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Effects of Insularity and Species Interactions Ec Beak shape variation in a hybrid species: effects of diet, insularity and species interactions Laura Piñeiro Fernández Master of Science Thesis 2015 Centre for Ecological and Evolutionary Synthesis Department of Biosciences Faculty of Mathematics and Natural Sciences University of Oslo, Norway © Laura Piñeiro Fernández 2015 Beak shape variation in a hybrid species: effects of diet, insularity and species interactions Laura Piñeiro Fernández http://www.duo.uio.no/ Trykk: Reprosentralen, Universitetet i Oslo. Beak shape variation in a hybrid species: effects of diet, insularity and species interactions Acknowledgments This thesis was written at the Centre for Ecological and Evolutionary Synthesis (CEES) at the Department of Biosciences, University of Oslo, under the supervision of Professor Glenn-Peter Sætre, Dr. Fabrice Eroukhmanoff and Dr. Anna Runemark. Glenn , I am really thankful to you from the day you decided to accept me as a student in the sparrow group. It has been a great pleasure to work with you in such an incredible environment. Thanks for all the suggestions, notes and tips during this process. Fabrice and Anna , I have no words to describe your amount of patient with me! Your doors were always open when I needed to go in with a new question. I couldn’t ask for a better team of supervisors, it is amazing how much I’ve learned from both of you, and how I’ve enjoyed your thoughts and comments. This thesis exists thanks to both of you; your 24/7 e-mail presence (I really thought at some point you would have just ended up blocking me) Statistics would still be my worst nightmare without your help. The rest of the Sparrow group, you have been the best team to work do research with. Thank you for the feedback and for being always so willing to help. My costudents and friends in Oslo, I can’t imagine this without you… we made it this far, and here we are! Shane, you have made every day easier, funnier, and even sweeter with all the ice-creams and chocolate! Katie , you’ve been more than a friend, a non-biological sister! Thanks for the meals, gossips and the talks, the trip to Spain, every single thing. Nacho “¡qué bueno que viniste!” Los cafés, meriendas y fútbol me salvaron más tardes de las que estoy dispuesta a admitir, y diciendo cosas sin sentido me ayudaste a no perder la poca cordura que me queda. A mis amigos, siempre estáis ahí, desde el momento que dije que me iba hasta ahora, con los brazos abiertos y una 1906 cada vez que volví de visita fugaz. Y por último, a mi familia, por el apoyo incondicional cada vez que esta cabeza loca decide emigrar a otro país. En especial a mi ahijado Bruno y a Alba , gracias por los ánimos, risas y sonrisas . Y por supuesto mis padres , no tengo palabras para daros las gracias, si no fuera por vosotros yo no sería quién soy ni estaría donde estoy ahora. Gracias por creer que podría hacerlo, por animarme siempre a hacer lo que yo quisiera, a volar y a descubrir. Gracias por aguantar mis momentos de estrés cuando pensaba que ya no podía más. Gracias por emocionaros conmigo. No podría tener una familia mejor. I II Abstract Beak morphology is an ecologically important trait for birds. Beak size and shape are crucial for adapting to new niches and for optimal utilization of existing food sources. The adaptive potential of a species can be limited by the amount of standing genetic variation and by genetic and ontogenetic constraints. Organisms of mixed ancestry, however, can be hypothesized to have a particularly high adaptive potential due to more genetic variation and the breakup of genetic constraints. Here, I take advantage of the homoploid hybrid origin of the Italian sparrow ( Passer italiae ), which resulted from hybridization between the Spanish sparrow ( P. hispaniolensis ) and the house sparrow (P. domesticus ), to study aspects of local adaptation in beak morphology. I investigate how beak shape differs from that of its parent species and which factors best explain beak shape variation in this species. I found significant differences in both beak size and shape between Passer sparrow populations within and between species. I also found that local diet, as estimated from stable isotopes, is a strong predictor of beak shape variation in the Italian sparrow, suggesting that the beak of the Italian sparrow readily adapts to local conditions. Further, comparing island and mainland populations of both Italian and Spanish sparrows, I found that insularity is another important factor affecting the size and shape of the beak in a parallel fashion in both species. Finally, I investigated possible effects of species interactions by comparing diet and beak morphology of Italian sparrows living sympatrically with Spanish sparrows with nearby allopatric