even if only laterally related to the problem, all research has the potential for making an important contri¬ bution to our understanding of the system, or even to result in an unexpected break¬ through that allows a better way of solving the problem. However, when resources are limiting (as they nearly always are), prioritation and optimization of research efforts becomes an unavoidable necessity. The case of the Blue-fronted Amazon in Argentina As an example of the complexities and implications of using a strategic approach in ornithology, I will briefly discuss the problem of exploitation of the Blue-fronted Ama¬ zon Amazona aestiva in Argentina. 258 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI This species was abundant in the Chaco savannas of Argentina until the beginning of the 1980s, when intensive trapping began as a result of a booming demand for the pet trade. Exportation of parrots from Argentina was declared legal and unrestricted due to the pest status that Argentine authorities assigned to almost all parrot species. As a consequence, about -204,000 Blue-fronted Amazons were exported from Argen¬ tina between 1981 and 1987 (Traffic Uruguay, unpublished report). Although accurate figures are not available, there is circumstantial evidence indicat¬ ing that in fact the population is being rapidly reduced both in numbers and in range (Bucher et al. unpublished report to the World Wildlife Fund). The potential for the species becoming endangered has been under close scrutiny by international conser¬ vation organizations, and the World Wildlife Fund has sponsored a research program aimed at verifying the sustainability of such intense exploitation. The strategic approach already described shows the following to be the main factors influencing the present decline of the Blue fronted Amazon in Argentina (Figure 2) (Beissinger & Bucher in press, Bucher in press): a) Strong demand: Parrots have become fashionable pets in Europe and the United States. As a result, a sustained demand and associated high prices transformed the parrot trade to a very profitable business. The trade chain extends from the campesino in the Chaco, where each fledgling sells for about US $7, to the pet stores in developed countries, where each Amazon sells for around $400. b) The legal pest status assigned by Argentinean authorities to the species allows for unlimited export quotas from Argentina. c) Severe nesting habitat destruction is caused by local campesinos who destroy the nesting cavity or even cut the tree in order to gain easy access to the nestlings. d) Land tenure problems on both public and private land facilitate unrestricted ex¬ ploitation of wildlife in the Chaco savannas, as well as the destruction of good tree habitat for parrots. e) There is a lack of expertise to implement sustainable exploitation schemes (ranch¬ ing) which could offer a viable alternative to the present irrational exploitation. f) Superimposed on the problem of parrot overexploitation is a continuing process of forest cutting in the area. This poses a serious long-term threat to the survival of the species in the region. Forest cutting averages about 300,000 ha/year af¬ fecting mostly mature forest, a preferred breeding habitat for Amazons. The com¬ bined action of these factors is simultaneously causing important breeding habi¬ tat losses to parrots and high mortality. These factors are the most likely causes of the observed decline. From the analysis of Figure 2, it becomes clear that in order to stop the present population decline, all factors require simultaneous con¬ sideration. Unless all of them are managed properly, advances in only some of them will probably not help to achieve the ultimate goal. For example, even if a total export ban is dictated, high international demand will continue to exert a pressure that may be solved via sustainable ranching or diverted into poaching and smuggling. Unsolved land use problems may also prevent otherwise well designed sustainable ranching programs from being successful. Furthermore, unless the problem of deforestation and land use are considered, long-term sur¬ vival of the species will continue to be threatened. In order to optimize efforts and resources, each of the detected factors can in turn be rated according to its im¬ portance in terms of achieving the final goal, and also the kind of action it requires be considered. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 259 Actions may include not only research, but also education, legal enforcement, regional development planning, etc. Although in many cases those actions are not within the capabilities and commitments of a research team, adequate dissemination and trans¬ fer of the findings to the interested sectors may result in an important contribution to the achievement of the final goal. Once the preliminary model has been obtained, and all the known main driving forces and key factors identified, each one can in turn be analyzed following the same pro¬ cedure, until each factor becomes equivalent to a research project. For example, a lack of know-how of sustainable management can in turn be subdivided into several interacting components. When this stage in the process is reached, the need for modelling and utilization of already existing experience becomes clear. By keeping each research project within the context of the matrix, it is much easier to delineate and conduct each one of the research projects in an interactive way. For example, parrot population models, vegetation regeneration models, and international demand estimates need to be highly interactive in order to estimate exportation quotas or pro¬ duction costs in a given year. Moreover, a careful analysis of experiences from similar situations in equivalent species or equivalent ecosystems may provide significant clues and allow at least preliminary approximations during the first stages of the project. The whole process can be iterated as long as necessary, not only for the initial phases of the management program, but also during its implementation in order to respond in a flexible and adaptive way to unexpected outcomes. Management recommendations can be produced at any moment during the process, which hopefully will become more and more refined and specific with time. For exam¬ ple, the preliminary assessment clearly indicates that a total ban on trade should be imposed immediately in order to stop a rapid process of habitat deterioration and exploitation, and the reopening of the market should be considered only after suffi¬ cient experience on sustainable management has accumulated. At the same time, research should be initiated on agricultural damage evaluation and non-lethal ways of controlling bird damage. Finally, information transfer to the sectors involved in for¬ est management and conservation is urgently required in order to deal with the long¬ term deforestation problem. A traditional approach to the same problem risks overemphasizing a few aspects of high academic interest or those that are easier to observe (probably within the bio¬ logical side), and may tend to ignore others that are more difficult or less interesting to analyze but equally important in terms of the final goal (such as migration, food availability, or land tenure problems). Concluding remarks 1) Applied research in ornithology has already made important contributions to man¬ agement. However, the present needs require increased efforts from ornitholo¬ gists to solve our dramatic environmental crisis. 2) Ornithologists should not allow specialization to narrow their interests to the point that they lose the possibility of interacting with the real world where the birds live. 3) Involvement in applied problems by ornithologists requires not only commitment but also adequate approaches and techniques. Applied research programs would 260 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI benefit from developing a strategic, interactive, goal oriented approach, both at the individual and (ideally) at the inter-disciplinary team level. However, it should not lead us to forget that techniques alone are not enough. The best of tech¬ niques, unless guided by a clear vision of the fundamental issues and by a con¬ cept that gives them form, can turn solutions into larger problems. 4) Interdisciplinary, interactive, goal oriented research requires changes of attitudes and policies of individuals, research institutions, and funding organizations, which may prove difficult to implement, although not impossible. However, neglecting the need to make applied environmental research more efficient in a rapidly changing world may have serious costs in misspent resources and lost opportu¬ nities. A final comment There is no doubt that the world is heading into more, greater, and perhaps unex¬ pected environmental problems. It is also clear that time is running out, since we are involved in processes that behave in an exponential fashion. We need new attitudes, more skills, and more commitment if we expect to deal with these problems with some success. We also need to play a more active role to force the real world to incorpo¬ rate systematic and objective analysis of environmental sciences. Certainly this is not an easy task. Coming from the neotropics, where the conservation battle is being rapidly lost, I cannot be very optimistic. However, we do not have many options left. Otherwise, we ornithologists will have to satisfy ourselves with providing detailed and useless descriptions of the causes of extinction of an increasing number of bird spe¬ cies. There is some room for hope if we remember that ornithologists have already shown their ability to provide important insight and influence in helping the world to become aware of the environmental problems. We can, and probably we must continue tak¬ ing the lead. Let us then try to find innovative ways of putting science and practice together for the benefit of birds and mankind. ACKNOWLEDGEMENTS I wish to thank Steven Beissinger, Terence Boyle, Peter Myers, and John A. Wiens for their useful discussions and comments when preparing the manuscript. I am also grateful to Brian Huntley for having given me the opportunity of learning from his out¬ standing skills and success when in charge of the South African Programme for Eco¬ system Research, which had great influence on my conceptions on goal-oriented ecology. This review was prepared while on sabbatical leave at Colorado State Uni¬ versity working with John A. Wiens. I thank the World Wildlife Fund U.S. and the Consejo Nacional de Investigaciones Cientificas y Tecnicas of Argentina for financ¬ ing my sabbatical. LITERATURE CITED BEISSINGER, S.R., BUCHER E.H. In Press. Sustainable harvesting of parrots for conservation. In Beissinger, S.R., Snyder, N.R.F. (Eds). New World parrots in crisis: solutions from conservation biol¬ ogy. Washington, DC, Smithsonian Institution Press. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 261 BELL, B.D. (1991). Recent avifaunal changes and the history of ornithology in New Zealand. Acta XX Congressus International^ Ornithologici. BUCHER, E.H. In press. Neotropical parrots as agricultural pests. In Beissinger, S.R., Snyder, N.R.F. (Eds). New World parrots in crisis: solutions from conservation biology. Washington, District of Colum¬ bia, Smithsonian Institution Press. EHRLICH, P.R. 1989. Discussion: ecology and resource management - is ecological theory any good in practice?. Pp. 306-318 in Roughgarden, J., May, R.M., Levin, S.A. (Eds). Perspectives in ecologi¬ cal theory. Princeton, Princeton University Press. HOLLING, C.S. 1978. Adaptive environmental assessment and management. John Wiley & Sons. New York. NAPPER, J.L. (compiler). 1989. Department of Conservation Science project summaries. Science and Research Internal Report No. 46. Parts 1 and 2. Wellington, New Zealand Department of Conserva¬ tion. MURTON, R.K. 1971. Man and birds. Collins. London. POPPER, K.R. 1965. The logic of scientific discovery. New York, Harper & Row. ROUGHGARDEN, J., MAY, R.M., LEVIN, S.A. (Eds). 1989. Perspectives in ecological theory. Princeton, New Jersey: Princeton University Press. SCHLESINGER, W.H. 1989. Discussion: ecosystem structure and function. Pp. 268-274 in Roughgarden, J., May, R.M., Levin, S.A. Perspectives in ecological theory. Princeton: New Jersey, Princeton University Press. SLOBODKIN, L.B. 1988. Intellectual problems of applied research. BioScience 38:337-342. WILLIAMS, M. 1990. Presidential address: Ecology in an advocate’s age. New Zealand Journal of Ecology 13:1-7. 262 ACTA XX CONGRESSUS INTERNATIONAL^ ORNITHOLOGICI - . ' / ' ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 263 PLENARY LECTURE RESPIRATION OF AVIAN EMBRYOS AT HIGH ALTITUDES CYNTHIA CAREY 264 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 265 RESPIRATION OF AVIAN EMBRYOS AT HIGH ALTITUDE CYNTHIA CAREY Department of Environmental, Population, and Organismic Biology, University of Colorado, Boulder, CO 80309 USA ABSTRACT. The ability of birds to colonize a wide diversity of habitats, including some of the most hostile ones of earth, depends in part upon the achievement of levels of gas exchange between the embryo and the environment within tolerance limits. The reduction in barometric pressure at high al¬ titude causes avian embryos to develop in gaseous conditions that differ considerably from those at which embryos develop at sea level. Conservation of water appears to be a more important priority at moderate altitudes, whereas improvement of oxygen delivery becomes a primary selective force at higher altitudes. Modifications of eggshell conductance to gases compensate in part for variation in gaseous diffusion, particularly at low altitude, but specializations in embryonic physiological properties, such as blood oxygen carrying capacities and capillary density of the chorioallantoic membrane, play increasingly important roles in fostering growth and development at high altitudes. Keywords: Avian egg, eggshell conductance, gaseous diffusion, pore area, avian reproduction. INTRODUCTION Birds are among the most successful vertebrates in terms of their ability to invade and to colonize a wide variety of habitats, including some of the most hostile ones. Their ability to do so has depended on their capacity to reproduce successfully in a variety of climatic conditions. Successful reproduction, particularly in hostile environments, requires a suite of specializations that foster survival of offspring through incubation, hatching, fledging, independence from the adults, and ultimately breeding themselves. Focusing specifically on embryos, survival to hatching depends importantly on whether the requirements of the embryo are met within its tolerance limits. Adults periodically turn eggs, provide heat, and defend eggs from predators (Drent 1975). The egg, prepackaged before laying, contains all the nutrients, water and ions the embryo needs for growth and development to hatching. The final embryonic require¬ ment during incubation is for gas exchange between the interior of the egg and the external environment. Twenty years ago, relatively little was known about embryonic gas exchange and its relation to avian biology. The pioneering work of Hermann Rahn, Charles Paganelli, Amos Ar, and their colleagues, who have combined to publish over 80 papers on the subject, has led to a veritable explosion of knowledge about avian eggs. The goal of this paper is to summarize a small part of this field by considering the interrelations of gas exchange, shell structure and embryonic physiological properties of species breeding at high altitudes. 266 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI GAS EXCHANGE AT SEA LEVEL Before dealing with the particular problems encountered by avian embryos at high altitude, it is appropriate to review briefly the process of gas exchange in lowland embryos. The embryo exchanges 02, C02, and water vapor with its environment (Wangensteen & Rahn 1970/71). These gases travel principally by the process of diffusion down partial pressure gradients (Wangensteen et al. 1970/71, Wangensteen 1972). The gradients for 02 and C02 are established by the metabolism of the embryo; as it consumes 02 and produces C02 inside the shell, partial pressures of these gases (p02 and pC02) inside the egg decrease and increase, respectively, relative to pres¬ sures outside the egg (Wangensteen 1972). A partial pressure gradient for water vapor (APh2o) exists across the eggshell because the interior of the egg is fully satu¬ rated with water vapor, whereas the nest environment of almost all species, except those laying eggs in soil or rotting vegetation (Seymour & Ackerman 1980), is not fully saturated (Wangensteen & Rahn 1970/71). Oxygen travels into the egg while C02 and water vapor move outward through common pores in the shell (Paganelli et al. 1978). The factors which determine the rate of diffusion of a gas (M, crrPSTPD.sec1) are described by a modification of the Fick equation (Wangensteen 1972; Paganelli et al. 1975): M = (D/RT) • (Ap/L) • AP (1) where D = binary diffusion coefficient (cm2. sec1), RT = gas constant and absolute temperature (cm3STPD.cm Ttorr1), Ap = effective pore area (cm2), L = length of dif¬ fusion path, or shell thickness (cm), and AP = partial pressure difference of gas across the shell (torr). The terms (D/RT). (Ap/L) are often combined into the term “G” (cm3. sec1. torr1) representing the conductance, or the diffusive capacity of the egg¬ shell to each gas (Ar et al. 1974). Therefore, eq. 1 can be rewritten as: M = G • AP (2) / Conductance is conventionally reported as standardized to 760 torr so that G of a species can be compared with that of other species under identical conditions of pres¬ sure. The fact that 02 diffuses in the opposite direction from C02 and water vapor places opposing, or mutually antagonistic, requirements on the structure of the shell. The conductance must be large enough to allow sufficient 02 to diffuse inward for support of metabolic requirements, but restrictive enough to prevent excessive losses of water vapor and C02 from the egg. Most workers have agreed that control of losses of water vapor and/or C02 has probably been a more important priority for eggshell design at low altitudes than has been maximization of 00 delivery (Vleck et al. 1979, Vleck et al. 1980, Ar & Rahn 1980, Paganelli & Rahn 1984). The role the shell plays in regulation of embryonic gas exchange is sufficiently impor¬ tant that G has apparently evolved in conjunction with two other interrelated traits: duration of incubation period (I) and egg mass (W). Ar & Rahn (1978) observed that ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 267 G is inversely proportional to I for a given W. If the eggs of two species have equiva¬ lent mass but different incubation periods, G will vary inversely with I with the result that total gas exchange of the two eggs will be comparable by the end of incubation (Ackerman et al. 1980). Conductance has also apparently evolved in conjunction with the gaseous environment in which the eggs of a particular species are laid. Average G of eggs laid in very humid, hypoxic, or hypercapnic environments is substantially larger than predicted on the basis of I and W (Lomholt 1976, Birchard & Kilgore 1980, Seymour & Ackerman 1980). Therefore, the average G of a given species was presumably selected by differential mortality of embryos in eggs laid with excessively high or low conductance. However, relatively little is known about the tolerance limits of avian embryos. Water loss during incubation achieves two important results: formation of an air cell at the blunt end of the egg and maintenance of relative hydration of contents. As water diffuses out of the egg, it is replaced by an equivalent volume of air (Romijn & Roos 1938). Just prior to pipping the shell, the embryo pips into the air cell and uses the volume to expand its lungs, thus beginning the conversion from diffusive to convec¬ tive respiration (Vince & Tolhurst 1974). The egg contents of altricial and precocial eggs at laying are approximately 75 and 85% water, respectively (Ar & Rahn 1980). These relative hydration levels are maintained through pipping by water loss. The amount of water lost compensates for both the reduction in solids catabolized and the production of metabolic water (Ar & Rahn 1980). Ar & Rahn (1980) have suggested that water loss from the egg must be precisely regulated within narrow limits for optimal hatching. This proposal is based on obser¬ vations that the hatchability of Domestic Chicken ( Gallus domesticus) embryos de¬ creased markedly if too much or too little water is lost during incubation (Lundy 1969; Tullett & Board 1982) and that eggs lose about the same average amount of mass as water vapor (17%) during incubation (Ar & Rahn 1980). Apparently, only one study currently exists testing tolerances of embryos of wild species to variation in water loss: Red-winged Blackbird ( Agelaius phoeniceus) embryos hatch successfully from eggs losing between 7.4 and 33.0% of initial mass and tolerate wider variations in G and daily water loss than are observed in populations of this species in the field (Carey 1976). The interrelation of G, W, and I also results in general similarities in 02 and C02 ex¬ change for most species. The amount of 02 consumed per g embryo (Ar & Rahn 1 978) and the Po2 and Pco2 across the eggshell at comparable stages of incubation are similar in almost all lowland eggs (Rahn et al. 1974, Ar & Rahn 1978). Addition¬ ally, the final levels of 02 and C02 in the air cell at pipping (Pao2 and Paco2) fall within narrow limits around 104 and 40 torr, respectively (Rahn et al. 1974, Hoyt & Rahn 1980). The importance of the latter two observations relates to the fact that the Pao2 and Pco2 prior to the onset of pulmonary respiration closely resemble those found in the lungs of hatchlings and adults (Tazawa et al. 1971, Wangensteen 1 972, Rahn et al. 1974). Therefore, G not only regulates appropriate levels of gas exchange but also fosters preparation of the embryo for the onset of aerial respiration (Wangensteen et al. 1970/71 , Rahn et al. 1974). 268 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI No experiments have yet tested the tolerances of embryos of wild species to varia¬ tion in ambient Po2 and Pco2, but hatchability of domestic fowl embryos severely de¬ clines if a small portion of the eggshell is blocked just prior to the onset of pulmonary respiration (Tazawa et al. 1971). Hatchability of domestic fowl eggs is severely re¬ duced below 15% 02 or above 40% C02 (Lundy 1969). Variation in ambient Po2 could be especially detrimental for avian embryos, because it forms the upper end of the gradient for diffusion of 02 through the shell and ultimately into the tissues. GAS EXCHANGE OF EMBRYOS AT ALTITUDE Biogeographers have suggested that existing montane populations of birds have de¬ scended from groups that have moved up and down altitudinal gradients as mountain ranges have risen or eroded and as global climatic changes have occurred (Mayr & Diamond 1976, Vuilleumier 1986). At least 27 species of birds are known to nest at altitudes between 4000 and 6550 m (Rahn 1977, Harris 1981, Carey et al. 1987, 1989a, b, 1990). Birds breeding at both high altitude and high latitudes face problems of cold, seasonality in food availability and snow cover (Carey 1988). The unique fea¬ tures of the montane environment are low barometric pressure and its associated con¬ sequences: hypoxia, low water vapor pressure, and intense ultraviolet light (Mani 1962). Aggazzotti (1913) measured weight loss of some chicken eggs at Turin, Italy, and then again after transport to 2900 m on Monte Rosa. He found that the eggs lost sub¬ stantially more weight at 2900 m than at low altitude and surmised that the excessive weight loss was due to increased water loss from the eggs. Rahn & Ar (1974) were the first to identify the cause of this phenomenon. As barometric pressure (PB) de¬ creases with increasing altitude, the diffusion coefficient (D in Eq. 1) increases in¬ versely (Reid & Sherwood 1966, Paganelli et al. 1975, Paganelli 1980). Therefore, if Eq.2 is rewritten with PB representing the barometric pressure at the breeding loca¬ tion: / G (760) M = - • AP (3) pB one can see that the “effective” conductance of an egg (G) for any gas will be higher at altitude than at sea level. If an egg is transported from sea level to roughly 5500 m, where the PB is half that at sea level, losses of C02 and water vapor will be twice the amount at sea level, and 02 will diffuse into the egg twice as rapidly (Paganelli et al. 1975). However, an increase in the diffusion rate of 02 into the egg will only par¬ tially compensate for the fewer total number of 02 molecules present in ambient air at low Pb (Visschedijk et al. 1980). Since the data available on tolerance limits of low altitude embryos suggest that vari¬ ation in gas exchange caused by the increase in D at altitude might prove detrimen¬ tal, or even lethal, to embryos at high altitude, it is interesting to explore what mecha¬ nisms might compensate for the effect of D on gaseous diffusion (Rahn & Ar 1974). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 269 If prevention of excessive water and/or C02 losses were the most important priority, one or more of the following might be implemented: 1. a decrease in G by increasing L or decreasing Ap, which would have the effect of increasing the resistance of the shell to gas diffusion; 2. a decrease in the APh2o and APco2 by raising Ph2o and Pco2 in the nest environ¬ ment. This could most easily be accomplished by increasing adult attentiveness on the nest and trapping these gasses in the nest microenvironment. (Note: use of either or both 1 and 2 will restrict availability of 02 to the embryo); 3. increasing the initial water content of eggs and varying the buffering capacity of blood. If optimization of 02 delivery were the most important priority one or more of the following might be utilized: 1) an increase in G by decreasing L or increasing Ap, which would increase the re¬ sistance of the shell to diffusion of 02, 2) an increase in APo2 by decreased nest attentiveness, (Note: use of either or both 1 and 2 would increase water and C02 loss), 3) an increase in 02 delivery to the cells by an increase in blood 02 carrying capac¬ ity, variation in hemoglobin oxygen affinity, increased capillarity of the chorioallantoic membrane, or other possible specializations at the cellular or bio¬ chemical level. Moderate altitudes The existing data comparing eggs laid by montane species with their conspecific or congeneric relatives nesting at sea level support the contention the conservation of water and/or C02 is the most important priority for birds colonizing montane habitats up to about 3600 m. A number of studies have indicated that the average conduct¬ ance to water vapor (Gh2o, standardized to 760 torr), of both domesticated chickens and montane populations of Red-winged Blackbirds Agelaius phoenicius and Robins Turdus migratorius breeding up to 3450 m progressively declined with PB (Figure 1) (Rahn et al. 1977, Carey et al. 1983, Leon-Velarde et al. 1984a). A few other stud¬ ies have shown either a decrease in Gh2o in montane eggs that did not parallel the reduction in PB or even an increase with altitude (Packard et al. 1977, Sotherland et al. 1980, Taigen et al. 1980), but these studies used eggs of unknown age. Since Gh2o changes in early incubation in a number of species (see Carey 1983), the results could have reflected more the effect of egg age than an adjustment to altitude (see Carey 1980). The decrease in Gh2o with increasing altitude to about 3600 m was largely attributable to a decrease in Ap than an increase in L (Rahn et al. 1977, Carey et al. 1983, Leon- Velarde et al. 1984). Shell thickness itself is under mutually antagonistic selective pressures; it must be strong enough to support the mass of the incubating adult, yet thin enough for the embryo to pip through the shell and hatch. It is likely that these constraints made it difficult to alter shell thickness as a mechanism for counteracting the change in gaseous diffusion coefficients at altitude. 270 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI By approximating the reduction in PB, Gh2o (standardized to 760 torr) compensates for the increase in D, with the result that the “effective” Gh2o of montane eggs on the breeding grounds is the same as in lowland eggs. As a result, daily water losses from naturally incubated eggs of Red-winged Blackbirds and White-crowned Sparrows Zonotrichia leucophrys are independent of altitude to 3050 and 3660 m, respectively (Carey et al. 1983, Carey, unpubl. data). Therefore, these embryos develop in the same hydric environment as lowland embryos; without the decrease in Gh2o (stand¬ ardized to 760 torr), water losses would be at least 30% higher than they are. Rela¬ tive water content of freshly laid eggs of Red-winged Blackbird and Robin eggs and adult attentiveness of Red-winged Blackbirds did not vary significantly from low alti¬ tude to about 2900 m (Carey et al. 1983). FIGURE 1 - Water vapor conductance (standardized to 760 torr) of eggs laid at various altitudes expressed as a proportion of the mean conductances of a sea level conspecific or congeneric species. The mean conductances of the sea level groups are set at 1.0. The line represents the predicted relative conductance at any altitude, assuming that the con¬ ductance equals 1 .0 at sea level and is reduced in exact proportion to the barometric pres¬ sure. The symbols represent the following publications in which actual data on these spe¬ cies can be obtained: filled circles = Agelaius phoenicius and Turdus migratorius (Carey et al. 1983); open circle in a filled square = Larus serranus and Plegadis ridgwayi (Carey et al. 1987); filled star = Fulica americana peruviana (Carey et al. 1989a); open circle = Anas versicolor puna (Carey et al. 1989b); open square = Chloephapa melanoptera, Anas flavirostris oxyptera, and Rollandia rolland (Carey et al., 1990); filled half circle= Zonotrichia leucophrys (Carey; unpubl. data); filled triangle = Larus serranus (Leon-Velarde et al. unpubl data). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 271 Since the “effective” G at altitude approximated the comparable value at sea level, yet the total number of 02 molecules in the ambient air at altitude are fewer, the air cell 02 tension (Pao2) of Red-winged Blackbird eggs dropped progressively from about 80 torr at low altitude to about 50 torr at 2900 m (Carey et al. 1982). The Pao2 serves as the upper end of the gradient for diffusion of 02 into the blood (Wangensteen 1982). Low Pao2 was associated with decreased oxygen consumption (Yo2), prolonged incu¬ bation periods, and decreased hatchling masses of chicken embryos at 3800 m (Wangensteen et al. 1974), but Yo2 of Red-winged Blackbird embryos at 2900 m was statistically indistinguishable from that at lower altitudes, and hatchling masses and incubation periods of Red-winged Blackbirds, White-crowned Sparrows, and Horned Larks Eremophila alpestris were independent of altitude to 2900, 3475, and 3600 m, respectively (Carey et al. 1982). Therefore, it appears that control of water loss is the most important priority for shell design at altitudes up to around 3600 m. Presumably, loss of C02 from the shell would also be maintained at sea level values, although no studies have investigated C02 losses directly in eggs at moderate altitudes. Although availability of 02 is limited by reduction of Ap, embryos apparently still get enough 02 for normal metabolism and growth. The question has been addressed whether the change in Gh2o with altitude results from detection by the female of variation in PB or one of its correlates, or whether the characteristic G of a population results from long-term selection for females laying a genetically-fixed shell appropriate for the physical conditions at the breeding location. Rahn et al. (1982) transported chickens from a breeding colony at 3800 m to 1200 m and found a significant increase in GH2othat approximated the increase in PB. How¬ ever, some statistical weaknesses existed in the data: unequal numbers of eggs from each female were pooled for comparison of low and high altitude groups, and data from females which didn’t produce eggs at the lower altitude were included in the montane average. Other studies have found no effect of variation in PB on Gh2o. Leon- Velarde et al. (1984b) found no significant difference between average Gh2o of eggs laid by chickens which had been transported from sea level to 2800 m within 24 hr after hatching and those laid by the sea level stock from which they had originated. Average Gh2o of eggs laid by Bengalese Finches Lonchura striata and quail Coturnix coturnix did not change between sea level and after transport to 2900 m, when eggs of the same female were compared before and after transport (Carey et al. 1984). Therefore, it is probable that the reduction in Gh2o observed in montane groups of chickens and wild birds most likely results from long-term selection for genetically fixed shell characteristics. Genetic control of shell features has also been documented by Sotherland et al. (1979) and Bucher & Barnhart (1984). High altitude We now move to consideration of the question concerning how birds breeding above 3600 m cope with the effect of increasing D on gaseous diffusion. The data suggest that priorities shift from conservation of water and/or C02 to improvement of 02 de¬ livery. Conductance (standardized to 760 torr) of eggs laid at higher altitudes is not decreased to the same extent as in barometric pressure at the breeding location (Fig¬ ure 1). In fact, G of some montane species exceeds that of their lowland counterparts, resulting in a curvilinear relation between G and PB. If G (standardized to 760 torr) of eggs laid at altitudes above 4000 m were reduced to the same proportion as PB so 272 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI that the effective G at altitude was the same as at sea level, the amount of 02 reach¬ ing the embryo would be inadequate to support costs of both maintenance and growth (Carey et al. 1989a). Most of the comparisons in Figure 1 for eggs laid above 4000 m were made with averages of montane eggs and those of similarly-sized eggs of lowland congeneric speqies (Carey et al. 1987, 1989b, 1990). These types of com¬ parisons were necessitated by the fact that most species breeding at very high alti¬ tudes do not breed over broad altitudinal gradients and lack conspecific lowland populations. Therefore, probably the most accurate indication concerning how Gh2o of a high altitude population compares with a lowland conspecific population is pro¬ vided by American Coots Fulica americana peruviana. Average Gh2o (standardized to 760 torr) of American Coot eggs laid at 4150 m was 107% of the sea level value, whereas PB at the montane location was 60% of that at sea level (Carey et al. 1989a). Since G of eggs of many species laid above 3600 m does not track the reduction in PB at the laying location and undercompensates for the increase in D, the “effective” G, caused by the influence of D at altitude, is higher than that at sea level. As a re¬ sult, water losses from naturally incubated eggs of American Coots at 4150 m and Andean Gulls Larus serranus and Puna Ibises Plegadis ridgwayi at 4400 m were sub¬ stantially greater than that of their sea level counterparts. The effect of the higher rate of water loss on the embryo is difficult to assess because data on incubation periods of these montane groups, needed to calculate the total water loss during incubation, are not available. If incubation periods of these montane groups are equivalent to those of their lowland counterparts, Andean Gull and American Coot eggs at 4400 and 4150 m, respectively, would lose about 20 and 18%, respectively, of their initial mass as water vapor during incubation (Carey et al. 1987, 1989a). These percentages are well within the range for total water loss established for species breeding at low alti¬ tudes (Ar & Rahn 1980). Incubation periods of gulls and coots would have to be sub¬ stantially prolonged for tolerance limits exhibited by Red-winged Blackbird embryos to be approached (Carey 1986). Relative water content of freshly laid and partially incubated Andean Gull and American Coot eggs fell within the range of values known for lowland precocial and semi-precocial eggs (Carey et al. 1987, 1989a). The lack of variation in relative water content of these montane eggs could result from one of at least two possible causes: 1) no selection has existed for adding extra water to the egg as a mechanism for offsetting increased water loss because embryonic tolerance limits are not taxed by the rates of water losses at these altitudes, or 2) dilution of egg content may cause mortality by interfering with cellular processes. Studies on eggs experiencing severe rates of water loss are necessary to discriminate between these or other possibilities. Despite the improvement in effective G by an increase in Ap in eggs of some species laid above 4000 m (Carey et al. 1987, 1989a, 1989b), montane embryos develop in a hypoxic environment. In fact, variation in shell characteristics plays an increasingly less important role at higher altitudes because no possible adjustment can create gaseous conditions similar to those at sea level. Instead, embryonic physiological properties become increasingly more important for adjusting to abnormal gaseous conditions inside the shell. The Pio2, or the “effective” 02 tension outside the egg at sea level is approximately 148 torr (Wangensteen & Rahn 1970/71). The Pao2 inside an egg declines from approximately that value in freshly laid eggs to about 100 torr, ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 273 or slightly less in small eggs, by the end of incubation due to the increased demand for 02 by the growing embryo (Wangensteen 1972, Vleck et al. 1979, Hoyt & Rahn 1980, Bucher & Barnhart 1984). The Pio2 at 4150 m is roughly 84 torr (Carey et al. 1989a). The Pao2 of the aircell of fresh montane eggs approximates 84 torr and then declines from that level during incubation (Carey et al. 1989a, 1991a). Therefore, even at the beginning of incubation, the Pao2 of montane eggs is substantially below that of lowland eggs at the end of incubation. Embryonic oxygen consumption (Yo2) in such hypoxic environments has exhibited two types of patterns. One pattern is illustrated by chicken and American Coot embryos at 3800 and 4150 m, respectively; Yo2 was depressed compared to that of their low¬ land counterparts (Wangensteen et al. 1974, Carey et al. 1989a). The growth rates of chicken embryos were slower and hatchling masses were smaller than those of their sea level relations (Wangensteen et al. 1974). Embryonic masses of American Coots at 4150 m were similar to, but their incubation periods were probably longer than those of lowland American Coots (Carey et al. 1989a). Wangensteen et al. (1974) have hypothesized that the depressed Yo2 of montane chicken embryos is an adaptation which maintains the Pao2 as high as possible with the result that the APo2 driving 02 into the blood as large as possible. However, since prolongation of the in¬ cubation period exposes eggs to greater risks of predation and since smaller hatchlings are often at a disadvantage for survival (Packard 1990), an alternative explanation for the low Yo2 is that montane chick and coot embryos are unable to obtain sufficient 02 for maintenance of normal growth and metabolism in the hypoxic environment inside the shell. Chicken embryos are remarkably sensitive to hypoxia. Hatchability decreases below sea level values at altitudes as low as 1600 m and drops to extremely low levels above 3000 m (Moreng 1983, Leon-Verlarde et al. 1984a). Coot embryos appear quite viable and hatchability seems high at 4150 m (Carey et al. unpublished data). Since American Coots breed over altitudinal gradi¬ ents of at least 4150 m in the Peruvian Andes, the depressed Yo2 of montane coot embryos may be symptomatic of the effect of gene flow from lower altitudes which may prevent genetic specializations to the physical environment at high altitudes. The other pattern in Yo2 is exhibited by Puna Teal Anas versicolor puna embryos. The level of Yo2 of these individuals was comparable to that of lowland chickens, and in fact, was higher at comparable embryonic masses than the Yo2 of lowland American Coots (Carey et al. 1989a, 1991a). While comparative data on hatchling masses of Puna Teal embryos are not available, the incubation period of montane eggs appears to be similar to that reported for a captive population held at sea level for several generations (Carey et al. 1991a). The distribution of Puna Teal is largely limited to the high Andes; they are replaced at lower altitudes by Silver Teal Anas versicolor ( Blake 1977). Their limited altitudinal distribution appears to have fostered development of specializations that improve abilities to maintain “normal” levels of Yo2 in very hypoxic environments. Puna Teal embryos are able to maintain higher Yo2 at all embryonic masses than American Coot embryos despite having a smaller pressure head ( Po2) for diffusion of 02 into the blood; Pao2 of 15 g Puna Teal and American Coot embryos was approxi¬ mately 43 and 55 torr, respectively (Carey et al. 1989a, 1991a). However, this differ- 274 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI ence in Po2 does not result in a difference in the Po2 of arterialzed blood (Pao2) of embryos of these two species, because Puna Teal eggs have a smaller resistance to 02 diffusion between the air cell and the chorioallantoic blood. Chicken embryos at sea level have a large PAo2-Pao2 difference of approximately 52 torr which can be de¬ creased to some extent in hypoxic conditions (Piiper et al. 1980). The cause of this large resistance is hypothesized to be either a small arteriovenous shunt or a water layer between the inner shell membrane and chorioallantoic membrane, in which 02 is relatively insoluble (Piiper et al. 1980). The PAo2-Pao2 difference of montane Ameri¬ can Coot and Puna Teal 15g embryos was about 25 and 10 torr, respectively (Carey et al. 1991a, 1991b). The mechanism by which this resistance is lowered in these two species is unknown. Vertebrates in hypoxic circumstances increase the oxygen-carrying capacity of blood with a variety of mechanisms. Compared with values from low altitude congeneric or conspecific groups, adult birds living all or part of the year at 3000 m exhibited sig¬ nificantly higher hematocrits, red-blood cell counts, and hemoglobin concentrations, although it is unclear whether these differences were a response to hypoxia, pro¬ longed cold, or both (Carey & Morton 1976). Chicken embryos significantly increased hematocrit after the shell was partially covered with a material impermeable to 02 (Tazawa et al. 1971). Hematocrits of montane Andean Coot embryos were slightly higher at all embryonic masses than were those of their lowland counterparts (Carey et al. 1 991b). Hematocrits of 1 6g Puna Teal embryos were higher than those reported for 18-day chicken embryos and for Canada Goose Branta canadensis and Bar¬ headed Goose Anser indicus embryos incubated at 1600 m (Snyder et al. 1982, Carey et al. 1991a). Other possible mechanisms of enhancing oxygen delivery to the cells, such as hemoglobin 02 affinity, genetic variation in hemoglobin, effect of organic phosphates on hemoglobin affinity, variation in blood flow, myoglobin concentrations, and capillary density of the chorioallantoic membrane, have not yet been investigated in embryos of wild birds at altitudes. Genetic differences in hemoglobin and hypoxia appear to be particularly likely features in which specializations of montane embryos could be identified (Baumann 1984) / Despite the specializations which may exist for maximizing 02-carrying capacity of the blood of montane American Coot and Puna Teal embryos, the Po2 of the venous blood returning from the tissues to the chorioallantoic membrane ranges from about 3 to 10 torr in mid- to late incubation (Carey et al. 1991a, 1991b). Since venous oxygen ten¬ sions are thought to reflect the Po2 of the tissues (Tenney 1974), these embryos have the remarkable ability not only to maintain life but also to grow at tissue oxygen ten¬ sions that would be lethal on a prolonged basis for adult birds and mammals. Specializations at the cellular and biochemical level which foster this ability have not yet been identified. Carbon dioxide production increases with embryonic mass throughout all or most of incubation as an end product of aerobic metabolism (Wangensteen 1972). The in¬ crease in D resulting in the higher effective G at high altitudes causes CO, to diffuse more rapidly from the egg than at sea level. As a result, the air cell Pco2 (Paco2) of montane American Coot eggs was about 10-20 torr below that of their lowland coun¬ terparts by the end of incubation (Carey et al. 1989a). The Paco2 of Puna Teal eggs ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 275 was higher at any given embryo mass than that of the coot eggs due to the higher rate of C02 production (Carey et al. 1991a). The Paco2 forms the lower end of the gradi¬ ent for diffusion of C02 from the blood into the air cell, and, as such, plays an impor¬ tant role in determining blood pH. Despite the differences between Paco2 of montane coot and teal and lowland coot and chicken values, blood pH of the montane embryos did not differ significantly from lowland values (Tazawa 1971, Carey et al. 1991a, 1991b). Achievement of constant blood pH at different Pco2 is probably due to a re¬ duction in the [HC03- ] in the blood of montane embryos, but the mechanism by which this occurs is unknown. CONCLUSIONS The modifications in the eggshell of eggs laid at moderate altitudes result in rates of water loss and probably C02 loss that are similar to those at low altitudes, at the cost of reduction in the Po2 inside the shell. Despite such hypoxia, embryos at moderate altitudes are able to maintain levels of Yo2 that promote normal development and hatching. At altitudes above 3600 m, apparently oxygen levels become limiting enough that selection promoted a relatively larger pore area than at moderate alti¬ tudes for some species. This adjustment increased availability of oxygen to the em¬ bryo at the cost of increased rates of losses of C02 and water vapor. The curvilinear relationship between G and PB is one of the few examples in biology in which a lin¬ ear environmental stress is associated with a reversal of an adaptation. This reversal results from a shift in priorities from conservation of water vapor and C02 at lower al¬ titudes to improvement of 02 availability at higher ones. At altitudes over 4000 m, no adjustments in shell features can produce gaseous conditions that are similar to those at sea level; as a result, embryonic physiological properties play a progressively im¬ portant role in promoting survival and growth in gaseous conditions that would be lethal for certain lowland species. No information is available on eggs or embryos of species breeding over 4600 m. Hopefully, further research will provide information on the remarkable specializations fostering successful reproduction at these altitudes which are certain to exist in these groups. ACKNOWLEDGEMENTS Research summarized herein by C. Carey was supported in part by National Science Foundation and National Geographic Society grants. LITERATURE CITED ACKERMAN, R.A., WHITTOW, G.C., PAGANELLI, C.V., PETTIT, T.N. 1980. Oxygen consumption, gas exchange, and growth of embryonic Wedge-tailed Shearwaters (Puffinis pacificus chlororhynchus). Physiological Zoology 53: 210-221. AGGAZZOTTI, A. 1913. Influenza dell’aria rarefatta sull’ontogenesi. Noata 1. La perspriazione delle ova di gallina durante lo sviluppo in alta montagna. Wilhelm Roux Arch. Entwicklungsmech. Org. 36: 633-648. 276 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI AR, A., PAGANELLI, C.V., REEVES, R.B., GREENE, D.B., RAHN, H. 1974. The avian egg: water vapor conductance, shell thickness and functional pore area. 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Respiration Physiology 14: 64-74. WANGENSTEEN, O.D., RAHN, H. 1970/1971. Respiratory gas exchange by the avian embryo. Res¬ piration Physiology 11: 31-45. WANGENSTEEN, O.D., RAFjN, H., BURTON, R.R., SMITH, A.H. 1974. Respiratory gas exchange of high altitude adapted chick embryos. Respiration Physiology 21: 61-70. WANGENSTEEN, O.D., WILSON, D., RAHN, H. 1970/1971. Diffusion of gases across the shell of the hen's egg. Respiration Physiology 11: 16-30. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 279 PLENARY LECTURE CONSTRAINTS ON REPRODUCTION IN ALBATROSSES J. P. CROXALL 280 * " ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 281 CONSTRAINTS ON REPRODUCTION IN ALBATROSSES J. P. CROXALL British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley Road, Cambridge CB3 OET, UK ABSTRACT. The interplay of ecological, behavioural and physiological (chiefly endocrine) factors in regulating the reproductive rate of albatrosses (chiefly Wandering, Black-browed and Grey-headed Al¬ batrosses at South Georgia) is reviewed. The focus is on three key processes: acquisition of breed¬ ing status, breeding frequency and breeding success. Hormonal and related physiological mechanisms preventing sexual maturity persist longer in females than males; acquisition of social maturity takes several more years. Age of first breeding depends on rate of pair formation which chiefly reflects the ability of males to spend time displaying at nests ashore (i.e. foregoing foraging). In basically biennial¬ breeding species endocrine mechanisms preclude females from breeding in years following success in rearing a chick; males are not restricted in this way but are constrained by behavioural factors. Even first-time breeders only show small reductions in breeding success and efficiency compared to expe¬ rienced birds. This is unusual amongst seabirds and suggests that most necessary experience has been acquired during the long periods of immaturity and pair formation. Costs of reproductive strate¬ gies and tactics in survival terms cannot be adequately assessed, partly because full life-time repro¬ ductive success data are unavailable and partly because of major demographic changes over recent decades directly affecting adult and juvenile survival, consequent on incidental mortality associated with fisheries. Keywords: Wandering Albatross, Black-browed Albatross, Grey-headed Albatross, Diomedea, D. exulans, D. melanophris, D. chrysostoma, ecology, behaviour, physiology, endocrinology, sexual ma¬ turity, pair-formation, social maturity, breeding frequency, breeding success, survival, South Georgia, lies Crozet. INTRODUCTION The principal decisions facing a bird contemplating reproduction are how to acquire a partner, at what age (or time) to start breeding, how often to breed, how many eggs to lay and how to maximise and/or optimise the success of each breeding attempt. Many of these problems are inter-linked and all involve interplay of a complex suite of ecological, ethological and physiological factors. Most detailed experimental inves¬ tigations of reproductive strategies and tactics have been conducted on small passerines (e.g. tits Paridae - McCleery & Perrins 1988, Nur 1988, Tinbergen & Daan 1990), which have the advantage of short generation times and relative ease of han¬ dling and measurement but the complication of large and variable clutch sizes. De¬ spite their considerable longevity and long generation times (greatly delaying acqui¬ sition of data on lifetime reproductive success), large non-passerines offer advantages of small and often fixed clutch size and their large size may enhance visibility, rec¬ ognition of individuals by observers and have advantages for certain types of experi¬ mental physiology in the field. Albatrosses, with their long delay before starting breeding, typically biennial breed¬ ing in several species, single-egg clutch and extreme longevity are often regarded as a paradigm of K-selected species. However, for seabirds in general and albatrosses 282 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI in particular there has been little synthesis of information with respect to these adap¬ tations. This paper is a preliminary review of the relationships between ecological, behavioural and physiological (chiefly endocrine) factors in regulating the reproduc¬ tive rate and performance of albatrosses. It focuses on three processes: acquisition of breeding status, breeding frequency, and breeding success. It concludes by con¬ sidering factors influencing survival - the most important characteristic of all in deter¬ mining productivity in long-lived birds. SITES AND SPECIES This paper draws chiefly on research conducted at Bird Island, South Georgia, sig¬ nificantly supplemented by material from French studies at lies Crozet. It features principally the Wandering Albatross Diomedea exulans and subsidiarily the Black- browed and Grey-headed Albatrosses Diomedea melanophris and D. chrysostoma (Table 1). The last two species belong to the main element of the genus Diomedea, often referred to as mollymawks. They are sexually monomorphic, breed in large, dense (often mixed) colonies on grassy slopes or cliffs at sub-Antarctic islands. Wan¬ dering Albatrosses are one of two-three species referred to collectively as great al¬ batrosses. They are twice the size of mollymawks, sexually dimorphic in size and plumage, and breed in dispersed colonies usually on flat or gently sloping grassland at sub-Antarctic islands. All species have very long reproductive seasons but only in the great albatrosses do these attain or exceed one year. TABLE 1 - Duration of breeding season events in Wandering, Black-browed, and Grey-headed Albatrosses at South Georgia. Species Male Weight Female Prelaying attendance Incubation Chick rearing Total Breeding attendance frequency Wandering Albatross 10.6 9.0 27 78 278 383 Biennial Black-browed Albatross 3.9 3.7 16 68 116 200 Annual Grey-headed Albatross 3.8 3.6 26 72 141 239 Biennial ACQUISITION OF BREEDING STATUS Albatrosses have one of the longer periods of sexual immaturity known in birds, with some individuals not joining the breeding population until older than 15 years of age (Croxall 1982, Weimerskirch & Jouventin 1987, Weimerskirch et al. 1987). Even the average individual returns to its natal colony some five years before its first breeding attempt. The role of physiological (endocrine) and behavioural factors in the process of maturation, pair-bond formation and breeding has been best studied in the Wan¬ dering Albatross but all the evidence indicates that it is broadly similar in the other species. Endocrine processes For male Wandering Albatrosses, Hector et al. (1986a) showed that the amplitude of the testicular cycle (in terms of both testis size and testosterone concentration in the blood) increases with age up to about age 10 years (Figure 1). However, for males of age four-five years, the size of the testis and levels of circulating testosterone at ACTA XX CONGRESSUS I NITER NATION ALIS ORNITHOLOGICI 283 the appropriate time are amply sufficient for them to be regarded as physiologically sexually mature. For females, however, birds younger than age seven years show high and variable progesterone levels and low concentrations of oestradiol and luteinising hormone (LH) and are physiologically incapable of breeding. In contrast, older breeding birds have consistently low levels of progesterone and high levels of oestradiol and LH (Hector et al. 1986a, 1990). -iiooo 70 « 60 E 50 £ 40 ' 30 20 10 0 CD E o TD £ -20 100 cr c i o TD CD O FIGURE 1 - (a) Maximum testicular length (•) (± standard error; n >3) and concentrations of circulating testosterone in different age classes of immature male Wandering Alba¬ trosses (± standard error of the mean; n >8). The age classes, in years, are given on the bottom axis. Data from breeding birds caught at copulation (C) and at the same time as the non-breeding samples (CJ are shown for comparison (n = 6 on both occasions), (b) Maximum diameter of largest follicle (*) and concentrations of circulating oestradiol and progesterone in different age classes of immature female Wandering Albatrosses (± stand¬ ard error; n >9). Categories C and C1 are as for males (n = 6 on both occasions). (After Hector et al. 1986a). These findings led to the suggestion (Hector et al. 1986a) that the ovary secretes progesterone rather than oestradiol to prevent vitellogenesis and egg formation, while regulating LH secretion by negative feedback. Subsequent experiments (Hector et al. 1990) showed that the progesterone was of ovarian origin and that, in response to LH injection, non-breeding birds secreted only progesterone but those birds about to lay eggs secreted both progesterone and oestradiol (Figure 2). Whatever the precise interactions within the hypothalamus-pituitary-gonad axis and the exact role of progesterone in the inhibition of egg-laying, it seems clear that physi¬ ological sexual maturity in female Wandering Albatrosses is not attained before seven years of age. This is entirely consistent with field data, whereby in 1500 records only 2% of females have laid at age seven years and none at an earlier age. Remarkably, this situation has been maintained even though the average age of first breeding has 284 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI been decreasing in recent years (Croxall et al. 1990). Thus the physiological barrier has been maintained in the face of ecological/behavioural pressure towards an ear¬ lier age for first breeding. Behavioural processes Physiological sexual maturity does not necessarily guarantee immediate maturity in the social context, because even albatrosses of appropriate age take several years to acquire partners and breed (Pickering 1989). 4 200 Control Saline Injec. (n = 4) 0 0 200 Non-Breeders (n = 23) 0 200 Last-Time Non-Breeders (n = 7) 0 / 4 200 First Time Breeders (n = 3) A PROG AE2-1 7/J M S/I ng/l FIGURE 2 - Change in progesterone and oestradiol concentration after a single injection of ovine LH in four separate groups of female Wandering Albatrosses. Values are mean ± standard error. (After Hector et al. 1990). FIGURE 3 - Attendance patterns of Wandering Albatrosses: (a) in relation to experience. Each horizontal bar spans the median arrival to median departure date. Its thickness is proportional to the number of days ashore. The median number of days ashore is shown within or above each block. Males (stippled), females (clear) (n = 393). (b) in relation to stage of pair bond formation. I: Inexperienced birds; ENP: Experienced non-pairing birds; EP: Experienced pairing birds; LTNB: Last-time non-breeders; F: Former breeders. Other conventions as in Figure 3a (n = 421). (After Pickering 1989). ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 285 uoubiujoj puoq-j|Bd jo 06bjs o k_ 03 .O 0) c 03 0 1 0 Q o <13 O > o z -C o k- 03 n 03 Ul c 03 o 03 o > o z 0 -*— > 0 Q UOSB0S 286 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Wandering Albatrosses of both sexes first return to Bird Island (and, indeed, to their natal area on the island) at an average age of five years (range 3-12 years). On their first visit they arrive some two months after the median laying date and stay for only three days on average. In each succeeding year they arrive earlier and stay longer (Figure 3a). In relation to their stage of pair-bond formation (Figure 3b), however, the critical change comes between experienced pairing birds (i.e. birds seen together regularly at a nest site), none of which arrive before the median laying date, and last¬ time non-breeders, whose pattern of arrival and attendance is similar to that of former breeding birds (Pickering 1989). At all ages and in all categories females spend sig¬ nificantly less time ashore than males. The actual process of pair formation proceeds through various stages. Initially, with increasing experience (time spent ashore), birds spend more time displaying and in¬ teract with more birds of the opposite sex (Figure 4). Subsequently displays are per¬ formed with increasing frequency with a decreasing number of partners (Figure 4) and increasingly often at the site of a nest built by the male bird (Pickering 1989). o +—> CO k. •O c 4 Wandering Fail (36) 63 17 8 Albatross (36-75) (7-43) (5-13) Success (64) 0 66 13 7 (55-75) (4-20) (5-9) Black-browed Fail (63) 75 9 5 Albatross (58-87) (4-25) (0-16) Success (37) 81 7 4 (57-93) (0-19) (0-18) Grey-headed Fail (54) 54 24 9 Albatross (26-80) (12-51) (3-23) Success (46) 1 68 10 7 (0-2) (31-88) (5-32) (0-19) * Values in parentheses are average percentage in each category. Thus for Black-browed Albatross there is no significant difference between the breed¬ ing frequency of birds that are successful or not in rearing chicks. A relatively small proportion of birds whose partners are still alive defer breeding by more than one year. The other two species exemplify the biennial strategy, with most successful and unsuccessful breeders making their next attempt two or one years later respectively but with a significant proportion in both cases deferring by longer than this. The only significant difference between Grey-headed and Wandering Albatrosses is the very small proportion (usually <1%) of successful Greyheaded Albatrosses which attempt breeding the next year. It has long been evident that successful Wandering 290 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Successful Fail ■o «L> 0) i_ JD O +■« o> c c u. 3 *-> TJ O E c o o a o 50 - 0 50 H 0 50 H 0 50 H 0 50 H 0 50 H 0 50 H 0 50 H 0 1=L U * . * ■ 1976 1977 1978 1980 1981 1982 1983 * 1984 12L 1234+ 123+ Years after breeding attempt / FIGURE 7 - Annual variation in proportion of South Georgia Wandering Albatrosses return ing to breed in subsequent years after success and failure in rearing a chick. Asterisk in dicates data incomplete. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 291 Albatrosses, with an incubation and chick-rearing period totalling 356 days, cannot breed in successive years (Tickell 1968); interest therefore centres on the nature of the mechanism precluding breeding in successful birds. With the mollymawks, how¬ ever, incubation plus chick-rearing and the total breeding-attendance period in Grey¬ headed Albatrosses lasts only 29 and 39 days longer, respectively, than in Black- browed Albatrosses. The small magnitude of these differences raises more fundamen¬ tal questions about the control and consequences of breeding frequency. WA BBA GHA Fas! Inc Rear Inc Rear nc ODFAJ AODFAJ AOD Female albatross FIGURE 8 - Diagrammatic representation of gonad (follicle diameter) and hormone (con¬ centrations of circulating progesterone and oestradiol) status in female Wandering (WA), Black-browed (BBA) and Grey-headed (GHA) Albatrosses throughout incubation (Inc) and chick-rearing (Rear) and, in GHA Success, while attending the breeding colony in a non¬ breeding year. (Data from Hector et al. 1986a, b, 1990). Endocrine influences Hector et al. (1986a,b) showed that male Wandering, Biack-browed and Grey-headed Albatrosses all have annual gonadal and hormone cycles. Successful female Wander¬ ing Albatrosses show completely undeveloped follicles throughout chick-rearing and an associated syndrome of relatively high progesterone and low oestradiol levels, 292 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI though the latter appear to increase significantly towards the end of chick-rearing (Figure 8). Neither successful nor failed birds reappear until the next breeding at¬ tempt, so the stage at which any hormone or gonad development occurs is unknown. Female Black-browed Albatrosses have annual gonadal and hormone cycles irrespec¬ tive of whether they were successful or not in rearing chicks; they show relatively high oestradiol and low progesterone concentrations during chick-rearing. When female Grey-headed Albatrosses which failed to rear a chick reappear to breed the next season, in follicle development and hormone profile they exactly resemble Black-browed Albatrosses. If they are then successful in rearing a chick, the circulat¬ ing progesterone concentration increases sharply late in the rearing season (Figure 8). Of these birds, those that return during the next breeding season (about 40% of the population) show little if any follicular development and high concentrations of circulating progesterone and low concentrations of oestradiol. None of these birds breeds. The key difference between the ability to breed or not in female Grey-headed Albatrosses seems to be correlated with the relative concentrations of oestradiol and progesterone late in chick-rearing - and possibly into the non-breeding season. This is a situation reminiscent of - or even analogous to - that operating in the development of sexual maturity. This discovery led to two immediate questions. First, why should such a mechanism be developed in Grey-headed and not Black-browed Albatrosses? Second, how can a few Grey-headed Albatrosses breed in the year after a successful season? The answer to these questions requires consideration of the ecological and behavioural context of the physiological mechanisms. 4.5 4.0 cn -C gi I 3.5 3.0 <> 0 • Black -browed Albatross (BBA) O Grey-headed Albatross (GHA) / \ \ \ 1 w~ Lay BBA GHA Hatch i r BBA GHA Fledge FIGURE 9 - Weights of breeding Black-browed and Grey-headed Albatrosses at South Georgia at different stages of the breeding cycle. (Data from Prince et al. 1981 and unpub¬ lished). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 293 Behavioural and ecological influences The endocrine data indicate that significant physiological events occur at the begin¬ ning and end of the breeding season and cycle; these are presumably linked to be¬ havioural and ecological events at similar times. The principal evidence available from such periods concerns change in condition (as indicated by body weight) during breeding and the timing of behaviour around the egg-laying period. Breeding Black-browed and Grey-headed Albatrosses essentially maintain weight during the period from egg-laying to hatching but lose weight throughout the chick¬ rearing period (Figure 9). Because successful Grey-headed Albatrosses lose weight for nearly one month longer than Black-browed Albatrosses, by the time the chick fledges they have lost 60% more weight (800g vs 500g). Thus Black-browed Alba¬ trosses have 20% more time to regain a much smaller proportionate weight loss (13% vs 20%), compared to Grey-headed Albatrosses. It is not surprising, therefore, that Grey-headed Albatrosses should experience greater difficulties than Black-browed Albatrosses in returning to breeding condition in time for the next season. Or that females, needing also to acquire additional energy and reserves for egg formation should be particularly affected. What is perhaps surpris¬ ing is that the situation is not one in which those birds which can regain breeding condition breed and those which cannot do not. Instead, Grey-headed Albatross fe¬ males successful in rearing chicks are apparently prevented from even attempting to breed through hormonal suppression of vitellogenesis. This suggests that even for those Grey-headed Albatross females in the best condition at the end of a success¬ ful breeding attempt, the cost of attempting to breed in five months time is sufficiently prohibitive for physiological mechanisms to have evolved to prevent it. For birds which fail, however, this block apparently does not exist and female condition in the following September/October presumably determines which birds actually attempt to breed. It should be noted that only 54% of pairs actually breed in the next year with 24% de¬ laying their next attempt for one further complete year (Table 2) . There are, however, no equivalent physiological constraints on male Grey-headed Al¬ batrosses. Nevertheless, almost all males remain faithful to their previous partner and do not breed in the year following one in which they rear a chick. The basis of this seems likely to relate to a combination of factors. First, physiological difficulty in re¬ gaining condition quickly enough to breed five months after successfully rearing a chick. Thus only 54% of failed birds manage to breed the next year; furthermore only 40% of successful males attend the colony and only a small proportion of these do so early enough to breed. Second, advantages deriving from continuing to breed with an existing familiar and presumably compatible partner. There are, however, two interesting features of the behaviour of Grey-headed Alba¬ trosses in their year ‘off’. First, the fact that a substantial proportion of birds visit the colony in this year. What is the purpose of this? It cannot be in order to make an extra breeding attempt (but see below) because all females are reproductively quiescent (see above). It is not in order to defend nest sites against incoming breeders because birds arrive too late to do this (Figure 10). The more likely explanations involve re¬ newal of pair bonds and reaffirmation that both partners are still alive. The advantages of doing this might relate on the one hand to improved breeding performance for pairs which meet during the year off and, on the other hand, in cases where only one mem¬ ber of the pair turns up, to receiving advance warning of the possible need to start the 294 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI potentially time-consuming task of seeking a new partner. Evidence for improved performance is slight; there are stronger indications of enhanced speed of re-pairing for birds whose partner fails to appear in the year off. Grey -headed Albatross Yolk formation Males Females Yolk formation Females starts arr. arr Ive stops return ♦ t H * t i 33 i 25 i i 19 18 i 11 1 i 2 0 Days before laying FIGURE 10 - Timing of events at a Grey-headed Albatross breeding colony in the period prior to egg-laying. (Data from Astheimer et al. 1985 and Prince unpublished). Second, the very small proportion of birds which breed immediately after a success¬ ful year (Table 2) comprises only the males from established pairings plus new female partners. This is made possible because males which were successful the previous year arrive significantly earlier than their partners (Figure 10) and overlap with the arrival of established breeding and last time non-breeding females, some of whom will discover that their partners have not turned up. Unless the males’ existing partner has disappeared, these pairings rarely last for more than one breeding season, the pre¬ vious partnership being re-established at the next breeding attempt. The extra breed¬ ing attempt by these established males is as successful, on average, as those of any other first time breeder - i.e. slightly, but not significantly less than those by estab¬ lished breeding pairs (see later). We have insufficient data to determine if there is a cost (in terms of reduced survival) to the extra breeding attempt but the advantage gained from this tactic seems small - except in the context of rapid acquisition of a new female partner in a population where competition for females exists. Within species, assessment of the relationship between breeding frequency and sur¬ vival requires much more data over complete reproductive lifetimes than we have available currently. However the substantial interspecies difference in mean annual survival between Black-browed Albatross (92%) and Grey-headed Albatross (95%) is consistent with an inverse relationship between breeding frequency and survival (Prince 1980, Croxall 1982, Weimerskirch et al. 1987). ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 295 BREEDING SUCCESS Breeding performance is obviously intimately related to attributes of the individual (e.g. inherent quality, acquired proficiency) but, especially in seabirds which are long- lived and monogamous, also involves a partnership effect (e.g. familiarity, compatibil¬ ity and the causes and consequences of divorce). There are, as yet, relatively few data on these topics for albatrosses; this section summarises some preliminary find¬ ings in relation to the results of studies of other seabird species. Long-term studies of seabirds have contributed significantly to the body of data on birds indicating that reproductive performance (almost invariably expressed in terms of breeding success) increases with breeding experience. In general, success in¬ creases through the first few years of reproduction, then levels off; subsequently there may or may not be a detectable decrease in the performance of the oldest birds. Im¬ proving performance is usually attributed to one or more of: a) Increasing experience of breeding; b) Increasing general competence (e.g. of feeding, avoiding predation, etc); c) Increasing reproductive effort to compensate for decreasing survival; d) Differential survival, favouring higher quality individuals. Nearly all seabird studies provide support for the roles of experience and competence (e.g. Thomas & Coulson 1988, Ollason & Dunnet 1988, Reid 1988, Pugesek 1984, Sydeman et al. 1991) but relatively few have been able to distinguish between the two and show that both operate (Nur 1984, Wooller et al. 1990). Although survival rates of many species decline with age and/or experience, the residual reproductive value hypothesis requires increased reproductive effort and success in older birds. The few studies adequately detailed to examine this have not supported the hypothesis (Reid 1988, Sydeman et al. 1991). Several studies (e.g. Coulson & Porter 1985, Bradley et al. 1989, Sydeman et al. 1991) have shown that long-lived individuals were more productive at some or all stages of their adult life i.e. that quality-related differentials exist. A few studies (e.g. Coulson 1966, Mills 1973, Bradley et al. 1990) have shown that breeding success increases with increasing pair-bond duration, independent of increased overall breed¬ ing experience. Divorces are often preceded by a higher than average failure rate (e.g. Richdale 1957, Mills 1973, Nelson 1978, Reilly & Cullen 1981, Shaw 1986, Ollason & Dunnet 1988, Bradley et al. 1990). New pairings frequently experience reduced breeding success initially (Coulson 1966, Mills 1973, Davis 1976, Brooke 1978, Ollason & Dunnet 1988, Bradley et al. 1990), though this rapidly improves if the pair stays together. If Wandering Albatrosses are typical, albatrosses may be most unusual amongst seabirds in that significant advantages of increasing experience and competence are barely detectable even when comparing birds breeding for the first time with birds of many years breeding experience. Thus for Wandering Albatrosses, first-time breed¬ ers are not significantly less successful than more experienced birds in their likelihood of raising a chick to fledge, although differences in hatching success, chiefly because of poor co-ordination of shift routine between the partners, are nearly significant (J P Croxall unpublished data). Lequette & Weimerskirch (1990) found that first time breed¬ ers were slightly less efficient at feeding their chicks (in terms of rate of delivery of 296 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI meals) than more experienced birds but that this did not persist beyond the early stage of chick-rearing. Thus chick growth rates and ages and weights of fledging was similar in all categories of experienced and inexperienced birds. These findings would support suggestions (e.g. Lequette & Weimerskirch 1990) that when albatrosses first breed they have attained essentially the same foraging competence as experienced breeders. However, such a conclusion raises some important questions, including whether the effect might simply be characteristic of Wandering Albatrosses and reflect unusually favourable circumstances (e.g. of food supply) while they are rearing chicks. First, the Wandering Albatross has one of the most consistently high levels of breed¬ ing success amongst seabirds (Weimerskirch & Jouventin 1987, Croxall et al. 1988). At South Georgia this ranges from 52% to 73% (mean 64%) over 15 years, whereas other albatrosses show lower and more variable breeding success (Weimerskirch et al. 1986, Croxall et al. 1988), some of which might relate to breeding experience. Clearly the situation in other species of albatross needs to be investigated. Second, other Procellariformes which show a similar degree of deferred sexual ma¬ turity also show improved reproductive success with greater breeding experience. Thus in the Northern Fulmar Fulmarus glacialis, in which the modal age of first breed¬ ing in males and females was eight and 12 years respectively, breeding success in both sexes improved until about their tenth breeding year, and thereafter remained approximately constant (Ollason & Dunnet 1988). In the similar Antarctic Fulmar F. glacialoides, breeding success increased significantly with the first three years of breeding experience but did not change significantly thereafter (Weimerskirch 1990). Other Procellariformes also show clear increases in reproductive success with breed¬ ing experience (e.g. Short-tailed Shearwater Puffinus tenuirostris (Wooller et al. 1990); Manx Shearwater Puffinus puffinus (Brooke 1990)) but these species breed at younger ages - modal age for P. tenuirostris is seven years (Bradley et al. 1989) - giving the possibility that acquisition of foraging competence might not be complete before birds start breeding. / Third, it is possible that, in rearing a chick successfully, inexperienced birds suffer a greater cost, (e.g. through working harder and finishing breeding in poorer condition) in terms of subsequently reduced survival or longer intervals between successive breeding attempts, than experienced birds. There is some indication from survival rate (though not from breeding frequency) that this might be so for Wandering Albatrosses (Croxall 1982). Such effects, however, should be more noticeable still in albatross species which breed annually and hence do not have a complete year to recover af¬ ter a successful breeding season. The hypothesis that very long-lived species with long-deferred sexual maturity delay breeding until they are fully competent foragers, in contrast to shorter-lived species in which breeding proceeds simultaneously with finishing acquiring foraging compe¬ tence, requires considerable further study. However it is clear that analysis of the relative roles of inherent quality, acquired proficiency and partner suitability and qual¬ ity in contributing to reproductive success will require consideration of events and processes prior to the start of breeding. Few such data are currently available, even for comprehensively studied species. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 297 SURVIVAL For most seabirds longevity is the single most important contributor to lifetime produc¬ tivity. The significance and relationships of many of the traits and adaptations dis¬ cussed above can only be adequately assessed in the light of information on their effects on survival and/or on the relationship between longevity and reproductive suc¬ cess. There are two particular problems in doing this with albatrosses. First, no study of sufficient scope has been conducted for long enough to have data on lifetime reproductive success for even one complete cohort of birds. Therefore most of the topics needing investigating, e.g. the balance of advantage and disadvan¬ tage (in terms of productivity and survival) between birds which breed at the young¬ est possible age and those which delay breeding even longer, cannot yet be studied. Second, the factors influencing survival of albatrosses (and, indeed, many other seabird species) may have changed drastically in recent decades. Thus for the South Georgia population of Wandering Albatrosses, mean annual survival of juveniles has decreased by 6% (from 90% to 84%) between the 1960s cohorts recruiting in the early 1970s and the 1970s cohorts recruiting in the early 1980s (Table 3). Similarly, mean annual survival of adults decreased by 2-3% (from 96% to 93-94%) from values re¬ corded in the 1960s to those current through the late 1970s and early 1980s (Table 4). These are major changes given the natural demography of albatross populations and may have been even greater for the same species at the Crozet Islands (Weimerskirch & Jouventin 1987) where adult survival averaged 92% and only 26% of juveniles survived to age 5 years (cf. 49% at South Georgia). TABLE 3 - Mean annual survival of juvenile Wandering Albatrosses at South Geor¬ gia, comparing cohorts in the 1960s with those in the 1970s. Sexes combined; based on data in Croxall et al. (1990) Year Age (years) surviving to Ringed Retrapped Mean annual survival 1962 9 400 174 0.912 1963 8 1000 399 0.892 Mean 0.896 1972 5 368 143 0.828 1973 5 75 27 0.815 1975 5 854 389 0.854 1976 5 847 337 0.832 1977 5 806 406 0.872 1978 5 871 462 0.881 1980 5 743 342 0.856 Mean 0.854 1972 - 1980 8 4564 1082 0.835 298 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI TABLE 4 - Mean annual survival of adult Wandering Albatrosses at South Georgia. Based on data in Croxall et al. (1990). Cohort Years 4 Survival (%) Male Female Combined 1958 1976-84 95.4 92.1 93.7 1962 1976-84 94.3 92.6 93.5 1963 1976-84 95.5 93.3 94.4 1958-63 1976-84 94.8 93.1 94.0 Study area 1961-63 95.7 Study area 1976-84 93.2 All the evidence (e.g. Croxall & Prince 1990, Brothers 1991) suggests that the main causes of these changes in survival were catches of albatrosses associated with tuna long-line fisheries, which developed in the 1970s. The fishery-related mortality was not random across sexes and age classes. Females are potentially at greater risk because of their more northerly at sea distribution (Weimerskirch & Jouventin 1987), taking them into the zone of the tuna fisheries. The reality of this effect is confirmed by analysis of ringing recoveries (Croxall & Prince 1990), movements of satellite tracked birds (P A Prince unpublished data) and by the significantly lower survival rates of females (Table 4; Croxall et al. 1990). In addition, adults seem to be more susceptible than juveniles (Croxall & Prince 1990), possibly because they are socially dominant to juveniles in the melees around fishing boats. However, birds of all sexes and ages were killed and in such numbers as to represent at least 50% of the total annual mortality of each sex and age class (Croxall et al. 1990). The imposition of such a level of mortality likely to be unrelated (or related in ways very different from previously) to characteristics like bird condition, quality, pre¬ vious breeding experience etc, is likely severely to compromise attempts to relate longevity and reproductive success to reproductive tactics and strategies. Failure to derive significant relationships in regard to survival (e.g. the lack of age-related sur¬ vival in the Wandering Albatross) may well not be typical of natural populations but simply reflect the results of recent artificial influences. CONCLUSIONS Many aspects of our studies of albatross biology and ecology have produced similar results to the often much longer term and more extensive studies of other seabird species. However, there are a number of unique results, some of which may prove to be characteristic only of albatrosses but others of which may have much broader applicability. One such feature is that there may be significant differences between the sexes in physiological and related behavioural and ecological adaptations. Thus in the Wan¬ dering Albatross one essential difference between the sexes in physiological terms is that males are sexually mature at an earlier age than females. This presumably re¬ flects the much greater magnitude (in terms of time spent in duties ashore) of the change between non-breeding and breeding status for females than for males and the ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 299 additional costs of egg formation in females. The costs of breeding too early in fe¬ males are presumably sufficiently great for a physiological mechanism to have evolved to prevent premature reproduction. In behavioural terms the most surprising finding was that despite the relatively highly evolved displays and the distinct sexual dimorphism in size and plumage in Wander¬ ing Albatrosses the principal male signal acted on by females appeared to relate to the duration and/or frequency of attendance ashore. In spending time ashore males are advertising their ability to cope with the demands of fasting - the key adaptation required once they start breeding in order to cope with the long incubation shifts. Female commitment to life ashore is significantly less than males, so in both endo¬ crine and behavioural terms females appear to be protected from sustaining undue stress. Because Black-browed and Grey-headed Albatrosses are the same size and have very similar timing and rather similar duration of breeding events, yet breed annually and biennially respectively, they are uniquely suitable for investigating control of breeding frequency. Even in a species whose sexes are of very similar size and structure, there may be significantly different physiological influences on the two sexes. Thus in the Grey¬ headed Albatross an endocrine-mediated mechanism prevents females, but not males, from breeding immediately following success in chick-rearing. Bienniality in males seems to be maintained by a combination of physiological difficulty in regain¬ ing condition and advantages of continuing reproduction with a familiar, experienced and presumably compatible partner. The absence of an endocrine constraint to breeding frequency in males may help them, in circumstances of actual or potential mate loss, to start re-pairing as soon as possible. This is desirable because there are usually fewer unpaired adult females than males. Informing their partners of their continued survival is presumably one reason why substantial numbers of Grey-headed Albatrosses of both sexes try to re¬ turn in their year off. Because albatrosses are essentially pelagic throughout their inter-breeding period, evidence for the proximate influence of condition on breeding ability and frequency will be very difficult to acquire. However, in addition to regaining body condition follow¬ ing breeding, albatrosses also need to undertake moult (no species moults during its breeding season). In the Wandering Albatross, Weimerskirch (in press) has demon¬ strated inverse relationships between body weight and moult (in males but not fe¬ males) and direct relationships between time between breeding attempts and moult (in females but not in males), which suggest that the energy costs of moult may not be insignificant and that the two sexes may experience different constraints in this regard. Although a strong interspecies correlation exists between breeding frequency and survival there are so far insufficient data to demonstrate this intraspecif ically . Breeding success in albatrosses appears to be much less influenced by experience than in other seabirds - perhaps because they have acquired sufficient experience during the lengthy periods of immaturity and pair formation prior to starting breeding. Much more 300 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI work is needed on this topic in albatrosses and seabirds generally. Except for the Wandering Albatross, which has shown consistently high breeding success over the last 15 years, variation in breeding success may relate significantly to the particular years in which breeding is attempted and especially to the avoidance of years of poor breeding success for the whole population. At South Georgia such years occur rela¬ tively frequently (Croxall et al. 1988), although they are often associated with smaller breeding populations and higher indices of deferred breeding than usual (Croxall & Rothery 1991). In any case the role of breeding frequency in determining overall pro¬ ductivity will be much less significant than the influence of longevity. Albatrosses live so long that data on lifetime reproductive success are very slow to accumulate and it is impossible at present to assess the consequence of reproductive tactics and strat¬ egies. However the magnitude of changes in Wandering Albatross survival over re¬ cent decades, due principally to human fishing activities, raises the strong possibil¬ ity that mortality from these sources may prevent detection of the subtler effects of individual differences in reproductive adaptations. There is also the broader concern, noted by Croxall & Rothery (1991), that because human influences include direct predation, competition for common resources, mor¬ tality through introduced animals, destruction of breeding habitat and pollution of feed¬ ing habitat, few, if any, data on seabird populations come from sites and systems unperturbed by man. There is a real danger that the responses of populations of seabirds, especially those species with long generation times and low productivities, may no longer be to a range of relatively constant and predictable natural factors but instead, so far as they are able, to a variety of rapidly changing and unpredictable constraints imposed through human interference. ACKNOWLEDGEMENTS I thank the many colleagues who have worked with albatrosses at South Georgia, but in particular Julian Hector, Simon Pickering and, most of all, Peter Prince, who also allowed me to use his unpublished data. Henri Weimerskirch kindly provided me with copies of forthcoming papers. LITERATURE CITED ASTHEIMER, L.B., PRINCE, P.A., GRAU, C.R. 1985. Egg formation and the pre-laying period of Black- browed and Grey-headed Albatrosses at Bird Island, South Georgia. Ibis 127: 523-529. BRADLEY, J.S., WOOLLER, R.D., SKIRA, I.J., SERVENTY, D.L. 1989. Age dependent survival of breeding Short-tailed Shearwaters. 