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IN BERMUDA by © Miguel Alberto Mejías HABITAT AND ECOLOGY OF BREEDING AND NON-BREEDING WHITE- TAILED TROPICBIRDS (PHAETHON LEPTURUS CATESBYI) IN BERMUDA By © Miguel Alberto Mejías A thesis submitted to the school of Graduate Studies in partial fulfilment of the requirements for the degree of Master of Science Department of Biology Memorial University of Newfoundland January 10th 2017 St. John’s, Newfoundland and Labrador ABSTRACT Pelagic seabirds have terrestrial and marine life stages and conditions at both sites can affect survival. On land, seabirds are threatened by destruction of nests and introduced predators. Assessments of threats at-sea, however, requires data on pelagic movement and behaviour, which is unknown for many seabirds. To mitigate degradation of breeding sites, artificial cavities are sometimes used. To identify pelagic activities geolocators (GLS) are a useful tool. However, data on nest-site selection, nest success and predation of artificial and natural seabird cavities are limited. In Bermuda, we monitored 158 natural and 178 artificial cavities of White-tailed Tropicbirds (Phaethon lepturus catesbyi) in 2013-2015. Tropicbirds favoured natural cavities with nesting sand, smaller entrances and on steeper cliffs. Artificial cavity occupancy varied by location, year and cavity depth. Clutch survival increased in deeper cavities, with smaller entrances and no rubble. Nestling survival varied by year and increased with age. American Crow (Corvus brachyrhynchos) predation was higher in natural cavities, and shallower cavities with larger entrances. We obtained GLS data from 25 recaptured tropicbirds in 2015 and learned that birds in the fall and winter ranged from Bermuda to the Mid-Atlantic Ridge. Non-breeders spent 95% of night periods and 59% of day periods wet. We found nest-cavity traits and temporal factors important for nesting tropicbirds. However, introduced rats were unaffected by cavity dimensions and should be removed. Lastly, we found, for the first time, marine areas where Atlantic tropicbirds are vulnerable and can be studied for pelagic threats. i ACKNOWLEDGEMENTS I would first like to thank my M.Sc. supervisor, Dr. Yolanda Wiersma, for giving me immense support when times were difficult. In addition to my supervisor, I would like to thank the remaining members of my committee, Dr. Ted Miller and Dr. Greg Robertson, for giving me enormous insight and encouraging me to think critically. Special thanks are due to my lab mates, Carley Schacter and Katherine Robbins, for helping me process and present my tropicbird tracking data. I am also indebted to Dr. Dave Schneider and fellow graduate students Greg Fury and Kyle Miller for statistical support. I thank my work colleagues in the Bermuda Department of Environment and Natural Resources, including Mr. Drew Pettit and Dr. Mark Outerbridge for issuing permits allowing tropicbird handling. I am also grateful for the love, support and assistance in the field provided by my loving mother and father, Patricia and Albert Mejías. Thank you both for encouraging me to have confidence in my competence. I also extend my thanks to the numerous friends and family members who assisted me with the capture and recapture of tropicbirds for my geolocator study. This field work was made possible due to the generous funding provided by the Bermuda Zoological Society and the NSERC grant of Dr. Ian Jones, the latter individual who originally proposed studying the at-sea movements of tropicbirds with geolocators. Lastly, I would like to thank the gracious funding I received through Graduate Studies Fellowships and Teaching Assistant position at Memorial University. ii Table of Contents ABSTRACT……………………………………………………………………………i ACKNOWLEDGEMENTS……………………………………………………………ii Table of Contents………………………………………………………………………iii List of Tables…………………………………………………………………………viii List of Figures………………………………………………………………………….xi List of Appendices………………………………………………………………...….xiii 1. Introduction and Overview……………………………………………………...…1 1.1 Thesis Background………………………………………………………………1 Seabird ecology………………………………………………………..................1 Seabird threats relating to human settlement…………………………..................2 Conservation of cavity-nesting seabirds.……………………………..…………..3 Nest-site selection in cavity-nesting seabirds…………….....................................6 Nest-success in cavity-nesting seabirds……………………………………...…...7 Introduced mammalian and avian predators……………….……….…………….9 1.2 Non-breeding distribution of seabirds……………………...….……………..11 iii What we currently know about post-breeding seabirds…..……….…………….11 Techniques used to study movement of non-breeding seabirds………………...12 1.3 Thesis hypothesis for nest-site selection, nest success and predation………………………………………………………...