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Download The THE INFLUENCE OF BROOD-SIZE ON REPRODUCTIVE SUCCESS IN TWO SPECIES OF CORMORANT, Phalacrocorax auritus & P_. pelagicus, AND ITS RELATION TO THE PROBLEM OF CLUTCH-SIZE by IAN ROBERTSON B.A., University of British Columbia, 1966 B.Sc, University of British Columbia, 1968 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department o r Zoology We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA August, 1971 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department The University of British Columbia Vancouver 8, Canada ii ABSTRACT Brood-rearing capabilities and their relation to the problem of clutch-size were investigated in two cormorant species, Phalacrocorax auritus and P_. pelagicus, at Mandarte Island, British Columbia. Extra chicks were added to normal sized broods (1 to 4 chicks) so that the range of brood-sizes studied was 1 to 8 in Double-crests (auritus) and 1 to 7 in Pelagics (pelagicus). Fledging success in Double-crests is generally 90 per cent or higher in broods of up to 6 chicks. In Pelagics fledging success drops sharply below 90 per cent in broods greater than 4. In no Pelagic nest did more than 4 chicks fledge, whereas up to 7 chicks fledged in Double-crest nests. Growth rates in both species are lower in supernormal broods. However, this decline amounts to only 12 per cent in Double-crest broods. It is 35 per cent in Pelagic broods. There is a strong relationship between mortality and the low growth rates of certain Pelagic chicks in supernormal broods. Slower growth in large broods is reflected in a fledging period which is longer by several days in both species. The return to Mandarte Island of yearling Double- crests (in 1970) indicates a post-fledging mortality of approximately 50 per cent in normal broods and 47 per cent in supernormal broods, although the latter sample includes only one brood of 7. This slight survivorship difference is not enough to offset the reproductive advantage of Marge broods. Observation of feeding rates in both species indicates an increase in feeding trips with increasing brood-size which is less than i i i proportional. As this does not appear to be enough to explain the ability of parent Double-crests to feed supernormal broods it was hypothesized that these parents return with more food. Several other aspects of feed• ing frequency were examined and found consistent with this hypothesis. Study of the success of individual feedings indicated no tendency for a higher success level in normal broods. The strain on parents raising large broods in both species was shown by the decreasing nest-site attend• ance as broods get larger and as the chicks grow older. The high brood-rearing capabilities of the Double-crests were dis• cussed and two problems on which there was little or no data were raised. It was concluded that Double-crest parents do not endanger their future survivorship in order to raise supernormal broods, although data is poor on this question. A possible breakdown in the feeding of broods greater than 5 may occur under certain conditions. The differing brood- rearing capabilities of the two species were tentatively explained in terms of their population status, i.e., growing versus stable, with the Double-crests the growing population. This idea corresponds with one raised by Lack (1965) who has suggested a correlation between higher than normal brood-rearing capabilities and a growing population. As such, the findings are consistent with Lack's clutch-size hypothesis. The occur• rence of most of the nestling mortality in the first two weeks of the nestling period suggest factors other than food may limit brood-rearing success. iv TABLE OF.CONTENTS Page LIST OF TABLES v LIST OF FIGURES vi Section INTRODUCTION 1 Egg laying and Incubation . 4 MATERIALS AND METHODS 6 RESULTS 8 Fledging success and Chick Production .... 8 Growth Rates .13 Length of Fledging Period 20 Post-fledging Survival ... 22 Feeding Rates , . 26 Nest-site Attendance .35 DISCUSSION 39 LITERATURE CITED 45 V LIST OF TABLES Table Page 1. The Frequency of Clutch-sizes, the Hatching Success, and the Frequency of the Brood-sizes at Hatching in the Pelagic and Double-crested Cormorants 5 2. Number of Broods studied during the 1969 and 1970 Field Seasons 8 3. Chick Mortality in Double-crested and Pelagic Cormorants related to Age and Brood-size 12 4. Growth rates of Individual Chicks, measured in grams per day increase, ranked from Fastest to Slowest Chick 18 5. Mortality of Pelagic Cormorant chicks correlated with Growth Rate 19 6. Weights of chicks from Normal and Supernormal Broods late in their First Week 35 vi LIST OF FIGURES Figure Page 1. The influence of brood-size on fledging success and chick production in the Double-crested and Pelagic Cormorants 10 2. Growth curves from normal broods of Double-crested Cormorant (upper line) and the Pelagic Cormorant (lower line) 14 3. The influence of brood-size on growth rate in Double-crested and Pelagic Cormorants . 