MORPHOLOGICAL DIAGNOSES OF GARRA (CYPRINIFORMES: CYPRINIDAE) FROM NORTH-EASTERN INDIA WITH FOUR NEW SPECIES DESCRIPTION FROM BRAHMAPUTRA RIVER

ARUNACHALAM M.1, NANDAGOPAL S.2, MAYDEN R.L.3*
1Sri Paramakalyani Centre for Environmental Sciences, Manonmaniam Sundaranar University, Alwarkurichi- 627 412, TN, India.
2Sri Paramakalyani Centre for Environmental Sciences, Manonmaniam Sundaranar University, Alwarkurichi- 627 412, TN, India.
3Department of Biology, Saint Louis University, Saint Louis, Missouri- 63103, USA.
* Corresponding Author : cypriniformes@gmail.com

Received : 17-08-2013     Accepted : 31-10-2013     Published : 03-12-2013
Volume : 4     Issue : 3       Pages : 121 - 138
J Fish Aquaculture 4.3 (2013):121-138
DOI : http://dx.doi.org/10.9735/0976-9927.4.3.121-138

Cite - MLA : ARUNACHALAM M., et al "MORPHOLOGICAL DIAGNOSES OF GARRA (CYPRINIFORMES: CYPRINIDAE) FROM NORTH-EASTERN INDIA WITH FOUR NEW SPECIES DESCRIPTION FROM BRAHMAPUTRA RIVER." Journal of Fisheries and Aquaculture 4.3 (2013):121-138. http://dx.doi.org/10.9735/0976-9927.4.3.121-138

Cite - APA : ARUNACHALAM M., NANDAGOPAL S., MAYDEN R.L. (2013). MORPHOLOGICAL DIAGNOSES OF GARRA (CYPRINIFORMES: CYPRINIDAE) FROM NORTH-EASTERN INDIA WITH FOUR NEW SPECIES DESCRIPTION FROM BRAHMAPUTRA RIVER. Journal of Fisheries and Aquaculture, 4 (3), 121-138. http://dx.doi.org/10.9735/0976-9927.4.3.121-138

Cite - Chicago : ARUNACHALAM M., NANDAGOPAL S., and MAYDEN R.L. "MORPHOLOGICAL DIAGNOSES OF GARRA (CYPRINIFORMES: CYPRINIDAE) FROM NORTH-EASTERN INDIA WITH FOUR NEW SPECIES DESCRIPTION FROM BRAHMAPUTRA RIVER." Journal of Fisheries and Aquaculture 4, no. 3 (2013):121-138. http://dx.doi.org/10.9735/0976-9927.4.3.121-138

Copyright : © 2013, ARUNACHALAM M., et al, Published by Bioinfo Publications. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution and reproduction in any medium, provided the original author and source are credited.

Abstract

Garra minimus is described from the Ranga River of upper Bramaputra River Drainage, Garra kimini is described from the tributary of Ranga river, Garra nigricauda is described from Pasighat and Garra alticaputus is described from Dikrong river at Boorum village from the Indian State of Arunachal Pradesh. Garra minimus is unique in its size in comparisons with all other species of Garra reported/published from North East India and also with more number of branched pectoral-fin rays. Garra alticaputus is diagnosed in having a deeper head and lacking any coloration on fins and body relative to other species from these drainages and the region that have distinctive colour patterns either on the body or in their the fins. Garra nigricauda is diagnosed from other species of Garra from this region with its distinctive robust body size and by the forked caudal-fin tipped black and the upper lobe of caudal-fin being pointed and shorter than the lower lobe. Garra kimini is diagnosed in moderately large headed, height less than its length; width almost equal to height with more pectoral-fin rays like the other three species described herein.

Keywords

Garra minimus, Garra alticaputus, Garra nigricauda, Garra kimini

Introduction

Since first described by Hamilton [1] in his book on the fishes of the Ganges River, the genus Garra (type species Cyprinus lamta (= Garra lamta) has been, and continues to be, one of the most enigmatic and taxonomically difficult cyprinid genera of the Asian region. As traditionally recognized the species occur from Borneo to West Africa through China, South and Southeast Asia, Middle East, Arabian Peninsula and East Africa. The genus, however, is an extremely difficult group for biodiversity studies and warrants extensive studies as to the morphology, taxonomy, and phylogenetics. Two of the most significant factors underlying difficulties with the group include a combination of its diversity and its taxonomic instability, both serving as major impediments for researchers. Similar complications exist even for the subfamily to which this genus is allocated, Labeonini, through the constant flux it is composition and internal taxonomy, two recalcitrant qualities, that are only to be compounded by current phylogenetic evidence that the genus is not monophyletic [2,3] .
In additional to the underlying issues outlined above, multiple, even more basic hindrances contribute to difficulties and/or complications for anyone working with or hoping to work on “Garra” [3] or Garra as traditionally recognized. These include the fact the traditionally recognized Garra is very widespread, has a great diversity of species (over 100) and many undescribed species, confusing morphological variation often seen in species without thorough evaluation across the genus, notably complex and frequently confusing literature on its diversity and taxonomy, and many issues surrounding types specimens for species and type species for genera in Cyprinidae. Some examples include the descriptions of or placement of species in different genera, including Discognathus Heckel [4] , Chondrostoma and Gobio [5] , Gonorhynchus and Platycara [6-8] and Gonorhynchus [9] . Day [10] subsequently allocated all of these species to Garra.
To date, 20 species of Garra have been reported to occur cross north and northeastern India. These include the type species for the genus, G. lamta [1] and the following 19 species: G. gotyla [12] , G. nasuta [7] , G. rupecula [8] , G. lissorhynchus [9] , G. kempi [13] , G. annandalei [13] , G. naganensis [13] , G. abhoyai [13] , G. manipurensis [14] , G. litanensis [15] , G. compressus [16] , G. elongata [17] , G. nambulica [18] , G. paralissorhynchus [19] , G. arupi [20] , G. kalpangi [21] , G. namyaensis [22] , G. dampaensis [23] , G. magnidiscus [24] .
Species so far reported from the Indian state of Arunachal Pradesh include G. annandalei, G. gotyla gotyla, G. kempi, G. lissorhynchus, G. mcclellandi, G. lamta and G. naganensis [25] . Recently, an additional species, Garra arupi was described from upper Brahmaputra basin in Arunachal Pradesh [20] and also Garra magnidiscus was described from the same basin [24] and is also a member of the “Garra fauna” of this region. In recent inventory surveys of fishes from this region we collected and purchased fishes from local villagers from Betapul colony near the Ranga River. Some of the Garra specimens obtained in these inventory efforts are morphologically distinctive and diagnosable, and are herein are described as new to science.

