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Hybridization in North Island Tree Weta Copyright is owned by the Author of the thesis. Permission is given for a copy to be downloaded by an individual for the purpose of research and private study only. The thesis may not be reproduced elsewhere without the permission of the Author. Hybridization in North Island Tree Weta Natasha Mckean A thesis submitted in fulfilment of the requirement for the degree of Master of Science in Zoology Massey University, March 2014 Abstract Abstract Hybridization has important implications for species concepts, understanding the speciation process, as well as the issue of sister species coexistence and conservation. Tree weta (genus Hemideina) are an interesting natural system for studying hybridization, as some species have multiple chromosome races that can interbreed, and all are broadly parapatric with at least one other species in the wild, so there are many opportunities for natural hybridization to occur. It is not known in many cases whether species pairs are hybridizing in the wild, or whether introgression occurs. This study focused on the interactions between H. thoracica and its two neighbouring species; H. crassidens and H. trewicki. Surveys of one area of sympatry between H. thoracica and H. crassidens, and one between H. thoracica and H. trewicki, were conducted to see if the parent species and putative hybrids could be easily distinguished, and to find out the relative ratios of the parents and putative hybrid individuals. Weta from the parent populations were studied in areas where they are sympatric as well as allopatric to look for evidence of possible divergence and/or introgression in sympatry. These studies showed that where these species pairs are sympatric, parent forms were predominant, with few morphological intermediates, despite parent species existing in similar proportions. Hemideina thoracica and H. trewicki differed in sympatry regarding both size and possibly life history, with H. thoracica females being larger and both sexes maturing later than H. trewicki. Hemideina thoracica and H. crassidens showed possible evidence of introgression, but no evidence of divergence in sympatry. Karyotypes, a mitochondrial locus, and eight nuclear loci were examined for evidence of introgression between the species pairs in sympatry. All putative hybrids (morphological intermediates) from both species pairs were found to be genetic hybrids, with strong evidence of being F1 hybrids. No evidence was found for introgression of karyotypes or of mitochondrial haplotypes in either case. No evidence of introgression was found at nuclear loci for H. thoracica and H. trewicki. However, H. thoracica and H. crassidens showed some overlap at nuclear markers in sympatry, suggesting a low level of introgression. There was also a sex-bias in the production of F1 hybrids, with most having a H. crassidens mother. Hemideina thoracica appears to interact differently with its two neighbouring species; H. crassidens and H. trewicki. Hemideina thoracica and H. trewicki appear to be reproductively isolated, and are possibly exploiting different niches. Hemideina thoracica and H. crassidens by contrast, showed no evidence of divergence and are presumably dealing with strong interspecific competition, as well as introgression where they meet. These two species are unusual in maintaining a bimodal hybrid zone in the apparent absence of assortative mating. They also contrast with Haldane’s rule, as F1 males have some level of fertility, while females are likely infertile. A sex-bias in the production of F1 hybrids may be due to ‘sexual exclusion’, and so could possibly provide an explanation of how H. thoracica has managed to displace H. crassidens from much of its former range. II III Acknowledgments Acknowledgements I would like to thank all of those who have helped me throughout the course of this project. Firstly, thanks to my supervisors Dr Mary Morgan-Richards & Dr Steve Trewick for guidance, feedback and enthusiasm. Thanks to my grandmother Ruth Mckean and other family and friends for support and encouragement. Thanks also to Shaun Neilson, Dr Mariana Bulgarella, Dylan Anderson, Emily Koot, Phoebe Donovan, Rose Gregerson, Lorraine Peoples, Tamera Clapperton and Ellen Williamson for help collecting live weta and field surveys, as well as Anne Kim and Louisa Sivyer for help feeding and caring for live weta. Dr Mariana Bulgarella for weta samples and helpful advice, Dr Priscilla Wehi and Niki Minnards for data on weta morphology, Dr Rashmi Kant for positive control samples for work on Wolbachia, and Dr Edwina Dowle for weta mitochondrial genomes and help with mapping against Wolbachia. Paul Barrett, Tracy Harris, Cleland Wallace and Trish McLenachan for equipment and help in the lab, Rachel van Heugten, Victoria Twort and Briar Smith for microsatellite and nuclear sequence primers, and Lizzie Daly and Michael Gemmell for their help with software. IV V Contents Contents Abstract .................................................................................................................................................. II Acknowledgements............................................................................................................................... IV List of figures .......................................................................................................................................... X List of tables ........................................................................................................................................ XIV Chapter 1: Introduction ......................................................................................................................... 1 1.1.1 Hybridization ....................................................................................................... 1 1.1.2 Hybrid zones........................................................................................................ 2 1.1.3 Selection for divergence ..................................................................................... 3 1.1.4 Species concepts ................................................................................................. 3 1.1.5 Conservation ....................................................................................................... 4 1.1.6 Tree weta ............................................................................................................ 4 1.1.7 Aims ..................................................................................................................... 6 1.2 References ........................................................................................................................................ 7 Chapter 2: Tree Weta in Sympatry ...................................................................................................... 13 2.1 Introduction ...................................................................................................................... 13 2.1.1 Issues with sympatry ........................................................................................ 13 2.1.2 Patterns in sympatric populations ................................................................... 13 2.1.3 Tree weta .......................................................................................................... 14 2.1.4 Aims ................................................................................................................... 16 2.2 Methods ............................................................................................................................ 17 2.2.1 Study sites ......................................................................................................... 17 2.2.2 Identification of species .................................................................................... 17 2.2.3 Ratio of parent species and hybrids ................................................................. 19 2.2.4 Morphological Characters ................................................................................ 19 2.3 Results ............................................................................................................................... 23 2.3.1 Ratio of Parent Species and Hybrids in Sympatry ........................................... 23 2.3.2 Colouration ....................................................................................................... 26 2.3.3 Prolateral hind tibia spines ............................................................................... 28 2.3.4 Size differences ................................................................................................. 29 2.3.5 Stridulatory ridges ............................................................................................ 30 2.4 Discussion .......................................................................................................................... 31 2.4.1 Morphological characters for the parent species and putative hybrids ......... 31 2.4.2 Ratio of parent species ..................................................................................... 32 2.2.3 Hemideina thoracica and Hemideina trewicki ................................................. 32 2.4.4 Hemideina thoracica and Hemideina crassidens ............................................. 33 2.4.5 Reproductive character displacement ............................................................. 34 2.4.6 Conclusion ........................................................................................................
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