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Det Här Verket Är Upphovrättskyddat Enligt Lagen (1960:729) Om Upphovsrätt Till Litterära Och Konstnärliga Verk 0 CM 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 12 11 10 9 8 7 6 5 4 3 2 1 0 INCH - - Lagen (1960:729) om upphovsrätt till litterära litterära till upphovsrätt (1960:729) om Lagen Lagen (1960:729) om upphovsrätt till litterära och och litterära till upphovsrätt (1960:729) om Lagen © It has been digitized with support of Kap. 1, 16 § första stycket p 1, for p 1, for stycket 1, 16 § första Kap. of been has digitized support with It . Det har digitaliserats med stöd av Kap. 1, 16 § första stycket p 1, för forsk p 1, för stycket 1, 16 § första Kap. av med stöd digitaliserats . Det har This work is protected by Swedish Copyright Law ( Law Swedish Copyright by protected is work This verk). konstnärliga och of copyright the consent without public to the no be dissiminated purpose,may and scientific holder. means This text. readable machine to converted and been OCR-processed have texts All printed to hard books are printed early Some the document. from text copy and search can you that it visually compare always should so one errors, contain may the text and correctly OCR-process correct. is what determine to the images with Det här verket är upphovrättskyddat enligt upphovrättskyddat är verket Det här verk konstnärliga upphovsrättsinehavarens till utan allmänheten vidare spridas inte och får ningsändamål, medgivande. söka och kopie kan du till Det text. betyder att maskinläsbar Alla OCR-tolkade tryckta är texter OCR-tolka med dåligt att tryck svåra vara kan dokument äldre Vissa från dokumentet. ra texten visuellt man bör och därför innehålla fel kan texten den OCR-tolkade att vilket medför korrekt riktigt. är vad som avgöra att för bilder med verkets jämföra Sexual selection before and after copulation in water striders Ingela Danielsson GÖTEBORG UNIVERSITY 2002 GÖTEBORGS UNIVERSITET Naturvetenskapliga fakulteten Sexual selection before and after copulation in water striders Ingela Danielsson AKADEMISK AVHANDLING som för avläggande av filosofie doktorsexamen i ekologisk zoologi, enligt Naturvetenskapliga Fakultetens beslut, kommer att offentligen försvaras fredagen den 26 april 2002, kl 10.00 i föreläsningsalen, Zoologiska institutionen, Medicinaregatan 18, Göteborg. Avhandlingen försvaras på engelska. Examinator är Professor Malte Andersson. Fakultetsopponent är Professor Leigh W. Simmons, Department of Zoology, The University of Western Australia, Nedlands WA 6907, Australia. Zoologiska Institutionen Göteborgs Universitet 2002 Danielsson, Ingela (2002) Sexual selection before and after copulation in water striders Animal Ecology, Department of Zoology, Göteborg University, Box 463, SE 405 30 Göteborg, Sweden. E-mail: [email protected] Abstract Post-copulatory sexual selection is now known to occur in most animal groups. Our knowledge of the causes and effects of variation in the relative fertilization success of the competing males is, however, still limited. Crucial questions concern which male traits are favoured by post-copulatory sexual selection, and whether such selection reinforces or counteracts pre-copulatory sexual selection. I address these questions by studying relative fertilization success among male water striders (Gerris lacustris and G. lateralis). There is large variation in paternity for two competing males, mated sequentially with the same female. Paternity is, however, repeatable across replicate females, showing that males differ consistently in their ability to achieve fertilization success. By analysing shape variation in male genital morphology, a relationship was discovered between the shape of male intromittent genitalia and relative paternity success. This result provides experimental support for the hypothesis that male genitalia evolve via post-copulatory sexual selection. A detailed analysis revealed that different aspects of male genitalia affect different fitness components. Certain traits affected male ability to obtain fertilization success while others affected male ability to avoid sperm precedence by subsequent males. The effect of the male genitalia shape varied with female morphology, suggesting that selection on male genitalia depends on the frequency distribution of female phenotypes. Male ejaculate size and fertilization success increased with copulation duration, suggesting that males are able to increase their relative fertilization success by transferring more sperm. Males appear to be sperm limited however, and transferred significantly less sperm after a short than after a long recovery period. Thus males apparently need some time to replenish sperm stores between matings, a constraint with implications for sexual selection of male body size. Mating is costly for females