Phylogenetic systematics of the viviparous halfbeak genera Dermogenys and Nomorhamphus (Teleostei: Hemiramphidae: Zenarchopterinae)

Zoological Journal of the Linnean Society (2001), 133: 199–283. With 69 figures doi: 10.1006/zjls.2000.0281, available online at http://www.idealibrary.com on

Phylogenetic systematics of the viviparous halfbeak genera Dermogenys and Nomorhamphus (Teleostei: Hemiramphidae: Zenarchopterinae) AMY DOWNING MEISNER∗ Department of Biological Sciences, The George Washington University, Washington D.C. 20052, USA Received August 2000; accepted for publication November 2000

Among the 13 genera and over 100 species of halfbeaks, three genera – Dermogenys, Nomorhamphus and Hemirhamphodon – are internally fertilized and viviparous. These genera belong to a more inclusive clade, the Zenarchopterinae, that also includes Zenarchopterus, inferred to be internally fertilized and to lay fertilized eggs, and the monotypic Tondanichthys, also inferred to be internally fertilized. Whereas the Hemiramphidae are distributed worldwide, internally fertilized halfbeaks are restricted to Southeast Asia. Recent data from histological surveys of the gonads of both males and females as well as embryonic modifications associated with viviparity have been combined here with osteological characters in a phylogenetic analysis. Results indicate overwhelming support for a sister-group relationship between Hemirhamphodon and (Dermogenys+Nomorhamphus). Monophyly of the Dermogenys+Nomorhamphus clade is also well supported. These results confirm earlier suggestions that Dermogenys, as previously defined, is paraphyletic. Within the Dermogenys+Nomorhamphus clade, two monophyletic clades are supported: one comprises ten species including four new species (Dermogenys bruneiensis, Dermogenys robertsi, Dermogenys palawanensis and Dermogenys collettei ) and the other comprises 13 species including three undescribed species (Nomorhamphus rossi, Nomorhamphus pinnimaculata and Nomorhamphus manifesta). Diagnoses for the species of Dermogenys and Nomorhamphus, as well as a natural classification for  2001 The Linnean Society of London the included species, are presented. ADDITIONAL KEY WORDS: viviparity – internal fertilization – Southeast Asia.

Hemirhamphodon, were placed in an unresolved polytomy with the monotypic genus, Tondanichthys (Fig. 1). Much of the uncertainty in generic relationships stems from the fact that sexually mature specimens of Tondanichthys are not available. Consequently, viviparity in Tondanichthys has been inferred, not observed. The purpose of this paper is to diagnose the genera of viviparous halfbeaks, propose species limits for Dermogenys and Nomorhamphus, describe seven new species, and present a phylogenetic hypothesis of the relationships among the species of Dermogenys and Nomorhamphus as well as the genera of internallyfertilized halfbeaks.

INTRODUCTION The family Hemiramphidae, or halfbeaks, comprises 13 genera and over 100 species of atherinomorph fishes distributed worldwide in both freshwater and marine habitats. Among the halfbeaks, three genera – Hemirhamphodon, Dermogenys and Nomorhamphus – are internally fertilized and viviparous. These genera have been proposed to belong to a more inclusive clade of halfbeaks, the Zenarchopterinae, including Zenarchopterus and Tondanichthys, also thought to be internally fertilized (Collette, 1995; Meisner & Collette, 1999). In a preliminary hypothesis of relationships (Collette, 1995), the three genera known to be viviparous, Dermogenys, Nomorhamphus and

TAXONOMIC HISTORY The genus Dermogenys Kuhl & van Hasselt in van Hasselt (1823) was first described in a letter from Java written by van Hasselt following the death of Kuhl. The genus was described for a new species, D. pusilla

∗ Current address: Department of Ichthyology, American Museum of Natural History, Central Park West @ 79th Street, New York, NY 10024, USA. E-mail: [email protected] 0024–4082/01/100199+85 $35.00/0

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A. D. MEISNER Hyporhamphus

Zenarchopterus

Tondanichthys

Hemirhamphodon

Dermogenys

Nomorhamphus

Figure 1. Hypothesis of relationships for the genera of internally-fertilized halfbeaks as proposed by Collette (1995).

(discovered during the lifetime of both Kuhl and van Hasselt). The validity of the genus remained unquestioned for nearly a century as nine new species were described. In the first revision of Dermogenys, Mohr (1936a) recognized four of the then ten nominal species, placing three in synonymy with D. pusilla, the type species of the genus. Since Mohr’s revision, seven additional species and three subspecies have been described (Table 1). In a subsequent revision, Brembach (1991) recognized ten species and three subspecies. In that revision, characters of the modified anal fin in males (andropodium) were used to distinguish species. However, also in that revision, the name D. pusilla was applied to populations that have consistent differences in the andropodium (Kottelat, 1992), and Mohr’s synonymy of several species was uncritically accepted. This has led to the widespread use of the name D. pusilla in museum collections and throughout the literature. The genus Nomorhamphus Weber & de Beaufort (1922) was originally described for two species, N. celebensis and N. hageni. No clear diagnostic characters were presented in the original description, but the only two species of viviparous southeast Asian halfbeaks lacking an elongate lower jaw were placed into the genus. No mention of the modified male anal fin was made in the original description of the genus. In a revision of Nomorhamphus, Mohr (1936b) recognized the two nominal species and also made no reference to any details of the modified anal fin (andropodium).

Since Mohr’s revision, five additional species and two subspecies have been described (Table 1). In a subsequent revision, Brembach (1991) recognized all nine nominal species and subspecies as valid. In that revision, differences in the morphology of the andropodium were noted but were not used to distinguish species. In a review of that revision, Kottelat (1992) found several diagnostic characters difficult to interpret and questioned the validity of some nominal species. In previous phylogenetic hypotheses, the sister group relationship between Dermogenys and Nomorhamphus was supported by the thickened first anal pterygiophore of males supporting the modified anal fin, the andropodium (Anderson & Collette, 1991; Collette, 1995). Furthermore, Nomorhamphus has been distinguished by lack of ‘true’ gill rakers on the first arch (Collette, 1995), a character interpreted here as a reduction of the gill rakers and also found to be more widely distributed among the viviparous halfbeaks (Meisner & Collette, 1999). Thus, there are no characters that distinguish Dermogenys from Nomorhamphus as currently constituted. Additionally, the taxonomy of these two genera has been confused further by the use of overlapping meristic and morphometric characters to diagnose species as well as the uncritical acceptance of the synonymy of several species. The taxonomy of both genera is confused and a well-supported hypothesis of the phylogenetic relationships for these species has never been proposed.

MATERIAL AND METHODS Institutional abbreviations follow Leviton et al. (1985) except CMK (Private Collections of Maurice Kottelat), MVT (uncatalogued collection of Tyson Roberts), and ZRC (Zoological Reference Collection of the Raffles Museum of Biodiversity Research, National University of Singapore). Terms used for describing the andropodium follow Brembach (1976) and Meisner & Burns (1997a). The term andropodium implies that it functions as an intromittant organ; the anal fin probably does not function directly in the transfer of sperm but may orient the elongate genital papilla toward the female gonopore during insemination (Meisner & Burns, 1997a). For convenience, however, the anterior five to seven anal-fin rays that become thickened and curved posteriorly are collectively referred to as the andropodium (Fig. 2). In some species, there is a distinct ‘knee’, the geniculus that is formed at the point of curvature along the second anal-fin ray. A pair of spines is present distally on this ray. The spiculus refers to the terminus of the second anal-fin ray distal to the paired spines. The cryptoplica, a fleshy covering that develops on both sides of the modified anal-fin rays, grows down and overlaps these modified rays

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Table 1. Nominal species, subspecies, and varieties of halfbeaks assigned to the genera Dermogenys and Nomorhamphus (in chronological order of description) and recognized equivalent species according to the results of this study. New combinations and species in bold

Dermogenys pusillus Kuhl & van Hasselt in van Hasselt, 1823 Hemiramphus fluviatilis Bleeker, 1850 Hemiramphus brachynotopterus Bleeker, 1853 Hemiramphus sumatranus Bleeker, 1853 Hemiramphus orientalis Weber, 1894 Dermogenys siamensis Fowler, 1934 Dermogenys burmanicus Mukerji, 1935 Dermogenys vogti Brembach, 1982 Dermogenys montanus Brembach, 1991 Dermogenys pusillus borneensis Brembach, 1991 Dermogenys pusillus borealis Brembach, 1991 Dermogenys bispina Meisner & Collette, 1998 Dermogenys bruneiensis Dermogenys robertsi Dermogenys palawanensis Dermogenys collettei

Dermogenys D. pusilla D. pusilla D. brachynotopterus D. sumatrana D. orientalis D. siamensis D. burmanica D. vogti D. orientalis D. sumatrana D. siamensis D. bispina D. bruneiensis sp. nov. D. robertsi sp. nov. D. palawanesis nov. D. collettei sp. nov.

Hemiramphus viviparus Peters, 1865 Hemirhamphus weberi Boulenger, 1897 Hemiramphus ebrardtii Popta, 1912 Hemiramphus hageni Popta, 1912 Nomorhamphus celebensis Weber & de Beaufort, 1922 Dermogenys nakeri Fowler & Bean, 1923 Dermogenys pectoralis Fowler, 1934 Dermogenys viviparus var. mindanensis Herre, 1944 Dermogenys philippinus Ladiges, 1972 Nomorhamphus towoetii 1972 Nomorhamphus brembachi Vogt, 1978 Nomorhamphus liemi liemi Vogt, 1978 Nomorhamphus liemi snijdersi Vogt, 1978 Dermogenys megarrhamphus Brembach, 1982 Nomorhamphus ravnaki ravnaki Brembach, 1991 Nomorhamphus ravnaki australis Brembach,1991 Nomorhamphus sanussii Brembach, 1991 Nomorhamphus kolonodalensis Meisner & Louie, 2000 Nomorhamphus manifesta Nomorhamphus rossi Nomorhamphus pinnimaculata

Nomorhamphus N. vivipara comb. nov. N. weberi comb. nov. N. ebrardtii comb. nov. N. hageni N. celebensis N. bakeri comb. nov. N. pectoralis comb. nov. N. vivipara comb. nov. N. philippina comb. nov. N. towoetii N. brembachi N. liemi N. liemi N. megarrhamphus comb. nov. N. brembachi N. brembachi N. brembachi N. kolonodalensis N. manifesta sp. nov. N. rossi sp. nov. N. pinnimaculata sp. nov.

ventrally. The physa is a membranous pouch that is situated between rays three and four and lies on one side of the fin, covering a small portion of the first and second anal-fin rays. To carry out a comprehensive phylogenetic analysis, specimens were cleared and stained according to the procedure of Dingerkus & Uhler (1977). Representatives of species not available for this type of preparation were X-rayed. Measurements were made with digital calipers to the nearest 0.1 mm and follow Anderson & Collette (1991). Counts were made from cleared and stained preparations and radiographs.

Precaudal vertebrae begin with the anterior most vertebra and end with the last vertebra without a hemal spine. Counts of caudal vertebrae begin with the first vertebra with a hemal spine and include the terminal half centrum. Following cladistic methodology outlined by Hennig (1966) and subsequently discussed and modified by others including Nelson & Platnick (1981), in which shared derived characters are used as evidence of phylogenetic relationship, a data matrix consisting of both reproductive and osteological characters was compiled and analysed using the branch and bound

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Geniculus

Spines

Spiculus

Andropodium

Figure 2. Diagrammatic representation of the modified anal-fin (andropodium) of Dermogenys and Nomorhamphus. Middle radials absent, ossified distal radials and cartilage not illustrated; bone is stippled. (Dermogenys bispina Meisner and Collette, USNM 345500, 28.5 mm SL, Sabah, Penang River). (Drawing by Keiko Hiratsuka Moore).

option in PAUP ver 3.1.1 (Swofford, 1993). In the absence of a priori knowledge of patterns or processes of evolution (e.g. gradualism), characters were treated as unordered and equally weighted. I believe that the most robust way of hypothesizing character state order results from a cladogram based on all available characters (see also Hauser & Presch, 1991). Characters were optimized using both ACCTRAN and DELTRAN options. Equally parsimonious optimizations are discussed in the text. Cladograms were rooted using the oviparous halfbeak genera, Hyporhamphus and Hemiramphus, as outgroups following Collette (1995). Characters are numbered in the phylogenetic analysis and species descriptions roughly according to their position along the anterior-posterior axis of the skeleton; reproductive characters are listed last. Counts for the holotype/lectotype are enclosed in square brackets. As many of the recognized species of Dermogenys and Nomorhamphus as possible were included in the phylogenetic analysis. However, for two species, N. bakeri and N. philippina, specimens were not available for cleared and stained preparations or histological study. For these species, many osteological and reproductive characters could not be scored for analysis. When analyses were run including these species, with unscored characters coded as ‘?’, a completely uninformative result was obtained. Therefore, these species were excluded from the phylogenetic analysis and their possible relationships are discussed in the text. (For index to genera and species, see Appendix 3.) MORPHOMETRIC ABBREVIATIONS

SL, standard length; LJL, lower jaw length; UJL, upper jaw length; UJW, upper jaw width; SN-P1, snout

to pectoral fin distance; SN-P2, snout to pelvic fin distance; P2-C, pelvic fin to caudal fin distance; HDL, head length; BDP1, body depth at pectoral fin base; BDP2, body depth at pelvic fin base; ORBL, bony orbital diameter; INTORBL, bony interorbital width.

OSTEOLOGICAL ABBREVIATIONS

APAL, autopalatine; CL, cleithrum; COR, coracoid; EB 1–4, epibranchial 1–4; END, endopterygoid; EP, epural; HY 1–5, hypural 1–5; HYO hyomandibula; MET, metapterygoid; PB 2–3, pharyngobranchial 2–3; PC3, postcleithrum 3; PHY, parhypural; POP, preopercle; PTT, posttemporal; QUA, quadrate; RAD, radials; SCA, scapula; SYM, symplectic; UN, uroneural.

CHARACTERS UPPER JAW

The upper jaws of halfbeaks examined in this study share characters proposed as synapomorphies for the Atherinomorpha (Rosen, 1964; Rosen & Parenti, 1981): rostral cartilage decoupled from the premaxilla; absence of crossed rostral ligaments; and a disc-like ethmoid. Dentition and development of the jaw bones was described for Dermogenys ‘pusilla’ (locality not stated) by Clemen, Wanninger & Greven (1997) and Greven, Wanninger & Clemen (1997). In Greven et al. (1997), the rostral cartilage is identified as the preethmoid. The upper jaw of all halfbeaks comprises two paired bones, the premaxilla and maxilla. The premaxillae form a flat, broad triangular plate. In Hyporhamphus, Hemiramphus, Tondanichthys and

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A

C

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B

A

B

C

D

E

F

D

Figure 3. Diagrammatic representation of the premaxillary plate and oral dentition in dorsal view; anterior to the top (characters 1, 2). A, Representation of the condition observed in Hyperhamphus, Hemiramphus, Tondanichthys, and Zenarchopterus. Hypohamphus unifasciatus USNM 196819 (73.0 mm). B, Dermogenys collettei CAS 139413, female, 40.0 mm. C, Nomorhamphus towoetii USNM 338352, female, 68.0 mm. D, N. liemi USNM 338351, male, 45.0 mm.

Zenarchopterus (Fig. 3A) the upper jaws are short and the shape of the premaxillary plate approximates an equilateral triangle. The oral teeth in these genera are arranged in two or three rows along the outer margins of the premaxillae and are either tricuspid or conical. The right and left premaxillary bones form a narrowbased, elongate triangular plate in Hemirhamphodon, Dermogenys, and the following Nomorhamphus species: N. hageni, N. megarrhamphus, N. weberi, N. towoetii, N. ebrardtii, N. bakeri, N. pectoralis, N. vivipara, N. philippina, N. rossi, N. manifesta, N. kolonodalensis and N. pinnimaculata (Fig. 3B,C). The upper jaw is much longer than wide. The oral dentition varies from one to four rows of conical teeth arranged in straight rows along the outer edge of the premaxillae (Fig. 3B,C). In Dermogenys (Fig. 3B), uniserial teeth extend medially in a concave row from the outer rows of teeth to a point at about one half the length of the premaxilla (Fig. 3B). This arrangement of oral teeth has been called a dentigerous bar by Clemen et al. (1997) and Greven et al. (1997). In N. liemi, N. brembachi and N. celebensis the upper jaw is short, roughly trapezoidal, and approximately as long as it is wide

Figure 4. Diagrammatic representation of the maxilla. Anterior to the left. A, Hyporhamphus unifasciatus USNM 196819 74.5 mm. B, Zenarchopterus rasori USNM 263456 female, 64.0 mm. C, Tondanichthys kottelati USNM 330491 male, 61.5 mm. D, Hemirhamphodon kuekenthali ROM 4209 female, 47.0 mm. E, Nomorhamphus liemi USNM 338351 male, 43.0 mm. F, Dermogenys orientalis USNM 338416 female, 54.0 mm.

(Fig. 3D). In N. liemi and N. brembachi (Fig. 3D) the anterior oral teeth are arranged into a large patch of five or six rows. The maxilla of Hemiramphus, Hyporhamphus, Zenarchopterus and Tondanichthys (Fig. 4A–C) is short with the distal half oriented dorsally. At a point approximately midway along its length, the proximal half of the bone is oriented slightly anteriorly so that the distal tip lies just ventral to the posterior aspect of the premaxillary plate. In Hemirhamphodon, Nomorhamphus and Dermogenys (Fig. 4D–F), the maxilla is more elongate and the distal half of the bone is angled slightly more posteriorly. LOWER JAW

Most beloniforms pass through a halfbeak stage at some point in development (Collette et al., 1984) (Fig. 5B). In needlefishes, this halfbeak stage is temporary and eventually growth of the upper jaw matches that of the elongate lower jaw (Boughten, Collette & McCune, 1991). In most halfbeaks, however, the disparity between the lengths of the upper and lower jaws is

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A

A

B

Figure 6. Dorsal view of the nasal fossa and nasal barbel. A, Hemirhamphodon kuekenthali. B, Zenarchopterus ornithocephala (reprinted with permission from Anderson and Collette, 1991).

B

Figure 5. Diagrammatic representation of the head showing variation in lower jaw length (after Brembach, 1991), previously used to distinguish Dermogenys from Nomorhamphus. A, reduction in length of lower jaw, a condition observed in some species of Nomorhamphus (character 5). B, ‘halfbeak’ condition.

retained throughout development (Fig. 5B). The condition found in needlefish, retention of elongate upper and lower jaws in the adult form, has been hypothesized as plesiomorphic for Beloniformes in general (Collette et al., 1984). Four species, N. towoetii, N. celebensis, N. brembachi and N. liemi, lack an elongate lower jaw and instead, have an upper and lower jaw approximately equal in length (Fig. 5A). In the remaining species of Dermogenys and Nomorhamphus, the lower jaw extends anteriorly past the length of the upper jaw (Fig. 5B). Coding of this character requires an hypothesis of homology of jaw length. The equal length of the jaws in needlefishes is formed by elongation of both upper and lower jaws. The condition found in these Nomorhamphus species differs from needlefishes in that neither the upper nor lower jaw is elongate. Therefore, the condition in Nomorhamphus is interpreted as a reduction in length of the lower jaw. This follows the transformation series proposed by Collette et al. (1984) in which loss of an elongate lower jaw is hypothesized as derived within the Exocoetoidei.

A

B

Figure 7. Diagrammatic representation of the left opercle. A, condition observed in Hemiramphus, Hyporhamphus, Zenarchopterus, and Tondanichthys. Hyporhamphus unifasciatus USNM 196819 74.5 nn. B, condition observed in Hemirhamphodon, Nomorhamphus, and Dermogenys. Dermogenys collettei FMNH 45867 female, 42.0 mm.

is short and spatulate. The nasal fossa in internally fertilized halfbeaks (Fig. 6), which are fresh and brackish water fishes, is much smaller and the nasal barbel is elongate, protruding out of the fossa. These characters have been proposed as synapomorphies for the internally-fertilized halfbeaks by Anderson & Collette (1991), Collette (1995), and Meisner & Collette (1999). OPERCULAR APPARATUS

NASAL REGION

In marine halfbeaks such as Hyporhamphus and Hemiramphus, the nasal fossa is large and the nasal barbel

The opercle of Hyporhamphus, Hemiramphus, Tondanichthys and Zenarchopterus is a large, flat, oval bone without spines or serrations (Fig. 7A). The opercle

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

A

B

C

D

E

F

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Figure 8. Diagrammatic representation of the left lacrimal. Anterior to the left. A, Hemiramphus brasiliensis USNM 294455 139.0 mm. B, Zenarchopterus buffonis USNM 173767 female, 110.0 mm. C, Tondanichthys kottelati USNM 330491 male, 61.5 mm. D, Hemirhamphodon kuekenthali USNM 330828 male, 50.0 mm. E, Dermogenys collettei FMNH 45867 female, 42.0 mm. F, Nomorhamphus megarrhamphus USNM 338501 female, 63.0 mm.

in these genera extends dorsally to contact the skull. The opercle in Hemirhamphodon, Dermogenys and Nomorhamphus (Fig. 7B) is much smaller, does not extend dorsally past the midpoint of the hyomandibula and lacks spines or serrations. Remaining bones in the opercular series are smooth and do not vary significantly among genera of halfbeaks examined. Therefore, the preopercle is illustrated only for Dermogenys (see Fig. 9). INFRAORBITAL SERIES

The infraorbital series consists of the dermosphenotic and lacrimal. This condition conforms to a general reduction in bones of the infraorbital series in atherinomorph fishes (Rosen, 1964; Rosen & Parenti, 1981). The dermosphenotic is a slender, curved bone extending ventrally near the midpoint of the hyomandibula. Size and shape of the dermosphenotic does not vary significantly among taxa examined in this study. In all genera, the lacrimal is a broad, flat bone. In Hemiramphus, Hyporhamphus, Zenarchopterus, Tondanichthys, Hemirhamphodon and Dermogenys (Fig. 8A–E), the lacrimal is round and approximately as long as it is wide. In Nomorhamphus (Fig. 8F), the lacrimal is more oval, being distinctly longer than it is wide.

JAW SUSPENSORIUM

The autopalatine in Hyporhamphus (Fig. 9B) and Hemiramphus is a slender bone with the head offset dorsally. It is elongate and extends ventrally past the midpoint of the slender endopterygoid. In Zenarchopterus (Fig. 9C), Tondanichthys and Hemirhamphodon, the autopalatine is slender and much shorter, not extending to the midpoint of the thin endopterygoid. The head of the autopalatine is reduced and the dorsal portion of the bone is offset anteriorly. The autopalatine in both Dermogenys (Fig. 9A) and Nomorhamphus (Fig. 9D) is expanded, when compared to the above genera, with an anterior flange along its entire length. In Nomorhamphus, the autopalatine is elongate and extends past the midpoint of the endopterygoid. In Dermogenys, the autopalatine is shorter with the distal half angled anteriorly. In Dermogenys and Nomorhamphus, the endopterygoid is thicker than that of other genera examined.