Materials and Methods

Abbreviations used herein include SL = standard length, HL = head length, ZSI/SRS = Zoological Survey of India/Southern Regional Station, F = Fish, FF = Freshwater Fish, MSUMNH = Manonmaniam Sundaranar University Museum of Natural History, (MSUMNH) in Alwarkurichi, India, MUMF = Manipur University Museum of Fishes and CMA = Collection of M. Arunachalam. Methods for measurements and counts follow those of Hubbs and Lagler [26] . Measurements were taken to the nearest 0.1 mm using digital calibers. Body measurements are expressed as percentage of Standard length (%SL); head measurements are expressed as percentage of head length (%HL). Distance between pectoral-fin origin and vent and distance between pelvic-fin origin and vent are also taken for the present study [Fig-1] .
Methods for PCA was employed using the sheared data of the morphometric measurements of the four species and types of other species were not included in this analysis because of such limited materials (N = 1 or 2 specimens).
Counts and morphometric characters such as disc length, disc width, central-pad length, central-pad width, post-dorsal length and body depth follow methods of Kullander and Fang [27] . As noted by Kullander and Fang [27] lip structures serve as important characters for the diagnosis and identification of most species of Garra and are also important for resolving their phylogenetic relationships. It is unfortunate that most descriptions of Garra, even recent, lack details of lip structure and are thus not useful for comparisons without having specimens, many of which are either difficult to obtain via loans due to institutional policies or the number of specimens are few.
Garra minimus, Garra alticaputus, Garra nigricauda and Garra kimini were compared with the related species Garra elongata (ZSI FF 4157- Paratype); Garra gotyla gotyla (ZSI F 198 / 2); Garra naganensis, (ZSI F 9970 / 1); Garra lissorhynchus (ZSI F 8098 / 1 - Holotype); Garra nambulica (ZSI 4139 - Paratype); Garra annandalei (ZSI F 6082 / 2 - Holotype); Garra paralissorhynchus (ZSI FF 4158 - Paratype); Garra kempi (ZSI FF 7716 / 1); Garra abhoyai (F 5307 / 1) from the Zoological Survey of India, Calcutta [Fig-2] . Specimens of G. compressus G. litanensis G. namyaensis G. manipurensis were examined from Manipur University Museum of Fishes, Manipur, India (MUMF) [Fig-3] . Holotypes are deposited in the Zoological Survey of India/Southern Regional Station (ZSI/SRS) at Chennai; paratypes are deposited at the Manonmaniam Sundaranar University Museum of Natural History, (MSUMNH) in Alwarkurichi, India, and in first author’s personal collections (Collection of M. Arunachalam, CMA).

New Species Descriptions

Garra minimus sp. nov. [Fig-4]
Holotype: ZSI/SRS F8577, 1 ex. Male. 52.9 mm SL, tributary of Ranga River, Lower Subanshri District, Arunachal Pradesh, India (N 270 21’ 359’’ E 93­­0 42’ 214’’), Collectors: M. Arunachalam, M. Raja, C. Vijayakumar and S. Nandagopal. 17 Jun 2011.
Paratypes: MSUMNH 59, 5 ex. 39.8-52.9 mm SL CMA 17, 14 ex, 35.6-53.1 mm SL. All other details same as Holotype.

Diagnosis

Garra minimus sp. nov. can be differentiated from other species of Garra by multiple homologous morphometric characters, the latter being relative sizes of homologous structures and distances between homologous landmarks. [Table-1] provides comparative meristic characteristics of type species of Garra type specimens from ZSI and MUMF.
Garra minimus is a very small species of Garra with smoothly rounded snout tip, a very distinct rostral lobe and without a proboscis. Barbels in two pairs, pre dorsal and venter scales present, breast scales absent, lateral-line scale rows 35-37, circumpeduncular scale rows 12, circumferential scale rows 18-20.
G. minimus is distinguished from G. lissorhynchus by fewer pre dorsal-fin scale rows (10-11 vs. 13), fewer circumpeduncular scale rows (12 vs. 16), greater anal-fin scale rows (4-6 vs. 3.5) and absence of W-shaped band on caudal-fin (vs. presence). It is distinguished from G. nambulica by fewer pre dorsal-fin scale rows (10-11 vs. 24), fewer circumpeduncular scale rows (12 vs. 16) and absence of W-shaped band on caudal-fin (vs. presence). The species is distinguished from G. annandalei by more branched pectoral-fin rays (14-16 vs. 13), more pre dorsal-fin scale rows (10-11 vs. 9); more lateral-line scale rows (35-37 vs. 34) and fewer circumpeduncular scale rows (12 vs. 16). Garra minimus differs from G. gotyla gotyla in having more lateral-line scale rows (35-37 vs. 33), fewer circumpeduncular scale rows (12 vs. 16), more anal scale rows (4-6 vs. 3), absence of scales on breast (vs. present) and snout without a proboscis (weakly developed vs. well developed median proboscis). It is distinguished from G. elongata in having more branched pectoral-fin rays (14-16 vs. 12) and fewer lateral-line scale rows (35-37 vs. 39) and from G. naganensis in having more branched pectoral-fin rays (14-16 vs. 13), fewer lateral-line scale rows (35-37 vs. 39) and fewer circumpeduncular scale rows (12 vs. 16). From G. paralissorhynchus it is distinguished by more pectoral-fin rays (14-16 vs. 11) and lateral-line scale rows (35-37 vs. 31) and fewer pre dorsal-fin scale rows (10-11 vs. 12) and circumpeduncular scale rows (12 vs. 16). It is distinct from G. kempi in having more pectoral-fin rays (14-16 vs. 12), fewer lateral-line scale rows (35-37 vs. 40) and fewer pre dorsal-fin scale rows (10-11 vs. 12). From G. arupi it is distinguished by more branched pectoral-fin rays (14-16 vs. 10-11) and fewer circumpeduncular scale rows (12 vs. 16). It is distinguished from G. gravelyi by more lateral-line scale rows (35-37 vs. 32-34); from Garra nasuta (McClelland) by more lateral-line scale rows (35-37 vs. 34).
It is diagnosed from G. nigricauda sp. nov. by size, scales on breast (naked vs. well developed) and fewer circumferential scale rows (18-20 vs. 22-24). From G. kimini sp. nov. it is diagnosed in having more lateral-line scale rows (35-37 vs. 33-34) and fewer circumpeduncular scales (12 vs. 16). It is distinguished from G. kalpangi in having more branched pectoral-fin rays (14-16 vs. 10-12), more lateral-line scale rows (35-37 vs. 32-33), fewer circumpeduncular scale rows (12 vs. 16) and more anal-fin scale rows (4-6 vs. 3). It is diagnosed from G. rupecula in having more branched dorsal-fin rays (8 vs. 6-7), more lateral-line scale rows (35-37 vs. 32-34) and fewer upper transverse scale rows (3.5 vs. 4.5-6.5). It is diagnosable from G. manipurensis in having more pectoral-fin rays (14-16 vs. 12), lateral-line scale rows (35-37 vs. 34), and pre anal-fin scale rows (14-18 vs. 12), as well as fewer transverse scale rows (3.5/2.5-3.0/2.5-3.0 vs. 4.5/3.5/4.5), circumpeduncular scale rows (12 vs. 16), and circumferential scales (18-20 vs. 24). From G. compressus it is distinguished in having more pectoral-fin rays (14-16 vs. 12) and fewer lateral-line scale rows (35-37 vs. 39); from G. litanensis in having more pectoral-fin rays (14-16 vs. 13) and lateral-line scale rows (35-37 vs. 32); from G. namyaensis in having more pectoral-fin rays (14-16 vs. 12), lateral-line scale rows (35-37 vs. 31) and fewer pre dorsal-fin scale rows (10-11 vs. 13) and circumpeduncular scale rows (12 vs. 16). It is differentiated from G. abhoyai in having more pectoral-fin rays (14-16 vs. 11), lateral-line scale rows (35-37 vs. 33) and fewer pre dorsal-fin scale rows (10-11 vs. 17) and circumpeduncular scale rows (12 vs. 16). It is diagnosable from G. magnidiscus in having more branched pectoral-fin rays (14-16 vs. 10-12), less pre dorsal-fin scale rows (10-11 vs. 12-15) and less lateral-line scales (35-37 vs. 40-42), as well as fewer transverse scale rows (3.5/2.5-3/2.5-3 vs. 4-4.4/3-3.5/3-3.5).

