University of Florida Thesis Or Dissertation Formatting

TAXONOMIC REVISION OF THE GENUS Acanthocobitis (TELEOSTEI: NEMACHEILIDAE) WITH RECOGNITION OF THE GENUS Paracanthocobitis (TELEOSTEI: NEMACHEILIDAE) AND THE DESCRIPTION OF FOUR NEW SPECIES

RANDAL ANTHONY SINGER

A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE

UNIVERSITY OF FLORIDA

2013

To my family for all their love and support

ACKNOWLEDGMENTS

Funding for this study provided by the All Cypriniformes Species Inventory project funded by the U.S. National Science Foundation (DEB 1022720). I thank Dr.

Lawrence Page for guidance and direction on this project, and for editorial review during the writing process. I would also like to thank Dr. William Smith-Vaniz and Dr. Akito

Kawahara for guidance and review on my committee. For specimen loans I would like to thank the following people: Dave Catania (CAS), Rob Robins (UF), Jeff Williams

(USNM), Doug Nelson (UMMZ), Florian Wicker (SMF), Kelvin Lim (ZRC), Mary Burridge

(ROM), Karsten Hartel (MCZ), Andy Bentley (KU), Peter Bartsch (ZMB), Uli Schliewen and Dirk Neumann (ZSM), Patrice Pruvost (MHNH), and Chaiwut Grudpan (UMNF). I would also like to thank Steven Grant for use of his photos, information regarding fishes, and his groundwork on the group. I thank Jesse Grosso for providing help with composing maps and georeferencing localities. I thank Dr. Daniel Lumbantobing for assisting in the osteology analysis and guidance. I thank Zachary Randall for assistance with photography and formatting. Rob Robins for help with editing and guidance in research process. I thank Bill Eschmeyer for his extensive work on the Catalog of

Fishes, which provided a great deal of information for this study. Lastly, I would like to thank my wife Corie Singer and my daughter Adelaide Singer for support and patience during my research and studies.

TABLE OF CONTENTS

page

ACKNOWLEDGMENTS ...... 4

LIST OF TABLES ...... 8

LIST OF FIGURES ...... 9

ABSTRACT ...... 11

INTRODUCTION ...... 12

Taxonomic History ...... 12 Natural History ...... 13

MATERIALS AND METHODS ...... 15

Institutional Abbreviations for Material Examined ...... 16 Material Examined ...... 17 Acanthocobitis pavonacea ...... 17 Paracanthocobitis abutwebi, n.sp...... 17 Paracanthocobitis adelaideae, n.sp...... 18 Paracanthocobitis botia ...... 18 Paracanthocobitis linypha, n.sp...... 18 Paracanthocobitis mackenziei ...... 19 Paracanthocobitis maekhlongensis, n.sp...... 22 Paracanthocobitis mandalayensis ...... 22 Paracanthocobitis mooreh ...... 23 Paracanthocobitis pictilis ...... 23 Paracanthocobitis rubidipinnis ...... 23 Paracanthocobitis urophthalma ...... 24 Paracanthocobitis zonalternans ...... 24

SYSTEMATIC ACCOUNTS ...... 28

Genus Acanthocobitis ...... 28 Diagnosis ...... 28 Description ...... 28 Distribution ...... 29 Remarks ...... 29 Acanthocobitis pavonacea (McClelland, 1839) ...... 30 Diagnosis ...... 30 Description ...... 30 Distribution ...... 31 Genus Paracanthocobitis ...... 31 Diagnosis ...... 31

Description ...... 32 Distribution ...... 32 Paracanthocobitis abutwebi, n.sp...... 32 Diagnosis ...... 33 Description ...... 33 Distribution ...... 34 Etymology ...... 34 Paracanthocobitis adelaideae, n.sp...... 34 Diagnosis ...... 34 Description ...... 35 Comparison ...... 35 Distribution ...... 35 Etymology ...... 36 Paracanthocobitis botia (Hamilton, 1822) ...... 36 Diagnosis ...... 37 Description ...... 37 Comparison ...... 38 Distribution ...... 38 Remarks ...... 38 Paracanthocobitis linypha, n.sp...... 39 Diagnosis ...... 39 Description ...... 39 Comparison ...... 40 Distribution ...... 40 Etymology ...... 40 Paracanthocobitis mackenziei (Chaudhuri, 1910) ...... 41 Diagnosis ...... 41 Description ...... 41 Comparison ...... 42 Distribution ...... 42 Remarks ...... 42 Paracanthocobitis maekhlongensis, n.sp...... 44 Diagnosis ...... 44 Description ...... 44 Comparison ...... 45 Distribution ...... 45 Remarks ...... 45 Etymology ...... 46 Paracanthocobitis mandalayensis (Rendahl 1948) ...... 46 Diagnosis ...... 46 Description ...... 46 Comparison ...... 47 Distribution ...... 47 Paracanthocobitis mooreh (Sykes, 1839) ...... 47 Diagnosis ...... 48 Description ...... 48

Comparison ...... 49 Distribution ...... 49 Remarks ...... 49 Paracanthocobitis pictilis (Kottelat 2012) ...... 50 Diagnosis ...... 50 Description ...... 50 Comparison ...... 50 Distribution ...... 51 Paracanthocobitis rubidipinnis (Blyth 1860) ...... 51 Diagnosis ...... 52 Description ...... 52 Comparison ...... 52 Distribution ...... 52 Paracanthocobitis urophthalma (Günther, 1868) ...... 53 Diagnosis ...... 53 Description ...... 53 Comparison ...... 54 Distribution ...... 54 Paracanthocobitis zonalternans (Blyth 1860) ...... 54 Diagnosis ...... 55 Description ...... 55 Comparison ...... 55 Distribution ...... 56 Remarks ...... 56

DISCUSSION ...... 105

KEY TO SPECIES OF PARACANTHOCOBITIS ...... 107

LIST OF REFERENCES ...... 110

BIOGRAPHICAL SKETCH ...... 113

LIST OF TABLES

Table page

3-1 Morphometric data and meristic counts for Paracanthocobitis abutwebi (N=35)...... 93

3-2 Morphometric data and meristic counts for Paracanthocobitis adelaideae (N=8)...... 94

3-3 Morphometric data and meristic counts for Paracanthocobitis botia (N=10)...... 95

3-4 Morphometric data and meristic counts for Paracanthocobitis linypha (N=10)...... 96

3-5 Morphometric data and meristic counts for Paracanthocobitis mackenziei (N=46)...... 97

3-6 Morphometric data and meristic counts for Paracanthocobitis maekhlongensis (N=12)...... 98

3-7 Morphometric data and meristic counts for Paracanthocobitis mandalayensis (N=23)...... 99

3-8 Morphometric data and meristic counts for Paracanthocobitis mooreh (N=30)...... 100

3-9 Morphometric data and meristic counts for Paracanthocobitis pictilis (N=16). . 101

3-10 Morphometric data and meristic counts for Paracanthocobitis rubidipinnis (N=11)...... 102

3-11 Morphometric data and meristic counts for Paracanthocobitis urophthalma (N=5)...... 103

3-12 Morphometric data and meristic counts for Paracanthocobitis zonalternans (N=26)...... 104

LIST OF FIGURES

Figure page

3-1 Acanthocobitis longipinnis. ZMB, 4795, Holotype, ventral view ...... 57

3-2 Acanthocobitis longipinnis. Lateral view of head, showing suborbital ...... 58

3-3 Head shapes in Acanthocobitis and Paracanthocobitis. A) Acanthocobitis ...... 59

3-4 Lateral views of Acanthocobitis type specimens. A) Acanthocobitis ...... 60

3-5 A) Cobitis pavonacea, SMF 9091, paralectotype, 121 mm SL female, ...... 61

3-6 Distribution of Acanthocobitis pavonacea...... 62

3-7 Paracanthocobitis maekhlongensis, UF 182864,58.9 mm SL. Ventral ...... 63

3-8 Paracanthocobitis mackenziei, KU 40459, 51.5 mm SL male, lateral ...... 64

3-9 Suborbital process shapes in Paracanthocobitis. A) Suborbital flap on P...... 65

3-10 Bones supporting the suborbital flap. LE (yellow) = lateral ethmoid; IO ...... 66

3-11 Paracanthocobitis abutwebi. UMMZ 208646, holotype, 46.1 mm SL ...... 67

3-12 Distribution of Paracanthocobitis abutwebi...... 68

3-13 Paracanthocobitis adelaideae. A) adult color pattern, USNM 378387, ...... 69

3-14 Distribution of Paracanthocobitis adelaideae...... 70

3-15 Paracanthocobitis botia. A) BMNH 1889.2.1.1562-1571, neotype, 45.3 ...... 71

3-16 Distribution of Paracanthocobitis botia...... 72

3-17 Paracanthocobitis linypha. A) USNM 378387, holotype, 34.9 mm SL ...... 73

3-18 Distribution of Paracanthocobitis linypha...... 74

3-19 A) Paracanthocobitis mackenziei, KU 29159, 67.1 mm SL male. B) ...... 75

3-20 Distribution of Paracanthocobitis mackenziei...... 76

3-21 Paracanthocobitis maekhlongensis. A) UF 182864, holotype ...... 77

3-22 Comparison of key morphological differences in Paracanthocobitis ...... 78

3-23 Distribution of Paracanthocobitis maekhlongensis...... 79

3-24 Paracanthocobitis mandalayensis. A) Specimen from Irrawaddy River ...... 80

3-25 Paracanthocobitis mandalayensis. Juvenile color pattern, UF 1811110, ...... 81

3-26 Distribution of Paracanthocobitis mandalayensis ...... 82

3-27 Paracanthocobitis mooreh. Lateral view, MCZ 52380, 44.4 mm SL ...... 83

3-28 Distribution of Paracanthocobitis mooreh ...... 84

3-29 Paracanthocobitis pictilis. A) UF 172927, 71.4 mm SL male. B) ...... 85

3-30 Distribution of Paracanthocobitis pictilis...... 86

3-31 Paracanthocobitis rubidipinnis. Lateral view, ZRC 43570, 55.5 mm SL ...... 87

3-32 Distribution of Paracanthocobitis rubidipinnis...... 88

3-33 Paracanthocobitis urophthalma. Lateral view, BMNH 1864.7.11, ...... 89

3-34 Distribution of Paracanthocobitis urophthalma...... 90

3-35 Paracanthocobitis zonalternans. A) ZSM 27468, neotype, 33.1 mm SL ...... 91

3-36 Distribution of Paracanthocobitis zonalternans...... 92

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 GENUS Acanthocobitis (TELEOSTEI: NEMACHEILIDAE) WITH RECOGNITION OF THE GENUS Paracanthocobitis (TELEOSTEI: NEMACHEILIDAE) AND THE DESCRIPTION OF FOUR NEW SPECIES

Randal Anthony Singer

August 2013

Chair: Lawrence M. Page Major: Zoology

Morphological variation was studied in populations referred to the loach genus

Acanthocobitis (Nemacheilidae). Two-hundred thirty-two individuals were examined across the range of the genus, including specimens of all recognized species.

Paracanthocobitis was recognized as a genus and distinguished from Acanthocobitis in

having an emarginate or truncate (vs. pointed) caudal fin; patch (vs. no patch) of

adipose tissue on anterior end of lateral line; 9-15 (vs. 17-19) branched dorsal-fin rays,

and a rounded (vs. more triangular-shaped) head.

Acanthocobitis is monotypic, containing only A. pavonacea. Twelve species of

Paracanthcobitis are recognized, including seven species currently in Acanthocobitis: P.

botia, P. mandalayensis, P. mooreh, P. pictilis, P. rubidipinnis, P. urophthalma, and P.

zonalternans; P. mackenziei removed from synonymy; and four new species: P.

abutwebi, P. adelaideae, P. linypha, and P. maekhlongensis. All species are diagnosed

and distributions mapped. A key is provided for identification.

CHAPTER 1 INTRODUCTION

Taxonomic History

In 1861, Peters described the genus Acanthocobitis and designated

Acanthocobitis longipinnis, a new species from the Ganges River basin in India, as the

type species. Menon (1987) recognized Acanthocobitis longipinnis as a junior synonym of Cobitis botia Hamilton 1822, a species allocated to Nemacheilus by Shrestha (1978), therein placing Acanthocobitis longipinnis in the synonymy of Nemacheilus botia.

Bănărescu & Nalbant (1968) treated Acanthocobitis as a subgenus of Noemacheilus

Kuhl & van Hasselt 1823 (a synonym of Nemacheilus Bleeker 1863), following which

Menon (1987) placed Cobitis mooreh Sykes 1839, C. rubidipinnis Blyth 1860, C.

urophthalmus Günther 1868, and C. zonalternans Blyth 1860 in Nemacheilus.

Kottelat (1989,1990) recognized Acanthocobitis as a genus and placed N. botia,

N. rubidipinnis and N. zonalternans in Acanthocobitis. Pethiyagoda (1991) subsequently

recognized Nemacheilus urophthalmus as a species of Acanthocobitis.

Menon (1999) retained the name Noemacheilus for all species of Acanthocobitis,

noting that the latter was better treated as a subgenus of Nemacheilus, rather than as a

genus. However, this classification has not been widely accepted.

Grant (2007) proposed the name Paracanthocobitis as a subgenus of

Acanthocobitis after observing the holotype of A. longipinnis and finding it to be

morphologically quite distinct from A. botia. Grant also distinguished A. longipinnis from

species he placed in Paracanthocobitis in having a pointed caudal fin, slender body,

long dorsal fin, and greenish coloration. These distinctions were based on observations

of the type specimen of A. longipinnis and live individuals in aquaria. He named Cobitis

zonalternans as the type species of the subgenus Paracanthocobitis. In the same paper, Grant examined the type specimens for Cobitis pavonacea McClelland 1839 and determined them to be identifiable as A. longipinnis, making A. longipinnis a junior

synonym of A. pavonacea.

Kottelat (2012) described Acanthocobitis pictilis as a new species, and elevated

Nemacheilus rubidipinnis mandalayensis from subspecies to species and placed it and

Nemacheilus mooreh in Acanthocobitis. In his remarks, he referenced Grant (2007), but

did not recognize the validity of the subgenus Paracanthocobitis stating that a proper

taxonomic study had not been done. A subsequent publication by Kottelat (2012), that

included treatments for all loaches of the world, did not recognize the subgenus

Paracanthocobitis. However, it did recognize A. longipinnis as incertae sedis and A.

pavonacea as a possible species.

The objectives of this study are to examine the morphological and geographical

variation in species referred to the Acanthocobitis. Paracanthocobitis is recognized as a

genus, and four new species of Paracanthocobitis are described.