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Reproductive endocrinology of the Black- browed Albatross Diomedea melanophris and the Grey-headed Albatross D. chrysostoma. Journal of Zoology 208: 237-254. HECTOR, J.A.L., PICKERING, S.P.C., CROXALL, J.P., FOLLETT, B.K. 1990. The endocrine basis of deferred sexual maturity in the Wandering Albatross Diomedea exulans. Functional Ecology 4: 59-66. JOUVENTIN, P., WEIMERSKIRCH, H. 1988. Demographic strategies in southern albatrosses. Pro¬ ceedings of the Nineteenth International Ornithological Congress: 857-865. LEQUETTE, B., WEIMERSKIRCH, H. 1990. Influence of parental experience on the growth of Wan¬ dering Albatross chicks. Condor 92: 726-731. McCLEERY, R.H., PERRINS, C.M. 1988. Lifetime reproductive success in the Great Tit Parus major. Pp. 136-153 in Clutton-Brock, T.H. (Ed.). Reproductive success. Chicago, University of Chicago Press. MILLS, J.A. 1973. The influence of age and pair bond on the breeding biology of the Red-billed Gull Larus novaehollandiae scopulinus. 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Population and energetic aspects of the relationship between Black-browed and Grey-headed Albatrosses and the Southern Ocean marine environment. Pp. 473-477 in Siegfried, W.R., Condy, P.R., Laws, R.M. (Eds). Antarctic nutrient cycles and food webs. Berlin, Springer-Verlag. PRINCE, P.A., RICKETTS, C., THOMAS, G. 1981. Weight loss in incubating albatrosses and its im¬ plications for their energy and food requirements. Condor 83: 238-242. PUGESEK, B.H. 1984. Age-specific reproductive tactics in the California Gull. Oikos 43: 409-410. REID, W.V. 1988. Age-specific patterns of reproduction in the Glaucous-winged Gull: increased effort with age? Ecology 69: 1454-1465. RICHDALE, L.E. 1957. A population study of penguins. Oxford, Oxford University Press. RICHDALE, L.E., WARHAM, J. 1973. Survival, pair-bond retention and nest site tenacity in Buller’s Mollymawk. Ibis 115: 257-263. SHAW, P. 1986. Factors affecting the breeding performance of Antarctic Blue-eyed Shags Phalacrocorax atriceps. Ornis Scandinavica 17: 141-150. 302 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI SYDEMAN, W.J., PENNIMAN, J.F., PENNIMAN, T.M., PYLE, P., AINLEY, D.J. 1991. Breeding per¬ formance in the Western Gull: effect of parental age, timing of breeding and year in relation to food availability. Journal of Animal Ecology 60: 135-150. THOMAS, C.S., COULSON, J.C. 1988. Reproductive success of Kittiwake Gulls Rissa tridactyla. Pp. 251-262 in Clutton-Brock, T.H. (Ed.). Reproductive success. Chicago, University of Chicago Press. TICKELL, W.L.N. 1968. The biology of the great albatrosses Diomedea exulans and D. epomophora. Antarctic Research Series 12: 1-55. TICKELL, W.L.N., PINDER, R. 1975. Breeding biology of the Black-browed Albatross Diomedea melanophris and Grey-headed Albatross D. chrysostoma at Bird Island, South Georgia. Ibis 117: 433-450. TINBERGEN, J.M., DAAN, S. 1990. Family planning in the Great Tit (Parus major): optimal clutch size as integration of parent and offspring fitness. Behaviour 114: 160-190. WEIMERSKIRCH, H. 1982. La strategie de reproduction de I’Albatros Fuligineux a Dos Sombre. Comite National Frangais de Recherches Antarctiques 51: 437-448. WEIMERSKIRCH, H. 1990. The influence of age and experience on breeding performance of the Ant¬ arctic Fulmar Fulmarus glacialoides. Journal of Animal Ecology 59: 867-875. WEIMERSKIRCH, H. In press. Sex-specific differences in molt strategy in relation to breeding in the Wandering Albatross. Condor. WEIMERSKIRCH, H., JOUVENTIN, P. 1987. Population dynamics of the Wandering Albatross, Diomedea exulans, of the Crozet Islands: causes and consequences of the population decline. Oikos 49: 315-322. WEIMERSKIRCH, H., JOUVENTIN, P., STAHL, J.-C. 1986. Comparative ecology of the six albatross species breeding on the Crozet Islands. Ibis 128: 195-213. WEIMERSKIRCH, H., CLOBERT, J., JOUVENTIN, P. 1987. Survival in five southern albatrosses and its relationship with their life history. Journal of Animal Ecology 56: 1043-1055. WEIMERSKIRCH, H., LEQUETTE, B., JOUVENTIN, P. 1989. Development and maturation of plum¬ age in the Wandering Albatross Diomedea exulans. Journal of Zoology, London 219: 411-421. WOOLLER, R.D., BRADLEY, J.S., SKIRA, I.J., SERVENTY, D.L. 1990. The reproductive success of Short-tailed Shearwaters Puffinus tenuirostris in relation to their age and breeding experience. Jour¬ nal of Animal Ecology 59: 161-170. / ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 303 SYMPOSIUM 1 BIOGEOGRAPHY AND SPECIATION IN NEOTROPICAL BIRDS Conveners K-L. SCHUCHMANN and F. VUILLEUMIER 304 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI SYMPOSIUM 1 * Contents INTRODUCTORY REMARKS: BIOGEOGRAPHY AND SPECIATION IN NEOTROPICAL BIRDS KARL L. SCHUCHMANN . 305 NEOTROPICAL AVIAN DIVERSITY AND RIVERINE BARRIERS A. P. CAPPARELLA . 307 DISJUNCT BIRD DISTRIBUTIONS ALONG THE WEST SLOPE OF THE PERUVIAN ANDES IRMA FRANKE . 317 SPECIATION IN PATAGONIAN BIRDS FRANQOIS VUILLEUMIER . 327 GENETIC DIFFERENTIATION IN YELLOWTHROATS (PARULINAE: GEOTHLYPIS) B. PATRICIA ESCALANTE-PLIEGO . 333 BIOGEOGRAPHIC PATTERNS IN BIRDS OF HIGH ANDEAN RELICT WOODLANDS J. FJELDSA . 342 CONCLUDING REMARKS: BIOGEOGRAPHY AND SPECIATION IN NEOTROPICAL BIRDS FRANQOIS VUILLEUMIER . 354 / ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 305 INTRODUCTORY REMARKS: BIOGEOGRAPHY AND SPECIATION IN NEOTROPICAL BIRDS KARL L. SCHUCHMANN Alexander Koenig - Zoological Research Institute and Museum, Adenauerallee 150-164, D-5300 Bonn 1, Germany The Neotropics are inhabited by the most diverse avifauna in the world. Roughly 3300 bird species have been documented, among them only 180 migrants from North America. This contrasts with about 1600 resident species within the Afrotropics, 961 in the Oriental Region and 906 in Australasia. Compared to the ornithofauna of tropi¬ cal Africa to Australia, the Neotropical avifauna is taxonomically very different and, due to a long-lasting geological isolation, comprises a high percentage of endemic bird families (30%). The high degree of taxonomic differentiation is illustrated by the species/family ratio, which is 33.9 for South America against 22.4 for Africa and 14.6 for Australia. During the past two decades Neotropical ornithology has benefitted greatly from new publications, especially regional bird lists, field guides and handbooks. This has im¬ proved the active research participation of amateur ornithologists and resulted in a cooperation with professionals similar to that in northern countries. Today, a wealth of new data has become available including information on abundance of birds in space and time, on geographical variation and ecological requirements of species inhabiting a given region. With this new knowledge as background numerous analyti¬ cal studies on zoogeography and speciation have been carried out. The following two questions will form the framework of our symposium: What are the historical roots of the present-day distribution of neotropical birds, and how can we explain speciation patterns? Since Darwin and Moriz Wagner, the biogeographical analysis of distribution has helped to identify the degree and patterns of isolation which are prerequisites to ge- netical differentiation of populations at the species level, during the process of allopatric speciation. Parapatric speciation is caused by genetical differentiation of continuously distributed populations influenced by ecological gradients, sympatric speciation by chromosome and other local changes within a population. However, parapatric and sympatric differentiation in vertebrates, especially in birds, probably had minor impacts on speciation. Geographical isolation of populations seems deci¬ sive for species differentiation in continental faunas. It is supposed to have been caused by two processes: fragmentation of formerly continuous areas (vicariance), and transgression of barriers by groups of individuals (dispersal, founder populations). These processes were probably influenced heavily by climatic effects. Two hypotheses are presently being discussed to explain the mosaic-like distribution patterns and the speciation events of Amazonian lowland-forest superspecies: the refuge hypothesis and the riverine hypothesis. The refuge model is based on 306 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI cyclical climatic fluctuations causing expansions and contractions of dry to humid terra-firme forest and of non-forest habitats, especially during Quaternary and Terti¬ ary, but not restricted to these epochs. The riverine model, on the other hand, as¬ sumes the fragmentation of bird ranges during the development of the extensive Amazonian river system as a consequence of the uplifting of the Andes. * For montane species similar hypotheses would be the refuge model and a mountain- range isolation model. Both of these would be modified by orogenic processes. The speciation events as a consequence of climatic fluctuations are theoretically con¬ vincing, but we still lack conclusive evidence to support the refuge idea. Moreover, we rarely know whether speciation events were caused by recent climatic cycles, earlier cycles, or both. As to unsolved problems connected with the riverine (or mountain- range) model, we have to deal with the origins of those lowland forest birds to which rivers hardly represent barriers and with those birds which have continuous distribu¬ tions in the headwater regions where the rivers cease to be barriers. Probably, dis¬ tribution and speciation result from both isolation types: rivers or mountain ranges and habitat refuges. With further biological and geological information, it may be possible to assess the general validity of these models. For the near future our work on biogeography and speciation of Neotropical birds will profit greatly if we continue our studies on phylogenetic affinities within species groups by cladistic methods, establishing phylograms based on geology and comparing them with cladograms based upon anatomical, behavioural, and biochemical information. Additionally, our studies of geographical variation would be facilitated if more compu¬ ter generated contour map studies of character scores and measurements would be¬ come available (Haffer and Fitzpatrick method). All our challenging projects can only be achieved if we can continue responsible collecting of specimens all over Central and South America. International collaboration projects would be very useful. A major objective of present activities of amateurs and professionals is to prepare an atlas of speciation in Neotropical birds. The first step needed to realize this ambitious plan is to publish detailed range maps of birds from those states and countries where the avifauna is known in great detail. The gazetteers of Latin America edited by R. Paynter and collaborators will be invaluable. Computerization of specimen collections and of records in the literature is highly desirable, especially lists of locations of type specimens as we discover unexpected sibling species or specific differences between birds previously thought to be subspecies. A variety of museum, field and laboratory studies will be needed and should receive support. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 307 NEOTROPICAL AVIAN DIVERSITY AND RIVERINE BARRIERS A. P. CAPPARELLA Department of Biological Sciences, Illinois State University, Normal, Illinois 61761, USA ABSTRACT. The high frequency with which rivers delimit phenotypically differentiated bird taxa (spe¬ cies, subspecies) is unique to Amazonia. The riverine barrier hypothesis (an alternative to the Pleistocene refugia hypothesis) posits that the high regional species richness in Amazonia resulted from the development of the riverine system. These hypotheses are evaluated in the light of the knowl¬ edge of patterns of phenotypic and genetic differentiation in birds and the current geological evidence. Predictions are developed by which to test these hypotheses, gaps in our current knowledge are pro¬ filed, and specific research programs are recommended. Keywords: riverine barriers, refugia, bird diversity, Amazonia. INTRODUCTION For birds, the Amazon basin (South America) has the highest alpha (single-point) and gamma (regional) species diversity (Amadon 1973). Hypotheses to explain this diver¬ sity are ecological and historical. Ecological hypotheses address the causes of sin¬ gle-point diversity, and in part regional. To explain fully regional diversity, historical hypotheses have been offered. Of the historical hypotheses advanced, the Pleistocene refugia hypothesis (Haffer 1969) is widely accepted (Prance 1982). It states that regional diversity is attributable to the periodic fragmentation and coalescence of the forest during Pleistocene climatic fluctuations. The isolation of forest fragments (refugia) is the major promoter of speciation, and rivers only constitute a partial barrier to species’ re-expansion follow¬ ing climatic amelioration (Haffer 1974). Another historical hypothesis, suggested by the congruence of many birds’ ranges with rivers, is that the formation of the Amazonian river system after the uplift of the Andes induced speciation in forest-dwelling birds by fragmenting their ranges and prohibiting gene flow. Under this hypothesis, regional diversity derives from riverine barriers serving as a vicariant mechanism interrupting gene flow (Sick 1967). Although these hypotheses are not mutually exclusive, there is a profound difference between them: the existence of the vicariant agent for the Pleistocene refugia hypoth¬ esis (isolated forest refuges) is inferred, while the vicariant agent for the riverine bar¬ rier hypothesis (riverine system) is known to exist. As discussed herein, recent geo¬ logical and biochemical systematic evidence have challenged the existence of Ama¬ zonian refugia. If refugia did not exist, then our understanding of speciation mecha¬ nisms in Amazonia must be re-evaluated. In this paper, I summarize some evidence bearing on the two hypotheses, develop predictions to test them, identify gaps in our knowledge, and offer research programs to improve our understanding of regional species diversity in Amazonia. Finally, I touch on the conservation implications of this issue. 308 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI GEOLOGICAL EVIDENCE The majority of geomorphological, palynological, and paleoclimatological data used to support refugia is derived from sites in the Andes or peripheral to forested Amazonia (Prance 1982). Such evidence from within the basin is equivocal (Salo 1987, Colinvaux 1989). One study directly challenges the refugia hypothesis by inter¬ preting pollen and megafossil remains from the western edge of the putative Napo refugium as harbouring montane vegetation, not lowland tropical forest, during a mid- Wisconsin interstade (Colinvaux 1987). The crux of the argument regarding the effects of glacially-induced climatic alteration is which effect — reduction of precipitation and/or reduction of temperature — most strongly affected the vegetation. (The geological timing of these events is a separate problem discussed under prediction 2.) Under the refugia hypothesis, substantial arid¬ ity caused the forest to be replaced by savanna, except in mesic areas (Haffer 1969). An alternative is that the reduction of precipitation was insufficient to fragment the forest; rather, forest vegetation zones contracted under the influence of reduced tem¬ perature so that peripheral open habitats expanded and the montane/lowland tropi¬ cal forest ecotone dropped in elevation (Colinvaux & Liu 1987). Therefore, the cen¬ tral question is whether the Amazonian lowland tropical rain forest fragmented or con¬ tracted. Due to the lack of data from within the basin, this question cannot be an¬ swered currently. For this reason, current patterns of species distributions are used to infer previous vicariant events. BIOLOGICAL EVIDENCE Phenotypic differentiation The observation that Amazonian rivers delimit the range of many taxa of volant birds is in textbooks (e.g. Wallace 1876, Mayr 1942), general avifaunal works (e.g. Hellmayr 1910, Snethlage 1913), and discussions of specific taxonomic groups (e.g. Todd 1927). Hellmayr (1910) reports that the Madeira River delimits the range of 67 taxa. The lower Amazon river delimits the range of 80 taxa, and three large tributaries of lower Amazonia — Tocantins, Xingu, Tapajoz — delimit 37, 22, and 12 taxa, respectively (Snethlage 1913). Three seasons of field work in the upper Amazon (Iquitos) docu¬ mented 4 taxa delimited by the lower Napo river and 24 delimited by the wider Ama¬ zon river (Capparella 1987). In presenting such tabulations, authors usually differen¬ tiate between two categories: 1) taxa which have an opposite bank replacement form, and 2) taxa which do not. In the case of the former it is assumed, but has not been demonstrated, that the two are sister taxa. Despite these tabulations, a complete published list of the number of taxa delimited by rivers in Amazonia is lacking, although such a list could be compiled from the data utilized by Haffer (1978) and the information in Peters (1934-1987). Problems with such a list stem from the uncertainty regarding the ranges of Amazonian birds (e.g. Parker & Remsen 1987) and the delimitation of taxon limits (e.g. Capparella & Rosenberg in press). Additional collections are needed, particularly in the headwaters, where the breakdown of river-delimited ranges are most likely (e.g. see Bates et al. 1989). One uncertainty in compiling such lists is illustrated by comparing the species collected at three sites within 80 km in contiguous forest along the north bank of the Napo and Amazon rivers (Capparella 1987). Although there is considerable ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 309 similarity in species composition and number, some species (e.g. Black-headed Antbird, Percnostola rufifrons ; all names from Meyer de Schauensee 1970) were col¬ lected at only one site. This complicates the determination of rivers delimiting species ranges because a sample at a single trans-river site may not detect the species, even though it is present at other sites on the same bank. Three kinds of differences, other than sampling error, may explain these among-site changes in understory avifauna: 1) microhabitat availability, 2) seasonal or mobile resources, and 3) bird density. These factors must be evaluated when determining the likelihood that a species is truly absent from a particular region and interpreting such absence as indicative that this species’ range is delimited by a river. For example, the Black-headed Antbird absence may be attributable to the first factor. On the other hand, some species appear to be absent even when there is not a clear-cut replace¬ ment taxon (e.g. White-plumed Antbird, Pithys albifrons, absent south of the Amazon). Genetic differentiation Genetic differentiation among river-separated populations of species that do not dif¬ fer in plumage has been demonstrated using protein electrophoresis. River-congru¬ ent genetic differentiation is known in five monomorphic, terra firme (not seasonally flooded), forest-understory birds (river given after name; A=Amazon, N=Napo): Wedge-billed Woodcreeper (Glyphorynchus spirurus ; Dendrocolaptidae; A, N), Stip¬ ple-throated Antwren ( Myrmotherula haematonota ; Formicariidae; A), Blue-crowned Manakin ( Pipra coronata ; Pipridae; A, N), Black-faced Antbird ( Myrmoborus myotherinus ; Formicariidae, A, N), White-plumed Antbird ( Pithys albifrons ; Formicariidae; N) (Capparella 1987,1988; Flackett & Rosenberg 1990). In addition, genetic differentiation among phenotypically differentiated, river-delimited taxa has been demonstrated for: Long-tailed Flermit ( Phaethornis superciliosus ; Trochilidae, A, F. Gill & J. Gerwin pers. comm.), Blue-crowned Manakin ( Chiroxiphia pareola napensis and C. p. regina ; Pipridae, A), Golden-headed/Red-headed Manakin allospecies ( Pipra erythrocephaia and P. rubrocapilla ; Pipridae; A) (Capparella 1987,1988). These taxa span a diversity of familial affinities and life history traits shar¬ ing only their occurrence in the understory of terra firme forest. I expect the congruence of genetic differentiation with rivers to be a general phenom¬ enon among understory terra firme forest birds, although the minimum size-class river that will show this effect is not known. The number of genetically differentiated forms delimited by rivers is clearly greater than that predicted from plumage differences alone, although vocal differences may tell a story akin to allozymes (T. A. Parker III, pers. comm.). Interestingly, the levels of differentiation across the Amazon for four non-phenotypically differentiated birds (Wedge-billed Woodcreeper, Stipple-throated Antwren, Black-faced Antbird and Blue-crowned Manakin) are comparable to the mean value for temperate zone avian species (Capparella 1987, Flackett & Rosenberg 1990). The examination of patterns of genetic differentiation among Amazonian birds is in its infancy. Unknown at present is if other general classes of birds, especially terra firme forest-canopy birds and varzea (seasonally flooded) forest-understory birds, will also show genetic differentiation congruent with rivers. The paucity of river-associated phenotypic differentiation in most canopy birds (e.g. Isler & Isler 1987) suggests there may not be river-associated genetic differentiation. If so, this would support the inter¬ pretation that neotropical birds inhabiting the dark understory of terra firme forest will 310 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI not cross light gaps such as rivers (Capparella 1987). Canopy species, occupying the well-lighted top of the forest, may not show such “negative phototaxis”. However, the presence of phenotypic differentiation in some canopy groups such as toucans (Ramphastidae) that is only partially congruent with rivers has been offered both in support of Pleistocene refugia (Haffer 1974) and against it (Cracraft & Prumm 1988). The lack of river-associated phenotypic differentiation in most varzea birds suggests there may not be river associated genetic differentiation in these species. This sug¬ gests that the transitory nature of varzea forest requires across-river dispersal or, alternatively, the formation of oxbows by rivers is causing passive dispersal of oppo¬ site bank varzea forest with the birds. A biochemical systematic approach is needed to characterize genetic differentiation within these bird groups. THE HYPOTHESES The riverine barrier hypothesis The riverine barrier hypothesis is implicit in earlier writings (e.g. Wallace 1876, Hellmayr 1910, Snethlage 1913) and was discussed by Sick (1967). It begins with the formation of the Amazonian lowlands due to the rise of the Andes, completed some 2 million years ago, leading to deposition filling in the lake occupying the centre of South America. This permitted the colonization of new land by forest and birds from sources located on the Guianan shields. The rise of the Andes initiated also the for¬ mation of the drainage pattern that has become the Amazon riverine system. As these rivers increased in width, they began to fragment the rain forest and the ranges of birds that inhabited them. The effects of glacially-induced climatic effects on this process are unclear. The fol¬ lowing have been proposed: 1) an Amazon river embayment which would widen the lower Amazon and its tributaries (Haffer 1974), 2) lakes and extensive flooding at various times (Campbell & Frailey 1984), and 3) lowering of the montane-lowland for¬ est ecotone deeper into the headwaters (Colinvaux & Liu 1987), thereby preventing gene flow. However, the central tenet in the riverine barrier hypothesis is that the riv¬ ers themselves were the primary vicariant mechanism, regardless of possible ancil¬ lary processes. The explanatory power of this or any other vicariant model is its ability to account for a substantial number of bird distributions. The assumption is that a common thread does explain the majority of distributions in Amazonia and that it is not the result of a multiplicity of forces that have operated independently on each taxon. This was the appeal of the Pleistocene refugia model. The appeal of the riverine barrier model is that the agents of vicariance — the rivers — are known to exist while the existence of refugia are as yet only inferred. Rivers versus refugia The riverine barrier and Pleistocene refugia hypotheses both state that rivers can serve as barriers, although they differ regarding their effectiveness. The former states that rivers cause differentiation whereas the latter views rivers as limiting the re-ex¬ pansion of taxa formerly isolated in refugia and/or providing a location (due to reduced gene flow) at which a contact zone between former refugia can stabilize. Although the causes are different, the resultant patterns would be similar so it is difficult to develop ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 311 predictions to distinguish between them. Following are four predictions that should permit testing these two hypotheses. Prediction 1. Terra firme forest-understory birds, regardless of taxonomic affinity or level of phenotypic differentiation (from none to substantial) will show significant geno¬ typic differentiation across the major rivers of Amazonia when examined by biochemi¬ cal methods. As described earlier, this has been shown for several species across the Napo and Amazon rivers. Interestingly, the level of differentiation was higher across the Amazon than across the Napo. Whether this is due to the recency of attainment of the present width of the Napo, the occurrence of more frequent oxbowing permit¬ ting passive transport, or the capability of headwater crossing remains to be deter¬ mined. If the latter is the case, one would expect to find clinal variation among sam¬ ples taken further upstream on both banks. Further sampling of taxa across other riv¬ ers is needed to confirm this general pattern and determine what minimum size-class river is associated with genotypic differentiation. Although confirmation of this predic¬ tion does not rule out refugia, it appears to obviate the need to postulate refugia for terra firme forest-understory birds. Prediction 2. The calibrated genetic distance value (i.e., a time estimate) between sister taxa separated by a river will be clustered around 1-2 million years ago (m.y.a.) under the riverine barrier hypothesis instead of a spread of values over a 6-million- year interval in the refugia hypothesis. The completion of the Andean uplift and con¬ comitant river formation was Late Pliocene-Early Pleistocene (some 2 m.y.a.). The glacially-induced climatic fluctuations affecting South America are now thought to have begun in the Late Tertiary as much as 6 million years ago (Van der Hammen 1985; Van Zinderen Bakker 1986), and this has led to a modification of the original Pleistocene refugia hypothesis (J. Haffer pers. comm.). The onset of refugia now dates to the Late Tertiary and there were pulses of refugia until the last glacial event in the Late Pleistocene. This modification is substantial because a major consequence of the original hypothesis (Haffer 1969) was that regional species diversity in Amazonia dates from the Pleistocene. The neutral mutation model of the evolution of electrophoretic characters states that they evolve in a roughly time-dependent manner. This theoretically permits the utili¬ zation of genetic distances to date divergence events. However, the existence of a proper calibration time remains controversial. Application of one commonly cited value (Gutierrez et al. 1983) to previously described Amazonian taxa reveals that sister taxa of suboscine Amazonian birds are, on average, much older than taxa of temperate zone birds (Hackett & Rosenberg 1990). For trans-Amazon taxa mentioned earlier, the median value of 1.6 m.y.a. supports the prediction of the riverine barrier hypothesis (Capparella 1987,1988, Hackett & Rosenberg 1990). Further calibrated values are needed to determine if there is a concentration around 2 m.y.a. or a spread of values over a 6-million-year interval. Prediction 3. Allozyme heterozygosity values and mitochondrial DNA clonal diversity values will decrease outward from the core of each refugium. The expansion of the formerly restricted taxa into newly arising forest would involve a stepwise series of founder events. A transect through a putative refugium should reveal a central core of high allozyme heterozygosity/mtDNA clonal diversity with a decrease as one moves away from the core, assuming that the expanding peripheral populations have not reached equilibrium. FIGURE 1 - Map of Amazonia with rivers between which are 14 interriverine “islands” predicted to contain concentrations of taxa on unique evolutionary trajectories. These faunal units are named: east Trombetas, Trombetas-Negro, Negro-Japura, Japura-Putumayo, Putumayo- Napo, Napo-Maranon, Maranon-Ucayali, Ucayali-Jurua, Jurua-Purus, Purus-Madeira, Madeira-Tapajoz, Tapajoz-Xingu, Xingu-Tocantins, east ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 312 PACIFIC OCEAN ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 313 This pattern is not expected under the riverine barrier hypothesis. Unfortunately, such a transect would be logistically difficult. An interesting variant of prediction 3 involves a consequence of forest re-expansion across rivers. For example, a black-bodied subspecies of the Blue-crowned Manakin is proposed to have originated on the north bank of the Amazon in the Napo refugium. It is now found on the south bank because when the forest re-expanded, the black¬ bodied subspecies reached and crossed the Amazon. Under this prediction, one ex¬ pects a difference in diversity of genetic markers in samples of the black-bodied form on both sides of the river. The south bank forms were founders, probably not freely crossing the river, and therefore should show less allozyme heterozygosity and mtDNA clonal diversity. However, if the Amazon river merely bifurcated the two ranges there should be equal amounts on both banks. Similar situations should ex¬ ist with other taxa and other rivers in Amazonia. Prediction 4. The number of rare alleles will increase outward from the core of the refugium. If the founder populations are still increasing in number, then theoretical models predict an excess of rare alleles in those populations (Maruyama & Fuerst 1984). This prediction also assumes that the peripheral populations have not re¬ bounded to reach an equilibrium. This pattern is not expected under the riverine bar¬ rier hypothesis. Large sample sizes and appropriate genetic markers are needed to test this prediction. Problems with the riverine barrier hypothesis When examining patterns of phenotypic differentiation in Amazonia two fundamental observations can be used to argue against the riverine barrier hypothesis: 1) the in¬ consistency of congruence between phenotypic differentiation and rivers, and 2) the existence of suture zones, i.e., congruent contact zones between phenotypically dif¬ ferentiated taxa that occur in the middle of forest unassociated with rivers (Haffer 1987). As discussed earlier, lack of phenotypic differentiation does not mean lack of genetic differentiation. Therefore, it is critical to determine the patterns of genotypic differentiation. The demonstration of extensive genetic differentiation among phenotypically monomorphic, terra firme, forest-understory birds separated by rivers suggests that there is no inconsistency in congruence between genetic differentiation and rivers; therefore, the first argument is weakened. Due to our incomplete knowledge of the distribution of Amazonian birds (e.g. Parker & Remsen 1987) and sampling problems that compromise the thorough inventory at any one site (see phenotypic differentiation section), putative suture zones may be an artifact. If not, then the existence of suture zones not associated with rivers is the strongest evidence for the refugia hypothesis. One way to invoke the riverine barrier hypothesis is to assume a major river course change permitting the contact of formerly river-delimited forms and/or to attribute suture zones to the meeting after re-expan¬ sion of forest beyond the headwaters of rivers. While evidence for major river course changes has been presented (Salo et al. 1986), the acceptance of frequent changes in river courses is a problem for the riverine barrier hypothesis because it could lead to the prediction of a homogenization of regional diversity as formerly isolated interriverine areas come into contact (unless sufficient time for differentiation has occurred). 314 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI An alternative explanation is suggested by the environmental gradient hypothesis (Endler 1982). Endler proposes that present-day ecogeographic factors permit the development of clinal variation that can lead to parapatric speciation without geo¬ graphic isolation. Therefore, a suture zone may be an abrupt step in a cline. This assumes that gene flow is not sufficient to overcome the selective influence of the environmental gradient responsible for promoting differentiation. This view challenges the assumption that strong geographic differentiation in birds can only evolve with complete geographic isolation and implies that these avian contact zones are the re¬ sult of primary, not secondary, introgression. Recent work on isozyme variation within Amazonian birds in continuous forest has revealed levels of genetic differentiation higher than expected from studies of temper¬ ate zone species (Braun & Parker 1985, Capparella 1988). It is therefore possible that the response to an environmental gradient would produce differentiation without allopatry in tropical birds. To determine this it is necessary to effect a transect across a suture zone. Because suture zones are in the middle of forest this is logistically difficult. However, a transect of this type has recently been accomplished across a putative suture zone in southeastern Peru (several Louisiana State University Mu¬ seum of Natural Science expeditions), and the pattern of genetic differentiation should clarify the reality and significance of suture zones. CONSERVATION The distinction between the riverine barrier and refugia hypotheses is important from a conservation perspective. One approach to identifying forest areas for reserves in¬ volves the determination of putative refugia because these are considered to be the source areas for present-day biotic diversity (Wetterberg 1976, Gentry 1986). The evidence presented earlier casts doubt on the necessity for postulating refugia when potential vicariant elements — rivers — are known to be present. If putative refugia are not the centres of biotic diversity, then the refugia method of identifying areas for pre¬ serves is not valid. If rivers are the chief agents enhancing regional species diversity, then only thorough inventory of many sites within the Amazon basin will permit the identification of regions of unique biotic diversity for preservation. Using the analogy of “an archipelago of islands” (Snethlage 1913), coupled with cur¬ rent knowledge of patterns of avian differentiation, it is possible to identify 14 “islands” of terra firme habitats bounded by rivers (Figure 1). These interriverine areas should contain entities (specifically, terra firme forest-understory birds) on unique evolution¬ ary trajectories ( sensu Cracraft 1983), so-called “evolutionary significant units” which should be the objects of conservation management (Barrowclough & Flesness in press). Therefore, these 14 faunal units should be targeted for reserves to maximize the preservation of Amazonian biotic diversity. ACKNOWLEDGMENTS I thank colleagues at the Louisiana State University Museum of Natural Sciences for support, discussions, and interest in neotropical avian diversity, George Barrowclough (American Museum of Natural History) for challenging discussions, Jurgen Haffer for ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 315 critical comments and information on the latest formulation of the refugia hypothesis, the American Ornithologists’ Union and Illinois State University for travel funds, and the conveners of this symposium (Karl L. Schuchmann, Francois Vuilleumier) for their invitation. LITERATURE CITED AMADON, D. 1973. Birds of the Congo and Amazon forests: a comparison. Pp. 267-277 in Meggars, B. J., Ayensu, E. S., Duckworth, W. D. (Eds). Tropical forest ecosystems in Africa and South America: a comparative review. Washington, D. C., Smithsonian Institution Press. BARROWCLOUGH, G. F., FLESNESS, N. R. In press. Species, subspecies, and races: the problem of the units of management in conservation. In Harris, H., Allen, M. (Eds). Wild mammals in captivity. Chicago, University of Chicago Press. BATES, J. M., GARVIN, M. C., SCHMITT, D. C., SCHMITT, C. G. 1989. Notes on bird distribution in northeastern Dpto. Santa Cruz, Bolivia, with 15 species new to Bolivia. Bulletin British Ornithological Club 109: 236-244. BRAUN, M. J., PARKER, III, T. A. 1985. Molecular, morphological, and behavioral evidence concerning the taxonomic relationships of “Synallaxis” gularis and other synallaxines. Ornithological Monographs 36: 333-346. CAMPBELL, K. E., FRAILEY, D. 1984. Holocene flooding and species diversity in southwestern Amazonia. Quaternary Research 21: 369-375. CAPPARELLA, A. P. 1987. Effects of riverine barriers on genetic differentiation of Amazonian forest undergrowth birds. Ph.D. dissertation. Louisiana State University. CAPPARELLA, A. P. 1988. Genetic variation in neotropical birds: implications for the speciation proc¬ ess. Acta Congressus Internationalis Ornithologici XIX: 1658-1664. CAPPARELLA, A. P., ROSENBERG, G. H. In press. A new subspecies of Percnostola rufifrons (Formicariidae) from northeastern Amazonia Peru. Wilson Bulletin. COLINVAUX, P.A. 1989. Ice-age Amazon revisited. Nature 340: 188-189. COLINVAUX, P.A. 1987. Amazon diversity in light of the paleoecological record. Quaternary Science Reviews 6: 93-1 1 4. COLINVAUX, P.A., LIU, K.-B. 1987. The late-Quaternary climate of the western Amazon basin. Pp. 113-122 in Bergen, U. H. (Ed.). Abrupt climate change. Dordrecht, Reidel. CRACRAFT, J. 1983. Species concepts and speciation analysis. Current Ornithology 1: 159-187. CRACRAFT, J., PRUMM, R. 1988. Patterns and processes of diversification: speciation and histori¬ cal congruence in some neotropical birds. Evolution 42: 603-620. ENDLER, J. 1982. Pleistocene forest refuges: fact or fancy? Pp. 179-200 in Prance, G. (Ed.). Biological diversification in the tropics. New York, Columbia University Press. GENTRY, A. H. 1986. Endemism in tropical versus temperate plant communities. Pp. 153-161 in Soule, M.E. (Ed.). Conservation biology. Sunderland, Massachusetts, Sinauer Associates. GUTIERREZ, R. J., ZINK, R. M., YANG, S. Y. 1983. Genic variation, systematic, and biogeographic relationships of some galliform birds. Auk 100: 33-47. HACKETT, S., ROSENBERG, K. V. 1990. A comparison of genetic and phenotypic differentiation in antwrens of the genus Myrmotherula. Auk 107: 473-489. HAFFER, J. 1969. Speciation in Amazonian forest birds. Science 165: 131-137. HAFFER, J. 1974. Avian speciation in tropical South America. Publication Nuttall Ornithological Club No. 14. HAFFER, J. 1978. Distribution of Amazon forest birds. Bonner Zoologische Beitrage 29: 38-78. HAFFER, J. 1987. Biogeography of neotropical birds. Pp. 105-150 in Whitmore, T. C., Prance, G. T. (Eds). Biogeography and Quaternary history in tropical America. Oxford, Clarendon Press. HELMAYR, C. E. 1910. The birds of the Rio Madeira. Novitates Zoologicae 17: 257-428. ISLER, M. L., ISLER, P. R. 1987. The tanagers. Washington, D.C., Smithsonian Institution Press. MARUYAMA, T., FUERST, P.A. 1984. Population bottlenecks and nonequilibrium models in popula¬ tion genetics. I. Allele numbers when populations evolve from zero variability. Genetics 108: 745-763. MAYR, E. 1942. Systematics and the origin of species. New York, Columbia University Press. MEYER DE SCHAUENSEE, R. 1970. A guide to the birds of South America. Philadelphia, The Acad¬ emy of Natural Sciences. PARKER, III, T. A., REMSEN, JR., J. V. 1987. Fifty-two Amazonian bird species new to Bolivia. Bul¬ letin British Ornithological Club 107: 94-107. 316 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI PETERS, J. L. 1934-1987. Check-list of birds of the world (15 volumes). Cambridge, Massachusetts, Cambridge University Press. PRANCE, G. (Ed.) 1982. Biological diversification in the tropics. New York, Columbia University Press. SALO, J. 1987. Pleistocene forest refuges in the Amazon: evaluation of the biostratigraphical, lithostratigraphical and geomorphological data. Annales Zoologici Fennici 24: 203-211. SALO, J., KALLIOLA, R., HAKKINEN, I., MAKINEN, Y., NIEMELA, P., PUHAKKA, M., COLEY, P. D. 1986. River dynamics and the diversity of Amazon lowland forest. Nature 322: 254-258. SICK, H. 1967. Rios e enchentes na Amazonia como obstaculo para a avifauna. Pp. 495-520 in Lent, H. (Ed.). Atas do simposio sobre a biota amazonica, vol. 5 (Zoologia). Rio de Janeiro, Conselho de Pesquisas. SNETHLAGE, E. 1913. Uber die verbreitung der vogelarten in unteramazonien. Journal fur Ornithologie 61:469-539. TODD, W. E. C. 1927. New gnateaters and antbirds from tropical America, with a revision of the ge¬ nus Myrmeciza and its allies. Proceedings Biological Society Washington 40: 149-178. VAN DER HAMMEN, T. 1985. The Plio-Pleistocene climatic record of the tropical Andes. Journal Geo¬ logical Society London 142: 483-489. VAN ZINDEREN BAKKER, E. M. 1986. African climates and paleoenvironments since Messinian times. South African Journal of Science 82: 70-71. WALLACE, A. R. 1876. The geographical distribution of animals, volume I. London, Macmillan and Company. WETTERBERG, G. B. 1976. An analysis of nature conservation priorities in the Amazon. Brasilia, Bra¬ zilian Institute for Forestry Development. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 317 DISJUNCT BIRD DISTRIBUTIONS ALONG THE WEST SLOPE OF THE PERUVIAN ANDES IRMA FRANKE Museo de Historia Natural, Universidad Nacional de San Marcos, Casilla 14-0434, Lima 14, Peru ABSTRACT. Within the steppe-like western slope of the Peruvian Andes forest patches (dry cloud for¬ est) occur from Ecuador to 13°S. Eight sites were studied between Huamba and Zarate. A total of 124 taxa were used to analyze disjunctions. Species numbers decrease from N to S along a gradient of decreasing rainfall. Group 1 taxa (102 species and subspecies) occur from Ecuador south. Nine dis¬ tributional limits exist. Group 2 taxa (22 species and subspecies) include endemics. Five areas of taxon replacement were identified. The major replacement area is near the Rio Santa Valley. Birds now liv¬ ing in dry cloud forests along the western Andes originated north of this region. Distribution gaps have played an Important role in the history of the taxa. Keywords: Zoogeography, disjunction, Andes, dry cloud forests, Peru. INTRODUCTION The Pacific slope of the Peruvian Andes is dominated by arid, steppe-like vegetation types. However, many small forest patches occur in this dry zone between 2400 and 3000 m, forming the richest communities of the western slope of the Peruvian Andes (Koepcke, H.W. 1961, Valencia & Franke 1980, Franke & Valencia 1984). In north¬ ern Peru these dry cloud forests occupy extensive and relatively continuous areas, but further south they occur in increasingly smaller and more isolated patches, to about 13°S (Koepcke, H.W. 1961, Valencia 1990). These disjunct forests are thought to represent fragments of a formerly more continuous forest zone (Koepcke, M. 1958; Koepcke, H.W. 1961). The floristic and faunistic affinities between west slope and east slope forests, as well as the existence of low passes in northern Peru, especially the Porculla Pass (2145 m), led to the hypotheses that east slope species (1) crossed the low passes to the west, and (2) dispersed southward through the forest belt (Koepcke, M. 1958, 1961b, Koepcke, H.W. 1 961 , Simpson 1 975). Biogeographers accept the idea that the Andean montane forests were more continuous when depressed altitudinally during cool, humid glacial periods. At such times temperate and subtropical zones were lo¬ cated along less dissected lower slopes. Conversely, the upward shift of climatic zones during interglacial periods led to increasing fragmentation of montane vegeta¬ tion (Haffer 1987), thus isolating bird populations and playing a role in speciation along the western Peruvian Andes. METHODS Field studies were made at 8 dry cloud forest sites between 2500 and 3000 m, from 4°41 ’S, near the Ecuadorian border, to 1 1 °55’S in central Peru: (1 ) Huamba, Depart¬ ment of Piura (4°41’S, 2900 m); (2) Chinama, Lambayeque (6°06’S, 2550 m); (3) Llaguen, La Libertad (7°42’S, 2600 m); (4) Cochabamba, Ancash (9°27’S, 2800 m); 318 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI (5) Winapajatun, Ancash (9°41’S, 2600 m); (6) Noqno, Ancash (10°03’S, 2850 m); (7) Linday, Lima (11°50’S, 3000 m), and (8) Zarate, Lima (11°55’S, 2850 m). Distributional records of the species observed at the 8 sites (Appendix 1) are based on the author’s field work, museum specimens (Museo de Historia Natural, Universidad de San Marcos, Lima; Louisiana State University Museum of Natural History, Baton Rouge; National Museum of Natural History, Washington; Field Mu¬ seum of Natural History, Chicago; and British Museum of Natural History, Tring), and the literature (Koepcke M. 1958, 1961a, b, 1965, Valencia & Franke 1980, Franke & Valencia 1984, Hellmayr, Conover & Cory 1918-1949, Zimmer 1931-1975, Fjeldsa & Krabbe 1990, Parker et al. 1985, Meyer de Schauensee 1966, Schulenberg 1987, Schulenberg & Parker 1981, Parker 1981, O’Neill & Schulenberg 1979, Plenge 1974). For each species the records were plotted on a map of the study area in order to determine the latitudinal limits of each species. After eliminating low altitude species that reach the dry cloud forests only occasionally, a list of 128 species was obtained. For 9 of these species only scattered records exist and these species were eliminated. In several of the 119 remaining spe¬ cies, more than one subspecies occur in the study area. Given relatively minor differ¬ ences between most subspecies, geographic variation may actually be clinal. Since subspecies present difficulties in biogeographic analysis (Cracraft 1985) they were avoided, with one exception. Four species have morphologically distinct subspecies ranging along the latitudinal gradient studied: Cranioleuca antisiensis (3 subspecies), Aglaeactis cupripennis (2), Lepthastenura pileata (2) and Atlapetes seebohmi (2). In spite of Cracraft’s (1985) caveat I have included the subspecies of these four species because they are well marked and present interesting distributional information. Thus, 124 taxa (1 16 species and 8 subspecies) were analyzed in the present paper. RESULTS The number of species in dry cloud forests decreases markedly from north to south, from 102 species in northwestern Peru to 56 in central Peru (Figure 1). The N - S de¬ crease in species numbers is significantly correlated (r=0.8723, P>0.001) with the gradient of decreasing rainfall. In turn vegetation characteristics of the dry cloud for¬ ests reflect the rainfall gradient (Valencia 1990). The 124 taxa are divided into two groups. In the first group I place 102 taxa that oc¬ cur in Ecuador and have a continuous or nearly continuous distribution southward to a certain point along the western slope. In the second group (22 taxa) I include spe¬ cies and subspecies that do not occur in Ecuador, but have a continuous or relatively continuous distribution along part of the western slopes (Appendix 1). DISTRIBUTION PATTERNS OF GROUP 1 TAXA The 102 taxa in group 1 can be divided into nine subgroups, with southern limits at the following areas: (1) Porculla Pass (7 taxa, 5°51’S); (2) Chinama (7 taxa, 6°06’S); (3) Rio Chancay Valley (4 taxa, 6°40’S); (4) Rio Saha Valley (26 taxa, 6°54’S); (5) Rio Jequetepeque Valley (2 taxa, 7°19’S); (6) Rio Chicama Valley (4 taxa, 7°29’S); ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 319 (7) Llaguen (8 taxa, 7°42'); (8) Cordilleras Blanca and Negra (5 taxa, ca. 10°S); (9) 39 taxa occurring at the southern limit of the study area (1 1°55’S) but not necessar¬ ily having their southern distributional limit there. A plot of numbers of taxa in group 1 against latitude shows a N - S decreasing trend (Figure 1). This trend is highly correlated with the rainfall gradient (r= 0.9128, P<0.001 ) and very similar to the trends obtained for vegetation parameters (number of woody species, density, basal area and vertical structure; Valencia 1990). These results strongly suggest that climate and habitat conditions are important factors in these distributions patterns. - 1 - 1 - 1 - ~~i - - 1 - 1 — - 1 - 1 5 6 7 8 9 1 0 11 12 LATITUDE (DEG.S) FIGURE 1 - Number of avian taxa occurring in dry cloud forests along the western slope of the Peruvian Andes, a) Total number of taxa b) Taxa with distributions extending from Ecuador southward. DISTRIBUTION PATTERNS OF GROUP 2 TAXA The second group consists of 22 taxa including species endemic to western Peru, several of which have a very restricted distribution. Nine subgroups of closely related taxa replace each other latitudinally along the study area (Figure 2). Three of these subgroups consist of subspecies: Cranioleuca antisiensis (3 subspecies), Lepthastenura pileata (2) and Aglaeactis cupripennis (2). Three subgroups consist of species that are often treated as subspecies: Anairetes nigrocristatus and A. reguloides, Saltator nigriceps and S. aurantiirostris, and Atlapetes seebohmi and A. nationi. Finally, the last three subgroups consist of congeneric species: Synallaxis elegantior and S. zimmeri, Ochthoeca piurae and O. leucophrys, and Diglossa humeralis and D. brunneiventris. Only two of these subgroups show distributional overlap ( Anairetes and Ochthoeca species), where the deep Rio Santa Valley sepa¬ rates the Cordilleras Blanca and Negra. 320 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 4- Sa 4. An + Db Lpp Acc Caz *OI Anr 5 6 7 8 9 10 11 12 LATITUDE ( DEG. S) FIGURE 2 - Latitudinal distribution of nine groups of related taxa that replace each other along the western slope of the Peruvian Andes. Abbreviations: Ann, Anairetes nigrocristatus ; Anr, A. reguloides ; Op, Ochthoeca piurae\ 01, O. leucophrys ; Cap, Cranioleuca antisiensis palamblae; Cab, C. a. baron'r, Caz, C. a. zaratensis ; Acp, Aglaeactis cupripermis parvulus ; Acc, A. c. caumatonotus; Lpc, Lepthastenura pileata cajabambae ; Lpp, L. p. pileata ; Dh, Diglossa humeralis ; Db, D. brunneiventris ; Ass, Atlapetes seebohmi seebohmi; An, Atlapetes nationi, Sn, Saltator nigriceps ; Sa, S. aurantiirostris ; Se, Synallaxis elegantior, Sz, S. zimmeri. Shaded areas: major areas of species or subspecies replacement. Five major areas of species or subspecies replacement or range limits (Figure 2) are listed below. (1) Around 5°20’S, near El Tambo, Piura; northern limit of Anairetes nigrocristatus and Ochthoeca piurae. (2) Around 6°30’S, near Llama and Chugur, Cajamarca, deep valley of Rio Chancay (upper Rio Reque); contact between Diglossa humeralis and D. brunneiventris (Graves 1982, Vuilleumier 1984), replacement of Saltator nigriceps by S. aurantiirostris and Cranioleuca antisiensis palamblae by C.a. baron i. (3) Around 7°30’S, near Pluacraruco, Sunchubamba and Llaguen, deep valleys of Rios Jequetepeque and Chicama; replacement of Synallaxis elegantior by S. zimmeri, southern limit of Aglaeactis cupripennis parvulus, northern limit on the western slope of Atlapetes seebohmi seebohmi, Lepthastenura pileata cajabambae, Anairetes reguloides, Polyonymus caroli, and Chrysoptilus atricollis. Se <+ Sn Dh <4* Acp Cap 4- ElTambo Rio RiosChicama& Rio Area Chancay Jequetepeque Santa ▼ ▼ ▼ ▼ O p Ann Cab, Ass:-' '■ L pc : Rio Pativilca ▼ Sz ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 321 (4) Between 8°30' and 9°S, Rio Santa Valley. Area no. 4 is particularly important. The Rio Santa is the largest river of the Pacific watershed and forms the largest val¬ ley of western Peru. The upper Rio Santa, running for 180 km parallel to the coast before turning westward separates the eastern and very high Cordillera Blanca from the western, lower Cordillera Negra. Avian distribution patterns in this area are complex. Most species occurring north of the Rio Santa Valley have a relatively extensive range and are present in both the Cordilleras Blanca and Negra. A few species, however, have only been reported from the Cordillera Blanca (e.g. Mecocerculus leucophrys, Ochthoeca rufipectoralis). Four of the groups of taxa (see above) show replacement or separation in the Rio Santa Valley. (1) Northerly Anairetes nigrocristatus occurs in the Cordillera Blanca, whereas southerly A. reguloides occurs in the Cordillera Negra. There is only one possible record of reguloides from the Cordillera Blanca (Frimer & Nielsen 1989). (2) Cranioleuca antisiensis baroni (northerly), occurs only in the Cordillera Blanca, whereas C. a. zaratensis (southerly), occurs in the Cordillera Negra. Fjeldsa and Krabbe (1990:358-359) consider baroni a full species. They further state that C. baroni baroni “crosses to Pacific slope of Cordillera Negra” in Ancash. However, the 9 specimens col¬ lected during my study in the Cordillera Negra (Ancash) have the same charac¬ ters as the 10 specimens of zaratensis from the Department of Lima and not the characters of baroni. (3) Aglaeactis cupripennis caumatonotus occurs on both cordilleras, but has not been recorded on the western slope north of the Santa Valley, where the specimens available correspond to A. c. parvulus. (4) Ochthoeca piurae and O. leucophrys are sympatric in the Cordillera Negra, but the latter species also occurs in the Cordillera Blanca. It is noteworthy that Synallaxis zimmeri, iike O. piurae an endemic species restricted to a small area of the Pacific slope, occurs along the Cordillera Negra and further north in Lambayeque. These taxa have not yet been recorded from Cordillera Blanca or any other locality within the Santa Valley. (5) Around 10°30’S, Rio Pativilca Valley; replacement of Lepthastenura pileata cajabambae by L. p. pileata and Atlapetes seebohmi seebohmi by A. nationi ; southern limit of Synallaxis zimmeri and Ochthoeca piurae . DISCUSSION Northern origin The distribution patterns described in this paper, especially the patterns of species replacement and/or species limits at given areas along the western slope of the Pe¬ ruvian Andes, supports the concept of a northern origin for most avifaunal elements of dry cloud forests. The N - S decrease in species numbers is also correlated to the decrease in rainfall and associated vegetation parameters (number of woody species, density, basal area and vertical structure, Valencia 1990). Thus, historical as well as ecological factors have had important effects on avian distribution. The complex distribution patterns found in Ancash also support the idea of a north¬ ern origin. If the species dispersed from north to south, they would be expected to have occupied the appropriate habitats in the Rio Santa Valley, including the 322 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Cordillera Blanca, before they dispersed to the Cordillera Negra. Several species with a southern limit in Ancash show this pattern, as do also the Cranioleuca and Anairetes groups. In these instances, the northern species (or subspecies) occupies the Cordillera Blanca, whereas the southern one occurs from the Cordillera Negra south¬ ward. » The distribution of Synallaxis zimmeri and Ochthoeca piurae does not correspond to this pattern, however. These 2 species occur north of the Santa Valley, in La Libertad and along the Cordillera Negra, but they are not recorded in the Santa Valley. The dif¬ ferences between the patterns in Cranioleuca and Anairetes on the one hand, and in S. zimmeri and O. piurae on the other, can be explained by the fact that species oc¬ cupying the interior of the Santa Valley, Cordillera Blanca, range up to around 4000 m, whereas Synallaxis zimmeri and Ochthoeca piurae occur lower, from 1500- 1800 m to about 2800 m. Both species occur in open, dry habitats as well as in dry cloud forests. It is thus not surprising that these species did not reach the upper Santa Valley, but instead dispersed southward along the lower slopes. Efficacy of barriers Although the efficacy of low passes in the western cordillera as barriers to dispersal of montane species has been questioned (Parker et al. 1985), it is intriguing to note that many range limits coincide with areas where the western Andes are lowest (Vuilleumier 1969, 1977) and with the deepest valleys of the Pacific drainage. This distribution gap could be an artifact resulting from incomplete sampling in the area. Only further investigation will resolve this issue unequivocally. However, a study of the distribution of 306 dry cloud forest patches shows that several important latitudinal gaps coincide with the low areas of the western Andes and with the deep valleys of the Pacific slope (Valencia 1990). If montane forests were more continuous in the past during cool and humid glacial periods, when they occupied the lower, more continu¬ ous mountain slopes, then the low passes and the deep valleys must have played an important role in fragmenting the forest zone as it shifted upward during interglacial periods. The increasing patchiness of the forest resulted probably not only from the more complex topography of the upper slopes (Haffer 1987), but also from the retreat of forest patches to favorable slopes. Valencia (1990) has shown the tendency of for¬ est patches to be restricted to more humid slopes. The low passes must also have had an effect in dissecting the forest area. As mentioned earlier, dry cloud forests occur generally between 2400 and 3000 m. Summits in the low Andes of northwest¬ ern Peru are below 3000 m, thus restricting considerably the potential extension of forest. Vuilleumier (1969, 1977, 1984) presented models of the effects of the Pleistocene depression in northern Peru. Main gaps Three gaps are especially important. The first gap, between 6°30' and 6°47’S, corre¬ sponds to the Rio Chancay (upper Rio Reque) Valley, Cajamarca, where the western Andes are low and narrow. The contact between Diglossa humeralis and Diglossa brunneiventris is found here (Graves 1982, Vuilleumier 1984). Koepcke (1961b) and Zimmer (1942) considered the low pass in this area to be the dispersal route by which Cranioleuca antisiensis baroni and Cyclarhis gujanensis reached the western slope. This area may have been more important for dispersal and speciation of birds than the Porculla Pass. Even though Porculla is the lowest pass in northwestern Peru, it occurs in a relatively linear area of the western Andes, thus presenting no special ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 323 problem for dispersal (Vuilleumier 1984 discussed the barrier effect of the Porculla Pass and surrounding areas). The second gap, between 8°28' and 8°49’S, correponds to the Rio Santa Valley and is wide (about 50 km). This valley separates Cranioleuca antisiensis baroni from C. a. zaratensis, and Anairetes nigrocristatus from Anairetes reguloides. In both cases the northern taxon occurs in the Cordillera Blanca, while the southern one is distrib¬ uted from the Cordillera Negra southward. The third gap, between 10°30’ and 10°45’S, corresponds to the Rio Pativilca Valley. This valley separates two pairs of taxa, Lepthastenura pileata cajabambae from L. p. pileata, and Atlapetes seebohmi seebohmi from Atlapetes nationi, and marks the southern limit of west slope endemics, Ochthoeca piurae and Synallaxis zimmeri. ACKNOWLEDGEMENTS I thank Karl-L. Schuchman and F. Vuilleumier for their invitation to participate in this symposium. Attendance at the I.O. C. was made possible by a special travel grant to F. Vuilleumier, a grant from the American Ornithologists Union, and assistance from the organizers of the 20th I.O.C. I am thankful to M.L. Gorman for his advice. For per¬ mitting me to examine specimens under their care and providing other courtesies, I am grateful to J. O’Neill and V. Remsen of Louisiana State University Museum of Natural History; J. Fitzpatrick and S. Lanyon of the Field Museum of Natural History; M. Foster of the National Museum of Natural History and A. Knox and P. Colston of the British Museum of Natural History. Field work was supported by the Frankfurt Zoological Society (project 905/81) and the Consejo Nacional de Ciencia y Tecnologia (CONCYTEC) of Peru. LITERATURE CITED CRACRAFT, J. 1985. Historical biogeography and patterns of differentiation within the South Ameri¬ can avifauna: areas of endemism. Pp 49-84 in Buckley, P.A., Foster, M.S., Morton, E.S., Ridgely, R.S., Buckley, F.G. (Eds). Neotropical Ornithology, Ornithological Monographs No. 36, Washington, D.C., American Ornithologists’ Union. FJELDSA, J., KRABBE, N. 1990. Birds of the High Andes. Zoological Museum, University of Copen¬ hagen. FRANKE, I., VALENCIA, N. 1984. Zarate: una unidad de conservacion. Report. Museo de Historia Natural, Universidad Nacional Mayor de San Marcos. FRIMER, O., NIELSEN, S.M. 1989. The status of Polylepis forests and their avifauna in Cordillera Blanca, Peru. Technical report from an inventory in 1988, with suggestions for conservation manage¬ ment. Copenhagen. GRAVES, G.R. 1982. Speciation in the Carbonated flower-piercer (Diglossa carbonaria ) complex of the Andes. Condor 84:1-14. HAFFER, J. 1987. Quaternary history of tropical America. Pp 1-18 in Whitmore, T.C., Prance, G.T. (Eds). Biogeography and Quaternary History in Tropical America. Oxford: Claredon Press. HELLMAYR, C.E., CONOVER, B. CORY, C.B. 1918-1949. Catalogue of birds of the Americas. 1 1 vols. Chicago: Field Museum of Natural History Publications 197; 223; 242; 266; 330; 347; 365; 381; 430; 514; 615; 616; 634. KOEPCKE, H.W. 1961. Synoekologishe Studien an der Westseite der peruanischen Anden. Bonner Zoologischer Beitraege 29: 1-320. KOEPCKE, M. 1958. Die Vogel des Waldes von Zarate (Westhang der Anden in Mittelperu). Bonner Zoologischer Beitraege 9:130-193. 324 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI KOEPCKE, M. 1961a. Birds of the western slope of the Andes of Peru. American Museum Novitates N. 2028. KOEPCKE, M. 1961b. Las razas geograficas de Cranioleuca antisiensis (FURNARIIDAE, AVES), con la descripcion de una nueva subespecie. Publicaciones del Museo de Historia Natural “Javier Prado”. Ser. A. Zoologia. No. 20. KOEPCKE, M. 1965. Zur Kenntnis einiger Furnariiden (Aves) der Kuste und des westlichen Andenabhanges Perus. Beitrag zur Neotropischen Fauna IV: 1 50-1 73. MEYER DE SCHAUENSEE, R. 1966. The species of birds of South America and their distribution. Narberth, Penn. Livingston Publishing Company. O’NEILL, J.P., SCHULENBERG, T.S. 1979. Notes on the Masked Saltator, Saltator cinctus, in Peru. Auk 96:610-613. PARKER, T.A.,III. 1981. Distribution and biology of the White-cheeked Cotinga Zaratornis stresemanni, a high Andean frugivore. Bulletin of the British Ornithologists’ Club 101: 256-265. PARKER, T.A., III; SCHULENBERG, T.S.; GRAVES, G.R., BRAUN, M.J. 1985. The avifauna of the Huancabamba region, northern Peru. Pp. 169-197 in Buckley, P.A., Foster, M.S., Morton, E.S., Ridgely, R.S., Buckley, F.G. (Eds). Neotropical Ornithology, Ornithological Monographs No. 36, Washington, D.C., American Ornithologists’ Union. PLENGE, M.A. 1974. Notes on some birds in west-central Peru. Condor 76: 326-330. SCHULENBERG, T.S. 1987. New records of birds from western Peru. Bulletin of the British Ornitholo¬ gists’ Club 107: 184-189. SCHULENBERG, T.S., PARKER, T.A.,111. 1981. Status and distribution of some northwestern Peruvian birds. Condor 83: 209-216. SIMPSON, B.B. Pleistocene changes in the flora of the high tropical Andes. Paleobiology 1: 273-294. VALENCIA, N. 1990. Ecology of forests on the western slopes of the Peruvian Andes. Ph. D. Thesis. Aberdeen University. VALENCIA, N., FRANKE, I. 1980. El bosque de Zarate y su conservacion. Boletin de Lima. 7: 76-86; 8: 26-35. VUILLEUMIER, F. 1969. Pleistocene speciation in birds living in the high Andes. Nature 223: 1179- 1180. VUILLEUMIER, F. 1977. Barrieres ecogeographiques permettant la speciation des oiseaux des hautes Andes. Pp. 29-51 in Descimon, H. (Ed. ). Biogeographie et evolution en Amerique tropicale Ecole Normale Superieure, Paris. Publications du Laboratoire de Zoologie, Numero 9. VUILLEUMIER, F . 1984 . Zoogeography of Andean birds : two major barriers; and speciation and tax¬ onomy of the Diglossa carbonaria superspecies. National Geographic Society Research Reports 16: 713-731. ZIMMER, J.T. 1931-1955. Studies of Peruvian birds. American Museum Novitates 500, 509, 523, 524, 538, 545, 558, 584, 646, 647, 668, 703, 728, 753,756, 757, 785, 819, 860, 861, 862, 889, 893, 894, 917, 930, 962, 963, 994, 1042, 1043, 1044, 1045, 1066, 1095, 1108, 1109, 1126, 1127, 1159, 1160, 1168, 1193, 1203, 1225, 1245, 1246, 1262, 1263, 1304, 1345, 1367, 1380, 1428, 1449, 1450, 1463, 1474, 1475, 1513, 1540, 1595, 1604, 1609, 1649, 1723. APPENDIX 1 Avian taxa recorded at 8 dry cloud forest sites along the western slope of the Peru¬ vian Andes. Code number indicates distribution southward from Ecuador. (1) To Porculla Pass, 5°51’S; (2) To Chinama, 6°06’S; (3) To Rio Chancay Valley, 6°40’S; (4) To Rio Saha Valley, 6°54’S; (5) To Rio Jequetepeque Valley, 7°19’S; (6) To Rio Chicama Valley, 7°29’S; (7) To Llaguen, 7°42’S; (8) To Cordilleras Blanca and Negra, ca. 1 0°S; (9) To Zarate, 1 1 °55’S. (10) Avian forms that do not occur in Ecuador; (1 1 ) Avian forms with scattered records from western Peru; (12) Low altitude species. Taxa are listed as species (family). * Migrant from North America; all other species are “resident”. Distribution pattern 1: Pipreola arcuata (Cotingidae), Myiotheretes fumigatus (Tyrannidae), Troglodytes solstitialis (Troglodytidae), Saltator cinctus (Cardinalidae), Hemispingus verticalis (Thraupidae), Trogon personatus (Trogonidae), Cyanolyca turcosa (Corvidae). ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 325 Distribution pattern 2: Geotrygon frenata (Columbidae), Piculus rivolii (Picidae), Cranioleuca antisiensis palamblae (Furnariidae), Phyllomyias uropygialis (Tyrannidae), Elaenia pallatangae (Tyrannidae), Pachyramphus albogriseus (Cotingidae), Catamblyrhynchus diadema (Catamblyrhynchidae). Distribution pattern 3: Diglossa humeralis (Coerebidae), Saltator nigriceps (Cardinalidae), Colibri thalassinus (Trochilidae), Lafresnaya lafresnayi (Trochilidae). Distribution pattern 4: Penelope barbata (Cracidae), Ciccaba albitarsus (Strigidae), Coeligena iris (Trochilidae), Ensifera ensifera (Trochilidae), Pharomachrus auriceps (Trogonidae), Lepidocolaptes affinis (Dendrocolaptidae), Margarornis squamiger (Furnariidae), Pseudocolaptes biossonneautii (Furnariidae), Automolus ruficollis (Furnariidae), Grallaria ruficapilla (Formicariidae), Phyllomyias nigrocapillus (Tyrannidae), Mionectes striaticollis (Tyrannidae), Contopus fumigates (Tyrannidae), Turdus serranus (Turdidae), Atlapetes leucopterus (Emberizidae), Atlapetes torquatus (Emberizidae), Tangara viridicollis (Thraupidae), Anisognathus lacrymosus (Thraupidae), Thraupis cyanocephala (Thraupidae), Hemispingus superciliaris (Thraupidae), Diglossa cyanea (Coerebidae), Myioborus melanocephalus (Parulidae), Basileuterus coronatus (Parulidae), Conirostrum sitticolor (Coerebidae), Cyclarhis gujanensis (Vireonidae), Vireo gilvus (Vireonidae). Distribution pattern 5: Accipiter striatus (Accipitridae), Synallaxis elegantior (Furnariidae). Distribution pattern 6: Mecocerculus stictopterus (Tyrannidae), *Catharus fuscater (Turdidae), Atlapetes rufinucha (Emberizidae), Tangara vassorii (Thraupidae). Distribution pattern 7: Adelomyia melanogenys (Trochilidae), Aglaeactis cupripennis parvulus (Trochilidae), Heliangelus viola (Trochilidae), Scytalopus unicolor (Rhinocryptidae), Hemispingus melanotis (Thraupidae), Myioborus miniatus (Parulidae), Basileuterus nigrocristatus (Parulidae), Basileuterus trifasciatus (Parulidae). Distribution pattern 8: Veniliornis fumigates (Picidae), Turdus fuscater (Turdidae), Nyctidromus albicollis (Caprimulgidae), Mecocerculus leucophrys (Tyrannidae), Ochthoeca rufipectoralis (Tyrannidae). Distribution pattern 9: Nothoprocta pentlandii (Tinamidae), Geranoaetus melanoleucus (Accipitridae), Buteo polyosoma (Accipitridae), Falco sparverius (Falconidae), Columba fasciata (Columbidae), Zenaida auriculata (Columbidae), Leptotila verreauxi (Columbidae), Aratinga wagleri (Psittacidae), Glaucidium jardinii (Strigidae), Caprimulgus longirostris (Caprimulgidae), Colibri coruscans (Trochilidae), Patagona gigas (Trochilidae), Lesbia nuna (Trochilidae), Metallura tyrianthina (Trochilidae), Ampelion rubrocristatus (Cotingidae), Elaenia albiceps (Tyrannidae), Anairetes parulus (Tyrannidae), Contopus cinereus (Tyrannidae), Ochthoeca jelskii (Tyrannidae), Myiotheretes striaticollis (Tyrannidae), Agriornis montana (Tyrannidae), Muscisaxicola maculirostris (Tyrannidae), Myiarchus tuberculifer (Tyrannidae), Notiochelidon murina (Hirundinidae), Notiochelidon cyanoleuca (Hirundinidae), Cinclus leucocephalus (Cinclidae), Troglodytes aedon (Troglodytidae), Turdus chiguanco (Turdidae), Zonotrichia capensis (Emberizidae), Phrygilus plebejus (Emberizidae), Catamenia analis (Emberizidae), Catamenia inornata (Emberizidae), Pheucticus chrysogaster (Cardinalidae), Thraupis bonariensis (Thraupidae), Piranga flava (Thraupidae), Thlypopsis ornata (Thraupidae), Diglossa sittoides (Coerebidae), Conirostrum cinereum (Coerebidae), Carduelis magellanica (Carduelidae). 326 ACTA XX CONGRESSUS INTERNATIONAL^ ORNITHOLOGICI Distribution pattern 10: Leptasthenura pileata pileata (Furnariidae), Atlapetes nationi (Emberizidae), Leptasthenura pileata cajabambae (Furnariidae), Synallaxis zimmeri (Furnariidae), Atlapetes seebohmi seebohmi (Emberizidae), Ochthoeca piurae (Tyrannidae), Aglaeactis cupripennis caumatonotus (Trochilidae), Cranioleuca antisiensis zaratensis (Furnariidae), Asthenes pudibunda (Furnariidae), Ampelion stresemanni (Cotingidae), Phrygilus fruticeti (Emberizidae), Poospiza rubecula (Emberizidae), Cranioleuca antisiensis baroni (Furnariidae), Polyonymus caroli (Trochilidae), Chrysoptilus atricollis (Picidae), Anairetes reguloides (Tyrannidae), Aeronautes andecolus (Apodidae), Metallura phoebe (Trochilidae), Saltator aurantiirostris (Cardinalidae), Diglossa brunneiventris (Coerebidae), Metriopelia ceciliae (Columbidae), Ochthoeca leucophrys (Tyrannidae). Distribution pattern 11: Claravis mondetoura (Columbidae), Bolborhynchus orbygnesius (Psittacidae), Otus koepckeae (Strigidae), Otus sp (Strigidae), Aegolius harrissi (Strigidae), Uropsalis segmentata (Caprimulgidae), *Catharus ustulatus (Turdidae), Atlapetes seebohmi simonsi (Emberizidae), Pipraeidea melanonota (Thraupidae). Distribution pattern 12: Columbina cruziana (Columbidae), Amazilia amazilia (Trochilidae), Thaumastura cora (Trochilidae), Myrtis fanny (Trochilidae), Piculus rubiginosus (Picidae), Myiopagis subplacens (Tyrannidae), Euscarthmus meloryphus (Tyrannidae), Pachyramphus homochrous (Cotingidae), Campylorhynchus fasciatus (Troglodytidae), Turdus reevei (Turdidae), Poospiza hispaniolensis (Emberizidae), Parula pitiayumi (Parulidae), Dives warszewiczi (Icteridae). / ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 327 SPECIATION IN PATAGONIAN BIRDS FRANQOIS VUILLEUMIER Department of Ornithology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA ABSTRACT. The 138 genera and 218 species of iandbirds and waterbirds breeding in Patagonia (south-temperate South America) were surveyed for evidence of speciation phenomena, especially presence of geographical isolates, secondary contact zones, and broad geographic overlaps between closely related congeners. Many of the speciation patterns revealed in this analysis can be correlated with late Pleistocene vicariance events associated with glaciation cycles. Keywords: South America, Patagonia, biogeography, avifauna, speciation, vicariance, Pleistocene events. INTRODUCTION In order to expand the geographic scope of my studies of speciation in Andean birds, I surveyed the 138 genera and 218 species of land- and waterbirds breeding in Patagonia (south-temperate South America from 36°-38°S to about 56°S). In this re¬ port I describe speciation phenomena in selected taxa of Patagonian birds, in an at¬ tempt to: (a) identify the ecological and geographical circumstances permitting speciation to take place today, and (b) verify the hypothesis that environmental changes in the late Pleistocene were conducive to speciation. The present paper is parallel to an essay on speciation in high Andean birds presented at the XVI Ith I.O. C. (Vuilleumier 1980) and is written in the same format to facilitate comparisons. METHODS AND MATERIAL Assumptions As in my survey of high Andean birds I assume that speciation is allopatric (Mayr 1963) and that vicariance events can be reconstructed after “one assesses phylogenetic relationships among species, maps their distribution, and documents the nature of isolation or sympatry among taxa of a given genus” (Vuilleumier 1980: 1256) Four kinds of phenomena are important in this regard (1) geographical isolates within species; (2) geographical isolates between semispecies and allospecies (Amadon 1966); (3) secondary contacts between formerly isolated species-level taxa; and (4) broad range overlaps between closely related congeners (sister species). Patagonian vegetation and avifauna Temperate rainforests dominated by beech ( Nothofagus ) in western Patagonia, and dry to arid steppes with tussock grass or scrub east of the Andes, are the two main vegetation types. Other vegetation formations (Magellanic moorland and alpine scrub) cover smaller areas (Hueck & Seibert 1972). I assign 229 species (1 1 oceanic, 53 littoral or freshwater, and 165 terrestrial) to the breeding avifauna. The 11 oceanic species (Spheniscidae, Diomedeidae, 328 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Procellariidae, Hydrobatidae, Pelecanoididae, Sulidae) are excluded from the survey, which deals with 218 species (53 aquatic, 165 terrestrial). The analysis is based on a combination of museum and literature research, supple¬ mented by extensive field work (February-March 1965, November-December 1985, February-March 1987, October 1987, January 1988, November 1988). RESULTS Geographical isolates within species Intra-Patagonian disjunctions (1) Phalacrocorax atriceps (Phalacrocoracidae). P. atriceps and P. albiventer are considered conspecific (Devillers & Terschuren 1978; Siegel-Causey 1986). Coastal areas. Isolated populations breed on inland lakes in northern Patagonia ( lacustris ; Navas 1970) and southern Patagonia (Reynolds 1934). These isolates are separated from littoral populations by landscapes including forest, steppe, and mountain. (2) Cinclodes oustaleti (Furnariidae). Alpine vegetation. The northern Patagonian populations ( oustaleti ) are separated from the southern ones in the Tierra del Fuego archipelago ( hornensis ) by a large hiatus (Vuilleumier unpubl.; map in Fjeldsa & Krabbe 1990). The nature of the barrier is unclear, since suitable looking habitat oc¬ curs in much of the intervening montane region. (3) Leptasthenura aegithaloides (Furnariidae). Open scrub and thorny bushes. An isolated population as discovered and studied by Vuilleumier (unpubl.) in 1985-1988 in NW Tierra del Fuego. The nearest known mainland population (Vuilleumier unpubl.) occurs across the Strait of Magellan in Magallanes. (4) Xolmis pyrope (Tyrannidae). Forest and forest edge, locally matorral (Tierra del Fuego). The Chiloe Island population ( fortis ) is weakly differentiated from the main¬ land one and separated by a narrow water gap (Vuilleumier 1985). (5) Cistothorus platensis (Troglodytidae). Marsh grassland. The Patagonian populations appear to consist of two disjunct groups (1 1 and 12 in Traylor 1988) that show weak differentiation. The nature of the apparent gap between these isolates may correspond to the area occupied by the Patagonian icecap. Extra-Patagonian disjunctions (1) Pterocnemia pennata (Rheidae). Grassy steppe. Patagonian populations ( pennata ) are isolated from high Andean ones living in the dry puna ( tarapacensis and garleppi) by a gap (map in Fjeldsa & Krabbe 1990) of about 800 km of montane terrain, some of which would appear suitable for occupation (Vuilleumier unpubl.). (2) Strix rufipes (Strigidae). Nothofagus forest. The Patagonian form ( rufipes ) is sepa¬ rated from the chaco form ( chacoensis ) (Short 1975) by dry monte woodland and Patagonian steppe. (3) Picoides lignarius (Picidae). Nothofagus forest and edge. Two morphologically in¬ distinguishable isolates, one in Patagonia and the other in Bolivia, are separated by about 1000 km of vegetation including woodland, scrub, and monte, partially occupied ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 329 by allospecies mixtus (Short 1982). Competitive exclusion could maintain the isola¬ tion. Geographical isolates between semi- or allospecies Intra-Patagonian disjunctions (1) Pteroptochos (Rhinocryptidae). The two Nothofagus- inhabiting taxa are separated by the Bio-Bio River, castaneus occurring north, and tarnii south of this barrier. I consider these taxa as members of a superspecies (Vuilleumier 1985) but others (Fjeldsa & Krabbe 1990) keep them conspecific. (2) Scytalopus (Rhinocryptidae). Northern taxon fuscus (found in more open situa¬ tions, contra Fjeldsa & Krabbe 1990) and southern one magellanicus (breeding in dense Nothofagus forest, again contra Fjeldsa & Krabbe 1990) appear to meet at or near the Bio-Bio River. Fjeldsa & Krabbe (1990) treat these two forms as species that show considerable sympatry. They are borderline cases. Whether there is sympatry seems open to question in the absence of specimen records. Extra-Patagonian disjunctions (1) Charadrius (Charadriidae). Patagonian taxon falklandicus (coastal pebble beaches but also inland areas) is closely related to high Andean alticola of the puna (borderline case: they are either subspecies or allospecies), and to New Zealand taxon bicinctus. Bock (1958) considers falklandicus and bicinctus to form a superspecies. Fie suggested (Bock 1958:88) that “ falklandicus and bicinctus differentiated from each other in Antarctica, one migrating north to South America and the other to New Zealand”. (2) Gallinago (Scolopacidae). Patagonian stricklandii (boggy areas of Tierra del Fuego and Cape Plorn archipelagoes) and high Andean jamesoni (paramos and wet edge of puna) are borderline cases, treated by some authors as subspecies (Fjeldsa & Krabbe 1990) and by others as species (Hayman et al. 1986). They show morphological dif¬ ferentiation and are separated by about 2000 km of largely unsuitable montane veg¬ etation (too dry). Similarity in aerial displays between South American species (some¬ times placed in genus Chubbia) and New Zealand snipe (genus Coenocorypha ) sug¬ gests “common ancestry for these two groups of southern hemisphere snipes” (Miskelly 1990). (3) Aphrastura (Furnariidae). Allospecies masafuerae (Juan Fernandez Islands) and spinicauda (forested mainland) are isolated by 600 km of water (Vuilleumier 1985). (4) Sicalis (Emberizidae). Patagonian allospecies lebruni (grassy steppes and dirt banks) and high Andean olivascens (scrub and rocky areas) are separated by about 400 km of montane vegetation partly occupied by congener S. auriventris, thus sug¬ gesting interspecific competition as a factor in maintaining isolation. Secondary contacts Parapatry (1) Thinocorus (Thinocoridae). Closely related and differentiated species orbignyianus (Andean) and rumicivorus (steppe) are largely al lopatric but share a 2000 km-long contact zone along the Andean foothills where overlap is narrow (Maclean 1969; map in Fjeldsa & Krabbe 1990). (2) Phrygilus (Emberizidae). Allospecies gayi (steppe) and patagonicus ( Nothofagus forest and edge) appear to share a 1500 km-long contact zone along the Andes at the ecotone between these vegetation types (Vuilleumier unpubl.). 330 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Hybridization (1) Catharacta (Stercorariidae). Limited hybridization between allospecies antarcticus and chilensis has been documented by Devillers (1978) in Santa Cruz. (2) Phrygilus (Emberizid^e). A zone of interspecific hybridisation between allospecies gayi and patagonicus is under study in NW Tierra del Fuego (Vuilleumier unpubl.). Marginal overlaps (1) Podiceps (Podicipedidae). The restricted range of gallardoi (Patagonia) is entirely subsumed in the extensive .range of occipitalis. Storer (1982) described a hybrid between the two species. The southern peripheral overlap between gallardoi and occipitalis is matched, in the high Andean puna, by the one between taczanowskii and occipitalis. Do these patterns represent double invasions? (2) Chloephaga (Anatidae). The small range of southern Patagonian rubidiceps is en¬ compassed within the range of its wider ranging sister species poliocephala. This could be an instance of double invasion. (3) Geositta (Furnariidae). The range of southern Patagonian antarctica is entirely enclosed within that of widespread cunicularia. Habitat co-occupancy has been dem¬ onstrated in NW Tierra del Fuego (Vuilleumier unpubl.). This could be another case of double invasion. (4) Melanodera (Emberizidae). The range of southern Patagonian melanodera (grassy steppe, moorland) is enclosed within the range of widespread xanthogramma (An¬ dean, maritime cliffs in Cape Horn area) (Vuilleumier unpubl.). Again, this could be a double invasion. Broad range overlaps (1) Haematopus (Haematopodidae). Three species overlap extensively along Patagonian coasts (wide ranging palliates, southern South American ater, Patagonian leucopodus). Occasional hybridisation has been demonstrated between ater and palliatus and between ater and leucopodus (Jehl 1978). / (2) Attagis (Thinocoridae). The broad overlap zone in the Patagonian Andes between sister species gayi and malouinus (Hayman et al. 1986, Fjeldsa & Krabbe 1990) re¬ mains to be demonstrated with specimen data. (3) Enicognathus (Psittacidae). There is broad overlap between ferrugineus and leptorhynchus in forests of northern Patagonia (Vuilleumier 1985). DISCUSSION Range discontinuities occur in a number of Non-Passerine, Non-Oscine, and Oscine species living in forest, steppe, alpine scrub, and along the coast of Patagonia. Most intra-Patagonian differentiation is weak, only a couple of instances existing at the superspecies level ( Pteroptochos , Scytalopus). On the other hand, many extra- Patagonian disjunction patterns involve major differentiation. Geographically, extra- Patagonian vicariance patterns include taxa distributed in other parts of the southern hemisphere as well as taxa distributed elsewhere in South America, especially the Andes. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 331 Whereas the numerous instances of disjunction suggest that speciation is currently very active in and around Patagonia, it is often difficult to pinpoint the nature of the barriers between isolates, both within Patagonia and between Patagonia and other regions. An exception may be the Bio-Bio River isolating Pteroptochos castaneus and P. tarnii, or various water gaps in Leptasthenura, Xolmis, and Aphrastura. In most other cases the nature of the gap cannot be specified at present (examples: Cindodes oustaleti, Cistothorus platensis). Some of the large gaps between Patagonian and extra-Patagonian taxa (especially Andean ones) are noteworthy (examples: Pterocnemia, Strix, Gallinago, Sicalis). The pattern shown by these and other taxa suggests a common causal factor for such huge and similar geographical gaps. That vicariance events led to the observed isolation seems clear. What is not clear is the nature of the events. Two possibilities can be mentioned. One is deteriorating ecologi¬ cal conditions in the gap area, especially an increase in aridity. The second is the presence of congeneric taxa in the hiatus zone: they could prevent contact through interspecific competition (examples: Picoides, Sicalis). Few secondary contacts have been studied in detail in Patagonian birds. Exceptions are Catharacta (Devillers 1978) and Phrygilus (Vuilleumier unpubl.). Other instances of narrow range overlaps remain elusive, and little concrete evidence actually docu¬ ments the nature of sympatry (and in Attagis, even its existence). Preliminary study of both marginal and broad range overlaps suggests that, as in some of the high An¬ dean cases studied earlier, “closely related species pairs may have only minor habi¬ tat differences in sympatry” (Vuilleumier 1980: 1260) An example is the ecological co¬ existence of Geositta cunicularia and G. antarctica in NW Tierra del Fuego. In other genera, however, ecological segregation is observed in the zone of sympatry ( Chloephaga , Haematopus, Melanodera). Several appealing models of vicariance involving late Pleistocene glaciation events have been proposed ( Phalacrocorax , Devillers & Terschuren 1978; Tachyeres, Livezey 1986; Catharacta, Devillers 1978). These reconstructions dealt with littoral taxa and could be applied to other ecologically similar groups ( Haematopus ). But surely different models must be sought for wholly terrestrial birds. In the past few years I have investigated several landbird genera in detail ( Phalcoboenus , Attagis, Geositta, Cindodes, Phrygilus). For two of them ( Geositta , Phrygilus) I now have enough data (distributional records, specimens) for attempts at reconstruction of their past histories. These will be presented elsewhere. The repeated pattern involving two sister species, with one wide-ranging taxon and another with a restricted range in southern Patagonia, found in taxonomically and ecologically diverse genera ( Chloephaga , Geositta, Melanodera), suggests double invasion as a common mode of speciation in Patagonia. It is tempting to speculate about the differentiation of small populations in peripherally isolated refuges south of the main icecap at times of glacial maxima, and about a subsequent phase of second¬ ary contact with sympatry, following deglaciation and recolonization by vegetation and consumers (Plumphrey & Pefaur 1979). ACKNOWLEDGEMENTS I thank the National Geographical Society and the Sanford Fund of the American Museum of Natural H istory for financial assistance, and the organizers of the 20th I.O. C. for their invitation to participate in this symposium. 332 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI LITERATURE CITED AMADON, D. 1966 The superspecies concept. Systematic Zoology 15: 245-249. BOCK, W.J. 1958. A generic review of the plovers (Charandriinae, Aves). Bulletin of the Museum of Comparative Zoology 1 18: 27-97. DEVILLERS, P. 1978. Distribution and relationships of South American skuas. Le Gerfaut 68: 374-417. DEVILLERS, P., TERSCHUREN, J.A. 1978. Relationships between the Blue-eyed Shags of South America. Le Gerfaut 68: 53-86. FJELDSA, J., KRABBE, N. 1990. Birds of the high Andes. Copenhagen, Zoological Museum of the University of Copenhagen. HAYMAN, P., MARCHANT, J., PRATER, T. 1986. Shorebirds. Boston, Houghton Mifflin Company. HUECK, K., SEIBERT, P. 1972 Vegetationskarte von Sudamerika. Stuttgart, Gustav Fischer. HUMPHREY, P.S., PEFAUR, J.E. 1979. Glaciation and species richness of birds on austral South American islands. Occasional Papers of the Museum of Natural History of the University of Kansas No. 80: 1-9. JEHL, J.R., Jr. 1978. A new hybrid oystercatcher from South America, Haematopus leucopodus x H. ater. Condor 80: 344-346. LIVEZEY, B.C. 1986. Phylogeny and historical biogeography of steamerducks (Anatidae: Tachyeres ). Systematic Zoology 35: 458-469. MACLEAN, G.L. 1969. A study of seedsnipe in southern South America. The Living Bird 8: 33-80. MAYR, E. 1963. Animal species and evolution. Cambridge, Massachusetts, Belknap Press of Harvard University Press. MISKELLY, C.M. 1990. Aerial displaying and flying ability of Chatham Islands Snipe Coenocorypha pusilla and New Zealand Snipe C. aucklandica. Emu 90: 28-32. NAVAS, J.R. 1970. La identidad de los cormoranes del Lago Nahuel Huapi (Aves, Phalacrocoracidae). Neotropica 16: 140-144. REYNOLDS, P.W. 1934. Apuntes sobre aves de Tierra del Fuego. Hornero 5: 339-353. SHORT, L.L. 1982. Woodpeckers of the world. Delaware Museum of Natural History, Greenville, Dela¬ ware. Monograph Series No. 4. SIEGEL-CAUSEY, D. 1986. The courtship behaviour and mixed-species pairing of King and Imperial Shags ( Phalacrocorax albiventer and P. atriceps). Wilson Bulletin 98: 571-580. STORER, R.W. 1982. A hybrid between the Hooded and Silver Crebes ( Podiceps gallardoi and P. occipitalis). Auk 99: 168-169. TRAYLOR, M.A., Jr. 1988. Geographic variation and evolution in South American Cistothorus platensis (Aves: Troglodytidae). Fieldiana Zoology New Series No. 48: i-iii, 1-35. VUILLEUMIER, F. 1980. Speciation in birds of the high Andes. Acta XVII Congressus Internationalis Ornithologici Berlin (1978): 1256-1261. VILLEUMIER, F. 1985. The forest birds of Patagonia: ecological geography, speciation, endemism, and faunal history. Pp. 255-304 in Buckley, P.A., Foster, M.S., Morton, E.S., Ridely, R.S., Buckley, F.G. (Eds). Neotropical ornithology. Monograph No. 36. Washington, D.C., American Ornitholgists’ Union. