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Taxonomic Revision of the Hill-Stream Loach Genus Homaloptera (Teleostei

Permanent Link: http://ufdc.ufl.edu/UFE0045469/00001

Material Information

Title: Taxonomic Revision of the Hill-Stream Loach Genus Homaloptera (Teleostei Balitoridae)
Physical Description: 1 online resource (159 p.)
Language: english
Creator: Randall, Zachary S
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2013

Subjects

Subjects / Keywords: balitoropsis -- homaloptera -- homalopteroides -- homalopterula -- loaches
Biology -- Dissertations, Academic -- UF
Genre: Zoology thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: Homaloptera van Hasselt 1823 as treated historically exhibits a large amount of morphological diversity and is paraphyletic. The diversity and phylogenetic relationships of Homaloptera are reviewed, and the following three genera are removed from the synonymy of Homaloptera: Homalopteroides Fowler 1905, Homalopterula Fowler 1940, and Balitoropsis Smith 1945. Homaloptera is distinguished from these genera based on a combination of characters including a unique color pattern, origin of the dorsal fin anterior to the origin of the pelvic fin, > 20 predorsal scales, and mouth morphology. The following six species are recognized in Homaloptera: Homaloptera ocellata van der Hoeven 1832, H. bilineata Blyth 1860, H. orthogoniata Vaillant 1902, H. ogilviei Alfred 1967, H. confuzona Kottelat 2000, and H. parclitella Tan and Ng 2005. Homalopteroides was investigated and is distinguished from Homaloptera, Homalopterula, and Balitoropsis based on a combination of characters including a unique mouth morphology,origin of the dorsal fin posterior to the origin of the pelvic fin, > 60 lateral-line scales, and > 30 predorsal scales. Species included in Homalopteroides are H. wassinkii (Bleeker 1853), H. modestus (Vinciguerra 1890), H. rupicola (Prashad & Mukerji 1929), H. smithi (Hora 1932), H. stephensoni (Hora 1932), H. weberi (Hora 1932), H. tweediei (Herre 1940), H. indochinensis (Silas 1953), H. nebulosus (Alfred 1969), H. manipurensis (Arunkumar 1998), and H. yuwonoi (Kottelat 1998), and a new species known to occur only in the Rajang River Basin, Sarawak, Malaysia,described herein. Homalopteroides sp. n. is distinguished from all other species of Homalopteroides by a larger gape width of 28.5-33.3% HL.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Statement of Responsibility: by Zachary S Randall.
Thesis: Thesis (M.S.)--University of Florida, 2013.
Local: Adviser: Page, Larry M.
Electronic Access: RESTRICTED TO UF STUDENTS, STAFF, FACULTY, AND ON-CAMPUS USE UNTIL 2015-05-31

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2013
System ID: UFE0045469:00001

Permanent Link: http://ufdc.ufl.edu/UFE0045469/00001

Material Information

Title: Taxonomic Revision of the Hill-Stream Loach Genus Homaloptera (Teleostei Balitoridae)
Physical Description: 1 online resource (159 p.)
Language: english
Creator: Randall, Zachary S
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2013

Subjects

Subjects / Keywords: balitoropsis -- homaloptera -- homalopteroides -- homalopterula -- loaches
Biology -- Dissertations, Academic -- UF
Genre: Zoology thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: Homaloptera van Hasselt 1823 as treated historically exhibits a large amount of morphological diversity and is paraphyletic. The diversity and phylogenetic relationships of Homaloptera are reviewed, and the following three genera are removed from the synonymy of Homaloptera: Homalopteroides Fowler 1905, Homalopterula Fowler 1940, and Balitoropsis Smith 1945. Homaloptera is distinguished from these genera based on a combination of characters including a unique color pattern, origin of the dorsal fin anterior to the origin of the pelvic fin, > 20 predorsal scales, and mouth morphology. The following six species are recognized in Homaloptera: Homaloptera ocellata van der Hoeven 1832, H. bilineata Blyth 1860, H. orthogoniata Vaillant 1902, H. ogilviei Alfred 1967, H. confuzona Kottelat 2000, and H. parclitella Tan and Ng 2005. Homalopteroides was investigated and is distinguished from Homaloptera, Homalopterula, and Balitoropsis based on a combination of characters including a unique mouth morphology,origin of the dorsal fin posterior to the origin of the pelvic fin, > 60 lateral-line scales, and > 30 predorsal scales. Species included in Homalopteroides are H. wassinkii (Bleeker 1853), H. modestus (Vinciguerra 1890), H. rupicola (Prashad & Mukerji 1929), H. smithi (Hora 1932), H. stephensoni (Hora 1932), H. weberi (Hora 1932), H. tweediei (Herre 1940), H. indochinensis (Silas 1953), H. nebulosus (Alfred 1969), H. manipurensis (Arunkumar 1998), and H. yuwonoi (Kottelat 1998), and a new species known to occur only in the Rajang River Basin, Sarawak, Malaysia,described herein. Homalopteroides sp. n. is distinguished from all other species of Homalopteroides by a larger gape width of 28.5-33.3% HL.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Statement of Responsibility: by Zachary S Randall.
Thesis: Thesis (M.S.)--University of Florida, 2013.
Local: Adviser: Page, Larry M.
Electronic Access: RESTRICTED TO UF STUDENTS, STAFF, FACULTY, AND ON-CAMPUS USE UNTIL 2015-05-31

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2013
System ID: UFE0045469:00001


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1 TAXONOMIC REVISION OF THE HIL L STREAM LOACH GENUS HOMALOPTERA (TELEOSTEI: BALITORIDAE) By ZACHARY SETH RAND A LL A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS F OR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2013

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2 2013 Zachary Seth Randall

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3 ACKNOWLEDGMENTS Funding for this study was provided by the All Cypriniformes Species Inventory Project funded by the U.S. National Scienc e Foundation (DEB 1022720). I would like to thank Lawrence Page for dire cting me to this topic of study and for his constructive criticism and guidance. I also thank my committee members Jonathan Bloch and Ed Braun for their guidance and helpful comments a nd David Reed for access to his camera setup funded by the U.S. National Science Foundation (DEB 0845392) The following individuals kindly provided photos for F igures: Mark Sabaj Prez for F igures 5 19 and 5 23, Lynne Parenti and Jeff Clayton for F igure 5 34, Renny Hadiaty for F igure 1 5A & D, Sandra Raredon for F igure 4 4, and Daniel Lumbantobing for F igure 1 5C. For specimen loans and access to institutional specimens I thank Mark Sabaj Prez (ANSP), James Maclain (BMNH), David Catania (CAS), Mary A. Rog ers (FMNH), Karsten Hartel (MCZ), Sirwan Suksri (NIFI), Prachya Musikasinthorn (RLIKU), Hernn Lpez Fernndez (ROM), Rob Robins (UF), Doug Nelson (UMMZ), Jeff Williams (USNM), and Kelvin Lim (ZRC) I would like to thank F. W. H. Beamish, C. Grudpan, J. Gr udpan, R. Plongsesthee, W. Tangjitjaroen, P. Chanintarapoom, S. Dakhunthod, W. Ngokkownok, R. Singer and Z. Martin for assist ance with collecting. I would also like to thank t he following volunteers for Georeferencing and data organization: Patrick Riggs, Lunide Orleus, Karen Alldridge, Ella Foster, and Chloe Hough. My study benefited from discussions on Homaloptera with L. M Page, W. E. Eschmeyer D. Lumbantobing, R. H. Robins Z. Martin, R. Singer, A.W. Thomson, and J. Grosso Permission to use the photog raphs seen in Figure 4 6 was arranged by David Catania of the California Academy of Sciences

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4 TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................ ................................ ................................ .. 3 LIST OF TABLES ................................ ................................ ................................ ............ 7 LIST OF FIGURES ................................ ................................ ................................ .......... 9 ABSTRACT ................................ ................................ ................................ ................... 13 CHAPTER 1 INTRODUCTION ................................ ................................ ................................ .... 15 Homaloptera van Hasselt 1823 ................................ ................................ ............... 17 Homalopteroides Fowler 1905 ................................ ................................ ................ 18 Homalopterula Fowler 1940 ................................ ................................ .................... 18 Bali toropsis Smith 1945 ................................ ................................ .......................... 20 Indian Group ................................ ................................ ................................ ........... 21 2 METHODS ................................ ................................ ................................ .............. 29 Morphometric And Meristic Methods ................................ ................................ ...... 29 Molecular Phylogenetic Methods ................................ ................................ ............ 30 Taxonomic Methods ................................ ................................ ................................ 31 3 MATERIALS EXAMINED ................................ ................................ ........................ 33 Institutional Abbreviations ................................ ................................ ....................... 33 Homaloptera ................................ ................................ ................................ ........... 33 Homalopteroides ................................ ................................ ................................ ..... 34 Homalopterula ................................ ................................ ................................ ........ 37 Balitoropsis ................................ ................................ ................................ ............. 37 Indian Group ................................ ................................ ................................ ........... 38 4 ON THE PARAPHYLY OF HOMALOPTERA VAN HASSELT 1823 ....................... 39 Homaloptera van Hasselt 1823 ................................ ................................ ............... 39 Homalopteroides Fowler 1905 ................................ ................................ ................ 42 Homalopterula Fowler 1940 ................................ ................................ .................... 45 Balitoropsis Smith 1 945 ................................ ................................ .......................... 47 Indian group ................................ ................................ ................................ ............ 49 5 TAXONOMIC REVISION OF HOMALOPTEROIDES FOWLER 1905 .................... 58 Homalopteroides Fowler 1905 ................................ ................................ ................ 58

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5 Homalopteroides wassinkii (Bleeker 1853) ................................ ............................. 58 Diagnosis ................................ ................................ ................................ .......... 58 Description ................................ ................................ ................................ ....... 59 Coloration ................................ ................................ ................................ ......... 61 Distribution ................................ ................................ ................................ ....... 62 Homalopteroides modestus (Vinciguerra 1890) ................................ ...................... 62 Diagnosis ................................ ................................ ................................ .......... 6 2 Description ................................ ................................ ................................ ....... 63 Coloration ................................ ................................ ................................ ......... 65 Distribution ................................ ................................ ................................ ....... 66 Remarks ................................ ................................ ................................ ........... 67 Homalopteroides rupicola (Prashad & Mukerji 1929) ................................ .............. 68 Diagnosis ................................ ................................ ................................ .......... 69 Description ................................ ................................ ................................ ....... 70 Distribution ................................ ................................ ................................ ....... 70 Homalopteroides smithi (Hora 1932) ................................ ................................ ...... 70 Diagnosis ................................ ................................ ................................ .......... 70 Description ................................ ................................ ................................ ....... 71 Distribution ................................ ................................ ................................ ....... 71 Remarks ................................ ................................ ................................ ........... 71 Homalopteroides stephensoni (Hora 1932) ................................ ............................ 72 Diagnosis ................................ ................................ ................................ .......... 72 Description ................................ ................................ ................................ ....... 73 Distribution ................................ ................................ ................................ ....... 73 Homalopteroides weberi (Hora 1932) ................................ ................................ ..... 74 Diagnosis ................................ ................................ ................................ .......... 74 Description ................................ ................................ ................................ ....... 75 Coloration ................................ ................................ ................................ ......... 76 Distribution ................................ ................................ ................................ ....... 77 Homa lopteroides tweediei (Herra 1940) ................................ ................................ 77 Diagnosis ................................ ................................ ................................ .......... 78 Description ................................ ................................ ................................ ....... 79 Distribution ................................ ................................ ................................ ....... 79 Homalopteroides indochinensis (Silas 1953) ................................ .......................... 79 Diagnosis ................................ ................................ ................................ .......... 80 Description ................................ ................................ ................................ ....... 80 Distribution ................................ ................................ ................................ ....... 80 Remarks ................................ ................................ ................................ ........... 80 Homalopter oides nebulosus (Alfred 1969) ................................ .............................. 81 Diagnosis ................................ ................................ ................................ .......... 81 Description ................................ ................................ ................................ ....... 82 Dis tribution ................................ ................................ ................................ ....... 82 Homalopteroides yuwonoi (Kottelat 1998) ................................ .............................. 83 Diagnosis ................................ ................................ ................................ .......... 83 Description ................................ ................................ ................................ ....... 84 Distribution ................................ ................................ ................................ ....... 84

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6 Remarks ................................ ................................ ................................ ........... 84 Homalopteroides m anipurensis (Arunkumar 1998) ................................ ................. 84 Diagnosis ................................ ................................ ................................ .......... 85 Description ................................ ................................ ................................ ....... 85 Dist ribution ................................ ................................ ................................ ....... 85 Remarks ................................ ................................ ................................ ........... 86 Homalopteroides n. sp ................................ ................................ ............................ 86 Diagnosis ................................ ................................ ................................ .......... 87 Description ................................ ................................ ................................ ....... 87 Coloration ................................ ................................ ................................ ......... 89 Distribution ................................ ................................ ................................ ....... 90 6 DISCUSSION ................................ ................................ ................................ ....... 148 LIST OF REFERENCES ................................ ................................ ............................. 154 BIOGRAPHICAL SKETCH ................................ ................................ .......................... 159

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7 LIST OF TABLES Table page 1 1 Morphometric measurements and meristic counts for Homaloptera Homalopteroides Homalopterula and Balitoropsis All measurements are expre ssed in mm. ................................ ................................ ............................... 22 1 2 External morphological characters for Homaloptera Homalopteroides Homalopterula and Balitoropsis ................................ ................................ ........ 23 2 1 G enbank accession numbers and sources for analyzed samples. ..................... 32 5 1 Diagnostic characters for species of Homalopteroides Species that may have a fully pigmented lower lobe of the caudal fin are represented by ^. .......... 91 5 2 Diagnostic characters for species belonging to Homalopteroides ..................... 93 5 3 Morphometric measurements and m eristic counts for Homalopteroides wassinkii (N=2). All measurements in mm. Number of individuals in parentheses. Paralectotype represented by a *. ................................ ................. 95 5 4 Morphometric measurements and meristi c counts for adult (N=58) and juvenile (N=10) Homalopteroides modestus All measurements in mm. Number of individuals in parentheses. ................................ ................................ 97 5 5 Morphometric measurements and meristic counts for p aratype of Homalopteroides rupicola ................................ ................................ .................. 99 5 6 Morphometric measurements (N=19) and meristic counts (N=21) for Homalopteroides smithi All measurements in mm. Number of individuals in parentheses Five syntypes are included in meristic counts. ............................ 101 5 7 Morphometric measurements and meristic counts for Homalopteroides stephensoni (N=5). All measurements in mm. Number of individuals in p arentheses. ................................ ................................ ................................ ..... 103 5 8 Morphometric measurements and meristic counts for Homalopteroides weberi (N=31). All measurements in mm. Number of individuals in parentheses. ................................ ................................ ................................ ..... 105 5 9 Morphometric measurements and meristic counts for Homalopteroides tweediei (N=7). All measurements in mm. Number of individuals in parentheses. Holotype represented by a *. ................................ ...................... 107 5 10 Morphometric measurements and meristic counts for Homalopteroides nebulosus (N=4). All measurements in mm. Number of individuals in parentheses. Holotype represented by a *. ................................ ...................... 109

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8 5 11 Morphometric measurements and meristic counts for holotype of Homalopteroides yuwonoi All measurements in mm. ................................ ...... 111 5 12 Morphometric measurements and meristic counts fo r Homalopteroides n. sp. (N=5). All measurements in mm. Number of individuals in parentheses. Holotype is represented by a *. ................................ ................................ ......... 112

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9 LIST OF FIGURES Figure page 1 1 Type localities of species belonging to the genus Homaloptera ( s.l ) .................. 24 1 2 Dorsal views of: (A) Homaloptera ocellata UF 161718, 64.1 mm SL; (B) Homalopteroides wassinkii UMMZ 1 55660, 46.2 mm SL; (C) Homalopterula ripleyi ANSP 188908, 43.1 mm SL; (D) Balitoropsis zollingeri UF 166094, 52.1 mm SL. Scale bars represents 30mm; B, C, and D share a scale bar. ....... 25 1 3 Lat eral views of: (A) Homaloptera ocellata UF 161718, 64.1 mm SL; (B) Homalopteroides wassinkii UMMZ 155660, 46.2 mm SL; (C) Homalopterula ripleyi ANSP 188908, 43.1 mm SL; (D) Balitoropsis zollingeri UF 166094, 52.1 mm SL. Scale bars represents 30mm; A, B and C share a scale bar. ....... 26 1 4 Ventral views of: (A) Homaloptera ocellata UF 161718, 64.1 mm SL, V represents mental pad; (B) Homalopteroides wassinkii UMMZ 155660, 46.2 mm SL; (C) Homalo pterula ripleyi ANSP 188908, 43.1 mm SL, asterisk represents mental lobe; (D) Balitoropsis zollingeri UF 166094, 52.1 mm SL. Scale bars equals 3mm; A, B, and C share a scale bar. ................................ .... 27 1 5 Late ral views of living (A) Homaloptera orthogoniata not cataloged; (B) Homalopteroides cf. smithi not cataloged; (C) Homalopterula cf. ripleyi not cataloged; (D) Balitoropsis zollingeri not cataloged. Photos A & D by Renny Kurnia Hadiaty. Photo C by Danie l Lumbantobing. ................................ ............. 28 4 1 Type localities for species belonging to Homaloptera ................................ ......... 51 4 2 Type localities for species belonging to Homa lopteroides ................................ .. 52 4 3 Type localities for species belonging to Homalopterula ................................ ...... 53 4 4 Balitoropsis bartschi USNM 107963 (holotype). Scale bar represents 30 mm. Photos by Sandra Raredon. ................................ ................................ ............... 54 4 5 Type localities for species belonging to Balitoropsis ................................ ........... 55 4 6 (A) Do rsal, lateral, and ventral views of Homaloptera montana CAS 139871, 46.4 mm SL; Scale bar represents 30mm;(B) ventral view of H. montana, CAS 139871. Photos by California Academy of Sciences Ichthyology Section. ................................ ................................ ................................ .............. 56 4 7 Type localities for species of Homaloptera ( s.l ................................ ................................ ................................ ................. 57

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10 5 1 Dorsal, lateral, and ventral views of Homalopteroides wassinkii, UMMZ 155660 46.2 mm SL; Indonesia, Java, Tjisokan, inland fishery. Scale bar represents 30 mm. ................................ ................................ ............................ 114 5 2 Distribution of Homalopteroides wassinkii Dot represents locality for specimens examined; asterisk represents type locality. ................................ ... 115 5 3 Homalopteroides modestus, Thailand, Kanchanaburi, Mae Khlong basin: A) dorsal, lateral, and ventral views of an adult, UF 181080, 37.2 mm SL, scale bar repre sents 30 mm; B) lateral view of a juvenile, UF 176377, 24.8 mm SL, scale bar represents 10 mm. ................................ ................................ ............ 116 5 4 Mouth of Homalopteroides modestus UF 181080, 37.2 mm SL. Scale bar represents 2 mm. ................................ ................................ .............................. 117 5 5 Fifth ceratobranchial of Homalopteroides modestus : A) adapted from Vinciguerra (1890), p. 331; B) UF 176454, 31.48 mm SL, left side, ventral view; C) UF 176454, 31.48 mm SL, right side, do rsal view. B and C scale bar represents 0.5 mm. ................................ ................................ ........................... 118 5 6 Distribution of Homalopteroides modestus Black dots represent localities for specimens examined; asterisk represents type locality. ................................ ... 119 5 7 Habitat of Homalopteroides modestus Thailand, Kanchanaburi, Mae Khlong basin, Mae Nam Kwae, Khayeng River, 30 Dec. 2011, 14.65 N, 98.566667 E. ................................ ................................ ................................ ...................... 120 5 8 Lateral view of Homalopteroides rupicola : A) paratype SU 28726, 21.3 mm SL, Myanmar, Myitkyina District, Sankha, a large hillstream, midway between Kamaing and Mogaung, Irrawaddy River basin. Scale bar represents 10 mm. B) ad apted from Prashad and Mukerji (1929), plate 8, Figure 3. ...................... 121 5 9 Type locality of Homalopteroides rupicola ................................ ....................... 122 5 10 Dorsal, La teral, and ventral views of Homalopteroides smithi : paratype USNM 109821, 50 mm SL, Thailand, Nakon Sritamarat, Klong Pong Tadi Stream; ? Pakphanang basin. Scale bar represents 20 mm. ........................... 123 5 11 Dorsal and lateral view of Homalopteroides smithi : UF 183330, 43.6 mm SL, Thailand, Nakon Sritamarat, Chang Klang Village, Tapee Basin. Scale bar represents 20 mm. ................................ ................................ ............................ 124 5 12 Distribution of Ho malopteroides smithi Dots represent localities for specimens examined; asterisk represents type locality. ................................ ... 125 5 13 Scatter plot of caudal peduncle depth for all species of Homalopteroides ....... 126

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11 5 14 Mouth of Homalopteroides stephensoni USNM 230254, 50.0 mm SL. Scale bar represents 2 mm. ................................ ................................ ....................... 127 5 15 Dorsal, lateral, and v entral views of Homalopteroides stephensoni, USNM 230254, 50.0 mm SL, Indonesia, West Kalimantan, Kapuas basin. Scale bar represents 20 mm. ................................ ................................ ............................ 128 5 16 Distribution of Homalopteroides stephensoni Dot represents locality for specimen examined; asterisk represents type locality. ................................ ..... 129 5 17 Mouth of Homalopteroides weberi ROM 82131, 33.7 mm SL. Scale bar represents 2 mm. ................................ ................................ .............................. 130 5 18 Dorsal, lateral, and ventral views of Homalopteroides weberi east Malaysia, Sarawak, Baram basin; A) ROM 70464, 40.8 mm SL; B) ROM 82131, 33.7 mm SL. Scale bars represent 20 mm. ................................ .............................. 131 5 19 Dorsal, lateral, and ventral views of syntype Homalopteroides weberi BMNH 1895.7.2.81; 41.8 mm SL east Malaysia, Sarawak, Baram basin; scale bar represents 10 mm. Photos by Mark Sabaj Prez. ................................ ............ 132 5 20 Distribution of Homalopteroides weberi Dots represent locality for specimens examined; asterisk represents type locality. ................................ ..................... 133 5 2 1 Dorsal, lateral, and ventral views of Homalopteroides tweediei ; A) holotype SU 33012; Malaysia, Johore, Mawai District (? Kota Tinngi), ? Ambat basin; B) UMMZ 238923 26.0 mm SL, Malaysia, Johore, Kahang River, Endau basin. Scale bar represents 10 mm. P hotos of A by CAS. ............................... 134 5 22 Distribution of Homalopteroides tweediei ................................ ......................... 135 5 23 Dorsal, lateral, and ventral views of Homalopteroi des indochinensis holotype represents 10 mm. Photos by Mark Sabaj Prez. ................................ ............ 136 5 24 Type locality of Homalopteroides indochinen sis ................................ .............. 137 5 25 Lateral views of Homalopteroides nebulosus A) Holotype ZRC 2020; 23.6 mm SL; Malaysia, Kelantan, Sok River, Kampong Sok, Kelantan basin; B) paratype BMNH 1967.11.15.15; 22.2 mm SL; same locality data as holotype. Scale bar represent 10 mm. ................................ ................................ ............. 138 5 26 Type locality of Homalopteroides nebulosus ................................ ................... 139 5 27 Hom alopteroides yuwonoi ; holotype MZB 5938, 38.6 mm SL; Indonesia, Borneo, Kalimantan Barat, Kapuas basin; A) lateral view, B) ventral view. ...... 140 5 28 Type locality of Homalopteroides yuwonoi ................................ ...................... 141

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12 5 29 Type locality of Homalopteroides manipurensis ................................ .............. 1 42 5 30 Scatter plot of gape width for all species of Homalopteroides ......................... 143 5 31 Dorsal, lateral, and ventral views of Homalopteroides n. sp holotype USNM 323875, 52.9 mm SL, Malaysia, Sarawak, Rajang basin. Scale bar represents 20 mm. ................................ ................................ ............................ 144 5 32 Mouth of Homalopteroides n. sp. holotype USNM 323875, 52.9 mm SL. Scale bar represents 3 mm. ................................ ................................ .............. 145 5 33 Distribution of Homalopteroides n. sp Dots re present locality for specimens examined; asterisk represents type locality. ................................ ..................... 146 5 34 Habitat of Homalopteroides n. sp east Malaysia, Sarawak, Balui River, Rajang basin, 02 Aug. 1991, 2.416667 N, 113.733333 E. Photo by Lynne Parenti. ................................ ................................ ................................ ............. 147 6 1 Phylogenetic relationships of the family Balitoridae from a Bayesian analysis of the Rag1 gene. An asterisk indicates a posterior probabilit y of 100%. ......... 153

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13 Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science TAXONOMIC REVISION OF THE HIL L STREAM LOACH GENUS HOMALOPTERA (TELEOSTEI: BALITORIDAE) By Zachary Seth Randall May 2013 Chair: Lawrence M. Page Major: Zoology Homaloptera van Hasselt 1823 as treated historically exhibits a large amount of morphological diversity and is paraphyletic. The diversity and phylogenetic relationships of Homaloptera are reviewed, and the following three genera are removed from the synonymy of Homaloptera : Homalopteroides Fowler 1905, Homalopterula Fowler 1940, and Balitoropsis Smith 1945. Homaloptera is dist inguished from these genera based on a combination of characters including a unique color pattern, origin of the dorsal fin following six species are recognized in H omaloptera : Homaloptera ocellata van der Hoeven 183 2, H. bilineata Blyth 1860 H. orthogoniata Vaillant 1902 H. ogilviei Alfred 1967, H. confuzona Kottelat 2000 and H. parclitella Tan and Ng 2005 Homalopteroides was investigated and is distinguished fr om Homaloptera Homalopterula and Balitoropsis based on a combination of characters including a unique mouth morphology, origin of the dorsal fin posterior to the origin of the pelvic fin, Homalopteroides are H. wassinkii (Bleeker 1853), H. modestus (Vinciguerra 1890), H. rupicola (Prashad

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14 & Mukerji 1929), H. smithi (Hora 1932), H. stephensoni (Hora 193 2), H.weberi (Hora 1932), H. tweediei (Herre 1940), H. indochinensis (Silas 1953), H. nebulos us (Alfred 1969), H. manipurensis (Arunkumar 1998), and H. yuwonoi (Kottelat 1998) and a new species known to occur only in the Rajang River Basin, Sarawak, Malay sia, described herein. Homalopteroides sp. n. is distinguished from all other species of Homalopteroides b y a larger gape width of 28.5 33.3% HL.