DORSAL GILL ARCHES

Pharyngobranchial one is absent in all halfbeaks examined, a character interpreted as plesiomorphic for Beloniformes and Cyprinodontiformes (Rosen, 1964; Rosen & Parenti, 1981). Pharyngobranchials two and

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APAL APAL QUA

HYO

MET

MET

END

QUA

SYM

HYO

END

POP

SYM A

B

APAL APAL HYO MET END

QUA

MET

HYO

QUA END

SYM C

D

SYM

Figure 9. Diagrammatic representation of the left side of the jaw suspensorium. All cartilage not illustrated; small stippling bone; large stippling cartilage. A, Dermogenys collettei, CAS 139413 female, 40.5 mm. B, Hyporhamphus unifasciatus, USNM 196819, 70.0 mm. C, Zenarchopterus rasori, USNM 263456 female, 64.0 mm. D, Nomorhamphus liemi, USNM 338351 male, 46.0 mm. Abbreviations listed on p. 202.

three are tooth-bearing and ossified. The right and left third pharyngobranchials are fused into a large plate. The development of the branchial skeleton is described by Clemen et al. (1997). An illustration of the gill arches is provided by Greven et al. (1997) for ‘D. pusilla’ (locality not specified) where the fused pharyngobranchials three and four are labeled as the dorsal tooth plate, pharyngobranchial two as the lateral tooth plate, and the fused ceratobranchial five as the ventral tooth plate. The fused pharyngobranchials in Hemiramphus and Hyporhamphus (Fig. 10A) form a plate that is round posteriorly and narrow along the anteriorposterior axis. The anterior processes of this gill arch element are in line with this axis. In internally fertilized halfbeaks, pharyngobranchial three is shorter along the anterior-posterior axis and expanded posteriorly. In Tondanichthys (Fig. 10B) and Zenarchopterus, anterior processes are in line with the anterior-posterior axis but in Hemirhamphodon, Dermogenys (Fig. 10C) and Nomorhamphus (Fig. 10D) they are offset laterally. Four epibranchials are present in all halfbeaks examined, the second and third epibranchials being smaller than the first and fourth. Reduction in size of

second and third epibranchials has been proposed as a synapomorphy for Beloniformes and Cyprinodontiformes (Rosen & Parenti, 1981). In Hemiramphus, Hyporhamphus, Tondanichthys and Zenarchopterus, the first epibranchial is the largest and extends anteriorly, lying ventral to the anterior extent of pharyngobranchial two (Fig. 10A,B). The base of epibranchial one is expanded only slightly relative to the width of the bone. In Hemirhamphodon, Dermogenys and Nomorhamphus, epibranchial one is short, approximating the length of epibranchial four (Fig. 10C,D). The base of epibranchial one in these genera is expanded greatly relative to the dorsal aspect. Epibranchial four in Hemiramphus, Hyporhamphus, Tondanichthys and Zenarchopterus is a short, straight bone with a short ramus on the posterior surface near the point of articulation with pharyngobranchial three (Fig. 10A,B). The articular surface of epibranchial four is expanded. In Hemirhamphodon, Dermogenys and Nomorhamphus the posterior ramus is absent. Epibranchial four is slender and either has the proximal and distal articular surfaces reduced or slightly expanded as in D. pusilla, D. burmanica, D. siamensis and D. collettei (Fig. 10B,C).

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

EB1

PB2

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EB1 PB2

EB2

PB3 EB2

EB3

PB3

EB4

EB3 EB4

A

B

EB1 PB2

EB1 PB2

EB2 EB2

PB3

PB3

EB3

EB3 EB4

EB4

C

D

Figure 10. Diagrammatic representation of the dorsal portion of the gill arches. Anterior towards the top. A, Hyporhamphus unifasciatus, USNM 196819, 74.5 mm. B, Tondanichthys kottelati, ZSM/CMK 7980 male, 61.5 mm. C, Dermogenys burmanica, CAS 134945 female, 38.1 mm. D, Nomorhamphus weberi, USNM 338506, female, 64.0 mm. Small stippling bone; large stippling cartilage. Abbreviations listed on p. 202.

GILL RAKERS

Gill rakers have been illustrated for several species of Dermogenys and Nomorhamphus (Brembach, 1991). Gill rakers on the first arch in Hemiramphus (Fig. 11A), Hyporhamphus, Zenarchopterus and Tondanichthys are elongate with many conical teeth along the length of lateral surfaces. In Hemirhamphodon, Dermogenys (Fig. 11B) and Nomorhamphus (Fig. 11C, D), gill rakers are short and lie within the connective tissue above the gill arch elements. In Hemirhamphodon, Dermogenys, N. pectoralis, N. rossi, N. manifesta, N. pinnimaculata and N. kolonodalensis, the gill rakers are tear-drop shaped and generally toothless (Fig. 11B); however, some gill rakers on the second and third arches have one to three teeth on the dorsal surface. In N. megarrhamphus, N. weberi, N. ebrardtii, N. towoetii and N. celebensis gill rakers are more or

less tear-drop shaped with many conical teeth on the dorsal surface (Fig. 11C). In N. liemi and N. brembachi, gill rakers have a wide base and many teeth on the dorsal surface (Fig. 11D).

CAUDAL FIN

The caudal skeleton of all halfbeak taxa examined comprises the terminal half centrum fused to the hypural plate consisting of hypurals 1–5, a parhypural, uroneural, and three epurals. In Hemiramphus, Hyporhamphus (Fig. 12A), Zenarchopterus (Fig. 12B) and Tondanichthys (Fig. 12C) the last two or three hemal spines are enlarged. In these genera, relatively large bony plates are present along the neural arch, being most prominent in the oviparous genera Hemiramphus

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Figure 11. Photographs of gill rakers (arrowed). A, Hemiramphus brasiliensis (USNM 294455, 91.0 mm); B, Dermogenys sumatrana (USNM 306569, male, 31.0 mm); C, Nomorhamphus towoetii (USNM 388352, female, 60.0 mm). D, Nomorhamphus liemi (USNM 338351, male, 49.0).

and Hyporhamphus (Fig. 12A). The epurals and uroneural are large bones, the latter fused to the dorsal hypural plate along most of its length. In all four genera, the parhypural is autogenous (Fig. 12A–C). In Hyporhamphus and Hemiramphus (Fig. 12A) the fused hypurals 1–2 are widely separated from fused hypurals 3–4. Hypural 5 is separate from the dorsal hypural plate along most of its length. In Zenarchopterus and Tondanichthys (Fig. 12B,C), the ventral hypural plate consists of fused hypurals 1–2 and the dorsal hypural plate consists of hypurals 3–5 that are separate along most of their lengths. In the viviparous genera, Hemirhamphodon, Dermogenys and Nomorhamphus, the bony mass of the caudal skeleton appears considerably reduced when compared to above genera (Fig. 12D–F). The neural plates are not expanded and the last hemal spine is only slightly expanded. Supernumerary hemal and neural spines are not unusual (as illustrated for Dermogenys Fig. 12E). In viviparous genera, the epurals are slender bones and the uroneural is inconspicuous and fused to the dorsal hypural plate. The parhypural is completely fused to the ventral hypural plate. The ventral hypural plate consists of fused hypurals 1–2 and the dorsal hypural plate of fused hypurals 3–4. Hypural 5 is separate along most of its length. In N. weberi, N. megarrhamphus, N. ebrardtii (Fig. 12F), N. towoetii, N. celebensis, N. brembachi and N. liemi, hypural 5 is completely fused to the dorsal plate.

DORSAL FIN

The origin of the dorsal fin in Hemiramphus, Hyporhamphus, Tondanichthys, Zenarchopterus and Hemirhamphodon is anterior to that of the anal fin. The first dorsal pterygiophore has a ventral process that extends near the neural spines (Fig. 13A–D). The dorsoanterior extension of this bone is slight. In Dermogenys and Nomorhamphus, origin of the dorsal fin is posterior to that of the anal fin. The first dorsal pterygiophore is a slender bone that extends anteriorly along the dorsal surface with a ventral process that is reduced or absent (Fig. 13E).

PECTORAL GIRDLE

In Hemiramphus and Hyporhamphus (Fig. 14A) no ramus is present on the post-temporal or it is present only as a nubbin. Orientation of the pectoral-fin rays relative to the posttemporal differs between oviparous halfbeaks and the Zenarchopterinae. In the oviparous genera, the cleithrum is oriented more vertically so that the pectoral radials are high-set (Fig. 14A). In Tondanichthys (Fig. 14B), Zenarchopterus, Hemirhamphodon (Fig. 14C), Dermogenys (Fig. 14D) and Nomorhamphus there is a ramus on the post-temporal near the point of articulation with the cleithrum. In the internally fertilized genera, the cleithrum is angled more posteriorly so that the radials are set lower on the body (Fig. 14B–D).

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

209

EP HY 5 UN

EP HY 5

HY 3 + 4

HY 3 + 4

HY 1 + 2

HY 1 + 2

PHY D

A

EP

EP

HY 5

HY 5 HY 4 HY 3

HY 3 + 4

HY 1 + 2 HY 1 + 2

PHY E

B EP HY 5 HY 4

EP HY 3 + 4 + 5

HY 3 HY 1 + 2 HY 1 + 2

PHY

C

F

Figure 12. Diagrammatic representation of the caudal skeleton. A, Hyporhamphus unifasciatus USNM 196819 74.5 mm. B, Zenarchopterus rasori USNM 263456 female, 64.0 mm. C. Tondanichthys kottelati ZSM/CMK 7980 male, 61.5 mm. D, Hemirhamphodon kuekenthali USNM 330828 male, 50.0 mm. E, Nomorhamphus viviparus CAS 137829 male, 34.0 mm. F, Nomorhamphus ebrardtii ZMH 7150 male, 50.0 mm. Cartilage not illustrated; bone is stippled. Abbreviations listed on p. 202.

PELVIC GIRDLE

The right and left pelvic bones in Hemiramphus, Hyporhamphus, Zenarchopterus and Tondanichthys are not separated as widely as in Dermogenys, Nomorhamphus and Hemirhamphodon (Fig. 15). The anterior tips are angled toward the midline so that they are in close proximity. In lateral view, the posterodorsal portion extends as a thin process. In ventral view, the ventromedial flange is bifurcate anteriorly (Fig. 15A). In Hemirhamphodon, Dermogenys and Nomorhamphus, the pelvic bones are also widely separated; however, there is a lateral reorientation of these bones so that the anterior tips are no longer in close proximity but lie parallel to the axis of the body (Fig. 15B,C).

The lateral view is identical to that described above except that in Dermogenys and Nomorhamphus an anterior spur is present (Fig. 15C). The ventromedial flange in Hemirhamphodon, Dermogenys and Nomorhamphus is entire (Fig. 15B,C).

MALE ANAL FIN (ANDROPODIUM)

Details of anatomy and development of the andropodium in Dermogenys and comparisons to other internally fertilized atherinomorphs were described by Meisner & Burns (1997a) and will only be summarized here. Details of variation at the species level are included in the systematic accounts with figures. The andropodium in

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A. D. MEISNER

A

B

D

C

E

Figure 13. Diagrammatic representation of the first dorsal pterygiophore. A, Hyporhamphus unifasciatus USNM 196819, 75.0 mm; B, Tondanichthys kottelati ZSM/CMK 7896, male, 61.5 mm; C, Zenarchopterus rasori USNM 263456, female 62.0; D, Hemirhamphodon kuekenthali ROM 4209, female, 48.0; E, Nomorhamphus megarrhamphus, USNM 338351, female, 63.0 mm. Bone stippled; anterior to the right.

Nomorhamphus is identical in overall morphology to that of Dermogenys. In all species, the first anal pterygiophore is thickened, presumably to support the musculature of the modified anal fin. However, in two species, N. weberi and N. megarrhamphus, the first anal pterygiophore is thinner and angled anteriorly at a sharper angle. In Zenarchopterus, ray five or rays five and eight are modified (Collette, 1982, 1985). In Hemirhamphodon, however, anal-fin rays five and eight are elongate and thickened (Anderson & Collette, 1991). Morphology of the andropodium in Tondanichthys is not known with certainty because mature specimens are not available in museum collections; nonetheless, from specimens available, it appears that rays six and seven are enlarged (Collette, 1995). Despite this uncertainty, it is clear from immature specimens that anal fin modification in Tondanichthys is different from that in Dermogenys and Nomorhamphus. A pair of spines is present distally on the second anal-fin ray in Dermogenys and Nomorhamphus. The tissue type of these spines is unknown, however. In all cleared and stained preparations, these spines remain unstained except in some species of Dermogenys, D. palawanensis, D. bispina, D. robertsi, a small section at the tip of the spines stains red with alizarin red. These findings suggest that the spines are not ossified but may be keratinized structures. INSEMINATION

Most Beloniformes, including Hemiramphus and Hyporhamphus, are oviparous (Collette et al., 1984), sperm

and eggs are broadcast into the water and fertilization and larval development are external. Three genera are known to be internally fertilized and viviparous, Dermogenys, Nomorhamphus and Hemirhamphodon (Roberts, 1993; Meisner & Burns, 1997b). For these genera, coding of characters associated with internal fertilization is unambiguous. Zenarchopterus, on the other hand, has been proposed to be internally fertilized and to lay fertilized eggs. Sperm has been observed within the ovary of at least one female examined indicating that these fishes are inseminated (pers. obs.), but whether fertilization actually takes place in the ovary requires further study. However, this information suggests that insemination does occur, and therefore this character is coded as present for Zenarchopterus. Due to the lack of mature specimens of Tondanichthys, all characters related to reproduction are coded as missing in this genus. SPERMATOGENESIS AND SPERM BUNDLE MORPHOLOGY

It has been suggested that in the formation of sperm bundles, an association is formed between spermatid nuclei and Sertoli cell cytoplasm within spermatocysts during spermiogenesis (Grier, Fitzsimmons & Linton, 1978). Details of spermatogenesis in Dermogenys, Nomorhamphus and Hemirhamphodon were described by Downing & Burns (1995) and for Zenarchopterus by Grier & Collette (1987). Hemiramphus and Hyporhamphus are oviparous, sperm packets are not formed, and there is no regular organization within

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

PTT

211

PTT SCA SCA RAD

CL

RAD

CL

PC3

PC3 COR

COR

A

B PTT PTT SCA SCA RAD

CL

CL

RAD

PC3 PC3

COR C

D

COR

Figure 14. Diagrammatic representation of the left pectoral girdle. A, Hyporhamphus unifasciatus USNM 196819, 74.5 mm; B, Tondanichthys kottelati USNM 330491, male, 61.5 mm. C, Hemirhamphodon kuekenthali USNM 330828, male, 50.0 mm; D, Dermogenys collettei FMNH 45867, female, 42.0. Small stippling bone; large stippling cartilage. Abbreviations listed on p. 202.

the spermatocysts (pers. obs.). In Hemirhamphodon and Zenarchopterus, spermatid nuclei become arranged unevenly around the periphery of the spermatocyst. In Dermogenys and Nomorhamphus spermatid nuclei become arranged evenly around the periphery of the spermatocyst in one or several layers (Downing & Burns, 1995). Again, because mature males of Tondanichthys are not available, characters associated with spermatogenesis are coded as missing. VIVIPARITY

As mentioned previously, Hemiramphus and Hyporhamphus are oviparous. Details of viviparity in Dermogenys and Nomorhamphus have been described elsewhere (Meisner & Burns, 1997b) and will only be summarized here. Viviparity in these genera has been classified into five types that appear to be unique among halfbeaks. In types I and II, development throughout most of gestation is intrafollicular. Here, ovulation coincides with parturition, meaning that embryos pass only briefly through the ovarian

lumen. An apparently intermediate form is found in two species, D. robertsi and D. palawanensis. In these species, morphological modifications are identical to other species with the type II form but a larger yolk reserve is present in early embryos. However, because external influences on yolking of eggs and variations in yolk utilization by early embryos have not been studied, this condition is combined with other species with type II viviparity based on similarity in overall modifications. In types III–V, embryos are evacuated into the ovarian lumen at an earlier stage of development and consequently, there is a long period of intraluminal development. The ovaries of specimens of N. pinnimaculata examined contained early cleavage embryos or yolky eggs only, leading to some ambiguity in histological interpretation. However, because yolky oocytes appeared to be contained within a discrete ovigerous ridge the condition in this species is included with type IV variants. In species with type III viviparity (N. pectoralis and N. rossi, N. manifesta) there was some variation in the extent of belly sac (expanded pericardial sac and coelomic

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(Bork & Mayland, 1998). It is difficult to determine the length of luminal gestation in Hemirhamphodon but viviparity appears to be different from that observed in Dermogenys and Nomorhamphus. Because no developing embryos have been observed within the ovaries, viviparity is coded as absent in Zenarchopterus and as missing information in Tondanichthys because no mature females are available. A

PHYLOGENETIC ANALYSIS

B

Twenty-eight taxa comprising ten species of Dermogenys, 13 species of Nomorhamphus, and five outgroup taxa were analysed. When a data matrix comprising 45 characters (coded and described in the Appendix) was analysed using the branch and bound option in PAUP version 3.1 (Swofford, 1993), 36 equally parsimonious trees were found with a length of 57, CI= 0.995, and RI=0.965. A strict consensus of these trees is presented in Figure 16. Defining characters of each species are listed in the systematic accounts. Node A: Zenarchopterinae Fowler, 1934a (Zenarchopterus+Tondanichthys+ Hemirhamphodon+Nomorhamphus+Dermogenys)

C

Figure 15. Ventral view of the right and left pelvic bones. A, representation of the condition observed in Hyporhamphus, Hemiramphus, Tondanichthys, and Zenarchopterus (Hyporhamphus unifasciatus USNM 196819, 74.5 mm); B, Hemirhamphodon kuekenthali USNM 330828, male, 50.0 mm; C, representation of the condition observed in Dermogenys and Nomorhamphus (Nomorhamphus kolonodalensis USNM 338490, female, 44.0 mm). Bone is stippled; anterior towards the top.

Monophyly of the group comprising the genera proposed to be internally fertilized (see Meisner and Collette, 1999) is supported by three characters: (6) nasal fossa small; (7) nasal papilla elongate, extending out of the nasal fossa; and (24) caudal fin round or emarginate. An additional character, (36) insemination, may also support this group (see also Meisner & Collette, 1999). The informativeness of this character is in question because insemination has been observed in Zenarchopterus but not in Tondanichthys. For Tondanichthys, insemination is inferred from the presence of a modified anal fin in males. Node B: Hemirhamphodon+(Nomorhamphus+Dermogenys)

cavity) hypertrophy. Yet, based on similarity in overall morphology the condition observed in these species is coded as identical. A somewhat unusual form was observed in N. ebrardtii (type V), details are provided by Meisner & Burns (1997b). Detailed information on viviparity in Hemirhamphodon is scarce. Some species appear to have a unique form of viviparity in which a significant yolk reserve is present as well as a unique form of superfetation (Roberts, 1989; pers. obs.). In this form, earliest-stage embryos are found anteriorly in the ovary with successively later stages posteriorly. Usually, there is only one embryo per developmental stage. It has also been reported from aquarium observations that H. tengah may lay fertilized eggs

The sister group relationship of Hemirhamphodon and (Dermogenys + Nomorhamphus) is supported by 11 characters: (1, state 1) right and left bones of premaxilla form a narrow-based triangular plate; (8) epibranchial one reduced to approximately length of epibranchial four, base expanded; (9) ramus absent on epibranchial four; (10, state 2) proximal and distal surfaces of epibranchial four reduced, each oriented posteriorly; (11) anterior process of pharyngobranchial three offset laterally; (13, state 1) gill rakers reduced, roughly tear-drop shaped, with one to three teeth; (15) opercle reduced, not extending dorsally past midpoint of hyomandibula; (20) pelvic bones widely separated, lateral reorientation of anterior processes to lie along axis of the body; (21) ventral medial flange of pelvic

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

213

Hyporhamphus Hemiramphus Zenarchopterus Tondanichthys Hemirhamphodon D. pusilla D. burmanica

A

E

D. siamensis D. collettei D. robertsi

D

D. palawanensis

B

D. orientalis F D. sumatrana D. bruneiensis D. bispina N. liemi K J C

N. brembachi N. celebensis

I N. towoettii N. ebrardtii

H

N. megarrhamphus L

N. weberi N. vivipara N. kolonodalensis N. pinnamaculata

G

N. manifesta M

N. rossi N. pectoralis

Figure 16. Strict consensus of 36 equally most parsimonious trees. Undescribed species indicated by locality. Length= 57; CI=0.995; RI=0.965.

bones entire; (25) reduction in bony elements of caudal skeleton; (26) parhypural fused to ventral hypural plate.

first anal pterygiophore in males thick; and (33, state 2) mode of spermatogenesis in which spermatid nuclei are oriented evenly around periphery of spermatocyst.

Node C: Dermogenys+Nomorhamphus

Node D: Dermogenys

Seven characters support monophyly of Dermogenys+Nomorhamphus: (16) endopterygoid expanded; (22) anterior lateral extension of pelvic girdle; (23) ventral process of first dorsal pterygiophore reduced or absent, dorsoanterior extension expanded; (28) origin of anal fin anterior to dorsal-fin origin; (29) analfin rays 1–7 modified into andropodium; (30, state 1)

Monophyly of Dermogenys s.s. is supported unambiguously by three characters: (2) uniserial conical teeth that extend medially in a concave row from outer row of teeth to a point at about one half the length of premaxilla; (38) melanophores anterior to anal fin arranged into a distinct spot in females; and (31) welldefined geniculus.

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A. D. MEISNER

Three additional characters are potential synapomorphies of Dermogenys s.s. depending on the optimization scheme. The first is (17, state 1) shortened, expanded autopalatine. When optimized using the DELTRAN option, this multistate character can be placed as a synapomorphy for this genus. However, an equally parsimonious optimization proposes it as a synapomorphy for Dermogenys+Nomorhamphus with a subsequent change to state 2 in Nomorhamphus. For the second character, (34, state 1) large spermatozeugmata, optimization follows that of the previous character. The third is intrafollicular gestation (39). When optimized with DELTRAN, this character is a synapomorphy for Dermogenys. However, an equally parsimonious optimization of this multistate character using ACCTRAN has state 1 as a synapomorphy for the viviparous halfbeaks with two subsequent changes one to state 3 in Hemirhamphodon and another to state 2 in Nomorhamphus. Much of this ambiguity results from a lack of detailed descriptions of viviparity in Hemirhamphodon and from the ambiguous polarity of these characters. Remarks. Dermogenys pusilla, the type species of Dermogenys, is within this clade. The name Dermogenys is therefore restricted to the 12 species that are members of this clade: D. pusilla, D. burmanica, D. siamensis, D. collettei, D. orientalis, D. sumatrana, D. bispina, D. palawanensis, D. robertsi, D. bruneiensis, D. brachynotopterus and D. vogti (see Systematic Accounts). Node E: Dermogenys pusilla – species group This clade is supported by two characters: (10, state 1) epibranchial four curved slightly at midlength with proximal articular surface slightly expanded; (40) type I form of viviparity.

