Taxonomic review of the genus Leptobrama Steindachner 1878 (Perciformes: Leptobramidae), with the resurrection of Leptobrama pectoralis (Ramsay and Ogilby 1887)
The leptobramid genus Leptobrama, distributed in southern New Guinea and northern Australia, is characterized by having a short head; a large mouth, with the posterior tip of the upper jaw extending beyond the level of the posterior margin of the eye; a single dorsal fin with four closely set spines, originating posterior to the level of the anal-fin origin; anal fin long, with three spines; pectoral fin short, not reaching to the level of the anus; ctenoid scales small, firmly adherent. The species of Leptobrama are revised taxonomically, and the genus comprises the species Leptobrama muelleri Steindachner 1878 and Leptobrama pectoralis (Ramsay and Ogilby 1887), the latter of which is resurrected herein. The diagnostic characters of these species are as follows: L. muelleri—head 21.7–22.7 % of standard length (SL), snout length 19.9–24.5 % of head length (HL), upper jaw length 53.0–61.3 % HL, vomerine tooth patch rhomboid, tip of pectoral fin almost reaching level of tip of appressed pelvic fin, lateral line gradually curved anteriorly, total gill rakers 13–18, scale rows below lateral line 13–14, prominent black spot distally on anterior dorsal fin; L. pectoralis—head 22.2–25.8 % SL, snout length 16.9–22.6 % HL, upper jaw length 61.8–66.1 % HL, vomerine tooth patch rounded, tip of pectoral fin extending beyond level of tip of appressed pelvic fin, lateral line steeply curved anteriorly, total gill rakers 6–12, scale rows below lateral line 15–18, no black spot distally on anterior dorsal fin.
KeywordsLeptrobramidae Leptobrama muelleri Leptobrama pectoralis Australia Papua New Guinea
The percoid fishes of the genus Leptobrama Steindachner 1878 are exclusively distributed in coastal waters and estuaries around northern Australia and southern New Guinea. The genus was established by Steindachner (1878) and characterized by numerous small pointed teeth on both jaws, vomer, palatines and pterygoids; the dorsal-fin base much shorter than the anal-fin base, and both fins covered with scales; slender spines closely adherent to the anterior dorsal and anal fins; and the body covered by coarse and closely fitting scales. At the same time, he originally described the type species of the genus, Leptobrama muelleri Steindachner 1878, on the basis of a single specimen collected at Townsville, Queensland, Australia. Only three years later, Macleay (1881) established the genus Neopempheris and described Neopempheris ramsayi Macleay 1881 based on a specimen collected from Rockingham Bay, Queensland, Australia. A second species of Neopempheris collected from the Aird River, Papua New Guinea, was described as Neopempheris pectoralis by Ramsay and Ogilby (1887), and was distinguished from N. ramsayi by differences in the coloration of the dorsal-fin tip, the shape of the lateral line, and the pectoral-fin size. Steindachner (1883) stated that N. ramsayi was the same species as L. muelleri. Ogilby (1913) synonymized both N. ramsayi and N. pectoralis with L. muelleri, based on the examination of four specimens and the holotypes of both species. Since that time, the genus Leptobrama has been considered as a senior synonym of Neopempheris, including the single species L. muelleri (see McCulloch 1914; Fowler 1928; McCulloch 1929; Fowler 1931; Munro 1967; Allen and Swainston 1988; Allen 1997; Mooi 2001; Hoese and Bray 2006; Nelson 2006).
During the senior and second authors’ field survey of marine fishes in eastern Indonesian waters, they found three individuals of unusual fishes caught in the Arafura Sea, at Ambon City Fish Market, Maluku. The fish had a silvery, glittering body, a pointed snout, a large mouth, a single dorsal fin, and subfalcate dorsal and anal fins. From these characters, we could easily recognize them as a species of Leptobrama, but the characters were not consistent with those of L. muelleri. Based on a detailed examination of 28 specimens belonging to the genus Leptobrama including the primary types of the three nominal species in its synonymy, we came to the conclusion that Leptobrama is not monotypic, but consists of two valid species. In the present account, these two species are distinguished, and their morphological characteristics are redescribed in detail.