Description

Body elongate, predorsum ascending very slightly with body depth and almost equal in dorsal and ventral profiles [Table-2] .
Snout rounded with transverse grooves. Orbit is nearer to preopercle than to tip of snout. Rostral lobe with many small erect tubercles; smaller individuals with only 5 tubercles. Snout with minute, scattered and erect tubercles. Two pairs of barbels present; rostral barbels reaching middle of rostral cap and placed inside a groove not connected with rostral caps. Central pad wider than long. Upper lip with a median notch and papilliferous along upper jaw. Lower labial fold covering only 40% of central pad.
Dorsal-fin origin 2 scale rows anterior to pelvic-fin origin; first and second branched rays longest; posterior margin of fin concave. Pectoral-fin rounded; when depressed not reaching base of pelvic-fin. Pelvic-fin with pointed tip; when depressed not reaching to origin of anal-fin. Anal-fin short; when depressed almost reaching base of caudal-fin. Upper lobe of caudal-fin slightly longer than the lower lobe.
Dorsal-fin rays ii, 8 (last ray branched at base); pectoral-fin rays i, 14-16; pelvic-fin rays i, 8-9; anal-fin rays ii, 5; caudal-fin rays 10+9 (17 branched); lateral-line complete, scale rows 35-37; dorsal-fin origin and lateral-line separate by 3.5 scale rows; lateral transverse scales between lateral- line and pelvic-fin origin 2.5-3.0 scale rows; and between lateral-line and anal-fin origin by 2.5-3.0 scale rows; pre dorsal-fin scale rows 10-11; circumpeduncular scale rows 12; circumferential scales 18-20; pre anal-fin scale rows 14-18 [Table-3] .

Scales

Breast area with V-shaped naked pattern; venter with scales up to pectoral-fin insertion.

Color in Live

Dorsal, lateral, and ventro-lateral scale rows, to one scale row above venter, with base color of olive brown; scales above lateral-line with dark dorsal and ventral blotches that confluence to form distinct dark lateral stripes from occiput where distinctly separated to anterior caudal peduncle and then converging near posterior extend of pigmented caudal peduncle; center of dorsum with scale dark and with narrow dark margins. On caudal peduncle, area around hypural plate and slightly posterior on bases of caudal-fin rays, dorsally and ventrally to anterior extent of procurrent rays cream-colored and without melanophores; base of caudal-fin with distinct black circular spot separate from body but continuous with black stripe present on median rays and membranes of caudal-fin.
Head dark olive brown dorsally and laterally. Powder blue spot present where opercle joins body dorsally; blue coloration encircling eye and on lower-anterior cheek, ventro-lateral edges of rostral barbels, and corner of mouth, notably contrasting with surrounding areas. No light region posterior to orbit or dark spots on cheek or upper opercle. Lower cheek and branchiostegal rays and membranes silver; above which tan. Snout light tan. Nares and barbels red. Ventrally, head pinkish to cream anteriorly; lips, barbels, and papillae similar in color.
Breast, belly, and posteror to anal-fin origin silver-white, immaculate. Ventrolaterally light green coloration extending at least two scale dorsally. Well-developed, distinct black spot present just behind operculum, on the first lateral-line scale.
Distally, all branched rays of dorsal-fin base dusky; short unbranched rays dark, creating darkened anterior to dorsal-fin; leading ray unpigmented; membranes of branched rays dusky to black medially and distally but reddish-yellow to orange all along base of fin; distal margin of fin clear anteriorly, becoming reddish-pink posteriorly. Dorsally, pectoral-fin pinkish-red and dusky, and with some darkened areas. Leading unbranched ray darkened medially and distally; clear basally. Membrane between leading ray and first branched ray dusky; membranes for rays 2-6 dusky. Branched rays 2-8 pinkish red, especially distally where most concentrated; basally, fin lighter in color but with similar coloration; interior membranes within a branched ray reddish. Narrow distal edge of fin powder blue. Pelvic-fin as in pectoral-fin except leading ray not dark and reddish coloration more intense. Caudal-fin with similar reddish coloration throughout, lightening posteriorly, except for extreme dorsal and ventral lobes; principal and branched rays lined with melanophores; median branched rays more darkly pigmented, creating a broad median stripe; posterior 1/3 of lower lobe of caudal-fin darkly pigmented; dark stripe on distal edge of upper and lower lobes. Ventrally, pectoral and pelvic-fins are as above black median rays and membranes, creating stipe from caudal spot, and narrow black lines along.

Color in Preservative

Dorsum and head dark. Dorsal-fin base with black spots; no spots or colouration in pelvic, pectoral or anal-fins.

Distribution and Habitat

Garra minimus inhabits an unregulated region of the Ranga River, 20 km downstream of a Hydro-electric project in the Lower Subanshri District of Arunachal Pradesh [Fig-5] . Sampling was conducted in the river at a local settlement area called Betapul Colony. At the sampling site, the right bank (looking upstream) was more stable and with rooted vegetation; the left bank was relatively unstable. The stream had high and fast flow during the sampling period due to heavy rain the previous day. Water was turbid and depth ranged from 0.5 m to 0.6 m with the deeper portions towards the right bank. Substrate types included bedrock, large boulders, small boulders, cobbles, gravel and silt. Water in this area was relatively undisturbed except for bathing and cloth washing by people of the colony. Two women from the colony used hook and line fishing and caught larger fish specimens and species, including species of Tor and Neolissochilus; four of the torine specimens were purchased from them. The women also fished using a round steel dining plate. Wheat flower paste with turmeric was made into small bead-sized balls and were placed on the plate and covered by a white cloth with small holes for catching loaches. Other species found at this locality included Devario aequipinnatus, Schistura sp., Barilius sp., Schizothorax sp., Neolissochilus hexagonolepis, Tor putitora, and Aborichythys sp.

Etymology

The name minimus is Latin, referring to the small body size of the species as compared to the other species reported from this region.
Garra alticaputus sp. nov. [Fig-6]
Holotype: ZSI/SRS F8578, 1 ex. Male. 71.6 mm SL, Dikrong River at Boorum Village (tributary of Ranga River), Lower Subanshri District, Arunachal Pradesh, India (N 270 09’ 026’’ E 930 42’ 227’’). Collectors: M. Arunachalam, M. Raja, C. Vijayakumar and S. Nandagopal. 16 Jun 2011.
Paratypes: MSUMNH 60, 3 ex. 57.6-61.6 mm SL; CMA 18, 4 ex, 57.6-71.6 mm SL. All other details same as Holotype.