Remarks. This clade comprises D. pusilla, D. siamensis, D. burmanica and D. collettei. Available data cannot resolve species relationships within this clade. Node F: Dermogenys orientalis – species group This clade is supported by two characters: (35) threadlike sperm nuclei; and (41) type II form of viviparity. Remarks. This clade comprises D. orientalis, D. sumatrana, D. bispina, D. palawanensis, D. bruneiensis and D. robertsi. Resolution of species relationships within this clade is affected by two homoplastic characters: (1) cap of bone on spines of second anal-fin ray in males (32), present in all species except D. orientalis and D. sumatrana; and (2) an elongate, pigmented genital papilla in females (37), absent in D. robertsi. Node G: Nomorhamphus Monophyly of Nomorhamphus is supported by five characters: (14) oval lacrimal; (17, state 2) elongate, expanded autopalatine; (34, state 2) small spermatozeugmata; (39) long period of intraluminal gestation; and (45) well-developed ovigerous ridge. Remarks. This study supports retaining the name Nomorhamphus for this clade because it contains N. celebensis, the type species of Nomorhamphus. These findings also support expanding generic limits to include seven species previously classified as Dermogenys, N. bakeri (see Systematic Accounts), N. ebrardtii, N. megarrhamphus, N. weberi, N. vivipara, N. pectoralis, N. philippina (see Systematic Accounts) as well as N. celebensis, N. towoetii, N. liemi, N. brembachi, N. hageni, N. kolonodalensis and N. rossi, N. manifesta and N. pinnimaculata.

KEY TO ZENARCHOPTERINAE (FOWLER, 1934a)

Diagnosis. Anal fin in males modified; nasal fossa small; nasal papilla elongate, protruding out of nasal fossa (Fig. 6); caudal fin round or emarginate. 1A. 1B. 2A. 2B. 3A.

3B.

Teeth along the length of the elongate lower jaw ..................................................... Hemirhamphodon Teeth along the length of the elongate lower jaw absent ................................................................... 2 Modified anal-fin rays in males without a fleshy covering (cryptoplica) ................... Zenarchopterus Modified anal-fin rays in males with a fleshy covering (cryptoplica) ................................................. 3 Second anal-fin ray in males without a distinct geniculus; uniserial teeth not extending medially in a concave row from outer row of teeth; melanophores anterior to the anal fin in females not forming a distinct spot ................................................................................................................ Nomorhamphus Second anal-fin ray in males with a distinct geniculus; uniserial conical teeth extend medially in a concave row from outer row of teeth to a point at about one half the length of the premaxilla (Fig. 3B); melanophores anterior to the anal fin in females forming a distinct spot ............ Dermogenys

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

Node H Monophyly of this clade is supported by two characters: (13, state 3) reduced gill rakers, tear-drop shaped with many teeth on dorsal surface; and (27) fifth hypural completely fused to dorsal hypural plate. Remarks. This clade comprises N. brembachi, N. liemi, N. celebensis, N. towoetii, N. ebrardtii, N. megarrhamphus and N. weberi.

215

upper jaw extending medially in a concave row from outer rows of teeth to a point at about one half the length of premaxilla (dentigerous bar of Greven et al., 1997; Clemen et al., 1997); expanded endopterygoid; shortened, expanded autopalatine; distinct geniculus on second anal-fin ray in males beginning at segment three or four; gestation entirely intrafollicular; large spermatozeugmata resulting from mode of spermatogenesis in which spermatid nuclei become arranged in one layer evenly around periphery of spermatocysts.

Node I Monophyly of this group is supported by one character: (5) reduction in length of lower jaw so that upper and lower jaws are approximately equal in length. Remarks. This clade comprises N. brembachi, N. liemi, N. celebensis and N. towoetii. Node J This clade is supported by one character: right and left bones of premaxilla form a broad trapezoidal plate that is about as long as wide (1, state 2). Remarks. This clade comprises N. brembachi, N. liemi and N. celebensis. Node K The sister group relationship between N. liemi and N. brembachi is supported by two characters: (3) oral teeth arranged into five or six rows anteriorly on upper jaw; and (13, state 2) reduced gill rakers, wide base, many teeth. Node L The sister-group relationship of N. megarrhamphus and N. weberi is supported by one character: (30, state 2) first anal pterygiophore in males thin, angled sharply anteriorly. Node M This group comprising N. pectoralis, N. manifesta and N. rossi is supported by one character: (42) type III viviparity.

SYSTEMATIC ACCOUNTS GENUS DERMOGENYS KUHL & VAN HASSELT IN HASSELT (1823) (Figures 17, 18; Table 1)

Dermogenys Kuhl & van Hasselt in van Hasselt, 1823: 131 (type species: Dermogenys pusilla Kuhl & van Hasselt in van Hasselt, 1823, by monotypy). Diagnosis. Dermogenys is distinguished from other genera in the Hemiramphidae by a row of teeth on

Description. Sexually dimorphic, maximum known standard lengths of species ranges from 17.3–66.0 mm for females and 15.0–36.7 mm for males; body elongate; lower jaw elongate (4.5–16.3 times in SL); biserial conical teeth on upper and lower jaws; no teeth along extended portion of lower jaw; oral teeth conical; autopalatine expanded; gill rakers on first arch tear-drop shaped, with 1–3 teeth; first epibranchial bone expanded at base; third pharyngobranchial bones fused medially; lower pharyngeal tooth plates fused; branchiostegal rays 9–13; upper jaw longer than wide (UJL/ UJW 1.1–1.9); nasal papilla elongate, extending out of nasal fossa; nasal fossa small; infraorbital series consisting of lacrimal and dermosphenotic; lacrimal round; high-set pectoral fins; pectoral-fin rays 9–12; pectoral fins pointed, not reaching the base of the pelvic fins; first pleural rib on third vertebra; pelvic bones widely separated with anterior processes oriented laterally; anterior spur on lateral extension of pelvic bones; pelvic-fin rays 6; anal-fin rays 1–7 in males modified into an andropodium; cryptoplica present; physa present; distinct geniculus on second analfin ray in males; genital papilla elongate in mature males, nearly reaching first anal-fin ray; anal-fin rays 14–17 (Table 5); first anal pterygiophore in males thickened; middle anal radials absent; distal anal radials ossified; origin of anal fin anterior to that of dorsal-fin origin; ventral process on first dorsal pterygiophore reduced or absent, anterior extension expanded; dorsal-fin rays 8–11 (Table 6); caudal fin oval or truncate, without distinct lobes; three epurals; parhypural fused to lower hypural plate; fifth hypural separate from the dorsal hypural plate along most of its length; uroneural and posterior neural plates reduced; principal caudal fin rays 7+8; precaudal vertebrae 21–27 (Table 2); caudal vertebrae 15–20 (Table 4); total vertebrae 38–44; viviparous; gestation entirely intrafollicular (Meisner & Burns, 1997b); large spermatozeugmata formed (Downing & Burns, 1995). Distribution. Dermogenys is broadly distributed throughout fresh and brackish waters of Southeast Asia (Figs 17, 18). At least two species are restricted to mainland Asia; D. burmanica lives in Burma [Myanmar] and Bangladesh and D. siamensis throughout

216

A. D. MEISNER China Bangladesh N Myanmar Philippines 15 Indo-China South China Sea Andaman Sea

Palawan Sulu Sea

Malay Peninsula

5

Borneo

Celebes Sea

Sulawesi Sumatra 5

Java

95

105

115

125

Figure 17. Selected localities illustrating the distribution of species in the Dermogenys pusilla-group. (Ε) D. siamensis; (Ο) D. burmanica; (Η) D. pusilla; (_) Dermogenys collettei.

Vietnam, Cambodia and Thailand. Dermogenys brachynotopterus, if a valid species, is found in India. A more widespread species, D. collettei occurs in the southern portion of the Malay Peninsula, Sumatra, and the northwestern portion of Borneo (Sarawak, Brunei and northwestern Kalimantan). Dermogenys sumatrana has been collected on both eastern and western portions of Kalimantan (southern Borneo) as well as western Sumatra. Dermogenys bispina occurs in Sabah (northeastern Borneo) and on the western Philippine island of Busuanga. Dermogenys pusilla is restricted to Java. Dermogenys bruneiensis is endemic to Brunei in northwestern Borneo. D. robertsi is found on the western Philippine islands of Culion and Busuanga as well as portions of northeastern Kalimantan (southern Borneo). Dermogenys palawanensis is found only on the southwestern Philippine island of Palawan. Dermogenys orientalis is endemic to the southwestern arm of Sulawesi (Sulawesi Selatan). Remarks. Authorship of Dermogenys and Dermogenys pusilla has variously been attributed to Kuhl & van

Hasselt, 1823, van Hasselt in Kuhl & van Hasselt, 1823 (by Roberts, 1989), Kuhl & van Hasselt in van Hasselt, 1823 (by Kottelat, 1987), and, most commonly, van Hasselt, 1823 (by e.g. Weber & de Beaufort, 1922). Much of the confusion stems from the fact that both the genus Dermogenys and the species D. pusilla were described in letters by van Hasselt after the death of Kuhl. In a translation of these letters (Alfred 1961: 85), it is stated clearly that “. . . a Hemiramphus which is found in the brooks around Buitenzorg and nearly all over Java has already during the lifetime of Kuhl been separated by us from that genus under the name Dermogenys . . . the species bears the name Pusillus . . . and has been figured by us”. In a more recent review of Kuhl & van Hasselt material, Roberts (1993) also concluded that the species was discovered and described while both were still living. Therefore, authorship of both the genus Dermogenys and the species D. pusillus is attributed to Kuhl & van Hasselt in van Hasselt, 1823 (as in Kottelat, 1987). In addition, Dermogenys is feminine and, therefore, the species name has been emended to D. pusilla (Kottelat, 1992).

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

217

China Bangladesh N Myanmar Philippines 15 Indo-China South China Sea Andaman Sea

Palawan Sulu Sea

Malay Peninsula

5

Borneo

Celebes Sea

Sulawesi Sumatra 5

Java

95

105

115

125

Figure 18. Selected localities illustrating the distribution of species in the Dermogenys orientalis-group. (Ε) Dermogenys sumatrana; (Φ) D. bruneiensis; (Η) D. bispina; (_) D. robertsi; (Ο) D. palawanensis; (Χ) D. orientalis.

Table 2. Frequency distribution of precaudal vertebrae in species of Dermogenys

D. D. D. D. D. D. D. D. D. D. D. D.

pusilla brachynotopterus sumatrana orientalis siamensis burmanica vogti bispina bruneiensis robertsi palawanensis collettei

21

22

23

24

25

26

27

Mean

N

— — — 1 — — 1 1 — — 1 —

2 — 1 7 — — — 11 — 21 3 6

19 — 2 8 3 1 2 5 2 6 14 14

3 — 4 10 16

— — — — — 2 — — 4 — — —

— 1 — — — 9 — — — — — —

— — — — — 6 — — — — — —

23.0 26.0 23.4 23.0 23.8 26.2 22.7 22.2 24.3 22.2 22.7 22.7

24 1 7 26 19 17 4 17 6 27 18 21

1 — — — — 1

218

A. D. MEISNER KEY TO SPECIES OF DERMOGENYS KUHL & VAN HASSELT IN VAN HASSELT, 1823

1A. 1B. 2A. 2B. 3A. 3B. 4A.

4B.

5A. 5B. 6A. 6B. 7A. 7B. 8A.

8B. 9A.

9B.

Melanophores anterior to anal fin in females arranged into a thin line or oval .................................. ..................................................................................................... (Dermogenys pusilla species-group) 2 Melanophores anterior to the anal fin in females arranged into a conspicuous U or V ...................... ................................................................................................................. (D. orientalis species-group) 5 Third or fourth segment in second anal-fin ray in males elongate, comprising the longest segment of this ray ................................................................................................................................................ 3 Third and fourth segment in second anal-fin ray in males slightly elongate, approximately equal in length ...................................................................................................................................................... 4 Thickening present along anterior edge of second anal-fin ray in males; spiculus straight, divided into 4–6 segments (Fig. 37) ................................D. collettei (Sumatra, Borneo, Peninsular Malaysia) No thickenings along anterior edge of second anal-fin ray in males; spiculus thin, divided into 3–4 segments (Fig. 24) ....................................................................................................... D. pusilla (Java) Segments just proximal to paired spines in second anal-fin ray in males thickened anteriorly; no serrations on anterior edge of second-anal fin ray in males; spiculus thick, straight, distal tip of spiculus without a distinct ventral orientation; spiculus distinctly segmented (Fig. 19) .................... ....................................................................................................... D. burmanica (Burma, Bangladesh) Segments just proximal to the paired spines in second anal-fin ray of males not thickened; serrations present along anterior edge of second anal-fin ray; spiculus sickle-shaped, distal tip oriented ventral; (Fig. 26) ......................................................................... D. siamensis (Thailand, Vietnam, Cambodia) Third or fourth segment in second anal-fin ray in males elongate, comprising the longest segment of this ray ................................................................................................................................................ 6 First segment in second anal-fin ray in males longest; third and fourth segments approximately equal in length ........................................................................................................................................ 9 Cap of bone present on paired spines (Fig. 2) ..................................................................................... 7 No cap of bone on paired spines ...................................................................... D. bruneiensis (Brunei) Two sets of spines on distal tip of spiculus (Fig. 2) ................................................................................. .............................................................. D. bispina (Sabah, Indonesia; Busuanga Island, Philippines) One set of spines on distal tip of the spiculus ..................................................................................... 8 Genital papilla in females elongate with black pigment at distal tip; in females melanophores anterior to anal fin arranged into a broad V with a wide base extending onto lateral body surfaces, directly adjacent to genital papilla and extending anteriorly past level of anus ... D. palawanensis (Palawan) Genital papilla in females short, unpigmented; in females, melanophores anterior to anal fin arranged into a wide, round spot darker anteriorly, diffuse toward anal-fin origin .......... D. robertsi (Culion) Spiculus short, curved ventrally, segmented to distal tip; three or four segments proximal to paired spines thickened along anterior edge (Fig. 22); in females, melanophores arranged into a narrow diffuse U extending anteriorly to level of anus ............................................. D. orientalis (Sulawesi) Spiculus enlarged ventrally, segmented, roughly sickle-shaped with distal tip oriented dorsally (Fig. 28); in females, melanophores anterior to the anal fin arranged into a narrow V extending anteriorly, away from the midline, past level of anus ............ D. sumatrana (Sumatra, Southern Kalimantan)

DERMOGENYS BRACHYNOTOPTERUS (BLEEKER, 1853a)

Hemiramphus brachjnotopterus Bleeker, 1853a: 146 (original description, Hooghly River, India. Dutch ‘j’ equivalent to ‘y’ in Latin). Hemirhamphus brachynotopterus. Gu¨nther, 1866: 275 (characters). Day, 1878: 517 (collection report). Mohr, 1936a: 55 (questioned placement in Dermogenys). Dermogenys brachynopterus. Brembach, 1991: 17 (misspelling). Dermogenys brachynotopterus. Talwar & Jhingram,

1991: 732 (characters, distribution). Kottelat, 1992: 268–269 (taxonomy, emended spelling). Zenarchopterus brachynotopterus. Jayaram, 1981: 289 (characters, distribution). Diagnosis. See Remarks. Description. SL: 45.3 mm. Morphometrics listed as per cent SL: SN-P1, 27.8, SN-P2, 66.9; P2-C, 36.6; HDL, 23.6; ORBL, 5.3. Meristics: dorsal-fin rays [10]; analfin rays [15]; precaudal vertebrae [26]; caudal vertebrae [17]; total vertebrae [43].

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

219

Figure 19. Dermogenys burmanica. A, male, T. Roberts uncatalogued material (MVT 7), 26.1 mm SL; Pegu Division, Bangladesh; B, female, 38.8 mm SL.

Distribution. Hooghly River, India. Colour in alcohol. unknown (see Remarks). Remarks. This species is known in museum collections only from the female holotype. The holotype, which is in poor condition, has 26 precaudal vertebrae. Another species, D. burmanica is characterized by a high number of precaudal vertebrae, generally 25 or more (Table 2), and I question whether the two are actually distinct species. Dermogenys brachynotopterus has been distinguished by an upper jaw that is wider than long. However, this character cannot be verified in the holotype because the upper jaw is missing. Additionally, no other specimens (available in museum collections) appear to have been collected from the Hooghly River. If it is not distinct from D. burmanica, D. brachynotopterus has priority and D. burmanica would be a junior synonym. Material examined. INDIA. 1 specimen (45.3 mm). RMNH 6964 (Holotype of Dermogenys brachynotopterus; female, 45.3 mm); Hooghly River. DERMOGENYS BURMANICA MUKERJI, 1935 (Figures 19, 20; Tables 2, 3, 4, 5, 6)

Dermogenys burmanicus Mukerji, 1935: 213–318 (original description, Delta district of Burma). Brembach, 1991: 155–156 (anatomy, relationships, distribution).

Dermogenys pusillus not of Kuhl & van Hasselt in van Hasselt, 1823. Herre, 1944: 45 (collection report). Downing & Burns, 1995: 330 (identified as Dermogenys pusillus populations from Burma and Bangladesh in testis study). Dermogenys burmanicus. Talwar & Jhingram, 1991: 732 (listed as a synonym of Dermogenys pusillus). Dermogenys burmanica. Kottelat, 1992: 269 (emended spelling). Dermogenys pusilla not of Kuhl & van Hasselt in van Hasselt, 1823. Meisner & Burns, 1997b: 298 (identified as Dermogenys pusilla populations from Myanmar and Bangladesh with Type I halfbeak viviparity). Differential diagnosis. Dermogenys burmanica is distinguished from other Dermogenys by a thick, distinctly segmented spiculus; segments in second anal-fin ray in males just proximal to paired spines thickened along anterior edge, appearing as two distinct thickenings (Fig. 20); higher precaudal vertebrae count, rarely fewer than 25 versus 25 or fewer in other Dermogenys species (Table 2); in females, melanophores anterior to anal fin arranged into a thick line extending from anal fin to genital papilla. Description. (Table 3) Slender-bodied (BDP1 <12% SL), vertebrae 42–44; precaudal vertebrae 25–27, rarely 23 or 24 (Table 2); caudal vertebrae 16–17 (Table 4); predorsal scales 38–45; lower jaw elongate (4.7–7.5 times in SL); upper jaw longer than wide (UJL/UJW

220

A. D. MEISNER

1.1–1.9); branchiostegal rays 9–11; pectoral-fin rays 10; second anal-fin ray in males with 9–11 segments proximal to the paired spines; geniculus beginning at third or fourth segment; second anal-fin ray in males, third or fourth segment slightly elongate; thick, distinctly segmented spiculus (Fig. 20); anal-fin rays 14–16 (Table 5); dorsal-fin origin over anal-fin ray 7 or 8; dorsal-fin rays 9–10 (Table 6); caudal fin oval; type I form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 19) Background colour tan or olive (fresh material); thin midlateral stripe from opercle to caudal fin; scattered melanophores on lateral surfaces; broad, diffuse spot anterior to pectoral fin; some females with black pigment on distal tips of anterior anal-fin rays; Burma [Myanmar] populations, males with black pigment on distal tips of posterior dorsal-fin rays; Bangladesh populations, males and females with black pigment on distal tips of pelvic rays. Distribution. Burma [Myanmar] and Bangladesh. Remarks. The original description was based on two specimens, one male and one female. Because the syntypes were not examined directly by the author, no lectotype is designated.

Figure 20. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Dermogenys burmanica. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. UMMZ 208482 (30.0 mm).

Material examined. BURMA [Myanmar]: 79 specimens from five collections (18.1–60.4 mm SL). ZSI F 117801/1 (syntypes of Dermogenys burmanicus, 1 female, 32.4, 1 male, 31.4, examined by Bruce B. Collette); Hanthawadd District, Kyauktan township, Myagaing village; 1934. CAS 134945 (32 females, 34.9– 60.4, 4 males 28.6–29.4, of which 2 males, 2 females

Table 3. Morphometrics of Dermogenys burmanica (A) female syntype of (ZSI 117891/1); (B) male syntype; (C) 17 females; and (D) 8 males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

32.4 7.5 1.6 30.9 62.7 35.5 27.5 9.6 9.6 5.9 5.9

31.4 5.5 1.9 32.8 61.1 38.2 29.0 9.9 9.9 6.4 5.1

C 30.9–60.4 5.6–7.0 1.1–1.5 28.5–34.3 60.7–65.1 35.2–39.2 22.2–28.7 8.4–10.6 8.4–10.6 3.1–5.5 4.9–6.8

D 28.3–32.9 4.7–5.2 1.1–1.7 31.0–35.0 59.8–65.3 37.2–39.9 24.4–29.9 8.7–11.2 8.7–11.2 8.0–5.6 5.1–6.8

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

221

Table 4. Frequency distribution of caudal vertebrae in species of Dermogenys

D. D. D. D. D. D. D. D. D. D. D. D.

pusilla brachynotopterus sumatrana orientalis siamensis burmanica vogti bispina bruneiensis robertsi palawanensis collettei

15

16

17

18

19

20

Mean

N

1 — 2 — 6 — — — — — — 1

15 — 5 — 12 2 4 — 4 3 — 2

9 1 2 8 — 16 — 4 2 18 — 17

— — 1 14 — — — 9 — 6 5 —

— — — — — — — 4 — — 12 —

— — — — — — — — — — 1 —

16.3 17.0 16.2 17.6 15.7 16.9 17.0 18.0 16.3 17.1 18.7 16.8

25 1 10 22 18 18 4 17 6 27 18 20

Mean

N

14.4 15.0 15.0 15.6 14.1 14.5 15.0 16.1 14.8 14.7 16.1 14.8

25 1 10 23 21 16 4 15 5 25 15 20

Table 5. Frequency distribution of anal-fin rays in species of Dermogenys

D. D. D. D. D. D. D. D. D. D. D. D.

pusilla brachynotopterus sumatrana orientalis siemensis burmanica vogti bispina bruneiensis robertsi palawanensis collettei

14

15

16

17

18

14 — — — 19 9 — — 2 6 — 6

11 1 8 9 2 6 4 2 2 19 1 13

— — 2 12 — 1 — 10 1 — 11 1

— — — 2 — — — 3 — — 3 —

— — — — — — — — — — — —

— — — — — — — — — — — —

Table 6. Frequency distribution of dorsal-fin rays in species of Dermogenys

D. D. D. D. D. D. D. D. D. D. D. D.

pusilla brachynotopterus sumatrana orientalis siamensis burmanica vogti bispina bruneiensis robertsi palawanesis collettei

8

9

10

11

12

13

Mean

N

1 — — — — — — — — — — —

12 — 1 — 13 10 1 5 — 3 — 3

12 1 8 10 8 6 2 9 4 19 1 15

— — 1 12 — — 1 — 1 3 15 4

— — — — — — — — — — — —

— — — — — — — — — — — —

9.4 10.0 10.0 10.5 9.4 9.4 10.0 9.6 10.2 10.0 10.9 10.0

25 1 10 22 21 16 4 14 5 25 16 22

have been cleared and counterstained); Rangoon; Herre; 1937. CAS 61340 (3 females, 18.4–28.4, 6 males 20.1–27.6); Pegu Div. Dayame Chaung (flows into Sittang River), 1 mi north of Daik-U; Roberts, 9 Mar

1985. USNM 342155 (10 females, 18.1–39.2, 10 males, 23.4–31.8); specific locality not specified; Ferraris; 1996. FMNH 51553 (12 females, 48.0–50.4); specific locality not specified; U, Tha, Myint; Jul 1951.