and they are generally reluctant to mate. This sexual conflict over mating generates directional pre-copulatory sexual selection for large males. When large and small males were allowed to compete for fertilizations, large males mated more often, but small males copulated longer and gained higher fertilization success per mating by transferring more sperm. Shorter copulations by large males were probably a result of a behavioural trade-off between mating frequency and ejaculate size caused by sperm limitation. The result of this antagonistic effect of pre- and post-copulatory sexual selection was that large and small males had similar reproductive success. Although large males had a clear mating advantage they had no fertilization advantage, and there appears to be no net selection for increased male size. Key words: sexual selection, sperm competition, cryptic female choice, fertilization success, mating success, evolution of genitalia, body size, copulation duration, ejaculate size, sperm transfer, Gerridae, Gerris, water striders Göteborg University 2002 ISBN 91-628-5164-0 Thesis for the Degree Doctor of Philosophy Sexual selection before and after copulation in water striders Ingela Danielsson GÖTEBORG UNIVERSITY Faculty of Science Animal Ecology, Department of Zoology Box 463, SE 405 30 Göteborg Sweden Göteborg, April 2002 The oral defence of this thesis will take place Friday 26 April 2002, at the Department of Zoology, Medicinaregatan 18 Göteborg, Sweden. Faculty opponent is Professor Leigh W. Simmons, Department of Zoology, The University of Western Australia, Nedlands WA 6907, Australia. Sexual selection before and after copulation in water striders Ingela Danielsson The author's address: Animal Ecology, Department of Zoology Göteborg University Box 463, SE 405 30 Göteborg, Sweden E-mail: [email protected] © 2002 Ingela Danielsson Parts of this thesis are reprinted with permissions from the publishers: Figure 1-3 in the thesis summary, by Brill Academic Publishers Paper 1, by Finnish Zoological and Botanical Publishing Board Paper 2, by Springer-Verlag Paper 3, by The Society for the Study of Evolution Paper 4, by Oxford University Press Paper 5, by The Royal Society ISBN 91-628-5164-0 Printed in Göteborg, Sweden by Kompendiet Cover picture by Jens Rydell & Ingela Danielsson, Layout by Eric Järpe Danielsson, Ingela (2002) Sexual selection before and after copulation in water striders Animal Ecology, Department of Zoology, Göteborg University, Box 463, SE 405 30 Göteborg, Sweden. E-mail: [email protected] Abstract Post-copulatory sexual selection is now known to occur in most animal groups. Our knowledge of the causes and effects of variation in the relative fertilization success of the competing males is, however, still limited. Crucial questions concern which male traits are favoured by post-copulatory sexual selection, and whether such selection reinforces or counteracts pre-copulatory sexual selection. I address these questions by studying relative fertilization success among male water striders (Gerris lacustris and G. lateralis). There is large variation in paternity for two competing males, mated sequentially with the same female. Paternity is, however, repeatable across replicate females, showing that males differ consistently in their ability to achieve fertilization success. By analysing shape variation in male genital morphology, a relationship was discovered between the shape of male intromittent genitalia and relative paternity success. This result provides experimental support for the hypothesis that male genitalia evolve via post-copulatory sexual selection. A detailed analysis revealed that different aspects of male genitalia affect different fitness components. Certain traits affected male ability to obtain fertilization success while others affected male ability to avoid sperm precedence by subsequent males. The effect of the male genitalia shape varied with female morphology, suggesting that selection on male genitalia depends on the frequency distribution of female phenotypes. Male ejaculate size and fertilization success increased with copulation duration, suggesting that males are able to increase their relative fertilization success by transferring more sperm. Males appear to be sperm limited however, and transferred significantly less sperm after a short than after a long recovery period. Thus males apparently need some time to replenish sperm stores between matings, a constraint with implications for sexual selection of male body size. Mating is costly for females and they are generally reluctant to mate. This sexual conflict over mating generates directional pre-copulatory sexual selection for large males. When large and small males were allowed to compete for fertilizations, large males mated more often,
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