Diagnosis

Garra alticaputus can be differentiated from other species of Garra by multiple homologous meristic characters, relative sizes of homologous structures and distances between homologous landmarks.
Garra alticaputus be diagnosed from G. lissorhynchus by the absence of a W-shaped dark band at the posterior end of caudal-fin, fewer lateral-line scale rows (33 vs. 34) and fewer pre dorsal-fin scale rows (10 vs. 13). From G. nambulica it is distinguished in having more branched pectoral-fin rays (13-15 vs. 12) and fewer pre dorsal-fin scale rows (10 vs. 24); from G. annandalei in having more pectoral-fin rays (13-15 vs. 13) and pre anal-fin scale rows (16-22 vs. 18); from G. gotyla gotyla in having subcutaneous scales on breast (vs. well developed) and proboscis on snout (not well developed vs. well developed median proboscis). The species is distinguished from G. elongata in lacking dark grey longitudinal stripe from gill-opening to caudal-fin base, transverse black bar near the free margin of dorsal-fin, black longitudinal mark on medium rays of caudal-fin and absence of colours in other fins, fewer lateral-line (33 vs. 39), pre dorsal-fin scale rows (10 vs. 13) and anal-fin scale rows (3-3.5 vs. 5.5). It is distinguished from G. naganensis in having fewer lateral-line (33 vs. 39), pre dorsal-fin scale rows (10 vs. 12) and anal-fin scale rows (3-3.5 vs. 6) and a greater number of pectoral-fin rays (13-15 vs. 13). From G. paralissorhynchus it differs in having more pectoral-fin rays (13-15 vs. 11) and lateral-line scale rows (33 vs. 31) and fewer pre dorsal-fin scale rows (10 vs. 12); from G. kempi in having more pectoral-fin rays (13-15 vs. 12) and circumpeduncular scale rows (16 vs. 12) and fewer lateral-line (33 vs. 40) and pre dorsal-fin scale rows (10 vs. 12).
The new species differs from G. kalpangi in having more pectoral-fin branched rays (13-15 vs. 10-12); from G. rupecula in having more branched dorsal-fin branched rays (8 vs. 6-7) and fewer upper transverse rows (4.5 vs. 4.5 to 6.5). In ventral side of G. rupecula, breast, belly and post pelvic-fin regions naked while in G. alticaputus well developed scales on belly region and subcutaneous scales on breast region. The species is diagnosed from G. manipurensis in having more pectoral-fin rays (13-15 vs. 12) and anal-fin scale rows (16-22 vs. 12), fewer lateral-line scale rows (33 vs. 34) and subcutaneous scales on breast region (vs. naked). From G. compressus in differs in having more pectoral-fin rays (13-15 vs. 12) and transverse (4.5/3.5/3.5 vs. 3.5/2.5/3.0) and circumpeduncular scale rows (16 vs. 12), as well as fewer lateral-line (33 vs. 39), pre dorsal-fin scale rows (10 vs. 12) and anal-fin scale rows (3-3.5 vs. 5). From G. litanensis it differs in having more lateral-line (33 vs. 32) and circumpeduncular scale rows (16 vs. 12). It is distinguished from G. namyaensis in having more pectoral-fin rays (13-15 vs. 12) and lateral-line scale rows (33 vs. 31), and fewer pre dorsal-fin scale rows (10-11 vs. 13). It is diagnosable from G. abhoyai it differs in having more pectoral-fin rays (13-15 vs. 11) and pre anal-fin scale rows (16-22 vs. 7), and fewer pre dorsal-fin scale rows (10 vs. 17); from Garra arupi in having more branched pectoral-fin rays (13-15 vs. 10-11) and fewer lateral-line (33 vs. 35-36), pre dorsal-fin scale rows (10 vs. 11-12) and anal-fin scale rows (3-3.5 vs. 6-8). It differs from G. gravelyi in having more pre dorsal-fin scale rows (10 vs. 8-9). It differs from G. nasuta (McClelland) in having fewer lateral-line scale rows (33 vs. 34) and poorly developed proboscis (vs. well developed trilobed proboscis). The species differs from G. minimus (described herein) in having fewer lateral-line (33 vs. 35-37) and anal-fin scale rows (3-3.5 vs. 4-6) and more circumpeduncular (16 vs. 12) and circumferential scale rows (22 vs. 18-20); from G. nigricauda sp. nov. in having fewer lateral-line scale rows (33 vs. 35-36); more circumpeduncular scale rows (16 vs. 12) and scales present on breast region (subcutaneous vs. well developed); from G. kimini in having subcutaneous breast scales (vs. naked), poorly developed proboscis (vs. present and well developed) and a shallow transverse groove on snout (vs. deep). It is diagnosable from G. magnidiscus in having more branched pectoral-fin rays (13-15 vs. 10-12), less pre dorsal-fin scale rows (10 vs. 12-15), less lateral-line scales (33 vs. 40-42), as well as more circumpeduncular scales (16 vs. 12-14).

Description

Body short and thin; dorsal profile not ascending. Snout rounded and with transverse groove. Orbit closer to preopercle that to snout [Table-4] . Rostral lobe distinct and with at least 16 pointed (erect, retrose, antrose) tubercles. Two pairs of barbels. Rostral barbel extending beyond rostral cap and placed inside a groove connected to rostral cap. Central pad wider than long. Upper lip without a median notch and upper jaw papilliferous. Lower fold covering 40% of central pad.
Dorsal-fin origin 2 scale rows anterior to pelvic-fin origin; first and second branched dorsal-fin rays longest; posterior margin of fin almost straight. Pectoral-fin rounded; when depressed not reaching pelvic-fin origin. Pelvic-fin with pointed tip; when depressed not reaching anal-fin origin. Anal-fin is very short; when depressed not reaching caudal base. Upper and lower lobes of caudal-fin equal.
Dorsal-fin rays ii, 8 (last ray branched at base); pectoral-fin rays i, 13-15; pelvic-fin rays i, 8; anal-fin rays ii, 5; caudal-fin rays 10+9 (17 branched); lateral-line complete, scales 33; lateral transverse scales 4.5 between dorsal-fin origin and lateral-line, 3.5 between lateral-line and pelvic-fin origin and 3.5 between lateral-line and anal-fin origin; pre dorsal-fin scale rows 10; circumpeduncular scale rows 16; circumferential scale rows 22; pre anal-fin scales 16-22 [Table-5] .

Scales

Belly scaled and subcutaneous scales in chest region.

Color in Live

Dorsum and dorso-lateral areas of body with light green base color to two scale rows below lateral-line; more ventrally base coloration silvery-white. Dorso-lateral scales with silvery center and dusky at margin and with large dark base. Lateral-line punctate; pores covered and encircled with melanophores, creating darker and more noticeable spots than as described for dorsal and dorso-lateral scales. Caudal peduncle posterior to posterior base of anal-fin greenish brown laterally, cream dorsally, and pinkish ventrally; laterally dorsal and ventral edges of scales with confluence of concentrated melanophores, creating 4 narrow stripes extending base of caudal-fin; base of caudal-fin with narrow dark band around edge; anterior to and separate from band central 3 scales at hypural plate with broad dark bark bar. Venter of head and body silver or yellow in life, cream-yellow following preservation.
Head pinking-brown dorsally; opercle, subopercle, cleithral area, extending to base of pectoral-fin insertion where spot is dark and large, greenish. Cheek, area encircling orbit, lacrimal area, and snout light pinkish-tan; lower cheek and branchiostegals, extending to venter, silvery. Well-developed, distinct black spot present just behind operculum, on the first lateral-line scale. Barbels whitish.
Dorsal-fin largely dusky with yellowish hue; leading unbranched rays and first 4 branched rays more darkly pigmented; bases of branched rays 2-6 black, creating spotted pattern on rays alone. Dorsally, pectoral-fin creamish-yellow basally on all rays and membranes; unbranched and 1st, 2nd, 3rd rays of branched rays light reddish; anterior 4-6 rays darkly pigmented posteriorly with thin line; ventrally basal 2/3 of fin yellow, light reddish distally. Pelvic-fin similar to pectoral-fin except unbranched and 1st and 2nd branched rays light reddish, remaining rays light greenish-yellow excluding last branched ray that is clear. Outer region of anal-fin whitish; base of fin with creamish-pink base for all rays; unbranched, 1st ray of branched ray, and distal end of 2nd branched ray light reddish; rays 2-5 yellow basally and extending distally to near edge of fin. Pectoral, pelvic and anal-fins are without black markings. Caudal-fin yellowish-cream, especially at base and along distal areas of lobes; ventral lobe with longer yellow extension; edges of rays and membranes dusky, slightly more dense centrally but not dark enough for any dark stripe extending from base of fin; tip of lower lobe with more dense concentration of melanophores in membranes and along rays, creating dark spot. Ventro-lateral scales yellowish; dorsal region of caudal peduncle yellowish in colour; middle region of caudal peduncle some scales are greenish in colour; 4th row of lateral scale rows have bunch of melanopores forming a green band up to the insertion of dorsal-fin. Melanopores concentration is more in tip of the scales; less in dorsal-fin insertion to end of the caudal peduncle region. Characteristically well-developed, distinct black spot present just behind operculum, on the first lateral-line scale. venter whitish; Barbels whitish. 3rd to 6th branched rays of dorsal-fin base with black dots. Outer region of pectoral-fin has whitish in colour; unbranched and 1st, 2nd, 3rd rays of branched rays are light reddish in colour; lower portion of the remaining rays are light yellowish in colour; upper portion has whitish in colour. Outer region of pelvic-fin have whitish in colour; unbranched and 1st and 2nd rays of branched rays are light reddish in colour; remaining rays are light greenish in colour excluding last branched ray. Outer region of anal-fin have whitish in colour; unbranched and 1st ray of branched rays are light reddish in colour; lower portion of the 2nd branched ray have light green in colour; upper portion of the 2nd branched ray have light reddish in colour; remaining rays are light green in colour excluding last branched ray. Pectoral, pelvic and anal-fins are without any black markings in the basal regions. Caudal peduncle with small black blotch. After preservation, there is no pigmentation in ventral surface.