Genus Leptobrama Steindachner 1878
Diagnosis. Head length 21.7–25.8 % SL; posterior tip of upper jaw extending beyond level of posterior margin of eye; dermal eyelid developed posteriorly; a single dorsal fin with four close-set spines originating posterior to level of anal-fin origin; anal fin long-based with three spines; pectoral fin not reaching to level of anus; scales small, ctenoid, firmly adherent.
Description. Body oblong, extremely compressed; dorsal and ventral contours of body convex; maximum depth at anal-fin origin. Head and snout short; mouth large, oblique; upper jaw slanted down posteriorly; dermal lips absent on both jaws. Teeth on both jaws small, sharp, slender, conical in shape, forming tooth patches extending to outer surface of jaws; teeth on the innermost row biggest in both jaws, those of lower jaw somewhat larger than those of upper jaw; small toothless area at anterior tips of both jaws; vomer, palatines, ectopterygoids, endopterygoids, basibranchials, hypobranchials, ceratobranchials, and tongue with tooth patches formed by minute conical teeth. Anterior nostril circular with small dermal flap at posterior margin, posterior nostril semicircular, somewhat larger than the anterior; eyes large. Opercle with no spines; pseudobranch developed; gill rakers rod-like or spatular; usually additional rudimentary stump-like gill rakers on distal parts of both upper and lower arches (Fig. 1). Dorsal-fin origin posterior to level of center of body; anal fin with three spines originating at level of center of body; anterior soft rays of dorsal and anal fins somewhat elongated, forming subfalcate fins; caudal fin deeply emarginate with nine + eight principal rays; pectoral fin short with two unbranched rays; pelvic fins with one spine and five soft rays. Head, body and all fins almost wholly covered by scales except for snout and opercular membrane; predorsal scaly area reaching to level of anterior margin of eye; lateral line complete. Vertebrae 10 + 14 = 24, nine supraneural bones.
Remarks. This strange-shaped fish has long attracted ichthyologists’ interest regarding its taxonomic position and phylogenetic relationships. Steindachner (1878) suggested that Leptobrama was close to the genus Brama Bloch and Schneider 1801 based on the morphology of fins and scales, although he stated that the shape of the body was similar to Scomberoides Lacepède 1801 or Lichia Cuvier 1816 (Carangidae). Macleay (1881) placed the genus Neopempheris (later synonymized with Leptobrama, as stated above) in the family Kurtidae, together with Pempheris Cuvier 1829. Subsequently, Ogilby (1913) established a subfamily Leptobraminae for the genus Leptobrama within the family Pempheridae. Tominaga (1963) redefined the family Pempheridae and excluded Leptobrama from that family. Consequently, Tominaga (1965) elevated the subfamily Leptobraminae to the family level as Leptobramidae, and defined the family in detail based on an elaborate examination of the internal morphology of Leptobrama, mainly the dorsal fin structure and its pterygiophores. Furthermore, he suggested a close relationship between the Leptobramidae and the Carangidae, especially the genus Scomberoides. Although most recent authors followed Tominaga’s concept of Leptobramidae, the family was arranged closer to Pempheridae than to Carangidae (Mooi 2001; Nelson 2006; Hoese and Bray 2006).