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15 CHAPTER 1 INTRODUCTION Homaloptera van Hasselt 1823 is the most species rich genus of the subfamily Balitor inae, comprising 35 valid species ( Eschmeyer & Fricke 2012) It is the type genus for the family Homalopteridae Bleeker 1859 which is a junior synonym of Balitoridae Swainson 1839 (Kottelat 1988). Van Hasselt distinguished Homaloptera from other genera by the horizontal position of its pectoral and pelvic fins; however, this character is applicable to several genera in Balitoridae. Through the years the genus has been distinguished from other genera of the subfamily Balitorinae in having smooth lips vs. lip s with papillae ( Balitora Gray 1830, Hemimyzon Regan 1911, Annamia Hora 1932, Sinogastromyzon Fang 1930, Metahomaloptera Chang 1944, Jinshaia Kottelat & Chu 1988), a single barbel at each corner of the mouth vs. more than one barbel ( Lepturichthys Regan 19 11, Neohomaloptera Herre 1944, Cryptotora Kottelat 1998), the gill opening extending to the ventral surface of the body vs. not reaching the ventral surface ( Bhavania Hora 1920), and absence vs. presence of two papillae between the lateral portions of the lower lip ( Travancoria Hora 1941). Other diagnostic characters for Homaloptera are 3 8 simple and 8 13 branched pectoral fin rays, and 2 simple and 7 8 branched pelvic fin rays. Species of Homaloptera occur in India, Myanmar, China, Thailand, Laos, Cambod ia, Vietnam, and south to Sumatra, Java, and Borneo ( Fig ure 1 1 ) The species have a diverse range in body size, with the smallest species, Homaloptera tweediei Herre 1940, reaching 26 mm SL (Herre 1940), and the largest, Homaloptera parclitella Tan and Ng 2005, reaching 102 mm SL (Tan & Ng 2005). Like most balitorids, species

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16 of Homaloptera are rheophilic and share a body plan designed for fast flowing water. Some members of the genus feed on invertebrate s and diatoms (Rainboth 1996). Historically, Homalo ptera has exhibited a large amount of morphological variation. This variation, likely due to the genus being paraphyletic, has been commented on by many authors (Fang 1930; Hora 1932; Kottelat 1998; Tan and Ng Homaloptera is a polyphyleti c genus. It has been variously divided into some subgenera, whose phyletic relationship h as not yet been well worked ou t... w hether they may be grouped under this genus or not is quite uncertain. As a whole the classification of Homaloptera at present requi res further investigations (Fang 1930, pg. 27 ). The subgenus mentioned by Fang (1930) is Homalopteroides Fowler 1905 treated as a subgenus by Weber and Beaufort (1916) Great confusion prevails regarding the taxonomy of the fishes assigned to the genus Homaloptera and for a long time the genus acted as a lumber room for the accommodation of Homalopterid species of diverse affinities. In recent years several species have been taken out of the genu s and rightly made the types of new genera; but a great dea l still remains to be done (Hora 1932, pg. 275 ) The new genera referred to by Hora (1932) are Hemimyzon Regan 1911, Lepturichthys Regan 1911, and Protomyzon Hora 1932 Kottelat (1998) proposed that Homaloptera consisted of five lineages, which were recog nized as subgenera by later authors, including Tan and Ng (2005) and Tan (2009). These lineages were Homaloptera van Hasselt 1823, Homalopteroides Fowler 1905, Homalopterula Fowler 1940, Balitoropsis Smith 1945, and a group from the Western Ghats of India which remains to be named. Dorsal, l ateral and ventral views of the type

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17 species of the genera proposed by Kottelat (1998) are shown in Figures 1 2,1 3, 1 4 & 1 5 All five groups are described in more detail below. Homaloptera van Hasselt 1823 (Figures 1 2A, 1 3A, 1 4A and 1 5A) Kottelat (1998) recognized Homaloptera ocellata van der Hoeven 1832, H. bilineata Blyth 1860, H. orthogoniata Vaillant, 1902 and H. ogilviei Alfred 1967 as being possibly a clade to the exclusion of species belonging to Homalopt era ( sensu lato ) (having in common reddish tints and similar dark markings on the head and the fins), a sl ightly compressed body, 56 65 lateral line scales and the dorsal fin origin situated in advance of the pelvic fin orig this diagnosis and the presence of an elongated head and a lateral line scale count of 61 77, Tan and Ng (2005) treated Homaloptera as a subgenus. They included H confuzona Kottelat 2000 in Homaloptera ( sensu stricto ), but neglected to includ e Homaloptera parclitella Tan and Ng, a species that shares all of the characters used by Tan and Ng (2005) to diagnose the subgenus. Kottelat (2012) recognized Homaloptera ( sensu stricto ) van Hasselt 1823 at the genus level, but gives no data or explanati on to support this decision. The only character provided by Kottelat (1998) that seems to be apomorphic for Homaloptera (sensu stricto) is the color pattern. All other characters provided by Kottelat (1998) and Tan and Ng (2005) are presen t in some or all species in Balitoropsis Smith 1945, Homalopterula Fowler 1940, and Homalopteroides Fowler 1905 (Tables 1 1 & 1 2).

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18 Homalopteroides Fowler 1905 (Figures 1 2B, 1 3B, 1 4B and 1 5B) Fowler (1905) removed Homaloptera wassinkii Bleeker 1953 from Homaloptera and placed it in a new genus Homalopteroides which was distinguished from Homaloptera by having the origin of the dorsal fin posterior, rather than anterior, to the origin of the pelvic fin This new genus, based on a single character, was given subgeneri c rank by later authors (Weber & de Beaufort 1916; Fowler 1940; Silas 1 953; Alfred 1969; Menon 1987). More recently, Homalopteroides was recognized as being peduncle, relativ ely long paired fins with the pectorals reaching and usually overlapping Tan and Ng (2005) treated Homalopteroides as a subgenus of Homaloptera Kottelat (2012) recogniz ed Homalopteroides as a genus, but gave no data or explanation to support this decision. None of the characters provided by Kottelat (1998) for Homalopteroides are apomorphic given that the three other potentially valid genera that he proposed (recognized as subgenera of Homaloptera by Tan & Ng 2005), Homaloptera ( sensu stricto ) Balitoropsis and Homalopterula include species that share one or more of these characters ( Tables 1 1 & 1 2 ). ter the origin of the dorsal fin in relation to the origin of the p elvic fin, also is not unique. All of the species belonging to subgenus Homalopterula and the group of Indian species from the Western Ghats also have the origin of the dorsal fin posterior to the origin of the pelvic fin. Homalopterula Fow ler 1940 (Figures 1 2C, 1 3C, 1 4C and 1 5C)

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19 The genus Homalopterula was created by Fowler for his new species Homalopterula ripleyi Fowler 1940 and was distinguished from other balitorids by the ted caudal and entirely naked Homalopterula was treated as a junior synonym of Homaloptera by Silas (1953) based on the illustration provided in garding the nature of the caudal and presence or absence of scales on the ventral surface of the abdomen in species of Homaloptera H. ripleyi (ANSP 68713) but thought it was best to consider Homalopterula as a junior synonym of Homaloptera This treatment by Silas (1953) was followed by Roberts (1989). The jaw shape F igured in Fowler (1940 Fig ure 6) is not consistently found in the H. ripleyi material collected by Ott (2009). Kottelat (1998) recognized Homalop terula as being possibly a distinct genus (2005) treated Homalopterula as a subge description for Homalopterula but did not recognize the group as a subgenus or genus. Kottelat (2012) treated Homalopterula Fowler 1940 at the genus level, but gave no supporting data or explanation to support this d escision. The only character provided by Kottelat (1998) that seems to be apomorphic for Homalopterula is caudal fin shape. Species belonging to the subgenera Homaloptera Homalopteroides or Balitoropsis share one or more of all other characters provided (Tables 1 1 & 1 2 ) A relatively wide mouth (gape) is found in some species belonging

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20 to Homalopteroides A more cylindrical body is found in species of Homaloptera Short paired fins are found in species of Homaloptera and Balitoropsis Balitoropsis Smith 1945 (Figures 1 2D, 1 3D, 1 4D and 1 5D) The genus Balitoropsis was created for the new species Balitoropsis bartschi Smith 1945 and was distinguished from Homaloptera ( sensu lato ) by having a deep preoral groove extending around the corners of the mouth and papillated lips. However, Kottelat and Chu (1988) noted that all members of Homaloptera have a preoral groove to a varying degree. Balitoropsis was further distinguished from other balitorids by Smith (1945) in having a cylindrical body, a greatly depr essed head, ventral fins extending past the anus which is closer to the base of the pelvics then to the anal fin, and a pectoral fin originating posterior to the eye and 4 simple and 10 branched rays. All of these characters are found in Homaloptera ( sensu lato ). Balitoropsis has been recognized by many authors as valid (Silas 1953; Chen 1978; Sawada 1982 ; Tang and Chen 2000; & Nguyen and Nguyen 2002 ) Kottelat (1998) recognized Balitoropsis elongate b ody, a slender caudal peduncle, carinated scales, short paired fins (pectorals usually not reaching pelvic base, pelvics not reaching anal), a dark body with a series of Tan and Ng (2005) treated Balitoropsis as a subgenus. Kottela t (2012) recognized Balitoropsis as a genus, but provides no data or explanation to support this descision. None of the characters provided by Kottelat (1998) for Balitoropsis are apomorphic; species of the subgenera Homaloptera and Homalopteroides may sha re one or more characters (Tables 1 1 & 1 2 ). Species belonging to Homaloptera have an

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21 elongate body, a shallow caudal peduncle, carniated scales and short paired fins. Species of Homalopteroides have dark bodies with a series of saddles along the back. I ndian Group Five species of Homaloptera ( sensu lato ) are known from the Western Ghats of India: H. montana Herre 1945, H. pillaii Indra and Rema Devi 1981, H. menoni Shaji and Easa 1995, H. santhamparaiensis Arunachalam et al 2002, and H. silasi Madhusoo dana Kurup and Radhakrishnan 2011. These species have been recognized as possibly constituting a group requiring a separate genus (Pethiyagoda & Kottelat 1994 [first two species] Kottelat 1998 [first three species] ), or subgenus (Tan & Ng 2005). Kottelat (2012) placed the Western Ghats species in Homaloptera The subgenera Homaloptera Homalopteroides Balitoropsis, and Homalopterula are elevated to genera b ased on their morphological diagnosability discussed in Chapter 4 and a taxonomic revision of Homalo pteroides will be given in C hapter 5.

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22 Table 1 1 Morphometric measurements and meristic counts for Homaloptera Homalopteroides Homalopterula and Balitoropsis All measurements are expressed in mm. M orphometrics Homaloptera ( n=36) Homalopteroides (n=136) Homalopterula (n=16) Balitoropsis (n=34) Standard Length 21.9 91.1 17.8 58.2 31.6 84.8 24.3 101.9 % of Standard Length Head length 22.1 30.0 22.2 30.6 21.8 28.4 18.5 25.6 Body depth 14.4 20.6 11.0 20.0 11.9 17.1 9.2 16.9 Body width 9.5 19.5 11.2 20.3 13.0 17.2 9.2 19 .0 Pectoral fin length 17.6 26.8 23.7 34.8 23.0 31.4 18.6 27.3 Pelvic fin length 14.7 22.9 16.9 25.7 18.2 26.1 16.3 23.5 Caudal peduncle length 13.8 21.6 12.1 18.4 7.4 17.2 13.9 21.5 C audal peduncle depth 6.2 10.1 6.1 11.5 8.6 11.1 4.6 9.2 % of Head Length Length of orbit 11.2 25.5 19.1 31.4 11.0 20.3 14.9 27.3 Interorbital width 23.7 35.7 19.2 34.2 31.5 37.1 28.6 37.1 Width of gape 14.5 23.5 17.3 33.3 24.4 38.5 15.6 23.9 % of Interorbital Width Length of orbit 35.3 85 70.1 146.7 30.6 64.3 43.8 78.9 M eristics Lateral line pore count 60 73 32 52 62 75 42 63 Predorsal scale count 20 27 14 25 32 56 13 19

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23 Table 1 2 External morpho logical characters for Homaloptera Homalopteroides Homalopterula and Balitoropsis C haracter Homaloptera ( s.s ) Homalopteroides Homalopterula Balitoropsis (n=36) (n=136) (n=16) (n=34) Dorsal fin origin relative to pelvic fin origin (Anterior/Posteri or) Anterior Posterior Posterior Anterior or above Pectoral fin extending to or past pelvic fin (Y/N) N Y/N Y/N N Pelvic fin extending to or past anus (Y/N) Y/N Y/N Y/N Y/N Caudal fin shape Forked Forked Truncate or deeply emarginated Forked Lips fin ely pappilated (Y/N) Y N Y/N Y Ventral surface of head with papillations (Y/N) Y N Y/N Y Isthmus foramen present (Y/N) N Y Y Y Scales keeled (Y/N) Y Y/N N Y

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24 Figure 1 1. Type localities of species belonging to the genus Homaloptera ( s.l )

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25 Figure 1 2. Dorsal views of: (A) Homaloptera ocellata UF 161718, 64.1 mm SL; (B) Homalopteroides wassinkii UMMZ 155660, 46.2 mm SL; (C) Homalopterula ripleyi ANSP 188908, 43.1 mm SL; (D) Balitoropsis zollingeri UF 166094, 52.1 mm SL. Scale bars represents 30mm ; B C and D share a scale bar.

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26 Figure 1 3. Lateral views of: (A) Homaloptera ocellata UF 161718, 64.1 mm SL; (B) Homalopteroides wassinkii UMMZ 155660, 46.2 mm SL; (C) Homalopterula ripleyi ANSP 188908, 43.1 mm SL; (D) Balitoropsis zollingeri UF 166094, 52.1 mm SL. Scale bars represents 30mm; A B and C share a scale bar.

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27 Figure 1 4. Ventral views of: (A) Homaloptera ocellata UF 161718, 64.1 mm SL, V represents mental pad; (B) Homalopteroides wassinkii UMMZ 155660, 46.2 mm SL; (C) Homalopte rula ripleyi ANSP 188908, 43.1 mm SL asterisk represents mental lobe; (D) Balitoropsis zollingeri UF 166094, 52.1 mm SL. Scale bars equals 3 mm; A B and C share a scale bar.

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28 Figure 1 5 Lateral views of living (A) Homaloptera orthogoniata not cata loged; (B) Homalopteroides cf. smithi not cataloged; (C) Homalopterula cf. ripleyi not cataloged ; (D) Balitoropsis zollingeri not cataloged. Photos A & D by Renny Kurnia Hadiaty Photo C by Daniel Lumbantobing.

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29 CHAPTER 2 METHODS Morphometric A nd Meris tic Methods Measurements and counts follow Hubbs and Lagler (2004 ) and Kottelat (1984) with exception of the following measurements defined below The terms origin and insertion refer, respectively, to the anterior and posterior points of the bases for eac h fin. Body depth at anus is the greatest body dimension at the anus. Prepectoral length is from tip of the snout to the origin of the pectoral fin Dis tance between anus and anal fin is from the anal opening to the origin of the first anal fin Snout t o nostril distance is from the tip of the snout to the anterior part of the nostril. Nostril to operculum distance is from the most posterior margin of the nostril to the hindmost part of the opercle. Internostril width is the narrowest distance between the nostrils. Int erorbital width is the least distance between the orbits including the orbital rim. Interrostral width is the narrowest distance between the rostral barbels. Inter lower lip width is the narrowest distance between the lateral portions o f the lower lips. All fin ray counts are given as follows: simple rays in Roman numerals followed by branched rays in Arabic numerals where dorsal fin and anal fin ray counts include the last ray split at the base represented by The caudal fin ray cou nt is the total number of branched rays. Small scales are counted as A single specimen for Homalopteroides modestus was cleared and double stained for bone and cartilage using the technique of Taylor and van Dyke (1985). Photographs were taken of prese rved specimens using Visionary Digital (Palmyra,

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30 Virginia) with Canon 40D and 5D cameras at UF and edited using Photoshop CS3 Coordinates were determined using maps and Google earth. Maps were constructed using Arc map version 9. 3. 1 in Arc GIS 9 th editi on. The abbreviation for alcoholic specimens is ALC and a ? represents lack of data or uncertainty of classification or locality Lengths were measured to the nearest 0.1 mm using digital calipers and taken o n the left side when possible. All measurments are given in millimeters (mm). Gape width is presented as a proportion of head length (HL), and eye length is presented as a proportion of interorbital width (IO). Head length and measurements of the body are given as proportions of standard length (SL). 4 00 individuals were examined in this study representing the 6 species belonging to Homaloptera 11 of 12 to Homalopteroides 3 of 4 to Homalopterula 6 of 9 to Balitoropsis Molecular Phylogenetic Methods Sequences of the nuclear Recombination Activating Gene 1 (Rag1) for phylogenetic analysis were compiled from Genbank, representing six genera and 10 species of Balitoridae (Table 2 1 ). Rag1 is the most species diverse gene for Homaloptera on Genbank. Outgroup taxa were the loaches Pangio elongata (Cobitidae) and Barbatula barbatula (Nemacheilidae). A multiple sequence alignment was assembled in Seaview 4 (Gouy et al. 2010) using a muscle algorithm (Edgar 2004). In assessing the systematic position of the genera of Balitorid ae, a Bayesian analysis was conducted using MrBayes 3.1 (Huelsenbeck & Ronquist 2001) with posterior probabilities estimated with a Mark ov chain Monte Carlo analysis. T o determine the best fit model of nucleotide evolution for the dataset, likelihood score s were compared in PAUP* ver. 4.b10 (Swofford 2002) and when scores were comparable, a Likelihood

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31 Ratio Test (LRT) was performed (Felsenstein 1981). Four Markov chain Monte Carlo reactions were run simultaneously for 1,000,000 gener ations under the GTR+I+ G model. The trees and likelihood scores were sampled at every 100 generations. The standard deviation for split frequencies was 0.003, and the overlay plot and trace files of both runs were analyzed in the program Tracer v1.5 (Rambaut & Drummond 2007). Th e MCMC runs reached converg ence after 100,000 generations. One thousand trees were discarded as burn in, and the remaining trees were used to construct a 50% majority consensus tree in Figtree v1.3.1 (Rambaut & Drummond 2010). Taxonomic Methods Species in this study are recognized using the phylogenetic species concept. Under this concept a species is the smallest monophyletic group that is diagnosable by unique character states.