222

A. D. MEISNER

Figure 21. D. orientalis. A, male, USNM 338416, 30.2 mm SL, Selatan, Gowa District, Limbung, Sulawesi; B, female, USNM 338416, 46.5 mm SL.

BANGLADESH: 43 specimens from three collections (19.2–47.6 mm SL). UMMZ 208497 (6 females, 23.7–45.4, 3 males, 25.0– 26.4); Chittagong, Ichamati River at mouth into Karnafuli River, Rainboth, Tsai, Hussain, and Quader; 27 Dec 1977. UMMZ 208482 (14 females, 19.2–47.6, 6 males, 23.4–32.9, of which 2 males, 2 females have been cleared and counterstained); Chittagong Hill Tracts, Sangu River and small trib. creek ca. 1 mi upstream from Bandarban; Rainboth, Tsait, Quader; 25 Dec 1977. MVT 7, uncatalogued (9 females, 25.3–39.6, 5 males, 22.9–25.8); Pegu Division, Kha-Yein Chang number 4 km NE of Hlegu on road to Pegu at Wah Knit Kone village; Roberts; 1985. DERMOGENYS ORIENTALIS (WEBER, 1894) (Figures 21, 22; Tables 2, 4, 5, 6, 7)

Hemiramphus orientalis Weber, 1894: 427 (original description, Celebes [Sulawesi]). Herre, 1953: 159 (listed as a synonym of Dermogenys pusillus). Hemirhamphus orientalis. Boulenger, 1897: 429 (characters, distribution). Herre, 1953: 159 (listed as a synonym of Dermogenys pusillus).

Dermogenys orientalis. Weber & de Beaufort, 1922: 137 (characters, distribution). Herre, 1931: 25 (collection report). Mohr, 1936a: 41 (placed in synonymy with Dermogenys pusillus). Herre, 1953: 159 (listed as a synonym of Dermogenys pusillus). Brembach, 1991: 162 (characters, relationships, distribution). Kottelat, 1992: 270 (taxonomy). Kottelat et al., 1993: 118 (distribution). Downing & Burns, 1995: 330 (testis study). Dermogenys montanus. Brembach, 1982: 54 (original description, Bantimurung, Sulawesi). Brembach, 1991: 161 (anatomy, relationships, distribution). Downing & Burns, 1995: 330 (testis study). Dermogenys montana. Kottelat, 1992: 269–270 (emended spelling). Kottelat et al., 1993: 118 (distribution). Dermogenys sp. (Sulawesi). Meisner & Burns, 1997b: 298 (identified as Dermogenys sp. (Sulawesi) with Type II form of halfbeak viviparity). Differential diagnosis. Dermogenys orientalis is distinguished from other Dermogenys by the short, thin spiculus, curved ventrally, segmented to distal tip (Fig. 22); in females, melanophores anterior to anal fin

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

223

1995); type II form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 21) Background colour white or tan; thin, midlateral stripe from opercle to caudal fin; black pigment along length of lower jaw; scattered melanophores on dorsal portion of body; fins with scattered melanophores; concentration of melanophores at base of pelvic fin; line of melanophores at base of pectoral fin; some with pigment on distal tips of pelvic fins. Distribution. Maros, Sulawesi Selatan, Indonesia. Remarks. An unambiguous diagnostic character, short, thin spiculus curved ventrally and segmented to its distal tip is found in males of this species, therefore, the largest male syntype examined is herein designated as the lectotype. In the original description of D. montanus (Brembach, 1982), here considered a junior synonym of D. orientalis, it was stated that the description was based on 25 specimens with no holotype designated. In a subsequent revision, it was reported that three lots were examined containing the holotype and 25 paratypes (Brembach, 1991). Based on the original description, this ‘holotype’, if separated, is actually the lectotype and the lots, which contain 30 specimens, are paralectotypes.

Figure 22. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Dermogenys orientalis. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. USNM 338416 (31.0 mm).

arranged into a narrow, diffuse U, extending anteriorly to level of anus. Description. (Table 7) Slender-bodied (BDP1 <16.1% SL); vertebrae 38–41 [40]; precaudal vertebrae 21–24 [22] (Table 2); caudal vertebrae 17–18 [18] (Table 4); predorsal scales 27–34; lower jaw elongate (4.5–9.1 times in SL); upper jaw longer than wide (UJL/UJW 1.1–1.4); branchiostegal rays 10–12; pectoral-fin rays 10–12; genital papilla in females elongate, pigmented; second anal-fin ray in males with 7–8 segments proximal to paired spines; segments 3 and 4 slightly elongate, approximately equal in length; three or four segments proximal to spines appear thickened along anterior edge; short, thin spiculus curved ventrally, segmented to distal tip (Fig. 22); anal-fin rays 15–17 [16] (Table 5); dorsal-fin origin over anal-fin ray 6 or 7 [7]; dorsal-fin rays 10–11 [10] (Table 6); caudal fin oval; thread-like sperm nuclei (Downing & Burns,

Material examined. SULAWESI SELATAN: 653 specimens from 20 collections (14.6–66.0 mm SL). Lectotype of Hemiramphus orientalis (ZMA 104.374) (herein designated): (male, 31.4). Measurements listed in Table 7. ZMA 123.573 (paralectotypes of Hemiramphus orientalis, 10 females, 23.1–39.2, 4 males, 21.3–27.8); ZMH 7145 (radiograph of lectotype of Dermogenys montana); Bantimurung; Brembach; 1978. ZMH 7147 (paralectotypes of Dermogenys montana, 9 females, 48.9– 66.0, 19 males, 30.4–34.9); collected with the lectotype. ZMH 7146 (radiograph of paralectotype of Dermogenys montana, 1 male). USNM 338416 (144 specimens, females, 19.0–49.8, males, 21.8–31.4, of which 2 males, 2 females have been cleared and counterstained); Gowa District, Limbung, approximately 5 km SE on road from Ujung Pandang to Patalasang; Louie, Parenti, and Amos; 3 Aug 1995. USNM 338499 (24 females, 18.0– 62.0, 1 male, 23.0); Maros, Bantimurung; Louie and Raftik; 22 Jul 1995. USNM 338417 (12 females, 14.6– 47.6, 5 males, 18.6–22.3); Maros, Tanralili, approximately 15 km SE of turnoff at Batanahse on road from Ujung Pandang to Maros; Louie and Parenti; 5 Aug 1995. USNM 338418 (127 specimens, females, 25.0–63.4, no mature males); Pangkep; Minasatene and Louie; 24 Jul 1995. USNM 338498 (1 female, 30.5,

224

A. D. MEISNER Table 7. Morphometrics of Dermogenys orientalis. (A) male lectotype of (ZMA 104.374); (B) three female paralectotypes; (C) one male paralectotype; (D) 20 females; and (E) 22 males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

D

E

31.4 — 1.4 36.1 60.7 40.3 34.1 13.7 13.7 6.1 6.1

30.9–39.2 — 1.1–1.3 29.5–32.1 61.2–63.3 39.3–41.8 25.9–29.9 11.3–12.2 11.3–12.2 5.6–6.3 6.5–7.1

27.8 — 1.3 34.5 58.6 40.7 27.7 11.5 11.2 6.1 6.8

39.3–66.9 5.1–9.1 1.1–1.4 32.8–35.1 59.8–64.3 36.6–42.8 25.3–31.0 11.9–13.9 11.9–15.1 4.3–5.7 5.5–7.7

26.7–34.7 4.5–6.4 1.1–1.4 34.8–37.6 58.7–63.9 37.7–41.3 29.6–34.2 12.3–16.1 12.3–16.1 5.6–6.9 5.8–7.9

11 males, 21.7–35.8); Maros, Bantimurung; Louie and Raftik; 22 Jul 1995. USNM 338496 (81 specimens, females, 26.8–35.8, males, 22.8–32.2, of which 2 males have been cleared and counterstained); Pangkep; Minasatene, Panekang, Louie and Amos; 24 Jul 1995. CMK 6109 (15 females, 21.1–60.8, 8 males, 15.0–30.0, 21 undet., of which 1 female, 3 males have been cleared and counterstained); Kabupaten Maros: above Bantimurung waterfall; Kottelat; 7 Jun 1988. CMK 6102 (9 females, 18.9–52.2, 17 males, 20.5–32.4, 36 undet., of which 2 males have been cleared and counterstained); Kabupaten Maros: Pattuang; Kottelat; 7 Jun 1988. CMK 6147 (13 females, 17.7–47.6); Sungei Leorang, km 3 on road from Enrekang to Pare Pare; Kottelat, and Kottelat-Kloetzli; 14 Jun 1988. CMK 6454 (7 females, 16.6– 51.9, 7 males, 15.7–27.9); Desa Kolukka, 4 km south of Bone on the road to Palopo; Kottelat and Werner; 10 Mar 1989. CMK 6455 (5 females, 24.0–57.9, 2 males, 24.9–32.7); km 11 on road from Malili to Wotu; Kottelat and Werner; 10 Mar 1989. CMK 6188 (5 females, 28.1– 40.6, 2 males, 28.3); small stream in swampy area close to Sungei Malili, about midway between and junction of S. Larona and S. Pongkaru; Kottelat and KottelatKloetzli; 19 Jun 1988. CMK 6445 (7 females, 17.6–29.3, 5 males, 21.4–30.4); Sungei Terak, km 19 on road from Enrekang to Pare Pare; Kottelat and Werner; 9 Mar 1989. CMK 6146 (8 females, 16.0–28.8, 6 males, 16.8– 23.6, 17 undet.); Sungei Nanggala at Marante, Desa Tandung, Kecamatan Rantepoa, Kabupaten Makale; Kottelat and Kottelat-Kloetzli; 12 Jun 1988. CMK 6119 (3 females, 37.4–59.3, 1 male, 29.0); Road from Ujung Pandang to Malino, Sungei Jeneberang at Desa Lanna, Kecamatan Parangloe, Kabupaten Gowa; Kottelat and Kottelat-Kloetzli; 9 Jun 1988. USNM 193012 (4 females, 22.9–48.1 mm, 3 males, 30.3–31.8, of which 1 female, 1 male have been cleared and counterstained); Banti

Moerong [Bantimurung], above falls; Smithsonian– National Geographic Expedition; 1937. DERMOGENYS PUSILLA KUHL & VAN HASSELT IN VAN HASSELT, 1823 (Figures 23, 24; Tables 2, 4, 5, 6, 8)

Dermogenys pusillus Kuhl & van Hasselt in van Hasselt, 1823: 131 (original description, Buitenzorg [Bogor], Java). Bleeker, 1865: 30–31 (characters, distribution). Bean & Weed, 1912: 595–596 (collection report). Weber & de Beaufort, 1922: 140 (characters, distribution). Weed, 1933: 49–50 (characters). Mohr, 1936a: 41–50 (revision). Herre, 1944: 43–45 (collection report). Herre, 1953: 159 (synonymy). Jayaram, 1981: 289 (distribution). Talwar & Jhingram, 1991: 732–733 (characters, distribution). Downing & Burns, 1995: 330 (testis study). Hemiramphus fluviatilis Bleeker, 1850: 95 (original description, Buitenzorg [Bogor], Java). Bleeker, 1865: 31 (listed as a synonym of Dermogenys pusillus). Weber, 1894: 427 (distribution). Hemirhamphus fluviatilis. Gu¨nther, 1866: 275 (characters). Dermatogenys fluviatilis. Jordan & Seale, 1908: 540 (characters, collection report). Mohr, 1936a: 41–50 (synonymy). Dermogenys pusillus pusillus. Brembach, 1991: 168 (characters, distribution). Dermogenys pusilla. Kottelat, 1992: 269–270 (emended spelling). Kottelat et al., 1993: 118 (characters, distribution). Meisner & Burns, 1997b: 298 (reproductive biology, viviparity). Differential diagnosis. The anal fin of Dermogenys pusilla is most similar to that of D. collettei. It is

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

225

Figure 23. Dermogenys pusilla. A, female, UMMZ 155726, 30.1 mm SL; circles are oocytes as seen through the body wall; B, neotype, male, UMMZ 237500, 22.6 mm SL; Buitenzorg [Bogor], Java.

distinguished from all other Dermogenys by its thin spiculus, generally divided into three or, at most, four segments (Fig. 24); andropodium distinguished from D. collettei by absence of the anterior thickening of segments 4–6 on the second anal-fin ray and a spiculus with fewer segments. Description. (Table 8) Slender-bodied (BDP1 <12% SL); lower jaw elongate (5.9–7.9 times in SL); precaudal vertebrae 22–24 [22] (Table 2); caudal vertebrae 15–17 [17] (Table 5); total vertebrae 38–40 [39]; predorsal scales 32–38; upper jaw longer than wide (UJL/UJW 1.1–1.6); branchiostegal rays 9–12; pectoral-fin rays 9–10; anal-fin rays 14–15 [14] (Table 5); second analfin ray in males with 7 segments proximal to paired spines, third or fourth segment elongate; spiculus thin, divided into three or four segments (Fig. 24); dorsalfin origin over anal-fin ray 6–7 [6]; dorsal-fin rays 8–10 [8] (Table 6); caudal fin oval; type I form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 23) Background colour tan or brown; diffuse midlateral stripe from opercle to base of caudal fin; in females, melanophores anterior to anal fin forming a wide oval extending from anal-fin origin to genital papilla; black pigment at base of

pelvic-fin rays in both males and females, distal tips of pelvic-fin rays and posterior dorsal-fin rays in males; concentration of melanophores at base of pectoral fins in both sexes. Distribution. Java, Indonesia. Remarks. In the first revision of the genus, Mohr (1936a) placed several species in synonymy with Dermogenys pusilla. In a subsequent revision, this synonymy was uncritically accepted and two additional subspecies of D. pusilla (D. pusillus pusillus, D. pusillus borneensis, here considered a junior synonym of D. sumatrana) were described (Brembach, 1991). This has led to the extensive use of Dermogenys pusilla in museum collections and throughout the literature. In an investigation of Kuhl and van Hasselt material collected in Java, Roberts (1993) reported that no specimens of the original collection of D. pusilla exist. As stated in the International Code of Zoological Nomenclature (1985), a neotype can be designated under “exceptional circumstances” referring to cases in which a neotype is critical for solving a complex zoological problem such as “the confused or doubtful identities of closely similar nominal species-group taxa . . .” (p. 157).

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Therefore, to stabilize the identity of D. pusilla, a neotype is herein designated from the type locality. Material examined. JAVA: 216 specimens from 14 collections (15.5–46.2 mm SL). Neotype (herein designated): UMMZ 237500 (male, 22.6); pond in botanical gardens, Buitenzorg [Bogor], Java; Thienemann; 15 Sep 1928. Measurements listed in Table 8.

Figure 24. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Dermogenys pusilla. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. UMMZ 155727 (25.0 mm).

BMNH 1866.5.229 (holotype of Hemiramphus fluviatilis; female, 46.2; Buitenzorg [Bogor]). UMMZ 155726 (16 females, 19.1–30.6, 7 males, 17.3–23.9, one undet.); pond in botanical gardens, Buitenzorg [Bogor]; Thienenmann; 15 Sep 1928. UMMZ 155820 (10 females, 17.9–41.3, 5 males, 19.7–24.8, 2 undet.); Lake Bagendit, near Garoet, elev ca 700 m; Hubbs and Arragon; 26 May 1929. UMMZ 155819 (1 female, 30.7, 1 male, 19.4, 2 undet.); Rice fields near Padalarong; Hubbs and Arragon; 28 May 1929. UMMZ 155815 (53, females, 33.2–40.5, males, 22.0–24.4, of which 2 males have been cleared and counterstained); Sitoe Tjiboeroej, near Padalarang, 20 km. W of Bandoeng, elev. ca 700 m; Hubbs and party; 22 May 1929. UMMZ 155814 (12 females, 27.8–34.7, 4 males, 22.5–22.6, 1 undet.); Tjiti’is (creek) just below road near mouth in Tjimanoek, 3 km N of Garoet, elev. nearly 700 m, Hubbs and Arragon, et al.; 26 May 1929. UMMZ 155816 (13 females, 19.5–37.3, 10 males, 15.5–25.8); Tijwalen (creek), trib. of Tjitandoej (River) at Godebak (kampong) between Panoembagen and Pandjaloe, elev. ca 465 m; Hubbs and Arragon; 27 May 1929. USNM 72572 (7 females 28.1–36.2, 1 male, 25.8); Buitenzorg [Bogor], Bryant and Palmer; 1909. USNM 72569 (1 male, 24.0); Buitenzorg [Bogor], Bryant and Palmer, 1909. CMK

Table 8. Morphometrics of Dermogenys pusilla. (A) male neotype (UMMZ 237500); (B) 11 females; and (D) 9 males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

22.6 6.1 1.6 40.3 65.9 38.5 31.0 11.1 11.1 6.2 5.8

26.5–40.5 6.0–7.9 1.1–1.5 30.5–34.2 56.7–62.9 37.8–42.8 23.9–28.5 9.1–11.0 10.5–12.7 4.3–6.2 5.7–6.9

21.0–24.9 5.9–7.2 1.1–1.4 32.6–36.3 56.4–61.4 39.7–44.1 25.6–28.0 9.0–11.6 10.8–13.1 5.8–7.1 6.0–7.0

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Figure 25. Dermogenys siamensis. A, male, CMK 10709, 26.7 mm SL; Trat Province, Thailand; B, female, CMK 10709, 48.8 mm SL.

6094 (5 females, 21.3–30.4, 5 males, 19.4–24.3, 1 undet.); Kabupaten Bogor, Situ [=lake] Sawangen, Viviaria Indonesia fish farm; Kottelat and Yuwono, 4 Jun 1988. BMNH 1912.12.10: 31–32 (2 females 27.7–32.4). CAS 120500 (8 females, 23.5–40.9, 3 males, 21.6–24.0), Buitenzorg [Bogor], Campbell. CMK 9187 (25 females, 27.2–27.8, 17 males, 20.7–26.3, 1 undet.); Timur, Kali Brantas basin: irrigation canal Kali Suko in Desa Pungging Kec. Pungging, Jembatan Ringgil, 7°30′S 112°30′E; Kottelat, 10 Jul 1992. DERMOGENYS SIAMENSIS FOWLER, 1934a (Figures 25, 26; Tables 2, 5, 6, 7, 9)

Dermogenys siamensis Fowler, 1934a: 144–145 (original description, Chieng Mai, North Siam [Thailand]). Brembach, 1991: 166 (comparison to D. pusillus borealis Brembach, 1991: 166). Kottelat, 1992: 270 (taxonomy). Meisner & Burns, 1997b: 298 (reproductive biology). Dermogenys pusillus not of Kuhl & van Hasselt in van Hasselt, 1823. Kottelat, 1985: 271 (collection report, Kampuchea). Downing & Burns, 1995: 330 (identified as populations from Vietnam and Thailand in a testis study).Dermogenys pusillus borealis. Brembach, 1991: 166 (original description, Bangkok, Thailand). Dermogenys pusilla borealis. Kottelat, 1992: 270 (taxonomy, emended spelling). Dermogenys pusilla not of Kuhl & van Hasselt in

van Hasselt, 1823. Meisner & Burns, 1997b: 298 (identified as populations from Thailand and Vietnam with the Type I form of halfbeak viviparity). Differential diagnosis. Dermogenys siamensis is distinguished from other Dermogenys by sickle-shaped spiculus, segmented distally to tip; serrations on anterior edge of second anal-fin ray and posterior edge of fourth anal-fin ray (Fig. 26). Description. (Table 9) Slender-bodied (BDP1 <13% SL); vertebrae 38–40 [40]; precaudal vertebrae 23–24 [24] (Table 2); caudal vertebrae 15–16 [16] (Table 4); 32–37 predorsal scales; lower jaw elongate (4.5–7.5 times in SL); upper jaw longer than wide (UJL/UJW 1.1–1.7); branchiostegal rays 9; pectoral-fin rays 9–10; second anal-fin ray in males with 6–8 segments proximal to paired spines, segments three and four slightly elongate, approximately equal in length; sickle-shaped spiculus, segmented distally to tip (Fig. 26); serrations on anterior edge of second anal-fin ray and posterior edge of fourth anal-fin ray; in females, concentration of melanophores anterior to anal fin varying from a thin oval extending from anteriormost anal-fin ray to genital papilla to a thin line or absent; anal-fin rays 14–15 [14] (Table 5); dorsal-fin origin over anal-fin ray 6 or 7 [6]; dorsal-fin rays 9–10 [9] (Table 6); caudal fin oval; type I form of halfbeak viviparity (Meisner & Burns, 1997b).

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the same lot (ANSP 59861–59875). In the original description, the largest specimen, having nine dorsalfin rays was designated as the holotype (Fowler, 1934a). Consequently, the largest specimen also with nine dorsal-fin rays from this combined lot is identified as the holotype. Material examined. THAILAND. 178 specimens from 21 collections (17.1–53.0 mm SL).

Figure 26. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Dermogenys siamensis. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. UMMZ 224700 (23.0 mm).

