Re-Description of Betta Rubra Perugia, 1893 (Teleostei: Osphronemidae), an Enigmatic Fighting Fish from Sumatra

Bulletin of Fish Biology Volume 13 Nos. 1/2 30.11.2011 21-32

Re-description of Betta rubra Perugia, 1893 (Teleostei: Osphronemidae), an enigmatic ﬁ ghting ﬁ sh from Sumatra

Wiederbeschreibung von Betta rubra Perugia, 1893 (Teleostei: Osphronemidae), einem rätselhaften Kampfﬁ sch von Sumatra

Ingo Schindler1 & Stefan van der Voort2

1Warthestr. 53a, D-12051 Berlin, Germany, [email protected] 2Beetzlaan 72K, NL-3762 CG Soest, the Netherlands, [email protected]

Summary: Betta rubra, occurring from Meulaboh to Sibolga in north-western Sumatra, is formally re-described based on freshly collected preserved and live specimens. The species can be differentiated from other members of the genus by the combination of the following characters and character states: an iridescent light blue patch at pectoral fi n origin, alternating black bars and red blocks ventrolaterally on the fl ank, presence of a dark triangular patch below the eye, absence of red twin bars on the opercle when preserved. Based on the differential diagnosis B. rubra is herein removed from the Betta foerschi species group, in which it was tentatively placed, and transferred into its own group. Uncertainties regarding its distribution are resolved. Biogeography and possible speciation of B. rubra are briefl y discussed.

Key words: Osphronemidae, Bettini, Betta rubra, Betta foerschi, species group, biogeography.

Zusammenfassung: Betta rubra wird anhand von lebenden und konservierten Exemplaren wiederbeschrie- ben. Die Art ist im nordwestlichen Sumatra von Meulaboh bis Sibolga verbreitet. Sie unterscheidet sich von den übrigen Arten der Gattung durch die Kombination folgender Merkmale und Merkmalsausprägungen: ein irisierender hellblauer Fleck am Ansatz der Brustfl osse, alternierend schwarze Balken und rote Zonen auf den ventralen Körperseiten, ein dunkler, dreieckiger Fleck unter dem Auge, Fehlen von roten, parallelen Balken auf dem Kiemendeckeln bei konservierten Exemplaren. Basierend auf den Ergebnissen (vergleichende Diagnose) wird die Art von der Betta foerschi-Artengruppe (in der sie vorläufi g eingeordnet wurde) in eine eigenständige Gruppe überführt. Unsicherheiten in Bezug auf das Verbreitungsgebiet werden geklärt. Die Biogeografi e und mögliche Artentstehung von B. rubra werden kurz diskutiert.

Schlüsselwörter: Osphronemidae, Bettini, Betta rubra, Betta foerschi, Artengruppe, Biogeographie.

1. Introduction was already published by REGAN (1910), B. rubra remained an elusive species due to the The Southeast Asian osphronemid genus lack of fresh material. As colouration of live Betta (generally called ‘fi ghting fi sh’) includes specimens (in particular in males) is an impor- nearly 70 species (TAN & NG 2005 a, b, 2006, tant diagnostic character and often a major SCHINDLER & SCHMIDT 2006, FROESE & PAULY characteristic feature to distinguish species 2012). Betta species are small fi shes (up to (TAN & TAN 1994, 1996, TAN & NG 2005 a, about 100 mm SL). The majority has been b), it was not possible to classify B. rubra with described in the last two decades. One of adequate certainty. The species name rubra the oldest species names in the genus is Betta was erroneously used for a specimen from rubra Perugia, 1893. Originally described from Singapore (HERRE 1940) and different forms the west coast of Sumatra, for a long time it of the Betta splendens group (DONOSO-BÜCHNER was known only from the type specimens. & SCHMIDT 1999) and has even been regard- Although a fi rst drawing of one of the types ed as a synonym of B. imbellis (see SCHMIDT