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32 T able 2 1. Genbank accession numbers and sources for analyzed samples. Spe cies Genbank No. Source Balitora annamitica EF056359.1 et al 2007 Barbatula barbatula EU711107.1 Mayden et al 2008 Barbucca diabolica EF056391.1 et al 2007 Beaufortia kweichowensis EF056362.1 et al 2007 Homalo ptera leonardi EU711130.1 Mayden et al 2008 Homaloptera parclitella EF056358.1 et al 2007 Homaloptera smithi EF056356.1 et al 2007 Homaloptera zollingeri EF056388.1 et al 2007 Pangio oblonga EF056346.1 htov et al 2007 Pseudogastromyzon cheni EF056357.1 et al 2007 Pseudogastromyzon fasciatus EF056376.1 et al 2007 Sewellia lineolata EU409609.1 Chen et al 2008

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33 CHAPTER 3 MATERIALS EXAMINED I nstitutional A bbreviations Institutional abbreviations follow Eschmeyer and Fricke (2012) and are defined below. ANSP Academy of Natural Sciences, Philadelphia BMNH Natural History Museum, London CAS California Academy of Sciences, San Francisco MZB Museum Zoologicum Bogori ense, Cibinong NIFI, National Inland Fisheries Institute, Fish Taxonomy Division, Bangkok; RLIKU, Research Laboratory of Ichthyology, Kasetsart University, Bangkok, Thailand ROM, Royal Ontario Museum, Department of Natural History, Toronto; SU, Stanfor d University (The collection now resides at CAS); UF, University of Florida, Florida Museum of Natural History, Gainesville; UMMZ, University of Michigan Museum of Zoology, Ann Arbor; USNM, Smithsonian Institution National Museum of Natural History, Dep artment of Vertebrate Zioology, Division of Fishes, Washington D.C.; ZRC, Zoological Reference Collection, Department of Life Sciences, Faculty of Science, National University of Singapore, Singapore. Homaloptera Homaloptera bilineata : Myanmar: Sagaing Di vision: Irrawaddy River: CAS 231723, 4 ALC. Homaloptera confuzona : Thailand: Changwat P rov. : Trat River: UF 169906, 1 ALC. Thailand: Chantaburi Prov.: INHS 93493, 5 ALC. Homaloptera ocellata : Indonesia: Sumatra: Lampung : UF 161605, 1 ALC. UF 166104, 2 ALC. UF

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34 166107, 4 ALC. UF 161718, 2 ALC. Indonesia: Sumat ra: South Sumatra : UF 166096, 4 ALC. UF 166106, 2 ALC. Homaloptera ogilviei : Malaysia: Negri Sembilan: BMNH 1966.9.26.1 (paratype), 1 ALC. ZRC 15 53 (paratype), 2 ALC. Malaysia: Terengganu: ZRC 1555 (holot ype), 1 ALC. Malaysia: Perak: Perak Riv er: ZRC 1554 (paratype), 2 ALC. Indonesia: Sumatra: Lampung : UF 161716, 2 ALC. UF 166091, 2 ALC. Thailand: Surat Thani: Tapi River: INHS 93605, 7 ALC. Indonesia: Sumatra: South Sumatra: Musi River: UF 166085, 2 ALC. H omaloptera orthogoniata : Indones ia: West Kalimantan : Kapuas Basin: CAS 49326, 1 ALC. Homaloptera parclitella : Malaysia: Terengganu Prov.: Terengganu B asin: ZRC 49257 (holotype), 1 ALC. ZRC 47167 (paratype), 1 ALC. Malaysia: Perak: Tapah Fisheries Station: SU 39390, 2 ALC 6 of 6 species Homalopteroides Homalopteroides indochinensis : Vietnam: Indo China (? Tonkin) : BMNH 1933 8 19 50 (holotype, unique), 1 ALC. Homalopteroides modestus : Thailand: Kanch anaburi Prov.: Mae Khlong basin: ANSP 179826, 5 ALC, 14.78 25 N, 98.669167 E. NIFI 4508, 1 ALC, 14.719444 N, 98.505833 E. NIFI 4517, 1 ALC, 14.659722 N, 98.533611 E. UF 172926, 1 ALC, 14.661944 N, 98.7125 E. UF 173067, 1 ALC, 15.070556 N, 98.580556 E. UF 176377, 10 ALC, 14.752778 N, 98.500556 E. UF 176 408, 2 ALC, same collection data as NIFI 4517. UF 176438, 8 ALC, same collection data as NIFI 4508. UF 176454, 4 ALC, 14.534444 N, 98.787778 E. UF 176544, 1 ALC, 14.680278 N, 9 8.527222 E. UF 176557, 8 ALC, 14.689444 N, 98.513611 E. UF 181080, 5 ALC, 14.65 N, 98.566667 E. UF 181160, 9 ALC, 14.633333 N, 98.8 E. UF 181141, 1 ALC, 14.633333 N, 98.8 E. ZRC 53385, 1 ALC, same collection data as UF 176557. ZRC 53386, 1 ALC, same collection data as

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35 UF 181160. Tha iland: Tak Prov.: Salween Basin: NIFI 378 6, 1 ALC, 16.994614 N, 98.499494 E (coordinates estimated). NIFI 4514, 1 ALC, 16.779344 N, 98.710069 E (coordinates estimated). ROM 51147, 2 ALC, 16.9666667 N, 98.5666667 E. ZRC 41272, 4 ALC, 16.296417 N, 98.712472 E. Myanmar: Tanin tharyi Region: T enasserim Basin: ZRC 22889, 1 ALC, 13 N, 98.75 E (coordinates estimated). Myanmar: (?) Kayin State: (?) Salween basin: BMNH 1893.2.16.50 (paralectotype of Helgia modesta ), 1 ALC, 16.555556 N, 98.24 E (coordinates estimated). Homalopteroides n. sp : East Malaysia: Sarawak : Batang Balui USNM 323875 (holotype of Homalopteroides n. sp. ) 1 ALC, 2.416667 N, 113.766667 E USNM 323875 (paratopotype of Homalopteroides n. sp. ), 1 ALC USNM 323878 (paratype of Homalopteroides n. sp. ), 1 ALC, 2.416667 N, 11 3.733333 E. USNM 323879 (paratype of Homalopteroides n. sp. ), 2 ALC. Homalopteroides nebulosus : Malaysia: Kelantan: Sok River: BMNH 1967.11.15.15 (paratype of Homaloptera nebulosa ), 1 ALC. SU 66428 (paratype of Homaloptera nebulosa ), 1 ALC. ZRC 2020 (hol otype of Homaloptera nebulosa ), 1 ALC. ZRC 1759 (par at ype of Homaloptera nebulosa ), 1 ALC Homalopteroides cf. nebulosus : Thailand: Narathiwat: Sungai Kolok Basin: NIFI 3613, 5 ALC. Indonesia : West Kalimantan: Kapuas Basin. USNM 230250, 3 ALC Central Kali mantan: BMNH 2000.10.25.133 8, 3 ALC. East Malyasia: Sarawak: ZRC 45888, 3 ALC. Indonesia: Sumatra: UF 166083, 1 ALC UF 166084, 1 ALC UF 166088, 1 ALC. Sumatra: Way Tulangbawang: UF 162161, 2 ALC UF 161560, 2 ALC. Sumatra: Air musi: UF 166087, 1 ALC. Ho malopteroides rupicola : Myanmar: Myitkyna District: Sankha River: SU 28726 (paraty pe of Chopraia rupicola ), 1 ALC Homalopteroides smithi : Thailand: Nakhon Srithammarat: Ban Kiriwong:

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36 BMNH 1934.12.18.34, 1 ALC. USNM 109821 (paratype of Homaloptera smithi ), 5 ALC. UF 183330, 3 ALC UF 183411, 2 ALC. UF 18 3915, 1 ALC. Surathani Prov.: NIFI 3030, 3 ALC. ANSP 179981, 2 ALC. Trang Prov.: ANSP 76851, 3 ALC ANSP 76852, 1 ALC. Homalopteroides cf. smithi : Thailand: Phang gna Prov.: UF 182834, 4 ALC. UF 183738, 1 A LC. UF 183762, 2 ALC. Ranong P rov.: UF 183391 1 ALC. UF 183825, 5 ALC. ZRC 42185, 3 ALC. Chiang mai Prov.: UF 183403, 1 ALC. UF 177809, 1 ALC. USNM 107941, 1 ALC. USNM 229308, 5 ALC. Maehongson P rov.: NIFI 4511, 2 ALC Nan P rov.: NIFI 4512, 3 ALC. NIFI 45 18, 3 ALC. RLIKU 1500, 3 ALC. RLIKU 1626, 1 ALC. UMMZ 195111, 1 ALC. Bangkok Prov.: ANSP 56409, 1 ALC. ANSP 56410, 1 ALC. Chanthaburi Prov.: USNM 288450, 5 ALC. UF 169910, 5 ALC. RLIKU 1807, 5 ALC. RLIKU 1804, 1 ALC. UF 169907, 3 ALC. UF 169908, 3 ALC. J ohore P rov.: ZRC 7156, 2 AL C. Kanchanaburi Prov.: ANSP 179825, 1 ALC. UF 176454, 1 ALC. NIFI 4513, 3 ALC. UF 172923, 6 ALC. UF 183914, 1 ALC. Loei P rov. : ZRC 41314, 3 ALC. Narathiwat P rov.: ZRC 4376 9, 3 ALC.: Prachuap Khiri Khan Prov.: UF 181056, 8 ALC. UF 181069, 1 ALC. UF 1 81056, 1 ALC. Sa Kaeo Prov.: ANSP 179900, 3 ALC. INHS 93533, 3 ALC. Son gkhlan prov.: ZRC 43784, 3 ALC. ZRC 43796, 3 ALC. Ubon Ratchatani Prov.: AUM 56074, 4 ALC. UMMZ 233755, 1 ALC. NIFI 3436, 1 ALC. Indonesia: Sumatra: Lampung: UF 1616 19, 3 ALC. Laos: Attapeu p rov. CAS 92438, 1 ALC. CAS 233493, 1 ALC. Homalopteroides stephensoni : Indonesia : West Kalimantan : Sungai Pinoh: USNM 230254, 5 ALC Homalopteroides tweediei : Malaysia: Johore: Mawai: BMNH 1938.12.1.132 (paratype of Homaloptera tw eediei ), 1 ALC. SU 33012 (holotype of Homaloptera tweediei ), 1 ALC. SU 33013 (Paratypes of Homaloptera tweediei ), 2 ALC Homalopteroides

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37 wassinkii : Indonesia: Java: Buitenzorg: Tjampea: BMNH 1866.5.2.52 (lectotype of Homaloptera wassinkii ), 1 ALC. Indonesi a: Java: Lab. Binnenvisscherij (fishery): UMMZ 155660, 1 ALC. Homalopteroides weberi : East Malaysia : Baram River : ANSP 68718 (misidentified as Homaloptera wassinkii ), 11 ALC. East Malaysia: Sarawak: Akar River: BMNH 1895.7.2.81 (syntype of Homaloptera webe ri ), 7 ALC. East Malaysia: Sarawak: BMNH 1933.8.9.7 1 ALC BMNH 1978.9.5.45 47 3 ALC ROM 70456, 3 ALC ROM 70458, 3 ALC ROM 70464, 3 ALC ROM 70466, 3 ALC ROM 82115, 1 ALC. ROM 82131, 3 ALC. East Malaysia: Sabah: FMNH 99366, 1 ALC UMMZ 238960, 1 ALC. B runei : Darussalam : FMNH 117636, 2 ALC Homalopteroides yuwonoi : Indonesia: Kalimantan Barat: Ka p u as basin: MZB 5938 (holotype of Homaloptera yuwonoi ), 1 ALC. Homalopterula Homalopter ula gymnogaster : Indonesia: Sumatra: Lake Meninju: BMNH 1866.5.2.49 (holoty pe), 1 ALC. Indonesia : Sumatra Utara: USNM 390062, 2 ALC. Homalopterula heterolepis: Indonesia: Sumatra: Aceh Prov.: ZMA 100999 (Syntype of Homalopterula heterolepis ), Photo only. Homalopter ula ripleyi : Indonesia: Sumatra: Aceh P rov. : ANSP 68713 (holotype) 1 ALC ANSP 187003, 2 ALC. ANSP 187004, 1 ALC. US NM 390027, 2 ALC. Ind onesia: Sumatra: Sumatera Utara : ANSP 188907, 1 ALC. ANSP 188908 4 ALC. Homalopter ula vanderbilti : Indonesia: Sumatra: Tripa River: ANSP 68688 (holotype), 1 ALC. Balitoropsis Balitoro psis batek : Indonesia : Central Kalimantan: Katingan basin: MZB 10990 (holotype), 1 ALC. ZRC 51743 (paratypes), 5 ALC. Balitoropsis leonardi : Malaysia: Pahang : ZRC 1753 (paratype) 1 ALC. ZRC 7166, 1 ALC. Balitoropsis maxinae :

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38 Thailand: Bangkok Prov.: Tachi n: ANSP 68004 (holotype), 1 ALC. Balitoropsis ophiolepis : Indonesia: Java: Bandong: BMNH 1866.5.2.48 (paralectotype), 1 ALC. Indonesia: Sumatra: Musi: UF 166101, 1 ALC. UF 166103, 1 ALC. Indonesia: Sumatra: Tulang bawang: UF 166109, 4 ALC. Indonesia: West Kalimantan Prov.: USNM 230251, 1 ALC. Balitoropsis sexmaculata : Thailand: Chiang Mai P rov. : ANSP 56374 (holotype), 1 ALC. ANSP 56375 (paratypes), 5 ALC. UF 177819, 3 ALC. UF 183358, 2 ALC. UF 183398, 3 ALC. Balitoropsis zollingeri : Indonesia: Java: Bandong : BMNH 1866.5.2.53 (syntype) 1 ALC Indonesia: Sumatra: Lam pung Prov.: UF 166105, 1 ALC. Indonesia: West Kalimantan Prov.: CAS 49331, 1 ALC. Malaysia: Negri Sembilan: Jelai river: SU 66420, 1 ALC. Indonesia: Sumatra: Musi: UF 166094, 1 ALC. UF166095, 2 AL C. Indonesia: Sumatra: Tulang bawang: UF 166102, 1 ALC. UF 161715, 3 ALC. Malaysia: Pahang Prov.: SU 66424 (p aratypes of Homaloptera nigra ), 2 ALC. Thailand: Nakhon Si Thammarat: UF 183727, 1 ALC. Thai land: Trang prov. USNM 107963 (h olotype of B. bartschi ). Ind ia n G roup Homaloptera montana : India: Anamallai Hills: Southern India: CAS 39871 (holotype of Homaloptera montana ), 1 ALC.

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39 CHAPTER 4 ON THE PARAPHYLY OF HOMALOPTERA VAN HASSELT 1823 Homaloptera van Hasselt 1823 (Fig ure s 1 2A, 1 3A, 1 4A and 1 5 A ) Homaloptera van Hasselt, 1823: 133. (type species: Homaloptera ocellata van der Hoeven 1832, by subsequent monotypy in van der Hoeven, 1830: 211). Gender feminine. Helgia Vinciguerra, 1890: 328. (type species: Helgia bilineata Blyth, 1860: 172, by subsequ ent designation by Jordan 1920: 448). Gender feminine. Two species names are listed for Homaloptera in van Hasselt (1823), H. javanica and H. fasciata but were without descriptions or F igures. Hora (1932) visited Leiden in an effort to find any F igures t hat might have been drawn by Kuhl or van Hasselt for the fishes of Java, but was unsuccessful. The species names H. javanica and H. fasciata therefore are nomina nuda since they lack any descriptions or F igures (Hora 1932). The first species described and F igured under Homaloptera was H. ocellata van der Hoeven 1832, making H.ocellata the type species for Homaloptera Homaloptera bilineata Blyth 1860 was placed in the genus Helgia Vinciguerra (1890) and was subsequently designated as the type species of Hel gia by Jordan (1920), making Helgia Vinciguerra 1890 a synonym of Homaloptera van Hasselt 1823. Homaloptera ocellata was placed in the subgenus Homalopteroides by Weber and Beaufort (1916) with a description of the species as having a lateral line scale co unt of 43 47 and an origin of the dorsal fin posterior to the origin of the pelvic fin. However, the specimen examined by Weber and Beaufort ( MNHN 0000 3122 ) was later examined by Hora (1932) and identified as Homaloptera wassinkii Bleeker 1853. This speci men was designated as the holotype for Balitora ocellata Cuvier and Valenciennes

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40 1846. The species name for B. ocellata is a homonym of H. ocellata van der Hoeven 1832. Diagnosis Homaloptera is distinguished from Homalopter oides Homalopterula and Balit oropsis by the following combination of characters: color pattern consisting of very large red to red brown saddles on dorsum and upper side, origin of the dorsal fin anterior to the origin of the pelvic fin, line scales keeled scales, ventral surface of head covered with papillations and a mouth morphology consisting of two thick rostral barbels in close proximity to one another, triangular shaped and finely papillated lips, and the presence of a mental pad (Fig ure 1 4A ) Based on these characters the following species are recognized in Homaloptera van Hasselt 1823: Homaloptera ocellata van der Hoeven 183 2, H. bilineata Blyth 1860 H. orthogoniata Vaillant 1902 H. ogilviei Alfred 1967, H. confuzona Kottelat 2000 and H. parclitella Tan and Ng 2005 The type localities for species of Homaloptera are shown in Figure 4 1 Homaloptera is distinguished from Homalopteroides by having a cream colored body and a color pattern consisting of very large red to red brown saddles on dorsum and upper side vs. a cream colored body with a darker pattern of greenish tints and black s addles in living specimens (Figure 1 5 B); an origin of the dorsal fin anterior, rather than posterior, to the origin of the pelvic fin; line scales vs. 47 for Homalopter oides wassinkii ( Bleeker 1853 ) 39 44 H. modest us (Vinciguerra 1890), 44 for H. rupicola (Prashad & Mukerji 1929), 32 42 for H. smithi ( Hora 1932 ) 44 52 for H. stephensoni ( Hora 1932 ) 42 49 for H. weberi ( Ho ra 1932 ) 34 38 for H. tweediei ( Herre 1940 ) 44 for H. indochinensis ( Silas 1953 ) 38 40 for H. nebulos us ( Alfred

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41 1969 ) 42 43 (Arunkumar, 1998) for H. manipurensis (Arunkumar 1998), and 40 for H. yuwonoi ( Kottelat 1998 ); ventral surface of head covered w ith papillations vs. not covered; thick barbels with the medial and lateral rostral barbels in close proximity to one another vs. barbels thin and widely separated from one another at the base, triangular rather than crescent shaped lips; finely papillate d lips vs. smooth lips; and the presence vs. absence of a mental pad. Homaloptera is distinguished from Homalopterula by having a cream colored body with a color pattern consisting of very large red to red brown saddles on dorsum and upper side in living specimens vs. a cream colored body without reddish tints and with black saddles ( Figure 1 5 C); an origin of the dorsal fin anterior, rather than 49 50 for H gymnogaster (Bleeker 1853), more than 50 (Weber & de Beaufort, 1916) for H. heterolepis ( Weber & de Beaufort 1916) 40 47 for H. ripleyi Fowler 1940 and 56 for H. vanderbilti (Fowler 1940); scales with keels vs. scales without keels; barbels with the medial and lateral rostral barbels in close proximity to one another vs. barbels widely separat ed from one another at the base; triangular rather tha n crescent shaped lips; and the presence of a mental pad vs. mental lobes (1 4C). Homaloptera is distinguished from Balitoropsis by having a cream colored body with a color pattern consisting of very large red to red brown saddl es on dorsum and upper side in living specimens vs. a dark colored body without reddish ti nts and with black saddles (Figure 1 5 13 15 for B zollingeri (Bleeker 1853), 13 15 for B ophiolepis (Bleeker 1853), 14 (Tan, 2009) for B tateregani i (Popta 1905), 13 19 for B sexmaculata (Fowler 1934), 16 for B maxinae (Fowler 1937), 16 for

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42 B leonardi (Hora 1941), ? for B yunnanensis Chen 1978, 14 17 for B vulgaris (Kottelat and Chu 1988), and 17 19 (Tan, 2009) for B batek (Tan 200 9). Homalopteroides Fowler 1905 (Figure s 1 2 B 1 3 B 1 4 B and 1 5 B ) Homalopteroides Fowler, 1905: 476. ( t ype species: Homaloptera wassinkii Bleeker 1853, by original designation [the type species wa s misidentified, see below.]). Gender masculine. Chopraia Prashad & Mukerji, 1929: 188 ( t ype species: Chopraia rupicola Prashad & Mukerji 1929, by or iginal designation). Gender feminine. Fowler (1905) designated Homaloptera wassinkii Bleeker 1853 as the type species of Homalopteroides The description given by Fowler for H. wassinkii was inaccurate and has caused confusion for authors, including Weber and de Beaufort (1916). The specimens (ANSP 68718) examined by Fowler are identifiable as Homaloptera weberi Hora 1932 when compared to the seven syntypes of H. we beri (BMNH 1895.7.2.81). They all have an orbital length greater than or equal to the interorbital width, and a pelvic fin that extends past the anus. Fowler inaccurately gave a total pectoral fin ray count of 14 (v, 9) and an inaccurate count of 38 latera l line scales from the gill opening to the base of the caudal fin. Correct counts are 16 (v, 10, i) rays and 45 47 scales. Article 70.3 of the International Code of Zoological Nomenclature (ICZN 1999) addresses misidentified type species. According to the provisions of the article, the species that will best serve stability and universality can either be the nominal species cited and misidentified (Art. 70.3.1, H. wassinkii Bleeker 1853) or the taxonomic species examined (Art. 70.3.2, H. weberi Hora 1932). The genus Homalopteroides is valid, and

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43 both of these species are members. Homaloptera wassinkii Bleeker 1853 remains the type species of Homalopteroides Diagnosis Homalopteroides is distinguished from Homaloptera Homalopterula and Balitoropsis by the following combination of characters: dorsal fin origin posterior to a large orbital length a central furrow at the isthmus and a mouth morphology consisting of two thin and wid ely separated rostral barbels on each side of the mouth, thin crescent shaped lips, the absence of any structures such as a mental pad or lobes between the lateral portions of the lower lip, and a chin that extends anterior to the lateral portions of the l ower lip. Based on these characters, the following species are recognized in Homalopteroides : H. wassinkii (Bleeker 1853), H. modestus (Vinciguerra 1890), H. rupicola (Prashad & Mukerji 1929), H. smithi (Hora 1932), H. stephensoni (Hora 1932), H.weberi (Ho ra 1932), H. tweediei (Herre 1940), H. indochinensis (Silas 1953), H. nebulos us (Alfred 1969), H. manipurensis (Arunkumar 1998), and H. yuwonoi (Kottelat 1998). The type localities for species of Homalopteroides are shown in Figure 4 2 Homalopteroides is distinguished from Homaloptera by having the origin of the dorsal fin posterior, rather than anterior, to the origi n of the pelvic fin; line pored scales vs. 60 67 for H. ocellata 6 6 73 for H. bilineata 65 for H. orthogoniata 60 63 for H. ogilviei 63 64 for H. confuzona and 66 70 for H. parclitella ; a larger orbital length (70.1 146.7% of interobital width [IO]) vs 44.9 55% IO for H. ocellata 65.4 69.2% IO for H. bilineata 44.4% IO for H. orthogoniata 66.7 83.3% IO for H. ogilviei 35.6 52.3% IO for H. confuzona and 35.3 42.9% IO for H. parclitella ; presence vs. absence of a central furrow at the isthmus ; thin barbels with the medial and lateral

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44 rostral barbels widely separated from one another at the base vs. barbels thick and in close proximity to one another; crescent rat her than triangular shaped lips; and the absence vs. presence of a mental pad Homalopt eorides is distinguished from Homalopterula lateral line pored scales vs. 64 66 for H. gymnogaster 63 70 (Weber & de Beaufort 1916) for H. heterolepis 70 74 for H. ripleyi and 75 for H. vanderbilti ; scales vs. 49 50 f or H gymnogaster more than 50 (Weber & de Beaufort 1916) for H. heterolepis 43 47 for H. ripleyi and 56 for H. vanderbilti ; a larger orbital length vs. 60% IO for H. gymnogaster ? for H. heterolepis 46.8 64.3% IO for H. ripleyi and 30.6% IO for H. v anderbilti ; thin vs. thick lips; the absence vs. presence of mental lobes between the lateral portions of the lower lip ; and a chin that extends anterior to, rather than up to, the lateral portions of the lower lip. Homalopteroides is distinguished from Ba litoropsis by having the origin of the dorsal fin posterior, rather than above or anterior to the origin of the pelvic fin ; a larger orbital length vs 60 76% IO for B. zollingeri 47.1 65.5% IO, 47.1 61.5 % IO f or B. ophiolepis 38.5% IO ( Tan, 2009) for B tateregani i 43.8 78.9% IO for B. sexmaculata 55% IO for B. maxinae 50 60% IO for B. leonardi ? for B yunnanensis ? for B. vulgaris and 41.0% IO (Tan, 2009) for B. batek The same mouth characters listed above for distinguishing Homalopteroides from Homaloptera distinguish Homalopteroides from Balitoropsis except lip shape, which can be crescent shaped as in B. zollingeri ( Fig ure 1 4D )

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45 Homalopterula Fowler 1940 (Figures 1 2C, 1 3C, 1 4C and 1 5 C ) Homalopterula Fowler, 1940: 379. (type species: Hom aloptera ripleyi Fowler 1940:379, by original designation. Gender feminine. Although H. vanderbilti was described in the same publication as Homalopterula ripleyi Fowler (1940) designated Homaloptera ripleyi Fowler as the type species of Homalopterula an d assigned H. vanderbilti to Homaloptera rather than Homalopterula Homalopterula vanderbilti and is one of four species in the genus, known only from Sumatra. No other generic names have been proposed for any of these Sumatran species. Diagnosis Homalopt er ula is distinguished from Homaloptera Homalopteroides and Balitoropsis by the following combination of characters: dorsal fin origin posterior to the origin of the pelvic fin, 60 lateral line scales, 30 predorsal scales, a central furrow at the isthmus larger gape width (24.4 38.5% head length [HL]), and a mouth morphology consisting of two thick and widely separated rostral barbels on each side of the mouth, thick crescent shaped lips, presence of mental lobes between the lateral portions of the lower lip, a chin that extends up to the lateral portions of the lower lip and a truncate or deeply emarginated caudal fin. Based on these characters the following species are reco gnized in Homalopterula : H. gymnogaster ( Bleeker 1853 ), H. heterolepis ( Weber and Beaufort 1916 ), H. ripleyi Fowler 1940, and H. vanderbilti ( Fowler 1940 ). The type localities for species of Homalopterula are shown in Figure 4 3. Homalopter ula is distingui shed from Homaloptera by having the origin of the dorsal fin posterior, rather than anterior to, the origin of the pelvic fin; 30 predorsal scales vs. 20 24 for H. ocellata 24 27 for H. bilineata 21 for H. orthogoniata 20 22 for

