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Developmental Trade-Offs and Resource Allocation in Caddis Flies

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Developmental Trade-Offs and Resource Allocation in Caddis Flies Stevens, David J. (2000) Developmental trade-offs and resource allocation in caddis flies. PhD thesis http://theses.gla.ac.uk/3764/ 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] Developmental Trade-Offs and Resource Allocation In Caddis Flies. David J. Stevens A thesis submitted for the degree ofDoctor ofPhilosophy to the Faculty of Science, University ofGlasgow. September 2000 Declaration I declare that the work presented in this thesis has been completed by myselfunless otherwise acknowledged in the text. No part ofthis work has been presented for any other qualification. David Stevens September 2000 Acknowledgements I could not have hoped for two better supervisors than Pat Monaghan and Mike Hansell. I wish to thank them not just for giving me my chance in the first place, but for giving me the freedom within the project to explore many avenues ofinvestigation. Most ofthese proved fruitful, ifonly to teach me not to do it again. The support, encouragement and advice they gave were ofimmeasurable help over the course ofthe project. The staffofthe University Field Station deserve special mention for the memorable times I had at Rowardennan. The following (Rhona, Vivian, Caroline, Ishbel and Peter) deserve mention for emptying the light trap, checking nets and cooking chicken curry. Chris Cutts, Nicola Bissett, Mat O'Hare, Scot Ramsey, Sarah Jarvis, Colin Bull, Ian McCarthy, Alistair & Heather Duguid, Jenny Devine, Alan Grant, Alan Campbell and the inimitable Colin Adams, made my days at UPS a lot offun (and also sometimes a lot ofhard work). Everyone who helped me on field-work may well regret it now, but thanks all the same. Kath Sloman, Amanda McLean, Mat Cottam, Alexandra Hay and especially Melanie Fletcher deserve credit for bringing caddis flies to a wider audience Special mention must go to June Freel. I couldn't have done it without her, and her bomb calorimetry, ether extraction and rollerskating skills are a sight to behold. Thanks also to Kim Wilson for the nitrogen analysis and Liz Denton for her patient revision offigures, which were only necessary because ofmy own complete incompetence. John Laurie gave erstwhile support to the mighty Hammers, and it shows. The staffofthe Graham Kerr Building have also been fantastic, whether academic or technical, but special thanks to Ruedi Nager, Neil Metcalfe and Graeme Ruxton for putting up with my stats related questions. Thanks also to Geoff Hancock for identifying the tricky ones . I would also like to thank my fellow inmates ofRoom 517, we all did a four-year stretch and none ofus were let out early for good behaviour. Thanks to Mark Cook, Vicky Heaney, Torn Sawyer, Ali McCluskie, and lately Tom Van Pelt ('cobblers'), Ellen Kalambach and Jae Burn Lee. Whatever happened to Paul though? No acknowledgements section could be complete without thanking Francis and Stuart, who in a strange way helped me keep my sanity (" III miss you guys"). Lunch at Jack McPhees? ...Lots ofothers made my time here memorable, especially Dave Fraser, Ian Morgan, Paul Parslow-Williams, Elvira Poloczanska, Fia Selmer, Jen Andrew, Andrea Aitken, Ben Ross and the shy and retiring Kenny Leung. Although not in Glasgow, the alumni ofthe Durham MSc course 1995/96 played a big part in the whole affair. Special thanks to Mark Eaton, Nicola Brown, Melanie Hunt, Steve Campbell, Nick Wilkinson, Steve Park, Mr and Mrs Ian King and Dave Hole. Kate Arnold did the impossible - she read the whole thing, cooked and helped with the printing. So, its all her fault really. Thanks Kate, I 'really' couldn't have done it without you. Finally I would like to thank my mum, Joyce Stevens, for supporting me through what must have seemed an eternity ofstudent life. The cheque's in the post. ABSTRACT Resource allocation and life history strategy are obviously interconnected for all organisms. Limited resources within individuals lead to allocation trade-offs between fitness traits, and constrain their evolution. Many trade-offs have been characterised, mainly concerning the costs ofreproduction. Trade-offs occurring between body parts during the growth and development oforganisms have received little attention. Such 'Developmental Trade-Offs' between body structures are predicted to have profound effects on an organism's subsequent life history traits. These effects will be particularly significant for organisms where part