NAT. NAT. HIST. B 札L. SIAM Soc. 42: 263-290 ， 1994

SYSTEMATIC REVISION OF TROPICAL ASIAN FRESHWATER GLASSPERCHES (AMBASSIDAE) ，WITH DESCRIPTIONS OF THREE NEW SPECIES

Tyson Tyson R. Roberts*

ABSTRACT

Continental Continental fresh w蹴 rs of 加 pical Asia 眠 inhabited by 5 genera and 15 freshwater species species of 山 predominantly marine percomorph far 凶 Iy Ambassidae (formerly known 錨 Chandidae). Chandidae). P紘 ist 組 h 箇 3 species; In dia has 6; Nepal ，Bangladesh ，B 町 ma ，司副 land ，and Cambodia Cambodia each have 4; and Indonesia has 6. 百 ese fish 巴 sare notably diver 冒e，including specialized two specialized monotypic genera of scale-eaters and one scale-less genus in which males

have have gr 官atly expanded and color 和1dorsal and anal fins. 羽田e new species are described ， Gymnochanda flamea from southern Borneo ， Parambassis tenasserimensis from the biogeographically biogeographically isolated Tenasserim basin in southeastern Burma ， and the elegant Parambassis Parambassis vollmeri 合om the Salween basin of Burma and 百lailand. Significant range extensions extensions in cJ ude Parambassis lala (previously known only from Pakistan ，In dia ， and Bangladesh) 仕om 血e Irra waddy basin and Parambassis apogonoides (previously known only only from Indone 唱ia) from the Pahang basin in peninsular Malaysia and the Mekong basin in in Cambodia and southern La os (below Le e Pee Waterfalls).

町 τ'R ODUCTION

Marine Marine and freshwater Am bassidae were last revised by FRASER-BRUNNER (1955). Sever a1 species have been discovered since then ， and much more has been learned about geographic a1 distribution and biology ，esp 配 ia1 1y of the f詑shwater forms. Endemic fresh- water water glassperches or Am bassidae occ 町 in Madag 部 car ， the Austr a1 ian region (泊c1 uding New Guinea) ，and 位。 pic a1 Asia.τ 'h ey are most numerous in In dia and Bomeo. Am bassidae is is the only secondary freshwater fru:凶 ly of percomo 甲h fishes in tropic a1 Asia in which 企eshwater forms 紅 'e morphologic a1 1y and biologically more diverse 白an m 訂加e ones. Th e freshwater forms in c1 ude 出 e largest and the sm a1 1est members of 白 e fatnily in Asia; the the only ones known with highly speci a1 ized dentition and feeding habits (i. e. ，出 e sc a1 e- eating eating genera Paradoxodacna and Chanda); the only known oral-brooding species (Parambassis (Parambassis apogonoides); and the only forms with filamentous fins or marked sexu a1 dimo 甲hism (Gymnochanda). Wh ereas nearly a1 1 marine species have large sc a1 es (usu a1 1y around around 30 in lateral series) ，freshwater forms range from the sc a1 eless Gymnochanda to Chanda with very fine sc a1 es (1 25 in lateral series). 百le r佃 ge in number of dors a1 and an a1 fin rays also is greater in freshwater forms ， as is the number ，size ， and distribution of serrations serrations on headbones.

*Research *Research Affi liate ，Smithsonian Tropical Research Institute

263 263 264 TYSON R. ROBER 百

Many In do-Pacific marine ambassid species with large scales and relatively short upωmed jaws enter 企'eshwater ， but only 泊 the lowermost or tidal portion of rivers and in relatively relatively isolated rivers (e.g. ，on islands). 百lU S 白ey 紅 e not characteristic elements of 血e continental continental Asian freshwater fish fauna. 百lI s paper is concemed only with the endemic freshwater freshwater forms. So far as 1 have b田 n able to ascertain ， the species treated herein do not inhabit inhabit marine ，estu 紅 ine or brackish waters ，although several sometimes occ 町 in strongly tidal tidal but still largely or entirely freshwater habitats.

MA'τERIALS AND METHODS

Specimens Specimens examined are deposited in the following institutions: ANSP ，Academy of Natural Natural Sciences ，Philadelphia; BMNH ，British Museum ofNatural Hi story ， London; CAS ， California California Academy of Sciences ，San Francisco; MCZ ，Museum of comp 紅 'ative Zoology ， Cambridge; Cambridge; MNHN ，Mus 色um National d'Histoire Naturelle ，P 釘 is; RMNH ， National Natuurhistorisch Natuurhistorisch Museum ，Le iden; MZB ，Museum Zo ologicum ， Bogor; USNM ， National Museum of Natural History ，Washington ， D.C.; ZMA ，Zo ological Museum ，Universitet V 佃 Amsterdam; and ZRCS ，Zo ological Reference Collections ， National University of Singapore. Singapore. Th ere has been considerable confusion about the distribution of f記shwater species of Am bassidae ，especially in India ， often due to misidentified specimens and erroneous 10- cality cality records. 1 have tried to r民 tify 偽is by collecting fresh material and by exami 凶ng and reidentifying older material in many museums. Unfortunately locality da ぬ form 叩 y lots lots is vague or dubious ，and 1 have felt obliged to exercise discretion in deciding whether to to report 血em.τbus BMNH has old lots of Chanda nama and Parambassis lala supposedly collected collected in Ceylon (Sri Lanka) ， where probably neither occurs. In some instances ， the co 町 ectness of the localities has been queried by earlier workers in the catalogues or on the bottle bottle labels. Many lots bearing relatively uniformative locality data such as “In dia" 釘 e not not included under material examined ，even 血ough 血ey were examined and re ・identified by me. Readers Readers familiar with systematic papers on Asian 企eshwater fishes w 出 notice 白e relatively relatively large amount of material 白at has been identified and reported upon. It is p紅白ul 釘 ly important to record well documented and identified material 合om numerous localities localities in view of the widespread disappear 組， ce of many species and the paucity ， of reliable reliable dis 凶butional information. For 白is reason also it is essential to record the date of collection collection and collector ，so 出at we can know not only where species occurred ， but when 出ey occurred 血ere.τbe name of the collector helps establish proven 姐 ce of the speci- mens and provides a means of confm 凶ng locality data and dates of collection. Ol der specimens usually are accompanied by minimal locality data. 百le most useful item item in reporting locality data for freshwater fishes generally is the name of the river basin ， but 血 is is seldom stated for older specimens. Instead ， there may be a poorly legible locality ， or unfar 凶li 紅 spelling of a place n創 ne. 1 tend to re 金ain from report 泊g sp 回出lens with with dubious or illegible locality data. An d while some editors insist on up-dating 1ωality data data on old museum specimens ，1 feel 出is is a serious waste of time and a misguided effort. effort. Modernizing old localities is fraught with potential misinterpretations and e町'O rs. In In many instances there is no modem geographical equivalent for old geographical terms. SYSTEMA TIC REVISION OF TROPICAL ASIAN FRESHW A TER GLASSPERCHES 265

Origin a1 loc a1 ity data (in c1 uding spelling) is part of the proven 組 ce of sp 田 imens ，and should should not be a1 tered at the whim of an author ，editor ， or data-banker. To cite just one particularly particularly egregious error ， an editor who insisted on using only the most modern geo ・ graphic a1 place names changed the spelling of 出e Me k1 0ng Ri ver to Mekong throughout one one of my papers. Fortunately it was possible to co 町 ect 血 is particular mistake in 白e page- proofs. proofs. Wh en it seems 出at up-dating or otherwise improving loc a1i ty data would be useful and and unlikely to introduce error 1 give the addition a1 information or interpretation 泊 brack- ets ， sometimes with a question m 釘k. Th e nomenclature for head serrations in Ambassidae was established by FRASER- BRUNNER (1 955). In Am bassidae the membranous tubules of 白e ceph a1 ic laterosensory system 訂 'e grea t1 y enlarged and exposed. 百le bony laterosensory can a1 s on 血e preorbi ta1， suborbit a1， and preopercul 紅 bones have become open troughs wi 出 both margins often be 訂泊g seπae (bony projections resembling saw-teeth). 百lesem 紅 gins 紅 ec a1 1ed edge and ridge ridge by FRASER-BRUNNER.

SYSTEMAτlC ACCOUNT

C加 nda Ha 凶lton ，1822

Chanda Har 凶lton ，1822:103 ，370 (placed on Offici a1 Li st of Generic Narnes in Zo ology;

type type sp 田 ies Chanda nama H 創 ni1 ton ，1822 ，by designation of ICZN: Opinion 1121). Hamiltonia Hamiltonia Swainson ， 1839:176 (type species Hamiltonia ovata Swainson ， 1839 = C. nama Harnilton ， 1822; placed on Offici a1 Index of Rejected and In valid Generic Narnes in Zoology: Zoology: Opinion 1121). Bogoda Bleeker ， 1853:89 (type species by absolute tautonomy Chanda bogoda Har 凶 ton 1822 1822 = Chanda nama Harnilton 1822).

Diagnosis Diagnosis (modified from ROBERTS ，1989:159-60). - The speci a1 ized lepidophagous or scale-eating scale-eating genus Chanda differs from a1 1 other arnb 部 sids in having a strongly projecting lower lower jaw; preorbit a1 edge with 4 serrae ，ridge non-se 町ate except for prim 紅 y seπa; cheek ， i11 gi11 cover ， predorsum above later a1 line and body just below base of dors a1 fm sc a1 eless; sc a1 es on body very sm a1 1， about 125 in later a1 series; posterior nostril vertic a1 1y ov a1， about about twice as large as anterior nos 凶1. It differs from a1 1 other arnbassid genera except the the sc a1 e-eating Paradoxodacna in having enlarged jaw teeth in two main rows ， teeth in outer outer row with strongly curved crowns directed later a1 1y and those in inner row with similarly similarly curved crowns directed medially. Further distinguishing characters are gi11 rakers on first gill arch 5+13 = 18; dors a1 fin soft rays 15 ・17 ，叩a1 14 ・17; pector a1 fin rays 13; a vertic a1 1y elongate humer a1 spot; vertebrae 10+15 = 25 (rarely 10+16 = 26). Only a single single species known.

Chanda nama H 創凶lton ，1822

Chanda nama Har 凶lton ， 1822: 109 ，371 ， p 1. 39 ，fig. 37 (type loc a1 ity “ ponds throughout Beng a1"). 266 TYSON R. ROBERTS

Chanda phula Hamilton ， 1822: 111 ， 371 (type locality “勺 nor 此th- 屯ea 邸st 飽.er 口m百1 P紅 t岱s of Bengal" 勺，ヲ) Chanda bogod αHamilton， ， 1822: 111 ， 371 (t 旬yp 戸e localit ザy ‘“‘、 no 託h- 毛ea 儲st 旬emp 紅 t臼so ぱf Bengal" 勺，ヲ) Amηlb 如assis め0 blong 伊α， Cu 町lvi 町e釘釘ri泊nCωuv 吋ie 釘r & Valenciennes ， 1828: 185 (no type locality). Ambassis Ambassis nama ， Cuvier in Cuvier & Valenciennes ， 1828: 185. Ambassis Ambassis phula ， Cuvier in Cuvier & Valenciennes ， 1828: 186. Ambassis Ambassis bogoda ， Cuvier in Cuvier & Valenciennes ， 1828: 187. Hamiltonia Hamiltonia ovata Swainson ， 1839: 176 (replacement name for Chanda nama Hamilton 1822). 1822). Placed by ICZN on Official Index of Rejected and Invalid Specific Names in Zoology Zoology (Opinion 1121). Bogoda nama ，Bleeker ， 1853: 89. Hamiltonia Hamiltonia nama ，Fraser-Brunner ， 1955: 209.

Type material examined. - ~在NI王N 2952 ， 49.7 mm (holotype C. oblonga).