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46 H. ogilviei 23 27 for H. confuzona and 24 for H. parclitella ; larger gape width (24.4 38.5% head length [HL]) vs. 19.6 23.5% HL for H. ocellata 15.1 18.1% HL for H. bilineata 20.3% HL for H. orthogoniata 14.5 20.6% HL for H. ogilviei 17.5 21.5% HL for H. confuzona and 17 .9 19.7%HL for H. parclitella ; presence vs. absence of a central furrow at the isthmus ; medial and lateral rostral barbels widely separated from one another at the base vs. barbels in close proximity to one another; crescent rather than triangular shaped lips; the presence of mental lobes vs. presence of a mental pad ; and a truncate or deeply emarginated caudal fin vs. forked Homalopter ula is distinguished from Homalopteroides by having 60 total lateral line pored scales vs. 47 for H wassinkii 39 44 f or H. modest us 44 for H. rupicola 32 42 for H. smithi 44 52 for H. stephensoni 42 49 for H. weberi 34 38 for H. tweediei 44 for H. indochinensis 38 40 for H. nebulos us 42 43 (Arunkumar 1998) for H. manipurensis and 40 41 for H. yuwonoi ; 30 predo rsal scales vs. 24 25 for H wassinkii 17 23 for H. modest us 19 for H. rupicola 16 22 for H. smithi 16 20 for H. stephensoni 14 21 for H. weberi 14 16 for H. tweediei ? for H. indochinensis 16 21 for H. nebulos us 18 21 (Arunkumar 1998) for H. mani purensis and 16 for H. yuwonoi ; t hick vs. thin barbels and lips; the presence vs. absence of mental lobes between the la teral portions of the lower lip; a chin that extends up to, rather than anterior to, the lateral portions of the lower lip ; and a trunc ate or deeply emarginated caudal fin vs. forked Homalopter ula is distinguished from Balitoropsis by having the origin of the dorsal fin posterior, rather than above or anterior, to the origin of the pelvic fin; 30 predorsal scales vs. 13 15 for B zollin geri 13 15 for B ophiolepis 14 (Tan 2009) for B

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47 tateregani i 13 19 for B sexmaculata 16 for B maxinae 16 for B leonardi ? for B yunnanensis 14 17 (Kottelat and Chu 1988) for B vulgaris and 17 19 (Tan 2009) for B batek ; larger gape width (24. 4 38.5% head length [HL]) vs. 19.3 23.9% HL for B zollingeri 15.6 20.2% HL for B ophiolepis ? for B tatereganii 18.5 22.1% HL for B sexmaculata 16.4% HL for B maxinae 22.2% HL for B leonardi ? for B yunnanensis ? for B vulgaris and ? for B batek ; medial and lateral rostral barbels widely separated from one another at the base vs. barbels in close proximity to one another; the presence of mental lobes vs. presence of a mental pad ; and a truncate or deeply emarginated caudal fin vs. forked. Balitoropsis Smith 1945 (Figures 1 2 D 1 3 D 1 4 D & 1 5 D ) Balitoropsis Smith 194 5 : 278 (type species: Balitoropsis bartschi Smith 1 945: 279 by original designation ) Gender feminine. ? Pseudohomaloptera Silas 19 53 : 204 (type species: Homaloptera tate reganii Popta 1905:180 by original designation ) Gender feminine. Kottelat and Chu (1988) recognized Balitoropsis bartschi as a junior synonym of H. zollingeri Although they never examined the holotype (USNM 107963), they concluded that B. bartschi is l ikely a gravid Homaloptera zollingeri Bleeker 1853. Based on photographs of USNM 107963 (Figures 4 4 ) and the type locality (Trang Province, Homaloptera tatereganii Popta 1905 was placed in the new genus Pseudohomaloptera by Silas (1953). Pseudohomaloptera was distinguished from Homaloptera by the presence of a deep rostral groove. Tan (2009) incorrectly interpreted Silas (1953) deep rostral groove for postoral groove and recognized

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48 Pseudo homaloptera as a junior synonym of Homaloptera (and by inference of Balitoropsis recognized as a subgenus of Homaloptera ) Balitoropsis is recognized here as a valid genus characterized by rostral and postoral grooves, and Pseudohomaloptera is a junior sy nonym. However, Tan (2009) gave a simple pelvic ray count of 3 for H. tatereganii whereas Silas (1953) gave a count of 2. Simple pelvic ray count is an informative character for the family Balitoridae. Most genera of the subfamily Balitorinae have 2 simple pelvic rays (some have > 2, e.g., Hemimyzon Regan 1911), (2009) count is accurate, the name Pseudohomaloptera may be valid. Until the holotype (RMNH 7632) can be examined H. tatereganii Popta 1905 is tentatively recognized in Balitoropsis Diagnosis Balitoropsis is distinguished from Homalopter a Homalopteroides and Homalopterula by the following combination of characters: a dark body with black saddles in living specim ens, origin of the dorsal fin anterior or above the origin of the pelvic fin, predorsal count 20, scales with keels ventral surface of head covered with papillations and a mouth morphology consisting of two thick rostral barbels in close proximity to one another, finely papillated lips, a nd the presence of a mental pad. Based on these character s the following species are recognized in Balitoropsis Smith 1945: B zollingeri (Bleeker 1853), B ophiolepis (Bleeker 1853), B tateregani i (Popta 1905), B sexmaculata (Fowler 1934), B maxinae (Fowler 1937), B leonardi (Hora 1941), B yunnanensis Chen 1978, B vulgaris (Kottelat and Chu 1988), and B batek (Tan 2009). The type localities for species in the subgenus Balitoropsis are shown in Figure 4 5

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49 Balitoropsis is distinguished from Homaloptera by having a dark body with black saddles in living spe cimens vs. a cream colored body with a color pattern consisting of very large red to red predorsal scales vs. 20 24 for H. ocellata 24 27 for H. bilineata 21 for H. orthogoniata 20 22 for H. ogilviei 23 27 for H. confuzona and 24 for H. parclitella Balitoropsis is distinguished from Homalopteroides by having an origin of the dorsal fin anterior to or above the origin of the pelvic fin vs. posterior to the origin of the pelvic fin ventra l surface of head covered with papillations vs. not covered, thick barbels with the medial and lateral rostral barbels in close proximity to one another v s. barbels thin and widely separate d from one another at the base finely papillated lips vs. smooth lips, and the presence vs. absence of a mental pad Balitoropsis is distinguished from Homalopterula by having an origin of the dorsal fin anterior to or above the origin of the pelvic fin vs. posterior to the origin of the pelvic s vs 49 50 for H gymnogaster more than 50 (Weber & de Beaufort 1916) for H. heterolepis 43 47 for H. ripleyi and 56 for H. vanderbilti ; scales with keels vs. scales without keels; barbels with the medial and lateral rostral barbels in close proximity to one another vs. barbels widely separate d from one another at the base; and the presence of a mental pad vs. mental lobes Indian group Five species placed in Homaloptera are known from the Western Ghats of India: H. montana Herre 1945, H. pillaii Indra and Rema Devi 1981, H. menoni Shaji and Easa 1995, H. santhamparaiensis Arunachalam et al 2002, and H. silasi Madhusoodana Kurup and Radhakrishnan 2011. These species have been recognized as possibly constituting a group requiring a separate genus (Pethi yagoda & Kottelat 1994 [first two

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50 species]; Kottelat 1998 [first three species] ); or subgenus (Tan & Ng 2005). Kottelat (2012) placed the Western Ghats species in Homaloptera Due to the inaccessibility of specimens from institutions in India, this group has had an unresolved taxonomic status. The holotype of H. montana (SU 39871) (Figure 4 6 ) can be distinguished from all species of Homaloptera by having the origin of the dorsal fin posterior to th e insertion of the pelvic fin. It has a mouth morphology ( Figure 1 8B) that is more similar to that of Homalopteroides and Homalopterula than to that of Homaloptera or Balitoropsis Homaloptera montana is most similar in morphology to the species of Homalopterula in having a chin that extends up to the lateral po rtions of the lower lip and a larger total lateral line pore count of 70 72. If all species of this group have the origin of the dorsal fin posterior to the insertion of the pelvic fin as does H. montana they are easily diagnosed from Homaloptera. The typ e localities for species in this Indian group are shown in Figure 4 7

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51 Figure 4 1. Type localities for species belonging to Homaloptera

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52 Figure 4 2. Type localities for species belonging to Homalopteroides

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53 Figure 4 3. Type localities for species belonging to Homalopterula

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54 Figure 4 4. Balitoropsis bartschi USNM 107963 (holotype). Scale bar represents 30 mm. Photos by Sandra Raredon.

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55 Figure 4 5. Type localities for species belonging to Balitoropsis

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56 Figure 4 6. (A) Dorsal, lat eral, and ventral views of Homaloptera montana CAS 139871, 46.4 mm SL; Scale bar represents 30mm; (B) ventral view of H. montana, CAS 139871. Photos by California Academy of Sciences Ichthyology Section

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57 Figure 4 7 Type localities for species of Homa loptera ( s.l

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58 CHAPTER 5 TAXONOMIC REVISION OF HOMALOPTEROIDES FOWLER 1905 Homalopteroides Fowler 1905 Homalopteroides consists of 12 species including a new species described herein. Known species of Homalopteroides occur in India, Myanmar, Thailand, Laos, Cambodia, Vietnam, Sumatra, Java, and Borneo All of these species share the following combination of charactres: origin of the dorsal fin posterior to the origin of the pelvic fin a large orbital length (70.1 146.7% IO), a central furrow at the isthmus and a mouth morphology consisting of two thin and widely separated rostral barbels on each side of the mouth, thin crescent shaped lips, the absence of any structure such as a mental pad or lobes between the lateral portions of the lower lip, and a chin that extends anterior to the lateral portions of the lower lip. Diagnoses and descriptions of species of Homalopteroides are given be low. Species are arranged chronologically by date of description. Diagnostic traits are given in Tables 5 1 and 5 2 Homalopteroides wassinkii ( Bleeker 1853 ) (Fig ure s 5 1 and 5 2; Table 5 3) Homaloptera wassinkii Bleeker 1853: 163,Natuurkundig Tijdschrift voor Nederlandsch Indi v. 4: 155 164. Lectotype. RMNH 4987; Tjampea (Ciampea), Buitenzorg (Bogor), Designated by Alfred 1961: 36; Bulletin of the Raffles Museum No. 30: 32 37. Paralectotypes. RMNH 1934 2 ALC, same designation as RMNH 4987 BMNH 1866.5.2 .52, 1 ALC, purchased from Bleeker; Java and Sumatra. Diagnosis Homalopteroides wassinkii is distinguished from all other species of Homalopteroides ( Table 5 1 and 5 2) by the combination of its doubly branched pelvic

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59 fin rays (3 or 4 distal points on one ray vs. 2), a total lateral line pore count of 47 20 circumpeduncular scales, presence of an axillary pelvic lobe 18 pectoral fin rays (v i, 11 i) and a pelvic fin not extending to the anus Homalopteroides wassinkii is morphologically most similar to H. smithi by having doubly branched pelvic fin rays, presence of an axillary lobe, 18 pectoral fin rays, 9 pelvic fin rays, and a pelvic fin not extending to the anus It is distinguished from H. smithi by having a higher total lateral line pore count of 4 7 vs. 35 42 and a higher circumpeduncular scale count of 20 vs. 16 18. It can be distinguished from all other species of Homalopteroides except for H. manipurensis (no specimens examined) by the presence vs. absence of doubly branched pelvic fin rays. It can further be distinguished by its higher total lateral line pore count of 47 vs. 39 44 for H. modestus 42 44 for H. rupicola 34 38 for H. tweediei 44 for H. indochinensis 38 40 for H. nebulosus 42 43 for H. manipurensis and 41 for H. yuwonoi ; the p resence of an axillary pelvic lobe vs. absence in H. modestus H. rupicola H. tweediei H. nebulosus and H. yuwonoi ; 18 pectoral fin rays vs. 15 for H. modestus 13 for H. tweediei 14 for H. nebulosus and 15 for H. manipurensis ; and a pelvic fin not ex tending to the anus vs. extending to the anus for H. stephensoni H. weberi H. indochinensis and H. yuwonoi Description Dorsal, lateral, and ventral views of an adult are shown in Figure 5 1. Measurements and meristic counts are given in Table 5 3 Homa lopteroides wassinkii is an average sized species for the genus reaching 46.2 mm SL. The body is arched predorsally, tapers posteriorly to the caudal fin base, and has a flattened ventral surface. When viewed dorsally the head is conical and is covered wit h small tubercles. The eyes are ovoid, positioned dorsolaterally midway between the snout tip and opercle

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60 edge, and smaller in length than the interorbital width. The nostrils are positioned closer to the snout tip than to the opercle edge. Origin of the d orsal fin is posterior to the origin of the pelvic fin and closer to the caudal fin base than to the snout tip. Pectoral fin reaches far past the pelvic fin origin and is longer than the head in length. Pelvic fin does not reach the anus and is doubly bran ched. Anal fin does not reach the caudal fin Axillary pelvic lobe present. Margin s of dorsal and anal fin s straight. The caudal fin is forked with rounded lobes where the lower lobe is slightly longer than the upper lobe. Body scaled except for the ventra l surface anterior to pelvic fin origin; scales between the pelvic fin insertion and origin are deeply embedded. Some scales, may have a small nipple at their posterior extremity. The total lateral line pore count is 47, predorsal count is 24 25, circumped uncular scale count is 20. Scale count s above and below the lateral line are 7 and 6, respectively. Scale count below the lateral line to pelvic fin origin is 7. Variations in fin counts are given in Table 5 3 ; modal numbers are given here. Dorsal fin ray s iii, 7; anal fin rays i, 5; pectoral fin rays vi, 11, i; pelvic fin rays ii, 7; total caudal fin ray count 17. The mouth (Figure 1 4B) is inferior with the lower jaw slightly visible. The lips are thin, smooth, crescent shaped, and continuous around th e corners of the mouth. The lateral portion of the lower lip is broad and the medial portion is thin. The chin extends anterior to the lateral portion of the lower lip. Rostral and postlabial grooves are present. Two pairs of rostral barbels and a pair of maxillary barbels are at each corner of the mouth. The medial rostral barbel reaches almost to the base of the lateral rostral barbel. The lateral rostral barbel does not reach the base of the maxillary barbel. The maxillary barbel reaches horizontally to a vertical up to the posterior nostril edge. The gill

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61 membrane is united to the isthmus with a large central furrow where the gills meet. The gill opening extends from the level of top of the orbit to the ventral surface of the body. Coloration In 70% e th anol: The general color is shown in Figures 5 1 and 1 4B. The faded. In dorsal view the base is cream and mottled brown. There are five brown dorsal saddles. The 1st is be tween the supraoccipital and dorsal fin origin, and is surrounded by 4 small blotches; 2nd is under the dorsal fin; 3rd is located between the dorsal fin insertion and anal fin origin; 4th spans most of the anal fin length; and 5th wraps the caudal fin bas e A preorbital bar extends from the lateral rostral barbel base to the eye. There is a brown mottled blotch anterior to the nares. There are brown blotches on the interorbital space arranged in a circular pattern. There are three large brown blotches ante rior to the supraoccipital ; one covers the post epiphyseal fontanelle and two are lateral to the fontanelle The lateral view shares the same base color pattern as the dorsal view. Series of mottled blotches extends from the preorbital bar to the top of th e head. The lateral line is covered by a solid brown stripe. A dark brown bar is present above the pectoral fin origin. The axillary pelvic lobe is mottled brown. In ventral view, the base is cream with mottled spots at the anal fin. The bases of the rostr al barbels are dark brown. The upper lip and the lateral aspect of the lower lip are mottled dark brown. All fins are hyaline with fine brown blotches. The following fins are banded brown: dorsal fin with 3, pectoral fin with 3, and pelvic fin with 2 bands The anal fin has 1 brown blotch at its origin and 2 bands. The caudal fin has 2 brown bands a proximal

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62 ba nd which reaches the superior extent of the upper lobe, and a distal V shaped band that is pointed anteriorly. The lower lobe of the caudal fin is n ot pigmented between the bands. The paired fins and caudal fin are brown at their bases, and the dorsal fin has a dark brown spot at its origin. Distribution Homalopteroides wassinkii was originally described from the ? Cisadane R iver basin, Ciampea Bogor Java (Figure 5 2). The other individual examined in this study is from an inland fishery at Tjisokan Homalopteroides modestus (Vinciguerra 1890) (Fig ure s 5 3, 5 4, 5 5, 5 6 and 5 7; Table 5 4) Helgia modesta Vinciguerra 1890: 330, Pl. 11 ( Figure 12); Annali del Museo Civico di Storia Naturale di Genova (Serie 2) v. 9. Lectotype. MSNG 15173A; Myanmar, Meekalan, Tenasserim; L. Fea, 1887. Designated by Tortonese 1961: 188; Catalogo del tipi de pesci del Museo Civico di Storia Naturale di Genova. (Parte I). Annali del Museo Civico di Storia Naturale `Giacomo Doria' v. 72: 179 191. Paralectotypes. BMNH 1893.2.16.50, 1 ALC; Myanmar, Meetan, Tenasserim; L. Fea, 1887. MSNG 15173B, 1 ALC (1, now 3); same collection data as BMNH 1893.2.16.50 ZMA 100982, 2 A LC; same collection data as BMNH 1893.2.16.50. Homaloptera modesta : Hora 1932: 288; Memoirs o f the Indian Museum, v. 12 (2). Diagnosis Homalopteroides modestus is distinguished from all other species of Homalopteroides (Table s 5 1 and 5 2) by the combi nation of its caudal fin color pattern that consists of a black proximal band a black distal band and a black median blotch, 15 pectoral fin rays (v, 9, i) (modal number, see Table 5 4 for variations in fin counts) a pectoral fin length greater than the head length, 5 6 scales above and 5 6 scales below the lateral line (to the pelvic fin), a total lateral line pore count of 39 44, absence

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63 of an axillary pelvic lobe, pelvic fin not extending to the anus, orbital length less than the inter orbital width ( an adult character, see Ch. 6 ), and a maxillary barbel that reaches to or slightly past the ante rior orbital rim. Homalopteroides modestus is morphologically most similar to H. nebulosus H. rupicola H. tweediei and H. manipurensis It is distinguished by having 15 pectoral fin rays vs. 13 14 in H. nebulosus 16 in H. rupicola and 13 in H. tweediei It can be distinguished further from H. nebulosus and H. tweediei by a pectoral fin length greater than head length vs. pectoral fin length less than head l ength, and from H. rupicola by an interorbital width greater than the orbital length vs. interorbital width less than or equal to the orbital length. It is distinguished from H. manipurensis by its lower scale count above the lateral line of 5 6 vs.8 and 7 branched dorsal fin rays vs. 6. Homalopteroides smithi and H. wassinkii are distinguished from H. modestus by the presence vs. absen ce of an axillary pelvic lobe. Homalopteroides indochinensis H. stephensoni H. weberi and H. yuwonoi are distinguished from H. modestus by a pelvic fin th at extends to or past the anus. Description Dorsal, lateral, and ventral views of an adult are shown in Figure 5 3A. Measurements and meristic counts are given in Table 5 4. Homalopteroides modestus is a moderate s ized s pecies reaching 44 mm SL. The body has a flattened ventral surface, is arched predorsally and tapers posteriorly to the caudal fin base. The head is conical when viewed dorsa lly and covered with tubercles. The orbits are small, ovoid, positioned dorsolater ally, and smaller in leng th than the interorbital width. The nostrils and orbital rim are in close pro ximity but are not in contact.

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64 The mouth (Figure 5 4) is inferior with both upper an d lower jaws slightly visible. The lips are thin, smooth, crescent s haped, and continuous ar ound the corners of the mouth. The lateral portion of the lower lip is broad and the medial portion is thin. The chin extends anterior to the lat eral portion of the lower lip. Posterior to the lower lip is a series of large tubercle s t hat extend onto the operculum. Rostral and p ostlabial grooves are present. Two pairs of rostral barbels and a pair of maxillary barbels are presen t at each corner of the mouth. The medial rostral barbels are separated from one another by a large lobe. T he rostral barbels are separated by a small lobe with a distance about equal to the lengt h of the medial rostral barbel. The lateral rostral barbel reaches the base of the maxillary barbel, and the maxillary barbel reaches horizontally to a vertical at or slightly posterio r to the anterior orbital rim. The gill opening extends to the ventral surface of the body, and the gill membrane is united to the isthmus with a large centra l furrow where the gills meet. The opercle reaches to or past the 2nd pectoral fi n ray. Body scaled except for the ven tral surface anterior to pelvic fin; scales between the anal fin origin and pelvic insertion are deeply embedded. In the cleared and stained specimen a medial row of scales from the pelvic fins reached to a small patch of sca les between the pectoral fins. Most scales, especially above the lateral line and at the dorsal fin origin, have a small nipple at their posterior extremity; up to six nipples are present on a scale. The total lateral line po re count is 39 44. Vari ations in fin counts are given in Table 5 4; modal numbers are given here. Dorsal fin has 3 simple and 7 branched rays (iii, 7 ) and originates posterior to the origin of the pelvic fin, closer to the caudal fin base than to the snout. Pectoral fin has 6 simple and 9 branched rays (v, 9, i), is longer than the head length and reac hes past the

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65 pelvic fin origin. Pelvic fin has 2 simple and 7 branched rays (ii, 7), lacks an axillary pelvic lobe, and does not reach the anus which is located closer to the anal fin origin tha n to the pelvic fin insertion. Anal fin has 1 simple and 5 branched rays (i, 5 ). The caudal fin is forked with rounded lobes and has a total of 17 branched rays; the lower lobe slightly longer than the upper. Total number of vertebrae is 3 1, compris ing 15 abdominal and 16 caudal. The fifth ceratobranchial ( Figure 5 5) bears a large transverse ventralis process a nd has a single row of 8 teeth. Positioning of the teeth varies in the left and right ceratobranchials in the cleared and stain ed s pecimen examined. Lateral view of a juvenile is shown in Figure 5 3B. Measurements and meristics for juveniles (individuals with an orbital length greater than or equal to the interorbita l width) are given in Table 5 4. The maximum size is 27 mm. Characte rs not sha red with adults are discussed. The body lacks a well defined predorsal arch and lobes between the barbels. Few tubercles may be pr esent by and between the nares. Nipples on the sca les are not prominently found. The pectoral fin reaches the pelvic fin, but does not always reach past it. Coloration In 70% e thanol: The general color is shown in Figures 5 3 and 5 4 The color pattern varies slightly between adults and juveniles. In dorsal view, the base color is cream with small mottled brow n blotche s. There are five black dorsal saddles. The 1st saddle may have smaller, incomplete saddles surrounding it and is between the supraoccipital and dorsal fin origin; 2nd is located at the dorsal fin origin; 3rd is located at the dorsal fin insertion between the dorsal fin and anal fin origins; 4th is dorsal to the anal fin; and 5th wraps the caudal fin base. The saddles often are faint and har d to

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66 distinguish in juveniles. A preorbital bar extends from the lateral rostral barbel base through the orbit and to the supraoccipital, which is outlined in black. The lateral view shares the same base co lor pattern as the dorsal view. A black bar extends from the medial rostral barbel ba se to or just before the nares. A black bar extends from the orbit to the posterio r edge of the opercle just below t he origin of the lateral line. The lateral line is marked by a solid black stripe that may contain circular blotches. One 4 large pigmented blotches may be present below the black stripe between the pectoral fin origin and t he anterior edge of the orbit. The 3rd and 4th saddles usually commence at the lateral line. Mottled brown blotche s sometimes coalesce to form distinct li nes above and below the lateral line and may form large blotche s between the pelvic and anal fins. In ventral view, the base is cream and, when mottled blotche s are present, they are restricted to a band at the anal fin origin. The barbel bases are black, and the lips are occasionally pigmented between the rostral barbels and at the maxillary barbels. All of the fins have black bands; the dorsal fin has 3 4 i n adults and 2 3 in juveniles. The pectoral fin has 2 3 bands, the pelvic fin has 1 2, and the anal fin has 1 2 in adults and 1 in juveniles. The caudal fin has 2 ba nds, a proximal band and a distal ba nd and a median blotch t hat may fuse to the distal band. The lower lobe of the caudal fin is pigm ented only at the caudal bands. The paired fins are black at their bases, and the unpaired fins have a black blotch at their origins. All fins have hyaline tips. Distribution Homalopteroides modestus occurs in the Salween, Mae Khlong, and Tenasserim basins ( Figure 5 6). The latter two basins drain the Tenasserim Range, a large

