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Parioglossus Osteology, Phylogeny and Biogeography of Parioglossus (Perciformes, Gobioidei) Rui Wang A thesis submitted in conformity with the requirements for the degree of Master of Science Graduate Department of Zoology University of Toronto O Copyright by Rui Wang (200 1) National Librafy Bibliothèque nationale 1+1 of Camda du Canada A uisitions and Acquisitioy et diqraphic Sewices services bibliographiques 395 WaUingtori Street 395. nie Wdlingtorr ûnawaON K1AON4 OttawaûN KlAûN4 Canada Canada The author has granteci a non- L'auteur a accordé une ticence non exclusive licence atlowing the exclusive permettant ii la National Libraiy of Canada to Bibliothbque nationale du Canada de reproduce, loaq distribute or seîi reproduire, prêter, distribuer ou copies of thi~thesis in microfom, vendre des copies de cette thèse sous paper or electronic formats. la forme de microiiche/film, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the &oit d'auteur qui protège cette thèse. thesis nor substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimes rejxoduced without the author's ou autrement reproduits sans son pemiission. autorisation. Abstract Osteology, Phylogeny and Biogeography of Parioglossus (Perciformes, Gobioidei) Rui Wang, Department of Zoology, M. Sc., 200 1, University of Toronto Pariogiossus (Percifoms, Gobioidei) is a group of small fishes that inhabits warm temperate to tropical regions of the western Pacific and Indian Oceans. Sixteen species and one undescribed species (P. sp A) were identified during this study. The taxonomy, extemal characters, osteology, phylogeny and biogeography of the genus were investigated. A panimony analysis of the 17 species generated three most parsimonious trees. AI1 trees support the following relationships between species: (((((P.dotui, P. neocaledonicus) P. nudus) P. marginalis) P. lineatus) P. triguetrus); which is the sister group of ((P. nudus, P. sp A) P. philippinus); the grouping of ((P. inferruptus.P. palus~ris)P. taeniatus);their relationship with P. raoi and P. sinensis is unresolved; P. verticaiis is the sister species of al1 other ingroup taxa. A biogeographical analysis of the genus suggests that western-Pacific was not the centre of origin, but rather has overlapping biotas fkom di fferent areas. Acknowledgements My life, over the past three years, had been really overwhelming. 1 am really grateful to those individuals who helped me through al1 these. The staff and students at the Royal Ontario Museum provided exceptional assistance through various ways. M. Rouse helped me with al1 in- and out-specimens, M. Bumdge helped me with clearing and staining specimens, W. King, M. Zur and E. Holm also provided technical help. F. Santini introduced some interesting topics for discussions of phylogeny. W. Kilburn kindly lent me his drawing tools so that 1 could finish rny drawing. Dm.R. Murphy and D. McLeman have been of great assistance in a multiplicity of ways. I thank them all. To the following individuals I express my gratitude for loans of material, permission to process and dissect specimens: M. McGrouther (AMS), D. Catania and J. Fong (CAS), Drs. G. Duhamel, P. Patrice and B. Séret (MNHN), K. Hatooka (OMNH), S. Jewett and L. Palmer (USNM), Prof H-L Wu (SFU) and Dr. J-X Yang (KIZ). My mother (Y. Liu), my sister (X. Wang) and my brother (J. Wang), as well as their families helped me in various ways. They helped me look after my daughter, so that I could concentrate on my thesis without worrying too much about her. 1 dedicate this thesis to al1 of hem, including my husband (M. Li) and my daughter (A. Li). Although my daughter is still too young to read it now, 1 wish when she grow up, she will know the value of al1 the sacrifices we encounter. Finally, 1 would like to thank the members of my thesis committee (Drs.R.W. Murphy, D.A. McLennan) and rny thesis supervisor, Dr. R. Winterbottom. The latter provided supervision and support, and demonstrated an amazing tolerance of the procedure which led to the final version of this thesis. iii This work was supported by several University of Toronto open scholarships, a Frederick P. Ide Graduate Award at University of Toronto, as well as an NSERC operating gant (OGP-0007619) to R Winterbottom. Table of contents ABSTRACT . i i ACKNOWLEDEGMENTS . iii LISTOFTABLE . vii B.. LIST OF FIGURES . vi11 CHAPTER 1. INTRODUCTION: . 1 CHAPTER 2. MATERIALS . 9 CHAPTER 3. METHODS AND TERMINOLOGY . 14 Theory and criteria used for the construction of data matrix . 14 CHAPTER 4. RESULTS . 19 Key for the species of Parioglossus . 19 The osteology of Purioglossus ruinfordi . 22 Intraspecific variation of osteology in Pariuglossus . 43 Variation of osteology and general morphology between species 46 Character list . 46 S y stematics of Parioglossus . 64 Confidence and accuracy of the topology . 65 Description of a new species: Parioglossus sp. A . 