ones. The beak of micro-allopatric Italian sparrows did not differ in size or shape from Spanish sparrows and they had greatly overlapping diets. In contrast, in sympatry Italian sparrows had a significantly different diet and beak shape supporting the hypothesis that sympatric Italian sparrows have adapted to a different niche to escape competition from its parent species (i.e. character displacement). In conclusion, the Italian sparrow appears to have a highly adaptable beak, and beak shape variation seems to be crucial for adaptation to novel niches and environments including niche shifts induced by a closely related competitor. III IV Table of contents ACKNOWLEDGMENTS ............................................................................................... I ABSTRACT .................................................................................................................. III INTRODUCTION .......................................................................................................... 1 MATERIALS AND METHODS ................................................................................... 7 BEAK MORPHOLOGY DIFFERENCES AMONG PASSER SPARROWS ....................................... 8 FACTORS EXPLAINING BEAK MORPHOLOGY IN A HYBRID SPECIES ................................... 9 EFFECTS OF INSULARITY ON BEAK MORPHOLOGY .......................................................... 11 INTERACTIONS BETWEEN THE HYBRID SPECIES AND ONE OF ITS PARENTAL SPECIES ...... 11 RESULTS ...................................................................................................................... 13 BEAK MORPHOLOGY DIFFERENCES AMONG PASSER SPARROWS ..................................... 13 FACTORS EXPLAINING BEAK MORPHOLOGY IN A HYBRID SPECIES ................................. 16 EFFECTS OF INSULARITY ON BEAK MORPHOLOGY .......................................................... 20 INTERACTIONS BETWEEN THE HYBRID SPECIES AND ONE OF ITS PARENTAL SPECIES ...... 24 DISCUSSION ................................................................................................................ 27 BEAK MORPHOLOGY DIFFERENCES AMONG PASSER SPARROWS ..................................... 28 FACTORS EXPLAINING BEAK MORPHOLOGY IN A HYBRID SPECIES ................................. 30 EFFECTS OF INSULARITY ON BEAK MORPHOLOGY .......................................................... 32 CHARACTER DISPLACEMENT IN PARENTAL SYMPATRY .................................................. 33 REFERENCES ............................................................................................................. 37 TABLES ........................................................................................................................ 45 APPENDIX ................................................................................................................... 57 FIGURES ........................................................................................................................ 57 TABLES ......................................................................................................................... 63 V VI Introduction “Natural Selection almost inevitably causes much extinction of the less improved forms of life and induces what I have called divergence of character” (Darwin 1859 (2004) p. 12). In 1859, Charles Darwin explained the principle of evolutionary divergence in On the Origin of Species , in which he postulates that the origin of species is the result of divergence of characters among plants and animals as a response to natural selection. More than a century later, speciation, defined as the emergence of reproductive barriers to gene flow between taxa (Coyne & Orr 2004), is still considered a major topic in evolutionary biology (Butlin et al. 2012). The role of ecological factors in the speciation process has received much attention in recent years. In birds, the most diverse terrestrial vertebrate group, one of the factors that has contributed to the diversity of niches they occupy is adaptive variation in beak morphology (Mallarino et al. 2012). Variation in beak size and shape has been shown to be important not only in husking food, but it also affects handling time (Benkman 1993; Benkman 2007). Beak shape can also affect song (Derryberry et al. 2012) and since song is subjected to sexual selection in many birds, beak morphology as a result be a target of sexual selection (Huber & Podos 2006). A classical example of beak morphology adaptation is the adaptive radiation of Darwin’s finches on the Galapagos Islands, where species differ dramatically in beak shapes and feeding niches. During his journey to the Galapagos Islands on the Beagle, Darwin noticed how a number of finch species exhibited variation in beak morphology,
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