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67 mountain chai n between Thailand and Myanmar. The eastern part of the Tenasserim Ra nge is crossed by the Mae Nam Kwae Noi River (Mae Khlong basin) and the southwestern part gives rise to the Tenasserim River. The Salween River flows north of the Tenasserim Range. Other species that have a distribution in both the Mae Khlong and Salween b asins are Schistura mahnerti (Kottelat 1990; Plongsesthee et al. 2011), and Acanthocobitis pictilis (Kottelat 2012). Most individuals of H. modestus have been collected in mountain streams in rubble and gravel riffles (Figure 5 7). Homalopteroides modestu s has been reported from Borneo by Boulenger (1894) and Popta (1906); however, these populations were subsequently described as H. weberi and H. stephensoni by Hora (1932). The description given for H. modestus from Manipur, India (Selim and Vishwanath 199 8), does not agree with the data from this study and probably refers to a different species. Remarks Homalopteroides modestus was placed in the genus Helgia along with Homaloptera ( Homaloptera ) bilineata Bl yth 1860 by Vinciguerra (1890). Jordan (1920) sub sequently designated H ( Homaloptera ) bilineata as the type species of Helgia making Helgia Vinciguerra 1890 a synonym of Homaloptera van Hasselt 1823. Fowler (1905) designated Homaloptera wassinkii Bleeker 1853 as the type species for Homalopteroides to which H. modesta belongs. Homalopteroides modestus originally was described from the Meekalan and Meetan rivers, Tenasserim Division (now known as the Tanintharyi Region, Mon State and southern Kayin State) of southern Myanmar, and type specimens subsequen tly were designated by Tortonese (1961). The only type specimen examined in this study (due to availability) was a paralectotype, BMNH 1893.2.16.50. This specimen is in

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68 extremely poor condition and only pectoral fin and pelvi c fin ray counts could be made. Vinciguerra (1890) gave a total pectoral fin ray count of 13 (v, 8) and a total pel vic fin ray count of 8 (ii, 6). However, the paralectotype examined has a total pectoral fin ray count of 15 16 (v, 10 11) (only one side could be counted) and a total pel v ic fin ray count of 9 (ii, 7). Since its original description, H. modestus has been redescribed by Silas (1953), Menon (1987), a nd Selim and Vishwanath (1998). Silas (1953) gave a total pectoral fin ray count of 13 14 (v vi, 8) and the same pelvic fin ray cou nt given by Vinciguerra (1890). Menon (1987) gave the same counts as Silas (1953), whereas Selim and Vishwanath (1998) fol lowed the original description. Based on all material examined in the present study, H. modestus has a total pectoral fin ray count of 14 16 (iv vi, 8 10, i) with a modal number of 15 (v, 9, i). Silas (1953), Menon (1987), and Selim and Vishwanath (1998) gave a lateral line scale count of 47 although Vinciguerra (1890) cisare con esattezza: line cannot b e precise: there are about 47). The 69 individuals sampled in this study had a lateral line pore count of 38 42 +1 2. Vinciguerra (1890) gave a pharyngeal tooth count of 8 and described the 2nd to 4th teeth as being well developed, and the other five as rudimentary. In one of the specimens examined in this study, the pharyngeal tooth count was 8, the first six teeth being large and the last two small (Figure 5 5). Homalopteroides rupicola (Prashad & Mukerji 1929 ) (Fig ures 5 8 and 5 9; Table 5 5) Chopraia rupicola P rashad & Mukerji, 1929: 188, Pl. 8 (Figures 3 and 6 ); Records of the Indian Museum (Calcutta), v. 31, (pt 3)

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69 Holotype. ZSI F 10879/1; Myanmar, Myitky ina District, small rocky streams around Kamaing. Paratype. SU 28726, 1 ALC; Myanmar, Myitkyina District, Sankha, a large hillstream, midway between Kamaing and Mogaung; B. N. Chopra, 1926; Type catalog Bhlke, J. E. 1953: 40. Homaloptera rupicola : Hora 1932: 288; Memoirs of the Indian Museum, v. 12 (2). Diagnosis Homalopteroides rupicola is distinguished from all other species of Homalopteroides (Table s 5 1 and 5 2) by the combination of its 16 pectoral fin rays (v 10, i), total lateral line pore coun t of 42 44, absence of an axillary pelvic lobe, and a pelvic fin not extending to the anus Homalopteroides rupicola is morphologically most similar to H. modestus, H. tweediei and H. nebulosus by not having an axillary pelvic lobe, and a pelvic fin not extending to the anus. It is distinguished from them by 16 pectoral fin rays vs. 15 for H. modestus 13 for H. tweediei and 14 for H. nebulosus and is further distinguished from H. tweediei and H. nebulosus by a higher total lateral line pore count of 47 vs. 34 38 and 38 40, respectively. It can be distinguished from all other species of Homalopteroides except for H. yuwonoi and ? H. manipurensis by the absence vs. presence of an axillary pelvic lobe It can further be distinguished by 16 pectoral fin ray s vs. 18 for H. wassinkii 17 for H. smithi 17 for H. indochinensis and 15 for H. manipurensis ; and a pelvic fin not extending to the anus vs. extending for H. stephensoni H. weberi H. indochinensis and H. yuwonoi

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70 Description Lateral view is shown i n Figure 5 8. Measurements and merist ic counts are given in Table 5 5. See Prashad & Mukerji (1929 ) and Menon (1987) for a description of H. rupicola Distribution Homalopteroides rupicola occurs in the Irrawaddy River basin, Myitkyina District Myanmar (F igure 5 9). The holotype is from a small rocky stream around Kamaing, upper Myitkyina District Myanmar. Homalopteroides smithi (Hora 1932 ) (Fig ure s 5 10, 5 11and 5 12; Table 5 6) Homaloptera smithi Hora, 1 932 : 286 Pl. 11 (Figure 3); Records of the India n Museum (Calcutta), v. 12 (2 ) Paratypes. KUMF 0165, 3ALC; Thailand, Nakon Sritamarat, Ban Kiriwong, Tadi River; Smith, 1928; Type catalog Monkolprasit 1969:5 USNM 109821, 5 ALC (originally 6, 1 individual given to Hora); Thailand, Nakon Sritamarat, K long Pong Tadi Stream; Smith, 1928. Diagnosis Homalopteroides smithi is distinguished from all other species of Homalopteroides ( Table 5 1 and 5 2) by the combination of its doubly branched pelvic fin rays a total lateral line pore count of 35 42, 16 ci rcumpeduncular scales, presence of an axillary pelvic lobe 1 7 pectoral fin rays (vi, 10, i) (modal number, see Table 5 6 for variations in pectoral fin ray counts), and a pelvic fin not extending to the anus Homalopteroides smithi is morphologically mos t similar to H. wassinkii ( s ee H. wassinkii diagnosis). It can be distinguished from all other species of Homalopteroides except for H. manipurensis (no specimens examined) by the presence vs. absence of doubly branched pelvic fin rays. It can further be distinguished by the presence of an

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71 axillary pelvic lobe vs. absence in H. modestus H. rupicola H. tweediei H. nebulosus and H. yuwonoi ; 17 pectoral fin rays vs. 15 for H. modestus 13 for H. tweediei 14 for H. nebulosus and 15 for H. manipurensis ; a nd a pelvic fin not extending to the anus vs. extending for H. stephensoni H. weberi H. indochinensis and H. yuwonoi Description Dorsal, lateral, and ven tral views of a paratype (USNM 109821) and dorsal and lateral views of a recently collected indivi dual are shown shown in Figures 5 10 and 5 11. Measurements and merist ic counts are given in Table 5 6. See Hora (1932) for a description of H. smithi Distribution Homalopteroides smithi occurs in the Pakphanang Tape, Trang, and Phum Duang River basins southern Thailand ( Fig ure 5 12 ). The type locality is the Tadi River Ban Kiriwong, Nakon Sritamarat The Tadi River or stream (Smith 1945) arises from the Khao Luang, draining into the gulf of Thailand (? Pakphanang basin ). Homalopteroides smithi has b een reported from the Chao Phraya ( Fowler 1934 Kottelat 2001) and Mekong basins (Rainboth 1996, Kottelat 2001, Rainboth et al. 2012), and from Chantaboon (Chantaburi), Southeastern Thailand (Silas 1953); however, based on the material examined in this stu dy, these populations are likely new species Remarks Kottelat (2012) synonymized three species with Homalopteroides smithi : Homaloptera maxinae (with question), Homaloptera lineata, and Homaloptera indochinensis (with question). The holotype of H. maxin ae (ANSP 68004) was examined in this study and belongs to the genus Balitoropsis Balitoropsis maxinae is most similar to B. zollingeri by having a lower total lateral line pore count of 45. A

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72 taxonomic revision is needed for Balitoropsis to determine the status of B. maxinae The holotype for H. lineata (USNM 199488) was examined in this study, however the specimen is in poor condition, and no counts could be made. The type locality for H. lineata is in Chiang Rai, northern Thailand. Specimens from that ar ea were examined in this study and preliminary findings suggest that H. lineata is a valid species, as recognized by Rainboth et al (2012). The holotype for H. indochinensis (BMNH 1933.8.19.50) was examined in this study. The specimen is in poor conditio n and only a few counts and measurements could be made. Homalopteroides indochinensis is distinguished from H. smithi by having a pelvic fin that extends to the anus vs. not extending to the anus absence vs. presence of doubly branched pelvic fin rays to tal lateral line pore count of 44 vs. 35 42, and an orbital length that is equal to the interorbital width vs. smaller than the interorbital width. Homalopteroides stephensoni (Hora 1932 ) (Fig ure s 5 14, 5 15, and 5 16; Table 5 7) Homaloptera stephensoni Hora, 1932: 281, pl. 11 Figure 1 (type locality Indonesia: Borneo: Kalimantan Timur: Upper Mahakam River River). Holotype. RMNH 7633, unique, Tan 2009: 61, Figure 8. Diagnosis Homalopteroides stephensoni is distinguished from all other species of Homalo pteroides (Tables 5 1and 5 2) by its shallow caudal peduncle width of 6.1 6.6%SL (Fig ure 5 13). It can further be distinguished by the combination of smaller medial rostral barbels, total pelvic fin ray count of 10, a pelvic fin that extends up to or

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73 past the anus, total lateral line pore count of 46 52, and a smaller interorbital width of 21.3 23.2% HL. Homalopteroides stephensoni is morphologically most similar to H. weberi and H. yuwonoi by having a total pelvic fin ray count of 10, a pelvic fin that ext ends up to or past the anus, a smaller interorbital width and a shallower caudal peduncle. It is distinguished from H. weberi by having smaller medial rostral barbels ( Figure 5 14) and a shallower caudal peduncle of 6.1 6.6% SL vs. 6.7 8.0 % SL. It is dist inguished from H. yuwonoi by having a larger total lateral line pore count of 46 52 vs. 41, and the absence vs. presence of forward ly directed saddles on the side of the body (See H. yuwonoi ) It can be distinguished from all other species of Homalopteroid es by a total pelvic fin ray count of 10 vs. 8 9 and, except for H. indochinensis, by a pelvic fin that reaches up to or past the anus vs. not reaching the anus It can further be distinguished by a smaller interorbital width of 21.3 23.2% HL vs. 30 30.9% HL for H. wassinkii 25.8 34.2 % for H. modestus 27.2% HL for H. rupicola 26.5% HL for H. indochinensis 25.6 30.3% HL for H. nebulosus and 25.5 31.8% HL for H. manipurensis Description Dorsal, lateral, and ventral views are shown in Figure 5 15. Meas urements and meristic counts are given in Table 5 7. See Hora (1932) and Tan (2009) for a description of H. stephensoni Distribution Homalopteroides stephensoni occurs in the Mahakam and Kapuas River basins, Kalimantan, Indonesia ( Figure 5 16). The type locality is the upper Mahakam River, East Kalimantan.

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74 Homalopteroides stephensoni has been reported from Sabah ( Chin 1990 ; Martin Smith and Tan 1998 ) and Sarawak ( Parenti and Lim 2005 ), Malaysia; however, these populations are likely H. weberi Homalopte roides weberi (Hora 1932 ) (Fig ures 5 17, 5 18, 5 19, 5 20 ; Table 5 8) Homaloptera weberi Hora, 1932: 284, pl. 11 Figure 2 (type locality Malaysia: Borneo: Akar River). Syntypes. BMNH 1895.7.2.81, 7 ALC ZMA 100990, 1 ALC ZSI F11292, 1 ALC. Type catalog : Menon and Yazdani (1968:118 119), Nijssen et al. (1982:29). Diagnosis Homalopteroides weberi is distinguished from all other species of Homalopteroides (Tables 5 1 and 5 2) by the combination of total pelvic fin ray count of 10, a pelvic fin that extend s up to or past the anus, non reduced medial rostral barbels, caudal peduncle depth of 6.7 8.0 % SL, total lateral line pore count of 44 49, and a smaller interorbital width of 20.3 25.6% HL. Homalopteroides weberi is morphologically most similar to H. step hensoni and H. yuwonoi by having a total pelvic fin ray count of 10, a pelvic fin that extends up to or past the anus, a smaller interorbital width, and a shallower caudal peduncle. It distinguished from H. stephensoni by having non reduced medial rostra l barbels ( Fig ure s 5 14 and 5 17 ) and a deeper caudal peduncle of 6.7 8.0 % SL vs. 6.1 6.6% SL ( Figure 5 13 H. yuwonoi by having a higher total lateral line pore count of 44 49 vs. 41, and the absence vs. presence of forward ly dire cted saddles Homalopteroides by a total pelvic fin ray count of 10 vs. 8 9, and except for H. indochinensis, by a pelvic fin

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75 that reaches past the anus vs. not reaching the anus It can further be distinguished by a smaller interorbital width of 20.3 25.6% HL vs. 30 30.9% HL for H. wassinkii 25.8 34.2 % for H. modestus 27.2% HL for H. rupicola 26.5% HL for H. indochinensis 25.6 30.3% HL for H. nebulosus and 25.5 31.8% HL for H. manipu rensis Description Dorsal, lateral, and ventral views are shown in Figures 5 18 and 5 19 Measurements and meristic counts are given in Table 5 8. Homalopteroides weberi is a large s ized species reaching 58.2 mm SL. The body has a flattened ventral surfa ce, is slightly arched predorsally and tapers posteriorly to the caudal fin base. The head is conical when viewed dorsa lly and covered with tubercles. The orbits are large ovoid, positioned dorsolaterally, and greater than or equal to the interorbital wid th. The mouth ( Figure 5 17 ) is inferior with lower and at times upper jaws slightly visible. The lips are thin, smooth, crescent shaped, and continuous ar ound the corners of the mouth. The lateral portion of the lower lip is broad and the medial portion is very thin. The chin extends anterior to the lat eral portion of the lower lip. Rostral and p ostlabial grooves are present. Two pairs of rostral barbels and a pair of maxillary barbels are presen t at each corner of the mouth. The rostral barbels are separ ated by a small lobe with a distance about equal to the lengt h of the medial rostral barbel. The lateral rostral barbel reaches just up to the base of the maxillary barbel, and the maxillary barbel reaches horizontally to a vertical up to the posterio r nos tril rim. The gill opening extends to the ventral surface of the body, and the gill membrane is united to the isthmus with a large centra l furrow where the gills meet. Body scaled except for the ventral surface anterior to pelvic fin; scales anterior to a nus are deeply embedded and can be seen between the pelvics on the syntypes

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76 ( Figure 5 19 ). Most scales have a small centered nipp le at their posterior extremity In some specimens no nipples could be seen, possibly an artifact of preservation. The total la teral line po re count is 44 49. Variations in fin counts are given in Table 5 8 ; modal numbers are given here. Dorsal fin has 3 simple and 7 branched rays (iii, 7 ) and originates posterior to the origin of the pelvic fin, closer to the caudal fin base t han to the snout. Pectoral fin has 6 simple and 10 branched rays (v, 10 i), is longer than the head length and reac hes just anterior to, at, or a little past the pelvic fin origin. Pelvic fin has 2 simple and 8 branched rays (ii, 8 ), axillary pelvic lobe present and reach es at or past the anus Anal fin has 1 simple and 5 branched rays (i, 5 ). The caudal fin is forked with rounded lobes and has a total of 17 branched rays; the lower lobe slightly longer than the upper. Coloration In 70% e thanol: The ge neral color is shown in Figures 5 17, 5 18 and 5 19 In dorsal view, the base color is cream with small mottled brow n blotches. There are five to six dark brown dorsal saddles. The 1st saddle which may be comprised of two small saddles or surrounded by fou r small saddles is between the supraoccipital and dorsal fin origin; 2nd is located at the dorsal fin origin; 3rd which may be comprised of two small saddles is located at the dorsal fin insertion between the dorsal fin and anal fin origins; 4th is dorsal to the anal fin; and 5th is at the edge of the caudal fin base. There is a brown mottled blotch between the nares. A brown bar extends from between the lateral rostral barbel and maxillary barbel bases, through the orbit to the supraoccipital The lateral view shares the same base co lor pattern as the dorsal view. A brown bar may extend from between the medial and lateral rostral barbel ba ses to or just before the nares. A brown bar may extend from the maxillary barbel base to underneath

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77 the orbit at time s followed by a large blotch. Another brown bar extends from the orbit to the posterior edge of the opercle just below t he origin of the lateral line. The lateral line is marked by either a solid brown stripe that may contain circular blotches or is marke d by large brown blotches. The last two saddles usually commence at the lateral line. A dark brown bar above the pectoral fin origin is present. In ventral view, the base is cream and, when mottled blotche s are present, they are restricted to a band at the anal fin origin. The barbel bases are brown and the lips may be mottled brown on its dorsal edge All of the fins have brown bands; the dorsal fin has 2 3; pectoral fin has 3 that originate with large blotches; pelvic fin has 2 that may originate with l arge blotches; anal fin has 1 with a blotch anterior to it The caudal fin has 2 brown bands, a proximal band which may not always reach the superior extent of the upper lobe, and a distal V shaped band that is pointed anteriorly. The lower lobe of the cau dal fin is only pigmented at the bands. The paired fins and caudal fin are brown at their bases, and the dorsal fin has a brown blotch at its origin. Distribution Homalopteroides weberi occurs in the Baram and Batang River basins, Sarawak, Malaysia; the Se gama and Padas River basins, Sabah, Malaysia; and the Belait River basin, Brunei (Figure 5 20 ). The type locality is the Akar ( ? Akah) R iver, Baram River basin, Sarawak, Malaysia This species has been reported to inhabit fast waters with deep mud bottoms w ith rocks and to feed on immature aquatic insects (Inger and Chin 1962). Homalopteroides tweediei (Herra 1940 ) (Fig ure s 5 21 and 5 22 ; Table 5 9)

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78 Homaloptera tweediei Herre 1940: 7, Pl. 1 (Figure 12); Bulletin of the Raffles Museum No. 16. Holotype SU 330 12; Malaysia, Johore, Mawai District (? Kota Tinngi ), shallow rapid creek about 40 miles north of Singapore; Herre, 1937. Paratypes. SU 33013, 2 ALC BMNH 1938.12.1.132, 1 ALC ; same collection data as holotype. Homaloptera tweediei : Alfred 1969: Zoolog ische Mededelingen (Leiden) v. 43 (18). Diagnosis Homalopteroides tweediei is distinguished from all other species of Homalopteroides (Table s 5 1 and 5 2) by the combination of its 4 4 scales above and 3 4 scales below the lateral line (to the pelvic fin), a total lateral line pore count of 34 38 13 pectoral fin rays ( iv, 8 i) (modal number, see Table 5 9 for variations in pectoral fin ray counts) a pectoral fin length less than the head length orbital length greater than the inter orbital width a bsence of an axillary pelvic lobe, 9 pelvic fin rays, pelvic fin not extending to the anus, and a maxillary barbel that reaches horizontally to a vertical up to the posterior orbital edge Homalopteroides tweediei is morphologically most similar to H. neb ulosus by having a small pectoral fin ray count, small pectoral fin length large orbital length, absence of an axillary pelvic lobe, 9 pelvic fin rays, pelvic fin not extending to the anus, and a maxillary barbel that reaches horizontally to a vertical to midorbit It is distinguished from H. nebulosus by having 4 4 scales above and 3 4 scales below the lateral line (to the pelvic fin) vs. 5 6 scales above and 5 6 scales below and a total lateral line pore count of 34 38 vs. 38 40. It can be distinguis hed from all other species of Homalopteroides by 1 3 pectoral fin rays vs. 18 for H. wassinkii 15 for H. modestus 16 for H. rupicola 17 for H. smithi 16 17 for H. stephensoni 16 for H. weberi 17 for H.