Colour in alcohol. (Fig. 25) Background colour white or tan; diffuse midlateral stripe from pectoral to caudal fin, becoming more prominent posteriorly; elongate spot present anterior to pectoral fin; black pigment along length of lower jaw; in females, large melanophores scattered ventrally from pelvic to anal fins; in males, black pigment at base and distal tips of pelvicfin rays and posterior dorsal-fin rays; in females black pigment at base of pelvic fins. Distribution. Thailand, Vietnam, Cambodia. Remarks. In the original description of D. siamensis, a holotype was designated and given a separate catalog number (ANSP 59860). However, the paratypes and holotype were not separated and were combined within

ANSP 59860 (holotype of D. siamensis, female, 47.4); Chiang Mai, on Me Nam Ping; de Schauensee; 22 Jan 1933. ANSP 59861–59875 (paratypes of Dermogenys siamensis, 7 females, 24.3–41.2, 8 males, 19.8–28.7, of which 1 male, 1 female have been cleared and counterstained); collected with holotype. ZMH 7612 (radiograph of holotype of Dermogenys pusillus borealis, male); Bangkok. ZMH 7138 (11 females, 26.8– 42.0, 5 males 23.9–27.3); Bangkok; 1965. USNM 119517 (29 females, 34.1–53.0, 2 males, 27.7); Meping River at Chiang Mai; Deignam; 22 Apr 1941. USNM 109702 (2 females, 42.4–45.6); Nonthaburi, Menanchao Phya; Smith; 2 Sep 1940. USNM 109802 (5 females, 20.5–36.5, 4 males, 20.6–25.1); Bangkok Canal; Roberts and Wongrat; 10 May 1973. USNM 229290 (3 females, 32.0–38.4, 1 male, 21.5); Klong Ta Pa at Ban Pong, W. of Bangkok; Roberts and Wongrat; 15 Apr 1973. UMMZ 195917 (11 females, 17.6–46.0, 15 males, 18.2–27.8, of which 2 males and 2 females have been cleared and counterstained); Chanthaburi, Trib. of Khlong Na Ngam, 1 km south of Ban Ang; Gulf of Thailand drainage; Lagler, Suvatti, and Boonbrahm; 6 Apr 1965. BMNH 1915.5.12: 15–17 (3 females, 36.5– 48.1, 1 male, 21.5, of which 1 female has been cleared and counterstained). CMK 5262 (6 females, 24.8–41.6, 6 males, 23.1–28.3); Chiang Mai Prov., Mae Nam Ping at Chiang Mai; Kottelat; 16 Apr 1985. CMK 5392 (5 females, 19.6–39.5, 1 male, 24.2); Ranong Prov., stream on road from Ranong to Kra Buri, km 37, 10°15′N 98°45′E; Kottelat; 24 Apr 1985. CMK 5352 (4 females, 18.8–50.4); Phangnga Prov., Khlong Khum Mui, west of Amphoe Takum Thung, road from Phangnga to Phuket, km 16, 8°23′N 98°27′E; Kottelat; 22 Apr 1985. CMK 10709 (6 females, 19.8–48.1, 3 males, 24.5–26.2); Trat Prov., Khlong Huai Raeng, km 18 on road 3271 from Trat to Bo Rai, 12°24′N 102°40′E; Kottelat, Lim, Tan and Kubota; 3 Dec 1993. CMK 10684 (5 females, 26.2–34.5, 4 males, 19.1–26.9); Trat Prov., stream near Ban Tha Kum, 9 km north of Ban Noen Sung on road 3271 from Trat to Bo Rai, 12°32′N 102°37′E; Kottelat, Lim, Tan, and Kubota; 3 Dec 1993. MCZ 47131 (1 male, 27.7); Kra Isthmus, Isthmus of Kra Creek Khlong Falamee, a swift muddy creek flowing into inner lake of Tale Sap, about 2 km W of Pak Payoon; Roberts; 20 Jun 1970. MCZ 47040 (3 females, 17.6–37.1, 3 males, 23.8–25.5, 1 undet.); Kotumban, ca. 30 km from Bangkok on the way to Samut Sarkorn; Roberts; 11 Jul 1970.

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Table 9. Morphometrics of Dermogenys siamensis. (A) female holotype (ANSP 59860); (B) three famele paratypes (ANSP 59861–59875); (C) four male paratypes (ANSP 59861); (D) 15 females; and (E) 12 males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

D

E

47.4 5.6 1.4 30.0 64.1 39.9 24.3 10.8 10.8 4.6 5.7

32.6–41.2 — 1.3–1.4 29.1–31.4 58.7–63.3 38.9–42.3 26.1–26.5 9.1–11.4 9.1–11.4 4.8–5.2 5.4–6.1

26.3–28.7 — 1.2–1.6 33.1–35.4 60.3–61.7 37.6–39.2 28.6–32.7 9.4–10.7 9.4–10.7 5.2–5.9 5.5–5.9

29.3–43.7 5.4–5.74 1.1–1.4 31.2–33.3 59.6–63.8 36.9–40.7 26.4–29.8 9.6–11.4 9.6–11.4 4.6–5.3 5.8–6.9

21.8–34.5 0.5–7.5 1.3–1.7 34.7–37.2 60.6–66.1 35.9–41.7 28.9–31.5 8.3–12.7 8.3–12.7 4.4–6.8 5.5–6.8

MCZ 47115 (4 females, 39.4–43.7, 2 males, 32.7–34.5); Chantiburi Prov., roadside ditch, 2 km. from Tah Chalap, on road from Chantiburi City; Roberts; 5 May 1970. MCZ 47114 (1 female, 34.2, 7 males, 22.0–24.4); Om-noy, ca. 26 km from Bangkok on the way to Samut Sarkorn; Roberts; 11 Jul 1970. MCZ 57394 (2 females, 23.6–24.2); Chantiburi, Khlong Kee Nawn behind Catholic Church, Chantiburi City; Roberts and Wongrat; 5 May 1970. MCZ 47041 (3 females, 31.6–34.5, 2 males, 30.6); Kotumban, ca. 30 km from Bangkok on the way to Samut Sarkorn; Roberts; 11 Jul 1970.

River drainage; Rainboth, Arden, and Minh; 22 Oct 1974. UMMZ 227298 (6 females, 18.4–38.2, 5 males, 21.6–23.9); North bank of Mekong River at mouth of canal 2 km below Vinh Long; Smith and Weidenbach; 22 Jun 1974. UMMZ 227297 (3 females, 32.9–39.9); Tidal pond near Vinh Long, Mekong River drainage; Fishermen; 22 Jun 1974. UMMZ 224704 (10 females, 27.1–45.2, 1 male, 21.3); An Giang Prov., Ministry of agriculture rice field ca. 10 km W of Long Xuyan on N.R. 10; Smith, Weidenbach, and Rainboth; 22 Jul 1974.

CAMBODIA. Nine specimens from five collections (16.6–34.0 mm SL). CMK 4776 (2 females, 16.6–31.6, 1 male, 27.9); O Po Kampon, Tham Pra Kam, d’Aubenton, 18 Jun 1964. CMK 4777 (3 males, 21.2–27.6); Road to Sianoukville, Tuk Sap; d’Aubenton; 14 Feb 1961. MNHN 1985-977 (1 female, 34.0); route de Sianoukville; Tuk Sap; 1962. MNHN 1985-989 (1 female, 31.3); route de Sianoukville, km 170; 1964. MNHN 1985-983 (1 female, 28.2, 1 male, 20.6); Bengkebal Damrey; d’Aubenton; 1962. VIETNAM. 80 specimens (18.4–45.2 mm SL).

from

six

collections

UMMZ 224825 (2 females, 31.5–32.1); Chan Coc, South end of Vinh Tuong Island, Bassae River; Rainboth, Arden, and Minh; 1974. UMMZ 224700 (21 females, 23.5–43.6, 17 males, 19.3–25.0, 1 undet., of which 2 females, 2 males have been cleared and counterstained); An Giang Prov., ca. 10 km W of Long Xuyen on NR10, Mekong Drainage; Smith, Weibenbach, and Rainboth; 22 Jul 1974. UMMZ 225194 (11 females, 22.6–39.1, 3 males, 20.1–21.8); An Giang Prov., 1 km SW of Hwy 10 in Hoa Binh Thanh district, Mekong

DERMOGENYS SUMATRANA (BLEEKER, 1853b) (Figures 27, 28; Tables 2, 4, 5, 6, 10)

Hemiramphus sumatranus Bleeker, 1853b: 526 (original description, Lake Manidjau, Sumatra). Herre, 1953: 159 (listed as a synonym of Dermogenys pusillus). Hemirhamphus sumatranus. Bleeker, 1865: 32 (listed as a synonym of Dermogenys sumatranus). Gu¨nther, 1866: 275 (characters). Herre, 1953: 159 (listed as a synonym of Dermogenys pusillus). Dermogenys sumatranus. Bleeker, 1865: 32 (characters, distribution). Weber & de Beaufort, 1922: 139–140 (characters, distribution). Mohr, 1936a: 41 (listed as a synonym of Dermogenys pusillus). Herre, 1953: 159 (listed as a synonym of Dermogenys pusillus). Brembach, 1991: 167 (description of andropodium). Dermogenys pusillus not of Kuhl & van Hasselt in van Hasselt, 1823. Roberts, 1989: 154 (collection report). Dermogenys pusillus borneensis. Brembach, 1991: 167 (original description, Pontianak, Kalimantan Western Borneo).

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Figure 27. Dermogenys sumatrana. A, male, RMNH 10435, 31.1 mm SL; Bukittinggi, Sumatra; B, female, RMNH 10435, 48.9 mm SL.

Dermogenys pusilla borneensis. Kottelat, 1992: 270– 271 (taxonomy, emended spelling). Dermogenys sumatrana. Kottelat, 1992: 269, 271 (taxonomy, emended spelling). Kottelat et al., 1993: 118 (distribution). Dermogenys sp. Downing & Burns, 1995: 330 (identified as Dermogenys sp. population from East Borneo in a testis study). Differential diagnosis. Dermogenys sumatrana is distinguished from other Dermogenys by the spiculus which is enlarged ventrally, segmented, roughly sickleshaped with distal tip oriented dorsally (Fig. 28); in females melanophores anterior to anal fin arranged into a narrow V extending anteriorly away from midline past level of anus. Description. (Table 10) Slender-bodied (BDP1 <13% SL); vertebrae 39–42 [42]; precaudal vertebrae 22–24 [24] (Table 2); caudal vertebrae 15–18 [18] (Table 4); predorsal scales 28–34; lower jaw elongate (4.7–6.2 times in SL); upper jaw longer than wide (UJL/UJW 1.2–1.6); branchiostegal rays 10; pectoral-fin rays 10– 11; genital papilla in females elongate, black pigment along its length; second anal-fin ray in males with nine segments distal to spines; segmentation approximately

equal; spiculus enlarged ventrally, segmented, roughly sickle-shaped, distal tip oriented dorsally (Fig. 28); anal-fin rays 15–16 [16] (Table 5); dorsal-fin origin over anal-fin ray 6–8 [8]; dorsal-fin rays 9–11 [10] (Table 6); caudal fin oval; threadlike sperm nuclei (Downing & Burns, 1995); type II form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 27) Background colour tan; thin midlateral stripe from opercle to caudal fin; scattered melanophores over dorsal surface; black pigment along length of lower jaw; black pigment at base of pelvic fins; all other fins dusky. Remarks. No lectotype is designated because the only remaining syntype is a female in poor condition and diagnostic characters are difficult to distinguish. Distribution. Kalimantan Barat and Bukittinggi (western Sumatra), Indonesia. Material examined. SUMATRA. Three specimens from two collections (31.7–51.6 mm SL). BMNH 1866.5.330 (syntype of Hemiramphus sumatranus, female, 51.6); Lake Manidjau. RMNH 10435 (1 female, 48.1, 1 male, 31.7); Fort de Kock [Bukittinggi]; Jacobson, 1920.

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169–170 (anatomy, relationships, distribution). Kottelat et al., 1993: 118 (distribution). Diagnosis. See Remarks. Description. (Table 11) Deep-bodied (BDP1>14% SL); vertebrae 38–41 [41]; precaudal vertebrae 21–24 [24] (Table 2); caudal vertebrae 17 [17] (Table 4); predorsal scales 46–54; lower jaw short (9.0–10.2 times in SL); upper jaw longer than wide (UJL/UJW 1.1); females with elongate, pigmented genital papilla; anal-fin rays 15 [15] (Table 5); dorsal-fin origin over anal-fin ray 7 [7]; dorsal-fin rays 9–11 [9] (Table 6); caudal fin truncate. Colour in alcohol. (Fig. 29) Background colour brown; scattered melanophores on lateral surfaces; thin midlateral stripe from opercle to base of caudal fin. Distribution. Topobulu, Sulawesi Selatan, Indonesia.

Figure 28. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Dermogenys sumatrana. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. USNM 306569 (31.0 mm).

KALIMANTAN BARAT. 27 specimens from three collections (23.1–53.0 mm SL). ZMA 112.571 (syntypes of Dermogenys pusillus borneensis, 4 females, 30.9–43.2, 5 males, 27.9–28.1, 4 undet.); Sungei Blakin upper course, south Meridian, mangrove vegetation; L. Rutten; circa 1911. USNM 306569 (6 females, 28.0–53.0, 5 males, 26.3–30.4, of which 2 males have been cleared and counterstained); Sungei Semosan about 50–60 km before the sea; Christensen; 13 Jul 1982. CAS 49428 (1 female 28.5, 2 males, 23.1–28.5) Kapuas River basin, Sungei Durian, non-mangrove tidal creek flowing directly into Kapuas Ketchil, 7–8 km WNW of Pontianak; Roberts, Woerjoatmodjo; 13 Jul 1976. DERMOGENYS VOGTI BREMBACH, 1982 (Figure 29; Table 2, 4, 5, 6, 11)

Dermogenys vogti Brembach, 1982: 54–55 (original description, Topobulu, Sulawesi). Brembach, 1991:

Remarks. Dermogenys vogti is known only from four female specimens and without males, diagnostic characters are elusive. However, this species is tentatively classified as a Dermogenys based on presence of an elongate, pigmented genital papilla in females. If in fact it is a valid species, its affinities may lie within the D. orientalis-clade (see discussion). In a previous revision of the genus (Brembach, 1991), the lectotype (designated by Brembach, 1991) was erroneously called a holotype. Material examined. Four specimens from two collections (50.7–61.7 mm SL). ZMH 7148 (radiograph of lectotype, 1 female, 61.7); Topobulu, Sulawesi. ZMH 7149 (paralectotypes, 3 females, 50.7–56.8); collected with holotype.

DERMOGENYS BISPINA MEISNER & COLLETTE, 1998 (Figures 30, 31; Tables 2, 4, 5, 6, 12)

Dermogenys pusillus not of Kuhl & van Hasselt in van Hasselt, 1823. Herre, 1944: 43–45 (in part, specimens from British North Borneo [Sabah], CAS 133611, CAS 133488). Inger, 1955: 66–67 (characters, biology, collection report). Inger & Chin, 1962: 152–154 (characters, biology). Downing & Burns, 1995: 330 (identified as population from North Borneo in a testis study). Dermogenys pusilla not of Kuhl & van Hasselt in van Hasselt, 1823. Meisner & Burns, 1997b: 298 (identified as population from Sabah with the Type II form of halfbeak viviparity). Differential diagnosis. Dermogenys bispina is distinguished from all other species of Dermogenys by

232

A. D. MEISNER Table 10. Morphometrics of Dermogenys sumatrana. (A) female syntype (BMNH 1866.5.330); (B) 10 females; (C) 6 males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202 A SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

B

51.6 — 1.5 26.9 62.7 40.5 24.4 10.0 10.0 5.6 6.6

30.7–53.2 5.3–6.2∗ 1.3–1.6 30.7–35.1 59.5–65.4 36.9–40.8 24.3–29.5 10.4–12.3 9.4–12.3 5.2–6.4 6.0–7.3

C 26.4–31.1 4.7–5.3∗∗ 1.2–1.6 31.8–38.9 61.7–64.7 38.7–40.5 26.4–32.4 9.9–11.4 9.9–11.4 5.5–6.4 6.1–7.3

∗ Proportion based on two specimens. ∗∗ Proportion based on three specimens.

Figure 29. Dermogenys vogti, paralectotype, female, ZMH 7149, 56.9 mm SL; collected with lectotype Tobobulu, Sulawesi.

Table 11. Morphometrics of three female paralectotypes of Dermogenys vogti (ZMH 7149). Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202 SL 1. SL/LJL 2. UJL/UJW 3. SN-P1 4. SN-P2 5. P2-C 6 HDL 7. BDP1 8. BDP2 9. ORBL 10. INTORBL

50.7–56.9 9.0–10.2 1.1 31.5–32.2 57.8–60.8 40.2–41.8 27.4–28.6 14.4–16.1 15.8–16.9 4.9–6.1 7.6–7.7

spiculus unsegmented, thick and curved dorsally; second set of smaller spines on distal tip of spiculus (Fig. 31); in females, melanophores directly anterior to the anal fin arranged into a wide U extending anteriorly, on either side of the genital papilla, to level of anus (see Meisner & Collette, 1998). Description. (Table 12) Slender-bodied (BDP1 <14% SL); vertebrae 39–42 [40]; precaudal vertebrae 21–23 [22] (Table 2); caudal vertebrae 17–19 [18] (Table 4); predorsal scales 25–34; lower jaw elongate (5.5–8.1 times in SL); upper jaw longer than wide (UJL/UJW 1.4–1.7); branchiostegal rays 10–11; pectoral-fin rays 11–12; second anal-fin ray in males with 6–7 segments proximal to paired spines, segment three elongate; spiculus unsegmented, thick, curved dorsally; second set of smaller spines on distal tip of spiculus; both sets of spines with a cap of bone at distal tip (Fig. 36);

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233

Figure 30. Dermogenys bispina. A, holotype, ZRC 40391, male, 24.9 mm SL; Danum Valley; Sg. Malua, Sabah. (Photograph by Ruth Gibbons).

genital papilla in females elongate, pigmented; analfin rays 15–17 [15] (Table 5); dorsal-fin origin over anal-fin ray 6 or 7 [7]; dorsal-fin rays 9–10 [11] (Table 6); caudal fin oval; threadlike sperm nuclei (Downing & Burns, 1995); type II form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 30) Background colour tan or brown; thin diffuse midlateral stripe extending from opercle to caudal fin; diffuse spot anterior to pectoral fin; males with black pigment on posterior dorsal-fin rays; some males with black pigment on lateral surfaces of elongate genital papilla; in females, melanophores directly anterior to the anal fin arranged into a wide U extending anteriorly on either side of the genital papilla to the level of anus (see Meisner & Collette, 1998). Distribution. Sabah, Indonesia; Busuanga, Philippines. Ecology. As is true of several other species of Dermogenys, D. bispina is most abundant in slightly brackish water but moves upstream into turbid freshwater (Inger & Chin, 1962). In eight stations where D. bispina was collected, a total of 53 other species were recorded (summarized from Inger’s field notes at FMNH). Four species were collected at four of these eight stations: two minnows, Nematabramis everetti and Puntius sealei, and two catfishes, Hemibagrus nemurus (Bagridae), and Clarias teijsmanni (Clariidae). Seven species were taken at three of the eight stations, 16 at two of the eight. Dermogenys bispina lives in small streams or in large rivers, where they may be seen swimming at the surface near the banks (Inger & Chin, 1962). Material examined. SABAH: 175 specimens from 17 collections (20.2–50.1 mm SL). Holotype ZRC 40391 (male, 24.9) Danum Valley, Sg.

Malua; Tan and Goh; Oct 1996. Measurements listed in Table 12. Paratypes: ZRC (from 40391) (2 females, 35.5–40.4); collected with holotype. ZRC 37643 (3 females, 35.2– 43.7; 4 males, 23.8–26.2); Kinbatagan basin at Kg. Batu Pateh; Lim et al.; Apr 1994. ZSM 27562 (3 females, 41.7–46.3, 3 males, 26.3–31.2); outskirts of Sandakan, Kettner, Krummenacher, and Witte; 1988. FMNH 51693 (13 females, 23.7–48.6, 10 males, 22.5–30.7, 4 undet.); East Coast Residency, Kinabatangan Dist.; tributary of the Little Kretam River (Sungei Gana) just above Nipa belt; Inger RFI 103 113; 12 May 1950. FMNH 44898 (3 females, 35.8 mm–30.3, 3 males, 28.0–29.5); Sandakan Dist., mile 8, North Rd; Fisheries Department; 1950. FMNH 44899 (1 female, 35.0, 1 male, 23.9); Kinabatangan Dist., Lake Bilit; Tubb; 1949. FMNH 68412 (3 females, 28.0–39.3, 1 male, 25.3); Kinabatangan Dist.; Dermakot Camp, unnamed stream crossing railroad trace; Inger and Chin RFI 1141–9; 25 Apr 1956. FMNH 68413 (1 male, 24.4); Kinabatangan Dist., Dermakot Camp; Chin RFI 1242–3; 30 Apr 1950. FMNH 68415 (1 male, 29.2); Tawau Dist., Kalabakan, Sungei Marikut; Inger RFI 1728–34; 16 Jun 1956. FMNH 101114 (2 females, 25.3–44.8, 3 males, 25.4–30.1); Sandakan Dist., Tenosa, near SKAN; Tubb; 1947. FMNH 51696 (3 females, 30.7–49.2, 1 male, 21.1); Kinabatangan Dist., forest trib. of Kretam Kechil; Inger RFI 122; 1950. USNM 345500 (11 females, 27.7–46.2, 7 males, 20.2– 28.5, 3 undet., of which 2 males have been cleared and counterstained); East Coast Residency, Kinabatangan Dist., Pinang River, trib. of Little Kretam River; Inger and Jones RFI 64–72; 10 May 1950. FMNH 68414 (2 males, 27.4–28.4); Tawau Dist., Palau Sebatik; Inger RFI 1420; 28 May 1956. CAS 133488 (10 females, 28.0–50.1, 4 males, 28.9–34.0, of which 2 females, 1 male have been cleared and counterstained); Sandakan Dist., Gum-Gum River; Herre; 11 Apr 1938. CAS 133611 (69, females, 29.7–49.5, males, 21.9–31.9, of

234

A. D. MEISNER Table 12. Morphometrics of Dermogenys bispina. (A) male holotype (ZRC 40391); (B) 12 female paratypes; and (C) 11 male paratypes. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9.

SL/LJL UJL/UJW SN-P1 SN-P2 HDL BDP1 BDP2 ORBL INTORBL

A

B

C

24.9 6.9 1.6 35.3 59.8 30.5 10.4 9.6 5.9 6.0

35.2–49.8 6.1–8.1 1.4–1.7 32.7–35.2 60.7–65.2 27.5–31.0 9.3–13.5 9.6–13.5 4.2–5.8 4.6–6.4

23.8–31.9 5.5–6.8 1.4–1.6 33.6–37.3 57.8–64.6 28.5–34.2 10.2–12.2 10.3–12.2 4.9–6.4 5.7–7.1

stream between Telupid and 42 km before intersection with road to Beluran; Kettner, Krummenacher, and Witte; 11 Mar 1988. OTHER MATERIAL EXAMINED

SABAH: 23 specimens (26.4–48.4 mm SL).

Figure 31. Diagrammatic representation of the modified anal-fin (andropodium) of Dermogenys bispina, paratype, USNM 345500, 28.5 mm SL. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. (Drawing by Keiko Hiratsuka Moore).

which 2 females, 2 males have been cleared and counterstained); Sandakan Bay drainage, Kabili R.; Herre; 29 Jan 1937. ZSM 27540 (1 female, 30.3, 2 males, 26.2, 1 undet.); road from Kota Kinbalu to Sandakan, larger

from

nine

collections

FMNH 51691 (7 females, 33.9–48.4); East Coast Residency, Kinabatangan Dist., fork of east and west Gaja, in west fork, trib. of Kretam Kechil R.; Inger RFI 170–174; 22 May 1950. FMNH 51695 (3 undet.); Kinabatangan Dist., clear water trib. of Kretam Kechil; Inger RFI 115; 12 May 1950. FMNH 51694 (2 females, 35.4–45.0, 1 undet.); Kinabatangan Dist., near mouth of Kretam Kechil R.; Inger RFI 56–57, 9 May 1950. FMNH 51699 (1 female, 45.8, 1 undet.); Kinbatangan Dist., mouth of Pinang River, a trib. of Little Kretam River, Inger RFI 197–198, 26 May 1950. FMNH 101113 (1 female, 43.2); Sandakan Dist., mile 5 North Road; Tubb, 1947. FMNH 51697 (2 females, 26.4–37.7); Sandakan Dist., Sapagaya Forest Reserve, trib. of Sapagaya River; Inger RFI 478–489; 20 Jul 1950. FMNH 44897 (1 female, 27.4); Kinabatangan Dist., Mintak; Tubb; 1949. FMNH 51698 (1 female, 43.5); East Coast Residency, Kinabatangan Dist., Little Kretam River, mouth of Ayer Terjun; Inger RFI 73–75; 10 May 1950. FMNH 44896 (3 females, 29.9–47.3); Sandakan Dist., mile 5 North Road and/or Tenosa, near SKAN, Tubb, 1947. BUSUANGA ISLAND: 69 specimens from one collection (29.2–55.6 mm SL). USNM 138667 (69, females, 29.2–55.6, males, 31.8– 36.7); Paugauron River, Port Caltour; Albatross; 1908.