Natural History

The recorded range of Paracanthocobitis extends from Pakistan in the Indus

River basin to western Thailand in the Mae Khlong River basin. Most species of

Paracanthocobitis can be found in slow to moderate-flowing water in piedmont areas

where mountains meet the lowlands. The preferred substrate is small- to medium-sized

rocks and gravel, with some species preferring areas with leaf litter. Color patterns vary

from species to species, but generally consist of a brown to yellow body with brown

bars, blotches, spots and stripes. All members of this genus have a spot or ocellus on

the caudal peduncle, which might act as a false eye to deter predators.

In Paracanthocobitis, secondary sexual dimorphism is present in the form of a suborbital flap or a suborbital groove in males. Additionally, females have deeper bodies, shorter pectoral fins, and suborbital grooves in some species Breeding males have tubercles on the head and pelvic fins.

Breeding in this genus has not been observed in nature. Breeding time can be inferred to be during the rainy season (typically, January-March), as wild-captured specimens of Acanthocobitis during this time have bright red fin coloration, and females are gravid. Some species of Paracanthocobitis are popular in the aquarium trade, and breeding has been observed in captivity. Paracanthocobitis botia, P. rubidipinnis and P. zonalternans have all been bred in captivity. They seem to prefer to breed over small gravel, and fry associate with aquatic vegetation.

There appears to be no published information on the diet of Acanthocobitis. The presence of highly papillated lips and a subterminal mouth suggests that they, similar to most loaches, feed by searching through the substrate for food. Examinations of stomach contents have revealed larval chironomids and ephemeropterans.

CHAPTER 2 MATERIALS AND METHODS

Measurements and counts follow Kottelat (1990) with the exception that dorsal- fin and anal-fin counts include the last branched ray as a separate ray included in the count, rather than denoting it as “½”. Counts and measurements were taken on up to 10 specimens from each sample (specimens from one locality and date). Juveniles

(specimens less than 25 mm SL) were excluded from examination because of potential ontogenetic changes in body shape. Measurements were made to the nearest 0.1 mm

using digital calipers and a Leica M80 dissection microscope. All measurements are

given in millimeters (mm). Two-hundred thirty-two individuals were examined, including

specimens of all 12 species of Paracanthocobitis and one species of Acanthocobitis.

Individuals of P. pictilis and P. maekhlongensis were cleared and stained

following Armbruster (in litt.), a modification of Potthoff (1984) and Taylor and van Dyke

(1985). Photographs were taken of preserved specimens using a Visionary Digital

(Palmyra, Virginia) with Canon 40D and 5D cameras, and edited using Adobe

Photoshop CS5.1. Geographic coordinates were determined using maps and Google

Earth. Maps were constructed using Arc GIS 3.3 and Adobe Photoshop CS3.

Fieldwork was conducted in Thailand using backpack electroshockers, seines

and dipnets. Specimens were euthanized in MS-2222, preserved in 10% formalin, and

transferred to 70% ethanol for storage. Live color photos were taken when possible

using a Canon Powershot digital camera.

Species were recognized using the Phylogenetic Species Concept. A species is

the smallest diagnosable cluster of individual organisms within which there is a parental

ancestry and descent (Cracraft 1987).

Institutional Abbreviations for Material Examined

Institutional abbreviations follow Eschmeyer and Fricke (2012) and are defined below.

• ANSP: Academy of Natural Sciences, Philadelphia, U.S.A.

• AUM: Auburn University Natural History Museum, Auburn, Alabama, U.S.A.

• BMNH – Natural History Museum, London, United Kingdom.

• CAS: California Academy of Sciences, San Francisco, California, U.S.A.

• KU: University of Kansas Natural History Museum and Biodiversity Research

Center, Division of Ichthyology, Lawrence, Kansas, U.S.A.

• MCZ: Museum of Comparative Zoology, Harvard University, Ichthyology

Department, Cambridge, Massachusetts, U.S.A.

• NIFI: National Inland Fisheries Institute, Fish Taxonomy Division, Bangkok,

Thailand.

• NRM: Naturhistoriska Riksmuseet, Department of Vertebrate Zoology,

Ichthyology Section, Stockholm, Sweden.

• ROM: Royal Ontario Museum, Department of Natural History, Toronto.

• SMF: Senckenberg Forschungsinstitut und Naturmuseum, Abteilung Marine

Zoologie, Sektion Ichthyologie, Frankfurt am Main, Hessen, Germany.

• UF: University of Florida, Florida Museum of Natural History, Gainesville, U.S.A.

• UMMZ: University of Michigan Museum of Zoology, Ann Arbor, U.S.A.

• UMNF: Ubon Ratchathani University Ichthyology Collection, Ubon Ratchathani,

Thailand.

• USNM: Smithsonian Institution National Museum of Natural History, Department

of Vertebrate Zioology, Division of Fishes, Washington D.C., U.S.A.

• ZMB: Humboldt-Universität, Museum für Naturkunde, Zoologisches Museum,

Vertebraten (Wirbekltiere), Ichthyologie, Berlin, Germany.

Material Examined

Acanthocobitis pavonacea

INDIA: BRAHMAPHUTRA DRAINAGE: SMF 68 (1; 95 mm SL) Assam

(26.436147, 92.910461); SMF 9070 (1; 101.6 mm SL) Assam (same coordinates as

SMF 68); SMF 9090-9091 (1; 106.8 mm SL) Assam same locality as SMF 68; SMF

9091-9092 (1;121.1 mm SL) Assam same locality as SMF 68

INDIA: GANGES DRAINAGE: ZMB 4795 (1ex.) (25.393661, 86.429443)

Paracanthocobitis abutwebi, n.sp.

BANGLADESH: MEGHNA BASIN: UMMZ 208782 (39, 10 ex; 40.7-53.3 mm SL)

Piyain Gang River at Songram Punji, 1.5 mi downstream from India (25.183, 89.983);

UF 178719 (3; 24.1-33.0 mm SL) Dhaka, Jamuna River Drainage at Dhaleswari River,

Nandakhali Ferry Ghat (23.799417, 90.24594); UF 178050 (2; 50.2-50.9 mm SL)

“Sunamgari Danio Site” (no coordinates); PARATYPES UF 172474 (2; 37.9-48.8 mm

SL) Sylhet, Changerkhal River (24.95439, 91.7438); CAS 227857 (1; 66.1 mm SL)

canal/swamp in Tongvar Haor; UMMZ 208626 (9; 35.5-52.0 mm SL) Sylhet (N Sylhet)

Piyain Gang River at Songram Punji, 1/4 mi downstream from Indian border

(25.101145, 91.754265); HOLOTYPE UMMZ 208646 (5; 39.5-51.0 mm SL) Sylhet,

Rangapani Khal (creek), 4 mi NNW of Jaintiapur on Sylhet - Shillong HWY (25.140001,

92.056103); CAS 233500 (2; 34.6-35.6 mm SL) Sylhet Ganges River Drainage Nur

Nody Stream upstream from Lubha River at Nihalpur Village (25.03615, 92.30607);

CAS 227855 (1; 50.8 mm SL) same locality as CAS 233500; AUM 50461 (1; 24.9 mm

SL) Savar Dhaka, Dhaleswari River Drainage, Dhaleswari River, ferry ghat near Nanda

Khali (Balu Ghat); UF 172551 (1; 50.8 mm SL) N Sylhet, Kanka River by road to

Jaintiapur from Sylet (24.99072222, 92.06019444);

Paracanthocobitis adelaideae, n.sp.

BURMA: IRRAWADDY BASIN: HOLOTYPE to be removed from USNM 378387

(7; 45.9-51.4 mm SL) Kha Wan, Kha Wan stream (25.085599, 96.96991); PARATYPES to be removed from USNM 378393 (1; 28.2 mm SL) Mayen stream (25.355, 97.063)

Paracanthocobitis botia

INDIA: BRAHMAPUTRA BASIN: NEOTYPE: BMNH 1889.2.1.1562-1571 (10;

37.1-57.7 mm SL), Assam (26.598351, 92.909546); BMNH 1889.2.1.1581-1582 (2;

51.9-55.4 mm SL), Bengal (25.712021, 89.769287); UMMZ 244583 (1; 19.9 mm SL),

India, West Bengal, Dolong River, tributary of Mansai River, vic. of Shildanga

(26.221628, 90.72784).

Paracanthocobitis linypha, n.sp.

BURMA: IRRAWADDY BASIN: PARATYPES: CAS 88918 (10; 38.8-42.9 mm

SL), Sagaing Division, Kalemyo fish markets (23.189521, 94.073045); HOLOTYPE: to

be removed from USNM 378387 (1; 34.9 mm SL), Kha Wan, Kha Wan stream

(25.085599, 96.96991); ex USNM 378384 (20, 10 ex; 29.9-36.3 mm SL), Nan-Kwe

Chaung (20.502602, 96.361713); ex. USNM 372168 (1; 21.3 mm SL), Kachin, Hpa-Lap,

stream of Nam Chim Chaung, of Nankwe Chaung, NW of Myitkyina (25.52666667,

97.37916667); USNM 378400 (58, 10 ex; 29.2-34.4 mm SL), Nan-Kwe Chaung

(20.502602, 96.361713); USNM 378406 (1; 36.6 mm SL), Nanmate stream

(25.38416667, 97.01055556); USNM 372182 (1; 22.5 mm SL), Kachin State, Nankwe

Chaung, W of Myitkyina, on both sides of highway bridge (25.3322, 97.28); ex. USNM

378393 (4; 23.9-26.4 mm SL), Mayen stream (25.355, 97.06333333).

Paracanthocobitis mackenziei

BANGLADESH: MEGHNA BASIN: UF 176610 (3; 37.9-48.8 mm SL), Dhaka,

Jamuna River Drainage at Dhaleswari River, Nandakhali Ferry Ghat

(23.799416666666, 90.2459444444444); AUM 50621 (1; 47.9 mm SL), Sylhet,

Sunamganj River drainage, Sunamganj River/floodplain (25.070051, 91.407423).

INDIA:INDUS BASIN: MCZ 18690 (3; 36.5-37.4 mm SL), Haryana plains, 70 mi SW of

Amballa (29.609282, 76.18721); BMNH 1932.2.20.30 (3; 39.9-58.1 mm SL), Jubbulpore

(Jabbalpur) (32.776883, 74.92527); CAS 41295 (6; 47-57.4 mm SL), Punjab, Kangra

Valley (32.161663, 76.294098).

INDIA: CENTRAL INDIA: BMNH 1889.2.1.1590 (Lectotype of Nemacheilus aureus), (1; 31.9 mm SL), Jabbulpore, India (23.104997, 79.93475); BMNH

1889.2.1.1577-1580 (4; 38.6-43.2 mm SL), Allahabad (25.423957, 81.85); BMNH

1889.2.1.1576 (1; 41.3 mm SL), Loodianah (30.993503, 75.839882).

INDIA: GANGES BASIN: BMNH 1867.5.12.44 (1; 56.0 mm SL), Morar River near

Geralior (26.265526, 78.062689); CAS 32369 (2; 34.9-38.6 mm SL), Bisrampau C.P.

(Connaught Place) (28.646607, 77.264242); CAS 34744 (2; 49.5-59.6 mm SL), Rewa,

Damgarh (Ramgarh?) taken by Nerbudda Survey (23.64075, 85.520096); CAS 41294

(3 ex.) Bihar, Bishrampur, (former Central Province), (27.58283333, 84.714); CAS

32369 (2; 34.9-38.6 mm SL), New Delhi, Bisrampau (Bisrampur?), near CP (Connaught

Place) (28.646607, 77.264242); MCZ 4253 (1; 45.3 mm SL), Haridwar (29.964585,

78.188468).

INDIA: MAHANADI RIVER BASIN: CAS 61839 (10; 41.8-58.8 mm SL), Orissa

State, fish market at Sone Pur (21.154157, 83.879728).

NEPAL: GANGES BASIN: KU 28527 (1; 35 mm SL), Sapari/Sunsari, just upstream from Koshi River Barrage (26.52499952, 86.93499756); KU28570 (5; 23-

40mm SL), Saptari/Sunsari, just downstream from Koshi River Barrage (26.5182991,

86.9266968); KU 28588 (2; 55-68mm SL), Sunsari, associated seepage at Koshi Tappu

Wildlife Refuge (26.62330055, 87.03330231); KU 28620 (5; 44-66mm SL) Sunsari

Bhantabari Seepage at Bhantabari (26.5249996, 86.9732971); KU 28631 (1;

54mm SL), Rautahat/Sarlahi, at Raj-Marg highway (27.10000038, 85.48329926); KU

28648 (8; 29-52mm SL), Dang Rajpur, at Rajpur (27.8432999, 82.5633011); KU 28665

(4; 35-46mm SL), Kanchanpur Brahamadev at Brahamadev (29.08169937,

80.14170074); KU 28681 (3; 33-44mm SL), Kanchanpur , confluence of 3 rivers

(Chaudhar, Bahuni, Gobraiya), at Royal Suklaa Phantaa Wildlife Reserve (28.7166996,

80.1999969); KU 28715 (3; 43-50mm SL), Kanchanpur Mahendranagar At Raj-Marg highway, 9km E of Mahendranagar (28.9517002, 80.2549973 ); KU 28734 (1;

41mm SL), Kailali Waters of Kailali district along Raj-Marg highway (28.7000008,

80.9000015); KU 28759 (1; 60mm SL), Bardiya/Kailali Chisapani at Chisapani

(28.64170074, 81.28170013); KU 28810 (1; 44mm SL), Banke Bhurigaon at

Madhuwaa/Kanchanpur (28.0699997, 81.8467026); KU 28821 (3; 49-57mm SL),

Nawalparasi Tribentighat at Tribentighat above barrage (27.4491997, 83.9216995); KU

28891 (1; 55mm SL), Chitwan/Nawalparasi at Kanahaa river confluence (27.48329926,

83.94000244); KU 28907(9; 36-58mm SL), Chitwan/Nawalparasi, at Gharial

Conservation Project Camp (27.55830002, 84.12000275); KU 28924 (5; 42-58mm SL),

Chitwan/Nawalparasi, just downstream from Lamichaur Military Camp (27.58169937,

84.13670349); KU 28942 (1; 55mm SL), Chitwan/Nawalparasi at Amaltarighat

(27.55330086, 84.11329651); KU 28964 (1; 54mm SL), Nawalparasi at Pier No. 13

(27.41670036, 83.85669708); KU 29077 (2; 48-50mm SL), Tanahun Khairenitar at

Khairenitar (28.0333004, 84.0667038); KU 29138 (4; 57-64mm SL), Jhapa,

Bhadrapur(26.53829956, 88.10169983); KU 29159 (2; 67-75mm SL), lam Mai Khola at

Mai Khola (26.8850002, 87.928299); KU 29180 (1; 55mm SL), Jhapa at Raj-Marg

highway (26.6583004, 87.8700027 ); KU 29190 (2; 45-47mm SL), Morang, Belbari

(26.65999985, 87.41169739); KU 29466 (2; 35-51mm SL), Bardiya at Chapang

(28.3567009, 81.71330261); KU 40585 (1; 48.1 mm SL), Narayani, Chitwan, Rapti

River upstream from confluence w/ Lothar on border of Chitwan Nat. Park

(27.56255556, 84.70475); KU 40367 (1; 42.4 mm SL), Mechi, Jhapa, Mechi River at

Karkarvitta, at bridge crossing at border with India (26.64597222, 88.16230556); KU