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79 indochinensis 15 for H. manipurenesis and 16 fo r H. yuwonoi; and total lateral line pore count of 34 38 vs. 47 for H. wassinkii 39 44 for H. modestus 42 44 for H. rupicola 35 42 for H. smithi 46 52 for H. stephensoni 44 49 for H. weberi 44 for H. indochinensis 42 43 for H. manipurenesis and 41 for H. yuwonoi Description Dorsal, lateral, and ven tral views of holotype (SU 33012) and lateral view of a recently collected individual are shown shown in Figure 5 21 Measurements and merist ic counts are given in Table 5 9. See Herre (1940) and Alfred (1969) for a description of H. tweediei Distribution Homalopteroides tweediei occurs in the ? Ambat and Endau basins, Johore, Malaysia (Fig ure 5 22 ). The type locality is a shallow rapid creek about 40 miles north of Singapore ? Ambat basin Mawai Distr ict (? Kota Tinngi ), Johore, Malaysia. Homalopteroides tweediei has been reported from the Mekong basin (Rainboth 1996, Kottelat 2001, and Rainboth et al. 2012); however, based on the material examined in this study, this population is likely a new speci es. Homalopteroides indochinensis (Silas 1953 ) (Fig ures 5 23 and 5 24 ) Homaloptera sp from Indo China. Silas 1951: 12; Journal of the zoological society of India, v.3 (1). Homaloptera indochinensis Silas 1953: 192, Pl. Figure 2; Records of the Indian M useum (Calcutta) v. 50 (2) expedition to Indo China

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80 Diagnosis Homalopteroides indochinensis is distinguished from all species of Homalopteroides ( Tables 5 1 and 5 2 ) by the combin ation of its 9 pelvic fin rays, pelvic fin extending to the anus, orbital length equal to the inter orbital width presence of a pelvic axillary lobe, 17 pectoral fin rays ( v, 10 i i ), a total lateral line pore count of 44 and a maxillary barbel that reac hes horizontally to a vertical up to posterior nostril edge Homalopteroides indochinensis is distinguished from H. stephensoni H. weberi and H. yuwonoi by having 9 vs. 10 pelvic fin rays; from H. wassinkii H. modestus H. rupicola H. smithi H. tweedi ei and H. nebulosus by having a pelvic fin that extend es to the anus ; and from H. manipurensis by having an orbital length equal to the inter orbital width vs. less than inter orbital width Description Dorsal, lateral, and ventral views of holotype BMNH 1933.8.19.50 are shown in Figure 5 23. N o T able for m easurements or meristics is given here since the holotype is in poor condition and only a small number of counts and measurements could be made See Silas (1951,1953) for a description of H. indochinens is Distribution The type locality for Homalopteroides indochinensis is Indo China (? Tonkin) northern Vietnam (Fig ure 5 24 ). See Remarks. Remarks The holotype of Homalopteroides indochinensis was collected by Delacour and Lowe during an expedition to I ndo China. The specimen is without locality information, but other specimens from the expedition were collected from Pekh, N. Tonkin, Vietnam; Xien Khonang or Ching chuang, Laos; and Kontum and Dak to in Annam, Vietnam

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81 (Silas 1951). Silas (1953) was unsure of where the specimen came from and designated Tonkin (?), Vietnam, as the type locality. Kottelat ( 2012 ) suggested that Tonkin is unlikely to be the type locality of H. indochinensis since no species of Homalopteroides is known from that region, and that Kontum or Dakto, Sesan basin a tributary of the Mekong River Vietnam where species of Homalopteroides are known to occur is more likely Until fresh material of H. indochinesis is collected Tonkin, northern Vietnam, will be recognized as the type loc ality. See remarks in the H. smithi account for comments on H. indochinensis as a junior synonym. Homalopteroides nebulosus (Alfred 1969 ) (Fig ures 5 25 a nd 5 26 ; Table 5 10) Homaloptera nebulosa Alfred 1969: 227, Pl. 1 ( Figure 3 4); Zoologische Mededeli ngen (Leiden) v. 43 (18). Holotype. ZRC 2020; Malaysia, Kelantan, Sok River, Kampong Sok; Alfred 1966. Paratypes. ZRC 1759, 1 ALC BMNH 1967.11.15.15, 1 ALC SU 66428, 1 ALC RMNH 25971, 1 ALC; same collection data as Holotype. Diagnosis Homalopteroi des nebulosus is distinguished from all other species of Homalopteroides ( Table 5 1 and 5 2) by the combination of its 5 6 scales above and 5 6 scales below the lateral line (to the pelvic fin) a total lateral line pore count of 38 40, a maxillary barbel that reaches horizontally to a vertical up to midorbit, 1 4 pectoral fin rays (modal number, see Table 5 10 for variations in pectoral fin ray counts) 16 circumpeduncular scales, absence of an axillary pelvic lobe, 9 pelvic fin rays, pelvi c fin not extend ing to the anus

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82 Homalopteroides nebulosus is morphologically most similar to H. tweediei (see H. tweediei diagnosis). It can be distinguished from other species of Homalopteroides except for H. rupicola and H. manipurensis (no specimens examined) by a ma xillary barbel that reaches horizontally to a vertical up to midorbit vs. not reaching up to midorbit. It can further be distinguished from other species of Homalopteroides by a total lateral line pore count of 38 40 vs. 47 for H. wassinkii 42 44 for H. r upicola 46 52 for H. stephensoni 44 49 for H. weberi 44 for H. indochinensis and 42 43 for H. manipurenesis ; 14 pectoral fin rays vs. 18 for H. wassinkii 15 for H. modestus 16 for H. rupicola 17 for H. smithi 16 17 for H. stephensoni 16 for H. web eri 17 for H. indochinensis 15 for H. manipurenesis and 16 for H. yuwonoi ; 16 circumpeduncular scales vs. 20 for H. wassinkii and 18 20 for H. modestus ; absence of an axillary pelvic lobe vs. presence for H. wassinkii H. smithi H. stephensoni H. webe ri and H. indochinensis ; 9 pelvic fin rays vs. 10 for H. stepehnsoni H. weberi and H. yuwonoi ; and a pelvic fin not extending to the anus vs. extending for H. stepehnsoni H. weberi H. indochinensis and H. yuwonoi Description Lateral view of holotyp e (ZRC 2020) and paratype ( BMNH 1967.11.15.15 ) are shown in Figure 5 25. Measurements and meristic counts are given in Table 5 10. See Alfred (1969) for a description of H. nebulosus Distribution Homalopteroides nebulosus occurs in the Kelantan basin Kel antan Malaysia (Figure 5 26 ). The type locality is the Sok River, Sok village, Kelantan Malaysia.

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83 Homalopteroides nebulosus has been reported from Borneo, Kapuas basin (Roberts 1989); however, based on the material examined in this study, this populatio n is a new species. Homalopteroides yuwonoi (Kottelat 1998 ) (Fig ure s 5 27 and 5 28 ; Table 5 11 ) Homaloptera yuwonoi Kottelat 1998: 267, Figure 1; Ichthyological Exploration of Freshwaters v. 9 (3). Holotype. MZB 5938; Indonesia, Borneo, Kalimantan Barat, vicinity of Danau Sentarum Wildlife Reserve: Sungai Hulu Leboyan at Keluwin: 108'51"N, 11215'32"E; Kottelat et al., 1995. Paratype. CMK 11704, 1 ALC; same collection data as holotype. Diagnosis Homalopteroides yuwonoi is distinguished from all species o f Homalopteroides ( Table s 5 1 and 5 2) by its forward ly directed saddles on the side of the body. It can further be distinguished by the following combination of characters: absence of axillary pelvic lobe, total lateral line pore count of 41, 10 pelvic fi n rays, a pelvic fin that extends up to or past the anus, shallow caudal peduncle, and smaller interorbital width of 20.6% HL. Homalopteroides yuwonoi is morphologically most similar to H. stephensoni and H. weberi by having a total pelvic fin ray count o f 10, a pelvic fin that extends up to or past the anus, a smaller interorbital width and a shallower caudal peduncle. It is distinguished from both of them by its forward ly directed saddles on the side of the body, the absence vs. presence of an axillary pelvic lobe, and a smaller total lateral line pore count of 41 vs. 46 52 for H. stephensoni and 44 49 for H. weberi It can be distinguished from all other species of Homalopteroides by a total pelvic fin ray count of 10 vs. 8 9, and except for H. indochin ensis, by a pelvic fin that reaches up to or past the

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84 anus vs. not reaching the anus It can further be distinguished by a shallower caudal peduncle of 7.3% SL vs. 8.4 9.0% SL for H. wassinkii 9.1 11.5 % SL for H. modestus 8.7% SL for H. rupicola 7.7 91% SL for H. smithi 8.0 9.4% SL for H. tweediei 8.7% SL for H. indochinensis and 9.3 9.8% SL for H. nebulosus Description Lateral view of holotype ( MZB 5938 ) is shown in Figures 5 27. Measurements and meristic counts are given in Table 5 11 See Kottela t ( 1998 ) for a description of H. yuwonoi Distribution H. yuwonoi is known only from the the Kapuas Basin, western Kalimantan, Indonesia ( Fig ure 5 28). The type locality is the Hulu Leboyan River at Keluwin, Kapuas basin, in the vicinity of Danau Sentarum Wildlife Reserve western Kalimantan, Indonesia. Remarks The holotype (MZB 5938) of H. yuwonoi was examined in this study T he following results given in this study conflict with those in the type description (Kottelat 1998): orbital length greater than i nterorbital width vs. orbital length less than interorbital width and pectoral fin length less than head length vs. greater than head length. Homalopteroides manipurensis (Arunkumar 1998 ) (Fig ure 5 29 ) Homaloptera manipurensis Arunkumar 1998 : 176, Figure 1; Uttar Pradesh Journal of Zoology v. 18 (3) Holotype. MUMF 3333/1A; India, Manipur, Chindwin basin Lokchao River near Moreh, 110 km from Imphal City; L. Arunkumar, 1994. Paratype. MUMF 3333, 3 ALC; India, Moreh, Moreh Bazar; L. Arunkumar, 1994.

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85 Diag nosis Homalopteroides manipurensis is distinguished from all other species of Homalopteroides ( Table s 5 1 and 5 2) by the combination of 6 branched dorsal fin rays, 8 scales above the lateral line, 15 pectoral fin rays (v 10 ) a total lateral line pore co unt of 42 43 orbital length greater than the inter orbital width and 8 9 pelvic fin rays. Homalopteroides manipurensis is distinguished from all other species of Homalopteroides by having 6 branched dorsal fin rays vs. 7. It is further distinguished fro m species of Homalopteroides by having 8 scales above the lateral line vs. 7 for H. wassinkii 5 6 for H. modestus 6 for H. rupicola 5 6 for H. smithi 6 7 for H. stephensoni 4 4 for H. tweediei 5 6 for H. nebulosus and 6 for H. yuwonoi ; 15 pect oral fin rays vs. 18 for H. wassinkii 16 for H. rupicola 17 for H. smithi 16 17 for H. stephensoni 16 for H. weberi 13 for H.tweediei 17 for H. indochinensis ,14 for H. nebulosus and 16 for H. yuwonoi ; H. stephensoni H. weberi and H. yuwonoi by ha ving 9 pelvic fin rays vs. 10; a total lateral line pore count of 42 43 vs. 47 for H. wassinkii 46 52 for H. stephensoni 44 49 for H. weberi 34 38 for H tweediei 44 for H. indochinensis 38 40 for H. nebulosus and 41 for H. yuwonoi ; orbital length gr eater than inter orbital width vs. lesser for H. rupicola H. stephensoni H. tweediei or equal for H. indochinensis ; and 8 9 pelvic fin rays vs. 10 for H. stepehnsoni H. weberi and H. yuwonoi Description No specimens available for examination; see Ar unkumar (1998). Distribution H. manipurensis is known only from the Lokchao River and lower portion of the Maklang River, Yu River Basin, Manipur, India (Figure 5 29 ).

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86 Remarks No specimens of H. manipurensis were examined in this study, and the above diag nosis is based only on the orginal description ( Arunkumar 1998). Kottelat (2012) treated H. manipurensis as a junior synonym of H. rupicola with question. Homalopteroides manipurensis shares with H. rupicola a similar branched pelvic fin ray count of 8 9 v s. 9 10 in all other species of Homalopteroides Homalopteroides manipurensis is distinguished from H. rupicola by having a pectoral fin ray count of 15 vs. 16, a branched dorsal fin ray count of 6 vs. 7, 8 scales above the lateral line vs. 6, and an orb ital length less than the interorbital width vs greater than the orbital length. H. manipurensis might not belong to Homalopteroides since Arunkumar (1998) described it as having equal caudal lobes, a trait not seen in other species of Homalopteroides wher e the lower lobe is longer. Until specimens of can be examined Homalopteroides manipurensis is provisionally placed in the genus. Homalopteroides n. sp (Fig ure s 5 31, 5 32, 5 33 and 5 34 ; Table 5 12) Holotype: USNM 32387 5 1 ALC, 52.9 mm SL; East Malaysi a, Sarawak Batang Balui, Trib. Stream, long tow, where it enters Batang Balui, just downstream from logging camp ; 2.416667 N, 113.766667 E ; L. R. Parenti, A. Among, K. Luhat, A. Luhat; 06 August 1991 Paratopotype: USNM 323875, 1 ALC, 50.5 mm SL. Same locality information as holotype. Paratypes: USNM 323878, 1 ALC, 39.5 mm SL, East Malaysia, Sarawak, Batang Balui Trib. Stream, Jangan Aya, f lowing i nto Batang Besua. 2.416667 N, 113.733333 E. Same collecter information as holotype USNM 323879, 2 A LC, 36.8 37.4 mm SL; East Malaysia, Sarawak, Batang Balui, Trib. Stream Kemtu; 2.38 N, 113.75 E. Same collecter information as holotype.

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87 Diagnosis Distinguished from all other species of Homalopteroides (Table s 5 1 and 5 2 ) b y a larger gape width ( Fig ure 5 30 ) of 28.5 33.3% HL vs. 22.3 24.2% HL for H wassinkii 20.0 28.4% HL for H. modest us 19.1% HL for H. rupicola 20.3 27.5% HL for H smithi 20.0 23.2% HL for H. stephensoni 18.6 26.9% HL for H. weberi 18.4 22.6% HL for H. tweediei 17.3% HL for H. indochinensis 17.4 21.2% HL for H. nebulos us ? for H. manipurensis and 18.0 19.0% HL for H. yuwonoi It is further distinguished by a deeper caudal peduncle, 10.1 10.9% SL, vs. 8.4 9.0% SL for H wassinkii 8.7% SL for H. rupicola 7.7 9.1% SL for H. smithi 6.1 6.6% SL for H. stephensoni 6.7 8.4% SL for H. weberi 8.0 9.4% SL for H. tweediei 9.3 9.8% SL for H. nebulos us and 6.8 7.2% SL for H. yuwonoi and a higher circumpeduncular scale count of 20 22 vs. 16 for H. rupicola H. stephensoni H. twe ediei H. nebulos us H. manipurensis H. yuwonoi and 16 18 for H. smithi and H. weberi It is further distinguished from H. stephensoni H. weberi and H. yuwonoi by having a total pelvic fin ray count of 9 vs. 10. Description Dorsal, lateral, and ventral views of an adult are shown in Figure 5 31 Measurements and meristic counts are given in Table 5 12. Homalopteroides n. sp is a large species for the genus reaching 52.9 mm SL. The body is slightly deeper than wide, is arched predorsally, tapers posteri orly to the anal fin insertion, and has a flattened ventral surface. When viewed dorsally the head is conical and is covered with small tubercles. The eyes are ovoid, positioned dorsolaterally midway between the snout tip and edge of the opercle, and small er in length than the interorbital width. The nostrils are positioned closer to the snout tip than to the edge of the opercle. Origin of the dorsal fin is posterior to the origin of the the pelvic fin, closer to the caudal fin base

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88 than to the snout tip. P ectoral fin reaches slightly past the pelvic fin origin and may be a little longer than the head length. Pelvic fin does not reach the anus and is not doubly branched. Anal fin does not reach the caudal fin base. Axillary pelvic lobe is absent. Margins of dorsal and anal fins are straight. The caudal fin is forked with rounded lobes; the lower lobe is slightly longer than the upper lobe Body scaled except for the ventral surface anterior to the pelvic fin insertion; scales between the anal fin origin and pe lvic fin insertion are deeply embedded. Most scales, especially just posterior to the supraoccipital and at the dorsal fin origin, have a small nipple at their posterior extremity; up to five nipples on one scale have been counted. The total lateral line p ore count is 43 45, predorsal count is 22 25, circumpeduncular count is 20 22. Scale counts above and below the lateral line are 6 8 and 6 7, respectively. Scale count below the lateral line to pelvic fin origin is 6 7. Variations in fin counts are giv en in Table 5 12. Dorsal fin rays iii, 7; anal fin rays i ii, 5; pectoral fin rays iv v, 7 10, i; pelvic fin rays ii, 7; total caudal fin ray count 16 17. The mouth (Figure 5 32 ) is wide (28.5 33.3 % HL) and inferior with the lower jaw visible. The lips are thin, smooth, crescent shaped, and continuous around the corners of the mouth. The lateral portion of the lower lip is broad and the medial portion is thin. The inter lower lip width is large (20.0 22.6 % HL), and the chin extends anterior to the later al portion of the lower lip. Rostral and postlabial grooves are present. Two pairs of rostral barbels and a pair of maxillary barbels are at each corner of the mouth. The medial rostral barbel reaches just anterior to or at the base of the lateral rostral barbels. The lateral rostral barbel reaches a little more than half way the distance to the base of the maxillary barbel. The maxillary barbel reaches horizontally to a vertical at the

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89 anterior orbital rim or to mid orbit. The gill membrane is united to th e isthmus with a large central furrow where the gills meet. The gill opening extends from the level of mid orbit to the ventral surface of the body. Coloration In 70% ethanol: The general color is shown in Figures 5 31 and 5 32 In dorsal view the base is cream and mottled brown. There are five to six brown dorsal saddles. The 1st is between the supraoccipital and dorsal fin origin, is irregular, and may consist of up to five blotches; 2nd spans most of the pelvic fin length; 3rd is located between the dors al fin insertion and anal fin origin; 4th spans most of the anal fin length; and 5th is at the caudal fin there is an extra saddle between the 3rd and 4th saddles. A dark brown preorbit al bar extends from in between the medial and lateral rostral barbel bases and shifts laterally to meet the eye. There is a brown mottled blotch between and anterior to the nares. This blotch and pre orbital bar form a cream colored Y outline when viewed anteriorly. The eye is outlined in dark brown, and there may be random dark brown blotch dorsomedial to it. A dark brown bar covers the post epiphyseal fontanelle. Post orbital bars extend to the supraoccipital and to the posterior opercle edge in alignmen t with the lateral line. The lateral view shares the same base color pattern as the dorsal view. A dark brown bar extends horizontally from the posterior edge of the lateral rostral barbel to the posterior orbital edge where it continues upward diagonally for a small distance. In some individuals the upward extension of the bar is an unconnected dark brown blotch. Over the lateral line is a solid brown stripe that contains circular blotches in larger individuals. Mottled brown blotches sometimes coalesce to form distinct lines above and

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90 below the lateral line and may form large blotches between the pelvic fin and caudal fin base in larger individuals. The venter is cream with mottled blotches restricted to a band at the origin of the anal fin. The bases of t he rostral barbels are dark brown. The upper lip and the lateral aspect of the lower lip are mottled dark brown. All fins are hyaline with fine brown blotches. The dorsal fin has 2, pectoral fin has 1 2, and pelvic fin has 1 2 brown bands. The anal fin has 1 brown blotch that in one individual is restricted to the second ray. The caudal fin has 2 brown bands, a proximal band which may not always reach the superior extent of the upper lobe, and has a distal V shaped band that is pointed anteriorly. The lower lobe of the caudal fin is fully pigmented up to the posterior edge of the distal V shaped band. The paired and caudal fins are brown at their bases. The dorsal fin has a dark brown blotch at its origin. Distribution Homalopteroides n sp. is known from on ly the Balui River a tributary of the Rajang River, Sarawak Malaysia (Figure 5 33 ) These individuals were collected in streams with bottoms of mud, leaf litter, gravel, and boulders (Figure 5 34) The species w as collected in 1992 by Lynne Parenti and colleagues around 40 km SW of the Bakun hydroelectric dam prior to its construction. The current population status of this species is unknown.

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91 T able 5 1. Diagnostic characters for species of Homalopteroides Species that may have a fully pigmented lowe r lobe of the caudal fin are represented by ^. Total pelvic ray count Pelvic fin extending to or past anus (Yes/No) Doubly branched pelvic fin rays (Yes/No) Axillary pelvic lobe present (Yes/No) Total pectoral ray count (Modal) Pectoral fin length, >, = to, or < than head length Total branched dorsal ray count Number of dark bands on caudal fin H. wassinkii 9 N Y Y 18 > 7 2 H. modestus 9 N N N 15 > 7 2 1/3 H. rupicola 8 9 N N N 16 > 7 2 H. smithi 9 N Y Y 17 > 7 2^ 4^ H. stephensoni 10 Y N Y 16 17 > 7 2 3 H. weberi 10 Y N Y 16 > 7 2 H. tweediei 9 N N N 13 < 7 2 3 H. indochinensis 9 Y N Y 17 > 7 2 3

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92 T able 5 1. Continued Total pelvic ray count Pelvic fin extending to or past anus (Yes/No) Doubly branched pelvic fin rays (Yes/No) Axillary p elvic lobe present (Yes/No) Total pectoral ray count (Modal) Pectoral fin length, >, = to, or < than head length Total branched dorsal ray count Numbe r o f dark bands on caudal fin H. nebulosus 9 N N N 14 7 2 1/3^ H. yuwonoi 10 Y N N 16 < 7 2 H. manip urensis 8 9 ? ? ? 15 > 6 3 H. n. sp. 9 N N N 15 7 2^

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93 T able 5 2 Diagnostic characters for species belonging to Homalopteroides Circumpe d unc ular scale count Total latera l line pore count Scale count above and below lateral line to pelvic fin Ca udal peduncle depth % SL Orbital length >, = to, or < than interorbital width Interorbital width % HL Gape width % HL Maxillary barbel reaches vertically to H. wassinkii 20 47 7/7 8.4 9.0 < 30.0 30.9 22.3 24.2 Posterior nostril edge H. modestus 18 20 3 9 44 5 6/5 5 9.1 11.5 < (Adult) 25.8 34.2 20.0 28.4 At or slightly past orbital rim H. rupicola 16 42 44 6/6 8.7 > 27.2 19.1 Past orbital rim H. smithi 16 18 35 42 5 6/5 6 7.7 9.1 < (Adult and juvenile) 27.5 29.7 19.1 27.3 Posterior nostril edge to o rbital rim H. stephensoni 16 46 52 6 7/8 6.1 6.6 > 21.3 23.2 20.0 23.2 Anterior nostril edge H. weberi 16 18 44 49 6 8/6 8 6.7 8.0 >, < 20.3 25.6 20.2 26.9 Anterior to posterior nostril edges H. tweediei 14 16 34 38 4 4/3 4 8.0 9.4 > 19.0 2 4.6 18.4 26.0 Up to posterior orbital edge H. indochinensis ? 44 ? 8.7 = 26.5 17.3 Posterior nostril edge

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94 T able 5 2 Continued Circumpe d unc ular scale count Total latera l line pore count Scale count above and below lateral line to pelvic fin Caudal peduncle depth % SL Orbital length >, = to, or < than interorbital width Interorbital width % HL Gape width % HL Maxillary barbel reaches vertically to H. nebulosus 16 38 40 5 6/5 6 9.3 9.8 >, < 21.0 24.2 17.4 21.2 Midorbit H. yuwonoi 16 41 6/5 7.3 > 21.6 18 19 Midnostril H. manipurensis ? 42 43 8/? ? < 25.5 31.8 ? ? H. n. sp. 20 22 43 45 6 8/6 7 10.1 10.9 < 24.4 30.1 28.5 33.3 Up to midorbit

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95 Table 5 3. Morphometric measurements and meristic counts for Homalopteroides wassinkii (N=2) All m easurements in mm. N umber of individuals in parentheses Paralectotype represented by a *. Character Paralectotype n=1 Mean M orphometrics Standard l ength 35.6 46.2 % of s tandard l ength Head length 26.4 26.0 26.2 Body depth 14.9 16.0 15.5 Bo dy width 14.3 19.0 16.7 Distance between pelvic fin and anal fin 33.1 36.4 34.8 Distance between anus and anal fin 5.6 6.5 6.1 Dorsal fin base length 12.9 13.9 13.4 Dorsal fin length 21.3 21.6 21.5 Pectoral fin length 31.7 31.8 31.8 Pelvic fin length 23.0 23.4 23.2 Anal fin length 14.9 14.7 14.8 Predorsal length 54.2 55.2 54.7 Caudal peduncle length 15.2 15.4 15.3 Caudal peduncle depth 9.0 8.4 8.7 % of h ead l ength Head width 66.0 81.7 73.8 Head depth 41.5 43.3 42.4 Snout length 41.5 45.8 43.7 Nostril to operculum distance 61.7 60.8 61.3 Internostril width 22.3 23.3 22.8 Length of orbit 20.2 19.2 19.7 Interorbital width 30.9 30.0 30.4

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96 Table 5 3 Continued Character Paralectotype n=1 Mean Width of gape 22.3 24.2 23 .3 Pectoral fin length 120.2 122.5 121.4 % of i nterorbital w idth Length of orbit 65.5 63.9 64.7 M eristics Dorsal fin ray count iii, 7 (2) Pectoral fin ray count vi, 11, i (*); vi, 11, i & vi, 12, i (1) Pelvic fin ray count ii, 7 (2) Anal fin r ay count i, 5 (2) Caudal fin ray count 17 (1); N/A (*) Lateral line pore count 45+ 2 on caudal fin Circumpeduncle scale count 20 Number of scale rows above / below lateral line 7 / 6 Number of scale rows below lateral line to pelvic fin origin 7 Pre dorsal scale count 24 25

