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Of Dippers Cinelus Einelus

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Of Dippers Cinelus Einelus I, Population ecology and lifetime reproductive success of dippers Cinelus einelus Thesis submitted for the degree of Doctor of Philosophy John W. Logie Avian Ecology Unit, Department of Biological and Molecular Sciences, University of Stirling. May 1998. Abstract Acidified catchments are known to hold significantly reduced dipper Cinclus cinclus populations throughout the year relative to circum-neutral rivers, although the processes leading to these declines remain unclear. This study considered the population ecology of dippers within the circum­ neutral River Devon catchment, Central Scotland, and focused primarily on determining the factors influencing survival, breeding probabilities and reproductive success. It aimed to examine the role of spatial variation in 'habitat quality' on the population (and meta-population) dynamics of dippers, based on measures of seasonal and lifetime reproductive success, and the balance between survival and reproduction; in particular, to assess if the reduced reproductive success of dippers on acid rivers is likely to lead to population declines. Within the Devon catchment, approximately 81 % of all adults survived from spring (March/April) to autumn (September/October), with 65% of these birds surviving from autumn to the following breeding season. Overall, these estimates predicted annual adult survival rates of c.53%, with no significant differences between years. Population density had no detectable effect on adult mortality rates, although juvenile over-winter survival was significantly lower than the adult rate at between 40 and 58%, and negatively related to the total size of the autumn population. There was no evidence of sex differences in juvenile over-winter survival, or any significant influence of weather or river flows on the rates for adults or juveniles. The local post-fledging survival of females was significantly lower than for males, however, apparently reflecting sex differences in post-natal dispersal. On average, less than 6.5% of all eggs laid, or 10.4-14.5% of male and 6.3-9.2% of female fledglings raised within the Devon catchment survived locally to breeding age. Juvenile, although not adult, recapture rates in spring were significantly lower than for birds known to have bred previously and negatively related to spring river flows. This suggested that with recapture dependent on a breeding attempt that was successful at least until laying, either more first year birds failed during the initial stages of nesting or that full breeding was not achieved at age one. The birds fledging the most young, both within a season and over a lifetime, all bred at 'prime' lowland sites characterised by wide, shallow rivers of intermediate gradient, although with less than 10% of all birds attempting to raise a second brood each year, no significant habitat differences were identified in any component of reproductive output measured until fledging. River width, altitude and gradient were all significantly inter-correlated and related to laying date, however, and post-fledging survival was significantly reduced for late fledged young. On average, lowland birds laid earlier than upland breeders, and were significantly more likely to produce autumn 'recruits' due to the enhanced post-fledging survival prospects of their young. This suggested that broad measures of river structure can provide a biologically appropriate classification of habitat qUality. The size of the breeding population of dippers within the Devon catchment appeared to be related to the availability of critical resources, most likely food, roost sites and ultimately breeding territories through density-dependent changes in over-winter mortality and recruitment. The relative importance of resource abundance and recruitment levels in determining autumn population densities on acid streams still remained unclear, although reference to published relationships between acidity and reproductive success suggested that with adult survival at the rate estimated for the Devon catchment, many dipper populations are unlikely to produce sufficient recruits to match all adult losses, and may only persist with continued immigration from more productive (circum­ neutral) catchments elsewhere. Acknowledgements Before a trip to Paris, I asked Dave Bryant if he could get hold of some computer software, so that I could look at it, and when I arrived, I wouldn't seem an like idiot. In true professorial style, he replied 'Idiot? Hmmmmm ... I don't think there's any easy way round it!'