Color in Preservative

Dorsum not dark; dorsum and venter light in colour. Dorsal-fin base with black dots. Pectoral, pelvic and anal-fins are without markings. Caudal peduncle with small black blotch.

Distribution and Habitat

This species has only been sampled from the Dikrong River at Boorum village [Fig-7] . This river is located near the Helipad of Itanagar in a village called Boorum. Along left bank (looking upstream) the river is shallow with a substrate of sand and cobble. The right river bank is highly confined and with more vegetation. Collections were made all along the shallow and deeper regions using cast nets, seines and hand nets. This area is densely populated with humans and the left bank is highly disturbed with dumping of litter. The extraction of cobbles from this area is major threat to fish habitats. Other species sampled from this site included Crossocheilus latius, Barilius bendelisis, Aspidoparia morar, Tor putitora, Chagunius chaugunio, Schistura sp., Psilorhynchus sp., Botia sp., Gagata cenia and Mystus sp.

Etymology

The name alticaputus is a Latin adjective referring to the deep head of the species.
Garra nigricauda sp. nov. [Fig-8]
Holotype: ZSI/SRS F8580, 1 ex. Male. 142.8 mm SL, Siang river, near Pasighat, Arunachal Pradesh, India. (N 270 55’ 22’’ E 950 05’ 54’’). Collectors: M. Arunachalam, M. Raja, C. Vijayakumar, M. Muralidharan. 13 Nov 2010.
Paratypes: MSUMNH 61, 5 ex. 102.2-142.8 mm SL, CMA 19, 9 ex, 92.6-114.7 mm SL. All other details same as Holotype.

Diagnosis

Garra nigricauda sp. nov. is distinguished from G. lissorhynchus in having more branched pectoral-fin rays (14-16 vs. 14); fewer pre dorsal-fin scale rows (9-10 vs. 13); fewer circumpeduncular scale rows (12 vs. 16); more circumferential scales (22-24 vs. 21); the lack of a W - shaped marking on caudal-fin (vs. Present); and a distinct black subterminal stripe in each caudal lobe. The species is distinguished from G. nambulica in having more branched pectoral-fin rays (14-16 vs. 12); fewer pre dorsal-fin scale rows (9-10 vs. 24); and fewer circumpeduncular scale rows (12 vs. 16); from G. annandalei by more pectoral-fin rays (14-16 vs. 13); fewer circumpeduncular scale rows (12 vs. 16); more pre anal-fin scale rows (22-24 vs. 18); and from G. gotyla gotyla by more lateral-line scale rows (34-36 vs. 33); fewer circumpeduncular scale rows (12 vs. 16).
It is diagnosed from G. elongata in having more branched pectoral-fin rays (14-16 vs. 12); fewer lateral-line scale rows (34-36 vs. 39); fewer pre dorsal-fin scale rows (9-10 vs. 13); and the presence of breast scales (vs. absent); from G. naganensis by fewer circumpeduncular (12 vs. 16) and pre dorsal-fin scale rows (9-10 vs. 12); from G. paralissorhynchus by more branched pectoral-fin rays (14-16 vs. 11); more lateral-line (34-36 vs. 31) and circumferential scale rows (22-24 vs. 20); fewer pre dorsal-fin scale rows (9-10 vs. 12); fewer circumpeduncular scale rows (12 vs. 16); from G. kempi sp. nov. in having more branched pectoral-fin rays (14-16 vs. 12); fewer lateral-line scale rows (34-36 vs. 40); and fewer pre dorsal-fin scale rows (9-0 vs. 12); from G. kalpangi in having more branched pectoral-fin rays (14-16 vs. 10-12); more lateral-line scale rows (35-36 vs. 32-33); and fewer circumpeduncular scale rows (12 vs. 16); from G. rupecula by more branched dorsal-fin rays (8 vs. 6-7) and lateral-line scale rows (35-36 vs. 32-34); fewer upper transverse scale rows (4.5 vs. 4.5-6.5) and well developed scaled on breast, belly, and post pelvic areas of venter (vs. naked breast, belly and post pelvic areas) from G. manipurensis by more pectoral-fin rays (14-16 vs. 12); lateral-line scale rows (35-36 vs. 34) and pre anal-fin scale rows (22-24 vs. 12) and fewer circumpeduncular scale rows (12 vs. 16); from G. compressus by more pectoral-fin rays (14-16 vs. 12) and transverse scale rows (4.5/3.0-3.5/3.5-4.0 vs. 3.5/2.5/3.0) and fewer lateral-line (35-36 vs. 39) and anal-fin scale rows (3-4 vs. 5). This species is distinguished from G. litanensis by more pectoral-fin rays (14-16 vs. 13) and lateral-line scale rows (35-36 vs. 32), and from G. namyaensis by more pectoral-fin rays (14-16 vs. 12); lateral-line scale rows (35-36 vs. 31) and fewer pre dorsal-fin scale rows (9-10 vs. 13) and circumpeduncular scale rows (12 vs. 16). It is differentiated from G. abhoyai in having more pectoral-fin rays (14-16 vs. 11) and lateral-line scale rows (34-36 vs. 33), and fewer pre dorsal-fin scale rows (10-11 vs. 17) and circumpeduncular scale rows (12 vs. 16). It is distinguishable from G. kimini sp. nov. by its overall size, more lateral-line (35-36 vs. 33-34) and pre anal-fin scale rows (22-24 vs. 17-21). It is diagnosable from G. magnidiscus in having more branched pectoral-fin rays (14-16 vs. 10-12), less pre dorsal-fin scale rows (9-10 vs. 12-15) and less lateral-line scales (35-36 vs. 40-42), as well as well developed scales on chest and belly (vs. Subcutaneous).

Description

Body elongate and robust, with rising dorsal profile. Ventral surface on head and breast flattened, abdomen slightly rounded [Table-6] .
Snout rounded, shallow, and with long transverse groove with rostral lobes marginally adnate. Tubercles present laterally on snout to level of nostrils. Two pairs of barbels. Rostral barbel reaching margin of rostral cap. Central pad wider than long. Between exposed lower jaw and lower lip short straight papilliferous fold extending medially from corner of mouth.
Dorsal-fin insertion 2.5 scale rows anterior to pelvic-fin origin. First and second branched rays of dorsal-fin longest; posterior margin of fin concave, longest ray of depressed fin vertical to pelvic-fin insertion; and vertically 5 scale rows. Pectoral-fin is rounded; branched rays 4-5 longest, when depressed extending to base of pelvic-fin. Pelvic-fin is rounded, first and second branched rays longest, when depressed almost reaching base of anal-fin. Anal-fin pointed, first branched ray longest, when depressed tip extending to base of caudal-fin base. Tips of caudal-fin lobes pointed, upper lobe shorter than lower lobe.
Dorsal-fin rays ii, 8 (last ray branched at base); pectoral-fin rays i, 14-16; pelvic-fin rays i, 8; anal-fin rays ii, 5; caudal-fin rays 10+9 (17 branched); lateral-line complete, scales 35-36; lateral transverse scales 4.5 between dorsal-fin origin and lateral-line; 3.0-3.5 between lateral-line and pelvic-fin origin; 3.5-4.0 between lateral-line and anal-fin origin; pre dorsal-fin scale rows 9-10; circumpeduncular scale rows 12; circumferential scale rows 22-24; pre anal-fin scale rows 22-24 [Table-7] .

