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(1994) Snow Buntings Plectrophenax Nivalis: the Behavioural Ecology and Site Use of an Itinerant Flock Species in the Non- Breeding Season

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(1994) Snow Buntings Plectrophenax Nivalis: the Behavioural Ecology and Site Use of an Itinerant Flock Species in the Non- Breeding Season Smith, Richard D. (1994) Snow buntings Plectrophenax nivalis: the behavioural ecology and site use of an itinerant flock species in the non- breeding season. PhD thesis. http://theses.gla.ac.uk/5739/ Copyright and moral rights for this thesis are retained by the author A copy can be downloaded for personal non-commercial research or study, without prior permission or charge This thesis cannot be reproduced or quoted extensively from without first obtaining permission in writing from the Author The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the Author When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given Glasgow Theses Service http://theses.gla.ac.uk/ [email protected] Sno,v Buntings PlectrophenllX nivalis: the Behavioural Ecology and Site Use of an Itinerant Flock Species in the Non-breeding Season Richard D. Smith This thesis is submitted in candidature for the Degree of Doctor of Philosophy, Department of Zoology, University of Glasgow. November 1994. ACKNOWLEDGEMENTS I am totally indebted to my supervisor Neil Metcalfe whose good nature must have been sorely tested in the past five years by the demands of a part-time absentee student. Not only was Neil brave enough (some might use a different adjective) to take me on, he was also primarily responsible for securing much of my funding. Neil's contribution to the direction and analysis of my work has been immense, and he has promptly and uncomplainingly passed extensive comments on the contents of this thesis and several other long and tedious manuscripts. I must also guiltily thank him for being my general dog's-body at Glasgow: printing, copying, procuring, administering, fabricating, liasing, etc., etc. way beyond the call of duty. Many thanks, Neil. Two other persons deserve special mention for making this thesis happen. Des Thompson had a pivotal role in obtaining funding for the initial work on wintering Snow Buntings and parallel studies on the species' breeding biology. He then gave me the confidence to focus future work towards obtaining a higher degree. Des has also given me access to many Scottish Natural Heritage staff and facilities, and provided many stimulating ideas and criticisms. I am also indebted to Mick Marquiss for generating me with enthusiasm, providing practical and statistical advice and discussion, and commenting extensively and intelligently on an earlier draft of this thesis. The study received funding from (in order of monetary value) the Natural Environment Research Council, Scottish Natural Heritage (formerly the Nature Conservancy Council, Scotland), Highlands and Islands Enterprise (formerly the Highlands and Islands Development Board), the Nuffield Foundation, the British Ecological Society, the Association for the Study of Animal Behaviour, Cairngorm Chairlift Company, Craignish Conservation Trust, the British Ornithologists' Union, and the British Trust for Ornithology. The Royal Society for the Protection of Birds provided accomodation during early 1988. I am grateful for the help of a number of colleagues at Glasgow University, especially Martin Bums for computing advice. Liz Denton, Roddie MacDonald, Nozrat Mirzai and Caroline Askew all provided technical support. Thanks also to everyone at Rowardenan Field Station. Scottish Natural Heritage staff have been extremely cooperative, notably Allan Brown who introduced me to the magical world of computing. Marianne Robson, computing staff in Edinburgh and office staff at Achantoul, Aviemore have also been very helpful. The Meteorological Office at Achantoul gave access to weather records. I also appreciate the contribution ofCairngorm Chairlift Company to this project. Tommy Paul, and latterly Tim Whittome, allowed me to work on the Ski Area, provided free chairlift tickets, and were generally responsive to my requests for help. Neil Baxter and the other rangers supplied daily weather records. The garage and road crews enabled me, and often my car, to get to my observation sites in some atrocious weather conditions. Thanks also to the bus drivers for their tolerance. and everyone who mentioned seeing birds. Thanks are also due to Highlands and Islands Enterprise. the Royal Society for the Protection of Birds. the ski companies at Glen Shee and The Lecht. and the farmers and keepers at the Cabrach and Corgarff for letting me work on their land. A number of friends and colleagues have given me assistance. advice, encouragement and floor space. I particularly wish to thank Adam Watson. Hector Galbraith, Phil Whitfield, Pat