Key to the species of the genus Leptobrama
Prominent black spot distally on anterior dorsal fin, anterior lateral line gradually curved, total gill rakers 13–18, scale rows below lateral line 13–14 … Leptobrama muelleri
No black spot distally on anterior dorsal fin, anterior lateral line steeply curved, total gill rakers 6–12, scale rows below lateral line 15–18 … Leptobrama pectoralis
Leptobrama muelleri Steindachner 1878
Counts and measurements of the two species of Leptobrama
Holotype SMNS 2415
Holotype of Neopempheris ramsayi
Holotype AMS I.1308
Standard length (mm)
104–242 (160, 11)
62–300 (178, 14)
Dorsal-fin soft rays
16–18 (17.5, 11)
17–19 (17.9, 14)
Anal-fin soft rays
26–28 (27.0, 11)
26–30 (28.1, 14)
Branched pectoral-fin rays
14–16 (14.9, 11)
14–16 (14.7, 13)
Gill rakers on upper arch
3–5 (4.6, 11)
1–4 (2.4, 13)
Gill rakers on lower arch
9–13 (10.8, 11)
5–9 (6.7, 13)
Total gill rakers
13–18 (15.5, 11)
6–12 (9.1, 13)
Scales on curved part of lateral line
30–37 (32.4, 11)
Scales on straight part of lateral line
42–46 (44.5, 11)
Total scales on lateral line
76–80 (76.9, 11)
74–78 (76.3, 12)
Scales above lateral line
10–12 (10.9, 11)
11–13 (11.9, 12)
Scales below lateral line
13–14 (13.7, 11)
15–18 (16.0, 13)
As % of standard length
21.7–22.4 (22.0, 6)
22.2–25.8 (23.3, 9)
57.0–60.9 (59.2, 6)
58.8–60.9 (59.9, 9)
Dorsal-fin base length
22.9–24.8 (24.1, 6)
23.8–37.9 (27.1, 9)
Anal-fin base length
33.8–38.8 (36.6, 6)
36.0–40.2 (38.9, 9)
Snout to pectoral-fin insertion
22.9–24.0 (23.4, 6)
22.4–26.1 (24.1, 9)
Snout to pectoral-fin tip
37.3–39.6 (38.5, 5)
39.6–45.8 (41.8, 8)
Snout to pelvic-fin insertion
26.4–29.3 (28.2, 6)
27.9–33.5 (29.3, 9)
Snout to anal-fin origin
47.7–56.6 (52.0, 6)
49.6–54.9 (52.4, 9)
Length of longest dorsal-fin ray
13.7–17.2 (15.5, 6)
15.1–19.1 (17.0, 9)
Length of longest anal-fin ray
13.7–15.6 (15.0, 6)
10.1–17.7 (15.1, 9)
14.3–17.2 (16.3, 5)
16.3–20.3 (18.4, 8)
11.3–13.1 (12.0, 6)
9.1–15.2 (11.5, 9)
Body depth at anal-fin origin
30.2–33.2 (30.9, 6)
31.1–35.2 (33.5, 9)
Caudal peduncle length
12.9–16.1 (14.3, 6)
13.2–17.0 (14.8, 9)
Caudal peduncle depth
9.5–10.7 (10.1, 6)
9.8–11.3 (10.7, 9)
As % of head length
21.0–24.5 (22.4, 6)
16.9–22.6 (20.6, 9)
Upper jaw length
56.5–61.3 (58.4, 6)
61.8–66.1 (64.5, 9)
24.3–32.5 (28.4, 6)
19.9–30.3 (25, 9)
Dermal eye opening
15.9–28.2 (21, 4)
10.4–25.6 (15.9, 9)
Postorbital head length
49.3–56.8 (52.4, 6)
48.5–64.1 (58.6, 9)
Length of longest gill raker
12.1–15.2 (13.3, 3)
7.8–9.2 (8.5, 2)
23.0–35.1 (27.1, 6)
22.0–26.9 (24.7, 9)
As % of upper jaw length
35.6–43.4 (38.3, 6)
27.3–34.4 (31.9, 9)
Leptobrama mülleri Steindachner 1878: 388 (type locality: Endeavour Strait, Queensland, Australia, erroneously given as Cleveland Bay, Townsville); Steindachner 1879: pl. 3, fig. 1 (figure of type only).
Neopempheris ramsayi Macleay 1881: 517 (type locality: Rockingham Bay, Queensland, Australia).
Leptobrama mülleri: Ogilby 1913: 64 (in part; Australia and southern New Guinea); McCulloch 1914: 214 (Australia); Fowler 1928: 191 (in part; Queensland, Australia and southern New Guinea); Fowler 1931 (in part; New Guinea and Queensland); Tominaga 1965: 33 (Port Hedland, Western Australia).