Non-type material examined. - MCZ 25166 ， 7:24.9-36.7 mm ，Chenab R. ， Indus basin ，Pu njab ，Pakistan ，M. M. Carlton; ZMA 115.948 ，2:41 ふ 48 .5 mm ，Pakistan ，Manha (Manka?) Canal ，Sep t. 1977 ，M. Sadiq; MCZ 9711 ，9:26.7 ・37.2 mm ，and 9712 ， 8: 17.0-50.8 17.0-50.8 mm ，Sursutiu R.， N India ，M. M. Carlton; BMNH 1889.9.26.1-9 ， 9: 38 .4 -45.1 mm ，Deoli ，R 勾putana ， F. Day; BMNH 1889.2. 1. 2831 ・35 ，6:39.0 ・71.1 mm ，Assam ， F. Day; BMNH 1889.9.26.1 ・9，9:37.6-45.0 mm ，Deoli ，Rajputana ，Co l. Biddulph; MCZ 9716-17 ，

12:16.6-32.3 12:16.6-32.3 mm ，Ghugger R.， M. M. Carlton; CAS 79070 ， 364: 13.1 幽 59.8 mm ，riverside pools pools and fish ponds ne 紅 Sonepur ，Mahanadi basin ， Orissa state ，20 ・21 Feb. 1985 ， T. R. Roberts; Roberts; BMNH 1938.2.22.129 ・131 ，3: 40.2 ・41. 3 mm ，Godavery R. headwaters ，Deolali ， Nasik Nasik dis t.， Bombay Pr esidency ，no date ，Bombay Na t. His t. Soc.; CAS 79071 ， 7: 40.8-

52.2 52.2 mm ， Tungabahdra Ri ver at H 創 npi ，Kri shna basin ， Karnataka state ，3 Feb. 1985 ，T. R. Roberts Roberts

Diagnosis. Diagnosis. - Same as for genus.

Comments. - As with all of Hamilton's fishes ，Chanda nama was described on 白e basis of of a drawing. Chanda phula and C. bogoda ，however ， were described by Hamilton without without the benefit of his original drawings of these two nominal species ，which ， with other other Hamilton drawings ，had been retained by the Govemment of Bengal (HORA ， 1929: 171).

Th us there are no holotypes for these three nominal species. Chanda phula ， C. bogoda 佃 d A. A. oblonga have been regarded as ju 凶or synonyms of Chanda nama (DAY ， 1875: 50; FRASER-BRUNNER ， 1955: 209). Day discussed the synonymy of C. phula and C. bogoda with with C. nama ，compar 担g Hamilton's original descriptions of the nominal taxa in some detail ， but apparently neither Day nor Fraser-Brunner saw Hamilton's original drawings of C. C. bogoda and C. phula ， nor did they examine the type material of Valenciennes's A. oblonga. oblonga. Hamilton's drawings of C. phula 組 d C. bogoda have been published in HORA ， 1929 ， p l. 20 ，figs. 1 姐 d 3. Having examined these figures and also the holotype of A. oblonga ，I confirm the synonymy given by Day.

Di stribution. - Chanda nama definitely occurs in the In dus ，Ganges ，Mahanadi ， and Kri shna ， and perhaps other river basins in India. Records from Burma probably e町oneous. SYSTEMA Tl C REVISION OF TROPICAL ASIAN FRESHWATER GLASSPERCHES 267

Records Records from Malaya erroneous.

Gymnochanda Fraser-Brunner ， 1955

Gymnochanda Fraser-Brunner ，1955:210 ，fig. 4 (type species Gymnochanda filamentosa Fraser-Brunner ，1955 ，by original designation and monotypy). Gymnochanda Boeseman ，1957:75 ，pl. 3 (type species Gymnochandafilamentosa Boeseman 1957 ，by original designation and monotypy).

Comment on name. - Marinus Boeseman informed me some years ago 伽 t he had quite independently independently of Fraser-Brunner chosen the same n釘 ne for his new fish. Given the most obvious obvious distinctive characters of the fish ，it is readily understandable how this could happen. happen.

Di agnosis. - Gymnochanda differs from all other ambassids in having body entirely scaleless ， and first four dorsal and anal fin rays of males greatly elongated or filamentous. Humeral spot absen t. Vertebrae 8+17 = 25. Two species ， one from the southem Malay peninsula 佃 d westem Bomeo (Kalimantan Barat) ， the other recently discovered in sou 白- em Bomeo (Kalimantan Tengah).

Gymnochanda filamentosa Fraser-Brunner ， 1955

Gymnochanda filamentosa Fraser-Brunner ， 1955: 210 ，fig. 4 (type locality “ southem Malaya"; Malaya"; aquarium specimens). Gymnochandafilamentosa Gymnochandafilamentosa Boeseman ， 1957: 75 ，pl. 3 (type locality “Singapore?"; aqu 釘 ium specimens). specimens).

Type material examined.-BMNH 1992.1 1. 23.1-6 ， 6: 14.5-18.5 mm (all males) ， “Sou 由 Malaya: presented by Cap t. M. A. W. Oavies and Raffles Museum" (types of G. filamentosa filamentosa Fraser-Brunner); RMNH 20902 ， 28.5 mm ，Singapore? (holotype G. filamentosa Boeseman); Boeseman); RMNH 20888 ，# 0: 13.1 ・25.6 mm ， Singapore? (paratypes G. filamentosa Boeseman). Boeseman).

Non-type Non-type material examined. - ZRCS 308 ， 4: 18.7 ・23.0 mm ，Johore ，Kl uang ，March 1955 ，H. K. Tong; CAS 49441 ，8:23.4-29.1 mm ， Sungai Melawi and mouth of Sungai Pinoh Pinoh at Nangapinoh ， Kapuas basin ， 21-26 July 1976 ， T. R. Roberts & Soetikno.

Di agnosis. - Gymnochanda filamentosa differs from G. flame a， new species ，its only congener ， mainly in coloration and size. Body 仕組sparent or translucent yellowish or yellowish-brown yellowish-brown in life ，filamentous rays of sexually mat 町 e males milk-white distally (ROBERTS ， 1989: 160); to 32 mm. According According to the original description by Fraser-Brunner ， the filamentous fins of G. filamentosa filamentosa are sexually dimorphic ，characteristic of the males only. 百lI s finding was 268 TYSON R. ROBER' 百

based based 'O n the 'O bservati 'O ns 'O f living fish by a Cap t. M. A. W. Davis ，including the c'O mment 血 at “the males displayed t 'O the females and t 'O each 'O ther by vibrating and spreading the unusual unusual rays 'O f the d 'O rsal and anal fins" (in FRAsER-BRUNNER ， 1955: 212).

Distribution. Distribution. - Gymnochanda filamentosa is kn 'O wn 'O nly fr 'O m the s'O uthernm 'O st p制'O f Malay penins 叫a (J 'O h'O re) and Kapu 槌 basin 'O f westem B 'O me 'O (Kalimantan Barat).

Gymnochanda flamea new species Figure Figure 1

Syntypes. Syntypes. - ZRCS 22861 ・66 ， 6: 14 .4 -18.8 mm ，MNHN 1992.1055 ， 78: 11 ふ 22.5 mm ，

Danau Sembuluh at B 佃 gkal ， Seruyan basin ， Kalimantan Teng 油， 12 ・13 June 1992 ， T.R. R 'O berts (syntypes sent t 'O MZB 組 d USNM fr 'O m MNHN series).

Di a伊 .osis. - Gymnochanda flamea differs 企om its 'O nly kn 'O wn c'O ngener ， G. filamentosa ， mainly mainly in c'O l'O rati 'O n and size. Live fish 紅 etr 叩 sp 紅 'ent 'O r translucent ，wi 血 little 'O rn 'O yell 'O wish tint; males have pelvic fms and filament 'O us extensi 'O ns 'O fd 'O rsal and anal fms red; red; c'O ncentrati 'O ns 'O f red chr 'O mat 'O ph 'O res als 'O 'O n sn 'O ut ，pred 'O rsum ，and spin 'O us d 'O rsal fin; fin; b'O dy depth 2.3-2.7; largest specimen (male) 'O nly 22.5 mm (vs 32 mm f'O r G. filamentosa); filamentosa); largest female 16.9 mm.

Sexual Sexual dimorphism. - Th e MNHN series 'O f 78 specimens represents ap 'O pulati 'O n sam- ple ple 'O f breeding fish. It c 'O nsists 'O f 39 males ，13.1 ・22.5 mm; 23 females ， 13.3 ・16.9 mm; 組 d 16 y'O ung ， sex undeterr 国ned ，I1. 6・14.0mm.τ 'h e 121 釘 gest specimens ，18.1-22 .5 mm ， 釘 'e all males with l'O ng filament 'O us extensi 'O ns 'O fb 'O血 d'O rsal and anal fms.τ 'h ey have up t 'O 5 町 6 thin vertical bars 'O n the b'O dy and bright red c'O l'O rati 'O n in 1百e. Al l males with filaments filaments 'O nb 'O th fins. Al l females app 紅 'ently gravid 'O r with ripening g'O nads; largest females females gravid 'O r at least with well devel 'O ped 'O varies: presence 'O f eggs veritied by dis- secti 'O n in ten specimens. A female 'O f 16.5 mm with 15 eggs 'O f ab 'O ut 0.6 mm and a few much smaller eggs. Females and y'O ung with 'O ut filament 'O us fms ， largely 仕組sparent ， with 'O ut faint vertical bars 'O r red c'O I'O r.

Food habits. - Dennis H. Murphy 'O f the Zo'O l'O gy Depar 卸 lent ，Nati 'O nal U 凶versi ザ'O f Singap 'O re ，組ndly examined st 'O mach c 'O ntents 'O ffive G.flamea ，17.7-19.8 mm ，合'O mDanau Sembuluh. Sembuluh. He f'O und 白ey had been feeding mainly 'O n midwater animals ，pred 'O min 阻 tly Cladocera Cladocera 'O f several species (present in st 'O machs 'O f all five). One had fed 'O n midwater c'O pep 'O ds (cal 姐'O ids) ， several 'O n midwater sw 加 m 泊g Hemiptera (c 'O rixids and pleids). Th e 'O nly 'O ther f'O'O d items f'O und were a single t'O p-fl 'O a出 g dipteran and a single b'O tt 'O m- living living mayfly. F 'O'O d habits 'O f G. filamentosa in na 何回 unkn 'O wn.

Distribution. Distribution. - Gymnochanda flamea is kn 'O wn 'O nly fr 'O m 由e type l'O cality ， Danau Sembuluh ， Seruyan basin ，s 'O uth B 'O me 'O. Thi s is a large ，shall 'O w ，dendritic lake ，with 'O ut tidal tidal fluctuati 'O ns; water slightly murky but definitely n'O t black; pH n'O t rec 'O rded. SYSTEMAT IC REV ISION OF TROP ICAL AS IAN FRESHW ATER GLASS PER C H ES 269

Figlll 巴 J. GYlll l1 ocha l1 da f7 alllea ， Danall Sembll JlI h (sy nt yp e)

Figllre Figllre 2. Paradoxoda cl1 a pirati ca ， Danall Sembll Joh 270 T YSON R. R OBERTS

Figure 3. Para111bassis apogonoides. 85 .1 111111 , Stung Treng.

Figure 4 . Para 111bassis /ala, 26 111111 , Mandalay. SYSTEMATIC REVISION OF TROPICAL ASIAN FRESHWATER GLASSPERCHES 271

Paradoxodacna Roberts ，1989

Paradoxodacna Roberts ， 1989: 160 (type species P. piratica Roberts ，1989 ，by original designation 叩 d monotypy).

Diagnosis Diagnosis (modified from ROBERTS ， 1989: 159-60). - A scale-eating ambassid with retrogna 出ous jaws (l ower jaw shorter 白叩 upper); dentition highly specialized ，different from that of all other ambassids except the markedly prognathous genus Chanda ，which it it superficially resembles (ROBERTS ，1989 ，fig. 125). Posterior nostril round ， about s創 ne size size as anterior nostril; gill rakers elongate ，5・ 7+15-16 = 20 ・23; cheek ，gill cover ，and body including including predorsum covered with scales; scales in lateral series 62 ・65; transverse scale rows rows 8-9/1: 20 ・22; dorsal fin soft rays 9・10 ， anal 9; pectoral fin rays 14 ・15; no humeral m 紅 k; vertebrae 10+14=24. One species known.

