Lumbantobing 2014 Four new species of Rasbora of the Sumatrana group (Teleostei: Cyprinidae) from...

ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2014 Magnolia Press

Zootaxa 3764 (1): 001–025

www.mapress.com/zootaxa/Article

http://dx.doi.org/10.11646/zootaxa.3764.1.1

http://zoobank.org/urn:lsid:zoobank.org:pub:CCCD2FDF-BDF9-48A0-ADDE-8017F9AE1943

Four new species of Rasbora of the Sumatrana group (Teleostei: Cyprinidae)

from northern Sumatra, Indonesia

DANIEL N. LUMBANTOBING1,2

1Department of Biological Sciences, The George Washington University, 2023 G Street. NW, Lisner Hall 340, Washington, D.C.,

20052. E-mail: [email protected] of Fishes, Smithsonian Institution, National Museum of Natural History, Washington, D.C., 20013

Abstract

Four new species of the minnow genus Rasbora of the Sumatrana group, R. arundinata, R. haru, R. maninjau, and R. bin-

dumatoga, are described from northern Sumatra. Rasbora arundinata is distinguished from all congeners in the Sumatrana

group by the black midlateral stripe overall forming a reed-leaf-like profile. Rasbora haru differs from its congeners in

the Sumatrana group in having the black midlateral stripe overall forming a stamen-like profile. The new species endemic

to Lake Maninjau in central west Sumatra, Rasbora maninjau, is unique among all the congeners in the Sumatrana group

in having a combination of the black midlateral stripe extending from the midhumeral region of uniform width, the prom-

inent acutely triangular basicaudal blotch, and the oval supra-anal pigmentation. Rasbora bindumatoga is distinguished

from all congeners in the Sumatrana group by a combination of the black rectangular subdorsal blotch, the absence of su-

pra-anal pigmentation, and the somewhat oval basicaudal blotch. Rasbora arundinata, R. maninjau, and R. bindumatoga

occur allopatrically in the northwestern coastal region of Sumatra, while R. haru is known from northeastern coastal area

of Sumatra. A new diagnostic character for the Sumatrana group is described: partial exposure of the upper lip due to a

submedial contact between the maxilla and the lower lip, which is marked posteriorly by a lachrymal groove.

Key words: Danioninae, Rasbora, new species, northern Sumatra, Sundaland

Introduction

Rasbora is a small-to-moderate-sized genus in the family Cyprinidae that lives throughout a vast geographical

range within Asia, including the Indian subcontinent, southern China, and Southeast Asia (Weber & Beaufort 1916;

Brittan 1954a). With currently 77 species, Rasbora constitutes the most species-rich genus in the cyprinid

subfamily Danioninae (Eschmeyer 2013; Froese & Pauly 2013). Taxonomically, Rasbora has been widely

considered as a catch-all group due to a lack of unique diagnostic characters (Brittan 1954a; Kottelat &

Vidthayanon 1993; Liao et al. 2010; Tang et al. 2010). In the first and most comprehensive revision of Rasbora,

Brittan (1954a) recognized three subgenera (Rasbora, Rasboroides, and Megarasbora), and further classified the

subgenus Rasbora into eight species complexes: the lateristriata, the sumatrana-elegans, the caudimaculata, the

trifasciata, the argyrotaenia, the daniconius, the einthovenii, and the pauciperforata complexes. Brittan’s species

complexes have been widely used as a practical system for the classification of the group and frequently revised by

many authors (Kottelat & Vidthayanon 1993; Siebert & Guiry 1996; Kottelat 2005; Liao et al. 2010). In their brief

revision of Rasbora, Kottelat & Vidthayanon (1993) replaced the category of species complex with ‘species

group,’ which has been widely used by most of the later workers.

Authors after Brittan (1954a) have created several new genera (Boraras, Brevibora, Horadandia, Kottelatia,

Rasboroides, Trigonopoma, and Trigonostigma) for some lineages within Rasbora (Kottelat & Vidthayanon 1993;

Kottelat & Witte 1999; Liao et al. 2010). Despite these newly created genera, many workers still have recognized a

larger assemblage equivalent to the concept of the genus Rasbora sensu Brittan (1954a), being frequently referred

to as “the genus Rasbora sensu lato” (hereafter Rasbora s. l.). Rasbora s. l. comprises the genus Rasbora sensu

stricto (hereafter Rasbora s. s.) and the new genera mentioned above (Kottelat & Vidthayanon 1993; Kottelat &

Witte 1999; Conway 2005; Liao et al. 2010; Tang et al. 2010).

Accepted by L. Page: 17 Dec. 2013; published: 7 Feb. 2014 1

Members of the highly pelagic genus Rasbora s. l. were traditionally identified in having a combination of the

following characters: a laterally compressed, elongate body; a symphseal knob of the dentary fitting into a

corresponding depression in the maxilla; the dorsal fin with two simple and seven branched rays and its origin

located about mid-point between the snout and the hypural notch; and anal fin with three simple and five branched

rays (Brittan 1954a). Using morphological characters, Liao et al. (2010) reconstructed phylogenetic relationships

of Rasbora s. l., along which eight new synapomorphies that were inferred for the first time to support the

monophyly of the genus. Nevertheless, with more danionine taxa as outgroups, a recent study by Liao et al. (2011)

confirmed only two of the eight characters uniquely diagnose Rasbora s. l.: the dark supra-anal pigmentation

combined with the subpeduncular streak; and the rasborin process on the fourth epibranchial.

One of the most diverse yet taxonomically problematic species groups of Rasbora is the Sumatrana group,

which was initially referred to as the R. sumatrana-elegans complex by Brittan (1954a). The Sumatrana group is

characterized by a unique black pigment pattern consisting of a reduced or modified midlateral stripe arranged

alongside other pronounced elements, such as the supra-anal pigmentation and the blotch on the caudal peduncle

(Brittan 1954a, 1954b; Kottelat & Vidthayanon 1993; Kottelat & Tan 2011, 2012). Unlike in the other species

groups of Rasbora, members of the Sumatrana group are relatively uniform in their overall appearance, which

includes their body size, shape, and life coloration.

The lateral pigmentation pattern formed by melanophores, which are readily observable after preservation,

provides a suite of characters useful in distinguishing species of the Sumatrana group, and several new species

recently have been described using these characteristics (Kottelat 2005; Kottelat & Tan 2011, 2012). Kottelat &

Tan (2012) examined different types of melanophores comprising the mid-lateral pigmentations, each of which

appears distinct, primarily as a consequence of its unique topographical position in the mid-frontal plane of the

skin. Nevertheless, as implied in the recent taxonomic studies of the Sumatrana group (Kottelat 2005; Kottelat &

Tan 2011, 2012), it is the lateral distribution of melanophores along the trunk that provides more diagnostic value,

which is manifested in highly interspecific variation in the shape, size, and intensity of the components of the black

lateral pigmentation pattern. In the present study, a thorough observation on the lateral distribution of

melanophores has revealed another array of distinct elements comprising the black lateral pigmentation pattern,

which shows high variability in the species level. Accordingly, a new terminology for such elements in the

Sumatrana group is established and described.

During an ichthyological survey of northwestern and central Sumatra in 2006, three species initially identified

as R. sumatrana (Bleeker, 1852), R. elegans Volz, 1903, and R. lateristriata (Bleeker, 1854) were collected.

However, further examination revealed that each of the three species possessed a distinct set of diagnostic

characters, which warranted each to be recognized as a new species described herein. The addition of these three

new species of the Sumatrana group, together with four newly described species of the Trifasciata group

(Lumbantobing 2010) and three other valid species (R. jacobsoni, R. reticulata, and R. vulcanus), brings the

number of the species Rasbora restricted to the western coast of Sumatra to ten, and demonstrates the high

endemism of the region. Another undescribed species of the group from the northeastern slope of Sumatra is also

described herein as a new species.

Material and methods

The name of the species group or subgroup follows a nomenclature convention applied by Springer and Allen

(2004), who used the capitalized and non-italicized species epithet for the group (e.g., the Sumatrana group; the

Lateristriata subgroup). This system is selected over systems used by previous authors (e.g., Kottelat &

Vidthayanon 1993; Siebert & Guiry 1996; Liao et al. 2010; Lumbantobing 2010) because it distinguishes group or

subgroup names clearly from the Linnaean binomial nomenclature of the species names.

Most of the specimens were collected between June and August 2006 by seining and electrofishing.

Institutional abbreviations follow Sabaj Pérez (2012). Morphometric measurements were recorded from the left

side of a specimen when possible following Kottelat (2001) and Lumbantobing (2010) using digital calipers to the

nearest tenth of a millimeter. Measurements are reported as a range of percentage of standard length. Fin-ray and

scale counts follow Kottelat (2001). Vertebral counts follow Siebert & Richardson (1997) and were taken from

radiographs. Terminology for body color patterns follows Brittan (1954a) and Lumbantobing (2010).

LUMBANTOBING2 · Zootaxa 3764 (1) © 2014 Magnolia Press

Several new terms for body pigmentation useful to diagnose the Sumatrana group are also described herein.

These delineate the details of the three primary diagnostic characters of the group: (1) the black midlateral stripe

(Fig. 1: MLS); (2) the supra-anal pigmentation (Fig. 1: SAP); and (3) the basicaudal blotch (Fig. 1: BCB).

In general, a complete black midlateral stripe in the Sumatrana group can be divided into three elements: (1)

the midhumeral diffuse patch (Fig. 1: MDP); (2) the subdorsal blotch (Fig. 1: SDB); and (3) the posterior-portion

stripe (Fig. 1: PPS). The midhumeral diffuse patch is formed by one or two rows of scales in the midlateral region

between the gill opening and the vertical through the dorsal-fin origin, the exposed portion of which is sparsely

speckled by melanophores. The speckled scale rows overall appear as a diffuse pigmented swath. The subdorsal

blotch is the ventrally widened portion of the black midlateral stripe located more anteriorly and somewhat below

the dorsal fin and dorsal to the pelvic fin. The posterior-portion stripe is the section of midlateral stripe, which

extends along the posterior half of the body and is dorsally bordered or traversed by the axial streak. Species of the

Sumatrana group may possess all of the three elements of the black midlateral stripe, whereas some species only

possess one or two elements, with other elements rudimentary or absent. The species of the Sumatrana group show

high levels of variation in the black midlateral stripe due to the combination of variation in shape, size, intensity,

position, and completeness of elements, which vary across species.

FIGURE 1. Left lateral views of body of the Sumatrana group showing: (A) general body pigmentation; (B) schematic

drawing of lateral body pigmentation. AS = Axial Streak; BCB = Basicaudal Blotch; BCS = Basicaudal Spot; BR = Basal

Reticulation; BTP = Basicaudal Triangular Patch; CP = Caudal Pigmentation; DLS = Dorsolateral Stripe; MDP = Midhumeral

Diffuse Patch; MLS = Midlateral Stripe; POP = Postopercular Pigmentation; PPS = Posterior-Portion Stripe; PR = Peripheral

Reticulation; SAP = Supra-anal Pigmentation; SDB = Subdorsal Blotch.

Zootaxa 3764 (1) © 2014 Magnolia Press · 3FOUR NEW SPECIES OF RASBORA FROM SUMATRA

The supra-anal pigmentation is the densely pigmented region dorsal to the anal fin. This pigmentation varies in

shape, from an oval to a roundish blotch to an elongated thin line along the anal-fin base. This pigmentation also

varies in its position relative to the anal fin. The third primary diagnostic character of the Sumatrana group is the

basicaudal blotch, which consists of two elements: (1) the triangular patch (Fig. 1: BTP), and (2) the basicaudal

spot (Fig. 1: BCS). Some species of the Sumatrana group can possess both elements, whereas others only possess

one element. Each element of the basicaudal blotch varies in size, shape, and intensity. The basicaudal triangular

patch can also vary on the basis of its apex position relative to the axial streak, which is either parallel or more

ventrally positioned.

Results

Common features. All species of the Sumatrana group from northern Sumatra share the following characters.