Scales

On venter, well developed scales in chest and belly region.

Color in Preservative

All the specimens were obtained from the market but were relatively fresh and had lost some of their wild colouration or it appeared somewhat muted; however, specimens were intact without any damage (damaged specimens were not included in the study). Dorsum and sides of head and body with base coloration grayish-brown; scales above lateral-line and on dorsum with light grayish-brown center and thin, darkly pigmented margins, creating a distinct scaled pattern. Below lateral-line base coloration lighter; some, but not all, scales with light center, as above lateral-line, but with broader marginal band on scales, creating a somewhat mottled color pattern. Mid-lateral stripe appearing lighter than coloration above or below lateral-line; stripe three scale rows deep anteriorly and narrowing to one scale row on caudal peduncle. Base of caudal-fin with dark round spot surrounded posteriorly, dorsally, and ventrally by creamish-colored area; posteriorly to cream area complete base of caudal-fin with broad dark bar extending from, but not including, dorsal and ventral procurrent rays. Caudal peduncle coloration same as anterior part of body; no narrow dark stripes from pigmentation on dorsal and ventral edges of scales. Venter of head immaculate; breast, belly, and region between pelvic-fin insertion and anal-fin origin immaculate to light cream; post anal-fin area of caudal peduncle creamish but also with some melanophores and appearing darker approaching caudal-fin.
Head with large dark and oval-shaped spot posterior to eye and directly above opercle and dorsal preopercle. Orbit encircled by broad, creamish-colored ring. Lacrimal region and tip of snout connected and light gray. Dorsal-fin with dusky unbranched and branched rays; bases of branched rays, beginning with second branched ray with distinct dark and elongate spots with melanophore covering the rays in this region; distal to dark part of ray, rays yellowing-red; beginning with second branched fin ray all membranes black. Distal tip of tallest unbranched and first branched rays yellowing-red. Dorsum of pectoral-fins are with narrow white distal band on leading unbranched ray and six branched rays; basally and medially rays reddish-orange and membranes dusky; leading ray darkly pigmented complete length and set apart from all other rays of fin; subdistally fin with broad black band, widest anteriorly and narrowing to only margin after 8th ray. Basal area at pectoral-fin insertion darkly pigmented and set apart from coloration of body. Ventrally, pectoral-fins with distinct yellowish colouration, beyond which fin is immaculate. Pelvic-fin with yellowish-red color widely distributed and dark pigmentation restricted to membranes, darkest in first membrane; distally, fin with broad light to clearish band. Ventrally pelvic-fin colored as in pectoral-fin. Anal-fin coloration with more reddish-yellow pigmentation basally and extending to medial area of fin; leading rays not darkly pigmented, colored reddish-yellow; distal tip of anterior few rays clear to creamish; distal to reddish-yellow coloration fin with broad black band. Caudal-fin rays and membranes dusky; median branched rays and membranes black, creating appearance of stripe in center of fin; first and second branched caudal rays and membranes of lower lobe black distally and becoming more gray anteriorly, creating stripe; upper and lower lobes of caudal-fin and distal edge black; black area largest in lower lobe. Fins grayish with markings along the outer margins of pectoral, pelvic and caudal-fins; caudal-fin also tipped with black with more concentrated on the lower lobe than upper lobe. Anal-fin with black streaks; edge of fin black.

Distribution and Habitat

This species has only been sampled from the Siang River, near Pasighat [Fig-9] . Siang River is a 5th order river with unstable bands on both sides. Substrate types is mostly cobbles and sand. The width of the river is 0.5-0.6 km. Except fishing activity there is no disturbance in this stretch where G. nigricauda is collected.

Etymology

The name is nigricauda is Latin referring to the black caudal-fin edges.
Garra kimini sp. nov. [Fig-10]
Holotype: ZSI/SRS F8581, 1 ex. Male. 83.4 mm SL, tributary of Ranga River, 7 km from Hola camp, Lower Subanshri District, Arunachal Pradesh, India (N 270 20’ 59’’ E 930 56’ 37’’). Collectors: M. Arunachalam, M. Raja, C. Vijayakumar and S. Nandagopal. 18 Jun 2011.
Paratypes: MSUMNH 62, 10 ex. 62.3-85.0 mm SL, CMA 20, 6 ex, 64.9-78.4 mm SL. All other details same as Holotype.

Diagnosis

Garra kimini is distinguished from G. lissorhynchus by fewer pre dorsal-fin scale rows (9-10 vs. 13); more circumferential scales (22-24 vs. 21); and lack of W - shaped marking on caudal lobe (vs. W - shape marking present). From G. nambulica it is distinguished by more branched pectoral-fin rays (14-15 vs. 12); fewer pre dorsal-fin scale rows (9-10 vs. 24); naked breast (vs. well-developed scales). It is distinguished from G. annandalei in having more pectoral-fin rays (14-15 vs. 13); circumferential (22-24 vs. 22); and pre anal-fin scale rows (17-21 vs. 18); proboscis with slight protrusion (vs. no proboscis); and deep transverse groove (vs. transverse groove absence). The species is distinguished from G. gotyla gotyla in having more lateral-line scale rows (33-34 vs. 33); naked breast (vs. well-developed scales); and slight protrusion for proboscis (vs. well-developed median proboscis); from the Garra elongata by more branched pectoral-fin rays (14-15 vs. 12) and fewer lateral-line (33-34 vs. 39) and less pre dorsal-fin scale rows (9-10 vs. 13); and poorly developed scales in breast region (vs. naked); from G. naganensis in having fewer lateral-line (33-34 vs. 39) and fewer pre dorsal-fin scale rows (9-10 vs. 12); and naked breast area (vs. reduced breast squamation); from G. paralissorhynchus by more branched pectoral-fin rays (14-15 vs. 11) and lateral-line (33-34 vs. 31) and circumferential scale rows (22-24 vs. 20) and fewer pre dorsal-fin scale rows (9-10 vs. 12).
The species is diagnosed from G. kempi in having more branched pectoral-fin rays (14-15 vs. 12) and fewer lateral-line (33-34 vs. 40) and pre dorsal-fin scale rows (9-10 vs. 12); from G. kalpangi in having more branched pectoral-fin rays (14-15 vs. 10-12). The species is further distinguished from G. rupecula in having more branched dorsal-fin rays (8 vs. 6-7); fewer upper transverse scale rows (4.5 vs. 4.5-6.5); and well developed belly scales (vs. breast area naked); and well developed scales on belly (vs. belly naked); from G. manipurensis by more pectoral-fin rays (14-15 vs. 12); fewer anal-fin scale rows (2.5-3.0 vs. 4) and pre dorsal-fin scale rows (11 vs. 9-10); and more pre anal-fin scale rows (17-21 vs. 12); from G. compressus by more branched pectoral-fin rays (14-15 vs. 12); fewer lateral-line (33-34 vs. 39), pre dorsal-fin scale rows (9-10 vs. 12) and circumpeduncular scale rows (16 vs. 12); from G. litanensis by more branched pectoral-fin rays (14-15 vs. 13); lateral-line scale rows (33-34 vs. 32); and fewer anal-fin scale rows (3.5 vs. 2.5-3.0). The species is diagnosed from G. namyaensis by more branched pectoral-fin rays (14-15 vs. 12); more lateral-line scale rows (33-34 vs. 31); fewer pre dorsal-fin scale rows (9-10 vs. 13); and from G. abhoyai by more pectoral-fin rays (14-15 vs. 11); fewer pre dorsal-fin scale rows (9-10 vs. 17) and more pre anal-fin scale rows (17-21 vs. 7). It is diagnosable from G. magnidiscus in having more branched pectoral-fin rays (14-15 vs. 10-12), less pre dorsal-fin scale rows (9-10 vs. 12-15) and less lateral-line scales (33-34 vs. 40-42), as well as more circumpeduncular scales (16 vs. 12-14), presence of tubercles on pectoral-fin (vs. absence).