Paradoxodacna piratica Roberts ，1989 Figure Figure 2

Paradoxodacna piratica Roberts ，1989:160 ，figs. 125 ・6 (type locality Kapuas basin ， Kalimantan Kalimantan Barat (westem Bomeo).

Di agnosis. - Same as for genus. Type material examined. - ZMA 112.376 ， 54.9 mm ，Mendalan rivier ，28 June 1908 ，H. A. Lorentz Lorentz (paratype).

Non-type material examined. 一 MNI 町 1992;1056 ，24: 17.9 田 40.1 mm ，Danau Sembul 曲， Sung 泊 Seruyan basin ， Kalimantan Tengah ， 12-13 June 1992 ， T. R. Roberts; ZMA 120.828 ， 12: 12: 52.5-75.5 mm ，Djambi ， Batang H 紅 i， 23 April 1909 ， P. E. Moolenburgh; BMNH 1959 .4 .23.19-20 ， 2: 56.9-68.7 mm ，Djambi ，Sumatra ， Mooloenburgh.

Di stribution. - Pr eviously known only from Kapuas basin in westem Bomeo ， P. piratica now is reported also from Seruyan basin of southem Bomeo (Kalimantan Tengah) and from Sumatra.

Parambass お Bleeker ，1874

Parambassis Parambassis Bleeker 1874: 86 (type species Ambassis apogonoides Bleeker ，1851 ，by original original designation). Pseudambassis Pseudambassis Bleeker ， 1874: 86 (type species Chanda lala Har 凶It on ，1822 ，by original designation designation and monotypy). Acanthoperca Acanthoperca Castelnau ， 1878: 44 (type species A. gulliveri Castelnau ，1878 ，by monotypy). Whitleyina Whitleyina Fowler & Be 肌 1930: vii ，163 ，334 (type species Ambassis wo 伊 Bleeker 1850 ， by monotypy).

Diagnosis. Diagnosis. - Parambassis comprises a diverse and possibly polyphyletic 釦 ay of 姐 ictly 272 TYSON R. ROBER 百

freshwater freshwater ambassids widely dis 仕ibuted in the Indian subcontinent ， mainland and insular Southeast Southeast Asia ， and the Australian region including New Guinea. They differ from all marine marine ambassids in having straight or only slightly uptumed rather than strongly uptumed jaws. jaws. Most also have much smaller scales. Scales in lateral series from 30 to about 100. Dorsal Dorsal fin rays 9-16 ， anal 9-17. Vertebrae 10+14 = 24 or 10+15 = 25. Preorbital bone with with serrations on ridge and edge. Despite ， or indeed because of ，diversity of body shape and meristic characters in Parambassis ，it is difficult to split 出is genus into meaningful smaller genera. 百le type species species of the three nominal genera here placed as junior synonyms of Parambassis Bleeker are are all quite distinct from P. apogonoides (type species of Parambassis) ， and may well represent represent good genera. But were these genera to be recognized at the present time it would be difficult or impossible to assign the rest of the known species to them with 組 y con- fidence fidence in the results. Parambassis can be divided into species with short dorsal and anal fins fins (with only 9-11 soft rays ，usually without humeral spo t) and long dorsal and anal fins

(12 ・ 17 soft rays ， usually with humeral spot) but these probably are not monophyletic groups. groups.

Parambass お apogonoides (Bleeker ， 1851) Figure Figure 3

Ambassis Ambassis apogonoides Bleeker ， 1851b:200 (type locality Banjermassing). Parambassis Parambassis apogonoides ，Bleeker ， 1874:103. Chanda apogonoides ，Fraser 品 unner ， 1955:204.

Type materiaI examined. - None.

Non 吻 pe materiaI examined. - LAOS: ZRCS 35117-18 ， 2: 33.6-38.3 mm ，Mekong R. below Lee Pee Waterfalls ，June-July 1993 ， T. R. Roberts; CAMBODIA: CAS 81541 ， 85.1 mm ， Stung Treng market ，Jan.-Feb. 1994 ， T. R. Roberts; MNHN 1992.1062 ，4:4 1. 5-55.2 mm ，Days km 9， Tonle Sap ， 31 Jan. 1961 ， F. d'Aubenton; MALAY PENINSULA: ZRCS 19603 ，78 .5 mm ，Tasek Bera ，Pahang ， June 1940 ，M. W. Tweedie; ZRCS 1313 ， 65.8 mm ， Lake Chini ，Pahang ，1967 ， 205th Squadron RAF Changi; BORNEO: MNHN 1982-717 ， 36.9 36.9 mm ， Sungai Engkayas ，Kapuas basin ， 16 July 1976 ， T. R. Roberts; ZMA 112.309 ， 62 .3 mm ，Mendalan rivier ，28 June 1908 ， H. A. Lorentz; SUMATRA: BMNH 1959 .4 .23.24- 25 ，2:42.8-72.9 mm ，Djambi ， Moolenburgh; ZMA 112.297 ， 67.7 mm ，Djambi ，1909 ， P.E. Moolenburgh. Moolenburgh.

Di agnosis (modified 合'O m ROBERTS ， 1989: 162). - Dorsal profile moderately sloping ，gently concave; concave; preorbital ridge smooth; orbital rim with 2-3 small seηae posteriorly ， otherwise smooth; smooth; upper limb of preopercular margin seπate; gill rakers 21-22; scales in lateral series series 40 ・47; cheek with 5 scale rows; predorsum entirely covered with scales; dorsal and anal anal fin branched rays 10-11; second anal fin spine only slightly larger 出印刷rd; pectoral fin fin rays 12-16; no humeral mark; vertebrae 10+14 = 24. SYSTEMATIC REVISION OF TROPICAL ASIAN FRESHWATER GLASSPERCHES 273

Serration Serration of 36.9 mm specimen from Kapuas. - Preorbital edge with 10 serrae ， ridge non-

se 汀 ate; suborbital and supraorbital without seηae; preopercular ridge with 4 large se 町 ae on horizontallimb ，no se 町 ae on verticallimb; preopercular edge with 10 spines on horizontal limb ，1 large and 19 small 田町田 on vertical limb. Specimens from lower Mekong (MNI 到 1992.1062) 叩 d Kapuas (MNHN 1982.717) have been comp 紅巴d directly 叩 d 由ey have very sirnilar serration.

Oral brooding. - Oral brooding of just over 100 young of 3-7 mm by a5 1. 6 mm P. apogonoides apogonoides is reported by ROBERTS ， 1989:162. There is no additional report of oral brooding brooding in 出is species or in any other member of the family Ambassidae.

Distribution. Distribution. - Pr eviously known only Sumatra and Borneo ， P. apogonoides is now re- corded corded from the Malay peninsula (Pahang basin) and from the Lower Mekong basin (southern (southern Laos below Lee Pee Waterfalls; Stung Treng ，Cambodia; and To n1 e Sap ，Cam- bodia). bodia).

Parambassis Parambassis baculis (Hamilton ， 1822)

Ch αnda baculis Hamilton ， 1822: 112 ， 371 (type localit 旬y ‘“‘ nor 此th- 屯ea 郁st 包ern p釘 t岱so ぱf Benga 必1") Amηlb 如assi おsb拘ac ωu叫l的iおs， Cuvier in Cuvier & Valenciennes ， 1828: 187; Day ， 1875: 51 ， p l. 15 ， fig. fig. 1). Parambassis Parambassis baculis ，Kottelat ， 1989: 17. Pseudambassis Pseudambassis baculis ，Talwar & Jhingran ， 1992: 803.

Type material examined. - None.

Non-type material examined. - CAS 24250 ，9:2 1. 8・30.8 mm ， Indus R. ne 紅 Sukkur ，4 Jan. Jan. 1969 ， E. S. Herald; MCZ 9711 ， 23.8 mm ，and MCZ 9712 ，3:16.8-20.5 mm ， Sursutiu R.， R.， N India ，M. M. Carlton; MCZ 9715 ， 24.6 mm ，Ghugger R.， M. M. Carlton; MNHN 9359 ， 35.0 mm ，Indoustan ， F. Day; BMNH 1889.2. 1. 2858 ， 34.8 mm ，Gowhatty ， F. Day; CAS ・SU 39980 ， 30.6 mm ，Kampong Duars & Siliguri Terai ， Tista R. drainage ，Nov. 1938 ， S. S. L. Hora; CAS 50345 ， 37.0 mm ， Naraini R. 10 rni W of Narangar ，26 April 1975 ，T .R.

Roberts; Roberts; CAS 50274 ，3:26.2 ・27.6 mm ，Naraini R. 4 rni E of Kh oria Mohan (b elow confluence with with Rapti R.)， 24 April 1975 ，T .R. Roberts; CAS 50188 ， 2: 30 ふ 35.3 mm ， Rapti R. and tributaries 11 tributaries rni SSE of Narang 紅， April 1975 ， T. R. Roberts.

Diagnosis. Diagnosis. - Parambassis baculis evidently is a sma l1 species (l argest specimen ex 創 ll- ined ined under 40 mm); body moderately elongate ， a characteristic shared among Asian freshwater freshwater Ambassidae only with th 巴 specialized scale-eaters Chanda nama and Paradoxod α， cna piratica; head shape different from all other freshwater Ambassidae; snout gently gently rounded ，jaws short and equal ，叩 d nape distinctly concave; gill rakers on first gill arch arch short and stout ， only 2-4+9 ・13 = 12 ・15; scales ve 町 small ，lateral series about 80-100; dorsal dorsal and anal fins with 13 ・14 branched rays ，pectoral fin rays 13; vertebrae 10+14=24. Serration Serration of 35.0 mm specimen from Indoustan. - Preorbit 叫 edge with 3 sma l1 upper 274 TYSON R. ROBBR' 首

and and 31arge lower se 町ae ，ridge wi 出 1紅 ge exposed primary se 町a plus 41 紅 'ge lower spioes; 00 suborbital se 町 ae; supraorbital with a short portioo without serrae anteriorly ， followed by 6 small serrae beginniog over middle of eye ， theo 31 ぽ ge se 町 ae posteriorly; ridge of preopercle preopercle ooo-serrate; horizootal limb of preopercle with 13 large serrae ，1制 just above angle; angle; vertical limb without serrae; an isolated serra above preopercle (posterior to last large large supraorbital serra; 00 se 町ae 00 exposed po 氏ioo of shoulder girdle.

Comment. 一Hamiltoo's drawiog of C. baculis has beeo published io HORA ，1929 ， pl. 20 ， fig. fig. 7.τ 'h is is oot so good as his other drawiogs of Am bassidae ， and less help 釦1 io ideotifyiog 由e species. 百le figure of C. baculis io DAY ， 1875 is good ， and leaves 00 doubt doubt as to which species he thought Hamiltoo had described. A lectotype should be desigoated ， but I prefer to wait uotil more material becomes available and the species c叩 be studied more c紅 'efully.

Distribution. Distribution. - Parambassis baculis is knowo only from the lodus and Ganges basios.

Paramb ωs お dayi (B leeker ， 1874)

Ambassis nalua ， Buch. Ham.? ，Day ， 1865a:12 (Cochin). Ambassis nalua ，Day ， 1865b:15 (p 釘 tim). Ambassis dayi Bleeker ， 1874:95 (type locality Cochio; based 00 Day's descriptioo of 'Ambassis ‘'Ambassis nalua ， Buch. Ham.?"); Day ， 1875: 54 ， p l. 15 ，fig. 7; Fraser-Bruooer ， 1955:198. 1955:198. Amb 卸assis ，悶v'alen 町z叱ciennii Va 剖aill 組.1， 1875: 34 (t 旬yp 戸e locali 句ザ “"ea 削ux 白.od 即u.ce 邸sd也e Ali 均pa 可y" 勺) Param ηtbassi 臼's ， da の:y 日li ，τT ‘'alwa 紅r& 拍nti 泊ogr ，姐， 1992: 80 1.