Bull. Fish Biol. 13 (1/2) 21 1996). TWEEDIE (1952) noted that the speci- between two critical points (as described in men identifi ed as B. rubra by HERRE (1940) SCHINDLER & SCHMIDT 2006). The distances are is probably Betta pugnax. The taxon B. rubra measured with a dial calliper reading to the near- was also confused or even synonymized with est 0.1 mm. Ratios are expressed as percentages Betta picta in the popular literature (VIERKE of standard length (SL). Counts were made as 1986). WITTE & SCHMIDT (1992) confi rmed described by WITTE & SCHMIDT (1992), except the taxonomic validity of B. rubra and pointed for the number of predorsal scales which are out that the characteristic triangular mark counted continuously (NG & KOTTELAT 1994). below the eye (WITTE & SCHMIDT 1992: fi g. Numbers in parentheses indicate the number of 8h) separates it from the members of the specimens examined with that particular count. Betta splendens- group. They placed B. rubra in Terminology of colour patterns follows TAN & its own species group. The photographs of NG (2005a) with modifi cations after SCHINDLER two type specimens by KOTTELAT (1993, KOT- & SCHMIDT (2006). The description follows the TELAT in SCHÄFER 1997) indicated that B. rubra general format used by TAN & NG (2005a) and differs in appearance from all other nominal SCHINDLER & SCHMIDT (2006). Besides the data Betta species. TAN & NG (2005b) provided a recorded herein data from previous descriptions fi rst short re-description of B. rubra based on were included (VIERKE 1979, SCHALLER & KOT- some of the types and a few new, but impro- TELAT 1989, TAN & NG 2005a, b, 2006). perly preserved specimens from Aceh Barat, The terminology and application of phy- collected in 1982; however, live colours were logenetics follows HENNIG (1966, see WÄGELE not recorded. TAN & NG (2005a, b) classifi ed 2004), i.e. that character polarity was estimated this species as a member of the Betta foerschi independently and prior to a tree construction. It group. The lack of freshly collected material is most parsimonious to treat characters, which and colour photographs of live specimens appear uniquely in the in group and closely re- made it impossible to confi rm their classifi - lated out groups, a priori as apomorphic unless cation and complete the description of the proven otherwise. taxonomically important colour pattern. The signifi cance of pairwise differences in Recently, a local fi sherman collected new morphometric data between examined speci- material of B. rubra at the west coast of mens of Betta rubra and Betta foerschi are tested Sumatra. Besides preserved specimens also with the Student’s t-test. To control the increased live fi sh were imported. Therefore it is now type I error in multiple comparisons the step- possible for the fi rst time to describe the live up false discovery rate procedure (BENJAMINI & colour pattern and to clarify the taxonomical HOCHBERG 1995) is used to indicate substantial position of B. rubra. dissimilarities. The purpose of this paper is to present a Museum acronyms: BMNH = Natural His- re-description of Betta rubra from preserved tory Museum, London; MSNG = Museo Civico material and photographs of live specimens and di Storia Naturale Giacomo Doria, Genova; consequently to place it in its own species group ZMB = Museum für Naturkunde, Berlin. based on apomorphic characteristics. We also Material examined: Betta rubra: ZMB 34533, 5, resolve the confusion regarding its distribution 30.5-41.1 mm SL, Indonesia, Aceh, west coast and provide an English translation of Perugia’s of northern Sumatra around Meulaboh, licensed original Italian description as an appendix. import and preserved 2010, don. Colin Dunlop. Collection of the authors, 1 male, 52.7 mm, SL, 2. Material & methods Meulaboh, Sumatra, Indonesia, preserved 2010. Observations of reproductive behaviour and The measurements taken follow WITTE & live colouration are based on fi ve specimens SCHMIDT (1992) with the modification that collected from the same locality which have been all measurements are taken as a straight-line kept in aquaria; not preserved. Betta foerschi: Col-

22 lection of the authors, 4 ex, 32.7-44.6 mm SL, position; Betta foerschi species group); TAN & licenced import from Borneo (no exact locality NG (2005b): 119, fi g. 1 (photograph of type available), preserved 2010, don. Colin Dunlop. specimens, taxonomy, B. foerschi species group). Type specimens: MSNG 13019 (10 specimens; 3. Results Sibolga); BMNH 1893.5.29.1 (1 specimen; Lake Toba). Betta rubra Perugia Diagnosis: Betta rubra differs from all other members of the genus by the presence of an References: Betta rubra Perugia, 1893: 242 iridescent light blue patch at pectoral fi n origin (original description; Lago Toba, Siboga); VOLZ in both sexes, an alternating pattern of black 1904: 493 (referred to PERUGIA (1893), no newly bars (with iridescent blue scales in live) and red collected specimens from Lake Toba); VOLZ blocks on fl ank ventrolaterally, and the presence 1907:127 (listed); REGAN 1910: 781, pl. 77, fi g. of a broad chin bar, which appears as a dark 1 (taxonomy; description of a syntype); WEBER triangular mark below eye. & DE BEAUFORT 1922: 361 (description, re-exa- Description: For general appearance see fi g- mination of the BMNH syntype); TORTONESE ures 1 and 2. Meristic and morphometric data 1963: 340 (MSNG syntypes listed); WITTE & are summarised in table 1. Maximum standard SCHMIDT 1992: 327, fi g. 8h (taxonomic status; B. length usually <50 mm. Body slender (body rubra species group); KOTTELAT et al. 1993: 163, depth at dorsal fi n origin <25.5 % of SL); pl. 76 (colour photograph of two representative head length comparatively short for a mouth syntypes); TAN & NG 2005a: 58 (systematic brooding species (head length usually <31 %

Fig. 1: Betta rubra, male (39 mm SL), preserved (ZMB 34533). Abb. 1: Betta rubra, Männchen (39 mm SL), konserviert (ZMB 34533).