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235

Figure 32. Dermogenys bruneiensis sp. nov. A, holotype, USNM 320493, male, 34.5 mm SL; Dolakin River, Brunei; B, allotype (=paratype), USNM 363184, 55.0 mm SL.

DERMOGENYS BRUNEIENSIS SP. NOV. (Figures 32, 33; Tables 2, 4, 5, 6, 13)

Differential diagnosis. Dermogenys bruneiensis is distinguished from other Dermogenys species by the thin, elongate, sickle-shaped spiculus that is segmented distally to tip; distal tip of spiculus oriented ventrally (Fig. 33). Description. (Table 13) Slender-bodied (BDP1 <12% SL); vertebrae 39–42 [39]; precaudal vertebrae 23–25 [23] (Table 2); caudal vertebrae 16–17 [16] (Table 4); 32–38 predorsal scales; lower jaw elongate (5.0–6.5 times in SL); upper jaw longer than wide (UJL/UJW 1.3–1.5); pectoral fin rays 10–11; dorsal-fin origin over anal-fin ray 6 or 7 [7]; anal-fin rays 14–16 [14] (Table 5); dorsal-fin rays 10–11 [10] (Table 6); genital papilla in females elongate, pigmented; melanophores anterior to anal fin in females arranged into a diffuse, narrow heart-shaped spot extending anteriorly to genital papilla; spiculus thin, elongate, sickle-shaped, segmented distally to tip; distal tip of spiculus oriented ventrally (Fig. 31); caudal fin oval; thread-like sperm nuclei (Downing & Burns, 1995); type II form of halfbeak viviparity. Colour in alcohol. (Fig. 32) Background colour white or olive; thin midlateral stripe from opercle to base of caudal fin, more distinct posteriorly; scattered melanophores over dorsal surface; males with black pigment on posterior anal-fin rays.

Habitat. Dermogenys bruneiensis has been collected in coastal, black waters throughout Brunei (summarized from L. R. Parenti’s field notes at USNM). Streams where specimens of D. bruneiensis were collected either ran through Nipah or dipterocarp vegetation. In seven stations where D. bruneiensis was collected, a total of 27 species were collected. Species collected at more than one of the stations include: the phallostethids Neostethus borneensis and N. bicornis; the gobies Stimatogobius borneensis, Brachygobius doriae, Mugilogobius rambaiae and Hemigobius sp.; the puffer Tetraodon leirus; and the eleotrids Bostryhus sp. and Ophiocara porocephala. A species of Zenarchopterus was also collected at one station. Etymology. Named after the country of Brunei where this species is endemic. Distribution. This species appears to be restricted to Brunei (northwestern Borneo). The eastern limit of its range is the Temburong District and the western limits are the Belait River on the western border. Another species, D. collettei, has also been collected at one locality in the Belait River district but the two have not, to date, been collected from the same locality. Material examined. BRUNEI. Six specimens from two collections (25.2–55.3 mm SL). Holotype: USNM 320493; male (34.3); Dolhakim River, trib. of the Brunei River near Bandar Seri Begawan;

236

A. D. MEISNER

Parenti, Wong et al.; 17 Aug 1991. Measurements are listed in Table 13. Allotype (=Paratype): USNM 363184; female (55.3); Dolhakim River, trib. of the Brunei River near Bandar Seri Begawan; Parenti, Wong et al.; 19 Aug 1991. Paratypes: USNM 320491 (1 male, 28.4, 1 undet.); Dolhakim River, trib. of the Brunei River near Bandar Seri Begawan; Parenti, Wong et al.; 19 Aug 1991. USNM 363185 (2 females, 25.2–41.6); Dolhakim River, trib. of the Brunei River near Bandar Seri Begawan; Parenti, Wong et al.; 17 Aug 1991. DERMOGENYS ROBERTSI SP. NOV. (Figures 34, 35; Tables 2, 4, 5, 6, 14)

Dermogenys viviparus not of Peters 1865. Herre, 1944: 48 (collection report). Herre, 1953: 160 (collection report). Meisner & Burns, 1997b: 298 (identified as Dermogenys viviparus populations with the Type I/ II form of halfbeak viviparity).

Figure 33. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Dermogenys bruneiensis sp. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. USNM 363185 (34.3 mm).

Table 13. Morphometrics of Dermogenys bruneiensis sp. nov. (A) male holotype (USNM 320493); (B) two female paratypes and (C) two male paratypes. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

34.3 5.0 1.4 36.4 63.0 40.8 31.8 11.4 15.8–16.9 6.1 7.0

41.6–55.3 5.1–6.5 1.3–1.5 33.1–34.4 62.0–62.7 39.8–40.1 28.4–29.3 10.5–12.0 10.5–12.0 5.1–5.3 6.3–6.9

C 28.4–34.3 5.0–5.6 1.4–1.5 33.8–36.4 59.9–63.0 39.8–40.8 27.8–31.8 10.9–11.4 10.9–11.4 4.9–6.1 6.3–8.8

Differential diagnosis. The anal fin of D. robertsi is similar to D. palawanensis. In these two species, the spiculus is thick, not clearly segmented with distal tip oriented ventrally (Fig. 35). Dermogenys robertsi is distinguished from D. palawanensis by the combination of melanophores anterior to anal fin in females arranged into a wide, round spot extending anteriorly just to level of genital papilla, darker anteriorly, diffuse towards anal-fin origin; genital papilla in females inconspicuous. Description. (Table 14) Slender-bodied (BDP1 <14% SL); vertebrae 38–41 [40]; precaudal vertebrae 22–23 [23] (Table 2); caudal vertebrae 16–18 [17] (Table 5); predorsal scales 30–37; lower jaw elongate (5.6–9.6 times in SL); upper jaw longer than wide (UJL/UJW 1.1–1.7); branchiostegal rays 11; pectoral-fin rays 10– 11; second anal fin of males with 8–10 segments proximal to paired spines, third or fourth segment elongate; spiculus thick, unsegmented, distal tip oriented ventrally (Fig. 33); cap of bone on spines; anal-fin rays 14–15 [14] (Table 5); dorsal-fin origin over anal-fin ray 6 or 7 [6]; dorsal-fin rays 9–11 [9] (Table 6); caudal fin oval; threadlike sperm nuclei (Downing & Burns, 1995); type II form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 34) Freshly fixed material olive/ whitish, older material tan or brown; diffuse midlateral stripe along posterior half of body, scattered melanophores laterally; diffuse oval spot anterior to pectoral fin. Etymology. Named for Tyson Roberts, Southeast Asian fish expert who provided specimens from his private collection for this study.

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237

Figure 34. Dermogenys robertsi sp. nov. A, holotype, CAS 137633, male, 26.0 mm SL; Wayan River, Busuanga; B, allotype (=paratype), CAS 69838, 33.1 mm SL; collected with holotype.

Distribution. Culion and Busuanga, Philippines and Kalimantan Timur (Borneo), Indonesia.

KALIMANTAN TIMUR. Two specimens from one collection (29.4–30.1 mm SL).

Material examined. BUSUANGA: 46 specimens from four collections (20.8–56.6 mm SL).

CMK 9550 (2 males, 29.4–30.1); Timur, Sungei Sebuku basin, Sungei Tikung at Apas; Kottelat; 16 Feb 1993.

Holotype: CAS 137633 male (26.0); Palawan Province, Busuanga Island, Wayan River at Bario San Nicolas; Herre; 21 Jun 1940. Morphometric measurements listed in Table 14.

Other material examined. CULION. Ten specimens from two collections (33.5–44.5 mm SL).

Paratypes (only lots containing mature males are designated paratypes). Allotype (=Paratype): CAS 169838 female (33.1); collected with holotype. CAS 169839 (18 females, 27.5–47.6, 1 male, 28.7) collected with holotype. CAS 137635 (8 females, 37.6– 56.6, 3 males, 29.3–33.4, of which 2 females have been cleared and counterstained); Herre; 28 Aug 1940. CAS 55259 (5 females, 22.5–28.2, 5 males, 20.8–25.5, 4 undet., of which 2 males and 2 females have been cleared and counterstained); Wayan Creek, Vicinity of San Nichols; Vindum; 29 Jun 1984. CULION. 16 specimens (23.1–54.8 mm SL).

from

two

collections

CAS 128481 (13 females, 31.5–50.7, 1 male, 31.1); Palawan Province, Baldat; Herre; 28 Apr 1931. FMNH 50935 (2 males, 23.1–31.0); Calamaines, San Pedro; Hoogstraal; 30 Mar 1947.

FMNH 47292 (5 females, 33.5–44.5); Baldat; Herre; 28 Apr 1931. UMMZ 100438 (5 females, 46.0–54.8); Herre; 28 Apr 1931. KALIMANTAN TIMUR. Five specimens from two collections (27.0–38.3 mm SL). CMK 9526 (2 females, 32.9–38.3); Timur, Sungei Sebuku Basin, Sungei Tulit, west shore and sand and gravel bank immediately upriver from Sekikilan; Kottelat and McKee; 12 Feb 1993. CMK 9503 (3 females, 27.0–32.4); Timur, Sungei Sebuku Basin, Sungei Tikung, upriver of Kekayap, 3°57′51′′N, 116°58′8′′E; Kottelat; 11 Feb 1993. DERMOGENYS PALAWANENSIS SP. NOV. (Figures 35, 36; Tables 2, 4, 5, 6, 15)

Differential diagnosis. The anal fin of D. palawanensis is similar to D. robertsi (Fig. 35). Dermogenys palawanensis is distinguished from D. robertsi by the combination of melanophores anterior to anal fin in females

238

A. D. MEISNER

threadlike sperm nuclei; type II form of halfbeak viviparity. Colour in alcohol. (Fig. 35) Background colour olive; concentration of melanophores on dorsal and lateral surfaces; thin midlateral stripe posteriorly; distinct oval spot anterior to pectoral fin; fins dusky. Etymology. Named for the southwestern Philippine island of Palawan, where this species is endemic. Distribution. Palawan, Philippines. Material examined. Fifty-two specimens from six collections (23.2–48.6 mm SL). Holotype: ZRC 46170 male (33.2); Estrella falls near Narra, trib. of Malatgao River, about 80–100 km south of Puerto Princesa, Margraf, 29 Sep 1994. Measurements listed in Table 15. Allotype (=Paratype): ZRC 46171 female (44.0); collected with holotype.

Figure 35. Diagrammatic representation of the anterior modified anal-fin rays of the andropodium in two species of Dermogenys robertsi and D. palawanensis spp. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. USNM 138673 (34.0 mm).

forming a broad V with a wide base, extending onto lateral surface, directly adjacent to genital papilla, extending anteriorly past level of anus; females with an elongate, pigmented genital papilla. Description. (Table 15) Slender-bodied (BDP1 <15% SL); vertebrae 40–43 [42]; precaudal vertebrae 21–23 [23] (Table 2); caudal vertebrae 18–20 [19] (Table 4); predorsal scales 29–33; lower jaw short (4.0–16.3 times in SL); upper jaw longer than wide (UJL/UJW 1.3–1.7); branchiostegal rays 11–13; pectoral-fin rays 10–12; second anal-fin ray in males with 7–8 segments proximal to paired spines, third or fourth segment elongate; spines with cap of bone; spiculus thick, few segments, distal tip oriented ventral (Fig. 33); genital papilla in females elongate, pigmented; anal-fin rays 15–17 [16] (Table 5); dorsal-fin origin over anal-fin ray 5 or 6 [5]; dorsal-fin rays 10–11 [11] (Table 6); caudal fin oval;

Paratypes: CMK 11972 (10 females, 23.3–48.6, 11 males, 23.2–36.3, 2 undet.); collected with holotype. USNM 138672 (7 females, 37.1–48.6, 1 male, 31.4); Stream near village at Chase Head, Endeavor Stream; Albatross; 22 Dec 1908. USNM 138673 (8 females, 25.7–44.9, 9 males, 25.6–32.1, of which 1 female and 2 males have been cleared and counterstained); River at Nakoda Bay; Albatross; 31 Dec 1908. USNM 150809 (1 male, 30.9); River at Nakoda Bay; Albatross; 31 Dec 1908. USNM 138668 (1 male, 29.2); Ulugan Bay; Albatross; 1945. DERMOGENYS COLLETTEI SP. NOV. (Figures 37, 38; Tables 2, 4, 5, 6, 16)

Dermogenys pusillus not of Kuhl & van Hasselt in van Hasselt, 1823. Herre, 1944: 45 (collection report). Dermogenys pusilla not of Kuhl & van Hasselt in van Hasselt, 1823. Meisner & Burns, 1997b: 298 (identified as Dermogenys pusilla populations from Binton, Singapore, and Johor with the Type I form of halfbeak viviparity). Differential diagnosis. The anal fin of D. collettei is most similar to that of D. pusilla. It is distinguished from D. pusilla by a combinations of: thicker spiculus divided into 4–6 segments (vs. <3); proximal segments 4–6 of second anal-fin ray in males noticeably thicker along anterior edge (Fig. 38). Description. (Table 16) Slender-bodied (BDP1 <14% SL); vertebrae 39–42 [39]; precaudal vertebrae 22–24 [23] (Table 2); caudal vertebrae 15–17 [16] (Table 4); predorsal scales 34–39; lower jaw elongate (5.0–8.8 times in SL); upper jaw longer than wide (UJL/UJW

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239

Table 14. Morphometrics of Dermogenys robertsi sp. nov. (A) male holotype (CAS 137633); (B) female allotype CAS 169838 (=paratype); (C) 18 female paratypes; and (D) 3 male paratypes. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentage of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

D

26.0 9.6 1.1 35.0 61.2 40.8 31.5 11.2 11.2 5.8 6.9

33.1 — 1.4 34.4 63.7 40.8 29.9 11.5 11.5 5.4 5.7

33.5–57.1 6.2–7.7∗ 1.3–1.7 32.3–35.9 59.7–64.0 37.6–43.0 28.5–31.8 11.3–14.0 11.3–14.0 5.2–6.0 6.8–7.8

29.4–33.4 5.6∗∗ 1.5–1.6 35.5–37.4 60.5–66.5 38.3–40.5 30.8–31.8 12.3–13.6 12.3–13.6 6.7–6.9 7.8–8.1

∗ Proportion based on three specimens. ∗∗ Proportion based on one specimen.

Figure 36. Dermogenys palawanensis sp. nov. A, holotype, ZRC 46170, male, 33.2 mm SL; Tributary of the Malatgao River, Palawan; B, allotype (=paratype) ZRC 46171, 44.0 mm SL, collected with the holotype.

1.2–1.7); branchiostegal rays 10–12; pectoral fin rays 10–11; second anal-fin ray in males with 8–10 segments proximal to paired spines, third segment generally elongate; spiculus divided into 4–6 segments; proximal segments 4–6 of the second anal-fin ray in males

noticeably thicker along anterior edge (Fig. 38); analfin rays 14–16 [14] (Table 5); dorsal-fin origin over anal-fin ray 6–8 [6]; dorsal-fin rays 9–11 [10] (Table 6); caudal fin oval; type I form of halfbeak viviparity (Meisner & Burns, 1997b).

240

A. D. MEISNER Table 15. Morphometrics of Dermogenys palawanesis. sp. nov. (A) male holotype (ZRC 46170); (B) allotype (=paratype) (ZRC 46171); (C) 16 female paratypes; and (D) eight male paratypes. Standard length in mm; measurements 1 and 2 proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

D

33.2 13.8 1.5 36.1 63.0 40.0 32.8 12.7 12.7 6.0 5.7

44.0 16.3 1.3 33.9 63.4 39.3 30.9 11.6 11.6 5.3 5.6

31.3–48.6 4.0–10.0 1.3–1.6 32.4–34.4 60.2–64.5 38.1–41.5 29.1–38.3 11.1–14.2 11.1–14.2 4.7–6.8 5.7–6.9

30.8–36.1 8.3–10.1 1.3–1.7 33.6–36.3 56.7–62.1 38.2–43.0 30.3–33.0 11.7–13.5 11.7–13.5 4.8–5.5 6.2–7.5

Figure 37. Dermogenys collettei sp. nov. A, holotype, ZRC 37790, male, 26.0 mm SL; Kuching, Sarawak; B, allotype (=paratype), ZRC 46161, 33.1 mm SL; collected with the holotype.

Colour in alcohol. (Fig. 37) The overall body pigmentation varies in intensity depending on locality. In Sumatra, Sarawak, Singapore, and Johor populations background colour is white or olive; concentration of melanophores on dorsal half of body giving a dusky appearance; black pigment on dorsal portion of caudal peduncle and along lower jaw; diffuse midlateral stripe from opercle to caudal fin, prominent posteriorly; small

concentration of melanophores anterior to pectoral fin; large melanophores scattered ventrally in throat region. Brunei populations are pigmented similarly but with greater intensity. In males, black pigment present on distal tips of posterior dorsal-fin rays and base and distal tips of pelvic-fin rays; in females, melanophores anterior to anal fin arranged into a narrow oval or triangular band.

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241

Material examined. SARAWAK: 64 specimens from 11 collections (19.0–49.3 mm SL). Holotype: ZRC 37790 (male, 31.4 mm) Sarawak, Kuching; Lim; 30 Jun 1994. Measurements listed in Table 16. Allotype (=Paratype): ZRC 46161 (female 42.8 mm); collected with holotype. Paratypes: ZRC 46162 (3 females, 35.8–49.3, 6 males, 30.2–32.3); Kuching; Lim; 30 Jun 1994. ZRC 25999– 26004 (3 females, 34.5–46.5, 3 males, 22.3–29.1); 7 km, Serian-Balai Ringam Road, Sungei Merian Bedup at Kampung Merian Bedup; Ng and Lim; 2 Jul 1992. ZRC 37955 (6 females, 20.6–37.5, 3 males, 19.0–27.7); Sungei Engkurah, km 8.5, Tebakang Mongkos road after branching from Serian-Tebedu road; Kottelat et al.; 16 Jun 1994. ZRC 37899 (1 female, 29.2, 1 male, 30.2); Sungei Ngeli, ca. km 6 Simunjan Balai Ringin road; Kottelat et al., 11 Jun 1994. ZRC 37882 (2 females, 19.3–42.9, 1 male, 29.1); Bau-Matang river, 16 km and Bau; Kottelat et al.; 9 Jun 1994. USNM 320501 (2 females, 40.8–43.5, 1 male, 30.0); Kuching, Sarawak River, north shore at Kampung Tupung; Parenti; 13 Aug 1991. FMNH 45867 (14 females, 23.2–41.8, 2 males, 29.3–29.6, of which 2 males, 1 female have been cleared and counterstained); Samarahan, Borneo; Harrison; 5 Jan 1949. FMNH 63037 (7 females, 33.6– 46.4, 1 male, 28.7); Niah, Sungei Soah; Medway; 17 Aug 1959. CMK 5684 (1 female, 42.8, 4 males, 32.7–33.9); Sungei Sebubut near Matang; Bader; 25 Aug 1986. MCZ 57957 (1 male, 29.3) Kuala Samunsam; Mittermier; 21 Apr 1979. Figure 38. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Dermogenys collettei sp. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. ZRC 37736 (31.0 mm).

Ecology. Specimens from Brunei were collected in mangroves near the mouth of the Belait River in black water with a recorded pH of 3.8 (Summarized from the field notes of L. R. Parenti at USNM). Other species collected with these specimens were: Megalops cyprinoides, Rasbora c.f. pauciperforata, Arius maculatus, Parambasis wolffii, Betta akarensis, Butis sp., Stigmatogobius brocki and Tetraodon nigroviridis (Parenti & Meisner, 1995). Etymology. Named for Bruce B. Collette, who initially brought the problem of halfbeak taxonomy to my attention. Distribution. Sumatra, Sarawak, and southern Malay Peninsula, Malaysia; Kalimantan (northwestern), Indonesia; Singapore; and Brunei.

SUMATRA. 52 specimens (16.9–48.9 mm SL).

from

five

collections

AMNH 9584 (2 females, 35.6–38.6); Bagan Si Api Api; Herbst; 1912. CMK 4505 (4 females, 40.6–48.9, 4 males, 22.7–27.2); Chinese temple near Perdagangan, Sumatera Barat; Bianco and Kottelat; 25 Nov 1984. ZRC 31404–31406 (2 females, 39.5–43.5, 1 male, 32.6); Pulau Lingga; Lim; 1993. ZRC 32848–32857 (5 females, 16.9–46.3, 6 males, 18.8–30.8); Riau archipelago, Pulau Binton; Tan et al.; 12 Jun 1993. ZRC 33824–33851 (1 female, 23.3, 8 males, 23.8–30.0, 19 undet., of which 2 males have been males cleared and counterstained); Riau archipelago, Pulau Binton; Sivasothi et al.; 13 May 1993. SINGAPORE: 66 specimens from two collections (17.8–44.4 mm SL). ZRC 37736 (24 females, 24.4–44.4, 24 males, 17.8–30.9, of which 2 males have been cleared and counterstained); Lim and Tan; 16 Jun 1994. CAS 133945 (11 females, 31.7–43.8, 7 males, 26.1–30.6, of which 2 males have been cleared and counterstained); Jurong; Herre; 8 May 1937. PENINSULAR MALAYSIA: 19 specimens from three collections (20.2–50.9 mm SL).