Body slender, elongate and laterally compressed (Fig. 2). Greatest body depth located between verticals through

pelvic-fin insertion and dorsal-fin origin. Dorsal profile of head overall posterodorsally slanted from margin of

upper lip to rear of head. Snout convex, somewhat turned upwards, and slightly concave along supranasal profile.

Dorsal profile of body overall slightly arched, convex from supraoccipital to dorsal-fin origin, and slightly

posteroventrally slanted from latter point to caudal-fin base. Ventral profile of body gently irregularly convex from

margin of lower lip to posterior terminus of anal-fin base, straight and nearly horizontal along caudal peduncle.

Mouth oblique and slightly superior. Tip of dentary forming anterior terminus of head. Symphyseal knob of

dentary strongly developed, slightly upturned, and fitting into corresponding well-developed symphyseal

indentation between premaxillae. Conspicuous obtuse depression on ventrolateral margin of upper jaw notched by

deep lachrymal groove. Lateral surface of upper lip discontinuously exposed, with part of anterior portion well-

exposed, submedial portion slightly covered due to contact point between maxilla and lower lip, and then exposed

again posteriorly from lachrymal groove to rictus (Fig. 3). Rictus situated slightly anterior to, or at vertical through,

anterior margin of eye. Isthmus marked with indistinct projection, situated at vertical through anterior margin of

pupil and forming indistinct obtuse angle along ventral profile of head.

Scales cycloid, moderately large with regularly imbricate arrangement, focus located more basally; some

specimens with semicircular flange on posterior margin. Lateral-line series with anterior portion posteroventrally

steeply inclined, becoming somewhat straight from 9th to 17th scale, and somewhat posterodorsally ascending

from 18th to last scale. All scales on lateral-line series pored. Long lancet-shaped axillary scale located dorsal to

pelvic-fin base and separated dorsally by one scale above from lateral-line series. Several lanceolate sheath scales

present along anal-fin base and medial portion of caudal-fin base. Dorsal-fin profile somewhat pointed,

subtriangular and with posterior margin slightly convex. First unbranched ray approximately one-third length of

second ray. Pectoral-fin profile slightly falcate. Droplet-shaped fleshy axillary lobe situated dorsal to base of first

unbranched ray. Pelvic-fin profile slightly falcate. Anal-fin profile acutely subtriangular with concave posterior

margin. Caudal fin deeply forked, with acutely pointed asymmetrical lobe and lower lobe slightly longer.

All species of the Sumatrana group from northern Sumatra also share several coloration shown in alcohol

specimens as follows (Fig. 2). Dorsolateral portion of body with dusky background and ventrolateral region

lacking dusky pigmentation. Dorsum of head dusky with meningeal covering of brain most intensely pigmented.

Opercle overall dusky due to somewhat sparse concentration of superficial melanophores. Submedial opercular

canal marked with deeply embedded dense melanophores forming obscure mid-opercular streak separating dorsal

and ventral portions of opercle. Occipital region demarcated by transverse streak. Javelin-shaped mid-dorsal stripe

about one-fourth scale wide and extending from nape to dorsal part of caudal peduncle. Dorsolateral and midlateral

region of body with reticulate pattern consisting of peripheral and basal reticulation. Reticulation most intense on

anterior portion of mid-lateral region, and decreasing gradually in intensity posteriorly and ventrally. All fins with

fin rays and interradial membranes bordered by lines of small melanophores resulting in fin rays with dark margins,

except for unpigmented distal portion of paired fins and posteriormost anal-fin rays. Each caudal-fin lobe with last

procurrent ray and five longest principal rays superficially pigmented with dense melanophores along

approximately basal one-third of each fin ray. Superficial pigmentation overall resulting in obscure triangular dark

patch on each caudal lobe.

All species of the Sumatrana group from northern Sumatra share common sexual dimorphisms as follows.

Females are more deep-bodied than males. Males have 1–2 rows of antrorse tubercles on the dorsoproximal side of

pectoral-fin rays.


FIGURE 2. (A) Rasbora arundinata, holotype, MZB17881, 74.5 mm SL, female; (B) Rasbora haru, holotype, MZB 17888,

65.4 mm SL, female; (C) Rasbora maninjau, holotype, MZB 17890, 66.9 mm SL, male; (D) Rasbora bindumatoga, holotype,

MZB 17889, 47.1 mm SL, female. Bars equal 1 cm.


FIGURE 3. Left lateral views showing lips. (A) the Sumatrana group: Rasbora volzi, ZRC 45656, 87.1 mm SL; (B) the

Einthovenii group: R. jacobsoni, ZRC 49140, 47.4 mm SL; (C) the Trifasciata group: R. bankanensis, USNM 230222, 43.7 mm

SL; (D) the Trifasciata group from northwestern Sumatra: R. tobana, CAS 68363, 19.1 mm SL. Each bar equals 2 mm. Arrow

points to lachrymal groove.

Rasbora arundinata, new species

(Figures 2A, 4A–C, 7A)

Rasbora lateristriata var. sumatrana (non Bleeker 1854): Weber & de Beaufort 1916.

Rasbora lateristriata (non Bleeker 1854): Wirjoatmodjo 1987; Kottelat & Vidthayanon 1993.

Rasbora sumatrana (non Bleeker 1852): Hadiaty 2005.

Rasbora cf. sumatrana (non Bleeker 1852): Lumbantobing 2010.

Holotype. MZB 17881 (ex. USNM 390034), female, 74.5 mm SL, Indonesia, Sumatra, Province of Nanggroe

Aceh Darussalam, Kabupaten Aceh Singkil: Road from Subulussalam to Singkil, Lae Petal River, 02°31’76”N,

098°02’64”E, 21 July 2006, D. N. Lumbantobing, D. Rudaya, and N. M. Ray.

Paratypes. Indonesia, Sumatra: collected with holotype: MZB 17882 (ex. USNM 390034), 1, 67.8 mm SL;

USNM 390034, 2, 64.5 and 68.5 mm SL. Province of Aceh: Kabupaten Gayo Lues: MZB 17885 (ex. USNM

390069), 1, 64.7 mm SL; USNM 390069, 1, 47.5 mm SL, Kampung Lintoh, a tributary of Tripa River, on the road

from Takengon to Blangkejeren, 04°02’06”N, 097°20’33”E; D. N. Lumbantobing, R. K. Hadiaty, D. Rudaya, and

N. M. Ray, 7 July 2006. Kabupaten Aceh Selatan: Kecamatan Kluet Timur: D. N. Lumbantobing, D. Rudaya and

N. M. Ray: 15 July 2006: MZB 17884 (ex. USNM 390053), 8, 42.7–86 mm SL; USNM 390053, 7 (1 CS), 45.2–

85.2 mm SL, Lawe Mokap River, tributary of Kluet River, 03°09’96’’N, 097°23’90’’E; ZRC 53196 (ex. USNM

390053), 3, 49.7–76.0 mm SL; USNM 401462, 4, 43.0–68.2 mm SL, Hari Pinem River (a tributary of Kluet River),

03°09’62N, 097°24’89”E; ZRC 53197, 2, 37.9–58.0 mm SL, Hari Pinem River (a tributary of Kluet River),

03°09’62N, 097°24’89”E. Kabupaten Aceh Singkil: MZB 17886, 5, 22.7–40.3 mm SL, road between Rimo and

Singkil, Laicuk Bridge, tributary of Alas River, 02°19’29”N, 097°55’61”E, D. N. Lumbantobing, D. Rudaya and


N. M. Ray, 21 July 2006. Kabupaten Aceh Tenggara: USNM 391607, 2, 54,6–62.8 mm SL, Ketambe, Alas River,

03º41’91’’N, 097º38’74’’E, D. N. Lumbantobing, R. K. Hadiaty, D. Rudaya, and N. M. Ray, 2 July 2006. Province

of Sumatera Utara (North Sumatra): MZB 17883, 2, 47.9–71.6 mm SL, Kabupaten Tapanuli Tengah, irrigation

canal of Aek Pinangsori River (tributary of Batang Lumut River) on road between Sibolga and Batangtoru,

01°33’59’’N, 098°54’62’’E, approximately 46 m above sea level, D. N. Lumbantobing, D. Rudaya, N. M. Ray, and

P. Simanjuntak, 4 August 2006.

Non-types. All from Indonesia: Sumatra: Province of Aceh: Kabupaten Aceh Barat: MZB 4646, 4, 71.1–85.9

mm SL, Krueng Ukam, Tadue, Kuala, A. Saim. Kabupaten Aceh Selatan: MZB 5656, 5, 77.0–80.0 mm SL, Alur

Serembaning, Ruding Lanak, Sungai Alas di hilir Sungai Gelombang, Soetikno W., 20 Feb 1984; MZB 5658, 4,

64.5–99.9 mm SL, Sungai Alas, 5 km from hulu Gelombang, Soetikno W. and D. Hardjono, 3 Feb 1983.

Kabupaten Aceh Singkil: MZB 17887, 1, 42.7, Dano, road between Gelombang and Subulussalam, small river

under bridge, 02°41’42”N, 097°59’70”E; D. N. Lumbantobing et al. 18 July 2006.; USNM 390145, 1, 26.9 mm

SL, same data as MZB 17887; USNM 406855, 1, 69.3 mm SL, fish market in Gelombang, D. N. Lumbantobing, D.

Rudaya, and N. M. Ray, 19 July 2006; USNM 401463, 20 (7 CS), 43.8–85.6 mm SL, swamp draining to Lae

Kumbi River (tributary of Alas River), 02º39’05’’N, 097º51’55’’E, D. N. Lumbantobing et al. 20 July 2006; ZMA

102.393, 10, 65.9–97.3 mm SL, Air Runding, Padang Benedenlanden, E. Jacobson, November 1913. Kabupaten

Aceh Tenggara: MZB 4505, 17, 39.7–78.7 mm SL, Ketambe, Sungai Jamur Geuleu (65 km from Kutacane), I.

Rachmatika, 7 Mar 1982; MZB 4516, 2, 47.5 and 50.2 mm SL, Ketambe, Sungai Alas, Ninik. S., 3 Mar 1982;

MZB 4518, 26, 31.6–77.2 mm SL, Ketambe, Sungai Alas, Soetikno and D. Hardjono, 9 Mar 1982; USNM 404352,

3, 51.7–62.7 mm SL, fish market in Kutacane, D. N. Lumbantobing et al. 2 July 2006; USNM 401211, 1, 63.0 mm

SL, Kampung Air Kelabu, Alas River near road between Kutacane and Blangkejeren, 03º42’69’’N, 097º38’02’’E,

D. N. Lumbantobing, R. K. Hadiaty, D. Rudaya, and N. M. Ray, 2 July 2006. Kabupaten Aceh Tengah: MZB 5364,

4, 62.5–81.9 mm SL, Krueng Owaq, Kecamatan Lingge, D. Wowor, 26 Jan 1984. Kabupaten Nagan Raya: USNM

401210, 1, 34.1 mm SL, Seumayam River, 03º58’15’’N, 096º39’13’’E, D. N. Lumbantobing et al. 11 July 2006.

Province of Sumatera Utara (North Sumatra): USNM 401209, 2, 16.2–20.7 mm SL, Kabupaten Tapanuli Selatan,

Kecamatan Batang Toru, Desa Garoga, Aek Garoga River, 01°30’95”N, 098°59’39”E, D. N. Lumbantobing, D.

Rudaya, and N. M. Ray, 25 July 2006; ZMA 102.395, 5, 59.3–66.2 mm SL, West Nias (Nias Island), Kleinoeg de

Zwaan.

Diagnosis. Rasbora arundinata is distinguished from all congeners in having a unique black midlateral stripe

overall forming a reed-leaf-like profile, which consists of: a subdorsal blotch that starts tapering anteriorly from the

vertical through the dorsal-fin origin where its dorsal margin appears to be slightly crested, and the tapering

extends further anteriorly reaching midhumeral region via a long pointed anterior tip; a midhumeral diffuse patch

in which melanophores concentrate right below the subdorsal blotch and extends anteriorly in a somewhat diffuse

fashion to reach the midhumeral region; and a posterior-portion stripe that terminates posteriorly at the area

immediately dorsal to the anterior apex of the triangular basicaudal blotch. Rasbora arundinata can be

distinguished further from its congeners by the combination of all the following characters listed in the description

section of this species.

