The Behaviour, Population Biology and Physiology of the Petrels

623 Pages·1996·36.25 MB·English

by  John Warham

The Behaviour, Population Biology
and Physiology of the Petrels
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The Behaviour, Population
Biology and Physiology
of the Petrels
JOHN WARHAM
Zoology Department
University of Canterbury
Christchurch, New Zealand
ACADEMIC PRESS
Harcourt Brace & Company, Publishers
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ACADEMIC PRESS LIMITED
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U.S. Edition Published by
ACADEMIC PRESS INC.
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This book is printed on acid free paper
Copyright © 1996 ACADEMIC PRESS LIMITED
All rights reserved
No part of this book may be reproduced or transmitted in any form or by
any means, electronic or mechanical including photocopying, recording,
or any information storage and retrieval system without the permission in
writing from the publisher
A catalogue record for this book is available from the British Library
ISBN 0-12-735-4158
Typeset by Paston Press Ltd, Loddon, Norfolk, UK
Printed and bound in Great Britain by the University Press, Cambridge
Contents
Preface vii
CHAPTER 1
Petrel Populations 1
CHAPTER 2
Petrels at Sea: Distribution, Dispersal and Migration 64
CHAPTER 3
Feeding and Foods 124
CHAPTER 4
Behaviour and Vocalizations: A General Introduction 173
CHAPTER 5
Behaviour of Albatrosses 209
CHAPTER 6
Behaviour and Vocalizations of Procellariidae, Hydrobatidae
and Pelecanoididae 257
CHAPTER 7
Physiology and Energetics 317
CHAPTER 8
Biochemistry 364
CHAPTER 9
Locomotion 388
CHAPTER 10
Anatomical Matters 424
v
vi BEHAVIOUR, POPULATION BIOLOGY AND PHYSIOLOGY OF PETRELS
CHAPTER 11
Evolution and Radiation 474
CHAPTER 12
Petrels and Man 506
References 542
Species Index 607
Subject Index 610
Preface
When beginning this companion volume to The Petrels back in 1990 I would have
been alarmed to know that 5 years' steady work would be needed to review topics
ignored or only touched upon in the earlier book. In practice, the amount of
information proved to be very extensive and scattered, added to which some
aspects, such as the study of tubenoses at sea, have been intensively researched
recently and it has been difficult to keep abreast of developments.
On account of page limitations the text is quite condensed, so that some topics
may seem to be treated rather summarily and discussion curtailed, while other,
perhaps lesser-known, aspects are given more prominence. However, I have tried to
ensure that ample referencing points the reader to the appropriate literature in such
instances.
I have been greatly helped by others who have provided data, tape-recordings,
and preprints of work in the press, and to many workers and their publishers who
have permitted the use of their illustrations: these are acknowledged in the text.
Colleagues in the Zoology Department have continued their support and I am also
indebted to our University Librarians for help over many years. A particular debt is
owed to colleagues who have commented on parts of the text. These include D. G.
Ainley, W. R. P. Bourne, V. Bretagnolle, J. C. Coulson, M. E. Forster, R. Gales, M. P.
Harris, J. A. L. Hector, M. J. Imber, J. Jacob, G. R. Martin, D. B. Peakall, L. McPherson,
K. A. Nagy, D. T. Parkin, A. M. Paterson, C. J. Pennycuick, P. A. Prince, D. R.
Thompson, B. M. Wenzel and G. C. Whittow. Errors are my own, and I hope not to
have passed any on—'from one naturalist to another as an heir-loom'—to use F. W.
Hutton's words from long ago!
My final gratitude goes to my wife Pat, who can now look forward to a husband
free to tackle rather more mundane tasks, such as painting the house, and even find
time for a holiday.
References to the earlier volume take the form The Petrels, p. 00. Statistical means
are ± 1 SD unless otherwise stated, and nomenclature remains that of the checklist in
The Petrels, pp. 424-A33, except that I have used Pseudobulweria for aterrima, rostrata
and macgillivrayi.
John Warham
Christchurch
vu
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CHAPTER 1
Petrel Populations
Contents
I Introduction 1 VI Mortality 26
II Colony sizes 3 A Loss of eggs 26
A Estimating the sizes of petrel B Loss of chicks 28
populations 3 C Adult losses on land 28
III Coloniality 6 VII Population changes 39
A Introduction 6 A Population increases 39
B Colony characteristics 8 B Population declines 42
C Colony development 8 VIII Population regulation 42
IV Population structure 10 IX Petrel communities on land 45
A Introduction 10 A Introduction 45
B Cory's Shearwater at Grand B Some major communities 46
Salvage Island 11 C Factors affecting distributions
The Sooty Albatross at Possession on land 49
Island 11 Habitat modification 50
D Non-breeders and prebreeders 11 A Introduction 50
Age structures of petrel B Effects on plants 51
populations 16 C Effects on soils 57
A Age at first return to the XI Shearwater 'greens' 58
colony 17 XII Effects of farming practices 60
B Age at first breeding 17 A Grazing animals 61
C Fecundity 19 B Fire 63
D Longevity and survival 20 XIII Seed dispersal 63
I Introduction
Judging by their worldwide distribution, numbers and biomass, the Procellarii-
formes are the most successful of all seabirds. Petrel populations vary in size from
the very small to the very large.
Biggest aggregations occur at high latitudes, particularly in the subantarctic.
Although some northern hemisphere birds such as the storm petrels Oceanodroma
leucorhoa and O.furcata reach large numbers, and Fulmarus glacialis about 4.3 million
pairs, most of the rather few species there have relatively small populations, perhaps
due to past and present competition from the auks. For example, Lensink (1984)
estimated that the three breeding Alaskan petrels totalled about 11 million birds, the
12 species of auks amounting to some 35.6 million birds. It seems more likely that
their competition has held down the northern petrel radiation rather than a lack of
protection from predators, since burrowing auks do well.
Some estimates of world populations of petrel species have been given, mostly of
the easily counted surface nesters. For instance, almost all Laysan Albatrosses
Diomedea immutabilis breed on islands of the Hawaiian Leeward Chain which,
according to Fefer et al. (1984), support 759 000 breeding birds plus an estimated
1 771000 non-breeders, that is, a total of about 2.5 million albatrosses (Table 1.1).