Description

Body elongate, slightly compressed laterally, becoming more extreme on caudal peduncle. Dorsal profile of body slightly convex from tip of occiput to dorsal-fin origin [Table-8] .
Base of dorsal-fin sloped slightly. Dorsal profile straight from posterior end of base of dorsal-fin to base of caudal-fin. Ventral profile of body almost straight from pectoral to pelvic-fin insertions and from pelvic-fin insertion to anal-fin origin. Posteroventally, anal-fin base nearly straight. Profile between posterior end of base of anal-fin to base of caudal-fin straight. Head moderately large, depressed and with a convex interorbital space; head height less than length; head width almost equal to height. Snout broadly rounded and with a narrow, shallow groove across snout tip, forming a transverse lobe with 12-13 small to large erect tubercles. Two pairs of barbels; rostral barbels located antero-laterally and longer than diameter of eye; maxillary barbels at corner of mouth and shorter than rostral barbels. Rostral cap well developed, moderately granulated and with a narrow papillated margin. Disc elliptical, length shorter than wide. Caudal-fin emarginated; lobes equal in length and tips slightly pointed.
Dorsal-fin rays ii, 8 (last ray branched at base); pectoral-fin rays i, 14-15; pelvic-fin rays i, 8; anal-fin rays ii, 5; caudal-fin rays 10+9 (17 branched); lateral-line complete, scale rows 33-34; 4.5 lateral transverse scale rows between dorsal-fin origin and lateral-line; 3.5 between lateral-line and pelvic-fin origin; and 3.5 between lateral-line and anal-fin origin; pre dorsal-fin scale rows 9-10; circumpeduncular scale rows 16; circumferential scale rows 22-24; pre anal-fin scale rows 17-21 [Table-9] .
Tubercles are absent on pelvic, anal and caudal-fins. Dorsal surface of pectoral-fin covered with spiny tubercles; each ray bearing numerous minute organs for its entire length. Leading unbranched ray with 3 rows of tubercles; first row with 27 tubercles, second row with 23, and third row with 38. First branched ray of pectoral-fin with 2 rows of tubercles; first row with 58 tubercles and second row with 30. The second branched ray also with 2 rows; first row with 52 tubercles and second row with 35. The third branched ray with 2 rows of tubercles; first row with 54 tubercles and second row with 37. Pectoral-fin with erect tubercles to 7th branched ray.

Scales

Scales on chest naked and Belly scaled.

Color in Preservative

In live specimens head, body, and dorsum grey with a blackish spot immediately posterior to dorsal gill-opening. Colours of dorsal, pectoral and pelvic-fins are not distinct.

Distribution and Habitat

Only one population is currently known for this species from a stream near Hola camp. This stream confluences with Ranga River in Lower Subanshri district [Fig-11] . With few human settlements in the area, the terrestrial habitats and stream appears to be relatively undisturbed. As a third-order stream it contains numerous rocks and large boulders. At the time of collection flow was not high and the stream. Morphology of stream valley “U”- shaped within this reach and not steep enough; slopes of both sides shallow. Because of the U-shaped valley, stream gradient was less than 5% and water velocity was low. Species of Garra was found in areas with a high velocity and the substrate types are mostly large boulders. Loaches were collected by local people in areas with cobbles and sand. Water was clear and appeared unpolluted, although there were visual signs of the stream being used by nearby villagers for bathing and washing clothes.

Etymology

The name kimini is a noun in apposition and refers to the nearby village where the new species was collected.

Comparisons of Species Using Morphometric Characters

Principal component analysis provides a summarization of the morphometric variation and comparisons of the four new species relative to one another [Fig-12] . This analysis demonstrates the distinctiveness of these species in morphometric characters serving to provide information regarding shape. Characters of high loading on PCI included head depth at occiput, head depth at pupil, peduncle depth, dorsal-fin insertion to pelvic-fin insertion; those on PCII included pelvic-fin length, disc length, disc width, central pad width. The summary of the shape variables in this analysis, as well as the data provided in diagnoses, descriptions, and comparisons, in combination with meristic characters, coloration, lip and disc morphology, demonstrate these four species as both diagnosable and distinct evolutionary lineages (sensu Evolutionary Species Concept). Using this summary of morphometric features G. minimus, G. alticaputus, G. nigricauda and Garra kimini are clearly distinct.

Discussion

characteristic feature of Asian Garra [27,28] . Species of Garra thus far reported or recently described from the Indian state of Arunachal Pradesh include: G. lissorhynchus, G. paralissorhynchus, G. annandalei, G. gotyla gotyla, G. kempi, G. lamta, G. naganensis, G. arupi, G. nasuta and G. rupecula [8] Garra minimus, Garra nigricauda, Garra elongata and Garra kempi have 12 circumpeduncular scale rows while all other species have 16 circumpeduncular scale rows, except for Garra alticaputus and Garra kimini that also has 16 circumpeduncular scale rows. G. minimus sp. nov. differs from G. manipurensis [14] by the presence of proboscis and transverse groove; showed variation in lateral-line scales and more pectoral-fin rays. Another described species, G. compressus [16] from Manipur also showed distinctive characters by the presence of proboscis and transverse groove and also by the scale less chest. Also described from the same locality, G. litanensis [14] is closely related to G. minimus, but with the differences is in more number of lateral-line scales. G. alticaputus sp. nov. is also distinguished from G. manipurensis by not well developed proboscis and shallow transverse groove; showed variation in pectoral-fin rays. Also G. alticaputus showed variation from G. compressus by more branched pectoral-fin rays; less lateral-line scales; less pre dorsal-fin scales and the presence of proboscis and transverse groove on snout with belly scaled and subcutaneous scales on chest region. G. alticaputus showed distinction with G. litanenais by the well developed scales on belly. G. nigricauda sp. nov. is closely related to G. manipurensis but the variations are in more number of pectoral-fin rays and the presence of proboscis and transverse groove and also presence of scales on chest and belly. It showed variation with G. compressus by also more pectoral-fin rays; less lateral-line scales; less pre dorsal-fin scales and the presence of transverse groove on snout and also the presence of scales on belly and chest. G. nigricauda is also distinguished from G. litanenais by more number of lateral-line scales. G. kimini sp. nov. is also distinct from G. manipurensis by more branched pectoral-fin rays; presence of proboscis and transverse groove on snout. G. kimini is also distinct from G. compressus by more branched pectoral-fin rays; less lateral-line scales; less pre dorsal-fin scales. However, G. elongata has 39 lateral-line scale rows while G. kempi has 40 lateral-line scale rows and both species have 1 vs. 2 unbranched dorsal-fin rays and 7 vs. 8 branched dorsal-fin rays, respectively. G. minimus has 14-16 branched pectoral-fin rays which is higher than any Garra species reported or described so far from India.
Arrangement of breeding tubercles on the head and body to be of taxonomic importance. Bailey and Suttkus [29] ; Robins and Raney [30] and earlier authors Gibbs [31] ; Hubbs and Black, [32] considered them to be valuable indications of phylogeny. The distribution of tubercles in the genus Garra has been reported only from the regions of snout and cheek and however, the distribution of ‘Pearl organs’ to other parts of body and fins can be considered as taxonomically important. A similar condition of the distribution of nuptial tubercles in males and sometimes in females from head to trunk and all of the fins up to caudal has been reported in some catostomid fishes from Kansas.
It was noted by Kullander and Fang [27] that lip structures are important for the identification of most species of Garra and are also important for resolving their phylogenetic relationships. It was lamented that most of the new descriptions of Garra are lacking the details of lip structure and hence are not comparable. However, Garra minimus and Garra alticaputus have distinct upper-lip structures formed by papillae. Garra rupecula McClelland [8] was described from the Mismi Mountains, Assam (now in Arunachal Pradesh) of the Bramaputra River basin and as having a W-Shaped band on the caudal-fin and absence of scales in the pre dorsal-fin areas [13] ; however, types of this species are not available for examination. McClelland described Garra nasuta under Gonorhynchus from a single specimen collected from the Kasya Mountains. However, the real identity of this species is not clear as the description was quite inadequate. Thus, Menon [33] described this species from specimens collected from Kasya mountains. However, the species described by Menon [33] turned out to be an another species, Gonorhynchus caudatus McClelland [34,35] .
Garra is one of the most diverse and taxonomically confusing groups of the Cyprinidae (Cypriniformes) that has yet to be examined for evidence of monophyly or examined a breadth of morphological and molecular traits. Given the distribution of this genus, information as to the genealogical relationships of the species would provide critical historical inferences as to the biogeographic connections both within and between Africa and Asia. However, the taxonomy of Garra is not mature enough to support such a well-developed hypothesis and awaits further studies using morphological and molecular data focused on species-group level questions. Many old descriptions exist and many names are available but resolution of species diversity and nomenclature within this putative monophyletic genus is in need of great attention by many researchers. The above example is only one of several cases wherein there is considerable confusion of appropriate names, lack of type materials, and unknown geographic variation that may or may not be considered species diversity. Phylogenetic studies are critically important even now with confused nomenclature but not until many of the types, names, and nomenclatorial issues remaining in the genus are resolved, taxonomic revisions and a well-resolved phylogeny must await more detailed examine and report and illustrate the same characters for comparisons and diagnoses. This would initially be well served if a prioritized and collaborative effort were conducted for collections of topotypes of species of Garra and dorsal, lateral, and ventral color photos were made available of the whole body, head and fins, as well as consistency in the collection of comparative characters.