242

A. D. MEISNER Table 16. Morphometrics of Dermogenys collettei sp. nov. (A) male holotype (ZRC 37790); (B) female allotype (=paratype); (C) 18 females; and (D) 18 males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

D

31.4 6.8 1.7 37.3 65.0 39.5 32.5 12.7 12.7 5.7 6.7

42.8 8.7 1.7 35.5 65.5 35.5 31.1 12.6 12.6 5.1 6.8

30.5–49.3 6.1–8.8 1.2–1.6 32.6–35.7 58.7–64.4 35.6–39.8 26.9–31.1 9.9–13.3 9.9–13.3 5.0–6.2 5.7–8.1

23.9–32.3 5.0–6.6 1.2–1.6 32.6–38.4 59.0–75.4 36.8–40.7 28.0–34.1 11.3–14.2 11.3–14.2 5.6–6.8 4.2–7.8

CAS 139413 (9 females, 28.3–43.3, 6 males, 20.2–27.2, 5 undet., of which 2 males have been cleared and counterstained); Johor State, Kota Tingga, Malaysia; Herre; 17 Oct 1940. ZSM 26957 (1 female, 36.7, 1 male, 27.8); Geck; 1988. CAS 53983 (2 females, 42.1–50.9); Johor State, Mawai Dist., 50–60 miles north of Singapore; Herre; 22 Mar 1934. BRUNEI: 11 specimens (26.8–44.1 mm SL).

from

one

collection

USNM 332450 (6 females, 26.8–44.1, 5 males, 27.2– 28.6, of which 1 male, 1 female have been cleared and counterstained); Daerah Belait; Sungei Melinu, tributary stream of Sungei Belait, third tributary stream on right side from Kuala Belait; Parenti and Downing et al.; 27 Jul 1993. WESTERN KALIMANTAN: 9 specimens from one collection (26.2–36.8 mm SL). CMK 6638 (1 female, 36.8, 8 males, 26.2–29.7, of which 1 male has been cleared and counterstained); Nyarungkup, about 10 km north of Singkawang; Kottelat et al.; 20 Apr 1990. GENUS NOMORHAMPHUS WEBER & DE BEAUFORT, 1922 (Figures 39, 40; Tables 1, 18–21)

Nomorhamphus Weber & de Beaufort, 1922: 141 (type species: Nomorhamphus celebensis Weber & de Beaufort, 1922 by subsequent designation of Jordan, 1923: 776). Mohr, 1936b: 55–58 (taxonomy). Dermogenys (Rhamphodermogenys) Fowler & Bean, 1922: 15 as a subgenus of Dermogenys, not of Kuhl & van Hasselt in van Hasselt, 1823 (type species:

Dermogenys bakeri Fowler & Bean, 1922: 15–16 by original designation and monotypy). Diagnosis. Nomorhamphus is distinguished from other genera in the Hemiramphidae by an oval lacrimal; elongate, expanded autopalatine; absence of a definite geniculus on second anal-fin ray in males; small spermatozeugmata resulting from a mode of spermatogenesis in which spermatid nuclei become arranged evenly into several layers around the periphery of the spermatocysts; ovary with well-defined ovigerous ridge; period of intraluminal gestation. Description. Sexually dimorphic, maximum known standard lengths of species ranges from 21.2–100.1 mm for females and 22.1–55.0 mm for males; body elongate; upper and lower jaws approximately equal in length or lower jaw elongate; when elongate measuring 3.7–22.2 times in SL; oral teeth conical; no teeth along extended portion of lower jaw; endopterygoid expanded; gill rakers on first arch reduced; first epibranchial bone expanded at base; third pharyngobranchial bones fused medially; lower pharyngeal tooth plates fused; branchiostegal rays 9–12; length of upper jaw variable (UJL/ UJW 0.7–2.3); nasal papilla elongate extending out of nasal fossa; nasal fossa reduced; infraorbital series consisting of only lacrimal and dermosphenotic; highset pectoral fins; pectoral-fin rays 10–13; first pleural rib on third vertebra; pelvic bones widely separated with anterior processes oriented laterally; anterior spur on dorsolateral extension of pelvic bones; pelvic fins rounded; pelvic-fin rays 6; pelvic-fin origin closer to pectoral fin than caudal fin; anal-fin rays 1–7 in males modified into an andropodium; cryptoplicae

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

243

2 Celebes Sea N Sulawesi Utara

0

Gulf of Tomini

Sulawesi Tengah

Lake Poso 2 Lake Matano Lake Mahalona

Sulawesi Selatan

Lake Towuti

4

Sulawesi Tenggara Gulf of Bone

6

Flores Sea

120

125

130

Figure 39. Distribution of Nomorhamphus on Sulawesi. (Ε) N. megarrhamphus; (Φ) N. weberi; (Μ) N. towoetti; (Χ) N. celebensis; (_) N. kolonodalensis; (Α) N. ebrardtii; (Β) N. brembachi and N. liemi. Dashed line indicates approximate boundaries.

present; physa present; genital papilla elongate in mature males, nearly reaching first anal-fin ray; analfin rays 13–19; first anal pterygiophore in males thickened; middle anal radials absent; distal anal radials ossified; anal-fin origin anterior to dorsal-fin origin; ventral process on first dorsal pterygiophore reduced or absent, anterior extension expanded; dorsal-fin rays 9–14; caudal fin truncate, without distinct lobes; fifth hypural either separate from the dorsal hypural plate along most of its length or fused along its length; three epurals; parhypural fused to lower hypural plate; uroneural and posterior neural plates reduced; principal caudal fin rays 7+8; precaudal vertebrae 20–27; caudal vertebrae 15–20; total vertebrae

36–46; viviparous; period of intraluminal gestation (Meisner & Burns, 1997b); small spermatozeugmata formed (Downing & Burns, 1995). Distribution. Species of Nomorhamphus are restricted to Sulawesi and islands of the Philippines. Nomorhamphus brembachi, N. liemi, N. hageni, N. megarrhamphus, N. weberi, N. towoetii, N. celebensis, N. ebrardtii and N. kolonodalensis are endemic to the island of Sulawesi (Fig. 39). An additional seven species, N. pinnimaculata, N. bakeri, N. pectoralis, N. vivipara, N. philippina, N. rossi, N. manifesta are distributed throughout the islands of the Philippines (Fig. 40).

244

A. D. MEISNER

Remarks. The subgeneric name, Rhamphodermogenys, was originally described for Dermogenys bakeri, a species here placed within Nomorhamphus, thus making this name available as a generic name for the latter genus. The original date of publication for Rhamphodermogenys is 28 July 1922 and that of Nomorhamphus is May 1922. Therefore, the name Nomorhamphus has priority and Rhamphodermogenys is herein considered a junior synonym.

N Luzon

15 South China Sea

NOMORHAMPHUS BAKERI (FOWLER & BEAN, 1922) COMB. NOV.

Busuanga

(Figures 41, 42; Tables 17–21)

Samar

Dermogenys bakeri Fowler & Bean, 1922: 15–16 (original description, Zamboanga, Philippines). Weed, 1933: 50 (characters, distribution). Herre, 1953: 160 (listed as a synonym of Dermogenys viviparus). Rhamphodermogenys bakeri. Mohr, 1936a: 51 (placed in synonymy with D. viviparus). Herre, 1944: 45 (placed in synonymy with D. viviparus). Herre, 1953: 160 (listed as a synonym of Dermogenys viviparus).

Panay

Leyte

Palawan

Cebu

10

Negros

Sulu Sea Mindanao

Jolo Celebes Sea

Borneo 120

125

Figure 40. Distribution of Nomorhamphus in the Philippines. (Ε) Nomorhamphus rossi; (Φ) N. manifesta; (Χ) N. pectoralis; (Β) N. vivipara; (_) N. bakeri; (Μ) N. pinnimaculata; (Α) N. philippina.

Table 17. Morphometrics of Nomorhamphus bakeri comb. nov. (A) male holotype (USNM 84275); (B) one female; and (C) one male. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

1. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

31.6 — 1.3 34.2 58.9 42.7 28.2 10.4 10.4 6.3 7.3

32.0 5.1 1.3 32.8 60.0 42.5 28.8 10.0 10.0 6.3 7.5

32.1 — 1.4 37.1 59.2 42.1 31.5 12.8 12.8 6.5 7.2

Differential diagnosis. The anal fin of Nomorhamphus bakeri is most similar to N. pectoralis, N. philippina, N. vivipara, N. pinnimaculata and N. manifesta in that the spiculus is elongate. It is distinguished from other species of Nomorhamphus by segmentation of the second anal-fin ray in males in which the fifth or sixth segment is elongate (Fig. 42). Description. (Table 17) Slender-bodied (BDP1 <13% SL); vertebrae 39–41 [40]; precaudal vertebrae 23–24 [24] (Table 18); caudal vertebrae 16–17 [16] (Table 19); 33–35 predorsal scales; lower jaw elongate (5.1 times in SL); upper jaw longer than wide (UJL/UJW 1.3–1.4); pectoral fin pointed reaching base of pelvic fin in males; pectoral fin rays 11 [11]; second anal-fin ray in males with 8–9 segments proximal to paired spines, fifth or sixth segment elongate; spiculus with 7–8 segments, terminal segment elongate, distal tip ventral, slight curvature to entire spiculus (Fig. 43); anal-fin rays 14–16 [15] (Table 20); dorsal-fin origin over anal-fin ray 6 or 7 [6]; dorsal-fin rays 10 [10] (Table 21); caudal fin truncate; fifth hypural separate from the dorsal hypural plate along most of its length. Colour in alcohol. (Fig. 41) Background colour tan or brown; thin midlateral stripe, prominent posteriorly; diffuse spot anterior to pectoral fin; black pigment on anterior dorsal-fin rays; black pigment arranged into diffuse spot on proximal portion of dorsal caudal rays, black pigment on distal tips of pelvic-fin rays. Distribution. Zamboanga, Mindanao, Philippines. Material examined. ZAMBOANGA. Five specimens from four collections (31.6 mm–38.8 mm SL).

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

245

Table 18. Frequency distribution of precaudal vertebrae in species of Nomorhamphus 20 N. N. N. N. N. N. N. N. N. N. N. N. N. N. N. N. N.

bakeri brembachi celebensis ebrardtii hageni liemi megarrhamphus pectoralis philippina towoetii vivipara weberi robertsi rossi kolonodalensis manifesta pinnimaculata

7

21

22

23

24

25

26

27

Mean

N

— — 1 18 7 1 — 1 2 — 6

— 5 4 6 4 7 — 13 — 2 7

1 8 — — — 5 — 4 — 1 9

— — — — — — 13 — — — —

2 — 2

6 — 5 2 —

1 12 9 6 —

2 3 — — — 1 8 — — — 1 1 1 1 — 1 —

— — — — — — 2 — — — — 11 — — —

— — — — — — 3 — — — — 16 — — —

—

—

23.6 22.8 21.5 21.3 21.3 22.5 25 22.2 21.0 22.3 22.2 26.5 22.1 23.0 22.4 23.1 20.6

3 16 5 24 11 14 26 18 2 3 23 28 10 13 16 10 17

10

USNM 84275 (holotype of Dermogenys bakeri; male, 31.6); USNM 138676 (1 male, 36.6). USNM 84273 (1 female, 38.8); Baker; 27 Feb 1914. USNM 138677 (1 male, 32.1, 1 female, 32.0); Albatross; 8 Oct 1909. NOMORHAMPHUS BREMBACHI VOGT, 1978a (Figures 43–45; Tables 18–22)

Nomorhamphus brembachi Vogt, 1978a: 227 (original description, Longron, Celebes [Sulawesi]). Meyer, 1985: 408 (characters, distribution). Brembach,

— — — 1 —

1991: 174–175 (anatomy, relationships, distribution). Kottelat et al., 1993: 121 (distribution). Nomorhamphus ravnaki ravnaki Brembach, 1991: 184–185 (original description, Ban Timurung, Sulawesi). Nomorhamphus ravnaki australis Brembach, 1991: 185–186 (original description, Bossolo, Sulawesi). Nomorhamphus sanussii Brembach, 1991: 187–188 (original description, Segoja, Sulawesi). Kottelat et al., 1993: 121 (distribution). Nomorhamphus sanussi. Downing & Burns, 1995: 330 (testis study).

Table 19. Frequency distribution of caudal vertebrae in species of Nomorhamphus

N. N. N. N. N. N. N. N. N. N. N. N. N. N. N. N. N.

bakeri brembachi celebensis ebrardtii hageni liemi megarrhamphus pectoralis philippina towoetii vivipara weberi robertsi rossi kolonodalensis manifesta pinnimaculata

15

16

17

18

19

20

Mean

N

— — — 2 — — — — — — 1 — — — 1

2 3 — 11 — — 6 — 3 — 3 1 — — 13 2 3

1 3 1 10 — 1 7 7 — 2 17 17 2 7 2 8 14

— 5 4 — 7 2 — 10 — 1 2 7 8 6 —

— 7 — — 3 1 — 1 — — — 1 — — —

— 1 — — — — — — — — — — — — —

—

—

—

16.3 18.0 17.8 16.3 18.3 18.0 16.5 17.7 16.0 17.3 16.9 17.3 17.8 17.5 16.1 16.8 16.8

3 19 5 23 10 4 13 18 3 3 23 26 10 13 16 10 17

—

246

A. D. MEISNER Table 20. Numbers of anal-fin rays in Nomorhamphus of species

N. N. N. N. N. N. N. N. N. N. N. N. N. N. N. N. N.

bakeri brembachi celebensis ebrardtii hageni liemi megarrhamphus pectoralis philippina towoetii vivipara weberi robertsi rossi kolonodalensis manifesta pinnimaculata

13

14

15

16

17

18

19

Mean

N

— — — — — — — — — 2 — — — — 3

1 — 2 — — — — — 2 1 1 — — 1 13 10 14

1 2 1 13 2 3 10 — 1 — 14 — 10 — —

1 10 1 10 5 1 3 4 — — 7 7 — 10 —

— 7 — — 1 — — 12 — — 1 13 — 1 —

— 1 — — — — — 2 — — — 7 — — —

— — — — — — — 1 — — — — — — —

—

—

—

—

—

15.0 16.4 14.8 15.4 15.9 15.3 15.2 17.1 14.3 13.3 15.4 17.0 10.0 15.0 13.6 10.0 13.9

3 20 4 23 8 4 13 19 3 3 23 27 10 12 16 10 16

2

Nomorhamphus australe. Kottelat, 1992: 271 (taxonomy, emended spelling). Kottelat et al., 1993: 120– 121 (distribution). Nomorhamphus ravnaki. Kottelat et al., 1993: 121 (distribution). Downing & Burns, 1995: 330 (testis study).

distal 1/4 slopes ventrally; spiculus thick, short, not segmented clearly (Fig. 45); anal-fin rays 15–18 [16] (Table 20); dorsal-fin origin over anal-fin ray 6 or 7 [6]; dorsal-fin rays 11–14 [13] (Table 21); caudal fin truncate; fifth hypural fused to dorsal hypural plate; fleshy mandibular appendage generally present in males, elongate and pigmented in some mature males (Fig. 44).

Differential diagnosis. Two species of Nomorhamphus (N. brembachi and N. liemi ) that live in the Maros region of Sulawesi Selatan (southwestern arm of Sulawesi) are characterized by a second anal-fin ray in males with 10 segments proximal to paired spines, length of segments approximately equal, curved at approximately 3/4 its length so that the distal 1/4 slopes ventrally; short, laterally expanded spiculus, not clearly segmented (Fig. 45). Nomorhamphus brembachi is distinguished from Nomorhamphus liemi by the combination of: fin pigmentation (see Colour in alcohol); and elongate, pigmented mandibular appendage in many large males (Fig. 44).

Colour in alcohol. (Figs. 43,44) Background colour tan or olive with a diffuse midlateral stripe; females with black pigment on anterior and posterior anal-fin rays and along posterior dorsal-fin rays; reduced mandibular appendage; some males with an elongate, black pigmented mandibular appendage varying in length and black pigment on distal half of unmodified analfin rays, distal half of dorsal- and pelvic-fin rays; caudal fin dusky.

Description. (Table 22) Deep-bodied (BDP1 >14% SL); vertebrae 39–43 [41]; precaudal vertebrae 22–24 [23] (Table 18); caudal vertebrae 16–20 [18] (Table 19); predorsal scales 44–46; length of lower jaw reduced, upper and lower jaws approximately the same length; upper jaw slightly wider than long (UJL/UJW 0.86– 0.97); patch of conical teeth anteriorly on upper jaw; gill rakers wide with many teeth; 10–11 branchiostegal rays; pectoral fin pointed, extending past pelvic-fin origin in males; 12–13 pectoral-fin rays; second analfin ray in males with 10 segments proximal to paired spines, size of segments approximately equal; second ray is curved at approximately 3/4 its length so the

Remarks. Nomorhamphus brembachi, N. ravnaki ravnaki, N. ravnaki australis and N. sanussii had previously been distinguished by length of the fleshy mandibular appendage in males (Brembach, 1991). The mandibular appendage in the only male paratype of Nomorhamphus ravnaki is similar to some male paratypes of Nomorhamphus ravnaki australe. One male in the paratype series of N. sanussii also has an elongate appendage. Additionally, there appears to be intraspecific variation in length of this appendage (Figs 43, 44). Due to overlap in meristics and morphometrics, there are currently no characters to unambiguously diagnose each of these nominal species.

Distribution. Maros, Sulawesi Selatan, Indonesia.

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS KEY TO SPECIES OF NOMORHAMPHUS WEBER AND DE BEAUFORT, 1922

1A. Segments in second anal-fin ray in males approximately equal in length ........................................ 3 1B. Segment five or six in second anal-fin ray in males elongate, longer than other segments ..............2 2A. Spiculus short, unsegmented distally; terminal segment short, thick (Fig. 68) N. rossi (Luzon, Philippines) 2B. Spiculus elongate, with 7–8 segments; terminal segment elongate, thin (Fig. 42) .............................. ..................................................................................... N. bakeri (Zamboanga, Mindanao, Philippines) 3A. First anal pterygiophore in males thin, angled sharply anteriorly ................................................... 4 3B. First anal pterygiophore thickened, not angled anteriorly ................................................................. 5 4A. Spiculus curved slightly ventral, without a distinct sickle shape; black pigment on distal tips of dorsal-fin rays (Fig. 53) ............................. N. megarrhamphus (Lakes Towuti, Mahalona, Sulawesi) 4B. Spiculus distinctly sickle-shaped; no black pigment on dorsal-fin rays (Fig. 64) ................................. ........................................................................................................ N. weberi (Lake Matano, Sulawesi) 5A. Spiculus short (Fig. 45) .......................................................................................................................... 6 5B. Spiculus elongate (Fig. 55) .................................................................................................................. 10 6A. Second anal-fin ray in males curved at approximately 1/4 its length so that the distal 1/2 slopes ventrally; spiculus short, thick, not clearly segmented (Fig. 45) ....................................................... 7 6B. Second anal-fin ray in males straight; spiculus straight, segmented, not noticeably thickened (Fig. 47) ............................................................................................................................................................ 8 7A. Females with black pigment on anterior and posterior anal-fin rays; males with black pigment on distal half of unmodified anal-fin rays and on dorsal portion of dorsal and pelvic-fin rays; many large males with an elongate, pigmented mandibular appendage (Figs. 43,44) ............................................ ................................................................................................................... N. brembachi (SW Sulawesi) 7B. Black pigment on at least distal half of all fin rays, many pigmented along their entire length; females and males with a reduced mandibular appendage; (Fig. 51) ......... N. liemi (SW Sulawesi) 8A. Upper jaw length as long or longer than upper jaw width ................................................................. 9 8B. Upper jaw length less than upper jaw width (UJL/UJW<1.0) (Fig. 46) ................................................ ........................................................................................................ N. celebensis (Lake Poso, Sulawesi) 9A. Lower jaw not extending significantly past the length of the upper jaw (Fig. 58); females dusky, most males completely melanistic ............................................ N. towoetii (Lakes Poso, Towuti, Sulawesi) 9B. Lower jaw extending past the length of the upper jaw; males olive or whitish (preserved material); males and females with black pigment on distal tips of dorsal-, anal-, and pelvic-fin rays (Fig. 63) . ....................................................................... N. kolonodalensis (Lake Towuti, Kolonodale, Sulawesi) 10A. Spiculus with 4–6 segments; distal tip of spiculus expanded laterally, curved dorsally to contact distal tip of third anal-fin ray (Fig. 49) ..................................................... N. ebrardtii (SE Sulawesi) 10B. Spiculus with 5–10 segments, terminal segment elongate, not curved dorsally ............................ 11 11A. Spiculus offset dorsally from the proximal portion of the second anal-fin ray, comprised of 5–6 segments (vs. 7–10), terminal segment of the spiculus, rarely curved (Fig. 55) ................................... ........................................................................................................... N. pectoralis (Luzon, Philippines) 11B. Spiculus in line with the proximal portion of the second anal-fin ray, comprising of 7–10 segments, terminal segment of the spiculus curved ventrally ........................................................................... 12 12A. Spiculus with 9–10 segments, terminal segment of the spiculus shorter than the segmented portion (Fig. 57); black pigment on distal tips of dorsal and anal fin rays ........................................................ ......................................................................................... N. philippina (Cebu, Mindanao, Philippines) 12B. Spiculus with 7–9 segments, terminal segment of the spiculus longer than the segmented portion; no black pigment on anal-fin rays ...................................................................................................... 13 13A. Segments 4–5–6 in spiculus elongate, terminal segment of spiculus elongate, slender, curved ventrally (Fig. 60) .......................................................................... N. vivipara (Luzon, Samar, Mindanao, Jolo) 13B. Segments in the spiculus approximately equal in length, terminal segment turned ventrally but shorter than in N. vivipara ................................................................................................................. 14 14A. Black pigment present on distal half of dorsal-fin rays, distal tips of pelvic-fin rays; distal tips of caudal-fin rays, and in females on distal tips of anterior anal-fin rays (Fig. 64) ................................. ................................................................................................. N. manifesta (Ilokos Norte, Philippines) 14B. Black pigment present along the first three dorsal-fin rays and distal tips of pelvic-fin rays in males; in females prominent black spot on anterior anal-fin rays (Fig. 66) ..................................................... .................................................................................................... N. pinnimaculata (Leyte, Philippines)

247

248

A. D. MEISNER Table 21. Frequency distribution of dorsal-fin rays in species of Nomorhamphus

N. N. N. N. N. N. N. N. N. N. N. N. N. N. N. N. N.

bakeri brembachi celebensis ebrardtii hageni liemi megarrhamphus pectoralis philippina towoetii vivipara weberi robertsi rossi kolonodalensis manifesta pinnimaculata

9

10

11

12

13

14

Mean

N

— — — — — — — — — — 3 — — — —

3 — — — — — 10 — — — 10 2 1 1 8

— 2 — — — — — — — — — — — — —

2

— 6 4 21 4 3 — 11 3 2 1 4 1 6 — 1 2

— 11 — — 6 — — — — 1 — — — — —

—

— 1 1 2 — 1 3 8 — — 9 20 9 5 8 9 12

—

—

10.0 12.7 11.8 11.9 12.6 11.8 10.2 11.6 12.0 12.3 10.3 11.1 11.0 11.4 10.5 11.1 11.0

3 20 5 23 10 4 13 19 3 3 23 26 11 12 16 10 16

Figure 41. Nomorhamphus bakeri comb. nov. USNM 138676, mle, 63.1 mm SL from Zamboanga, Mindanao, Philippines.