Fig. 2: Betta rubra, female (38 mm SL), preserved (ZMB 34533). Abb. 2: Betta rubra, Weibchen (38 mm SL), konserviert (ZMB 34533).

Bull. Fish Biol. 13 (1/2) 23 of SL); snout slightly pointed. Dorsal and anal Suborbital stripe dark, merging with dark chin fi ns pointed (more in males); dorsal fi n situated bar. Dark triangular patch below eye, merging comparatively far posteriorly (predorsal length with suborbital stripe and chin bar. Lower lip 64-66 % of SL); caudal fi n rounded in females; mostly black. Black spots on dorsum of head. rounded, pointed or lanceolate in males; anal An alternating pattern of 5-7 black bars (with fi n base length about half of SL or more (anal iridescent blue scales) and red blocks on the fi n base 49-55 % of SL); dorsal fi n base length fl anks ventrolaterally. Faint caudal peduncle 10-13 % of SL; pelvic fi ns with long fi lamen- spot present. Caudal, dorsal and anal fi ns red, tous ray (length up to 34 % of SL); pelvic fi n outer margins light-blue or white. Anal fi n and spine comparatively short (usually <5 % of SL); central part of caudal fi n with whitish-green pectoral fi ns rounded (length 18-20 % of SL). streaks. Pelvic fi ns red with faint to dominating Dorsal fi n rays I, 7 (3), I, 8 (2) or II, 8 (1). Anal whitish-green streaks near origin, fi lament white. fi n rays III, 22 (1), III, 23 (2), III, 24 (2) or III, Pectoral fi ns hyaline. Presence of iridescent light 25 (1). Pectoral fi n rays 12 or 13. Pelvic fi n rays blue patch at pectoral fi n origin. I, 1,4. Lateral scales 29 (3) or 30 (3). Transverse Description of female: Body background scales at dorsal fi n origin 9 (1), 9.5 (3) or 10 (2). dorsally dark brown, ventrally lighter. Some Predorsal scales 19-22; postdorsal scales 9-10. fl ank scales highlighted with iridescent blue. Preserved colouration: See fi gures 1 and 2. Opercle with faint golden shine, red twin bars Body brown or brownish-olive, dorsum dark virtually absent or small remnant of posterior brown, ventral parts light brown or whitish. On bar present. Orbital stripes, chin bar and lower ventral part of body 5-7 irregular, spaced dark lip colour as described above. Dark triangular bars (more distinct in males than in females) patch below eye present. Dorsum of head with with reddish blocks in between. Ventral part black spots. Single horizontal fl ank stripe on of head light brown or greyish; lower lip dark. lower half of body just below centre, merging Chin bar broad, reduced to a conspicuous dark with caudal peduncle spot. Blue iridescence on triangular mark below eye; pre- and postorbital dorsolateral scales faint or absent. Dark fl ank stripes present, posterior part of the latter bars and red blocks absent. Unpaired fi ns hyaline on opercle appearing as broad patch (more to brownish. Pelvic fi ns faint red, fi lament white. conspicuous in males than in females); opercle Presence of whitish-green streaks same as for with reddish margin. Dorsal and caudal fi ns males, albeit fainter. Pectoral fi ns hyaline. Pec- reddish-brown without transverse bars, anal fi n toral fi n origin with iridescent light blue patch, grey-brown, distally reddish-brown, pelvic fi ns slightly weaker than in males. reddish-brown to reddish. Pectoral fi n hyaline, Colour intensity mood-dependant. No particu- with a dark patch at origin. Upper stripe on lar male brood colouration was observed. Female fl ank absent; central stripe faint in males, more individuals before and during spawning with lighter conspicuous in females, running from pectoral brown stripe running along dorsum of head to fi n base to lower caudal fi n base, not connected dorsal fi n and from dorsal fi n to caudal fi n. with caudal fi n base spot. Caudal fi n base spot Reproductive behaviour: Observations in faint or absent in males, more conspicuous in captivity showed that Betta rubra is a paternal females. Second central stripe absent or strongly mouth brooder. Spawning always commences reduced. on a substrate ranging from fl oating leaf to the Live colouration: See fi gures 3 and 4. Descrip- bottom. After spawning the male goes essen- tion of male: Body background dark brown tially into hiding until releasing the fry. dorsally, light brown ventrally. Dorsolaterally Distribution: Lowlands of north western Su- some scales highlighted with iridescent blue. matra; from Meulaboh in the north to Sibolga Red twin bars on opercle, posterior bar angling in the south (fi g. 5). down along gill plate. Opercle with faint golden Etymology: PERUGIA (1893) did not explain shine. Dark pre- and postorbital stripes distinct. the species name. It is doubtlessly derived from

24 Fig. 3: Live colouration of Betta rubra, adult male (not preserved). Abb. 3: Lebendfärbung von Betta rubra, adultes Männchen (nicht konserviert).