Type material examined. - MNl王 N 9351 ，3:107-125 mm ，A1i pay [port 00 Malabar coast of of lodia ， south of Cochio; also spelled Al epay ，Al eが， Al leppi] ，1835 ，J. J. Dussumier (s 戸ltypes A. valenciennesi).

Diagnosis. Diagnosis. - Mouth eloogate and horizootal ，moreso 白血泊佃y other member of the far 凶ly Am bassidae with possible exceptioo of P. thomasi; gill rakers 00 frrst gill arch 27 ・ 28; 28; scales large ， very deciduous ， predorsum eotirely covered with scales; lateral scale series series 34 ・39 ，佐'ansverse series 5/1114; anal fm scale sheath with only a single row of scales; scales; dorsal 岨 d 組 al fio soft rays 10-12; 00 humeral spot; vertebrae 10+14 = 24 (types of of A. valenciennesi).

Se 町 ation of 107 mm sp 即加.en from Alipay. - Pre orbital edge with 18 serrae of about equal equal size; ridge with 8 small ， poorly developed serrae; suborbital ridge with about 10 small small weak serrae; preopercular edge with 10 s位oog se 町ae 00 horizootal limb ，4 stroog spioes spioes 00 verticallimb just above angle ，血 eo rest of edge smooth; preopercular ridge with

18 18 equally 1釘 ge serrae 00 v即位al limb ，4 00 vertical limb just above angle ， and rest of verticallimb verticallimb with 37 very fioe ， evenly sp 釦 ed serrae; ioteropercle wi 白 9 serrae; supraorbital eotire eotire except for 2 stroog se 町'ae at posteriormost poiot; exposed shoulder girdle without SYSTEMA TIC REVISION OF TROPICAL ASIAN FRESHWA'τ 芭R GLASSPERCHES 275

serrae; serrae; s住'O ng posteriorly directed serra on skull bone just above articulation of preopercle.

Color Color of fresh specimens. ー“ Back greyish green with minute brown points ，abdomen silvery ， with a silver longitudinal band ， and having pinkish reflections; opercles silvery ， fins fins greyish ，frrst and second spine of frrst dorsal with a blackish outer m 紅 'gin; fine grey points points on fin membrane more especially between second and 也ird spines. Caudal minutely spotted. spotted. Eye silvery" (DAY ， 1865b: 16).

Comments. - Fraser-Brunner did not examine material of A. dayi ，姐 d did not mention A. A. valenciennii (for which the correct spelling is A. valenciennes i). Parambassis dayi is mo 中hologically most similar to P. thomasi ，but 血ey are certainly different species. Parambassis Parambassis dayi h出 a more elongate body ， and slightly more gi1l rakers.

Distribution. Distribution. - Parambassis dayi occurs in several SW Indian coastal rivers.

Parambassis Parambassis 1a1a (Hamilton ， 1822) new comb. Figure Figure 4

Chanda lala Har 凶lton ， 1822: 114 ，371 ，p1. 16 ，fig. 39 (type locality fresh waters of Gangetic provinces; provinces; description based only on males). Ambassis Ambassis alta 印刷im) Cuvier in Cuvier & Valenciennes ， 1828: 183 (type locality Bengale). Ambassis Ambassis lala ， Cuvier in Cuvier & Valenciennes ， 1828: 184 (after Hamilton ， 1822). Pseudambassis Pseudambassis lala ，Bleeker ， 1874: 86; Talwar and Jh ingran ， 1992: 804.

Typ e material examined. 一MNl王 N 9341 ，3: 23.7 ・28.9 mm ，Bengale ， Duvaucel (syntypes A. A. alta).

Non-type material examined. - MCZ 60878 ， 25.3 mm ， Sind Tatta ，SE Karachi ， 19 July 1955 ，Peabody Mus. Exped.; MCZ 9714 ， 3: 17.2-20.0 mm ，Su t1 ej R. at Loodiana ，M. M. Carlton; Carlton; MNHN 9342 ，5:16 .4 -2 1. 8 mm ，Calcutta ， Dussumier; MNHN unca t. (ex MNI 町 9356) ，4:2 1. 0・25.6 mm ，Bengale ， Day; BMNH 1934.10.17.97 ・99 ， 3: 19.0-22.9 mm ， Am adpur ，Bengal ， Das; CAS-SU 33338 ，16:13.9-38.2 mm ，Pu lta ，Ganges delta ， 10 April 1937 ， A. W. He 町'e; CAS-SU 33335 ， 2: 19.4-26.3 mm ，Utt 釘 bag ，Ganges delta ， 14 April 1937 ， A. W. Herre; CAS 50374 ， 8: 20 ふ 26.0 mm ，Kh ailaya market ，Terai ，Nepal ，1 May 1975 ， T. R. Roberts; CAS 79072 ， 132: 11. 0-28.2 mm ，riverside pools and fish ponds ne 紅

Sonepur ，Mahanadi basin ， Orissa state ，20 ・21 Feb. 1985 ， T. R. Roberts; CAS 81542 ， 96: 13.6-28.0 13.6-28.0 mm ，Mandalay market ，13 ・25 April 1993 ， T. R. Roberts; CAS 81543 ， 3: 20.0- 25.5 25.5 mm ，swampy s釘eam in rice paddy on western outskirt of Mandalay ，20 Apri11993 ， T. R. Roberts.

Di agnosis. - Parambassis lala is distinguished from all other Asian freshwater amb 部 sids by having males with very distinctive coloration of body and fins and from all 0 白ers except except Gymnochanda by very small adult size. Sexually active males with a single large ctenus ctenus projecting from each scale on body above anal fin base 佃 d on scaly shea 血 of anal fin. fin. 276 TYSON R. ROBERTS

Gill Gill rakers on first gill arch 16 ・18; scales in lateral series 66; dorsal fin branched rays 13 ・15 ，組al 14 ・17; pectoral fin rays only 10-12; scales in lateral series 66; supraoccipital portion portion of predorum without scales; humeral spot present; vertebrae 10+14 = 24 (10+15 = 25 in most other Indian f回shwater Am bassidae). An al fin scale shea 出 with up to 5 rows of scales. In males scales of anal fin sh 伺白 血 d adjacent p制 of body each wi 由 a single posteriorly projecting ctenus or contact org 組 (sensu (sensu WILEY & COLLET 四， 1970; COLL 回百， 1977); contact organs otherwise unknown 加Am bassidae. Preorbital Preorbital edge with 7 moderately large serrae ， ridge with 3 small downwardly di- rected rected se 町 ae (no prim 紅 y or major spine on dorsal p制 of ridge); supraorbital with anterior half half smooth ，posterior half with 6 small spines; suborbital without se 町 ae; preopercular ridge ridge horizontal limb with 9 moderately 1紅 ge serrae ，1 serra just above angle ，otherwise verticallimb verticallimb non-serrate; preopercular edge with 12 nearly small equal serrae on horizontal portion ，vertical portion with 2 small 田町田just above angle and 7 very small serrae on upper upper half; no serrae on interopercle or shoulder girdle.

Coloration. Coloration. - Li ve fish translucent greenish ，yellow ， or even orangish. Adult males with two broad ，vertical dark bands ，composed of numerous fine and close-set melanophores ， clearly clearly visible even in discolored specimens 150 years old; distal portion of first and second second dorsal fins and entire pelvic fins black.

Discussion. Discussion. - Although cle 釘 ly distinguished and well fig 町 'ed by HAMILTON ，1822 ，白 is very very distinctive but tiny species w 剖 placed 部 a synonym of A. ranga by DAY ， 1875: 51 and left there by FRASER-BRUNNER ， 1955. And whereas Cuvier (in CUVIER & VALENCIE 附 ES ， 1828) recognized Chanda lala as valid on 白.e basis of 悦 original de- scription scription by Hamilton ， he failed to recog 凶ze 白e species in his own material. Cuvier utilized utilized two samples collected by Duvaucel in B 阻 gale for the description of Ambassis alta. One sample ，MNI 王N 9342 ， comprises a 23.7-mm sexually ma 加re male and two gravid females ，28 ふ 28.9 mm ， of Parambassis lala. 百le 血ree specimens have only 10 ・12 pec- toral toral fm rays ， anal fin sheath with up to 5 scale rows ， and 10+14 = 24 vertebrae. Th e other sample sample used by Cuvier to describe A. alta ，MNI 王N 9340 ，3:35. 4- 48.3 mm ，is of Parambassis ranga. ranga. Until 恥 freshwater ambassids of Asia have been studied more thoroughly ，I prefer a conservative approach to their generic classification. Th us Pseudoambassis is not recognized recognized as a valid genus here ，even 白ough its type species ， P. lala ， is distinctive and may merit generic sep 釘 ation 針。 m the other species here assigned to Parambassis.

Distribution. Distribution. - Parambassis lala is known from lowlands of the Ganges-Brahmaputra and and Mahanadi basins in In dia and the Irr awaddy basin in Myanmar.

Parambassis Parambassis macrolepis Bleeker ， 1857 Figure Figure 5

Bogoda macrolepis Bleeker ， 1857: 13 (type locality Pontianak ，Kapuas R ふ SYS T E M A T IC RE VI SION OF T ROP ICA L AS IAN FRES HW A T ER G LASSPE R C H ES 277

Fig ul 巴 5. Parambass is m acrolep is，72 111m ，D anau Se l11 bul oh

Figur e 6. Para mb assis I百 Il ga ， 64.5 mm ，Lake In le 278 TYSON R. ROBERTS

Figure 7. Parambassis siamensis, Toni e Sap.

Figure 8. Parambassis tenasserimeiiSis, 65.7 mm, Tapoleh Kloh (holotype). SYSTEMA TIC REVISION OF TROPICAL ASIAN FRESHW A TER GLASSPERCHES 279

Ambassis Ambassis (Bogoda) microlepis von Martens ， 1868: 9 (Kapuas; unwarranted spelling emen- dation). dation). Parambassis Parambassis microlepis ，Bleeker ， 1874: 104. Ambassis Ambassis microlepis ，Vaillant ， 1893: 110 (Kapuas). Chanda macrolepis ，Fraser-Brunner ， 1955: 205. Parambassis Parambassis macrolepis ，Roberts ， 1989: 162 ，fig. 128 (Kapuas).

Type material examined. - None.

Non-type material examined. - ZMA 112 .3 77 ， 3: 66.5 ・72.3 mm ，Boenoet ，Bomeo ，26

June June 1909 ，H. A. Lorentz; MNHN 1992.1057 ， 5: 60.8 明 78.0 mm ，Danau Sembuluh ， Sungai Seruyan Seruyan basin ， Kalimantan Tengah ， 12-13 June 1992 ，T. R. Roberts; BMNH 1959 .4 .23.18 ， 74.2 74.2 mm ，Djambi ，Sumatra ， Moolenburgh.

Diagnosis Diagnosis (modified from ROBERTS ， 1989: 162). - Body moderately deep; dorsal profile strongly strongly concave ，moreso than in any other Parambassis; preorbital ridge without seπae; gill gill rakers on first gill arch 6-7+11-12 = 17-18; scales in lateral series 50 ・59; cheek with 13 ・15 scale rows; predorsum entirely covered with scales; predorsal scales more than 30; transverse transverse scales rows 9/1130; anal fin scale sheath with up to 3 or 4 scale rows; dorsal and anal fin soft rays 9・ 11; pectoral fin rays 13 ・ 15; no humeral spot; vertebrae 10+14=24(5) (Sembuluh). (Sembuluh).

Serration Serration (sample from Sembuluh). - Preorbital edge with 10 well developed serrae ， ridge ridge without any 田町田; suborbital nor トserrate; supraorbital non 開田町ate except for small terminal terminal 田町a; preopercul 訂 edge horizontal limb 12 se 町ae and 2 ne 訂叩gle ，vertical limb otherwise otherwise non-serrate; preopercular ridge horizontal and vertical limb non-serrate except for for 2 large 田町田 near angle; interopercle 7 weak se 町 ae; shoulder girdle non- 田町ate.