71 CHAPTER 5. BIOGEOGRAPHY OF THE GEWS. 74 CHAPTER 6. CHARACTERS OF THE INGROUP SPECIES 95 LITERATURE CITED 1O3 APPENDIX 1 . 118 APPENDIX 2 . 120 APPENDIX 3 . APPENDIX 4 . LIST OF TABLE TABLE 1. Classification of families and subfamilies in Gobioidei in the last 30 years œ 7 vii LIST FIGURES FIGURE 1. Dorsal view of the cranium. 24 FIGURE 2. Ventral view of the cranium. 25 FIGURE 3. Left lateral view of the cranium. 26 FIGURE 4. Left lateral view of superficial bones, jaws and suspensorium. 29 FIGURE 5. Media1 view of urohyal and left hyoid arch. 32 FIGURE 6. Dorsal view of gill arches and basihyal . 34 FIGURE 7. LeR lateral view of pectoral girdle . 37 FIGURE 8. Dorsal view of pelvic girdle. 38 FIGURE 9. Diagrammatic leR lateral view of Parioglossus, showing vertebrae, ribs, epineurals, and pteregiophores of first dorsal fin . 40 FIGURE 10. LeA lateral view of caudal skeleton. 41 FIGURE 11. Intraspecific variation of caudal cornplex. 44 FIGURE 12. Premaxilla variation. 45 FIGURE 13. GillopeningofPurioglossus . 49 FIGURE 14. Head pore tenninology. 50 I iGURE 15. Degree of ossification of rostral cartilage . 52 FIGURE 16. Degree of ossification of pelvis. 55 FIGURE 17. Variation in procurrent cartilage of Parioglossus. 56 FIGURE 18. Extemal morphological characters of Parioglossus. 59 FIGURE 19. Variation of preopercular process . 63 FIGURE 20. Phylogeny of Porioglossui. 66 FIGURE 2 1. Fi@ percent majority rule consensus of bootstrap portion . 69 viii FIGURE 22. Map of areas used in BPA nrialysis . 78 FIGURE 23. Distribution of Parioglossus aporos, P. neocaledonicus and P. sinensis 79 FIGURE 24. Distribution of Parioglossus dotui, P. marginalis and P. sp A. 80 FIGURE 25. Distribution of Parioglossw fornosus . 81 FIGURE 26. Distribution of Parioglossus internrprus . 82 FIGURE 27. Distribution of Porioglossus lineutus, P. tliquetnrs and P. verticalis . 83 FIGURE 28. Distribution of Parioglossus nudus . 84 FIGURE 29. Distribution of Parioglossuspalusiris . 85 FIGURE 30. Distribution of PariogIossus philippinus 88 FIGURE 3 1. Distribution of Parioglossus rainfordi . a 87 FIGURE32.DistributionofPariogIosstl~raoi . a 88 FIGURE 3 3. Distribut ion of Parioglossus taeniatus . 89 FIGURE 34. Area relationships suggested by the phylogeny and distribution of Parioglossus . a 90 CHAPTER 1. INTRODUCTION: The gobioid fish genus Parioglossus (Teleostei: Gobioidea) was erected by Regan (1 9 12) for the type species P. taeniatus, and now has 16 recognized species. Most of these species inhabit wmtemperate to tropical regions of the western Pacific and Indian Oceans, north to southem Korea (Kim and Han, 1993), south to southem Australia, West to Madagascar and east to Fiji (Rennis and Hoese, 1985). Only P. taeniatus, P. philippinus and P. moi are distnbuted in both the western Pacific and the Indian Oceans. Members of the genus are normally found around the roots of mangroves, or around algae in estuaries and coastal coral reefs. Seven of the 16 species (P. formosur, P. nudus, P. palustris, P. philippinus, P. rainfordi, P. raoi and P. taeniatus) appear to be geographically widespread (Rennis and Hoese, 1985). One species seems to be confined to fresh water (P. neocaledonicus; Dingerkus and Séret, 1992). Tomiyama (1958, 1959) reviewed the genus and recognized four species (P. taeniatus Regan 19 12, P. rainfordi McCulloch, 192 1, P. borneensis Koumans, 195 3 and P. ddui Tomiyama, 1958). Miller (1971) studied the systematic position and occurrence of Parioglossus in Lndian waters and recognized the same four species as Tomiyama. He also found that the number of soft rays of the second dorsal fin and anal fin had little value for species definition. He pointed out the possibility that al1 the four nominal species could be color phases of a single species, perhaps linked with sexual dimorphism in nuchal ridge and degree of elongation of first dorsal fin developrnent. Remis and Hoese (1985) revisited the genus, recognized 14 species (including six new species and one undescnbed as P. sp), detailed the synonyms, general morphology of the genus and osteology of some species, including the diagnoses, descriptions, distributions, cornparisons, and photographs of each species, and provided a key to the species. Parioglossus borneensis Koumans 1953 as viewed by Tomiyama (1958,1959) and Miller (197 1) was regarded by Rennis and Hoese (1 985) as a synonym of P. palustris. Rennis and Hoese (1 985) also discussed the similarities between Parioglossus and related genera in the subfamily Ptereleotrinae. Since then, two additional new species have been described: P. neocdedonicus Dingerkus and Séret, 1992 and P. sinensis Zhong, 1994. The P. sp of Rennis and Hoese was subsequently described as P. interruptur by Suzuki and Senou (1994). They compared their new species with other described species, and also provided a key to the genus. Suzuki et al. (1994) reviewed the species of Purioglossus in Japan (including their distribution), and provided a key to the Japanese species.
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