16 4. The influence of brood-size on age at fledging in Double-crested Cormorants 21 5. The influence of brood-size on age at fledging in Pelagic> Cormorants 21 6. First year survivorship (upper line) and progeny alive at one year (lower line) in Double-crests as a function of brood-size at fledging 25 7. Daily feeding trips as a percentage of fourth week feeding trips in Double-crested and Pelagic Cormorants 28 8. Daily brood growth as a function of brood-size in Double-crested and Pelagic Cormorants 28 9. Daily number of feeding trips as a function of brood- size in Double-crested and Pelagic-Cormorants 30 10. Daily number of feeding trips per chick as a function of brood-size in Double-crested and Pelagic Cormorants 30 11... .The. influence ..of. brood-size on the number of individual feedings per feeding trip in Double-crested and Pelagic,Cormorants 33 12. The influence of brood-size on the number of feedings per day per chick in Double-crested and Pelagic Cormorants 33 vii Figure Page 13. Frequency of unsuccessful feedings as a function of age in normal and supernormal broods of the Double-crested Cormorant 34 14. Nest-site attendance in the Double-crested Cormorant as a function of age and size of brood . .37 15. Nest-site attendance in the Pelagic Cormorant as a function of brood-size 38 ACKNOWLEDGMENTS I am deeply indebted to my research advisor, Dr. R. H. Drent, whose advice and encouragement were a con• stant inspiration during the course of this study. Dr. J. F. Bendell and Dr. J; M. Taylor kindly read the manu• script and offered many helpful suggestions. For permission to carry out the study at Mandarte Island I am thankful to the East Saanich band of Indians. On the island my work load was eased considerably by the help of a number of colleagues and research assistants. These include A. F. Koelink, J. 0. Anvik, N. Aitchison, J. 6. Ward, B. A. Henderson, and C. Inkster. To the Inksters and Mrs. R. Mathews of Sidney I wish to extend my appreciation for their hospitality. This study was supported by a Canadian Wildlife Service Research contract and by National Research Council Grants to.Dr. Drent. THE INFLUENCE OF BROOD-SIZE ON REPRODUCTIVE SUCCESS IN TWO SPECIES OF CORMORANT, Phalacrocorax auritus AND P. pelagicus, AND ITS RELATION TO THE PROBLEM OF CLUTCH-SIZE INTRODUCTION This study investigates the role played by brood-size in the reproductive success of birds which feed their young (nidicolous birds). The purpose of this inquiry is to gain understanding of the relation between clutch-size and brood-rearing success at various brood-sizes, including experimentally enlarged broods. According to Lack's theory, the broodrsize contributing the most progeny to the breeding population should correspond to the most common clutch-size (Lack, 1954). Presum• ably, this is the result of selection for the clutch-size which gives rise to the most reproducing offspring. The theory predicts that broods larger than normal, either artificially enlarged or natural, will not produce more reproducing progeny than that produced by the normal brood- size^). This means that even if the large broods produce more fledged offspring than normal, post-fledging mortality will be higher in the former thus eliminating the original advantage of the large broods. This is precisely what Lack (1948) found in his study of the Starling, Sturnus vulgaris. In the Great Tit, Parus major, the large broods pro• duce' even fewer progeny than"normal'broods (Lack, Gibb, and Owen, 1957). Only in the Glaucous-winged Gull, Larus glaucescens, is there, evidence that supernormal broods do not suffer a higher post-fledging mortality than normal (Vermeer, 1963). Owing to the difficulties in obtaining reliable estimates of post- fledging survival , studies on the clutch-size problem have been largely restricted to the incubation and nestling periods. Most of these studies 1 2 have shown that the brood-size producing the most fledged offspring cor• responds to the most common clutch-size, even when a sample of broods is experimentally enlarged (Rice and Kenyon, 1962; Nelson, 1966a; Huntington, in Lack, 1966; Harris, 1966; and Summers, 1970).
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