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The Systematics of the Freshwater Crabs (Decapoda: Brachyura: Potamidae) of Southern China, and Freshwater Zoogeography of China

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The Systematics of the Freshwater Crabs (Decapoda: Brachyura: Potamidae) of Southern China, and Freshwater Zoogeography of China The systematics of the freshwater crabs (Decapoda: Brachyura: Potamidae) of southern China, and freshwater zoogeography of China Chao Huang Supervisors: Malte C. Ebach, Shane T. Ahyong, Shawn W. Laffan A thesis in fulfilment of the requirements for the degree of Doctor of Philosophy School of Biological Earth and Environmental Sciences Faculty of Science 1 February 2020 2 3 4 5 Tiwaripotamon sp., a miniature undescribed species from the karstic forests of Guangxi, China. 6 Preface This thesis is a compilation of my works during the course of my PhD candidature, conducted under the guidance of my supervisors Malte C. Ebach, Shane T. Ahyong and Shawn W. Laffan. It consists of one introductory chapter which is partly derived from a book chapter (Chapter 1), six published research papers (Chapters 2-7) and a summary chapter (Chapter 8). The research paper chapters are presented as they would appear in a journal except the acknowledgements and references have been moved to the start and end of the thesis, respectively. The introduction of the research paper in chapter 8 has been moved under chapter 1. I conceptualized the research, conducted field collections, gathered data, performed analyses, wrote the manuscripts and prepared the tables and figures. The contributions of my supervisors/co-authors are as follows: Chapter 1: This Chapter is partially derived from my part of the book chapter S. T. Ahyong & C. Huang (2020). Colonization, adaptation, radiation, and diversity in fresh water. In: Thiel, M. & Poore, G.C.B. (eds.), The Natural History of the Crustacea, Volume 8, 319: Evolution and Biogeography. Oxford University Press. STA provided some reference material and gave advice in the preparation of the manuscript. Chapter 2: Huang, C., H. T. Shih and P. K. Ng 2017. A new genus and new species of Potamidae (Crustacea: Decapoda: Brachyura: Potamoidea), the first stygomorphic cave crab known from China and East Asia. Zootaxa, 4232 (1). HTS conducted the phylogenetic analysis and contributed to the “DNA analyses and discussion” part of the paper. PKN and STA gave advice in interpretation of data, morphological comparison and in the preparation of the manuscript. Chapter 3: Huang, C., S. T. Ahyong and H. T. Shih 2017. Cantopotamon, a new genus of freshwater crabs from Guangdong, China, with descriptions of four new species (Crustacea: Decapoda: Brachyura: Potamidae). Zoological Studies, 56 (41). STA gave advice in interpretation of results, scientific illustrations and in the preparation of the manuscript. 7 HTS conducted the phylogenetic analysis and contributed to the “phylogenetic analysis” and “discussion” parts of the paper. Chapter 4: Huang, C. 2018. Revision of Yarepotamon Dai & Türkay, 1997 (Brachyura: Potamidae), freshwater crabs endemic to southern China, with descriptions of two new genera and four new species. Journal of Crustacean Biology, 38 (2). HTS and STA gave advice in the interpretation of data and preparation of the manuscript. Chapter 5: Huang, C., H. T. Shih and S. T. Ahyong 2018. Two new genera and two new species of narrow-range freshwater crabs from Guangdong, China (Decapoda: Brachyura: Potamidae). Journal of Crustacean Biology, 38 (5). HTS conducted the phylogenetic analysis and contributed to the “phylogenetic analysis and discussion” part of the paper. STA gave advice in interpretation of data, scientific illustrations and in the preparation of the manuscript. Chapter 6: Huang, C., K. C. Wong and S. T. Ahyong 2018. The freshwater crabs of Macau, with the description of a new species of Nanhaipotamon Bott, 1968 and the redescription of Nanhaipotamon wupingense Cheng, Yang, Zhong & Li, 2003 (Crustacea, Decapoda, Potamidae). ZooKeys, 810. KCW accompanied me in the field collection and contributed some ecological observations. STA gave advice in interpretation of data, scientific illustrations and in the preparation of the manuscript. Chapter 7: Huang, C., S. Z. Huang and Z. X. Shen 2020. A new long-legged terrestrial freshwater crab, Calcipotamon puglabrum n. gen., n. sp. (Crustacea, Decapoda, Potamidae), from Hainan Island, China. Zootaxa, 4766 (3). SZH first discovered the new species and contributed to the “habitat” section of the description. ZXS organized the collaboration and contributed to the “introduction” section. Chapter 8: Huang, C., M. C. Ebach and S. T. Ahyong 2020. Bioregionalisation of the freshwater zoogeographical areas of mainland China. Zootaxa, 4742 (2). MCE gave advice 8 in conceptualizing the research and in the preparation of the manuscript. STA gave advice in the structure and preparation of the manuscript. Disclaimer: The new taxonomic names in this work are disclaimed for nomenclatural purposes and as such are not available from this thesis. 