Leptobrama muelleri: McCulloch 1929: 235 (in part; Queensland, Western Australia and New Guinea); Mooi 2001: 3210 (in part; southern Papua New Guinea and northern Australia); Hoese and Bray 2006: 1280 (? in part; Northern Territory, Queensland, Western Australia, New Guinea).
Holotype. SMNS 2415 (formerly the smaller specimen of SMNS 2415), 132 mm SL, sex unknown, Endeavour Strait, Queensland, Australia, collected by F. von Müller, Aug. 1878.
Other materials. Twelve specimens, 104–242 mm SL. AMS I.1396, holotype of Neopempheris ramsayi, 181 mm SL, male, Rockingham Bay, Queensland, Australia; AMS I.13134, 242 mm SL, female, Port Hedland, Western Australia, Australia; AMS IB.1583, 240 mm SL, Onslow, Western Australia, Australia; NMW 76432, 127 mm SL, sex unknown, ?Townsville, Cleveland Bay, Queensland, Australia; QM I1015*, 209 mm SL, Moreton Bay, Queensland, Australia; QM I27567*, 104 mm SL, sex unknown, Norman River (near mouth), Queensland, Australia; QM I28467*, 182 mm SL, sex unknown, Freshwater Bay Beach, Queensland, Australia; QM I39510*, two specimens, 124–128 mm SL, sex unknown, Karumba Point Beach, Queensland, Australia; SMNS 2293, two specimens, 120–145 mm SL, sex unknown, Townsville, Cleveland Bay, Queensland, Australia; SMNS 27071 (formerly the larger specimen of SMNS 2415), 143 mm SL, sex unknown, Endeavour Strait, Queensland, Australia (asterisks indicate the specimens for which only some count data are available).
Materials for distributional records. Eleven specimens. QM I2007, Cape Bowling Green, Queensland, Australia; QM I4568, Cleveland Bay, Queensland, Australia; QM I6753, Upstart Bay, Queensland, Australia; QM I6767, Cape Cleveland, Queensland, Australia; QM I11200, southeast corner of Gulf of Carpentaria, Queensland, Australia; QM I12518, Gladstone, Queensland, Australia; QM I16551, Murray River mouth, Queensland, Australia; QM I22719, I22770, Sarina Inlet, Queensland, Australia; QM I24019, Cleveland Bay, Queensland, Australia; QM I39146, O’Connell River, Queensland, Australia.
Frequency distributions of gill rakers, lateral-line scales and scale-rows above and below lateral line in the two species of Leptobrama
Description. Counts and measurements of the holotypes of Leptobrama muelleri and Neopempheris ramsayi, and other specimens are shown in Table 1. Body somewhat slender; dorsal contour round, gradually convex; ventral contour anterior to anal-fin origin weakly convex but anal-fin base straight; head and mouth relatively small, snout rather long; lower margins of lachrymal and preopercle finely serrated (including holotype) or almost smooth; posterior margin of preopercle entire; normal gill rakers long and slender, rod-like; supraneural and pterygiophore insertion pattern 0/0/0//0/0/0/0/0/0/1 (two specimens), 0/0/0/0//0/0/0/0/0/1 (four, including holotype), 0/0/0/0//0/0/0//00/1 (one), 0/0/0/0//0/0//00/0/1 (one). Maximum recorded length 242 mm SL (AMS I.13134).
Color of preserved specimen. Head and body generally brownish dorsally, silvery white ventrally; snout and all fins pale yellowish brown.