Monaghan. Keith Duncan. Dave Horsfield. Rab Rae. Colin and Margaret Daveson. Phil Atkinson, Steve Buckland, Dave Thompson, Chris Thomas. Stuart Mackay, Stuart Rae and Juan Carlos Senar. I also wish to thank all those who helped me trap birds. or contributed to my summer observations of Snow Buntings. Trapping data from other North-east Scottish sites were kindly provided by Rab Rae, Raymond Duncan, Mike Kimber. Ian Mackay. Mick Marquiss and Bob Procter. Colour-rings were copiously and promptly supplied by Mike Nicol and were very successful. Kevin Baker administered ringing activities at the B.T.O. I am also grateful to Alan and Tilly Smith for keeping their Reindeer from eating too much of the food that I provided for Snow Buntings. and Charlie Walpole. Ron Grant. George Macdonald Sr. and Jr .• and Dai Davies for generally putting up with and helping me during the last few winters. I must also thank the birds themselves for some great entertainment. Many, especially those which remained to breed in the Caimgorms. became unforgettable individuals. Hopefully, this thesis will go some way towards justifying their trapping and marking. Finally, and by no means least. I would like to express my gratitude to the family and friends who have put up with and supported me through this 'writing-up' stage of the project. Phone calls and visits were more than welcome distractions, which kept me going in the dark months when everything was a chore and the tunnel seemed to have no end. Now, however. the real world beckons .... To my folks, Dorothy and David DECLARATION I declare that the work described in this thesis has been carried out by myself unless otherwise cited or acknowledged. It is entirely of my own composition and has not, in whole or in part, been submitted for any other degree. RICHARD DAVID SMITH November 1994 TABLE OF CONTENTS SUMMARY p6 Chapter 1. INTRODUCTION p9 1.1 Social organisation and winter site choice p9 1.2 Biology of the Snow Bunting pl4 1.3 Aims of this study pIS Chapter 2: STUDY AREA, WEATHER and GENERAL METHODS pl7 2.1 Cairn Gorm Study Area pl7 2.2 Other trapping sites pl8 2.3 Measurement of weather variables pl9 2.4 Definition of a snowfall p20 2.5 Trapping and marking techniques p2 I 2.6 Winter immigrants and local breeding birds p22 2.7 Juveniles and adults p23 2.8 The non-breeding season p23 2.9 Statistical treatment p23 Chapter 3: AGE DETERMINATION, WING-FEATHER COLOUR AND \VING-LENGTH CHANGE p24 3.1 INTRODUCTION p24 3.2 METHODS p25 3.3 RESULTS p26 .1 Evidence for reliability of age determination p26 .2 Amount of white on the flight-feathers p27 .3 Wing-length change p27 3.4 DISCUSSION p29 3.5 APPENDIX: Age determination p30 Chapter 4: AGE AND SEX VARIATION IN CHOICE OF WINTERING SITE: THE EFFECT OF ALTITUDE p32 4.1 INTRODUCTION p32 4.2 METHODS p33 4.3 RESULTS p35 4.4 DISCUSSION p35 1 Chapter 5: POPULATION SIZE, COMPOSITION AND MOVEl\1ENT PATTERNS AT HIGH ALTITUDE SITES p38 5.1 INTRODUCTION p38 5.2 METHODS p39 5.3 RESULTS p41 .1 Age/sex composition of catches p41 .2 Trapping bias p43 .3 Age/sex composition of catches versus sightings p44 .4 Numbers on Cairn Gorm p45 .5 Population composition in relation to snowfalls p46 .6 Movements of ringed birds p48 5.4 DISCUSSION p51 .1 Annual fluctuations in population size and composition pSI .2 Altitudinal preferences at Cairn Gorm p53 .3 Differential migration following snowfalls p53 Chapter 6: FORAGING SUCCESS pS6 6.1 INTRODUCTION p56 6.2 METHODS p58 .1 Analysis of time budgets p59 6.3 RESULTS p62 .1 General activity during time budgets p62 .2 Peck-rate whilst feeding p62 a Treatment of variables p62 b Multiple regression of peck-rates whilst feeding p63 c Individual improvement of foraging success over the winter p65 .3 Percentage of time spent feeding p66 a Multiple regression of time spent feeding p66 b Loglinear analysis of time spent feeding p67 .4 Access to food and feeding positions p70 a Flock-size choice p70 b Priority of access p70 c Ability to remain in a feeding position p71 d Quality of position within the flock p72 6.4 DISCUSSION p73 .1 Influences on feeding efficiency p73 .2 Applicability of the results p73 .3 Extrinsic influences on foraging success p74 2 .4 Intrinsic influences on foraging success p7S .5 Variation in access to the best foraging conditions p76 .6 Ecological implications p77 Chapter 7: AGGRESSION AND DOMINANCE p78 7.1 INTRODUCTION p78 7.2 METHODS p80 .1 Definition of a contest p80 .2 Sources of data p82 .3 Comparisons of dominance status p82 7.3 RESULTS p83 .1 Factors affecting the rate of aggressive interactions p83 a Results from feeding budgets p83 b Results from patch budgets p86 .2 The consequences of aggression p87 a Influence of aggression on feeding behaviour p88 b Effect of aggression on feeding location p88 .3 Factors affecting individual differences in dominance p89 a The role of sex, age and experience
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