Barbodes montanoi ( Sauvage, 1881 )

Hui, Tan Heok & Husana, Daniel Edison M., 2021, Barbodes pyrpholeos, new species, the first cave-dwelling cyprinid fish in the Philippines, with redescription of B. montanoi (Teleostei: Cyprinidae), Raffles Bulletin of Zoology 69, pp. 309-323 : 315-322

Barbodes montanoi ( Sauvage, 1881)

( Figs. 8–11 )

Puntius montanoi Sauvage, 1881: 103 ; Bertin & Estève, 1948: 29. Barbodes montanoi (Sauvage) – Herre, 1924: 292; 1953: 125;

Kottelat, 2013: 79.

Material examined. Syntypes — MNHN 3398 , 1 ex., ca. 90 mm TL; Philippines: Mindanao , Simulao River; Montano, 1880 [low resolution image only; https://science.mnhn.fr/ institution/mnhn/collection/ic/item/list?scientificName=punt ius+montanoi] . — MNHN 3399 , 4 ex. (of 12); Philippines: Mindanao [low resolution images and digital radiographs only; https://science.mnhn.fr/institution/mnhn/collection/ic/ item/a-3399?listIndex=2&listCount=2] .

Non-type material — PNM 15652, 6 ex. , ZRC 61233, 6 ex., 17.5–87.2 mm SL; Philippines: Mindanao, Agusan drainage; Enchanted Falls ; coll. D. E. Husana, 14 May 2014 . — ZRC 61234, 2 ex., 50.0– 67.2 mm SL; Philippines: Mindanao, Agusan drainage; pools at Ugnop; coll. Prosperidad Mountaineering Club, 1 May 2012 .

Diagnosis. Entire body yellowish-brown dorsally and cream ventrally, with faint brown reticulate pattern corresponding to anterior margin of exposed part of body scales; body pattern consisting of four to six black dots or dashes (dependent on ontogenetic stage), two positioned at base of dorsal fin and anal fin (anal-fin spot missing or faint in adults), one at dorsal part of opercular opening (missing or faint in adults), one at mid-body below dorsal-fin origin, one at mid-body above anal-fin origin, one at middle of caudal peduncle anterior to caudal fin; unpaired fins anterior half golden-yellow, posterior half reddish-orange; paired fins yellowish or hyaline; eyes present and pigmented; last unbranched dorsal-fin ray spinous, with 8–12 weak serrations on specimens smaller than 62 mm SL (see Fig. 8 ); lateral scale series 24–27 (mode 24); predorsal scales 8.

Barbodes montanoi is chiefly distinguished from other selected Philippine and Sundaic congeners as follows: from B. amarus , B. lateristriga , and B. kuchingensis by having a pattern of black spots on the body (vs. body without black markings in B. amarus ; pattern of two black bars on anterior half of body with central black stripe on posterior half of body in B. lateristriga and B. kuchingensis ); from B. manguaoensis and B. palavanensis by having larger black spots, covering 2–3 scales (vs. smaller spots, restricted to single scale); from B. banksi , B. bunau , and B. rhombeus by having a small round black spot below dorsal-fin base (vs. black triangular marking in B. banksi and B. bunau ; short black bar in B. rhombeus ); from B. binotatus by having fewer black spots on body when young (ca. 4 vs. ca. 9); from B. dunckeri by the absence (vs. presence) of orange pigment on the paired fins in life; from B. everetti by having distinct black spots on body in sub-adults and adults (vs. black spots on body connected by black lateral stripe in sub-adults and adults); and from B. sealei by having only 8–12 weak serrations on last unbranched dorsal-fin ray in specimens smaller than 62 mm SL (vs. last unbranched dorsal-fin ray with 13–21 welldeveloped serrations in specimens larger than 60 mm SL).

Description. See Figs. 9–11 for general impression, and Table 1 for meristic and morphometric data.