Fig. 4: Live colouration of Betta rubra, adult female (not preserved). Abb. 4: Lebendfärbung von Betta rubra, adultes Weibchen (nicht konserviert).

Bull. Fish Biol. 13 (1/2) 25 Fig. 5: Distribution of Betta rubra; dots: (1) Meulaboh; (2) Sibolga (type locality). Abb. 5: Verbreitung von Betta rubra. Punkte: (1) Meulaboh, (2) Sibolga (Typuslokalität). the Latin adjective ‘ruber’, meaning red, which blue iridescent opercle in life (versus present), was most likely assigned to the species for its lack of upper and second central stripes (versus colouration. present); from the Betta akarensis group by lack Comparative notes: TAN & NG (2005a, b) of transverse bars in caudal fi n (versus usually tentatively placed Betta rubra in the Betta foerschi present and always well-defined in mature group. It differs from the species of this group males), absence of faintly coloured greenish- (Betta foerschi, Betta strohi and Betta mandor) by the gold scales on opercle (versus present); from possession of an iridescent light blue patch at the Betta unimaculata group by having a re- pectoral fi n origin (versus absent), presence of latively small head and mouth (versus large a dark triangular patch beneath the eye (versus blunt head and large mouth), pelvic fi ns with absent), a prominent central stripe on lower half long fi lament ous ray up to 34 % of SL (versus of body just below midline (no upper or second short pelvic fi n fi lament); from the Betta picta central stripe present), bars on fl ank compara- group by absence of upper and second central tively broad (versus usually narrower), presence stripes (versus present), pointed fi ns (versus of red blocks between bars ventrolaterally (ver- rounded), anal fi n and ventral part of caudal sus such pattern absent) and absence of red twin fi n lacking dark margin (versus broad dark bars on opercle when preserved (versus present). margins present); from the Betta splendens group See table 1 for differences in morphometric data by oral brooding (versus bubblenest brooding), between examined specimens. lack of upper central stripe (versus present); Besides the diagnostic characters provided from the Betta coccina group by having a larger above, B. rubra can be distinguished from all adult size (usually >38 versus <38 mm SL), other species groups as recognized in TAN & multi-coloured body (versus a uniform red or NG (2005a, b) as follows: from the Betta bellica black), oral brooding (versus bubblenest broo- group by lateral scales 29 or 30 (versus 33), oral ding); from the Betta waseri group by absence brooding (versus bubblenest brooding); from of dark throat markings (versus present), lack the Betta pugnax group by absence of a green or of distinct dark markings on opercle (versus

26 Tab. 1: Morphometric data (in percent of standard length; SL in mm) of Betta rubra and B. foerschi. Mean = arithmetic mean; min = lowest value; max = highest value; sd = standard deviation; t-test = results (uncor- rected p-values) of Student’s test; * = p < 0.05; ** = p < 0.01. Tab. 1: Morphometrische Daten (in Prozent der Standardlänge; SL in mm) von Betta rubra und B. foerschi. Mean = arithmetischer Mittelwert, min = kleinster Wert, max = höchster Wert, sd = Standardabweichung, t-test = Ergebnisse (unkorrigierte p-Werte) des Student-Test, * = p < 0,05, ** = p < 0,01.

present); from the Betta edithae group by fewer we here follow the historic and current concept anal fi n rays (22-25 versus 28-30); from the (e.g. REGAN 1910, BRITZ 2001, TAN & NG 2005a, Betta anabatoides group by pelvic fi ns with long b, RÜBER et al. 2006) of the genus. fi lamentous ray (versus pelvic fi n fi lamentous The name of the subfamily that includes the ray usually not extended), head without two genus Betta is listed as Macropodusinae (e.g. BRITZ light spots behind eye (versus present), smaller 2001, RÜBER et al. 2006). However, the family body depth (<25.5 versus 30-33 % of SL); group names Trichogastrini and Bettini pro posed from the Betta albimarginata group by having by BLEEKER (1879) and Parophiocephalidae fewer anal fi n spines (