Description. General appearance shown in Fig. 2A. Morphometric and meristic data given in Table 1.

Dorsohypural distance equal to distance from dorsal-fin origin to area between vertical through anterior margin and

posterior margin of eye. Limit between head and trunk indistinct in lateral view. Cephalic tubercles absent. Lateral

line complete (all scales perforated; 24–26 + 3–5). Dorsal-fin origin over 13th lateral-line scale. Tip of adpressed

pectoral fin barely reaching vertical through pelvic-fin insertion. Pelvic fin inserted below 12th lateral-line scale

and distinctly anterior to vertical through dorsal-fin origin. Tip of adpressed pelvic fin extending past anal opening

almost to anal-fin origin; in larger specimens reaching anal-fin origin. Anal-fin origin below 18th or 19th lateral-

line scale.

Coloration in alcohol. General body coloration in alcohol preservation shown in Fig. 2A. Schematic lateral

body pigmentation shown in Fig. 7A. Lachrymal region superficially pigmented with scattered small melanophores

more concentrated peripherally. Dusky gular pigmentation reaching posteriorly to vertical through rictus. Occipital

region with two superficial lines and one deeply-embedded solid line in between. Post-opercular streak thick and

prominent with dense melanophores; situated posterior to and running along pectoral girdle and reaching ventrally

to axillary lobe of pectoral fin. Axillary lobe pigmented with more sparsely-distributed and stellate melanophores

as far as subdistal portion. Mid-dorsal stripe one-fourth scale wide and extending from nape to dorsal part of caudal

peduncle.


TABLE 1. Morphometric and meristic data for Rasbora arundinata and R. haru.

Rasbora arundinata

n = 31

Rasbora haru

n = 25

Range Mean ± SD Range Mean ± SD

Standard length (mm) 42.7–86.3 68.8 ± 14.2 26.5–80.3 56.7 ± 13.8

Percentage of standard length

Total length 135.0–142.7 139.2 ± 1.7 136.6–144.3 140.0 ± 1.7

Head length 24.7–28.3 26.3 ± 1.0 26.2–31.7 28.5 ± 1.3

Predorsal length 53.7–60.0 56.9 ± 1.4 52.0–60.1 56.4 ± 1.6

Preanal length 69.4–75.6 72.1 ± 1.5 68.8–76.5 73.6 ± 1.9

Prepelvic length 48.0–54.4 50.6 ± 1.4 47.7–54.7 51.7 ± 1.6

Dorsal depth 21.9–27.6 24.8 ± 1.3 24.6–30.5 27.1 ± 1.5

Body depth 22.1–28.9 26.3 ± 1.5 25.5–32.4 29.7 ± 1.6

Caudal-peduncle depth 12.8–15.3 13.9 ± 0.7 13.3–16.1 14.9 ± 0.7

Caudal-peduncle length 15.0–18.9 16.9 ± 1.0 12.9–17.8 15.2 ± 1.3

Dorsal-fin base length 10.9–12.6 11.7 ± 0.6 11.1–15.1 13.1 ± 0.9

Anal-fin base length 10.6–13.1 11.8 ± 0.6 10.8–14.1 12.5 ± 0.8

Pelvic-fin length 19.3–23.6 21.6 ± 1.2 19.0–25.3 21.7 ± 1.4

Pectoral-fin length 23.1–29.5 25.2 ± 1.3 22.1–28.7 26.1 ± 1.4

Upper caudal lobe length 30.4–40.6 35.0 ± 2.0 33.4–40.0 36.7 ± 1.8

Median caudal length 14.3–19.7 16.3 ± 1.4 14.1–20.0 17.7 ± 1.4

Lower caudal lobe length 32.2–40.5 37.3 ± 1.9 34.6–41.6 38.4 ± 1.9

Dorsohypural distance 44.8–52.1 47.5 ± 1.6 47.8–54.5 50.4 ± 1.6

Percentage of head length

Eye diameter 26.6–35.0 29.7 ± 1.9 26.5–37.1 30.4 ± 2.5

Snout length 28.3–33.7 30.7 ± 1.3 25.0–32.5 28.9 ± 1.6

Head width 47.9–58.0 52.6 ± 2.3 45.2–55.2 52.2 ± 2.7

Head depth 59.7–72.9 66.4 ± 3.3 64.2–71.8 68.4 ± 1.9

Interorbital width 25.5–33.8 29.3 ± 1.8 23.7–33.6 29.3 ± 2.7

Range Mode (n) Range Mode (n)

Meristics

Gill rakers on 1st gill arch 11–12 12 (6) 10–11 10 (3)

Pharyngeal teeth formula 5,4,2 5,4,2 (6) 5,4,2 5,4,2 (3)

Dorsal-fin rays i, 7½ i, 7½ (12) i, 7½ i, 7½ (10)

Pectoral-fin rays i, 11–13 i, 12 (12) i, 12–13 i, 12 (10)

Pelvic-fin rays i, 7–9 i, 8 (12) i, 8 i, 8 (10)

Anal-fin rays ii, 5½ ii, 5½ (15) ii, 5½ ii, 5½ (15)

Dorsal procurrent rays of caudal fin 7–8 8 (12) 7–8 8 (10)

Dorsal principal rays of caudal fin 9 9 (12) 8–9 9 (10)

Ventral procurrent rays of caudal fin 8 8 (12) 8 8 (10)

Ventral principal rays of caudal fin 6–8 8 (12) 8 8 (10)

Scales of lateral-line series 24–25 + 3–4 25 + 4 (15) 23–25 + 3–5 24 + 4 (15)

Predorsal scales 12–13 12 (15) 10–12 11 (15)

Transverse scales ½4, 1, 2½ ½4, 1, 2½ (15) ½4, 1, 2½ ½4, 1, 2½ (15)

Circumpeduncular scales 12 12 (15) 12 12 (15)

Vertebrae (total) 33–34 33 (15) 32–33 33 (10)

Prehaemal vertebrae 16–17 16 (10) 16 16 (10)

Haemal vertebrae 17 17 (10) 16–17 17 (10)

Peduncular vertebrae 8–9 9 (10) 8 8 (10)


FIGURE 4. Photographs of living specimen of Rasbora arundinata, USNM 391607 (paratype).

Peripheral reticulation distinct and covering at maximum four longitudinal scale rows along dorsolateral

portion of body. Basal reticulation distinct and covering up to five longitudinal scale rows and also dorsal scale

row, with network of independent chevron-shaped bars. Peripheral and basal reticulation overlapping on first to

fourth longitudinal scale rows and also on dorsal scale row. Black midlateral stripe prominent, more intense on

central portion forming somewhat wedge-shaped subdorsal blotch, and overall forming reed-leaf-like profile.

Stripe slightly angled anteroventrally and strongly tapering anteriorly until becoming obscure thin line reaching

mid-humeral region. Stripe slightly attenuating posteriorly with ventral margin slightly ascending and extending to

black basicaudal blotch. Axial streak posteriorly overlapping dorsal margin of black midlateral stripe, but streak

separate from stripe in area above 12th or 13th lateral-line scale. Streak decreasing in intensity anteriorly until

diffusing above 7th lateral-line scale. Longitudinal light area indistinct, most visible along area adjacent to

posterodorsal portion of axial streak and black midlateral stripe, but covered with reticulation.

Deeply-embedded diamond-shaped black basicaudal blotch confluent anteromedially with black midlateral

stripe. Blotch consisting two elements: basicaudal triangular patch and basicaudal spot. Basicaudal triangular patch

confluent anteriorly with black midlateral stripe, originating posterior to hypural notch, and flaring posteriorly. In

some specimens, triangular expansion not well-developed. Basicaudal spot confluent anteriorly with and appearing

darker than triangular patch, and terminating anterior to medial sheath scale of caudal fin. Supranal pigmentation

distinct appearing as a somewhat tear-shaped black patch smaller than pupil, originating slightly posterior to

vertical through anal-fin origin, and terminating at vertical through base of third branched anal-fin ray.

Subpeduncular pigmentation dusky, slightly decreasing in intensity anteriorly. Distal edge of caudal fin pigmented

with scattered melanophores resulting in narrow dusky striped margin.

Coloration in life. Ground coloration of dorsolateral surface of head and body pale brown with slightly silvery

sheen, ventral surface barely pigmented with whitish reflective guanine (Fig. 4A–C). Dorsum of head largely

dusky, with scattered yellowish reflective patches and streaks on snout, lateral-line canals, meningeal layer, and

supraorbital. Anterodorsomedial portions of operculum and upper end of gill slit with reflective yellowish patches

(Fig. 4C). Reflective midlateral stripe metallic yellowish, juxtaposed ventrally by black midlateral stripe.

Reflective stripe prominent on anterior portion of trunk, continuously extending from humeral region, slightly

widening posteriorly until above 9th or 10th lateral-line scale, then tapering posteriorly until appearing as thin

reflective line below dorsal fin, and terminating at hypural notch (Fig. 4A–B). Black peripheral reticulation

bordered anteriorly by metallic yellowish sheen overall appearing as reticulated pattern of yellowish reflective

crescents. All fins hyaline. Dorsal and caudal fins with yellowish sheen on subdistal portion of branched rays.

Habitat and distribution. The specimens of Rasbora arundinata were collected in various types of habitat,


such as gravel-bottomed mountain streams, moderate-flowing turbid rivers, and muddy backwater pools. This

species is known from the Tripa Jaya, Kluet, and Alas rivers that flow into the Indian Ocean in the southern part of

northwestern Sumatra (Fig. 8). Rasbora arundinata was collected sympatrically with Rasbora api, R. kluetensis

(only in the Kluet River), R. jacobsoni, and R. truncata (only in the Alas River).

Etymology. The specific name, arundinata, an adjective from the Latin ‘arundo’ for reed, refers to the shape

of the black midlateral stripe of the species appearing like a reed leaf.

Rasbora haru, new species

(Figures 2B, 7B)

Rasbora lateristriata var. sumatrana (non Bleeker, 1854): Weber & de Beaufort 1916.

Rasbora spilotaenia (non Hubbs and Bleeker, 1954): Kottelat & Vidthayanon 1993; Ott 2009.

Holotype. MZB 17888 (ex. ZRC 51986), female, 65.4 mm SL, Indonesia, Sumatra, Province of Sumatera Utara,

Kabupaten Karo, Lau Kawar (catch from local anglers), 03º11.816’N, 98º23.420’E, 1441 m asl, T. Sim et al. 12

April 2009.

Paratypes. All from Indonesia: collected with holotype: ZRC 51986, 4, 22.4–61.8 mm SL. Sumatra: BMNH

1889.11.12.81, 1, 47.1 mm SL, District of Deli; ZMA 102.394, 25, 25.8–58.6 mm SL, Deli, de Bussy; ZMA

102.403, 2, 58.4 and 76.9 mm SL, Tandjong, December 1984; ZMA 102.402, 1, 57.7, Battak hooglande by

Rampong Brastagei, de Bussy; ZMA 102.404, 2, 49.9–63.4 mm SL, Serdang, Sei Poetih, V. Dedem, 10 September

1909; ZMA 119.515, 13, 31.2–74.4 mm SL, Boven Langkat, Gloegoer River, small creek with sandy clay bottom,

upstream of Bohorok, J. E. A. den Doop, Aug 1917. Province of Sumatera Utara: Kabupaten Karo: Bianco and M.