Total length and counts of dorsal- and anal-fin soft rays of “syntypes” of Leptobrama muelleri
Total length (mm)
Dorsal-fin soft rays
Anal-fin soft rays
Smaller specimen of SMNS 2293
Larger specimen of SMNS 2293
Steindachner (1878) did not provide a figure of L. muelleri in the original description, which was an abbreviated abstract of the later paper by Steindachner (1879) who presented the figure with the caption “Fig. 1. Leptobrama Mülleri Steind. (s. Steind. Ichthyol. Beitr. VIII. Heft)” (Fig. 3a). Although the figured specimen was not specified as the holotype, we considered that as the holotype of L. muelleri because the dorsal- and anal-fin soft rays of the figured specimen can be counted with 17 and 27, respectively, and the total length of the figure measured ca. 165 mm TL. Also, the holotype was the only specimen available to Steindachner at the time, as noted in the original description, and must therefore have been illustrated in the figure. Steindachner’s figures are usually printed in the same size as the figured specimens (T. Yoshino, personal communication). The shape of holotype also agrees well with the figure. Concerning Steindachner’s (1879) figure, Ogilby (1913) stated as a personal communication from McCulloch that the figure might have been made from Ramsay’s specimen (the holotype of Neopempheris ramsayi) because the figure agreed with the holotype so well. However, we believe the figure was made from the holotype of L. muelleri because of the above-mentioned facts. The holotype of N. ramsayi measuring 181 mm SL [223 mm from the tip of the snout to the tip of the lower lobe of caudal fin (upper lobe of caudal fin broken), 9 inches in the original description] is much longer than the specimen in Steindachner’s figure.
Neopempheris ramsayi (Figs. 1c, 3b) was described by Macleay (1881) on the basis of a single specimen from Rockingham Bay, Queensland. Shortly after the description of N. ramsayi was published, Steindachner (1883) synonymized N. ramsayi with L. muelleri. Because all morphological characters of the holotype of N. ramsayi agree well with those of L. muelleri (Table 1), both nominal species are here considered as conspecific, with L. muelleri the senior synonym.
Leptobrama pectoralis (Ramsay and Ogilby 1887)
Neopempheris pectoralis Ramsay and Ogilby 1887: 388 (type locality: Aird River, Gulf Province, Papua New Guinea).
Holotype. AMS I. 1308, 189 mm SL, female, 30 miles up the Aird River from its mouth, Papua New Guinea.
Other materials. Fourteen specimens, 62–300 mm SL. AMS I. 12816, 203 mm SL, female, Moreton Bay, Queensland, Australia; AMS I.15557-188, two specimens, 105–106 mm SL, sex unknown, Gulf of Carpentaria, Queensland, Australia; AMS I.34303-009, three specimens, 266–297 mm SL, females, Shoalwater Bay, Sabina Beach, Queensland Australia; AMS IB.1258, 62 mm SL, sex unknown, Fitzroy River, Mackenzie Island, Queensland, Australia; FRLM 23463, 23464, 204–225 mm SL, females, Arafura Sea (purchased in Ambon, Indonesia); QM I27586*, three specimens, 117–124 mm SL, sex unknown, Karumba Point beach (near Norman River mouth), Queensland, Australia; QM I39511*, 105 mm SL, sex unknown, Karumba Point beach (near Norman River mouth), Queensland, Australia; USNM 518542*, 300 mm SL, sex unknown, Parama I., Papua New Guinea (asterisks indicate the specimens for which only some count data are available).
Materials for distributional records. Thirteen specimens. QM I1017, Rockhampton area, Queensland, Australia; QM I4164, I4165, Yeppoon, Queensland, Australia; QM I5502, Gladstone, Queensland, Australia; QM I6752, Upstart Bay, Queensland, Australia; QM I8115, Proserpine River, Queensland, Australia; QM I25066, I25067, I25068, I25809, I25810, Arthurs Creek mouth, Gulf of Carpentaria, Queensland, Australia; QM I28395, Sabina Point, Queensland, Australia; QM I39147, Repulse Creek, Queensland, Australia.
Diagnosis. A species of Leptobrama defined by the following combination of characters: head length 22.2–25.8 % SL, snout length 16.9–22.6 % HL, upper jaw length 61.8–67.3 % HL, snout length 27.3–34.4 % of upper jaw length (Table 1); total gill rakers 6–12 (Table 2); scale rows below lateral line 15–18; vomerine tooth patch rounded; tip of pectoral fin extending beyond level of tip of appressed pelvic fin; lateral line steeply curved anteriorly (Fig. 2c, d); no black spot distally on anterior dorsal fin.