40575 (1; 43.5 mm SL), Narayani, Chitwan, Lothar River at bridge crossing highway

through Piple Village (27.58819444, 84.73313889); KU 40408 (1; 39.3 mm SL), Mechi,

Jhapa, Biring River at highway (26.64183333, 87.93730556); KU 40433 (1; 29.3 mm

SL), Kosi, Morang, Lohandra River at Belbari town (26.66288889, 87.40813889); KU

40616 (3; 42.6-45.0 mm SL), Narayani, Chitwan, Narayani River at Narayanghat picinic

spot (27.69547222, 84.40994444); KU 40600 (12; 23.4-33.4 mm SL), Narayani,

Chitwan, Khair Khola at bridge crossing highway E of Ratnanagar Town (27.61841667,

84.53266667); KU 40594 (10; 15.8-26.6 mm SL), Narayani, Chitwan, Sim Pani Ghol

(stream?) at Sim Pani Village (27.58283333, 84.714); KU 40459 (41, 10 ex. 37.7-72.5

mm SL), Kosi, Sunsari, Koshi River diversion, 1km N of highway, around 2 km before

bridge crossing Koshi (26.60402778, 87.02411111); KU 40630 (6; 27.7-46.8mm SL),

Narayani, Chitwan, Budrapti Kohla at Bachauli-3 Baghmara (27.549246, 84.51236); KU

40507 (8; 32.4-63.1mm SL), Beri, Bardiya, mainstem of Karnali River, downstream of

Chisapani (28.6332222, 81.27511111); KU 40647 (5; 21.9-45.3 mm SL), Narayani,

Chitwan, tributary of the Rapti River, Chitwan National Park (27.57241667,

84.49638889); KU 40489 (9; 40.8-52.7mm SL), Kosi, Sunsari, Sunsari River at Inarua

(26.61072222, 87.12744444).

Paracanthocobitis maekhlongensis, n.sp.

THAILAND: MAE KHLONG BASIN: UF 176571 (3; 54.4-57.7 mm SL)

Kanchanaburi, Mae NamKwae, Huay Ban Rai (14o42’48.7”N, 98o31’25.3”E);

HOLOTYPE UF 182864 (2; 56.4-58.9 mm SL) Kanchanaburi, Thong Pha Phum, Ban

Rai, NW of Huai Khayeng close to SW side of reservoir, off road 327 (14o41’25”N,

98o31’16”E); UF 176387 (1; 58 mm SL) Kanchanaburi, Mae Nam Kwae (14o45’10”N,

98o30’2”E); UF 176542 (1; 54.2 mm SL) Kanchanaburi, Huay Pak Kok (14o40’49”N,

98o31’38”E); UF 176437 (1; 51.6 mm SL) Kanchanburi, Mae Name Kwae, Huay Ban

Rai (14o43’9.6”N, 98o30’21”E); PARATYPES UMMZ 209462 (4; 61.3-75 mm

SL)Kanchanaburi Kwai Yai River (14.227113,99.233551)

Paracanthocobitis mandalayensis

BURMA: IRRAWADDY DRAINAGE: ex. USNM 378387 (3; 45.9-51.4 mm SL)

Kha Wan, Kha Wan stream (25.085599, 96.96991); ex. USNM 372168 (1; 42.3 mm SL)

Kachin, Hpa-Lap stream, of Nam Chim Chaung, of Nankwe Chaung, NW of Myitkyina

(25.52666667, 97.37916667); USNM 378384 (4; 50.5-64.2 mm SL) Nan-Kwe Chaung

(20.502602, 96.361713); CAS 88890 (4; 61.9-88.4 mm SL) Kachin, from Myitkyina

Market (25.382495, 97.403297); USNM 378409 (4; 45.7-61.4 mm SL) Nanmate stream

(25.38416667, 97.01055556); USNM 344646 (3; 69.6- 82.4 mm SL) Kachin, Irrawaddy

drainage, Nankwe stream, 10 mi SW of Myitkyina along main N-S road (25.33361111,

97.28083333).

THAILAND: WANG RIVER DRAINAGE: UF 181110 (14, 7 ex.; 34.4-60.5mm SL)

Lampang, Mae Nam Wang Nam Mae Pan, bridge on Hwy 4025 1 km E of Ban Thung

Pong (18.78194444, 98.49111111)

THAILAND: PING RIVER DRAINAGE: UMMZ 209453 (5; 41.8-65.9 mm SL) Chiang

Mai Province (18.795, 99.008056)

Paracanthocobitis mooreh

INDIA: KRISHNA RIVER BASIN: CAS 32619 (4; 34.2-37.1 mm SL),

Maharashtra, Deolali (19.470153, 74.620242); MCZ 52384 (10 of 22 ex.; 34.9-47.8 mm

SL), Vithal Wadi River, Poona (18.392972, 74.002419); MCZ 52380 (11; 40.2-50.5 mm

SL), Poona, Mula River near Chintamani Temple, Theur (18.522585, 74.041486);

BMNH 1889.2.1.1587-1589 (3; 43.2-48.3 mm SL), Poona(18.53789, 73.865723); MCZ

52377 (1; 44.7 mm SL,) Khadakwasle Dam (18.433333, 73.766667).

INDIA: KAVERI RIVER BASIN: BMNH 1870.5.2.17-18 (2; 40.7-41.0 mm SL).

Wynaad (11.423326, 76.24423). Godavari

Paracanthocobitis pictilis

BURMA: ATARAN BASIN: UF 172927 (5; 59.0-72.1 mm SL; 1 cleared and stained, 76.4 mm SL), Ataran drainage, from aquarium fish dealer, Kamphol

Udonmritthiruj, in Pathumithani; UNMF-P 07667 (52, 11 ex: 55-81.2 mm SL), Kraeng

Tor waterfall, Pyathony Su Mon State (15.3884, 98.298281).

Paracanthocobitis rubidipinnis

BURMA: IRRAWADDY BASIN: CAS 89008 (1; 62.5 mm SL), Mandalay fish markets, (21.981255, 96.056099); CAS 231702 (1; 62.2 mm SL), Mandalay/Sagaing

Division, Irrawaddy River, 2 km upstream of upper bridge in Sagaing (21.889854,

95.998821); CAS 231849 (1; 58.5 mm SL), Mandalay Division, Mandalay, Irrawaddy

River, upstream of Mandalay (22.036639, 96.034813); CAS 231517 (1; 48.5 mm SL),

Irrawaddy River, near Ywar Thit Gyi, downstream from Mandalay (21.912, 95.7741);

CAS 231687 (2; 55.9-57.5 mm SL), Irrawaddy River, near Amarapura, downstream

from Mandalay (21.90265, 96.00985); ZRC 43570 (1; 55.5 mm SL), Bago, Irrawaddy

(from fish dealer) (17.433855, 96.484566); BMNH 1893.2.16.52 (1; 71.2 mm SL),

Mandalay (21.987622, 96.039505).

BURMA: ATARAN BASIN: USNM 378401 (3; 46.4-62.7 mm SL), Kawkareik

market (Hlaing Bwe River?), (16.507529, 98.244152).

Paracanthocobitis urophthalma

SRI LANKA: KALU GANGA BASIN: CAS 41298 (1; 45.9 mm SL), "Sri Lanka"

from Colombo museum 1940; CAS 30147 (1; 29.1 mm SL), Ratnapura (6.691887,

80.369568); BMNH 1981.1.21.320 (1; 33.4 mm SL), Gin Ganga, Upper Nanikita Ela

(6.369894, 80.480118); BMNH 1864.7.11.35-36 9, syntypes of Nemacheilus

urophthalmus, (2; 45.8-48.8 mm SL), South Ceylon (6.087642, 80.184202).

Paracanthocobitis zonalternans

BANGLADESH: MEGHNA BASIN: UMMZ 208629 (1 ex.), Sylhet (N Sylhet),

Piyain Gang River at Songram Punji, 1/4 mi downstream from Indian border

(25.101145, 91.754265); ex CAS 233500 (1 ex.), Sylhet, Ganges River Drainage, Nur

Nody stream, upstream from Lubha River, at Nihalpur Village (25.03615, 92.30607);

UMMZ 208629 (1 ex.), UMMZ 208784 (17 ex.), Sylhet (N Sylhet), Piyain Gang River at

Songram Punji, 1.5m downstream from India border, Surma (Meghna) drainage

(25.18333333, 89.98333333).

BURMA: IRRAWADDY BASIN: CAS 34752 (2 ex.). 9 mi NW of Hlegu,

Ngamoeyeik Creek (17.218528, 96.200695); CAS 32368 (13 ex.). Irrawaddy

River/drainage. near Indaw (In Daw?), Chaung and Karnaing (24.22552, 96.143146);

USNM 288452 (26 ex.).

THAILAND: SALWEEN BASIN: Mae Hong Son Province, 5 km W of Mae

sariang (Salaeng?), (18.157772, 97.886098); USNM 378390 (1 ex.) ,Yin Monetan of

Naung Cho township, Northern Shan State (22o14’8”N, 96o39’59”E); USNM 372182 (1

ex.), Kachin State, Nankwe Chaung, W of Myitkyina, on both sides of highway bridge

(25o19’56”N, 97o16’48”E); USNM 378439 (~75; 31.5-39.7 mm SL), Kayin stream, near

Sittallang River, E of Taungoo (18.939088, 96.454711); ZRC 43453 (10 ex.), Shan

State, Ma Gawe River, along Kalaw-Thazi Hwy between Mandalay and Nampantet

(21.396819, 96.244812); UF 30191 (2 ex.), Gyobu Reservoir, 9 km N of Taik-Kyi Taik-

Kyi township, Rangoon (16.831276, 96.25); USNM 378397 (2 ex.), Mandalay Division,

Irrawaddy drainage, aritificial pond in Toegyi Village, near Pyin Oo Lwin on the road

Mandalay-Hsipaw (21.97666667, 96.39).

MALAYSIA: MALAY PENINSULA: MCZ 47226 (1 ex.), Satun Malay Peninsula,,

Boripatra River, on road from Satun to Hat Tai. (6.814626, 99.974098); ZRC 41103 (1 ex.), Perlis Sintok-Pdg. Senai Rd. (6.456424,100.490949); ZRC 46671 (3 ex.), Kedah

Kg. Tandop about 6-7 mi from Baling on road to Weng (5.665334, 100.901245).

THAILAND: MAE KHLONG BASIN: ANSP 179829 (2 ex.), Ulong River, tributary of Mae Nam Khwae Noi, at route 323 bridge, 5-10 km from Thong Pha Phum

(14.782518, 98.669386); UF 176449 (35 ex.), Kanchanaburi, Mae Nam Khwae, Huay

Ban Rai (14.71933333, 98.50583333); UF 176554 (23 ex.), same locality as UF

176449; UMMZ 209461 (7 ex.), Kanchanaburi, Kwai Yai River (14.227113, 99.233551);

ZRC 42002 (2 ex.), Ratchaburi Prov., Suan Pheng Dist., Suan Pheng Waterfall

(13.524024, 99.239007); ROM 48972 (2 ex.), Ranong Province, N of Bang Kaeo, off highway 4 close to border of Burma (10.25, 98.75); UF 182821 (13 ex.), Kanchanaburi,

Thong Pha Phum, near Rt. 3272 (14.69027778, 98.52111111); NIFI 3087 (4 ex.),

Kanchanaburi, Thong Pha Phum Huay Ulong (14.759777, 98.638315)

THAILAND: PENINSULAR THAILAND: UF 182833 (4 ex.), near Phang Nga, off

Rt. 4090 (8.571416667, 98.41783333); UF 183331 (2 ex.), Nakhon Si Thammarat, near

Rt. 4015 (8.348861111, 99.6921111); UMMZ 238962 (2 ex.), Phuket, Nam Tok Ton Sai

(8.026255, 98.363285); ZRC 45717 (8 ex.), Phuket, H.H. Tan, field # THH0006

(7.957237, 98.316936); ZRC 42198 (3 ex.), Ranong, Khlong Phrae, Sai Ban Kraeo Noi

(Bang Kaeo?), km 8 on rd. branching E. 32 km of Ranong on route to Kra Buri

(10.183836, 98.719597); ZRC 42016 (1 ex.), Prachaup Khiri Khan, Thaup Sakae

District W of Ang Thong (11.44855556, 99.48283333); ZRC 42184 (8 ex.), Ranong stream, N of Khura Buri, 100 km S of Ranong (9.199376, 98.412724); ZRC 41974 (5 ex.), Ranong King, Amphae Suk Sam Lan (Ran?), Ton Koi Waterfall (9.353185,

98.429385); SMF 3966 (holotype of Noemacheilus phuketensis), Phuket Island, off the west coast of the south peninsula Thailand (7.991918, 98.342628); SMF 3967 (paratype of Noemacheilus phuketensis ), (1 ex.), same locality as SMF 3966; SMF 4244

(paratype of Noemacheilus phuketensis ) (1 ex.), same locality as SMF 3966.

THAILAND: SALWEEN BASIN: ZSM 27468 (Neotype of Cobitis zonalternans,

33.1 mm SL), Huay Me Charno, 4 km south of Amphoe Mae Romat on rd. 1085; CAS

35775 (6 ex.), Nam Mesarieng, near Mesarieng (Mae Sariang), northern Thailand

(18.151316, 97.934432); CAS 35776 (2 ex.), same locality as CAS 35775; CAS 36042

(2 ex.), same locality as CAS 35775; NIFI 00874 (14; 28.0-32.5 mm SL), Tak Province,

Huai Mae Kamui, Aunphae, Tha Song Yang (17.230005, 98.229046); NIFI 02181 (2;

27.0-27.1 mm SL), Surat Thani Province, Suratthani Tapi Basin, Khlong Sok at Ban

Kheng Sok (8.90678, 98.522415); NIFI 01435 (13 ex.), Kanchanaburi, Khao Noi

(13.966887, 99.589834); NIFI 02057 (7 ex.), same locality as NIFI 01435; NIFI 6393 (1 ex.), Mae Hong Son Province, Mae Sanga River (18.238482, 97.960739); ZRC 41258

(8 ex.), Tak, Salween Basin, Mae Nam Moi, S of Phop Phra (16o17’25”N, 98o42’20.4”E);

ROM 51139 (5 ex.). Tak Province, N of Mae Charao (16.9667, 98.5667).