Kottelat: CMK 4429, 6, 49.4–56.1 mm SL, Lau Santam, about 1 km from Pernangenem (5–10 km South of Penen),

17 November 1984; CMK 4447, 1, 103.3 mm SL, Sungai Bluei above Segugi, 18 November. 1984; CMK 4461, 14,

12.8–43.8 mm SL, Rindu River at Permandin, 30 km south of Medan on the road to Kabanjahe, 19 November

1984. Kabupaten Langkat: D. Wowor: Sekundur: Sungai Besitang: MZB 4468, 7, 62.8–79.9 mm SL, Alur Sungai

Tenang;, 25 October 1981; MZB 4476, 16, 25.8–74.2 mm SL, 26 October 1981; MZB 4535, 7, 58.1–87.2 mm SL,

23 October 1981. Sungai Bohorok: MZB 4494, 7, 42.7–92.5 mm SL, 6 November 1981. Kecamatan Bohorok:

Desa Bukit Lawang: Bohorok River: Haryono and Saptono: MZB 11848, 43, 36.6–63.6 mm SL, 16 December

1999; MZB 11849, 21, 26.5–69.2 mm SL, 16 December 1999; USNM 408393 (ex. MZB 11849), 10 (3 CS), 40.9–

55.6 mm SL; ZRC 53198 (ex. MZB 11849), 10, 31.7–54.2 mm SL; MZB 11850, 16, 33.0–56.9 mm SL, a sewer

near the rubber plantation, 12 December 1999; MZB 11852, 22, 31.9–59.7 mm SL, a sewer near Izumi, 16

December 1999. Desa Timbang Lawan: Haryono: MZB 11851, 3, 54.3–63.6 mm SL, 15 December 1999.

Diagnosis. Rasbora haru is distinguished from congeners in having the black midlateral stripe slightly

descending anteriorly along its length until abruptly ending above the 6th or 7th lateral-line scale, attenuating

posteriorly, barely reaching the black triangular basicaudal blotch, overall forming a stamen-like or a wedge-

shaped profile with a posterior apex slightly tapering. Rasbora haru further differs from three other species of the

Sumatrana group in northern Sumatra (Rasbora arundinata, R. maninjau, and R. bindumatoga) in having: fewer

gill rakers on the first gill arch with a mode of 10 (vs. 11–13); a distinct limit between the head and the trunk

characterized by abrupt convexity along the anterior portion of the trunk; a relatively deep body with a mean of

29.7% SL (vs. 24.1–27.3%); and a relatively deep caudal peduncle with a mean of 14.9% SL (vs. 11.9–14.0%).

Rasbora haru can be distinguished further from its congeners by the combination of all the following characters

listed in the description section of this species.

Description. General appearance shown in Fig. 2B. Morphometric and meristic data given in Table 1.

Dorsohypural distance equal to distance from dorsal-fin origin to area between tip of snout and vertical through

anterior margin of nostril. Limit between head and trunk distinguishable by abrupt convexity of anterior predorsal

profile relative to slant of head profile. Cephalic tubercles observable from one specimen of male, relatively small

and very few, distributed on dorsal surface of head especially on supraorbital area. Lateral line complete (all scales

perforated; 23–25 + 3–5). Dorsal-fin origin situated over 11th or 12th lateral-line scale. Tip of adpressed pectoral

fin extending beyond vertical through pelvic-fin insertion. Pelvic fin inserted below 10th or 11th lateral-line scale

and distinctly anterior to vertical through dorsal-fin origin. Tip of adpressed pelvic fin reaching anal-fin origin.

Anal-fin origin located below 17th or 18th lateral-line scale.


Coloration in alcohol. General body coloration in alcohol preservation shown in Fig. 2B. Schematic lateral

body pigmentation shown in Fig. 7B. Lachrymal region superficially pigmented with dorsal portion more densely

pigmented. Dusky gular pigmentation extending posteriorly to vertical through rictus. Fleshy opercular flap

pigmented with scattered small melanophores. Background pigmentation overall dusky and reticulated. Peripheral

reticulation relatively thick and very distinct and covering at maximum 41/2 longitudinal scale rows along

dorsolateral and midlateral portions of body, and also dorsal scale row. Basal reticulation relatively thick and

prominent, covering up to 5 longitudinal scale rows and also to dorsal scale row, with network of continuous

parenthesis-shaped bars. Peripheral and basal reticulations not overlapping along lateral-line scales.

Black midlateral stripe prominent, more intense on central portion, and overall appearing as stamen-like

profile. Stripe slightly narrowing anteriorly until abruptly ending over 6th or 7th lateral-line scale, attenuating

posteriorly until appearing as thin line of melanophores and extending to form trace of scattered melanophores

along caudal peduncle reaching black basicaudal blotch. Black basicaudal blotch consisting two elements:

basicaudal triangular patch and deeply-embedded darker basicaudal spot. Triangular patch of black basicaudal

blotch originating posterior to hypural notch and flaring posteriorly at maximum to distance equal to one-half of

caudal-peduncle depth until becoming confluent with basicaudal spot. Basicaudal spot terminating anterior to last

sheath scale of caudal fin. Axial streak prominent and posteriorly bordering dorsal margin of black midlateral

stripe, but streak separate from stripe below area between posterior terminus and mid-portion of dorsal-fin base.

Streak decreasing in intensity anteriorly until disappearing above 7th lateral-line scale. Dusky dorsolateral stripe

very faint, most visible along anterior half of trunk. Longitudinal light area indistinct, most visible along anterior

half of trunk ventral to axial streak. Supra-anal pigmentation distinct, appearing as thin stripe originating slightly

posterior to anal-fin origin and terminating at vertical through base of last branched anal-fin ray. Subpeduncular

pigmentation barely visible. Distal border of caudal fin pigmented with scattered melanophores.

Habitat and distribution. This species is known from the Bohorok River that flows into the Malacca Strait in

the northeastern part of Sumatra (Fig. 8).

Etymology. The specific name, haru, a noun in apposition, is derived from the name of an old kingdom in the

area currently known as Deli, where the species lives.

Rasbora maninjau, new species

Figures 2C, 5A–B, 9B

Rasbora lateristriata var. sumatrana (non Bleeker, 1854): Weber & Beaufort 1916.

Rasbora lateristriata (non Bleeker, 1854): Kottelat & Vidthayanon 1993.

Holotype. MZB 17890, 66.9 mm SL, Indonesia, Province of Sumatera Barat (West Sumatra), Kabupaten Agam,

Kecamatan Tanjung Raya, by Hotel Danau Maninjau, 6 August 2006.

Paratypes.Same data as holotype: MZB 21120, 1, 57.0 mm SL; USNM 406859, 1 (CS), 58.8 mm SL.

Non-types. ZMA 102.400, 8, 31.3–66.0 mm SL, Meer van Manindjau, M. Weber, 1888.

Diagnosis. Rasbora maninjau is distinguished from congeners by the combination of the following characters:

the black midlateral stripe extending from the midhumeral region with a relatively uniform width and lacking the

subdorsal blotch; the prominent acutely triangular basicaudal blotch; and the oval supra-anal pigmentation.

Rasbora maninjau is further distinguished from the three other species of the Sumatrana group in northern Sumatra

(Rasbora arundinata, R. haru, and R. bindumatoga) in having: 26–27 + 3–4 lateral-line scales (vs. 24–25 + 4;

Table 2); 13 gill rakers on the first gill arch (vs. 10–12); 34 vertebrae (vs. 32–33); a relatively narrow body with a

mean of 24.1% SL (vs. 26.3–29.7%); and a relatively narrow caudal peduncle with a mean of 11.9% SL (vs. 13.9–

14.9%). Rasbora maninjau can be distinguished further from its congeners by the combination of all the following

characters listed in the description section of this species.

Description. General appearance shown in Figure 2C. Morphometric and meristic data given in Table 2.

Dorsohypural distance equal to distance from dorsal-fin origin to nostril. Limit between head and trunk distinct in

lateral view. Cephalic tubercles present on males. No female specimen examined. Tubercles relatively small,

distributed over most of head surface, extending onto nape region as far as second predorsal scale and to several

scales on anteroventral region of body. Lateral line complete (all scales perforated; 26–27 + 3–4). Dorsal-fin origin


situated over 13th lateral-line scale. Tip of adpressed pectoral fin barely reaching vertical through pelvic-fin

insertion. Pelvic fin inserted below 12th lateral-line scale and distinctly anterior to vertical through dorsal-fin

origin. Tip of adpressed pelvic fin barely reaching vertical through anal opening. Anal-fin origin located below

18th lateral-line scale.

TABLE 2. Morphometric and meristic data for Rasbora maninjau and R. bindumatoga.

Rasbora maninjau

n = 7

Rasbora bindumatoga

n = 22


Standard length (mm) 57.3–82.7 68.4 ± 9.0 23.3–72.0 48.6 ± 12.5

Percentage of standard length

Total length 130–136.3 134.5 ± 2.1 137.1–141.9 139.0 ±1.3

Head length 24.3–26.0 24.9 ± 0.7 24.1–31.3 27.7 ± 1.5

Predorsal length 52.7–56.2 54.2 ± 1.1 51.6–57.0 54.3 ± 1.3

Preanal length 68.9–73.7 71.0 ± 1.5 68.7–74.1 71.9 ± 1.5

Prepelvic length 47.1–52.7 49.6 ± 2.2 47.7–72.5 51.7 ± 4.9

Dorsal depth 21.4–24.1 23.0 ± 1.0 23.1–28.6 26.0 ± 1.7

Body depth 21.3–26.5 24.1 ± 1.6 25.0–29.9 27.3 ± 1.3

Caudal-peduncle depth 11.0–12.6 11.9 ± 0.5 12.0–15.1 14.0 ± 0.7

Caudal-peduncle length 16.5–19.4 18.3 ± 1.1 14.5–17.9 16.3 ± 0.9

Dorsal-fin base length 11.1–12.6 11.9 ± 0.5 11.4–15.0 12.9 ± 0.9

Anal-fin base length 11.0–12.6 12.0 ± 0.6 11.0–13.1 12.3 ± 0.6

Pelvic-fin length 17.1–20.3 18.7 ± 1.5 17.6–22.0 19.9 ± 1.1

Pectoral-fin length 21.0–22.5 21.7 ± 0.5 22.2–25.7 24.3 ± 1.0

Upper caudal lobe length 30.2–34.0 32.5 ± 1.5 32.8–40.1 36.7 ± 2.0

Median caudal length 14.0–16.3 15.5 ± 0.9 16.1–21.0 18.3 ± 1.2

Lower caudal lobe length 30.7–36.8 34.1 ± 2.5 35.4–42.8 37.4 ± 1.8

Dorsohypural distance 48.0–51.3 50.0 ± 1.3 48.1–54.7 50.8 ± 1.7

Percentage of head length

Eye diameter 27.1–32.2 29.2 ± 2.0 23.9–33.1 30.5 ± 2.0

Snout length 26.7–30.5 29.2 ± 2.0 26.8–31.3 29.0 ± 1.4

Head width 47.1–52.1 50.5 ± 1.8 49.0–58.2 54.1 ± 2.6

Head depth 63.4–69.2 65.2 ± 1.9 63.3–74.1 68.1 ± 3.0

Interorbital width 25.3–28.7 26.7 ± 1.2 25.7–32.6 29.7 ± 2.1

Range Mode (n) Range Mode (n)

Meristics

Gill rakers on 1st gill arch 13 13 (1) 11–12 11 (4)

Pharyngeal teeth formula 5,4,1 5,4,1 (1) 5,4,2 5,4,2 (4)

Dorsal-fin rays i, 7½ i, 7½ (7) i, 7½ i, 7½ (10)

Pectoral-fin rays i, 11–13 i, 12 (7) i, 11–13 i, 12 (10)

Pelvic-fin rays i, 8 i, 8 (7) i, 8 i, 8 (10)

Anal-fin rays ii, 5½ ii, 5½ (7) ii, 5½ ii, 5½ (10)

Dorsal procurrent rays of caudal fin 7 7 (7) 7–8 8 (10)

Dorsal principal rays of caudal fin 9 9 (7) 9 9 (10)

Ventral procurrent rays of caudal fin 8 8 (7) 8 8 (10)

......continued on the next page


FIGURE 5. Photographs of living specimen of Rasbora maninjau, MZB 17890 (holotype).

Coloration in alcohol. General body coloration in alcohol in alcohol preservation shown in Fig. 2C.

Schematic lateral body pigmentation shown in Fig. 9B. Lachrymal region superficially pigmented with peripheral

portion more densely pigmented. Dusky gular pigmentation decreasing in intensity posteriorly until reaching to

anterior portion of branchiostegal flaps. Peripheral reticulation very prominent and covering at maximum 51/2

longitudinal scale rows along dorsolateral, midlateral, and ventrolateral portions of body. Basal reticulation very

prominent and covering at maximum 5 longitudinal scale rows, with network of continuous parenthesis-shaped

bars. Peripheral and basal reticulation maximally overlapping on first to fifth longitudinal scale rows and also on

dorsal scale row.

