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Osteology, Taxonomy, Phylogeny, and Body Shape Changes of Eocene Catostomid and Problematic

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Osteology, Taxonomy, Phylogeny, and Body Shape Changes of Eocene Catostomid and Problematic Catostomid Fishes by Juan Liu A thesis submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Systematics and Evolution Department of Biological Sciences University of Alberta © Juan Liu, 2016 Abstract Extant catostomid fishes occupy diverse aquatic ecosystems and niches in North America (NA). Less than 3% of catostomid taxonomic richness, or two species, are found outside of NA in Asia. Such adjunct and unbalanced distribution pattern has been established since the late Oligocene, when catostomids disappeared from Asia and became common fishes in NA. Eocene catostomids, representing the oldest fossil records of this family, reached relatively high taxonomic richness in both NA and Asia, and thus are critical to understand evolution of catostomids. This thesis is a study of about 3000 specimens of Eocene catostomids and problematic catostomids. Osteology of 33 extant catostomid species and 11 other otophysans are examined for comparison. Morphological characters are acquired through microscopic examination and CT scans. Cladistic analyses using the parsimony criterion are performed in PAUP 4b10 and TNT 1.5, and explored in MrBayes on XSEDE (3.2.6) using model-based Bayesian inferences through the CIPRES Science Gateway. Analyses of body shape changes and variations of Eocene catostomids are performed using the landmark-based geometric morphometric method. Body shapes of laterally preserved complete fish are captured using surface scanning and digitized with landmarks in TPS utility and TPS dig2, then analysed in MorphoJ 1.06C. First, taxonomy is clarified for Eocene catostomids and problematic catostomids. Of the NA taxa, ii Amyzon gosiutense, a previously hypothesized junior synonym of A. aggregatum, retains its specific status based on a suite of osteological features. “NewGenus” brevipinne is revised from the nominal Amyzon brevipinne and represents the first known shallow bodied, small sized, fluvial-sediment derived, and loach-like catostomid genus. Of the Asian taxa, Jianghanichthys hubeiensis, which was firstly assigned to Osteochilus of Cyprinidae and later Catostomidae, is suggested to be a stem cypriniform based on osteological comparison and phylogenetic analysis. The first fossil family of Cypriniformes, Jianghanichthyidae, is erected to contain the species. Additional problematic catostomids, which were described as Eocene Osteochilus from south China, are assigned to Jianghanichthyidae and combined into two species: Jianghanichthys sanshuiensis and J. linliensis. New species Amyzon kishenehnicum and Jianghanichthys huachongensis are described from NA and Asia, respectively. Second, the systematic position and intrarelationships of Eocene catostomids were largely unresolved before this study, although they are critical to understand the evolution of the family and Cypriniformes. Three sets of phylogenetic analyses based on different character sets are performed. The first set vouches that Jianghanichthys doesn't belong to any known cypriniform family, but is a basal clade. The second set finds that Amyzon is a basal clade of Catostomidae instead of a member in the Ictiobinae. The third suite of analyses is optimized by maximally utilizing Eocene species and creating an osteological character list. These analyses suggest that most Eocene catostomids (except Plesiomyxocyprinus) were stem taxa of Catostomidae, and iii “NewGenus” brevipinne represented the most basal clade. Also, known species of Amyzon are not monophyletic, with Asian A. hunanense and NA A. commune being distinctive basal clades. Another Asian genus Plesiomyxocyprinus is sister to all extant catostomids except ictiobines. Third, Eocene catostomids possess similar general appearances and have highly overlapped meristic and morphometric characters, which caused had taxonomic confusion. Quantitative anatomical comparison in this study (geometric morphometrics) finds significant body shape differences between jianghanichthyids and sampled catostomids, A. aggregatum versus A. gosiutense, and A. aggregatum versus A. kishenehnicum. Regression analysis on Procrustes coordinates and centroid sizes shows that allometry does not significantly affect body shapes differences among well-sampled catostomids and jianghanichthyids. Phylogenetically mapped morphospace indicates evolutionary trajectories and trends of body shape changes are congruent to phylogenetic branches of Eocene catostomids and jianghanichthyids. In sum, all unambiguous oldest cypriniforms are included for discussion (Catostomidae, and Jianghanichthyidae of this study plus Cyprinidae). The only area