Food habits. - Stomach contents of 72-mm specimen from Danau Sembuluh consist of three three Rasbora-like cyprinid fishes 15 ・20 mm standard length.

Di stribution. - Parambassis macrolepis is known only from Sumatra (Batang Hari) 組 d Bomeo (Kapuas in Kalimantan Barat; Seruyan in Kalimantan Tengah).

Parambassis Parambassis ranga (Hamilton ， 1822) Figure Figure 6

Chanda ranga Hamilton ， 1822: 113 ，371 ， p l. 16 ，fig. 38 (type locality “ fresh waters of all parts parts of 血e Gangetic provinces"). Ambassis Ambassis ranga ， Cuvier in Cuv. & Va l.， 1828: 183; Day ， 1878: 51 ， p l. Ambassis Ambassis barlovi Sykes ， 1841: 350 ， p l. 60 fig. 1 (type locality Beema River at Pairgaon). ?Ambassis ?Ambassis indica McClelland ， 1842: 585 ， p l. 4 fig. 1 (type locality Loodian 油). Ambassis Ambassis notatus Blyth ， 1860: 138 (type locality Sittang Ri ver at Pegu). Pseudambassis Pseudambassis notatus ，Guha & Talwar ， 1983: 15 ，fig. 1 (Bly 出 type specimens redesribed). 280 TYSON R. ROBERTS

Pseudambassis Pseudambassis ranga ， Talwar &拍 ingran ， 1992:805. Parambassis Parambassis ranga ，Kottelat ， 1989: 17.

Type material examined. 一MNl町 9340 ，3: 35 .4・ 48.3 mm ，Beng a1 e，Duvaucel (s 戸 types A. A. alta); BMNH unca t.， 2:34.7 ・4 1. 2 mm ，Dukhun ， Sykes (wi 白 printed label “Equula Bilineata Bilineata Dukhun Sykes" [unpublished name]; probable syntypes A. barlov i).

Non-type material exa 凶 ned.-PAKISTAN: ZMA 115.934 ， 2: 43.2 ・47.2 mm ， Indus R.，

Kasur dis t.， 1 March 1977 ，M. Saddiq; INDIA: MCZ 9710 ， 11: 30.5 ・35.6 mm ，Ghugger R. ，M.M. Carlton; BMNH 1889.2. 1. 2844-45 ， 2: 46.9-57.5 mm ，Assam ， F. Day; 阻畑町 9356 ， 50.7 mm ，Beng a1 e，Day; CAS 50308 ， 4: 37.7δ0.1 mm ， Rapti R. at Darbar ， 25 April

1975 ，T. R. Roberts; CAS 50352 ， 4: 43.1 ・55.6 mm ， Naraini R. 10 mi W of Narang 紅， 26 April April 1975 ，T. R. Roberts; CAS ・SU 39991 ， 43.0 mm ，Kalimpong Duars & Siliguri Terai ， Tista Tista R. drainage ，Nov. 1938 ， A. W. Herre; CAS 79077 ，37 .4 mm ，riverside pools 姐 d fish ponds ponds near Sonepur ，Mahanadi basin ， Orissa state ，20 ・21 Feb. 1985 ， T. R. Roberts; BMNH 1938.2.22.132-141 ， 33: 28.7-48.6 mm ，Darna R.， Deol a1 i， Nasik dis t.， Bombay prov. ，Bom- bay Natur a1 History Society; CAS 61947 ， 3: 23.6-37.0 mm ，Tungabahdra R. at Kampli ，2 Feb. Feb. 1985 ，T. R. Roberts; CAS 79076 ， 127: 17.2-63.7 mm ，Tungabahdra R. at Hampi ， Kri shna basin ， Karnataka state ，3 Feb. 1985 ，T .R. Roberts; CAS-SU 39992 ， 32.3 mm ， K a1 avacheda ，Vizagapatam dis t.， Madras pres. ，6 Sept 1939 ，Fisheries Research station at Vizagapatam; Vizagapatam; CAS-SU 39990 ， 35.9 mm ，Madras ，3 Jan. 1941 ， A. W. Herre; M Nl王 N 9350 ， 54.9 54.9 mm ，Coromandel ， Belanger; CAS-SU 33341 ， 7: 30.9 ・45.1 mm ，Cuddapah ， S. India ， April ，1937 ， A. W Herre; MYANMAR: AMNH 13795 ，Kaung Hein ，Vemay-Hopwood

Chindwin Exp.; ZMA 112 .4 08 ， 53.7 mm ，Irr awaddy at Bhamo; CAS uncat. ，11:24.2 ・67.6 mm ，Irr awaddy R. at Indaw ，Chaung and Kamaing ，27 March 1956 ， J.A. Tubb; MNl王 N 9343 ， 4: 38 .3 -44.1 mm ，Ir awadi ，1829 ， Reynaud; CAS 53385 ， 45.6 mm ， Saung-hka Chaung near near Kamaing ，Burma ，27 March 1956 ，J. A. Tubb; CAS 53383 ， 36.9 mm ，Mogaung Chaung at Mogaung ，Burma ，28 March 1956 ， J. A. Tubb; CAS unca t.， Lake Indawgyi ， upper upper Burma ，24 March 1956 ，J. A. Tubb; CAS 81660 ， 2: 54.8-64.5 mm ， Inle Lake ，5 March 1994 ， T. R. Roberts; ZMA 112.312 ，2: 46 ふ 55.0 mm ，Mand a1 ay market ，Ann an da1 e; MNHN 9344 ，2:46.7-48.2 mm ，Rangoon ，1829 ， Reynaud; CAS-SU 40233 ， 3: 25.0 ・44.0 mm ，Rangoon ，1940 ， G. E. Gates; CAS ・SU 33333 ， 3: 52.3 ・66.5 mm ，and CAS-SU 33339 ， 8:45.6-63.2 8:45.6-63.2 mm ，R 担 goon ， 30-31 March 1937 ，A. W. Herre; CAS 79078 ， 11: 34.7δ7.0 ， and CAS 79148 ， 12: 43.7 ・62.9 mm ，Rangoon market ，5 ・ 12 March 1985 ， T. Roberts;

BMNH 189 1.11. 30.9-18 ，27:37.6-6 1. 7 mm Sittang R.， E.W. Oates; CAS 79147 ， 5: 21. 6・ 55.0 55.0 mm ，Kh a-Yein Chaung ， at Wah Ki t Kh one vil1 age ， about 4 km NE of Hlegu on road to to Pegu ，Sittang basin ，9 March 1985 ， T. R. Roberts; THAILAND: CAS 79079-80 ， 5: 22.2 ・ 58 .3 mm ，S a1 ween R. mainstream near Mae Sam Leap ，百 lail 組 d， 21 April 1989 ， T. R. Roberts; Roberts; CAS 79081 ，3: 40.3-44.6 mm ，Menam Moei 129 km N of Mae Sot on highway 1085 ，Salween basin ，27 April 1991 ， T. R. Roberts; BMNH 1893.2.16.1-3 ， 3: 50.7-7 1. 5 mm ，Kokarit ，Sa1 ween basin ， L. F，回; BMNH 1889.2. 1. 2859-60 ， 2: 44.3 ・53.0 mm ，Moulmein ， F. F. Day.

Di agnosis. - A moderately deep bodied species; gil1 rakers 6・8+15 ・20 = 22-28; sc a1 es in later a1 series about 56-70; transverse sc a1 e rows 9・1111 /3 0・33; an a1 fin shea 血 with up to SYST 勘仏百C REVISION OF TROPICAL ASIAN FRESHW Aτ 'E R GLASSPERCHES 281

3 scale rows; dorsal fin soft rays 14-16 ，analI4-17; pectoral fin rays 12 ・ 13; humeral mark present; present; vertebrae 10+15 = 25.

Morphometrics. Morphometrics. - Morphome 凶cs of two specimens of P. ranga from Inle Lake are given in in Table 1. This sample was chosen for measurement because it appeared to be morphometrically morphometrically typical of many samples of P. ranga.

Serration. Serration. - 49.1-mm specimen from Bengale: Preorbital ridge with 7 moderately large serrae ， edge with very large exposed primary or major serra and 7 smaller serrae; suborbital with with 9 very small se 町'ae; supraorbital with 11 serrae ，largest posteriorly; preopercular ridge with with 8 serrae on horizontal limb and 1 just above angle ，vertical limb otherwise smooth; preopercul 釘 edge with 12 seπae on horizontal limb and 15 ・ 16 small se 町ae on vertical limb; limb; exposed p副 of shoulder girdle with 4 small upper and 2 or 3 very small lower se 町'ae.

Serration Serration of 54.9 ・mm specimen from Coromandel: Preorbital edge 8 se 町'ae (l ower 3 very very small) ， ridge large primary and 4 moderate 田町ae; suborbital non-se 町 ate; supraorbital smooth smooth anteriorly ，4 moderate seπae posteriorly; preopercular edge entirely smooth except for for 2 moderately large serrae ne 紅 angle; preopercul 紅 ridge with 13 equal sized se 町 ae on horizontal horizontal limb and 25 tiny serrae on vertical limb; interopercle no seηae; pectoral girdle with with 6 weak upper and 7 veηsmall lower seπae. Serration Serration of 52.9-mm specimen from Rangoon: Pr eorbital edge 8 large serrae ， ridge

large large prim 紅 y plus 3 moderately large se 町ae; pr ωpercular， edge horizontal limb 13 and verticallimb verticallimb 19 serrae ，preopercular ridge horizontallimb 4 seπae plus 1 at 組 gle ，vertical limb limb no serrae; interopercle no serrae; shoulder girdle feebly se 町 ate ，2 upper and 1 lower small small serrae.

Coloration. 一HAMILTON ， 1822:113-14 described color of live Gangetic P. ranga as follows: “… bright green and silver gloss. 百le whole ， except the gill covers ， which are silvered ，is diaphanous ， showing the muscles ，five ribs on each side the backbone ， a silvery

S住ipe accomp 佃 ying the latter ， and a silvery membrane 白at lines the cavity of the abdo ・ men. men. Th e upper parts 紅 e dotted with black... the scales on the body and fms are minute ， and and so transparent ，that 血ey can sc 紅白ly be seen without a magnifying glass: 白eyare marked with concen 凶 c lines." Specimens 合om the clear Lake Inle are particularly colo 凶Il (Fig. (Fig. 6). Samples from muddy water are much less color 白1.

Distribution. Distribution. - Parambassis ranga is widely dis 凶buted in Pakistan ，India ， and Myanmar ， including including but not limited to the Indus ，Ganges-Brahmaputra ，M 油 anadi ，Gadavery ，Kri shna ， Cauvery ，Irrawaddy ，Sittang ， and Salween river basins.

Parambassis Parambassis siamensis (F owler ， 1937) new comb. Figure Figure 7

Chanda siamensis Fowler ， 1937: 230 ，自 g. 228 (type locality B 組 gkok). Chanda punctulata Fraser-Brunner ， 1954: 206 ，fig. 3 (type locality Tasek Bera ， Pahang). 282 TYSON R. ROBER 百

Parambassis Parambassis punctulatus K 'O t旬lat ， 1989: 17.

Type material examined. - ANSP 68233 ， 45.2 mm ，Bangk 'O k，1936 ，R. M. deSchauensee (h 'O l'O type C. siamensis); ANSP 68234-41 ， 8:26.8-36.9 mm ，Bangk 'O k，1936 ， R. M. deSchauensee deSchauensee (p 紅 atypes C. siamensis); BMNH 193 1. 7.20.56-65 ，15:1 1. 8-34.7 mm ， Tasek Bera ，Pah 組 g，Dep t. Fisheries Singap 'O re (presumably 34.7 ・mmsp 田 imen is h'O l'O type and 'O ther 14 釘 e paratypes 'O f C. punc ωlata).