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 333 GENETIC DIFFERENTIATION IN YELLOWTHROATS (PARULINAE: GEOTHLYPIS) B. PATRICIA ESCALANTE-PLIEGO Department of Ornithology, American Museum of Natural History, City University of New York, Central Park West at 79th Street, New York, NY 10024, USA and Museo de Zoologia de la Facultad de Ciencias de la Universidad Nacional Autonoma de Mexico ABSTRACT. Allozyme variation is analyzed within and among populations of Neotropical and Nearctic species of Yellowthroats (Parulinae: Geothlypis). Results indicate that the populations assayed have levels of variability comparable with those of other birds. As indicated by FST values, heterogeneity tests, and genetic distances among populations, species in the north (G. trichas) are less differenti¬ ated genetically than Neotropical taxa. The Baja California populations of G. beldingi are somewhat differentiated; more so are the Middle American populations of G. poliocephala. Differentiation among the Neotropical populations of the G. aequinoctialis complex is substantial and coincides with populations having disjunct ranges. It is likely that there is more than one species in this taxon. These results, along with previous findings on Amazonian forest species, indicate that Neotropical avifaunas are more genetically differentiated than are their Nearctic relatives, and suggests that they could be older evolutionarily. Keywords: Geothlypis, parulid warblers, Neotropics, genetic structure, speciation. INTRODUCTION As part of a study on the phylogeny of Yellowthroats (Parulinae: Geothlypis), I used allozyme electrophoresis to survey genetic variation in a series of populations of nine taxa. This technique provides data for phylogenetic analyses among closely related species, and for estimating parameters of population differentiation within species. Studies of the genetic structure of natural populations provides information critical to an understanding of population dynamics, modes of speciation, and phenotypic diver¬ sity (Templeton 1980, Barrowclough 1983, Zink & Remsen 1986). The study of genetic diversity within and among Neotropical species of birds is in its infancy. In the few species examined to date, workers have found that populations of avian species in Amazonian forests are more structured spatially than are populations of temperate species, but have equivalent levels of genetic variability (Braun & Parker 1985, Capparella & Lanyon 1985, Capparella 1987, Gerwin & Zink 1989, Hackett & Rosenberg 1990, Gill & Gerwin 1989). To place these findings in a broader context, it is critical to investigate species associated with Neotropical habitats other than low¬ land rainforests. Because of their wide distribution in open wetlands, warblers of the genus Geothlypis offer an opportunity to compare genetic variability, population struc¬ ture, and habitat restriction in both temperate and tropical America. In this paper, I summarize data obtained from an electrophoresis analysis of members of this complex. My objective is to evaluate the nature of allozymic variation at the population and geographic levels in Neotropical and Nearctic Yellowthroats in the light of our knowledge of other species. 334 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI THE YELLOWTHROAT COMPLEX As currently recognized (American Ornithologists’ Union 1983), Geothlypis is com¬ posed of nine species, most of which are allopatric. Geothlypis taxa inhabit marshes, but most forms have broader habitat tolerances, and also occupy other wetlands where tall grass, brush, second growth and sugarcane grow. In North America, G. trichas (Common Yellowthroat) is widely distributed (Figure 1) with populations ranging from completely migratory in the north, to sedentary in cen¬ tral Mexico. In this latter area they co-occur in marshes of the interior valleys with G. speciosa (Black-polled Yellowthroat). In the southernmost part of its range (South Central Mexico), G. trichas also overlaps with G. poliocephala (Gray-crowned Yellowthroat); they both breed in sugarcane fields that have replaced natural marshes. None of these instances of sympatry seem to involve close relatives, based on an analysis of plumage characters (Escalante-Pliego, in prep.). San Jos* Chapala □ trichas n beldingi flavovelata rostra ta FIGURE 1 - Distributional range of four Geothlypis species and sample localities mentioned in the text. US: Rhode Island (not in map); Florida, Osceola. Bahamas, Abaco. Mexico: Hidalgo, Zupitlan ; Morelos, Yautepec, Michoacan, Lago de Cuitzeo ; Jalisco, Lago de Chapala East; Baja California Norte, San Telmo ; Baja California Sur, San Ignacio ; Baja California Sur, San Jose; Tamaulipas, Altamira. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 335 Besides G. speciosa, forms with restricted ranges also associated with marshes are G. beldingi (Belding’s Yellowthroat) in the Peninsula of Baja California, and G. flavovelata (Altamira Yellowthroat) in northeastern Mexico. Although sharing an affin¬ ity for wet habitats, other Yellowthroats have broader habitat tolerances. G. nelsoni (Hooded Yellowthroat) is found in Mexico along the humid slopes of the eastern mountains. G. rostrata (Bahama Yellowthroat) occurs in bush and abandoned sugarcane fields in some of the Bahama Islands. In the lowlands of Mexico, Central America, and western South America south to Ecuador, G. poliocephala and G. semiflava (Olive-crowned Yellowthroat) replace each other from north to south near streams in marshy and humid brushy areas of second growth lowland forests. In Nica¬ ragua, Costa Rica, and western Panama, where their ranges are in contact, G. poliocephala is found in drier areas (Figure 2). The South American representative of the group, G. aequinoctlalis (Masked Yellowthroat) is widely distributed, and occurs in several kinds of wetlands, including marshes. The geographic range of the aequinoctialis complex is made up of four dis¬ junct areas (Figure 3): the peripheral form of western Costa Rica and western Panama ( chiriquensis ), sometimes regarded as a separate species (AOU, 1983); the trans- Andean or Pacific slope form (auricularis group); and two cis-Andean forms, one north (nominate aequinoctialis group), and one south of the Amazon Basin ( velata group). Cartago u speciosa nelsoni m semiflava e poliocephala CuKzao Yautapac FIGURE 2 - Distributional range of four Geothlypis species and sample localities mentioned in the text. Mexico: Michoacan, Lago de Cuitzeo ; Morelos, Yautepec; Puebla, Huauchinango. Costa Rica, Cartago ; El Limon, Puerto Viejo. 336 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI METHODS Study areas and samples Samples were collected on breeding grounds in Mexico, Costa Rica, Venezuela, Florida (USA), and the Bahamas, between 1987 and 1989 (Figures 1, 2, and 3). Ex¬ act localities are available from the author. The habitat visited in Chapala, Cuitzeo, and Zupitlan is marshland. The marshes of the interior basins of Mexico have been much reduced in the last 500 years through alluviation, desiccation and artificial drain¬ ing (Tamayo and West, 1964). Aside from marshes, G. trichas was found in sugarcane fields around Cuitzeo, and Yautepec. In the arid peninsula of Baja Califor¬ nia, marshes are widely scattered. In northern Baja California, G. trichas was found along the streams that originate in the western slope of the Sierra de Juarez (San Telmo). In the central desert of the peninsula G. beldingi was found in the springs of San Ignacio, and at the southern tip of the Peninsula (San Jose). The extensive marsh at Altamira (Laguna Champayan) receives a large discharge from the Panuco River. Lambayeque FIGURE 3 - Branching diagram from distance Wagner procedure using Rogers’ distances for the four samples of G. aequinoctialis superimposed in distributional ranges (Ridgely & Tudor 1989). The lines are not drawn to scale. Localities: Costa Rica, Puntarenas, San Vito. Venezuela, Miranda, Caucagua. Peru, Lambayeque, Las Pampas. Bolivia, Santa Cruz, Velasco. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 337 The lake and inundated area around it extends several hundred square kilometers. At San Vito (Costa Rica) and Caucagua (Venezuela), the habitats where Geothlypis was found, are flooded seasonally but are not true marshes; grassy meadows in San Vito, brush and sugarcane fields in Caucagua. The remainder of the localities have a range of open and humid lands or scrub along streams. Following collection, liver, heart, and breast muscle were preserved in liquid nitrogen until transported to the laboratory, where they were stored at -70°C. Additional samples from the USA (Rhode Island), Peru, and Bolivia were obtained from other frozen tissue collections (see Acknowledgements). Electrophoresis Allozyme variation was assayed for 271 Yellowthroat tissue samples for 32 enzymatic loci following standard techniques (Selander et al. 1971, Harris & Hopkinson 1976, Richardson et al. 1986). The 32 loci scored are: ACON-1, ACON-2, ADA, ADH, DIA, EAP, EST-1 , ESTD, FUM, GDA, GDH, GOT-1, GOT-2, GPDH, G-3-PDH, G-6-PDH, IDH-1 , IDH-2, LAP, LDH-1 , LDH-2, MDH-1 , MDH-2, ME, NP, PEP-A, PEP-B, PEP-C, PGM-1 , PGM-2, PGM-3, and SDH. Parameters of genetic variability and differentiation were estimated using several computer programs: BIOSYS-1 (Swofford and Selander 1989) for allelic frequencies, heterozygosities, percent of polymorphic loci, chi square tests of heterogeneity, HardyWeinberg equilibrium tests, Nei’s (1978) and Rogers’ (1972) genetic distances; NEI2 for heterozygosities and genetic distances, and BOOTFST which provides Wright (1978) FST values with their confidence intervals calculated through a bootstrap approach. RESULTS Genetic Variability. Of the 32 loci scored, 10 were monomorphic for all populations, 6 were private polymorphisms unique to individual populations, and the other 16 showed shared polymorphisms among populations and/or species. For samples with over five indi¬ viduals, none of the variable loci showed significant (P<.001) departures from Hardy¬ Weinberg equilibrium. Detailed allelic frequencies will be reported elsewhere (Escalante-Pliego, in prep.). Table 1 shows the average heterozygosity per locus, and the percent of polymorphic loci for each population. Measures of interpopulational differentiation were calculated for species with more than one representative sample. Population Differentiation in Yellowthroat species. The samples of the G. trichas used in this study comprise a considerable part of the total geographic range. Tests of heterogeneity were not significant for 15 of 19 vari¬ able loci. Two loci at which electrophoresis detected substantial geographic differen¬ tiation were PGM-1 and PEP-B. At the PGM-1 locus (P<.001), the western samples, including San Telmo and three localities of west-central Mexico, shared a polymorphism which was absent in the eastern samples (Rhode Island, Florida, and Zupitlan). At the PEP-B locus the heterogeneity test among populations was signifi¬ cant (P<. 009); all the samples have the same common allele in frequencies ranging 338 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI from 0.5 to 1 .0; the frequencies of the less common allele showed no apparent geo¬ graphic trend. At the G-6-PDH locus, allelic frequencies were significantly different (P<.045) because of the absence of an uncommon allele in two populations (Zupitlan and Rhode Island). At SDH a significant difference (P<.015) was due to the presence of an allele in the northeasternmost population assayed (Rhode Island). This allele is shared with Bahamian’G. rostrata. The mean FST values in the trichas complex was 0.077 (95% confidence interval: 0.01 2-0.1 1 2). Averaged Nei’s genetic distance for all G. trichas populations was 0.004. TABLE 1 - Genetic variation in populations of Geothlypis species calculated over 32 loci. Habitat N Het SE P.L. Geothlypis trichas Rhode Island (USA) s 13 .0411 (.0160) 22 Osceola (Florida, USA) s 13 .0449 (.0146) 31 Zupitlan (Mexico) m 15 .0298 (.0206) 13 Yautepec (Mexico) c 15 .0322 (.0156) 19 Cuitzeo (Mexico) m,c 14 .0585 (.0213) 28 Chapala (Mexico) m 12 .0531 (.0242) 19 San Telmo (Mexico) s 15 .0509 (.0226) 19 Geothlypis beldingi San Ignacio (Mexico) m 18 .0384 (.0194) 9 San Jose (Mexico) m 36 .0442 (.0248) 13 Geothlypis flavovelata Altamira (Mexico) m 7 .0633 (.0307) 16 Geothlypis nelsoni Huauchinango (Mexico) s 14 .0500 (.0233) 19 Geothlypis rostrata Abaco (Bahamas) s 17 .0481 (.0211) 19 Geothlypis sped os a Cuitzeo (Mexico) m 23 .0106 (.0080) 9 Geothlypis semiflava Puerto Viejo (Costa Rica) s 19 .0343 (.0155) 28 Geothlypis aequinoctialis San Vito (Costa Rica) m 9 .0577 (.0285) 13 Santa Cruz (Bolivia) m? 2 .0469 (.0345) 6 Lambayeque (Peru) m? 3 .0521 (.0309) 9 Caucagua (Venezuela) s 8 .0656 (.0242) 25 Geothlypis poliocephala Cartago (Costa Rica) s 5 .0250 (.0196) 6 Yautepec (Mexico) c 13 .0478 (.0212) 22 Habitat: s=shrubby and along streams; m = marsh; c = cane fields. N = sample size. Het = Average heterozygosity per locus. P.L. = per cent of polymorphic loci (99% of frequency criterion). Only the GDA locus showed a significant difference (P<.005) between the two sam¬ ples of G. beldingi on the Peninsula of Baja California. The FST value was 0.059 (95% confidence interval 0.001-0.171); Nei’s (1978) genetic distance was 0.011. The two single samples of G. poliocephala came from widely separated areas of their distribution that represent some of the well differentiated subspecies. Tests of hetero¬ geneity were significant for two loci: PEP-C (Pc.001), and PGM-2 (P<.07); the FST ACTA XX CONGRESSUS I NTER NATION ALIS ORNITHOLOGICI 339 value was 0.199 (95% confidence interval 0.027-0.281), and Nei’s (1978) genetic dis¬ tance was 0.01 1 . In G. aequinoctialis of South America and Costa Rica, fixed differences were found between various populations at the GDA and ME loci. Differences were also large at other variable loci, giving significant tests of heterogeneity at 8 of 12 loci (P<.005 for five loci, P<.02 in three loci). This consistent differentiation across loci is reflected in a high FST value of 0.553 (95% confidence interval: 0.312-0.751). Averaged Nei (1978) genetic distances among the four populations were 0.1245, but the sample from Ven¬ ezuela had a higher average value (0.1445). The use of Rogers’ distances provided similar results. A clustering of Rogers’ distances superimposed in the distribution of the G. aequinoctialis complex is shown in Figure 3. DISCUSSION Levels of intrapopulation variability (Table 1) in Geothlypis species are within the range of those previously reported for birds (Braun & Parker 1985, Capparella 1987, Nevo et al. 1984). Due to differences in sample size, and because heterozygosities have large standard errors, a statistical test cannot be performed among the populations associated with different habitats. Flowever, it is noticeable that G. speciosa had low heterozygosity and few polymorphic loci. Some populations that are restricted to marshes, such as G. beldingi and G. trichas of Zupitlan, also have low numbers of polymorphic loci, but similar heterozygosity scores. A dissimilar pattern in other marsh inhabitants is observed in Table 1. The sample of G. trichas from Zupitlan shows some signs of isolation: Zupitlan’s av¬ erage genetic distance of 0.0052 is higher than that of the combined average of all the populations in the trichas complex (0.0036). These values are due to significant differences in frequencies at the PGM-1 and PEP-B loci, as mentioned earlier. Zink and Klicka (1990) examined allozyme variation in the trichas complex using samples from presumed sedentary (Texas), and migratory (Minnesota) populations, and found an Fst value of 0.04. An equivalent comparison using San Telmo, Rhode Island and Florida samples also yields an FST value of also 0.04. Considering the sedentary populations in the south only, the FST value increases to 0.08, but if we ignore Zupitlan, the three remaining populations yield a value of 0.034, similar to the north¬ ern part of the trichas complex. Because they colonize and breed in sugarcane fields, populations in the rest of the Mexican Plateau perhaps have maintained a larger amount of gene flow in contrast to those of the more arid part of the Plateau where Zupitlan is located. For other forms, such as G. beldingi of Baja California, the scattered distribution of marshes seems to have affected population structure and restricted gene flow. The slight phenotypic differentiation between the two populations is concordant with the slight genetic differentiation in these two populations. The estimates of FST and Nei’s (1978) genetic distance for the Middle American G. poliocephala were approximately twice those in the trichas complex and in beldingi. This genetic divergence is paralleled by phenotypic distinctiveness. 340 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI The results for the aequinoctialis complex were more surprising as they suggest more advanced differentiation than the current classification reflects. Very high FST values, significant tests of heterogeneity, and large genetic distances show a pattern of diver¬ gence between the northern Amazonian form aequinoctialis and the other three forms. This pattern contrasts with the phenotypic differentiation observed. Plumage patterns are conservative in the complex, except for a reduction of the mask patch in the Pa¬ cific slope form ( auricu laris ). In this first study of a non-forest group, the Neotropical species of Geothypis showed increasing levels of population differentiation from north to south. FST values of Nearctic populations of the trichas complex compare well with other values found in warblers of North America, such as 0.03 in well marked subspecies of Dendroica coronata (Barrowclough 1980). FST values increased in the Middle and South Ameri¬ can Geothlypis species. The genetic distance values obtained here correlate well with values obtained for Parulids (Barrowclough & Corbin 1978), with an average Nei’s (1978) genetic distance of 0.100 for species level differentiation. In this context, the four allopatric forms of the aequinoctialis complex would seem to deserve species rank. ACKNOWLEDGEMENTS Financial support was provided by the Frank M. Chapman Memorial Fund (American Museum of Natural History), the Alexander Wetmore Fund (American Ornithologists’ Union), the Graduate Studies Committee of the Biology Program (City University of New York), and Sigma Xi. I thank officers and colleagues who helped me to obtain collecting permits: Secretaria de Desarrollo Urbano y Ecologia (Mexico), Ministerio de Recursos Naturales, Energia y Minas (Costa Rica), Department of Agriculture (Baha¬ mas), State of Florida Game and Fresh Water Fish Commission, and the U. S. Fish and Wildlife Service. Many people shared in the field work with me and made much of my investigation possible. I am especially grateful to Robert M. Zink and Jay Pitocchelli for providing me with partial tissues for assays. I thank George F. Barrowclough, Francois Vuilleumier, Town Peterson, Charles Myers, and Angelo Capparella for advice and help. I am indebted to my friends and colleagues at the Museo de Zoologia de la Facultad de Ciencias de la UNAM (Mexico) for uninterrupted moral support. LITERATURE CITED AMERICAN ORNITHOLOGISTS’ UNION. 1983. Check-List of birds of North America. American Orni¬ thologists’ Union. 6th Edition. BARROWCLOUGH, G. F. 1980. Genetic and phenotypic differentiation in a Wood Warbler (genus Dendroica) hybrid zone. Auk 97:655-669. BARROWCLOUGH, G. F. Biochemical studies of microevolutionary processes. Pp. 223-261, in Per¬ spectives in Ornithology, A. H. Brush, G. A. Clark, (Eds). New York, Cambridge University Press. BARROWCLOUGH, G. F., CORBIN, K.W. 1978. Genetic variation and differentiation in the Parulidae. Auk 95:691-702. BRAUN, M. J., PARKER, T.A. III. 1985. Molecular, morphological, and behavioral evidence concern¬ ing the taxonomic relationships of "Synallaxis" gularis and other synallaxines. Pp. 333-346 in Neotropical Ornithology, P. A. Buckley, M. S. Foster, E. S. Morton, R. S. Ridgely, F. G. Buckley, (Eds). Ornithological Monographs No. 36. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 341 CAPPARELLA, A. 1987. Effects of riverine barriers on Genetic Differentiation of Amazonian Forest Un¬ dergrowth Birds. PhD. thesis. Louisiana State University and Agricultural and Mechanical College. CAPPARELLA, A. P., LANYON, S.M. 1985. Biochemical and morphometric analysis of the sympatric, Neotropical, sibling species Mionectes macconnelli and M. oleagineus. Pp. 347-355 in Neotropical Ornithology, P. A. Buckley, M. S. Foster, E. S. Morton, R. S. Ridgely, F. G. Buckley, (Eds). Ornitho¬ logical Monographs No. 36. GERWIN, J. A., ZINK, R.M. 1989. Phylogenetic patterns in the genus Heliodoxa (Aves: Trochilidae): An allozymic perspective. Wilson Bulletin 101: 525-544. GILL, F. B., GERWIN, J.A. 1989. Protein relationships among Hermit Hummingbirds. Proceedings of the Academy of Natural Sciences of Philadelphia 141:409-421. HACKETT, S. J., ROSENBERG, K.V. 1990. Comparison of phenotypic and genetic differentiation in South American Antwrens (Formicaridae). Auk 107: 473-489. HARRIS, H., HOPKINSON, D.A. 1976. Handbook of enzyme electrophoresis in human genetics. Ox¬ ford: North Holland Publications Company. NEI, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of in¬ dividuals. Genetics 89: 583-590. NEVO, E., BEILES, A., BEN-SHLOMO, R. 1984. The evolutionary significance of genetic diversity: eco¬ logical, demographic and life history correlates. Pp. 13-213 in Evolutionary Dynamics of Genetic Di¬ versity. G. S. Mani, (Ed.). New York, NY, USA. RICHARDSON, B. J., BAVERSTOCK, P.R., ADAMS, M. 1986. Allozyme Electrophoresis. Sydney, Aus¬ tralia. Academic Press. RIDGELY, R. S., TUDOR, G. 1989. The Birds of South America. Vol 1. University of Texas Press, Austin, Texas, USA. ROGERS, J. S. 1972. Measures of genetic similarity and genetic distance. Studies in Genetics. Univ. Texas Publ. 7213: 145-153. SELANDER, R. K., SMITH, M.H., YANG, S.Y., JOHNSON, W.E., GENTRY, J.B. 1971. Biochemical polymorphism and systematics in the genus Peromyscus. I. Variation in the old-field mouse ( Peromyscus polionotus). Studies in Genetics VI. University of Texas Publication 7103: 49-90. SWOFFORD, D. L„ SELANDER, R.B. 1989. BIOSYS-1. A Computer Program for the Analysis of Allelic Variation in Population Genetics and Biochemical Systematics. Champaign, Illinois, USA. TAMAYO, J. L., WEST, R.C. 1964. The hydrography of Central America. In, Handbook of Middle Ameri¬ can Indians. Vol. 1. Natural Environment and Early Cultures. Univ. Texas Press. Austin, Texas, USA. TEMPLETON, A. 1980. Modes of speciation and inferences based on genetic distances. Evolution 34: 719-729. WRIGHT, S. 1978. Evolution and the Genetics of Populations. Volume 4. Variability Within and Among Natural Populations. University of Chicago Press, Chicago, Illinois, USA. ZINK, R. M., KLICKA, J.T. 1990. Genetic variation in the Common Yellowthroat and some allies. Wilson Bull. 102:514-520. ZINK, R. M., VAN REMSEN, J. 1986. Evolutionary processes and pattern of geographic variation in birds. Current Ornithology vol. 4 R. F. Johnston, (Ed.). Plenum Press, NY, pp. 1-69. 342 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI BIOGEOGRAPHIC PATTERNS IN BIRDS OF HIGH ANDEAN RELICT WOODLANDS J. FJELDSA Zoological Museum, Universitetsparken 15, DK-2100 Copenhagen, Denmark ABSTRACT. Woodlands, especially Polylepis, form patches in steppe-like Andean habitats at 3500- 4500 m. High-elevation woodland is marginal habitat for a number of lower-elevation birds, but there is no direct evidence that colonization at the edge of the ecological range leads to specialization to the new habitat. This happens only after an initial period of entrapment, located away from the founding area, usually on the upper Pacific slope of the Peruvian Andes. Many specialists had east-slope hu¬ mid forest origins. Other source areas were the eastern premontane woodlands of Bolivia and Argen¬ tina, as “entrapment” in the Cochabamba basin was followed by dispersal across Peru. Several lev¬ els of differentiation of populations suggest cycles of dispersal and vicariance, but only in a few gen¬ era were there speciation events after the initial adaptation to Polylepis. Keywords: Biogeography, Andes, vicariance, refuge theory, Polylepis woodlands. INTRODUCTION Primary woodlands, especially with Polylepis , or with Gynoxys, Buddleia and various scrubs, form small patches well above the Andean treeline. This habitat may always have been rather localized, but thousands of years of human activity have reduced it to vestiges (Figure 1; Ansion 1986). Most bird species specialized for this habitat thus have relict distributions, and some are extremely rare (Collar et al. MS). This paper discusses biogeographic patterns of these birds. MATERIALS / Fligh-elevation woodlands in Peru and Bolivia were charted since 1987 (Fjeldsa 1987, in press, Fjeldsa & Krabbe 1990: 846-7, Frimer & Moller 1989), and this was supple¬ mented with literature surveys and data from museum specimens. Current views on the species’ distributions and systematic relationships are in Fjeldsa & Krabbe (1990). Relationships, needed to correlate distributions of sister taxa, were evaluated from assumed shared derived character states. In a few cases phylogenic hypotheses were developed for larger species groups. THE BIRDS OF POLYLEPIS WOODS Table 1 lists over 100 species living in Polylepis woodland. This includes species in adjacent habitats that invade Polylepis woodlands locally. Additional hummingbirds, spinetails and tanagers of the humid montane forest visit Polylepis seasonally, espe¬ cially in the northern Andes. Furthermore, a number of birds of prey and some paramo and puna birds use the more open, bushy parts of the woodlands, or roost there. There is continuous variation from these marginal species to those having their ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 343 adaptive peaks in high-elevation woodlands (in the following marked with *, or ** for the most specialized). Most specialists are attached to an elevational zone rather than to specific trees, but a few are adapted for searching insects in the multi-layer struc¬ ture of Polylepis bark ( Oreomanes fraseri**, Leptasthenura xenothorax**; two Cranioleuca species*), searching its small-leaved canopy ( Anairetes alpinus**), eat¬ ing Polylepis seeds ( Carduelis crassirostris**) or eating sugary secretions and aphids from associated Gynoxys bushes ( Chalcostigma stanleyi*, Xenodacnis parina*, Poospiza alticola**). Most specialists are suited for biogeographic studies because of their sedentary habits. They seem able to remain in their tiny “habitat islands” even when snowstorms sweep the highlands (Fjeldsa in press b), in contrast to the wide¬ spread birds of open highland habitat. FIGURE 1- High-elevation woodlands in Peru and Bolivia. Black areas have patches of Polylepis, mainly at 3500-4500 m elevation. Outside these areas are found few Polylepis patches more than a few hectares in area. The shaded zone on the eastern Andean scarps shows treeline at c. 3500 m, sometimes with Polylepis admixed. Based on field studies, maps, LANDSAT imagery, and ground surveys. Faunal provinces Figure 2 gives examples of patchy, disjunct (relict), and endemic distributions of Polylepis birds. Such maps were used to produce summary maps of densities of spe¬ cies richness (Figure 3). The highest species richness is found along the edges of the highlands and in cordilleras intersected by deep valleys. In Peru and Bolivia, many specialists also inhabit altiplanos far from lower-elevation habitat (Figure 3), in con¬ trast to Polylepis woodlands north of the North Peru Low (Ecuador), which serve as 344 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI TABLE 1 - Species living in Polylepis wood at least in part of their range. Legend: High elevation H, * and ** being specialists of woods in this zone. Species with their main distribution on lower elevation or in open habitat are marked ++ if also well established in Polylepis, and + if so locally. Habitats: 0= open land, U= ubiqui¬ tous, S= scrub, Pa= paramo, W= wood, HF= humid forest. Brackets mark assumed sister taxa. Species Nothoprocta taczan. N. omata Columba maculosa Bolborhynchos aymara B. aurifrons B. orbygnesius Caprimulg. longirost. Colibri coruscans Oreotroch. estella O. adela Patagona gigas Aglaeac.cupri. A. castelnaud. Lesbia victoriae L. nuna Sappho sparganura Metallura phoebe Metallura baroni * Chalcostigma stanleyi Ch. olivaceum Ch. heteropogon Ch. herrani Oreonympha nobilis Veniliomis nigriceps V. fumigatus Colaptes atricollis C. rupicola Upucerthia serrana U. andaecola U. ruficauda Cincl. (exc.) aricomae Leptasthenura andic. L. striata L. pileata L. xenothorax L. (aegi.) berlepschi L. fuliginiceps L. yanacensis Cranioleuca baroni C. albicapilla Asth.(dorbig.) semisp Asthenes pudibunda A. heterura A. ottonis A. virgata A. urubambense Phacell. striaticeps Grallaria andicola Scytalopus magellan. Acroptemis orthonyx Ampelion rubrocrist. A. stresemanni Mecocerc. leucophrys Habitat Range _ +W Apurimac +OH Peru-NW Arg +W Peru-Bol. + +S Bol-WArg. +OH Peru-NW Arg + + W Peru-Bol. ++U Andean +W Ven-NW Arg. + +OH Peru-Bol. + +S Cochabamba + +S Ecu-W Arg. + +W-HF Col-Peru * or ++W Apurimac +S-W Col-Peru + +S Col-N Bol. + +S-W Bol-WArg. + +S-W W Peru or ++W Azuay, Ecu. *-Pa Ecu-N Bol. +OH C.Blanca + Real / + +Pa Ven-Col. + +Pa Col-N Peru +S-W Apurimac +HF Col-N Bol. +HF Andean HF +W W Peru-Maranon ++OH Peru- AW Arg. * NW Peru Bol-NW Arg. S Peru-Arg. Abancay-C.Real Ven-NW Bol. S W Peru N and W Peru Abancay-Cuzco W Bol. Bol-NW Arg. C.Blanca, Aban N-NW Peru Apurimac SW Peru W Peru Cochabamba Apurimac C Peru E ridge Peru-Bol C Peru/NW Arg. Peru -t- + PaW in Peru part *-HF Ven-N Peru + W-HF Ven-Bol. *(W) C.Blanca-Lima + HF Andean HF + +S +s-o ** H*OPa *-S *-W ** +OH + +S **(Bol:*S) *-W *-w *(S.O) + +S-W + +S + +S-W + Pa *-Pa *-SW •(Pa) } } Anairetes alpinus A. nigrocristatus A. reguloides A. parulus Ochthoeca rufipect. O. fumicolor O. oenanthoides O. leucophrys Cnemarcus erythro. Polioxolmis rufipenn. Agriomis montana A. andicola Notiochelidon murina N. flavipes Troglodytes aedon Turdus fuscator T. chiguanco Myioborus brunniceps Conirostrum cinereum C. tamarugense C. ferrugineiventre C. sitticolor Oreomanes fraseri Diglossa brunneiventr. D. carbonaria Xenodacnis parina Thraupis bonariensis Thtypopsis ruficeps Saltator aurantii. S. rufiventris Catamenia inomata Phrygilus atriceps P. punensis P. fruticeti P. unicolor P. plebejus Atlapetes rufinucha A. fulviceps A. schistaceus A. rufigenis A. nationi Zonotrichia capensis Poospiza boliviano P. alticola P. hypochondria P. rubecula P. caesar P. baeri P. garleppi Carduelis crassirostris C. magellanica C. atrata C. uropygialis + +S +S + +S +W-HF +Pa-HF •(OH) +S *-HF •(O) +OH *-OH *-Pa +Pa-HF ++U +W-HF + +OH +HF-W ++U *(S) ++HF +Pa-HF ** S + +S + +S *(HF) + +S +S-HF +W-S *-HF +S *-OH +OH + +S *-OH +OH-S +HF + +HF + Pa-HF *-W + +W-S +u +s } } } ** C.Blanca C.Real C.Blanca SW Peru S Col-S Chile Col-Bol. Ven-Bol. Peru-Bol. SW Ecu-N W Arg. patchy Ven-Bol. Peru-Bol. S Col-S Arg. 1 rare Ecu-NW Arg .J Ven-N Bol. V Ven-N Bol. / Americas Ven-N Bol. Ecu-W Arg. Bol-NW Arg. S Col- N Bol. S Peru/N Chile/ C Peru-N Bol. Ven-N Bol. Col-Bol. (Col.)Peru-N Bol-1 Cochabamba (Ecu.) Peru Ecu-Arg. C Peru-NW Arg. Peru-Arg. Cochabamba Ven-W Arg. SW Peru-NW Arg.\ Peru-NW Bol. Peru-S Arg. Ven-S Arg. Ecu-W Arg. Col-N Bol. Cochabamba Ven-Ecu., C Peru C.Blanca + Apurimac^ SE Ecu-W Peru / Ven-S Arg. Cochabamba '} s} 1 ** C.Blanca +S Bol-W Arg. } + + W-S W' Peru + + W-S Apurimac *(S) Tucuman *(S) Cochabamba ** Peru-NW Arg. +S-W Col-Arg. *-OH Peru-NW Arg. -t-O-S W Peru-Chile-Arg. } } ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 345 FIGURE 2 - Three examples of distributions (adapted from Fjeldsa & Krabbe 1990). Wide¬ spread but patchy species (A-C), disjunct species (D-F), and endemic species with re¬ stricted distribution (G-l). marginal habitat for species of lower-elevation humid forest or paramo habitat. Oreomanes fraseri**, the only genuine Polylepis bird in this area, may have come from Peru (Figure 2b). Since the patches of high-elevation woodland in the northern Andes, always in proximity of lower-elevation forest, apparently lack specialists, this area is excluded from further analysis. 346 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Some Polylepis specialists are almost as widespread as the fragmented habitat itself ( Polioxolmis rufipennis*, Ochthoeca oenanthoides*, Oreomanes fraseri**, Carduelis crassirostris**; compare Figures 2 and 3), which obscures their biogeographic origin. Oreomanes may be perfectly adapted for surviving in small populations far away from lower-elevation refuge habitat. Other species maintain a wide range by wandering in the non-breeding seaso'n. Other species and megasubspecies have more restricted ranges, some being endemic to a small area (Figure 2). Subcenters in the West Cordillera. Many species occur along almost the entire West Peruvian Andean Subcenter (Cracraft 1985), but others are more local (see density peaks in Figure 3a). The highest concentration of endemic species is in semihumid canyons from the Cordillera Blanca draining into the headwaters of the Rio Maranon. This subcenter is squeezed between a center of endemism on the upper Pacific slope of Lima, and one in the upper Maranon valley, with endemics both in the arid subtropi¬ cal part (Cracraft 1985) and in temperate scrub in side valleys. The C. Blanca and Lima subcenters are connected by mountain chains which have Polypelis patches scattered almost all the way to Oyon in northern Lima. Upucerthia serrana*, Ampelion stresemanni*, Xenodacnis parina petersi** and bella* and Poospiza rubecula* span both subcenters. However, megasubspecies pallidior* of Mecocerculus leucophrvs, Anairetes nigrocristatus, Atlapetes r. rufigenis* and Poospiza alticola** are centered in C. Blanca; Leptasthenura striata*, Anairetes reguloides and Atlapetes nationi/ seebohmi are centered on the Pacific Slope. Strong differentiation of C. Blanca 4-6 high elevation specialist y.g SpGC'lGS 10-14 FIGURE 3 - Density of species living in Polylepis woodlands (all Table 1 species, A), and of high-elevation specialists (starred in Table 1, B). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 347 centered and Lima-centered subspecies of Leptasthenura pileata* and Cranioleuca baroni*, and weak differentiation in some other species, suggest periods of past iso¬ lation. Some racial shifts also exist further south, and to this can be added an un¬ named semispecies* of the Asthenes dorbignyi group (see Fjeldsa & Krabbe 1990:365-70) and Conirostrum tamarugense. The Apurimac center. Many species are centered in the Andahuaylas-Abancay area in Apurimac, but three of these range as far south as Puno, in tiny rainshadow areas within the Cordillera Real. High-elevation endemics are Nothoprocta taczanowskii, Aglaeactis castelnaudii*, Oreonympha nobilis, Asthenes ottonis and virgata, Cranioleuca albicapilla*, an unnamed Scytalopus, Xenodacnis p.parina*, Atlapetes rufigenis forbesi* and Poospiza caesar. Four of these show slight to well-marked subspecific differentiation within this range. The species inhabit patches of semihumid woodlands on the edge of the puna in otherwise arid valleys, but A. virgata is typical of paramo /Polylepis ecotones across central Peru. The Apurimac center was not recognized by Cracraft (1985), who pooled most of its species with the humid-forest birds of an East Peruvian Andes Subcenter, but two with humid-forest forms in the South Peruvian Subcenter. Cinclodes (excelsior)aricomae** and Leptasthenura xenothorax ** of his South Peruvian Subcenter have tiny populations in C. Real and southeast of Abancay (Fjeldsa & Krabbe 1990:846-7). Cochabamba center. The intermontane basin of Cochabamba is the center for Oreotrochilus adela, Asthenes heterura, Diglossa carbonaria, Saltator rufiventris*, Atlapetes fulviceps, Poospiza boliviana and garleppi* (other endemics inhabiting open habitat or lower-elevation forest). Some of these extend south to the Andean scarp of northern Argentina. Surprisingly Cracraft (1985) did not treat Cochabamba as an area of endemism, but pooled its species with the Austral Andean Center, and one species in the South Peruvian Andean Subcenter. The situation is in fact complex. Besides Cochabamba endemics, other taxa are differentiated in Bolivian and Argentine semispecies. Sappho sparganura, Upucerthia andaecola and Leptasthenura fulviceps span both centres. There is probably an independent center for birds of prepuna and arid puna on ad¬ jacent altiplanos. Relationships between faunas Lessons from tracks connecting well differentiated taxa. Of the seven widespread Polylepis specialists (Table 1), five have their closest relative in the humid montane forest. Most of these relatives are widespread along the east Andean slope, but Cnemarchus erythropygius* {re lated to Polioxolmis rufipennis*) is rare and local, and Conirostrum ferrugineiventre (possibly related to Oreomanes fraseri**) is disjunct (east slope of central Peru and from Cuzco to Cochabamba). Carduelis magellanicus (“an¬ cestor” of C. crassirostris**), is widespread, but is absent from the arid puna of north¬ ern Chile, where C. crassirostris** may have originated in isolation in the Polylepis zone above the Atacama desert. The last species of this widespread group {Leptasthenura andicola *) belongs in a group of southern origin. More narrowly distributed endemic species show various faunal connections. Some tracks connect the West Andean Subcenter or parts of this subcenter with Apurimac, or the Bolivian slopes, and connect this latter area with adjacent lowlands. Among 348 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Oreotrochilus hummingbirds, the Cochabamba form adela seems more primitive than its higher-elevation congeners. A close C. Blanca-Apurimac relationship is shown for the megasubspecies of Atlapetes rufigenis*, and a close Apunmac-Cochabamba re¬ lationship for Asthenes ottonis/heterura. Some strange long-distdnce relationships exist also: Oreonympha nobilis (Apurimac) apparently is the sister taxon of Oxypogon guerinii (Colombia); Cinclodes (excelsior) aricomae** (Abancay-C. Real) is the sister taxon of C. excels ior spp. of Ecuador-Co- lombia. Most specialists live characteristically well isolated from the ranges of nearest lower-elevation taxa, or overlap only marginally. No Polylepis specialist for which the East Cordillera seems to be optimal has a close relative in the adjacent zone of hu¬ mid forest. Asthenes urubambensis*, restricted to the east Andean treeline ecotone has its nearest relative ( Asthenes flammulata group*) patchily distributed on higher elevations. Thus A. urubambensis* has adapted to this ecotone habitat from above rather than from below. Interpretations based on specific phylogenetic hypotheses. The fact that few Polylepis specialists form superspecies or species groups (Table 1) suggests that much of this fauna arose by independent events with little subsequent opportunity for speciation. A few exceptions exist, though. FIGURE 4 - Density of endemic species with restricted distributions, and of megasubspecies found in Polylepis woodlands (A) and of Polylepis species having disjunct populations (B). ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 349 Leptasthenura spinetails (Figure 5a) apparently originated in Argentina or southern Brazil, two groups invading the highlands. Possibly L. yanacensis arose early by iso¬ lation in Cochabamba, and then spread north, its Peruvian populations** becoming narrowly attached to Polylepis. The other group may have ascended via the southern “monte” and “pre-puna” habitats and speciated in Peru after having first adaptated to Andean shrubsteppe and Polylepis. The three top branches^-**) of the phylogeny can clearly be interpreted in terms of isolation on the Pacific slope of Peru, subsequent isolation in the north, and finally colonization to mountains of Apurimac/Cuzco. Atlapetes brushfinches are typical of humid Andean forest. One branch (Figure 5b) possibly crossed the North Peru Low and became isolated in pockets of semihumid habitat on the Pacific slope, one clade spreading south in western Peru, in part in Polylepis. One population then became isolated on the Maranon slope of C. Blanca (A. r. rufigenis*; with secondary contact and casual hybridization with A. (nationi) seebohmi in Cajamarca) and subsequent spread to Apurimac (A. r. forbesi*). The warblingfinches, Poospiza, are confusing, and my phylogeny (Figure 4c) is ten¬ tative and unrooted. This group clearly radiated in the lowlands of Argentina and Bo¬ livia, but two clades adapted to highland conditions occur as far as C. Blanca. A Leptasthenura striata W “ yanacensis*** C. Blanca 1 Abancay- Bol. fuliginiceps Bol.-NW striolata SE Brazil pileata** nc/W Peru xenothorax*** Abancay-Cuzco andicola** Ven.-N Bolivia aegithaloides* Patagonia-WB platensis Argentina B Atlapetes albiceps Tumbesi forbesi f* j leucopterus IW Ecuador Apurimac rufigenis** C. Blanca seebohmi* NW Peru nationi W Peru setaria SE Brazil baeri** Tucuman hypochondria^ Bol. NWArg. C Poospiza caesar* Apurimac garleppi** Cochabamba whittii Bol.- NWArg. nigrorufa La Plata I boliviana* Cochabamba thoraci SE Brazil rubecula** Lima alticola***' C. Blanca hispanolensis W Peru dresseri Tumbes pallidiceps SW Ecuador schistaceus* N Andes, C Peru rufinucha group humid Andes xorquata -Chaco cinerea Cerrado melanoleuca Chaco erythrophrys Bol.-NW Arg. ornata Arg. FIGURE 5 - Phylogenetic hypotheses for Leptasthenura spinetails (A), the Atlapetes schistaceous group of brushfinches (B) and Poospiza warblingfinches (C). One to three asterisks (*) indicate increasing association with Polylepis. 350 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Disjunct distributions suggesting young range fragmentations. Many species inhabit widely separated areas (Figures 2d-f and 4b). Since the populations are weakly dif¬ ferentiated a fairly recent range fragmentation can be inferred. For birds of Polylepis near the humid treeline ecotone, breaks have taken place mainly between Ecuador and Peru, and between C. Blanca-adjacent Pasco and Abancay-C. Real. This south¬ ern gap consists of low ridges with limited cloud-cover interrupted by many dry val¬ leys. These ridges may be poor for these birds. Cycles of dispersal and range fragmentation in central Peru can be invoked to explain the differentiation of two megasubspecies of Xenodacnis parina*, and of Chalcostigma Stanley /* and olivaceum, now partly sympatric. For birds of semiarid wood/scrub, range disjunctions exist along the Pacific slope and toward small populations in the valleys of Huancavelica-Ayacucho and the Titicaca basin. Remnants of a track from Cochabamba to central Peru can be seen for Phacellodomus striaticeps. This species inhabits xerophytic thickets in the lower tem¬ perate zone from northern Argentina to La Paz ( striaticeps ) and in valleys of Cuzco- Apurimac ( griseipectus ). An unnamed subspecies* inhabits a large Polylepis wood¬ land near Lampa (Lake Titicaca area). Lessons from current dispersal. Distribution details showing how species locally ex¬ pand out of their normal life-zone to enter another may exemplify how dynamic biogeographic changes could have happened in the past: Many lowland species of arid woodland as well as humid shrub enter the Cochabamba basin, reaching the Polylepis woodlands or almost the puna edge. I suggest that the mosaic of rainshadows and moisture-capturing ridges, together with the expected rise in temperatures in rainshadow areas, create opportunities for low¬ land birds of various ecological requirements to use Bolivian valleys as gates to the highlands. Some of the Cochabamba populations are differentiated as subspecies, so we are not watching effects of man-made habitat changes. In Apurimac, some of the more adaptable humid-forest birds colonize Polylepis- dot¬ ted scarps fringing arid valleys (e.g., Ampelion rubrocristatus). Distributional data also show some exchange of species across the pass between Huanuco and the Upper Maranon C. Blanca (Figure 3a) or over to northern Lima (isolated populations of Coeligana violifer, Pterophanes cyanoptera and Pipraeidea melanonota). Humid-forest birds like Veniliornis fumigatus and Basileuterus nigrocristatus occur above 4000 m in C. Blanca. Similar colonizations may also have taken place long ago, judging from the presence here of a highland megasubspecies* of the widespread humid-forest Mecocerculus leucophrys. Many birds of the arid Pacific slope make considerable vertical movements, season¬ ally or in response to an el nino cycle. Isolated populations of some west slope birds in some arid, east-draining valleys suggest crossings of the West Cordillera in north Peru, in C. Blanca (where Colaptes atricollis lives above 4000 m) and from the Rio Canete drainage in southern Lima. The high-elevation birds Grallaria andicola* and Xenodacnis p. parina* colonize humid treeline habitat in the eastern Andes. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 351 CONCLUSIONS Correlations of areas of endemism can be used to corroborate or falsify hypotheses about patterns of dispersal. In the present case the suggested routes are in good agreement with current patterns of dispersal, suggesting that biogeographic proc¬ esses opperating today also did so, periodically, in the past. An important track went from the foothills in Bolivia to Cochabamba, and then across the Altiplano to the Pa¬ cific slope of Peru, or to Apurimac and from there to C. Blanca and Lima. Also, dis¬ persal of birds of humid montane forest to C. Blanca and the Pacific slope of north¬ ern Peru is important, and may have happened in the distant past judging from the fact that most specialized (old) Polylepis birds have their counterparts on the humid eastern Andean slope. After specializing to high elevation woodlands they may have had periodic opportunities to disperse from C. Blanca to Abancay-C. Real along the humid eastern ecotone (Figure 3b), and to speciate after range interruption. However, only part ot the speciations share geographic histories, other events being unique, as species have used a number of different ways to colonize high-elevation woods. FIGURE 6 - Hypothesized species-generating areas. Stippled is maximum area of Pleistocene glaciers. East-facing slopes that may have remained wooded are shaded. 352 ACTA XX CONGRESSUS I NITER NATION ALIS ORNITHOLOGICI Many species of Andean valleys and slopes, endemic and widespread forms alike, ascend to Polylepis on scarps toward adjacent highlands, but apparently this situa¬ tion at the margin of the ecological range does not give the break in geneflow nec¬ essary for specialization. Specializations to highest-elevation woodlands are shown only by forms living far from the possible founding areas (or in part living far away, permitting the interpretation that proximity is secondary). This pattern suggests that the specialization was allopatric rather than peripatric. (However, restriction to high elevation habitat could be reinforced by successive colonizions of an area, because of competitive interaction from the more recent colonizer, as in Aglaeactis cupripennis/ castelnaudii* and Metallura williamilbaroni*, see Fjeldsa & Krabbe 1990: 258 and 282). I postulate that specialization to high-elevation woodland required interruption of geneflow with lower-elevation stocks, especially if opportunities to spread from the humid East Andean slope to C. Blanca or Lima were followed by habitat contraction. Isolated on the Pacific slope, these birds found the most permanently humid habitat in the Polylepis zone. A few species also arrived at the Pacific slope from the scrubby valleys of Bolivia directly across the altiplano or via Apurfmac. Incipient specialization to Polylepis could have been reinforced by periods of isolation in Polylepis woodland on the altiplano, far from Andean slopes and valleys (Vuilleumier 1986). The pattern of endemism does not support the view that woodland birds survived glacial periods in ice-free refugia around Lake Junfn (Fjeldsa 1981, Hansen et al. 1984) and on the altiplano (Simpson 1975, Servant & Fontes 1978). The suggested patterns fit well with the assumed Pleistocene cycles of contraction and expansion of forest habitat (Simpson & Haffer 1978, Flenley 1979, Prance 1982). When discussing potential Pleistocene refugia it is worth mentioning the topographic situation of C. Blanca, the Abancay area and the Cochabamba basin (Figure 6). All are isolated by mountain ranges from the humid slopes, but sufficiently close for some rainfall from the clouds that often surround the snow-covered peaks at the headwa¬ ters of the Maranon, where the Apurfmac Canyon intersects the highland north of Abancay, and in the Tunari Range isolating the Cochabamba basin. Because of posi¬ tive effects of glaciers on the climate in immediately adjacent refuge areas (Lindroth 1965), and large past wetlands in the Cochabamba basin, these areas are likely to have maintained some woodland throughout the glacial cycles and have entrapped populations of a variety of once-widespread forms, thus generating new species. Steep elevational gradients from the cool and humid zone near the glaciers to the hot and arid climate at the bottom of rain-shadow valleys would permit wide adaptational diversity in these refuges. Subsequently, lower-elevation endemics could spread through the montane valley, while Polylepis specialists could follow moisture-capturing ridges across central Peru or pass from Abancay to C. Real. Simpson (1975) suggests the presence of a Polylepis zone on the Pacific slope of Peru during the glacial pe¬ riods. It is hard to accept Cracraft’s (1985) view that present areas of endemism are discrete pre-Pleistocene units inhabited by subspecies and species alike, irrespective of habi¬ tat, as opposed to “refuges”. The attempt is weakened by his above-mentioned inacuracies when allocating species to specific areas. Nevertheless, the hypothesis of speciation resulting from Pleistocene climatic cycles needs testing using “molecular evolutionary clocks” such as DNA base-sequencing, predicting that Pleistocene cy¬ cles should result in rather constant nodal distances across several species groups. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 353 ACKNOWLEDGEMENTS For companionship and help during the data-gathering I am indebted to Niels Krabbe (the many other persons who assisted were listed in Fjeldsa & Krabbe 1990). I thank Peter Arctander, Jan Fisher Rasmussen and Francois Vuilleumier for their comments on the manuscript. LITERATURE CITED ANSION, I. 1986. El arbol y el bosque en la sociedad andina. Instituto nacional Forestal y Fauna - FAO. COLLAR, N.J., GONZAGA, L.P., KRABBE, N. MS. Threatened birds of the Americas. Cambridge, U.K.: ICBP and IUCN. CRACRAFT, J. 1985. Historical biogeography and patterns of differentiation within the South Ameri¬ can avifauna: areas of endemism. Pp. 49-84 in Buckley, P.A., Foster, M.S., Norton, E., Ridgely, R.S., Buckley, F.G. (Eds) Neotropical Ornithology. Washington, D.C., AOU Ornithological Monographs No. 36. FJELDSA, J. 1981. Comparative ecology of Peruvian grebes - a study of the mechanisms of evolution of ecological isolation. Videnskabelige Meddelelser fra dansk naturhistorisk Forenening 143: 125-249. FJELDSA, J. 1987. Birds of relict forests in the high Andes of Peru and Bolivia. Copenhagen: Zoologi¬ cal Museum. FJELDSA, J. in press. The activity of birds during snow-storms in high-elevation woodlands in Peru. Bulletin of the British Ornithologists Club. FJELDSA, J., KRABBE, N. 1990. Birds of the High Andes. Copenhagen: Zoological Museum, and Svendborg: Apollo Books. FLENLEY, J.R. 1979. The rainforest: a geological history. London: Butterworths. FRIMER, O., NIELSEN, S.M. 1989. The status of Polylepis forests and their avifauna in Cordillera Blanca, Peru. Copenhagen: Zoological Museum. HANSEN, B.C.S., WRIGHT, H.E., BRADBURY, J.P. 1984. Pollen studies in the Junin area, central Peruvian Andes. Geological Society of America Bulletin 95: 1454-65. LINDROTH, C.H. 1965. Skaftafell, Iceland - a living glacial refugium. Oikos(SuppL): 1-142. PRANCE, G.T. 1982. A review of the phytogeographic evidence for Pleistocene climate changes in the neotropics. Annals of the Missouri Botanical Garden 69:594-624. SERVANT, M., FONTES, J.C. 1978. Les lacs quaternaires des hauts plateaux des Andes boliviennes. Premieres interpretations paleoclimatiques. Cahiers ORSTOM, Ser. Geol. 10:9-23. SIMPSON, B. 1975. Pleistocene changes in the flora of the tropical Andes. Paleobiology 1: 273-94. SIMPSON, B.B., HAFFER, J. 1978. Speciation patterns in the Amazonian forest biota. Annual Review of Ecology and Systematics 9: 497-518. VUILLEUMIER, F. 1986. Origins of the tropical avifaunas of the high Andes. Pp. 586-622 in Vuilleumier, F., M. Monasterio (Eds) High altitude tropical biogeography. New York: Oxford Univ. Press. 354 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI CONCLUDING REMARKS: BIOGEOGRAPHY AND SPECIATION IN NEOTROPICAL BIRDS FRANQOIS VUILLEUMIER Department of Ornithology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA INTRODUCTION The papers in this symposium cover the Neotropics from its northern limit (Escalante) to its southern extremity (Vuilleumier), including its main mountain system (Franke, Fjeldsa), and its largest river (Capparella). These papers also offer several ap¬ proaches to biogeography and speciation. Thus, emphasis varies from consideration of a single taxon ( Geothlypis ), to analysis of an entire fauna (Patagonia). Ecologically, the habitats represented include open vegetation, lowland rainforests, and montane woodlands. The geographical focus on the high altitude woodlands is either specific (Peru) or general (tropical Andes). Finally, a single question is treated in the case of Amazonia, whether rivers of this basin act as barriers. The diversity of biological ques¬ tions and of research strategies used in these papers reflects the ecological, taxonomical, and evolutionary diversity of the Neotropical Region itself. HOW MUCH IS BEING DONE The sample of research provided in this symposium shows the degree of detail pur¬ sued today. At the local level, Franke has surveyed dozens of dry cloud forest patches along the western Andes of Peru, whereas at the global level Fjeldsa has explored dozens of relictual woodlands over the entire length of the tropical Andes. Key taxa have been assayed with electrophoresis for a better understanding of population structure in areas as different as Baja California (Escalante) and the heart of Amazonia (Capparella). And parapatric contacts between sister taxa are studied at the southernmost tip of the region (Vuilleumier). HOW MUCH REMAINS TO BE DONE With such a vast region and such a rich fauna, there is almost no end to the number of questions that need answering. The authors of these papers point out specific av¬ enues for future work. Thus Capparella urges the use of biochemical data for a bet¬ ter assessment of areas that should be protected in Amazonia. The practical appli¬ cation of scientific results from speciation analysis to conservation cannot be more clear. On a more academic level, Capparella’s research shows the need for more paleoecological data in lowland rainforests, so that better correlations can be made between history inferred from electrophoretic information and history inferred from dated sequences of fossil pollen. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 355 Franke’s work emphasizes the need tor good, well labeled series of specimens from many localities in order to make accurate assessments of detailed distribution pat¬ terns. Vuilleumier’s review reveals the uniqueness of the Patagonian avifauna. Southernmost South America appears to be both a diversity sink (relict taxa) and a diversity generator (vicariance in situ). Escalante makes tantalizing suggestions about fine grained differences between warblers that live in wetlands and others that live in drier habitats. In spite of hard work, still larger samples will be needed before one can elucidate the switch from marsh to non-marsh in Geothlypis. Fjeldsa mentions taxa that can be singled out for study of colonization of relict woodlands at very high alti¬ tudes in the Andes. CONCEPTUAL ADVANCES At the XVI Ith IOC in Berlin, Short convened a symposium on speciation in South American birds, with papers on upper Amazonia, the high Andes, suboscine versus oscine radiation, woodpecker speciation, and flycatcher speciation. Geographically the present symposium reveals a finer level of analysis than the one in Berlin. Methodo¬ logically, the blend of morphological, ecological, and biochemical kinds of evidence is now routine in the Neotropics. The liquid nitrogen tank is today a standard piece of equipment in expeditions. What about conceptual advances provided by this more versatile approach? In terms of the refuge theory, I worry that the advances made by paleobotanists, such as those discussed by Huntley (1988) and Webb (1988) at the 19th IOC in Ottawa, have not yet been incorporated by ornithologists in their daily thinking. In terms of geographical ecology we have made good progress away from a quasi- typological thinking focused on subspecies (embodied in Zimmer’s work on geographi¬ cal variation in birds from the Peruvian Andes), as is evident when reading Franke’s paper. Subspecies are a tool for distributional analysis, no longer an end in them¬ selves. In terms of speciation, many more cases have been studied since the 1978 IOC in Berlin. Hence research on the course of speciation (Vuilleumier 1980) has progressed to the point where enough taxa have now been analyzed, and from so many more areas of the Neotropics, that one can see at once, and in much richer detail, both the unity of the process in different taxa and different areas, and the diversity of its de¬ velopment. Nevertheless, more research is needed on parapatric taxa, in order to sort out the confusing array of potential causal factors that have produced parapatry. In other words, we have not yet obtained enough evidence to fully refute Endler’s (1982) claims that at least some parapatric patterns could be the result of speciation along dines that are or were parallel to environmental gradients. LITERATURE CITED ENDLER, J. 1982. Pleistocene forest refuges: fact or fancy? Pp. 179-200 in Prance, G. (Ed.) Biologi¬ cal diversification in the tropics. New York, Columbia University Press. HUNTLEY, B. 1988. European post-glacial vegetation history: a new perspective. Acta XIX Congressus International^ Ornithologici Berlin (1978): 1061-1077. VUILLEUMIER, F. 1980. Reconstructing the course of speciation. Acta XVII Congressus International^ Ornitholgici Berlin (1978): 1296-1301. WEBB III, T. 1988. Vegetational change in eastern North America from 18000 to 500 BP. Acta XIX Congressus International^ Ornithologici Ottawa (1986): 1050-1060. 356 ACTA XX C US INTERNATIONALIS ORNITHOLOGICI ' - ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 357 SYMPOSIUM 2 ORIGINS AND EVOLUTION OF THE AUSTRALASIAN AVIFAUNA Conveners R. SCHODDE and L. CHRISTIDIS 358 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI SYMPOSIUM 2 Contents INTRODUCTORY REMARKS: ORIGINS AND EVOLUTION OF THE AUSTRALASIAN AVIFAUNA LES CHRISTIDIS . 359 GEOLOGICAL EVOLUTION AND BIOTIC LINKS IN THE MESOZOIC AND CENOZOIC OF THE SOUTHWEST PACIFIC G. R. STEVENS . . 361 THE ORIGIN AND RADIATION OF AUSTRALASIAN BIRDS: PERSPECTIVES FROM THE FOSSIL RECORD WALTER E. BOLES . 383 BIOCHEMICAL EVIDENCE FOR THE ORIGINS AND EVOLUTIONARY RADIATIONS IN THE AUSTRALASIAN AVIFAUNA: THE SONGBIRDS LES CHRISTIDIS . 392 THE EVOLUTIONARY HISTORY OF PARROTS AND COCKATOOS: A MODEL FOR EVOLUTION IN THE AUSTRALASIAN AVIFAUNA DOMINIQUE HOMBERGER . 398 THE DEVELOPMENT OF MODERN AVIFAUNULAS RICHARD SCHODDE and DANIEL P. FAITH . 404 CONCLUDING REMARKS: ORIGINS AND EVOLUTION OF THE AUSTRALASIAN AVIFAUNA RICHARD SCHODDE . 413 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 359 INTRODUCTORY REMARKS: ORIGINS AND EVOLUTION OF THE AUSTRALASIAN AVIFAUNA LES CHRISTIDIS Department of Ornithology, Museum of Victoria, 71 Victoria Crescent, Abbotsford, 3067 Australia. Zoogeographically, Australasia stretches from Sulawesi in the west to the Hawaiin Islands and Easter Island in the east. Its limits are not clear-cut and the boundary between the Australasian and Oriental regions has not been resolved conclusively, perhaps because it cannot be. The geological history of the boundary is complex, with Sulawesi formed from both Gondwanan (eastern half) and Eurasian (western half) elements (Audley-Charles 1981). The habitats of Australasia cover the whole range of global environments, from semi-arid deserts to tropical rainforests, alpine grasslands and temperate scleromorphic forests. These are interspersed amongst continental land masses, volcanic islands and islands which are part of continental plates. All these factors combined have produced in Australasia a very rich and unique avifauna, with a greater species diversity per area than any other zoogeographical region. Over 90 percent of the land and freshwater bird species are unique to the region. Not suprisingly, the Australasian avifauna - its origins and radiations - has continued to capture the attention of biologists since the time of A.R. Wallace. Considerable interest and controversy has centred on the relationships of the land bird fauna. Tra¬ ditional theory has had it that Australasia received its avifauna through a series of successive waves of inmigration from Eurasia over the last 30 to 40 million years (Keast 1984). Advances in plate tectonics, paleontology and molecular biology have challenged this view. The timing of continental drift has revealed that when Australia was supposed to be getting its early stocks of Eurasian birds, it was in fact thousands of kilometres further south towards Antarctica (Rich 1975). Moreover, fossil evidence demonstrates the presence of birds in Australia 110 MYBP and passerines at least 30 MYBP (Rich and Baird 1986). These facts have led to the conclusions that some of the major orders of birds are Gondwanan in origin. Which groups are Gondwanan, however, is still some way from final resolution (Olson 1988). By far the greatest controversy centres around the molecular data from DNA-DNA hy¬ bridization (Sibley & Ahlquist 1985). The results so far suggest that the Australasian passerines are autochthonous and have undergone marsupial-like radiations. The radiations themselves seem to be old, lending support to the view that the passerines originated in Gondwana. The interest generated from these studies is reflected in the fact that the origins and subsequent radiations in the Australasian avifauna have been popular as subjects of symposia and papers at three of the last six International Ornithological Congresses. The venue of the 20th congress here in the Antipodes adds to the topicality of yet 360 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI another symposium on the subject, here drawing on its information from the follow¬ ing aspects: regional palaeogeographic history, the regional fossil record, molecular contributions on the origins of the passerines, adaptive radiations in the parrots and cockatoos, and the biogeographical history of regional avifaunal assemblages. LITERATURE CITED AUDLEY-CHARLES, M.G. 1981. Geological history of the region of Wallaces Line. Pp. 24-35 in Whitemore, T.C. (Ed.). Wallace's Line and Plate Tectonics. Oxford Monographs in Biogeography 1. Oxford: Oxford University Press. KEAST, A. 1984. Contemporary ornithogeography: The Australian avifauna, its relationships and evo¬ lution. Pp 457-468 in Archer, M., Clayton, G. (Eds). Vertebrate zoogeography and evolution in Australa¬ sia. Perth: Hesperian Press. OLSON, S.L. 1988. Aspects of global avifaunal dynamics during the Cenozoic. Pp. 2023-2029 in Ouellet, H. (Ed.). Acta XIX Congressus International^ Ornithologici. Ottawa: University of Ottawa press. RICH, P.V. 1975. Antarctic dispersal routes, wandering continents and the origin of Australasia's non- passeriform avifauna. Memoirs National Museum Victoria. 36: 63-126. RICH, P.V., BAIRD, R. 1986. History of the Australian avifauna. Pp. 97-139 in Johnston, R.F. (Ed.). Current Ornithology 4. New York: Plenum Press. SIBLEY, C.G., AHLQUIST, J.E. 1985. The phylogeny and classification of the Australo-Papuan pas¬ serine birds. Emu 85: 1-14. / ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 361 GEOLOGICAL EVOLUTION AND BIOTIC LINKS IN THE MESOZOIC AND CENOZOIC OF THE SOUTHWEST PACIFIC G. R. STEVENS DSIR Geology & Geophysics, Department of Scientific and Industrial Research, P.O. Box 30368, Lower Hutt, New Zealand. ABSTRACT. From middle Jurassic times earth movements began to create considerable areas of land in the New Zealand region. By late Jurassic and early Cretaceous times, a large landmass had been developed, extending north towards New Caledonia, east to Chatham Islands, west to Lord Howe Rise and south to the edge of Campbell Plateau. Development of this landmass, and its associated terres¬ trial links, together with the warm-temperate conditions then apparent over large areas of Gondwana, provided favourable conditions for the spread of Gondwanan elements in the region. In early and mid¬ dle Cretaceous Australia, New Zealand, New Caledonia and Antarctica were characterised by cool- temperate climates, and these regions shared many cool-temperate biotic elements; land links provided access and exchange for early angiosperms. However, all land links between New Zealand/New Cal¬ edonia and Australia/Antarctica were broken after 85 Ma. From this time onwards the ancestral Tasman Sea and Southern Ocean became effective barriers to overland dispersal between southeastern Gondwana and New Zealand/New Caledonia. Subsequently, and consequently all terrestrial colonists had to arrive by flying, swimming or floating. Many birds did so, but no terrestrial snakes and no mam¬ mals, except bats. Keywords: Jurassic, Cretaceous, Cenozoic, Rangitata Orogeny, Gondwana, New Caledonia, Papua New Guinea, Australia, New Zealand, Antarctica, South America. INTRODUCTION Throughout early Mesozoic time immediately preceding the advent of birds, the SW Pacific region underwent a series of major changes in geography, climate and topog¬ raphy that set the scene for the biotic developments that took place in the late Mesozoic and Cenozoic. During this time, the major landmasses that now comprise the SW Pacific region (i.e. Australia, Antarctica, New Zealand and New Caledonia) were integral parts of the southeastern edge of the supercontinent Gondwana. In Carboniferous and Permian times Gondwana had slowly drifted across the South Pole, and large areas of Australia and Antarctica had been scoured by continental ice sheets (McKerrow & Scotese 1990). During these times the areas now occupied by New Zealand and New Caledonia (then virtually one landmass) were largely under the sea, except for ephemeral volcanic islands, and as a consequence escaped the ef¬ fects of glaciation (Stevens 1985). During the Triassic the same global movements continued that had earlier rotated the continents of southeastern Gondwana towards the South Pole. However, in the Triassic these movements had the net effect of gradually moving southeastern Gondwana into mid-latitudes. Nonetheless, New Zealand (and New Caledonia) re¬ mained in high latitudes (70°-80°S) for much of Triassic time. But, as it is likely that the world was ice-free during the Triassic (Hallam 1985), New Zealand and New Cal¬ edonian climates were probably not colder than cool- or cold-temperate (Stevens 1980a, 1985). 362 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI FIGURE 1 - Diagrammatic reconstruction of the southeastern margin of Gondwana in the middle and late Jurassic, ca. 165-145 Ma. Base map for this and the succeeding diagrams Figures 2-3 modified from Grunow, Dalziel & Kent (1987), Grunow, Kent & Dalziel (1987), Lawver & Scotese (1987), de Wit et al. (1988). The diagonal line pattern indicates regions in which extensive areas of land were present at the time, although its exact distribution is not known. At this time, movements of the Rangitata Orogeny were creating new land in the New Zealand region. Development of this new land provided opportunities for New Zealand to share in the terrestrial biota of adjacent Gondwana lands, as indicated by the solid arrows. Biota that populated New Zealand at this time (and perhaps also earlier in the Triassic, when land links were also present), included at least some of the ancestral stocks of the “archaic" elements present in the modern New Zealand flora and fauna. Representa¬ tives of such stocks are shown along the right hand edge of the diagram: (from top to bot¬ tom) the Kauri Agathis australis (araucarian stocks): podocarp stocks; tree-fern stocks; the New Zealand Frog Leiopelma; Weta Deinacrida heteracantha (Rhaphidophoridae), Tuatara Sphenodon punctatus; Peripatus; the giant New Zealand Land Snail Paryphanta. Ratite birds, including the ancestors of the New Zealand Moa (Dinornithiformes) may also have populated Gondwana lands at this time, assuming that walking rather than flying was their preferred mode of locomotion. The boxes highlight the factors (e.g. “Rangitata Orogeny”) that influenced biotic developments in the New Zealand region at the times covered by the reconstructions. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 363 The Triassic saw a carry-over of the general sedimentary conditions of the Permian, with much of New Zealand and New Caledonia continuing to be sites of active island arc systems, flanking the coastline of southeastern Gondwana. However, judging from sequences of non-marine beds, there was probably more land present in New Zea¬ land and New Caledonia in the Triassic than in the Permian (Stevens & Suggate 1978, Retallack 1987). The new land areas that developed may have established more per¬ manent land links to southeastern Gondwana (in contrast to the probably more ephemeral links of the Permian), and provided access for ancestral plant stocks, in¬ cluding ancestors of groups such as psilotales, ferns, lycopods, araucarians, podocarps and ginkgos (Fleming 1975, p. 10, 1977a, 1978). JURASSIC Rotation of eastern Gondwana in a direction away from the South Pole continued in the Jurassic, changing New Zealand’s geographical position to 60°-70°S by middle Jurassic times (Stevens 1980a, 1985). As a result of the progressive movement of land away from the polar regions, world climates continued to improve, and conse¬ quently, Jurassic climates were appreciably more equable than those of the present day (Stevens 1980a, Hallam 1975, 1985, Frakes 1986). Evolution of Greater New Zealand Throughout Jurassic times the earth movements of the Rangitata Orogeny, the initial stirrings of which were evident in the middle and late Triassic, became even more marked (Fleming 1967a, 1970, Stevens 1978, Suggate, 1978, Bradshaw et al. 1981). These movements progressively folded and elevated above sea level much of the material that had been deposited in the basin structures of the Carboniferous/Triassic arcs in the New Zealand - New Caledonian region. By the middle Jurassic, and ex¬ tending into the earliest Cretaceous, a large new landmass had developed, which extended far beyond the shorelines of modern New Zealand: northwards at least as far as New Caledonia, westwards to the Lord Howe Rise; eastwards to the Chatham Islands; and southwards to the southern edge of the Campbell Plateau (Figure 1). As rifting had not yet begun on what was to become the site of the Tasman Sea and Southern Ocean, such a landmass was virtually continuous with Australia and Antarc¬ tica, which at that time were largely emergent (e.g. Storey et al., 1987, St. John 1984, White 1986). Long fingers of land and interlinked chains of islands probably also ex¬ tended northwards from the new landmass towards Papua New Guinea and Indone¬ sia, although exactly how far is not known. The formation of such large areas of land in the middle/late Jurassic and earliest Cretaceous probably constituted the greatest development of new land that ever occurred in the New Zealand region. Dispersal of “archaic” biotic elements to New Zealand Development of “Greater New Zealand” and its links with the lands of southeastern Gondwana, when taken together with the equable climatic conditions that were then widespread, provided optimal opportunities for the spread of terrestrial organisms. At this time, the ancestors of many of the “archaic” elements present in the modern New Zealand terrestrial biota became established assuming that their stocks had originally developed in other areas. Among plants, these ancestral stocks may have included araucarians and podocarps, following on from the Triassic. Among animals, ances¬ tral stocks of at least some of the following became established in New Zealand in the 364 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI FIGURE 2 - Diagrammatic reconstruction of the southeastern margin of Gondwana in the early Cretaceous, ca. 1 20-1 1 0 Ma. The main uplift phases of the Rangitata Orogeny were continuing during this time. The early Cretaceous, together with the late Jurassic, consti¬ tutes the greatest extension of land in New Zealand’s geological history - the development of the New Zealand microcontinent or “Greater New Zealand” (indicated by the speckled pattern). In the Aptian-Albian a rifting phase (indicated by thick lines) commenced along the west coast of New Zealand, along the western side of the Lord Howe Rise, and in the Bounty Trough. Major rifting also commenced between the African and South American plates (indicated by the toothed pattern). Ancestral angiosperms appeared at about this time and radiated throughout the Gondwana margins. Because of the disruption of land routes by the onset of rifting along the western edge of “Greater New Zealand”, heralding formation of the Tasman Sea, the ancestral angiosperms may have used a land route into New Zealand via Antarctica (as indicated by the solid arrow). Ancestral ratite birds, includ¬ ing ancestral moas, may have also used this route (as also in the late Jurassic, Figure 1), if they were flightless at this time. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 365 Jurassic: the New Zealand frogs, Leiopelma, the Tuatara Sphenodon, skinks and leptodactylid geckos; onychophorans (Peripatus); and other groups of freshwater and terrestrial invertebrates (Keast 1973, Fleming 1975, Bull & Whitaker 1975, King 1987, Worthy 1987 a, b) (Figure 1). Although contrary opinions are held (e.g., Eskov 1987, Craw 1985, 1988), it is con¬ sidered that ancestors of these modern organisms were originally widely distributed across the lands of eastern Gondwana and that establishment of land links between “Greater New Zealand” and the cratonic areas of Australia and Antarctica, then largely above the sea, facilitated their spread into New Zealand and probably New Caledo¬ nia. There is no fossil record to support such a proposal (Fleming 1975) but Gondwanan relationships are known, for example, for sphenodontian reptiles (Fraser 1986, Benton 1986), leptodactylid geckos (King 1987) and taxa related to Leiopelma (Estes & Reig 1973) . It is generally accepted that the Rangitata Orogeny reached its climax in the late Jurassic and early Cretaceous, (Suggate 1978). “Greater New Zealand” probably also reached its maximum size during this time. Concurrently, continuing rotation of Gondwana swung New Zealand even further northwards from its middle Jurassic position. From late Jurassic to possibly also earliest Cretaceous times, New Zealand was lying at about 55°S and was influenced by sub-tropical/warm-temperate condi¬ tions (Stevens 1980a, 1985). This period is considered to have provided the greatest (as well as probably the last) opportunity for subtropical/warm-temperate terrestrial or¬ ganisms to move from southeastern Gondwana to New Zealand and New Caledonia along continuous land routes. The early birds, newly differentiated from their ancestral reptilian stocks (Feduccia 1980, Olson 1985, Cracraft 1988), first began to occupy various ecological niches throughout the world in the late Jurassic and earliest Cretaceous. Judging from oc¬ currences of fossil feathers, they were present in Australia at least as early as 120 Ma (Talent et al. 1966, Rich et al. 1989, Rich 1976, 1979, Rich & Baird, 1986). At this time, substantial areas of land existed in southeastern Gondwana, comprising ances¬ tral Australia, Antarctica, New Zealand and New Caledonia. These areas probably formed one continuous landmass, with broad links to the remainder of Gondwana. Furthermore, the climate was reasonably uniform and equable, ranging from subtropi¬ cal to warm-temperate. It is likely therefore that the early birds found few if any bar¬ riers, geographic or climatic, to their spread throughout eastern Gondwana in those times, providing the most propitious circumstances for at least some of the ancestral stocks of ratite birds to have become established in New Zealand (notably those of the Moa, Dinornithiformes; Cooper et al. 1990). Flowever, this scenario assumes that the ancestral ratites were already virtually flightless and needed continuous land routes for dispersion (Fleming 1982, Stevens 1985, 1989). EARLY CRETACEOUS The phase during which extensive land connections existed between “Greater New Zealand” and southeastern Gondwana was brought to an end in the early Cretaceous. During this time an extensive rift system developed along the western margin of New Zealand and the western flank of the Lord Howe Rise (Figure 2). The onset of this rifting marked the beginning of movements that eventually, in the late Cretaceous, led 366 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI / FIGURE 3 - Diagrammatic reconstruction of southeastern Gondwana in the middle Cre¬ taceous, ca. 100-95 Ma. By this time the altitude and extent of the “Greater New Zealand” landmass uplifted by the Rangitata Orogeny (Figures 1, 2) had been markedly reduced by thermal subsidence and by erosion, both marine and terrestrial; marine transgression was commencing along the east coast of New Zealand. Active rifting, accompanied by devel¬ opment of finger-like marine embayments along the rift zones, was occurring along the western margin of “Greater New Zealand”, marking the sites of the future Tasman Sea and Southern Ocean (indicated by the toothed pattern). The approximate distribution of land in the New Zealand region is indicated by the speckled pattern. The ancestral stocks of Southern Beech (Nothofagus) and Proteaceae were radiating to southern lands at about this time, using Antarctica as a stepping stone. Entry of these stocks to New Zealand was probably via a southern route, as indicated by the solid arrow. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 367 to development of oceanic crust in the Tasman Sea and Southern Ocean. The rifting movements began ca. 120 Ma and became particularly evident by ca. 110 Ma (Laird 1981, Whitworth et al. 1985, Sporli 1987, Tulloch & Kimbrough 1989). As the rifts progressively widened and deepened, they were invaded by long fingers of sea and pre-existing land connections became disrupted. During the late Jurassic and early Cretaceous, continental movements began in west¬ ern Gondwana that were to have major consequences in eastern Gondwana. The separation of Antarctica and India from Africa to form the Indian Ocean, and the sepa¬ ration of Africa from South America to form the South Atlantic Ocean, reversed the movement of eastern Gondwana that throughout the Triassic and Jurassic had been in a direction away from the South Pole (Stevens 1980b). As a result, the geographi¬ cal position of southeastern Gondwana (and more particularly, that of eastern Aus¬ tralia and New Zealand) changed from mid-latitude in the Jurassic to high latitude in the early Cretaceous (Stevens 1980a, 1985, Rich et al. 1989). It is likely that the cli¬ matic change accompanying this rotation was equivalent to a change from sub-tropi- cal/warm-temperate in the Jurassic to cool-/cold-temperate in the early Cretaceous (Stevens 1971, 1980a, 1985). Although ice may have been present at the North Pole during at least some of the time of the early Cretaceous (Kemper 1987), there are no indications that Cretaceous climates in New Zealand or Australia were ever colder than cold-temperate, despite New Zealand being within 5°-10° of the South Pole in the mid-Cretaceous (Oliver et al. 1979). Cool humid conditions occurred in New Zea¬ land, South Australia and Victoria in the early Cretaceous, with a varied biota, includ¬ ing forest vegetation, living in markedly seasonal climates with winter temperatures dropping close to freezing (Stevens & Speden 1978, Raine et al. 1981, Rich & Rich 1988, Frakes & Francis 1988, Rich et al. 1989, Douglas 1990, Francis & Frakes 1990). The extensive landmass of “Greater New Zealand” became exposed at this time to the effects of peneplanation and marine erosion, and of thermal subsidence of the lithosphere (Sporli & Ballance 1989). By mid-Cretaceous times, large areas of the land had been worn down to such low levels that widespread marine transgression was taking place, particularly along the eastern margin (Stevens & Suggate 1978, Stevens & Speden 1978). Erosion and diminution of the “Greater New Zealand” landmass, together with the development of active rifting zones between Australia and the west¬ ern margins of New Zealand and the Lord Plowe Rise, contributed to the steady de¬ terioration of the Gondwanan land links that had been a feature of Jurassic times. Plowever, judging from the presence of ancestral angiosperms in New Zealand from ca. 105 Ma onwards (Fleming 1975, Raine et al. 1981) (Figure 2), and of ancestral Southern Beech (Nothofagus) from ca. 90/85 Ma onwards (Dettmann, 1989, Dettmann et al. 1990) (Figure 3), some land links persisted, particularly towards the south, through western Antarctica to South America. The close affinities of New Zealand and South American Nothofagus (Poole 1987) suggest strong connections at this time for cool- or cold-temperate floras (Case, 1988, Dettmann, 1989). Rifting and subsidence had also commenced along the south coast of Australia at about 125 Ma (Hegarty et al. 1988) or 140 Ma (Williamson et al. 1990). By about 90 Ma it gave way to an episode of slow sea-floor spreading between Australia and Ant¬ arctica (Cande & Mutter 1982, Veevers 1986, 1987, Flegarty et al. 1988, Powell et al. 1988). However, the onset of rapid seafloor spreading and substantial separation of 368 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI FIGURE 4 - Diagrammatic reconstruction of southeastern Gondwana in the late Creta¬ ceous, ca. 75 Ma. Base map modified from Kamp (1986a). Rifting and slow spreading had been occurring between Australia and Antarctica since ca. 95 Ma (indicated by the toothed pattern). Active sea-floor spreading had been occurring since ca. 85 Ma in the Tasman Sea and in the sector of the Southern Ocean south of the southern edge of the Campbell Pla¬ teau (as indicated by the generalised sea-floor spreading pattern). Although land connec¬ tions between the New Zealand region and the remainder of Gondwana had been broken, it was still possible for volant animals to overfly the developing oceans before they became too wide and too stormy (as indicated by the two broad striped arrows). Biochemical data suggest that ancestral stocks of the New Zealand “wrens” may have become established in New Zealand at about this time. The approximate distribution of land in the New Zea¬ land region is indicated by the speckled pattern. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 369 the two continents did not begin until 55 Ma (Weissel & Hayes 1972). The impact of this tectonic activity on biotic interchanges between Australia and Antarctica (and by extension, South America) between 90 and 55 Ma is difficult to evaluate because of the extremely sparse terrestrial fossil record. However, occurrences of fossil Proteaceae, Nothofagus and other plants (Dettmann & Thomson 1987, Pocknall & Crosbie 1988, Dettmann & Jarzen 1988, Dettmann 1989, Dettmann et al. 1990) indi¬ cate that terrestrial connections persisted across the area of rifting, probably in the region between Victoria/Tasmania and North Victoria Land, Antarctica (Veevers, 1987, Wilson et al. 1989) and that these links continued into at least the latest Cretaceous, 85-75 Ma. The distribution of fossil marsupials in the late Eocene is also indicative of the persistence of southern land connections then, presumably linking Australia, Ant¬ arctica and South America (Woodburne 1982, Woodburne & Zinsmeister 1982, 1983, 1984, Case et al. 1988, Case 1988), but not including New Zealand, where indigenous marsupials are unknown. LATE CRETACEOUS Expansion of the Atlantic and Indian Oceans continued at a steady rate throughout late Cretaceous time. These movements affected many sectors of southern Gondwana and had the effect of swinging New Zealand northwards away from the South Pole, so that by the late Cretaceous it lay between 65°-55°S, compared with 85°S in the early-middle Cretaceous (Stevens 1980a, b, 1985). The same movements had also rotated West Antarctica, so that it now straddled the South Pole. Nonetheless, climatic conditions were evidently still not favourable for the accumulation of ice, as traces of late Cretaceous glaciation are unknown. On the contrary, there is abundant fossil evidence that large areas of Antarctica were prob¬ ably clothed in forests (presumably of Nothofagus and other cool-temperate taxa; Dettmann, 1989, Dettmann et al. 1990). Therefore, as in early and mid-Cretaceous times, the climate of New Zealand, Australia and other “southern” lands was cool- or cold-temperate and had well-defined seasons (Jefferson 1982, 1983, Francis 1986, Parrish & Spicer 1988a, b, Case 1988, Pirrie & Marshall 1990). Rifting movements that continued around the primaeval New Zealand landmass throughout early and mid-Cretaceous times culminated about 85 Ma with the estab¬ lishment of open ocean in the Tasman Sea and in the Southern Ocean south of New Zealand (Figure 4). The development of open oceanic conditions brought to an end any likelihood of land connections to New Zealand and New Caledonia from Australia and the remainder of southeastern Gondwana. As the Tasman Sea and the Southern Ocean opened, and as sea-floor spreading moved “Greater New Zealand” away from the remainder of southeastern Gondwana, its surface continued to be lowered by erosion and thermal subsidence (Kamp 1986a, b, Sporli 1987, Korsch & Wellman, 1988, Tulloch 1990). By late Cretaceous times much of it had been reduced to low relief and marine incursion was widespread. Biochemical data indicate that the ancestral stocks of the New Zealand “wrens” (Xenicidae: Rifleman, Bush Wren, Rock Wren, Stephens Island Wren; Bull & Whitaker 1975, Fleming 1982) were established by 85 to 90 Ma (Sibley, Williams & Ahlquist 370 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI FIGURE 5 - Diagrammatic reconstruction of southeastern Gondwana in the late Paleocene, ca. 57 Ma. Base map modified from Kamp (1986a). Active sea-floor spread¬ ing had ceased in the Tasman Sea at ca. 60 Ma. Sea-floor spreading was continuing in the Southern Ocean south of the Campbell Plateau and commenced at an accelerated pace between Australia and Antarctica at ca. 55 Ma (as indicated by the generalised sea-floor spreading pattern). Although land connections between New Zealand and the remainder of Gondwana had been broken in the late Cretaceous, ca. 85 Ma, it is likely that flying animals could still cross the developing oceans (as indicated by the two broad striped ar¬ rows). Such animals may have included the ancestors of the New Zealand Short-Tailed Bat Mystacina, the winged ancestors of the New Zealand Kiwi Apteryx, and the stocks that gave rise to the endemic families of New Zealand passerine birds. The drawings along the right edge of the diagram are of representatives of families that probably became estab¬ lished in New Zealand during Paleocene times. From top to bottom: Saddleback Philesturnus carunculatus, Huia Heteralocha acutirostris, New Zealand Thrush Turnagra capensis, New Zealand Short-Tailed Bat Mystacina tuberculata, Kokako or Blue-Wattled Crow Callaeas cinerea, Rock Wren Xenicus gilviventris, Kiwi Apteryx. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 371 1982, Sibley & Ahlquist, 1983). Although such biochemical “dating” should be re¬ garded with caution (cf. Houde 1987, Lewin 1988a, b, Kirsch & Krajewski 1988, Erwin 1989), it nevertheless suggests that the establishment of the ancestral stocks of the New Zealand “wrens” may have occurred just before the commencement of sea floor spreading in the Tasman Sea and the Southern Ocean, and could have been co-in- herited from eastern Gondwana. Given the later rifting of Australia from Antarctica, so also could the ancestral stocks of Australo-Papuan songbirds - the menceroids, meliphagoids and corvoids - have been co-inherited from Gondwana if they did not begin to diverge from one another and from Eurasian groups until about 55 or 50 Ma, as postulated by Sibley and Ahlquist (1985). PALEOCENE During the late Paleocene, ca. 60 Ma, the Tasman Sea had probably reached its present width (1950 km), or slightly more (as its overall width was subsequently re¬ duced by the late Cenozoic spreading of the Lord Howe Ridge, Stock & Molnar 1982). However, sea floor spreading continued south of New Zealand, between Antarctica and the southern edge of the Campbell Plateau. From this time onwards, the ances¬ tral Tasman Sea and Southern Ocean became effective barriers to overland routes into New Zealand, and all terrestrial colonists had to arrive by either flying or swim¬ ming/floating. Many birds did so, but no terrestrial snakes and no mammals except bats (Figure 5). Despite its increasing isolation, the changing archipelago of New Zealand continued to receive bird colonists from nearby lands especially from Australia, where a rich and diverse avifauna was developing throughout the Cenozoic (Rich 1976). After the ratites, which may have come by land (see above), the next bird colonists known to have arrived were the endemic families of New Zealand passerines - the New Zealand “wrens”, wattlebirds and “thrushes” (Xenicidae, Callaeatidae and Turnagridae) of uncertain relationships, but some showing “Gondwana” affinities with the ovenbirds and antbirds of South America, the lyrebirds of Australia and the pittas of the Old World tropics (Bull & Whitaker 1975, Fleming 1977b, 1982). The ancestral stocks of these endemic bird families may have arrived in the Paleocene, before the seas then opening around New Zealand became too wide and stormy. However, as mentioned earlier in this paper biochemical data point to the possibility that the an¬ cestral stocks of at least one endemic bird family - the New Zealand “wrens” - may have radiated to New Zealand in late Cretaceous times (95-90 Ma, according to Sibley & Ahlquist 1 983). At the same time as the ancestral stocks of the New Zealand endemic passerines were traversing the seas then opening up around New Zealand, the ancestors of the New Zealand Short-Tailed Bat Mystacina may have made a similar journey. Biochemi¬ cal studies of Mystacina have demonstrated that it has close phylogenetic affinities with Central and South American phyllostomoid bats and have suggested (using the assumption that bats originated in the Paleocene) that the lineages separated about 35 Ma (Daniel & Baker 1986, Pierson et al. 1982, 1986). It is therefore likely that the ancestral stocks of Mystacina followed a route from South America via Antarctica, and although as noted earlier such biochemical “dates” should be used with caution, ra¬ diation may have occurred sometime in the early Paleogene. 