Bull. Fish Biol. 13 (1/2) 27 rists and importers alike, and appeared in the from the species of the Betta foerschi group by ornamental fi sh trade for the fi rst time in 2007 the apomorphic characters or character states (LINKE 2007). The reddish colour elements of given in the diagnosis and comparative notes the type specimens lead to the misconception (see above). WITTE & SCHMIDT (1992) placed B. that B. rubra is part of the Betta splendens group rubra in a group of its own. They based their (WITTE 1997: 12) or even regarded as conspe- conclusion mainly on the unusual broad chin cifi c with Betta imbellis Ladiges, 1972 (SCHMIDT bar, which appears as a triangular mark below 1996, DONOSO-BÜCHNER & SCHMIDT 1999). The the eye in preserved specimens. Beside this cha- latter species is well known by its bright reddish racter (considered here as apomorphic) we fi nd colour in life (LINKE 2007, DONOSO-BÜCHNER the bright iridescent light blue patch at pectoral & SCHMIDT 1999). Specimens labelled ‘B. rubra’ fi n origin in both sexes, the alternating black bars in the popular literature of the early twentieth (with iridescent blue scales in life) and red blocks century were actually a reddish form of Betta on fl ank ventrolaterally and (as compared with splendens (see RACHOW 1937). Betta rubra shares other species) the more ventrally situated central with the species of the Betta splendens group the stripe (in correlation with the lack of the upper lack, or at least strong reduction, of a second central and second central stripe) to be unique in central stripe and a comparatively small head the genus. These characters (or character states) length. However, the additional description of are treated as apomorphic. The rising number B. rubra presented herein and discussion below of autapomorphies increases the necessity to make it apparent that it is not a member of recognize a taxonomically separate unit (STUESSY the Betta splendens group. Besides the characters 1997). Since B. rubra is excluded from all the given in the diagnosis, B. rubra differs in being other proposed species groups as distinguished a mouthbrooder (see reproductive behaviour by TAN & NG (2005a) by key characters (see above), while the species of the B. splendens comparative notes), we argue that the patristic group are bubblenest builders (SCHMIDT 1996). distance (counting apomorphies separating The genus Betta is divided into several species taxa), and thus the pheno-cladistic distinctness groups (TAN & NG 2005a, b). The groups are of B. rubra to the other species groups is suf- not necessarily based on synapomorphies but fi cient to place it (in accordance with WITTE & rather on superfi cial similarity of their members SCHMIDT 1992) in a separate group. (TAN & KOTTELAT 1998). TAN & NG (2005a, The only other known species in the genus b) placed B. rubra in their Betta foerschi species possessing a comparable patch at the pectoral group. Besides B. rubra, the Betta foerschi group fin origin is Betta compuncta from the upper currently consists of the following species (TAN Mahakam basin in Kalimantan Timur, Bor- & NG 2005a, 2006): B. foerschi Vierke 1978, B. neo; a member of the Betta unimaculata species strohi Schaller & Kottelat 1989 and B. mandor group (TAN & NG 2006). In B. rubra, the patch Tan & Ng 2006. Eidonomic features used in is iridescent light blue in life and black when TAN & NG (2005a, b) to include B. rubra in the preserved; in B. compuncta, it is black in both foerschi group are: presence of two vertical lighter live and preserved colouration. Hence, it seems coloured bars on opercle, vertical bars on body more likely that both species have evolved this when preserved, general body shape. These, patch independently. however, are trivial (vertical bars on opercle Caudal fi n shape in B. rubra appears to be and body are present in many other species and variable (see fi gs 1 and 3 and description above). likely plesiomorphic) or imprecise (no propor- Male specimens with rounded and lanceolate tions of body shape are provided that separate caudal fi ns have been collected from the same these species from other species groups). Ta- localities. Thus, the caudal fi n shape in B. rubra xonomically these features are impractical for is consequently not diagnostic, only informative. diagnosing a Betta foerschi group that includes B. Male caudal fi n shape has been repeatedly used rubra. Instead, the latter can be differentiated as a diagnostic character to distinguish species