Body slim, deepest at dorsal-fin origin (31.8–35.7% SL), tapering to its narrowest at caudal peduncle (13.8–15.3% SL). Head pointed (head depth 68.9–73.9% HL, head width 54.5–60.7% HL), relatively short, less than ⅓ of standard length (head length 27.4–29.5% SL). Mouth sub-terminal, two pairs of maxillary barbels, barbel not longer than eye diameter, posterior pair longer than anterior, reaching past middle of eye. Eye relatively large (eye diameter 21.9–29.4% HL). Dorsal-fin origin and pelvic-fin origin at mid-body (predorsal-fin length 53.1–56.1% SL; pre-pelvic fin length 51.8–54.5% SL), anal-fin origin at ¾ length of body (preanalfin length 74.7–76.9% SL). Dorsal and anal fins triangular in shape, with a smooth and straight anterior margin, straight posterior margin. Caudal fin forked with upper lobe slightly longer (upper caudal-fin lobe length 30.7–33.2% SL; lower caudal-fin lobe length 29.1–30.4% SL). Pelvic fin triangular with straight margin, pectoral fin rounded. Lateral series with 24–27 scales (mode 24), 8 predorsal scales, 4½.1.3 transverse scales, 12 circumpeduncular scales, 4½ scales between lateral series and pelvic-fin origin. Vertebral count 17–19 + 13–14 = 31–32 (mode = 32; n = 9).

Live colouration. See Fig. 9 for fresh colouration. Head and body dorsum brownish, sides golden, venter silverish to cream. Body scales with anterior portion with narrow, dark brown edge, giving a reticulation pattern. Body with distinct black to dark brown dots or dashes (number of dots variable dependent on maturation stage), one dot just above dorsal part of opercle opening, one dot at dorsal-fin base, one dot along mid-body below dorsal-fin origin, one dot along mid-flank above anal-fin origin, one dot at anal-fin base, dot spot along middle of caudal peduncle before caudal-fin base. Unpaired fins with anterior portions yellowish and middle to posterior portions orange to red. Paired fins yellowish-brown.

Preserved colouration. See Figs. 10 , 11 for preserved colouration. Head and body brownish, darker on dorsum and cream on ventrum. Four to six distinct dark brown/ black dots or dashes along flank, all larger or equal to eye diameter. Faint brown blotch on body just posterior to edge of opercle. First distinct dark brown spot at anterior base of dorsal fin, second dark brown spot on mid-flank below dorsal-fin origin, third dark brown tiny blotch along mid-flank above anal-fin origin, fourth dark brown spot on middle of caudal peduncle before caudal-fin base. A faint brown spot can sometimes be visible above the anal-fin base. Some individuals (50–60 mm SL) with interrupted dark brown lateral stripe, running from opercle edge to caudal-fin base. All fins hyaline.

In a juvenile ( Fig. 11 ; 32.2 mm SL), six distinct black to dark brown spots can be observed. The locations of the spots as above, except the spot on body just posterior to opercle edge and above the anal-fin origin are very distinct. All spots equivalent to or slightly larger than eye diameter.

The ontogenic change in body pattern is exhibited where the number of body spots decreases with age, as noted by Kottelat (2013: 482).

Distribution. Reported from Mindanao only. This species may be restricted to the Agusan River basin in Mindanao, Philippines. Further surveys are necessary to clarify the true distribution of this species.

Field notes. Syntopic fish species from Enchanted Falls locality include: Nomorhamphus cf. bakeri (Zenarchopteridae) , Gulaphallus cf. panayensis (Phallostethidae) , and multiple species of goby ( Gobiidae ).

Remarks. The type locality for B. montanoi is Simulao River, in central Mindanao ( Sauvage, 1881; Kottelat, 2013); and not Simulac River as stated by Fricke et al. (2021). The specimens of B. montanoi examined herein are from the water pools at Ugnop and Enchanted Falls, and likely represent true B. montanoi .