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97 Table 5 4. Morphometric measurements and meristic counts for adult (N=58 ) and juvenile (N=10) Homalopteroides modestus All measu rements in mm N umber of individuals in parentheses. Character Range(j uvenile) Range(a dult) Mean %SD(j uvenile) Mean%SD (a dult) M orphometrics Standard l ength 18.6 27.0 23.1 44.2 % of s tandard l ength Head length 25.4 30.6 24.1 28.7 27.31.55 26.51.10 Body depth 13.5 15.1 14.6 20.0 14.30.60 17.01.19 Body width 11.2 13.9 11.6 17. 9 12.20.83 14.71.14 Distance between anus and anal fin 3.8 6.0 3.0 5.4 4.80.83 4.10.64 Dorsal fin base length 9.1 11.5 9.0 13.8 10.40.68 11.51.10 Dorsal fin length 20.5 23.6 19.5 23.7 21.90.88 21.51.00 Pectoral fin length 28.2 31.6 26.6 32. 5 30.01.17 29.61.28 Pelvic fin length 19.4 22.2 19.1 23.8 21.20.92 21.51.26 Anal fin length 14.5 17.2 14.0 18.8 15.80.77 16.30.92 Predorsal length 52.6 57.7 52.4 57.4 54.01.56 54.81.21 Caudal peduncle length 13.5 16.8 12.1 17.4 15.01.30 14 .32.97 Caudal peduncle depth 9.1 10.4 9.2 11.5 9.70.42 10.50.52 % of head l ength Head width 57.2 74.0 61.3 76.5 67.46.08 69.83.58 Head depth 39.7 48.3 42.7 54.1 44.02.33 48.72.95 Snout length 32.1 45.7 37.0 50.5 41.23.92 44.72.29 Nostr il to operculum distance 60.5 71.5 58.0 69.6 66.33.10 64.52.60 Length of orbit 28.0 31.4 19.1 29.6 29.41.25 24.52.04 Interorbital width 25.8 31.4 26.1 34.2 28.81.70 29.81.71

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98 Table 5 4. Continued Character Range( j uveni le) Range( a dult) Mea n%SD( j uvenile) Mean%SD ( a dult) Inter lower lip width 13.9 17.2 11.9 19.9 15.91.09 15.31.75 % of interorbital width Length of orbit 100 113.6 70.1 95.9 102.34.27 82.97.08 Meristics Dorsal fin ray count iii, 7 (62); iii, 7 (5); iii, 8 (1) Pectoral fin ray count v, 9, i (45); v, 10, i (5); iv, 9, i (1); v, 9, ii (1); v, 10 (1); vi, 9, i (1); iv, 9, i & iv, 10 (1); v, 9, i & iv, 10 (1); v, 9, i & iv, 11 (1); v, 9 & v, 8, i (1);v, 9, i & v, 9 (2); v, 9, i & v, 10 (1); v, 9, i & v, 10, i (2 ); v, 9, i & vi, 8, i (1); v, 9, i & vi, 10 (1); vi, 9 & v, 9, i (1); vi, 9 & v, 10 (1); vi, 9, i & v, 10 (1) Pelvic fin ray count ii, 7 (61); ii, 6, i (5); ii, 5, ii (1); ii, 7, i & ii, 6, i (1) Anal fin ray count i, 5 (55); ii, 5 (11); N/A (2) Meri stics Caudal fin ray count 17 (59); 15 (5); 16 (1); N/A (2) Lateral line pore count 38 42 + 1 2 on caudal fin Lateral line pore at pelvic fin origin 14 17 Lateral line pore at dorsal fin origin 16 20 Lateral line pore at anal fin origin 29 33 Caudal peduncle scale count 17 20 Number of scale rows above / below lateral line 5 6 / 4 6 Number of scale rows below lateral line to pelvic fin origin 5 6 Predorsal scale count 17 22

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99 Table 5 5. Morphometric measurements and meristic counts for paratyp e of Homalopteroides rupicola Character Paratype Morphometrics Standard l ength 21.3 % of standard l ength Head length 27.7 Body depth 15.9 Body width 11.5 Distance between pelvic fin and anal fin 32.0 Distance between anus and anal fin 6.1 Dorsal fin base length 12.5 Dorsal fin length 22.7 Pectoral fin length 28.1 Pelvic fin length 20.3 Anal fin length 15.6 Predorsal length 53.9 Caudal peduncle length 14.6 Caudal peduncle depth 8.7 % of head l ength Head width 68.4 Head de pth 47.9 Snout length 40.9 Nostril to operculum distance 66.4 Internostril width 23.8 Length of orbit 28.9 Interorbital width 27.2 Width of gape 19.1

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100 Table 5 5 Continued Character Paratype % of interorbital w idth Length of orbit 106.2 Meristics Dorsal fin ray count iii, 7 Pectoral fin ray count v, 10, i Pelvic fin ray count ii, 6 Anal fin ray count i, 5 Caudal fin ray count N/A Lateral line pore count 40 42 + 2 on caudal fin Circumpeduncle scale count 16 Number of s cale r ows above / below lateral line 6 / 5 Num ber of scale rows below lateral line to pelvic fin origin 6 Predorsal scale count 19

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101 Table 5 6. Morphometric measurements ( N =19) and meristic counts (N =21) for Homalopteroides smithi All measurements in mm. N umber of individuals in parentheses Five s yntypes are included in meristic counts Character Syntypes (n=3) Juveniles(n=3) Adults(n=13) Mean%SD (n=19) Morphometrics Standard l ength 42.9 49.7 17.8 23.2 26.7 51.2 % of standard l ength Head len gth 23.9 27.9 28.4 29.7 25.4 28.9 27.21.49 Body depth 13.5 15.3 13.6 14.4 12.6 16.1 14.51.00 Body width 15.8 17.3 11.5 12.4 11.6 17.6 14.81.96 Distance between pelvic fin and anal fin 34.0 35.5 31.7 33.3 30.7 35.7 33.31.36 Distance between anus and anal fin 6.1 6.6 6.0 7.7 5.2 7.1 6.30.60 Dorsal fin base length 12.6 13.2 12.2 12.9 11.3 15.5 12.91.07 Dorsal fin length 21.3 22.9 21.7 23.0 20.4 23.0 22.00.85 Pectoral fin length 30.1 34.8 25.9 31.6 27.8 34.5 31.32.41 Pelvic fin length 2 1.7 25.7 19.0 22.8 18.6 25.2 22.11.9 Anal fin length 14.3 15.7 15.8 16.4 13.4 15.9 14.90.78 Predorsal length 52.6 59.2 53.1 58.3 53.1 57.9 55.01.92 Caudal peduncle length 13.2 16.8 13.8 17.6 13.3 17.2 15.01.42 Caudal peduncle depth 7.7 8.5 7.9 9.1 7.9 8.8 8.40.42 % of head l ength Head width 72.1 81.3 55.7 62.8 59.6 76.1 68.06.43 Head depth 43.6 45.9 42.8 44.9 38.8 45.9 43.72.21 Snout length 45.6 49.2 40.2 41.7 38.1 48.2 44.42.77 Nostril to operculum distance 59.3 63.2 65.3 67.6 59 .2 66.7 63.12.45 Internostril width 19.8 28.5 23.6 25.4 18.0 24.0 22.12.49 Length of orbit 20.5 22.8 25.8 28.5 20.2 27.2 23.42.24 Interorbital width 27.5 29.2 27.8 29.7 24.4 31.1 27.91.45 Width of gape 24.2 27.3 22.0 22.3 19.1 26.0 23.52.04

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102 T able 5 6 Continued Character Syntypes (n=3) Juveniles(n=3) Adults(n=13) Mean%SD (n=19) % of i nterorbital w idth Length of orbit 74.5 79.0 87.8 97.4 64.9 95.1 84.18.32 Meristics Dorsal fin ray count iii, 7 (21) Pectoral fin ray count vi, 10, i (9); vi, 11, i (4); vi, 10, i & v, 11, i (3); vi, 11, i & v, 12, i (1); vi, 10, i & vi, 9, i (2); vi, 10, i & vi, 11, i (1); vi, 10, i & vi, 10, ii (1) Pelvic fin ray count ii, 7 (21) Anal fin ray count i, 5 (19*); ii, 5 (2) Caudal fin ray count 17 (21) Lateral line pore count 34 41 + 1 on caudal fin Circumpeduncle scale count 16 (20*); 18 (1) Number of scale rows above / below lateral line 5 6 / 4 6 Number of scale rows below lateral line to pelvic fin origin 5 6 Predorsal scale count 16 22

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103 Table 5 7. Morphometric measurements and meristic counts for Homalopteroides stephensoni ( N = 5 ) All measurements in mm. N umber of individuals in parentheses Character Range Mean%SD Morphometrics Standard l ength 42.1 55.4 % of standard l eng th Head length 23.5 25.0 24.70.67 Body depth 11.6 13.0 12.30.56 Body width 13.6 18.2 15.01.91 Distance between pelvic fin and anal fin 31.6 33.3 32.60.66 Distance between anus and anal fin 11.4 14.0 12.91.04 Dorsal fin base length 13.1 1 5.4 14.30.94 Dorsal fin length 21.4 23.6 22.00.94 Pectoral fin length 26.1 27.6 26.90.64 Pelvic fin length 19.7 21.4 20.70.70 Anal fin length 13.2 14.0 13.50.32 Predorsal length 51.0 51.9 51.50.32 Caudal peduncle length 15.3 16.7 16.10.5 8 Caudal peduncle depth 6.1 6.6 6.40.21 % of h ead l ength Head width 61.9 67.0 64.62.45 Head depth 40.0 42.6 41.11.28 Snout length 44.7 48.5 46.51.44 Nostril to operculum distance 56.9 63.8 61.52.64 Internostril width 15.2 20.2 17.91.96 Length of orbit 22.3 25.5 24.21.21 Interorbital width 21.3 23.2 21.90.75 Width of gape 20.0 23.2 21.21.33

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104 Table 5 7 Continued Character Range Mean%SD % of interorbital w idth Length of orbit 103.4 120.0 110.67.09 M eristics Dorsal fin ray count iii, 7 (5) Pectoral fin ray count v, 11 (2); v, 11, i (1); v, 11, i & vi, 11, i (1); v, 10, i (1) Pelvic fin ray count ii, 8 (5) Anal fin ray count i, 5 (5) Caudal fin ray count 17 (5) Lateral line pore count 44 51 + 1 2 on caudal fin Ci rcumpeduncle scale count 16 Number of scale rows above / below lateral line 6 7 / 5 7 Number of scal e rows below lateral line to pelvic fin origin 7 8 Predorsal scale count 16 20

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105 Table 5 8. Morphometric measu rements and meristic counts for Homalopte roides weberi ( N = 31 ) All measurements in mm. N umber of individuals in parentheses Character Syntypes (n=7) Range(n=24) Mean%SD (n=31) Morphometrics Standard l ength 23.6 26.7 28.1 58.2 % of s tandard l ength Head length 23.6 26.7 22.2 28.3 25 .31.30 Body depth 11.5 12.8 11.0 15.7 12.50.98 Body width 12.2 15.3 12.4 17.7 14.51.31 Distance between pelvic fin and anal fin 31.2 35.2 30.3 35.0 33.31.29 Distance between anus and anal fin 11.9 13.5 10.7 16.4 12.51.19 Dorsal fin base lengt h 12.3 15.0 12.2 14.9 13.60.77 Dorsal fin length 20.9 24.4 21.1 24.2 22.60.96 Pectoral fin length 24.9 30.9 25.9 32.5 28.71.71 Pelvic fin length 20.5 23.4 19.9 23.2 21.50.96 Anal fin length 13.7 15.8 13.2 15.2 14.30.63 Predorsal length 51.0 52.5 49.1 54.7 51.90.96 Caudal peduncle length 14.4 18.2 12.3 17.2 15.31.28 Caudal peduncle depth 6.9 7.8 6.7 8.0 7.20.30 % of h ead l ength Head width 61.7 72.5 59.8 77.9 67.54.44 Head depth 38.3 43.5 33.3 52.6 41.83.85 Snout length 41.9 4 8.9 43.2 52.3 46.32.43 Nostril to operculum distance 59.0 65.5 58.7 67.9 61.31.92 Internostril width 17.0 20.7 16.7 21.6 18.71.27 Length of orbit 23.4 26.1 20.7 27.0 24.11.85 Interorbital width 20.6 25.5 20.3 25.6 22.61.40 Width of gape 22.0 25.8 20.2 26.9 22.91.59

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106 T able 5 8 Continued Character Syntypes (n=7) Range(n=24) Mean%SD (n=31) % of Interorbital Width Length of orbit 100 120 90.6 133.3 107.29.59 M eristics Dorsal fin ray count iii, 7 (31) Pectoral fin ray count v, 10, i (14); v, 10, i & v, 9, i (2); v, 10, i & v, 11, i (2); v, 10, i & vi, 10, i (2); v, 11, i (1); v, 10, i & vi, 9 (1); v, 11, i & vi, 10, i (1); v, 10, i & vi, 10 (1); v, 10, i & v, 8, i (1); vi, 9, i & vii, 9, i (1); v, 10. i & v, 10, ii (1); v, 10, i & v, 9 (1); v, 10, i & iv, 11, i (1) Pelvic fin ray count ii, 8 (31) Anal fin ray count i, 5 (16); ii, 5 (13) Caudal fin ray count 17 (29);16 (2) Lateral line pore count 42 47 + 1 2 on caudal fin Circumpeduncle scale count 16 18 Number of scale rows above / below lateral line 6 8 / 5 7 Number of scale rows below lateral line to pelvic fin origin 6 8 Predorsal scale count 15 21

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107 Table 5 9. Morphometric measu rements and meristic counts for Homalopteroides tweediei ( N = 7 ) All measurements in m m. N umber of individuals in parentheses Holotype represented by a *. Character Holotype Range (n=6) Mean%SD (n=6) Morphometrics Standard l ength 26.6 23.1 26.0 % of standard l ength Head length 27.1 28.1 30.4 28.80.82 Body depth 14.3 13.4 1 5.7 14.31.05 Body width 11.7 11.8 14.1 12.70.86 Distance between pelvic fin and anal fin 30.7 33.2 32.30.95 Distance between anus and anal fin 6.6 8.6 7.70.78 Dorsal fin base length 11.0 12.3 11.80.49 Dorsal fin length 21.3 25.5 23.21.4 6 Pectoral fin length 23.7 23.7 30.0 26.02.43 Pelvic fin length 19.9 16.8 19.6 18.31.14 Anal fin length 13.6 16.2 15.01.08 Predorsal length 51.1 52.0 55.3 54.11.23 Caudal peduncle length 17.3 13.7 17.3 15.21.45 Caudal peduncle depth 9.4 8 .0 9.3 8.60.47 % of head l ength Head width 61.1 57.2 62.5 59.52.12 Head depth 41.1 46.6 43.42.17 Snout length 30.6 35.6 39.4 38.11.32 Nostril to operculum distance 63.0 68.5 65.91.97 Internostril width 25.0 20.9 26.0 23.32.16 Length of orbit 27.8 25.7 28.8 27.31.10 Interorbital width 20.8 19.0 24.6 22.22.29 Width of gape 19.4 18.4 26.0 20.92.72

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108 T able 5 9 Continued Character Holotype Range (n=6) Mean%SD (n=6) % of i nterorbital w idth Length of orbit 133.3 110.6 142.7 124 .212.5 M eristics Dorsal fin ray count iii, 7 (5); N/A (1*) Pectoral fin ray count iv, 8, i (4); iv, 9 (1); iv, 9, i & iii, 10 (1); N/A (*) Pelvic fin ray count ii, 7 (7) Anal fin ray count i, 5 (4); N/A (2*) Caudal fin ray count 17 (5); N/A (1*) Lateral line pore count 33 37+ 1 on caudal fin Circumpeduncle scale count 16 (5*), 14 (1) Number of s cale rows above / below lateral line 4 4 / 3 4 Num ber of scale rows below lateral line to pelvic fin origin 3 4 Predorsal scale count 14 16

PAGE 109

109 Table 5 10. Morphometric measurements and meristic counts for Homalopteroides nebulosus ( N = 4 ) All measurements in mm. N umber of individuals in parentheses Holotype represented by a *. Character Holotype Range (n=3) Mean%SD (n=3) Morphometrics Stan dard l ength 23.6 21.2 22.2 % of s tandard l ength Head length 28.0 28.0 29.7 29.00.74 Body depth 15.3 15.1 16.0 15.40.41 Body width 12.7 11.3 13.2 12.40.80 Distance between pelvic fin and anal fin 32.2 29.2 32.3 31.11.46 Distance between a nus and anal fin 6.8 4.7 8.0 6.31.42 Dorsal fin base length 12.2 14.0 13.11.28 Dorsal fin length 24.0 24.1 24.00.10 Pectoral fin length 27.5 25.2 29.9 27.51.91 Pelvic fin length 21.2 19.7 21.2 20.40.67 Anal fin length 16.9 14.5 16.9 15.7 1.01 Predorsal length 53.4 53.2 54.7 53.70.67 Caudal peduncle length 13.1 13.1 14.5 13.80.61 Caudal peduncle depth 9.3 9.3 9.8 9.60.22 % of h ead l ength Head width 60.6 64.5 62.62.76 Head depth 42.4 40 46.0 42.22.74 Snout length 34.8 40 .3 37.63.87 Nostril to operculum distance 66.7 66.5 67.7 67.20.67 Internostril width 24.2 21.0 24.2 22.61.38 Length of orbit 27.3 27.3 27.4 27.30.08 Interorbital width 30.3 25.6 30.3 27.72.35 Width of gape 21.2 17.4 21.2 19.31.55

PAGE 110

110 T able 5 10 Continued Character Holotype Range (n=3) Mean%SD (n=3) % of i nterorbital w idth Length of orbit 90 94.4 106.5 99.38.37 M eristics Dorsal fin ray count iii, 7 (1*); N/A (2) Pectoral fin ray count v, 9 (*); iv, 9,i (1); v, 7, i (1); v, 8, i ( 1) Pelvic fin ray count ii, 7 (2*); N/A (1) Anal fin ray count ii, 5 (*); i, 5 (1) N/A (2) Caudal fin ray count 17 (1); N/A (2*) Lateral line pore count 36 39+ 1 2 on caudal fin Circumpeduncle scale count 16 (4) Number of scale rows above / below la teral line 5 6 / 4 5 Num ber of scale rows below lateral line to pelvic fin origin 5 6 Predorsal scale count 16 21

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111 Table 5 11. Morphometric measurements and meristic counts for holotype of Homalopteroides yuwonoi All measurements in mm. Characte r Holotype Morphometrics Standard l ength 38.6 % of s tandard l ength Head length 26.4 Body depth 15.0 Body width 15.3 Pectoral fin length 25.4 Pelvic fin length 22.3 Anal fin length 15.3 Predorsal length 54.1 Caudal peduncle length 18.4 Caudal peduncle depth 7.3 % of Head Length Head depth 52.0 Length of orbit 22.5 Interorbital width 21.6 % of i nterorbital w idth Length of orbit 104.5 M eristics Dorsal fin ray count iii, 7 Pectoral fi n ray count v, 10 i Pelvic fin ray count ii, 8 Anal fin ray count i, 5 Caudal fin ray count 17 Lateral line pore count 39 + 2 on caudal fin Circumpeduncle scale count 16 Predorsal scale count 16

PAGE 112

112 Table 5 12. Morphometric measurements and me ristic counts for Homalopteroides n. sp (N =5) All measurements in mm. N umber of individuals in parentheses. H olotype is represented by a Character Holotype Range (n=4) Mean%SD (n=4) Morphometrics Standard l ength 52.9 36.8 50.5 % of s tandard l ength Head length 25.8 25.9 28.0 26.80.94 Body depth 19.0 15.5 17.1 16.30.72 Body depth at anus 13.7 13.0 13.8 13.30.34 Body width 17.2 13.9 17.2 15.51.60 Predorsal length 53.9 52.9 54.3 53.80.62 Prepectoral length 20.4 20.3 21.3 20.8 0.46 Prepelvic length 46.3 45.1 46.1 45.70.52 Preanal length 79.0 77.0 79.0 78.20.85 Pre anus length 73.5 71.7 73.4 72.30.75 Distance between anus and anal fin 5.1 5.5 7.1 6.00.75 Dorsal fin base length 12.6 12.0 13.2 12.40.53 Dorsal fin le ngth 21.5 20.0 21.0 20.60.42 Pectoral fin base length 9.8 10.8 11.8 11.10.48 Pectoral fin length 28.3 26.9 28.0 27.40.47 Pelvic fin base length 6.4 6.5 6.8 6.60.20 Pelvic fin length 20.6 19.8 20.4 20.10.30 Anal fin length 15.9 13.3 14.8 14. 30.69 Caudal peduncle length 12.4 12.6 14.4 13.70.77 Caudal peduncle depth 10.1 10.2 10.9 10.50.30 % of h ead l ength Head width 69.6 59.0 68.1 63.23.92 Head depth 48.4 45.0 51.4 47.82.66 Snout length 43.3 42.8 45.9 44.51.31 Snout to nostr il distance 30.1 32.1 33.5 32.70.63

PAGE 113

113 T able 5 12 Continued Character Holotype Range (n=4) Mean%SD (n=4) Nostril to operculum distance 66.7 65.9 68.1 66.91.00 Internostril width 19.1 19.0 22.1 20.41.34 Length of orbit 21.3 19.9 24.3 22.02.13 Interorbital width 25.7 24.4 30.1 26.82.38 Length of maxillary barbel 12.9 11.3 13.4 12.40.94 Length of lateral rostral barbel 11.9 11.1 13.0 11.80.81 Width of gape 33.3 28.5 30.7 29.60.90 Inter lower lip width 22.6 20.0 21.4 20.60.67 % of i nterorbital w idth Length of orbit 82.9 78.7 88.1 82.34.06 M eristics Dorsal fin ray count iii, 7 (5) Pectoral fin ray count v, 8, ii & v, 8, i (*); v, 9, i & iv, 10, i (2); v, 9, i (1); v, 7, ii & v, 8, i (1) Pelvic fin ray count ii, 7 (5) Ana l fin ray count ii, 4, i (*); i, 5 (3); ii, 5 (1) Caudal fin ray count 17 (4*);16 (1) Lateral line pore count 41 44 + 2 1 on caudal fin Lateral line pore at pelvic fin origin 15 18 Lateral line pore at dorsal fin origin 18 20 Lateral line pore at an al fin origin 32 35 Circumpeduncle scale count 20 22 Number of s cale rows above / below lateral line 6 8 / 6 7 Number of scale rows below lateral line to pelvic fin origin 6 7 Predorsal scale count 22 25

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114 Figure 5 1. D orsal, lateral, and v entral views of Homalopteroides wassinkii UMMZ 155660, 46.2 mm SL; Indonesia, Java Tjisokan, inland fishery S cale bar represents 30 mm.

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115 Figure 5 2. Distribution of Homalopteroides wassinkii Dot represent s localit y for specimens examined; asterisk r epresents type locality.

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116 Figure 5 3 Homalopteroides modestus, Thailand, K anchanaburi, Mae Khlong basin: A) dorsal, lateral, and ventral views of an adult, UF 181080, 37.2 mm SL, scale bar represents 30 mm; B) lateral view of a juvenile, UF 176377, 2 4.8 mm SL, scale bar represents 10 mm.

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117 Figure 5 4. Mouth of Homalopteroides modestus UF 181080, 37.2 mm SL. Scale bar represents 2 mm.

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118 F igure 5 5. Fifth ceratobranchial of Homalopteroides modestus : A) adapted from Vinciguerra (1890), p. 331; B) UF 1 76454, 31.48 mm SL, left side, ventral view; C) UF 176454, 31.48 mm SL, right side, dorsal view. B and C scale bar represents 0.5 mm.

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119 Figure 5 6. Distribution of Homalopteroides modestus Black dots represent localities for specimens examined; asterisk represents type locality.

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120 F igure 5 7. Habitat of Homalopteroides modestus Thailand, Kanchanaburi, Mae Khlong basin, Mae Nam Kwae, Khayeng River, 30 Dec. 2011, 14.65 N, 98.566667 E.