Non 吻 pe material examined. - MEKONG BASIN: CAS 79087 ，25:2 1. 7・42.9 mm ，Huay Nam Ngam and Mek 'O ng mains 田 am 1 km d'O wnriver 仕om 百lat Phan 'O m ，1J 姐. 1989 ， T. R. R. R 'O berts; CAS 79149 ， 14: 10.6 ・37.6 mm ，Si-Saket market ，NEτ 'h ailand ，29 March 1990 ， T. T. R. R 'O berts; CAS 79090 ， 18: 27.1 ・50.3 mm ，Ub'O I Ratchatani market ， 15 Dec. 1992 ， T. R. R. R 'O berts; CAS 79150 ，16:24.1-40.5 mm ，Menam D 'O m 'O n highway 2248 ， 17 km E 'O f Nam Yun ，ab 'O ut 100 km S 'O f Ub'O I Ratchatani ， 21 Dec. 1990 ， T. R. R 'O berts; CAS 79085 ， 5: 5: 17.1 ・40.1 mm ，dammed backwater 'O f Menam Chi at Agricult 町'a1 C 'O llege ，Maha Chana Chai ， 14 March 1991 ， T. R. R 'O berts; CAS 79084 ， 64: 23.1-38 .3 mm ，n 'O rthem end 'O f Sirinth 'O m Reserv 'O ir ， near Phibun Mansahan ， 14 Sep t. 1990 ， T. R. R 'O berts; MNl到 1927 .1 81 ・ 2， 20: 29 .5 -39 .4 mm ， Prek Phnan ，Cambodge ， 27 Dec. 1923 ， A. Kr empf; CHANTABURI BAS 町: CAS 79098 ， 49.6 mm ，1紅 ge river on highway 3245 ， 19 km fr 'O m juncti 'O n with highway 3， 11 Dec. 1990 ， T. R. Roberts; CHAO PHRAYA BASIN: CAS 79151 ， 3: 25.8 ・ 31. 6 mm ，irrigati 'O n can a1 ab 'O ut 6 km E 'O f Pi chit ，Cha 'O Phraya basin ，30 N 'O v. 1990 ， T. R. R. R 'O berts; CAS 79089 ， 6: 32.0-43.4 mm ，Nak 'O m Sawan market ， 4-5 Feb. 1989 ， T. R. R 'O berts; BANGPAKONG BASIN: CAS 79088 ， 3: 32.0 ・33.7 mm ，Kl'O ng Waeh Kam'O ng at at highway 304 ，9 km by r'O ad N 'O f junction with highway 33 at Kab inburi ，22 March 1989 ， T. R. R 'O berts; CAS 79152 ， 32: 20.5 ・31. 2 mm ，'O pen channel in 1紅 ge swamp ab 'O ut 9 km SSW 'O f Pr achinburi ，1 ・2 Dec. 1990 ， T. R. R 'O berts; MEKLONG BASIN: CAS 79092 ，Huai Kh a Kh aeng at Kh a 'O Bandai (near c 'O nfluence with Huay Mae Di); CAS 79091 ， 190: 13.0 ・33.6 mm ，Huay Kh a Kh aeng arm of Srinakarin Reserv 'O ir at Gr eung Grai ， Mekl 'O ng bas 泊， 22 J釦. 1991 ， T. R. R 'O berts; CAS 79153 ， 4: 17 .4 -25.1 mm ，Sangklaburi market ，Mekl 'O ng b部泊， 12 Feb. 1989 ， T. R. R 'O berts; CAS 79093 ， 3: 29.6-37.5 mm ，Huai M a1 ai ， 21 km WNW 'O f Sangklaburi ， 12 Feb. 1989 ， T. R. R 'O berts; CAS 79094 ，Huay Chee ， about about 25 km SW 'O f Kanchanaburi ， 23 Feb. 1990 ， T. R. R 'O berts; TAPI BASIN: CAS 79095 ， 23: 20.6-3 1.1 mm ，Kl'O ng Ipan ，'O n highway 40372 km S 'O f juncti 'O n with highway 4035 ，4 March 1989 ， T. R. R 'O berts; CAS 79154 ， 12: 26.6-34.8 mm ，CAS 79096 ， 13: 25.9- 35.5 35.5 mm ，Kl'O ng S'O k5 km W 'O f Phan 'O m on highway 410 ， 27 Feb. 1989 ， T. R. Roberts; PATTANI BASIN: CAS 79155 ， 45: 25.2 ・36 .4 mm ，CAS 79097 ， 36: 24.2-3 1.1 mm ，Pattani

R. m 泊nstream 佃 d 凶but 紅 y 35 km by road N of Betong on highway 410 ， 11 March 1989 ， T. T. R. Roberts; PAHANG BASIN: CAS-SU 30405 ， 2: 33.2-34.7 mm ，Tasek Bera ， 15 March 1934 ， A. W. Herre; ZRCS 9466 ・67 ，2:29.1 ・36.2 mm ，Lak e Chini ，no date ，205 Sqn

Changi; Changi; CAS 79062 ， 2: 37.7 ・40 .5 mm ， Tasek Chini ，no date ，Roy a1 Air Force Changi; ZRCS 2522 ， 7: 25.0-28.7 mm ，Ku a1 a Tahan ，1948 ， C.S. Ogilvie; ZRCS 12793 ， 23.5 mm ， Ku a1 a Li pis ， 21 Jan. 1958 ， Speakman; JOHORE: ZRCS 12794 ，38 .3 mm ，Semberong West

Dam ， 14 April 1990 ， V. Yong and K. Yong; ZRCS 12790 ・91 ， 2: 37.5-39.1 mm ， near

Layang-Lay 佃 g， 14 March 1990 ，P. K.L. Ng. SYSτ 芭MATIC REVISION OF TROPICAL ASIAN FRESHWATER GLASSPERCHES 283

Diagnosis. Diagnosis. - Parambassis siamensis is very sirnilar to P. ranga but generally attains a smaller smaller size and tends to have fewer scales. Gill rakers on first gill arch 5・7+17 ・18 = 22 ・ 25; 25; predorsum fully scaled; scales in lateral series about 55-60 ，transverse 11/1/ 22-23; cheek cheek with 5 scale rows; scale shea 血 on anal fin with 2 scale rows; humeral mark very weakly weakly developed or absent; vertebrae 10+14-15 = 24 (1) ，25 (19).

Comments. - Fraser-Brunner distinguished his C. punctulata 企om all other species of Asian Asian freshwater ambassids mainly by color ，i.e. ， the puncωlate marginal coloration of almost almost every scale on the body (FRASER-BRUNNER ，1954 ，fig. 3). Identical punc 佃 lation occurs occurs in some samples of P. ranga and 泊 P. siamensis ，especially in those collected 合om reservoirs. reservoirs. In the material exarnined of P. siamensis ， specimens 合om Srinakarin and Sirinthorn Sirinthorn reservoirs show 白is coloration particularly well. It is noteworthy that Fr; 錨 er- Brunner Brunner reported ap 訂 'atype of C. punctulata from Lahore; I have not exarnined 血is specimen ， but presumably it is of P. ranga. Th us no useful character w 部 provided by

Fraser-Brunner ， nor is any known ， to distinguish between P. punctulata 佃 d P. siamensis. At most localities P. siamensis ma 伽res at a relatively small size ， under 30 mm ，and does does not grow much larger. In a few localities it attains a somewhat larger size ， to 40 or 50 mm ， but nowhere does it attain the sizes commonly attained by P. ranga ，i.e. 60 or even 70mm.

Distribution. Distribution. - Parambassis siamensis occ 町'S in the Middle and Lower Mekong basins of of Th ailand ，Laos ，Cambodia ，and Vietnam; and in lowland rivers throughout Th ailand and the the Malay pe 凶nsula (Mekong ，Chantaburi ，Bangpakong ，Chao Phraya ，Meklong ，Tapi ， Pattani ，Pahang ，Perak ，and lesser basins).

Paramb 田 sis tenasserimensis new species Figure Figure 8

Holotype. Holotype. - CAS 79082 ， 65.7 mm ， Tapoleh Kl oh ， Tenasserim b出 in ， 13 March 1992 ， T. R. Roberts.

Paratypes. Paratypes. - BMNH 1992.1 1.1 6.14 ・15 ， 2: 46.9-47.7 mm ， Tenasserim R.， March 1992 ， T. R. Roberts; CAS 79083 ，4:49.1-52.7 mm ， Kayethoo Kl oh ， Tenasserim basin ， 12 March 1992 ，T. R. Roberts; CAS 81657 ， 49.8 mm ， Tenasserim R.， March 1992 ，T. R. Roberts.

Dia 伊倒is. - Parambassis tenasserimensis is 血e most deep-bodied species of Paramb ωsis ， with with steep predorsal profile and short ， deep caudal peduncle. G i1l rakers on first gill arch 5・8+16 = 21-24; dorsal fin soft rays 14 ・17 ， anal 14 ・18 ，pectoral 12-13; predorsum ne 紅 rnidline rnidline scaleless; scales in lateral series 53 ，tr 叩 sverse 8/1/23; anal fin shea 白 with on1 y two scale rows anteriorly 組 d one row posteriorly; oval humeral spot present; vertebrae 10+15 10+15 = 25.

Morphometrics. - Morphometrics of P. tenasserimensis 紅 'e comp 紅 ed with P. ranga and P. vollmeri in Table 1. Head length and body dep 血 are greater 白an in P. ranga. 284 TYSON R. ROBER'τ 百

Dep 白 of the caudal peduncle ，4.0 -4 .6 times in st 組白rd length ， probably is greater 白佃in any any other species of Parambassis. Th e peduncle also probably is shorter 血組 in other Parambassis ，but 白 is not re f1 ected in 由es 旬ndardme 部町ment of peduncle leng 也 to 血e middle middle of the caudal fm base because 泊白is species it measures mainly the increased height height of the peduncle.

Se 町 atioo. - 50.8-mm specimen: Pr eorbi taI edge 5 serrae ， ridge major serra and 4 small 田町ae; suborbi taI non-serrate; supraorbital with 4 serrae beginning over posterior p紅 t of eye; eye; preopercular edge lower limb with 9 discrete serrae ，vertical limb about 10 weak 田町ae; interopercle non- 田町ate; shoulder girdle wi 血 3 upper and 3 lower serrae.

Coloratioo Coloratioo of 骨髄bsp 田 imens. - Head ， body and fms dusky olive gr 田 n overall; preopercle and and opercle silvery ，abdomen whitish. spinous dorsal ，anterior p釘 ts of dorsal and anal ， outer outer margin of pelvic ，and upper lobe of caudal fin yellowish; p田 toral fm clear; humeral mark black; scale margins outlined by fme melanophores.

Discussioo. ー百lI s species is close to P. ranga ， but I have not observed any P. ranga (including (including many large sp 即 imens not preserved) with a comparably d田 p body and caudal peduncle ，similar steep predorsal profile ， or coloration quite like these Tenasserim speci- mens. mens.