This species was named after the celebrated French traveller and collector Joseph Montano (1844–1914), who passed through Mindanao in 1880 ( Herre, 1924: 293). The original description of Puntius montanoi ( Sauvage, 1881: 103, 104) is based on 13 individuals ( Kottelat, 2013). The MNHN currently contains specimens labelled as both holotype (MNHN 3398) and paratypes (MNHN 3399, 12 individuals) of Puntius montanoi , likely following Bertin & Estève (1948). Following ICZN article 72.1, these 13 specimens must be referred to collectively as syntypes for the following reasons: 1) Sauvage (1881) did not explicitly designate a holotype for Puntius montanoi ; and 2) no lectotype designation has taken place in the years subsequent to the original description.

Sauvage (1881) stated that the third bony dorsal ray is smooth (presumably without serrations) and nearly the length of the head. Based on our examination of the recently collected specimens from Enchanted Falls and radiographs of nine additional specimens (ZRC 61233 and PNM 15652), specimens smaller than 62 mm SL have 8–12 weak serrations on the spinous last unbranched dorsal-fin ray, and specimens larger than 65 mm SL have no visible serrations. This could possibly be attributed to ontogenetic changes with size. The radiograph of four specimens listed as MNHN 3399 shows no signs of serrations on the spinous last unbranched dorsalfin ray, either indicating that they are mature individuals or the digital radiograph is of a resolution grade insufficient to show such fine details. Herre (1924: 293) noted that the absence of dorsal-fin ray serrations would make this species unique among lowland cyprinids found in Mindanao Island. General notes on Barbodes . There are at least 27 other Barbodes species recorded from the Philippines, most of them poorly known and either not depicted with figures when originally described or badly illustrated. Within Mindanao Island in the Philippines, six other Barbodes have been recorded from riverine habitats (fide Kottelat, 2013 — B. cataractae , B. joaquinae , B. montanoi , B. quinquemaculatus , B. resimus , and B. umalii ). There are 17 endemic species of Barbodes that occur only from within Lake Lanao and the single outflow river— B. amarus , B. baoulan , B. clemensi , B. disa , B. flavifuscus , B. herrei , B. katolo , B, lanaoensis, B. lindog , B. manalak , B. pachycheilus , B. palaemophagus , B. palata , B. sirang , B. tras , B. truncatulus , and B. tumba (fide Herre, 1924; Myers, 1960; Kottelat, 2013). From Mindoro, B. hemictenus is recorded; from Balabac, B. ivis is known; from Palawan, B. manguaoensis and B. palavanensis are listed (see Kottelat, 2013).

From Borneo, B. bunau is described from Sesayap basin in North Kalimantan; B. banksi , B. everetti , B. kuchingensis , and B. lateristriga are recorded from the southern half of Borneo; B. xouthos is known only from Lake Merimbun in Tutong District, Brunei Darussalam, northwestern Borneo; and B. sealei has been collected from throughout Borneo.

From Peninsular Malaysia, B. banksi , B. dunckeri , B. lateristriga , and B. rhombeus are found. From Java, B. binotatus sensu stricto is now assumed to be restricted to western Java; B. microps is known only from the Gunung Sewu cave system, Central Java (according to Weber & de Beaufort, 1916: 186).

Amongst all these species of Barbodes , B. montanoi looks superficially most similar to B. sealei (only known from Borneo; see Fig. 12 ), but differs in the following characters: body spots tend to be stretched longitudinally thus appearing ovoid (vs. taller and rounder spots for B. sealei ); both species with faint dark brown reticulate pattern on body, but with different extent of pigmentation— B. montanoi with only anterior portion of non-embedded body scales with brown pigments, vs. B. sealei with crescentic anterior margin of non-embedded body scales with brown pigments; dorsal and anal fins hyaline (vs. presence of narrow dusky black posterior margins for B. sealei ); slimmer upper and lower caudal-fin lobes (vs. broad caudal-fin lobes for B. sealei ); fewer vertebral count than B. sealei (31–32 [mode = 32, n = 9], vs. 33 [n = 4]); fewer and weak serrations on last unbranched dorsal-fin ray (8–12, vs. 13–21 stronger serrations for B. sealei ).