28 (e.g. B. pallida versus B. prima and B. pulchra ver- coast line from the north to the south. North- sus B. pugnax) or species groups (TAN & TAN western Sumatra harbours a rich endemic fi sh 1996, TAN & NG 2005a, KOTTELAT & NG 1994, fauna and is treated as a separate ichthyofaunal SCHINDLER & SCHMIDT 2006). As variation in province (LUMBANTOBING 2010). The examined caudal fi n shape is described also for B. schalleri specimens of B. rubra were collected in the (see KOTTELAT & NG 1994), the taxonomical vicinity of Meulaboh (north-western Sumatra, value of the feature deserves to be critically Aceh). In consideration of the information gi- reviewed per case. ven by TAN & NG (2005a, b) and in accordance A lectotype for the taxon B. rubra is listed with WITTE & SCHMIDT (1992) and WITTE in TAN & NG (2005a, b). However, none of (1997), we conclude that B. rubra is restricted these works contain a valid designation of a to the lowlands of north-western Sumatra (fi g. lectotype. Likewise, neither TAN & NG (2005a, 5). Hence, B. rubra is another example of an b) nor ESCHMEYER & FRICKE (2012) provide endemic species for this ichthyofaunal province. any reference to a work, in which a lectotype is The ichthyogeographic isolation from the validly designated, and we were unable to locate species of the Betta foerschi group supports one as well. From a taxonomical point of view the view of the evolutionary uniqueness of it is not necessary to select a lectotype because B. rubra. The ichthyofaunal province (north- there is no indication that the type series of B. western Sumatra; LUMBANTOBING 2010) corre- rubra includes more than one species. sponds to a presumed tropical forest refugium The type locality given by PERUGIA (1893) during the Last Glacial Maximum (MEIJAARD is Lago Toba, Siboga. ‘Siboga’ is obviously 2003). Forest refugia are believed to have a alternative spelling for Sibolga (cf. KOTTELAT major impact on the allopatric speciation in et al. 1993), a seaport at the west coast of the Amazonia (HAFFER 2008). It is suspected that province Sumatera Utera. REGAN (1910) and such refugia have played a major role in the WEBER & DE BEAUFORT (1922) listed ‘Lago evolution of species groups and subgenera Toba, Sumatra’ as type locality. This enforced within the tropical aplocheiloid killifi sh fa- the incorrect impression that B. rubra inhabits milies (HUBER 1998). Since killifi sh occupy the volcanic lake Toba at an altitude of over 900 similar ecological niches in tropical forest m a.s.l. (e.g. VIERKE 1978). TORTONESE (1962) areas of South America and West Africa as restricted the locality to ‘Siboga [= Sibolga], do members of the genus Betta in Southeast Sumatra occidentale [= western Sumatra]’. As Asia (HUBER 1998, pers. obs.), it is not unlikely pointed out by TAN (2008), Lake Toba is not that such macro-ecological processes force the type of habitat where fi ghting fi sh species allopatric speciation and the development of are expected to occur. In fact, Betta species can independent evolutionary assemblages within usually be found in the lowlands or at the foot- the genus Betta as well. The biogeography, and hills of low mountain ranges. Betta rubra has not in particular the fl uctuating sea levels over been identifi ed in subsequent collections from geological time, palaeo-climates, landform the Lake (VOLZ 1904), and WITTE (1997) argued developments and the distribution of rain- that B. rubra does not occur in Lake Toba. forest refugia and swamp areas (see VORIS Besides its taxonomical distinctness, B. rubra 2000, MEIJAARD 2003) are not yet included in is also remarkable from a zoographical point of taxonomic studies on Southeast Asian fi ghting view. It is distributed in the lowlands of north- fi shes. It is well known that such processes western Sumatra where all major drainages infl uenced the evolution of species and spe- empty into the Indian Ocean (see distribution cies groups in Southeast Asia (e.g. MEIJAARD provided above; WITTE & SCHMIDT 1992, TAN & GROVES 2004, ALFARO et al. 2004). Hence, & NG 2005b). This region is hydrographically the incorporation of such data may support isolated from the eastern parts of the island by the hypotheses about the taxonomy and spe- the Barisan Mountains which run parallel to the ciation within Betta.