372 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Judging from deep sea drilling (e.g. Barker et al. 1987, Kerr, 1987) and palynological studies (e.g. Truswell 1984, Dettmann et al. 1990), large parts of Antarctica were forested during the Paleogene, until at least the early Oligocene. These forested ar¬ eas may have offered a trans-Antarctic connection with South America for ancestral Mystacina which, on encountering the steadily widening oceanic gaps then present around New Zealand, ffew across them to colonise the “Greater New Zealand” land- mass. Such a passage may have been via the Campbell Plateau, areas of which were emergent at this time (Figure 5). An alternative route may have involved radiation across Antarctica, thence into Australia and then via a route into New Zealand by means of island-hopping southwards along the Lord Howe Rise. The possible use of this alternative route is suggested by the presence on Mystacina of a tick normally parasitic on Australian bats (Daniel 1979). The volant immigrants that came to New Zealand in early Paleogene times by flying across the newly created surrounding oceans may have also included at least some of the flying ancestors of the ratites, particularly those of the Kiwi (Apteryx), if the views of Houde (1986), Howgate (1986) and the biochemical data of Sibley and Ahlquist (1987) and Cooper et al. (1990) are accepted. The opening-up of ocean between New Zealand and the southeastern margin of Gondwana had the effect of moving New Zealand and New Caledonia northwards away from the South Pole, so that in the Paleocene they were in latitudes of 50°-45°S compared with 65°-55°S in the late Cretaceous (Stevens 1985). The effects of this northwards movement are evident in the New Zealand marine fau¬ nas, as indicated by increasing influxes of warm-water Malayo-Pacific molluscan taxa beginning in late Paleocene times (Fleming 1967b, 1975, Beu & Maxwell 1968, Beu et al. 1990, Hornibrook 1978). EOCENE Sea-floor spreading finished activity in the Tasman Sea in the late Paleocene, ca. 60 Ma. Subsequently, sea-floor spreading became focussed in the Southern Ocean south of New Zealand and Australia. Although rifting between Australia and Antarctica had been occurring since the late Cretaceous, ca. 95 Ma (Cande & Mutter 1982; Veevers 1986, 1987), active sea-floor spreading did not commence until early Eocene, ca. 55 Ma. After this time, Australia and Antarctica moved apart, although contact via the South Tasman Rise (extending southwards from Tasmania) continued until ca. 35 Ma (Kennett et al. 1975). As Antarctica moved southwards, Australia and New Zealand moved northwards. New Zealand’s geographic position changed from 50°-45°S in the Paleocene to 45°-40°S latitude in the late Eocene (Stevens 1985). The gradual build-up of ice on Antarctica, commencing on East Antarctica in the ear¬ liest Oligocene (or earlier in highland valleys) and on West Antarctica in the late Miocene (Barker et al. 1987; Barron et al. 1988), combined with the opening-up of surrounding seaways put an end to Antarctica’s role as a stepping stone for south¬ ern migrants. At the same time, these developments led to the establishment of Circum-Antarctic wind and ocean current systems (Kennett et al. 1975, Barker & Burrell 1977). As these systems reached their optimum development (typified by the ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 373 West Wind Drift), increasing numbers of plants and animals adapted to dispersal became distributed around the southern lands (the “Neoaustral” elements of Fleming 1975), including shore and pelagic birds such as penguins (Figure 6). TRANS-TASMAN CENOZOIC LINKS Although Australia has contributed to New Zealand’s flora and fauna throughout most epochs of geological history, trans-Tasman migration was greatly strengthened by the advent of the West Wind Drift, following the separation of Australia and Antarctica. Consequently, from Miocene times onwards there was a marked increase in the FIGURE 6 - By late Eocene times, ca. 42 Ma, widening oceanic gaps had made the pos¬ sibility of shallow-water marine links between the southern lands very remote. Southern seas were beginning to be stirred by marine current systems that were the forerunners of the modern Circum-Antarctic Current, although full development of an integrated system was delayed until the South Tasman Rise had cleared the east Antarctic shelf at ca. 34 Ma (Kennett et al. 1975) and the Drake Passage had opened up between South America and the Antarctic Peninsula at ca. 29 Ma (Barker & Burrell 1977). Nonetheless, sufficient mi¬ gration opportunities were available for ancestral penguins to achieve a broad southern distribution and many other marine groups with good dispersal capabilities, either as adults or juveniles achieved similar distributions by utilising the developing Circum-Antarctic marine currents. 374 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI immigration into New Zealand of Australian plants and animals, both marine and ter¬ restrial. A notable feature of the late Cenozoic was the arrival of many land birds of Australian origin, via storm-generated westerly gales. As a result, the modern New Zealand bird fauna has a strong Australian aspect, generated by multiple colonisation. The Takahe, for example, is derived from an old (Miocene-Pliocene?) immigration of Porphyrio stock and has diverged considerably, whereas the Pukeko, which is indis¬ tinguishable from Australian forms, is evidently from a very recent (Holocene?) migra¬ tion (Fleming 1982). A feature of those birds that became isolated in New Zealand were adaptions to flightlessness and gigantism. The New Zealand environment, without flesh-eating predators, tended to encourage flightlessness and also the development of ground¬ nesting habits. Loss of flight in turn paved the way to giantism, and as the birds be¬ came larger other changes occurred: the feathers grew heavier and sparser, the legs stouter and shorter, and the brain became relatively small in relation to the beak, jaws, and cheek muscles. This was a response to the adoption of a grazing habit. The Takahe, Kakapo, and Extinct Goose (Cnemiornis) are examples of late Cenozoic land FIGURE 7 - Reconstruction of Australia and South-east Asia at ca. 40 Ma (late Eocene). At about this time a major arc-continent collision commenced just north of New Guinea and the new land areas that resulted provided opportunities for the interchange of terrestrial floras and faunas between Australia and southeast Asia. India began to collide with south¬ ern Tibet at about 44 Ma. Figures 7 & 8 are based on Audley-Charles et al. (1988). The modern continental outlines are for reference only. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 375 W. Sulawesi Java Sumatra Banda Arc FIGURE 8 - Reconstruction of Australia and Southeast Asia at ca. 30 Ma (late Oligocene). The continuing northward movement of Australia has reinforced the extent of the South¬ east Asian connections seen in Figure 7. SOUTH-EAST ASIAN LINKS Although there had been links between Australia and South-east Asia extending back into Mesozoic time, they were largely ephemeral, via volcanic islands that appeared and disappeared in concert with waxing and waning volcanic activity (Cox 1990, Audley-Charles et al. 1988, Audley-Charles 1988). Substantial links did not begin to appear until the early Neogene (Figure 7). As Australia moved northwards, the fringes of island arcs, depositional troughs and allochthonous terranes along its northern boundary (that were later to give rise to Papua New Guinea and adjacent islands) began to interact with the southern edge of the Indonesian arc, as Indonesia and southeast Asia moved southwards in response to the opening and continued expan¬ sion of the South China Sea (Audley-Charles 1987, 1988, Audley-Charles et al. 1988, Hutchison, 1989, Pigram & Davies 1987, Truswell et al. 1987). Such interaction soon developed into a major collisional situation that progressively deformed and coalesced the former volcanic arcs, terranes and the contents of depositional troughs. 30° s 60° S 30° N Collision Zone E.Sulawesi N. Guinea 376 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Middle Miocene 16.5-10.5 million years 20* S 30° S 40° S Late Miocene 10.5-5.0 million years FIGURE 9 - As Australia moved northwards the areas affected by aridity progressively increased, as indicated in this diagram by the dot pattern. The growth of aridity is recorded by the changing composition of clay minerals in sediments laid down on the Lord Howe Rise, the clay minerals changing in response to increase in the amount of wind-blown material derived from the growing areas of desert and semi-desert. The drilling sites along the Lord Howe Rise that provided the clay mineral information and dating are indicated by stars. Modified from Stein and Robert (1986, Figure 12). The first indications of earth movements related to contact between Papua New Guinea and the Indonesian/Southeast Asian arc appeared about 40 Ma ago (Figure 7). Then, as a response to major changes in the direction of motion in the Pacific Plate (Fortuin et al. 1988, Hall 1987, Kroenke 1984, Audley-Charles 1987, Audley- Charles et al. 1988), earth movements continued from the late Eocene to middle ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 377 Miocene (Figure 8). As a consequence, a large block of new land was raised, roughly aligned along the site of modern Papua New Guinea, to form a northern edge to the Australian continent (Whitmore 1982) . Later, in the late Miocene and Pliocene, a seaway developed between Papua New Guinea and Australia, but intermittent land connections were renewed during the glacio-eustatic low sea levels of the Plio- Pleistocene. Such connections remained through to the Last Glacial, when a broad plainland existed across large areas of the sites of the modern Arafura Sea and Torres Strait (e.g. Walker 1972, Heatwole, 1987). This land connection was finally broken by the Flandrian sea level rise beginning about 10,000 years ago. THE DESERTIFICATION OF AUSTRALIA The northwards movement of Australia in the late Cenozoic brought it into progres¬ sively drier climatic belts, with the result that the widespread humid vegetation of the early Cenozoic gave way to desert and semi-desert between the late Oligocene/early Miocene (27-24 Ma) and middle Miocene (15 Ma) (Chamley 1986, Kemp 1978, 1981, Truswell & Harris 1982, Truswell 1990, Bowler 1976) (Figure 9). In northern and cen¬ tral Australia levels of aridity progressively built up to reach maxima coinciding with major cooling events in the late Miocene (5 Ma) and late Pliocene (2.5 Ma) (Locker & Martini 1986, 1989, Stein & Robert 1986); and concurrently, climates in southern Australia underwent alternating humid and semi-arid cycles (Stein & Robert 1986, Kennett & von der Borch 1986). Consequently, as large parts of northern and central Australia progressively became dominated by desert and semi-desert, the humid fauna, notably the birds, took refuge in the rain forest areas that still persisted in pock¬ ets along the east coast (Schodde 1982). ACKNOWLEDGEMENTS I wish to thank Dr Dick Schodde for the invitation to contribute this paper to the sym¬ posium “Origins and Evolution of the Australasian Avifauna” and for his valuable edi¬ torial assistance. The paper was reviewed by Dr A. Beu and N. de B. Hornibrook of DSIR Geology & Geophysics, whom I wish to thank for their kind assistance. DSIR Geology & Geophysics provided facilities for the preparation of this paper, the word processing of which was undertaken by Irene Galuszka. The diagrams accompany¬ ing this paper have been prepared by my wife, Diane Stevens, and the inset drawings of fauna and flora were contributed by Ron Brazier, Palaeontological Artist, DSIR Ge¬ ology & Geophysics, Lower Hutt. 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ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 383 THE ORIGIN AND RADIATION OF AUSTRALASIAN BIRDS: PERSPECTIVES FROM THE FOSSIL RECORD WALTER E. BOLES Australian Museum, 6-8 College Street, Sydney, New South Wales 2000, Australia ABSTRACT. The avian fossil record of Australasia is patchy in its temporal and faunistic represen¬ tation, much of it restricted to the Pleistocene. Australia has by far the longest avian record, dating from the Cretaceous, still richest for the Quaternary but growing for the Tertiary. The record is too scanty and uncoordinated between major regional landmasses to trace connections between them, and it says more on the radiations, early diversity and previous distribution than on origins. Few groups have a rich enough record for useful phylogenetic studies. Nonetheless the record is starting to provide crucial documentation for a broader picture of Australasian bird evolution and an interpretation of climatic and environmental changes, and should increasingly provide tests of theories on origins, colonisation routes and divergence times and document changing distributions and early faunal connections. Keywords: Fossil record, Australia, New Zealand, New Caledonia, New Guinea, Australasia, evolu¬ tion, radiation, colonisation, Tertiary, Quaternary. INTRODUCTION This summary is fortunate in following several recent reviews of the avian fossil record in Australasia: New Zealand (Fordyce 1982), New Guinea (Rich & van Tets 1982), New Caledonia (Balouet & Olson 1989) and Australia (Rich & van Tets 1982, Rich & Baird 1986, Baird in press, Rich in press). It presents an overview of the region and updates some aspects raised by these authors, but cannot attempt their level of de¬ tail. References are generally restricted to studies that have appeared since these reviews. For more thorough discussions and references not cited in the text, consult these reviews and Olson (1985). FOSSIL RECORD OF MAJOR REGIONAL LANDMASSES (Figure 1) Australasia has a patchy record, both in temporal and faunistic representation and in palaeontological research. Although richer and longer for Australia than for New Zea¬ land, New Caledonia and New Guinea, the record is poor compared with those of Eu¬ rope and North America. New Zealand The early fossil record of birds in New Zealand is scanty. Most Palaeogene taxa are Sphenisciformes, with scattered seabirds and some as of yet unstudied material. Al¬ though the record extends from at least the Palaeocene, other than for penguins it is sparse and uninformative. Moas are first recorded from Pliocene deposits, but reached their greatest richness in the Pleistocene, when they became a dominant component of the record until the Recent; two families, six genera and 11 species are currently recognised (Cracraft 1976, Worthy 1988 and references therein). 384 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Material from the Late Pleistocene forms the vast majority of the known record, much referable to living taxa. Genera no longer present in New Zealand include a range of anseriforms, some extinct and others living but now extralimital; several large raptors; a large owlet-nightjar; the extinct Apterornithidae, gruiforms of uncertain affinities; and a greater diversity and distribution of the New Zealand wrens (Acanthisittidae) (Fordyce 1982, Millener 1988). New Guinea The only two sites providing avian fossils, one Pliocene, one Pleistocene, have yielded cassowaries Casuarius (Rich & van Tets 1982, Rich et al. 1988). New Caledonia Published deposits are Late Quaternary, mostly Late Holocene, and indicate a greater diversity of taxa than that on the island today. The best known is the large flightless Sylviornis, originally thought to be a ratite, and later considered to be galliform (Poplin FIGURE 1 - Localities producing fossils birds in Australasia. Black triangles, Mesozoic; black circles, Tertiary; white circles, Quaternary. Late Pleistocene and Holocene sites rep¬ resented only by moas (Dinornithiformes) have been omitted. Adapted primarily from Rich and Baird (1986, Figures 1, 2), Fordyce (1982) and Balouet and Olson (1989, Figure 1). ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICi 385 & Mourer-Chauvire 1985, Balouet & Olson 1989), possibly a giant megapode. Balouet & Olson (1989) reported on 32 nonpasserine species, including 11 now extinct. Among the latter were representatives of extant genera previously unknown from New Caledonia, but currently distributed elsewhere, e.g., northern Melanesian islands, smaller islands off New Zealand, and Asia and neighbouring islands. Information on the passerines has not yet been published. Australia Australia has by far the longest avian fossil record in Australasia, dating from the Cretaceous. The Quaternary is richest, both numerically and taxonomically, but the Tertiary record is growing. Cretaceous records consist of a enantiornithine tibiotarsus from Queensland (Molnar 1986) and five small indeterminate feather impressions from Victoria. Avian remains from the extensive gap between 100-22 myBP, when birds were differentiating and southern land masses were connected, lack taxonomic diver¬ sity, are restricted geographically, or are yet unstudied. The Eocene record comprises penguins, which continue into the Oligocene as virtually the only birds of this age. Of the few other remains until the Late Oligocene, the most significant are recent finds from Murgon, southeastern Queensland (54 myBP) that await study. There are only two major intervals with rich, diverse avian records. The first, starting in the Late Oligocene and continuing through the Neogene, is represented in the present Great Artesian Basin (Lake Eyre and Lake Tarkarooloo sub-basins) of cen¬ tral Australia and in northern Australia (Figure 1); western Australian sites have not yet been exploited. Diverse groups of non-marine birds were present in Australia at least as far back as the mid-Tertiary. Because the deposits are fluviatile/lacustrine accumulations, the taxa dominating at almost every site are waterbirds, augmented by some terrestrial, often flightless, forms and scattered volant non-aquatic forms. Smaller birds are repre¬ sented in central Australian sites, but by and large, most preserved pre-Quaternary taxa were medium to large, leading Rich & Baird (1986) to comment “no single locality in Australia in the pre-Pleistocene . . . has produced a diverse avian microfauna.” Important recent finds include the Riversleigh deposits, northwestern Queensland, currently under study by W. Boles. This area is significant because of its array of both water- and forest-dwelling forms, including numerous passerines. The earliest depos¬ its are considered ?Late Oligocene-Early Miocene, with sites known through the Pleistocene; most are Miocene (Archer et al. 1989). The second important fossil-bearing interval consists mainly of Late Quaternary cave and aeolian-fluvial deposits, principally across the southern Australia. The Dromornithidae ( Genyornis ), giant megapodes Progura and some living families now absent from Australia (flamingoes, Phoenicopteridae; Rich & van Tets 1982) persisted until the Pleistocene. Most remains are referable to modern genera or species, whose distributions were often more extensive than at present (e.g. Tasmanian Native-hen Gallinula mortierii ; Baird 1986). Rich & van Tets (1982: Figure 27) exhibited the known fossil record of Australian bird families. Figure 2 shows the extensions in the temporal ranges revealed since, most attributable to finds from Riversleigh. FIGURE 2 - See caption opposite page. 386 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 387 Australia’s Tertiary fossil record suffers from several deficiencies. The different taphonomic forces of the Quaternary have yielded more complete remains than the usually incomplete and disassociated Tertiary material. Differences in the types of preservation and accumulating factors between pre-Quaternary and Quaternary de¬ posits oftens renders them incomparable. Another drawback of the pre-Quaternary record is the paucity of radiometrically dated sites. Most dates have been assigned through tenuous sequences correlating remains between a series of sites (Rich & Baird 1986). TAXONOMIC SCOPE OF THE FOSSIL RECORD The fossil record reveals more on the radiations, early diversity and previous distri¬ butions than it does on the origins of different groups of Australasian birds. Rich & van Tets (1982) considered Australian Neogene taxa to be a mixture of archaic and mod¬ ern forms: (a) possible remnant Palaeogene radiations, (b) representatives of fami¬ lies extant elsewhere, (c) representatives of families now extinct, and (d) more primi¬ tive members of extant families in Australia. The records of these groups vary con¬ siderably, and most suffer from deficiencies that restrict their interpretation. Too many families that are abundant in the modern avifauna have a scanty or non¬ existent fossil record, or one that is restricted to the Pleistocene. Therefore, little can be said about their early history in Australasia (e.g., Columbiformes, Psittaciformes). Other groups have a relatively good Tertiary representation but have yet to receive the necessary attention. In contrast to the well documented anseriforms of Australia’s and New Zealand’s Quaternary, those of the Tertiary still require extensive study, as do passerines from as far back as the Late Oligocene. FIGURE 2 - Geologic ranges of major groups of Australasian birds, modified from Rich and van Tets (1982, Figure 27) for Australia, to which have been added data from New Zea¬ land and New Caledonia. Dashed lines are chronological extensions based on material from Riversleigh, Queensland; dotted lines are extensions of ‘Australasian’ groups from the Phosphorites du Quercy (Mourer-Chauvire 1982, 1989). 1, enanthiornithine birds; 2, Koonwarra feathers; 3, unidentified bones, Murgon, Queensland, Australia; 4, unidentified bones, Waimakariri River, New Zealand (Fordyce 1982); 5, Sphenisciformes (penguins); 6, Diomedeidae (albatrosses); 7, Pelagornithidae (=Odontopterygidae auct) (bony-toothed birds); 8, unidentified bones, Waihao River, New Zealand (Fordyce 1982) ; 9, Dromornithidae (mihirungs) ; 10, Dromaiinae (emus); 11, Podicipedidae (grebes); 12, Pelecanidae (pelicans); 13, Phalacrocoracidae (cormorants); 14 , Anatidae (waterfowl); 15, Accipitridae (hawks, eagles); 16, Rallidae (rails); 17, Burhinidae (stonecurlews); 18, Laridae (gulls); 19 , Palaelodidae ( Palaelodus)-, 20, Phoenicopteridae (flamingoes); 21, Columbidae (pigeons); 22, Aegothelidae (owlet-nightjars); 23, Passeriformes (songbirds); 24, Dinornithiformes (moas); 25, Casuariinae (cassowaries); 26, Ciconiidae (storks); 27, Apterygiidae (kiwis); 28, Procellariiformes (petrels, shearwaters); 29, Anhingidae (darters); 30, Ardeidae (herons); 31, Threskiornithidae (spoonbills, ibis); 32, Falconidae (falcons); 33, Megapodiidae (megapodes); 34, Phasianidae (quail) ; 35, Turnicidae (button-quail); 36, Gruidae (cranes); 37, Rhynochetidae (kagus); 38, Apterornithidae ( Apterornis ); 39, Otididae (bustards); 40, Pedionomidae (plains wanderers); 41, Charadriiformes (waders); 42, Psittaciformes (parrots, cockatoos); 43, Cuculidae (cuckoos); 44, Tytonidae (barn owls); 45, Strigidae (typical owls); 46, Podargidae (frogmouths); 47, Apodidae (swifts); 48, Alcedinidae (kingfishers). 388 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Some groups have temporally restricted records that are well studied but not exten¬ sive enough to allow useful extrapolation beyond those particular fossils, e.g., the pelicans of Australia, Quaternary rails and waterfowl of Australia and New Zealand, the megapodes Progura of Australia, and Sylviornis of New Caledonia. A few families have diverse, well-documented records, which provide information on radiation and diversity of the groups — but no useful clues to their origins. The moas (Dinornithidae, Emeidae) have no doubt the most detailed record of any Australasian group, but it is so restricted in time that it offers little on their early history. Similar comments apply to the Dromornithidae, which have a longer but less representative record, and the Sphenisciformes, which have one of the longest fossil records of any extant group. Of considerable interest are groups that demonstrate early avifaunal connection with other continents: either their earliest record is not Australasian or similarly-aged re¬ mains are known from both areas. Prominent examples of the latter are the flamin¬ goes, known in central Australia from the mid-Tertiary until the Pleistocene, a record paralleled in Africa and the Northern Hemisphere, and the now extinct Palaelodidae (Baird & Rich in press). Three groups considered among the most characteristic of the Australasian region (megapodes, Megapodiidae; frogmouths, Podargidae; and owlet- nightjar, Aegothelidae) have their earliest records in the Eo-Oligocene deposits of France and Germany (Mourer-Chauvire 1982, 1989, Peters this symposium); only the last has pre-Quaternary remains from Australasia (Figure 2). Because Australasia has no Palaeo-Eocene record, the first real occurrence of these groups is open to ques¬ tion and the significance of these European records cannot be evaluated. ORIGINS AND EXTINCTIONS The pre-continental drift concept of Australasian bird origins derived them from five major ‘waves’ of northern emigrants: the earliest colonisations giving rise to the most differentiated taxa, through to the most recent, which are only subspecifically distinct from otherwise Old World taxa (Mayr 1944). Acceptance of continental drift and plate tectonics turned biogeographers efforts to identifying which modern Australasian taxa may have had southern origins (e.g. Cracraft 1972, Rich 1975, Schodde 1982). Evi¬ dence for a group’s Gondwanan origins included its degree of endemism, distribution of its closest relatives, and richness and diversity on each continent. Biochemical techniques identified several higher taxa that may have had their origins in Gondwanaland (reviewed by Christidis, this symposium). Such compilations have been generally based on nonpalaeontological criteria. Olson (1989) attempted to identify such groups based at least in part on the fossil history. He suggested that many groups characteristic of the modern Australasian avifauna may be relictual, as were possibly most avian groups with principally southern distri¬ butions, but nominated five groups that could have had austral beginnings: grebes, ducks, pigeons, parrots and passerines. The Australasian fossil record is too inadequate to be useful for the first four groups. Its passerine record, however, is of comparable age with the oldest known from the Northern Hemisphere (Mourer-Chauvire et al. 1989). One Australian mid-Miocene ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 389 specimen cannot be separated generically from the logrunners Orthonyx (Boles, unpub.), among Australasia’s most distinctive living endemic songbird genera. The fossil record does not deny, even if it does not confirm, the antiquity of the Australa¬ sian land bird fauna and the older endemic radiation in songbirds postulated by cur¬ rent molecular studies. However, it can in some instances test suggested divergence times emerging from these studies. The Australasian avifauna may have had multiple origins, but the portions attributable to dispersal and to vicariance are unclear. To make more significant contributions, the fossil record needs additional material from crucial times, supported by a better un¬ derstanding of avian relationships (Rich & van Tets 1982). As more remains are found and identified, and earlier misidentifications corrected (e.g. van Tets & Rich 1990), the extent of Pleistocene extinctions can be more accurately assessed. Habitat deterioration starting in the Miocene and climatic oscillations of the Pleistocene caused loss of extensive inland lakes in Australia. Waterbirds, particularly flamingoes, and possibly other families, presumably died out as a result. Older bird groups that radiated in the Palaeogene were already showing a post-Tertiary reduc¬ tion in diversity and may have also been terminally affected (e.g. Dromornithidae, Rich & van Tets 1982). Gigantism was prominent during the late Tertiary and Quaternary and large taxa, such as moas, Genyornis, Progura and Sylviornis, appeared foremost among the extinctions. Perhaps this is over emphasized because of the preferential preservation and relative ease of finding large bones and a preference of paleo-or- nithologists for studying them. The growing record of small birds may show that the emphasis on gigantism is somewhat artefactual. The effects of human predation on the megafauna is much debated. In Australia it is suspected of having had a major influence on the extinction of these animals, but there is no direct evidence from middens (however, that dromornithids obviously had the attention of Australia’s pre-European inhabitants is shown by cave paintings dating from 26 000 years BP). In contrast, the detrimental effects of human predation and habitat alteration on moas in New Zealand is well documented (Trotter & McCulloch 1984). Many biogeographic studies have not considered fossil evidence of previous distribu¬ tions. Overlooking evidence of major disruptions to many Pacific island avian commu¬ nities by non-European humans can lead to serious underestimations of species rich¬ ness (Steadman 1989). In Australasia there is evidence of such disruptions in New Zealand and New Caledonia. Historical biogeographic interpretations for taxa not usefully represented in the fossil record can be at best only tentative. CONCLUSIONS What can be said about the fossil record of birds in Australasia? Much is an indica¬ tion of what it lacks. It is too patchy and uncoordinated between the major regional land masses to trace the connections among radiations, although the growing Pleistocene record in New Zealand, New Caledonia and Australia will permit an inter¬ esting comparative picture. Even in Australia, with its longer, richer record, some cru¬ cial times are not represented, although new sites, such as Murgon, will help fill these gaps. 390 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI There are critical geographical areas not represented, such as western Australia and most of New Guinea. There is an overwhelming need for accurately and directly dated sites. The taxonomic composition of the record is biased at certain times and certain localities. The waterbird-dominated record of Australia’s Tertiary needs to be supple¬ mented by non-aquatic taxa. The record of no group is rich enough in its taxonomic and chronological coverage for extensive phylogenetic studies, much less useful speculation on its origins. Despite these deficiencies, the fossil record makes a useful contribution toward the understanding of Australasia’s avifauna. It provides crucial documentation needed for building a broad picture of avian evolution on these southern land masses by docu¬ menting changing distributions and early faunal connections with other continents; questioning or corroborating previous ideas of Australasian endemicity; testing theo¬ ries on origins, colonisation routes and divergence times of various bird groups; and aiding the interpetation of changes in climate and environment. The record is still scanty, but will assume growing importance. Greater accessibility to known and fu¬ ture fossil localities, increased funding for field work and preparation, and the grow¬ ing number of Australasian-based palaeo-ornithologists can not help but fruitfully build on the existing knowledge of the avian fossil record. ACKNOWLEDGEMENTS I am grateful to P.V. Rich for reading and discussing the manuscript and providing me with preprints of relevent papers. LITERATURE CITED ARCHER, M., HAND, S.J., GODTHELP, H., MEGIRIAN, D. 1989. Fossil mammals of Riversleigh, northwestern Queensland: preliminary overview of biostratigraphy, correlation and environmental change. Australian Zoologist 25: 29-65. BAIRD, R.F. 1986. The Pleistocene distribution of the Tasmanian Native Hen, Gallinula mortierii. Emu 84: 119-123. BAIRD, R.F. In press. Avian fossils from the Quaternary of Australia. In Rich, P.V., Baird, R.F., Mona¬ ghan, J., Rich, T.H. (Eds). Vertebrate palaeontology of Australasia. Melbourne, Thomas Nelson. BAIRD, R.F., RICH, P.V. In press. Palaelodus (Aves: Palaelodidae) from the Late Cenozoic of Aus¬ tralia. Natural History Museum of Los Angeles County Contributions in Science. BALOUET, J.C., OLSON, S.L. 1989. Fossil birds from Late Quaternary deposits in New Caledonia. Smithsonian Contributions to Zoology 469: 1-38. CHRISTIDIS, L. This symposium. Biochemical evidence for the origins and evolutionary radiations in Australasian avifauna: the songbirds. CRACRAFT, J. 1976. The species of moa (Aves: Dinornithidae). Smithsonian Contributions to Paleobiology 27: 189-205. FORDYCE, E. 1982. The fossil vertebrate record of New Zealand. Pp. 629-698 in Rich, P.V., Thompson, E.M. (Eds). The fossil vertebrate record of Australasia. Clayton, Monash University Offset Printing Unit. OLSON, S.L. 1985. The fosill record of birds. Pp. 79-238 in Farner, D.S., King, J.R., Parkes, K.C. (Eds). Avian biology 8. New York, Academic Press. OLSON, S.L. 1989. Aspects of global avifaunal dynamics during the Cenozoic. Acta XIX Congressus Internationalis Ornithologici: 2023-2029. PETERS, D.S. These proceedings. Zoogeographical relationships of the Eocene avifauna from Messel (Germany). ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 391 POPLIN, F., MOURER-CHAUVIRE, C. 1985. Syiviornis neocaledoniae (Aves, Galliformes, Megapodiidae), oiseau geant eteint de I’lle des Pins (Nouvelle Caledonie). Geobios 18: 73-97. RICH, P.V. 1975. Antarctic dispersal routes, wandering continents, and the origin of Australia’s non- passeriform continental avifauna. Memoirs of the National Museum of Victoria 36 63-126. RICH, P.V. In press. The Mesozoic and Tertiary history of birds on the Australian plate. In Rich, P.V., Baird, R.F., Monaghan, J., Rich, T.H. (Eds). Vertebrate palaeontology of Australasia. Melbourne, Thomas Nelson. RICH, P.V., BAIRD, R.F. 1986. History of the Australian avifauna. Current Ornithology 4: 97-139. RICH, P.V., PLANE, M.D., SCHROEDER, N. 1988. A pygmy cassowary ( Casuarius lydekkeri) from Late Pleistocene bog deposits at Pureni, Papua New Guinea. BMR Journal of Australian Geology and Geophysics 10: 377-389. RICH, P.V., VAN TETS, G.F. 1982. Fossil birds of Australia and New Guinea: their biogeographic, phylogenetic and biostratigraphic input. Pp. 235-384 In Rich, P.V., Thompson, E.M. (Eds). The fossil vertebrate record of Australasia. Clayton, Monash University Offset Printing Unit. SCHODDE, R. 1980. Origin, adaptation and radiation of birds in arid Australia. Pp. 191-224 in Barker, W.R., Greenslade, P.J.M. (Eds). Evolution of the flora and fauna of arid Australia. Freville, Peacock Publications. STEADMAN, D.W. 1989. Fossil birds and biogeography in Polynesia. Acta XIX Congressus International^ Ornithologici: 1526-1534. TROTTER, M.M., McCULLOCH, B. 1984. Moas, men, and middens. Pp. 708-727 in Martin, P.S., Klein, R.G. (Eds). Quaternary extinctions. Tucson, University of Arizona Press. VAN TETS, G.F., RICH, P.V. 1990. An evaluation of De Vis’ fossil birds. Memoirs of the Queensland Museum 28: 165-168. WORTHY, T.H. 1988. A re-examination of the moa genus Megalapteryx. Notornis 35: 99-108. 392 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI BIOCHEMICAL EVIDENCE FOR THE ORIGINS AND EVOLUTIONARY RADIATIONS IN THE AUSTRALASIAN AVIFAUNA: THE SONGBIRDS LES CHRISTIDIS Department of Ornithology, Museum of Victoria, 71 Victoria Crescent, Abbotsford, 3067, Australia ABSTRACT. DNA-DNA hybridization indicates that all Australasian-centred families of Passeriformes (songbirds) represent an endemic radiation, contrary to their conventional interpretation as diverse derivatives of Eurasian origin. Independent evidence based on protein electrophoresis is presented which corroborates many of the DNA findings but seriously calls into question the supposed links be¬ tween Menuridae (lyrebirds), Ptilonorhynchidae (bowerbirds) and Climacteridae (treecreepers). The lyrebirds are isolated consistently as a sister lineage to the remaining Australasian songbirds. An ap¬ parently autochthonous radiation, confined largely to the Australo-Papuan continental plate, comprises the Acanthizidae (thornbill-warblers), Pardalotidae (pardalotes), Meliphagidae (honeyeaters), Epthianuridae (Australian chats), Eopsaltriidae (Australasian robins), Orthonychidae (logrunners) and, arguably, Maluridae (fairy-wrens). The biochemical results are also compared with data from microcomplement fixation. A Gondwanan origin for the Passeriformes is canvassed. Keywords: Protein electrophoresis, biochemical evolution, phylogeny, passerines, songbirds, cladistic analysis, Australo-Papuan. INTRODUCTION Relationships of the Australo-Papuan songbirds (Passeriformes) have been the centre of much scientific interest following the DNA-DNA hybridization studies of Sibley and Ahlquist (summarized in Sibley et al. 1988). These identified a major dichotomy amongst the oscinine passerines: the Corvida which includes all the major Australo- Papuan lineages, and the Passerida which is centred in Eurasia (Figure d). The Corvida itself comprised three primary lineages: the lyrebirds and allies (Menuroidea), the honeyeaters and allies (Meliphagoidea), and the crows, monarch flycatchers and allies (Corvoidea). The first two of these superfamilies are all but endemic to Australia- New Guinea, and the last, although almost cosmopolitan, is nevertheless most diverse in Australasia. Although the results are exciting, concern has been raised regarding the reliability of the data and the methods of analysis. The lack of complete matrices, assumption of constant rate of molecular change, and limited tree building methodologies have all been points for criticism (see Mayr 1989, Sibley 1989, for recent commentary). One strategy to assess the results objectively is to compare them with those from other sets of data. To this end, the higher order relationships among Australo-Papuan song¬ birds are here examined by protein electrophoresis and, to a lesser extent, by the im¬ munological technique of microcomplement fixation. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 393 METHODS Data from protein electrophoresis and microcomplement fixation were taken from the studies of Christidis and Schodde (1991) and Baverstock et al. (in press), respectively, and compared with the phylogenies derived from DNA-DNA hybridization (Sibley & Ahlquist 1985). In the electrophoretic study, allelic variation at 18 loci was examined by UPGMA, distance Wagner and PAUP procedures in 42 species (38 indigenous, 4 introduced of Australo-Papuan passerines, covering all 14 major elements of the Australo-Papuan-centred families and representatives of two to five families in each of the primary Eurasian assemblages as recognized by DNA-DNA hybridization (Sibley & Ahlquist 1985, Sibley et al. 1988). Here, the protein data are analyzed fur¬ ther by the HENNIG 86 cladistic program (Farris 1988) because it is particularly ef¬ fective in finding the most parsimonious trees (Platnick 1989). In it, alleles were treated as characters and their presence or absence as states. Trees were outgroup rooted by the suboscinine Pitta. The principal procedures used were mhennig* and tread with branch-breaking and successive weighting (Platnick 1989). Microcomplement fixation data is taken from a series of oneway comparisons using one species from each of the Meliphagidae, Epthianuridae, Grallinidae and Climacteridae. Albumin antibody from the four was then compared against albumin antigens obtained from a number of species representing several other Australo- Papuan- and Eurasian centred families. Meliphagidae Acanthizidae Maluridae Eopsaltriidae Orthonychidae Pomatostomidae Corvidae Climacteridae Menuridae Ptilonorhynchidae Muscicapoidea * Sylvioidea * Passeroidea * Meliphagoidea y Corvoidea - Menuroidea >- Corvida r Passerida FIGURE 1 - Relationships of Australo-Papuan passerines as determined by DNA-DNA hybridization (based on Sibley & Ahlquist 1985). Only the 3 superfamilies within the Passerida are depicted and these are indicated by asterisks. 394 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI RESULTS For the further cladistic analysis of electrophoretic data, the mhennig* procedure pro¬ duced the most parsimonious trees; their consensus is depicted in Figure 2. Similar topologies were also produced for the major clades by the tread method. Both pro¬ cedures separate thejyrebird ( Menura ) consistently as a sister group to the remain¬ ing songbirds. Several stepwise polychotomies and dichotomies follow, only a few of which are consistent between the trees derived from the two procedures. One that is ■ Pitta - Menura Monarcha Struthidea □ Corvus Corvus Psophodes i — Rhipidura - ' — Cormobates Daphoenositta — Pachycephala - Cinclosoma - Pomatostomus - Chaetorhynchus □ Sphecotheres Coracina CDicaeum Zosterops - Orthonyx - Acanthiza - Pardalotus □ Melanodryas Microeca ■ Phylidonyris ■ Epthianura j— Philemon - « — Myzomela I — Ailamus -* — Pn.ltnns - ' — Peltops Cracticus - — uracticu - Ptilonorhynchus ■ Ptiloris Nectarinia Anthus r— Passer - * — Acrocephalus ■ Malurus - Hirundo FIGURE 2 - Strict consensus tree derived from an analysis using the HENNIG program (Farris 1988) version 1.5, based on allelles as characters and their presence or absence as states. The mhennig* procedure with branch-breaking and successive weighting was employed. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 395 comprises the meiiphagine honeyeaters ( Myzomela , Phylidonyris, Philemon), Austral¬ ian chats ( Epthianura ), thornbill-warblers ( Acanthiza ), pardalotes ( Pardalotus ), eopsaltriine robins ( Microeca , Melanodryas), logrunners ( Orthonyx ) and fairy-wrens ( Malurus ). A second assemblage encompasses Eurasian and African centred families included in the Passerida by DNA-DNA hybridisation (Nectarinia to Turdus in Figure 2). The Apostlebird, Struthidea, (Corcoracidae) is aligned with them anomalously only in the mhennig* tree. A third smaller assemblage comprising the butcherbirds ( Cracticus ), woodswallows and allies ( Artamus , Peltops), bowerbirds ( Ptilonorhynchus ) and birds of paradise ( Ptiloris ) is identified by the tread procedure only. Microcomplement fixation distances within the Meliphagoidea, as identified in Figure 1, average 17. Those between the Meliphagoidea and both the Corvoidea and Menuroidea are around 25, and the Passerida are still further distant at values of 35 to 55. DISCUSSION The concept of an endemic radiation among the major Australo-Papuan-centred groups of songbirds is supported by protein allelic data and less forcefully by immu¬ nological distance. The Australo-Papuan robins (Eopsaltriidae), whistlers (Pachycephalidae), monarch flycatchers (Monarchidae), thornbill-warblers (Acanthizidae) are only distantly related to their supposed Old World counterparts in the Muscicapidae ( Turdus ) and Sylviidae ( Acrocephalus ). This is entirely consistent with DNA-DNA hybridization data (Sibley & Ahlguist 1985). So too is the separation of eopsaltriine robins from the whistlers, with which they have often been linked (Boles 1979). Moreover the association between the families Acanthizidae, Pardalotidae, Meliphagidae and Epthianuridae (Australian chats), identified by DNA-DNA hybridiza¬ tion studies (Figure 1), is shown also in the allelic and immunological data. The data sets are, however, not congruent regarding the positions of the fairy-wrens (Maluridae), eopsaltriine robins and logrunner ( Orthonyx ) along the Australo-Papuan- centred families. Sibley and Ahlquist (1985) aligned the fairywrens with the Meliphagoidea (along with Acanthizidae and Meliphagidae) and recorded the latter two as outlying lineages of the Corvoidea. On allellic data, the eopsaltriine robins and logrunner are linked to the Acanthizidae and Meliphagidae instead. Their positioning of the fairy-wrens is just as ambivalent. Although the HENNIG analyses associated them with the Eopsaltriidae - Meliphagidae assemblage, this is not confirmed by PAUP and distance-based analyses (Christidis & Schodde 1991). Flowever, microcomplement fixation supports the association between the Maluridae, Meliphagidae and Acanthizidae but data for the Eopsaltriidae is lacking. The remaining Australo-Papuan songbirds included in the Corvoidea of Sibley and Ahlquist (1985) appear as an assemblage in the UPGMA and PAUP analysis of Christidis and Schodde (in press) but not in any of the protein-based FIENNIG trees. There are also some differences between the DNA-based and protein-based topologies, particularly in the positions of Cormobates (Climacteridae), Struthidea (Corocoracidae), Pomatostomus (Pomatostomatidae), and Ptilonorhynchus (Ptilonorhynchidae). Nevetheless, both DNA-DNA hybridisation and allellic data align Oriolidae ( Specotheres ) with Campephagidae ( Coracina ) and Artamidae ( Artamus ) with Cracticidae ( Cracticus , Peltops). Moreover, microcomplement fixation results 396 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI confirm that the Magpie-lark Grallina is a monarch flycatcher (Monarchidae), distant from Struthidea and Corcorax (Baverstock et al. in press). Although the Eurasian-centred families included in the Passerida of Sibley and Ahlquist (1985) appear to be closer to one another than to any Australo-Papuan-cen- tred families, the clear cut separation between the Passerida (Eurasian centred) and Corvida (Australo-Papuan centred) is not as clear cut in the protein data. Neverthe¬ less, microcomplement fixation data (Baverstock et al. in press) do to a limited extent support the separation between the Corvida and Pas-serida. One significant area of discord between the allellic and DNA data sets involves the relationships of the lyrebirds (Menuridae), treecreepers (Climacteridae) and bowerbirds (Ptilonorhynchidae). The present protein data do not support their inclu¬ sion as a single superfamily as found by Sibley and Ahlquist (1985). Instead, Ptilonorhynchus (bowerbird) consistently clusters with the Cracticidae (butcherbirds), Artamidae (woodswallows) and Paradisaeidae (birds of paradise). Such a result is more consistent with morphological data (Bock 1963). Cormobates (treecreeper) has no obvious relatives, while Menura (lyrebird) is always isolated as a sister lineage to the remaining songbirds. Moreover, microcomplement fixation data also fails to align the treecreepers with the lyrebirds. Despite conflicting evidence of relationships among these last three families, there is enough agreement between the DNA-DNA hybridization, protein and limited immuno¬ logical data sets to substantiate the hypothesis of an endemic radiation amongst the Australo-Papuan passerines. Moreover, the Corvida can be separated into four as¬ semblages. The first comprises the Acanthizidae, Paradalotidae, Meliphagidae, Epthianuridae, Eopsaltriidae, Orthonychidae and Maluridae. Two further assemblages are monofamilial and small: Menuridae and Climacteridae. These three assemblages are, with few exceptions, confined to the Australo-Papuan continental plate and can therefore be considered autochthonous. Judged by genetic distances (Christidis & Schodde in press) they appear to be descendants of the oldest branches of the song¬ bird radiation. The final assemblage is equivalent to the Corvoidea (Sibley & Ahlquist 1985) including the Ptilonorhynchidae but excluding Eopsaltriidae and Orthonychidae. These families are more widespread and can be considered a link between the first three assemblages and the Passerida. Genetic distance data (Christidis & Schodde 1991) suggests that they, along with the Passerida, are of a more recent origin. The framework outlined would argue against a Eurasian origin for the songbirds as a whole (cf. Wilson 1988, Mayr 1989). The meliphagine - eopsaltriine group along with the Climacteridae and Menuridae do not have links with the Passerida and appear to represent an early songbird radiation in Australasia. Of particular significance is the possible separation of the Menuridae as a distinct sister group to the remaining oscines. This is consistent with the conclusions of Feduccia and Olson (1982) based on morphological analyses. They suggest that the Menuridae represent the most primitive oscine lineage with links to the primitive South American suboscines, Rhinocryptidae. Such connections suggest that the relationships of the sub-oscinine New Zealand wrens (Acanthisittidae) also need to be re-examined by protein electrophoresis and other techniques (cf. Raikow 1987). From the divergences nearly uncovered among the passerines, a Gondwanan origin for the Passeriformes now becomes a plausibe hypothesis with the order splitting ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 397 there in two major radiations: a sub-oscinine radiation in western Gondwana (South America) and a primary oscinine radiation in eastern Gondwana (proto-Australasia). The ancestors of the Passerida could then have spread into Eurasia and Africa where they underwent subsequent radiations. It is unclear as to whether the Corvoidea (as defined here) underwent their primary radiation in Australasia and then spread into Asia and Africa, or first radiated in Asia and then re-spread into Australasia. Support for a southern origin for the Passeriformes can also be found in the fossil record. Olson (1988) reports that passerines do not appear in the Northern hemisphere un¬ til the latest Oligocene and only become common in the Miocene but with no indica¬ tion of an evolutionary origin. ACKNOWLEDGMENTS The author thanks F. Knight (CSIRO) for the illustrations and T. Galariniotis (Museum of Victoria) for typing the manuscript, LITERATURE CITED BAVERSTOCK, P.R., SCHODDE, R., CHRISTIDIS, L„ KRIEG, M., SHEEDY, C. In press. Evolution¬ ary relationships of the Australian mud-nesters (Grallinidae, Corcoracidae): immunological and osteo- logical evidence. Proceedings XX International Ornithological Congress. BOCK, W.J. 1963. Relationships between the birds of paradise and the bowerbirds. Condor 65: 91-125. BOLES, W.E. 1979. The relationships of the Australo-Papuan flycatchers. Emu 79: 107-1 10. CHRISTIDIS, L., SCHODDE, R. 1991. Relationships of Australo-Papuan songbirds (Aves: Passeriformes)- Protein Evidence. Ibis 133. FARRIS, J.S. 1988. Hennig 86 Reference, Version 1.5. Available from J.S. Farris (41 Admiral Street, Post Jefferson Station, New York). FEDUCCIA, A., OLSON, S L. 1982 Morphological similarities between the Menurae and Rhinocryptidae, relict passerine birds of the Southern Hemisphere. Smithson. Contrib. Zool. 366:1-22. MAYR, E. 1989. A new classification of the living birds of the world. Auk 106: 508-512. OLSON, S.L. 1988. Aspects of global avifaunal dynamics during the Cenozoic. Pp. 2023-2029 in Ouellet, H. (Ed.). Acta XIX Congressus Internationalis Ornithologici. Ottawa: University of Ottawa press. PLATNICK, N.l. 1989. An empirical comparison of microcomputer parsimony programs, II. Cladistics 5: 145-161. RAIKOW, R.J. 1987. Hindlimb myology and evolution of the Old World suboscine passerine birds Acanthisittidae, Pittidae, Philepittidae, Eurylaimidae). Ornithological Monographs 41. Washington: American Ornithologists Union. 81 p. SIBLEY, C.G., 1989. Response to E. Mayr. Auk 106: 512-515. SIBLEY, C.G., AHLQUIST, J.E. 1985. The phylogeny and classification of the Australo-Papuan pas¬ serine birds. Emu 85: 1-14. SIBLEY, C.G., AHLQUIST, J.E., MONROE, B.L. 1988. A classification of the living birds of the world based on DNA-DNA hybridization studies. Auk 105: 409-423. WILSON, A.C. 1988. Time scales for bird evolution. Pp. 1912-1917 in Ouellet, H. (Ed.). Acta XIX Congressus Internationalis Ornithologici Ottawa: University of Ottawa Press. 398 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI THE EVOLUTIONARY HISTORY OF PARROTS AND COCKATOOS: A MODEL FOR EVOLUTION IN THE AUSTRALASIAN AVIFAUNA DOMINIQUE HOMBERGER Department of Zoology & Physiology, Louisiana State University, Baton Rouge, LA 70803, USA ABSTRACT. The Psittaciformes represent a well-defined avian order that originated probably in the Australasian sector of Gondwana and which has had a long and independent evolutionary history lead¬ ing to its present pan-austral distribution. Despite their unity in several fundamental characteristics, the Psittaciformes exhibit a great diversity of size, plumage, biology, ecology, and both external and in¬ ternal morphology. This diversity has been explored from many sides in systematic studies through the application of a great variety of techniques and approaches. As a result, the systematics of the Psittaciformes has acquired the potential to provide the most corroborated classification of any Aus¬ tralasian avian order. Through the review and synthesis of data from the latest eco-geographical, func¬ tional-morphological and biochemical studies of parrots and cockatoos, the evolutionary history of the Psittaciformes will be constructed, both for its own sake and as a model for interpreting the evolution of other avian groups in the Australasian avifauna. Keywords: Parrots, cockatoos, functional morphology, ecological morphology, ecological biogeogra¬ phy, biochemical systematics, Australasia, phylogenetic reconstruction. INTRODUCTION The Psittaciformes, which form the third-largest non-passerine avian order and have a broadly pan-austral distribution, display their greatest morphological and ecological diversity in Australasia. Representing an old avian lineage, they have demonstrated a remarkable propensity towards adapting to widely differing ecological conditions, while at the same time retaining a set of ancestral psittaciform characters. Hence, Psittaciformes form a clearly defined avian order that has (a) no marginal taxa that could suggest phylogenetic affinities to other avian orders; (b) numerous taxa that are relicts of early radiations, and the genealogical relationships of which have become obscured; and (c) many taxa resembling one another because of convergently ac¬ quired characters in response to similar environmental conditions. In recent years, parrot systematics has shifted in focus from classifications based on similarities in standard characters as indicators of genealogical relationships towards the reconstruction of the evolutionary history. This reorientation was made possible by new developments in a variety of fields. Comparative functional morphology has provided a technique for distinguishing convergent characters from homologous ones and ancestral characters from derived states, thereby allowing the reconstruction of the evolutionary history of various characters (e.g., Glenny 1955, Homberger 1980, 1990, Guntert 1981, Homberger & Schodde in prep.). Biochemical systematics has generated new data for testing earlier hypotheses (e.g., Adams et al. 1984, Christidis et al. in press). Comparative ecological morphology of the feeding adaptations of parrots has also generated data that elucidate the nature of the linkage between the morphology of parrots and the ecology of their environment (e.g., Saunders 1974, Homberger 1990, unpubl. obs., Homberger & Schodde in prep.). Advances in the his¬ torical and ecological biogeography of Australasia, in their turn, have provided data ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 399 on the history of environmental changes and of associated changes in the geographi¬ cal distribution of Australasian parrots (e.g., Schodde & Calaby 1972, Schodde 1982). In the following, three Australasian psittaciform taxa are selected as examples to il¬ lustrate how a synthesis of such data can improve our understanding of the evolution¬ ary history of an avian order. RESULTS The Australasian true parrots The true parrots (Psittacinae sensu Homberger 1980) of Australasia are usually di¬ vided into the platycercine (broad-tailed) and psittaculine (red-billed) parrots (e.g., Smith 1975; Homberger 1980). Four genera, however, are not easily classified as they show different taxonomic affinities according to the characters considered. These genera are Alisterus (king parrots), Aprosmictus (red-winged parrots), Polytelis (long¬ tailed parrots), and Prosopeia (shining parrots) (Homberger 1980). Homberger (1980) provides a set of data and testable hypotheses as a basis for com¬ parison with other studies. Based on functional-morphological characters of the feed¬ ing apparatus, Alisterus, Aprosmictus, and Polytelis were assigned to the psittaculine parrots. Furthermore, these three genera were found to share not only a unique lin¬ gual surface with Eclectus and some species of Tanygnathus (great-billed parrots) but also a derived condition of the soft palate with Eclectus, Tanygnathus, Geoffroyus, and Psittacula (ring-necked parrots). Thus, the Australo-Papuan genera Eclectus, Alisterus, and Aprosmictus were interpreted as having evolved from an oriental psittaculine stem and to have given rise, via an Aprosmictus- 1 ike ancestor, to the Australian Polytelis, of which Alexandra’s Parrot P. alexandrae displays several uniquely derived characters. In contrast, Prosopeia was assigned to the platycercine parrots on the basis of functional-morphological characters of the feeding apparatus, and was shown to share numerous lingual and palatal characters with Platycercus (rosellas), Cyanoramphus (island parakeets), and Eunymphicus (horned parakeets). Because of the reduced surface structure of its hard palate, Prosopeia was interpreted as the Fijian end point of an evolutionary line that led from the Australian Platycercus via the Pacific Cyanoramphus to the New Caledonian Eunymphicus. Guntert’s (1981) data on the morphology of the proventriculus support the distinction between the psittaculine and platycercine parrots, but also show that Prosopeia has the most ancestral condition among the platycercine parrots. Like Geoffroyus, Tanygnathus, Eclectus, Alisterus, and Aprosmictus, it has also a very long intestine, which is an ancestral character within the Psittaciformes (Guntert 1981). Glenny’s (1955) study on the aortic arch system shows that the A-1 carotid pattern, which is found in the above psittaculine parrots, is the ancestral condition, whereas the A-2-s carotid pattern, which is found in the above platycercine parrots, is derived. The electrophoretic data by Christidis et al. (in press) support the close relationship among the platycercine parrots (in particular between Platycercus and Cyanoramphus) and between the psittaculine Eclectus and Geoffroyus. However, they depart from the data of the three previous studies by suggesting that Alisterus and Polytelis - as well as Psittacella (tiger-parrots) which is conventionally grouped with the psittaculine parrots - are more closely related to the platycercine parrots than to the psittaculine ones. 400 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI The biogeographical studies by Schodde & Calaby (1972) and Schodde (1982) show that birds with a present-day Tumbunan distribution in the rainforests of montane New Guinea and coastal eastern Australia are remnants of an avifauna that lived and di¬ versified in subtropical rainforests that covered most of Australia in mid-Tertiary times. This Tumbunan avifauna has been a reservoir for several more derived avifaunae: the Irian avifauna of the rainforests of lowland New Guinea and Cape York Peninsula, and the northern Torresian and southern Bassian avifaunae of wet sclerophyll forests and woodlands in Australia. The Eyrean avifauna of arid central Australia developed largely out of the Bassian avifauna, especially its western part. None of the above sets of data is complete and none of the resulting interpretations is final. But a synthesis of all of the above data can nevertheless provide an evolu¬ tionary scenario that may well bring us a step or two closer to the true phytogeny of parrots. Alisterus, having a Tumbunan distribution, is probably a descendant of a Tertiary rainforest-dwelling psittaculine ancestor from which it retained the ancestral A-1 ca¬ rotid pattern and long intestine, as well as the already derived lingual surface pattern and soft palate configuration. From such an ancestor, Eclectus is likely to have evolved by entering the Irian rainforests and by changing little, except its plumage. Geoffroyus, Tanygnathus, and Psittacula probably branched off the psittaculine stem separately from one another and before Eclectus did, as is suggested by their vari¬ ous ancestral features in the feeding and digestive systems. Psittacula has had the longest independent evolution as is suggested by electrophoretic data and the derived characters in its digestive tract. An Alisterus- like ancestor also gave rise to two other lineages: Aprosmictus in the Torresian north and Polytelis in the Bassian south (Schodde 1982). In its turn, the latter gave rise to the Eyrean Polytelis alexandrae (Schodde 1982) which displays many adaptations to its arid environment (Homberger 1980, Guntert 1981). That the present-day platycercine parrots may also have evolved from a Tertiary, rain¬ forest-dwelling ancestor, which had not diverged much yet from the psittaculine an¬ cestor, is suggested not only by the electrophoretic data which postulate a closer re¬ lationship between Alisterus and the platycercine parrots - but also by Psittacella. This Tumbunan genus has an ancestral A-1 carotid pattern (Smith 1975) and is presently restricted to montane New Guinea. Its platycercine affinities are suggested not only by electrophoretic data but also by its plumage (Guntert 1981, Schodde in lift. 1988, Christidis et al. in press). The only other platycercine parrots with an A-1 carotid pat¬ tern are the Australian genera Melopsittacus (Budgerigar), Neopsephotus (Bourke’s Parrot) and Neophema (grass parrots), which are thought to have branched off the main platycercine stem at an early stage of its evolution and adapted to arid condi¬ tions (Homberger 1980, Schodde 1982, Christidis et al. in press). The rainforest-dwelling platycercine ancestor then acquired the derived characters of the feeding apparatus that are shared by its present-day descendants (see below), but must have had retained a long intestine. The blue-cheeked rosellas (i.e., Platycercus elegans and P. caledonicus) in the rainforests and woodlands of Eastern Australia and Tasmania, respectively, represent one branch that may not have changed much from the ancestral stem, though it acquired a shorter intestine and also gave rise to sev¬ eral more arid-adapted species (Schodde 1982). Cyanoramphus is another branch ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 401 which may have changed little as it spread over the islands of the Tasman Sea and Pacific Ocean, though it also acquired a short intestine and a reduced surface pattern on the hard palate (Homberger 1980, Guntert 1981, Christidis et al. in press). The New Caledonian Eunymphicus may have evolved from a Cyanoramphus like ances¬ tor, from which it diverged little apart from a further reduction in the surface pattern of its hard palate and some modification to its plumage (Homberger 1980, Guntert 1981, Rinke 1989a), even so, a direct derivation from a rainforest-dwelling platycercine stem cannot be discounted at this point. Prosopeia is unlikely to have evolved directly from an Eunymphicus-Wke or Cyanoramphus- like ancestor (cf. Homberger 1980, Rinke 1989a). Because of its long intestine and more primitive proventriculus (Guntert 1981), it must have branched off an early Cyanoramphus or Eunymphicus stock that still had retained these ancestral characters or have arisen directly from the rainforest-dwelling platycercine stem. In either case, Prosopeia seems to have changed little from its immediate ancestor; but, given that the Fijian avifauna is derived from Papuasia (see Rinke 1989a), the direct derivation of Prosopeia from a northern Australo-Papuan rainforest-dwelling platycercine stem would make it less of a zoogeographical puzzle. Its plumage char¬ acters, which indicate a temperate ancestry (Rinke 1989b), are inconclusive. The Australasian cockatoos Cockatoos form a distinct group (Cacatuidae) within the Psittaciformes and are re¬ stricted to Australasia (e.g., Smith 1976, Homberger 1980, Adams et al. 1984), but their intrafamilial relationships have been little understood until recently. Glenny (1955, in lift. 1990) showed that the ancestral A-1 carotid pattern is found in the black cockatoos ( Calyptorhynchus spp.), the Palm Cockatoo ( Probosciger aterrimus), the Gang-gang Cockatoo ( Callocephalon fimbriatum), the Galah ( Eolophus roseicapillus), and the Cockatiel ( Leptolophus hollandicus), whereas the derived unicarotid B pattern is found in all other species ( Cacatua spp.). Dyck (1977) also found that cockatoo feathers lack a spongy ultrastructure and, therefore, lack blue, green and purple colours. The electrophoretic data of Adams et al. (1984) confirmed the monophyly of the Calyptorhynchus species and identified Leptolophus as an early offshoot from the cockatoo stem, but were unable to resolve the phylogenetic positions of Callocephalon, Eolophus, Cacatua galerita (Sulphur-crested Cockatoo), and Cacatua leadbeateri (Pink Cockatoo). On the basis of skull characters, Baird (1985) was able to separate Cacatua from Calyptorhynchus and Callocephalon. The study of skull characters by Homberger & Schodde (in prep.) confirmed this division, but also revealed similarities between Calyptorhynchus and Leptolophus, between Callocephalon and Eolophus, between Callocephalon and the red-tailed black-cockatoos ( Calyptorhynchus banksii and C. lathami), and between Calyptorhynchus banksii and Probosciger. My own eco-morphological data (Homberger 1990, unpubl. obs.) reveal that Callocephalon and most species of Calyptorhynchus (except C. banksii samueli) dis¬ play at least four of the five following ancestral psittaciform characters: arboreality, a pincer-like bill, a lack of filing ridges on the inner surface of the upper bill tip, poor 402 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI seed-shelling ability, and a diet of significant proportions of wood-boring or gall-form¬ ing insect larvae. In contrast, all the other cacatuid genera, as well as Calyptorhynchus banksii samueli, are characterized by the following derived, but of¬ ten convergently acquired characters: filing ridges and a transversal step on the in¬ ner surface of the upper bill tip, a projecting upper bill tip, the use of the lower man¬ dible as the instrumental part of the jaw apparatus, an excellent seed-shelling ability, and a diet of mainly seeds. A synthesis of the above data with biogeographical information suggests the follow¬ ing evolutionary history of cockatoos. Calyptorhynchus, Callocephalon, and Probosciger ev olved probably from a Tertiary, subtropical rainforest-dwelling Calyptorhynchus-Wke ancestor which had retained sev¬ eral proto-psittaciform characters, such as the A-1 carotid pattern, a lack of blue-green feather colours and a pincer-like bill that was used primarily to extract wood-boring or gall-forming insect-larvae. Today, no cacatuid descendants survive in subtropical rainforests. Calyptorhynchus and Callocephalon may have changed little, except in the morphology of their feeding apparatus, the present diversity of which is the result of different adaptations towards seed predation. Most species entered and remained tied to the Bassian forests and woodlands, except for some subspecies of Calyptorhynchus banksii, which entered the Torresian woodlands, and for Calyptorhynchus banksii samueli, which adapted to the arid Eyrean region by becom¬ ing more terrestrial and granivorous. Probosciger aterrimus, which has an Irian distribution, is probably a descendant of a Papuan population of the dark plumaged ancestral calyptorhynchid stem and has become a specialized seed-predator of large tree fruits. The Eyrean Leptolophus is probably derived from an early Calyptorhynchus-Wke ancestor, while Eolophus and Cacatua leadbeateri are more likely to have evolved from a Callocephalon-Wke ances¬ tor, though independently from each other. The origins of the typical white cockatoos are still unclear, but they comprise two separate lineages: The black-billed white cockatoos (i.e., Cacatua galerita, C. sulphurea, C. moluccensis, C. alba) and the white-billed cockatoos or corellas (e.g., Cacatua sanguinea, etc.). All these lineages probably have acquired their seed-shelling specializations independently of each other. Eolophus and the Australian representatives of the corellas and of Cacatua galerita have become adapted secondarily to exploit nutritious plant parts in or on the ground as well. Thus, the present diversity in cockatoos comprises largely the remnants of separate, early radiations from a central pool of forest-dwelling Calyptorhynchus- 1 ike ancestors towards more granivorous, arid-adapted, and terrestrial forms. The Papuan Pesquet’s Parrot ( Psittrichas fulgidus) The Pesquet’s Parrot has diverged so far from all other Psittaciformes that its origin and phylogenetic relationships have become obscure. It clearly evolved from a seed¬ shelling ancestor, but has become frugivorous and has acquired the derived A-2-s ca¬ rotid pattern (Glenny 1955, Homberger 1980, Guntert 1981). Because its feathers lack a spongy ultrastructure, Dyck (1977) suggested that Psittrichas and the cockatoos may have evolved from a common ancestor that had diverged from the main psittaciform stem before the evolution of a spongy ultrastructure in the feathers. Yet Psittrichas does not resemble the cockatoos in any way except in plumage, so that ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 403 its ancestor had probably already diverged from the Tertiary rainforest-dwelling an¬ cestor that gave rise to the cockatoos. This fits its present distribution, for Psittrichas occurs in the hill rainforests in New Guinea where many mid-Tertiary Tumbunan rel¬ icts survive today. CONCLUSIONS This reconstruction of the phylogeny of three Australasian psittaciform taxa suggests that the Psittaciformes radiated out of the subtropical rainforests of the Australasian sector of Gondwana. It also supports a well-established hypothesis that parrots evolved from an arboreal ancestor, because they are characterized by a zygodactylous foot and white egg shells, and because they usually nest in tree holes. Given the great difficulties and possible pitfalls inherent in any systematic study, but especially in a study of an old group with a poor fossil record, systematics will have to depend increasingly on collaborative, multi-disciplinary and non-dogmatic ap¬ proaches rather than on the use of single character complexes, techniques or meth¬ ods to unravel the evolutionary history of birds. LITERATURE CITED ADAMS, M., BAVERSTOCK, P.R., SAUNDERS, D.A., SCHODDE, R., SMITH, G.T. 1984. Biochemi¬ cal systematics and the Australian cockatoos (Psittaciformes: Cacatuinae). Australian Journal of Zo¬ ology 32: 363-377. BAIRD, R.R. 1985. Avian fossils from Quaternary deposits in “Green Waterhole Cave”, south-eastern South Australia. Records of the Australian Museum 37: 353-370. CHRISTIDIS, L., SCHODDE, R., SHAW, D.D., MAYNES, S.F. In press. Biochemical systematics within the Australo-Papuan parrots, lorikeets and cockatoos (Aves: Psittaciformes). DYCK, J. 1977. Feather ultrastructure of Pesquet’s Parrot Psittrichas fulgidus. Ibis 119: 364-366. GLENNY, F.H. 1955. Modifications of patterns in the aortic arch system of birds and their phylogenetic significance. Proceedings of the United States National Museum 104: 525-621. GUNTERT, M. 1981. Morphologische Untersuchungen zur adaptiven Radiation des Verdauungstraktes bei Papageien (Psittaci). Zoologische Jahrbucher, Abteilung fur Anatomie und Ontogenie der Tiere 106: 471-526. HOMBERGER, D.G. 1980. Funktionell-morphologische Untersuchungen zur Radiation der Ernahrungs- und Trinkmethoden der Papageien (Psittaci). Bonner zoologische Monographien, No. 13. HOMBERGER, D.G. 1990. Filing ridges and transversal step of the maxillary rhamphotheca in Aus¬ tralian cockatoos (Psittaciformes: Cacatuidae): A homoplastic structural character evolved in adapta¬ tion to seed shelling. Pp. 43-48 in van den Elzen, R., Schuchmann, K.-L., Schmidt-Koenig, K. (Eds). Proceedings of the International 100th Deutsche Ornithologen- Gesellschaft meeting: Current topics in avian biology. Bonn, Deutsche Ornithologen-Gesellschaft. HOMBERGER, D.G., SCHODDE, R. In prep. Taxonomic relationships within the cockatoos. In Saunders, D. (Ed.). Australian cockatoos: Their biology in the wild and in captivity. RINKE, D. 1989a. The relationships and taxonomy of the Fijian parrot genus Prosopeia. Bulletin of the British Ornithologists’ Club 109: 185-195. RINKE, D. 1989b. The reproductive biology of the Red Shining Parrot Prosopeia tabuensis on the is¬ land of ‘Eua, Kingdom of Tonga. The Ibis 131 : 238-249. SAUNDERS, D.A. 1974. Subspeciation in the White-tailed Black Cockatoo, Calyptorhynchus baudinii, in Western Australia. Australian Wildlife Research 1: 55-69. SCHODDE, R., CALABY, J.H. 1972. The biogeography of the Australo-Papuan bird and mammal fau¬ nas in relation to Torres Strait. Pp. 257-300 in Walker, D. (Ed.). Bridge and barrier: The natural and cultural history of Torres Strait. Canberra, Australian National University. SCHODDE, R. 1982. Origin, adaptation and evolution of birds in arid Australia. Pp. 191-224 in Barker, W.R., Greenslade, P.J.M. (Eds). Frewville, South Australia, Peacock Publications. SMITH, G.A. 1975. Systematics of parrots. Ibis 1 17: 18-68. 404 ACTA XX CONGRESSUS INTERN ATIONALIS ORNITHOLOGICI THE DEVELOPMENT OF MODERN AVIFAUNULAS RICHARD SCHODDE and DANIEL P. FAITH CSIRO Division of Wildlife and Ecology, P.O. Box 84, Lyneham, ACT 2602, Australia ABSTRACT. The development of the Australasian avifauna until its modern speciation is explored by area cladistic analysis and phylogenetic relationships among marker taxa. Eight basic avifaunal ele¬ ments are identified, all autochthonous except for the New Guinean alpine element. Comparison of the present structure and distribution of these elements against regional palaeogeography leads to the fol¬ lowing reconstructions. The minor New Zealand and New Caledonian elements comprise a limited Gondwanan base, augmented by extensive colonization from Australia-New Guinea. Australia and New Guinea share a common avifaunal history rooted in a subtropical rainforest (Tumbunan) faunula wide¬ spread in Australia through the mid Tertiary. The dessication of Australia and raising of the New Guinean cordillera at the close of the Tertiary caused the Tumbunan faunula to withdraw to east coastal and montane refuges and spurred radiation of its elements adapting to tropical rainforest and scleromorphic vegetation. Keywords: Australasian avifauna, causal biogeograhy, cladistic analysis, phylogenetic radiation, faunal elements, palaeogeography. INTRODUCTION Core Australasia comprises the east Gondwanan fragments of Australia, New Zea¬ land, New Guinea and New Caledonia. The bird faunas of these fragments have much more in common with one another than with those of any other continent: a suite of closely allied ratites ( Dromaius , Casuarius, Apteryx), a diversity of parrots and cocka¬ toos, primitive songbirds linked to the oscines ( Menura , Atrichornis, Xenicus ), and a dominance of honeyeaters, monarchine flycatchers and acanthizid warblers among passerines. It is this distinctive structure that binds the Australasian avifauna into a single regional unit, one formed from the same sources. From its origins to its speciation today, this avifauna has undergone long and involved radiations, invasions and extinctions, the courses of which have never been traced in any comprehensive way. Instead, ornithogeographers have relied on presumed source stocks and recent speciation for describing regional bird geography (Mayr 1944, Falla 1953, Kikkawa & Pearse 1969, Diamond 1972, 1985, Cracraft 1973, 1986, Bull & Whitaker 1975, Keast 1981, Pratt 1982, Ford 1987). What developed in be¬ tween, from the earlier mid Tertiary? What were the regional elements from which the avifauna of today radiated? Where do they survive? When and how did they diverge from one another? These are questions of pattern which, both spatial and temporal, need resolution before the processes assembling the present Australasian avifauna can be appreciated fully. METHODS In the absence of a comprehensive fossil record (Boles, this symposium), clues for tracing faunal development come from residual zoogeographical pattern. Two ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 405 FIGURE 1A - Regional habitats sampled for avifaunal comparison: 1-2, New Zealand sub¬ tropical rainforest and stream-sides; 3-4, New Zealand alpine moor; 5, New Caledonian rainforest and maquis; 6, New Caledonian paperbark ( Melaleuca ) savannah; 7-8, New Guinean alpine moor; 9-11, New Guinean montane rainforest above 1500 m a.s.l.; 12-16, New Guinean lowland rainforest below 500m a.s.l.; 17-18, New Guinean swamp grassland; 19-20, New Guinean tropical eucalypt woodland; 21-23, Australian tropical rainforest (in¬ cluding mangroves); 24-26, Australian subtropical rainforest; 27-32, Australian tropical eucalypt woodland; 33-38, Australian temperate/tableland eucalypt forest and heath; 39- 41, Australian mallee; 42-44, Australian desert ranges/basins. approaches to determine it are combined here. One assesses biogeographic congru¬ ence among faunal assemblages; the other unravels phylogenies of taxa on a geo¬ graphic base. Our assessment of assemblages attempts to identify the major avifaunal elements in Australasia. That of phylogeographic radiation in taxa is extrapolated to explore how those elements are related to one another. Our regional assemblages are drawn from the regional ornithogeographies and check¬ lists of Kinsky (1970) and Bull and Whitaker (1975) for New Zealand, Delacour (1966) for New Caledonia, Pratt (1982) and Beehler and Finch (1985) for New Guinea, and Condon (1975), Schodde (1975, 1982a) and Ford (1987 and references therein) for Australia. To avoid the typological constraints identified by Vuiileumier (1975), the assemblages are habitat-based and limited to species that breed in situ and occur on 406 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI BASIC AVIFAUNAL ELEMENTS Australo-Papuan subtropical-montane rainforest (Tumbunan) Australo-Papuan tropical-lowland rainforest (Irian) New Guinea alpine Australo-Papuan tropical eucalypt woodland (Torresian) Australo-Papuan temperate eucalypt forest-heath (Bassian) Australian arid-mallee eucalypt formations (Eyrean) New Zealand New Caledonian FIGURE IB - Basic avifaunal elements of Australasia and their present distribution. land and the land edge of freshwater. Only major natural habitats were used for de¬ limiting assemblages, eight being identified as representative for Australasia : sub- tropical-montane rainforest, tropical lowland rainforest, alpine moor, tropical eucalypt woodland, temperate-montane eucalypt forest-heath, eucalypt mallee, swamp grass¬ land, and desert formations. Component taxa were then scored for each habitat in 44 areas throughout the region to ensure that all significant centres of endemism were covered and that habitats were replicated regionally (Figure 1, caption). Genera and species were examined separately to add a temporal dimension. In all, 338 genera and 918 species were included, covering 96% and 94% respectively of Australasian breeding land and freshwater birds. To assess the relationships among regional assemblages, the component taxa from all sample areas were compared by two hierarchical procedures, one phenetic, the other cladistic. The phenetic program, UPGMA (as in PATN, Belbin 1987), employed Manhattan distances; in clustering areas according to overall similarity, it is biogeographically neutral insofar as it does not weight vicariance and dispersal over one another. The two cladistic packages used, PAUP version 3.0 (Swofford 1990) and Hennig 86 version 1.5 (Farris 1988), stress shared-presence of taxa in grouping ar¬ eas and employ parsimony to discriminate among vicariant explanations for the data. Because there is at present no indisputable outgroup for the regional assemblages, the area cladograms produced were not rooted. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 407 As a result, the cladograms provide no firm indication of ancestry and derivation among faunal assemblages. For tracing this, we drew evidence from phylogenetic case histories published for autochthonous marker taxa that have speciated in all or most of the presumed avifaunal elements in Australasia. RESULTS Although the UPGMA phenogram for genera grouped the 44 assemblages into clus¬ ters largely consistent with previous descriptive studies (e.g. Schodde 1982a) and the cladistic analyses (Figure 2), that for species produced absurd associations. Options (e.g. ‘B = -0.1’, Belbin 1987) to tighten the clusters produced large changes in topol¬ ogy indicative of instability. Accordingly the UPGMA analyses were discarded. Both cladistic programs for genera produced five equally short trees of 844 steps (consistency index 0.39) and identical topology. A strict consensus tree derived from the PAUP cladograms is presented in Figure 2, in which terminal branches for indis¬ putably close sister assemblages have been collapsed for better appreciation. The New Zealand rainforest and New Guinean and New Zealand alpine assemblages are separated first as outlying elements. Then the New Guinean swamp grassland com¬ ponent is split off, along with all Australo-Papuan and New Caledonian rainforest assemblages on one side and those of the Australian eucalypt or scleromorphic veg¬ etation on the other. Significantly for rainforest elements, those in montane New Guinea cluster with Australian subtropical assemblages, and those in tropical Australia with lowland New Guinea. The assemblages of scleromorphic vegetation are aligned as follows: New Guinean tropical with Australian tropical, then a cluster comprising the temperate assemblages of southern and montane eastern Australia, which are linked in turn to those of the Australian mallee and arid zone. There is one noteworthy anomaly. The north Queensland tableland assemblage, which includes many temperate species, is linked more closely to the tropical assemblages than the lowland central east coast assem¬ blage which itself includes many tropical-centred species. When their branches are reversed, the tree is lengthened by only two steps. For species, only HENNIG 86 coped with the vast matrix of 918 species partitioned among 44 areas. It produced six equally short trees of 1733 steps (consistency index 0.51). A strict consensus tree derived from them, in which terminal branches for close sister assemblages have again been collapsed, is presented in Figure 2. It is little different in topology from the tree for genera, the Australo-Papuan rainforest and scleromorphic vegetation elements separating once more, and the New Zealand and New Caledonian assemblages lying further out. The only noteworthy variations are the linking of Australian subtropical rainforest assemblages to those in tropical Australia instead of montane New Guinea; and the clustering of the north Queensland table¬ land eucalypt assemblage now with temperate or Bassian elements. Autochthonous Australasian marker taxa that have speciated through all or most re¬ gional avifaunulas are few and limited largely to Australo-Papua; and those that have been assessed phylogenetically are still fewer. Among all there is a consensus that rainforest-inhabiting members, particularly in montane New Guinea and subtropical Australia, represent ancestral forms from which those in scleromorphic vegetation 408 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI < _i < dc I— C/5 D < cu cn 05 Q. X 0) c c o Q_ 05 o CM LU DC O LL South-west Australian eucalypt torest/heath ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 409 have been derived. This has been found in the bronze-cuckoos, Chrysococcyx (Marchant 1972), and in the Australo-Papuan thornbills, scrubwrens, robins and fairy- wrens, Acanthiza, Sericornis, Petroica, Malurus (Mayr & Serventy 1 938,Christidis et al. 1988, Fleming 1950, Schodde 1982b). The single exception, stressed by Ford (1986) in acanthizid Gerygone, is a misinterpretation. Although Ford found no evi¬ dence for a subtropical rainforest (Tumbunan) origin, his two ancestral species- groups, Gerygone mouki and G. fusca, are respectively core elements of the subtropi¬ cal rainforests of Australia-New Zealand and represented by a root member, Gerygone ruficollis, in montane New Guinea. DISCUSSION Synthesis of ornithogeographical elements The congruence between the area cladograms for genera and species with descrip¬ tive ornithogeographies (e.g. Schodde 1982a) is consistent with a long history of en¬ demic vicariant development in the Australasian avifauna. This is particularly so for Australia-New Guinea, exploding misconceptions still current in the Australian litera¬ ture. Australia and New Guinea are not twin faunas (cf. Keast 1981); they share the same avifauna split fortuitously at Torres Strait by global climate. Its rainforest facet dominates in New Guinea and its scleromorphic vegetation facet in Australia, but outliers of both are present vicariously in both lands. Any ornithogeography which does not distinguish them in Australia is flawed (cf. Kikkawa and Pearse 1969). The avifaunas of New Zealand and New Caledonia, and of the limited environments of swamp grassland and alpine moor, are more distantly divergent and ambivalent in their links. Judged by their depauperate composition, they are built largely by disper¬ sive taxa. Because of the age of its members, the area cladogram of generic assemblages prob¬ ably represents basic Australasian avifaunal elements the better. When paired Aus¬ tralian and New Guinean assemblages are grouped and extrapolated through their vegetational environments today, eight major avifaunal elements result with a distri¬ bution as given in Figure 1. The five largest approximate closely to the Tumbunan, Irian, Torresian, Bassian and Eyrean avifaunulas already identified in earlier empiri¬ cal studies (Schodde 1982a and references therein), and are shared by Australia-New Guinea. The other three are the New Guinean alpine and New Zealand and New Caledonian elements. Significant endemism is characteristic of all eight; lack of it dis¬ counted the New Guinean swamp grassland fauna. Reconstruction of avifaunal development From their structure and links, the likely development of these avifaunal elements can now be traced on a palaeogeographical base. The small disharmonic New Zealand avifauna is layered in character, comprising a Gondwanan base upon which succes¬ sive waves of immigrants, mostly from Australia, have built (cf. Fleming 1962). The FIGURE 2 - Left. Strict consensus area cladogram of Australian avifaunal assemblages for genera, based on five PAUP trees produced using the procedures “Heuristic search, Hold = 5, Addition sequence = simple, Heuristic search with addition sequence = random”, and “TBR” branching swapping. Right. Strict consensus area cladogram of Australian avifaunal assemblages for species, based on six HENNIG 86 trees produced using the procedures “mhennig*” and “bb*” to apply extended branch swapping. 