There are five Barbodes species from the main Sundaic landmass which may look similar to B. montanoi in terms of having a spotted body pattern. They include B. banksi ( Fig. 13 ), B. binotatus ( Fig. 14 ), B. rhombeus ( Fig. 15 ), B. dunckeri ( Fig. 16 ), and B. everetti ( Fig. 17 ). The common feature of these five taxa is that at the juvenile stage, the body pattern consists of numerous black spots and blotches (see Kottelat, 2013: 482). For B. banksi , B. binotatus , and B. rhombeus , the small spots are lost upon maturity, the adult only retaining the black spot or blotch below the dorsal-fin base, supra-anal region, and caudal peduncle (see Figs. 13–15 ). For B. dunckeri and B everetti , these spots and blotches enlarge with age (see Figs. 16 , 17 ). Another common feature for all five taxa is the presence of serrations on the spinous last unbranched dorsal-fin ray at both juvenile and mature stages. Barbodes montanoi also exhibits a slenderer body when compared to B. banksi , B. rhombeus , B. dunckeri , and B. everetti .

Herre (1924, 1935) and Myers (1960) suggested that the ancestral stock of the Barbodes species flock in Lake Lanao is Barbodes binotatus . They also indicated that B. binotatus is found throughout the island of Mindanao except within Lake Lanao. At that juncture, the species of origin for the Lake Lanao species flock was not doubted. In today’s context, B. binotatus sensu stricto is believed to be restricted to the island of Java (M. Kottelat, pers. comm.). All other populations of B. binotatus sensu lato are believed to be other named taxa (currently in the synonymy of B. binotatus ) or undescribed taxa (M. Kottelat, ongoing research).

Comparative material examined

Philippines.

Barbodes amarus — ZRC 51648, 2 ex., 79.5–125.6 mm SL; Mindanao : Lake Lanao; April 2007 .

Barbodes manguaoensis — ZRC 60530, 10 ex., 25.2–54.8 mm SL; Palawan: Taytay, Lake Manguao ; 13 October 1988 . Barbodes palavanensis — ZRC 60529, 10 ex., 41.2–57.6 mm SL; Palawan, Irawan ; 18 November 1988 .

Sundaic area.

Barbodes banksi — ZRC 43643, 10 ex., 23.1–49.7 mm SL; Sarawak: Lundu, Sungai Sebiris, 8.7 km to Sematan on Lundu-Sematan road; 2 October 1998 . — ZRC 39387, 5 ex., 58.5–72.5 mm SL; Sarawak: Kampung Matang, 1.3 km to Sungai Cina Matang; 4 September 1995 . — ZRC 60155, 9 ex., 35.5–91.6 mm SL; Sarawak: Song District : Rejang River basin: Katibas River : Ulu Katibas, Sungei Merating, draining into Sungei Bloh, 01°38.045′N, 112°17.350′E, 121 m asl; 19 May 2018 . — ZRC 60076, 18 ex., 14.8–96.0 mm SL; Sarawak: Song District : Rejang River basin: Katibas River : Ulu Katibas , Sungei Satap, ca. 25 min upriver of Nanga Bloh Field Station, 01°39.240′N, 112°15.108′E, 117 m asl; 24 May 2018 .

Barbodes binotatus — ZRC 40127, 13 ex., 21.2–72.2 mm SL; Java: Bogor, tributary of Cipinang Gading ; 10 July 1996 . — ZRC 40141, 8 ex., 33.6–61.0 mm SL; Java: Desa Cinangueng, Kecamatan Ciampea ; 10 July 1996 .