Bull. Fish Biol. 13 (1/2) 29 Acknowledgements and Bornean fi shes. The Bulletin of the Raffl es Museum 16, 27-61. We thank Colin DUNLOP (Carluke, Scotland) for HUBER, J.H. 1998. Comparison of the old world and donation of specimens and providing a photo- the new world tropical Cyprinodonts. Société Française d’Ichthyologie, Paris. graph, Stefano VALDESALICI (Viano, Italy) for the KOTTELAT, M. 1997. Betta rubra. In: Aqualog, all translation of the original description, Gerhard Labyrinths – Bettas, Gouramis, Snakeheads and OTT (Flensburg, Germany) and Hans-Jürgen Nandids (SCHÄFER, F., ed.). Verlag ACS, Mörfel- ENDE (Halle/Saale, Germany) for information den-Walldorf. about hobbyist literature. We are grateful to KOTTELAT, M., A.J. WHITTEN, S. N. KARTIKASARI & S. Richard VAN DER LAAN (Almere, the Nether- WIRJOATMODJO. 1993. Freshwater fi shes of West- lands) and Rico MORGENSTERN (Zethau, Ger- ern Indonesia and Sulawesi. Periplus Editions, many) for proofreading the manuscript and Hongkong. providing valuable corrections and suggestions. KOTTELAT, M., & P. K.L. NG. 1994. Diagnoses of fi ve new species of fi ghting fi shes from Bangka and Borneo (Teleostei: Belontiidae). Ichthyological Literature Exploration of Freshwaters 5, 65-78. LINKE, H. 2007. The genus Betta. Betta News Journal ALFARO, M.E., D.R. KARNS, H.K. VORIS, E. (special edition), 5-30. ABERNATHY, & S.L. SELLINS. 2004. Phylogeny of LUMBANTOBING, D.N. 2010. Four new species of the Cerberus (Serpentes: Homalopsinae) and phylo- Rasbora trifasciata group (Teleostei: Cyprinidae) geography of Cerberus rynchops: diversifi cation of from northwestern Sumatra, Indonesia. Copeia a coastal marine snake in Southeast Asia. Journal 2010, 644-670. of Biogeography 31, 1277-1292. MEIJAARD, E. 2003. Mammals of southeast Asian BENJAMINI, Y., & Y. HOCHBERG. 1995. Controlling islands and their late Pleistocene environments. the false discovery rate: a practical and powerful Journal of Biogeography 30, 1245-1257. approach to multiple testing. Journal of the Royal MEIJAARD, E., & C.P. GROVES. 2004. The biogeo- Statistical Society. Series B 57, 289-300. graphical evolution and phylogeny of the genus BLEEKER, P. 1879. Mémoire sur les poissons à pharyn- Presbytis. Primate Report 68, 71-90. giens labyrinthiformes de l’Inde Archipélagique. NG, P.K.L., & M. KOTTELAT. 1994. Revision of the Verhandelingen der Koninklijke Akademie van Betta waseri species group (Teleostei: Belontiidae). Wetenschappen 19, 1-56. Raffl es Bulletin of Zoology 42, 593-611. BRITZ, R. 2001. The genus Betta, monophyly and in- PERUGIA, A. 1893. Di alcuni pesci raccolti in Suma- trarelationships, with remarks on the subfamilies tra dal Dott. Elio Modigliani. Annali del Museo Macropodinae and Luciocephalinae (Teleostei: Civico di Storia Naturale di Genova (Serie 2a) Osphronemidae). Ichthyological Exploration of 13, 241-247. Freshwaters 12, 305-318. POPTA, C.M.L. 1905. Suite des descriptions prélimi- DONOSO-BÜCHNER, R., & J. SCHMIDT. 1999. Kampffi - naires des nouvelles espèces de poissons recueillies sche – Wildformen. Bede Verlag, Ruhmannsfelden. au Bornéo central par M. le Dr. A. W. Nieuwen- ESCHMEYER, W.N., & R. FRICKE. 2012. Catalog of huis en 1898 et en 1900. Notes from the Leyden Fishes (Online Version, 15 March 2012). Avail- Museum 25, 171-186. able from http://researcharchive.calacademy.org/ RACHOW, A. 1937. Betta rubra. In : Aquarienfi sche research/Ichthyology/catalog/fi shcatmain.asp in Wort und Bild (HOLLY, M., H. MEINKEN, & FROESE, R., & D. PAULY (eds). 2012. FishBase. World A. RACHOW, eds). Loseblattsammlung, Liefe- Wide Web electronic publication. www.fi shbase. rung 24,10; 163/165. Verlag Gustav Wenzel ; org, version (11/2011). Braunschweig. HAFFER, J. 2008. Hypotheses to explain the origin of REGAN, C.T. 1910. The Asiatic fi shes of the fam- species in Amazonia. Brazilian Journal of Biology ily Anabantidae. Proceedings of the Zoological 68, 917-947. Society of London 1909, 767-787. HENNIG, W. 1966. Phylogenetic Systematics. Univer- RÜBER, L., R. BRITZ, & R. ZARDOYA. 2006. Molecular sity of Illinois Press, Illinois. phylogenetics and evolutionary diversifi cation HERRE, A.W.C.T. 1940. Additions to the fi sh fauna of of labyrinth fi shes (Perciformes: Anabantoidei). Malaysia and notes on rare or little known Malayan Systematic Biology 55, 374-397.