Black midlateral stripe prominent, more intense posteriorly, and of uniformly approximately one-third scale

wide along posterior of flank until reaching vertical through mid-point of dorsal-fin base. Stripe slightly tapering

anteriorly below dorsal fin with ventral margin becoming diffuse with less concentrated melanophores until

completely replaced by swath of stellate melanophores reaching pectoral girdle along almost all of anterior half of

flank. Dorsal edge of stripe bordered by axial streak from hypural notch to midhumeral region. Deeply-embedded

diamond-shaped black basicaudal blotch confluent with black midlateral stripe anteromedially and terminating at

anterior sheath scales of caudal fin. Blotch consisting of two elements: basicaudal triangular patch originating

posterior to hypural notch and flaring posteriorly to maximum one-fourth of caudal-peduncle depth; and darker

TABLE 2. (Continued)

Rasbora maninjau

n = 7

Rasbora bindumatoga

n = 22


Ventral principal rays of caudal fin 7 7 (7) 7–8 8 (10)

Scales of lateral-line series 26–27 + 3–4 27 + 4 (7) 24–25 + 3–4 24 + 4 (15)

Predorsal scales 12–13 12 (3) 11–12 12 (15)

Transverse scales ½4, 1, 2½ ½4, 1, 2½ (7) ½4, 1, 2½ ½4, 1, 2½ (15)

Circumpeduncular scales 12 12 (7) 12 12 (15)

Vertebrae (total) 34 34 (3) 32–33 33 (10)

Prehaemal vertebrae 17 17 (3) 16 16 (10)

Haemal vertebrae 17 17 (3) 16–17 17 (10)

Peduncular vertebrae 9 9 (3) 8 8 (10)


basicaudal spot terminating anterior to sheath scale of caudal fin. Axial streak posteriorly juxtaposed dorsal margin

of black midlateral stripe, but separating from stripe in area above 11th lateral-line scale. Streak decreasing in

intensity anteriorly until disappearing on mid-humeral region. Dusky dorsolateral stripe somewhat distinct, most

prominent and with maximum depth of almost one scale wide along anterior portion of trunk; slightly tapering

posteriorly and bordered ventrally by longitudinal light area. Longitudinal light area somewhat obscure, masked by

dusky pigmentation on each anterior portion of scale, extending longitudinally between dusky and black lateral

stripes, most prominent along mid-point of body to caudal peduncle. Supra-anal pigmentation originating posterior

to anal-fin origin and terminating at base of 2nd branched anal-fin ray, appearing as distinct ellipsoidal blotch.

Subpeduncular pigmentation distinct, most intense along mid-point of subpeduncular region. Longest ray of each

caudal-fin lobe covered by dense melanophores along its length. Distal edge of caudal fin pigmented with scattered

melanophores resulting in narrow dusky striped margin.

Coloration in life. Ground coloration of dorsolateral surface of head and body pale brown, midlateral and

ventral surfaces whitish to silvery due to presence of reflective guanine (Fig. 5A–C). Dorsum of head largely

dusky, with several faint scattered yellowish patches and streaks on snout, lateral-line canals, meningeal layer, and

occipital region (Fig. 4B–C). Black pigmentation on lateral body somewhat obscure, most visible posteriorly, with

metallic greenish to bluish sheen. Reflective midlateral stripe metallic orange-reddish, extending from humeral

region to caudal peduncle with relatively equivalent width, juxtaposed ventrally by obscure black midlateral stripe,

and terminating at distinct black basicaudal blotch. Black reticulation pattern relatively distinct. All fins hyaline.

Habitat and distribution. Specimens of Rasbora maninjau were collected in Lake Maninjau, a crater lake in

central western Sumatra draining to the Indian Ocean (Fig. 8). This species is a favorite among local people for

food and is known as ‘ikan badar’ in Minang, the native language of the area.

Etymology. The specific name, maninjau, a noun in apposition, refers to the name of the type locality, Lake

Maninjau.

Rasbora bindumatoga, new species

Figures 2D, 6A–C, 10C

Holotype. MZB 17889, female, 47.1 mm SL, Indonesia, Sumatra, Province of Sumatera Utara (North Sumatra),

Kabupaten Tapanuli Selatan, Batang Angkola River, 01°09’90”N, 099°24’83”E, D. N. Lumbantobing and D.

Rudaya, 4 August 2006.

Paratypes. All from Indonesia: collected with holotype: MZB 21117, 7, 23.3–60.6 mm SL; USNM 404351, 7

(3 CS), 41.5–49.1 mm SL; ZRC 53199, 4, 25.2–46.6 mm SL. Sumatra: Province of Sumatera Utara (North

Sumatra): CMK 4528, 16, 21.0–67.6 mm SL, road from Porsea to Pulau Raya, 33 km before Pulau Raya, Bianco

and M. Kottelat, 27 November. 1984; MZB 21118, 1, 57.6 mm SL, Kabupaten Tapanuli Utara, Aek Dahasan, on

the road from Onanhasang to Sipirok, 01°52’25”N, 099°03’64”E, D. N. Lumbantobing, D. Rudaya and N. M. Ray,

26 July 2006; MZB 21119, 1, 72.0 mm SL; Kabupaten Toba Samosir, Aek Silang River (a tributary draining to

Lake Toba), PLTA Pandan, 02°18’50”N, 098°44’51”E, D. N. Lumbantobing, D. Rudaya and N. M. Ray, 31 July

2006; USNM 390346, 1, 29.5 mm SL, Kabupaten Humbang-Hasundutan, near Doloksanggul, upstream of Aek

Sibundung River, 02º15.90’N/98º43.75’E, D. N. Lumbantobing, D. Rudaya and N. M. Ray, 31 July 2006; USNM

401208, 1, 30.2 mm SL, Kabupaten Mandailing Natal, Aek Siburiang River (a tributary of Batang Gadis River),

00°41’13”N, 099°40’11”E, D. N. Lumbantobing, D. Rudaya and N. M. Ray, 5 August 2006.

Diagnosis. Rasbora bindumatoga is distinguished from congeners by the following combination of characters:

the black midlateral stripe is only represented by a rectangular subdorsal blotch without a midhumeral diffuse patch

and the posterior-portion stripe, the absence of supra-anal pigmentation, and the basicaudal blotch appearing

somewhat oval and lacking the basicaudal triangular patch. Rasbora bindumatoga can be distinguished further

from its congeners by the combination of all the following characters listed in the description section of this

species.

Description. General appearance shown in Figure 2D. Morphometric and meristic data given in Table 2.

Dorsohypural distance equal to distance from dorsal-fin origin to area between anterior portion of snout and

posterior margin of pupil. Limit between head and trunk indistinct in lateral view, but more visible in smaller

specimens (~40 mm SL). Cephalic tubercles present on males, relatively small and few in number. Tubercles


distributed on dorsal surface of head from snout to occiput. Lateral line complete (all scales perforated; 24–26 + 3–

4). Dorsal-fin profile blunt with first branched ray being longest dorsal-fin ray. Dorsal-fin origin situated over 12th

lateral-line scale. Tip of adpressed pectoral fin nearly reaching vertical through pelvic-fin insertion. Pelvic fin

inserted below 11th lateral-line scale and distinctly anterior to vertical through dorsal-fin origin. Tip of adpressed

pelvic fin extending past anus and in some specimens reaching to anal-fin origin. Anal-fin origin located below

17th lateral-line scale.

FIGURE 6. Photographs of living specimen of Rasbora bindumatoga, MZB 21117 (paratype).

Coloration in alcohol. General body coloration in alcohol in alcohol preservation shown in Fig. 2D.

Schematic lateral body pigmentation shown in Fig. 10C. Lachrymal region superficially pigmented with more

peripherally-distributed melanophores. Dusky gular pigmentation decreasing in intensity posteriorly until reaching

to vertical through rictus. Peripheral reticulation distinct and covering at maximum 41/2 longitudinal scale rows

along dorsolateral and midlateral portions of body. Basal reticulation prominent and covering up to five

longitudinal scale rows and also to dorsal scale row, with network of somewhat continuous chevron-shaped bars.

Peripheral and basal reticulation overlapping at maximum on first to fifth longitudinal rows and also on dorsal

scale row.

Black midlateral stripe rudimentary and replaced by black subdorsal blotch dorsally in contact with anterior

portion of axial streak. Blotch rectangular or trapezoid, deeply embedded under approximately two scales at its

maximum length and 11/2 scales at its maximum depth. Distinct black axial streak extending from hypural notch to

area above subdorsal blotch, anteriorly diffusing above anterior margin of blotch, and terminating above pelvic-fin

insertion. Interspersed swath of melanophores extending along and bordered dorsally by axial streak resulting in

faint dusky midlateral stripe. Deeply embedded black basicaudal blotch situated on medial portion of posterior

margin of hypural plate with about two scales deep and 11/2 scales long resulting in somewhat ovoid mark. Dusky

dorsolateral stripe indistinct and visible along posterior portion of trunk bordered ventrally by axial streak.

Longitudinal light area indistinct, most visible along posterodorsal portion interspersing by reticulated

pigmentation. Supra-anal pigmentation absent. Subpeduncular faint. Distal edge of caudal fin pigmented with

scattered melanophores resulting in narrow dusky striped margin.

Coloration in life. Ground coloration of dorsal surface of head and body pale brown, ventrolateral surface of

head and body grey to whitish with silvery sheen due to guanine (Fig. 6A–C). Dorsum of head largely dusky, with

scattered yellowish to greenish reflective patches and streaks on snout, lateral-line canals, meningeal layer, and

supraorbital. Anterodorsomedial portions of operculum and upper end of gill slit with reflective yellowish patches

(Fig. 6C). Black pigmentation on lateral body very faint and appearing bluish grey, except for reticulated pattern

and basicaudal blotch relatively distinct. Yellowish reflective middorsal stripe present and distinct on dorsum of


body, extending from nape to dorsal caudal peduncle (Fig. 6B). Reflective midlateral stripe absent. All fins hyaline.

Dorsal, anal, and caudal fins with yellowish sheen on subdistal portion of branched rays.

Habitat and distribution. The specimens of Rasbora bindumatoga were collected in gravel-bottomed

mountain streams and moderate-flowing turbid rivers. This species is known from the Aek Sibundung, Batang

Toru, and Batang Gadis Rivers that flow into the Indian Ocean in the southern part of northwestern Sumatra, and

also from a tributary draining to Lake Toba (Fig. 8). It was collected sympatrically with R. api.

Etymology. The species epithet, bindumatoga, a noun in apposition, is derived from the term ‘Bindu Matoga’

in the two native languages of northern Sumatra, Mandailing and Toba, which are widely spoken throughout the

distribution range of the species. By the local Mandailing and Toba people, the term ‘Bindu Matoga’ is commonly

used as the name of a unique traditional ornament that has a rectangle outline. The epithet, thus, refers to the

rectangular shape of the black subdorsal blotch of the species.

FIGURE 7. Schematic drawing of left lateral view showing the three primary diagnostic lateral pigmentation characteristics of

members of the Hosii subgroup: (A) Rasbora arundinata; (B) Rasbora haru; (C) R. bunguranensis; (D) R. notura.

Discussion

Taxonomic problems in the Sumatrana group. The taxonomy of the Sumatrana group has been subjected to a

high rate of synonymy among species, primarily due to unclear species delineation and a lack of appreciation in

using highly variable characters potential for species diagnosis, which consequently lead to conflicting

interpretations by different workers. Brittan (1954a, 1954b) considered several nominal species of Rasbora as the


‘geographical races’ of R. sumatrana s.s.. He recognized only three valid species: R. sumatrana, R. volzi, and R.

elegans. Subsequent authors highlighted the variation of lateral pigmentation patterns as useful diagnostic

characters for delimiting species. As a result, several junior synonyms of R. sumatrana according to Brittan

(1954a), R. aurotaenia, R. calliura, R. paviana, R. hosii, and R. vulgaris, were resurrected as valid species (Kottelat

1986; Kottelat & Lim 1995; Kottelat 2001, 2005; Tan & Kottelat 2009). Kottelat & Vidthayanon (1993) increased

the number from three to 13 species.