CHAPTER 3 SYSTEMATIC ACCOUNTS

Genus Acanthocobitis

Acanthocobitis Peters 1861:712. Type-species: A. longipinnis Peters 1861 by

monotypy.

Diagnosis

Member of Nemacheilidae. Distinguished from all other genera in family except

Paracanthocobitis in having the lower lip with a large papillated pad on either side of a

medial interruption; upper lip with two to five rows of papillae and continuous with lower lip (Fig.3-1); conspicuous black spot with white outline on upper half of caudal-fin base.

Acanthocobitis is distinguished from Paracanthocobitis in having a pointed (vs. emarginate or truncate) caudal fin; enlarged, fleshy rostral barbels (Fig. 3-1); absence of patch of adipose tissue on anterior end of lateral line; long dorsal fin (> 33% of SL with

17-19 branched rays vs. 9-15 in Paracanthocobitis), triangular-shaped (vs. rounded) head (Fig.3-3), and vertically (vs. horizontally) oriented suborbital flap (Fig.3-2).

Acanthocobitis is monotypic containing only A. pavonacea.

Description

Heavily papillated lips; oval-shaped ocellus; ossified sub-orbital flap in male oriented vertically; 5-6 concentric rows of spots in dorsal fin; 10-11 transverse bands in caudal-fin; lateral line complete; axillary pelvic lobe absent; color pattern variable with

18-20 thin saddles continuing down side to below lateral line, small spots sometimes between saddles and bars; spots and dashes on head and opercle, interrupted stripe from orbit to tip of snout. Branched dorsal-fin rays 17-19; branched pectoral-fin rays 12-

13; branched pelvic-fin rays 8; branched anal-fin rays 6; branched caudal-fin rays 9+8 or

9+9.

Distribution

Acanthocobitis is known from the Ganges and Brahmaputra river basins of northern India and Bangladesh.

Remarks

Menon (1987) treated Acanthocobitis longipinnis, type species of Acanthocobitis, as a synonym of Acanthocobitis botia. Grant (2007) suggested that A. longipinnis was a valid species distinct enough to be placed in its own subgenus, and placed all other species of Acanthocobitis in a new subgenus, Paracanthocobitis with type species

Cobitis zonalternans Blyth 1860. Differences separating Acanthocobitis from

Paracanthocobitis included caudal-fin shape (pointed), suborbital flap orientation and

size (vertical and enlarged), and body shape (slender).

The same characters described by Grant (2007) were observed in the present

study as were other characters separating Acanthocobitis from Paracanthocobitis:

enlarged fleshy rostral barbels and lack of a patch of adipose tissue on the lateral line.

The two subgenera of Grant are treated herein as genera.

After observing types of Acanthocobitis longipinnis and Cobitis pavonacea, Grant

(2007) suggested that A. longipinnis from the Ganges River basin is a junior synonym of

A. pavonacea, from the Brahmaputra River basin. The type specimens of both nominal

species were observed in the present study and found to be in poor condition (Fig. 3-4,

3-5). The synonymy proposed by Grant (2008) will be upheld until, and if, fresh material

is collected and the two forms can be diagnosed.

Acanthocobitis pavonacea (McClelland, 1839)

Cobitis pavonacea McClelland, 1839, Asiatic Researches v.19 (pt 2) 305, 437, pl.52 (fig.

1). Assam, India Brahmaputra Basin.

Acanthocobitis longipinnis Peters 1861, Monatsberichte der Königlichen Preuss[ischen]

Akademie der Wissenschaften zu Berlin 712. Ganges River, India.

Noemacheilus pavonaceaus.—Menon 1987, The fauna of India and the adjacent

countries. Pisces. Vol. IV. Teleostei Cobitoidea. Part 1. Homalopteridae. Zoological

Survey of India. i-x + 1-259.

Acanthocobitis pavonacea.— A new subgenus of Acanthocobitis Peters, 1861

(Teleostei: Nemacheilidae). Ichthyofile No. 2: 1-9.

Diagnosis

See account for Acanthocobitis.

Description

Body deepest under dorsal fin or predorsally; head triangular-shaped when

viewed laterally; dorsal fin long (> than 33% of SL); caudal fin lanceolate-shaped with

12-13 transverse bands; male with ossified suborbital process; lateral line complete;

rostral barbels thick and fleshy; axillary pelvic lobe absent; branched dorsal-fin rays 17-

19, branched pectoral-fin rays 12-13; branched caudal-fin rays 8+8 or 9+8. Maximum

SL = 121.1 mm female.

Color pattern consisting of thin bars, much narrower than interspaces, extending

from dorsum to below lateral line until distal extremity of dorsal fin, then touching venter

post-dorsal, sometimes with small spots between. Head covered in irregular-shaped

blotches, with interrupted stripe, broken in center extending from orbit to top of snout.

Distribution

Acanthocobitis pavonacea is known from northeastern India in the Ganges River

basin and western Assam, India, in the Brahmaputra River basin (Fig.3-6).

Genus Paracanthocobitis

Paracanthocobitis Grant 2007:3. Type-species: Cobitis zonalternans Blyth, 1860 by

original designation.

Diagnosis

Member of Nemacheilidae. Distinguished from all other genera in family except

Acanthocobitis in having heavily papillated lips, upper lip with three to five rows of

papillae and continuous with lower-lip pads (Fig. 3-7); conspicuous black spot with white

outline on upper half of caudal-fin base.

Paracanthocobitis is distinguished from Acanthocobitis in having an emarginate

or truncate (vs. pointed) caudal fin; rostral barbels uniform in size; patch of adipose

tissue on anterior end of lateral line (Fig. 3-8); 9-15 branched dorsal-fin rays (17-19 in

Acanthocobitis), rounded (vs. triangular-shaped) head. Paracanthocobitis has a horizontally oriented suborbital process formed from a fusion of the lateral ethmoid and

the first infraorbital (Fig. 3-10). In adult males of some species of Paracanthocobitis,

there is a free fleshy extension called the suborbital flap on the lateral ethmoid. In other

species there is no free flap. Instead, the skin covering the lateral ethmoid is fused

dorsally and posteriorly but not ventrally, creating a suborbital groove. A well-defined suborbital groove, visibly associated with the lateral ethmoid, is found only in adult males. However, in species with a groove, the females often have a shallower, less well-defined groove, and the lateral ethmoid is not visible (Fig. 3-9).

Description

Heavily papillated lips; dark circular or sliver-shaped caudal spot with light outline; suborbital flap or groove in male; 3-6 concentric rows of spots in dorsal fin, 2-3 in anal fin; 4-11 transverse bands in caudal-fin; lateral line complete or incomplete; axillary pelvic lobe present or absent; color pattern variable intraspecifically; all species with dark spots and blotches on head and opercle, uninterrupted stripe from orbit to tip of snout; nasal cavity significantly larger (at least twice as large) than naris. Branched dorsal-fin rays 10-15; branched pectoral-fin rays 10-13; branched pelvic-fin rays 8; branched anal-fin rays 6-7; branched caudal-fin rays 9+8 or 9+9.

Distribution

Paracanthocobitis is known from eastern Pakistan in the upper Indus basin, east to Cambodia and Laos in the Mekong basin (Rainboth 2013). The northern limit of the range is the Ganges River basin in Nepal, with the southern limit in peninsular Sri Lanka and Malaysia.

Paracanthocobitis abutwebi, n.sp.

Holotype

UMMZ 208646 (46.1 mm SL male) Sylhet, Bangladesh, Rangapani Khal (creek), 4 mi

NNW of Jaintiapur on Sylhet - Shillong HWY (25.140, 92.056), W Rainboth, A Rahman, and S Ahmed, 19-February-1978.

Paratypes

UF 172474 (2; 37.9-48.8 mm SL) Sylhet, Bangladesh, Changerkhal River (24.954,

91.744), RL Mayden, 05-December-2007.

Diagnosis

(Fig. 3-11, Table 3-1)

Paracanthocobitis abutwebi differs from all other Paracanthocobitis in having combination of complete lateral line; color pattern of alternating large and small dark spots along lateral line, small dark dorsal saddles, speckled pattern on upper side between dorsal saddles; 4-5 dark bands in caudal-fin of adult; and 6-7 anal-fin rays; axillary pelvic lobe absent.

Description

Body depth more or less uniform, slightly deeper at dorsal-fin origin; branched dorsal-fin rays 12-13 (usually 12); branched pectoral-fin rays 10-13; branched pelvic-fin rays 8; branched anal-fin rays 6-7 (usually 7); branched caudal-fin rays 9+9; transverse bands in caudal fin 4-5; suborbital flap in male; axillary pelvic lobe absent; lateral line complete with 87-103 pores. Maximum SL = 53.3 mm female.

Color pattern consisting of 10-13 small dorsal saddles not extending to lateral line and becoming closer together towards the head sometimes with small spots or dashes in-between. Ten-16 dark spots along lateral line, spots not extending onto venter. Speckled pattern on upper side and sometimes extending onto lateral line.

Comparison

Paracanthocobitis abutwebi can be distinguished from all other

Paracanthocobitis, except P. botia, P. urophthalma, P. linypha and P. mooreh, in not having an axillary pelvic lobe. Paracanthocobitis abutwebi differs from P. linypha and P. mooreh in having a complete lateral line, and differs from P. urophthalma in not having dark bars on the side that extend across the lateral line onto the venter with narrower

interspaces. Paracanthocobitis abutwebi differs from P. botia in having alternating large

and small dark spots along the lateral line and a speckled pattern on upper side, 4-5

(vs. 6-7) dark bands on the caudal fin in the adult, and 6-7 (usually 7, vs. 6 in P. botia)

anal-fin rays.

Distribution

Paracanthocobitis abutwebi is known from the Meghna and lower Brahmaputra

and Ganges river drainages of Bangladesh (Fig. 3-12).

Etymology

Named for Professor Abu Tweb Abu Ahmed for his contributions to the study of

hillstream fishes of Bangladesh.

Paracanthocobitis adelaideae, n.sp.

Holotype

USNM 378387 (50.6 mm SL male) Kha Wan, Kha Wan stream, Myanmar (25.0856,

96.970), R. Britz, 02-April-2003.

Paratypes

ex. USNM 378393 (1; 28.2 mm SL) Mayen stream, Myanmar (25.355, 97.063), R. Britz,

02-April-2003.

Diagnosis

(Fig. 3-13, Table 3-2)

Paracanthocobitis adelaideae differs from other Paracanthocobitis in having

complete lateral line, axillary pelvic lobe, 12 dorsal rays, and presence of a free

suborbital flap in males, dark dark brown dorsal saddles connected to large squarish

blotches on side along and below lateral line. Caudal spot irregular shaped and located

more dorsally; head wide (interorbital 33.3 % of SL).

Description

Body depth wider pre-dorsally than post dorsally; branched dorsal-fin rays 12;

branched pectoral-fin rays 10-12 (usually 11); branched pelvic-fin rays 8; branched

anal-fin rays 6; branched caudal-fin rays 9+9; transverse bands in caudal fin 4-5;

suborbital flap present in male; axillary pelvic lobe present; lateral line complete with 72-

88 pores. Maximum SL= 51.4 mm female.

Color pattern consisting of a heavily pigmented wave or m-shaped pattern

sometimes appearing in juveniles as a checkerboard along the lateral line (Fig. 3-13).

Dorsal is considerably darker than ventral. Caudal spot irregular shaped and located

more dorsally than laterally. Head pigmented with spots, dashes or large dark blotches.

Comparison

Paracanthocobitis adelaideae is distinguished from P. mooreh, P. zonalternans,

and P. linypha in having a complete lateral line. It is differentiated from P.

mandalayensis, P. maekhlongensis, and P. pictilis in having a suborbital flap (vs.

suborbital groove) in males. It is differentiated from P. botia, P. urophthalma, and P.

abutwebi in having an axillary pelvic lobe. Paracanthocobitis adelaideae and P.

rubidipinnis differ in number of branched dorsal rays (12 in P. adelaideae and 14-15 in

P. rubidipinnis). Paracanthocobitis adelaideae is distinguished from P. mackenziei in

having large squarish blotches on side along and below lateral line vs. small dark

blotches on side just below lateral line, a greater interorbital width (33.3% vs 28.3% SL),

and in a dorsally positioned, irregular-shaped caudal spot.

Distribution

Paracanthocobitis adelaideae is known from the Irrawaddy River drainage of

northern Burma (Fig. 3-14).

Etymology

Named for Adelaide Singer, the daughter of the author. She was born during the revision.

Paracanthocobitis botia (Hamilton, 1822)

Cobitis botia Hamilton, F., 1822:350, 394, An account of the fishes found in the river

Ganges; northeastern Bengal. No types known.

Cobitis bilturio Hamilton, F., 1822:358, 395, An account of the fishes found in the river

Ganges; Brahmaputra River [at Goalpara], India. No types known.

Cobitis turio Hamilton, F., 1822:358, 395, An account of the fishes found in the river

Ganges; Brahmaputra River at Goalpara, India. No types known.

Cobites (Acoura) argentata Swainson, W., 1839:310, The natural history and

classification v. 2; Based on Ham. 358. No. 10 -- which is Hamilton 1822:358, "10th

Species.-- Cobitis turio." Unneeded replacement name for Cobitis turio Hamilton 1822.

Cobitis bimucronata McClelland, J., 1839:304, 435, Pl. 51 (fig. 4), Asiatic Researches v.

19 (pt 2); northeastern Bengal. Based on Hamilton manuscript drawings (Menon

1987:141).

Cobitis gibbosa McClelland, J., 1839:304, 436, Pl. 52 (fig. 7b), Asiatic Researches v. 19

(pt 2); Brahmaputra River [at Goalpara], India. Unneeded replacement name for Cobitis

turio Hamilton 1822.

Cobitis ocellata McClelland, J., 1839:304, 436, Pl. 51 (fig. 6), Asiatic Researches v. 19

(pt 2); Brahmaputra River [at Goalpara], India. No types known. Unneeded replacement name for Cobitis bilturio Hamilton 1822.

Cobitis arenata Valenciennes, A., in Jacquemont 1840:, Pl. 15 (fig. 1), Voyage dans

l'Independant les années 1828 à 1832; India. Holotype: MNHN 0000-3811.

Nemacheilus botia.—Shrestha 1978:37, Fish fauna of Nepal. Journal of Natural History

Museum Tribhuvan University v. 5 (1-4): 33-43.

Acanthocobitis botia.—Kottelat, M., 1990:28, Indochinese nemacheilines. A revision of

nemacheiline loaches (Pisces: Cypriniformes) of Thailand, Burma, Laos, Cambodia and

southern Viet Nam. Verlag Dr. Friedrich Pfeil, München. 1-262.

Acanthocobitis (Paracanthocobitis) botia.—Grant, S., 2007, A new subgenus of

Acanthocobitis Peters, 1861 (Teleostei: Nemacheilidae). Ichthyofile No. 2: 1-9.