that all oldest cypriniforms are found is south Asia, which suggests that area is the hotspot of origin and cladogenesis of Cypriniformes. Central Asia, including north China, Mongolia, Kazakhstan, and Fareast Russia, where the last remains of catostomid fossils in Asia recorded cyprinids' dispersal route, is a gateway leading to the dispersal events eastward to NA through Beringia and westward to iv Europe. The mosaic distribution of the Asian taxa across the phylogeny suggests multiple episodes of intercontinental dispersals during the early evolution of catostomids, probably through the freshwater connections of Beringia before the middle Eocene. v Preface This thesis contains published journal articles. Except the introduction and conclusion chapters, Chapters 2 and 3 are published, and Chapters 4 through 7 are original research being prepared for submission to peer-reviewed journals. In addition, five refereed conference abstracts and presentations have derived from projects of this thesis. Chapter 2 of this thesis has been published as Liu, J., M.-M. Chang, M. V. H. Wilson, and A. M. Murray. 2015. A new family of Cypriniformes (Teleostei, Ostariophysi) based on a redescription of †Jianghanichthys hubeiensis (Lei, 1977) from the Eocene Yangxi Formation of China. Journal of Vertebrate Paleontology e1004073:23. I collected the data, designed and performed the analyses, and wrote the paper. Wilson co-designed and discussed the analyses. Chang CT-scanned the specimen. Wilson and Chang provided funding support. All three co-authors (Chang, Wilson, and Murray) provided supervisory and editorial supports. Chapter 3 has been published as Liu, J., M. V. H. Wilson, and A. M. Murray. 2016. A new Catostomid fish (Ostariophysi, Cypriniformes) from the Eocene Kishenehn Formation and remarks on the North American species of †Amyzon Cope. Journal of Paleontology. FirstView: 1–17. I was responsible for the data collection and analysis, as well as the manuscript vi composition. Wilson helped with collecting data, co-designed analysis and provided funding support. The co-authors (Wilson and Murray) collected fossil materials and provided supervisory, editorial, and laboratory equipment supports. The following published abstracts and presentations are derived from or partially from the thesis: Liu J. and Mark V. H. Wilson, 2016. Eocene fossil record of catostomids (Teleostei: Cypriniformes) from North America and discovery of the first loach-like catostomid. Supplement to the online Journal of Vertebrate Paleontology. 76th Annual Meeting Society of Vertebrate Paleontology, Program and Abstracts. Salt Lake City, Oct., 2016: 175. (oral presentation) Liu J., Mark V. H. Wilson, and Alison M. Murray, 2015. Growth rates during ontogeny of early Eocene catostomids from British Columbia, Canada. Supplement to the online Journal of Vertebrate Paleontology. 75th Annual Meeting Society of Vertebrate Paleontology, Program and Abstracts. Dallas, Oct., 2015: 165. (oral presentation) Liu J., Mark V. H. Wilson, Alison M. Murray, and Z. Jack Tseng, 2014. A new sucker (Teleostei, Catostomidae) from the Eocene Kishenehn Formation of Montana and the systematic position of Amyzon. Supplement to the online Journal of Vertebrate Paleontology. 74th Annual Meeting Society of Vertebrate Paleontology, Program and Abstracts. Berlin, Nov., vii 2014: 169. (poster presentation) Liu J., M. V. H. Wilson, and A. M. Murray, 2013. North American Eocene suckers and their implications for the systematics of Catostomidae (Ostariophysi, Cypriniformes). Supplement to the online Journal of Vertebrate Paleontology. 73rd Annual Meeting Society of Vertebrate Paleontology, Program and Abstracts. Los Angeles, Oct., 2013: 163. (poster presentation) Liu J., Z. J. Tseng, M. V. H. Wilson, and A. M. Murray, 2012. Body shape differences between North American and Asian fossil catostomids and ontogenetic change in early cypriniforms. Supplement to the online Journal of Vertebrate Paleontology. 72nd Annual Meeting Society of Vertebrate Paleontology, Program and Abstracts. Raleigh, Oct., 2012: 128. (poster presentation) Liu J., M. V. H. Wilson, M-m. Chang, and A. M. Murray, 2011. The early diversity of non-cyprinid cypriniforms (Ostariophysi, Cypriniformes) in the Eocene of East Asia and North America. 71st Annual Meeting Society of Vertebrate Paleontology, Program and Abstracts. Las Vegas, Nov., 2011: 145. (oral presentation) Liu J., M-m. Chang, M. V. H. Wilson, 2010. The fossil catostomid Jianghanichthys from China and implications for the evolution of basal catostomids (Cypriniformes, Actinopterygii). 70th viii Anniversary Meeting Society of Vertebrate Paleontology, Program and Abstracts. Pittsburgh, Oct., 2010: 123A. (oral presentation) Of the published abstracts above, I was responsible for collecting data, designing and performing
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