Barbodes dunckeri — ZRC 60671, 1 ex., 79.6 mm SL; Malaysia: Johoe , Kahang, Gunung Belumut Recreational Forest; 11 June 2018 . — ZRC 37602, 1 ex., 58.4 mm SL; Malaysia: Johoe , near Kota Tinggi falls; 1993 .

Barbodes everetti — ZRC 54289, 13 ex., 12.6–71.6 mm SL; Sarawak: Sematan outskirts, Kampung Poeh, hill stream at base of Gunung Poeh; 5 November 2013 .

Barbodes rhombeus — ZRC 55401, 7 ex., 36.6–60.7 mm SL; Singapore: Mandai Track 15 stream; 18 June 2015 . — ZRC 54686, 20 ex., 25.6–87.2 mm SL; Singapore: Mandai Track 15 stream; 28 July 2015 .

Barbodes sealei — ZRC 40411, 3 ex., 46.6–70.1 mm SL; Malaysia: Sabah: Danum Valley, Sungei Bilong ; 3 October 1996 . — ZRC 61238, 5 ex., 37.6–94.0 mm SL; Malaysia: Sabah: Kalabakan basin, un-named stream en route to SAFE camp; 21 April 2017 . — ZRC 56188, 1 ex., 65.4 mm SL; Sabah: Sandakan, Kabili-Sepilok ; 27 April 2017 . — ZRC 60536, 5 ex., 59.6–90.5 mm SL; Sabah: Kalabakan Forest Reserve, Brantian basin, 5 m buffer stream; 4 May 2015 . — ZRC 60359, 18 ex., 29.0–80.0 mm SL; Sarawak: Sarawak Kiri basin, Annah Rais, Sungai Timouh ; 28 September 2018 .

Figures

Fig. 8. Barbodes pyrpholeos (left), 104.1 mm SL, showing smooth last unbranched dorsal-fin ray (damaged, broken mid-way but healed); Barbodes montanoi (right), 62.8 mm SL, showing last unbranched dorsal-fin ray with weak serrations. (Images by Tan Heok Hui)

Fig. 9. Live colouration of Barbodes montanoi, about 90 mm SL, from Enchanted Falls. (Photograph by Daniel Edison M. Husana)

Fig. 10. Barbodes montanoi, ZRC 61233, 87.0 mm SL, adult, Enchanted Falls. Top: dorsal view; middle: lateral view; bottom: radiograph. (Photographs and radiograph by Tan Heok Hui)

Fig. 11. Barbodes montanoi, ZRC 61233. Top: 55.0 mm SL sub-adult; middle: 52.9 mm SL sub-adult; bottom: 32.2 mm SL juvenile; illustrating ontogenetic variation of black spot pattern (not to scale). (Photographs by Tan Heok Hui)

Fig. 12. Barbodes sealei, ZRC 40411, Sabah: Danum. Top and middle: 68.3 mm SL sub-adult (note strong serrations on dorsal-fin spine); bottom: 46.6 mm SL juvenile (both images right-side reversed). (Photographs and radiograph by Tan Heok Hui)

Fig. 13. Barbodes banksi, Sarawak: Lanjak. Top: 79.0 mm SL; bottom: 19.4 mm SL (to scale). (Photographs by Tan Heok Hui)

Fig. 14. Barbodes binotatus, Java: Bogor. Top: 40.8 mm SL; bottom: 16.8 mm SL (to scale). (Photographs by Tan Heok Hui)

Fig. 15. Barbodes rhombeus, Singapore (non-native introduction). Top: 70.2 mm SL; bottom: 19.1 mm SL (to scale). (Photographs by Tan Heok Hui)

Fig. 16. Barbodes dunckeri, Malaysia: Kahang. Top: 81.1 mm SL; bottom: aquarium trade, ca. 30 mm SL (to scale). (Photographs by Tan Heok Hui)

Fig. 17. Barbodes everetti, Sarawak: Poeh, ZRC 54289. Top: 72.0 mm SL; bottom: 17.8 mm SL (to scale). (Photographs by Tan Heok Hui)