Diagnosis

(Fig. 3-15, Table 3-3)

Paracanthocobitis botia is distinguished from other species of Paracanthocobitis

by the combination of complete lateral line; large, dark patch on opercle; suborbital flap

not terminally attached in male, axillary pelvic lobe absent. Color pattern with small

dorsal saddles and 8-10 dark blotches, some connected by pigment, along and just

below lateral line.

Description

Body depth slightly increasing from head to dorsal-fin origin, then decreasing to

caudal peduncle; branched dorsal-fin rays 11-12; branched pectoral-fin rays 10 -11;

branched pelvic-fin rays 8; branched anal-fin rays 6; branched caudal-fin rays 9+9; transverse rows of spots in dorsal fin4-5 (usually 5); dark bands in caudal fin 5-7

(usually 5); male with suborbital flap; axillary pelvic lobe absent; lateral line complete with 83-100 pores. Maximum SL= 57.7 mm female.

Color pattern with 8-10 dark spots along side just below lateral line. Head covered in medium to large spots with large dark patch on opercle. Small dorsal saddles narrower than, or equal to, interspaces not extending to lateral line. Irregular pigmentation between saddles and lateral line. Ocellus and caudal spot extending onto caudal fin.

Comparison

Paracanthocobitis botia differs from all other species of Paracanthocobitis except

P. urophthalma, P. abutwebi, P. mooreh and P. linypha by the absence of an axillary

pelvic lobe. Paracanthocobitis mooreh and P. linypha can be distinguished by the

presence of an incomplete lateral line. Paracanthocobitis urophthalma differs in having

bars on side of body that extend from dorsum to ventor. Paracanthocobitis abutwebi

differs from P. botia in having 11-16 spots along the lateral line (vs. 8-10) sometimes

with a speckled pattern along lateral line, 4-5 bands in the caudal fin (vs. 5-7), and 6-7

(usually 7) anal-fin rays.

Distribution

Paracanthocobitis botia is known from the mainstem, and possibly tributaries of

the Brahmaputra River, in Assam, India (Fig. 3-16).

Remarks

Paracanthocobitis botia has been the most widely used name for species of

Paracanthocobitis. The original description of Cobitis botia by Hamilton (1822) states

that it was found in rivers of northeastern Bengal; no types are known. After a review of

a map of “The British Indian Empire” from the Imperial Gazetteer of India (1909) and a

map of “The Bengal Presidency at its fullest extent in 1858,” it can be said with

confidence that “northeastern Bengal” is located in what is modern-day Assam, India, in

the Brahmaputra basin. A specimen from BMNH 1889.2.1.1562-1571 is here selected

as the neotype (Fig. 3-15).

Paracanthocobitis linypha, n.sp.

Holotype

USNM 378387 (34.9 mm SL male) Kha Wan, Kha Wan stream, Myanmar (25.0856,

96.970) ), R. Britz, 02-April-2003.

Paratypes

CAS 88918 (10; 38.8-42.9 mm SL) Sagaing Division,Myanmar Kalemyo fish markets

(23.190, 94.073), CJ Ferraris, D Catania, U Myint Pe, 25-April-1996.

Diagnosis

(Fig. 3-17, Table 3-4)

Paracanthocobitis linypha can be distinguished from all other Paracanthocobitis

in having a color pattern consisting of thin 10-14 dark bars much narrower than

interspaces sometimes alternating between a shorter and longer bar, incomplete lateral

line not extending past distal extremity of pelvic fin; axillary pelvic lobe absent.

Description

Body depth widest under dorsal fin; branched dorsal-fin rays 11-12; branched

pectoral-fin rays 10-12; branched pelvic-fin rays 8; branched anal-fin rays 6-7 (usually

6); branched caudal-fin rays 9+9; transverse bands in caudal fin 5-7; suborbital flap

present in male; axillary pelvic lobe absent; lateral line incomplete with 22-42 pores.

Maximum SL = 42.9 mm male.

Color pattern consisting of 10-14 small, dark triangular saddles connecting to bars along lateral side that extend from the upper side to lower side and alternating back and forth between long bars that extend past lateral line to shorter line or dash not extending past lateral line. Interspaces much larger than stripes. Pigment on dorsal portion of head darker than ventral.

Comparison

Paracanthocobitis linypha can be distinguished from all Paracanthocobitis except

P. zonalternans and P. mooreh in having an incomplete lateral line extending at most to the distal extremity of pelvic fin. Paracanthocobitis linypha differs from P. zonalternans in not having an axillary pelvic lobe. It differs from P. mooreh in having a color pattern consisting of alternating bars rather than a spotted pattern along lateral line.

Distribution

Paracanthocobitis linypha is known from the Irrawaddy drainage in northern

Burma (Fig. 3-18).

Etymology

From the Latin noun linyphus (m) (linen weaver). The narrow bars on the side of

this species is reminiscent of a sewing needle in motion.

Paracanthocobitis mackenziei (Chaudhuri, 1910)

Nemachilus mackenziei Chaudhuri, B. L., 1910:183, Records of the Indian Museum

(Calcutta) v. 5 (pt 3); northern India (several localities). Syntypes: (11) ZSI F2017/1 (1),

4170-4171 (2), 4172/1-4173/1 (2).

Nemacheilus botia.—Shrestha 1978:37, Fish fauna of Nepal. Journal of Natural History

Museum Tribhuvan University v. 5 (1-4): 33-43.

Diagnosis

(Fig. 3-19, Table 3-5)

Paracanthocobitis mackenziei is distinguished from all other Paracanthocobitis in having complete lateral line, axillary pelvic lobe, and thin dark dorsal saddles connected to small dark blotches on side just below lateral line.

Description

Large size (adults reaching over 70 mm SL); body deeper pre-dorsally than post- dorsally; branched dorsal-fin rays 12-13 (usually 12); branched pectoral-fin rays 10-12

(usually 11); branched pelvic-fin rays 8; branched anal-fin rays 6-7 (usually 6); branched caudal-fin rays 9+9; transverse bands in caudal fin 5-7; suborbital flap present in male; axillary pelvic lobe present; lateral line complete with 80-100 pores. Maximum SL= 75.1 mm female.

Color pattern consists of 11-13 thin dorsal saddles that connect to blotches along lateral line forming a “zig-zag” pattern dipping below the lateral line. Pattern variable in this species with some having larger zig-zag pattern, disconnected saddles, or long s- shaped saddles that extend below lateral line.

Comparison

Paracanthocobitis mackenziei is distinguished from P. mooreh, P. zonalternans,

and P. linypha in having a complete lateral line. It is differentiated from P.

mandalayensis, P. maekhlongensis, and P. pictilis in not having a suborbital groove in

the male. Paracanthocobitis mackenziei is differentiated from P. botia, P. urophthalma,

P. linypha, P. mooreh and P. abutwebi in having an axillary pelvic lobe.

Paracanthocobitis mackenziei and P rubidipinnis differ in their number of branched

dorsal-fin rays (12-13 in P. mackenziei and 14-15 in P. rubidipinnis). It differs from P.

adelaideae in having a narrower interorbital width (28.3 % vs 33.3 % SL), and small

dark blotches on side just below lateral line vs. large squarish blotches on side along

and below lateral line.

Distribution

Paracanthocobitis mackenziei is the most widely distributed species of

Paracanthocobitis, and is known from the Ganges River basin of Nepal and northern

India, the Meghna River basin in eastern Bangladesh, the Mahanadi River basin in

eastern India, Narmada River basin in central India, and the upper Indus River basin of

northern India and eastern Pakistan (Fig. 3-20).

Remarks

Nemachilus mackenziei was described by Chaudhuri in 1910 from several

localities in the Ganges River basin in the modern states of Uttar Pradesh and Bihar in

northern India. The description gave detailed information about color and other

morphological characteristics of the type specimens, including presence of a suborbital

flap in male, complete lateral line, 5 or fewer bands in the dorsal fin, 6 bands in the

caudal fin, and a color pattern on the body consisting of black patches and spots

interspersed irregularly with 12-13 wedge-shaped blackish brown, inverted cone-like markings some of which extend below the lateral line and are unattached from one another. The description also noted that this pattern varied among individuals. Although this species was treated as a synonym of Paracanthocobitis botia (Hamilton) by Menon

(1987) and subsequent authors, it is distinct and resurrected from the synonymy of P. botia.

Nemacheilus aureus was described by Day (1872) from specimens collected at

Jabbalpúr (Jabalpur on modern maps), India. Day noted that the specimens he examined, from September 1871, had an incomplete lateral line and "were full of ova."

Whitehead and Talwar (1976) suggested that BMNH 1889.2.1.1587-90 and ZSI 2574

are syntypes of N. aureus, and that the specimens at ZSI are lost.

BMNH 1889.2.1.1587-89 consists of three specimens from Poonah (now Pune),

India. All three have an incomplete lateral line ending approximately above the origin of the pelvic fin, and the two largest have been dissected and ova are visible. BMNH

1889.2.1.1590 contains one specimen from Jabbalpúr; it has not been dissected and has a lateral line extending to the caudal-fin base. Given that one lot (BMNH

1889.2.1.1587-89) contains specimens with the characters described by Day, and the other (BMNH 1889.2.1.1590) is from the locality cited, it seems likely that Day included all of these specimens in the description as suggested by Whitehead and Talwar

(1976). Although Day mentions a suborbital flap in N. aureus, none is present in the specimens at BMNH. A flap must have been present on ZSI 2574, now lost.

With an incomplete lateral line, 11-12 dorsal-fin rays, and dark spots and blotches on the side of the body, specimens from Pune (BMNH 1889.2.1.1587-89) are

referable to P. mooreh. Pune is on the Bhima River in the Krisha River basin about 100 km southeast of Mumbai.

The one specimen from Jabalpur (BMNH.2.1.1590) has a complete lateral line,

vertical bars on the side, and is here selected as the lectotype of Nemacheilus aureus,

and is treated as incertae sedis given that the specimen is faded and sufficient

characters to diagnose its identity are not present. Jabalpur is in the state of Madhya

Pradesh on the Narmada River, which flows into the Arabian Sea at Bharuch.

Paracanthocobitis maekhlongensis, n.sp.

Holotype

UF 182864 (58.9 mm SL female) Kanchanaburi Province, Thailand, Thong Pha Phum,

Ban Rai, NW of Huai Khayeng close to SW side of reservoir, off road 327 (14o41’25”N,

98o31’16”E), R Plongsesthee, R Singer, Z Martin, and Z Randall, 31-December-2011.

Paratypes

UMMZ 209462 (4; 61.3-75 mm SL)Kanchanaburi Province, Thailand, Kwai Yai River

(14.227, 99.234), J Karnasuta, March-1975.

Diagnosis

(Fig. 3-21, Table 3-6)

Paracanthocobitis maekhlongensis differs from all other Paracanthocobitis in

having a suborbital groove in males, complete lateral line,13-14 branched dorsal-fin

rays, and color pattern consisting of dorsal saddles connecting to oval-shaped spots

along lateral line.

Description

Body deepest at nape, gradually decreasing to caudal peduncle; branched

dorsal-fin rays 13-14; branched pectoral-fin rays 12-13 (usually 12); branched pelvic-fin

rays 8; branched anal-fin rays 7; branched caudal-fin rays 9+9; transverse bands in

caudal fin 7-8; suborbital groove in adult male; axillary pelvic lobe present; lateral line

complete with 95-108 pores. Maximum SL = 75 mm male.

Color pattern consisting of 12-13 dark-brown triangular-shaped dorsal saddles;

saddles wider than interspaces, most have light-colored spot in center. Sliver or oval-

shaped blotches in between each saddle and along the lateral line. Pectoral- and pelvic-

fin base with red band and red coloration in dorsal and caudal fins. Top of head dark

with a well-defined interorbital stripe. Dark, round caudal peduncle spot.

Comparison

Paracanthocobitis maekhlongensis differs from all other Paracanthocobitis

except P. pictilis and P. mandalayensis in having a suborbital groove in males.

Paracanthocobitis maekhlongensis can be differentiated from P. mandalayensis in

having 13-14 (vs. 12) dorsal-fin rays. Paracanthocobitis maekhlongensis differs from P.

pictilis in not having enlarged dorsal pterygiophores and neural spines (Fig. 3-22) and a

round (vs. sliver-shaped) caudal spot. Additionally, P. pictilis has 6 (vs. 7) anal-fin rays, and a shorter head (6.5% vs 9.8% SL in P. maekhlongensis).

Distribution

Paracanthocobitis maekhlongensis is known from the Mae Khlong River basin,

Kanchanaburi Province, Thailand (Fig. 3-23).

Remarks

Kottelat 2012 described P. pictilis from the Ataran River basin of Burma and the

Mae Khlong River basin of Thailand. However, the Mae Khlong population is easily

distinguished from that in the Ataran basin. The type locality for P. pictilis is in the

Ataran, and specimens identified as P. pictilis from the Mae Khlong by Kottelat (2012) are P. maekhlongensis.

Etymology

Named for the Mae Khlong River in western Thailand, where this species appears to be endemic.

Paracanthocobitis mandalayensis (Rendahl 1948)

Nemacheilus rubidipinnis mandalayensis Rendahl, 1948, Die Süßwasserfische Birmas.

I. Die Familie Cobitidae. Arkiv för Zoologi v. 40 A (no. 7): 1-116. Holotype: NRM 13179

Nemacheilus rubidipinnis mandalayensis.—Menon, 1987, The fauna of India and the

adjacent countries. Pisces. Vol. IV. Teleostei - Cobitoidea. Part 1. Homalopteridae.

Zoological Survey of India. i-x + 1-259, Pls. 1-16.

Acanthocobitis mandalayensis.—Kottelat, 2012, Acanthocobitis pictilis, a new species of

loach from Myanmar and Thailand (Teleostei: Nemacheilidae). Zootaxa No. 3327: 45-

52.

Diagnosis

(Fig. 3-24, Table 3-7)

A species of Paracanthocobitis distinguished from congeners in having a long

suborbital groove, 11-12 branched dorsal rays, long caudal fin (~22.9% of SL) and 7-10

dark bands on caudal fin.

Description

Body deepest at nape; branched dorsal-fin rays 11-12; branched pectoral-fin rays

11-13; branched pelvic-fin rays 7-8; branched anal-fin rays 6-7; branched caudal-fin

rays 9+9; transverse rows of spots in dorsal fin 6-7; transverse bands in caudal fin 8-10;

axillary pelvic lobe present; lateral line complete with 67-103 pores. Maximum SL= 88.4

mm male.

Color pattern changes ontogenetically with juveniles having thin dorsal saddles

that extend just beyond the lateral line and connect to spots along lateral line (Fig. 3-

25). As the fish matures the saddles recede from the lateral line leaving oval or square-

shaped blotches wider than or equal to interspaces. Dorsum much darker than below

lateral line, with small saddles narrower than interspaces.