…………….18 1.4 Thesis objectives and predictions for post-breeding distribution and at-sea activities of seabirds……….......……………………………………………….22 1.5 Focal Species: White-tailed Tropicbirds (Phaethon lepturus catesbyi) ………………………………………………………….……………................25 White-tailed Tropicbird distribution and breeding biology…………………………………………………………………………..25 Threats facing White-tailed Tropicbirds across range…………………………………………………………………………….25 White-tailed Tropicbird colony in Bermuda: Threats and Conservation……...…………………………………………………………….27 Seasonal residency of Bermudian White-tailed Tropicbirds……………………………………………………………………..29 1.6 Statement of Authorship……………………...…….…………………………..30 Co-authorship statement…………………………………………………….....30 1.7 Literature Cited…………....................................................................................31 iv 2. Nest-cavity Selection and Nesting Success of Bermudian White-tailed Tropicbirds (Phaethon lepturus catesbyi)………………………….………………………….43 2.1 Abstract………………………………………………………………………...43 2.2 Introduction…………………………………………………………………….44 2.3 Methods……………………………………………………………...…………47 Study area……………………………………………………………………….47 Nest-cavity searching………………………………………………………...…48 Nest-cavity monitoring……………………………………………….................50 Nest-cavity characteristics………………………………………….…………...51 Statistical analysis……………………………………………………………….54 Nest-cavity characteristics…………………………………………..…………..54 Nest-site selection…………………………………………………...…………..54 Clutch and nestling survival……………………………………….……………55 Black Rat and American Crow predation……………………………………….56 2.4 Results………………………………………………………………..……….57 Nest-cavity characteristics………………………………………………………57 Nest-cavity selection…………………………………………………………….57 Clutch survival………………………………………………………………..…60 Nestling survival………………………………………………………………...66 Black Rat and American Crow predation………………………....…………….69 2.5 Discussion…………………………………………………………………….69 Conservation implications………………………………………………………76 v 2.6 Literature Cited……………………………………………………………….77 3. Non-breeding Distribution and at-sea Behavior of Bermudian White-tailed Tropicbirds (Phaethon lepturus catesbyi) in the North Atlantic………………………...………………………………………………………83 3.1 Abstract…………………………………………………...………….……….83 3.2 Introduction………………………………………………..………………….84 3.3 Methods……………………………………………………………...……….87 GLS logger development and retrieval……………………….…………..87 GLS logger programming and analysis……………………….………….89 Statistical analysis………………………………………………..……….92 3.4 Results………………………………………………………………………...93 Retrieval details, body condition and body measurements……………….93 Non-breeding distribution from GLS loggers………………..…………...95 Distribution of male and female tropicbirds………………………...…..102 At-sea activity patterns………………………………………….………102 3.5 Discussion……………………………………………………………...........107 Conservation implications………………………………………………113 vi 3.6 Literature Cited……………………………………………………………...............114 4. Summary and General Discussion…………………………………………………....121 4.1 Thesis Summary………………………………………………….…………121 4.2 Limitations………………………………………………………...……...…124 4.3 Conservation Implications………………………………………...………...126 Implications for breeding White-tailed Tropicbirds…………………….126 Implications for non-breeding White-tailed Tropicbirds………………..129 4.4 Future Research Directions………………………………………………….130 4.5 Literature Cited……………………………………………………………...133 vii List of Tables Table 2-1. Variables recorded from White-tailed Tropicbird nest-cavities expected to influence cavity occupancy, nest success and predation between 2013– 2015. Expected increase (↑) and decrease (↓) of response variables for the appropriate independent variables are shown. Cavity selection variables only applied to natural cavities are denoted with (*). Independent variables that were not tested towards a specific dependent variable are denoted with (X)……………………………………..……….52 Table 2-2. Descriptive statistics of natural and artificial cavities used by White-tailed Tropicbirds breeding in Bermuda between 2013– 2015. The mean ± SD for cavity types are shown outside parentheses and values inside parentheses show range and sample size…………………………………………………………………..………...58 Table 2-3. The total number of clutches, nestlings and nest success of White-tailed Tropicbird nests in Bermuda in 2013 – 2015. Numbers inside and outside brackets correspond to natural and artificial cavities respectively………………………………………………..................…61 Table 2-4. Causes of nesting failures among eggs and nestlings of breeding White-tailed Tropicbirds in Bermuda in 2013 – 2015. Codes for letters: C, American Crows; R, Black Rats; A, Argentine Ants; U, unknown predators (rat or crow)..………………..62 viii Table 2-5. Model selection results for testing hypotheses about the effects of nest-site characteristics and breeding season on Bermudian White-tailed Tropicbird clutch survival.
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