Distribu “00. - Parambassis tenasserimensis is known only from the Tenasserim basin in southeastem southeastem Myanm 低

Table Table 1. Mo 甲home 凶 c comparison of three sp 即 ies of Parambassis.

ranga ranga tenassertmensts vollmeri n: n: standard length mm 2: 54.8-64.5 5: 47.8-65.7 3: 54 .4・ 67.1 locality locality Inle Tenasserim Mae Sa Nga head head 3.5同 3.9 3.0-3 .4 3.6 snout snout 9.6-10.9 8.6-1 1. 2 12.1-13.2 eye eye 11. 3-1 1. 8 7.9-9.5 8.6-9.0 interorbital interorbital 12.7-13.9 11. 5-13.1 11. 0-11.8 body dep 血 2.3-2.4 1. 8-2.0 1. 8-2.0 caud. caud. ped. length 6.0-6.3 7.0-8.7 7.0-7.7 caud. caud. ped. depth 8.3-9.1 6.8-7.9 8.0-8.7 1st 1st dor. fin base 4.8-5 .4 4.3-4.7 4.2-4 .4 1st 1st dor. fin height 4.9 ・5.5 4.0-4.6 5.1-5.2 2nd dor. fin base 3.5 3.5-3.7 3.1-3 .4 2nd dor. fin height 5.1-5.9 4.7-5.7 5.5 ・6.0 anal anal fm base 2.8 2.5-2.7 2.6-2.8 創 lal fm height 3.2 -4 .6 4.2-5.1 5.5-5.8 SYSTEMATIC REVISION OF TROPICAL ASIAN FRESHWATER GLASSPERCHES 285

Parambassis Parambassis thomasi (Day ， 1870)

Ambassis Ambassis thomassi Day ， 1870: 369 (type loc a1 ity Calicut and Mangalore); Day ， 1875: 52 ， l. pl. 15 ，fig. 2) Ambassis Ambassis thomassi (nec Day ，1870) ，Hora ， 1923 (Menam Chao Phya at Bangkok and Nontaburi ，官lailand; specimens not re-examined ， but identification certainly eηone- ous). ous). Chanda thom ωi，Smith ， 1945: 482 (corrected spelling of species name; mistakenly reported from Thailand). Chanda thomassi ，Fraser-Brunner ， 1955: 204. Parambassis Parambassis thomassi ，Kottelat ， 1989: 17; Talwar & Jhingran ， 1992: 802.

Type material examined. - BMNH 1870.5.2.2-3 ， 2: 57.9-82.9 mm ，Wynaad ，Day (probable (probable syntypes A. thomasi); MNI 王N 9358 ， 61 mm ，Comorin ，Day (probable syntype A. A. thomasi).

Non-type material examined. - BMNH 1889.2. 1. 2861 ，58 .4 mm ，M a1 abar ， F. Day; MCZ

4310 ，99 .4 mm ，Canara ， F. Day; BMNH 1889.2. 1. 2862 ・63 ，2: 55.3 ・68.8 mm ，Canara ， F. Day

Note on spelling of name. - The spelling Ambassis thomassi appears at the beginning of the the species description (DAY ， 1870: 369) 佃 d is not repeated in the tex t. Day gave the co l1 ector's name ， H. E. 官lomas (p. 369) ，and stated (p. 370) “1 have named the species after after Mr.τ 'h omas." Fo l1 owing Art icle 32 (c) (ii) of the Internation a1 Code of Zoologic a1 Nomenclature Nomenclature (1985 ed.) the spelling should be corrected to Ambassis thomasi.

Diagnosis. Diagnosis. - Mouth relatively elongate ，nearly horizon 凶 (as in Parambassis dayi); lower jaw slightly projecting; gill rakers on first gill arch 8-9+16-19 = 24-28; predorsum fu l1 y scaled; scaled; cheek with 5 scale rows; scales in lateral series 39 ・40 (35 ・41 according to DAY ， 1875: 1875: 52); later a1 line sc a1 es 40; transverse scale rows 6/1113; dorsal and an a1 fin sc a1 y sheaths sheaths with two scale rows; predorsum entirely cover 巴d with sc a1 es; dorsal and an a1 soft fin fin rays 10-11; second an a1 fin spine very slightly 1釘 ger than third; pector a1 fm rays 15; no humer a1 spot; vertebrae 10+14 = 24.

Serration. Serration. - 61.0-mm specimen from Comorin: Preorbit a1 edge with 11 evenly spaced ， rather rather sm a1 1 se 町 ae distributed on entire length ， ridge entirely without serrae ，no prim 釘 y serra serra dors a1 1y; suborbi ta1: with about 12 very small serrae; supraorbit a1 with 4 moderately large large se 町ae posteriorly ， otherwise entire; preopercular ridge with 7 sma l1 se 汀 ae horizon- ta1 1y ，and one sm a1 1 one just above angle ，vertic a1 limb otherwise non-serrate; preopercul 釘 edge edge with 13 sm a1 1 seηae on horizont a1 portion and 28 sma l1 serrae on vertical po 抗ion; a serra serra above preopercular articulation; interopercle with 8 uniformly sma l1 se 町 ae; shoulder girdle girdle non-serrate.

Coloration. Coloration. -“ Greenish ， shot with silvery; no lateral silvery band" (DAY ， 1870: 370); “Greyish ， spotted with silvery ，there 訂 e a1 so brownish basal marks on many of the sc a1 es ， 286 TYSON R. ROBERl 宮 more especial1y along the back" (DAY ， 1875: 53).

Distribution. Distribution. - Kn own on1y from fresh water on Malabar (sou 血west) coast of India; “自 e co 槌 ts of Canara as 10w 酪 Cochin: it is found some distance in1 and even in e1evated 10calities" 10calities" (DAY ， 1875: 52).

Parambassis Parambassis voUmeri new sped 凶 Fig 町 e9

Ambassis Ambassis ranga Vincigue rra， 1890: 35 (p 紅白n; ooe spec 加 eo 仕'O m Meetan ，up 戸 rHouogarao [Salweeo [Salweeo basio]).

Holotype. Holotype. - CAS 79220 ， 55.5 mm ，Th ai1and ， Salweeo basio ，rapids 泊 Menam Moei at km 129 N of Mae Sot 00 highway 1085 ，26 Apri1 1991 ，T. R. Roberts.

Paratyp 回.ー CAS 79074 ， 70: 31.1 -54.2 mm ，collected with h010type CAS 79075 ， 139: 13.7-40.8 13.7-40.8 mm ，Th ai1and ， Salweeo basio ， oight seioe 00 s佃 dy beach ，Meoam Moei at 129 km N of Mae Sot 00 highway 1085 ， 27 Apri1 1991 ， T. R. Roberts; CAS 79073 ， 3: 28.5- 38 .3 mm ，and ZMA 120.862 ， 3: 32.9-36.2 mm ，Meoam Moei at Kl erko ， about 100 km N of of Mae Sot ， 16 Apri11989 ，T. R. Roberts; CAS 81659 ， 14: 31. 7・38.8 mm ，T ik1 0 Me1aah ， smal1 smal1 grave1-bottom s田 amflowing 企'O m Burma ioto Meoam Moei opposite Kl erko ， about 100 100 km N of Mae Sot ，24 Apri1 1989 ， T. R. Roberts; CAS 81658 ， 3: 54 .4・ 67.1 mm ，

百1ai1 組 d， Salweeo b邸泊， N 創 n Mae Sa Nga at Ban Huai Kh an ，24 km N of Mae Hoog Soo ， 16 16 Feb. 1991 ， T. R. Roberts; CAS 79221 ， 49.1 mm ，Th ai1and ， Salweeo basio ，Huay Kh oog just just be10w mouth of Huay Lo og ， about 45 km ENE of Mae Hoog Son ，20 Feb. 1991 ，T. R. Roberts. Roberts.

Diagnosis. Diagnosis. - Parambassis vollmeri is distinguished 企om al1 other amb 拙 sids (marioe and 仕eshwater) by its bri1 liant ，1紅 ge1y iridesceot c010ratioo and distioctive b1ack marks 00 body and fios. G i11 rakers 00 frrst gi11 arch on1 y3 -4 +10 ・12 = 13 ・16. Pr edorsum oaked; 1ateral1ine 1ateral1ine scale series about 50 ，佐組sverse 811/18; dorsal and anal fio scale sheath mos t1 y with with on1 ya s 泊gle scale row ， incomp1ete posterio r1 y esp 即 ial1y 00 dorsal fio (some smal1 scales 叩 terio r1 y formiog a secood row 00 bo 出 fios). Dorsal and anal fio branched rays 16-17; 16-17; pectoral fm rays 12 ・13. Vertebrae 10+15 = 25.

Morphometri 回 .-Morphome 凶 cs of P. vollmeri 紅 e comp 紅 ed with those of P. ranga and P. tenasserimensis ，出e species of Parambassis with geographical r組 ges ove r1 app 泊g or or nearby its range ， io Tab1e 1. Th e SDOut of P. vollmeri ， with 1ength 12 .4同 13.2 times in S刷 d制 1eo 伸， probab1y is shorter 血叩 io any other species of Parambassis. Head 1eo. 紳， 3.0 ・3.4 ，is sligh t1 y greater 由加泊P. ranga or P. tenasserimensis. 百1e more posterior dorsal and anal fio soft rays of P. vollmeri appe 訂 to be shorter 白an io other Parambassis ， al 血ough 白is is not reflected in the height of the 100gest e1ements at the beginning of the fios. fios. SYSTEMATIC REVISION OF TROPICAL ASIAN FRESHWATER GLASSPERCHES 287

Serration. Serration. - 49.9-mm specimen from Huay Kh ong: Preorbital edge with 8 moderate serrae ， ridge with large major seηa directed posteriorly and 4-5 small serrae projecting ventrally; ventrally; suborbital non-serrate; supraorbital with 9 se 町 ae of increasing size beginning over over middle of eye; preopercular edge lower limb with about 5 weak se 町 ae plus 1 just above above angle ，vertical limb otherwise non-serrate; preopercular ridge with 11 seηae on

horizontal horizontal and 18 on vertical limb; a strong posteriorly directed se 町 a immediately pos 自 terior terior to last supraorbital se 汀 a and above origin of preopercle; interopercle without se 町ae;

shoulder shoulder girdle with 2 small upper and 6 small lower se 汀 ae.

Distribution. Distribution. - Parambassis vollmeri is known only from large ，clearwater tributaries of the the Salween basin (Nam Mae Sa Nga ，Huay Khong ，Menam Pai and Menam Moei in Thailand ， upper Houngarao in Myanmar).

Etymology. ー- Parambassis vollmeri is named after the late Emest Vollmer ，J r.， of San Luis Luis Obispo ，Califomia ， for his interest in ichthyological exploration.

Parambassis Parambassis wo lJ fii (Bleeker 1851) Figure Figure 10

Amb αssis wo !ffi i Bleeker ， 1851a: 9 (type locality Bandjermassin). Ambassis Ambassis robustus Guocality Bomeo). Ambassis Ambassis boulenger Volz ， 1903: 553 (type locality Palembang ， Sumatra). ? Ambassis thomassi ， nec Day Hora ， 1923 Ambassis Ambassis wo !ffi i，Weber & de Beaufort ， 1929:401 Acanthoperca Acanthoperca wo 伊 i，Fowler ， 1937: 230 (Thailand) Chanda wo 伊， Smith ， 1945: 482 (Thailand) Parambassis Parambassis wo !ffi， Allen ， 1982: 166

Type material examined. - None.

Non-type material examined. - MNHN 1989.1344 ， 2: 5 1. 9幽 69 .5 mm ，Days km 9，Cam- bodia ，30 Jan. 1960 ， F. d'Aubenton; MNHN 1992.1058 ， 94.5 mm ，Sungai 恥1entaya near Sampit ， Kalimantan Tengah ， 10 June 1992 ， G. Lucas.

Di agnosis. - A large ，deep 司 bodied species ， with veηsteep ，straight predors a1 profile; second second anal fin spine greatly enlarged ， moreso than in 担 y other Asian freshwater ambassid; gill gill rakers on first gill arch 27 ・29; dorsal and an a1 branched fin rays 10 ・11; predorsum fully scaled; scaled; cheek with 9-10 scale rows; scales in lat 巴ral series 4テ46; no humer a1 spot; verte- brae brae 10+14 = 24.

Serration. Serration. - 94.5-mm specimen from Mentaya: Preorbital edge 11 serrae ， ridge no major 田町a8 small 田町民; preopercular edge 17 strong se 町 ae on horizontallimb and 3 just above angle ，verticallimb otherwise non-serrate; preopercular ridge 12 se 町 ae on lower limb plus

3 just above angle ，vertical limb otherwise non-se 汀 ate; supraorbital entire anteriorly ，7 288 TYSON R . ROB ERTS

Figure 9. Parambassis vollmeri, 55.5 mm, Menam Moei (holotype).