Comparison

Paracanthocobitis mandalayensis differs from all other Paracanthocobitis except

P. pictilis and P. maekhlongensis in having a suborbital groove in males.

Paracanthocobitis pictilis and P. maekhlongensis differ from P. mandalayensis in having

13-14 dorsal rays and lack the dark dorsal pigmentation with small saddles.

Distribution

Paracanthocobitis mandalayensis is known from the Irrawaddy drainage of

northeastern Burma, and the Wang and Ping rivers in the Chao Phraya River basin of

northwestern Thailand (Fig. 3-26).

Paracanthocobitis mooreh (Sykes, 1839)

Cobitis mooreh Sykes, 1839, On the fishes of the Deccan. Proceedings of the

Zoological Society of London 1838 (pt 6): 157-165. No type known.

Nemacheilus sinuatus Day, 1870, Notes on some fishes from the western coast of

India. Proceedings of the Zoological Society of London 1870 (pt 2): 369-374 [1-6]

Nemacheilus moreh.—Menon, 1987, The fauna of India and the adjacent countries.

Pisces. Vol. IV. Teleostei - Cobitoidea. Part 1. Homalopteridae. Zoological Survey of

India. i-x + 1-259.

Noemacheilus moreh.—Talwar & Jhingran, 1991, Inland fishes of India and adjacent

countries. In 2 vols. Oxford & IBH Publishing Co., New Delhi, Bombay, Calcutta. v. 1-2:

i-xvii + 36 unnumbered + 1-1158, 1 pl, 1 map. [V. 1, i-liv + 1-541, 1 map; v. 2, i-xxii +

543-1158, 1 pl.].

Acanthocobitis moreh.—Jayaram, 1999, The Freshwater fishes of the Indian region.

Narendra Publishing House, Delhi, xxvii+1-551.

Acanthocobitis mooreh.—Kottelat, 2012, Acanthocobitis pictilis, a new species of loach from Myanmar and Thailand (Teleostei: Nemacheilidae). Zootaxa No. 3327: 45-52.

Holotype: MHNG 2727.066

Acanthocobitis (Paracanthocobitis) mooreh.— Grant, 2007, A new subgenus of

Acanthocobitis Peters, 1861 (Teleostei: Nemacheilidae). Ichthyofile No. 2: 1-9.

Diagnosis

(Fig. 3-27, Table 3-8)

A species of Paracanthocobitis distinguished from other Paracanthocobitis in having an incomplete lateral line ending at the origin of the pelvic fin, lack of axillary pelvic lobe, 4-5 transverse bands in caudal fin, and a color pattern consisting of spots that become farther apart from anterior to posterior.

Description

Body depth wider pre-dorsally than post-dorsally; branched dorsal-fin rays 11-12

(usually 12); branched pectoral-fin rays 11-12; branched pelvic-fin rays 8; branched anal-fin rays 6-7 (usually 6); branched caudal-fin rays 9+9; transverse rows of spots in

dorsal fin 3-5; transverse bands in caudal fin 4-5; male with suborbital flap; axillary

pelvic lobe absent or rarely rudimentary; lateral line incomplete, ending at pelvic-fin

origin with 28-39 pores. Maximum SL= 50.5 mm female.

Color pattern consists of a series of small spots along lateral line, equal to or

narrower than interspaces, becoming more distant towards caudal peduncle. Smaller spots interspaced irregularly in-between lateral-line spots. Wedge-shaped dorsal

saddles, narrower than interspaces, occasionally with irregular blotches in-between.

Comparison

Paracanthocobitis mooreh differs from all other Paracanthocobitis except P.

zonalternans and P. linypha in having an incomplete lateral line. Paracanthocobitis

mooreh differs from P. linypha in having spots along the lateral line versus alternating

bars. It is differentiated from P. zonalternans in having 12 vs.10 dorsal-fin rays, and P.

zonalternans has a well-defined axillary pelvic lobe where P. mooreh usually lacks one.

Distribution

Paracanthocobitis mooreh is found in the Godavari, Krishna, and Kaveri river

basins of western and southern India (Fig. 3-28).

Remarks

Nemacheilus sinuatus Day, 1870 was described from Wynaad (now Wayanad)

on the Bhavani River in the Kaveri River basin, Karnataka, India. Although this locality is

much further south than other known localities for P. mooreh, the two syntypes of N.

sinuatus (BMNH 1870.5.2.17-18; Whitehead & Talwar 1976:157) are assignable to P.

mooreh.

Paracanthocobitis pictilis (Kottelat 2012)

Acanthocobitis pictilis Kottelat, 2012, Acanthocobitis pictilis, a new species of loach from

Myanmar and Thailand (Teleostei: Nemacheilidae). Zootaxa No. 3327: 45-52. Holotype:

MHNG 2727.066

Diagnosis

Paracanthocobitis pictilis can be distinguished from all other Paracanthocobitis in

having the combination of a small, sliver-shaped ocellus on the caudal peduncle and a

suborbital groove in males.

Description

(Fig. 3-29, Table 3-9)

Body depth deeper pre-dorsally than post dorsally; branched dorsal-fin rays 13-

14 (usually 13); branched pectoral-fin rays 11-13 (usually 13); branched pelvic-fin rays

8; branched anal-fin rays 6; branched caudal-fin rays 9+9; transverse rows of spots in dorsal fin 4-6; transverse bands in caudal fin 8-11; males with suborbital groove; axillary

pelvic lobe present; lateral line complete with 94-97 pores. Maximum SL= 81.2 mm

female.

Color pattern consisting of 12-15 thin dorsal saddles lacking a stripe of

pigmentation in the center, and not extending below lateral line. Lateral line with

numerous speckles and dashes above and below the lateral line. Top of head dark with

a well-defined interorbital stripe. Spot on caudal peduncle small and sliver shaped.

Comparison

Paracanthocobitis pictilis differs from all other Paracanthocobitis except P.

maekhlongensis and P. mandalayensis in having a suborbital groove in males.

Paracanthocobitis pictilis can be differentiated from P. maekhlongensis and P.

mandalayensis in having small spots and speckles along the lateral line (vs. large oval or square-shaped spots) and having a sliver-shaped ocellus on the caudal peduncle.

Paracanthocobitis maekhlongensis can be further differentiated from P. pictilis in not

having enlarged dorsal pterygiophores and neural spines (Fig.3-22), 7 (vs. 6) anal-fin rays, and a longer snout (9.8% vs 6.5% SL).

Distribution

Paracanthocobitis pictilis is known from the Ataran River drainage in eastern

Burma extending to Chedi Sam Ong, Three Pagoda Pass (Fig. 3-30).

Paracanthocobitis rubidipinnis (Blyth 1860)

Cobitis rubidipinnis Blyth, E., 1860:170, Report on some fishes received chiefly from the

Sitang River and its tributary streams, Tenasserim Provinces. Journal of the Asiatic

Society of Bengal v. 29 (2):138-174. Neotype: NRM 13743 (see Kottelat 1990:35).

Cobitis semizonata Blyth, E., 1860, same reference as for Cobitis rubidipinnis. Neotype:

NRM 13743 (see Kottelat 1990:35).

Nemacheilus rubidipinnis.—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. i-x + 1-259.

Acanthocobitis rubidipinnis.—Kottelat, M. 1990, Indochinese nemacheilines. A revision of nemacheiline loaches (Pisces: Cypriniformes) of Thailand, Burma, Laos, Cambodia and southern Viet Nam. Verlag Dr. Friedrich Pfeil, München. 1-262. Neotype: NRM

13743.

Acanthocobitis (Paracanthocobitis) rubidipinnis.—Grant, S., 2007, A new subgenus of

Acanthocobitis Peters, 1861 (Teleostei: Nemacheilidae). Ichthyofile No. 2: 1-9.

Diagnosis

(Fig. 3-31, Table 3-10)

Paracanthocobitis rubidipinnis is distinguished from congeners by having a

combination of 14-15 branched dorsal-fin rays, complete lateral line, suborbital flap in

male, and round dark spot on caudal peduncle.

Description

Body slender; branched dorsal-fin rays 14 -15; branched pectoral-fin rays 11-12;

branched pelvic-fin rays 8, branched anal-fin rays 6-7 (usually 7); branched caudal-fin

rays 9+9; 4-6 transverse rows of spots in dorsal fin; transverse bands in caudal-fin 6-8;

male with suborbital flap; axillary pelvic lobe present; lateral line complete with 90-110

pores. Maximum SL=71.2 mm female.

Color pattern consisting of 13-15 well-defined dorsal saddles extending onto side

as dar bars almost to lateral line. Squarish dark spots just below lateral line narrower

than interspaces; some connected to dark bars. Fins with red tint in life.

Comparison

Paracanthocobitis rubidipinnis differs from all species of Paracanthocobitis,

except P. pictilis and P. maekhlongensis, in having 14-15 (usually 15) branched dorsal-

fin rays vs. 10-13 in all other species. Paracanthocobitis rubidipinnis differs from P.

pictilis and P. maekhlongensis by the presence of a terminally unattached suborbital

flap in the male (vs. suborbital groove). In addition, P. pictilis has a sliver-shaped ocellus

on the caudal-fin base; the caudal spot is round in P. rubidipinnis.

Distribution

Paracanthocobitis rubidipinnis is known from the Irrawaddy and Ataran River

basins in Burma (Fig. 3-32).

Paracanthocobitis urophthalma (Günther, 1868)

Nemachilus urophthalmus Günther, 1868, Catalogue of the fishes in the British

Museum. Catalogue of the Physostomi, containing the families Heteropygii, Cyprinidae,

Gonorhynchidae, Hyodontidae, Osteoglossidae, Clupeidae thru Halosauridae, in the

collection of the British Museum. v. 7: i-xx + 1-512. Syntypes: BMNH 1864.7.11.35-36

Noemacheilus urophthalmus.—Talwar & Jhingran, 1991, Inland fishes of India and

adjacent countries. In 2 vols. Oxford & IBH Publishing Co., New Delhi, Bombay,

Calcutta. v. 1-2: i-xvii + 36 unnumbered + 1-1158, 1 pl, 1 map. [V. 1, i-liv + 1-541, 1

map; v. 2, i-xxii + 543-1158, 1 pl. Also reprinted, Balkema, Rotterdam, 1992.]

Acanthocobitis urophthalmus.—Pethiyagoda, 1991, Freshwater fishes of Sri Lanka. The

Wildlife Heritage Trust of Sri Lanka, Colombo. i-xiii + 1-362.

Acanthocobitis (Paracanthocobitis) urophthalmus Grant, 2007, A new subgenus of

Acanthocobitis Peters, 1861 (Teleostei: Nemacheilidae). Ichthyofile No. 2: 1-9.

Diagnosis

(Fig. 3-33, Table 3-11)

Paracanthocobitis urophthalma can be distinguished from all other species of

Paracanthocobitis in having dark bars on side of body that extend from dorsum to

venter, bars wider than interspaces; axillary pelvic lobe absent.

Description

Body deeper before dorsal fin, narrowing to caudal peduncle; branched dorsal-fin

rays 12-13; branched pectoral-fin rays10-12; branched pelvic-fin rays 8, branched anal-

fin rays 6; branched caudal-fin rays 9+9; transverse rows of spots in dorsal fin 4-6;

transverse bands in caudal fin 5-7; males with suborbital flap; axillary pelvic lobe

absent; lateral line complete or reaching at least to anal-fin origin with 59-71 pores.

Maximum SL= 48.8 mm male.

Comparison

Paracanthocobitis urophthalma is differentiated from all other species of

Paracanthocobitis in having dark bars wider than interspaces that extend from dorsum

to venter.

Distribution

Paracanthocobitis urophthalma is endemic to the Kalu Ganga basin on the island

of Sri Lanka (Fig. 3-34).

Paracanthocobitis zonalternans (Blyth 1860)

Cobitis zonalternans Blyth, 1860, Report on some fishes received chiefly from the

Sitang River and its tributary streams, Tenasserim Provinces. Journal of the Asiatic

Society of Bengal v. 29 (no. 2):138-174. Neotype: ZSM 27468.

Nemacheilus zonalternans.—Menon, 1987, The fauna of India and the adjacent

countries. Pisces. Vol. IV. Teleostei - Cobitoidea. Part 1. Homalopteridae. Zoological

Survey of India. i-x + 1-259, Pls. 1-16.

Noemacheilus phuketensis Klausewitz, 1957, Neue Süßwasserfische aus Thailand.

Senckenbergiana Biologica v. 38 (nos 3/4):193-204, Pls. 17, 17a, 18. Holotype: SMF

3966.

Acanthocobitis zonalternans.—Kottelat, 1990, Indochinese nemacheilines. A revision of

nemacheiline loaches (Pisces: Cypriniformes) of Thailand, Burma, Laos, Cambodia and

southern Viet Nam. Verlag Dr. Friedrich Pfeil, München. 1-262.

Acanthocobitis (Paracanthocobitis) zonalternans Grant, 2007, A new subgenus of

Acanthocobitis Peters, 1861 (Teleostei: Nemacheilidae). Ichthyofile No. 2: 1-9.

Diagnosis

(Fig. 3-35, Table 3-12)

A species of Paracanthocobitis distinguished from congeners in having an incomplete lateral line ending near dorsal-fin origin, 9-11 branched dorsal-fin rays, 6 anal-fin rays, and 4-5 transverse bands on the caudal fin.

Description

Body depth nearly uniform with some individuals being deeper-bodied pre- dorsally than post-dorsally; branched dorsal-fin rays 10-11; branched pectoral-fin rays

12-13; branched pelvic-fin rays 8; branched anal-fin rays 6; branched caudal-fin rays

9+8 or 9+9 (usually 9+8); transverse bands in caudal fin 4-6; suborbital flap present in male; axillary pelvic lobe present; lateral line incomplete, reaching at most to dorsal-fin origin with fewer than 50 pores. Maximum SL= 39.7 mm female.

Color pattern variable, with dark spots along lateral line sometimes mediated by a lateral stripe covering spots; dorsal pattern ranging from large to small dorsal saddles.

Combinations of patterns the aforementioned patterns can be present amoung individuals within a population. Pigment on dorsal portion of head darker than on ventral portion.

Comparison

Paracanthocobitis zonalternans differs from all other Paracanthocobitis except P. mooreh and P. linypha in having an incomplete lateral line. Paracanthocobitis zonalternans differs from P. mooreh and P. linypha in having an axillary pelvic lobe and

10-11 (vs. 12) dorsal-fin rays.

Distribution

Paracanthocobitis zonalternans is known from several drainages in Bangladesh,

Burma, Thailand, and Peninsular Malaysia including the Brahmaputra, Meghna,

Irrawaddy, Salawin, and Mae Khlong. The known range extends from Peninsular

Malaysia north to northern Burma. It is not found in the Chao Phraya basin and the western extent of its range is in northwestern Bangladesh (Fig.3-36).