9 Acknowledgements I thank my supervisors, Malte Ebach, Shane Ahyong and Shawn Laffan, for their support and guidance throughout my candidature. I also thank the people in the School of Biological, Earth and Environmental Sciences of UNSW, the Australian Museum and Sun Yat-sen University Biology Museum of SYSU for their support and advice during my research. I thank the Australian Government for providing the tuition fees for my research degree, and also to the Graduate Research School and PANGEA at UNSW for funding some of my overseas field trips and conference attendances. I thank my previous supervisor at SYSU, Jian Rong Huang, and Peter Ng from NUS for initiating me into the field of carcinology. I thank carcinologists Hsi-Te Shih, Xian Min Zhou, Jie Xin Zou, Hong Ying Sun and many others for their support in my research. I also thank my many naturalist/researcher friends who have accompanied me in the field and/or provided information or specimens for me, especially Zhuo Cheng Zhou, who helped me build connections within the Chinese naturalist and researcher community. I am also grateful to the colleagues who have reviewed the papers included in this thesis. Their constructive criticisms have improved the quality of my research. Last but not least, I would like to give thanks to my parents for their many years of love and financial support, to my eight-year- old younger sister whom I strive to be a good role model for and to my loving wife who has always been supportive of me. 10 Abstract Freshwater crabs make up close to 20% of brachyuran diversity with over 1300 species. China is the most species rich country for freshwater crabs with two families, 49 genera and 321 species, yet there are still large areas that are unsurveyed for them. To fill in these blanks, multiple field surveys have been carried out during the last few years which have resulted in the accumulation of new distribution records and the discovery of 13 new species (Calcipotamon puglabrum, Cantopotamon hengqinense, Cantopotamon shangchuanense, Cantopotamon yangxiense, Cantopotamon zhuhaiense, Diyutamon cereum, Luteomon spinapodum, Megapleonum ehuangzhang, Nanhaipotamon macau, Qianguimon elongatum, Qianguimon splendidum, Yarepotamon meridianum, Yarepotamon fossor) and the establishment of seven new genera (Calcipotamon, Cantopotamon, Diyutamon, Eurusamon, Luteomon, Megapleonum, Qianguimon). These new taxa have since been described and are included herein. Freshwater crabs are also good candidates for biogeographical research due to their very limited dispersal abilities. The large scale freshwater zoogeographic bioregionalisation studies of China were revised. In order to study the freshwater zoogeographical areas of mainland China, a dataset of close to ten thousand distributional records of freshwater animals that covers 80% of the amphibians, 60% of the freshwater fish and 90% of the freshwater crab species of mainland China was compiled. According to the results from cluster analysis and network analysis, four subregions, three dominions and five provinces were found, of which one dominion and two provinces were newly proposed. In addition, the endemic areas of each animal group were individually studied and were all found to reflect the bioregionalisation at the subregion level but differed from each other in the dominion and province levels. The first quantitative, multi-taxon based bioregionalisation of the freshwater zoogeographic areas of mainland China is herein proposed. 11 Table of contents Preface ................................................................................................................................... 7 Acknowledgements ............................................................................................................. 10 Abstract ............................................................................................................................... 11 Table of contents ................................................................................................................ 12 Chapter 1. Introduction ..................................................................................................... 15 General introduction to freshwater crabs ......................................................................... 15 Research history of systematics of the Chinese Potamidae ............................................. 18 History of freshwater zoogeographic bioregionalisation of China .................................. 23 Research aims................................................................................................................... 33 Thesis structure ................................................................................................................ 34 Chapter 2. A new genus
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