FIGURE 8. Distribution of species of the Sumatrana group in northwestern Sumatra and adjacent regions: Rasbora arundinata

(blue diamonds), Rasbora haru (green circles), Rasbora maninjau (yellow star), Rasbora bindumatoga (red asterisks), and R.

vulgaris (magenta squares). Symbols with black outline are the localities of holotypes. One filled symbol may represent more

than one locality.

Kottelat (2005) moved several members of the Sumatrana group into a new group, the R. paviana-group (the

Paviana group hereafter), comprised of five species from Indochina and the Malay Peninsula: R. paviana, R.

vulgaris, R. notura, R. hobelmani, and R. dorsinotata. This group is diagnosable based on the possession of a black

midlateral stripe terminated as a blotch at the caudal peduncle. However, the definition of ‘blotch’ (here termed

basicaudal blotch following Lumbantobing 2010) is not well circumscribed. This was the only diagnostic character

used by Kottelat (2005) for the new group, and he merely emphasized the tendency for the members of the group to

have a diamond-shaped basicaudal blotch (his “blotch”). Contrary to Kottelat (2005), who proposed that such a

blotch is restricted to the species in mainland Asia, it is present in nearly all the species of the Sumatrana group,

including the Sundaland species. These exhibit high level of variation in the shape of the blotch, its intensity, and its

position relative to the black midlateral stripe. Considering the broad distribution and variability of the basicaudal

blotch across all species in the group and the limited knowledge on the nature of this character, all species of

Rasbora with such caudal pigmentation, including the five species of the Paviana group sensu Kottelat (2005),


should be classified in the Sumatrana group. Future comparative studies focusing on the details of basicaudal blotch

as well as other details of lateral body pigmentation (e.g., black midlateral stripe and supra-anal pigmentation) will

be critical in solving taxonomic problems in the Sumatrana group and in delimiting species group.

Liao et al. (2010) revised Brittan’s species groups, including the Sumatrana group, according to a tree-based

hierarchy. Given its placement with species of the Sumatrana group, , the Caudimaculata group sensu Brittan

(1954a) was combined with the Sumatrana group. At the same time, Liao et al. (2010) excluded R. elegans from

the Sumatrana group and transferred it to the Einthovenii group, because it was embedded in a clade with R.

einthovenii and R. cephalotaenia, the only two representatives of the Einthovenii group in their analysis. The

inclusion of the Caudimaculata group in the Sumatrana group seems warranted in that all species of the

Caudimaculata group sensu Brittan (1954a), except R. dorsiocellata, possess the three primary diagnostic

characters of the Sumatrana group. The inclusion of R. elegans into the Einthovenii group, however, is not well

supported as there is only one reversed synapomorphy supporting the pertinent clade in the phylogeny of Liao et al.

(2010). Moreover, Liao et al. (2010) overlooked the significance of the basicaudal blotch as a potential

synapomorphy. They did not include it as a character in their phylogenetic analysis, despite the fact that Brittan

(1954a) and Kottelat (2005) considered the blotch to be one of the major diagnostic characters of the Sumatrana

group. Rasbora elegans should be retained in the Sumatrana group until more conclusive relationships following a

robust phylogenetic analysis realized.

Diagnosis and limit of the Sumatrana group. The specimens of Rasbora arundinata, R. haru, and R.

maninjau were identified initially as R. sumatrana, a species with uncertain limits and pronounced polymorphisms

across many different geographical regions. The type locality of R. sumatrana sensu stricto (hereafter R.

sumatrana s. s.) is Solok (West Sumatra), in the Indragiri drainage, from where a series of topotypes were collected

and figured in Kottelat et al. (1993) and Tan & Kottelat (2009). Tan & Kottelat (2009) reported that this species is

restricted to the fast-flowing streams of the interior of Sumatra, and is most abundant in the Batang Hari and

Indragiri River drainages. Nevertheless, because the number of examined specimens representing different

populations, especially from Sumatra, was so limited, Tan & Kottelat (2009) tentatively classified all morphs as R.

sumatrana sensu lato. Based on additional specimens of Sumatran Rasbora from the 2006 expedition and various

older museum collections, the high variability across four allopatric populations of R. sumatrana s. l., each of

which shows consistently distinct differences in the three primary diagnostic characters of the Sumatrana group,

are confirmed herein and, thus, recognized as new species.

All new species of the Sumatrana group living in northern Sumatra are distinguished from the members of

other species groups of Rasbora in the region (R. api, R. jacobsoni, R. kluetensis, R. meinkeni, R. nodulosa, R.

reticulata, R. truncata, and R. vulcanus) by the following characters: the black midlateral stripe is barely confluent

anteriorly with the post-opercular pigmentation; the basicaudal pigmentation is always present and larger than the

pupil [“blotch” sensu Kottelat (2001); Fig. 2]; and the lateral surface of the upper lip is discontinuously exposed

with the submedial portion slightly covered due to the contact between the maxilla and the lower lip (Fig. 3). The

arrangement between the maxilla and the lips, which can be clearly observed through the degree of exposure of the

upper lip in combination with the presence of lachrymal groove (Fig. 3), varies remarkably among the species

groups of Rasbora. In species of the Sumatrana group, the submedial portion of the lateral surface of upper lip, an

area just anterior to the lachrymal groove, is partially covered by the fleshy overlap of the anteroventral portion of

the maxilla with the mediodorsal portion of the lower lip. The lateral surface of the upper lip is partially exposed

along its anteriormost portion (the fleshy anterior portion of the premaxilla) and along its posterior area between

the distinct lachrymal groove and the rictus, a condition that is unique to the species of the Sumatrana group (Fig.

3A). Species of the Einthovenii group have an upper lip that is also partially covered by the anteroventral portion of

the maxilla that is in contact with the mediodorsal portion of lower lip; however, no lachrymal groove borders the

dorsal portion of the upper lip (Fig. 3B).

In further contrast, some species of the Trifasciata group (Rasbora bankanensis, R. ennealepis, R. hubbsi, R.

johannae, R. lacrimula, R. sarawakensis, R. trifasciata, and R. tuberculata) have an upper lip whose anteriormost

portion is covered entirely by the anteroventral portion of the maxilla. Overall, the lateral surface of the upper lip is

only halfway exposed along its posterior portion from the lachrymal groove to the rictus (Fig. 3C). Some other

species of the Trifasciata group (Rasbora api, R. kluetensis, R. meinkeni, R. nodulosa, R. tobana, R. truncata, and

R. vulcanus), all known from northwestern Sumatra, have an upper lip that is entirely exposed laterally because

there is no contact point between the maxilla and the lower lip (Fig. 3D). Given the distinct variation among


species groups of Rasbora, the degree of lateral coverage of the upper lip is useful for diagnosing species groups of

Rasbora. The discontinuity in lateral exposure of the upper lip, in combination with the presence of distinct

lachrymal groove (Fig. 3A), is a new diagnostic character of the Sumatrana group along with the three diagnostic

characters of lateral pigmentation.

Species subgroups in the Sumatrana group. On the basis of similarity in shape of the black midlateral stripe,

the Sumatrana group can be categorized into three subgroups: (1) the Hosii subgroup (Figs. 7 and 11); (2) the

Lateristriata subgroup (Fig. 9); and (3) the Elegans subgroup (Fig. 10). The Hosii subgroup is characterized by the

black midlateral stripe with the subdorsal blotch (the anterior subdorsal portion of the black midlateral stripe that is

wider than the posterior-portion stripe) always present (Figs. 7, 11), and contains 13 species: Rasbora arundinata

(Fig. 7A), R. aprotaenia (Fig. 11C), R. atranus, R. bunguranensis (Fig. 7C), R. cryptica, R. dorsinotata, R. haru

(Fig. 7B), R. hobelmani, R. hosii (Fig. 11A), R. notura, R. spilotaenia (Fig. 11B), R. sumatrana s. s., and R. volzi.

The Lateristriata subgroup is characterized by the black midlateral stripe tapering anteriorly and reaching the

midhumeral region, but lacking any distinct subdorsal blotch (Fig. 7). Thirteen species of the Sumatrana group

belong to the Lateristriata subgroup: R. atridorsalis, R. baliensis, R. calliura, R. caudimaculata, R. elberti, R.

lateristriata (Fig. 9A), R. maninjau (Fig. 9B), R. paviana (Fig. 9D), R. rasbora, R. subtilis, R. tawarensis, R.

trilineata, and R. vulgaris (Fig. 9C). In the Elegans subgroup, the black midlateral stripe transforms into the

rudimentary form lacking the posterior-posterior stripe, which appears as a semirectangular subdorsal blotch below

the dorsal fin (Fig. 10). The Elegans subgroup comprises of three species: R. bindumatoga (Fig. 10C), R. elegans

(Fig. 10A–B), and R. nematotaenia. Due to the poor condition of species examined or unavailability of the

specimens, three species of the Sumatrana group (R. leptosoma, R. taeniata, and R. unicolor) cannot be classified

in any of the three subgroups for certain.

Differential diagnosis using pigmentation pattern. Among the four new species from northern Sumatra

described herein, R. arundinata and R. haru resemble each other the most. Rasbora arundinata (Figs. 2A, 7A) is

distinguished from R. haru (Figs. 2B, 7B) in having the black midlateral stripe tapering anteriorly to form a reed-

leaf-like profile extending posteriorly as a uniformly wide stripe and broadly confluent with the triangular

basicaudal patch (vs. terminated abruptly anteriorly forming a stamen-like profile attenuating posteriorly until

disappearing on the hypural notch and barely confluent with the triangular basicaudal patch) and the supra-anal

pigmentation in the form of a tear-shaped black patch (vs. appearing as a thin elongate streak). Rasbora maninjau

(Figs. 2C, 9B) is similar overall to R. arundinata and R. haru, but readily distinguishable by having a thin black

midlateral stripe of relatively uniform width and lacking a subdorsal blotch (vs. an anteriorly broadening midlateral

stripe with the subdorsal blotch present). Rasbora bindumatoga is the most distinct new species and is

distinguished from the other three species by having the rudimentary form of black midlateral stripe appearing

solely as a black, somewhat rectangular subdorsal blotch, and by lacking supra-anal pigmentation.

In comparison with the other species of the Sumatrana group, R. arundinata (Fig. 7A) is similar overall to R.

bunguranensis from Natuna Island (Fig. 7C), and R. hosii from Bornean Sarawak (Fig.11A), R. notura from

eastern peninsular Malaysia (7D), and R. spilotaenia from southern Sumatra (Fig. 11B). It is distinguished from all

four of these species in having an anteriorly tapering black midlateral stripe, reaching the midhumeral region with

a long pointed tip (vs. terminated anteriorly in a more or less blunt tip without reaching the midhumeral region) and

an axial streak terminating above and barely confluent with the apex of the basicaudal triangular patch (vs.

confluent and terminating at the apex of the basicaudal triangular patch). Rasbora arundinata is also distinguished

from R. notura and R. hosii in having the supra-anal pigmentation appearing as a tear-shaped black patch (vs.

appearing as a thin elongate streak). Rasbora arundinata differs from R. spilotaenia by having a half-scale deep

black midlateral stripe along the posterior half of the body (vs. a thin black line along the posterior half of the

body). Rasbora arundinata is distinguished from R. bunguranensis in having the supra-anal anal pigmentation that

is prominent black (vs. grayish and inconspicuous).

Rasbora haru was initially identified as R. lateristriata (Fig. 9A). Nevertheless, the type locality of R.

lateristriata is West Java, and after comparison with specimens from West Java, the specimens of R. haru are

readily distinguishable from R. lateristriata in having a subdorsal blotch deeper than the posterior-portion stripe

(vs. lacking a subdorsal blotch) and the posterior tip of the black midlateral stripe discontinuous from the anterior

tip of the basicaudal blotch (vs. confluent with the basicaudal blotch). Rasbora haru also superficially resembles R.

notura, R. spilotaenia, R. bunguranensis, and R. hosii, but is distinguishable from all four in having the axial streak

terminating above and barely confluent with the apex of the basicaudal triangular patch (vs. confluent and

terminating at the apex of the basicaudal triangular patch) and the posterior tip of the black midlateral stripe


discontinuous with the anterior tip of the basicaudal blotch (vs. confluent with the basicaudal blotch). Rasbora

haru is further distinguished from R. spilotaenia and R. bunguranensis by the presence of supra-anal pigmentation

forming a thin, elongate streak (vs. an ellipsoidal blotch).