30 SCHALLER, D., & M. KOTTELAT. 1989. Betta strohi sp. TWEEDIE, M.W.F. 1952. Notes on Malayan fresh-water n., ein neuer Kampffi sch aus Südborneo (Osteich- fi shes. 3, The Anabantoid fi shes; 4, New and thyes: Belontiidae). Die Aquarien und Terrarien- interesting records; 5, Malay names. The Bulletin Zeitschrift 43, 31-37. of the Raffl es Museum 24, 63-95. SCHINDLER, I., & J. SCHMIDT. 2006. Review of the VAN DER LAAN, R. 2010. Freshwater fi sh list. Fifth mouthbrooding Betta (Teleostei, Osphronemidae) CD edition. Verlag Natur & Wissenschaft; from Thailand, with descriptions of two new spe- Solingen. cies. Zeitschrift für Fischkunde 8, 47-68. VIERKE, J. 1978. Labyrinthfische und verwandte SCHMIDT, J. 1996. Vergleichende Untersuchungen Arten. Engelbert Pfriem Verlag, Wuppertal- zum Fortpflanzungsverhalten der Betta-Arten Elberfeld. (Belontiidae, Anabantoidei). Bibliothek Natur & VIERKE, J. 1979. Betta anabatoides und Betta foerschi Wissenschaft Bd. 7. Verlag Natur & Wissenschaft, spec. nov., zwei Kampffi sche aus Borneo. Das Solingen. Aquarium 13, 386-390. STUESSY, T.F. 1997. Classifi cation: more than just VIERKE, J. 1986. Labyrinthfi sche – Arten, Haltung, branching patterns of evolution. Aliso 15, 113- Zucht. Kosmos-Vivarium. Franckh’sche Verlags- 124. buchhandlung, Stuttgart. TAN, H.H. 2008. Betta rubra unveiled. AAGB Laby- VOLZ, W. 1904. Fische von Sumatra, gesammelt von rinth No. 152 March 2008, 5-7. Herrn G. Schneider. Revue Suisse de Zoologie TAN, H.H., & M. KOTTELAT. 1998. Two new species 12, 451-493. of Betta (Teleostei: Osphronemidae) from the VOLZ, W. 1907. Catalogue of the fi shes of Sumatra. Kapuas Basin, Kalimantan Barat, Borneo. The Natuurkundig Tijdschrift voor Nederlandsch- Raffl es Bulletin of Zoology 46, 41-51. Indie 66, 35-250. TAN, H.H., & P.K.L. NG. 2005a. The fi ghting fi shes VORIS, H. 2000. Maps of Pleistocene sea levels in (Teleostei: Osphronemidae: genus Betta) of Sin- Southeast Asia: shorelines, river systems and gapore, Malaysia and Brunei. The Raffl es Bulletin time durations. Journal of Biogeography 27, of Zoology Suppl. 13, 43-99. 1153-1167. TAN, H.H., & P.K.L. NG. 2005b. The labyrinth fi shes WÄGELE, J.-W. 2004. Hennig’s phylogenetic systemat- (Teleostei: Anabantoidei, Channoidei) of Sumatra, ics brought up to date, p. 101-125. In: Milestones Indonesia. The Raffl es Bulletin of Zoology Suppl. in Systematics (WILLIAMS, D.M., & P.L. FOREY, 13, 115-138. eds). CRC Press, Boca Raton, FL, USA. TAN, H.H., & P.K.L. NG. 2006. Six new species WEBER, M., & L.F. DE BEAUFORT. 1922. The fi shes of of fighting fish (Teleostei: Osphronemidae: the Indo-Australian Archipelago. IV. Heteromi, Betta)from Borneo. Ichthyological Exploration Solenichthyes, Synentognathi, Percesoces, Laby- of Freshwaters 17, 97-114. rinthici, Microcyprini. E. J. Brill, Leiden. TAN, H.H., & S. H. TAN. 1994. Betta miniopinna, a new WITTE, K.-E. 1997. Key to the genus Betta Bleeker, species of fi ghting fi sh from Pulau Bintan, Riau 1850. Pp 13-15 in: Aqualog, all Labyrinths – Archipelago, Indonesia (Teleostei: Belontiidae). Bettas, Gouramis, Snakeheads and Nandids Ichthyological Exploration of Freshwaters 5, (SCHÄFER, F. ed.). Verlag ACS, Mörfelden- 41-44. Walldorf. TAN, H.H., & S.H. TAN. 1996. Redescription of WITTE, K.-E., & J. SCHMIDT. 1992. Betta brownorum, a the Malayan fi ghting fi sh Betta pugnax (Teleostei: new species of anabantoids (Teleostei: Belontii- Belontiidae), and description of Betta pulchra, new dae) from northwestern Borneo, with a key to the species from peninsular Malaysia. The Raffl es genus. Ichthyological Exploration of Freshwaters Bulletin of Zoology 44, 419-434. 2, 305-330. TORTONESE, E. 1963. Catalogo dei tipi di pesci del Museo Civico di Storia Naturale di Genova. (Parte III). Annali del Museo Civico di Storia Naturale Received: 14.10.2011 ‘Giacomo Doria’ 73, 333-350. Accepted: 10.11.2011

Bull. Fish Biol. 13 (1/2) 31 Appendix of the head length and equal to the interorbital width. The scales on the head have the same size We provide a translation of the original de- as the scales on the body; on the pre-operculum scription by PERUGIA 1893, originally published are three series, all haired. in Italian. The dorsal fi n, with eight soft rays, has the penultimate ray elongate and when folded back Betta rubra, n. sp. reaching to the middle of the caudal fi n. The anal fi n also has the last rays elongated. The D. 8. A. 22. Ll. 30. pectorals are as long as the head. The ventral Lake Toba. Siboga. fi ns have the outer ray elongate and reach the The body height is a quarter of the total length, 6th or 7th ray of the anal fi n. the head length is one third of the total length. Lateral line absent. The mouth is shaped obliquely, jaws of the The general colouration of the body and the same length and with very small teeth. The eye fi ns is red with fi ve to six large dark blotches diameter is relatively large; it is about a quarter on the fl anks.