410 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Gondwanan base - of kiwis, the recently extinct moas, nestorine parrots, and perhaps the xenicid wrens - is unique. Initial Tertiary immigrants probably included the ances¬ tors of cyanoramphine parrots, mohouines, crow-honeyeaters and callaeatids. Last to arrive were members of modern Australian genera and species, establishing them¬ selves in Gondwanan rainforests that have changed little since the mid Tertiary and been culled to an uncertain extent by tectonic events and Pleistocene glaciations. With 71 species, New Caledonia also has a small and disharmonic avifauna, in marked contrast to its continental flora rich in Gondwanan elements. The only likely living Gondwanan relict is the Kagu, Rhynochetos jubatus. Nearly all other avian taxa are of genera and species widespread in Australia-New Guinea and New Zealand today, indicating that the New Caledonian avifauna too has been built by dispersal, principally from Australian and New Guinean centres. Generic endemism is less than in New Zealand and there is little faunal layering, suggesting a later, more com¬ pressed history of colonization. The development of the great, harmonic Australian and New Guinean avifaunas, which today share 900 of the breeding land and freshwater birds in the region, is re¬ constructed here together because their lands have a common palaeogeographical history and share a rich Gondwanan base of ratites, megapodes and Psittaciformes, and possibly ground pigeons, tytonid owls, podargids, wood kingfishers and even passerines. By the mid Tertiary, New Guinea had not formed. Its present southern plain was part of the northern rim of the stable Australian plate and its northern sec¬ tor a skein of fringing islands. Australia itself was vegetated extensively with subtropi¬ cal rainforests that included Gondwanan laurels, myrtles, podocarps, cunoniads and Antarctic Beech ( Nothofagus ) found in montane New Guinea today (Kemp 1978; Barlow 1981). This correlation suggests that the subtropical, Tumbunan avifaunas now present in montane New Guinea were widespread in Australia then. The north New Guinean islands, in contrast, appear to have been clothed in rich rainforests of Malesian origin (Axelrod & Raven 1982). Two coincident events from the later Tertiary into the Pleistocene shaped subsequent developments. One was the raising of the massive central New Guinean cordillera to join the northern islands with the southern plain; New Guinea took its present form from then on (Pieters 1982). The other was the onset of aridity in Australia (Bowler 1982). The austral rainforest biota evidently withdrew before it, taking refuge ulti¬ mately in humid pockets on east coast ranges and in montane New Guinea. As Archer and Fox (1984) paraphrase, a walk up a New Guinea mountain is a walk back into time. On the caps of the New Guinean cordillera developed a novel alpine moor, opening new niches colonized itinerantly more by dispersing Eurasian taxa - phasianids, thrushes, pipits and munias - than Australian elements. The tropical Malesian rainforests of the north New Guinean islands now spread south around the cordillera into far northern Australia, to be occupied primarily by birds of Australian origin. This biogeographical paradox, which holds for mammals as well, still needs a full accounting (Gressitt 1982). In Australia itself, intensifying aridity spurred a radiation in already incipient scleromorphic vegetation avifaunulas (cf. Kikkawa et al. 1979; Barlow 1981). These drew their ancestral stocks mainly from autochthonous rainforests through adaptation ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 411 (Schodde 1982a). Regional topography and climatic pattern focussed aridity in the centre of the continent, in effect splitting the avifaunas of eucalypt forests and wood¬ lands into northern tropical (Torresian) and southern temperate (Bassian) belts. There they have since diverged from one another, as is evident in many vicariant pairs of taxa at generic (e.g. Geophaps - Phaps, Aprosmictus - Polytelis) and species (e.g. Melithreptus albogularis - M. lunatus, Artamus minor - A. cyanopterus) level. The arid centre itself drew its fauna from Bassian (primarily) and Torresian sources, building on trains of adaptive shifts that were culled repeatedly by climatic oscillations into the Pleistocene (Schodde 1982a). The scene had now been laid for the modern regional speciation documented so ably in current taxonomic literature. ACKNOWLEDGEMENTS We are indebted to Mr G. Richards and Ms R. Dunlop of CSIRO Division of Wildlife and Ecology for respectively preparing the figures and typing the manuscript. LITERATURE CITED ARCHER, M., FOX, B. 1984. Background to vertebrate zoogeography in Australia, 1-15. Vertebrate Zogeography and Evolution in Australsia. Perth, Hesperian Press. AXELROD, D.I., RAVEN, P.H. 1982. Paleobiogeography and origin of the New Guinea flora, 919-941 . Monographiae biologicae 42, Biogeography and Ecology of New Guinea. The Hague, W. Junk BARLOW, B.A. 1981. The Australian flora: its origin and evolution, 25-75. Flora of Australia, 1. Can¬ berra, Australian Government Publishing Service. BEEHLER, B.M., FINCH, B.W. 1985. Species-checklist of the Birds of New Guinea. Melbourne, Royal Australian Ornithologists Union. BELBIN, L. 1987. PATN: pattern analysis package. Canberra, CSIRO Division of Wildlife and Ecology. BOWLER, J.M. 1982. Aridity in the late Tertiary and Quaternary of Australia, 35-45. Evolution of the Flora and Fauna of Australia. Adelaide, Peacock Press. BULL, P.C., WHITAKER, A.H. 1975. The amphibians, reptiles, birds and mammals, 231-276. Monographiae biologicae 27, Biogeography and Ecology in New Zealand. The Hague, W. Junk. CHRISTIDIS, L., SCHODDE, R., BAVERSTOCK, P.R. 1988. Genetic and morphological differentiation and phylogeny in the Australo-Papuan scrubwrens ( Sericornis , Acanthizidae). Auk 105: 616-629. CONDON, H.T. 1975. Checklist of the Birds of Australia, 1 Non-passerines. Melbourne, Royal Australa¬ sian Ornithologists Union. CRACRAFT, J. 1973. Continental drift, paleoclimatology, and the evolution and biogeography of birds. J. Zool., Lond. 169: 455-545. CRACRAFT, J. 1986. Origin and evolution of continental biotas: speciation and historical congruence within the Australian avifauna. Evolution 40: 977-996. DELACOUR, J. 1966. Guide des oiseaux de la Nouvelle Caledonie et de ses dependences. Neuchatel, Delachaux et Niestl. DIAMOND, J.M. 1972. Avifauna of the eastern highlands of New Guinea. Nuttall Orn. Club Publ. 12. DIAMOND, J.M. 1986. Evolution of ecological segregation in the New Guinean montane avifauna, 98- 135. Community Ecology. New York, Harper and Row. FALLA, R. 1953. The Australian element in the avifauna of New Zealand. Emu 53: 36-46. FARRIS, J.S. 1988. HENNIG 86 Reference, Version 1.5. J.S. Farris, New York State University. FLEMING, C.A. 1950. New Zealand flycatchers of the genus Petroica Swainson (Aves). Trans. Proc. Roy. Soc. N. Zeal. 78:14-47, 127-160. FLEMING, C.A. 1962. History of the New Zealand bird fauna. Notornis 9: 270-274. FORD, J. 1986. Phylogeny of the acanthizid warbler genus Gerygone based on numerical analyses of morphological characters. Emu 86: 12-22. FORD, J. 1987. Hybrid zones in Australian birds. Emu 87: 158-178. GRESSITT, J.L. 1982. Zoogeographical summary, 897-918. Monographiae biologicae 42, Biogeogra¬ phy and Ecology of New Guinea. The Hague, W. Junk. 412 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI KEAST, A. 1981. The evolutionary biogeography of Australian birds, 1585-1635. Monographiae biologicae 41, Ecological Biogeography of Australia. The Hague, W. Junk. KEMP, E.M. 1978. Tertiary climatic evolution and vegetation history in the southeast Indian Ocean region. Palaeogeogr. Palaeoclim. Palaeoecol. 24: 169-208. KIKKAWA, J. , PEARSE, K. 1969. Geographical distribution of land birds in Australia - a numerical analysis Aust. J. Zool. 17: 821-840. KIKKAWA, J., INGRAM, G.J., DWYER, P.D. 1979. The vertebrate fauna of Australian heathlands - an evolutionary perspective, 231-279. Ecosystems of the World 9A, Heathlands and related shrublands. Amsterdam, Elsevier. KINSKY, F.C. 1970. Annotated checklist of the birds of New Zealand. Wellington, Reed. MARCHANT, S. 1972. Evolution of the genus Chrysococcyx. Ibis 114:219-233. MAYR, E. 1944a. Timor and the colonization of Australia by birds. Emu 44:1 13-130. MAYR, E., SERVENTY, D.L. 1938. A review of the genus Acanthiza Vigors and Horsfield. Emu 38: 245-292. PIETERS, P.R. 1982. Geology of New Guinea, 15-38. Monographiae biologicae 42, Biogeography and Ecology of New Guinea. The Hague, W. Junk. PRATT, T.K. 1982. Biogeography of birds in New Guinea, 815-836. Monographiae biologicae 42, Bio¬ geography and Ecology of New Guinea. The Hague, W. Junk. SCHODDE, R. 1975. Interim list of Australian songbirds, Passerines. Melbourne, Royal Australasian Ornithologists Union. SCHODDE, R. 1982a. Origin, adaptation and evolution of birds in arid Australia, 191-224. Evolution of the Flora and Fauna of arid Australia. Adelaide, Peacock Press. SCHODDE, R. 1982b. The Fairy-wrens. A Monograph of the Maluridae. Melbourne, Lansdowne Editions. SWOFFORD, D.L. 1990. PAUP: Phylogenetic analysis using parsimony, version 3.0. Champaign, Illinois Natural History Survey. VUILLEUMIER, F. 1975. Zoogeography, 421-496. Avian Biology 5. New York, Academic Press. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 413 CONCLUDING REMARKS: ORIGINS AND EVOLUTION OF THE AUSTRALASIAN AVIFAUNA RICHARD SCHODDE CSIRO Division of Wildlife and Ecology, P.O. Box 84, Lyneham, ACT 2602, Australia It is my personal pleasure to open my concluding remarks by dedicating this sympo¬ sium to the late Australian systematic ornithologist, Dr Julian Ralph Ford. Julian Ford died in tragic circumstances at the age of 54 only months after attending the last Congress in Ottawa. Just as distressingly, the root cause - sectional antipathy to col¬ lecting, no matter how carefully planned - still remains. A driving man, Julian Ford had a meteoric effect on systematic ornithology in Australia in his 25 years of involvement, rising to become the leading exponent of geographical variation and speciation in Australian birds. The principal contribution that symposia such as this make to hypothetical faunal ori¬ gins is their up-dating of perception in the light of current knowledge. Advances in perception, in their turn, can only be judged from comparison with earlier reconstruc¬ tions. The last coherent syntheses of origins and radiation in the Australasian avifauna were brought together by Ernst Mayr (1944, 1965) and Allen Keast (1961, 1981) through the 1950s to 1970s. They saw the Australasian avifaunas as built up by im¬ migrant dispersal from Eurasian sources, the ancestral stocks arriving in waves over Indonesian archipelagic stepping stones. Those stocks that arrived first radiated far¬ thest, such as the Australasian ratites, parrots, lyrebirds and honeyeaters; and those that came last changed least, such as the region’s few Eurasian thrushes, pipits, larks, dicaeids. In Australia itself, the bird fauna of scleromorphic eucalypt vegetation was interpreted as the older and that of the rainforests as much younger, arriving from New Guinea in the Pleistocene. Such a reconstruction fitted palaeogeographic and phylogenetic understanding of the times, that the earth’s crustal blocks were fixed, that the morphological similarity be¬ tween Eurasian and Australasian passerines reflected intra-familial connections, and that the subtropical Australian rainforests were Malesian in origin and recent in ad¬ vent (Burbidge 1960). Despite the growing acceptance of continental drift over the last two decades, concessions to Gondwanan elements in the regional avifauna remained few. Correlations among the diverse reviews presented today compel a different percep¬ tion. There is now strong evidence of a massive endemic radiation in Australasian oscines, a radiation of such diversity that it must extend deep into the Tertiary. Con¬ sistent with this are the newly discovered fossils of oscines in inland Australia - among the oldest in the world - at a time when Australia had barely rifted from Antarctica and was still thousands of kilometers south of Eurasian island arcs. Their ancestral habi¬ tats, as exemplified in the parrots, were the subtropical rainforests that survive today in montane New Guinea and in pockets in coastal eastern Australia. 414 ACTA XX CONGRESSUS INTERNATIONAL^ ORNITHOLOGICI When integrated, these correlations suggest that the base elements of the autochthonous Australasian avifauna - its megapodes, parrots, halcyonine kingfish¬ ers, frogmouths and owlet-nightjars, and members of its Anseriformes, pigeons and particularly songbirds - as likely as not came from Gondwana. These elements domi¬ nate the Australasian land bird fauna today. The focus of its radiation was the stable Australian plate through the earlier Tertiary. Apart from their few surviving Gondwanan relicts, the avifaunas of New Zealand and New Caledonia have been built by overseas dispersal within the region. The core of the Australian land bird fauna developed in subtropical Gondwanan rainforest that covered much of Australia through the Tertiary ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 415 as Australia dessicated at its close, the rainforest avifauna withdrew to pockets on the east coast and newly-forming montane New Guinea, spawning scleromorphic vegeta¬ tion-adapted elements in the process, in reverse to sequences postulated previously (Figure 1). The only undisputable Eurasian immigrants are members of several near cosmopoli¬ tan waterbird and raptor genera, a handful of strigids, swifts and coraciids, and, in passerines, rarely more than one or two species in yet another handful of Eurasian- centred families: motacillids, laniids, turdids, dicaeids, hirundinids etc. Together they comprise less than 20% of Australasia’s modern land and freshwater avifauna. This reconstruction has its own circumstantial difficulties, the most contentious of which is the assumption of a prevailingly Gondwanan origin for the region's modern avifauna. As Boles (this symposium) stresses, elements of Gondwanan and Eurasian origin still cannot be distinguished with certainty. The issue is exemplified by the passerines. If they arose in Gondwana, as Olson (1989) and McLean (1990) postu¬ late, were (1) Australasia’s primary stocks of oscines inherited from Gondwana, from where they budded off dispersing stocks to Eurasia; or (2) did oscines first radiate in Africa, from where just one or two founders dispersed overseas very early to Australa¬ sia to establish the endemic Australasian radiation; or (3) were ancestral oscines split vicariantly into east (Australasian) and central (African) Gondwanan elements by the separation of Africa ca. 100 mya, spawning the radiation of holarctic oscines from African sources. Arguments can be mounted for and against all of these hypotheses; none resolve them. Here the distinction of Gondwanan distribution from Gondwanan origin is crucial. Olson (1989) points out that a number of relictual, presumably Gondwanan groups in the southern hemisphere today also have a fossil record of late Cretaceous-early Tertiary age in the northern hemisphere. So too do the marsupials (Fox 1987). Even allied nondispersive genera of modern flowering plant families have such a bihemispheric distribution: Fagus and Nothofagus. Yet Nothofagus forest and the marsupials are classical examples of a relictual Gondwanan distribution today. Match¬ ing them are contemporary distributional connections among the casuariids, kiwis and tinamous (Bock & Buhler 1988), parrots, plains wanderers and seed snipe, magpie geese and screamers, and megapodes and cracids (Sibley & Ahlquist 1990). What¬ ever their origins, Australasian members of these groups surely came from Gondwana. Indeed, virtually all land bird elements present in Australasia up to the mid Tertiary 35 mya were eastern Gondwanan, because until that time Australasia was in touch with then-forested Antarctica, not Eurasia. As explanations, the diverse immi¬ gration tracks into Australasia - Eurasian, Antarctic, northern, southern, Malesian and Gondwanan - presently in vogue in Australian biogeographical literature (e.g. Rich 1975) are as misleading as they are redundant. Although Eurasian elements may have arrived via such routes, Gondwanan elements were inherited in situ. The con¬ cept of migration routes implies dispersal, confusing the difference between it and vicariance at continental level and the differing contributions that both processes have made to shaping the regional avifauna. That Eurasian immigration of any substance did not begin until the later Tertiary, within the last 15-20 mya, is suggested as well by several ecological and palaeofloristic correlations in passerines. Those passerines of indisputable Eurasian 416 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI origin fall into two groups in Australasia. Either they are widespread members of Eura¬ sian species that exploit niches unoccupied by autochthonous passerines (e.g. Long¬ tailed Shrike Lanius schach, and Pied Chat Saxicola caprata, in anthropogenic grass¬ land in New Guinea, and Richard’s Pipit Anthus novaezelandiae of bare fields eve¬ rywhere). Or they comprise small groups which have undergone endemic radiation to generic level in the regional grasslands and savannahs (e.g. estrildine finches, sylviid warblers). Common to the habitats occupied by all of them is novelty and disturbance. Thus the savannahs and shrub steppes in which Australasian estrildines, sylviids and pipits are found are comparatively new to the region, their palaeogeographic record being no older than the late Tertiary when Australia began to dessicate (Barlow 1981). Corre¬ spondingly few autochthonous Australian passerines occur in these habitats, just as few Eurasian taxa have entered the endemic Australasian rainforests and scleromorphic vegetation. Here is historical evidence for the ecogeographical principle that the building of faunas by dispersal depends not just on ability to disperse but also the availability of niches and the ability to establish. It is a principle illustrated graphi¬ cally in New Zealand today, in the contrast between the introduced European birdlife of its pastures and the indigenous avifauna of its native forests. From an antipodean outpost, the Australasian avifauna has now moved towards the centre of the world’s ornithogeographic stage. It holds historic clues and perhaps answers to such global questions as the relationships between continental avifaunas, the origin of the passerines, and the ecology of dispersal. In conclusion, however, a note of caution. Only a decade ago a completely different scenario for its origin and radiation was being espoused with equal conviction. Whether the perceptions ad¬ vanced here today will stand in the future depends, quite simply, on continuing broad- based research into the phylogeography and palaeontology of the regional avifauna. LITERATURE CITED BARLOW, B.A. 1981. The Australian flora: its origin and evolution, 25-75. Flora of Australia, 1. Can¬ berra, Australian Government Publishing Service. BOCK, W.J., BUHLER, P. 1988. The evolution and biogeographical history of the palaeognathous birds. Proc. Int. 100 DO-G Meeting, Current topics avian biol., Bonn 1988. BURBIDGE, N.T. 1960. The phytogeography of the Australian region. Aust. J. Bot. 10:75-209. FOX, R.C. 1987. Paleontology and the early evolution of marsupials, 161-169. Possums and Opos¬ sums; Studies in Evolution. Chipping Norton, New South Wales, Surrey Beatty. KEAST, A. 1961. Bird speciation on the Australian continent. Bull. Mus. Comp. Zool., Harv. 123:303- 495. KEAST, A. 1981. The evolutionary biogeography of Australian birds, 1585-1635. Monographial biologicae 41, Ecological Biogeography of Australia. The Hague, W. Junk. MCLEAN, G.L. 1990. Evolution of the passerines in the southern hemisphere. Transvaal Museum Bull., Suppl.22. MAYR, E. 1944. The birds of Timor and Sumba. Bull. Amer. Mus. Nat. Hist. 83:123-194. MAYR, E. 1965. What is a fauna? Zool. Jb. Syst. 92:473-481. OLSON, S.L. 1989. Aspects of global avifaunal dynamics during the Cenozoic. Acta XIX Congressus Internationalis Ornithologici: 2023-2029. RICH, P.V. 1975. Antarctic dispersal routes, wandering continents and the origin of Australia’s non- passeriform avifauna. Mem. Nat. Mus. Vic. 36:63-126. SIBLEY, C.G., AHLQUIST, J.E. 1990. Phylogeny and Classification of Birds. New Haven, Yale Univer¬ sity Press. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI SYMPOSIUM 3 ORNITHOGEOGRAPHY OF THE PACIFIC REGION Convener A. KEAST 418 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI SYMPOSIUM 3 Contents INTRODUCTORY REMARKS: ORNITHOGEOGRAPHY OF THE PACIFIC REGION ALLEN KEAST . . . 419 THE CONTRIBUTION OF FOSSILS TO KNOWLEDGE OF HAWAIIAN BIRDS H. F. JAMES . . . . . . . 420 ECOLOGICAL IMPACT OF THE HUMAN DEPLETION OF FRUGIVORUS BIRDS IN POLYNESIA DAVID W. STEADMAN . 424 BIOGEOGRAPHY OF NEW GUINEA BIRDS: A RE-EVALUATION IN LIGHT OF NEW SYSTEMATIC AND ECOLOGICAL INFORMATION THANE K. PRATT . 425 AVIAN EVOLUTION OF SOUTHERN PACIFIC ISLAND GROUPS: AN ECOLOGICAL PERSPECTIVE ALLEN KEAST . . . 435 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 419 INTRODUCTORY REMARKS: ORNITHOGEOGRAPHY OF THE PACIFIC REGION ALLEN KEAST Department of Biology, Queen's University, Kingston, Ontario, K7L 3N6, Canada The Pacific Region has figured importantly in biogeographic and evolutionary studies. Many major concepts of island evolution have been generated there. Hawaii contains the world’s most exuberant and diversified archipelago fauna. The large island of New Guinea sports a degree of physical and biotic diversity approaching that of the best continental lowland rainforest and montane forest areas. New Zealand, an archaic continental relict, contains a mixture of old endemics and modern colonizers from Australia. The islands of the southwest Pacific show faunal attenuation and are ac¬ tive areas of insular speciation, as the studies of Mayr, and others, have demon¬ strated. The objective of this symposium is to examine newer data relative to evolution in the Pacific Basin area. Of particular significance are the findings, from cave fossil data, that former avifaunas were distinct from modern ones. This necessitates a rethinking of basic concepts of Pacific ornithogeography. It is important to take another look at New Guinea now that more modern data on vertical and horizontal distribution, and on the ecology of major components, is available. It is appropriate to draw contrasts between the avifaunas of tropical New Guinea and cool temperate New Zealand, with its relatively small avifauna. Small insular avifaunas of the Pacific invite review in a community and ecological perspective. HAWAII GALAPAGOS NEW GUINEA MARQUESAS SOLOMONS SAMOA COOK IS PITCAIRN NEW CALEDONIA ZEALAND NORFOLK I 420 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI THE CONTRIBUTION OF FOSSILS TO KNOWLEDGE OF HAWAIIAN BIRDS H. F. JAMES Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA ABSTRACT. Fossils of Hawaiian birds reveal at least eight additional avian colonizations of the archi¬ pelago, and about 45 additional species of resident birds, thereby raising the number of species in the Holocene avifauna to about 92 and the number of avian colonizations to 20 or more. The colonizing species can be categorized as either waterbirds, raptors, or passerines. Flightlessness and terrestriality developed in over half of the waterbird lineages, adaptations for ornithophagy occur in half of the rap¬ torial lineages, and adaptive radiation occurred in two out of five passerine lineages. The extinction of approximately 61 species of resident Hawaiian birds is attributed to the impact of human settlement of the islands. Keywords: Hawaiian Islands, fossils, evolution, extinction, human impacts. INTRODUCTION Preliminary reports of rich avian fossil deposits in the Hawaiian Islands were published during the 1980’s (Olson & James 1 982a, b, 1984, James 1987, James et al. 1987). Descriptions of seven new genera and 32 new species of birds represented in these fossil collections are in press (James & Olson in press, Olson & James in press), and three fossil genera and species were described earlier (Wetmore 1943, Olson & Wetmore 1976). Knowledge of the fossils has advanced enough to permit a reevaluation of evolutionary and zoogeographic trends in the resident, Holocene avifauna, taking into consideration both the fossil and historically known birds. Besides the 35 extinct species referred to above, Hawaiian fossil collections/include less diagnostic specimens that may represent up to 21 additional new species. A conservative estimate of the number of resident birds that became extinct before the historic period is 45 species, compared to 47 resident species that either still survive or became extinct during the historic period. Thus, there were originally at least 92 resident species in the Holocene avifauna of the main islands, 31 of which still sur¬ vive, all but 10 of which are now thought to be threatened with extinction (Pyle 1988). These figures exclude species that probably did not colonize until after humans set¬ tled in the islands and created appropriate habitat for them (e.g., Black-crowned Night Heron, Nycticorax nycticorax* , and Short-eared Owl, Asio flammeus). The above figures also do not include an estimate of the number of extinct species for which fossils have not yet been found. Productive fossil deposits are known from only four of the eight main islands (Kauai, Oahu, Molokai, and Maui); a lesser amount of bone material is available from the island of Hawaii, mainly from archaeological ‘Taxonomy follows James & Olson (in press) and Olson & James (in press); common names follow Pyle (1988). ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 421 contexts; and little or nothing is available from the other islands. Thus, fossils have added 28 species to the avifauna of Maui, 24 to the avifauna of Oahu, 17 to the avifauna of Molokai, 1 1 to the avifauna of Kauai, four to the avifauna of Hawaii, one to the avifauna of Lanai, and none to the avifaunas of Kahoolawe or Niihau. More new species and distributional records can be expected as representative fossil collections become known from the poorly collected islands. COLONIZATION The historically known resident avifauna can be traced to 14 colonizing species (Mayr 1943), or, subtracting N. nycticorax and A. flammeus because they are probably post¬ human arrivals, to 12 natural colonizations of the main islands. Fossil evidence con¬ tributes at least eight additional colonizations, bringing the total to 20. This figure is a minimum, as it excludes two obscure fossil geese ( Geochen rhuax and the Super¬ numerary Oahu Goose), and it incorporates the following assumptions: 1) the four species of flightless goose-like ducks ( Thambetochen and relatives) are derived from a single colonization, 2) the flightless rails ( Porzana ) are derived from three coloni¬ zations, 3) the three species of Hawaiian crows are derived from one colonization, and 4) the two genera of Hawaiian honeyeaters (Meliphagidae: Chaetoptila and Moho) are derived from one colonization. The 20 colonizers were an ibis (Plataleidae), two true geese (Anserinae), a dabbling duck {Anas), a second dabbling duck or a shelduck (Anatinae), three rails {Porzana), a gallinule {Gallinula), a coot {Fulica), a stilt {Himantopus), a hawk {Buteo), a sea eagle {Haliaeetus), a harrier {Circus), an owl (Strigidae), a crow {Corvus), a flycatcher (Myiagridae), a thrush {Myadestes), a honeyeater (Meliphagidae), and a finch (Carduelinae). The species that managed to colonize are divisible into three general categories: 11 were waterbirds, four were raptorial species, and five were passerines. Continental relatives of many of the resident Hawaiian birds are recorded as occa¬ sional visitors in the islands (e.g., Plegadis chihi (White-faced Ibis), many species of Anas (dabbling ducks), Branta canadensis (Canada Goose), Anser albifrons (White- fronted Goose), Haliaeetus pelagicus (Steller Sea-Eagle), Circus cyaneus (Northern Harrier), and Carduelis flammea (Common Redpoll) (Pyle 1988). Although these spe¬ cies are not necessarily ancestral to related taxa in the resident fauna, their occa¬ sional arrival in the islands illustrates how colonization may have taken place. The possibility also exists of colonization by island-hopping along the hot-spot islands of the Hawaiian-Emperor chain, most of which were submerged long ago and became seamounts (Jackson et al. 1972). This could result in the preservation of relictual taxa from as long ago as the late Cretaceous, the apparent age of the oldest seamount in the Emperor Chain (Scholl & Creager 1973, Worsley 1973). However, despite the growing fossil record there are still no obvious relicts in the Hawaiian avifauna, hence no evidence that avian lineages of extreme antiquity have been preserved by island¬ hopping from the older seamounts. Colonization of the main islands from the still subaerial Northwestern Hawaiian Islands has been proposed for the Hawaiian finches (Drepanidini) (Sibley & Ahlquist 1982). 422 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI EVOLUTION Waterbirds Eleven (55%) of the birds that colonized the main islands were waterbirds. An eco¬ logical shift from aquatic or semi-aquatic habitats to terrestrial habitats apparently occurred in seven of these lineages ( Apteribis , Branta, a large goose from the island of Hawaii (Anserinae)-, Thambetochen and relatives, and the rails derived from all three colonizations by species of Porzana). Fossils of these birds are found in locali¬ ties that are not near wetlands. Branching speciation, in which more than one endemic species has developed from a single colonization, occurred in at least 6 of the 7 ter¬ restrial lineages, giving rise to 21 or 22 endemic species. Flightlessness is the rule among the terrestrial waterbirds, with all but one of the species in this group being completely (19-20 species) or nearly (one species) incapable of flying. Terrestrial waterbirds are the only group of Hawaiian endemics in which flightlessness evolved. The four resident waterbirds that remained in aquatic habitats are Anas wyvilliana (Hawaiian Duck), Fulica alai (Hawaiian Coot), Gallinula chloropus (Common Moor¬ hen), and Himantopus knudseni (Hawaiian Stilt). So far, no instances of flightlessness or branching speciation have been documented for these lineages, but there are few Holocene fossils of wetland birds. Raptors The Holocene fauna of the main islands includes seven raptorial species: Haliaeetus sp., Buteo solitarius (Hawaiian Hawk), Circus sp., and four strigid owls derived from a single colonization. The harrier and the four owls have long legs and short wings compared to related species outside the archipelago. Similar body proportions occur in bird-catching hawks of the genus Accipiter. Long-leggedness in Hawaiian raptors has been interpreted as an adaptation for hunting forest birds (Olson & James 1982b, in press). Passerines Fossils cast a new light on the spectacular adaptive radiation of the Drepanidini by adding 14 new species, descriptions of which are in press, and up to 8 additional undescribed species that are known from less diagnostic specimens (James & Olson in press). This raises the number of species in the radiation from 34, including three known historically from the Northwestern Hawaiian Islands, to between 48 and 56. Fossils also add to the diversity of the Hawaiian Corvidae and Meliphagidae, with two new species of crows and apparently two undescribed species of honeyeaters. EXTINCTION The approximately 45 Holocene extinctions recorded by fossils are attributed to the impact of prehistoric human settlement of the islands (Olson & James 1982a,b,1984, James et al. 1987), which commenced about 1500 years ago (Kirch 1974). Human impacts during the historic period caused 16 additional extinctions of endemic spe¬ cies (Pyle 1988). The prehistoric and historic period extinctions should be viewed as one ongoing “extinction event”, which so far has removed roughly as many species of birds from the Hawaiian Islands as there were mammals lost from North America at the end of the Pleistocene (see Anderson 1984 for a list of extinct Pleistocene mammals). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 423 In the Hawaiian Islands, terrestrial waterbirds suffered almost universal extinction. All of the 19-20 flightless species are extinct, as is the one goose that was a weak flier at best. It is no coincidence that the sole surviving member of this group, Branta sandvicensis, is also the only species capable of sustained flight. Waterbirds that re¬ mained in wetland habitats fared better: these four species are still extant. However, the fact that no extinctions have been documented for wetland birds so far may be due to inadequate fossil sampling of wetland habitats. All but one raptorial species ( Buteo solitarius) suffered extinction. Contributing factors may have included low population numbers, disappearance of important prey species, and predation by humans. It is also possible that the raptors, like the flightless birds, were ground nesters. Ground nesters would be vulnerable to nest predation by rats, pigs, and dogs that were introduced by the Polynesians (Olson & James in press). Among drepanidines, at least 12 (80%) of the species with finchlike bills were lost from the main islands, and 16 (50%) of the species with more derived bill shapes were lost, either during the historic or prehistoric periods. The relatively dry, lowland for¬ ests, which are now virtually absent from the islands, may have provided essential habitat for these birds. ACKNOWLEDGEMENTS Allen Keast and Storrs Olson offered useful comments on the manuscript. I am also grateful to Allen Keast for organizing the symposium and to Storrs Olson for collabo¬ rating in the Hawaiian fossil research. LITERATURE CITED ANDERSON, E.A. 1984. Who’s who in the Pleistocene: a mammalian bestiary. Pp. 40-89 in Martin, P.S., Klein, R.G. (Eds). Quaternary extinctions: a prehistoric revolution. Tucson: University of Arizona Press. JAMES, H.F. 1987. A Late Pleistocene avifauna from the island of Oahu, Hawaiian Islands. Documents des Laboratoires de Geologic de la Faculte des Sciences de Lyon 99: 221-230. JAMES, H.F., OLSON, S.L. in press. Descriptions of 32 new species of birds from the Hawaiian Is¬ lands: Part 2, Passeriformes. Ornithological Monographs. JAMES, H.F., STAFFORD, T.W., JR., STEADMAN, D.W., OLSON, S.L., MARTIN, P.S., JULL, A.J.T., McCOY, P.C. 1987. Radiocarbon dates on bones of extinct birds from Hawaii. Proceedings of the National Academy of Sciences USA 84: 2350-2354. KIRCH, P.V. 1974. The chronology of early Hawaiian settlement. Archaeology and Physical Anthropol¬ ogy in Oceania 9: 110-119 . MAYR, E. 1943. The zoogeographic position of the Hawaiian Islands. Condor 45: 45-48. OLSON, S.L., JAMES, H.F. 1982a. Fossil birds from the Hawaiian Islands: evidence for wholesale extinction by man before western contact. Science 217: 633-635. OLSON, S.L., JAMES, H.F. 1982b. Prodromus of the fossil avifauna of the Hawaiian Islands. Smithsonian Contributions to Zoology 365: 1-59. OLSON, S.L., JAMES, H.F. 1984. The role of Polynesians in the extinction of the avifauna of the Ha¬ waiian Islands. Pp. 768-780 in Martin, P.S., Klein, R.G. (Eds). Quaternary extinctions: a prehistoric revo¬ lution. Tucson: University of Arizona Press. OLSON, S.L., JAMES, H.F. in press. Descriptions of 32 new species of Hawaiian birds: Part 1, non¬ passerines. Ornithological Monographs. OLSON, S.L., WETMORE, A. 1976. Preliminary diagnoses of two extraordinary new genera of birds from Pleistocene deposits in the Hawaiian Islands. Proceedings of the Biological Society of Washington 89: 247-258. 424 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI PYLE, R.L. 1988. Checklist of the birds of Hawaii - 1988. ‘Elepaio 48: 95-106. SCHOLL, D.W., CREAGER, J.S. 1973. Geologic synthesis of Leg 19 (DSDP) results; far north Pacific, and Aleutian Ridge, and Bering Sea. Initial Reports of the Deep Sea Drilling Project 19: 897-913. SIBLEY, C.G., AHLQUIST, J.E. 1982. The relationships of the Hawaiiian Honeycreepers (Drepaninini) [sic] as indicated by DNA-DNA hybridization. Auk 99: 130-140. WETMORE, A. 1943. An extinct goose from the island of Hawaii. Condor 45: 146-148. WORSLEY, T.R. 1973. Calcareous nannofossils: Leg 19 of the Deep Sea Drilling Project. Initial Re¬ ports of the Deep Sea Drilling Project 19: 741-750. ECOLOGICAL IMPACT OF THE HUMAN DEPLETION OF FRUGIVOROUS BIRDS IN POLYNESIA DAVID W. STEADMAN New York State Museum, 3140 CEC, Albany, New York 12230, USA ABSTRACT. The extinction and extirpation that accompanied the human colonization of Polynesia involved many obligate frugivores (pigeons, parrots) and partial frugivores (megapodes, flightless rails, certain passerines). On most islands, frugivorous species of the forest canopy have been reduced in number, while those of the forest understorey and floor have been eliminated. At first human contact, most islands in the Marquesas supported 2 or 3 rails, 6 pigeons and doves, 3 parrots, and a starling. Only 1 to 3 frugivorous species survive on the same islands today. Similar depletion occurred in the Society and Cook Islands. Even isolated Henderson Island has lost a ground-dove and two pigeons since Polynesians arrived 800 years ago. In Tonga, human impact has eliminated a megapode, 2 flight¬ less rails, 5 pigeons and doves, 2 parrots, and a thrush on ‘Eua. Similar losses must have occurred throughout the region. Because of the decline in frugivorous birds, particularly columbids capable of ingesting large fruits, some Polynesian forest trees may be unable to disperse naturally within or be¬ tween islands today. This situation, now aggravated by mechanized forest clearing, may threaten sur¬ vival of the forest trees. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 425 BIOGEOGRAPHY OF NEW GUINEA BIRDS: A RE-EVALUATION IN LIGHT OF NEW SYSTEMATIC AND ECOLOGICAL INFORMATION THANE K. PRATT US Fish & Wildlife Service, Hawaii Research Group, P. 0. Box 44, Hawaii National Park, HI 96718, USA ABSTRACT. New Guinea supports an enormous and biogeographically distinct assemblage of birds, one of the earth’s four great tropical avifaunas. Past continental connections with Australia enabled the avifaunas of these two land masses to develop as a single one. As an equatorial mountainous island, New Guinea fosters the continued survival and evolution of the Australian rainforest avifauna, which, prior to the Pliocene, inhabitated much greater areas on the now arid continent. Following the impli¬ cations of current systematic and palaeontological studies, much greater endemicity for the Australo- Papuan region is now recognized. Current evolution of new species in New Guinea follows two broad patterns. For montane birds, speciation appears to take place by isolation and differentiation of new forms along the Central Ranges. Diversity of lowland and hill forest species is centered on three broad coastal plains where rainfall reaches a maximum for the island. These areas probably served as refugia for lowland populations during arid periods of the Pliocene and Pleistocene. Research of this impor¬ tant avifauna lags far behind that of other tropical regions. Keywords: Biogeography, avian palaeontology, speciation, New Guinea, Australia. INTRODUCTION Since the explorations of Alfred Russel Wallace (Wallace 1869), New Guinea has been known as the centre for an enormous and biogeographically distinct assemblage of birds, one of the earth’s four great tropical avifaunas. Although only a tenth the size of Australia, the island of New Guinea supports a larger fauna of resident land and freshwater birds than its continental neighbour to the south. These totals currently stand at 578 species for New Guinea (Beehler and Finch 1985, Beehler et al. 1986) and ca. 535 for Australia (Keast 1981a). Geographically the two land masses seem so different: New Guinea, a densely forested island of swampy plains divided by a mountainous spine 2,000 km long and reaching altitudes of 5,000 m; Australia, an arid continent fringed with wooded hills. Yet, perched on the continental shelf of Australia, New Guinea is separated from its southern counterpart only by a shallow sea and political boundaries, which serve more as barriers to dispersal of ornithologists than of birds. Past continental connections with Australia have enabled the avifaunas of these two land masses to develop as a single one. Further, the combined tropical avifauna of Australia and New Guinea, the Australo-Papuan Region (Sclater 1858, Keast 1981a), has had a profound influence on the development of avifaunas of the Oriental region and of oceanic islands in the tropical Pacific. Despite the importance of the New Guinea avifauna, its significance to Australian ornithology is frequently for¬ gotten. In part, this is because studies of New Guinea birds are few. For example, in this 20th IOC, the present paper is the only one of 140 symposia papers specifically addressing New Guinea birds. In this paper, I will review the biogeography of the New Guinea avifauna with emphasis on (1) development of the avifauna in relation to neighbouring regions and (2) abundant speciation on this now isolated island. 426 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI TABLE 1 - Distribution of selected families of land and freshwater birds breeding in the Australo-Papuan and Oriental Regions. Shown are numbers of genera and spe^ cies breeding in Australia proper, New Guinea, Australia and New Guinea combined, “Oriental Region” including only peninsular SE Asia, and numbers of genera shared between the Australo-Papuan Region and Oriental Regions. Families of passerines are ordered artificially for comparison of ecological counterparts, between regions. Data from Beehler & Finch 1985, King et al. 1975 modified from Keast 1990). Family Australia proper New Guinea 1 Australia & N.G. Oriental & Aust/N.G. Oriental Anseranidae magpie goose Dendrocygnidae 1-1 1-1 1-1 — — whistling ducks 1-2 1-2 1-3 1 1-2 Anatidae ducks, geese 8-16 4-7 11-16 2 4-4 Megapodidae megapodes Phasianidae 3-3 3-7 5-9 — — pheasants, quail 1-3 2-3 2-4 1 20-39 Columbidae pigeons 11-21 14-44 19-58 7 9-30 Psittacidae parrots 20-52 21-45 33-88 1 3-9 Tytonidae barn owls 1-5 1-4 1-5 1 2-3 Strigidae typical owls Podargidae* 1-4 3-6 3-7 2 7-19 Austral frogmouths Batrachostomidae* 1-3 1-2 1-3 — — Asian frogmouths Aegothelidae — — — — 1-5 owlet-nightjars Eurostopodidae* 1-1 1-6 1-6 — — eared nightjars 1-2 1-2 1-4 1 1-2 Caprimulgidae* nightjars 1-1 1-1 1-1 1 1-4 Hemiprocnidae tree swifts — 1-1 1-1 1 1-3 Apodidae swifts Cerylidae* 1-1 2-6 2-7 2 5-10 cerylid kingfishers Alcedinidae* — — — — 1-2 alcedinid kingfishers Dacelonidae* 1-2 2-4 2-4 2 2-6 dacelonid kingfishers 3-8 5-17 5-20 1 3-8 Meropidae bee-eaters 1-1 1-2 1-2 1 2-6 Coraciidae rollers 1-1 1-1 1-1 1 2-2 Bucerotidae hornbills Certhiidae — 1-1 1-1 1 8-13 Northern treecreepers Climaceridae — — — — 1-4 Australian treecreepers 2-6 1-1 2-6 — — The old muscicapi d assemblage • Maluridae fairywrens Acanthizidae 3-20 3-5 5-25 — — Australian warblers Eopsaltridae 9-40 4-20 9-54 1 1-1 Australian robins 7-20 11-25 12-38 — 1-1 Orthonychidae logrunners Pomatosomatidae 1-2 1-1 1-2 — — Australian babblers 1-4 1-2 1-5 — — ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 427 Cinclosomatidae whipbirds Monarchidae 2-8 4-6 5-14 monarchs & magpie larks 5-14 5-20 5-22 — 3-4 Rhipiduridae fantails 1-4 1-12 1-12 1 1-5 Pachicephalidae whistlers 5-14 6-26 8-34 1 1-1 Muscicapidae — 1-1 1-1 1 17-59 Turdidae “Sylviidae” and “Timaliidae” Old World warblers 1-2 2-2 2-4 1 5-16 & babblers 4-8 3-7 5-9 3 ca. 200 ‘familial arrangements proposed by Sibley & Ahlquist (1985) WHY THE NEW GUINEA AVIFAUNA IS AUSTRALIAN— WALLACE S SOLUTION TO A BIOGEOGRAPHIC PARADOX Recognition of New Guinea’s role as principal refuge and source of tropical avifauna for Australia and Oceania dates back to initial exploration of the island by Wallace. Apparent at that time and since has been the striking discontinuity between faunal assemblages as one passes from Asia across the Malay Archipelago to New Guinea. Together with other vertebrate groups and insects, an Asian bird fauna gives way to an Australian one, in contrast to the vegetation which shows little change across the same gradient. Wallace noted that the discontinuity coincided with the geographic position of the islands: those islands on the Asian continental shelf supported an Asian fauna; landbridge islands off Australia supported an Australian fauna. Wallace hypothesized that seas had flooded the continental shelves, isolating the islands and their biotas. For birds, the transition was especially evident among galliforms, most arboreal nonpasserine families and certain passerine families (Table 1). Groups that have ra¬ diated spectacularly in forests of New Guinea and Australia include pigeons, parrots, kingfishers, honeyeaters, birds of paradise, and bowerbirds. Cassowaries, megapodes, owlet-nightjars, and sittellas, though not so numerous, nevertheless are found only here. Up until ten years ago, other large groups of passerines were be¬ lieved to be shared between Asia and Australia. Similar morphologies tied the aus¬ tral warblers, flycatchers, thrushes, babblers, and others to the “muscicapid assem¬ blage” of Asia (Mayr and Amadon 1951) and supported the hypothesis that Australia derived much of its avifauna through waves of colonization by Asian immigrants. That these shared groups were passerine corroborated the idea that passerines worldwide had evolved and radiated recently (mid-Tertiary) relative to other bird families — an¬ other reason Australia would be principally at the receiving end of faunal interchange with Asia. Avian systematic studies of the past decade brought about the dissolution of the muscicapid assemblage (Mayr & Cottrell 1979, Sibley & Ahlquist 1985, Sibley et al. 1988) and further reshuffled other superficially similar passerine taxa in Australia and Asia, with the result that few of the large families are now considered shared between the two continental regions. Table 1 outlines the current taxonomic arrangement of the relevant passerines and nonpasserines, recognizing that relationships among these groups require further study. 428 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Thus, a visit now to Wallace’s realm finds an even greater division between the bird faunas of the Oriental and Australian Regions. The only large passerine families (>10 species in both region) with more or less equal representation in both regions are cuckoo-shrikes, starlings, orioles, white-eyes, and mannikins. Of these groups, the starlings, white-eyes, and mannikins clearly originated outside of Australia and, through their dispersal to much of the Pacific, colonized the island continent as well. All other large groups belong to one or the other continent with very thin representa¬ tion across “Wallace’s Line.” Australian groups with a toe-hold on Asia include, at one species each, the Australian warblers, robins, logrunners (if you count Eupetes), whistlers, and woodswallows. Likewise, the family list for Australia is padded by Asian add-ons: bee-eaters, 2 species; rollers, 1; larks, 1; pipits, 2; sunbirds, 2; flowerpeckers, 2. Confusing the issue of endemicity now is the family Corvidae as proposed by Sibley and Ahlquist (1985). Included in this assemblage besides the corvids proper are the cuckoo-shrikes, orioles, monarchs, fantails, drongos, cracticids, woodswallows, and birds of paradise; of these 9 closely related groups, 7 have their own history of exchange between Asia and Australia. Systematists who enjoy taxo¬ nomic and biogeographic tangles should pick at this one. NEW GUINEA BIRDS AND PLATE TECTONICS Wallace figured out that the land-bridge islands of the Oriental and Australian regions were just high ground on their respective continents and that rising oceans had cut them off. While this explained why the island biotas belonged to one continent or the other, broad differences between the biotas of the tropical Asian and Australo-Papuan regions remained unresolved until the advent of plate tectonics. Debate over the origins of the Australo-Papuan avifauna has focused on two hypoth¬ eses: (1) successive colonization of birds from Asia followed by radiation of particu¬ larly successful groups and (2) development of entirely or largely endemic families on the continent itself, with the few founding lineages either having arrived very long ago or having been there since the origin of modern birds. Because of Australia’s present isolation and the obvious recent arrival of certain Asian species, the colonization hy¬ pothesis was the first proposed (Mayr 1953, Darlington 1957). Beginning in the late 1960s (Dietz & Holden 1969), the theory of plate tectonics pro¬ vided an opportunity to consider the origins of the Australo-Papuan biota in a new historical context (Raven & Axelrod 1972, Keast 1981b). At the end of the Mesozoic, Australia was far south of its present position and, with Antarctica, formed a much larger continent. Seas separated this combined continent from South America and Africa, and some interchange of biota may still have been possible. Broadleafed ev¬ ergreen forests predominated. Early in the Tertiary (53 million years ago), Australia spilt from Antarctica and moved northward, reaching its present position in the Miocene, approximately 10 million years ago. By this time, ice had spread over Ant¬ arctica obliterating the diverse terrestrial communities that must once have existed there and eliminating faunal exchange with South America and Africa. New Guinea’s paleogeographic history began when island arcs formed in front of northward-moving Australia. Geological evidence (Loffler 1977, Doutch 1972) confirms that these early islands were always situated on the Australian continental shelf and were not an in¬ dependent land mass joined later. New Guinea never had the opportunity to evolve ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 429 a biota independently and has always been part of Australia (Schodde & Calaby 1972). Recognition of the former proximity of Australia to Antarctica, South America, and Africa has invited the search for affinities among their avifaunas. Unfortunately the evolutionary and geological timetables discourage such an interpretation, for nearly all modern bird lineages did not appear until the Palaeocene or Eocene (Olson 1988). The great extinction event at the Cretaceous/Tertiary boundary that felled the dino¬ saurs and other elements of the Cretaceous fauna can hardly be expected to have spared the birds, though its effect cannot be evaluated as yet. Perhaps the number of bird taxa on each continent was much reduced. The quest for shared elements among the southern avifaunas (Cracraft 1976, Rich 1975a, b) have produced ambigu¬ ous results, the potential candidates from Australia and New Guinea being the emu and cassowaries (as ratites), waterfowl, megapodes, button-quails, pigeons, parrots, and owlet-nightjars, and other less likely groups. In the past, passerines were dis¬ counted as their lack of a fossil record before the Miocene suggested an ascendence long after the break-up of southern continents. New interpretations of avian systematics (Mayr & Cottrell 1985, Sibley & Ahlquist 1985) have pushed back the age of passerines and suggest (1) that passerines are very old indeed, probably dating as far back as the presumed initial radiation of mod¬ ern birds in the Paleocene and Eocene, and (2) that perhaps all groups of old Australo-Papuan passerines are derived from a single, endemic radiation. This inter¬ pretation puts passerines in the running as charter members of the avifauna shared among the southern continents (Feduccia & Olson 1982). The most exciting evidence for origins of the Australo-Papuan avifauna comes from recent palaeontological work. Eocene sites in Europe and North America yielding large numbers of avian fossils, including associated skeletons, are currently being studied. Among the taxa identified are a number of modern groups formerly believed to be restricted to the southern hemisphere. Certain Australian taxa have turned up, including ratites, parrots, and owlet-nightjars. Olson (1988) has suggested that their modern distribution is relictual: these birds survived in Australia and died out else¬ where. Northern Eocene faunas differ also from modern ones in the taxa absent. The majority of ancient arboreal birds have been assigned to the Coraciiformes and Piciformes; none to the Passeriformes. Missing also are pigeons and, maybe, parrots. Among the waterbirds, the Anseriformes do not appear until the Miocene. As these groups are believed to date back this far, they may have originated in the Southern Hemisphere and spread to the north later. Unfortunately, contemporaneous sites have yet to be discovered in the southern continents. The oldest Australian bird fossils are of Miocene age. One wonders of the Australian Eocene: did the forests ring with the calls of coraciiforms and piciforms or the songs of passerines? RELATIONSHIPS BETWEEN THE PAPUAN AND AUSTRALIAN AVIFAUNAS In complete contrast to its present landscape of desert and woodland, Australia dur¬ ing the Tertiary supported vast tracts of mesic forest. This situation changed in the Pliocene as the continent became more arid. With the spread of xeric communities, rainforest persisted only in the eastern coastal ranges and, most importantly, in New 430 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Guinea. The fossil record for plants and mammals documents the local extinction of rainforest biota across the continent (Schodde & Calaby 1972, Keast 1981b); for birds, a comparable record is being discovered (Boles 1991). Unquestionably, the survival and continued evolution of Australia’s tropical avifauna has depended almost entirely on the continued presence of rainforest in New Guinea. Australian endemic lineages centered in New Guinea include the cassowaries, megapodes, fruit-pigeons, lories, pygmy and fig parrots, owlet-nightjars, dacelonid kingfishers, birds of paradise, berry-peckers, and longbills. Widespread Australian groups rich in Papuan genera and species include cuckoos, bowerbirds, robins, honeyeaters, acanthizid warblers, whistlers, monarchs, and others. Australo-Papuan rainforest birds outside of New Guinea inhabit the long archipelago of forest patches that run down the east coast of Australia. The most species-rich patch, at the base of the Cape York Peninsula, supports only half the species diver¬ sity of equivalent forest in New Guinea. Many Australian rainforest bird species are shared with New Guinea and probably arose there. Relationships between the tropical avifaunas of Australia proper and New Guinea have been studied extensively (Walker 1972, Schodde & Calaby 1972, Kikkawa et al. 1981). Differences between the avifaunas on either side of the Torres Strait can be accounted for by disproportionate extent of habitats rather than the present isolating effect of the Strait itself. For example, while the rainforest avifauna of Australia is improverished in comparison to that of New Guinea, the woodland bird community of southern New Guinea supports only half the species of Cape York, even though the two communities constituted a single one less than 10,000 years ago when the Torres Strait did not exist and xeric woodland penetrated much further inland from the New Guinea coast. Many woodland birds of Cape York are presumed to have then inhab¬ ited New Guinea as well, and if so have since become extinct, as predicted by the model of MacArthur and Wilson (1967). Kikkawa et al. (1981), in a detailed analysis of the distribution of bird communities along the Cape York Peninsula, concluded that size and distribution of habitat patches determined bird distribution. A further process was a “filtering effect”, a gradual attrition of rainforest species leading away from and woodland species towards New Guinea. DISTRIBUTION AND SPECIATION WITHIN NEW GUINEA Rainforests, by nature of their complex structure, moderate seasonality, and great diversity of food resources for birds, sustain far larger bird communities than other habitats. Relative to Australia proper, New Guinea maintains its diversity of tropical birds through (1) the immense expanse of its forests, which can support low-density bird populations large enough to escape extinction (MacArthur & Wilson 1967), (2) habitat diversity across gradients of moisture and altitude (Diamond 1972, Beehler 1982) and (3) extensive geographical regionalism (Diamond 1972, Pratt 1982). Area effect That larger areas hold more kinds of animals than smaller ones has been well docu¬ mented. This certainly applies to birds in New Guinea, where, for example, larger mountain ranges hold more species than smaller ones. Most New Guinea birds live ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 431 in low population densities; in other words, they are rare. Any birder knows this who tries in a day to fill a species list of birds recorded from a particular area. Rareness is manifest not only in low numbers for widespread species. Patchy distributions, re¬ sulting presumably from local extinction, are characteristic of many uncommon spe¬ cies. The Banded Yellow Robin and White-rumped Robin are two examples of hill forest species recorded only from a dozen or so localities and then only at species- specific altitudes. Local, or complete, extinctions are poorly understood for continental avifaunas; however, large expanses of habitat appear to provide a buffer for reduced populations and to offer more opportunities for survival in hard times. Altitudinal specificity Structuring of avian communities in New Guinea is a complex topic and beyond the scope of this short review. However, one mechanism that deserves mention, because of its hypothesized role in speciation and development of the avifauna, is the distri¬ bution of birds along altitudinal gradients (Diamond 1972). As one passes from the coastal plain to the mountain summits, bird species characteristic of low elevations drop behind while closely related species of higher elevations take their place. Spe¬ cies turnover may happen more than once; as many as six species (e.g., robins) can be stacked along an elevational gradient. Because some species leave off without replacement, overall species richness declines. Each species characteristically spe¬ cializes in an altitudinal band that remains fairly constant along the length of moun¬ tain range. While some species overlap broadly, others appear not to trespass appar¬ ent boundaries with neighbouring species. Speciation and geographical regionalism Local extinction and altitudinal segregation also bear on two broad patterns of regional endemism and speciation within New Guinea. For montane birds, the 2,000 km axis of the central ranges and 15 smaller outlying ranges provide ample opportunity for regional differentiation. Some ranges, such as the Vogelkop and Huon Peninsulas have accumulated a small assemblage of endemic species and a host of subspecies mixed in with the usual component of ubiquitous montane taxa (Diamond 1972, 1985). In addition, many species show patchy distributions; some being absent from seem¬ ingly suitable habitat. A proposed mechanism for speciation suggests that local ex¬ tinction somewhere along the middle of a species’ range is followed by differentiation of the two remaining separated populations. Upon reinvasion of the vacant middle ground, they again meet but do not interbreed because of newly acquired isolating behaviours. The two taxa now invade each other’s range, but they split their altitudinal habitat, so that one passes below the other. Various stages of this scenario can be found among examples of New Guinea montane birds (e.g. White-throated Treecreeper, various Parotia species). Principal centres of diversity for lowland birds occur in the south (Tran-Fly), west (Vogelkop) and North (Mamberamo/ldenburg). An analysis I conducted of geographi¬ cal speciation for lowland birds proposes that regional differentiation of widespread forms takes place in these three refugia (Pratt 1982). Barriers for dispersal between the refugia are scarcely apparent today, for rainforest follows the coastal plain around the island. However, the dispersal corridors lie in regions of lesser rainfall (Paijmans 1976, McAlpine & Keig 1983). Scenarios for Pleistocene climatic regimes suggest that these channels literally dried up (Nix & Kalma 1972), becoming woodland or savanna, habitats totally unsuitable for most birds of lowland closed forest. This arid period also 432 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI transformed into woodland the outer coastal margins of the rainforest refugia. Upon return of more mesic conditions and expansion of rainforest the separated taxa met again. The few cases of sympatry also show that altitudinal specialization serves as a segregating mechanism. For example, in southern New Guinea the Brown-collared Brush-Turkey only occurs in hillforest while the Black-billed Brush-Turkey occupies the lowland plains; in northern New Guinea, where the former species occurs alone, it occupies both the lowlands and hill-forest. CONCLUSIONS If there is a single message to conclude this presentation, it is that Australia does not end at the Torres Strait. New Guinea holds the greater part of Australia’s avifauna, and it also supports environments and taxa once far more widespread on the Austral¬ ian mainland. Despite the diversity of New Guinea birds and their relevance to avian research in the Australasian region, ornithology of New Guinea lags far behind that of other tropical avifaunas. Great opportunities await researchers on this insular frontier. ACKNOWLEDGMENTS Many thanks to Allen Keast, Bruce Beehler, Stors Olson, Charles Sibley, and Burt Munroe for discussion and literature sent during the preparation of this paper. Bruce Beehler came to the rescue and generously loaned all slides used for the presenta¬ tion while mine were in storage. Paul Banko, Bruce Beehler, Steve Fancy, and Allen Keast reviewed and commented on drafts of the paper. Special thanks to Allen Keast for involving me in the symposium and to Jim Jacobi for granting me time to prepare the paper and for seeking the means by which I could attend. LITERATURE CITED BOLES, W.E. 1991. The origin and radiation of Australasian birds: perspectives from the fossil record. Acta XX Congressus Internationalis Ornithologici. BEEHLER, B. M. 1982. Ecological structuring of forest bird communities in New Guinea. Pp 837-861 in Gressitt, J. L. Biogeography and ecology of New Guinea. Junk, The Hague. BEEHLER, B. M., FINCH, B. W. 1985. Species-checklist of the birds of New Guinea. R. A. O. U. Mono¬ graphs No. 1, Melbourne. BEEHLER, B. M., PRATT, T. K., ZIMMERMAN, D. A. 1986. Birds of New Guinea. Princeton Univer¬ sity Press, Princeton. CRACRAFT, J. 1976. Avian evolution on southern continents: influence of palaeogeography and palaeoclimatology. Pp. 40-52 in Frith, H. J., Calaby, J. H. (Eds). Proceedings of the 16th International Ornithological Congress. Australian Academy of Science, Canberra. DARLINGTON, P. J. 1957. Zoogeography: the geographical distribution of animals. Wiley, New York. DIAMOND, J. M. 1972. Avifauna of the eastern highlands of New Guinea. Nuttall Ornithological Club, Cambridge, Mass. DIAMOND, J. M. 1985. New distributional records and taxa from the outlying mountain ranges of New Guinea. Emu 85: 65-91. DIETZ, R. S., HOLDEN, J. C. 1969. The break-up of Pangea. Sci. Am. 233: 30-41. DOUTCH, H. F. 1972. The paleogeography of northern Australia and New Guinea and its relevance to the Torres Strait area. Pp. 1-10 in Walker, D. (Ed.). Bridge and Barrier: the Natural and Cultural His¬ tory of Torres Strait. Dept. Biogeography and Geomorphology Publication BG/3, Australian National University, Canberra. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 433 FEDUCCIA, A., OLSON, S. L. 1982. Morphological similarities between the Menurae and Rhinocryptidae, relict passerine birds of the Southern Hemisphere. Smithson. Contrib. Zool. 366: 1-22. KEAST, A. 1981a. The evolutionary biogeography of Australian birds. Pp.1 586-1 635. in Keast, A. (Ed.) Ecological biogeography of Australia. Junk, The Hague. KEAST, A. 1981b. Distributional patterns, regional biotas, and adaptations in the Australian biota: a synthesis. Pp 1891-1997 in Keast, A. (Ed.) Ecological biogeography of Australia. Junk, The Hague. KIKKAWA, J., MONTEITH, G.B., INGRAM, G. 1981. Cape York Peninusla: major region of faunal in¬ terchange. Pp 1695-1742 in Keast, A. (Ed.) Ecological biogeography of Australia. Junk, The Hague. LOFFLER, E. 1977. Geomorphology of Papua New Guinea. Australian National University Press, Can¬ berra. MCALPINE, J. R., KEIG, G. 1983. Climate of Papua New Guinea. CSIRO. Australian National Univer¬ sity Press, Canberra. MACARTHUR, R. H., WILSON, E. O. 1967. The theory of island bio-geography. Princeton University Press, Princeton. MAYR, E. 1953. Fragments of a Papuan ornithogeography. Proc. 6th Pacific Science Congress 4: 11-19. MAYR, E. , AMADON, D. 1951. A classification of recent birds. Amer. Mus. Nat. Novit. 1496:1-42. MAYR, E., COTTRELL, G.W. (Eds). 1979. Checklist of. birds of the world, vol. 1 (2nd ed.). Cambridge: Mus. Comp. Zool. NIX, H. A., KALMA, J. D. 1972. Climate as a dominant control in the biogeography of northern Aus¬ tralia and New Guinea. Pp. 61-91 in Walker, D. (Ed.). Bridge and Barrier: the Natural and Cultural History of Torres Strait. Dept. Biogeography and Geomorphology Publication BG/3, Australian National University, Canberra. OLSON, S. L. 1988. Aspects of global avifauna dynamics during the Cenozoic. in Ouellet, H. (Ed.) Acta XIX Congressus Internationalis Ornithologici. University of Ottawa Press. PAIJMANS, K. 1976. Vegetation. In Paijmans (Ed.). New Guinea vegetation. Australian National Uni¬ versity Press, Canberra. PRATT, T. K. 1982. Biogeography of birds in New Guinea. Pp. 815-836 in Gressitt, J. L. Biogeogra¬ phy and ecology of New Guinea. Junk, The Hague. RAVEN, P. H., AXELROD, D. I. 1972. Plate tectonics and Australasian paleobiogeography. Science 176: 1379-1386. RICH, P. V. 1975a. Antarctic dispersal routes, wandering continents, and the origin of Australia’s non- passeriform avifauna. Mem. Nat. Mus. Victoria 36: 63-126. RICH, P. V. 1975b. Changing continental arrangements and the origin of Australia’s non-passeriform continental avifauna. Emu 75: 95-112. SCHODDE, R., CALABY, J. H. 1972. The biogeography of Australo-Papuan bird and mammal faunas in relation to the Torres Strait. Pp. 257-306 in Walker, D. (Ed.). Bridge and Barrier: the Natural and Cultural History of Torres Strait. Dept. Biogeography and Geomorphology Publication BG/3, Austral¬ ian National University, Canberra. SCLATER, P. L. 1858. On the general geographical distribution of members of the Class Aves. Jour¬ nal Proceedings of the Linnean Society of Zoologists 2: 130-145. SIBLEY, C. G., AHLQUIST, J. E. 1985. The phylogeny and classification of the Australo-Papuan passerine birds. Emu 85: 1-14. SIBLEY, C. F., AHLQUIST, J. E., MUNROE, B. L. JR. 1988. A classification of the living birds of the world based on DNA-DNA hybridization studies. Auk 105: 409-423. WALKER, D. (Ed.). 1972 Bridge and Barrier: the Natural and Cultural History of Torres Strait. Dept. Biogeography and Geomorphology Publication BG/3, Australian National University, Canberra. WALLACE, A. R. 1869. The Malay Archipelago. Dover Publications, New York. Reprint of 1869 edition (1962). 434 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI ' ■ ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 435 AVIAN EVOLUTION OF SOUTHERN PACIFIC ISLAND GROUPS: AN ECOLOGICAL PERSPECTIVE ALLEN KEAST Department of Biology, Queen’s University, Kingston, Ontario, K7L 3N6, Canada ABSTRACT. The west-east avifaunal attenuation from biologically rich New Guinea to the impoverished islands of the central Pacific embraces three zones: continental New Guinea, the Melanesian Arc is¬ lands (Solomons - Fiji, plus Samoa), and the isolated Polynesian (Pacific Plate) islands to the east. The pattern varies with level in the taxonomic hierarchy and taxonomic group. Groups differ in capacity to establish eastwards. Birds-of-prey, megapodes, and nectar-feeding meliphagids do not occur beyond Fiji-Samoa. Frugivores are widespread and make up a high proportion of the avifaunas of eastern Poly¬ nesia. Communities of the Melanesian islands, though numerically impoverished, are ‘continental’ in ecomorphological types present, division of feeding substrates, and feeding behaviours. Members of a community studied in the field (Taveuni, Fiji) did not have wider ecological niches or greater ecologi¬ cal overlap values than those of an ‘equivalent’ continental rainforest (Macpherson Range, Queens¬ land). Central Pacific communities, by contrast, are depauperate and aberrant; some species show marked niche shifts. Newer geological data confirms that the Solomons, New Caledonia, and Fiji, the major centres of endemism, have occupied their present isolated positions through the Tertiary. Avifaunal origins by over-water dispersal, not vicariance, are confirmed. Keywords: Avifaunas, bird distributions, communities, island evolution, Pacific biogeography. INTRODUCTION Although it has long been appreciated that the southwest Pacific region is one of the most interesting areas of avian speciation and evolution (e.g., Mayr 1931, 1934, 1940a,b, 1941, Galbraith 1956) the area has received only limited attention in later years (but see Diamond 1970, Diamond & Mayr 1976, Diamond & Marshall 1976). This paper provides an updated ornithogeography of that section from eastern New Guinea to the Marquesas, considering faunal attenuation and community structure in an ecological context. METHODS The review is based on the literature, measurements of museum skins, and field work in Tahiti (August, 1984), Taveuni, Fiji (January, 1982; October, 1990); Macpherson Ranges, Queensland (October, 1983; December, 1986; November, 1988). Feeding behaviour, use of feeding substrates, and foraging habits were quantified using a modified version of the methods of Holmes & Recher (1986), and Recher & Gebski (1990). Feeding events (prey attack counts) were limited to five per individual bird and 20-25 individuals monitored over periods of nine or more days. Ecological diversities and overlaps were calculated by the formula of Levins (1968) and Schoener (1970), respectively. For fuller details see Keast (1991), some results of which are incorpo¬ rated here. 15 10 5 80 60 40 20 2 1 2 1 20 15 10 5 2 1 6 4 2 20 15 10 5 15 10 5 20 15 10 5 20 15 10 5 — ) le is , t ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Rarot. Tahiti N.G.E. GUADAL. N.CALED TAVEUNI I SAVAII I MANG. HUAH HlVA OA I ■ 1 I L ■ ! SPECIES/GENUS 1.4 1.4 LL* ■ ■■IB ACCIPITRIDAE & FALCONIDAE B 1 MEGAPODIIDAE RALLIDAE PSITTACIDAE SYLVIIDA E, ACAN THIZIDAE MONARCHIDAE, RHI 1.5 I I 'IDURIDAE, EOPSALTRIIDAE MELIPHAGIDAE Hender¬ son ivifaunal attenuation, numerical representation of families, species, etc., and )f avian families, eastern tip of New Guinea to Henderson Island. Additional ;hown in white. New Caledonia has two extinct families, Hiva Oa one; vo Ducula pigeons (Steadman 1988, 1989). For degree of statistical differ- the Melanesian Arc and Pacific Plate avifaunas see Appendix 1. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 437 The major islands The major islands (see Figure in Introduction to the symposium) fall into three catego¬ ries. New Guinea is large, continental, physiographically diversified and ornithologically very rich. Of modest size in the Oligocene most of it is of Pleistocene age. The Melanesian Arc islands (Solomons, New Caledonia, Fiji, New Hebrides) lo¬ cated towards the leading edge of the Australian Plate, are of intermediate size (e.g. Guadalcanal, 6,475 km2; New Caledonia, 16,750 km2); and also of mixed geological origin. All have been subaerial since the Miocene; New Caledonia possibly since the Cretaceous (Kroenke 1984). The Pacific Plate islands are small (Tahiti, 1,042 km2, Hiva Oa and Rarotonga, 200 and 67km2), volcanic, and have ages of 1-12 million years (Springer 1982). Their elevated areas (e.g. Tahiti reaches 2,238 m) are too restricted to support highland bird species. Inter-island distances between the major Arc islands are 800-1200 km; between the Plate islands, 1,200-2,100 km. RESULTS West-east avifaunal attenuation Features of the major island avifaunas from west to east are shown in Figure 1. To eliminate the compounding effects of allopatry within archipelagos data is given only for major islands. Figures are adjusted to incorporate extinct forms (Steadman 1988). The following conclusions emerge: (1) Drop-off in numbers of families is relatively uniform from New Guinea to Samoa, then falls precipitously. At the species level, by contrast, the drop occurs in three steps, between New Guinea and the Solomons, and east of Samoa. Genus/family ratios exceed 2.0 in New Guinea, Guadalcanal, and New Caledonia; are about 1.5 in Taveuni and Savaii; and 1.0 to the east. Species/genus ratios are 1.0 on all islands east of the Solomons, including New Caledonia, but are 1.2-1. 6 for the plate islands if fossils are included. Statistical tests are in Appendix 1. (2) The various avian groups differ markedly in eastwards distributions. Birds-of-para- dise are confined to New Guinea; hawks, megapodes, meliphagids extend as far as Fiji/Samoa (Mayr 1978). Rails, parrots, pigeons, kingfishers, swifts, the warbler Acrocephalus and monarch flycatcher Pomarea, by contrast, occur throughout the central Pacific. (3) Increasingly ‘unbalanced’ avifaunas result. Predators are absent east of Samoa. Frugivores (parrots, pigeons, starlings) make up an increasing percentage of the avifauna: vide eastern New Guinea, 40; Guadalcanal, 43; and for Rarotonga, Tahiti and Hiva oa (including fossil forms), 48-57. The reverse applies to insect-eaters. Varying ecological needs, as well as distance effects, presumably explain the differences. Avian community structure, insectivores, Melanesian islands Rain forest insectivore communities of Savaii and Taveuni at the eastern limits of Melanesia are compared numerically and in body sizes with ‘equivalent’ continental ones of the Macpherson Ranges, eastern Queensland, and Brown River, New Guinea (latter data from Bell (1982)) in Figure 2. Tarsus and bill lengths are plotted (scale, 10-40 mm). The former, of single measurements, is the best indicator of the bird’s body size (Rising & Somers 1989). The latter reflects size of dominant prey being consumed (Hespenheide 1973). Both Tahitian species are large. Savaii and Taveuni, with fewer species, show the same body and bill size spreads as the continental communities. 438 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI INSECT-EATERS PLUS MELIPHAGIDS, ZOSTEROPIDS, STURNIDS 6 4 2 0 TAHITI TARSUS LENGTH BILL LENGTH i i* i - CO LLJ o LU CL CO 6 4 2 0 mm BROWN RIVER NEW GUINEA i 9 10-11 12-13 14-15 16-17 18-19 20-21 22-23 24-25 26-27 28-29 30-40 LENGTHS (mm) FIGURE 2 - Community structure of insect-eaters and part-insect-eaters, based on lengths of tarsus and bill: Tahiti (Society Islands), Savaii (Samoa), Taveuni (Fiji), Macpherson Range, southeastern Queensland, and Brown River, Papua. Melanesian Island Arc avifaunas have the same range of body size forms as continental ones, though species numbers are much smaller. How does use of ‘ecological space’ compare? Taking Taveuni as the Melanesian is¬ land example, substrate use and foraging behaviours in six species are compared with six equivalent Macpherson Range forms (Figure 3). The Taveuni community separates into species that feed by gleaning from, and ‘fluttering at’, the foliage [Mayronis lessonii, Petroica multicolor, Myaigra azureocapilla)] aerial feeder ( Rhipidura spilodera, Myiagra vanikorensis)\ a trunk/branch and substratum feeder ( Lamprolia victoriae ); and an undergrowth-thicket feeder ( Vitia ruficapilla). (The lat¬ ter is not shown on the diagram.) These findings closely match those found by Holyoake (1979) for other seasons. It parallels the way feeding substrates are divided in the Macpherson Ranges community (Figure 3). How, then, are the greater number of species accommodated in the latter?. In part it is by a finer vertical partitioning of a feeding zone (Figure 3). That of the Taveuni Rhipidura spilodera is divided between R. fuliginosa and R. rufifrons, and Petroica multicolor between P. rosea and Tregellasia capito. In other cases two ecologically comparable species co-occur and are separated on microhabitat (e.g. Acanthiza pusilla and Gerygone mouki). For sig¬ nificance tests see Appendix 2. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 439 TAVEUNI MACPHERSON RA AERIAL FOLIAGE GL HANG GL FLUT AT VINES BRANCH OUTER INNER TRUNK UPPER LOWER BR, TR FLUT LOW SHRUB GROUND AERIAL FOLIAGE GL HANG GL FLUT AT VINES BRANCH OUTER INNER TRUNK UPPER LOWER BR, TR FLUT LOW SHRUB GROUND AERIAL FOLIAGE GL HANG GL FLUT AT VINES BRANCH OUTER INNER TRUNK UPPER LOWER BR, TR FLUT LOW SHRUB GROUND AERIAL FOLIAGE GL HANG GL FLUT AT VINES BRANCH OUTER INNER TRUNK UPPER LOWER BR, TR FLUT LOW SHRUB GROUND 0 10 20 30 40 0 10 20 30 40 % % MAYRORNIS LESSONII N=104 "I - \ - I | I - | - I" i GERYGONE MOUKI N=1 14 i - 1 - 1 - 1 - 1 - 1 - r 0 10 20 30 40 0 10 20 30 40 LAMPROLIA VICTORIAE N=97 i - 1 - - - | - 1 - 1 - r 0 10 20 30 40 0 10 20 30 40 % FIGURE 3 - Use of different substrates in foraging (numbers represent individual prey attacks), and foraging methods, six Taveuni and six Macpherson Range populations of species. For statistical treatment see Appendix 2. 440 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI How do members of the Taveuni insectivore bird community compare with the conti¬ nental one in (a) degree of ecological diversity exhibited (feeding substrate utilization and prey attack manoeuvres) and, (b) in degree of niche overlap? Figures of these are given in Appendices 3 and 4. Average diversity figures for members of the Taveuni community (10 species) for substrate use are 3.09 and for the Macpherson community (12 species), 3.00. For prey attack behaviours they are 1.64 and 1.85. Average figures for ecological overlap for the Taveuni community (9 paired compari¬ sons) are 0.53 for substrate use and for the Macphersons ones (11 comparisons) 0.36. For prey attack behaviours they are 0.63 and 0.67. The sets of figures are re¬ markably similar. The Taveuni community, hence, has a ‘continental-type’ structure in terms of these features, which presumably represent a stable state. Note, however, that niche shifts in island birds in the absence of competitors sometimes involves habitat changes (Diamond 1970). Pacific Plate island communities With their few species these communities contain striking examples of ecological shift and niche broadening. Acrocephalus is a short-billed reedbed and thicket insectivore in Asia and Australia. The long-billed insular derivates of Polynesia, by contrast, con¬ sume a wide range of insect types, plus lizards, fruit and nectar (Holyoak & Thibault 1984). Kingfishers ( Halcyon ) are shoreline and ground-feeding forest forms in Aus¬ tralia. In Polynesia they are predominantly arboreal rain forest forms feeding on in¬ sects in the branches and foliage, and occasionally in the air; they also take crabs and hunt in streams. The flycatcher Pomarea apparently has a narrower niche, feeding largely in the foliage and not also the air (Holyoak & Thibault 1984). This means that there is no true forest aerial feeder. Swifts do not deviate from their typical role. Acrocephalus and Pomarea commonly separate on habitat, arguing that resources are limited (D. Holyoak, personal communication). Grant (1968), and others, have noted the frequency with which island insectivores have longer bills, possibly an adaptation for taking of a wider range of prey. The long bill of Acrocephalus could be regarded as an extreme example of this. ‘Ecological release’, expansion of the niche in the ab¬ sence of competitors (Diamond 1970) could also be involved. Centres of endemism and Pacific geological history Endemic avian genera occur as follows: Solomons, 7; New Caledonia, 5; Fiji, 4; Sa¬ moa, 1 (Mayr 1978). Seven others occur on two or more island groups, indicating secondary spread. New Caledonia has an endemic family (Rhynochetidae), and a specialized fossil galliform ( Sylviornis ) of unknown affinities (Balouet & Olson 1989). Did these endemics arise by long distance dispersal, as the evidence suggests, or by vicariance (breakup of formerly continuous ranges)? Newer geological histories of the Pacific, incorporating plate tectonics (Coleman 1980, Kroenke 1984) show that whilst the Southwest Pacific has been highly dynamic tectonically the major island groups (Solomons, New Caledonia, Fiji) developed in their present position and have not, at least since the early Tertiary, been closer to each other and to the larger land masses than they are today. This confirms avian origins by over-water dispersal. DISCUSSION Diamond (1972) calculated the west-east attenuation rate for species from New Guinea into the central Pacific as representing a factor of two per 2,575 km. Earlier ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 441 Mayr (1940a, b) drew attention to the considerable avifaunal richness of the Melanesian islands compared to the impoverished Polynesian islands. This survey shows the picture to be more complicated. Attenuation patterns differ at the family, genus (Keast 1991), and species levels, in genus/family, and species/genus ratios, and according to taxonomic group. Whilst many ecological and taxonomic types are either confined to New Guinea, and/or the Melanesian islands, others extend to, and are highly successful in, the central Pacific. Parallel west-east attenuation patterns to those characterizing birds occur in many plant and animal groups. They are marked in mammals (Carter et al. 1946), reptiles, and many groups of plants and insects (list¬ ing in Thorne 1 963). What is the ecological meaning of decreasing species/genus ratios from west to east? This is to be expected because in any series such ratios decrease with decreasing species pool (Williams 1964). They may also indicate progressively reduced capac¬ ity to support closely related species (Grant 1966). Simberloff (1970) tested these alternative explanations for a large series of island groups. He found that deviations from predictions based on chance were not strongly correlated with island area, maxi¬ mum elevation, and distance from the source. He concluded also that similarities between congeneric species may also be a factor limiting (or permitting) co-occur¬ rence. See discussions of this subject in Peilou (1979, p. 225-226), and Brown and Gibson (1983, p. 526-527). The avifaunas of the southwest and central Pacific have been exclusively developed from the west by long distance dispersal (Mayr 1940a,b). This has also been accepted for virtually all animal and plant groups. There is a considerable literature, some of it quite speculative, as to how various groups achieved their range patterns (Thorne 1963). The newer data on geological history of the Pacific cements the case for avian origins by dispersal rather than vicariance. Note, however, that Raven & Axelrod 1974) have argued that origin of the unique floras of New Caledonia and Fiji would require these lying close to Australia in the Cretaceous or Early Tertiary. West/east attenuation patterns in the Pacific have presumably been built up over a long period of time and are now relatively stable. The occurrence of centres of ende¬ mism along the gradient do not change the basic pattern. Gradients are not unduly disrupted by differences in island sizes, or varying island distances. Large New Cal¬ edonia has an avifauna smaller than expectations. The Polynesian islands all have similar avifaunas despite their wide separations. Classic species/area effects do ap¬ ply within the Solomons archipelago (Diamond & Mayr 1976), but not to the Polynesian island groups (Steadman 1988, 1989). ACKNOWLEDGEMENTS This manuscript is based on research carried out under a research grant awarded by the National Science and Engineering Research Council of Canada and I should like to express my thanks to this body for its support. Measurements are based on skins in the collection of the American Museum of Natural History, New York and apprecia¬ tion is expressed to this body for making this material available to me. 442 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI LITERATURE CITED BALOUET, J.C., OLSON, S.L. 1989. Fossil birds from late Quaternary deposits in New Caledonia. Smithsonian Contributions to Zoology 469: 1-38. BELL, H.L. 1982. A bird community of lowland rainforest in New Guinea. 1. Composition and density of the avifauna. Emu 82: 24-41. CARTER, T.D., HILL, J.E., TATE, G.H.H. 1945. Mammals of the Pacific world. MacMillan, New York. COLEMAN, P.J. 1980. Plate tectonics background to biogeographic development in the southwest Pacific over the last 100 million years. Palaeogeography, Palaeoclimatology, Palaeoecology 31: 105-121. DIAMOND, J.M. 1970. Ecological consequences of island colonization by southwest Pacific birds. 1. Types of niche shifts. Proceedings National Academy of Science, U.S.A. 66: 529-536. DIAMOND, J.M. 1972. Biogeographic kinetics: estimation of relaxation times for avifaunas for South¬ west Pacific Islands. Proceedings National Academy of Sciences U.S.A. 69: 3199-3203. DIAMOND, J.M., MARSHALL, A.J. 1976. Origin of the New Hebridean avifauna. Emu 76: 187-200. DIAMOND, J.M., MAYR, E. 1976. Species-area relation for birds of the Solomon Archipelago. Proceed¬ ings National Academy of Science, U.S.A. 73: 187-266. GALBRAITH, I.C.J. 1956. Variation, relationships and evolution in the Pachycephala pectoralis superspecies (Aves, Muscicapidae). Bulletin of the British Museum (Natural History) Zoology 4: 133-222. GRANT, P.R. 1966. Ecological compatibility of bird species on islands. American Naturalist 100: 451-462. GRANT, P.R. 1968. Bill size, body size and ecological adaptations of bird species to competitive situ¬ ations on islands. Systematic Zoology 17: 319-333. HESPENHEIDE, H.A. 1973. Ecological inferences from morphological data. Annual Review Ecology and Systematics 4: 213-229. HOLMES, R.T., RECHER, H.F. 1986. Search tactics of insectivorous birds in an Australian eucalypt forest. Auk 103: 515-530. HOLYOAK, D.T. 1979. Notes on the birds of Viti Levu and Taveuni, Fiji. Emu 79: 7-18. HOLYOAK, D.T., THIBAULT, J.C. 1984. Contribution a I’etude des oiseaux de Polynesia orientale. Memoires Museo d’Histoire Naturelle 127: 1-209. KEAST, A. 1991. Insular evolution relative to environment and feeding niche shift in southwest Pacific populations of the ‘robin-flycatcher’ genus Petroica (Aves). Journal of Biogeography (submitted). KROENKE, L.W. 1984. Cenozoic tectonic development of the southwest Pacific. Technical Bulletin No. 6, United Nations Economic and Social Commission for Asia and the Pacific, Committee Co-ordinating Joint Prospecting for Mineral Resources in South Pacific offshore areas : 1 - 1 23. LEVINS, E.L. 1968. Evolution in changing environments. Princeton University, Press, Princeton. MAYR, E. 1931. Notes on Halcyon chloris and some of its subspecies. American Museum Novitates 469:1-10. MAYR, E. 1934. Notes on the genus Petroica. Birds collected during the Whitney South Sea Expedi¬ tion 29. American Museum Novitates 714: 119 . MAYR, E. 1940a. Borders and subdivisions of the Polynesian region based on our knowledge of the distribution of birds. Proceedings 6 Pacific Science Congress 4: 191-195. MAYR, E. 1940b. The origin and history of the bird fauna of Polynesia. Proceedings 6 Pacific Science Congress 4: 197-216. MAYR, E. 1978. Birds of the southwest Pacific. A field guide to the birds of the area between Samoa, New Caledonia, and Micronesia. Charles E. Tuttle, Rutland, Vermont. PIELOU, E.C. 1979. Biogeography. John Wiley, New York. RAVEN, P.H., AXELROD, D.l. 1974. Angiosperm biogeography and past continental movements. An¬ nals Missouri Botanical Gardens 61: 539-673. RECHER, H.F. , GEBSKI, V. 1990. Analysis of the foraging ecology of eucalypt forest birds: sequen¬ tial versus single-point observations. Pp. 174-180 in Morrison, F.M., Jehl, J.H., & Ralph, B.J. Studies in Avian Biology, Cooper Ornithological Society, Los Angeles. RISING, J.D., SOMERS, K.M. 1989. The measurement of overall body size in birds. Auk 106: 666-674. SCHOENER, T. 1970. Non-synchronous spatial overlap of lizards in patchy habitats. Ecology 51: 408-418. SIMBERLOFF, D.S. 1970. Taxonomic diversity of island biotas. Evolution 24: 23-47. SPRINGER, V.G. 1982. Pacific plate biogeography, with special reference to shorefishes. Smithsonian Contributions to Zoology 367: 1-182. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 443 STEADMAN, D.W. 1988. Fossil birds and biogeography in Polynesia. Pp. 1526-1534 in Ouellet, H. (Ed.). Proceedings 19th International Ornithological Congress, Ottawa. STEADMAN, D.W. 1989. Extinction of birds in Eastern Polynesia: a review of the record, and comparions with other Pacific Island groups. Journal of Archaeological Science 16: 177-205. THORNE, R.F. 1963. Biotic distribution patterns in the tropical Pacific. Pp. 311-350. in Gressitt, J.L. (Ed.). Pacific basin biogeography. Bishop Museum Press, Honolulu. WILLIAMS, C.B. 1964. Patterns in the balance of nature and related problems in quantitative ecology. Academic Press. APPENDIX 1 Numbers of families, genera, and species, are significantly different for a series of 12 Melanesian Arc islands relative to 7 Pacific Plate islands (that include those in Fig¬ ure 1): Mann-Whitney test (U = 84; P<0.0001 for extant forms; U = 84; P<0.0001, if the Plate faunas are supplemented by extinct forms). Genus/family and species/ge¬ nus ratios are significantly different between the two series for living forms (U = 66, P = 0.04; U = 65, P = 0.05). These ratios are marginally non-significant between the two series if extinct forms from the Plate islands are added (U = 60; P = 0.13; u = 61 ; P = 0.11). APPENDIX 2 Statistical comparisons (x2) for division of feeding effort between substrates for four cases of Tavenui species seemingly being represented by two in the Macpherson Ranges (Figure 2) were made on the basis of one randomly chosen observation per individual (using random numbers table). Results are: Mayrornis lessonii and Gerygone mouki (%2 = 1 .66; P = 0.8); M. lessonii and Acanthiza pusilla (x2 = 3.33; P = 0.5); G. mouki and G. pusilla (x2 = 2.2; P = 0.7); Rhipidura spilodera and R. fuliginosa (x2 = 6.7; P = 0.5); R. spilodera and R. rufifrons (x2 = 10.6; P = 0.2); R. fuliginosa and R. rufifrons (x2 = 16.9; P = 0.02); Petroica multicolor and P. rosea (x2 = 13.0; P = 0.07); P. multicolor and Tregellasia capito (x2 = 42.8; P = 0.0001); P. rosea and T. capito (x2 = 24.4; P = 0.001); Lamprolia victoriae and Sericornis magnirostris (x2 = 3.7; P = 0.7); L. victoriae and S. citreogularis (x2 = 30.6; P = 0.0001); S. magnirostris and S. citreogularis (x2 = 141.6; P = 0.0001). Thus Rhipidura fuliginosa and R. rufifrons; Petroica rosea and Tregellasia capito, and S. magnirostris and S. citreogularis (mainland species pairs); and Petroica multicolor (island) and T. capito (mainland), and L. victoriae (island) and S. citreogularis (mainland), are signifi¬ cantly different. On this basis only in one case ( Rhipidura ) does substrate use by the island form closely bridge the roles of two mainland species. 444 ACTA XX CONGRESSUS INTERNATIONAL^ ORNITHOLOGICI APPENDIX 3 Diversity indices, substrate utilisation and prey attack manoeuvres, formula of Levin's (1968). Species Substrate utilisation Prey attack (a) Tavenui manoeuvres Petroica multicolor 2.87 2.29 Mayrornis lessonii 1.58 1.83 Pachycephala pectoralis 2.41 1.11 Myiagra vanikorensis 3.26 2.89 Myiagra azureocapilla 2.40 1.81 Rhipidura spilodera 2.34 2.15 Lalage maculosa 2.12 1.06 Vitia ruficapilla 4.17 1.00 Lamprolia victoriae 5.95 1.00 Zosterops explorator 1.80 1.29 (b) MacPherson Ranges Petroica rosea 3.88 2.54 Tregellasia capito 4.89 2.82 Pachycephala pectoralis 2.78 2.46 Rhipidura rufifrons 3.51 2.76 Rhipidura fuliginosa 2.03 1.99 Sericornis magnirostris 5.53 1.00 Sericornis citreogularis 1.68 1.00 Sericornis frontalis 3.19 1.00 Gerygone mouki 1.73 2.22 Acanthiza pusilla 1.82 1.89 Climacteris leucophaea 3.64 1.00 Zosterops lateralis 1.70 1.51 / ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI x CD ~o C o h- 03 CD c CD O _C o CO CO CD > =3 CD O C 03 E X o z LU Q. Q. < j*. o 05 4—* 03 X 0) Q. 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E CO o o > CC _J > _l 445 446 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI .to o 03 .5 05 05 m -c | CO LO 00 05 ■ 05 CO C3 E ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 447 SYMPOSIUM 4 SYSTEMATICS AND BIOGEOGRAPHY OF AFROTROPICAL BIRDS Conveners T. M. CROWE and C. H. FRY 448 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI SYMPOSIUM 4 Contents INTRODUCTORY REMARKS: SYSTEMATICS AND BIOGEOGRAPHY OF AFROTROPICAL BIRDS T. M. CROWE . 