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University of Cape Town, South Africa The moult and migration strategies of Lesser Sand Plover, Greater Sand Plover and Terek Sandpiper Town Cape of Colin Jackson UniversityJCKCOL003 Thesis presented for the Degree of Doctor of Philosophy in Zoology, Department of Biological Sciences, University of Cape Town, South Africa March 2017 The copyright of this thesis vests in the author. No quotation from it or information derivedTown from it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes Capeonly. of Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University Supervisors Professor Les G. Underhill Animal Demography Unit Zoology Department of Biological Sciences University of Cape Town Dr. Clive Minton Australasian Wader Study Group Melbourne Australia Plagiarism declaration I know the meaning of plagiarism and declare that all of the work in this thesis, save for that which is properly acknowledged, is my own. Signature:________________________ Date:________14th March 2017______ Contents Abstract ii Acknowledgements iv Chapter 1 Introduction 1 Chapter 2 The moult of Lesser Sand Plovers in Kenya, India and Australia 22 Chapter 3 The moult of Greater Sand Plovers in Kenya, India and Australia 60 Chapter 4 The moult of adult Terek Sandpipers in Kenya, India and Australia 89 Chapter 5 The moult of immature Terek Sandpipers in Kenya, India and 114 Australia Chapter 6 The moult of Terek Sandpipers in South Africa 144 Chapter 7 The status and migration of the Lesser Sand Plover and the Greater 156 Sand Plover Chapter 8 The status and migration of the Terek Sandpiper 191 Chapter 9 A comparison of primary moult strategies in migrant waders 209 Conclusion and recommendations 228 “Even the stork in the sky knows her appointed seasons, and the dove, the swift and the thrush observe the time of their migration.” Jeremiah 8:7 i Abstract Moult in birds is a critical yet often underrated component of a birds’ annual life cycle together with breeding and migration. A good understanding of the moult strategy of a species is important for its conservation because key moulting grounds must be protected to allow this component of the life cycle to be undertaken. Lesser Sand Plovers, Greater Sand Plovers and Terek Sandpipers are among the least known of the commonly encountered long-distance migrant waders in Africa and Australasia. This is particularly true for the populations which spend the non-breeding season on the East African coast. This study aims to describe the primary moult strategies for the three species from five locations at different latitudes: south-eastern India, Kenya, South Africa, north-western Australia and south- eastern Australia. Differences in moult strategy between species, age and latitude are highlighted and explanations are proposed for the differences in the light of the breeding behaviour, migration time and routes, food availability and local climatic conditions. Results from this study are combined with other moult studies on migrant waders to present hitherto undescribed patterns in wader moult strategies at a global scale. A synthesis is presented of published and available unpublished information about the different populations for the three study species, their distribution, status and migration strategies with a focus on those visiting the East African coast. The three species follow a similar basic moult strategy in adults where primaries are moulted soon after arriving on the non-breeding grounds. Immature moult strategies are more complex and differ between species and latitude. In general adults in all three species at the more northern non- breeding sites start moult earlier than those travelling farther south probably as a result of greater proximity to breeding grounds. Lesser Sand Plovers have a relatively long duration of moult (152– 154 days) across all locations where data were sufficient; Greater Sand Plovers have a short duration (80 days) in India and similar, comparatively longer durations (132–134 days) at the other latitudes. The Indian site is used mainly by this species as a site to stopover and moult while on migration to more southerly non-breeding grounds. Terek Sandpipers in Kenya have a relatively short duration of moult (99 days) when compared to India and Australia (121–130 days) and tend to have a rapid start to moult which then slows down after the first few primaries are moulted. Young Lesser Sand Plovers in India mostly undergo a complete primary moult while in Kenya and Australia birds generally do not moult in their first year. First year Greater Sand Plovers mostly all undergo a complete primary moult irrespective of latitude. Terek Sandpipers in their first year do not moult in India in the north, mostly undergo a partial moult of outer primaries in Kenya on the ii equator, and do a complete moult in south-eastern Australia. The