. What follows is a 200 page testament to that, but I think, Dave, we just about got a way with it, and now seems an appropriate time to name all those responsible for getting me where I am today. Special thanks are due to David Bryant, Steve Newton and Anne Newton for collecting data in the years prior to 1992, and giving complete access to all their unpublished work. The Charles and Barbara Tyre Trust, Argyll, provided funding and paid my fees throughout. Without their generosity and goodwill, this work would not have been possible. Jean Clobert and Francois Sarrazin, at the Institute D'Ecologie, provided excellent training in the use of SURGE. To Dave Bryant. Shortly, after I started my PhD, you were promoted to Professor. Coincidence? Who can say? But Dave, I'd like to think that I played no small part in forming and spreading your reputation around the department. Throughout the years, you always gave excellent advice, whether I liked it or not, and never was a combination of faith, hope and charity more evident than during my time in Stirling. Without the unique Bryant funding policy, I wouldn't have had two pennies to rub together, but with it, I did have two pennies to rub together. I know you're not one for excessive praise, so in true Bryant-speak, I can only say that, as a supervisor, you were "probably good enough". In fact, you were much more, and I'm grateful for your help and advice over the years, but I'm not prepared to say that. To everyone in the Ecology Group, past and present, thanks for your valuable, and often invaluable advice. Robert, you cheered me with sunny disposition, infectious enthusiasm and jaunty haircut. In the early years, Maimie, Ross, Lulu, Sari, Rosli, Chris and Ian, and later Karen, Louise, Mandy, Sue, Laura and Brian cheered my just by being my paaaais. And Brian, long after you've cropped your waist-length dreadlocks and slipped back into middle-class conformity, I'll think of you, when ever I see a wet dog. Special thanks also to Su for all those carefully choreographed attempts to make me look good, including setting fire to your Goretex jacket halfway down Merryhills Culvert and pretending you couldn't tell the difference between a dipper and a wet sock. On the subject of making me look good, thanks too, lase, for that Raptor Research paper. In reply to Godfrey, 1998 (or whenever), lase, repetitive insults? Never! In spite of your unerring ability to immediately find the slightest fault in 200 pages of text, I'd say I let you off easy! So Jase, the split infinitive in the last sentence, and all other errors throughout the text, are there just for you. Special thanks also to Graham Dyson and Andrew Hall (postgraduate research's answer to Dougal and Father Jack) for creating a monster. You both provided valuable empirical evidence on the relationships between body size, energetic demand and reproductive "success", a legacy that will live on long after you've gone. Thank you (I think)! In spite of persistent references to the time it took me to finish, 1 hope you all know that when your time comes, I'll be available to give good advice (see my fact sheet, 'How to write a general discussion in three easy years'). To everyone that came and went at 55 King Street. Don, mate. You've got to hit the target, but you're never going to trouble the keeper from there. The boy done well. And Rosh. Although, you frightened me (a lot), I loved you all the same. Thanks also for confirming that the ultimate kitchen sin really does involve rubber gloves and warm soapy water. Finally, thanks to Catherine, Kevin, Vivien et al. in the Anxiety and Stress Research Centre, and everyone involved with the Prison Survey for putting things into some kind of perspective. Five years for a PhD? I know now I'd get less than that for armed robbery. He is a singularly joyous and loveable little fellow. He is the mountain streams ' own darling, the humming bird of the blooming waters, loving rocky ripple slopes and sheets offoam as a bee loves flowers, as a lark loves sunshine and meadows. Among all the mountain birds, none has cheered me so much in my lonely wanderings, none so unfailingly. For both in winter and summer he sings, sweetly, cheerily, independent alike of sunshine and of love, requiring no other inspiration than the stream on which he dwells. While water sings, so must he, in heat or cold, calm or storm, ever attuning his voice in sure accord; low in the drought of summer and the drought of winter, but never silent. John Muir (1894). Contents Chapter 1 Introduction 1 1.1. Introduction 1 1.1.1. Stream acidity and dipper populations 5 1.1.2. Factors likely to determine popUlation size in dippers 7 1.2. Aims 8 1.3. The suitability of dippers for population and 'lifetime' studies 10 1.3.1. The study history and previous work on the dippers of the Devon catchment 11 1.3.2. The study species 13 1.4. Thesis organisation 14 Chapter 2 General Methods 16 2.1. The study sites 16 2.1.1. Upland and lowland sites 18 2.2. General field methods 21 2.2.1. Autumn census techniques 21 2.2.2.
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