Remarks

Given the large geographic distribution, habitat variation, variability of color pattern across geographic ranges, and morphometric differences, it is likely that P. zonalternans is a species complex with several undescribed species.

Figure 3-1. Acanthocobitis longipinnis. ZMB, 4795, Holotype, ventral view showing mouth morphology. Photograph by Peter Bartsch.

Figure 3-2. Acanthocobitis longipinnis. Lateral view of head, showing suborbital flap, ZMB 4795, holotype. Photograph by Peter Bartsch.

Figure 3-3. Head shapes in Acanthocobitis and Paracanthocobitis. A) Acanthocobitis pavaonacea, live. B) Paracanthocobitis maekhlongensis, live.

Figure 3-4. Lateral views of Acanthocobitis type specimens. A) Acanthocobitis longipinnis ZMB 4795, holotype. B) radiograph of A. longipinnis, ZMB 4795, holotype. C) Cobitis pavonacea, SMF 68, lectotype, 95 mm SL female.

Figure 3-5. A) A. pavonacea, live (photo by Steven Grant). B) Cobitis pavonacea, SMF 9091, paralectotype, 121 mm SL female, preserved.

Figure 3-6. Distribution of Acanthocobitis pavonacea.

Figure 3-7. Paracanthocobitis maekhlongensis, UF 182864,58.9 mm SL. Ventral view of head, showing mouth morphology.

Figure 3-8. Paracanthocobitis mackenziei, KU 40459, 51.5 mm SL male, lateral view. A) patch of adipose tissue on anterior end of lateral line. B) Close up of adipose patch.

Figure 3-9. Suborbital process shapes in Paracanthocobitis. A) Suborbital flap on P. adelaideae, USNM 378387, 62.7mm SL male. B) Suborbital groove on P. mandalayensis, CAS 88890, 87.8 mm SL male.

Figure 3-10. Bones supporting the suborbital flap. LE (yellow) = lateral ethmoid; IO 1 ( blue) = infraorbital 1.

Figure 3-11. Paracanthocobitis abutwebi. UMMZ 208646, holotype, 46.1 mm SL male.

Figure 3-12. Distribution of Paracanthocobitis abutwebi.

Figure 3-13. Paracanthocobitis adelaideae. A) adult color pattern, USNM 378387, holotype, 50.6 mm SL male. B) juvenile color pattern, USNM 378393, paratype, 28.5 mm SL.

Figure 3-14. Distribution of Paracanthocobitis adelaideae.

Figure 3-15. Paracanthocobitis botia. A) BMNH 1889.2.1.1562-1571, neotype, 45.3 mm SL male. B) BMNH 1889.2.1.1562-1571, 57.7 mm SL female.

Figure 3-16. Distribution of Paracanthocobitis botia.

Figure 3-17. Paracanthocobitis linypha. A) USNM 378387, holotype, 34.9 mm SL male. B) CAS 88918, paratype, 42.9 mm SL female.

Figure 3-18. Distribution of Paracanthocobitis linypha.

Figure 3-19. A) Paracanthocobitis mackenziei, KU 29159, 67.1 mm SL male. B) Nemacheilus aureus, BMNH 1889.2.1.1590, lectotype, 31.9 mm SL female.

Figure 3-20. Distribution of Paracanthocobitis mackenziei.

Figure 3-21. Paracanthocobitis maekhlongensis. A) UF 182864, holotype 58.9 mm SL female, preserved. B) ex UF 182864, paratype 56.4 mm SL female, live.

Figure 3-22. Comparison of key morphological differences in Paracanthocobitis pictilis and P. maekhlongensis. A) P. pictilis showing enlarged pterygiophores and dorsal neural spines, UF 172927, 76.4 mm SL male. B) P. pictilis showing sliver- shaped caudal spot, UNMF-P 07667, 81.2 mm SL female. C) P.maekhlongensis showing normal pterygiophores and dorsal neural spines, UF 176571, 54.4 mm SL female. D) P. maekhlongensis showing circular caudal spot, UF 182864, holotype, 58.9 mm SL female.

Figure 3-23. Distribution of Paracanthocobitis maekhlongensis.

Figure 3-24. Paracanthocobitis mandalayensis. A) Specimen from Irrawaddy River basin, USNM 344646, 81.1 mm SL female. B) Specimen from Wang River basin, UF 181110, 60.5 mm SL male.

Figure 3-25. Paracanthocobitis mandalayensis. Juvenile color pattern, UF 1811110, 24.5 mm SL.

Figure 3-26. Distribution of Paracanthocobitis mandalayensis

Figure 3-27. Paracanthocobitis mooreh. Lateral view, MCZ 52380, 44.4 mm SL male.

Figure 3-28. Distribution of Paracanthocobitis mooreh

Figure 3-29. Paracanthocobitis pictilis. A) UF 172927, 71.4 mm SL male. B) UNMF-P 07667, 79.5 mm SL male.

Figure 3-30. Distribution of Paracanthocobitis pictilis.

Figure 3-31. Paracanthocobitis rubidipinnis. Lateral view, ZRC 43570, 55.5 mm SL female.

Figure 3-32. Distribution of Paracanthocobitis rubidipinnis.

Figure 3-33. Paracanthocobitis urophthalma. Lateral view, BMNH 1864.7.11, syntype, 48.8 mm SL male.

Figure 3-34. Distribution of Paracanthocobitis urophthalma.

Figure 3-35. Paracanthocobitis zonalternans. A) ZSM 27468, neotype, 33.1 mm SL male. B) ZRC 41258, topotype, 36.1 mm SL male.

Figure 3-36. Distribution of Paracanthocobitis zonalternans.

Table 3-1. Morphometric data and meristic counts for Paracanthocobitis abutwebi (N=35). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 12-13 12 Branched pectoral-fin rays 10-13 12 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6-7 7 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 80-103 89.5±6.53

Morphometrics

Standard length 31.4-53.3

% of standard length Predorsal length 40.2-47.4 42.9±0.17 Head length 17.5-22.1 19.3±0.11 Snout length 5.5-10.0 8.1±0.01 Pre-pelvic length 45.5-50.9 48.1±0.013 Pre-anal length 69.1-80.7 76.1±0.021 Body depth (@D1) 12.6-18.0 15.4±0.012 Caudal-peduncle depth 9.8-13.2 11.1±0.007 Pectoral-fin length 17.1-24.4 19.9±0.018 Pelvic-fin length 13.6-19.9 17.0±0.013

% of head length Eye diameter 24.7-40.9 32.31±0.03 Interorbital width 20.1-35.4 27.4±0.034

Table 3-2. Morphometric data and meristic counts for Paracanthocobitis adelaideae (N=8). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 12 12 Branched pectoral-fin rays 10-12 11 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6 6 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 72-88 78.8±6.06

Morphometrics

Standard length 45.9-51.4

% of standard length Predorsal length 45.8-49.6 48.0±0.13 Head length 19.6-22.3 21.1±0.01 Snout length 9.1-10.8 9.6±0.01 Pre-pelvic length 52.3-54.9 53.5±0.01 Pre-anal length 77.8-80.8 79.2±0.01 Body depth (@D1) 18.4-20.7 19.8±0.01 Caudal-peduncle depth 11.3-13.4 12.7±0.008 Pectoral-fin length 19.8-23.5 22.0±0.012 Pelvic-fin length 16.8-19.2 17.9±0.01

% of head length Eye diameter 23.6-29.6 26.0±0.02 Interorbital width 28.4-36.5 33.3±0.03

Table 3-3. Morphometric data and meristic counts for Paracanthocobitis botia (N=10). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 11-12 12 Branched pectoral-fin rays 10-11 11 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6 6 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 83-100 92±5.36

Morphometrics

Standard length 37.1-57.7

% of standard length Predorsal length 45.7-50.7 48.1±0.17 Head length 18.7-21.0 19.8±0.01 Snout length 7.8-9.0 8.5±0.004 Pre-pelvic length 48.6-56.4 52.1±0.03 Pre-anal length 76.0-80.7 78.4±0.01 Body depth (@D1) 12.6-18.4 14.7±0.02 Caudal-peduncle depth 10.5-12.5 11.9±0.01 Pectoral-fin length 16.5-24.3 20.6±0.02 Pelvic-fin length 14.5-16.6 15.6±0.01

% of head length Eye diameter 28.1-35.3 31.1±0.02 Interorbital width 23.2-31.1 26.3±0.03

Table 3-4. Morphometric data and meristic counts for Paracanthocobitis linypha (N=10). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 11-12 12 Branched pectoral-fin rays 10-12 11 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6-7 6 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 20-42 27.5±6.17

Morphometrics

Standard length 26.1-42.9

% of standard length Predorsal length 40.4-50.7 46.5±0.03 Head length 18.8-22.9 20.6±0.01 Snout length 6.0-9.0 7.9±0.01 Pre-pelvic length 47.6-54.1 50.8±0.02 Pre-anal length 72.7-79.7 76.9±0.02 Body depth (@D1) 13.7-18.5 16.1±0.02 Caudal-peduncle depth 7.9-12.4 10.4±0.01 Pectoral-fin length 17.5-21.8 22.0±0.02 Pelvic-fin length 15.5-18.0 17.6±0.01

% of head length Eye diameter 24.0-30.9 31.4±0.04 Interorbital width 24.1-35.3 30.4±0.04

Table 3-5. Morphometric data and meristic counts for Paracanthocobitis mackenziei (N=46). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 12-13 12 Branched pectoral-fin rays 10-12 11 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6-7 6 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 73-101 85.1±7.99

Morphometrics

Standard length 28.8-72.7

% of standard length Predorsal length 40.2-49.1 45.3±0.02 Head length 18.2-23.3 20.7±0.01 Snout length 7.1-11.1 9.5±0.01 Pre-pelvic length 45.1-54.3 50.9±0.02 Pre-anal length 66.7-80.3 77.0±0.02 Body depth (@D1) 11.9-21.5 17.6±0.02 Caudal-peduncle depth 8.5-13.9 12.1±0.01 Pectoral-fin length 18.0-24.9 21.3±0.02 Pelvic-fin length 13.3-18.7 16.4±0.01

% of head length Eye diameter 21.4-31.8 27.8±0.03 Interorbital width 21.4-33.9 28.3±0.03

Table 3-6. Morphometric data and meristic counts for Paracanthocobitis maekhlongensis (N=12). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 13-14 14 Branched pectoral-fin rays 12-13 12 Branched pelvic-fin rays 8 8 Branched anal-fin rays 7 7 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 94-108 99.1±4.19

Morphometrics

Standard length 51.6-75

% of standard length Predorsal length 42.8-47.6 44.1±0.01 Head length 19.4-20.9 20.3±0.004 Snout length 9.0-10.5 9.8±0.004 Pre-pelvic length 49.1-53.2 50.8±0.01 Pre-anal length 74.3-80.6 77.4±0.02 Body depth (@D1) 14.6-18.4 17.3±0.01 Caudal-peduncle depth 11.3-13.3 12.4±0.01 Pectoral-fin length 16.9-20.7 18.8±0.01 Pelvic-fin length 15.5-16.9 16.3±0.003

% of head length Eye diameter 20.5-26.1 23.3±0.01 Interorbital width 24.5-32.9 30.0±0.02

Table 3-7. Morphometric data and meristic counts for Paracanthocobitis mandalayensis (N=23). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 11-12 12 Branched pectoral-fin rays 11-13 12 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6-7 6 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 80-103 87.2±7.37

Morphometrics

Standard length 41.8-88.4

% of standard length Predorsal length 43.3-48.3 45.2±0.01 Head length 18.7-24.7 20.7±0.01 Snout length 8.0-10.8 9.4±0.01 Pre-pelvic length 47.8-54.6 50.8±0.02 Pre-anal length 74.1-81.2 77.1±0.02 Body depth (@D1) 14.4-19.2 17.4±0.02 Caudal-peduncle depth 10.6-12.9 12.1±0.01 Pectoral-fin length 15.4-22.9 19.6±0.02 Pelvic-fin length 14.2-20.4 16.6±0.01

% of head length Eye diameter 18.4-30.8 23.2±0.03 Interorbital width 24.9-36.3 29.9±0.03

Table 3-8. Morphometric data and meristic counts for Paracanthocobitis mooreh (N=30). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 11-12 12 Branched pectoral-fin rays 11-12 11 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6-7 6 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 25-39 33.8±4.89

Morphometrics

Standard length 34.2-50.5

% of standard length Predorsal length 44.5-49.3 46.5±0.01 Head length 19.6-22.3 20.9±0.01 Snout length 7.8-10.1 9.0±0.01 Pre-pelvic length 52.0-54.3 53.1±0.01 Pre-anal length 76.4-82.5 79.1±0.02 Body depth (@D1) 15.2-21.4 19.2±0.02 Caudal-peduncle depth 10.9-13.5 12.7±0.01 Pectoral-fin length 17.8-21.6 20.0±0.01 Pelvic-fin length 14.9-17.9 16.1±0.01

% of head length Eye diameter 26.8-30.0 28.2±0.01 Interorbital width 18.2-30.0 25.2±0.04

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Table 3-9. Morphometric data and meristic counts for Paracanthocobitis pictilis (N=16). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 13-14 13 Branched pectoral-fin rays 11-13 13 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6 6 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 94-98 96.2±1.64

Morphometrics

Standard length 55.0-81.2

% of standard length Predorsal length 39.8-42.7 41.1±0.01 Head length 16.0-20.1 17.7±0.01 Snout length 5.3-10.1 7.4±0.02 Pre-pelvic length 35.5-50.8 47.6±0.01 Pre-anal length 75.1-78.7 77.3±0.02 Body depth (@D1) 9.1-16.7 13.2±0.02 Caudal-peduncle depth 7.1-13.2 9.6±0.01 Pectoral-fin length 12.7-19.4 16.3±0.02 Pelvic-fin length 9.5-17.8 12.9±0.01

% of head length Eye diameter 21.1-25.8 24.1±0.02 Interorbital width 23.9-33.1 27.5±0.03

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Table 3-10. Morphometric data and meristic counts for Paracanthocobitis rubidipinnis (N=11). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 14-15 15 Branched pectoral-fin rays 11-12 12 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6-7 7 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 90-110 96.8±6.38

Morphometrics

Standard length 46.4-71.2

% of standard length Predorsal length 39.1-46.6 42.6±0.02 Head length 16.4-21.8 20.3±0.01 Snout length 6.9-10.9 9.0±0.01 Pre-pelvic length 43.4-52.4 47.6±0.03 Pre-anal length 71.3-80.8 76.0±0.03 Body depth (@D1) 12.5-17.4 16.0±0.01 Caudal-peduncle depth 9.6-12.8 11.6±0.01 Pectoral-fin length 15.8-22.1 18.9±0.02 Pelvic-fin length 15.2-17.9 16.3±0.01