449 TRENDS IN THE LITERATURE OF AFROTROPICAL ORNITHOLOGY W. R. SIEGFRIED, R. K. BROOKE and A. J. ARMSTRONG . 450 RADIATION IN AFRICAN CANARIES (CARDUELIDAE): A COMPARISON OF DIFFERENT CLASSIFICATORY APPROACHES RENATE VAN DEN ELZEN and HANS-L. NEMESCHKAL . 459 PICIFORM AFRO-ASIAN ZOOGEOGRAPHY AND SPECIATION L. L. SHORT and J. F. M. HORNE . 468 GEOGRAPHICAL MORPHOMETRIC VARIATION IN BIRDS OF THE LOWLAND EQUATORIAL FOREST OF AFRICA M. LOUETTE . 475 MORPHOMETRICS OF THE FALCONIFORMES: AN OVERVIEW A. C. KEMP and T. M. CROWE . 483 / ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 449 INTRODUCTORY REMARKS: SYSTEMATICS AND BIOGEOGRAPHY OF AFROTROPICAL BIRDS T. M. CROWE FitzPatrick Institute, University of Cape Town, Rondebosch 7700, South Africa Africa has a very special place in the history of mankind. If the mounting palaeonto¬ logical and biochemical evidence is correct this splendid continent was the evolution¬ ary cradle for both the genus Homo and for modern humans. Yet, Homo sapiens, man the wise, is in the process of destroying much of the fabulous biota to which his own future is inextricably linked. Even the most optimistic conservation scenarios for Af¬ rica predict that a large number of species are already doomed to extinction. There will be just too little room in the ever shrinking ark. Thus, like it or not, man the wise, or perhaps more appropriately, man the inconsiderate and short-sighted 'wise guy', will soon have to make some very tough decisions as to which taxa and populations should be saved. It is up to us as systematic biologists and biogeographers to pro¬ vide the evidence necessary to help make correct decisions in this terrible triage. Providing decision-makers with species inventories is not enough. At higher taxo¬ nomic levels, we need to identify the major evolutionary lineages that need to be pre¬ served to maximise evolutionary potential. At the lowest taxonomic levels, we need to know what patterns of within-species variation are needed to preserve a species' genetic integrity and thus ensure its long-term survival. From a biogeographical per¬ spective, how many reserves will be needed and how large and far apart must they be to preserve the surviving biota. I believe that the papers presented at this symposium will be valuable contributions in this regard. However, before I ask the first speaker to begin, I would like you to please stand up and join me in a brief moment of silence to protest the ill-conceived, ill-considered and ill-advised decision of the executive of the British Natural History Museum to abolish ALL of the research-oriented posts in its Sub-department of Or¬ nithology and charge visiting scientists substantial daily 'bench fees'. 450 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI TRENDS IN THE LITERATURE OF AFROTROPICAL ORNITHOLOGY W. R. SIEGFRIED, R. K. BROOKE, and A. J. ARMSTRONG FitzPatrick Institute, University of Cape Town, Rondebosch 7700, South Africa ABSTRACT. 13,662 citations in the Zoological Record from 1864 to 1980 inclusive which deal wholly or substantially with the birds of the Afrotropical Region (but not Madagascar) were identified and analyzed for various aspects. The predominance of citations dealing with taxonomy in the first 90 years was not maintained. A rise in the number of citations dealing with studies of living birds began in the 1950s and has been maintained since then. Biogeography was studied chiefly in the two decades 1951-1970 and systematics chiefly in the two decades 1941-1960. The Falconiformes and Charadriiformes (among others) were understudied before the 1950s, but this has since been more than remedied. There has been a recent decrease of interest in the Caprimulgiformes and Piciformes. The Passeriformes has never attracted as much attention as the number of its species warrants. This is a worldwide phenomenon. Keywords: Afrotropical ornithology, history, taxonomy, distribution, Caprimulgiformes, Charadriiformes, Falconiformes, Galliformes, Passeriformes, Piciformes, Procellariiformes. INTRODUCTION This paper is a contribution to a full history of ornithology in the Afrotropical Region, a work that has yet to be undertaken. Here we report on trends in topics of ornitho¬ logical study, based on the Afrotropical literature as recorded in the Zoological Record Aves from 1864 to 1980. There has been no detailed study or analysis of the whole contents of the Zoological Record. However, aspects have been noted by Besterman (1966 cited by Bridson 1968), Dadd (1971) and Chisman (1990). Only Besterman (1966) alluded even briefly to birds. Before the Zoological Record was established by Alfred Gunther, there were earlier attempts to list or catalogue zoological publications, the most important of which was A.F.A. Wiegmann’s Archiv fur Naturgeslchte from 1 835 to 1914 which included birds, but not one of them was as complete as the Zoological Record (Bridson 1968). This is still true (Chisman 1990). As far as Afrotropical ornithology is concerned, the most comprehensive listing of its literature for the 19th century is the 43 pages in Reichenow (1900). The next most important listing is the 71 pages in Chapin (1954). METHODS Each year’s Zoological Record Aves was scanned for citations dealing with Afrotropical birds, and the geographical breakdown provided therein was examined to check that all citations had been found. Each citation was given a unique identifying number. Since most of the literature is based on political entities or subdivisions thereof, the Afrotropical Region (chiefly Africa south of the Sahara Desert) was taken to include the whole of Mauritania, Mali, Niger, Chad and Sudan. Southwestern Ara¬ bia and oceanic islands, of which the largest is Madagascar, commonly regarded as Afrotropical, were excluded. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 451 Citations (separate entries, usually numbered, in the author index of each issue of the Zoological Record) were regarded as Afrotropical if it appeared from their titles or the geographical breakdown provided in the Zoological Record that they dealt entirely or substantially with Afrotropical birds within the region or in zoos, aviaries or laborato¬ ries elsewhere. Physical examination of citations was not normally attempted owing to time constraints and, with some of the 19th century literature, because it is not readily available. Identified citations were coded for computer analysis. Aspects coded included year of publication, principal topic of the citation (its main theme or purpose), important subsidiary topics of the citation where present, and the Wetmore (1960) order of birds covered when only one avian order was involved. Wetmore’s system was used be¬ cause it is the best known, not because we endorse it vis a vis other systems. All categories used are mutually exclusive in both statistical and logical senses, but the 38 topics defined in Appendix 1 are mutually exclusive only in the statistical sense. Citations were grouped into decades, e.g. 1891-1900, for analysis and detection of trends. Chi-square or, alternatively, log-likelihood ratio tests of independence for con¬ tingency data were undertaken, using SAS statistical software where appropriate (SAS User’s Guide 1985); the null hypothesis tested for each aspect being no change in relative citation frequencies between 1864 and 1980 inclusive. The null hypothesis for each aspect was rejected at the 5% significance level. FIGURE 1 - The relationship between actual (solid line) and expected (dashed line) numbers of citations for the four most studied principal topics (defined in Appendix 1) by decades. 452 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI RESULTS 13,662 citations were identified and analyzed. The main results concerning taxonomy/ systematics, distribution/biogeography and attention shown to different orders are given below. Principal topics The most frequent principal topics of study (the main theme or purpose of a citation) have been distribution, taxonomy, behaviour and breeding, in descending order of frequency (Figure 1). Each of these topics has frequencies an order of magnitude greater than the next most investigated topic, diet. However, only in the 1800s and in the decades after 1960 were there more distribution papers than expected. Breed¬ ing only became prominent in the 1960s, whereas behaviour became prominent in the 1930s. Taxonomy was the leading principal topic between the last decade of the 1800s and the 1950s, as the small piciforms and passerines were extensively sam¬ pled. The relative number of taxonomic papers decreased markedly in the 1960s, and taxonomy has not regained its status as the leading principal topic since then. Biogeography, for which there are only 108 citations, was most often studied between 1941 and 1980, although it was only in the 1950s and 60s that the number of citations exceeded expectation. Systematics, for which there are only 81 citations, was most often studied between 1941 and 1980, although it was only in the 1940s and 50s that the number of citations exceeded expectation. DECADES FIGURE 2 - The relationship between actual (solid line) and expected (dashed line) num¬ bers of citations for topics grouped into four major themes by decades. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 453 Figure 2 shows the results of amalgamating the 38 principal topics into four: taxonomy (including systematics), distribution (including biogeography), natural history and all topics not previously specified. Changes in the attention given to taxonomy and dis¬ tribution have been noted in the preceding paragraph. Natural history did not even approach its expected number of citations until the 1940s, and it was only from the 1960s onwards that its citations exceeded expectation, i.e. when taxonomy fell below it. All topics not previously specified are too heterogeneous for comment. Subsidiary topics The four most important subsidiary topics (major elements of a citation which are not its main theme or purpose) are the same as the four most important principal topics (Figure 3). The pattern between these four subsidiary topics broadly complements that of the principal topics. Behaviour is the most important in terms of citations, but only became prominent around the beginning of the 20th century as, prior to that, work¬ ers usually concentrated on taxonomy and distribution. Taxonomy is the second most prominent subsidiary topic, followed by breeding and distribution. Taxonomy and dis¬ tribution are highly correlated as topics, occurring together in many citations: hence the inversion of importance between these two topics from that found in studying the principal topics. Taxonomy was the most frequent subsidiary topic in the 1800s, but decreased markedly in the 1910s. Distribution was at its height as a subsidiary topic between 1921 and 1959, then decreased, which taxonomy did as a principal topic. _ DECADES _ _ FIGURE 3 - The relationship between actual (solid line) and expected (dashed line) num¬ bers of citations for the four most studied subsidiary topics (defined in Appendix 1) by decades. 454 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI The number of citations dealing with breeding as a subsidiary topic generally paral¬ lel the expected number, never reaching any great height. Orders The null hypothesis that members of orders were written about in proportion to their numbers in the Afrotropical avifauna (1,965 species, including vagrants) proves to be substantially true for six orders, viz. Podicipediformes, Gruiformes, Columbiformes, Strigiformes, Trogoniformes and Coraciiformes. The Struthioniformes and Galliformes have always attracted disproportionately great attention, whereas disproportionately little attention has been given to the Procellariiformes and Passeriformes (Figure 4; Table 1). Nine orders (Sphenisciformes, Pelecaniformes, Ciconiiformes Anseriformes, Falconiformes, Charadriiformes, Psittaciformes, Apodiformes and Coliiformes) have moved from expectation or below it to above expectation at one time or another in the 20th century. Three orders (Cuculiformes, Caprimulgiformes and Piciformes) have recently moved from expectation or above it to below expectation. c n sz; o DECADES FIGURE 4 - The relationship between actual (solid line) and expected (dashed line) num¬ bers of citations referring to four orders by decades. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 455 TABLE 1 - Percentage of worldwide citations in Wildlife Review dealing with passerines as opposed to other orders: Passeriformes constitute 58.4% of all bird species (Morony et al. 1975). Year Percentage of passerine citations Year Percentage of passerine citations 1976 29.5 1983 30.6 1977 26.4 1984 31.3 1978 31.1 1985 28.5 1979 30.7 1986 26.6 1980 30.2 1987 27.4 1981 31.5 1988 28.2 1982 31.1 1989 30.0 DISCUSSION The Zoological Record provides an operational baseline for studies such as this, even though it is known to be incomplete and to contain errors, not to mention changes in editorial policy over the years. Based on over 50 years combined experience of study¬ ing and writing on Afrotropical birds, we believe that the Zoological Record lists well over 90% of the significant literature on them, both refereed and unrefereed, as well as some of marginal significance. This opinion is supported by Mrs M.J. Thorne (Edi¬ torial Manager of BIOSIS, U.K., in lift. May 1990) and is not contradicted by Chisman’s (1990) findings. The decrease of interest in and publications on systematics and taxonomy of the Afrotropical biota has recently attracted attention (e.g. Ribbink & Greenwood 1988, Bruton 1989, Crowe et al. 1989, and, on a wider scale, Wilson 1985). Avian system¬ atics has never received much attention in the Afrotropical Region, partly because there are relatively few endemic taxa above the generic level, and partly because much of what has been done has been part of wider studies carried out in the north¬ ern hemisphere. In our experience, most of the taxonomic work on Afrotropical birds has been carried out by people unaffected by developments in the last 30 years in the theory and techniques of systematics. Biogeography, as opposed to purely distributional studies of particular taxa or areas, has never been widely practiced in the Afrotropical Region, perhaps because most workers potentially interested in this field have considered that the work of Chapin (1932) was close to adequate. In the period covered by this review (up to the end of 1980) the principal workers who have dealt with biogeography have been Reg Moreau (e.g. 1966) and Jack Winterbottom (e.g. 1974): neither produced a radical critique of Chapin’s work or used new techniques of study. 456 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Natural history or studies of living birds in the wild only approached expectation in the 1940s and exceeded it in the 1960s. This reflects the growing number of ornitholo¬ gists spending long periods in the Afrotropics or living there permanently, as well as an increasing interest in such matters, as opposed to taxonomic and distributional questions. The attention paid to six orders (Podicipediformes, Gruiformes, Columbiformes, Strigiformes, Trogoniformes and Coraciiformes) has been substantially in proportion to the number of species they contribute to the Afrotropical avifauna. The tendency has always been for large, conspicuous and economically important groups to receive more attention, either rrom the 1860s in the case of the Struthioniformes and the Galliformes or from some period in the 20th century. More curious perhaps is the recent decrease of interest in the Cuculiformes, Caprimulgiformes and Piciformes. With the rise of field studies considerable attention has been given to the Cuculinae breeding parasitical ly on members of the Passeriformes. Since orders have been included in this analysis when only one is involved in a citation, it is clear that citations on parasitic breeding have reduced the apparent attention given to the Cuculiformes and Passeriformes. This factor does not materially affect the relative lack of attention given to the Passeriformes. Most of the literature on the Caprimulgiformes has been taxonomic and since this aspect of Afrotropical ornithology is no longer dominant, this order has attracted lit¬ tle attention, not least because field studies on nocturnal birds are difficult to carry out. We are not sure why interest in the Piciformes has decreased. The relative neglect of the Passeriformes has been a constant in the Afrotropical lit¬ erature (cf. Craig 1987, 1988). Relative neglect of the Passeriformes is, in fact, a worldwide phenomenon. The similar neglect of the Procellariiformes is due to the fact that all are nonbreeding migrants to the coastal waters and beaches of the Region, mostly in the south. It was only in the 1970s that serious attention began to be given to nonbreeding seabirds in the Region: the African seabird journal Cormorant V\ rst appeared in 1976. The rise in the attention given to nine orders (Sphenisciformes, Pelecaniformes, Ciconiiformes, Anseriformes, Falconiformes, Charadriiformes, Psittaciformes, Apodiformes and Coliiformes) is a rectification of earlier lack of attention. In respect of the Psittaciformes there is a large body of literature based on laboratory and cap¬ tive studies. Another aspect which favours attention to large and conspicuous species is the relative ease in many cases with which statistically significant samples of data may be obtained. This is not usually true for members of the Falconiformes but here interest, often conservation based, has been focused on species at the top of food chains whose numbers and/or ranges have decreased markedly in many cases. ACKNOWLEDGEMENTS We, and all ornithologists, are obliged to our predecessors who made time to con¬ struct the Zoological Record Aves each year in addition to their research work and other duties (Bridson 1968). In chronological order they are: A. Newton 1864-1869, ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 457 H.E. Dresser 1870 (part), R.B. Sharpe 1870 (part), 1871-1874, 1882-1883, 1890- 1909, 0. Salvin 1875-1876, H. Saunders 1877-1881, A.H. Evans 1884-1889, W.L. Sclater 1910-1943, W.P.C. Tenison 1944-1963, A.I. Ivanov 1958-1963 (part). WRS thanks the Foundation for Research Development and the University of Cape Town’s Research Committee for financial support. We thank Mrs M.J. Thorne, Editorial Man¬ ager of BIOSIS, for photocopies of relevant literature. LITERATURE CITED BESTERMAN, T. 1966. World bibliography of bibliographies. Geneva [not seen - cited in Bridson 1968]. BRIDSON, G.D.R. 1968. The Zoological Record - a centenary appraisal. Journal of the Society for the Bibliography of Natural History 5: 23-34. BRUTON, M.N. 1989. Does animal systematics have a future in South Africa? South African Journal of Science 85: 348. CHAPIN, J.P. 1 932. The birds of the Belgian Congo Part 1 . Bulletin of the American Museum of Natural History 65: 1 756. CHAPIN, J.P. 1954. The birds of the Belgian Congo Part 4. Bulletin of the American Museum of Natural History 75B: 1-846. CHISMAN, J.K. 1990. Zoological Record, Biological Abstracts and Biological Abstracts/RRM\ a com¬ parison of overlap. RQ Winter 1989 pp. 242-247. CRAIG, A.J.F.K. 1987. Editorial. Ostrich 58: 23. CRAIG, A.J.F.K. 1988. Editorial - sixty years of Ostrich. Ostrich 59: 83-84. CROWE, T.M., KEMP, A.C., EARLE, R.A., GRANT, W.S. 1989. Systematics is the most essential, but most neglected, biological science. South African Journal of Science 85: 418-423. DADD, M.N. 1971. The Zoological Record - current developments. Biological Journal of the Linnaean Society 3: 291-294. MOREAU, R.E. 1966. The bird faunas of Africa. London, Academic Press. MORONY, J.J., BOCK, W.J., FARRAND, J. 1975. Reference list of the birds of the world. New York, American Museum of Natural History. REICHENOW, A. 1900. Die Vogel Afrikas, Vol. 1. Neudamm, Neumann. RIBBINK, A.J., GREENWOOD, P.H. 1988. Whither systematics: to wax or to wither? South African Journal of Science 84: 872-873. SAS User’s Guide 1985. Basics. Cary, SAS Institute. WETMORE, A. 1960. A classification for the birds of the world. Smithsonian Miscellaneous Collections 139(11): 1-37. WILSON, E.O. 1985. Time to revive systematics. Science 230: 1227. WINTERBOTTOM, J.M. 1974. The zoogeography of the South African avifauna. Annals of the South African Museum 66: 109-149. APPENDIX 1 DEFINITIONS OF TOPICS USED TO ANALYZE THEMES IN THE AFROTROPICAL AVIAN LITERATURE NB These definitions are not always logically mutually exclusive: see Methods. * indicates topics grouped for the category ‘natural history’ in Figure 2. Anatomy - studies of the structure of the soft parts of birds. ‘Behaviour - studies on the behaviour of birds, except for restricted studies on breeding, diet, etc. Biochemistry - studies of the chemistry and chemical functioning of birds’ bodies. Biogeography - studies of the patterns of geographical distribution of birds. Biometrics - studies of mensural data on parts of birds’ bodies or on diet or behaviour. ‘Breeding - studies of any aspect of reproduction. ‘Communication - studies of information transfer by voice or behaviour. ‘Community - studies of coexisting species in particular habitats or restricted areas. 458 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI ‘Conservation - studies of threats to bird populations, methods of mitigating such threats, and birds of conserved areas. Culture - human responses to birds not based on economic necessities. Development - studies of morphological or behavioural development of young birds, including in the egg. ‘Diet - studies of what birds eat. Distribution - data and studies on where bird species occur. Ecology - studies of birds’ relations with biotic and abiotic surroundings, usually quantified (seldom used). Economics - studies of relations between activities concerned with human livelihoods and birds. Environmental Change - studies of the effects of changing environments on birds and their populations. Evolution - studies of the effects of natural selection on characters of birds and their speciation. Genetics - studies of chromosomal and DNA structures and inheritance of characters. ‘Habitat - studies of faunas or communities associated with particular vegetation assemblages. History - history of ornithological activities in the Afrotropical region, from obituaries to ringing reports. Identification - aids to identifying bird species in the field. Integument - descriptions and studies on the skin and feathers of birds: see also Moult and Oology . ‘Locomotion - studies of flight, diving, walking and other means of movement. Migration - studies of migratory movements of birds: often subsumed in Distribution q.v. Moult - studies of moult of plumages and feather tracts. ‘Nutrition - studies of the value of different foods eaten by birds. Oology - studies of eggs and their external characteristics. Osteology - studies of bones and bone systems. Paleontology - studies of fossil birds, including archaeological remains. ‘Parasites - studies of external and internal parasites of birds and of birds breeding parasitically on others. ‘Pathology - studies of what kills birds or makes them sick, including countervailing treatments. ‘Pollution - studies of the side effects on birds of economic activities, including pesticides and oil spills. Physiology - studies of how living birds’ bodies work. ‘Population - studies of bird numbers, sex ratios and breeding productivity. ‘Ringing - studies arising from analyses of ringing recoveries, reports of ringing recoveries. Systematics - studies of avian relationships above the generic level. Taxonomy - studies of avian taxa at the generic level and below and the names to be applied thereto. Techniques - descriptions and studies of methods of achieving research results. / ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 459 RADIATION IN AFRICAN CANARIES (CARDUELIDAE): A COMPARISON OF DIFFERENT CLASSIFICATORY APPROACHES RENATE VAN DEN ELZEN1 and HANS-L. NEMESCHKAL2 1 Alexander Koenig Zoological Research Institute and Zoological Museum, Adenauerallee 150-164, D-5300 Bonn, Germany 2 Zoological Institute, University of Vienna, Althanstrase 14, A-1090 Vienna, Austria ABSTRACT. Phenetic analyses of skeletal measurements in 44 carduelid species identified three broad phenotypes. Since coefficients of variation are highest in bill measurements, it can be concluded that phenetic relationships are founded primarily on divergence in bill morphology. Each of the three groups extracted has both Afrotropical and non-Afrotropical representatives, indicating that the phenotypes are based on convergent similarity and are thus ecotypes. Phylogenetic analyses based on plumage fea¬ tures and behaviours split the genus Serinus ( sensu lato) into two sister groups, one consisting of Afrotropical seedeaters (genera Poliospiza and Dendrospiza ), the other including Afrotropical seedeaters and canaries ( Ochrospiza and Crithagra), Palaearctic canaries of the genus Serinus ( sensu stricto), rosefinches ( Erythrina ), siskins ( Spinus , Carduelis) and greenfinches ( Chloris ). Plesiomorphic character states in plumage features, as reflected in immature birds, prevail in the Afrotropical Region. Keywords: Phenetic relationships, cladistic relationships, skeletal variations, plumage colouration, behaviours, Carduelidae. INTRODUCTION The Carduelidae is comprised of about 140 seedeating species, with an almost world¬ wide distribution. Species richness is greatest in Holarctic (mostly Palaearctic) wooded and bushland biotopes. Siskins ( Spinus ) reach the Neotropics, and a few seedeaters occur in the Oriental Region. Nearly a third of the species occur within the Afrotropical Region, inhabiting dry and moist savannas. Of these, about 35 species are usually lumped within the genus Serinus ( sensu lato) together with several Palaearctic species. Attempts to sort out African Serinus spp. into species-groups have met with some success. On the basis of behavioural characters, five subgroups have been differen¬ tiated, calling into question the monophyly of Serinus (Nicolai 1960; van den Elzen 1985). The canaries ( Serinus ) are thus mainly distributed in the Palaearctic, one spe¬ cies ( Serinus canicollis) reaching the southern parts of the Afrotropical Region. Of the remaining species, 27 form (according to courtship display, nestbuilding behaviour and begging calls of the nestlings) four species groups, sometimes separated as four genera: Ochrospiza (10 species: leucopygia, menachensis, reichenowi, xantholaema, atrogularis, citrinipectus, mozambica, dorsostriata, xanthopygia and rothschildi)', Dendrospiza (6 species: citrinelloides, hyposticta, koliensis, scotops, capistrata, frontalis ); Crithagra (4 species: flaviventris, sulphurata, donaldsoni, albogularis ), Poliospiza (7 species: striolata, burtoni, leucoptera, reichardi, mennelli, gu laris, tristriata), and two genera of uncertain affinity, Pseudochloroptila and Alario. Genetic distances based on biochemical analyses of proteins and enzymes in seedeating passerines (Stempel 1986) confirmed the closer relationships of Afrotropical species ( Ochrospiza leucopygia, O. reichenowi, O. mozambica, Poliospiza leucoptera), and 460 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI included with them the Eurasian Greenfinch Chloris chloris and Bullfinch Pyrrhula pyrrhula, rather than two traditionally recognized Serinus spp. ( serinus and canaria var. dom.). In order to elucidate systematic relationships between African species groups, their affinities to the European stock and their adaptive radiation within the Afrotropical Region, two approaches were applied. Phenetic analyses of skeletal measurements were used to study osteological divergence, and phylogenetic analyses of plumage pattern and colouration (including some soft parts) were used to reconstruct phylogenetic histories. METHODS Phenetic methods Phenetic relationships were estimated using a UPGMA (unweighted pair group method using arithmetic averages), Q-mode cluster analysis (Sneath & Sokal 1973). Matrices of correlation coefficients between 44 skeletal measurements of 44 species (49 OTUs, including three species with two subspecies each and two crossbreds) were the basis of all analyses. Matrices were either derived from raw or transformed data (logs and/or partly z-standardized), with all measurements being divided by fe¬ mur length as a standard. Arithmetic averages of measurements represented species. For more details on morphometries, see van den Elzen et al. (1987) and Nemeschkal & van den Elzen (1990). Phylogenetic methods For the reconstruction of phylogenetic relationships, cladistics is the method of choice. According to cladistic principles only synapomorphies (shared derived character states) contain information about speciation events, and thus are indicators of monophyly (Hennig 1966, Wiley 1981, Ax 1984). Synapomorphies are defined by character polarizations, that is the direction each particular character is thought to have taken during evolution. Two approaches are widely accepted in determining character polarity (Watrous & Wheeler 1981, Crowe 1988): ontogenetic and out-group comparison. Character states possessed by immature phases and/or by sister groups (taxa of more distant relationship than the unit under study) are thought to represent plesiomorphic conditions. Our analyses mainly employed the ontogenetic approach. Nestling plumages in carduelids coincide in some general characteristics, e.g. heavy striation, brownish colours, light bars on the wings, two light spots at cheeks and under the eyes, wing and tail feathers bordered with light margins. They are thought to represent plesiomorphic character states, the condition present in the hypothetical ancestor. Forty-four plumage and one egg colouration characters were employed in this analysis. Thirty-three of the plumage characters were polarized on the basis of nestling phenotype. The remaining characters were unordered. For comparative pur¬ poses, only species were included that were also analyzed in the phenetic approach. Character polarizations are summarized in Table 1. Analyses were done with the computer programs PAUP (Swofford 1985) and HENNIG86 (version 1.5: developed by J.S. Farris), that use the parsimony criterion to derive an estimated tree of mini¬ mum character transformation steps (Wiley 1981). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 461 RESULTS AND DISCUSSION Phenetics The UPGMA analysis clustered species into three broad groupings (Figure 1). Clus¬ ters A and B represent smaller and medium sized species (wing lengths 62-80 mm). The first cluster of smaller species (A), comprising a phenotype with blunt bills and shorter legs, is further subdivided into two subclusters, one of Palaearctic species ( Serinus ( sensu stricto), including the Afrotropical S. canicollis and Alario, the Linnet Acanthis cannabina and the Long-tailed Rosefinch Uragus sibiricus)), the other smaller Afrotropical species ( Ochrospiza spp. and Poliospiza mennelli). Cluster B unites small to medium sized species characterized by pointed bills and longer hindlimbs. It includes both Palaearctic and Neotropical siskins ( Spinus spp.), the siskin-like serins ( Dendrospiza ), several Poliospiza spp., the Cape Siskin Pseudochloroptila totta, the Redpoll Acanthis fiammea and the European Goldfinch Carduelis carduelis. Cluster C unites all species with heavy bills and pronounced mandibles. All larger species are located within this cluster together with smaller, but massively billed species with wing lengths ranging from 66-105 mm. Palaearctic rosefinches ( Erythrina spp.) and greenfinches ( Chloris spp.) are combined in several subunits with Afrotropical seedeaters ( Crithagra and Poliospiza ) and the Pirol Finch Linurgus olivaceus. Character divergence Phenetic distance (Camin & Sokal 1965) measures character divergence by the range of the character over the group under study. To verify the amount each character contributes to the phenetic analysis, we compared the coefficients of variation (stand¬ ard deviation as percentage of the arithmetic mean of all species in each measure¬ ment) of all measurements. Generally, the skeletal measurements under considera¬ tion can be assigned to three functional complexes: bill and skull elements to the feed¬ ing complex, leg and pelvis elements to the hindlimb-locomotion complex, and ele¬ ments from wing and shoulder girdle to the flying-locomotion complex. Within these functional complexes, coefficients of variation (CV) were highest in the feeding com¬ plex and, within this complex, in bill measurements. The CVs of bill measurements (13.40%, average of 7 measurements) were double those of skull measurements (7.80%, average of 11 measurements), pectoral measurements (6.99%, average of 8 measurements) and pelvis measurements (6.83%, 4 measurements). They were almost three times as large as the CVs of leg measurements (4.98%, 3 measure¬ ments), and as large as those for measurements of wing bones (13.78%, 6 measure¬ ments). Thus, variation in bill dimensions plays a very important role in the carduelid phenotype. Carduelids are thought to be a phylogenetically young clade, considered by some authors as a family mainly characterized by their feeding habits, especially the abil¬ ity to cut open and husk dicotyledon seed. Siskin-types employ several foraging tech¬ niques and, especially the smaller species, can cling to vegetation. In the species sample chosen, the number of Afrotropical species almost equals the number of non- Afrotropical carduelids. They are also evenly distributed within the three main clus¬ ters of the UPGMA analysis. (Cluster A: 1 1 Afrotropical vs 7 non-Afrotropical species; cluster B: 8 vs 7; C: 9 vs 8). A Komolgorov-Smirnov Two-Sample Test was employed to test whether measurements in the Afrotropical samples had the same statistical distribution as measurements in residual species. Significant congruence exhibited 462 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI TABLE 1 - Plumage features, egg colour and ethological characters used in a phylogenetic analysis of carduelid species. character 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 species 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 2 3 3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 5 5 5 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 Dhypo 0 1 1 0 1 1 1 0 1 2 0 1 6 0 0 1 0 ■0 2 0 0 0 0 0 0 1 1 0 0 0 0 2 2 1 0 0 1 0 2 1 0 1 0 1 1 1 3 1 2 1 1 0 Dscot 1 1 1 0 1 1 1 0 1 2 0 1 4 0 1 1 0 0 2 0 0 0 0 0 0 1 1 0 0 0 0 2 2 2 0 0 1 0 1 1 0 1 0 1 1 1 3 1 2 1 1 0 Dcapi 1 2 1 0 1 1 1 0 1 2 0 1 2 0 0 1 1 0 2 0 0 0 0 0 0 1 1 0 0 0 0 2 2 2 0 0 1 0 1 1 0 1 0 9 9 1 3 9 9 9 9 0 PStot 0 0 0 1 1 1 1 0 2 3 0 1 4 1 1 2 1 0 1 1 1 1 1 1 0 3 1 0 0 0 1 2 1 1 0 0 0 0 1 2 1 1 0 1 1 1 3 1 0 1 0 1 Pstri 0 0 0 0 0 0 0 0 0 2 0 1 1 1 1 0 0 0 3 0 0 0 0 0 0 1 1 0 0 0 0 2 2 3 0 0 0 0 2 1 1 1 0 0 2 0 3 1 2 4 0 0 Ptris 0 0 0 1 1 1 1 0 0 2 0 1 1 0 1 0 1 0 3 1 0 1 1 0 0 4 1 0 0 0 0 2 3 1 0 0 0 0 2 2 1 1 0 0 2 0 3 1 2 4 0 0 Pleuc 0 0 0 1 1 1 1 0 0 2 0 1 1 1 1 0 1 0 3 0 0 0 1 0 0 1 1 0 0 0 0 2 2 1 0 0 0 0 2 2 9 9 9 0 2 0 3 9 9 4 9 0 Pmenn 0 0 0 1 1 1 1 0 0 2 0 1 1 0 1 0 0 0 3 1 0 0 0 0 0 1 1 0 0 0 0 2 3 1 0 0 0 0 1 1 9 9 9 0 2 0 3 9 9 4 9 0 Pburt 0 0 0 2 1 1 1 1 0 2 0 1 6 0 0 0 1 0 3 1 0 0 1 0 0 1 1 0 0 0 0 2 2 1 0 0 1 0 2 2 9 9 9 9 9 9 9 9 9 4 9 0 SEfla 1 2 1 0 0 0 0 0 1 2 1 2 6 1 0 1 1 0 1 1 0 0 0 0 0 1 1 0 0 0 0 2 2 3 0 0 0 0 1 1 1 1 0 0 2 0 3 1 0 4 0 0 Calbo 0 0 0 0 0 0 0 0 0 2 1 2 1 1 1 0 1 0 2 1 0 0 0 0 0 1 1 0 0 0 0 2 2 3 0 0 1 0 2 2 1 1 0 0 2 0 3 1 0 4 0 0 Csulp 1 2 1 0 0 0 0 0 1 2 1 2 4 1 1 1 1 0 2 1 0 0 0 0 0 1 1 0 0 0 0 2 2 3 0 0 1 0 2 2 1 1 0 0 2 0 3 1 0 4 0 0 Cdona 0 1 1 0 1 1 1 0 1 2 1 2 6 1 0 1 0 0 2 1 0 0 0 0 0 1 1 0 0 0 0 2 2 1 0 0 0 0 2 1 9 9 9 9 9 9 9 9 9 4 9 0 Spusi 2 2 2 3 1 1 1 1 1 1 1 2 2 0 1 1 0 0 4 0 1 0 0 0 0 1/ 1 0 1 0 0 2 2 1 0 0 0 0 2 2 0 1 1 1 1 1 2 0 0 2 1 0 Sseri 2 2 1 0 0 0 0 0 1 1 1 2 4 0 0 1 0 0 4 0 1 0 0 0 0 1 1 0 1 0 0 2 2 3 0 0 0 0 2 2 1 1 1 1 1 1 2 0 0 2 1 0 Ssyri 2 2 0 0 1 0 1 1 1 1 1 2 4 1 1 1 1 0 4 1 1 1 1 0 0 1 1 0 1 0 0 2 2 2 0 0 9 9 2 2 9 1 9 9 9 1 2 0 0 2 9 0 Scani 2 2 0 3 1 0 1 1 1 2 1 2 6 1 0 1 1 0 4 1 1 1 0 0 0 1 1 0 1 0 0 2 2 1 0 0 0 0 2 2 1 1 9 1 1 1 2 0 0 2 1 0 Scana 2 2 1 0 0 0 0 0 1 1 1 2 6 0 0 1 1 0 4 0 1 0 0 0 0 1 1 0 1 0 0 2 2 3 0 0 0 0 2 2 1 1 1 1 1 1 2 0 0 2 1 0 Aalar 2 2 2 3 1 1 1 1 2 2 0 1 2 1 2 2 1 0 2 1 0 1 1 1 0 4 1 0 1 1 0 3 3 2 0 0 0 0 1 1 1 1 9 1 1 1 3 0 0 2 0 0 Oatro 0 0 0 0 0 0 1 0 0 2 1 2 1 0 0 0 0 1 2 1 0 0 0 0 0 1 1 0 000212000 Oleuc Omoza Odors 0 0 0 0 1 2 0 0 1 2 0 0 0 0 0 0 0 0 1 2 1 0 1 0 0 2 0 2 0 1 1 1 0 1 2 3 0 0 2 3 1 0 0 0 0 2 0 2 1 2 1 0 1 1 0 0 0 0 3 1 3 0 0 1 0 0 1 0 0 1 0 0 1 1 0 0 0 0 0 1 1 0 1 1 1 0 1 2 0 1 2 4 2 2 2 2 1 0 1 1 1 0 1 0 1 4 0 0 1 4 0 0 1 0 0 0 0 0 2 2 0 0 1 1 2 2 2 1 2 1 2 0 0 1 0 2 2 2 1 2 1 2 2 1 4 0 3 10 1 1 1 4 0 0 0 0 0 1 1 0 0 3 10 1 0 0 0 0 0 1 1 0 1 0 0 0 0 1 1 0 1 2 1 0 1 1 0 0 1 0 1 0 3 10 1 0 0 0 0 4 Oxant 1 0 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 463 CHchIO 0 0 2 1 1 1 1 1 3 1 2 6 1 1 1 1 0 2 1 1 1 1 0 1 2 2 1 1 2 0 2 2 1 1 1 1 0 2 2 1 1 9 0 1 0 2 1 0 1 1 0 CHsinl 2 0 3 1 1 0 1 2 3 1 2 6 1 1 2 1 0 2 1 1 1 1 2 1 3 2 1 1 2 0 2 2 1 1 1 9 9 2 2 1 1 9 0 1 0 2 1 0 1 1 0 CHspi2 2 2 3 0 0 1 0 1 3 1 2 6 1 0 1 1 0 2 1 1 0 1 2 1 2 2 1 1 2 0 2 2 3 1 1 1 0 1 2 1 1 9 0 1 0 2 1 0 1 1 0 Loliv 2 2 2 3 1 1 1 1 1 3 0 1 2 1 0 1 1 0 1 1 0 0 1 0 0 3 1 0 0 0 0 2 3 2 1 1 9 9 1 2 9 9 9 9 9 9 9 9 9 9 9 0 SPcuc 2 2 2 3 1 1 1 1 2 2 1 2 2 1 0 2 1 0 4 1 0 0 0 2 1 2 2 1 1 2 0 2 2 2 0 0 1 1 1 2 1 1 1 1 1 1 2 0 1 3 1 0 SPbar 2 2 2 3 1 1 1 0 1 1 1 2 3 0 0 1 0 0 4 0 0 0 0 2 1 2 2 1 1 2 0 2 2 1 0 0 0 0 1 2 1 1 1 1 1 1 2 0 1 3 1 0 LEarc2 2 0 3 1 1 1 1 2 4 1 3 2 2 1 2 2 0 2 0 0 0 0 0 0 2 4 0 1 0 0 3 3 1 1 0 0 1 1 2 1 0 9 9 9 9 9 9 9 9 9 1 Eeryt 2 2 2 3 1 1 1 1 2 4 1 3 5 0 1 2 0 0 2 0 0 0 0 0 0 3 4 0 0 0 0 3 3 1 0 1 0 0 1 1 1 0 0 0 1 1 3 0 0 1 9 2 Emexi 2 2 0 2 1 1 1 0 2 4 1 3 5 0 1 2 0 0 2 0 0 0 0 0 0 4 4 0 0 0 0 3 3 1 0 0 0 0 1 1 1 0 0 0 1 1 3 0 0 1 0 2 Usibi 2 2 0 0 1 1 1 1 0 1 1 3 6 0 1 2 0 0 2 0 0 0 0 0 0 4 1 0 O O 0 2 1 1 0 0 0 0 1 1 1 0 0 0 1 9 3 0 0 1 9 2 PRvinO 2 2 2 1 1 1 0 2 4 1 3 6 2 0 2 2 0 1 0 1 1 1 1 0 3 1 0 0 0 0 3 3 1 0 0 0 0 1 1 9 9 9 9 9 9 9 9 9 9 9 2 COcco 2 2 2 3 1 1 1 1 2 3 0 1 2 1 0 2 1 0 1 1 0 1 1 0 0 4 3 0 0 2 1 3 1 1 1 1 0 1 2 2 0 0 0 0 9 0 2 0 0 1 9 0 Aflam 2 2 0 0 1 1 1 1 0 1 1 3 3 0 0 0 0 0 2 0 0 0 0 0 0 2 1 0 1 0 0 2 1 1 0 0 0 0 2 1 0 1 1 1 1 0 3 0 0 1 1 0 Acann 2 2 0 3 0 0 1 1 2 1 1 3 6 0 1 2 2 0 2 0 0 1 1 0 0 2 3 0 1 0 0 1 1 1 0 0 0 0 1 2 9 0 9 0 1 9 9 9 9 9 9 0 CArdu 2 2 2 3 1 1 1 1 2 3 1 4 5 1 2 1 0 2 1 0 1 2 1 3 2 1 1 2 1 3 3 2 1 1 0 0 2 2 1 1 9 1 1 0 3 0 1 3 9 0 Characters and character polarization (9= no comparison): 1. forehead (0-uniform); 2. forehead colouration (0-brown); 3. crown colouration (0-brown); 4. crown pattern (O-striated); 5. ear patch (0-light spot); 6. eye ring (0-present); 7. moustachial stripe (0-present); 8. supercilium (0-present); 9. back colouration (0-brown); 10. back pattern (O-striated); 11. uppertail coverts (0-like back); 12. uppertail coverts colouration (unordered); 13. throat pattern (unordered); 14. flanks (O-striated); 15. breast (0- not contrasting throat and belly); 16. breast colouration (0-brown); 17. breast pattern (O-striated); 18. necklace (0-not present); 19. belly colouration (unordered); 20. undertail coverts (O-striated); 21. undertail coverts colouration (0-not contrasting with belly); 22. wing bar (0-present); 23. wing coverts (O-striated); 24. colouration of primaries (0-uniform brown); 25. pattern of primaries (0-uniform); 26. pattern of secondaries (rear edge) (unordered); 27. colouration of secondaries (unordered); 28. pat¬ tern of secondaries (unordered); 29. tail feathers (0-equal); 30. tail pattern (0-uniform brown); 31. rear edge of tail (0-plain); 32. pattern in single tail feathers (unordered); 33. tail colouration (unordered); 34. facial pattern (unordered); 35. bill colouration (0-dark horn); 36. leg colouration (0-dark horn); 37. colouration of juvenile plumage (0-brown); 38. pattern of juvenile plumage (0-heavy striated); 39. sexual dimorphism (unordered); 40. female plumage (unordered); 41. gape colouration of nestlings (0-red- blue-spots); 42. gape rictal colouration of nestlings (0-whitish); 43. bill pattern of nestlings (0-no pat¬ tern); 44. nestbuilding (0-solely female); 45. display (unordered); 46. courtship display (0-fluffing-tail up); 47. wing posture during courtship display (unordered); 48. begging calls (0-simple); 49. location call (0-simple); 50. contact call (unordered); 51. nest sanitation (0-nest clean); 52. egg colouration (0- blueish-white). Abbreviations and species list: Dhypo - Dendrospiza hyposticta and (Dkol) D. ko!iensis\ Dscot - D. scotops ; Dcapi - D. capistrata: PStot - Pseudochloroptila totta\ Pstri - Poliospiza striolata and (Pgul) P. gularis ; Ptris - P. tristriata ; Pleuc - P. leucoptera ; Pmenn - P. mennelli ; Pburt - P. burtoni ; SEfla - Serinops flaviventris ; Calbo Crithagra albogularis ; Csulp - C. sulphurata ; Cdona (Cdonbuc) C. (donaldsoni) buchanani ; Spusi - Serinus pusillus', Sseri - S. serinus\ Ssyri - S. syriacus', Scani - S. canicollis and (Scit) S. citrinella; Scana - S. canaria ; Aalar - Alario alario ; Oatro Ochrospiza atrogularis and (Orei) O. reichenowi ; Oleuc - O. leucopygia ; Omoza - O. mozambica ; Odors - O. dorsostriata ; Oxant - O. xanthopygia ; CHchl- Chloris chloris ; CHsin - C. sinica; CHspi - C. spinoides ; Loliv - Linurgus olivaceus ; SPcuc - Spinus cucullatus ; 464 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI SPbar - S. barbatus and (SPspin) S. spinus ; LEarc (LEarclit) - Leucosticte ( arctoa ) littoralis ; Eeryt - Erythrina erythrina ; Emexi - E. mexicana\ Usibi - Uraqus sibiricus ; PRvin Procarduelis vinacea\ COcco - Coccothraustes coccothraustes ; Aflam - Acanthis flammea ; Acann - Linaria cannabina ; CArdu (CAcar, CAcartschu) - Carduelis carduelis, C. c. tschusii. between all measurements (P < 0.001) demonstrates that adaptive trends of morpho¬ logical traits in the two groups converge, despite their inhabiting two biogeographical regions with different biotopes. This confirms that adaptive radiation in carduelids apparently always involves bill diversification. For example, grosbeak-ecotypes are represented in the Afrotropical Region as well as in the Palaearctic (European Grosbeak Coccothraustes coccothraustes and African grosbeak seedeaters Crithagra donaldsoni , C. d. buchanani). This applies also to siskin-like birds (European Gold¬ finch Carduelis carduelis, siskins Spinus spp. versus African Citril Dendrospiza hyposticta and Cape Siskin Pseudochloroptila totta). Thus, phenetic relationships as exhibited in the UPGMA analysis reflect mainly adaptive trends in morphology, prima¬ rily bill dimensions, of carduelid species. Species in cluster A exhibit the serin- ecotype, cluster B the siskin-ecotype and C the seedeater-grosbeak-ecotype. That dif¬ ferences in size and bill morphology are the main characters that allow sympatric species to coexist has been verified by principal components analyses (van den Elzen et al. 1987, Nemeschkal & van den Elzen in press). Phylogenetics A first analysis of unweighted plumage and egg-colour characters yielded a cladogram with a very low consistency index, suggesting that some of the features coded are adaptations to similar ecological conditions and are plagued by convergence to a simi¬ lar extent as are skeletal traits. Loss of distinct plumage patterns, marked colours and striation is apparently favoured in open-country biotopes. For example, in several areas of the Afrotropical and Palaearctic Regions, uniformly grey to brownish birds are most commonly found in open habitats, green to yellow coloured species in denser vegetation. Only in this latter group are plumage patterns exhibited in courtship dis¬ plays, with savanna and semidesert-dwelling species accentuating rather postures and possibly vocalizations. In order to overcome this high level of homoplasy, plumage characters with high con¬ sistency indices were weighted 3-5 greater than those with low CIs. To this data set were added seven behavioral characters (Nicolai 1960, van den Elzen 1985 - Table 1) four of which were polarized using the ontogenetic criterion. This analysis resulted in a cladogram (Figure 2) which is thought to illustrate the best assumption of phylogenetic relationships between the species studied, since species groups of known affinities are united and the cladistic distribution pattern of species (with the exception of C. coccothraustes) coincides with biogeographical distributional patterns of the species and genera included (Hall & Moreau 1970). Behaviours and plumage patterns rank Poliospiza and Dendrospiza species together with Coccothraustes coccothraustes and three monotypic African genera, Alario, Pseudochloroptila and Linurgus as a sister group of the remaining Afrotropical, Palaearctic and Neotropical species (Figure 2). Within this second sister group, both greenish Ochrospiza species ( mozambica and dorsostriata) are separated from the remaining greyish-brown mem¬ bers of that genus. They form the sister group to Crithagra and the remaining Ochrospiza species on the one hand, and all members of the Palaearctic-Neotropical stock (Serin us, Acanthis, Leucosticte, Erythrina, Procarduelis, Spinus, Carduelis and Chloris) on the other. Neither phylogenetic analysis (using several species as outgroups) split Serinus ( sensu lato) into the distinct species groups indicated by a ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 465 comparison of unpolarized behavioural parameters, or placed African species closer to Chloris than to Serinus as indicated by genetic distances (Stempel 1986). Behav¬ ioural characters place Alario within Serinus. The two other monotypic genera linked to it in the cladogram are of unknown affinity according to their autapomorphic ( Pseudochloroptila ) or unknown ( Linurgus ) ethological attributes (van den Elzen 1985). S pus Sser Ssyr Scit Sser x Scana Scani Aala Scana A cann Usib O rei Oatr Oleu O moz O xan Pmen Odor x Omoz Odor Dhyp 1 PStott Pgul Ptri A flam SPspin SPbar CAcar CA cartschu Pstrio Calb Pleu Loli PR vin Cdonbuc Csul Csul Pbur Ccoc Cfla LE arc lit Eery CH chi CHsin CHspin E mex r 0 0.5 1 FIGURE 1 - Phenogram of 49 carduelids (44 species, 3 subspecies, 2 crossbreds; UPGMA based on skeletal measurements). 466 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI — hypanc Pstrio — Pmen Ptri Pleu — Pbur E Dhyp — PStott Dcap D sco L oti E A ala Ccoc E Oatr — Calb Oxan O leu Csul E Cfla C don E Odor O moz Scana Sser — Ssyr Scani — S pus — A flam - LE arc - Eery - E mex - U sib - PRvin - Acann SP bar r— SPcuc CA car CH spin E CH chi CH tin FIGURE 2 - Cladogram of 39 carduelids (five redundant species excluded; HENNIG 86 consensus tree based on plumage characters and behaviours.) .1ml Biogeographical and taxonomic considerations In the carduelid species investigated here, plesiomorphic plumage features and be¬ haviours prevail in the Afrotropics, the least anagenetic evolution being found in Poliospiza, and partly in Ochrospiza and Crithagra spp. Apomorphic plumage and behaviours prevail in non-Afrotropical species, being most distinctive in siskins and greenfinches. There are essentially two contradictory approaches employed to corre¬ late evolutionary character states of species with their distributional patterns. Accord¬ ing to the classical Darwinian centre of origin concept and the progression rule, ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 467 phylogenetically older species, exhibiting plesiomorphic character states are distrib¬ uted in ancestral centres (Myers & Giller 1988). In the vicariance model, species bear¬ ing plesiomorphic features represent ancestral forms, but apomorphic character states of species predominate in developmental centres (Croizat et al. 1974). Ecological concepts link plesiomorphic phenotypes of species with ancestral, unchanged ecologi¬ cal conditions within biotas. The Carduelidae may thus have either dispersed from the Afrotropical Region into the Palaearctic or invaded from the Palaearctic and retained within the Afrotropics, ancestral-like habitats. At least two invasions affected Africa. This paper does not aim at an update of nomenclature in carduelids. In general, ge¬ neric names applied coincide with species sets in the cladogram, with the exception of Ochrospiza. The grey-coloured species are disassociated from their greenish con¬ geners 0. mozambica and 0. dorsostriata, which have been placed between Crithagra and Serinus. Based on differences in colouration, the two species have been sepa¬ rated as Microserinus (Roberts 1922). Cladistic analyses support conclusions drawn from comparative ethology, that the genus Serinus ( sensu lato) as usually applied is paraphyletic, involving at least two species assemblages. LITERATURE CITED AX, P. 1984. Das phylogenetische System. Systematisierung der lebenden Natur aufgrund ihrer Phylogenese. Stuttgart, Gustav Fischer Verlag. CAMIN, J.H., SOKAL, R.R. 1965. A method for deducing branching sequences in phylogeny. Evolu¬ tion 19: 311-326. CROIZAT, L., NELSON, G., ROSEN D.E. 1974. Centers of origin and related concepts. Systematic Zoology 23: 265-287. CROWE, T.M. 1988. Molecules vs morphology in phylogenetics: a non-controversy. Transactions of the Royal Society of South Africa 46: 317-334. ELZEN, R. VAN DEN 1985. Systematics and evolution of African canaries and seedeaters (Aves:Carduelidae). Pp. 435-451 in Schuchmann, K.-L. (Ed.). Proceedings of the International Sym¬ posium on African Vertebrates. Bonn, Museum Alexander Koenig. ELZEN, R. VAN DEN, NEMESCHKAL, H.L., CLASSEN, H. 1987. Morphological variation of skeletal characters in the bird family Carduelidae: I. General size and shape patterns in African canaries shown by principal component analyses. Bonner Zoologische Beitraege 38: 221-239. HALL, B.P., MOREAU, R.E. 1970. An atlas of speciation in African passerine birds. London, Trustees British Museum (Natural History). HENNIG, W. 1966. Phylogenetic systematics. Urbana, University of Illinois Press. MYERS, A. A., GILLER, P.S. 1988. Analytical biogeography. New York, Chapman & Hall. NEMESCHKAL, H.L., VAN DEN ELZEN, R. 1990. Funktionskreise im Skelettsystem der Cardueliden - Ein morphometrischer Ansatz. Pp. 37-41 in van den Elzen, R., Schmidt-Koenig, K., Schuchmann, K.-L. (Eds). Current Topics in Avian Biology. Proceeding of the 100th Meeting of the Deutsche Ornithologen-Gesellschaft. Bonn, Deutsche OrnithologenGesellschaft. NEMESCHKAL, H.L., VAN DEN ELZEN, R. in press. Morphological divergence, distribution and sys¬ tematics of canaries of the Genus Ochrospiza. Proceedings of the Seventh Pan-African Ornithologi¬ cal Congress. NICOLAI, J. 1960. Verhaltensstudien an einigen afrikanischen und palaearktischen Girlitzen. Zoologisches Jahrbuch fuer Morphologie und Systematik 87: 317-362. SNEATH, P.H.A., SOKAL, R.R. 1973. Numerical taxonomy. San Francisco, Freeman and Co. STEMPEL, N. 1986. Proteinvariation und Taxonomie krnerfressender Singvoegel. Zeitschrift fuer Zoologische Systematik und Evolutionsforschung 25: 281-308. SWOFFORD, D.L. 1985. Phylogenetic analysis using parsimony, version 2.4. Champaign, Illinois Natural History Survey. WATROUS, L.E., WHEELER, Q.D. 1981. The out-group comparison method of character analysis. Systematic Zoology 30: 1-11. WILEY, E.O. 1981. Phylogenetics. New York, John Wiley & Sons. ij I 468 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI PICIFORM AFRO-ASIAN ZOOGEOGRAPHY AND SPECIATION L. L. SHORT1 and J. F. M. HORNE2 1 Ornithology Department, American Museum of Natural History, Central Park West at 79th Street, New York, N.Y. 1 0024-51 92, USA 2 Ornithology Section, National Museums of Kenya, P.O. Box 40658, Nairobi, Kenya ABSTRACT. Three piciform families present markedly different tropical Asian-African relations. 26 tropi¬ cal Asian barbets (three genera, two monotypic) have no near relatives among 42 Afrotropical barbets (seven endemic genera, one monotypic); connections are old and involve Calorhamphus and Megalaima with Gymnobucco, Stactolaema and Pogoniulus. The Afrotropics have 15 honeyguides of two tribes (four genera, three endemic, one monotypic); tropical Asia has but two species apt to be of recent origin, representing advanced species of Indicator. Picids number 52 in tropical Asia (14 gen¬ era, nine endemic, three monotypic), versus 26 Afrotropical species (six genera, three endemic, one monotypic); the regions share a forest genus (Sasia), wrynecks (Jynx), and species of Picoides (lat¬ ter two reach other regions). The two regions have remarkably different picofaunas, with two recent invasions indicated. Both regions have as strong or stronger ties to the Neotropics than to each other. Keywords: Piciformes, woodpeckers, barbets, honeyguides, systematics, zoogeography, tropical Asia, Afrotropics, Picidae, Capitonidae, Indicatoridae. INTRODUCTION Piciform birds including the woodpeckers (Picidae), barbets (Capitonidae) and honeyguides (Indicatoridae) occur in tropical Africa and Asia. Of these, the Indicatoridae are Palaeontropical, the Capitonidae are pan-tropical and the Picidae are Holarctic, Palaeontropical and Neotropical. The comparison of tropical African and Asian (we use “Asiotropics” to designate the Oriental or Indomalayan Region, as a simplification in accord with use of Afrotropics and Neotropics in the last decade) picofaunas is useful because of occurrence of the three families in both regions, and because these regions do share taxa, even down to the level of species, xthe lion ( Panthera leo) being one of the most notable. All families in the group treated herein are universally regarded as closely related, although relations of families are in dis¬ pute. All except the nest-parasitic honeyguides nest and roost in usually self-exca¬ vated cavities in trees, termitaria or the ground. We consider the barbets to be near the stem of the piciforms that we treat. The honeyguides have specialized features unique to them (e.g. bill hooks in hatchlings), but their behaviour is generally barbet¬ like, and they have none of the derived features marking the specialized woodpeck¬ ers (bill and tail modifications, foraging mode); if they evolved from woodpeckers (e.g. Sibley & Ahlquist 1985), then they did so at a time when ancestral picids had not yet become specialized, i.e. before the origin of the modern subfamilies Jynginae, Picumninae and Picinae of the Picidae. A general comparison shows these numbers of species in the two regions, with the Afrotropical number given first: Capitonidae (42 vs 26); Indicatoridae (15 vs 2); and Picidae (26 vs 51). Our taxonomy is that of Short & Horne (1985) and Morony et al. (1975) for barbets, Short & Horne (1988b) for honeyguides and Short (1982) for the woodpeckers. We consider no subgroups of barbets to be sufficiently derived to be ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 469 separated at the subfamily (Prum 1988) or family (Sibley et al. 1988) levels, except possibly the toucans (Ramphastinae) if they are placed in the same family as the barbets. RESULTS Capitonidae The barbets occur strictly within the limits of the tropics. Their diversity and numbers are greatest in the Afrotropics, unless toucans are considered as barbets, in which case the Neotropics virtually match the Afrotropics. There are seven endemic Afrotropical capitonid genera ( Gymnobucco , Stactolaema, Pogoniulus, monotypic Buccanodon, Tricholaema, Lybius and Trachyphonus) with species ranging in size from the 10 g tinkerbirds ( Pogoniulus ) to the large-billed, 108 g Lybius rolled and L. dubius (Short & Horne 1988a). Noteworthy is their diversity of foraging habits and correlated bill structure (pointed bill, mistletoe-eating Pogoniulus spp. to tooth-billed, frugivorous species of Lybius, the large species of which have heavy, grooved bills, and the long, narrow, pointed bill of omnivorous Trachyphonus spp.). Habitats occu¬ pied by Afrotropical barbets are also diverse, with over half the species shunning for¬ ests (which are the major habitat of Asiotropical and Neotropical barbets), and some occupying near desert situations (e.g. Tricholaema melanocephala), and bushed grassland, where typically feeding on the ground (ground-barbets, Trachyphonus spp.). Asiotropical barbets are far less diverse, with but three genera ( Megalaima , with 24 species and monotypic Calorhamphus and Psilopogon). These are forest species largely frugivorous in habits; a few species (e.g. M. haemacephala) occur in open second-growth woods, and more species forage in fruiting trees outside of forests in once-forested regions. They range in size from 32 g to 300 g (respectively M. haemacephala and M. virens). Songs of these barbets are simple hoot, pop or trill¬ ing repetitive, often loud notes, as in Afrotropical species of Gymnobucco, Stactolaema, Pogoniulus, Buccanodon, and Tricholaema ; none sings complex, often duetting songs such as those of Afrotropical Trachyphonus or Lybius spp. No genera of barbets are shared between these regions, and all genera are endemic to their region; thus, there have been no recent barbet movements between these regions, and connections are apt to have involved ancestral species of Asiotropical Calorhamphus and Meqalaima with those of Afrotropical Gymnobucco, Stactolaema and Pogoniulus at some unknown time in the past. Whether similarities in bill struc¬ ture and colouration between Calorhamphus and Gymnobucco are reflective of close relationship or convergence is open to question (Goodwin 1964, Prum 1988). Speciation events within Asiotropical barbets have involved past barriers, as for ex¬ ample the water barriers among Sundaland islands and Southeast Asia (e.g. the Megalaima armillaris and M. chrysopogon superspecies), insular separations (e.g. Ceylon from India, M. flavifrons and its relatives), and Himalayan montane forest separations (e.g. western M. zeylanica from eastern M. lineata by separation about Nepal). In the Afrotropics, there have been speciation events resulting from past sepa¬ ration of forest blocks by alternating wet and dry cycles (Mayr & O’Hara 1986) within Gymnobucco {peli and sladeni ) and Pogoniulus ( coryphaeus , leucomystax, simplex), but more important by far have been separation of woodland and bushland areas surrounding the forest to the north, east and south, i.e. subregion II and its provinces 470 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI and districts of Crowe & Crowe (1982). Most speciation in Stactolaema (a woodland offshoot of an ancestor in common with Gymnobucco), Tricholaema, Lybius and Trachyphonus, and even some species of Pogoniulus ( pusillus and chrysoconus, Short & Horne 1988a) has resulted from separation by forests or by xeric conditions of woodland segments north of the forest in West Africa, to the east of it in the Horn of Africa to East Africa, and to the south of it in southern Africa. Lake Chad in its past enlarged form likely separated Lybius dubius west of it and L. rolled \o its east. Vari¬ ous species of the Lybius toraquatus group show signs of evolution through past vicariance events (Short & Horne 1985, Crowe & Kemp 1988) around the forest block from West Africa ( vieilloti ) to the east ( leucocephalus , quifsobalito) and southward ( rubrifacies , chaplini, torauatus). Tricholaema leucomelas of southern bushlands in¬ terbreeds with T. frontatus of the miombo, and, very like leucomelas, allopatric diademata occurs in East Africa north of the other two. Indicatoridae Four genera and 15 species of honeyguides are found in the Afrotropics. The Asiotropics are home to but two species of honeyguides. Friedmann (1955) and Short & Horne (1988b) discussed relationships among species of this largely Afrotroplcal group. The honeybirds ( Prodotiscus ) are unlike other honeyguides except perhaps little known Melignomon in obtaining food as wax from the exudate of scale-insects (Hemiptera: Coccoidea), whereas species of Indicator ( nine) and virtually congeneric Melichneutes feed considerably on beeswax. Asiotropical honeyguides represent only the likely derived (yellow plumage features, rump markings, heavy bill, beeswax de¬ pendent) Indicator group. The somewhat different bill shape of Himalayan /. xanthonotus and Southeast Asian /. archipelagicus led Friedmann (1976) to conclude that they represent different invasions of the subgenera Melianothes ( xanthonotus ) and Indicator (archipelagicus) from Africa and Wong (1984) and Payne (1986) fol¬ lowed this up with vocal data, purportedly showing archipelagicus related to the varieaatusindicator complex of the Afrotropics. Foraging habits, displays and popu¬ lation dynamics of Afrotropical I. indicator, I. varieaatus, I. minor, and I. meliphilus (Short & Horne 1990) suggest that variegatus is distinctive, and the honeyguiding /. indicator is related as closely, or more so, to /. minor than it is to /. variegatus. Fur¬ thermore, /. indicator shares only with the two Asian species derived features of col¬ our pattern, namely, enhanced yellow-gold in the plumage, the golden yellow wrist patch (of indicator and archipelagicus, unique in Piciformes), and the rump patch (of immature /. indicator, in which the rump is white to creamy yellow-white, and of xanthonotus, in which it is orangish). Parsimony is better served by allowing for one honeyguide invasion of the Asiotropics (while honeyguides could have arisen in the Asiotropics, this would demand wholesale extinctions of all but the most advanced of honeyguides in that region), with differences between xanthonotus and archipelagicus arising through in situ divergence in the shifting Himalayan forests and lowland South¬ east Asian forests. Afrotropical honeyguides remain little known. Indicator pumilio and Melignomon eisentrauti have been described, /. narokensis elevated to species status then synonymized with I. meliphilus, I. conirostris separated from I. minor, and I. willcocksi separated from I. exilis, all within the past 35 years, and the breeding and hosts of one-third of the species are unknown, as indeed is the case for the Asiotropical spe¬ cies. Thus, speciation patterns within the Afrotropics are unclear, with only broad for¬ est-woodland allopatric speciation, if complete, accepted for I. conirostris-minor, and /. maculatus-variegatus, although such derivation also is likely for ancestral ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 471 Prodotiscus insignis-zambesiae (forest) and P. regulus (woodland). Since the major feeding of most Afrotropical honeyguides is on beeswax produced by a single subspe¬ cies of the Common Honeybee ( Apis mellifera), which could have arrived in Africa relatively recently (in geological terms), rapid speciation among Indicator and Melichneutes is a possibility, if not indeed a likelihood. As for using honeyguide songs as a key to phylogeny, this is not an easy character with which to deal. Songs are variable, and exceedingly alike between some species (e.g. Indicator conirostris-mi- nor, I. maculatus-variegatus , I. willcocksi meliphilus), but quite different in others (e.g. I. variegatus indicator, I. variegatus-minor, Short & Horne 1988b). Reversions to sim¬ ple songs, such as a trill, commonly occur in closely related barbets and, when they do take place, they afford no bases for comparisons (e.g. the simple trilled songs-du- ets of Trachyphonus vaillantii are unlike those of all its congeners, and the trilled, buzzy songs of Lybius minor and L. bidentatus provide no points of comparison with congeners, Short & Horne 1988a). These are vocally equivalent to totally melanic or albinistic plumage shifts with regard to comparisons of plumage pattern among con¬ geners. It seems likely that vocalizations other than songs may be more useful. As examples the chattery guiding call of /. indicator is heard from males of I. variegatus after copulation, and aggressive calls of I. minor rather closely resemble those of /. variegatus (pers. obs). Picidae Alone among these three piciform families, the Picidae occur in temperate areas, and show a range of adaptations remarkable when one considers their arboreal speciali¬ zation, for some species are successful on treeless plains (Short 1982). Nearly cos¬ mopolitan, they nonetheless are more speciose and more generally diverse in the tropics than in temperate areas, with the exception of the Afrotropics. They do not cross major water barriers, but they regularly reach near-shore islands, and are di¬ verse on oceanic West Indian islands not reached by barbets. A few picids ( Colaptes spp., Sphyrapicus spp,, Picoides hyperythrus, Jynx torquilla) are highly migratory, as are no barbets or honeyguides. Asiotropical picids are diverse, representing 14 genera and 52 species, compared with the other major centre of diversity, the Neotropics, with 1 1 genera in 88 species (Short 1982). There is a sub-centre of picid diversity in the Himalayan area, especially rep¬ resenting Picus and Picoides, and also Dryocopus, that are extensive in distribution throughout the Palaearctic {Picus) and even to the Neotropics ( Picoides , Dryocopus), as well as in Asiotropical mountains and lowlands (Short 1983). All three subfamilies are represented in the Asiotropics and the Afrotropics, but the latter has only one piculet ( Sasia africana, Picumninae) and its 24 species of Picinae represent only four genera, two of which ( Geocolaptes , Picoides) have but one species in that region. Shared species between the Afrotropics and Asiotropics include only the mainly Palaearctic, migratory Jynx torquilla and, among genera, only Sasia and Picoides (Short 1980, 1982; Short & Horne 1988c). The speciose Afrotropical genera Campethera and Dendropicos, and endemic monotypic Geocolaptes have no close Asiotropical relatives except for Picoides, likely derived from Dendropicos (Short 1980), but rather are tribally related to Neotropical woodpeckers (Short 1985). In par¬ ticular, Asiotropical Campephilini ( Dryocopus ), Picini {Picus, Dinopium Chrysocolaptes, Gecinulus, Sapheopipo, Blythipicus and Reinwardtipicus), endemic Meiglyptini {Meiglyptes, Hemicircus, Mulleripicus), and Celeus of the Colaptini, among the Picinae, and Picumnus among the Picumninae have no close relatives in the Afrotropics (Short 1982). 472 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Speciation among Afrotropical woodpeckers has been discussed elsewhere (Short 1980, Short & Horne 1988c). Woodland-grassland versus forest (e.g. Campethera abingoni superspecies, Dendropicos goertae superspecies, Dendropicos abyssinicus superspecies, and megasubspecies within Dendropicos fuscescens), east-west for¬ est block separation ( Dendropicos pyrrhogaster superspecies, Campethera maculosa superspecies and megasubspecies of C. cailliautii), montane forest isolation (megasubspecies of C. tullbergi), and woodland-woodland isolation ( C . punctuligera superspecies) represent major patterns in recent speciation, and these are in substan¬ tial agreement with barriers indicated by guineafowl (Crowe & Crowe 1982), by galliforms generally and by hornbills (Crowe & Kemp 1988), and by other diverse or¬ ders of non-passerines (Fry 1988). DISCUSSION Afrotropical problems We will not dwell on these, as all have been mentioned and some discussed (Short 1971, 1980, 1982; Short & Horne 1988c). Serious problems are: 1) the Neotropical relationships of Afrotropical Picidae in contrast to a) sparse relations of the latter with those of the Asiotropics, b) distant relations of Neotropical and Afrotropical barbets, and c) lack of honeyguides in the Neotropics; 2) the presence of a restrictedly forest piculet ( Sasia africana) having congeneric relatives in the Asiotropical forests, in con¬ trast to sparse picine and distant capitonid relations between the Afrotropics and Asiotropics; and, 3) the apparently recent derivation in the Asiotropics, as far away as Malaysia (where but 13% of lowland forest avian genera are held in common with the wet lowland Afrotropical forest, Wells 1988), from a speciose, advanced genus of Afrotropical honeyguides, with no concomitant barbet, and few possible picid connec¬ tions, even by dry woodland and scrub taxa. Here we concentrate on those problems concerning Afrotropical-Asiotropical connec¬ tions. In the case of the piculet Sasia africana, morphology and behaviour certainly indicate that it is congeneric with Asiotropical S. abnormis and S. ochracea despite its having a (very thin) hallux that is lacking as a derived feature of the Asiotropical species. One assumes that their common ancestor occupied woodland or bushland habitat (unless something of an array of forest sites occurred in an “Arabian pier”, Lees-Smith 1986:83), unlike modern species but not unknown in Neotropical species of related Picumnus (e.g. cirratus, Short 1982). This begs the question, to which we have no answer, of why, if Sasia could invade the Afrotropics from Asia, there were not more picid and at least some capitonid invasions of Asia from Africa, given the xeric habitats in the Afrotropics of species of Dendropicos, Campethera, Tricholaema, Pogoniulus and Trachyphonus ? We believe that Picoides originated from Afrotropical Dendropicos, as we discuss below. The only Afrotropical species of Picoides (P. obsoletus) occupies wooded grassland and woodland of the Sahel and highland East Africa, but its entry into Africa must have been relatively recent for its morphology is that of the canicapillus-minor group of Picoides and is unlike that of more Dendropicos- 1 ike P. temminckii and P. maculatus (see below). It seems that: 1) Jynx, if it evolved in the Afrotropics, where both species occur, was able to exit to the Palaearctic and fringe of the Asiotropics; 2) an early form of Dendropicos entered the Asiotropics from Africa and, after further evolution there, ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 473 gave rise to a species able to enter (re-enter) the Afrotropics; 3) Afrotropical- Asiotropical barbet connections occurred long ago, perhaps at the time when Dendropicos managed to reach the Afrotropics as nascent Picoides spp.; and, 4) likely one species of Indicator was able to reach the Asiotropics rather recently, there to become adapted to forest conditions as it spread and differentiated. All of these are subject to review when more data are available on the palaeobotany of the region from the Horn of Africa and the Arabian Peninsula to Pakistan. Lowland piciforms endemic to that critical area number only the picids Picoides dorae of the Arabian Peninsula and P. assimilis of Iran-Pakistan, both of which represent the advanced, redbellied, major group of Picoides (Short 1982). Asiotropical problems The Asiotropical barbets are not related closely to Afrotropical barbets, but all have the repetitive, simple vocalizations typical of Afrotropical Gymnobucco, Stactolaema, Tricholaema and Pogoniulus (but not of Lybius and Trachyphonus). Connections are deemed to be ancient, and perhaps are no closer than they are with Neotropical barbets (Short 1985). The honeyguides present only the problem of one or two inva¬ sions from the Afrotropics, and, as we have discussed above, evidence supports the view of one, relatively recent invasion, with subsequent adaptation to forests, xanthonotus those of the Himalayas, and rare archipelagicus, of rainforests in Greater Sundaland, now restricted to the lowlands. We note that Asiotropical barbets and woodpeckers are larger on average than those of the Afrotropics, and thus many are immune to honeyguide parasitism. The enigmatic presence in the Asiotropics of a typically patterned (the tail pattern is unique in piciforms) species of otherwise Neotropical Picumnus (22 species there), and of a species of otherwise Neotropical (10 species) Celeus evades easy explana¬ tion and seems to require relatively recent, ex-tropical distribution in the Palaearctic and Nearctic. Palaearctic Dryocopus martius and Asiotropic D. iavensis have Nearctic (one) and especially Neotropical (three) relatives. Restriction of Afrotropical relations to Sasia africana (discussed above) and Picoides obsoletus is difficult to explain. The latter relates to derivatives of Dendropicos in Sulawesi Picoides temminckii and Phil¬ ippine P. maculatus, both yellow-shafted in part, a feature found otherwise among piciforms only in many Neotropical Colaptini and Afrotropical Campetherini, including Dendropicos. P. obsoletus itself seems to represent a less barred and streaked de¬ rivative of the common ancestor of these species of Picoides, and of P. canicapillus- moluccensis-minor. » ■ • . '* - ■ i An intrinsic Asiotropic problem involves, appropriate to this Australasian Congress, the proximity of piciforms to the Asiotropical-Australasian boundary. A number of barbets reach Sumatra, Borneo and Java; only Megalaima haemacephala crosses Wallace’s line, reaching the Philippines, but it does not occur in Borneo. Four genera of picids have one species reaching the Philippines (widespread Chrysocolaptes lucidus and Dryocopus javensis and endemic Picoides maculatus and Mulleripicus funebris ); the endemics have as their nearest relatives endemic Sulawesi Picoides temminickii (forms a superspecies with P. maculatus) and M. fulvus. Both Sulawesi endemic piciforms thus are related to Philippine species (the Picoides situation went unnoticed by Cracraft 1988). The Philippine-Sulawesi connection is noteworthy zoogeographically, as otherwise Chysocolaptes and Dryocopus of the Philippines relate to Borneo, as does Palawan Dinopium javanense. No piciform reaches the Sahul edge of Australasia. 474 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI ACKNOWLEDGEMENTS We are grateful to our colleagues at the American Museum of Natural History and the National Museums of Kenya for help in many ways, to the L.C. Sanford Fund of the American Museum and Mrs. Marianna Collins for financial assistance, and to the Gallmann Memorial Foundation. LITERATURE CITED CRACRAFT, J. 1988. From Malaysia to New Guinea: Evolutionary biogeography within a complex continent - island arc contact zone. Acta XIX Congressus International^ Ornithologici, pp. 2581-2593. CROWE, T.M., CROWE, A. A. 1982. Patterns of distribution, diversity and endemism in Afrotropical birds. Journal of Zoological Society of London 198: 417-442. CROWE, T.M., KEMP, A.C. 1988. African historical biogeography as reflected by galliform and hornbill evolution. Acta XIX Congressus International^ Ornithologici, pp. 2510-2518. FRIEDMANN, H. 1955. The honey-guides. Bulletin of U.S. National Museum No. 208. FRIEDMANN, H. 1976. The Asian honeyguides. Journal of Bombay Natural History Society 71: 426-432. FRY, C.H. 1988. Speciation patterns in eight orders of Afrotropical land birds. Acta XIX Congressus International^ Ornithologici, pp 2528-2536. GOODWIN, D. 1964. Some aspects of taxonomy and relationships of barbets (Capitonidae). Ibis 106: 198-220. LEES-SMITH, D.T. 1986. Composition and origins of the southwest Arabian avifauna: A preliminary analysis. Sandgrouse 7: 70-91. MAYR, E., O’HARA, R.J. 1986. The biogeographic evidence supporting the Pleistocene forest refuge hypothesis. Evolution 40: 55- 67. MORONY, J.J., JR., BOCK, W.J., FARRAND, J., JR. 1975. Reference list of the birds of the world. New York, American Museum of Natural History. PAYNE, R.B. 1986. Bird songs and avian systematics. Current Ornithology 3: 87-110. PRUM, R.O. 1988. Phylogenetic interrelationships of the barbets (Aves: Capitonidae) and toucans (Aves: Ramphastidae) based on morphology with comparisons to DNA-DNA hybridization. Zoological Journal of the Linnaean Society of London 92: 313-343 SIBLEY, C.G., AHLQUIST, J.E. 1985. The relationships of some groups of African birds, based on comparisons of the genetic material, DNA Pp. 115-152 in Schuchmann, K-L. (Ed.). Proceedings of the International Symposium on African Vertebrates. Bonn, Zoologisches Forschungsinstitut und Museum Alexander Koenig. SIBLEY, C.G., AHLQUIST, J.E., MONROE, B.L., JR. 1988. A classification of the living birds of the world based on DNA-DNA hybridization studies. Auk 105: 409-423. SHORT, L.L. 1971. The affinity of African with Neotropical woodpeckers. Ostrich, Supplement 8: 35-40. SHORT, L.L. 1980. Speciation in African woodpeckers. Proceedings of the IV Pan-African Ornithologi¬ cal Congress, pp. 1-8. SHORT, L.L. 1985. Neotropical-Afrotropical barbet and woodpecker radiations: A comparison. Ameri¬ can Ornithologists’ Union Monograph No. 36, pp. 559-574. SHORT, L.L., HORNE, J.F.M. 1985. Social behavior and systematics of African barbets. (Aves: Capitonidae). Pp. 255-278 in Schuchmann, K-L. (Ed.). Proceedings of the International Symposium on African Vertebrates. Bonn, Zoologisches Forschungsinstitut und Museum Alexander Koenig. SHORT, L.L., HORNE, J.F.M. 1988a. Family Capitonidae. Pp. 413-486 in Fry, C.H., Keith, S., Urban, E.K. (Eds). The Birds of Africa, Volume III. London, Academic Press. SHORT, L.L., HORNE, J.F.M. 1988b. Family Indicatoridae. Pp. 486-512, ibid. SHORT, L.L., HORNE, J.F.M. 1988c. Family Picidae. Pp. 512-556, ibid. SHORT, L.L., HORNE, J.F.M. 1990. Behavioural ecology of five sympatric Afrotropical honeyguides. in van den Elzen, R., Schmidt-Koenig, K., Schuchmann, K.-L. (Eds). Current Topics in Avian Biology. Proceedings of the 100th Meeting of the Deutsche Ornithologen-Gesellschaft. Bonn, Deutsche Ornithologen-Gesellschaft. WELLS, D.R. 1988. Comparative evolution of Afro-Asian Equatorial-forest avifaunas. Acta XIX Congressus Internationalis Ornithologici, pp. 2799-2809. WONG, M. 1984. Behavioural indication of an African origin for the Malaysian honeyguide Indicator archipelagicus. Bulletin of the British Ornithologists’ Club 104: 57-60. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 475 GEOGRAPHICAL MORPHOMETRIC VARIATION IN BIRDS OF THE LOWLAND EQUATORIAL FOREST OF AFRICA M. LOUETTE Koninklijk Museum voor Midden-Afrika, B-3080 Tervuren, Belgium ABSTRACT. Intraspecific geographical morphometric variation (based on information extracted from the literature) is studied for 216 stenotopic African birds which inhabit lowland equatorial forest. Stand¬ ard measurements (wing, tail and bill length) were made on samples of selected species from Upper Guinea, Cameroon, western Zaire and eastern Zaire.. Geographical variation is present in quite a few species and variation is clinal in some cases, but without any consistent trend. Variation in other spe¬ cies is much more discontinuous. Therefore, geographical variation in the birds studied is unlikely to be an effect of a common ecogeographical rule. Haphazard divergence within forest fragments isolated over geological time seems to be a better causal explanation for this variation. Character displacement is unlikely to have influenced geographical variation in Bleda and Malimbus, two polyspecific stenotopic forest passerine genera with a different number of species in the several regions. Keywords: Ecomorphology, geographical variation, Africa, rainforest birds. INTRODUCTION Intraspecific geographical morphometric variation occurs in many birds which range widely in latitude (or in altitude) within tropical savanna as well as in temperate re¬ gions. For North American species, this variation is sometimes correlated with tem¬ perature and humidity, and is often explained (although not in all cases convincingly) by Bergmann’s and Allen’s ecogeographical rules (Zink & Remsen 1986). The contemporary climate of lowlands in equatorial tropical Africa, despite a certain amount of regional variation (especially concerning the length of the dry season), is thought to be basically stable (Leroux 1983). Therefore, among the 216 stenotopic lowland forest birds which are resident in this area (Louette 1990), a common ecogeographical explanation for morphometric variation over this narrow latitudinal zone (with negligible seasonal variation in daylength) is very unlikely. But differences in wing (= wing chord), tail and bill (= culmen) length have been documented for 26 nonpasserines (Brown et al. 1982, Urban et al. 1986, Fry et al. 1988) and for at least 20 forest passerines (White 1960, 1961, 1962, 1963) which occupy this area. The presence of such variation, moreover, is also confirmed by the fact that such differ¬ ences have been used repeatedly as subspecific characters for the species in question. In the absence of a likely climatic basis for this geographical variation, possibly intraspecific competition could promote ecomorphological variation through charac¬ ter displacement. In species-rich genera with closely related species, the hypothesis that the presence of more sympatric species in a given area should result in more differentiation should be examined. Possible candidates are members of the morpho¬ logically homogeneous and stenotopic forest genera Bleda (three species in the west - found in one locality (Louette 1981) versus two in the east) and Malimbus (seven 476 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI species in the centre - locally sympatric (Brosset 1978) versus five in the west and six in the east). This paper investigates the nature and possible causes of intraspecific geographical morphometric variation in stenotopic, lowland forest birds of equatorial Africa. TABLE 1 - Means in mm (N = 8) for measurements made on African birds of lowland tropical forest. Significant (P< =0.02 in Mann-Whitney U-test) differences are indicated by Species that are absent or rare (e.g. Alethe poliocephala ) are marked as Species Measure¬ ment Sex Regions A B C D Nectarinia wing m 69.2* 74.8 75.3 76.3 superba f 65.3* 68.6 70.1 70.4 tail m 43.7* 46.4 47.3 47.2 f 41.1 42.6 43.9 42.6 bill m 32.4* 36.6 36.6 36.6 f 32.2* 35.2 36.7 35.6 Andropadus wing m 78.8* 83.8 82.4* 88.1 latirostris f 78.1 77.5 78.3* 83.3 tail m 74.1 75.6 75.5* 82.5 f 73.6 70.6* 74.3* 77.3 bill m 17.3 17.9 18.1 17.8 f 16.4 16.9 17.6 17.0 Alethe wing m 93.3 91.8 - 89.1 poliocephala f 90.1 88.5* 85.4 tail m 60.8* 58.1* - 55.6 f 58.3* 55.3 - 53.8 bill m 20.6 20.3* - 19.6 f 20.0 19.8 - 19.4 Bleda eximia wing m 108.2* 96.3* 105.8 104.9 f 102.2* 91.3* 97.6 99.4 tail m 97.4* 86.0* 95.4 95.4 f 90.6* 80.8* 89.0 89.4 bill m 28.8* 23.3* 24.8 23.2 f 25.8* 20.9* 23.2 22.7 Bleda syndactyla wing m 110.3 111.9* 107.0 106.9 f 103.5 105.8* 101.3 98.6 tail m 93.8* 98.0* 92.8 93.5 f 88.6* 94.5* 88.7 87.0 bill m 29.0 29.0 27.9* 26.1 f 25.9 25.3 26.4* 24.5 Malimbus nitens wing m 89.8 88.4* 93.4* 86.7 f 82.3 81.8* 87.3* 81.7 tail m 57.2* 51.5* 54.5* 49.4 f 52.9 49.4 51.0* 47.6 bill m 23.3* 25.4 24.0 23.3 f 22.2 23.5 23.8* 22.1 Malimbus malimbicus wing m 87.7* 82.8* 89.3* 92.4 f 82.8* 77.1* 82.4* 85.3 tail m 58.5 56.1 55.3 55.7 f 57.6* 51.6 51.2 50.4 bill m 20.4 20.8* 22.3 22.8 f 19.6 19.9* 21.9 22.3 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 477 TABLE 1 - Continued Species Measure¬ ment Sex Regions B Mai im bus scutatus wing tail bill Mai im bus cassini wing tail bill Mali m bus rubricollis wing tail bill Mai im bus coronatus wing tail bill m m m m m m m m m m m m 88.2" 87.9 51.9 50.4 19.2" 19.1 106.5* 101.4 64.4" 61.6" 26.0" 24.1 92.1 90.1 51.3 51.1 20.8 19.9 89.6 88.7 52.3* 52.7* 19.8 19.0 103.2 100.4* 59.5 58.4* 23.8 24.3* 85.9 84.4 52.8* 53.0* 19.3 90.5 88.9 49.8 49.3 20.2 19.9 100.2 94.3 58.6 54.4 22.8 22.5 85.1 83.8 49.3 48.5 19.1 92.3 90.1 49.4 48.8 20.3 19.4 99.3 93.9 58.1 54.1 22.1 22.0 87.2 84.9 50.1 47.8 19.1 FIGURE 1 - Position of the lowland rainforest in tropical Africa and the locations of the four regions studied. 478 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI METHODS Wing, tail and bill length statistics were extracted from a range of sources cited be¬ low, especially “Birds of Africa” and the publications of C.M.N. White. Table 1 lists the species measured by me from four regions in equatorial Africa (see Figure 1) in Tervuren and in the British Natural History Museum: A: Liberia and surroundings; B: lowland Cameroon and eastern Nigeria; C: Equateur Province in Zaire; D: lowlying areas in Kivu Province in Zaire. These regions cover a rough transect across the African lowland equatorial forest. The three eastern regions are fairly equidistant from one another. However, palaeobiological evidence indicates that region A was (and still is) separated much more effectively from the others than the remaining three have been from one an¬ other. These four regions also correspond to the core areas of the important histori¬ cal forest refugia as defined by Prigogine (1988). The taxa selected here include a species absent in region A and a superspecies with allospecies in contact within re¬ gion B. All localities used are below 1000 m a.s.l., with specimens taken over the last sev¬ eral decades, lessening possible altitudinal and temporal bias in mensural variation. Specimens were admittedly prepared by different individuals, but all specimens meas¬ ured were well-prepared, making differences in measurements resulting from varia¬ tion in preparation methods unlikely. Sample size consisted of eight male and eight female specimens (adult skin specimens only) from each region. Wing length was measured using a stopped ruler, tail length and bill length were measured with calipers; all to the nearest 0.5 mm. Thorpe (1976) and Zink & Remsen (1986) have reviewed approaches to the study of geographical variation. In this study, because of the relatively small sample sizes in¬ volved, interregional and intersexual comparisons were made with a non-parametric test (Mann-Whitney U-test, critical value at P<0.02). RESULTS From results published in the literature (Table 2), it appears that geographical morphometric variation is not limited to particular taxonomic groups nor to a single measurement. However, intraspecific variation in wing length is most often noted, probably because it is one of the most frequently published measurements, differing by up to 10%. Moreover, there is no consistent ‘trend’ in the variation detected (Ta¬ ble 2), although parallel trends (not necessarily both statistically significant) were found for most species in both sexes, with males nearly always being markedly larger. A geographical cline? The results in Table 2 suggest that there may be some general trend which could be interpreted as clinal, with longer billed, winged and tailed birds in the eastern regions; but is this trend real? For example, for Nectarinia superba (a sexually dichromatic species) the sample from region A is smaller in all three measurements than those from the three eastern samples, between which there are no significant differences. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 479 TABLE 2 - Lowland Afrotropical forest bird species in which there is marked intraspecific geographical variation in at least one body measurement. West < East Accipiter castanilius: wing A-C East Tauraco macrorhynchus : wing A>B Centropus leucogaster superspecies: wing, bill A>B-C>D Raphidura sabin i: wing A>B-D Dendropicos gabonensis: wing A>B-D Alethe poliocephala : wing A-C>D (White 1962) Malimbus nitens: wing, bill AD (Chapin 1954) Complex Phoeniculus bollei : wing A-CD Ploceus aurantius: wing A>B N * = e e where h2 is the heritability of the trait under study, z is the difference in mean pheno¬ types, a is the standard deviation of the trait, and Ne, is the effective size of the colo¬ nizing population. For morphometric characters of birds, heritabilities typically range between 60% and 70%, and we have used the lower figure to produce a conserva¬ tive estimate of Ne‘. To obtain a representative estimate of o for the four species, pooled standard deviations were computed separately for each species over all New Zealand populations. DIVERGENCE OF DESCENDENT AND ANCESTRAL POPULATIONS Morphometric divergence The magnitude of morphometric divergence of New Zealand descendent populations of Chaffinches, House Sparrows, and Starlings from their English ‘ancestral’ populations is relatively small. Deviations of means of skeletal morphometric charac¬ ters between descendent populations and their most similar ancestral populations average 0.81%, 1.44%, and 0.73% respectively for the three species. New Zealand populations of Common Mynas, however, have diverged more noticeably from their Indian ancestral stock, the average divergence for skeletal morphometric characters being 1.77%. Unlike the other species, the descendent New Zealand populations of Common Mynas are smaller than their most similar ancestral population in all skel¬ etal characters (Calcutta). In Chaffinches, the New Zealand populations are on aver¬ age larger in all morphometric characters except lengths of leg bones, in which they are smaller than their morphometrically closest English population (Wareham). Simi¬ larly, the New Zealand populations of House Sparrows and Starlings are also on av¬ erage larger than their most similar United Kingdom population (Nottingham) in most skeletal characters. The magnitude of morphometric divergence between ancestral and descendent populations can best be appreciated multivariately using plots of the first two princi¬ pal components (Figure 1). For all four species divergence has occurred mainly in general size (PC I), though some shape changes have also occurred (PC II). The di¬ vergence of Common Mynas in general size is especially evident, and this includes the extant population in Melbourne, Australia, from which the New Zealand populations were derived about 10 years after the initial introduction of birds from ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 507 CD OJ CD O Q. {%V0Z) Z Od (%Z'S0 2 Od (%6'U.) 2 Od (%0'6) 2 Od FIGURE 1 - Principal component plots of morphometric variation in skeletal characters of four colonizing species of passerines. Population centroids are connected by a minimum spanning tree of average taxonomic distances. Ancestral populations are denoted with solid circles and descendent populations with solid squares. 