north-west Australian population was divided into two almost equal groups: those that undergo a complete primary moult, and those that only moult a few outer primaries. A difference in winglength between these two groups suggests that there may be two distinct populations involved, most likely originating from distinct breeding grounds. In-depth study is needed to confirm this. The overarching picture of moult in adult migrant waders is that at high latitudes moult starts early and is kept short. With decreasing latitude, the timing of moult is delayed with increasing distance from breeding grounds. In the tropics the lack of constraints of approaching harsh winter weather and long distances from breeding grounds allow species to adapt a wide range of moult strategies. As birds migrate farther south before they moult, constraints on time available to moult due to large distances travelled mean duration is reduced in order to fit it in. An upper limit in duration of moult in migrant waders irrespective of the size of the bird appears to be set at c. 160 days. Small birds have a wide variation in duration while as body size increases over 100 g, moult duration is necessarily longer. With harsh winters in the north, large waders are forced to migrate south in order to moult. This study suggests that irrespective of body size the ‘gold standard’ for duration of moult is around 120 days (four months). While timing of migration is relatively well-known for the three species in Australia, it was hitherto unknown that juvenile Lesser Sand Plovers do not reach Australia until February or March in contrast to all other migrant wader species where first years arrive in September/October. It is unknown where they stop en route or indeed why they delay in this way. All three species in Kenya peak at a maximum population in November and early December after which numbers reduce as birds start to depart for the breeding grounds. Resightings of colour-flagged Lesser and Greater Sand Plovers confirmed the north-western coast of India as an important stopover site for migrants heading to western China and Mongolian breeding grounds from Kenya. A review of country population estimates for the three species suggested non-breeding populations in Africa of 200,000 Lesser Sand Plovers, 25,000–35,000 Greater Sand Plovers and 115,000–200,000 Terek Sandpipers. Mida Creek is shown to be a key stronghold for non-breeding waders in Kenya and its importance for their continued conservation is thus confirmed. iii Acknowledgements My first thanks are to my supervisors, Prof Les Underhill and Dr Clive Minton, who have made this whole amazing adventure into wader research possible. Les, being ‘at hand’, has borne the bulk of guiding me through the labyrinth of undertaking a Ph.D. and in particular the ins and outs of moult analysis and interpretation. I have hugely appreciated his ‘groundedness’ in relation to statistical results and their application to biology (“all models are wrong, some are useful” rings in my ears…) and beyond that to the real importance of life continuing over and above writing a thesis. Clive has similarly been encouraging from the start and I have appreciated his ability to question results hard and help me reassess them more carefully. Our discussions on wader research in Australia and ideas for the Ph.D. way back at Ngulia were an inspiration to me and helped me to get stuck in to the work. I am particularly grateful to the Australasian Wader Study Group including the Victorian Wader Study Group and to Prof. Balachandran at the Bombay Natural History Society for the provision of all their ringing data on my study species – and putting up with and responding to my pestering with regards to questions on the data. I know how hectic it can be amidst a very busy life to have to answer such fine detail questions – and really appreciate it. David Pearson also gave me all his data from waders ringed at Mida Creek from over 15 years of ringing prior to my ringing there and has taught me huge amounts about ageing and moult in birds over the years. Our lengthy discussions on ageing of birds in the hand certainly contributed enormously to my delving into and understanding moult and ageing of waders. Jon Coleman and the Queensland Wader Study Group also provided some wader data for comparison and reference for which I am grateful. Magda Remisiewicz and Humphrey Sitters gave some extremely useful input to my analyses and asked hard questions of some results which helped in refining them substantially. Magda also over the years gave some very appreciated advice, insights, ideas and guidance on how to analyse moult data – thank you. Magda, Humphrey, Danny Rogers, Chris Hassell and Clive also allowed me to use some as yet unpublished results from their other wader research which I greatly appreciated.
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