Description. Counts and measurements of the holotype of Leptobrama pectoralis and other specimens are shown in Table 1. Body rather deep; dorsal contour rather linear or triangular, ascending to dorsal-fin origin, then descending to caudal peduncle; ventral contour anterior to anal-fin origin convex, but anal fin base straight; head relatively large, snout very short, mouth large; lower margin of lachrymal very finely serrated; lower margin of preopercle with obvious serration, posterior margin finely serrated; normal gill rakers short, spatular in shape; supraneural and pterygiophore insertion pattern 0/0/0//0/0/0/0/0/0/1 (one specimen), 0/0/0/0//0/0/0/0/0/1 (eight, including holotype), 0/0/0/0/0//0/0/0/0/1 (two). Maximum recorded length 300 mm SL (USNM 518542).
Color of thawed specimen. Head and body somewhat dark bluish silver dorsally, brilliant silver ventrally; tip of snout with a black spot; fins somewhat dusky (based on the photographs of FRLM 23463 and 23464).
Color of preserved specimen. Head and body generally brownish dorsally, silvery white ventrally; snout and all fins pale yellowish brown; tip of snout with a black spot.
Distribution and ecological note. This species is hitherto known from southern Papua New Guinea (Aird River, Gulf Province and Parama I., Western Province), Queensland, Australia (Gulf of Carpentaria to Moreton Bay), Northern Territory, Australia (Adelaide River; http://www.fishbase.org/photos/PicturesSummary.php?ID=4338&what=species; accessed 15 Jan. 2016) and the Arafura Sea (Fig. 4). Also, there is little information about the ecology of this species. The holotype was obtained when it jumped into the boat at about 30 miles up the Aird River, Papua New Guinea (Ramsay and Ogilby 1887). This indicates that the fish is a pelagic species entering or inhabiting the lower reaches of rivers.
Remarks. As stated above, Ramsay and Ogilby (1887) established their new species, Neopempheris pectoralis (=Leptobrama pectoralis) on the basis of the absence of a conspicuous black spot on the dorsal fin, the shape of lateral line and the much larger pectoral fin, and they stated that N. pectoralis was easily distinguishable from N. ramsayi (=L. muelleri) by these characters. Of these characters, the shape of the lateral line and size of pectoral fin are diagnostic characters for these two species as indicated here in “Diagnosis” of both species, but Ramsay and Ogilby (1887) did not conclusively elucidate the differences between the species. Although Ogilby (1913) accurately understood the morphological differences between N. pectoralis and L. muelleri, namely the spotted dorsal-fin form (=L. muelleri) with more slender body, shorter pectoral fin and less arched lateral line than spotless form (=N. pectoralis), he considered that both forms were conspecific based on the examination of four non-type specimens (one male with a dark spot on the dorsal fin and three females with a uniform dorsal fin) and both holotypes of N. ramsayi and N. pectoralis. He believed that those differences were attributable to sexual dimorphism. However, as stated above, we conclude that those differences are not due to sexual or intraspecific variation but to interspecific variation, as we found both sexes of L. muelleri (=spotted dorsal-fin form) in this study.
We are deeply indebted to M. A. McGrouther (AMS) for the loan of the AMS specimens to the first author, J. Johnson (QM) for permission to add his data to the present article and critical comments and suggestions on the manuscript, S. Merker (SMNS) for the loan of SMNS specimens, J. T. Williams and S. Raredon (USNM) for their examining and photographing a USNM specimen, E. Mikschi and H. Wellendorf (NMW) for their help during the first author’s visit to NMW and examination of NMW 76432, K. Hatooka (OMNH) for photographs and measurements of NMW 76432, and T. Yoshino (Okinawa Churashima Research Center) for information about Steindachner’s figures. This study was supported in part by JSPS KAKENHI Grant Numbers 19570084 and 23570114 awarded to the first author, and by the Multilateral Cooperative Research Program (Coastal Oceanography) and Asian Core Program of the JSPS.
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