Figure I 0. Parambassis wo/jfii, 93.5 mm, Sungai Mentaya. SYSτ 芭MATIC REVISION OF TROPICAL ASIAN FRESHW ATER GLASSPERCHES 289

weak se 町 ae p'O steri 'O rly st 紅 ting 'O ver p'O steri 'O rp 紅 t 'O f eye; sub 'O rbital margin irregular ， but n-serrate; n'O n-serrate; inter 'O percle with 12 str 'O ng se 町ae; sh 'O ulder girdle n'O n- 田町ate.

Di stribution. - Parambassis wolffii is dis 凶 buted disc 'O ntinu 'O usly in mainland and insu- lar lar S 'O utheast Asia. On the mainland it is kn 'O wn '0凶 y fr 'O m the Cha 'O Phraya and Mek 'O ng basins ，and evidently is absent fr 'O m the entire Malay peninsula. In出 .e islands it is kn 'O wn 企'O m Suma 住a (Palembang) and B 'O rne 'O (Kapuas ， Mentaya ，Barit 'O， K 油吋 an). A rec 'O rd 合'O m Baram Ri ver by WEBER & DE BEAUFORT ， 1929: 402 is d'O ub ぜu1.

ACKNOWLEDGMENTS

Th e auth 'O r especially wishes t 'O thank Francine Berk 'O witz ， Office 'O f In ternati 'O nal Pr'O grams ，Smiths 'O nian In stituti 'O n，f 'O r 紅 rang 泊g 凶 ps t 'O Nepal ，In dia ，and Myanmar d町ー ing ing which m 叩 y 'O f the specimens rep 'O rted in 血is paper were c'O llected. Special thanks f'O r l'O ng c'O ntinued supp 'O rt are due als 'O t 'O Ir a Rubin 'O ff ，R 'O ss Sim 'O ns ，and Frank Talb 'O t 'O f the Smiths 'O nian. Fieldw 'O rk in India was facilitated by Madhav Gadgil ，Bangal 'O re Institute 'O f Science ，and Kaza Rama Ra 'O 'O f the Zo'O l 'O gical Survey 'O f India. Fieldw 'O rk in Th ailand

W 部 d'O ne with the permissi 'O n 'O f the Nati 'O nal Research C 'O uncil 'O f Th ailand. Gene Lucas ， aquarist aquarist and Betta enthusiast ， 'O rganized ac 'O llecting trip t 'O Kalimantan Tengah. Visits t 'O 白e Zo'O l 'O gical Research C 'O llecti 'O ns 'O f the Nati 'O nal University 'O f Singap 'O re were fac i1i- tated tated by Peter Ng ， Kelvin Li m ，組d Yang Chang Man. Kelvin L 加 brought t 'O my attenti 'O n specimens specimens 企'O m Pahang he had identified 邸 Parambassis apogonoides. Martin van 0 討en and K 'O s van Egm 'O nt facilitated my w 'O rk in Leiden ， as did Oliver Crimmen ， Patrick Campbell ，and G 'O rd 'O nH 'O wes in L 'O nd 'O n. Radi 'O graphs were prep 紅 'ed by Jeanine Abel in P副 s， Eugenie Bohlke in Philadelphia ， and Dave Catania in San Francisc 'O. M 'O st 'O f the research and writing 'O f the paper w 出 d'O nein 由e Lab 'O rat 'O ire d' Ic hty 'O l'O gie ， Museum National d' Hi st 'O ire Naturelle ， and Dep 釘 tment 'O f Ichthyol 'O gy ，Califo 口語 a Acad- emy 'O f Sciences. I血ankMarie ・Lo凶 se Bauch 'O t，R 'O l組 dBill 紅 d，M 紅 t凶.e Des 'O utter ，Guido Di ngerkus ，Guy Duhamel ， and Jean-Claude Hureau in P剖 s and Tomio Iwamoto ，W i1l iam N. N. Eschmeyer 組 dDaveCa 匂凶a in San Francisco. Funding w ぉ provided by 血eSmi 血s'O凶 m Institution ， Smithsonian Tropical Research In stitute ，Institute for Biological Exploration ， and Committee f'O r Research and Expl 'O rati 'O n of the Nati 'O nal Geographic S 'O ciety (gr 姐 t

5141 ・93).

REFERENCES

ALL 剖， G. R. 1982. Parambassis altipinnis ，a new species of freshwa 旬 r glassfish from wes 匂rn New Guinea. Bull. Bull. Zo ol. Mus. Univ. Arn sterdam 8: 165-169. B 凶 E阻 R ， P. 1851a. Bijdrage tot de kennis der ichthyologische fauna van Borneo ，met beschrijving van 16

nieuwe nieuwe s∞由n van zo 巴twatervisschen. Nat. Tijds. Ned. 100. 1: 1・16. BLEE 阻 R， P. 1851b. Vierde bijdrage tot de kennis der ichthyologische fauna van Borneo ，met besc 耐甘 ving van

関凶ge nieuwe soorten van zoetwatervisschen. Nat. Tijds. Ned. 100. 2: 193 ・208. BLEE 阻 R ， P. 1853. Nalezingen op de ichthyologische fauna van Bengalen en Hi ndostan. Verh. Batav. Genootsch. Kunst. Kunst. Wet. ，25: ト164 ，1 pl. B 国阻R ， P. 1857. Tiende bijdrage tot de kennis der ichthyologische fauna van Bornω. Visschen van de rivi 巴m 290 290 τ'v SON R. ROBE R:官官

Barito ，Kah aj 組 en Ka p∞ as. Ac t. Sω. Sc i. Indo-Neerl. 2: 1・21. B 回蹴ER ， P. 1874. Revision des es 悼ces d' Amb assis et de Parambassis de I'In de archipelagique. Nat. Verh. Holl. Maatsch. Maatsch. Wetensch. ， 3. Verz. n， no. 2: 83-106. BL YTH ，EJ. 1861. Report on some fishes received chiefly from 恥 Sitang Ri ver and its 肘butary s回 ams. よAs iatic. Soc. ，Calcu 伽， 29[18 印]: 138 ・174. Bo 国 EMAN ，M. 1957. 伽 a collection of e鎚 t Asian fishes. Zo ol. Me d. 35: 69 ・78 ，pl. 3. CAS 叩凶AU ，F. L. 1878. No 回 on 血e fishes of 曲e Norman 尉ver. Proc. Li nn. Soc. New South W 匂les 3: 41-5 1. COL 日 ITE ， B.B. 1977. Epidermal breeding 帥 ercles and 加ny contact org 組 s in fishes. あP岬'. Zo ol. Soc. Lo ndon

39: 39: 225 ・268. CUVl 殴， G. ，AND A. VAU 到 αENNES. 1828. Histoire naturelle des Poissons ，vo l. 2. Paris. xxi 叫 90 pp. ，pls. 9・ 40. 40. DAY ， F. 1865a. on 血e fishes of Cochin ，on 恥 Malabar co 制 of In dia. Proc. Zo ol. Soc. Lo ndon 1865 (p t. 1): 558-565. 558-565. DAY ， F. 1865b. Th e fishes 01 Malabar. Lo ndon ，Bemard Quaritch. xxxu+293 pp. ，20 pls. DAY ， F. 1870. Notes on some fishes from the wes 飽mco ，制 of In dia. Proc. Zo ol. Soc. Lo ndon ， 1870: 369 ，・ 374. DAY ， F. 1875. Th e fishes olIndia; being a natural histo η olfishes 初 own ω in 加 bit the seas and fresh waters olIndi a， Burma ，and Ceylon. Lo ndon. Part 1: 1・168 ，pls. 1-40; FOWLER ，H.W. 1937. Z00 logical results of the 曲凶de Schauensee Siamese ex 戸 dition. Part 8. Fish 凶 ob 凶 ned in in 1936. Proc. Acad. Nat. Sci. Philadelphi a， 89: 125-264.

FOWLER ，H.W. ，AND B.A. BEAN. 1930.τbe fishes of 血 e families Ami idae ，Ch 組 didae ，Duleidae ，飢 d Serr 副也e， ob 旬inl 叫 by 伽 U 凶旬d Sta 蜘 Bureau of Fisheries Steamer “Alba 回 S8" 泊 1907 to 1910 ， chiefly in 血e Philippine Philippine Islands and adjacent seas. Bull. U. S. Natl. Mus. 1∞(1)， ix+ 334 pp. FRAs ER-BRUNNER ， A. 1955. A synopsis of 恥 cen 加 pomid fishes of 血e subfamily Ch andinae ，wi 由 description

of of a new genus 飢 d two new species. Bull. Ra jJl es Mus. 25: 185 ・213. Gu 貼， S. ，AND P.K. T，札， WAR. 1983. On 血巴 type specimens of Ambass おnota 加 Bly 也， 1861 (Pi sc 砿Am bassidae) ， wi 血 a redescription of the species. Bull. Zo ol. Surv. India 5: 15 ・19. G 加古IER ， A. 1859. Catalogue 01 the fishes in the British MIω 'eu m. Vol. 1. Catalogue olthe acanthopterygian

fishes fishes in the collection (ず the British Museu m. Gasteroste 油 e，Be ηci， 伽 ，Perc 油 e，Ap hredoderidae ， Pris ゆomatidae ，Mullidae ，Sparidae. Lo ndon. xxxi+524 pp. 日制ll. TON ， F. 1822. An account 01 the fishes lound 仇 the river Ganges and its branches. 国 inburgh and Lo ndon. Text Text vu +40 5 pp ・， Atlas 39 pls. Ho 貼， S. L. 19 叩.白川 collection of fish from Si am. J. Siam Soc. ， Nat. Hist. S叩，opl. 6: 143 ・184 ，pls. 10- 12. HORA ， S.L. 1929. An aid to the 蜘 dy of Hamil 蜘 Buchanan's “Gangetic Fishes" ，Me m. Indian Mus. ， 9: 169 ・ 192 ，pls. 13 ・23. KOTIELAT. KOTIELAT. M. 1989. ゐ ogeography of fishes from Indωh 担問e inland wat 悶 with 組 annota 旬d check-lis t. Bull. Zo ol. M ω'. ， Univ. Ams ter 幼:zm， 12: 1・， 56. M cC日 LLAND ，J. 1842. On 恥合唱sh-water fishωcoU 印刷byW 出iam Gri ffi 血， Es q. ， F.L.S. M 拍車sM 叫 cal Sぽ vice ， during his 回， vels und 町首le orders of the Supreme Go vernment of In dia from 1835ω1842. Calcutta よ Nat. Hist. 2: 5印・ 589 ，pls. 4- 6，15 ，18 ， 2 1. ROBERTS ，T. R. 1989.τbe freshwa 飽 r fishes of westem Bomeo (Kalinlan tan Bar 札In donesia). Mem. Califomia Acad. Acad. Sci. 14 ，凶 +210 pp. SMITH ，H.M. 1945. 百le 企官 shwater fishes of Siam ， or Th ailand. Bull. U. ぶ Nat. Mus. 188.

T 札，WAR ，P. K.， AND A. G. JH lN GRAN. 1992. lnland fishes ollndia and adjacent 印刷tries. 2. A.A. Balkema ， Rot 旬:rdam.

V AlL LANT ， L. 1875. Remarques s町 la famille des Bogod 恒i ，Bleeker. Bull. Soc. Philo m. Paris (6)12: 32 ・34. V 岨ム刷r， L. 1893. Con' 凶， bution a l'蜘 de de la faune ichthyologique de Bomeo. Nouv. Arc h. Mus. Hist. Natl. Paris Paris (3) 5: 23 ・114 ，2 pls. VOLZ ，W. 1903. Neue Fische aus Suma回 • Zo ol. A nz. α ..e ipzig) 26: 553 ・559. WEBER ，M. ，AND L. F. dE B 臥 UFORT. 1929. The fishes 01 the lndo ・Austral ，加 archipelago. 5. E.J. Brill ，Le iden. xiv +4 58 pp.

W n.E Y，M.L. ，AND B.B. CO LLETrE. 1970. Breedingωbercles and contact org 組 s in fishes: their occ 町 rence ，

S町ucture ，and sigr 泊白 C 組 ce. Bull. Amer. Mus. Nat. Hist. 143 :143 ・216.