% of head length Eye diameter 18.1-30.6 23.7±0.03 Interorbital width 22.0-32.3 27.1±0.03

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Table 3-11. Morphometric data and meristic counts for Paracanthocobitis urophthalma (N=5). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 12-13 12 Branched pectoral-fin rays 10-11 11 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6 6 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 59-71 65.3±6.02

Morphometrics

Standard length 29.1-48.8

% of standard length Predorsal length 44.0-47.2 45.4±0.01 Head length 19.9-22.5 21.4±0.01 Snout length 9.0-11.1 10.0±0.01 Pre-pelvic length 50.5 -53.8 52.3±0.01 Pre-anal length 74.9-78.0 76.9±0.01 Body depth (@D1) 18.2-22.7 19.6±0.02 Caudal-peduncle depth 12.4-13.5 13.0±0.003 Pectoral-fin length 20.4-23.4 21.8±0.01 Pelvic-fin length 15.8-20.9 18.7±0.02

% of head length Eye diameter 28.2-35.0 30.7±0.02 Interorbital width 23.8-28.1 26.2±0.02

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Table 3-12. Morphometric data and meristic counts for Paracanthocobitis zonalternans (N=26). Character Range Mean±%SD

Meristics

Branched dorsal-fin rays 10-11 11 Branched pectoral-fin rays 10-13 11 Branched pelvic-fin rays 8 8 Branched anal-fin rays 6-7 6 Branched caudal-fin rays 9+9 9+9 Lateral-line pore count 40-49 44.5

Morphometrics

Standard length 26.7-39.7

% of standard length Predorsal length 39.0-50.0 46.4±0.02 Head length 18.9-25.0 21.7±0.01 Snout length 7.8-10.6 8.8±0.01 Pre-pelvic length 46.6-5 55.9 50.5±0.02 Pre-anal length 74.0-81.2 77.8±0.02 Body depth (@D1) 14.5-25.0 18.6±0.03 Caudal-peduncle depth 11.1-13.8 13.0±0.01 Pectoral-fin length 17.3-26.1 21.9±0.01 Pelvic-fin length 16.3-21.5 19.7±0.01

% of head length Eye diameter 26.0-41.0 30.9±0.03 Interorbital width 27.0-42.6 36.4±0.03

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CHAPTER 4 DISCUSSION

Acanthocobitis contains only A. pavonacea, a species restricted to the

Brahmaputra and Ganges river basins of northern India and Bangladesh. Although A. longipinnis is considered to be a synonym of A. pavonacea, the only available specimen is the poorly preserved holotype. It is possible that the collection of fresh material may allow the two forms to be diagnosed from one another.

Prior to this revision, seven species of Paracanthocobitis, P. botia, P. mandalayensis, P. mooreh, P. pictilis, P. rubidipinnis, P. urophthalma, and P. zonalternans, were recognized, although geographic distributions of most were uncertain. Paracanthocobitis mackenziei, considered a synonym of P. botia by several authors, including Menon (1987) and Kottelat (1990), was removed from synonymy, and three new species were described from material previously identified as P. botia: P abutwebi, P. adelaideae, and P. linypha. Populations recently described as P. pictilis

(Kottelat 2012) were found to be based on specimens of two species, with P. pictilis restricted to the Ataran River basin of Burma, and a new species, P. maekhlongensis, in the Mae Khlong basin of Thailand. With the completion of this revision, the number of recognized species within Paracanthocobitis is 12. A key for identification of species of

Paracanthocobitis appears below.

From the material examined, Paracanthocobitis can be divided into three morphological, and perhaps monophyletic, species groups: the P. botia group (with a free suborbital flap in the adult male), the P. zonalternans group (small, with 10 or fewer dorsal rays), and the P. mandalayensis group (with a suborbital groove in the adult male) group. Species within the P. botia group are: P. abutwebi, P. adelaideae, P.

105

linypha, P. mooreh, P. rubidipinnis, and P. urophthalma. This group occurs in India, Sri

Lanka, Bangladesh, and Burma. The P. mandalayensis group includes: P. maekhlongensis, P. mandalayensis, and P. pictilis and is restricted to Burma and western Thailand. Paracanthocobitis zonalternans is the only species within the P. zonalternans species group, but is highly variable and likely contains undescribed species. A phylogenetic analysis will be necessary to confirm the monophyly of these groups.

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APPENDIX KEY TO SPECIES OF PARACANTHOCOBITIS

• 1a. Lateral line incomplete, reaching at most to distal extremity of pelvic fin; free

suborbital flap in male……….……….…………………………..…....…...…...... 2

• 1b. Lateral line complete or at least reaching anal-fin origin; suborbital flap or

groove in male…..………………..………………………...…..……….………………4

• 2a. 10 or fewer branched dorsal-fin rays; lateral line ending at distal extremity of

dorsal fin...….………..……….…….……………… Paracanthocobitis zonalternans

• 2b. 11-13 branched dorsal-fin rays; lateral line ending past distal extremity of

dorsal fin...... 3

• 3a. 12-20 alternating long and short bars (much narrower than interspaces)

extending above and below lateral line, sometimes connected

to triangular dorsal saddles; 5-7 dark bands on caudal fin

…….………...…………………..……………………Paracanthocobitis linypha, n.sp.

• 3b. 12-14 small triangular dorsal saddles disconnected from spots, varying in

size, along the lateral line; 4-5 (usually 4) dark bands on caudal fin; ocellus

on caudal peduncle extends onto caudal fin; dark patch on opercle

…………………………….…..………………………...…..Paracanthocobitis mooreh

• 4a. Large, dark bands from dorsum to venter

…...………………………..……………………….….Paracanthocobitis urophthalma

• 4b. Blotches, spots, or speckles along lateral line; …………...….……...……...... 5

• 5a. Axillary pelvic lobe absent.……….……………………………..……………..…..6

• 5b. Axillary pelvic lobe present..………….………..………………………..…...... 7

• 6a. 10-13 small dorsal saddles; 10-16 larger spots alternating with smaller spots

along lateral line; 4-5 (usually 4) dark bars on caudal fin; 6-7 (usually

107

7) branched anal-fin rays ………………...…….Paracanthocobitis abutwebi, n.sp.

• 6b. 8-10 dark spots along side just below lateral line, narrowing ventrally without well-defined spots between; 5-7 (usually 5) dark bars on caudal fin; 6 branched anal-fin rays...... ….Paracanthocobitis botia

• 7a. Small, dark triangular dorsal saddles connecting ventrally to thick, wave-like

pattern formed by connected blotches along lateral line, forming more

of a checkerboard pattern in juvenile.

…………………………………………….…..….Paracanthocobitis adelaideae, n.sp.

• 7b. Color pattern on side not a wave-like pattern formed by connected, dark

blotches along lateral line ...………...……….……………..………….………..….....8

• 8a. Sliver-shaped caudal spot; adult male with suborbital groove

…………………………………….…..…………………….Paracanthocobitis pictilis

• 8b. Round caudal spot; adult male with suborbital flap or groove….………….…..9

• 9a. 12-13 branched dorsal-fin rays; 6-7 (usually 6) branched anal-fin rays; dorsal

saddles irregular, no central light area ……………………….……………….…....10

• 9b. 13-15 branched dorsal-fin rays; 6-7 (usually 7) branched anal-fin rays;

dorsal saddles uniform with a central light spot or line……………………….……11

• 10a. Suborbital flap in male; curved dorsal saddles connected to blotches along

lateral line, sometimes forming a prominent zigzag pattern on dorsal surface; 6-7

dark bands on caudal fin...... Paracanthocobitis mackenziei

• 10b. Suborbital groove in male; small dorsal saddles disconnected from

large spots along lateral line (sometimes connected in juvenile); 5-7 dark bands

on caudal fin..…………..……….…...……….…...Paracanthocobitis mandalayensis

• 11a. Suborbital groove in male; 13-14 (usually 13) branched dorsal-fin rays;

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dorsal saddles much wider dorsally than ventrally, dorsal portion of saddles

wider than interspaces; irregular pigment between saddles

.…………………………………...………..Paracanthocobitis maekhlongensis, n.sp.

• 11b. Suborbital flap in male; 14-15 (usually 15) branched dorsal-fin rays;

dorsal saddles more uniform in shape with interspaces similar in size to

saddles; no pigment between saddles……….……..Paracanthocobitis rubidipinnis

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LIST OF REFERENCES

Blyth, E. 1860 Report on some fishes received chiefly from the Sitang River and its tributary streams, Tenasserim Provinces. Journal of the Asiatic Society of Bengal v. 29 (no. 2): 138-174.

Cracraft, J. 1987. Species concepts and the ontology of evolution. Biology and philosophy (2.3):329-346.

Day, F. 1872 Monograph of Indian Cyprinidae. Parts 4-6. Journal of the Asiatic Society of Bengal v. 41 (pt 2, nos 1-4): 1-29, 171-198; 318-326.

Grant, S. 2007 (29 Oct.) A new subgenus of Acanthocobitis Peters, 1861 (Teleostei: Nemacheilidae). Ichthyofile No. 2: 1-9.

Günther, A. 1868 (14 Mar.) Catalogue of the fishes in the British Museum. Catalogue of the Physostomi, containing the families Heteropygii, Cyprinidae, Gonorhynchidae, Hyodontidae, Osteoglossidae, Clupeidae,... [thru]... Halosauridae, in the collection of the British Museum. v. 7: i-xx + 1-512.

Hamilton, F. 1822 An account of the fishes found in the river Ganges and its branches. Edinburgh & London. i-vii + 1-405.

Jacquemont, V. 1835-44 Voyage dans l'Inde pendant les années 1828 à 1832, publié sous les auspices de M. Guizot, Ministre de l'instruction publique. Journal. 4 vols. text, 2 vols. Atlas. Paris. (1835-1844).

Jayaram K.1999, The Freshwater fishes of the Indian region. Narendra Publishing House, Delhi, xxvii+1-551.

Klausewitz, W. 1957 (1 Apr.) Neue Süßwasserfische aus Thailand. Senckenbergiana Biologica v. 38 (nos 3/4): 193-204.

Kottelat, M. 2012 (29 May) Acanthocobitis pictilis, a new species of loach from Myanmar and Thailand (Teleostei: Nemacheilidae). Zootaxa No. 3327: 45-52.

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Kottelat, M. (1990) Indochinese nemacheilines: A revision of nemacheiline loaches (Pisces: Cypriniformes) of Thailand, Burma, Laos, Cambodia and southern Vietnam. Germany: Verlag Dr. Friedrich Pfeil.

Kottelat, M. (1989) Zoogeography of the fishes from Indochinese inland waters with an annotated check-list. Bulletin Zoölogisch Museum, Universiteit van Amsterdam, 12, 1-55.

McClelland, J. 1839 Indian Cyprinidae. Asiatic Researches v. 19 (pt 2): 217-471, Pls. 37-61.

Menon, A. G. K. 1987 (May) The fauna of India and the adjacent countries. Pisces. Vol. IV. Teleostei - Cobitoidea. Part 1. Homalopteridae. Zoological Survey of India. i-x + 1-259.

Peters, W. (C. H.) 1861 Über zwei neue Gattungen von Fischen aus dem Ganges. Monatsberichte der Königlichen Preuss[ischen] Akademie der Wissenschaften zu Berlin 1861: 712- 713.

Pethiyagoda, R. 1991 Freshwater fishes of Sri Lanka. The Wildlife Heritage Trust of Sri Lanka, Colombo. i-xiii + 1-362.

Potthoff, T. 1984. Clearing and staining techniques. In: Moser, H.G. (Ed.) Ontogeny and Systematics of Fishes. Special publication American Society of Ichthyologists and Herpetologists, vol. 1. Allen Press, Lawrence, KS, USA, pp. 35-37.

Rainboth, W. J., Vidthayanon, C., & Dinh Yen, M. (2012).Fishes of the greater mekong ecosystem with species list and photographic atlas. (1st ed.). Ann Arbor, Michigan: Museum of Zoology, University of Michigan. p. 68.

Rendahl, H. 1948 Die Süßwasserfische Birmas. I. Die Familie Cobitidae. Arkiv för Zoologi v. 40 A (no. 7): 1-116.

Swainson, W. 1839 On the natural history and classification of fishes, amphibians, & reptiles, or monocardian animals. Spottiswoode & Co., London. v. 2: i-vi + 1-448.

Sykes, W. H. 1839 On the fishes of the Deccan. Proceedings of the Zoological Society of London 1838 (pt 6): 157-165.

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Talwar, P. K. and A. G. Jhingran 1991 Inland fishes of India and adjacent countries. In 2 vols. Oxford & IBH Publishing Co., New Delhi, Bombay, Calcutta. v. 1-2: i-xvii + 36 unnumbered + 1-1158, 1 pl, 1 map. [V. 1, i-liv + 1-541, 1 map; v. 2, i-xxii + 543-1158.

Taylor, W.R., van Dyke, G.C. 1985. Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium 9: 107-119.

Whitehead, P.J.P. and P.K. Talwar 1976. Francis Day (1829-1889) and his collections of Indian fishes. Bulletin of the British Museum (Natural History) Historical Series v. 5 (no.1):1-189.

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BIOGRAPHICAL SKETCH

Randal A. Singer was born in Ft. Myers, Florida. The first of two children born to his parents, Randy enjoyed frequent fishing trips with his father and summers on the

Mississippi gulf coast with his uncle that began his lifelong interest in fishes. Randy graduated from the University of Georgia in 2008 with a B.S. in ecology as a part of the inaugural class from the first stand-alone school of Ecology in the world. Upon graduating, Randy began working as a technician for the Georgia Museum of Natural

History Ichthyology Collection and the Jacqueline Mohan lab, focusing on the effects of climate change on fine root growth in trees. During this time, Randy’s eyes were opened to the wonders of the world of deep-sea fishes when he participated in the 2010 NOAA

MAR-ECO cruise to the mid-Atlantic Ridge. Following the completion of his positions at

UGA, Randy moved to Gainesville, Florida, to work for the Florida Museum of Natural

History Ichthyology Collection. On May 21st, 2011 Randy married Corie Ritchie in the mountains of Cleveland Georgia. In the same year, Randy began work on his master’s degree in zoology at the University of Florida focusing on fish taxonomy, during which time he frequently conducted fieldwork in Southeast Asia relevant to this thesis. On

October 9th 2012, Randy’s daughter Adelaide C. Singer was born. With the love and support of his family, Randy went on to finish his master’s degree in the summer of

2013. Going forward, Randy hopes to continue work with deep-sea fishes, hopefully in close proximity to a museum of natural history, enjoy time with his family, and explore and describe the wonderful diversity of the world’s fishes.

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