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121 F igure 5 8. Lateral view of Homalopteroides rupicola : A ) paratype SU 28726 21.3 mm SL, Myanmar, Myitkyina District, Sankha, a large hillstream, midway between Kamaing and Mogaung Irrawaddy River basin S cale bar represents 1 0 mm. B ) adapted from Prashad and Mukerji (1929 ), p late 8, Figure 3

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122 F igure 5 9. Type loca lity of Homalopteroides rupicola

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123 F igure 5 10. Dorsal, Lateral, and ventral view s of Homalopteroides smithi : paratype USNM 109821 50 mm SL, Thailand, Nakon Sritamarat, Klong Pong Tadi Stream ; ? Pakphanang basin S cale bar represents 20 mm.

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124 F igure 5 11. Dorsal and lateral view of Homalopteroides smithi : UF 183330 43.6 mm SL, Thailand, Nakon Sritamarat Chang Klang Village, Tapee Basin. S cale bar represents 20 mm.

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125 F igure 5 12. Distribution of Homalopteroides smithi Dots represent localities fo r specimens examined; as terisk represents type locality

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126 F igure 5 13. Scatter plot of caudal peduncle depth for all species of Homalopteroides

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127 F igure 5 14. Mouth of Homalopteroides stephensoni USNM 230254, 50.0 mm SL. Scale bar represents 2 mm.

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128 F igure 5 15. Dorsal, lateral, and ventral views of Homalopteroides stephensoni, USNM 230254, 50.0 mm SL, Indonesia, West Kalimantan, Kapuas basin S cale bar represents 20 mm.

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129 F igure 5 16. Distribution of Homalopteroides stephensoni D ot represent s loc alit y for specimen examined ; as terisk represents type locality.

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130 F igure 5 17 Mouth of Homalopteroides weberi ROM 82131, 33.7 mm SL. Scale bar represents 2 mm.

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131 F igure 5 18 Dorsal, lateral, and ventral views of Homalopteroides weberi east Malaysia Sarawak, Baram basin ; A) ROM 70464, 40.8 mm SL; B) ROM 82131, 33.7 mm SL Scale bar s represent 2 0 mm.

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132 F igure 5 19 Dorsal, lateral, and ventral views of syntype Homalopteroides weberi BMNH 1895.7.2.81; 41.8 mm SL east Malaysia Sarawak, Baram basin ; sc ale bar represents 10 mm. Photos by Mark Sabaj Prez

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133 Figure 5 20 Distribution of Homalopteroides weberi Dots represent localit y for specimens examined; as terisk represents type locality.

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134 F igure 5 21 Dorsal, lateral, and ventral views of Homalopte roides tweediei ; A) h olotype SU 33012; Malaysia, Johore, Mawai District ( ? Kota Tinngi), ? Ambat basin; B) UMMZ 238923 26.0 mm SL, Malaysia, Johore Kahang River, Endau basin Scale bar represents 10 mm. Photos of A by CAS.

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135 F igure 5 22 Distribution of Homalopteroides tweediei

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136 F igure 5 23 Dorsal, lateral, and ventral views of Homalopteroides indochinensis holotype BMNH 1933.8.19.50; bar represents 10 mm. Photos by Mark Sabaj Prez.

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137 F igure 5 24 Type locality of Homalopteroides indochinensis

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138 F igure 5 25. Lateral views of Homalopteroides nebulosus A) Holotype ZRC 2020; 23.6 mm SL; Malaysia, Kelantan, Sok River, Kampong Sok Kelantan basin ; B) paratype BMNH 1967.11.15.15 ; 22.2 mm SL; same locality d ata as holotype. Scale bar represent 10 mm.

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139 F igure 5 26 Type locality of Homalopteroides nebulosus

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140 Figure 5 27 Homalopteroides yuwonoi ; holotype MZB 5938, 38.6 mm SL; Indonesia, Borneo, Kalimantan Barat, Kapuas basin ; A) lateral view, B) ventr al view.

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141 F igure 5 28 Type locality of Homalopteroides yuwonoi

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142 Figure 5 29. Type locality of Homalopteroides manipurensis

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143 Figure 5 30. Scatter plot of gape width for all species of Homalopteroides

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144 F igure 5 31. D orsal, lateral, and ventra l views of Homalopteroides n. sp holotype USNM 323875, 52.9 mm SL, Malaysia Sarawak, Rajang basin S cale bar represents 2 0 mm

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145 Figure 5 32. Mouth of Homalopteroides n. sp. holotype USNM 323875 52.9 mm SL. Scale bar represents 3 mm.

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146 Figure 5 33. Distribution of Homalopteroides n. sp Dots represent locality for specimens examined; asterisk represents type locality.

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147 Figure 5 34. H abitat of Homalopteroides n. sp e ast Malaysia, Sarawak, Balui River Rajang basin, 02 Aug. 1991 2.416667 N, 113 73 3333 E. Photo by Lynne Parenti.

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148 CHAPTER 6 DISCUSSION Many authors have noted the possible paraphyly of the genus Homaloptera (Fang 1930; Hora 1932; Kottelat 1998; Tan & Ng 2005; Tan 2009), and several genera have been created for species previously in H omaloptera ( Hemimyzon Regan 1911, Lepturichthys Regan 1911, Protomyzon Hora 1932, and Cryptotora Kottelat 1998) or elevated from use as a subgenus ( Neohomaloptera Herre 1944). The morphological characters described in Chapter 4, as well as the phylogenetic analysis of the Rag1 gene discussed in more detail below, demonstrate the paraphyly of Homaloptera The Rag1 tree ( Figure 6 1 ) shows 98% posterior probability (pp) support for the family Balitoridae, weak support of 74% pp for Gastromyzoninae, a nd 100% p p for the Balitorinae. The low pp value for the Gastromyzoninae is likely due to the inclusion of the enigmatic genus Barbucca Roberts 1989, whose systematic position is questionable et al. 2007) and which was recognized in the monotypic Barbuc cidae by Kottelat (2012). In Balitorinae, Homaloptera is paraphyletic, with a 100% pp that Homaloptera smithi Hora 1932 is sister to a clade including Balitora annamitica Kottelat 1998, Homaloptera zollingeri Bleeker 1853, H. leonardi Hora 1941, and H. pa rclitella Tan and Ng 2005. Balitora annamitica is sister to these three specie s of Homaloptera with 100% pp. lechtov et al. (2007) found similar results with H. smithi sister to a clade containing Balitora and Homaloptera Based on these results and its morphological diagnosability, described in Chapter 4 Homalopteroides in the phylogeny represented by H. smi thi is removed from the synonymy of Homaloptera T he gen us Balitoropsis is not as well defined. Balitoropsis leonardi in the phylogeny represented by H. leonardi

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149 is sister to Homaloptera parclitella (87% pp), and both of them are sister to Balitoropsis zollingeri in the phylogeny represented by H. leonardi with a weak support of 59% pp. The inclusion of more species of Homaloptera and Balitoropsis into the phylogenetic analysis will likely show strong support for the monophyly of both genera. Balitorops is is recognized as valid based on its morphological diagnosability described in Chapter 4. Homaloptera consists of six species and is known to occur in south Myanmar, Thailand, Penninsular Malaysia, Sumatra, Java, and Borneo. Homaloptera parclitella and H. ogilviei are the only species belonging to this genus known to occur sympatrically, and only in the Endau Drainage, Malaysia (Ng and Tan 1999). Homalopterula is only known to occur in Sumatra and consists of four species, three of which, Homalopterula h eterolepis H. ripleyi and H. vanderbilti, were described from Aceh Province, Sumatra. There have been no reports of species of Homalopterula as occurring sympatrically. Balitoropsis consisits of nine species and is known to occur in China, Laos, Cambodi a, Thailand, Peninsular Malaysia, Sumatra, Java, and Borneo. There have been no reports of any species of Balitoropsis as occurring sympatrically. Balitoropsis zollingeri likely occurs sympatrically with other species of Balitoropsis since it has the wides t distribution among species of the genus, being found throughout the range of Balitoropsis except for China. The type locality for B. zollingeri is in Java. There have been two subsequent species described from northern Thailand and the Malay Peninsula, B alitoropsis bartschi (Smith 1945) and Homaloptera nigra (Alfred 1969), respectively. Both of these species have been recognized as junior synonyms of B.

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150 zollingeri (Kottelat and Chu 1988 and Kottelat 2001). A more in depth study on the taxonomy of B. zolli ngeri is needed to test the validy of these synonyms. problematic since it does not share the unique combination of characters diagnosing any of the four genera ( Homaloptera Homalo pteroides Homalopterula and Balitoropsis ) recognized in this study. A member of this group, Homaloptera montana can be distinguished from all species of Homaloptera by having the origin of the dorsal fin posterior to th e insertion of the pelvic fin. Bas ed on this unique character, a new genus will be created for Homaloptera montana (Herre 1945) in another publication. origin of the dorsal fin posterior to th e insertion of the pelvic fin) is shared among them, they will be tentatively placed in the new genus because they are morphologically more similar to H. montana then to any other species of Homaloptera Homalopteroides currentl y consist of 12 species and is known to occur in northeastern India, Myanmar, Thailand, Vietnam, Peninnsular Malaysia, Java, and Borneo. There are several populations in northern and eastern Thailand, Laos, Cambodia, Sumatra, and Borneo that have been iden tified as Homalpteroides smithi H. tweediei and/or H. nebulosus These identifications are inaccurate and refer to as many as five new species of Homalopteroides The following species of Homalopteroides are known to occur sympatrically: H. nebulosus and H. tweediei in the Endau Drainage (Ng and Tan 1999), and H. modestus and H. cf. smithi in the Mae Khlong Drainage (pers observation). Tan (2009) has recorded H. stephensoni from the Sarawak and Sabah of

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151 H. weberi since the two species are morphologically very similar. More specimens for both of these species need to be examined before it can be concluded that they are found in the same drainage and are sympatric. Some species of Homalopteroides were originally described only from small specimens a practice that confuses later comparisons among species when adults are examined due to ontogenetic change. For example, the types for H. rupicola were 24 27 mm SL (Prashad and Mukerji 1929) and for H. tweediei were 23 26 mm S L (Herre 1940) Based on the material examined in this study both H. rupicola and H. tweediei have an o rbital length greater than the interorbital width, a character used for distinguishing species of Homalopteroides To test whether relative orbital leng th is a reliable diagnostic character or one that changes ontogentically, adults and juveniles of H. modestus and H. smithi were compared Adults of both of these species had an orbital length less than th e interorbital width (Tables 5 4 and 5 6), but juve niles of H. modestus had an orbital length greater than or equal to t he interorbital width (Table 5 4). Since the states of this character (orbital length greater or lesser than interorbital width) are dependent on the life stage being examined, its use as a diagnosible character should be limited to adults, at least when distinguishing species of Homalopteroides This demonstrates the effect that ontogentic change can have on character assemblage, and the importance of sampling both adults and juveniles wh en describing a species. Based on the material examined, there are two groups within Homalopteroides the H. wass inkii group and the H. stephensoni group. The H. wassinkii group is diagnosed from the H. stephensoni group by the following combination of cha racters: a

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152 total pelvic fin ray count of 8 9 vs.10, and a deeper caudal peduncle, 8.0 10.9% SL, vs. 6.1 8.0% SL. Based on these characters, the following nine species of Homalopteroides belong to the H. wassinkii group: H. wassinkii H. modestus H. rupico la H. smithi H. tweediei H. indochinensis H. nebulosus H. manipurensis and H. sp. n ; and 3 species, only known from Borneo, belong to the H. stephensoni group: H. stephensoni H. weberi and, H. yuwonoi The H. wassinkii group has a wider distributio n than the H. stephensoni group, being also found in Borneo but not restricted to it. A more comprehensive phylogentic analysis is needed to test the timing of divergence for Homaloptera Homalopteroides Homalopterula Balitoropsis his analysis will help us better understand the geological history of Indochina and Sundaland, and the evolutionary relationships of these genera to other balitroids, a family that consists of a large amount of diversity that remains to be described.

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153 Fi gure 6 1. Phylogenetic relationships of the family Balitoridae from a Bayesian analysis of the Rag1 gene. An asterisk indicates a posterior probability of 100%.

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154 LIST OF REFERENCES Alfred, E.R. ( 1969) The Malayan cyprinoid fishes of the family Homalopterid ae. Zoologische Mededelingen (Leiden) 43, 213 237. Arunkumar, L. ( 1998) Homaloptera manipurensis a new homalopterid fish from Manipur, India. Uttar Pradesh Journal of Zoology 18, 175 179. Bhlke, J. E. (1953) A catalogue of the type specimens of r ecen t fishes in the Natural History Museum of Stanford University. Stanford Ichthyological Bulletin 5, 1 168. Boulenger, G.A. (1894) Descriptions of new freshwater fishes from Borneo. Annals and Magazine of Natural History 13, 245 251. Chen, Y. Y. (1998) The fishes of the Hengduan Mountains region. The series of the scientific expedition to the Hengduan Mountains of the Qinghai Xizang Plateau 1 364. Chen, W. J., Miya, M., Saitoh, K. and Mayden, R.L. (2008) Phylogenetic utility of two existing and four n ovel nuclear gene loci in reconstructing Tree of Life of ray finned fishes: The order Cypriniformes (Ostariophysi) as a case study. Gene 423, 125 134. Cuvier, G. & Valenciennes A. (1 846 ) Tome dix huitime, s uite du livre dix huitime, Cyprinodes, Livre dix neuvime, Des soces ou Luciodes Histoire naturelle des poisons 18, 1 505 Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32, 1792 1797. Eschmeyer, W.N. & Fricke, R. (Eds) Ca talog of Fishes electronic version. Available from http://research.calacademy.org/research/ichthyology/catalog/fishcatmain.asp (accessed 30 January 2012). Fang, P. W. ( 1930) New and inadequately known homalopterin loaches of China, with a rearrangement an d revision of the generic characters of Gastromyzon, Sinogastromyzon and their related genera. Contributions from the Biological Laboratory of the Science Society of China 6, 25 43. Felsenstein, J. (1981) Evolutionary trees from DNA sequences: a maximum likelihood approach. Journal of Molecular Evolution 17, 368 376. Fowler, H.W. ( 1905) Some fishes from Borneo. Proceedings of the Academy of Natural Sciences of Philadelphia 57, 455 523. Fowler, H. W. (1934) Zoological results of the third De Schauense e Siamese Expedition, Part I. -Fishes. Proceedings of the Academy of Natural Sciences of Philadelphia 86, 67 163.

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155 Fowler, H.W. ( 1940) Zoological results of the George Vanderbilt Sumatran Expedition, 1936 1939. Part II. -The fishes. Proceedings of the Aca demy of Natural Sciences of Philadelphia 91, 369 398. Gouy, M., Guindon, S. and Gascuel, O. (2010) SeaView version 4: a multiplatform graphical user interface for sequence alignment and phylogenetic tree building. Molecular Biology and Evolution 27 22 1 224. Herre, A.W.C.T. ( 1940) New species of fishes from the Malay Peninsula and Borneo. Bulletin of the Raffles Museum 16, 5 26. Hora, S.L. ( 1932) Classification, bionomics and evolution of homalopterid fishes. Memoirs of the Indian Museum 12, 263 330 Hubbs, C.L. & Lagler, K.F. (2004) Fishes of the Great Lakes region, with a new preface. University of Michigan Press, Ann Arbor, Michigan, 276 pp. Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogeny. Bioinformatics 17, 754 755. ICZN (1999) International code of zoological nomenclature [the Code] Fourth edition. The International Trust for Zoological Nomenclature, Natural History Museum, London, 306 pp. Inger, R.F. & Chin, P.K. ( 1962 ) The fresh water fishes of North Borne o. Fieldiana Zoology 45, 1 268. Jordan, D.S. (1920) The genera of fishes, part IV, from 1881 to 1920, thirty nine years, with the accepted type of each. A contribution to the stability of scientific nomenclature Leland Stanford Jr. University Publicatio ns, 43, 411 576. Kottelat, M. ( 1984) Revision of the Indonesian and Malaysian loaches of the subfamily Noemacheilinae. Japanese Journal of Ichthyology 31, 225 260. Kottelat, M. ( 1988) Indian and Indochinese species of Balitora (Osteichthyes: Cyprinifo rmes) with descriptions of two new species and comments on the family group names Balitoridae and Homalopteridae. Revue Suisse de Zoologie 95, 487 504. Kottelat, M. (1990) Indochinese nemacheilines. A revision of nemacheiline loaches (Pisces: Cypriniform es) of Thailand, Burma, Laos, Cambodia and southern Viet Nam Verlag Dr. Friedrich Pfeil, Mnchen, 262 pp. Kottelat, M. ( 1998) Homaloptera yuwonoi a new species of hillstream loach from Borneo, with a new generic name for H. thamicola (Teleostei: Balito ridae). Ichthyological Exploration of Freshwaters 9, 267 272.

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156 Kottelat, M. (2012) Acanthocobitis pictilis a new species of l oach from Myanmar and Thailand (Teleostei: Nemacheilidae). Zootaxa 3327, 45 52. Kottelat, M. ( 2012 ) Conspectus Cobiti d um: an inventory of the loaches of the world (Teleostei: Cypriniformes: Cobitoidei). The Raffles Bulletin of Zoology Supplement no 36, 1 199. Madhusoodana Kurup, B. & Radhakrishnan, K.V. (2011) Fishes of the genus Homaloptera van Hasselt, 1823 in Kerala, with description of a new species Homaloptera silasi Journal of the Bombay Natural History Society 107, 224 226. Martin Smith, K.M. & Tan, H.H. (1998) Diversity of freshwater fishes from eastern Sabah: annotated checklist f or Danum Valley and a consideration of inter and intra catchment variability. Raffles Bulletin of Zoology Supplement no. 46 573 604 Mayden, R.L., Tang, K.L., Wood, R.M., Chen, W. J., Agnew, M.K., Conway, K.W., Yang, L., Li, J., Wang, X., Saitoh, K., M iya M., He, S., Liu, H., Chen, Y., and Nishida, clade of freshwater fishes: implications of varied taxon and character sampling. Journal of Systematic Evolution 46, 424 438. Menon, A.G.K. ( 1987) The fauna of India and the adjacent countries. Pisces. Vol. IV. Teleostei Cobitoidea. Part 1. Homalopteridae. Zoological Survey of India 1 259. Menon, A.G.K. & Yazdani, G.M. (1968) Catalogue of type specimens in the Zoo logical Survey of India, Fishes. Records of the Zoological Survey of India 61, 91 190. Monkolprasit, S. P. (1969) Type specimens of fishes at the Kasetsart University Museum of Fisheries. Notes from Faculty of Fisheries, Kasetsart University, Bangkok 5, 1 8. Nijssen, H., van Tuijl, L., and Isbrcker, I.J.H. ( 1982) A catalogue of the type specimens of Recent fishes in the Institute of Taxonomic Zoology (Zologisch Museum), University of Amsterdam, The Netherlands. Verslagen en Technische Gegevens, Instit uut voor Taxonomische Zoogie Universiteit van Amsterdam. 33,1 173. Ng, H.H. & Tan, H. H. (1999) The fishes of the Endau drainage, Peninsular Malaysia with descriptions of two new species of catfishes (Teleostei: Akysidae, Bagridae). Zoological Studies 38, 350 366. Nguyen, V. H. & Nguyen, H. D. ( 2002 ) Mot giong moi va mot loai moi thuoc ho Balitoridae (Cypriniformes) tim thay o huyn Dien Bien, tinh Lai Chau. (A new genus and a new species of freshwater fish Balitoridae, Cypriniformes from the Dienbien Dis trict, Laichau Provoince, Vietnam). Tap Chi Sinh Hoc (J. Biol.) 24, 9 14.

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157 Parenti, L.R. & Lim, K.K.P (2005) Fishes of the R ajang basin, Sarawak, Malaysia. Raffles Bulletin of Zoology Supplement no. 13, 175 208. Pethiyagoda, R. & Kottelat, M (1994) Three new species of fishes of the genera Osteochilichthys (Cyprinidae), Travancoria (Balitoridae) and Horabagrus (Bagridae) from the Chalakudy River, Kerala, India. Journal of South Asian Natural History 1, 97 116. Plongsesthee, R., Page L.M., and Be amish, W. (2011) Schistura aurantiaca a new species from the Mae Khlong basin, Thailand (Teleostei: Nemacheilidae). Ichthyological Exploration of Freshwaters 22, 169 178. Popta, C.M.L. (1906) Rsultats ichthyologiques des voyages scientifiques de Monsi eur le Professeur Dr. A. W. Nieuwenhuis dans le centre de Borno (1898 et 1900). Notes from the Leyden Museum 27, 1 304. Prashad, B. & Mukerji, D.D. (1929) The fish of the Indawgyi Lake and the streams of the Myitkyina District (Upper Burma). Records of the Indian Museum 31, 161 223. Rainboth, W.J., Vidthayanon, C., and Yen, M. D. (2012) Fishes of the Greater Mekong e cosystem w ith s pecies l ist and p hotographic a tlas. Miscellaneous Publications, Museum of Zoology, University of Michigan 201 1 172 R ambaut, A. & Drummond, A. (2010) FigTree v1.3.1 Institute of Evolutionary Biology, University of Edinburgh, Edinburgh, United Kingdom. Rambaut, A. & Drummond, A.J. (2007) Tracer v1.4 Available from http://beast.bio.ed.ac.uk/Tracer. Selim, K. & Vishwan ath, W (1998) A new record of Homaloptera modesta (Vinciguerra): Cyprinidae from Manipur. Journal of the Bombay Natural History Society 95, 352 354. Silas, E.G. ( 1953) Classification, zoogeography and evolution of the fishes of the cyprinoid families H omalopteridae and Gastromyzonidae. Records of the Indian Museum 50, 173 263. and Tan, H.H. (2007) Families of Cobitoidea (Teleostei: Cypriniformes) as revealed from nuclear genetic data and the position of the mysterious genera Barbucca Psilorhynchus Serpenticobitis and Vaillantella Molecular Phylogene tics and Evolution 44, 1358 1365. Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods) Sinauer Associates, Sunderland, Massachusetts.

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158 Tan, H.H. ( 2009) A new species of hill stream loach (Teleostei: Balitoridae) from central Kalimantan, with redescriptions of Homaloptera tateregani Popta and Homaloptera stephensoni Hora. Zootaxa 2171, 48 64. Tan, H.H. & Ng P.K.L. ( 2005) Homaloptera parclitella a new species of torrent loach from the Malay Peninsula, with redescrip tion of H. orthogoniata (Teleostei: Balitoridae). Ichthyological Exploration of Freshwaters 16, 1 12. Tang, W. Q. & Chen, Y. Y. (2000) Study on taxonomy of Homalopteridae. Journal of Shanghai 9, 1 9. Taylor, W.R. & van Dyke, G.C. (1985) Revised proced ures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium 9, 107 119 Tortonese, E. ( 1961) Catalogo del tipi de pesci del Museo Civico di Storia Naturale di Genova. (Parte I). Annali del Museo Civico di Storia Naturale `Giacomo Doria' 72, 179 191. Vinciguerra, D. (1890) Viaggio di Leonardo Fea in Birmania e regioni vicine. XXIV. Pesci. Annali del Museo Civico di Storia Naturale di Genova 9, 129 362. Weber, M. & de Beaufort L.F. (1916) The fishes of the Indo Australian Archipelago. III. Ostariophysi: II Cyprinoidea, Apodes, Synbranchi. The Fishes of the Indo Australian Archipelago 3, 1 455.

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159 BIOGRAPHICAL SKETCH Zachary S. Randall was born in Miami, Florida. One of four children he was int erested in the natural world at a young age and has pursued hobbies in music an d photography. He received his a ssociate s degree from Miami Dade College in 2006 where he studied fine art photography He then received h is Bachelor of Arts in a nthropo logy from the University of Florida, where he graduated Cum Laude in 2008 He then worked as a full time research assistant at the Florida Museum of Natural History in the D epartments of Vertebrate Paleontology, Ornithology, and Ichthyology. He enrolled as a graduate student in the D epartment of Biology at the University of Florida in 2011, with a research assistantship from the All Cypriniformes Species Inventory Project funded by the U.S. National Science Foundation. He will continue to work at the Florid a Museum of Natural History as a technician and researcher for the next year and is currently investigating programs for a PhD