508 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI India to Melbourne in 1862. Although ancestral and descendent populations of Star¬ lings have diverged in general size, the Nelson sample in central New Zealand is morphometrically very similar to its United Kingdom counterparts. The New Zealand populations of Chaffinches are very similar in size to the two United Kingdom sam¬ ples, especially the sample from Wareham in southern England, where at least some of the transplanted birds were taken (Ince et al. 1980). The larger multivariate size of New Zealand populations of House Sparrows relative to northeast English populations from around Norwich and Nottingham is also clearly depicted in Figure 1. Rate tests of the divergence of the morphometric means of the New Zealand descend¬ ent populations from their most similar ancestral population for each species revealed that the hypothesis of random drift of essentially neutral characters cannot be rejected as the cause of their divergence. For House Sparrows, Chaffinches, and even the more divergent Common Mynas, all skeletal characters yield estimates of Ne* » Ne in the founder populations. In Starlings, one character out of 14 (cranium depth) has Ne* < Ne, and another (premaxilla length) approaches the boundary estimate. Thus for these two characters the divergence of the descendent populations from the ances¬ tral populations appears to be too great to be accounted for solely by drift. Genetic divergence The genetic divergence of ancestral and descendent populations of the four species of colonizing passerines is summarized in Table 1 for the number of loci showing sig¬ nificant geographic variation, the among population component of genetic variance (Fst), and mean Rogers’ genetic distance (DR). For all species, the establishment of colonizing populations in New Zealand has led to an increase in the average among- population genetic distances. Relative to the ancestral populations, levels of genetic subdivision (as judged by Fst values) in each species have also increased due to the divergence of the colonizing populations in New Zealand. The greater morphometric divergence of ancestral and descendent populations of Common Mynas compared to the other three passerines is paralleled by correspondingly greater genetic divergence in this species (Table 1). TABLE 1 - Morphometric and genetic divergence within and among ancestral and de¬ scendent populations of four colonizing species of birds. Species Comparison Morphometries Na Va d Nb Genetics F., Dr House Sparrow N.Z. & England0 8 6.2 0.034 8 0.015 0.026 Within N.Z. 11 5.3 0.013 4 0.005 0.020 Within England 6 6.9 0.019 3 0.010 0.021 Starling N.Z. & U.K.d 9 5.1 0.013 6 0.038 0.025 Within N.Z. 5 3.1 0.009 6 0.032 0.019 Within U.K. 9 4.1 0.011 1 0.010 0.008 Chaffinch N.Z. & U.K.e 1 2.3 0.013 9 0.034 0.031 Within N.Z. 2 2.8 0.010 6 0.040 0.026 Within U.K. 2 6.4 0.014 1 0.006 0.016 Common Myna N.Z. & India' 11 22.2 0.022 18 0.059 0.035 Within N.Z. 10 5.3 0.011 4 0.016 0.105 Within India 13 30 8 0.021 2 0.032 0.023 a Number of geographically variable characters. b Number of geographically variable loci. c Seven lo¬ calities in northeastern England (Parkin & Cole 1985). d Six localities in England and Scotland (Ross 1983). e One locality in England and one in Scotland, ' Six localities in India. zoo ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI (%8’8I.) 2 QdOOOd (%9'9 I.) 2 QdOOOd (%ZLl) 2 QdOOOd (% L’9 0 2 QdOOOd 509 FIGURE 2 - Principal coordinate plots of genetic differentiation among ancestral and descendent populations of four species of colonizing passerines. Population centroids are connected by a minimum spanning tree of Rogers’ (1972) genetic distances. Ancestral populations are denoted with solid circles and descendent populations with solid squares. 510 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Multilocus graphic representations of the genetic divergence of ancestral and de¬ scended populations are provided by the principal coordinates plots for each species in Figure 2. These plots clearly indicate that the descendent populations are geneti¬ cally differentiated from their ancestral counterparts in all species, with the exception that the Nelson sample of Starlings groups closely with the United Kingdom samples. Interestingly, in Common Mynas, the most genetically divergent sample is the one from Melbourne, indicating that this population has diverged at a faster rate than the derivative New Zealand populations. POPULATION DIFFERENTIATION IN COLONIZING POPULATIONS IN NEW ZEALAND Morphometries The magnitude of population differentiation in morphometric characters among colo¬ nizing populations of House Sparrows and Starlings is similar to that in the ancestral populations from which they are derived (Table 1). However, ancestral populations of Chaffinches and Common Mynas are more differentiated than their respective de¬ scendent populations in New Zealand. In Common Mynas, average taxonomic dis¬ tances among populations in India are about double those in New Zealand, and in House Sparrows they are about 50% larger in the northeast England populations to those over the whole of New Zealand. There are clear differences among species in the degree of geographic variation they have developed. In particular, Chaffinches and Starlings are much less differentiated morphometrically among populations than are the other two species. House Sparrows and Common Mynas have differentiated significantly within New Zealand in more than half the 14 skeletal characters measured in this study, whereas Chaffinches and Star¬ lings have only developed significant geographic variation in two and five characters, respectively. In New Zealand, the average added variance component (VA) of indi¬ vidual characters, the portion of the total morphometric variance partitioned among localities, is only about half as large in Chaffinches and Starlings as in the other spe¬ cies (Table 1 ). Genetics At the level of apparently neutral allozyme loci, population differentiation is qualita¬ tively different to that in morphometric characters. For Starlings, Chaffinches, and Common Mynas, statistically significant heterogeneity in allele frequencies was de¬ tected at twice or more the number of loci in the New Zealand populations than in their ancestral counterparts (contingency x2 tests, P < 0.01). In House Sparrows, however, similar numbers of geographically variable loci were found in ancestral and descend¬ ent populations, though in both New Zealand and the United Kingdom only a restricted part of the total ranges were sampled. Genetic structuring is only weakly developed in ancestral and descendent populations in all four species. In both Starlings and Chaffinches, values of F are lower in the ancestral populations in the United Kingdom (Table 1). For Common Mynas and House Sparrows the situation is reversed, with the ancestral populations having higher Fst values. Multilocus genetic distances show the same pattern of population ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 511 differentiation, with the exception that ancestral and descendent populations of House Sparrows are about equally differentiated (Table 1). DISCUSSION Processes of population differentiation in colonizing species The four species of passerine birds studied here have successfully colonized a range of habitats in New Zealand in about 100-130 years since their introduction last cen¬ tury. In this short period of time, small scale population differentiation in morphometric characters of the skeleton and in protein-encoding nuclear genes has developed in this novel environment. The genetic and morphometric shifts that these colonizing populations have undergone relative to their ancestral stock are also quite small, but nevertheless they are mostly about equal or exceed in magnitude the levels of popu¬ lation differentiation that have presumably developed over many millenia in connected continental demes. Exploratory rate tests of the divergence of means of morphometric characters in all four species cannot reject the null hypothesis of random drift as the cause of this di¬ vergence, even in Common Mynas where all characters are smaller in descendent populations in New Zealand. Although these tests do not definitively rule out a role for natural selection in the divergence of ancestral and descendent populations, it is clear that extremely small selection coefficients would be involved and thus charac¬ ters would be effectively neutral or nearly so (Baker et al. 1990a). This conclusion highlights the conceptual advances in our thinking about microevolutionary diversifi¬ cation of populations that have occurred since Mayr’s 1965 paper on colonizing spe¬ cies of birds. Adaptive narrative explanations, so characteristic of this earlier period when the new synthetic theory was dominating evolutionary biology, are no longer ac¬ cepted in lieu of more compelling supporting evidence for causal mechanisms. Earlier analyses of population differentiation in New Zealand populations of House Sparrows showed that size variation was ordered approximately clinally, with birds averaging larger in warmer northern localities. Because this variation in size was as¬ sociated with an essentially N-S climatic gradient, and because the sexes have re¬ sponded differently to environmental variation, it was argued that natural selection for ecoclimatic adaptation was occurring in the New Zealand populations (Baker 1980). A similar argument was also advanced by Johnston & Selander (1971, 1973) to ex¬ plain the continental scale of geographic differentiation in North American populations of House Sparrows. New Zealand populations of Common Mynas show the same ten¬ dency as House Sparrows to be larger in northern localities (Baker & Moeed, 1979). Conversely, populations of Starlings and Chaffinches show no such trend, and instead morphometric variation among localities is haphazard with respect to environmental gradients (Ross & Baker 1982, Baker et al. 1990a). Geographic surveys of allele frequencies in allozymes were undertaken to directly reveal underlying patterns of genetic differentiation, rather than relying on inferences about genetic changes based on morphometric characters. These studies revealed that genetic drift has played an important role in population differentiation, and there¬ fore implied that gene flow among localities in New Zealand is not sufficient to over¬ come stochastic differentiation, at least in Starlings and Chaffinches (Ross 1983, 512 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Baker et al. 1990a). Geographic subdivision of the New Zealand landmass by water barriers, mountains, and native forests that are almost impenetrable for these passerines have apparently limited gene flow much as in native species (see compan¬ ion paper in this symposium by Daugherty and Triggs). This finding raises the ques¬ tion of whether the N-S trend of increasing size in House Sparrows and Common Mynas in New Zealand is really ordered by natural selection for climatic adaptation, especially since it is the reverse of that predicted by Bergmann’s rule. An alternative explanation is that these body size trends are wholly or partly environmentally in¬ duced, as has been demonstrated in Red-winged Blackbirds Agelaius phoeniceus by James (1983). Transfer experiments among localities in New Zealand with different climates are urgently needed to test this hypothesis. The realization that a large en¬ vironmental component in birds can be misinterpreted as selection for local adapta¬ tion is another important conceptual advance since the 1965 symposium. The different patterns of population differentiation in New Zealand by Chaffinches and Starlings on the one hand, and House Sparrows and Common Mynas on the other, almost certainly implicate agencies other than climatic ones. Future studies need to address morphological responses to niche width, food particle size, and the depauperate matrix of competitor species encountered by colonizing species in New Zealand relative to ecological conditions in their ancestral communities. Ecological variation potentially can be a potent force in population differentiation via selection for optimal body size (Case 1978). Finally, more attention needs to be focussed on the possible role of ecological and sexual selection on body size of males in colonizing species in New Zealand, since it is now well established that males have developed greater levels of population differentiation than have females in all four species thus far studied. As noted by Baker (1980), New Zealand provides a wonderful natural laboratory in which to conduct such important studies. ACKNOWLEDGEMENTS We gratefully acknowledge the staff of the Department of Ornithology in the Royal Ontario Museum, who over the past 18 years have supported all phases of our work. Special thanks are due to Mark Peck, Carol Edwards, Marg Goldsmith, and Brad Millen for excellent technical and field assistance. Financial support for our studies was provided by the Natural Sciences and Engineering Research Council of Canada (Grant A0200 to AJB). LITERATURE CITED BAKER, A. J. 1980. Morphometric differentiation in New Zealand populations of the House Sparrow (Passer domesticus). Evolution 34: 638-653. BAKER, A. J., MOEED, A. 1979. Evolution in the introduced New Zealand populations of the Common Myna, Acridotheres tristis (Aves: Sturnidae). Canadian Journal of Zoology 57: 570-584. BAKER, A. J., MOEED, A. 1980. Morphometric variation in Indian samples of the Common Myna, Acridotheres tristis (Aves: Sturnidae). Bijdragen tot de Dierkunde 50: 351-363. BAKER, A. J., MOEED, A. 1987. Rapid genetic differentiation and founder effect in colonizing populations of Common Mynas ( Acridotheres tristis). Evolution 41: 525-538. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 513 BAKER, A. J., PECK, M. K., GOLDSMITH, M. A. 1990a. Genetic and morphometric differentiation in introduced populations of Common Chaffinches ( Fringilla coelebs) in New Zealand. Condor 92: 76-88. BAKER, A. J., DENNISON, M. D., LYNCH, A., LE GRAND, G. 1990b. Genetic divergence in periph¬ erally isolated populations of chaffinches in the Atlantic Islands. Evolution 44: 564-582. BARROWCLOUGH, G. F., JOHNSTON, N. K., ZINK, R. M. 1985. On the nature of genic variation in birds. Pp. 135-154 in Johnston, R. F. (Ed.). Current Ornithology, Volume 2. New York: Plenum. CASE, T. J. 1978. A general explanation for insular body size trends in terrestrial vertebrates. Ecology 59: 1 -1 8. FLEISCHER, R. C., WILLIAMS, R., BAKER, A. J. 1991. Genetic differentiation in Hawaiian populations of Common Mynas ( Acridotheres tristis). Journal of Heredity, in press. GIBSON, A. R., BAKER, A. J., MOEED, A. 1984. Morphometric variation in introduced populations of the Common Myna ( Acridotheres tristis ): An application of the jackknife to principal component analy¬ sis. Systematic Zoology 33: 408-421. INCE, S. A., SLATER, P. J. B., WEISMANN, C. 1980. Changes with time in the songs of a population of Chaffinches. Condor 82: 285-290. JAMES, F. C. 1983. Environmental component of morphological differences in birds. Science 221: 184-186. JOHNSTON, R. F., SELANDER, R. K. 1971. Evolution in the House Sparrow. II. Adaptive differentia¬ tion in North American populations. Evolution 25: 1-28. JOHNSTON, R. F., SELANDER, R. K. 1973. Evolution in the House Sparrow. III. Variation in size and sexual dimorphism in Europe and North and South America. American Naturalist 107: 373-390. LANDE, R. 1976. Natural selection and random genetic drift in phenotypic evolution. Evolution 30: 314-334. LYNCH, M. 1988. The divergence of neutral quantitative characters among partially isolated populations. Evolution 42: 455-467. LYNCH, M., HILL, W. G. 1986. Phenotypic evolution by neutral mutation. Evolution 40: 915-935. MAYR, E. 1965. The nature of colonizations in birds. Pp. 29-43 in Baker, H. G., Stebbins, G. L. (Eds). The genetics of colonizing species. New York: Academic Press. PARKIN, D. T., COLE, S. R. 1985. Genetic differentiation and rates of evolution in some introduced populations of the House Sparrow, Passer domesticus, in Australia and New Zealand. Heredity 54: 15-23. POWER, D. M. 1971. Statistical analysis of character correlations in Brewer’s Blackbirds. Systematic Zoology 20: 1 86-203. ROGERS, J. S. 1972. Measures of genetic similarity and genetic distance. University of Texas Publi¬ cations 7213: 145-153. ROSS, H. A. 1983. Genetic differentiation of Starling ( Sturnus vulgaris: Aves) in New Zealand and Great Britain. Journal of Zoology, London 201: 351-362. ROSS, H. A., BAKER, A. J. 1982. Variation in size and shape of introduced Starlings, Sturnus vulgaris (Aves: Sturnidae), in New Zealand. Canadian Journal of Zoology 60: 3316-3325. SNEATH, P. H. A., SOKAL, R. R. 1973. Numerical taxonomy. San Francisco: W. H. Freeman. TURELLI, M., GILLESPIE, J. H., LANDE, R. 1988. Rate tests for selection on quantitative characters during macroevolution and microevolution. Evolution 42: 1085-1089. 514 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI MITOCHONDRIAL DNA AND AVIAN MICROEVOLUTION JOHN C. AVISE and R. MARTIN BALL, JR. Department of Genetics, University of Georgia, Athens, GA 30602, USA ABSTRACT. Mitochondrial (mt) DNA provides a rich source of uniparentally-transmitted genetic mark¬ ers especially useful in the study of matriarchal phytogeny over microevolutionary timescales. Various applications of mtDNA data to avian taxa are reviewed, including the assessment of: (1) magnitudes of intraspecific polymorphism; (2) genetic distinctions between sibling species; and (3) genetic dis¬ tances and phytogenies among congeners. Special attention is focused on patterns of geographic dif¬ ferentiation within avian species, where conspecific populations have proved to exhibit a variety of mtDNA phylogeographic structures. The deep and geographically structured subdivisions observed in the mtDNA genealogies of some avian species probably evidence the effects of Pleistocene biogeographic separations, while the additional, shallower mtDNA subdivisions and haplotype fre¬ quency shifts distinguishing populations of most species probably reflect more recent restrictions on gene flow. In general, mtDNA data offer a novel, phylogenetic perspective on microevolutionary proc¬ esses, and allow provisional interpretation of contemporary population structure in terms of historical demography. Keywords: Mitochondrial DNA, phylogeography, population structure, genetic markers, gene flow, historical demography. INTRODUCTION Because of its rapid evolution and uniparental inheritance, mitochondrial (mt) DNA offers novel perspectives on microevolutionary processes in higher animals (Avise et al. 1987, Wilson et al. 1985). A major motivation for mtDNA surveys of birds has been the need for critical reassessment of the processes governing genetic differentiation among conspecific populations. Few topics have generated greater discussion in re¬ cent years, yet there remain conflicting opinions on the magnitude of contemporary gene flow and historical connectedness among avian populations. On the one hand, most birds have high dispersal potential (because of flight), and many species have broad geographic distributions, suggesting that gene flow is high and population dif¬ ferentiation minimal. On the other hand, many species exhibit strong tendencies for nest-site philopatry (Greenwood 1980), and commonly show obvious geographic vari¬ ation in size, song, or plumage (often leading to the description of subspecies), sug¬ gesting that inter-populational gene flow is normally quite low. But since the genetic bases and evolutionary forces governing most morphological or behavioral differences are poorly understood, their relevance to estimating gene flow and magnitude of ge¬ netic divergence is unclear. Clearly, a reappraisal of avian population structure that capitalizes on the resolving power of mtDNA, and the historical perspective it provides, is appropriate. BACKGROUND, AND CALIBRATION OF MTDNA EVOLUTION IN BIRDS Presumably, mtDNA inheritance is predominantly or exclusively maternal in birds, as in most other multicellular animals (Avise & Vrijenhoek 1987). However, only a sin¬ gle direct test of avian mtDNA transmission appears available: Watanabe et al. (1985) ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 515 showed that progeny of a cross between a female Japanese quail and a male chicken exhibited the mtDNA genotype of the mother. Glaus et al. (1980) and Shields & Helm-Bychowski (1988) review evidence that the molecular features of avian mtDNA variation are similar to those of other vertebrate classes. Thus the molecule is about 16-20 kilobase-pairs in length, evolves primarily through nucleotide substitutions, and is normally homoplasmic (occurs as a single pre¬ dominant genotype within an individual — but see Avise & Zink 1988). The complete sequence of one avian mtDNA recently has been obtained (Desjardins & Morais 1990). In comparison to mammals and the clawed frog, whose mtDNAs are identical in gene content and order, chicken mtDNA lacks sequences associated with the light- strand replication origin, and exhibits a transposition of two loci. Only two explicit tests of mtDNA evolutionary rate appear available for birds. First, based on her own mtDNA data and those of Glaus et al. (1980), Helm-Bychowski (1984) plotted an initial rate of about 2% sequence divergence per million years be¬ tween several Galliforme species. Second, Shields & Wilson (1987) observed about 9% sequence divergence between Anser and Branta geese that from fossil evidence last shared a common ancestor about 4-5 million years ago. Both studies thus yielded evolutionary rate estimates close to the conventional mtDNA “clock” calibration for non-avian vertebrates (Brown et al. 1979). The rapid pace of mtDNA evolution has raised hope that the molecule can provide a rich source of genetic markers for closely related avian species (some of which are difficult to distinguish by morphological or allozyme traits). Initial results, summarized in Table 1, indicate that many fixed restriction site differences do indeed distinguish some sibling taxa of birds. TABLE 1 - Genetic differences between avian sibling taxa observed in conventional surveys of allozymes and mtDNA. Data for the first four comparisons come from Avise & Zink (1988); those for the other Parus from Braun & Robbins (1986; allozymes) and Mack et al. (1986; mtDNA); Ficedula, Gelter (1989). Taxon pair No. characters No. fixed Genetic distance surveyed differences allozyme mtDNA allozyme mtDNA allozyme mtDNA loci sites alleles sites3 £7 P Rallus elegans v R. longirostris 38 68 0 5 0.004 0.006 Limnodromus scolopaceus v L. griseus 36 77 2 24 0.060 0.082 Quiscalis major v Q. mexicanus 38 80 0 10 0.001 0.016 Parus bicolor bicolor v P.b. atriscristatus 36 83 0 5 0.063 0.004 Parus atricapillus v P. carolinensis 35 80 0 17 0.001 0.040 Ficedule hypoleuca v F. albicollis 35 214 0 >8 0.001 0.010 a Conservative estimate, assuming that fragment profile differences not due to a single restriction site gain/loss are due to only two such changes. b Nei's (1978) distance measure. 516 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI In a few cases, closely related species could not be separated cleanly on the basis of mtDNA genotype. For example, two genetically distinct arrays of mtDNA haplotypes were present in Mallard Anas platyrhynchos populations, while only one of these arrays was represented among the assayed Black Ducks A. rubripes (Avise et al. 1990). One possibility is that the distinct mtDNA genotypes were transferred between species via secondary hybridization. Alternatively, the phylogenetic split in the mtDNA gene tree may have predated the speciation event, such that the Mallard-Black com¬ plex now exhibits a “paraphyletic” relationship in terms of mtDNA genealogy. Table 2 summarizes mtDNA genetic distances between non-sibling avian congeners. Levels of sequence divergence range between 0.4 and 1 1%. Most such estimates of avian mtDNA sequence divergence are considerably lower than mean values for other vertebrate genera such as Lepomis sunfish and Hyla treefrogs, and thus parallel the conservative pattern of divergence in avian allozymes relative to these non-avian taxa (Kessler & Avise 1985). However, some non-avian vertebrate genera exhibit means and ranges of mtDNA p quite comparable to those reported for birds (review in Shields & Helm-Bychowski 1988). MtDNA distances among avian congeners are well within the expected linear portion of a curve relating sequence divergence to time (Brown et al. 1979). Thus the mtDNA distances may be of considerable utility in estimating phylogeny among closely related avian species, a suggestion strongly supported by the results of three multi-species systematic studies. Within Anas and Aythya ducks (Kessler & Avise 1984), Platycercus rosellas (Ovenden et al. 1987), and Ammodramus sparrows (Zink & Avise 1990), phylogenies inferred from mtDNA data agree well with those based on inde¬ pendent sources of genetic and/or other biological information. Levels of mtDNA polymorphism within avian species are usually high (Table 3). In a typical survey involving about 100 restriction sites scored per organism, a random pair of conspecific individuals differs detectably in mtDNA genotype with probability 0.70 or higher (genotypic diversity), and exhibits a sequence divergence (nucleotide diver¬ sity) of about 0.3%. Nonetheless, mtDNA polymorphism remains far lower than pre¬ dicted under neutrality theory, given suspected rates of sequence divergence and current day population sizes (Avise et al. 1988). For example, Figure 1 plots the ob¬ served and expected times to common mtDNA ancestry in the Red-winged Blackbird ( Agelaius phoeniceus). The observed mean distance between mtDNA lineages is more than two orders of magnitude lower than might have been anticipated from the census size of breeding females. Either the rate of mtDNA evolution is decelerated in such species, and/or evolutionary effective population sizes are vastly smaller than present-day population sizes. In principle, the latter could result from historical fluc¬ tuations in female numbers, or from a periodic positive directional selection on mtDNA genotypes that would have a similar net effect of channeling mtDNA through fewer female ancestors (Avise 1989). PHYLOGEOGRAPHIC DIFFERENTIATION WITHIN AVIAN SPECIES Because the phylogenies of haplotypes can be inferred from mtDNA data, considera¬ tions of historical as well as contemporary aspects of population structure are possi¬ ble. A straightforward method for summarizing the information content of mtDNA 517 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI o o c 0) a> H— a> LO o 03 00 CO 00 ■M- LO CO o 00 -M- CM CO O o -I— 03 q q q q < d d d o d d d d 2 d CO d o 1 LO T- d d o CO CM CM ■M- o CM co CM 1 — o o O o q q q q q E a d d d d d d d d d (0 3 TJ <0 > ■2 "D "D w £ .£ ■ O 1- O o 0) 2 (0 CL 0) E o Q. o 2 (A O (A 3 C a> O lo m 00 00 CD CD CD CD CD CD > > < < oQ oQ 0) _CD CD CD CD CD CD CD O CD CO 00 'r- CD CD ■,_ CD _• ‘> 03 < — - CD oC o C 03 N ^ T- CD CD N ID CD CO N- 00 CO ^1- 00 00 CD CD h- D- 00 CO 03 03 ^ 00 C o CD CD -c; = — 03 CD CD CD CD > > < < o3 o3 08 o3 £ £ JD 0) CD CD CD CD CD CD CD C CD "D C CD o o > a) co O CD CD T3 TD CD CD 't ^ ID ID CD lo lo h~ h~ lo LO lo co O 00 00 CO T- CM CO CO CO CM LO CD CD 3 E CU C\J 03 m E -2 .N 5 O 5 rt= ~0 c a o CD o P E CD CD E CD CD 2 CD P 03 ^CDdjEctsD- ^ Q 5 ^ CL E 03 0l CD 3 O x- ^ O CO :£ ^ S) c C C is ^ ^ CQ a. 518 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI involves superimposing on a distributional map the genealogies inferred from the mu¬ tational differences among mtDNA haplotypes. The outcomes can then be classified into “phylogeographic” categories, interpreted against neutrality expectations, and compared among studies (Avise et al. 1987). Redwings 16.7 33.3 50.0 66.7 83.3 100.0 116.7 133.3 GENERATIONS (x I03) FIGURE 1 - Observed and expected frequency distributions of times to common ancestry for mtDNA haplotypes in the Red-winged Blackbird. Expected times generated from in- breeding theory (see text) are given for each of two conditions: 1) N( = 20,000,000 (a reasonable guess for the current breeding population of female redwings; shown in the inset); and 2) Nf(e) = 36,700 (a value which yields a mean expected divergence time equal to that inferred from the mtDNA data). Observed times were derived from the data of Ball et al. (1988), using a conventional mtDNA “clock” calibration and a generation length of three years. [Notice the difference in scale along the abscissas of the inset and main graph]. For example, genetic gaps (involving many mutational steps) might appear in the mtDNA phylogeny itself, and the clades thus identified could either be confined to particular geographic regions (phylogeographic category I) or broadly sympatric (cat¬ egory II). Alternatively, all mtDNA genotypes might be closely related, and particular haplotypes either geographically localized (categoiy III), or widespread (category IV). If most avian populations have been connected historically by relatively high levels of gene flow (through high per-generation migration rates and/or periodic cycles of ex¬ tinction and recolonization of local populations from a source), then most avian spe¬ cies should fall into phylogeographic categories II or IV, likely depending in part on whether the species-wide effective population sizes were large or small, respectively. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 519 2 co 03 CD _Z3 03 iS > 03 CD "O E a) o £ w ® LCD ^ T_ 03 II O r- 03 c n c »_ 03 O sz ~ 03 CD .E as 'c: 2 £ 03 C/3 CD J= S*_ -•— » CD m- £ O 03 03 CD £ N -J3 03 03 o. r E 03 03 03 CD O 'sz O .!= CD 4— » CD CL E o o c 03 CD 03 X3 !5 ^ >+_ 03 O 03 03 TD CD o O 03 03 £ .E CD _C W t: o '> 03 =* o > CD ■O CD o CL .Ed® CD E' -Q _ .2 x: 3 ^ 03 ^ - o < _>» c Z c Q) 9 ° O E B Z O E c 03 X .9 03 o E 03 LD CO o - . 1^ 03 a- -5 8- 3 o 0 — 03 03 sz c. ~ iu .2 o J W ^ CD > >, < 2 ca H a. 03 CD o c 03 J— a> >*- a> 0 !E c CD c < z 03 § C/3 if) ID CO ID c 03 c < Z < z a CD -* — < c < c v J a c c c c 4 _ c k. 03 03 0 0 O CD SZ i _ 0) Q3 -4 — ' 2 0 0 0) sz a5 O O (0 O < > ■ -t — 1 0 CO ca CO 03 to CD < 0 t/3 03 O) (0 Z Z c CD § $ Z c $ 03 2 w 03 i_ o ® 3 .2 C T3 O a Q. 3T •d CO s i 03 c 03 T3 03 n E 3 c 0) I So? »_ CO c % © g> ^ S •- 2 2 Z. 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CO Is- o o Is- d co 00 00 LO .co CO c 03 to ca c co CJ -2 c 2 CQ CD 1- O O 9 9 d d O 03 00 d d co co 03 03 Is- co o o CM CM co o -c 0 c -c CO 03 CL -d - ca -Q a 2 co co cts 03 c c genotypic diversity, G = (n/n- 1) (1- e ff), where fi is the frequency of / th mtDNA haplotype in a sample of n individuals; and nucleotide diversity, p = (n/n- 1) e flp^, where fi and f are the frequencies of the / th and y th mtDNA haplotypes and p(/ is their estimated nucleotide sequence divergence. 520 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Alternatively, severe restrictions on gene flow should result in patterns falling into phylogeographic categories I or III, depending on whether the restrictions were in force over long-term (evolutionary) versus short-term (ecological) timescales, respec¬ tively. Various intermediate situations may also exist. For example, ancestral mtDNA genotypes might be geographically widespread, while closely related, newly arisen mutations were localized (phylogeographic category V). One explanation for such an outcome might be that a species had expanded recently from a single refugium, but that its populations were currently experiencing limited genetic contact. The following are brief synopses of the studies conducted to date on the mtDNA pat¬ terns observed in birds, as classified into the various phylogeographic categories of Avise et al. (1987). Phylogeographic categories IV and V The Red-winged Blackbird ( Agelaius phoeniceus) was the first avian species assayed extensively in terms of mtDNA phylogeny (Ball et al. 1988). In a continent-wide sur¬ vey involving 75 restriction sites and 127 individuals, 34 mtDNA clones were ob¬ served. Most genotypes were closely related (mean p = 0.002), and many of the mtDNA genotypes and clades were geographically widespread, suggesting consider¬ able historical connectedness among populations, either through gene flow, and/or retention of genotypes from a recent ancestral population. On the other hand, some mtDNA genotypes and clades were rare or absent from particular regions, indicating a mild population structure. The evidence for this mtDNA structure is further supported by the form of the frequency distribution of haplotype distances, which departs sig¬ nificantly from the geometric distribution expected under the idealized “high gene flow” model (Figure 1). One interpretation of the mtDNA data is that redwings recently colonized North America from a much smaller source population, perhaps since retreat of the last Pleistocene glacier some 13,000 years ago. If so, all redwing populations are closely related genetically, notwithstanding contemporary limits to dispersal and gene flow that allowed development of a mild genetic population structure across the continent. If this mtDNA-based scenario is correct, the extensive morphological differentiation among redwing populations (as reflected in the recognition of at least 23 subspecies) has occurred within the context of relatively shallow evolutionary separations. Perhaps some of the morphological differences are ecophenotypic [not based entirely on ge¬ netic differences, as is suggested by the nestling transplantation experiments of James (1983)]. Alternatively, genes responsible for morphological traits may have evolved so rapidly that geographic differentiation arose over a time-scale too short for clear detection by corresponding differences in the mtDNA assays. Other avian species reported to exhibit phylogeographic category IV or V population structure include the Song Sparrow Melospiza melodia in the western U.S. (Zink 1991), and the Downy Woodpecker Picoides pubescens across North America (Ball & Avise in prep). Phylogeographic category I A survey of 40 Seaside Sparrows ( Ammodramus maritimus) collected from New York to Louisiana revealed 1 1 mtDNA clones belonging to two distinct phylogenetic groups between which mean sequence divergence (after correction for within-group variation) was, p ~ 0.01 (Avise & Nelson 1989). One mtDNA array included all Atlantic coast ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 521 birds, while the other involved all Gulf coast specimens (Figure 2). This phylogeographic pattern probably evidences a long-term separation of Atlantic and Gulf coast populations, a scenario further supported by Funderburg & Quay’s (1983) zoogeographic reconstruction for the species, and by the remarkable geographic con¬ cordance between the mtDNA partitions in the Seaside Sparrow and those of several other coastal species in the southeastern U.S. (Avise & Ball 1990). The Atlantic-Gulf phylogenetic distinction in the Seaside Sparrow is not, however, recognized in the subspecies designations that provided the basis for population management decisions (Avise & Nelson 1989). Other avian species reported to exhibit a phylogeographic category I population struc¬ ture include the Sharp-tailed Sparrow Ammodramus caudacutus (Avise & Rising in prep), Fox Sparrow Passerella iliaca (Zink 1991), and Canada Goose Branta canadensis (Van Wagner & Baker 1990). In the latter species, two phylogenetically distinct mtDNA groups (differing by a mean sequence divergence of p * 0.018) were observed to correspond exactly to a subdivision of recognized subspecies into large¬ bodied versus small-bodied forms. 3 o LU CC Li. LU LU cr 0.8 ~ 0.7 - 0.6 - 0.5 - 0.4 - 0.3 - 0.2 - 0.1 - 0 SEASIDE SPARROW within Atl. or Gulf “ I - 1 - 1 - 1 - 1” 0.2 0.4 0.6 between Atl. and Gulf n / -1 - 1 - - - f=t 0.8 1.0 1.2 "I 1 .4 SEQUENCE DIVERGENCE (%) FIGURE 2 - Frequency distribution of mtDNA genetic distances observed in 780 pairwise comparisons among 40 Seaside Sparrows collected from localities along the Altantic and Gulf coasts. Phylogeographic category II A phylogeographic pattern until recently not reported in any animal species is that in which highly distinct mtDNA clones or clades co-occur as polymorphisms over a broad geographic area. We recently have found two such examples, involving the Snow Goose Anser caerulescens (Avise & Alisauskis in prep) and the Mallard Duck Anas platyrhynchus (Avise et al. 1990). In the Snow Goose, a survey of major breeding lo¬ cales across the species’ range in Arctic Canada revealed two distinct mtDNA clades (p = 0.01 1 ) shared by all surveyed populations (albeit in significantly different frequen¬ cies). Similarly, populations of Mallard from California and Manitoba shared distinct mtDNA clades differing by p = 0.009. 522 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI The origins of such mtDNA differences are unclear, although they may indicate effects of past population disjunction and recent admixture. The results are somewhat sur¬ prising, because most gene flow in waterfowl (Anatidae) is thought to be mediated by males, who pair with females on wintering grounds or during migration, and move to the female’s natal colony to breed (Cooke et al. 1975). Nevertheless, the mtDNA re¬ sults suggest recent connectedness among colonies involving females. The finding that mtDNA lines are shared between locales in some waterfowl, where dispersal is presumably male-biased, makes even more impressive the sharp mtDNA phylogeographic discontinuities observed in other avian species (such as the Seaside and Sharp-tailed Sparrows). CONCLUSIONS ABOUT AVIAN POPULATION STRUCTURE The mtDNA phylogeny of any species represents but one realization of the process of gene lineage sorting through an organismal pedigree, and hence must be inter¬ preted with caution as an indicator of overall genomic (nuclear) history. In the absence of concordant support from independent evidence, significant partitions in an mtDNA phylogeny cannot necessarily be assumed to evidence fundamental historical subdi¬ visions at the population level (Avise and Ball 1990). One form of support can come from a correspondence between mtDNA phylogenetic subdivisions and boundaries between historical biogeographic provinces (a good example involves the Seaside Sparrow). Or the support may derive from concordance of subdivisions in the mtDNA gene tree with those registered in other attributes such as morphology (a good exam¬ ple involves the large- and small-bodied forms of the Canada Goose - Van Wagner and Baker 1990). In other cases, confidence must rest on plausibility arguments - for example, given the current widespread distribution of the Red-winged Blackbird, its potential for long-distance movement, and the fact that major portions of its range were uninhabitable only 13,000 years ago, the species may indeed have expanded its range and population size in recent evolutionary times, as the mtDNA data strongly suggest. In interpreting gene phylogenies, a distinction should also be drawn between gener¬ alized conclusions about the magnitude of population structure, versus the evidence for particular historical population separations. In species whose populations exhibit contemporary isolation by distance in the absence of long-standing biogeographic barriers to dispersal, phylogenies of independent genes may each reveal significant population subdivision, yet exhibit little concordance in the population units identified. Such results imply generalized restrictions on gene flow, but the particular populations recognized by any gene may be of little evolutionary consequence. However, when populations or regions are concordantly identified by independent genes, long-stand¬ ing biogeographic separations are strongly implicated (Avise & Ball 1990). With these caveats in mind concerning inferences from mtDNA (or other) gene trees, the following conclusions about avian population structure can be drawn from the mtDNA data currently available: 1) Most avian species exhibit a wealth of intraspecific polymorphism, although still far less than predicted under neutrality theory given current population sizes and presumed rate of mtDNA evolution. Thus either: (a) directional selection favoring particular genotypes has served periodically to reduce the level of mtDNA ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 523 polymorphism; or (b) evolutionary effective population sizes are comparatively low for other reasons involving historical demography (such as fluctuations in num¬ bers of successfully-breeding females). 2) Populations of some avian species, such as the Red-winged Blackbird, are char¬ acterized by considerable historical connectedness over recent evolutionary time, as judged by limited mtDNA phylogeographic structure. Within some such spe¬ cies, extensive morphological differentiation appears to have arisen against a relatively shallow phylogenetic backdrop. 3) Other avian species, such as the Seaside Sparrow, consist of regional populations exhibiting strong phylogenetic separation. Inter-regional genetic dis¬ tances are much larger than those observed within regions, and the phylogenetic gaps tend to correlate with likely biogeographic partitions. Thus historical popu¬ lation separation (rather than retention of ancient lineages within a large, non- subdivided population) probably accounts for such major mtDNA phylogeographic discontinuities. 4) In some species (such as the Snow Goose, and perhaps the Mallard Duck), dis¬ tinct mtDNA lineages are shared by widely separated breeding populations. Wa¬ terfowl are unusual among birds in the strength of female site philopatry and de¬ gree of male-biased dispersal between breeding areas. Since extant populations of such species can exhibit relatively high levels of connection in a matriarchal genealogy, estimates of contemporary female movement (e.g., through results of direct observations or banding returns) may sometimes be quite misleading as indicators of the magnitude of historical interpopulation gene flow. 5) Avian populations appear to be structured at a variety of evolutionary timescales. In addition to the deep mtDNA phylogeographic structures of some species, which probably evidence long-term biogeographic separations, many populations also show significant haplotype frequency differences, suggesting further restrictions on gene flow over ecological timescales. Such contemporary structure is consist¬ ent with the limited realized vagiiity of most bird species relative to the size of their respective ranges, and probably represents a general isolation by distance rather than long-term vicariant separations of singular significance. LITERATURE CITED AVISE, J.C. 1989. Gene trees and organismal histories: a phylogenetic approach to population biol¬ ogy. Evolution 43: 1192-1208. AVISE, J.C., ANKNEY, C.D., NELSON, W.S. 1990. Mitochondrial gene trees and the evolutionary re¬ lationship of Mallard and Black Ducks. Evolution, 44: 1 109-1 1 19. AVISE, J.C., ARNOLD, J., BALL, R.M., BERMINGHAM, E., LAMB, T., NEIGEL, J.E., REEB, C.A., SAUNDERS, N.C. 1987. Intraspecific phylogeography: the mitochondrial DNA bridge between popu¬ lation genetics and systematics. Annual Review of Ecology and Systematics 18: 489-522. AVISE, J.C., BALL, R.M. 1990. Principles of genealogical concordance in species concepts and bio¬ logical taxonomy. Oxford Surveys in Evolutionary Biology 7: 45-67. AVISE, J.C., BALL, R.M., ARNOLD, J. 1988. Current versus historical population sizes in vertebrate species with high gene flow: a comparison based on mitochondrial DNA lineages and inbreeding theory for neutral mutations. Molecular Biology and Evolution 5: 331-344. AVISE, J.C., NELSON, W.S. 1989. Molecular genetic relationships of the extinct Dusky Seaside Spar¬ row. Science 243: 646-648. 524 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI AVISE, J.C., VRIJENHOEK, R.C. 1987. Mode of inheritance and variation of mitochondrial DNA in hybridogenetic fishes of the genus Poeciliopsis. Molecular Biology and Evolution 4: 514-525. AVISE, J.C., ZINK, R.M. 1988. Molecular genetic divergence between avian sibling species: King and Clapper Rails, Long-billed and Short-billed Dowitchers, Boat-tailed and Great-tailed Grackles, and Tufted and Black-crested Titmice. The Auk 105: 516-528. BALL, R.M., JR., FREEMAN, S., JAMES, F.C., BERMINGHAM, E„ AVISE, J.C. 1988. Phylogeographic population structure of Red-winged Blackbirds assessed by mitochondrial DNA. Proceedings of the National Academy of Sciences USA 85: 1558-1562. BALL, R.M., JR., NEIGEL, J.E., AVISE, J.C. 1990. Gene genealogies within the organismal pedigrees of random mating populations. Evolution 44:360-370. BRAUN, M.J., ROBBINS, M.B. 1988. Extensive protein similarity of the hybridizing chickadees Parus atricapillus and P. carolinensis. The Auk 103: 667-675. BROWN, W.M., GEORGE, M., JR., WILSON, A.C. 1979. Rapid evolution of animal mitochondrial DNA. Proceedings of the National Academy of Sciences USA 76: 1967-1971. COOKE, F., MACINNES, C.D., PREVETT, J.P. 1975. Gene flow between breeding populations of the lesser snow goose. The Auk 92: 493-510. DESJARDINS, P., MORAIS, R. 1990. Sequence and gene organization of the chicken mitochondrial genome. J. Mol. Biol. 212: 599-634. FUNDERBURG, J.B., JR., QUAY, T.L. 1983. Distributional evolution of the Seaside Sparrow. Pp. 19- 27 in Quay, T.L., Funderburg, J.B., Jr., Lee, D.S., Potter, E.F., Robbins, C.S. (Eds). The Seaside Spar¬ row, Its Biology and Management. Raleigh, North Carolina State Museum of Natural History. GELTER, H.P. 1989. Genetic and behavioural differentiation associated with speciation in the flycatch¬ ers Ficedula hypoleuca and F. albicollis. Doctoral dissertation. Sweden, Uppsala University. GLAUS, K.R., ZASSENHAUS, H.P., FECHHEIMER, N.S., PERLMAN, P.S. 1980. Avian mtDNA: struc¬ ture, organization and evolution. Pp. 131-135 in Kroon, A.M. & Saccone, C. (Eds). The Organization and Expression of the Mitochondrial Genome. Amsterdam, Elsevier/North Holland. GREENWOOD, P.J. 1980. Mating systems, philopatry and dispersal in birds and mammals. Animal Behaviour 28: 1140-1162. HELM-BYCHOWSKI, K.M. 1984. Evolution of nuclear and mitochondrial DNA in Gallinaceous birds. Unpub. PhD dissertation. University of California, Berkeley. JAMES, F.C. 1983. Environmental component of morphological differentiation in birds. Science 221: 184-186. KESSLER, L.G., AVISE, J.C. 1984. Systematic relationships among waterfowl (Anatidae) inferred from restriction endonuclease analysis of mitochondrial DNA. Systematic Zoology 33: 370-380. KESSLER, L.G., AVISE, J.C. 1985. A comparative description of mitochondrial DNA differentiation in selected avian and other vertebrate genera. Molecular Biology and Evolution 2: 109-125. MACK, A.L., GILL, F.B., COLBURN, R., SPOLSKY, C. 1986. Mitochondrial DNA: a source of genetic markers for studies of similar passerine bird species. The Auk 103: 676-681. NEI, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of in¬ dividuals. Genetics 89: 583-590. OVENDEN, J.R., MACKINLAY, A.G., CROZIER, R.H. 1987. Systematics and mitochondrial genome evolution of Australian rosellas (Aves: Platycercidae). Molecular Biology and Evolution 4: 526-543. SHIELDS, G.F., HELM-BYCHOWSKI, K.M. 1988. Mitochondrial DNA of birds. Pp. 273-295 in Johnston, R.F. (Ed.). Current Ornithology, Vol. 5. New York, Plenum Press. SHIELDS, G.F., WILSON, A.C. 1987. Calibration of mitochondrial DNA evolution in geese. Journal of Molecular Evolution 24: 212-217. TEGELSTROM, H. 1987. Genetic variability in mitochondrial DNA in a regional population of the Great Tit ( Parus major). Biochemical Genetics 25: 95-1 10. VAN WAGNER, C.E., BAKER, A.J. 1990. Association between mitochondrial DNA and morphological evolution in Canada geese. Journal of Molecular Evolution 31: 373-382. WATANABE, T., MIZUTANI, M., WAKANA, S., TOMITA, T. 1985. Demonstration of the maternal in¬ heritance of avian mitochondrial DNA in chicken-quail hybrids. Journal of Experimental Zoology 236: 245-247. WILSON, A.C., CANN, R.L., CARR, S.M., GEORGE, M., JR., GYLLENSTEN, U.B., HELMBYCHOWSKI, K.M., HIGUCHI, R.G., PALUMBI, S.R., PRAGER, E.M., SAGE, R.D., STONEKING, M. 1985. Mitochondrial DNA and two perspectives on evolutionary genetics. Biological Journal of the Linnaen Society 26: 375-400 . ZINK, R.M. 1991. Geography of mitochondrial DNA variation in two sympatric sparrows. Evolution, in press. ZINK, R.M., AVISE, J.C. 1990. Patterns of mitochondrial DNA and allozyme evolution in the avian genus Ammodramus. Systematic Zoology 39: 148-161. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 525 POPULATION DIFFERENTIATION IN NEW ZEALAND BIRDS CHARLES H. DAUGHERTY1 and SUSAN J. TRIGGS2 1 School of Biological Sciences, Victoria University, P.O. Box 600, Wellington, New Zealand 2 Science and Research Division, Department of Conservation, P.O. Box 10-420, Wellington, New Zealand ABSTRACT. Because New Zealand is an archipelago of three main islands with numerous offshore islands, many species of birds are fragmented into disjunct populations. The destruction of habitats and the introduction of mammalian predators by humans in the last 1000 years has further fragmented populations and reduced population sizes. Avian species that were once widely distributed, or still are, on the main islands often show clinal (Little Blue Penguin) or regional (Brown Kiwi, Yellow-crowned Parakeet) genetic differentiation. Main island species with naturally limited dispersal (Blue Duck) show local differentiation, as do some island isolates (Red-crowned Parakeet, Brown Teal, Yellow-eyed Penguin). Environmental alteration appears to contribute significantly to inter-specific hybridisation threatening some native species (Black Stilt, Forbes Parakeet, Grey Duck). Geographic patterns of genetic variation do not support present taxonomic classifications of some species. Keywords: New Zealand birds, allozymes, systematics, geographic variation, population differentia¬ tion, hybridisation. INTRODUCTION New Zealand has been isolated from all other landmasses since its separation from Gondwanaland about 80 million years ago (Stevens et al. 1988). This has led to a distinctive avifauna of approximately 300 species (Robertson 1985), including one endemic order, five endemic families, and 24 endemic genera. About 50 more spe¬ cies, including many endemics and such spectacular species as the moas (Dinornithiformes) and the Giant Eagle ( Harpagornis ), have become extinct since the arrival of humans about 1000 years ago. Ancient endemics such as Kiwi1 Apteryx, Kakapo Strigops habroptilus, New Zealand Wrens (Xenicidae), and Wattlebirds (Callaeidae) are not closely related genetically to other birds and have evolved unique morphological and behavioural features. On the other hand, much of the avifauna arrived in New Zealand in more recent evolution¬ ary times (e.g., Parakeets, Cyanoramphus spp.; Stilts, Himantopus spp.), and these species clearly show their affinities to overseas relatives. Since about 1850 some Australian species have established, apparently without human assistance (e.g., Welcome Swallow, Hirundo tahitica; Silvereye, Zosterops lateralis), and other species have been human-assisted settlers (e.g., House Sparrow, Passer domesticus; Star¬ ting, Sturnus vulgaris ; Indian Myna, Acridotheres tristis). In this paper we summarise and discuss patterns and levels of intra-specific differen¬ tiation of the extant New Zealand avifauna, focusing on native species for which 1 Common and scientific names follow Kinsky (1970) or Robertson (1985), unless otherwise stated. 526 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI genetic (usually allozyme) data are available. We relate these patterns to the recent ecological history of New Zealand and inferred historical patterns of distribution. Allan Baker and co-workers have assessed in detail the population differentiation of as¬ sisted immigrants (e.g., House Sparrow, Baker 1980; Indian Myna, Baker and Mooed 1979,1987; Starling, Ross 1983); we do not deal with these species here. SYSTEMATICS AND POPULATION DIFFERENTIATION Taxonomies describe the distribution of variation in natural populations, both within and between species (Corbin 1983). In recent decades the primary sources for tax¬ onomy of New Zealand birds have been Oliver (1930, 1955) and Kinsky (1970). These landmark volumes and the sources they draw upon often suffer from the general prob¬ lems described by Avise (1989) in identifying species and subspecies on the basis of few morphological traits, often not explicitly specified, and using outdated or unstated species or subspecies concepts. Taxonomy is the essential foundation for conservation practice, and instances of in¬ correct taxonomies have had costly consequences for conservation (Avise 1989, Avise & Nelson 1988, Daugherty et al. 1990a). Systematic analyses of New Zealand reptiles (Daugherty et al. 1990a, 1990b, Patterson & Daugherty 1990, R.A. Hitchmough, pers. comm.) using both allozyme and morphological characters have led to discovery of numerous cryptic species and re-assessment of the conservation status of many taxa. Taxonomic classifications of many New Zealand bird taxa have not been subjected to analysis using genetic data and contemporary systematic meth¬ odologies and may thus require significant revision. Genetic data have already revealed inadequacies in some longstanding taxonomic as¬ sessments of New Zealand birds. For example, Meredith and Sin (1988) demon¬ strated clinal variation in both allozyme and morphological characters of Blue Pen¬ guins Eudyptula minor along the east coast of both main islands, a pattern conflict¬ ing with the present division of Blue Penguins into five subspecies (Robertson 1985). Similarly, genetic data do not support the present taxonomy of Brown Kiwi. As many as eight species of kiwis were described last century. Oliver (1930) accepted four species, including two species of Brown Kiwi: Apteryx mantelli, the North Island Brown Kiwi, and A. australis, the South and Stewart Island Brown Kiwi. However, Oliver (1955) merged al! Brown Kiwis into a single species, A. australis, with three subspe¬ cies (North Island, South Island, Stewart Island). The pattern of allozyme variation (C.H. Daugherty et al., unpubi. data) agrees with the former taxonomy in defining two main geographic groups; a “North Island” type occurs as far south as Okarito on the west coast of the central South Island, and a second type occurs in Fiordland and Stewart Island. Parakeets provide a third instructive case. Allozyme variation in Cyanoramphus para¬ keets demonstrated that the endangered Chatham Island Yellow-crowned (or Forbes’) Parakeet is not a subspecies of the Yellow-crowned Parakeet C. auriceps as classi¬ fied by Oliver (1930,1955), but is instead a full species C. forbesi as originally de¬ scribed (Triggs & Daugherty, in press). Forbes’ Parakeet is most closely related to the Red-crowned Parakeet C. novaezelandiae novaezelandiae , with which it co-occurs. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 527 A complete study of differentiation in either Red-crowned or Yellow-crowned Para¬ keets could thus not be undertaken without first knowing the relationships of Forbes’ Parakeet. These examples demonstrate the necessity for basing conservation practice on ac¬ curate taxonomies. Surveys of population differentiation, even in taxa of the highest conservation significance, may often produce surprising results (Daugherty et al. 1990a). We expect that the use of formal systematic methodologies and new types of data, especially genetic data, will identify further inadequacies in the taxonomy of New Zealand birds. GEOGRAPHIC DISTRIBUTION AND POPULATION DIFFERENTIATION Until the arrival of humans, forest habitats were continuous over much of mainland New Zealand (Stevens et al. 1988). Human activity has reduced the forest cover of the two main islands from perhaps 85% to less than 25%. Farmlands and exotic for¬ est plantations have replaced indigenous forests. Lowland forest habitats are mostly small, isolated fragments. These changes have made the landscape more suitable for many introduced species and less satisfactory for natives. Many introduced mamma¬ lian species are efficient predators on native birds, especially the flightless ones. In¬ troduced birds are probably often effective competitors of native species. All these factors have altered distributions of native bird species. In this section we relate present patterns of geographic distribution to data on population differentiation. Species are arbitrarily categorised into five groups, on the basis of increasingly frag¬ mented levels of historical geographic distribution. Thus, the extent of differentiation of conspecific populations could be expected to increase with each group. GeographicalEy widespread, relatively abundant species Despite fragmentation of habitats, some native species such as Pukeko Porphyrio porphyrio, Tui Prosthemadera novaeseeiandiae , and Bellbird Anthornis melanura still occur widely throughout one or both islands. These species have adapted to modi¬ fied habitats and, in the case of the Pukeko that occupies farmland, may be far more widely distributed than in pre-human times. Two species in this category are the Blue Penguin and Brown Kiwi, both discussed earlier. The Blue Penguin appears to be highly variable over its entire range, but breeding populations diverge in a gradual, clinal fashion (Meredith & Sin 1988), as might be expected for abundant, geographically continuous populations. Brown Kiwi probably occurred continuously throughout all three main islands until the arrival of Europeans. Still limited to forests, they occur widely only where forests are extensive as in Fiordland and Stewart Island. In the North Island, Brown Kiwis are sometimes abundant in exotic pine plantations. Preliminary allozyme studies of small numbers of individuals from widely scattered locations throughout the entire range of Brown Kiwis (Daugherty et al., unpubi. data) reveal the two major geographic groups noted earlier. Taxonomic status of the two groups remains to be determined, but the level of differentiation (Nei’s D = 0.05) is about that often associated with speciation in birds (Barrowclough & Corbin 1978, Avise et al. 1980, Barrowclough 1983). 528 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Sample sizes are too small to resolve patterns of local variation. Nonetheless, polymorphisms are sufficiently common to suggest that genetic structuring may occur within the two main groups of Brown Kiwi. Geographically widespread species; populations once continuous, but now fragmented due to recent reductions in numbers or habitats Species in this category include mainland populations of Red-crowned Parakeets, Yellow-crowned Parakeets, and Weka Gallirallus australis. Genetic studies of Weka may be particularly revealing, as this species is presently separated into four subspe¬ cies that show significant colour variation (Robertson 1985). There may be some ur¬ gency to study Weka, as numbers appear to be diminishing throughout its present range. Genetic data for Yellow-crowned Parakeets (Triggs & Daugherty, in press) show that the species is apparently divided into two main geographic groups, one from the North Island and northern South Island, and a second from the central and southern South Island, a pattern generally resembling that of Brown Kiwis. The level of differentiation (D = 0.02) is roughly that associated with subspecies and sometimes species of birds. Variation in Yellow-crowned Parakeets must also take into account the extremely rare Orange-fronted Parakeet, originally described as a separate species C. malherbi but considered by many recent workers to be a colour morph of the Yellow-crowned Para¬ keet (Holyoak 1974, Taylor et al. 1986). Differences in allozyme frequencies exist between sympatric populations of Orange-fronted and Yellow-crowned Parakeets, but sample sizes are too small to allow statistical testing (Triggs & Daugherty in press). In view of the relatively limited genetic differentiation commonly found between con¬ generic species of birds (e.g., Barrowclough & Corbin 1978; Avise et al. 1980), reduc¬ tion of the taxonomic status of Orange-fronted Parakeets is premature. Formerly widespread species; numbers now greatly reduced Many species restricted to diminishing habitats such as forests are now greatly re¬ duced in numbers and distribution, e.g., Kokako Callaeas cinerea, Little Spotted Kiwi Apteryx owen/7, Kakapo, Blue Duck Hymenolaimus malacorhynchos, Saddleback Philesturnus carunculatus , and Stitchbird Notiomystis cincta. All were widespread until European times; some (Little Spotted Kiwi, Saddleback, Stitchbird) now survive only on a few offshore islands. The Little Spotted Kiwi, once found widely on both main islands, now survives in one population of some hundreds on Kapiti Island, and in several much smaller island populations originating from translocations of Kapiti Island birds. Comparison of allozymes of a single wild individual discovered on D’Urville Island with birds from Kapiti Island showed no difference (C.H. Daugherty & R. Colbourne, unpubl. data). Fewer than 50 Kakapo now survive. Comparison of allozyme frequencies of a popu¬ lation in Fiordland, where numbers have been reduced to just a few individuals, with a population from Stewart Island numbering about fifty birds, showed no significant differences (Triggs et al. 1989). Average heterozygosity in the Stewart Island popu¬ lation (H = 0.04) was similar to the average for other avian species (Barrowclough 1983, Barrowclough et al. 1985), while that of the Fiordland population was somewhat lower (H = 0.02), possible evidence of the genetic bottleneck. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 529 Blue Duck populations have been studied by both allozyme and DNA methodologies (Triggs et al. in press). This species was widespread in pre-European times, and its riverine distribution was essentially one-dimensional. DNA fingerprinting studies indi¬ cate that local populations consist of highly interrelated individuals, a result of strong natal philopatry and perhaps also recent isolation into river headwaters. Thus, exten¬ sive genetic differentiation occurs over relatively short distances in this species. Geographically widespread, naturally fragmented populations This type of distribution characterises species that occur on the outlying islands of New Zealand, especially the sub-Antarctic islands. They might be expected to show higher levels of population differentiation than species with continuous or recently fragmented distributional patterns. Very different levels of divergence characterise the two taxa examined genetically thus far. Red-crowned parakeets occur on most major outlying island groups, and variation in blood allozymes has been investigated in detail (Triggs & Daugherty in press). Ge¬ netic differentiation between island groups is low (D < 0.02), in line with levels found between subspecies and conspecific populations of other birds (Barrowclough 1983). The single individuals sampled of the Kermadec Parakeet C. n. cyanurus and Reischek’s parakeet C. n. hochstetteri from the Antipodes Islands, over 2000 km dis¬ tant from one another, are identical at 21 loci examined. Populations of the New Zea¬ land Red-crowned Parakeet C. n. novaezelandiae on three island groups near the North Island (Three Kings Islands, Poor Knights Islands, Little Barrier Island) are highly polymorphic at five of these loci but little differentiated, clustering as each oth¬ er’s closest relatives in support of their present taxonomic classification. The low lev¬ els of genetic differentiation may reflect recent occupation of islands by this species, or it may indicate recurrent gene flow and large population sizes. Allozyme variation in one of the world’s rarest penguins, the Yellow-eyed Penguin ( Megadyptes antipodes; fewer than 3000 breeding individuals), have been compared from three widely separated locations (the east coast of the South Island, Campbell Island, and Enderby Island in the Auckland Island group). Differentiation among these groups (Fst = 0.24, Triggs & Darby 1989) was about an order of magnitude higher than among other geographically widespread species of birds (Barrowclough 1983), indicating that gene flow among the three locations is extremely limited. In marked contrast to the relatively undifferentiated Red-crowned Parakeets, however, the high levels of genetic differentiation in Yellow-eyed Penguins are not reflected in any taxo¬ nomic subdivision into subspecies. Species with naturally limited distributions Many island endemic populations of species, such as the rare Chatham Island Black Robin Petroica traversi, may always have been small. Studies of these island isolates may reveal little differentiation from their closest relatives, as found, for example, among some island populations of the Red-crowned Parakeets. The Forbes’ Parakeet shows limited differentiation (D = 0.01) from its probable closest relative, the sympatric Chatham Island Red-crowned Parakeet (Triggs & Daugherty in press). This suggests a recent origin for the taxon, although the situation is confounded by recent inter¬ breeding between the species. Levels of differentiation can also be high. Brown Teal Anas aucklandica, for example, are presently separated into three subspecies (mainland New Zealand, Auckland Is- 530 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI lands, and Dent Island in the Campbell Island group). Allozyme divergence among these subspecies (D = 0.07-0.15) substantially exceeds the level usually associated with subspecies, suggesting that these taxa deserve full specific recognition (M.J. Williams et al . , unpubl. data). HYBRIDISATION In addition to geographic distribution, hybridisation also affects differentiation within genera or species of some New Zealand birds. Three cases of hybridisation, all origi¬ nally recognised by the presence of morphologically intermediate birds, have now been investigated using allozymes. All are associated with human modification of natural habitats and are of conservation significance. Grey Duck In recent decades, numbers of the introduced Mallard, Anas platyrhynchos, have in¬ creased throughout New Zealand at the expense of the native Grey Duck, Anas superciliosa. Mallards now outnumber Grey Duck by 4:1 throughout the country (Robertson, 1985). Allozyme analysis (Hitchmough et al. 1990) showed virtually no polymorphism and thus no detectable genetic difference between birds characterised morphologically as Grey Ducks, Mallards, or hybrids. The absence of polymorphism in Mallards is surprising in view of relatively high heterozygosity (H = 0.05) in North American populations (Ankney et al. 1986). Grey Ducks may persist because hybrids mate prefentially with Mallards. On the other hand, the numerical imbalance favour¬ ing Mallards may eventually lead to the demise of Grey Ducks. Black Stilts The Black Stilt, Himantopus novaezelandiae, has experienced a severe decline fol¬ lowing introduction of numerous mammalian predators, to which it seems especially susceptible, and extensive habitat changes associated with European settlement. The latter may have favoured the Pied Stilt, Himantopus h. leucocephalus which has greatly expanded its range and numbers (Robertson 1985). The two species have hybridised extensively (Pierce 1984). Allozyme studies showed that the few (~50) remaining Black Stilts are genetically distinct from Pied Stilts despite the extensive history of hybridisation (Green 1988); that is, Blacks are not simply a colour morph of Pied Stilts. Captive breeding pro¬ grammes to save the Black Stilt are now a central focus of its management. Parakeets The best known instance of hybridisation among New Zealand birds involves Forbes’ and Red-crowned Parakeets on Mangere and Little Mangere Islands in the Chatham Islands group. Habitat changes that reduced optimum habitat for Forbes’ Parakeets may be the cause of hybridisation that threatened the Forbes’ Parakeet, which oc¬ curred only on these two small islands (Cade 1983). The New Zealand Wildlife Serv¬ ice responded with a programme to remove (by shooting) all Red-crowned Parakeets and hybrids from these islands, and to re-establish suitable habitat for the Forbes’. Allozyme studies (Triggs & Daugherty in press) showed that morphologically “pure” Forbes’ Parakeets have retained their genetic distinctness from Red-crowned Para¬ keets, probably because hybrids breed mainly with Red-crowned Parakeets. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 531 Barriers to hybridisation may be low among New Zealand Cyanoramphus Parakeets. Hybrids of Red-crowned and Yellow-crowned Parakeets have now been identified on Little Barrier Island (Triggs & Daugherty in press), and most birds on the Auckland Islands have recently been reported to be hybrids of these two species (Taylor 1985, G. Elliott, pers. comm.). The demise of the Orange-fronted Parakeet, if it is truly a distinct species, could also be the result of hybridisation with the Yellow-crowned Parakeet (Triggs & Daugherty in press). CONCLUSIONS Few studies of geographic variation and population differentiation in New Zealand birds have used either contemporary systematic methodologies or genetic data. Ex¬ isting taxonomies often do not adequately describe variation for taxa where such stud¬ ies have been conducted, e.g., Brown Kiwi* Blue Penguins, Yellow-crowned Para¬ keets, Yellow-eyed Penguins, and Brown Teal. Systematic studies of geographic vari¬ ation can be expected to contribute much to conservation of species yet to be exam¬ ined similarly, such as Weka and Great Spotted Kiwi Apteryx haastii. Human habitation of New Zealand has greatly reduced forest habitats for birds. At the same time, new habitats (farmlands, pine plantations) have been occupied either by self-introduced species from Australia or by human-introduced species that may com¬ pete with native species. Populations of native birds have been reduced and their dis¬ tributions fragmented. Nonetheless, patterns of differentiation of native birds can often be related to prob¬ able pre-human patterns of distribution. Prior to human occupation of New Zealand, species such as the Brown Kiwi and Yellow-crowned Parakeet, for example, occurred continuously over wide areas. Present limited differentiation among populations of the North island and northern South Island supports an inference of high recent levels of gene flow. Similarly, the Blue Penguin, distributed more or less continuously down the east coast of both main islands, shows clinal differentiation in morphology and aliozymes. Some geographic barriers seem to have had limited effect in isolating populations. Cook Strait, for example, appears to have been no barrier to gene flow in either Yel¬ low-crowned Parakeets or the flightless Brown Kiwi, a pattern also found in the widely distributed New Zealand common skink, Leiolopisma nigriplantare (Daugherty et al. 1990b). Lack of significant population differentiation in Red-crowned Parakeets be¬ tween some oceanic islands may reflect either recency of invasion of these locations or recent gene flow and large population sizes. On the other hand, social structure (Blue Duck) and geographic barriers (Yellow-eyed Penguins, Brown Teal) may function to limit gene flow and lead to infra-specific dif¬ ferentiation of some avian species. Finally, recent habitat alterations are associated with hybridisation, primarily among taxa that were poorly differentiated to begin with. For Forbes’ Parakeets, Black Stilts, and Grey Ducks, the weakness of barriers to hybridisation places their future in doubt. 532 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI ACKNOWLEDGEMENTS We thank R.A. Hitchmough and D.G. Newman for constructive commentary that much improved an early draft of this manuscript. LITERATURE CITED ANKNEY, C.D., DENNIS, D.G., WISHARD, L.N., SEEB, J.E. 1986. Low genic variation between black ducks and mallards. Auk 103: 701-709. AVISE, J.C. 1989. A role for molecular genetics in the recognition and conservation of endangered species. Trends in Ecology and Evolution 4: 279-281. AVISE, J.C., NELSON, W.S. 1988. Molecular genetic relationships of the extinct dusky seaside spar¬ row. Science 243: 646-648. AVISE, J.C., PATTON, J.C., AQUADRO, C.F. 1980. Evolutionary genetics of birds II. Conservative protein evolution in North American sparrows and relatives. Systematic Zoology 29: 323-334. BAKER, A.J. 1980. Morphometric differentiation in New Zealand populations of the house sparrow {Passer domesticus). Evolution 34: 638-653. BAKER, A.J., MOOED, A. 1979. Evolution in the introduced New Zealand populations of the common myna, Acridotheres tristis. Canadian Journal of Zoology 57: 570-584. BAKER, A.J., MOOED, A. 1987. Rapid genetic differentiation and founder effect in colonizing populations of common mynas {Acridotheres tristis). Evolution 41: 525-538. BARROWCLOUGH, G.F. 1983. Biochemical studies and microevolutionary processes. Pp. 223-261 in Brush, A.H, Clark, G.A. (Eds). Perspectives in ornithology. Cambridge, England, Cambridge Univer¬ sity Press. BARROWCLOUGH, G.F., CORBIN, K.W. 1978. Genetic variation and differentiation in the Parulidae. Auk 95: 691-702. BARROWCLOUGH, G.F., JOHNSON, N.K., ZINK, R.M. 1985. On the nature of genic variation in birds. Pp. 135-154 in Johnston, R.F. (Ed.). Current ornithology, vol. 2. New York, Plenum Press. CADE, T.J. 1983. Hybridization and gene exchange among birds in relation to conservation. Pp. 288-309 in Schonewald-Cox, C.M., Chambers, S.M., MacBryde, B., Thomas, W.L. (Eds). Genetics and conservation. Menlo Park, California, The Benjamin/Cummings Publishing Company, Inc. CORBIN, K.W. 1983. Genetic structure and avian systematics. Pp. 211-244 in Johnston, R.F. (Ed.). Current ornithology, vol. 1. New York, Plenum Press. DAUGHERTY, C.H., CREE, A., HAY, J.M., THOMPSON, M.B. 1990a. Neglected taxonomy and con¬ tinuing extinctions of tuatara {Sphenodon). Nature 347: 177-179. DAUGHERTY, C.H., PATTERSON, G.B., THORN, C.J., FRENCH, D.C. 1990b. Differentiation of mem¬ bers of the New Zealand Leiolopisma nigriplantare complex (Sauria: Scincidae). Herpetological Mono¬ graphs 4: 61-75. GREEN, B.S. 1988. Genetic variation and management of black stilts, Himantopus novaezelandiae, and pied stilts, Himantopus himantopus leucocephalus (O. Charadriiformes). M.Sc. (Hons) Thesis, Victoria University of Wellington. HITCHMOUGH, R.A., WILLIAMS, M., DAUGHERTY, C.H. 1990. A genetic analysis of mallards, grey ducks and their hybrids in New Zealand. New Zealand Journal of Zoology 17: 467-472. HOLYOAK, D.T. 1974. Cyanoramphus malherbi, is it a colour morph of C. auriceps ? Bulletin of the British Ornithological Club 94: 4-9. KINSKY, F.C., Convener, The Checklist Committee, Ornithological Society of New Zealand. 1970. Annotated checklist of the birds of New Zealand, including the birds of the Ross Dependency. Welling¬ ton, A.H. & A.W. Reed for the Ornithological Society of New Zealand. MEREDITH, M.A.M., SIN, F.Y.T. 1988. Genetic variation of four populations of the little blue penguin, Eudyptula minor. Heredity 60: 69-76. OLIVER, W.R.B. 1930. New Zealand Birds. 1st ed. Wellington, Fine Arts (NZ). OLIVER, W.R.B. 1955. New Zealand Birds. 2nd ed. Wellington, A.H. & A.W. Reed. PATTERSON, G.B., DAUGHERTY, C.H. 1990. Four new species and one new subspecies of skink, genus Leiolopisma (Reptilia: Lacertilia: Scincidae) from New Zealand. Journal of the Royal Society of New Zealand 20: 65-84. PIERCE, R.J. 1984. Plumage, morphology and hybridisation of New Zealand stilts Himantopus spp. Notornis 31 : 106-130. ROBERTSON, C.J.R., (Ed.). 1985. The Reader's Digest Complete Book of New Zealand Birds. Syd¬ ney, Reader’s Digest Services Pty Limited. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 533 ROSS, H.A. 1983. Genetic differentiation of starling ( Sturnus vulgaris: Aves) populations in New Zea¬ land and Great Britain. Journal of Zoology, London 201: 351-362. STEVENS, G., MCGLONE, M., MCCULLOCH, B. 1988. Prehistoric New Zealand. Auckland, Heinemann Reed. TAYLOR, R.H. 1985. Status, habits and conservation of Cyanoramphus parakeets in the New Zealand region. Pp. 195-211 in Moors, P.J. (Ed.). Conservation of island birds. Cambridge, ICBP Technical Publication No. 3. TAYLOR, R.H. HEATHERBELL, E.G., HEATHERBELL, E.M. 1986. The orange-fronted parakeet ( Cyanoramphus malherbi) is a colour morph of the yellow-crowned parakeet (C. auriceps). Notornis 33: 17-22. TRIGGS, S.J., DARBY, J. 1989. Genetics and conservation of yellow-eyed penguin: an interim report. Science and research internal report no. 43, New Zealand Department of Conservation, Wellington. TRIGGS, S.J., DAUGHERTY, C.H. In press. Conservation and genetics of New Zealand parakeets. In Bell, B.D. (Ed.). The management of threatened species. ICBP Technical Bulletin. TRIGGS, S.J., POWLESLAND, R.G., DAUGHERTY, C.H. 1989. Genetic variation and conservation of kakapo ( Strigops habroptilus: Psittaciformes). Conservation Biology 3: 92-96. TRIGGS, S.J., WILLIAMS, M.J., MARSHALL, S., CHAMBERS, G.K. In press. Genetic relationships within a population of Blue Ducks ( Hymenolaimus malacorhynchos). Wildfowl 42. 534 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI PHENETIC VARIATION IN NEARCTIC EMBERIZINAE JAMES D. RISING Department of Zoology, University of Toronto, Toronto, Ontario, M5S 1A1, Canada ABSTRACT. I examined phenetic relationships, based on 24 skeletal measures, of 42 taxa (of 36 cur¬ rently recognized species) in 15 genera of North American Emberizinae. In spite of the taxonomic di¬ versity of the group, they seem to be remarkably similar in proportions, differing among taxa primarily in size. Nonetheless, taxa that live in short-grass or short tundra habitats, in which high winds are characteristic, seem to have relatively long wings, short toe bones (though long claws), and relatively short, conical bills. Marsh-dwelling species, on the other hand, tend to have relatively short wings, long legs and toes, and long bills. In general, species within a genus cluster together, although there is commonly a great deal of overlap among genera. The variability among populations of the Savannah Sparrow is as great as that among species in other genera. Closely related species that co-occur in the same habitat often are quite similar in size and proportions, indicating that interspecific competi¬ tion has not played an important role in the evolution of morphological differences among species in this group. Keywords: Emberizinae, phenetic similarity, habitat overlap. INTRODUCTION The Emberizinae (Emberizidae), or Old World Buntings and New World Sparrows, contain 65 recognized genera and 279 species, and thus are one of the largest sub¬ families of birds (containing about 3% of all avian species) (Bock & Farrand 1980). The group is well represented in the Nearctic: 17 genera and 47 species breed and occur regularly north of Mexico (AOU Check-list of North American Birds, 1983). Here I examine phenetic variation among a wide range of taxa of North American sparrows, buntings, and longspurs, and relate this variability to the interspecific and, in two instances, the intraspecific variation in typical habitat use and behaviour of the populations. METHODS The species and populations measured for the phenetic analyses are listed in Table 1 . Most of the specimens measured were collected recently and prepared as skin-and- skeleton specimens, and donated to the Royal Ontario Museum, but additional ma¬ terial was borrowed from several other museums (see list in Acknowledgements) to augment samples so that at least 10 individuals were measured for all but one of the taxa. For this paper, only data on males are reported; all of these species are sexu¬ ally dimorphic in size (Rising in prep.), and numbers of specimens of females avail¬ able are generally substantially smaller than numbers of male specimens. I measured the 24 features listed in Table 2 on the prepared skeleton of each speci¬ men. Measurements were taken to the nearest 0.1 mm, and were taken as illustrated by Robins & Schnell (1971), with the following exceptions: premaxilla length is from ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 535 the anterior edge of the narial opening; premaxilla depth is maximal depth of the premaxilla; and synsacrum width is the distance between the antitrochanters. In gen¬ eral, interpopulational variation in size is not considered in this study. Nonetheless, because of considerable geographic variation in size in many of these species, care was taken to use only specimens from a limited region for conspicuously variable species (Table 1). Several populations of some notably polytypic species were used when sufficient material was available. Specifically, four populations of Savannah Sparrows ( Passerculus sandwichensis s. /.) and Sharp-tailed Sparrows ( Ammodramus caudacutus s. /.) were used. In both instances these represent well differentiated taxa that have, at various times, been recognized as different species or at least as well- marked subspecies. These were the “Ipswich” Sparrow (P. s. princeps), “typical” Sa¬ vannah Sparrow (P. s. labradorius), “Belding’s” Sparrow (P. s. beldingi), and “Large¬ billed” Sparrow (P. s. rostratus ); and “typical” Sharp-tailed Sparrow (A. c. caudacutus ), “James Bay” Sharp-tailed Sparrow (A. c. altera), “Nelson’s” Sparrow (A. c. nelsoni), and “Acadian” Sharp-tailed Sparrow (A. c. subvirgata). TABLE 1 - Species, numbers, and samples of Emberizine sparrows, buntings, and longspurs measured for phenetic analyses Species N Localities from which specimens taken Green-tailed Towhee Pipilo chlourus 8 Az(5); Tx(2); ld(1) Rufous-sided Towhee P. erythrophthalmus 11 Ks(4); De(3); Ont(3); Ma(1); Mo(1) Bachman’s Sparrow Aimophila aestigalis 10 Tx(4); Fl(4); Ky(2) Cassin’s Sparrow A. cassini 17 Tx(9); NM(5); Ks(3) Rufous-winged Sparrow A. carpal is 19 Mex(1 1 ); Az(8) Rufous-crowned Sparrow A. ruficeps 10 Tx(5); Mex(3); NM(1); Az(1) American Tree Sparrow Spizella arborea 10 Man(5); Ont(4); Ak(1 ) Chipping Sparrow S. passerina 11 Ont(IO); Mo(1 ) Clay-colored Sparrow S. pallida 15 Sask(1 5) Brewer’s Sparrow S. breweri 11 Az(3); Tx(1 ); Aita(1); Co(1); Ok(1); Mt(1); Mex(1 ) Field Sparrow S. pusilla 11 Ks(5); Ont(4); Ar(1); Tx(1) Vesper Sparrow Pooecetes gramineus 35 Sask(1 1 ); Tx(6); Alta(4); Az(3); ND(3); Ont(3); WVa(2); BC(1); Ca(1); Nv(1) Lark Sparrow Chondestes grammacus 14 Tx(6); Ks(4); Wy(2); Ca(1); NM(1) Black-throated Sparrow Amphispiza bilineata 10 Az(5); Tx(3); NM(2) Lark Bunting Calamospiza meianocorys 32 Tx(1 6); Sask(9); Ks(5); Ok(2) Savannah Sparrow Passerculus sandwichensis labradorius 49 Moosonee, Ontario 536 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI TABLE 1 - continued Species N Localities from which specimens taken “Ipswich” Sparrow P. s. princeps 17 Sable Is., Nova Scotia “Belding’s” Sparrow P. s. beldingi 21 Bahia de San Quintin, Baja Cal. N. “Large-billed” Sparrow P. s. ro st rat us 20 Bahia Kino, Sonora Baird’s Sparrow Ammodramus bairdii 29 Sask(14); ND(12); Grasshopper Sparrow A. savannarum 21 ND(8); Tx(4); WVa(4); Az(2); Ks(1 ); N B( 1 ) ; Ont(1) Henslow’s Sparrow A. henslowii 15 Mi(5); Ks(4); Md(2); Oh(2); Ont(2) LeConte’s Sparrow A. leconteii 31 ND(1 0); Ont(10); Ks(8); Ky(1); Man(1 ) Sharp-tailed Sparrow A. c. caudacutus 21 Prime Hook Refuge, Delaware “Acadian” Sharp-tailed Sparrow A. c. subvirgata 17 Sackville, New Brunswick “James Bay” Sharp-tailed Sparrow A. c. altera 20 Moosonee, Ontario “Nelson’s” Sharp-tailed Sparrow A. c. nelsoni 14 Upham, North Dakota Seaside Sparrow A. maritimus 31 De(1 7); NC(7); NJ(6); NY(1) Fox Sparrow Passerella iliaca 19 Ont(9); Ks(6); Song Sparrow Melospiza melodia 16 Ont(6); NB(3); Que(3); Man(2); Ok(1); WVa (1) Lincoln’s Sparrow M. lincolnii 13 Ont(10); Que(2); Nfld(1) Swamp Sparrow M. georgiana 12 Ont(7); Mi (4) ; Nfld(1) / White-throated Sparrow Zortotrichia albicollis 10 Ont(10) Golden-crowned Sparrow Z. atricapilla 11 BC(7); Ca(4) White-crowned Sparrow Z leucophrys 72 NWT(29); Que(25); Man(18) Harris’s Sparrow Z querula 12 Ks(8); NWT(4) Dark-eyed Junco Junco hyemalis 33 Ont(33) McCown’s Longspur Calcarius mccownii 18 Sask(8); Mt(5); Alta(3); Ks(2) Lapland Longspur C. lapponicus 19 Coppermine, NWT Smith’s Longspur C. pictus 14 Ont(7); NWT(4); Man(3) Chestnut-collared Longspur C. ornatus 21 Alta(6); Sask(6); ND(4); Mt(2); Tx(2); Ks(1 ) Snow Bunting Plectrophenax nivalis 10 Mi (6) ; Ont(3); Ak(1) ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 537 TABLE 2 - Correlations between 24 variables and principal component 1 and prin¬ cipal compontent 2 scores for four Emberizid samples (correlations < 0.35 are not Printed) Principal Component 1 Principal Component 2 Variable All Aa Bb Cc All A B C Skull Length .91 .68 .95 .97 -.36 -.72 Skull Width .91 .95 .96 .95 Premaxilla Length .68 .56 .90 .83 -.54 -.83 .42 Premaxilla Depth .86 .94 .93 Narial Width .93 .75 .98 .96 .49 Premaxilla Width .87 .95 .85 .59 .52 Interorbital Width .77 .45 .85 .89 -.55 Mandible Length .89 .62 .94 .95 -.39 -.77 Gonys Length .61 .52 .90 .66 -.68 -.79 .67 Mandible Depth .86 .41 .93 .85 Coracoid Length .96 .94 .93 .85 Scapula Length .92 .88 .97 .89 .36 .45 -.42 Femur Length .91 .84 .94 .95 -.47 Femur Width .91 .93 .93 .95 Tibiotarsus Length .89 .76 .92 .91 -.51 Tarsometatarsus Length .86 .82 .89 .88 -.35 Humerus Length .95 .95 .98 .83 .39 Ulna Length .80 .76 .85 .83 .55 .62 .49 Carpometacarpus Length .82 .76 .90 .61 .54 .52 .40 -.73 Hallux Length .48 .75 .82 -.49 -.50 Sternum Length .93 .84 .96 .98 .35 .52 Sternum Depth .77 .65 .84 .88 .56 .71 .51 Keel Length .87 .74 .92 .96 .45 .64 .36 Synsacrum Width .93 .87 .98 .68 -.68 % Variance Explained 72.6 54.4 85.5 75.0 13.2 26.0 7.7 11.4 aPasserculus & Ammodramus ; bAimophila, Spizella, Chondestes, Amphispiza, Passerella, Melospiza, Zonotrichia, & Junco: cPooecetes, Calamospiza, Calcarius, & Plectrophenax Averages were calculated for each measured character and each taxon, and these averages were used in phenetic analyses. Thus, variability within the samples is not assessed here. I used Principal Components Analysis (PCA) (NT-SYS Factor; Rohlf 1985), operating on a matrix of correlations among non-standardized or transformed measurements to extract multivariate measures of size and shape variation among species. This is one of several more-or-less equally suitable methods that could be used for our objectives (Rising & Somers 1989). In addition to summarizing variation among all 42 samples (of 36 species recognized by the 1983 AOU Check-list), I subdivided the samples into three smaller sets in or¬ der to focus attention on the variability within those groups. These groups were: (A) The Passerculus and Ammodramus group, several presumably closely-related species that live in grassland or in sedge or grass marshes. (B) The Aimophila, Chondestes, Amphispiza, Spizella, Passerella, Melospiza, Zonotrichia and Junco group, 20 species that occur in a variety of different Long Bills Long Wings - Long Toes p£ ^ ^ Long Wings -> & Legs PC 2 (13.2%) _ 538 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Chestnut-col. Smith’s Snow Bunting I Chipping Lapland i Brewer’s i Field Savannah Clay-colored Junco Black-thrt.® Rufous- ^Baird s winged Cassin’s^ Lincoln's A "James Henslow’s # Bay • • i "Nelson’s" ^ i Vesper 0 McCown’s • Lark S. "Ipswich” ® White-crowned Fox Lark B. Grasshopper White-throated I Harris’s LeConte’s I'Belding’s" "Large-billed" i Golden-crowned Bachman’s R •• Rufous-cr. ►Swamp "Acadian" Sharp-tail ► Green-tailed Towhee i Song Rufous-sided Towhee # Seaside All Taxa PC 1 (SIZE) (72.6%) Juncos & Allies • Lark S. 9 Chipping Fox 0 • Field £ Brewer s Black-throated £ # Junco Am. Tree w Clay-colored w Rufous-winged Harris’s 0 0 White-crowned £ White-throated • Cassin’s # Golden-crowned • Song Lincoln’s % ^ Bachman’s • Swamp # Rufous-crowned PC 1 (SIZE) 85.5% FIGURE 1 - The relationships among four different sets of Emberizids in the space defined by principal components 1 and 2 (PCI and PC2). The axes are explained in the text. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 539 3 1/5 0£ C £ o i c o A v> £ ■O C3 2 co J § * Grassland Emberizids ® Baird’s 0 Savannah 0 Grasshopper 0 LeConte’s 0 Henslow’s "Nelson’s" # #”James Bay 0 "Acadian" 0 Sharp-tail Seaside 0 PC 1 (SIZE) (54.4%) A V) £ OX) c o £ Tf jo - 5 CL P CL > 3 ~o C co — c -O o c CO ^ 0 ™ 0 iS C 2 o ■*"’ 0 o ‘5 0 _Q E 0 CD C o - § I 0 CO ^ 1 * o LU c _ 0 CD 0 < P I- ~o o- CD CD CD CO CD CD CD CD CD CD CD CD 1 T— CO Is- Is- D- Is- 1"- r- h- Is- h- Is- Is- 1 T— CM O o o O o o o 0 0 0 0 1 in uo T— CM CM y— O o o O o o o 0 0 0 1 uo in T— CM CM o O o o O o o o 0 0 1 0 uo uo CM CM O) O o o o o o o 0 1 0 0 in uo CM CM CO O o o o o o o 1 0 0 0 in uo CM CM 1^ o o o o o o 1 0 0 0 0 uo uo CM CM CO o o o o o 1 CM CM CM CM 00 CM in uo CM CM in o o o o 1 o CM CM CM CM CO CM in m CM CM o- o o o 1 _ ^ _ 00 00 00 CO •O’ 00 uo uo CM CM CO o o 1 o o CM CM CM CM CO CM uo in CM CM CM o 1 o o o CM CM CM CM CO CM uo uo CM CM 1 o o 1 — o o CM CM CM CM 00 CM m in CM CM c <1> m CD CO CO T- CM Is- 00 in CD in CD CM I-. o o CD CD 00 00 ■0- O’ 00 00 Is- CM 00 E 00 00 in in I — 1 — o 0 0 0 TCt’ •O’ CD Is- o in uo in in LO in •O’ •^ •M- m m uo w 0 D - D — 0 ~ D —> “0 “3 —> “0 —3 —3 —3 - 5 a 3*3 CO co z z Z z z z z z z z z z z 0 > 0 _0 LL — "0 0 >* E a I 0 _c > c 0 a (0 0 0 E 0 0 3 CD 0 ■4 — ' o o JT5 9 -C TO 0 jC 0 a 0 CO c 7a 4 — • 0 > o 0 0 0 _c 0 3 0 J*. _i CD O CL 0 cr E z 0 CO .0 .0 c 0 c 0 c: 0 o 0 0 ■0 CL 0 _ _ 0 0 c: 0* 0 ra 0 0 -Q 0 o 0 c: 0 0 0 0 0 0 c LS C -L; ,c o 0 0 0 a> X -0 -Q -Q -Q co -D (0 • - 0 1- T3 < CM CO ■O' m CD 00 CD 0 T_ CM CO •O’ 1 — T” 1 — 1 — 1 — 606 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI TABLE 4 - Average percent nucleotide difference among sequences of a 239-288 base pair segment of cytochrome b from taxa differentiated at several taxonomic levels.3 Taxonomic level Number of pairwise comparisons Average percent nucleotide difference*3 Range Average transition: transversion biasc lntrapopulationd 6 0.1 •0.0 - 0.4 0.3:0 Interpopulation within subspecies 12 0.4 0.0 - 1.0 1:0 Intraspecific (subspecies) 49 0.6 0.0 - 1.7 2:0 A. belli 6 48 0.6 0.0 - 1.0 2:0 A. bilineata 1 1.7 — 4:1 Interspecific congeners' A. belli v A. bilineata 24 10.9 10.8 - 11.1 25:7 Interfamilial Amphispiza v Corcorax 14 13.9 13.4 - 15.9 17:17 Amphispiza v Pomatostomas 56 17.4 15.5 - 18.8 22:19 Interordinal Amphispiza v Gallus 14 19.7 19.1 - 20.1 26:31 3 Comparisons within Amphispiza and between Amphispiza and Gallus (Desjardins & Morais 1990) are based on 288 bp sequences. Interfamilial comparisons between Amphispiza and either Pomatostomas or Corcorax are based on 239 bp sequences (Kocher et al. 1989) homologous to a segment of the Amphispiza sequences. Four species of Pomatostomus (P. ruficeps, P. superciliosus, P. temporalis, P. isidori) and Corcorax melanorhamphos (Kocher et al. 1989) were used in the interfamilial compari¬ sons. b Calculated as the average number of pairwise nucleotide differences (transitions + transversions) divided by the total number of bases compared. c Calculated as the number of transitions:transversions averaged over all pairwise comparisons. d Pairwise comparison of two individuals of A. belli from each of six populations (Beegum, Castaic, Panoche Hills, Chalfant Valley, Rattlesnake Flat, Plush). e Pairwise comparison of two populations (two individuals from each) representing three subspecies of the Sage Sparrow {A. b. belli, A. b. canescens, A. b. nevadensis). ' Pairwise comparison between three subspecies of the Sage Sparrow and two subspecies of the Black- throated Sparrow (A. b. deserticola, A. b. opuntia). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 607 Nucleotide divergence between the two subspecies of Black-throated Sparrow (Geno¬ types 5 and 6) was greater than that found among subspecies of the Sage Sparrow, in keeping with their greater geographic separation. Five variable sites were detected (Table 2), all of which were silent sites in the third position of the codon. Four of these changes were transitions and one was a tranversion (Table 3). Position in Codon □ Amphispiza belli §§ A. bilineata FIGURE 2 - Average percent base composition of 1st, 2nd, and 3rd positions in codons for 12 cytochrome b sequences of the Sage Sparrow and two sequences of the Black-throated Sparrow. A total of 96 codons (288 nucleotides) was examined for all 14 sequences. Interspecific mtDNA variation. In contrast to the weak degree of intraspecific diver¬ gence, base sequences of the Sage Sparrow and Black-throated Sparrow showed numerous differences. Sequences of the two species varied at 35 sites (Table 2), 27 of which were silent. Transitions predominated over transversions, with 25 transitions and six-seven transversions occurring between pairs of individuals representing thespecies (Table 3). Of the 35 variable sites, seven (4 transitions, 3 transversions) were in the first position of the codon, one (transition) was in the second position, and 27 (23 transitions, 4 transversions) were in the third position. Finally, we noted the in¬ sertion of a guanine between the 64th and 65th codons in the Black-throated Spar¬ row that was lacking in the Sage Sparrow. Relationship between nucleotide variation and taxonomic level Percentage mtDNA sequence divergence increased with taxonomic level (Table 4). Pairs of individuals or taxa in the genus Amphispiza differed from 0.1% (intrapopulation) to 10.9% (interspecific congeners). Interfamilial comparisons be¬ tween Amphispiza (Emberizidae) and Corcorax (Grallinidae) or Pomatostomas (Muscicapidae) (Kocher et al. 1989) yielded an average percent nucleotide difference of 13.9% to 17.4%, respectively. In interordinal comparisons between Amphispiza (Passeriformes) and Gallus (Galliformes) (Desjardins & Morais 1990), the average percent nucieotide difference was 19.7%. The ratio of transitions to transversions showed a corresponding trend with taxonomic level (Table 4). A bias toward transitions prevailed in all comparisons of sequence divergence within Amphispiza. At the interfamilial level, approximately equal numbers of transitions and transversions accounted for the differences among taxa. Transversions outweighed transitions when comparing sequences between orders. 608 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Percent base composition of sequences of Amphispiza The percent base composition of codons, by position, is presented in Figure 2 for the 12 Sage Sparrow sequences and the two Black-throated Sparrow sequences. The four nucleotides occurred in approximately equal proportions in first position sites in both species. Second position sites were biased toward thymine (39.6%), with no dif¬ ference in the percent base composition of second positions between the Sage Spar¬ row and Black-throated Sparrow. A stronger compositional bias was observed in third position sites, where both guanine (2.1 -4.5%) and thymine (7.3-13.5%) occurred in low percentages relative to adenine (27.8-33.9%) and cytosine (50.5-60.4%). The bias against guanine at third position sites is a general feature of vertebrate mtDNA (Kocher et al. 1989). However, the deficiency of thymine at the same position appears to be an unusual characteristic of avian mtDNA (Kocher et al. 1989). DISCUSSION Nucleotide sequence difference versus allozymic divergence Johnson & Marten (in prep.) surveyed eletrophoretic variation within and among 24 populations of the Sage Sparrows and Black-throated Sparrow, including the 8 populations analysed in the present study. Nei’s (1978) genetic distances within sub¬ species of the Sage Sparrow were 0.001 for Beegum versus Castaic (A. b. belli), 0.005 between Panoche Hills and Chalfant Valley (A. b. canescens ), and 0.000 be¬ tween Rattlesnake Flat and Plush ( A . b. nevadensis). Electrophoretically, the popu¬ lation from Chalfant Valley was essentially identical to populations from both Rattle¬ snake Flat (D = .001) and Plush ( D = .000), indicating a closer affinity to A. b. nevadensis than to A. b. canescens (Johnson & Marten in prep.). The data on nucleotide sequence divergence in cytochrome b support the conclusion from the allozyme analysis that Sage Sparrows from the northern Owens Valley are best placed with A. b. nevadensis. Based on the same subset of populations (excluding Chalfant Valley), average Nei’s D values between subspecies of the Sage Sparrow were 0.0005 for A. b. belli versus A. b. canescens, 0.0105 between A. b. belli and A. b. nevadensis, and 0.0075 between A. b. canescens and A. b. nevadensis. Both the allozyme and mtDNA data clearly indicate that A. b. belli is more closely allied to A. b. canescens than either is to A. b. nevadensis. Quantitative comparisons of electrophoretic variability with mtDNA sequence diver¬ gence offer insight into evolutionary relationships. Thus, we calculated the average pairwise nucleotide difference between populations (= number of transitions + transversions [from Table 3] divided by the number of comparisons in each set of populations) and computed a simple correlation coefficient (r) between each of those values and the corresponding value for Nei’s D. The correlation between the two matrices was impressively high (r = 0.984), suggesting that both techniques provided a robust estimate of the actual amount of genetic difference present in Amphispiza. When the same two matrices were subjected to a Mantel test, t = 3.9908 and P< .001, similarly indicating a highly significant association. Utility of mtDNA base sequence analysis for assessing relationships at different taxonomic levels Despite fundamental changes in the organization of the avian mtDNA genome (Desjardins & Morais 1990), birds, like mammals (Bibb et al. 1981, Wilson et al. 1985, ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 609 Howell 1989), show differences in the inherent variability of regions within and be¬ tween genes in the mtDNA (Kocher et al. 1989). Such variation would profoundly in¬ fluence the relative utility of different portions of the mtDNA for phylogenetic analy¬ sis at various taxonomic levels. Kocher et al. (1989) demonstrated the usefulness of cytochrome b for the study of evolutionary relationships at the species level and above. For intraspecific studies, however, less conserved areas of mtDNA, such as the displacement-loop (control) region (Bibb et al. 1981, Wilson et al. 1985), might offer better information for assessing genetic divergence within and between populations. Nonetheless, the strong concordance between patterns of variation in allozymes and cytochrome b within Amphispiza suggests that cytochrome b se¬ quences, at least in the region chosen here, can be useful for assessing relationships in this group down to and including populations (Wilson et al. 1985). The degree to which cytochrome b sequences might elucidate intraspecific relationships in other avian taxa remains to be studied. ACKNOWLEDGMENTS We are indebted to Margaret F. Smith and Cristian Orrego for indispensable techni¬ cal advice and assistance, to Scott V. Edwards and Francis X. Villablanca for help¬ ful discussion, and to C. Scott Wood for collecting a valuable specimen in Oklahoma. Karen Klitz drafted the final versions of the figures and skillfully prepared the pen and ink sketches that adorn the map. Ellen Prager generously provided a gel dryer when ours failed. LITERATURE CITED AMERICAN ORNITHOLOGISTS’ UNION. 1957. Check-list of North American birds, Fifth edition. Bal¬ timore, The Lord Baltimore Press, Inc. ANDERSON, S., BANKIER, A.T., BARRELL, B.G., DE BRUIJN, M.H.L., COULSON, A.R., DROUIN, J., EPERON I.C., NIERLICH, D.P., ROE, B.A., SANGER, F., SCHREIER, P.H., SMITH, A.J.H., STADEN, R., YOUNG, I.G. 1981. Sequence and organization of the human mitochondrial genome. Nature 290: 457-465. BIBB, M.J., VAN ETTEN, R.A., WRIGHT, C.T., WALBERG, M.W., CLAYTON, D.A. 1981. Sequence and gene organization of mouse mitochondrial DNA. Cell 26: 167-180. CABOT, E.L., BECKENBACH, A.T. 1989. Simultaneous editing of multiple nucleic acid and protein sequences with ESEE. Computer Applications in the Biosciences 5: 233-234. DESJARDINS, P., MORAIS, R. 1990. Sequence and gene organization of the chicken mitochondrial genome. Journal of Molecular Biology 212: 599-634. GRINNELL, J., MILLER, A.H. 1944. The distribution of the birds of California. Pacific Coast Avifauna 27: 1-608. HOWELL, N. 1989. Evolutionary conservation of protein regions in the protonmotive cytochrome b and their possible roles in redox catalysis. Journal of Molecular Evolution 29: 157-169. JOHNSON, N.K., MARTEN, J.A. in prep. Genetic population structure, gene flow, and morphologic divergence in the Sage Sparrow. KOCHER, T.D., THOMAS, W.K., MEYER, A., EDWARDS, S.V., PAABO, S., VILLABLANCA, F.X., WILSON, A.C. 1989. Dynamics of mitochondrial DNA evolution in animals: amplification and sequencing with conserved primers. Proceedings of the National Academy of Sciences, USA 86: 6196- 6200. KOCHER, T.D., WHITE, T.J. 1989. Evolutionary analysis via PCR. Pp. 137147 in Erlich, H.A. (Ed.). PCR technology - principles and applications for DNA amplification. New York, Stockton Press. MANIATIS, T., FRITSCH, E.F., SAMBROOK, J. 1982. Molecular cloning - a laboratory manual. New York, Cold Spring Harbor Laboratory. 610 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI MANTEL, N. 1967. The detection of disease clustering and a general regression approach. Cancer Research 27: 209-220. MARX, J.L. 1988. Multiplying genes by leaps and bounds. Science 240: 1408-1410. NEI, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of in¬ dividuals. Genetics 89: 583-590. SANGER, P., NICKLEN, S.A., COULSEN, A.R. 1977. DNA sequencing with chain-terminating inhibi¬ tors. Proceedings of the National Academy of Sciences, USA 74: 5463-5467. WHITE, T.J., ARNHEIM, N., ERLICH, H.A. 1989. The polymerase chain reaction. Trends in Genetics 5: 185-189. WILSON, A.C., CANN R.L., CARR, S.M., GEORGE, M., GYLLENSTEN, U.B., HELM-BYCHOWSKI, K.M., HIGUCHI, R.G., PALUMBI, S.R., PRAGER, E.M., SAGE, R.D., STONEKING, M. 1985. Mitochondrial DNA and two perspectives on evolutionary genetics. Biological Journal of the Linnean Society 26: 375-400. / ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 611 MICROCOMPLEMENT FIXATION: PRELIMINARY RESULTS FROM THE AUSTRALASIAN AVIFAUNA P. R. BAVERSTOCK1, R. SCHODDE2, L. CHRISTIDIS3, M. KRIEG4 and C. SHEEDY4 1 School of Resource Science and Management, University of New England Northern Rivers, P.O. Box 157, Lismore, NSW 2480, Australia 2CSIRO Division of Wildlife and Ecology, P.O. Box 84, Lyneham, ACT 2602, Australia 3 Ornithology Department, Museum of Victoria, 71 Victoria Crescent, Abbsford, VIC 3067, Australia 4 Evolutionary Biology Unit, South Australian Museum, North Terrace, Adelaide, SA 5000, Australia ABSTRACT. Microcomplement fixation is being used to test conflicting phylogenetic hypotheses rel¬ evant to the evolution of various groups of Australasian birds. The studies are as yet incomplete, but preliminary results suggest that: 1) Albumin immunologic evolution in the Zebra Finch Taeniopygia guttata has been far from clock-like. 2) The Australian passerines are indeed a monophyletic assem¬ blage, as indicated by DNA/DNA hybridisation, but the timing of the origin of this group remains incon¬ clusive. 3) Albumin evolution in the parrots (Psittaciformes) has either been relatively slow, or the parrots are not a Gondwanan group. 4) The Australian Honeyeater (Meiphagid) radiation is monophyletic with respect to Melilestes and Myzomela. 5) The Australian chats Ephthianura and the Gibber Bird Ashbyia are sister taxa having close affinity with the Australian meliphagids. 6) The Aus¬ tralian “mudnesters” are not monophyletic, the Mudlark Grallina being a monarch flycatcher, closely allied to Myiagra. 7) The Australian treecreepers Climacteris are a monophyletic group that includes Cormobates leucophea, and that has affinities with the Meliphagidae. Keywords: Albumin, microcomplement fixation, passerines, parrots, finches, treecreepers, chats, mudnesters, honeyeaters. INTRODUCTION Microcomplement fixation is a quantitative immunological procedure used for estimat¬ ing the number of amino acid substitution differences between homologous proteins (Wilson et al. 1977). It has been used extensively to probe evolutionary relationships among taxa, mainly in the higher vertebrates (e.g. Sarich 1969, Maxson et al. 1982, 1988). For some proteins, the procedure has revealed a high level of rate uniformity within many vertebrate groups, and hence has been used as a molecular dating device (e.g. Sarich 1969, Maxson et al. 1982). Moreover, these studies have suggested that, in general, most vertebrates seem to have similar rates of molecular evolution for a given protein (e.g. Maxson et al. 1975). A possible exception is birds. Microcomplement fixation studies using mainly albumin and transferrin have been interpreted as indicating a slower rate of protein evolution in birds (Prager et al. 1974), a view supported by DNA/DNA hybridisation studies (Sibley & Ahlquist 1986). How¬ ever, this view has not received unanimous spport (e.g. Wilson 1988). Over the past several years, a group of us have been using microcomplement fixa¬ tion of albumin and to a lesser extent transferrin to probe evolutionary problems in the Australian avifauna. These studies are as yet incomplete, but this symposium pro¬ vides an opportunity for us to present preliminary data that are relevent not only to the 612 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI origins and evolution of the Australian avifuana, but also to our understanding of pro¬ tein immunologic evolution in birds. Because detailed analyses will be presented elsewhere, we simply present herein representative data that illustrate the approach. METHODS The microcomplement fixation procedure used follows closely that described by Champion et al. (1974) and Maxson & Maxson (1990). Antisera were raised in rab¬ bits (three per antigen) over a period of three months, and the purity of antisera checked by immunoelectrophoresis. All results are expressed as immunological dis¬ tances - ID (Champion et al. 1974). RESULTS AND DISCUSSION Unequal rates of albumin immunologic evolution As part of a study of molecular relationships among finches of the subfamily Estrildinae, we discovered an unusual case of rapid albumin immunologic evolution in the Zebra Finch, Taeniopygia guttata (Baverstock et al. 1990a). A sample of the relevant data is shown in Table 1 . Within the Estrildinae, Neochmia and Taeniopyaia are members of the grassfinch tribe Estrildini, and Lonchura of the tribe Lonchurini, whereas Passer is a member of a related subfamily Passerinae, sparrows. Antiserum to Neochmia temporalis albumin recognises the close similarity of albumin of Double- barred Finch T. bichenovii (ID = 3) and of Lonchura (ID = 2), but the albumin of T. guttata is immunologically quite distinct, with an ID of 38, exceeding that of Passer domesticus (ID = 24). Antiserum to albumin of T. guttata gives a similar reciprocal response, with IDs of 32 to 39 to other estriidines including its congener T. bichenovii. There are two possible explanations for these results: either the taxonomy of T. guttata at the generic, tribal and subfamilial level is in error, or the albumin of T. guttata has undergone a rapid immunological change. Chromosomal and allozymic data (Christidis 1986a,b, 1987a,b) indicate the latter. Indeed, microcomplement fixa¬ tion studies of transferrin, a protein that typically evolves at about twice the rate of al¬ bumin (Prager et al. 1974), yielded the expected result, with T. bichenovii closest to T. guttata (Table 1 ). Whatever the explanation for the dramatic change in the immunological properties of T. guttata albumin (Baverstock et al. 1990), the data clearly indicate that rates of albumin immunologic evolution are not always uniform among lineages of birds. Monophyly of the Australian passerine radiation Based upon DNA/DNA hybridisation studies, Sibley & Ahlquist (1985) concluded that many Australian passerine groups had been incorrectly placed among northern hemi¬ sphere-centred families, and that they in fact represented an endemic radiation within Australia (Parvorder Corvi). They further suggested that this group had a Gondwanan origin which, based on a DNA/DNA molecular clock, separated from the Parvorder Muscicapae (= Passerida) 55 to 60 mya. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 613 TABLE 1 - Immunological distances (IDs) of selected finches to rabbit anti-albumin of Neochmia temporalis{Nt), Taeniopygia guttata ( Tg ) and Lonchura oryzivora ( Lo ) and to rabbit anti-transferrin of T. guttata. Antigen Antibody Anti-albumin Lo Anti-transferrin Nt Tg Pd Tg Neochmia temporalis (Nt) 0 39 5 23 12 Taeniopygia guttata { Tg) 38 0 31 52 0 Lonchura oryzivora {Lo) 2 33 0 26 14 Passer domesticus {Pd) 24 52 22 0 17 Neochmia ruficauda 2 32 - - 9 Taeniopvgia bichenovii 3 34 - - 9 We have tested these conclusions with microcomplement fixation of albumin. Our data (Table 2) are consistent with the hypothesis that the Parvorder Corvi of Sibley and Ahlquist (1985) is a monophyletic assemclage. Moreover the data are, with few ex¬ ceptions, consistent with the branching relationships they proposed among the fami¬ lies. The timing of the separation of the Parvorders Corvi and Passerida by albumin immu¬ nology is more problematic. The relationship T = 0.6 D (where T = time in millions of years and D = albumin ID) has been used extensively for the “albumin clock” (e.g. Wilson et al. 1977), but Sarich (1985) has since suggested that the relationship T = 0.35 D is more appropriate for the placental mammal orders. The relationship between T and D for birds is even more controversial. Initial results (e.g Prager et al. 1974) led to the conclusion that in birds the rate of albumin evolution, and indeed of the total genome (Sibley & Ahlquist 1986), was about half that of other vertebrates, although Wilson (1988) has subsequently questioned the calibration of these clocks. Our data have corroborated the findings of Prager et al. (1974) that albumin IDs be¬ tween the orders of birds are about 40 to 50. If the orders of birds were already present in the late Cretaceous (Olsen 1985, Feduccia 1980), then this is a clear in¬ dication of a slower rate of albumin evolution in birds than in other vertebrates by about one half. Applying a formula of T = 1 .2 D to the passeriform data, we obtain a divergence time between the Parvorders Corvi and Passerida of about 50 mya. Such a timing is compatible with a Gondwanan origin for the Parvorder Corvi, in accordance with the proposal of Sibley & Ahlquist (1985). Applying the more usual formula gives a separation time of about 25 mya, which is too short for a Gondwanan origin. Albumin evolution in the parrots (Psittaciformes) The order Psittaciformes presently occurs in Australia and New Guinea, New Zealand, South America, Africa, India and Southeast Asia, precisely the distribution expected of a Gondwanan group (Smith 1975), notwithstanding a meagre European fossil record in the Tertiary. We therefore undertook an albumin immunologic study of this group in the hope of shedding further light on rates of molecular evolution in birds. A representative set of results is shown in Table 3. 614 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI TABLE 2 - Immunological distances of selected Passeriforms to rabbit anti-albumin of the honeyeater Anthochaera carinata (Ac), the chat Ephthianura tricolor ( Et ), and the treecreeper Climacteris picumnus ( Cp ). Classification follows Sibley and Ahlquist (1985). Ac Et Pc Parvorder Corvi Meliphagidae Anthochaera paradoxa 0 13 16 Anthochaera chrysoptera 0 - - Acanthagenys rufogularis 0 - - Phylidonyris novaehollandiae 6 9 21 Lichenostomus penicillata 4 13 14 Acanthorhynchus tenuirostris 2 - - Certhionyx variegatus 8 - 27 Myzomela sanguinolenta 21 18 38 Melilestes megarhychus 26 16 35 Ephthianuridae Epthianura tricolor 22 0 18 Epthianura albifrons - 3 - Epthianura aurifrons - 2 - Ashbyia love ns is - 2 - Acanthizidae Pardalotus striatus 15 17 18 Acanthiza chrysorrhoa 26 36 32 Maluridae Malurus pulcherrimus 16 20 17 Corvoidea Corcorax melanorhamphos 27 42 15 Grallina cyanoleuca 22 21 24 Menuroidea Menura novaehollandiae 21 - 23 Climacteris picumnus 23 27 0 Climacteris affinis - - 7 Climacteris erythrops - - n Climacteris leucophaea - - 10 Parvorder Passeri Turdidae Turdus merula 39 - 33 Hirundinidae Hirundo neoxena 33 - 31 Passeridae Passer domesticus 41 45 36 Zosteropidae Zosterops lateralis 47 43 34 Alaudidae Anthus novaeseelandiae 55 40 36 The immunological distances are again compatible with a slow rate of albumin evo¬ lution in birds if the parrots are indeed Gondwanan in origin. The phylogenetic picture revealed by these data is as compatible with a vicariant Gondwanan origin as with dispersal from Eurasia. If dispersal were the prime mechanism underlying the distri¬ bution of parrots, we might expect Indian parrots to be more closely related to ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 615 TABLE 3 - Immunological distances of selected parrots (Psittaciformes) to rabbit anti¬ albumin of the Bluebonnet Northiella haematogaster ( Nh ), the Budgerigar Melopsittacus undulatus {Mu), the Indian Ringneck Psittacula krameri ( Pk ), and the Peachface Lovebird Agapornis roseicollis ( Ar ). Antigen Nh Antibody Mu Pk Ar Northiella haematogaster (Aust) 0 20 9 27 Melopsittacus undulatus (Aust) 20 0 15 33 Psittacula krameri (India) 9 15 0 26 Agapornis roseicollis (Africa) 27 33 26 0 Australia Glossopsitta concina 3 14 10 37 Cacatua roseicapilla 7 18 13 28 New Zealand Strigops habroptilus 16 29 17 29 Nestor meridionalis 8 24 16 28 Cyanorhamphus novaezealandiae 9 16 8 25 India Psittacula alexandriae 9 15 0 27 America Deroptyus accipitrinus 4 16 7 27 Ara ararauna 8 20 9 26 Pyrrhura picta 9 - 13 26 Rhynchopsitta pachyrhyncha 5 19 9 26 Africa Psittacus erithacus 14 20 14 27 Poicephalus meyeri 13 29 12 30 Australian or to African parrots than to the parrots of South America and New Zea¬ land. Yet there is no evidence for either in the data. A curious result is the highly divergent albumin of the Budgerigar Melopsittacus undulatus from the other Australian parrots. Outgroup analysis revealed that here, as in the Zebra Finch, the high divergence is due to rapid evolution of the immunologic properties of Budgerigar albumin. Interestingly the albumin of Agapornis roseicollis is also highly divergent from the other African parrots compared. We were unable to determine the basis for this divergence. Relationships within the honeyeaters Meliphagidae Our protein data (Table 2) show that the New Guinean-centred genera Myzomela and Melilestes fall well outside other meliphagids represented, or that their albumins have had unusually fast rates of albumin evolution. DNA/DNA hybridisation data (Sibley & Ahlquist 1985) also suggest that these genera were distantly related to each other and to other meliphagids. Relationships of the Australian chats Ephthianuridae Two genera are typically included in the Ephthianuridae, Ephthianura and Ashbyia. They have usually been considered monophyletic, although Ashbyia has at times 616 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI been associated with the pipits (Motacillidae). The affinities of the family have been considered to lie with the Australian warblers, although aspects of morphology (Parker 1973) and DNA/DNA hybridisation (Sibley & Ahlquist 1985) place it with the Meliphagidae. Our data (Table 2) unequivocally align Ashbyia with Ephthianura, and they are con¬ sistent with monophyletic links with the Meliphagidae. The relationships of the Australian mud-nesters The Australian mud-nesters have traditionally included the Magpielark Grallina cyanoleuca, the Apostle bird Struthidea cinerea, and the White-winged Chough Corcorax melanorhamphos. A fourth species, the Torrent-lark Grallina bruijni, occurs in Papua New Guinea. The three genera have variously been combined in the same family, split into three separate families, or Struthidea and Corcorax have been grouped in one family and Grallina in another. More recently, DNA/DNA hybridisation studies have corroborated the last arrangement, indicating that Grallina is not related to the other two mudnesters but to the monarchine flycatchers, notably Monarcha and Myiaora (Sibley & Ahlquist 1985). Relevant microcomplement fixation data (Table 4) are unequivocal with regard to Grallina, clearly aligning it with the monarchine, Myiagra. TABLE 4 - Immunological distances of selected Corvoidea to rabbit anti-albumin of the Mudlark Grallina cyanoleuca. Grallinidae Grallina cyanoleuca 0 Grallina bruijni 0 Corcoracidae Corcorax melanorhamphus 29 Struthidea cinerea 20 Monarchidae Myiagra (average 4 spp.) 2 Monarcha (average 6 spp.) 8 Rhipidura (average 5 spp.) 10 Corvidae Corvus bennetti 22 Cracticidae Gymnorhina tibicen 24 The relationships of the Australian Treecreepers, family Climacteridae The Australian treecreepers have been variously aligned with holarctic creepers Certhiidae (Gadow 1883), with the honeyeaters Meliphagidae (Harrison 1969, Parker 1982), as an Australo-Papuan group with no close affinities (Schodde 1975, Orenstein 1977, Ames 1987), or distantly related to the lyrebirds Menura and bowerbirds Ptilonorhynchidae (Sibley & Ahlquist 1985). Moreover, Parker (1982) suggested that the C. leucophaea superspecies was sufficiently distinct from other climacterids as to be paraphyletic with an independent meliphagid ancestry. Albumin data (Table 2) are consistent with Climacteris sensu lato being a monophyletic unit, in which leucophaea nevertheless lies outside the other three spe¬ cies. There is no evidence for an especially close relationship of Climacteris to any ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 617 other group; the closest group appears to be the Meliphagidae, but the Menuridae are well distant. The future Much of the speculation presented herein is based upon one-way immunological re¬ actions and therefore must be treated with caution, especially in view of the indica¬ tions of grossly unequal rates of albumin evolution in the Zebra Finch. Nevertheless one-way reactions are useful for distinguishing markedly different phylogenetic hy¬ potheses, and for highlighting taxonomic problems warranting additional investigation. There is no doubt that recent developments in molecular technology are bringing nucleic acid sequencing into the realm of phylogenetic hypothesis testing as a prac¬ tical method. It is apparent that sequence data are far superior to immunological tech¬ niques such as microcomplement fixation for phylogenetic analysis. Nevertheless, information from microcomplement fixation data has given a wealth of background information on phylogenetic relationships among higher vertebrates, and provided the basis for considerable debate regarding rates of molecular evolution, times of phylogenetic divergence, and modes and tempos of morphological, behav¬ ioural and chromosomal evolution. ACKNOWLEDGEMENTS We are grateful to the many people who contributed blood samples for these studies, especially Jack Bourne and Ron Brown. We thank P. Altman and B. Young for care of rabbits. Robert Zink provided valuable criticism of the manuscript. LITERATURE CITED AMES, P.L. 1987. The unusual syrinx morphology of Australian treecreepers Climacteris. Emu 87: 192-195. BAVERSTOCK, P.R., CHRISTIDIS, L., KRIEG, M., BIRRELL, J. 1990. Albumin evolution in the estrildine finches (Aves: Passeriformes) is far from clock-like. Australian Journal of Zoology (in press). CHAMPION, A.B., PRAGER, E.M., WATCHER, D., WILSON, A.C. 1974. Microcomplement fixation. Pp. 337-416 in Wright, C.A. (Ed.). Biochemical and immunological taxonomy of animals. London, Academic Press. CHRISTIDIS, L. 1986a. Chromosomal evolution within the Estrildidae (Aves). 1. The Poephilae. Genetica 71 : 99-1 1 3. CHRISTIDIS, L. 1986b. Chromosomal evolution within the Estrildidae (Aves). 11. The Lonchurae. Genetica 71 : 1 14-128. CHRISTIDIS, L. 1987a. Biochemical systematics within palaeotropic finches (Aves:Estrildidae). Auk 104: 380-392. CHRISTIDIS, L. 1987b. Chromosomal evolution within the Estrildidae (Aves). 111. The Estrildidae. Genetica 72: 92-100. FEDUCCIA, A. 1980. The Age of Birds. Harvard Univ. Press, Cambridge. GADDOW, H. 1883. Certhiomorphae. Catalogue of birds in the British Museum, 8. British Museum, London. HARRISON, C.J.O. 1969. The possible affinities of the Australian treecreepers of the genus Climacteris. Emu 69:161-168. MAXSON, L.R., SARICH, V.M., WILSON, A.C. 1975. Marsupials, frogs and continental drift. Nature (London) 255: 397-400. 618 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI MAXSON, L.R., TYLER, M.J., MAXSON, R.D. 1982. Phylogenetic relationships of Cyclorana and the Litoria aurea species-group (Anura: Hylidae): a molecular perspective. Australian Journal of Zoology 30:643-651. MAXSON, L.M., MOLER, P.E., MANSELL, B.W. 1988. Albumin evolution in salamanders of the genus Necturus. Journal of Herpetology 22: 231-235. MAXSON, L.M., MAXSON, R.D. 1990. Immunological methods in Hillis, D., Moritz, C. (Eds). Molecu¬ lar systematics. Sinauer, Bonn. OLSON, S.L. 1985. The fossil record of birds. Pp. 79-238 in Farner, D.S., King, J.R., Parkes, K.C. (Eds). Avian Biology. Academic Press, New York and London. ORENSTEIN, R.l. 1977. Morphological adaptation for bark foraging in the Australian treecreepers (Aves: Climacteridae). Ann Arbor, Michigan: University Microfilms International. PARKER, S.A. 1973. The tongues of Epthianura and Ashbyia. Emu 73: 19-20. PARKER, S.A. 1982. The relationships of the Australo-Papuan tree-creepers and sitellas. South Aus¬ tralian Ornithologist 28: 197-200. PRAGER, E.M., BRUSH, A.H., NOLAN, R.A., NAKANISHI, M., WILSON, A.C. 1974. Slow evolution of transferrin and albumin in birds according to micro-complement fixation analysis. Journal of Molecu¬ lar Evolution 3: 243-263. SARICH, V.M. 1969. Pinniped origins and the rate of evolution of carnivore albumins. Systematic Zo¬ ology 18: 286-295. SARICH, V.M. 1985. Rodent macromolecular systematics. Pp. 423-452 in Luckett, W.P. , Hartenberger, J.L. (Eds). NATO adv. Stud. Inst. Series A; Life Sciences, Vol. 92. SCHODDE, R. 1975. Interim list of Australian songbirds. Melbourne, RAOU. SIBLEY, C.G., AHLQUIST, J.E. 1985. The phytogeny and classification of the Australo-Papuan birds. Emu 85:1-14. SIBLEY, C.G., AHLQUIST, J.E. 1986. Reconstructing bird phylogeny by comparing DNA’s. Scientific American 254: 82-92. SMITH, G.A. 1975. Systematics of parrots. Ibis 117: 18-66. WILSON, A.C. 1988. Time scale for bird evolution. Proceedings of the International Ornithological Congress 19: 1912-1917. WILSON, A.C., CARLSON, S.S., WHITE, T.J. 1977. Biochemical evolution. Annual Review of Biochem¬ istry 46: 573-639. / ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 619 AVIAN SYSTEMATICS BY SEQUENCE ANALYSIS OF mtDNA PETER ARCTANDER Institute of Population Biology, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen 0, Denmark ABSTRACT. Sequence data are qualitatively different from other DNA data in giving precise knowledge of the nature of genetic differences. This facilitates a functional analysis and differential weighting of characters based on empirical knowledge of the mode of evolution of the particular gene and of more general molecular constraints. The main disadvantage is that for practical reasons only minute parts of a genome can be sequenced. The power of resolution and the limitations for DNA sequence analysis are discussed and exemplified by 42 sequences of 287 base pairs (bp) in the cytochrome-b (cyt-b) gene of mitochondrial DNA (mtDNA) from 30 different bird species covering a wide taxonomic range. Resolving power is good at the species level, roughly within 1 - 30 million years since a common an¬ cestor (mya). However, the 287 bp are insufficient when analyzing more closely related forms, e.g. intra-specific relations, as well as more distant relationships. To resolve relationships at these dis¬ tances, DNA fragments more variable than cyt-b and/or longer stretches of DNA have to be sequenced. Because of very slow rates of amino-acid replacing substitutions, phylogenetic information is retained even over 600 mya. Keywords: Avian systematics, mitochondrial DNA, DNA sequence analysis, polymerase chain reac¬ tion, sequence evolution. INTRODUCTION Comparative morphology and anatomy have been the main tools for reconstructing avian evolution. Difficulties arise for this approach when dealing with very close rela¬ tives because of few characters, and on all systematic levels because of the difficul¬ ties in discovering convergence and parallelism. Overcoming these hurdles is the pri¬ mary reason for the interest in molecular data and especially DNA, which promises to provide informative, comparable and ‘objective’ data. There are currently three, quite different, DNA techniques in use: DNA/DNA hybridi¬ zation, which aims to compare whole genomes; restriction fragment length polymorphism analysis (RFLP), scoring part of the substitutions in a small fragment of the DNA; and sequencing analysis, revealing the actual genetic code of a DNA fragment. In the following I will concentrate on DNA sequence analysis (for survey see Arctander & Fjeldsa in press, Sibley & Ahlquist 1986, Quinn & White 1987). Technical achievements in biotechnology, such as the development of the polymerase chain reaction (PCR), have made it feasible to sequence homologous DNA segments from a wide variety of organisms. Sequence data differ qualitatively from other DNA data, giving precise knowledge of the nature of genetic differences. This facilitates a functional analysis and differential weighting of characters based on empirical knowl¬ edge of the mode of evolution of the particular gene and of the existence of molecu¬ lar constraints. The DNA sequence data illustrate principles of molecular evolution and, therefore, clarify the relationship between phenotypic and molecular evolution. Sequence data are superior in these ways and, therefore, highly desirable. The main 620 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI disadvantage is that for practical reasons only minute fragments of the genome can be sequenced. The present paper addresses the question: Can a few hundred base pairs provide reliable historical information? And if so, what is the power of resolution and what are the limitations of this approach? The data analyzed are all from mitochondrial DNA (mtDNA), but except for one case (Laniarius) nuclear DNA could as well have been analyzed. A more detailed presentation and discussion of the data will appear else¬ where. THE POWER OF RESOLUTION AT DIFFERENT SYSTEMATIC LEVELS Resolution at the species level Clear demarcation of species can be difficult, especially when dealing with allopatric populations. The biological species definition is operatively unhandy (McKitrick & Zink 1988, Haffer 1986). Species limits are often ambiguous because traditional methods provide few characters, and because analysis can be hampered by subjective char¬ acterization; concealed parallelism and convergence, especially when most charac¬ ters are in terms of color hues, body size, and so on, also confound analysis. The following two examples illustrate the potential of DNA sequence data at this level. A new species of Laniarius? Last year in Bulo Burti, Somalia, a Boubou Laniarius was captured (Smith et al. in press). From morphology, it was judged to be a new species but in spite of months of observations only this individual was recorded. This naturally raises the question: Could it be a rare result of a hybridization or a color freak? These questions can be answered by analysis of mtDNA. MtDNA is maternally inherited so any hybrid or morph will have the mtDNA from the mother species. Comparing mtDNA from this individual with species that could possibly have provided the mother should resolve the question. Another advantage of DNA analysis is that small amounts of tissue (blood or feather pulp; Arctander 1988, Arctander & Fjeldsa in press) are needed for analysis, and collection of this rare bird was unnecessary. A 295 base pair (bp) fragment of the mtDNA cytochrome b (cyt-b) gene was sequenced from 6 different Laniarius species and one Telophorus (outgroup). This data set contains 65 variable sites of which 44 are phylogenetically informative. DNA sequence cannot determine species status in its strict biological sense. How¬ ever, in this case we concluded that the mother of the Bulo Burti Boubou did not come from any of the examined species, ruling out hybridization and color morphism. The number of substitutions in pairwise comparisons including this taxon (average transversions 4.5, transitions 22.4 [transversion: purine to pyrimidine, or vice versa; transition: purine to purine or pyrimidine to pyrimidine]) corresponds well with the amount of variation observed within the other examined Laniarius, which are all con¬ sidered good species (mean transversions 4.3, transitions 23). Only three substitu¬ tions (two transitions and one transversion, all silent) were found between the two L. luhderi populations 400 km apart from Sudan and from Kenya, respectively. Only one replacement substitution was observed in the whole data set. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 621 TABLE 1 - Species analysed, the source of DNA and status of publication. Refers to the number of sequences from different individuals of the particular species. DNA was obtained from several different sources, blood samples collected and preserved ac¬ cording to Arctander (1988), blood quills from alcohol-preserved feathers (Smith et al. in press), muscle tissue first frozen, later alcohol-preserved (e.g. Edwards et al. in press) and from small fragments of old museum skins (e.g. Smith et al. in press, Paabo 1989). DNA was extracted following standard methods (Arctander 1988, Smith et al. in press). A 287 bp fragment of the mtDNA cyt-b gene spanning 95 amino ac¬ ids was analyzed for all species in this study. They cover the positions 14842 to 15128 (enumerated according to the corresponding human sequence, Anderson et al. 1981). The Gallus, Bos, Xenopus and Paracentrotus sequences are taken from the literature. Prior to sequencing of all the other species the DNA was amplified via the polymerase chain reaction (PCR) (Erlich 1989, Saiki et al. 1988). Double stranded amplifications were followed by single stranded ones, whereby sufficient copies of the desired DNA fragment were obtained for dideoxy termination sequencing (Sanger 1977, Gyllensten & Erlich 1988). The Primers HL14841 and HH15149 (Kocher et al. 1989) were used for the amplifications, which were performed in 50 pi reactions running 30-35 cycles, and with a 1:100 dilution of one primer in the single-stranded reaction. For the phylogenetic analysis the PHYLIP 3.3 programs DNAPARS, DNABOOT and DNAML (Felsenstein 1990) and CMP (Siegismund, unpublished) were used. Samples No. DNA source Publication 1 Laniarius, unnamed 1 feather Smith et al. in press 2 L. aethiopicus 1 skin 3 L.turatii 1 blood 4-6 L. luhderi 3 skin 7 L. ruficeps 1 skin 8 L. barbarus 1 blood 9 Pomatostomus temporalis 1 tissue Edwards et al. in press 10 P. ruficeps 1 tissue 11 P. isidori 1 tissue 12-17 Scytalopus magellanicus ssp. 6 blood/tissue unpublished 18-20 S. unicolor 3 blood 21-22 S. femoralis 2 blood 23 S. latebricola 1 blood 24-26 S. unnamed 3 blood/tissue 27 Melanoparaia maximiliani 1 blood 28 Grallaria andicola 1 blood 29 Ampelion stresemanni 1 blood Edwards et al. in press 30 A. rubrocristatus 1 blood unpublished 31 Phytotom a rutila 1 blood 32 Pireola arcuata 1 blood 33 Pipra coronata 1 skin 34 My i arcus tubercu lifer 1 blood 35 Elaenia pallatangae 1 blood 36 Pachyramphus validus 1 skin 37 Mionectes striaticollis 1 blood 38 Asthenes dorbignyi 1 blood 39 Pitta sordida 1 blood Edwards et al. in press 40 Catharus guttatus 1 blood 41 Ale a torda 1 blood unpublished 42 Gallus gallus 1 Desjardins & Morais 1990 43 Bos taurus 1 Anderson et al. 1 982 44 Xenopus laevis 1 Roe et al. 1 985 45 Paracentrotus lividus 1 Cantatore et al. 1 989 622 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI In the light of the DNA analysis the Bulo Burti Boubou is as genetically distinct as any of the other examined Laniarius species. Apart from answering our primary question the analysis places the new bird in the phytogeny of the Laniarius. The topology of the resulting phylogeny was in good agreement with conclusions based on morphology. Scytalopus,. morphologically indistinguishable. Tapaculos of the genus Scytalopus are small, short-tailed, blackish sedentary suboscines widespread in the Andes and in southeastern Brazil. The currently accepted taxa are extremely difficult to distinguish morphologically. Observation that the songs and male responses to play-back were strongly differentiated (N. Krabbe pers. com., Fjeldsa & Krabbe 1990) revealed that the number of biological species probably has been underestimated. Comparisons based on the 287 bp’s covering the same gene fragment of mtDNA cyt- b as used for Laniarius show approximately the same numbers of transitions and transversions as for Laniarius (mean transitions 26, transversions 4. 4, see Table 2). Sequencing several individuals originating from the same geographic region in all cases except one revealed no differences (the one individual was out of a group of 10 and had two silent 3rd position transitions). Once again, the sequence data do not automatically define these taxa as good biological species, but show that many allopatric forms that scarcely differ morphologically, but do so vocally, are clearly dis¬ tinguishable when analyzing DNA, and are as distinct as morphologically very sepa¬ rate Laniarius species, for example. These preliminary results indicate that several small isolated populations should be ranked as species and that a complete revision of the group probably is necessary. These Laniarius and Scytalopus sequences support taxonomic conclusions at the within-genus level. TABLE 2 - Transitions, transversions and amino acid replacements at different systematic levels. cyt-b 287 bp Transitions Transversions Replacements Range Mean % Range Mean % Range Mean % / N Bush shrikes 15-28 22 7.7 2-7 4.1 1.4 0-1 0.1 0.1 6 Austr. babblers1 23-33 29 10.1 1-7 4.6 1.6 0-7 4.7 4.9 3 Tapaculos 8-40 26 9.1 1-11 4.4 1.5 0-2 0.2 0.2 15 Cotingas 37 37 12.9 12 12.0 4.2 5 5 5.3 2 Intra specific 0-3 0.3 0.0 0-1 0.1 0.0 0 0 0 1 92 Intra genus 8-40 23.2 8.1 1-12 4.5 1.6 0-7 0.3 0.4 263 Inter family 22-44 29.6 10.3 14-33 27.9 9.7 2-9 5.5 5.8 84 Higher taxa 19-29 24.6 8.6 11-28 22.0 7.7 4-9 6.7 7.1 55 To Cow 19-33 27.5 9.6 42-58 50.0 17.4 20-23 21.2 22.6 I6 1 The relatively high number of substitutions are almost entirely due to Pomatostomus isidori. 2 Substitutions were only detected between two of the 19 individuals. 3 Representing four genera with 2, 3, 6 and 15 species respectively; see first part of the table. 4 The following sub-families and families are compared: Cotinginae, Piprinae, Tityrinae, Tyranninae, Mionectidae, Furanidae, Rhinochryptidae and Formicaridae. 5 Single species from the following higher categories: Tyrannides, Eurylamides, Passeres, Picae and Charidriformes. 6 One Cow compared to the species listed in note 5 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 623 Resolution at the family level Phytotoma - own family? Phytotoma, the plantcutters, were previously placed in a monotypic family because of their conspicuously serrated mandibular edges and syringeal structure (e.g. Warter 1965 [cited in Lanyon & Lanyon 1989], Ames 1971). Sibley and Ahlquist (1985) placed Phytotoma within the Cotinginae but did not include them in their DNA analysis, whereas Lanyon and Lanyon (1989) retain the family Phytotomidae. Using either the principle of maximum parsimony or maximum likelihood (Felsenstein 1990), a genealogical tree based on 287 bp of cyt-b from the mtDNA of 12 different Tyrannides (and a Pitta included as outgroup; Table 1:12, 28-39) clearly places Phytotomas within the Cotinga family, near the genus Ampetion. The data show a varying degree of resolving power concerning the other groups of the Tyranni. The Funarii represented by Asthenes, Scytalopus and Grallaria is very well supported as a monophyletic group. Other deeper nodes are not nearly as well resolved, indicating insufficient phylogenetic information probably due to back-muta¬ tions in positions that are relatively free to vary. The basic radiation of these groups might have happened over a very short period of time as suggested by Sibley and Ahlquist’s data (1988), which would make the resolution of these nodes difficult. Resolution at the level of the basal radiation of an order Oscine-suboscine split; monophyly of Australian oscines? In collaboration with S. V. Edwards and A. C. Wilson we investigated whether phylogenetic information is re¬ tained over even longer evolutionary distances. The Oscine - Suboscine dichotomy and the Sibley and Ahlquist proposed division of Passeriform birds into Passerida and Corvida (Sibley & Ahlquist 1988) were examined using 924 bp of cyt-b from 13 differ¬ ent Oscines and Suboscines and with a Picidae as outgroup for phylogenetic analy¬ sis. By raising the number of bp’s compared to 924 even nodes as deep as the Oscine/Suboscine split can be well resolved. The resulting phylogeny has almost the same topology as that obtained by Sibley and Ahlquist using DNA/DNA hybridization, but the sequence data strongly unite Australian babblers (Pomatostomidae) and thrushes (Turdinae) and these form a sister group with Sylvioidea (Parus)\ these re¬ sults contradict the monophyly of Australian oscines, Corvida, as proposed by Sibley and Ahlquist. SEQUENCE VARIATION OVER TIME An important reason for choosing the cyt-b gene is the apparent functional invariance of this gene mirrored in the conservative amino acid composition and pattern of vari¬ ation in all studied organisms (Edwards et al. in press, Irving et al. in press, Howel 1989). The pattern of variation supports the idea that the selective pressure on this gene will not vary much between different taxa especially when closely related forms are compared. This is important because comparative analysis could be seriously flawed because of different rates of substitutions in different lineages. In the oscine/ suboscine study, the suboscine branch reveals a significantly higher number of T's at third position in the reading frame than the oscine branch. This probably has nothing to do with phaenotypic selection, as practically all third position substitutions are 624 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Sequence variation over time 28 7 base pairs cyt-b mtDNA FIGURE 1 - Patterns of nucleotide substitutions spanning 600 million years for 287 base pair mtDNA cyt-b sequence for each position in the codon triplet and specified for transi¬ tions and transversions, respectively. Based on comparisons of: two Scytalopus (1 million years since common ancestor (mya)), their closest outgroup Melanoparaia (5 mya), Ampelion (30 mya), Catharus (50 mya), Gallus (100 mya), Bos (300 mya), Xenopus (400 mya), Paracentrotus (600 mya)(for references see Table 1). silent, but is rather an example of how molecular evolutionary mechanisms can influ¬ ence the pattern of substitution. Despite its conservatism, cytochrome b of the mtDNA contains information facilitat¬ ing analysis of phylogenetic relationships from the species to the ordinal level in birds. However, informative characters are few between closely related forms (e.g. only three substitutions separate Laniarius luhderi from Kenya and Sudan) so either very large amounts of sequence or a less evolutionary conservative region must be sequenced to study population structures, for example. This latter could for instance be the so called d-loop region of the mtDNA, which has been shown to be highly vari¬ able (Brown 1985). The basic pattern of variation according to codon positions is as follows: 1st position: all substitutions result in amino acid replacements except C/T when the amino acid is leucine; 2nd position: all substitutions result in replacements; 3rd position: most substitutions do not result in replacements; those that do are transversions. As the amino acid sequence is very conservative few replacements occur and most variation is observed at 3rd position and for leucine also at 1st position. Over time, the variable ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 625 positions are likely to be substituted again and again (‘multiple hits’) destroying phylogenetic information by causing ‘saturation’. All substitutions are not equally likely, and in avian mtDNA transitions seem to be about 20 times as likely as transversions at least at 3rd position (own unpublished results; S. Edwards pers. com.). Therefore, transversions will be phylogenetically informative for a much longer period of time. Sequence variation over time 287 base pairs cyt-b mtDNA ^ FIGURE 2 - The first 1/20 of the graph in Figure 1 in order to resolve the first 30 million years. These relationships are illustrated in Figures 1, 2 and 3; “million years since common ancestor” refers to rough estimates based on current systematic/paleontologic views. Replacements, all 2nd position changes and transversions at 1st position, are linear functions, and the remaining curves seem to consist of three parts. At first a steep linear function occurs, exhibiting “free” evolution with no back mutations; then follows a more or less protracted curve indicating the ‘multiple hit zone’; and, finally, a slowly rising linear function. The slight amplitude can be caused by slowly accumulating amino acid substitutions and evolved differences in molecular evolutionary mecha¬ nisms, because saturation phenomena alone would result in a horizontal curve. The above mentioned higher number of T’s at third position in the suboscines could serve as an example of this. Third position substitutions especially exhibit an ‘overshoot’ of transitions in the beginning of the ‘multiple hit zone’ within the first 30 million years. This is due to the differences in substitution rates between transitions and transversions, where the delay in transversion substitutions postpone their inhibitory influence on transitions. 626 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI For phylogenetic analysis, these curves reveal where phylogenetic information starts to fade out. This happens extremely fast for transitions at 3rd and 1st positions (the silent leucine substitution is a transition). Transversions at 3rd position do also run into the ‘multiple hit zone’ quickly, but significantly more slowly than transitions. All transversions at 1st and 2nd positions result in replacements as do all transitions at 2nd position. At these latter positions mtDNA does not appear saturated even after 600 million years, and phylogenetic information is apparently retained. This is also the case for amino acid replacements (Figure 3). Table 2 depicts this span of variation in relation to the systematic ranks of the different bird species. TABLE 3 - Amino acid comparison of chicken mitochondrial protein genes with mam¬ malian and Xenopus laevis homologous genes (%). * For each gene, mean values are quoted for the comparison of the chicken sequence with that from all three following mammals: mouse, cow and man. Redrawn from Desjardins and Morais (1989). Gene Chicken/mammals* Chicken /Xenopus ND1 70 70 ND2 45 57 COI 86 86 COM 67 70 ATPase 8 27 47 ATPase 6 55 68 com 75 80 ND3 57 67 ND4L 47 41 ND4 59 61 ND5 56 62 ND6 27 42 Cyt. b 74 75 Phylogenetic analysis of distantly related taxa must therefore rely mostly on replace¬ ment substitutions and transversions. The numbers of these, especially replacement substitutions, are small for cyt-b, which necessitates long stretches of sequence. Table 3 shows the relative degree of conservation for different vertebrate mitochondrial genes. Cyt-b is one of the most conservative, whereas NADH- dehydrogenase subunit 6, (ND6), is more than twice as variable and might therefore be a good choice. Although the number and rate of substitutions at 3rd position are expected to be the same as for cyt-b 2, its range of application may be larger. There are currently significant differences in the ways we can analyze protein-coding mtDNA genes such as cyt-b and non-protein coding as the d-loop and ribosomal genes. There are fairly good hypotheses of functional domains in the d-loop sequence and of stem and loop structures of the ribosomal gene products. Patterns of conser¬ vation seem to follow this with higher degree of conservation in stems, for example (Gadaleta et al. 1989). With the growing knowledge of the pattern of variation within these parts of the mtDNA, more information can be obtained. However a problem in analyzing sequences like this is that alignment at greater evolutionary distances (e.g. between the Tyranniid families mentioned above, own unpublished results) can be ambiguous because of deletion/insertion events. ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 627 Sequence variation over time Amino acid replacements, cyt-b mtDNA Mill, years since common ancestor FIGURE 3 - Amino acid replacements over time. Based on the same data set as in Figure 1. Inserted is an enlargement of the first 30 million years. ACKNOWLEDGEMENTS Jon Fjeldsa is thanked for his always helpful criticism and Robert Zink for numerous linguistic corrections. This study has been supported by The Carlsberg Foundation and The Danish National Research Council. LITERATURE CITED AMES, P.L. 1971. The morpholoqy of syrinx in passerine birds. Bulletin of Peabody Museum of Natural History 37:1-194. ANDERSON, S., BANKIER, A.T., BARREL, B.G., DE BRUIJN, M.H.L., COULSON, A.R., DROUIN, J., EPERON, I.C., NIERLICH, D.P., ROE, B.A., SANGER, F., SCHREIER, P.H., SMITH, A.J.H., STADEN, R., YOUNG, I.G. 1982. Sequence and organization of the human mitochondrial genome. Nature 290:457-465. ARCTANDER, P. 1988. Comparative studies of avian DNA by restriction fragment length polymorphism analysis: convenient procedures based on blood samples from live birds. Journal fur Ornithologie 129:205-216. ARCTANDER, P., FJELDSA, J. in press. DNA studies for avian systematics - Techniques, virtues and perspectives of DNA collections. In Edelstam, C., Mina, M. (Eds), Museum research in vertebrate zo¬ ology. Swedish Museum of Natural History, Stockholm. BROWN, W.M. 1985. The mitochondrial genome of animals. Pp. 62-88 in MacIntyre, R. (Ed.). Molecu¬ lar evolutionary genetics. New York, Plenum Press. 628 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI CANTATORE, P., ROBERTI, M., RAINALDI, G., GADALETA, M.N., SACCONE, C. 1989. The complete nucleotide sequence, gene organization, and genetic code of the mitochondrial genome of Paracentrotus lividus. Journal of Biological Chemistry 264:10965-10975. DESJARDINS, P., MORAIS, R. 1990. Sequence and gene organization of the Chicken mitochondrial genome, a novel geneorder in higher vertebrates. Journal of Molecular Biology 212:599-634. EDWARDS, S.W., ARCTANDER, P., WILSON, A.C. in press. Mitochondrial resolution of a deep branch in the genealogical tree for perching birds. Proceedings of the Royal Society London, series B. FELSENSTEIN, J. 1990. Phylip manual. Version 3.3. University Herbarium, Berkeley, California. FJELDSA, J., KRABBE, N. 1990. Birds of the high Andes. Zoological Museum, University of Copen¬ hagen. Apollo Books. GADALETA, G., PEPE, G., DE CANDIA, G., QUAGLIORIELLO, C., SBISA, E., SACCONE, C. 1989. The complete nucleotide sequence of the Rattus norvegicus mitochondrial genome: Cryptical signals revealed by comparative analysis between vertebrates. Journal of Molecular Evolution 28:497-516. HAFFER, J. 1986. Superspecies and species limits in vertebrates. Zeitschrift fur zoologische Systematik und Evolutionsforschung 24:169-190. HOWEL, N. 1989. Evolutionary conservation of protein regions in the proton-motive cytochrome b and their possible roles in redox catalysis. Journal of Molecular Biology 29:169-190. IRWIN, D.M., KOCHER, T.D., WILSON, A.C. in press. Evolution of the cytochrome b gene in mammals. Journal of Molecular Evolution. LANYON, S.M, LANYON, W.E. 1989. The systematic posibon of the plantcutters, Phytotoma. Auk 106: 422-432. MCKITRICK, M.C., ZINK, R.M. 1988. Species concepts in ornithology. Condor 90:1-14. QUINN, T., WHITE, B.N. 1987. Analysis of DNA sequence variation. Pp.103-198 in Cooke, F., Buckley, P.A. (Eds). Avian Genetics. London, Academic Press. ROE, B.A., MA, D.-P., WILSON, R.K., WONG, J.F.-H. 1985. The complete nucleotide sequence of the Xenopus laevis mitochondrial genome. Journal of Biological Chemistry 260:9759-9774. SIBLEY, C.G., AHLQUIST, J.E. 1985. Phylogeny and classification of New World suboscine passer¬ ine birds (Passeriformes: Oligomyodi: Tyrannides). In Buckley, P.A. et al. (Eds). Neotropical ornithol¬ ogy. Ornithological Monographs No. 36, Washington D.C., American Ornithologists’ Union. SIBLEY, C.G., AHLQUIST, J.E. 1986. Reconstructing bird phylogeny by comparing DNAs. Scientific American 254:82-92. SIBLEY, C.G., AHLQUIST, J.E., MONROE, B.L., JR. 1988. A classification of the living birds of the world based on DNA-DNA hybridization studies. Auk 105:409-423. SMITH, E.F.G., ARCTANDER, P., FJELDSA, J., AMIR, O.G. in press. A new species of shrike (Laniidae: Laniarius) from Somalia verified by DNA sequence data from the only known individual. Ibis. WARTER S.L. 1965. The cranial osteology of the new world Tyrannoidae and its taxonomic implica¬ tions. Ph.D. dissertation, Baton Rouge, Lousianna State Univ. TWIGS AND BRANCHES ON THE AVIAN TREE, AS REVEALED BY DIRECT SEQUENCING OF MTDNA VIA THE POLYMERASE CHAIN REACTION S.V. EDWARDS, T.W. QUINN and A.C. WILSON Division of Biochemistry and Molecular Biology, University of California, Berkeley, California 94720, USA ABSTRACT. The polymerase chain reaction (PCR) and direct sequencing are rapidly becoming the methods of choice for obtaining mitochondrial DNA sequences. Using several pairs of versatile spe¬ cific primers to amplify sequences from the cytochrome b gene, we have established that in birds, as in other vertebrates, there is a fast accumulation of silent transitions and a slower accumulation of silent transversions and amino acid replacements. This range of rates has allowed us to address phylogenetic questions at several temporal levels. We shall present trees which relate (1) species within the babbler genus Pomatostomus, (2) these Australian babblers to Palearctic thrushes rather than to other Australian "Corvida" and (3) flamingoes to Ciconiiformes or Charadriiformes rather than to Anseriformes. The merits of this approach will be compared to that of bulk hybridization of the “sin¬ gle-copy” fraction of nuclear DNA, with emphasis on the notion of statistical testing. ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 629 CONCLUDING REMARKS: MODERN BIOCHEMICAL APPROACHES TO AVIAN SYSTEMATICS ROBERT M. ZINK Museum of Natural Science, Louisiana State University, Baton Rouge, Louisiana 70803, USA ABSTRACT. This symposium showcased modern molecular approaches to avian systematics. Gelter and Tegelstrom illustrated the use of multiple techniques in understanding the evolution of closely related species. One of the most exciting methodological developments in molecular systematics is the polymerase chain reaction (PCR), which allows easy and rapid amplification of specific sequences of DNA, which can then be easily sequenced. Johnson and Cicero, Arctander, and Edwards, Quinn and Wilson used PCR and sequencing to study mitochondrial DNA (mtDNA), which allowed them to inves¬ tigate a variety of questions regarding geographic variation, speciation, phytogeny and the evolution of mtDNA itself. Baverstock and colleagues illustrated the use of immunological methods for inferring systematic relationships. Because the field of molecular evolution is large and rapidly expanding, these concluding remarks will deal not only with the contributions to this symposium, but with aspects not emphasised by the participants. Keywords: molecular methods, mitochondrial DNA, phylogeny inference, geographic variation. INTRODUCTION Molecular techniques have been widely heralded in systematics because they expose genetic variation directly. Advantages of molecular methods include: the apparent uniform rate of evolution of particular genes within lineages, the existence of explicit genetic models of character evolution, the apparent selective neutrality of many mo¬ lecular traits, the likelihood of genetic independence of characters, and the opportu¬ nity to evaluate homologous regions of DNA across broad taxonomic groups. Because we think of evolution ultimately in genetic terms, molecular methods are of obvious value. For instance, geographic variation has evolutionary significance if geographic patterns are genetically influenced. Speciation requires a conversion of genetic vari¬ ation from within to between taxa, and a phylogeny is a genetic trace of the history of such conversion events. Thus, molecular methods provide information critical to addressing these fundamental issues. CRITIQUE OF MOLECULAR METHODS IN AVIAN SYSTEMATICS Hillis and Moritz (1990) summarized the value of various molecular methods for ques¬ tions at different tiers in the taxonomic hierarchy. I have provided a similar set of opin¬ ions for avian systematics (Table 1). Issues considered include cost and time relative to the level and accuracy of resolution provided. Below I discuss the rationale behind my opinions. Protein electrophoresis I see little value in continuing protein electrophoretic studies of geographic variation in temperate-breeding birds, unless pilot studies indicate otherwise. Most studies 630 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI discovered little or no genetic differentiation despite often considerable morphologi¬ cal variation (Barrowclough 1983, Barrowclough & Johnson 1988). Either morphologi¬ cal differences are environmentally induced or the genes surveyed by protein electrophoresis do not reflect patterns of genetic differentiation underlying morphologi¬ cal variation. Intraspecific allozymic differentiation probably is lacking because of re¬ cency of population expansion and insufficient time for the evolution of concomitant allozyme markers. If allozyme alleles are selectively neutral (Barrowclough et al. 1985), then the substitution rate equals the mutation rate independent of effective population size in constant populations. Genetic differentiation might require a long period of time. Populations recently isolated will appear to be exchanging genes, but this is an illusion caused by retention of ancestral polymorphisms (Slatkin 1985). Thus, allozyme evolution is probably not sufficiently rapid on average to track frag¬ mentation of temperate avian populations (Gelter & Tegelstrom, this symposium). Protein electrophoresis does provide useful data for inference of phylogenetic rela¬ tionships among congeneric species. However, with only a single data set (of any kind) one has only inferential means of ascertaining confidence in a phylogenetic estimate (e.g. Felsenstein 1985). Congruence of genetically independent data sets provides an index of a phylogeny’s reliability. For example, cladograms derived from variation in allozymes and mtDNA within the genus Ammodramus were highly congru¬ ent, which would not be expected by chance alone (Zink & Avise 1990). The most likely factor producing such congruence is phylogeny, or common descent. Additional comparisons of mtDNA and allozymes in the genera Zonotrichia and Pipilo have in¬ dicated significant congruence (Zink et al. in press, Zink & Dittmann in press), sug¬ gesting that both types of data usually contain a discernable phylogenetic signal. At some level of divergence allozymes will provide no useful data for construction of avian phytogenies, although that limit is unknown (Lanyon & Zink 1988). Protein electrophoresis is relatively fast and inexpensive and provides information on multi¬ ple gene loci (Table 1 ). TABLE 1 - Molecular techniques and levels of resolution in avian systematics. RFLP = restriction fragment length polymorphism. RAPD = randomly amplified polymorphic DNA. Geographic Variation Pattern of Speciation Method Recent Ancient Protein Electrophoresis +/- Yes +/- Nucleic Acids MtDNA - RFLP Yes Yes +/- MtDNA Sequences Yes* Yes Yes Nuclear RAPDs Yes +/- No Nuclear Sequences Yes* Yes* Yes Immunology No +/- +/- DNA/DNA Hybridization No +/- +/- +/- = depends on question and cost No = not appropriate Yes = appropriate and cost effective Yes* = appropriate but expensive ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 631 Nucleic acids Mitochondrial DNA. Both restriction (endonuclease) fragment length polymorphisms (RFLPs; Gelter & Tegelstrom, this symposium) and sequences (Johnson & Cicero, Arctander, and Edwards, Quinn, & Wilson; this symposium) are being used frequently in systematic studies (Moritz et al. 1987) including those dealing with birds (Shields & Helm-Bychowski 1988). Within species, mtDNA RFLPs have greater resolving power than allozymes (Avise & Zink 1988, Zink in press, Gelter and Tegelstrom this symposium), although not all species exhibit geographic differentiation. In the Fox Sparrow Passerella iliaca significant differences in mtDNA, morphology and breeding habitats occur at the boundary of the Sierra Nevada and Great Basin, whereas no allozyme markers were discovered (Zink in press). In many instances, mtDNA differ¬ entiation mirrors morphological breaks (Johnson & Cicero, Arctander; this sympo¬ sium), but not all morphological breaks are accompanied by mtDNA differentiation. MtDNA and morphology may evolve at roughly similar rates in bird species. A further advantage of mtDNA is the potential to construct mtDNA allele phylogenies. With allozymes, one can estimate gene frequencies, but not phylogenetic relation¬ ships among alleles segregating at individual gene loci. Alleles at a locus evolve in a hierarchical fashion, as do species in a lineage. Because one can construct a phylogenetic network of mtDNA alleles found in different geographic samples, analy¬ sis of geographic variation is facilitated (termed “phylogeography” by Avise et al. 1987). Slatkin and Maddison (1989) show how mtDNA allele phylogenies can be used to estimate gene flow. For example, in a study (Zink et al. manuscript) of mtDNA vari¬ ation within the Common Grackle Quiscalus quiscula , four individuals at one site each possessed a unique mtDNA clone. These four “alleles” were found nowhere else, suggesting limited gene exchange. However, phylogenetic analysis of restriction site differences revealed that the four clones did not share a most recent common clonal ancestor but rather had nearest clonal relatives in other geographic samples. Signifi¬ cant gene flow is indicated because clones at this locality do not trace their common ancestry to a mother that existed at that site (Slatkin & Maddison 1989). Sequencing and RFLP analyses of mtDNA permit inference of phylogenetic relation¬ ships among avian taxa (Zink & Avise 1990; Arctander, and Edwards et al. this sym¬ posium), although I stress that mtDNA analysis provides a single “gene genealogy” embedded within the organismal phylogeny. The entire mtDNA molecule, although encoding several “genes,” has a single history because the entire mtDNA molecule is matrilineally inherited as a single linkage group. One expects that single gene genealogies will depart from the organismal phylogeny (Pamilo & Nei 1988), espe¬ cially in the interval of 4N (N = population size) generations after the nuclear gene pools become isolated (Neigel & Avise 1986, Ball et al. 1990). Comparison of mtDNA- based cladograms with those derived from other character sets are necessary to measure confidence that any mtDNA tree (even if the entire molecule has been sequenced) reflects the organismal tree. Nuclear Genes. To complement mtDNA analyses, a pressing need is for similar stud¬ ies (especially sequencing) of nuclear genes. At this point, few have attempted sequencing or RFLP studies in avian systematics (Mindell & Honeycutt 1989, Gelter & Tegelstrom this symposium). The amplification of nuclear DNA using random 632 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI primers offers the exciting potential to detect genetic markers for use in intraspecific studies (Williams et al. in press). Analyzing nucleic acids. Sequence comparisons will become one if not the major tool of molecular systematists, as revealed in this symposium. Although DNA sequencing is of interest at all levels, it is sometimes too expensive (at present) relative to other methods that can answer the same question. Also, with only four bases possible, nucleotide positions become “saturated” with base substitutions, and beyond a cer¬ tain level of divergence some sequences are uninformative because multiple substi¬ tution events at a position cannot be discovered. Insertions and deletions in DNA, recognized as “gaps” in sequence alignments, are frequent phenomena that present serious analytical problems (Felsenstein 1988). Nonetheless, comparison of DNA sequences represents the major advance in systematics. Immunology and DNA-DNA Hybridization These (distance) techniques are those that produce a single measure of similarity or difference between pairs of taxa. Immunological techniques, addressed in this sym¬ posium by Baverstock and colleagues, produce interesting information. These tech¬ niques probably are not appropriate for phylogeny inference (Table 1; Baverstock et al. this symposium) because they suffer several biases such as non-reciprocity. Fur¬ thermore, immunological techniques measure genetic differentiation at a single locus, and as discussed above, single genes might often be discordant with organismal phylogenies (Pamilo & Nei 1988). Immunological distance values can be mapped onto a phylogeny to reveal differences in evolutionary rates. However, there are too many reasons why a distance tree can be incorrect for them to be primary tools for phylogeny inference, and these reasons are difficult to study when all data are com¬ bined into a single distance measure (Swofford & Olsen 1990). DNA-DNA hybridiza¬ tion is also a secondary tool for phylogeny inference because of the many correction factors that are required for the technique to yield reliable phylogenies (Werman et al. 1990; see Krajewski [1989] for another opinion), except perhaps for anciently di¬ verged taxa (Table 1), where sequence comparisons are compromised by insertions and deletions. It would be of comparatively greater interest to map DNA-DNA hybridi¬ zation values onto a phylogeny than those derived from immunological (single gene) comparisons because the former surveys many genes. Techniques versus questions Although the phenomena investigated by systematists are inherently genetical, mo¬ lecular methods are not a panacea. Many have assumed that having genetic informa¬ tion superseded attention to data analysis and to evolutionary questions themselves. Some aspects of the evolutionary process will be difficult to resolve with molecular (or any) techniques. Some speciation patterns, those with closely spaced nodes occur¬ ring a relatively long time ago, cannot be discerned with characters that evolve at constant rates (Lanyon 1988). Thus, what was heralded as the virtue of molecular characters, their uniform rate of change, insures that they will be unable to resolve certain types of phylogenetic patterns. Characters involved in “key innovations” might be required to document monophyly of lineages that diverged close in time - such characters might be morphological or biochemical (e.g. protein or RNA configura¬ tions). Although it was once thought that molecular methods would expose the “ge¬ netics of speciation” it is now realized that unless one can study directly the genes that influence ranking traits (those that delimit species boundaries) we will still only ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 633 have indirect knowledge of processes of speciation (Cracraft 1989, Zink in press). Consequently, molecular characters will not be the data of choice for all questions in systematics and evolutionary biology. Nonetheless, a powerful and growing battery of molecular techniques permits robust inferences of evolutionary patterns and proc¬ esses. RATES OF MOLECULAR EVOLUTION A reason for early enthusiasm about molecular methods was the postulated existence of a molecular clock. Ensuing years have indicated that rate constancy is likely lim¬ ited to clades and molecules (Vawter & Brown 1986). Attempts to calibrate multi-lo¬ cus allozyme clocks reached little agreement (Avise & Aquadro 1982). The one at¬ tempt to calibrate an avian molecular clock (Gutierrez et al. 1983) was based upon a fragment of quail skeleton. This fossil was taken as the minimum age of a particu¬ lar lineage and led to the suggestion that one unit of Nei’s (1978) genetic distance accrued in 26.3 million years. Marten and Johnson (1986) challenged this fossil’s dating and suggested a calibration of 19.7 million years. Clearly, a calibration based on one data point is dubious at best. Calibration of mtDNA evolution in birds, namely 2% sequence divergence per million years, agrees with the value estimated for pri¬ mates (Shields & Wilson 1987). Opportunities to calibrate molecular divergences among avian taxa are limited by lack of information on relevant fossils. With inadequate information, one is left with indirect means of estimating confidence in calibration factors. I compared estimates of divergence dates for several avian taxa using allozyme and mtDNA data. On average, plots of allozyme and mtDNA estimates of differentiation should yield a slope of 1 .0 if the two are equally good at measuring evolutionary separation (indeed, if they are not, then even an “approximate” clock does not exist). This does not address the reliability of the calibrations based on quail and goose fossil remains. If the two independent calibrations gave the same diver¬ gence date for pairs of taxa, the slope should not only be one, but the regression line should pass through the origin. I plotted mtDNA (RFLP) and allozyme estimates of dates of divergence for shorebirds (Dittmann & Zink, unpubl. data), towhees (Zink & Dittmann in press) and sparrows (Kessler & Avise 1985, Zink et al. in press). Using the calibration for allozymes of 26.3, the y-intercept was 0.37 and the slope was 1.44. The calibration of 19.7 (Fig¬ ure 1) yields a slope of nearly 1.0 and a y-intercept of 0.27; 65% of the variance is explained. Although there is an apparent saturation effect in the mtDNA data (due mostly to the shorebird data), these two independent estimates provide very similar estimates of divergence dates, warranting further study. ARE MOLECULAR METHODS BETTER THAN MORPHOLOGICAL ONES FOR INFERENCE OF PHYLOGENIES? The literature has seen numerous debates on molecules versus morphology. Hillis (1987) noted that this is an unfortunate contrast, as most dichotomies usually are. For study of geographic variation and speciation, I consider molecular methods necessary for a study to be complete. In phytogeny inference, however, the roles of 634 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI morphological and molecular data are unclear. There are differences in the nature and analysis of molecular and morphological data. For example, morphologists study vari¬ ation in their taxa and define characters a priori, which can mitigate the effect of homoplasy and invariant characters. Molecular systematists choose a region of the genome, which is a deliberate rather than a random choice, but then the characters (e.g., bases in a DNA sequence) are evaluated without prejudice. In the analysis of molecular data the number of character states is often limited, whereas there may be many states in a morphological character (Mickevich & Weller 1990). Also, morpho¬ logical characters likely have significant genetic covariation, yet they are analyzed as though genetically independent, whereas molecular characters, if from unlinked genes, are likely independent in genetic transmission. In my opinion, a molecular method is the method of choice if the broad limits of the group are already known (probably based on morphology!), multiple genes can be sequenced, these genes exhibit appropriate rates of evolution, and the evolutionary history of the group in question does not consist of closely spaced ancient nodes. Even then, if the phylogenetic estimate does not strongly favor one particular branching structure, one should not be overconfident in the pattern. Similarly, one should not be overconfident in a gene tree (e.g., mtDNA gene lineage) no matter how robust it is, because it simply might not reflect the species tree (Pamilo & Nei 1988, Nei 1987). Many might feel that molecular methods are inherently superior for phylogeny infer¬ ence because they expose genetic variation directly. Comparisons of independent FIGURE 1 - Plot of mtDNA versus allozyme dates of divergence for towhees (P/'p/'/o), spar¬ rows ( Zonotrichia , Junco, Melospiza), and shorebirds ( Phalaropus , Calidris, Tringa, Limnodromus, Recurvirostra). ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 635 molecular estimates of the same sets of taxa will establish the robustness and levels of homoplasy in molecular data sets (the same is true for morphology). If multiple molecular data sets support a robust phytogeny, morphological data sets can be ex¬ amined to determine their reliability for phylogeny inference in that group. We can anticipate several outcomes. First, the problems with analysis of molecular data might be too great for them to be consistently “superior.” Second, it might prove that mor¬ phological data sets are best mapped onto a phylogeny derived from molecular data sets, thus revealing the nature of morphological evolution in particular lineages. Third, morphology might prove equally efficient (and cheaper) at recovering information about hierarchical relations among organisms. Fourth, morphology and molecules might be equally ambiguous, indicating that the nature of the evolutionary history is recalcitrant to its recovery (Lanyon 1988). It is not yet apparent which of these out¬ comes is most consistent in general with the results of evolution. Multiple data sets are complementary and are needed for a complete understanding of evolutionary patterns and processes. CONCLUSIONS Many aspects of avian systematics have proved difficult to understand when studied by traditional methods, and the field of avian systematics has for the most part languished during the past 30-40 years. Molecular methods have infused new enthu¬ siasm into the field that is demonstrated in this symposium. The future will, I think, yield important insights into avian population structure and evolutionary processes, and the phylogenetic relationships of many avian groups. Systematics is indeed alive and well, and should be an increasingly important field of study for ornithologists. ACKNOWLEDGMENTS I thank G. F. Barrowclough, J. M. Bates, K. J. Burns, D. L. Dittmann, S. J. Hackett, D. P. Pashley, A. T. Peterson, and S. J. Weller for thoughtful discussions on the topics presented herein. D. L. Dittmann and W. L. Rootes provided excellent laboratory assistance. 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ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 637 AUTHOR INDEX Abs M . 1903 Alatalo R V . 1418 Alisauskas R T . . .....2170 Ambrose S J . .............1988 Amundsen T . . . . ....1707 Andors A V . ..553 Andors A V............ . . . . 563 Andors A V . . . . . 587 Angelstam P K.... . 2292 Ankney C D . . . . 2170 Arcese P.. . 1514 Arctander P . . . ............619 Armstrong A J . . . ..........450 Arrowood P C . . . ...653 Arrowood P C . . 666 Arrowood P C . . 697 Atkinson I A E . 127 Avise J C . 51 4 Bacon P J . 1500 Baines D . 2236 Bairlein F . 2149 Baker A J . 493 Baker A J . 504 Ball G F . 984 Ball R M... . 514 Baptista L F . 1243 Barrett R T . 2241 Barrowclough G F . 493 Barrowclough G F . 495 Bateson P . 1 054 Baverstock P R . . . 591 Baverstock P R . 61 1 Beason R C . 1803 Beason R C . 1813 Beason R C . 1845 Beissinger S R . 1727 Bell B D . 5 Bell B D . 65 Bell B D . 193 Berkhoudt H . 897 Berruti A . 2246 Berthold P . 780 Biebach H . 773 Bird D M . 2429 Birkhead T R . . . 1345 Birkhead T R . . . 1347 Blokpoel H . .........2361 Blokpoel H . . . .....2372 Blokpoel H . . 2396 Bock W J . . 84 Boersma P D . .962 Boles W E . . 383 Bradley J S . . . . . ....1657 Brandl R . 2384 Braun E J . . . 21 30 Briggs S V . . .843 Brisbin I L Jr . 2473 Brisbin I L Jr . . . 2503 Brisbin I L Jr . .....2509 Brooke M de L . . . 1091 Brooke M de L . 1 1 13 Brooke R K . 450 Brown R G B . 2306 Brummermann M . 1785 Bryant D M . 1989 Bucher E H . 247 Bucher E H . 681 Bull P C . 62 Burgess E C . 2353 Burley N T . 1367 Burley N T . 1373 Burns M D . 2257 Butler P J . 1 875 Cadiou B . 1641 Calder W A . 800 Capparella A P . 307 Carey C . 263 Carey C . 800 Carpenter F L . 1 1 56 Carpenter F L . 1 1 88 Cassidy ALE V . 1514 Catterall C P . 1 204 Cawthorn M . 1 229 Chambers G K . 554 Chandola-Saklani A . 2030 Cherel Y . 2177 Christidis L . 359 Christidis L . 392 638 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Christidis L . 61 1 Cicero C . 600 Cimprich D A . 1 432 Clayton N S . 1252 Clout M N . 1 61 7 Cockrem J F . 2092 Collier K J . 860 Collins B G . 1 1 39 Collins B G . 1 1 66 Cooke F . 1666 Cooper A . 554 Coulson J C . 2365 Craig J L . 231 Craig J L . 251 3 Craig J L . 2546 Craig J L . 2553 Craig J L . 2561 Crowe T . 449 Crowe T . 483 Croxall J P . 279 Croxall J P . 1393 Cullen D P . 1229 Curry R L . 1 333 Custer T W . 2474 Cuthbert F J . 2401 Daan S . 1976 Danchin E . 1641 Daugherty C H . 525 Davidson N C . 2228 Davis L S . 1352 Dennison M D . 504 De Laet J . 1436 Derrickson S R . 2402 Dhondt A A . 141 7 Dhondt A A . 1436 Drent R . 761 Droge D L . 932 Dumbell G S . 251 3 Dumbell G S . 2561 Dunnet G M . 1639 Dyer A B . 1061 Eadie J M . 1 031 Ebihara S . 2015 Edwards S V . 628 Eens M . 1003 Ellis D H . 2403 Elzanowski A . 1921 Elzanowski A . 1938 Emmerton J . 1 837 Ens B J . 889 Erikstad K E . 2272 Escalante-Pliego P . 333 Estrella R R . 1 641 Evans PR . 2197 Evans P R . 2228 Evans P R . 2236 Evans R M . 1 734 Faith D P . 404 Feduccia A . 1 930 Feinsinger P . 1480 Feinsinger P . 1605 Fivizzani A J . 2072 Fjeldsa J . 342 Forbes L S . 1720 Ford H A . 826 Ford H A . 1 141 Ford H A . 1470 Ford H A . 1568 Franke I . 31 7 Freed L A . 1214 Friend M . 2323 Friend M . 2331 Friend M . 2356 Furness R W . 1678 Furness R W . 2239 Furness R W . 2241 Gaston A J . 2306 Gee G F . 2403 Gelter H P . 592 Gentle M J . 191 5 Gerstberger R . 21 1 4 Gnam R S . 673 Goldsmith A R . 2063 Goslow G E Jr . 701 Goslow G E Jr . 716 Goslow G E Jr . 748 Goss-Custard J D . 2199 Goto M . 2015 Goudie R 1 . 81 1 Gowaty P A . 932 Grant M . 2236 Grant P R . 1333 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI 639 Gray D A . 2105 Gray R D . 2553 Griffin J M . 1273 Groscolas R . 21 77 Grubb T C Jr . 1432 Gustafsson L . 1425 Gwinner E . 2005 Gwinner E . 2022 Haig S M . 2410 Haila Y . 2286 Handrich Y . 2177 Hasegawa M . 201 5 Haskell M . 860 Hausberger M . 1262 Hay J R . 2523 Heinsohn R G . 1309 Heldmaier G . 2042 Henderson I M . 860 Hinsley S A . 1757 Hirth K-D . 722 Hixon M A . . . 1156 Hochachka W M . 1514 Hogstedt G . . 1584 Holmes R T . 1542 Holmes R T . 1559 Homberger D . 398 Horne J F M . 468 Howe R W . 903 Hughes M R . 2138 Hulscher J B . 889 Hummel D . 701 Hummel D . 730 Hummel D . 748 Hunt G L Jr . 2272 Hunter F M . 1347 Hunter M L Jr . 2283 Hussell D J T . 947 Ilyichev V D . 91 Imber M J . 1377 Imber M J . 1402 Imber M J . 141 3 Innes J G . 2523 Irons D B . 2378 Isenmann P . 2384 Jackson D B . 2236 Jackson S . 1378 Jaksic F M . 1480 James F C . 2454 James F C . 2469 James H F . 420 Jarvinen O . 1479 Jenkins P F . 1262 Jenkins P F . 1285 Jimenez J E . 1480 Johnson N K . 600 Jones D R . 1 893 Karasov W H . 21 59 Kato A . 1393 Keast A . 41 9 Keast A . 435 Kemp A C . 483 Kempenaers B . 1 436 Kenagy G J . 1976 Kerlinger P . 1 1 22 Ketterson E D . 1229 Kikkawa J . 578 Kikkawa J . 1 195 Kikkawa J . 1 204 Kikkawa J . 1 240 King J R . 2186 Klinke R . 1805 Klomp N 1 . 1678 Komdeur J . 1 325 Kooyman G L . 1887 Korf H-W . 2006 Korpimaki E . 1528 Kreig M . 61 1 Kruijt J . 1068 Lal P . 2030 Lambeck R H D . 2208 Lamey T C . 1 741 Lank D B . 1 666 Lawler W G . 843 Laybourne R . 2454 Le Maho Y . 1 777 Le Maho Y . 21 77 Lebreton J D . 2384 Lee S C . 1734 Lee W G . 1617 Levey D J . 1624 Louette M . 475 Lovvorn J R . 1 868 640 ACTA XX CONGRESSUS INTERNATIONALE ORNITHOLOGICI Lundberg A . 1360 Lynch A . 1244 Lyon B E . 1023 Majer J D . 1 568 Mann N I . 1074 Martella M B . 681 Martin G R . 1091 Martin G R . 1 1 30 Martin G R . 1830 Martin L F . 681 Martin T E . 1 595 Masman D . 1976 Maurer B A . 826 Maurer B A . 835 May R M . 1012 McDonald M V . 1245 McFarland D C . 1 1 41 McKinney F . 868 McKinney F . 876 McKinney F . 885 McLean I G . 1273 McNab B K . 860 McNee S . 1166 McNeil R . 1098 Meathrel C E . 2390 Meire P M . 2219 Melancon M J . 2474 Mench J A . 1905 Merton D . 2514 Migot P . 2365 Millener P R . 127 Mills J A . 1522 Mills J A . 2390 Minot E O . 929 Minot E O . 992 Mock D W . 1703 Mock D W . 1741 Moermond T C . 903 Moller A P . 1001 Moller A P . 1041 Monaghan P . 2257 Monaghan P . 2365 Monnat J-Y . 1641 Montevecchi W A . 2246 Moore F R . 753 Moore F R . 787 Moore F R . 1 1 22 Moorhouse R J . 690 Mumme R L . 1317 Murphy M E . 2186 Murray K G . 1605 Nachtigall W . 722 Nagy K A . 793 Naito Y . 1393 Navarro J L . 681 Nee S . 1012 Nemeschkal H-L . 459 Nesmith C . 2454 Nettleship D N . 87 Nettleship D N . 2239 Nettleship D N . 2263 Newton I . 1689 Newton I . 2487 Newton L . 860 Nolan V Jr . 1229 Norman F 1 . 876 Obst B S . 793 Obst B S . 920 Oehme FI . 737 Ohlendorf H M . 2474 Oring L W . 2072 Ormerod S J . 2494 Oshima I . 2015 Owen M . 1 105 Paabo S . 554 Pain D J . 2343 Pant K . 2030 Parkin D T . 2435 Parkin D T . 2469 Part T . 1425 Paton D C . 1 1 56 Paton D C . 1611 Pearson D L . 1462 Pellatt E J . 1347 Perrins C M . 1499 Perrins C M . 1 500 Peters D S . 572 Peterson R W . 91 Petrie M . 1001 Petrie M . 1041 Philip HRH The Prince . 107 Piatt J F . 791 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 641 Piatt J F . 81 1 Piatt J F . 2272 Pienkowski M W . 2228 PlERSMA T . 761 Piersma T . 2228 PlNXTEN R . 1003 Place A R . 913 Place A R . . . 1378 Plunkett G . 1244 Ponganis P J . . 1887 Potter M A . . . 2092 Powell A N . 2423 Power D M . 545 Pratt T K . 425 Price D K . . . 1367 Prince P A . 1113 Prinzinger R . 1755 Prinzinger R . 1799 Pruter J . 2365 Quinn T W . 628 Quinn T W . . . 2441 Quinn T W . 2469 Ralph C J . . 1 444 Rapoport E H . 826 Rattner B A . 2474 Rayner J M V . 702 Rebelo A G . 1 1 80 Recher H F . 1470 Recher H F . 1 568 Reed C . 2514 Reinertsen R E . 1755 Reinertsen R E . 1799 Ricklefs R E . 929 Ricklefs R E . 992 Ridoux V . 1392 Risebrough R W . 2480 Rising J D . 534 Ristau C A . 1937 Robertson H A . 1617 Robertson R J . 974 Robin J-P . 21 77 Rockwell R F . 1666 Rogers C M . 1514 Root T . 817 Russell R W . 1 1 56 Rusterholz K A . 903 Saitou T . 1 1 96 Saunders D A . 653 Saunders D A . 658 Saunders D A . 697 Scharf W C . 2372 Schodde R . 404 Schodde R . . . 41 3 Schodde R . 61 1 Schuchmann K-L . 305 Semm P . . 1813 Sheedy C . 61 1 Sherry T W . 1542 Sherry T W . 1559 Short L L . 468 Sibley C G . 61 Sibley C G . 1 09 Siegfried W R . 450 Silverin B . 2081 Simon E . 21 05 Slagsvold T . 1703 Slagsvold T . 1707 Slater P J B . 1074 Smith D G . 2403 Smith J N M . 1514 Sorenson L G . 851 Spaans A L . 2361 Spaans A L . 2365 Spaans A L . 2396 Spurr E B . 2534 Stangel P W . 2442 Stangel P W . 2469 Steadman D W . 424 Stevens G R . 361 Stoleson S H . 1727 Strijkstra A M . 1 976 Sturges F W . 1 559 Suhonen J . 141 8 Sullivan K A . 1 957 Sutherland W J . 2199 Tegelstrom H . 592 Temeles E J . 1 1 56 Temple S A . 2298 ten Cate C . 1 051 ten Cate C . 1081 Terrill S B . 751 Terrill S B . 752 642 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI Thaler E . 1791 Thiollay J-M . 1489 Thiollay J-M . 1576 Thomas D H . 1757 Thomas D H . 2122 Thomas N J . 2331 Thornhill R . 1361 Tiebout H M III . 1175 Tiebout H M III . 1605 Triggs S J . 525 Triggs S J . 860 Uttley J D . 2257 Van Horne B . 2313 VAN DEN ELZEN R . 459 van Noordwijk A J . 2433 van Noordwijk A J . 2462 van Noordwijk A J . 2469 Vauk G . 2365 Veltman C J . 860 Verheyen R F . 1003 Vermeer K . 2378 Vickery J A . 2494 VUILLEUMIER F . 327 Vuilleumier F . 354 VUILLEUMIER F . 553 Vuilleumier F . 578 Vuilleumier F . 587 Wallace M P . 2417 Watts C . 1012 Weathers W W . 1957 Webster M D . 1765 Wenzel B M . 1820 Wetton J H . 2435 Whitehead M D . 1384 Whiteley P L . 2338 Wiens J A . 1461 Wiersma P . 761 Wiley J . 2417 Williams J B . 1964 Williams M J . 841 Williams M J . 860 Williams M J . 876 Williams T D . 1393 Willson M F . 1630 Wilson A C . 554 Wilson A C . 628 Wilson A C . 2441 Wilson J B . 1617 Wilson R P . 1853 Wiltschko W . 1803 Wiltschko W . 1845 Wiltscho R . 1803 Wiltscho R . 1845 Wingfield J C . 2055 Wobeser G . 2325 Wobeser G . 2356 Wolf L . . . 1229 Wooller R D . 1657 Wooller R D . 2390 Worthy T H . . 555 Yamagishi S . 1 1 95 Yamagishi S . 1220 Yamagishi S . 1240 Young B E . 1605 Yuill T M . 2338 Zack S . 1301 ZlEGENFUS C . 1229 Zink R M . 591 Zink R M . 629 Zweers G A . 897 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI 643 NOTES 644 ACTA XX CONGRESSUS INTERNATIONALIS ORNITHOLOGICI NOTES / QL671 .17 1990 \ita \\ Congress! is Intcriiationulis 3 2044 062 442 769 DATE DUE DEMCO, INC 38-2931