Comparative Materials Examined

Garra lissorhynchus Topotype: ZSI Calcutta, FF 8098 / 1; 73.05 mm SL; Collected by: L. Kosygin. Garra nambulica Paratype: ZSI Calcutta, 4139; 50.41 mm SL; Irengloic Stream flowing to Nambul River, Shingala Village, Imphal West District, Manipur, India. 03rd Feb. 2004. MUMF 8002; Type series examined from Manipur University Museum of Fishes. Garra annandalei Holotype: ZSI Calcutta, F 6082 / 2-1; 60.17 mm SL; Kokha nallah, Koshi river, District: Barabakshetra. India. 30th Jan.1946. Garra gotyla gotyla ZSI Calcutta, F 198/2; 121.92 mm SL; Kumaon Hills survey, Kosi River Kosi village, Almorah, 07th June1948. Garra elongata ZSI Calcutta, FF 4157; 81.28 mm SL; Hill stream near Tollai, Ukhrul district, Manipur, India, 12th Nov. 1997. Garra naganensis ZSI Calcutta, F 9970 / 1; 89.93 mm SL; Senapathi Stream, Naga hills, Assam, India). Collector L. Kosygin; Type series examined from MUMF. Garra paralissorhynchus Paratype: ZSI Calcutta, 4158; 52.35 mm SL; Location: Khuga River, Churachandrapur district, Manipur, India. 25th July 2000. Garra kempi Holotype: ZSI Calcutta, F 7716 / 1; 88.51 mm SL; Sharjon River, below Damda, the Abor hills, India. 25th July 2000 [Fig-2] . Garra abhoyai Type series examined from MUMF. Garra manipurensis Type series examined from MUMF. Garra compressus Type series examined from MUMF. Garra litanensis Type series examined from MUMF. Garra namyaensis Type series examined from MUMF [Fig-3] . Garra kalpangi data from Nebeshwar, et al [20] . Garra arupi data from Nebeshwar, et al [21] . Garra nasuta data from Monograph [33] of the Cyprinid fishes of the genus Garra Hamilton-Garra rupecula data from Talwar and Jhingran [36] . Garra dampaensis data from Lalronunga, et al [23] . Garra magnidiscus data from Tamang [24] .

Acknowledgements

This research was supported in part by the Sri Paramakalyani Centre for Environmental Sciences, (DST-Government of India-S/FIST/ESI-101/2010), Manonmaniam Sundaranar University, Saint Louis University, and the USA National Science Foundation Grants EF-0431326, DEB-1021840 and DBI-0956370 to RLM. This work was initiated, intended and completed as part of collaborative international initiatives to develop infrastructure in the taxonomy and systematics of Cypriniformes, likely the most diverse group of freshwater fishes in the world. The two initiatives Cypriniformes Tree of Life and All Cypriniformes Global Biodiversity Initiative (www.cyprinifor mes.org) have aided in this mission. We acknowledge the help of Mr. S. S. Mariappan for line drawing. Special thanks to Dr. K. Venkataraman, Director, ZSI Kolkatta; Shri. K.C Gopi, Scientist E, ZSI Kolkatta; Dr. K. Illango, Scientist D, ZSI/ SRS Chennai; Dr. B.H.C Murthy, Scientist B, ZSI Kolkatta; Dr. Jayasree Thilak, Scientist C, ZSI/ SRS Chennai for access to the collection of fishes and examination of species of Garra and Dr. W. Viswanath, Professor, Manipur University for the loan of specimens for comparative purposes.

Conflicts of Interest

None declared.

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Images
Fig. 1- Morphometric measurement of Garra
Fig. 2- Type specimens of Garra from ZSI, Kolkatta
Fig. 3- Type specimens of Garra examined from MUMF, Manipur. (A to E)
Fig. 4- Garra minimus sp. nov.
Fig. 5- Ranga River, the type locality of Garra minimus sp. nov.
Fig. 6- Garra alticaputus sp. nov.
Fig. 7- Dikrong River, the type locality of Garra alticaputus sp. nov.
Fig. 8- Garra nigricauda sp. nov.
Fig. 9- Siagng River, the type locality of Garra nigricauda sp. nov.
Fig. 10- Garra kimini sp. nov.
Fig. 11- 7 km away from Hola camp, the type locality of Garra kimini sp. nov.
Fig. 12- Principal Component plot of axes 1 and 2 for pooled morphometric data derived from four new species of G. minimus, (Cross) G. alticaputus, (Square) G. nigricauda (filled square) and G. kimini (diamond) from Arunachal Pradesh, India.
Table 1- Comparative meristic characteristics of species of Garra from ZSI and MUMF, with Garra minimus, Garra alticaputus, Garra nigricauda and Garra kimini sp. nov.(counts)
Table 2- Morphometric characteristics of Garra minimus sp. nov. from Ranga river, Arunachal Pradesh
Table 3- Morphometric characteristics of Garra alticaputus sp. nov. from Dikrong river, Arunachal Pradesh
Table 4- Morphometric characteristics of Garra nigricauda sp. nov. from Siang river, Pasighat, Arunachal Pradesh
Table 5- Morphometric characteristics of Garra kimini sp. nov. from 7 km away from Hola camp, Arunachal Pradesh.
Table 6- Meristic characteristics of Garra minimus sp. nov. from Ranga river, Arunachal Pradesh
Table 7- Meristic characteristics of Garra alticaputus sp. nov. from Dikrong river, Arunachal Pradesh.
Table 8- Meristic characteristics of Garra nigricauda sp. nov. from Siang river, Pasighat, Arunachal Pradesh
Table 9- Meristic characteristics of Garra kimini sp. nov. from 7 km away from Hola camp, Arunachal Pradesh.