Rasbora maninjau appears most similar to R. vulgaris from western peninsular Malaysia among the other valid

species of the Sumatrana group, but is distinguished from R. vulgaris by the presence of a supra-anal pigmentation

in the form of an ellipsoidal blotch (vs. a thin elongate streak) and a shallow, somewhat triangular basicaudal blotch

(vs. a very deep diamond-shaped blotch). Two other species that resemble R. maninjau are R. lateristriata from

Western Java (Fig. 9A) and R. paviana from the Malay Peninsula (Fig. 9D). Rasbora maninjau can be

distinguished from R. lateristriata by the possession of a black midlateral stripe terminating anteriorly on the

midhumeral region (vs. terminating anteriorly on the postopercular pigmentation) and a dot-shaped supra-anal

pigmentation smaller than the pupil (vs. a semicircular supra-anal pigmentation larger than the pupil).

The specimens of R. bindumatoga were initially identified as R. elegans, a species widely distributed

throughout Peninsular Malaysia and Sundaland (except Java). Rasbora bindumatoga (Fig. 10C) differs from R.

elegans (Figs. 10A–B) in the lack of supra-anal pigmentation (vs. present), the basicaudal triangular patch absent

(vs. present), and the basicaudal spot somewhat diffuse (vs. prominent). Rasbora bindumatoga also appears similar

overall to R. sumatrana s. s. [photograph in Tan & Kottelat (2009)] in color pattern, but is readily distinguishable

from this type species by the absence of the midhumeral diffuse tract (vs. present, continuous with the rectangular

subdorsal patch), the absence of the posterior midlateral stripe (vs. present, but diffuse), and the absence of the

supra-anal pigmentation (vs. present, as a thin, elongate line).

FIGURE 9. Schematic drawing of left lateral view of three primary diagnostic lateral pigmentation characteristics of members

of the Lateristriata subgroup: (A) R. lateristriata; (B) Rasbora maninjau; (C) R. vulgaris; (D) R. paviana.


FIGURE 10. Schematic drawing of left lateral view of three primary diagnostic lateral pigmentation characteristics of

members of the Elegans subgroup: (A) R. elegans from West Johor, Peninsular Malaysia; (B) R. elegans from East Johor,

Peninsular Malaysia; (C) Rasbora bindumatoga.

FIGURE 11. Schematic drawing of left lateral view of three primary diagnostic lateral pigmentation characteristics of

members of the Hosii subgroup: (A) R. hosii; (B) R. spilotaenia; (C) R. aprotaenia.


Key to the species of Rasbora of the Sumatrana group in northern Sumatra

1a. Posterior-portion stripe and midhumeral diffuse patch of black midlateral stripe absent; supra-anal pigmentation absent (Figs.

2D, 9C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rasbora bindumatoga

1b. Posterior-portion stripe and midhumeral diffuse patch of black midlateral stripe present; supra-anal pigmentation present . . . 2

2a. Black midlateral stripe not reaching midhumeral region anteriorly; posterior-portion stripe not confluent posteriorly with ante-

rior tip of basicaudal blotch; supra-anal pigmentation elongated (Figs. 2B, 7B); relatively deep-bodied (approximately 30%

SL) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rasbora haru

2b. Black midlateral stripe reaching midhumeral region anteriorly; posterior-portion stripe posteriorly confluent with anterior tip

of basicaudal blotch; supra-anal pigmentation ovoid; relatively slender-bodied (approximately 24–27% SL) . . . . . . . . . . . . . 3

3a. Subdorsal blotch present; black midlateral stripe extending anteriorly and confluent with subdorsal blotch (Figs. 2A, 7A); 23–

25 + 3–4 scales of lateral-line series; 11–12 gill rakers on first gill arch . . . . . . . . . . . . . . . . . . . . . . . . . . . .Rasbora arundinata

3b. Subdorsal blotch absent; black midlateral stripe tapering anteriorly, reaching midhumeral region and without subdorsal blotch

(Figs. 2D, 10B); 26–27 + 3–4 scales of lateral-line series; 13 gill rakers on first gill arch . . . . . . . . . . . . . . .Rasbora maninjau

Material examined

Rasbora api: MZB 16457, holotype, 45.6 mm SL, Indonesia, Sumatra, Province of Sumatera Utara (North

Sumatra), Kabupaten Tapanuli Tengah, irrigation canal of Aek Pinangsori River (tributary of Batang Lumut

River) on road between Sibolga and Batangtoru, 01°33’59’’N, 098°54’62’’E; USNM 391737, paratypes, 20

(alc.), 6 (CS), 6 (AZ), 5 (HIS), 21.3–43.2 mm SL, same locality data as holotype.

Rasbora bankanensis: USNM 230222, 5, 34.5–43.7 mm, Indonesia, Borneo, West Kalimantan, Sungai Melai and

mouth of Sungai Pinoh at Nangapinoh, 00°19’30’’S, 111°44’E.

Rasbora bunguranensis: ZRC 49887, 25, 13.0–79.0 mm SL, Indonesia, Natuna Island, Air Tayan River,

surrounding areas at Ranai, stream next to Natuna Hotel, 3º56.70’N/108º22.04’E.

Rasbora einthovenii: USNM 328027, 110, 21.0–44.6 mm, Brunei, District of Belait, Sungai Bau, tributary stream

of Sungai Belait River, upstream from Kampung Melilas.

Rasbora elegans: RMNH 26350, 2, 50.3 and 62.3, Malaysia, Johore, Bukit Serampang; RMNH 26352, 3, 62.8–

73.8 mm, Malaysia, Johore, Sungei Semalok, Nawai; ZRC 14054–14065, 12, 15.2–56.0 mm SL, Indonesia,

Riau Archipelago, Batam Island, northwestern part; ZRC 7080–7084, 5, 47.8–70.5 mm SL, Malaysia,

Malacca, Bemban stream, next to hot spring; ZRC 14217–14246, 30, 19.5–62.7 mm SL, Malaysia, Johor,

KotaTinggi, foothill of Gunung Panti; ZRC 39950, 2, 59.6 and 73.5 mm SL, Malaysia, Johor, Kota Tinggi,

Tementang River; ZRC 40248, 3, 71.6–80.4 mm SL, Malaysia, Johor, tributary of Batang Pinang River, 272

km towards Kluang; ZRC 12885–12893, 9, 23.9–67.2 mm SL, Singapore, Seletar Reservoir Park, stream

outside Nee Soon Rifle Range; ZRC 19436–19444, 9, 22.6–67.2 mm SL, Malaysia, Johor, Semberong River, 6

km from Kluang; ZRC 38270, 2, 47.1–54.9 mm SL, Indonesia, Riau Archipelago, Batam Island, 1º3’N,

103º59’E; ZRC 34625–34628, 4, 39.0–63.3 mm SL, Singapore, Rifle Range Road; ZRC 37743, 4, 27.9–50.5

mm SL, Singapore, stream of Lagoon Banir; ZRC 45685, 5, 22.2–60.1 mm SL, Indonesia, Borneo, Kalimantan

Timur, Mahakam.

Rasbora ennealepis: USNM 230230, 4, 29.1–36.7 mm, Indonesia, Borneo, West Kalimantan, Sungai Tebelian,

small forest stream 3–4 m wide and 50 cm deep, flowing into Sungai Pinoh River, 19 km S of Nangapinoh,

0°30’S, 111°45’E.

Rasbora hubbsi: BMNH 1997.5.13.18-15, 17, 26–49.1 mm, Malaysia, Sabah, Sungai Lonpodas.

Rasbora kluetensis: MZB 16470 (holotype), 30.9 mm SL, Indonesia, Sumatra, Province of Nanggroe Aceh

Darussalam, Kabupaten Aceh Selatan, irrigation canal of Kluet River, 03º09’14’’N, 097º24’43’’E; USNM

391747 (paratypes), 50 (alc), 3(AZ), 4 (cs), 2 (HIS) 17.9–31.0 mm SL, same locality data as holotype.

Rasbora jacobsoni: ZRC 49140, 2, 46.5 and 47.4 mm, Sumatra, Padang Panjang, road from Padang to Bukittinggi;

USNM 390325, 13.3–25.8 mm, Kabupaten Aceh-Singkil, Kecamatan Sultan Daulat, Desa Namo Buaya, road

between Gelombang and Subulussalam.

Rasbora johannae: MZB 6094 (holotype), 38.1 mm, Indonesia, Kalimantan Tengah, Barito River drainage,

tributary of Sungai Busang River upstream from Project Barito Ulu basecamp on Sungai Busang.

Rasbora hosii: RMNH 7752, 3, 35.3–66.1, East Indian Archipelago, Borneo, Siniai River; ZRC 25933–25938, 6,

14.3–45.2 mm SL, Malaysia, Borneo, Sarawak, 99 km from Kuching, Jaguh River, after Balai Ringin; ZRC

39379, 2, 47.7 and 59.8 mm SL, Malaysia, Borneo, Sarawak,10 mile rock into Kuap turnoff, 1º25’8.9”N,


110º20’52.9”E; ZRC 39486, 1, 39.2 mm SL, Malaysia, Kuching, roadside ditch between Serian and

Tebenkang; ZRC 39469, 2, 24.4–45.4 mm SL, Malaysia, Sarawak,15 km to Serikin after turnoff from Bau

main road; ZRC 39502, 1, 34.8 mm SL, Malaysia, Sarawak, towards Bau, Stom Muda River; ZRC 667, 25,

12.1–70.7 mm SL, Malaysia, Sarawak, Kampong Pangkalan Kuap, Bukit Setigang, 7 miles south of Kuching;

ZRC 38826, 6, 35.1–58.1 mm SL, Indonesia, Kalimantan Barat, Kapuas Basin, Lanjak River, 1 km east of

Lanjak; ZRC 26022–26023, 2, 39.1 and 39.7 mm SL, Malaysia, Sarawak, drain-off on road to Simunjan; ZRC

27808–27810, 4, 14.2–20.7 mm SL, Malaysia, Sarawak, 200 m west of Balai Ringin, Simunjan Road.

Rasbora kottelati: USNM 328136, 12, 23.7–44.2 mm, Brunei, District of Belait, stream through Kerangas (heath)

forest, tributary of Ulu Ingei River, between markers 1028 and 1029.

Rasbora lateristriata: BMNH 1866.5.2.153 (paralectotype),1, 54.7 mm SL, Sumatra or Java, P. Bleeker; RMNH

4969 (lectotype), 40.9 mm SL, East Indies, Java; RMNH 9089 (paralectotypes), 3, 45.1–54.9 mm SL, Sumatra

or Java; RMNH 10316, 1, 53.9 mm SL, Java, Soekaboemi, Indonesia, West Java; MZB 10578, 4, 51.8–66.6

mm SL, Kabupaten Bogor, Kecamatan Cijeruk, Kampung Lengkong, upstream of Cisadane River; MZB

10582, 5, 11.5–52.9 mm SL, Sukabumi, Kecamatan Curug, Desa Nanggerang, Kampung Gintung, Cileuleuy

River; MZB 10584, 25, 11.3–51.8 mm SL, same location as MZB 10582; MZB 18012, 32, 34.3–74.7 mm SL,

Kabupaten Bogor, Kecamatan Nanggung, Desa Malasari, Kampung Central (Citalahab), upstream of Cikaniki

River.

Rasbora meinkeni: ZMA 100259, 2, 36.4 mm (male) and 45.6 mm (female), locality unknown; MZB 16689, 31

(alc.), 2 (cs), 17.3–30.2 mm SL, Indonesia, Sumatra, Province of Nanggroe Aceh Darussalam, Kabupaten

Aceh Tengah, Kecamatan Lut Tawar, Kampung One-one, Lake Laut Tawar, Hotel Ringgali, 04°36’72”N,

096°51’85”E.