ofthe life cycle is spent in a non-feeding form. In such complex life cycles, a complete reorganisation ofthe body form occurs, often associated with a dramatic change of habitat. Resources accrued in the feeding life history stages are used to build the body and fuel the activities ofthe non-feeding form. Various factors can lead to variation in the resources available at body reorganisation, which will have a cascade effect into the next life history stage, through the action ofdevelopmental trade-offs and strategic allocation Caddis flies (Order Trichoptera) are one such example of an organism with a complex life cycle. They are holometabolous insects, which means that they undergo a complete metamorphosis, where the aquatic larval form is completely broken down and reconstructed into a terrestrial adult during pupation. Nutrient deficiencies at pupation are predicted to lead to developmental trade-offs, particularly between somatic and reproductive investment. This thesis is an examination ofdevelopmental trade-offs occurring in the pupal stage ofcaddis flies, using both an experimental and correlational approach. Resources available at metamorphosis were manipulated by forcing caddis fly larvae, Odontocerum albicorne, to divert more silk, and hence protein, into their larval cases. The resultant adult forms had shorter wings and lighter thoraces (somatic investment) than control individuals, but abdomen mass (reproductive investment) was not affected. The results ofthis experiment could be due to mechanistic constraints ofthe developmental pathways during pupation, because resources from the breakdown ofcertain structures may be 'earmarked' for incorporation into specific adult structures. Equally, the observed response could represent a strategic preservation ofreproductive reserves. 0. albicorne is a short lived species which mates soon after adult emergence and so sacrificing somatic invetsment may have less drastic affects on fitness. To examine whether resource allocation pathways during development ofthe adult form are flexible in caddis flies, the same resource depletion experiment was performed on the caddis fly species, Glyphotaelius pellucidus. Adult life span is much longer in this species, which is an adaptation to loss oflarval habitat for many months ofthe year. Here, somatic investment would be predicted to be ofmuch greater importance in the life history, and so ifthe response to the nutrient deficiencies was strategic, somatic reserves would be expected to be preserved at the expense ofreproductive potential. The pattern ofresource allocation in the resultant adults was in line with predicted life history requirements, as abdomen mass was sacrificed so as to maintain thoracic mass. Wing length was again shorter, suggesting these patterns were not as a result ofstrategically maintaining flight ability. This study shows that allocation patterns are not fixed, and that differential response to depletion are possible. Life history strategy is poorly known for adult caddis flies, particularly those from temporary habitats. An investigation was undertaken to catalogue the Limnephi1id species collected over a three-year period from a light trap on Loch Lomondside, and to document their flight periods. In total, 1376 individuals from eleven genera were collected. A range ofspecies were recovered between March and December, and species emerged from a variety oftemporary and permanent freshwater larval habitats. Flight periods indicated that life spans ranged from days to months. Eleven ofthe twenty species caught had not been documented in the area prior to this study. A few species are only commonly found in Scotland, and their biology is relatively poorly studied. Many ofthe flight periods differed from those published in the literature as typical for these species. Adults from the first year ofthe light trap study were used to measure resource dynamics within the adult body. Despite the fact that in animals with complex life cycles many non-feeding stages do not grow, they may show some evidence of 'phenotypic flexibility', and are able to reallocate some body resources to other functions during the life span. Evidence from butterflies suggests individuals can break down flight muscle mass and use the nitrogen this liberates in reproduction. Patterns ofthoracic mass change over time (= age) for the males of seven species (and for females from three ofthese) from the light trap indicated the same may occur in caddis flies. Evidence from one species reared in captivity, Limnephilus rhombicus, demonstrates
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