Rasbora nodulosa: MZB 16465 (holotype), 34.6 mm SL, Indonesia, Sumatra, Province of Nanggroe Aceh

Darussalam, Kabupaten Aceh Barat Daya, Kecamatan Tangan-tangan, Tangan-tangan River on road between

Blang Pidie and Tapaktuan, 03º39’09’’N, 096º54’83’’E; USNM 391743 (paratypes), 29 (alc.), 1 (AZ), 2 (cs), 4

(HIS), 16.4–34.9 mm SL, same locality data as holotype.

Rasbora notura: UF 173501, 17, 16.7–44.1 mm SL, Malaysia, Peninsular Malaysia, Johor Drainage, Sungai Siam,

small stream south of Layang-layang on main road towards Keluang; RMNH 26349, 2, 36.2–37.5 mm SL,

Trengganu, Sungai Bukit Lima, Jerangau Road

Rasbora paviana: ZRC 25091–25102, 12, 26.4–59.6 mm SL, Malaysia, Trengganu. Ulu Besut, Kemia River, 6 km

from Kampung Keruak; ZRC 40845, 10, 37.9–59.6 mm SL, Thailand, Trat Province, Nam Tok Saphan Hin

(waterfall), near Ban Saphin Hin (village), 12º06’02”N, 102º42’50”E.

Rasbora reticulata. ZMA 109.585 (paralectotype), 1, Indonesia, Nias Island, 5 hours southwest of Gunungsitoli.

Rasbora rutteni. ZMA 102.360, 1 of 23 paratypes examined, 30 mm, Borneo, Sungai Wain, rivulet near Bontang.

Rasbora sarawakensis. USNM 230234, 7, 1 CS, 24.5–45.2 mm, Indonesia, Borneo, West Kalimantan, Sungai

Tamang, tributary of Sungai Pinoh River opposite mouth of Sungai Kelawai River, 0°35’S, 111°44’E.

Rasbora spilotaenia: MZB 14060, 5, 27.7–52.8 mm SL, Indonesia, Sumatra, Lampung, Way Pemerihan.

Rasbora sumatrana: RMNH 7038 (lectotype), 75.2 mm SL, East Indies, Sumatra, Solok, P. Bleeker, 1879, ca.

1851; RMNH 8909 (paralectotype), 63.7 mm SL, East Indies, Sumatra, Solok, P. Bleeker, 1879, ca. 1851;

CMK 4642, 10, Indonesia, Sumatra, Sumatera Barat, Solok.

Rasbora tobana: CAS 68363 (syntypes), 4, 18.4–26.1 mm SL, Indonesia, Sumatra, Toba-See (Lake Toba).

Rasbora trifasciata: RMNH 7623, 1 of 9 syntypes examined, 41.8 mm, Indonesia, Borneo, 4 August 1900; ZMA

109274, 1 syntype, 31.9 mm, Borneo, Bo River.

Rasbora truncata: MZB 16678 (holotype), 26.6 mm SL, Indonesia, Sumatra, Province of Nanggroe Aceh

Darussalam, Kabupaten Aceh Tenggara, Kampung Air Kelabu, Alas River near road between Kutacane and

Blangkejeren, 03º42’69’’N, 097º38’02’’E; USNM 391744 (paratypes), 29 (alc), 6 (cs), 2 (HIS), 19.7–35.3 mm

SL, same locality data as holotype.

Rasbora tuberculata: MZB 5902 (holotype), 26.8 mm, Indonesia, Borneo, Kalimantan Barat, Kapuas basin, Sungai

Pala at Pala Hulu, Kecamatan Siberuang, Kampung Renyai Hulu, km 99 on road from Sintang to Putussibau,

0°21’42’’N, 111°55’47’’E.

Rasbora volzi: ZRC 45656, 12, 26.2–88.4 mm SL, Indonesia, Borneo, Kalimantan Timur, Kayan basin, Bahau,

Lalut Birai next to Lalut Birai Field station, feeder stream to En’ggeng B’io, 2°52’8”N, 115°49’19”E.

Rasbora vulcanus: MZB 9317 (holotype), 40.1 mm, Indonesia, West Sumatra, Painan, Batang Si Joontour,


approximately 53 km to Painan on Padang-Painan road, 1°04’52.8”S, 100°27’26”E.

Rasbora vulgaris: BMNH 1905.5.6.4–5 (syntypes), 2, 37.7 and 34.8 mm SL, Malaysia, Selangor, Kuala Lumpur;

RMNH 24858, 20, 68.1–78.1 mm SL, Malaysia, Malacca, Penang, Baya Lepas; RMNH 26373, 5, 21.7–50.4,

Malaysia, Selangor, 2 km Batu Tiga to Subang Road.

Acknowledgment

I express my thanks to L. Parenti (USNM) for her support and guidance throughout the project. I gratefully

acknowledge J. Burns (GWU) for his supervision and critical comments on the text. I thank R. Vari (USNM) and L.

Page (UF) who critically reviewed the manuscript. I am indebted to R. Hadiaty (MZB) who guided me in the field

and facilitated the permits. I am grateful to M. Kottelat and H. H. Tan (ZRC) for the insightful discussion and

constructive criticisms on the taxonomy of Rasbora. I acknowledge the following at USNM for their invaluable

assistance: J. Clayton, J. Williams, J. Finan, S. Raredon, and D. Cole (base map). I thank the following for access to

material under their care: M. Sabaj Pérez (ANSP), R. Britz, O. Crimmen and P. Campbell (BMNH), M. Kottelat

(CMK), K. Grossenbacher and S. Hertwig (NMBE), R. de Ruiter and M. van Oijen (RMNH), R. Robins (UF), R.

Vonk and H. Praagman (ZMA), and K. Lim and H. H. Tan (ZRC). I am grateful to D. Rudaya, N. M. Ray, D.

Syahril, and V. Simanjuntak for their fieldwork assistance. The 2006 ichthyofaunal survey in northern Sumatra was

financially supported by the Leonard P. Schultz Fund, Division of Fishes, USNM. This article constitutes part of

my dissertation submitted in partial fulfillment of the degree of Doctor of Philosophy to the Department of

Biological Sciences, the George Washington University.

Literature cited

Brittan, M.R. (1954a) A revision of the Indo-Malayan fresh-water fish genus Rasbora. Monographs of the Institute of Science

and Technology, Manila, 224 pp + 3 maps.

Brittan, M.R. (1954b) The cyprinid fish genus Rasbora in Malaya. The Bulletin of the Raffles Museum, 25, 129–156.

Conway, K.W. (2005) Monophyly of the genus Boraras (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters, 16,

249–264.

Eschmeyer, W.N. (Ed.) (2013) Catalog of Fishes electronic version. Available from: http://research.calacademy.org/research/

ichthyology/catalog/fishcatmain.asp (accessed 11 November 2013)

Froese, R. & Pauly, D. (Eds.) (2013) FishBase, World Wide Web electronic publication. Available from: http://

www.fishbase.org (accessed 1 October 2013)

Hadiaty, R.K. (2005) Keanekaragaman jenis ikan di Suaq Balimbing dan Ketambe, Taman Nasional Gunung Leuser, Provinsi

Nanggroe Aceh Darussalam. Jurnal Biologi Indonesia, 3, 379–388.

Kottelat, M. (1986) A review of the nominal species of fishes described by G. Tirant. Nouvelles Archives du Muséum d'Histoire

Naturelle de Lyon, 24, 5–24.

Kottelat, M. (2001) Fishes of Laos. Wildlife Heritage Trust, Colombo, 198 pp.

Kottelat, M. (2005) Rasbora notura, a new species of cyprinid fish from the Malay Peninsula. Ichthyological Exploration of

Freshwaters, 16, 265–270.

Kottelat, M., Whitten, A.J., Kartikasari, S.N. & Wirjoatmodjo, S. (1993) Freshwater fishes of Western Indonesia and Sulawesi.

Periplus, Hong Kong, 259 pp + 84 plates.

Kottelat, M. & Vidthayanon, C. (1993) Boraras micros, a new genus and species of minute freshwater fish from Thailand

(Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters, 4, 161–176.

Kottelat, M. & Lim, K.K. (1995) Freshwater fishes of Sarawak and Brunei Darussalam: a preliminary annotated check-list. The

Sarawak Museum Journal (New Series), 48, 227–256.

Kottelat, M. & Tan, H.H. (2011) Rasbora atranus, a new species of fish from central Borneo (Teleostei: Cyprinidae).

Ichthyological Exploration of Freshwaters, 22, 215–220.

Kottelat, M. & Tan, H.H. (2012) Rasbora cryptica, a new species of fish from Sarawak, Borneo (Teleostei: Cyprinidae).


Liao, T.Y., Kullander, S.O. & Fang, F. (2010) Phylogenetic analysis of the genus Rasbora (Teleostei: Cyprinidae). Zoologica

Scripta, 39, 155–176.

http://dx.doi.org/10.1111/j.1463-6409.2009.00409.x

Liao, T.Y., Kullander, S.O. & Fang, F. (2011) Phylogenetic position of rasborin cyprinids and monophyly of major lineages

among the Danioninae, based on morphological characters (Cypriniformes: Cyprinidae). Journal of Zoological

Systematics and Evolutionary Research, 49, 224–232.


http://dx.doi.org/10.1111/j.1463-6409.2009.00409.x

http://dx.doi.org/10.1643/ci-09-155


http://dx.doi.org/10.1111/j.1439-0469.2011.00621.x

Lumbantobing, D.N. (2010) Four new species of the Rasbora trifasciata-group (Teleostei: Cyprinidae) from Northwestern

Sumatra, Indonesia. Copeia, 2010, 644–670.


Ott, G. (2009) Redescription of Homaloptera ripleyi (Fowler, 1940) from Sumatra, Indonesia (Teleostei: Balitoridae). Bulletin

of Fish Biology, 11, 73–86.

Sabaj Pérez, M.H. (Ed.) (2012) Standard symbolic codes for institutional resource collections in herpetology and ichthyology:

an Online Reference. Version 3.0. American Society of Ichthyologists and Herpetologists, Washington, DC. Available

from: http://www.asih.org/ (23 February 2012)

Siebert, D.J. & Guiry, S. (1996) Rasbora johannae (Teleostei: Cyprinidae), a new species of the R. trifasciata-complex from

Kalimantan, Indonesia. Cybium, 20, 395–404.

Siebert, D.J. & Richardson, P.J. (1997) Rasbora laticlavia, a new cyprinid from Kalimantan, Indonesia, and lectotype

designation for R. vaillantii. Ichthyological Exploration of Freshwaters, 8, 89–95.

Springer, V.G. & Allen, G.R. (2004) Escenius caeruliventris and E. shirleyae, two new species of bleniid fishes from Indonesia,

and new distribution records for other species of Ecsenius. Zootaxa, 791, 1–12.

Tan, H.H. & Kottelat, M. (2009) The fishes of the Batang Hari drainage, Sumatra, with description of six new species.


Tang, K.L., Agnew, M.K., Hirt, M.V., Sado, T., Schneider, L.M., Freyhof, J., Sulaiman, Z., Swartz, E., Vidthayanon, C., Miya,

M., Saitoh, K., Simons, A.M., Wood, R.M. & Mayden, R.L. (2010) Systematics of the subfamily Danioninae (Teleostei:

Cypriniformes: Cyprinidae). Molecular Phylogenetics and Evolution, 57, 189–214.

http://dx.doi.org/10.1016/j.ympev.2010.05.021

Weber, M. & de Beaufort, L.F. (1916) The fishes of the Indo-Australian Archipelago. III. Ostariophysi: II Cyprinoidea, Apodes,

Synbranchi. E. J. Brill, Leiden, 404 pp.

Wirjoatmodjo, S. (1987) The river ecosystem in the forest area at Ketambe, Gunung Leuser National Park, Aceh, Indonesia.

Archiv für Hydrobiologie-ergebnisse der Limnologie, 28, 239–246.


http://dx.doi.org/10.1016/j.ympev.2010.0

http://dx.doi.org/10.1016/j.ympev.2010.0

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