Material examined. SULAWESI SELATAN: 93 specimens from 11 collections (24.0 mm–82.7 mm SL). ZMH 7165 (radiograph of holotype of N. brembachi ); stream near village of Longron; Vogt; 1978. ZMH 7166 (paratypes of N. brembachi, 1 male, 38.8, 1 female, 36.7); stream near village of Longron; Vogt; 1978. ZMH 7161 (radiograph of holotype of Nomorhamphus ravnaki australis, female); mountain stream near Bossolo, highlands of Maros; Vogt; 1979. ZMH 7163 (paratypes of Nomorhamphus ravnaki australis, 13 females, 28.9– 51.9, 9 males, 42.7–53.4, of which 1 female, 1 male have been cleared and counterstained); mountain stream near Bossolo, highlands of Maros; Vogt; 1979. ZMH 7158 (radiograph of paratype of Nomorhamphus

ravnaki ravnaki, female); mountain stream near Ban Timurung, highlands of Maros; Brembach; 1978. ZMH 7159 (radiograph of holotype of N. ravnaki ravnaki, 1 male, 55.0); mountain stream near Ban Timurung, highlands of Maros; Brembach; 1978. ZMH 7160 (paratypes of N. ravnaki ravnaki, 5 females, 57.0–77.6, 4 males, 34.2–39.0, 4 undet.); mountain stream near Ban Timurung, highlands of Maros; Brembach; 1978. ZMH 7615 (radiograph of holotype of Nomorhamphus sanussii, male); mountain stream near Sepoja; Brembach; 1978. ZMH 7616 (paratypes of N. sanussii, 9 females, 43.8–71.5, 11 males, 41.5–48.6, of which 1 female, 1 male have been cleared and counterstained); mountain stream near Sepoja; Brembach; 1978. USNM 338497 (6 females, 39.5–55.3, 11 males, 24.0–40.8, 2

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141–142 (original description, Lake Poso, Celebes [Sulawesi]). Weed, 1933: 49 (characters). Mohr, 1936b: 56 (characters, distribution). Meyer, 1985: 409 (characters, distribution). Brembach, 1991: 175– 176 (anatomy, relationships, distribution). Kottelat et al., 1993: 121 (distribution). Downing & Burns, 1995: 330 (testis study). Differential diagnosis. Three species of Nomorhamphus, N. celebensis, N. kolonodalensis and N. towoetii, that live in the Malili lakes region of Sulawesi Selatan and the Lake Poso region of Sulawesi Tengah are characterized by the morphology of the second analfin ray in males in which there are 10 segments proximal to the paired spines, segments approximately equal in length, and a distinctly segmented, straight spiculus that is not noticeably thickened (Fig. 47). Among these species, N. celebensis is distinguished by a distinctly wider and shorter upper jaw (UJL/UJW <1.0, vs. 1.0–1.3 in N. towoetii and 1.1–1.8 in N. kolonodalensis).

Figure 42. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus bakeri comb. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. USNM 84275 (31.6 mm).

undet., of which 2 males and 1 female have been cleared and counterstained); Maros, Bontomanai, 31 km SW of main Nadin Maros; Louie et al.; 22 Jul 1995. AMNH 35379 (6 females, 37.8–82.7; 6 males 42.7–47.6, of which 1 male has been cleared and counterstained).

Description. (Table 23) Deep-bodied (BDP1 >15.7% SL); vertebrae 39–40 [40]; precaudal vertebrae 21–22 [22] (Table 18); caudal vertebrae 17–18 [18] (Table 19); 37–44 predorsal scales; length of lower jaw reduced; upper and lower jaws approximately equal in length; upper jaw wider than long (UJL/UJW 0 .70–0.94); four rows of conical teeth on upper and lower jaws; gill rakers tear-drop shaped with many teeth; branchiostegal rays 8; small, lightly pigmented mandibular appendage in both sexes; pectoral fin pointed, not reaching base of the pelvic fin; 12–13 pectoral-fin rays; second anal-fin ray of males with 10 segments proximal to paired spines; length of segments approximately equal; spiculus straight, thin, and distinctly segmented (Fig. 48); anal-fin rays 14–16 [16] (Table 20); dorsalfin origin over anal-fin ray 4, 5, or 6 [6]; dorsal-fin rays 11–12 [12] (Table 21); caudal fin truncate; fifth hypural fused to dorsal hypural plate. Colour in alcohol. (Fig. 46) Background colour olive; diffuse midlateral stripe; black pigment on distal portion of anterior anal-fin rays, pronounced in females; black pigment on distal tips of pelvic-fin rays. Distribution. Lake Poso, Sulawesi Tengah, Indonesia.

NOMORHAMPHUS CELEBENSIS WEBER & DE BEAUFORT, 1922 (Figures 46, 47; Tables 18–21, 23)

Remarks. None of the five syntypes is a mature male. Therefore, the largest female specimen (ZMA 104.377, female) is herein designated as the lectotype.

Hemiramphus (Dermatogenys) sp. Weber, 1913: 202 (collection report). Nomorhamphus celebensis Weber & de Beaufort, 1922:

Material examined. SULAWESI TENGAH: 13 specimens from three collections (28.7–80.2 mm SL).

250

A. D. MEISNER

Figure 43. Nomorhamphus brembachi. A, paratype of N. brembachi, ZMH 7166, male, 39.0 mm SL, Longron, Sulawesi Selatan. B, paratype of N. sanussi, ZMH 7616, female 43.8 mm SL, Sepoja, Sulawesi Selatan.

Figure 44. Photographs of paratypes of nominal species and subspecies here considered junior synonyms of Nomorhamphus brembachi showing variation in development of the mandibular appendage in males: A, paratype of N. ravnaki, ZMH 7160, 54.8 mm SL, Ban Timurung, Sulawesi Selatan. B, paratype of N. sanussi, ZMH 7616, 39.9 mmSL, Sepoja, Sulawesi Selatan. C, paratype of N. ravnaki australe, ZMH 7163, 54.5, Bossolo, near Maros, Sulawesi Selatan. D, paratype of N. ravnaki australe, ZMH 7163, 48.5, Bossolo, near Maros, Sulawesi Selatan.

Lectotype (herein designated): ZMA 104.377 (lectotype of Nomorhamphus celebensis, 1 female, 66.8); Lake Poso; Sarasin (Table 16).

ZMA 104.376 (paralectotypes of Nomorhamphus celebensis, 1 female, 29.5, 1 immature male, 28.7, 2 undet.); Lake Poso; Kruyt. CMK 8013 (5 females,

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synonym of Nomorhamphus ebrardti ). Downing & Burns, 1995: 330 (testis study). Nomorhamphus ebrardti. Meyer, 1985: 410 (characters, distribution). Dermogenys ebrardtii. Brembach, 1991: 157–158 (anatomy, relationships, distribution). Kottelat et al., 1993: 118 (distribution). Meisner & Burns, 1997b: 298 (reproductive biology, viviparity). Differential diagnosis. Nomorhamphus ebrardtii is distinguished from all other Nomorhamphus by the shape of the spiculus which is elongate, distal tip expanded, curved dorsally, contacting distal tip of third anal-fin ray (Fig. 49).

Figure 45. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus brembachi and Nomorhamphus liemi. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. (Nomorhamphus liemi USNM 33835, 48.0 mm).

Description. (Table 24) Deep-bodied (BDP1 >14% SL); vertebrae 37–39 [38]; precaudal vertebrae 21–22 [21] (Table 18); caudal vertebrae 15–17 [17] (Table 19); 31–39 predorsal scales; lower jaw short (13.4–15.9 times in SL); upper jaw longer than wide (UJL/UJW 1.1–1.7); biserial conical teeth on upper and lower jaws; gill rakers tear-drop shaped, 1–3 teeth; branchiostegal rays 9–10; pectoral fin pointed, not reaching base of pelvic fin; pectoral-fin rays 12–13; second anal-fin ray in males with 10 segments proximal to paired spines, length of segments approximately equal (see Fig. 49); elongate spiculus, distal tip expanded, curved dorsally, contacting distal tip of third anal-fin ray (Fig. 49); anal-fin rays 15–16 [15] (Table 20); dorsal-fin origin over anal-fin ray 6 or 7; dorsal-fin rays 11–12 [11] (Table 21); fifth hypural fused to dorsal hypural plate; caudal fin truncate; type IV form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 48) Background colour tan; thin midlateral stripe from pectoral to caudal fin; distinct oval spot anterior to pectoral fin; black pigment along lower jaw; scattered melanophores on dorsal and ventral surfaces.

46.2–80.2, 3 males, 33.7–36.6); Poso River, 4 and 10 km downriver of Lake Poso; Bleher; 4 Nov 1991.

Distribution. Kabaena, Penango, and Rumbia, Sulawesi Tenggara, Indonesia.

NOMORHAMPHUS EBRARDTII (POPTA, 1912)

Material examined. SULAWESI TENGGARA. 35 specimens from four collections (33.7–60.0 mm SL).

COMB. NOV.

(Figures 48, 49; Tables 18–21, 24)

Hemiramphus (Dermatogenys) ebrardtii Popta, 1912: 187–190 (original description, Kabaena, Penango, and Rumbia Southeast Celebes [Sulawesi]). Meyer, 1985: 410 (listed as a synonym of Nomorhamphus ebrardti ). Dermogenys ebrardti. Weber & de Beaufort, 1922: 139 (characters, distribution). Mohr, 1936a: 50–51 (characters, distribution). Meyer, 1985: 410 (listed as a

SMF 6495 (radiograph of lectotype, 1 female); Kabaena, Penango, and Rumbia southeast Celebes [Sulawesi Tenggara]. RMNH 10535.36 (paralectotypes of Dermogenys ebrardtii, 2 females, 52.3–54.6). AMNH 208333 (paralectotypes of Dermogenys ebrardtii, 2 females, 39.9–47.5, 1 male, 37.4). ZMH 7150 (18 females, 39.1–60.0, 11 males, 33.7–48.5, of which 2 males have been cleared and counterstained); Insel Wawoni; Vogt; 1978.

252

A. D. MEISNER Table 22. Morphometrics of synonyms of Nomorhamphus brembachi. (A) one female paratype of N. brembachi; (B) one male paratype of N. brembachi; (C) five female paratypes of N. ravnaki ravnaki; (D) five male paratypes of N. ravnaki ravnaki; (E) three female paratypes of N. ravnaki australis; and (F) 11 males of N. ravnaki australis. Standard length is expressed in mm; measurement 1 is a proportion; measurements 2–9 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. UJL/UJW 2. SN-P1 3. SN-P2 4. P2-C 5. HDL 6. BDP1 7. BDP2 8. ORBL 9. INTORBL

A

B

37.5 — 34.7 61.1 40.8 32.0 15.5 15.5 6.1 8.5

39.0 0.81 36.4 61.3 43.1 32.8 16.7 16.7 6.7 8.7

C

D

56.6–78.1 0.88–0.93 33.4–35.1 61.7–64.1 39.8–42.1 31.6–33.4 14.7–16.5 14.7–16.5 5.3–5.7 7.7–8.5

35.2–55.0 0.87–0.97 35.9–37.6 59.1–63.3 41.1–43.4 33.0–35.1 14.2–16.9 14.2–16.9 5.5–6.3 7.1–8.0

E 7.6–51.8 0.86–0.90 33.5–34.8 58.8–61.4 41.2–42.0 31.7–32.8 15.1–15.6 15.1–15.6 5.3–5.6 7.8–8.4

F 42.8–56.5 0.89–0.96 35.0–39.0 6.0-–63.7 40.5–42.6 32.0–35.9 15.0–19.6 15.6–20.1 5.7–6.6 7.4–8.9

Figure 46. Nomorhamphus celebensis. A, male, CMK 8013, 36.6 mm SL; Poso River, Sulaesi Tengah; B, female 80.2 mm SL, CMK 8013.

NOMORHAMPHUS HAGENI (POPTA, 1912) (Figure 50; Tables 18–21, 25)

Hemiramphus hageni Popta, 1912: 190–192 (original description, Penango, Celebes [Sulawesi]). Nomorhamphus hageni Weber & de Beaufort, 1922:

142 (collection report); Mohr, 1936b: 56–58 (characters, distribution). Meyer, 1985: 411 (characters, distribution). Brembach, 1991: 177–178 (anatomy, relationships, distribution). Kottelat et al., 1993: 121 (distribution).

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anal-fin rays 15–17 (Table 20); dorsal-fin origin over anal-fin ray 4; dorsal-fin rays 12–13 (Table 21). Distribution. Penango, Sulawesi Tenggara, Indonesia. Remarks. Nomorhamphus hageni is known only from the original collection made in 1909. The type series is in poor condition, females have been dissected and all fin rays have deteriorated (Fig. 50). The identity of this species will likely remain in question until fresh material from the type locality is available. The original description was based on 36 specimens, lectotype designation by Brembach 1991: 177 (SMF 6521). Material examined. SULAWESI TENGGARA: 11 specimens from two collections (60.5–93.1 mm SL). AMNH 20834 (paralectotypes, 3 females, 66.8–93.1); Penango. SMF 6522 (paralectotypes, 8 females, 60.5–83.0).

Figure 47. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus celebensis; N. towoetii, and N. kolonodalensis. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. (Nomorhamphus kolonodalensis, USNM 338490, 39.0 mm).

Hemirhamphus hageni. Meyer, 1985: 411 (listed as a synonym of Nomorhamphus hageni ). Diagnosis. (See Remarks). Description. (Table 25) Deep-bodied (BDP1 >16%); vertebrae 39–41; precaudal vertebrae 21–22 (Table 18); caudal vertebrae 18–19 (Table 19); predorsal scales 45–57; upper jaw longer than wide (UJL/ULW 1.0–1.3);

NOMORHAMPHUS LIEMI VOGT, 1978 (Figures 45, 51; Tables 18–21, 26)

Nomorhamphus liemi Vogt, 1978b: 8 (original description, Maros, Celebes [Sulawesi]). Meyer, 1985: 412 (characters, distribution). Brembach, 1991: 179– 180 (anatomy, relationships, distribution). Kottelat, 1992: 271 (taxonomy). Nomorhamphus liemi liemi. Meyer, 1985: 412 (listed as a subspecies). Brembach, 1991: 179 (listed as a subspecies). Kottelat, 1992: 271 (taxonomy). Nomorhamphus liemi snijdersi Vogt, 1978a: 224 (original description, Maros, Celebes). Meyer, 1985: 412 (characters, distribution). Brembach, 1991: 179–182 (anatomy, relationships, distribution). Kottelat, 1992: 271 (taxonomy). Nomorhamphus liemi. Kottelat et al., 1993: 121 (distribution). Downing & Burns, 1995: 330 (testis study).

Table 23. Morphometrics of Nomorhamphus celebensis. (A) female lectotype (ZMA 104.377); (B) one female paralectotype; (C) one male paralectotype; (D) five females; and (E) three males. Standard length is expressed in mm; measurement 1 is a proportion; measurements 2–9 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. UJL/UJW 2. SN-P1 3. SN-P2 4. P2-C 5. HDL 6. BDP1 7. BDP2 8. ORBL 9. INTORBL

A

B

C

66.8 0.94 34.9 64.7 40.3 31.7 16.6 16.6 4.8 8.8

28.7 0.88 31.4 56.5 42.5 30.7 15.7 17.1 6.2 7.7

29.5 0.79 32.5 58.0 45.7 30.2 17.6 19.0 6.8 7.5

D 28.7–80.2 0.70–0.88 31.9–33.5 62.8–66.9 41.0–43.5 29.1–31.1 15.8–17.1 15.8–17.1 4.7–6.0 8.0–8.6

E 29.5–36.6 0.70–0.90 31.9–34.4 61.2–61.7 41.0–43.0 29.1–32.0 15.8–18.3 15.8–18.3 4.7–6.8 8.0–8.6

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A. D. MEISNER

Figure 48. Nomorhamphus ebrardtii comb. nov. A, male, ZMH 7150, 45.5 mm SL; Wawoni, Sulawesi Tenggara; B, female, ZMH 7150, 60.2 mm SL.

Differential diagnosis. Two species of Nomorhamphus (N. brembachi and N. liemi ) that live in the Maros region of Sulawesi Selatan (southwestern arm) are characterized by the second anal-fin ray in males with 10 segments proximal to the paired spines, length of segments approximately equal; the second ray is curved at approximately 3/4 its length so that the distal 1/4 slopes ventrally; short, thick spiculus not clearly segmented (Fig. 46). Nomorhamphus liemi is distinguished from N. brembachi by fin pigmentation in which at least the distal half of all fins is pigmented, many completely pigmented (Fig. 51). Description. (Table 26) Deep-bodied (BDP1 >12% SL); vertebrae 39–42 [39]; precaudal vertebrae 21–24 [21] (Table 18); caudal vertebrae 17–19 [19] (Table 19); 36–43 predorsal scales; length of lower jaw reduced, upper and lower jaw approximately same length; upper jaw about as wide as long (UJL/UJW 0.8–1.0); four rows of teeth on upper and lower jaws; patch of conical teeth on upper jaw extending posteriorly; gill rakers wide with many teeth; branchiostegal rays 9–10; pectoral fin ray pointed nearly reaching base of pelvic fin in males; pectoral-fin rays 11–13; second ray curved at approximately 3/4 its length so that the distal 1/4 slopes ventrally; spiculus thick, short and not clearly segmented (Fig. 45); anal-fin rays 15–16 [15] (Table

20); dorsal-fin origin over anal-fin ray 5 or 6 [6]; dorsalfin rays 11–12 [12] (Table 21); caudal fin truncate; fifth hypural fused to dorsal hypural plate. Colour in alcohol. (Fig. 51) Background colour tan or brown; males and females with a reduced, pigmented mandibular appendage; distinct midlateral stripe from pectoral to caudal fin; diffuse spot anterior to pectoral fin; some populations with dark pigment on ventral surface of lower jaw; females with black pigment on distal 3/4 of pelvic-fin rays, along entire length of dorsal-fin rays 1–3, on distal half of middle anal-fin rays, along entire length of posterior anal-fin rays, and along entire length of anterior anal-fin rays 1–3, caudal fin dusky; similar pigmentation in males except pigment on anterior anal-fin rays absent; all fins with at least half black pigment, fin of some populations completely pigmented. Distribution. Maros, Sulawesi Selatan, Indonesia. Remarks. Nomorhamphus liemi liemi and N. liemi snijdersi were distinguished on the basis of fin pigmentation. However, amount of fin pigmentation seems to vary within each subspecies. In all specimens examined, all fins had black pigment along at least half of their length. Because these two subspecies

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255

12 males, 36.9–44.7); collected with holotype. USNM 338351 (7 females 31.2–88.2; 7 males 35.6–44.3, 1 undet., of which 2 males, 1 female have been cleared and counterstained); Maros, Malawa, Bentenge 3 km into the mountains from Bentenge; Louie, Ama, Azia; 23 Jul 1995. NOMORHAMPHUS MEGARRHAMPHUS (BREMBACH, 1982) COMB. NOV. (Figures 52, 53; Tables 18–21, 27)

Dermogenys megarrhamphus Brembach, 1982: 55 (original description, Lake Towoeti (Towuti) central Sulawesi [Selatan]). Brembach, 1991: 158–159 (anatomy, relationships, distribution). Kottelat et al., 1993: 118 (distribution). Meisner & Burns, 1997b: 298 (reproductive biology, viviparity). Differential diagnosis. The anal fin of Nomorhamphus megarrhamphus is most similar to N. weberi except that the spiculus is only slightly curved, without a distinct sickle shape (Fig. 53).

Figure 49. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus ebrardtii comb. nov. ZMH 7150 (45.0 mm). Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled.

are found sympatrically and because the counts and measurements overlap, these two subspecies are here recognized as synonyms. Material examined. SULAWESI SELATAN: 55 specimens from six collections (31.2–88.2 mm SL). ZMH 7617 (radiograph of holotype of Nomorhamphus liemi liemi, female); rivers near Malawa, in the highlands of Maros; Vogt; 1978. ZMH 7619 (paratypes of Nomorhamphus liemi liemi, 3 females, 66.9–74.0); collected with holotype. ZMH 7618 (paratype of N. liemi liemi, 1 male, 40.7); collected with holotype. ZMH 7155 (radiograph of holotype of Nomorhamphus liemi snijdersi, female); rivers near Malawa in the highlands of Maros; Vogt; 1978. ZMH 7157 (paratypes of Nomorhamphus liemi snijdersi, 22 females, 37.6–69.6,

Description. (Table 27) Slender-bodied (BDP1 <13% SL); vertebrae 40–41 [40]; precaudal vertebrae 24–27 [24] (Table 18); caudal vertebrae 16–17 [16] (Table 19); predorsal scales 43–51; lower jaw elongate (4.0–5.7 times in SL); upper jaw longer than wide (UJL/UJW 1.3–2.3); biserial or triserial conical teeth on upper and lower jaws; gill rakers teardrop-shaped, many teeth on dorsal surface; branchiostegal rays 9–11; pectoral fin pointed, not reaching base of pelvic fin; pectoral-fin rays 10–13; second anal-fin ray in males, 10 segments proximal to spines, length of segments approximately equal, distal portion of entire ray curved ventrally; spiculus slightly curved, without a distinct sickle shape (Fig. 53); first anal pterygiophore thin, angled anteriorly; anal fin-rays 15–16 [15] (Table 20); dorsal-fin origin over anal-fin ray 5–6 [5]; dorsal finrays 10–11 [10] (Table 21); caudal fin truncate; fifth hypural fused to dorsal hypural plate; type IV form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 52) Background colour olive or whitish ventrally; concentration of melanophores on dorsal surface; diffuse midlateral stripe, most prominent posteriorly; black pigment at base and distal tips of pelvic-fin rays; black pigment on distal tips of dorsalfin rays; all other fins dusky. Distribution. Lakes Towuti and Mahalona, Sulawesi Selatan, Indonesia. Remarks. The original description of this species was based on 25 specimens with no holotype designated (Brembach, 1982). In a subsequent revision (Brembach, 1991), ZMH 7151 was listed as the holotype. However,

256

A. D. MEISNER Table 24. Morphometrics of Nomorhamphus ebrardtii comb. nov. (A) four female paralectotypes (AMNH 208333; RMNH 10535.36); (B) one male paralectotype (AMNH 208333); (C) six females; and (D) four males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

D

39.9–54.6 — 1.3–1.7 30.4–34.3 61.4–65.2 39.6–41.3 25.6–32.3 14.8–16.2 14.8–16.2 5.7–6.3 6.3–7.7

37.4 — 1.7 33.7 62.3 39.6 31.8 16.3 16.3 6.4 6.4

50.2–60.0 14.2–15.9 1.1–1.2 32.7–36.7 62.4–66.6 37.0–39.6 34.0–30.8 15.1–17.1 15.1–17.1 5.7–6.9 8.4–9.4

38.8–48.5 13.4–13.9 1.1–1.5 35.1–37.7 61.8–64.2 39.0–40.2 33.6–35.1 16.1–17.5 16.1–18.6 6.4–7.2 8.7–9.3

Figure 50. Nomorhamphus hageni, paralectotype, female, SMF 6522 63.0 mm SL; Penango, Sulawesi Tenggara.

Table 25. Morphometrics for nine female paralectotypes of Nomorhamphus hageni (SMF 6522–9; AMNH 208334). Standard length is expressed in mm; measurement 2 is a proportion; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

61.9–93.1 1.0–1.3 33.0–35.2 58.6–64.2 40.0–44.3 31.2–37.3 16.4–18.9 17.4–18.9 4.7–5.4 7.9–9.0

as no type was designated in the original description, ZMH 7151 is herein designated as the lectotype. In that same revision (Brembach, 1991), 25 paratypes (= paralectotypes) are listed contained in two lots, ZMH 7152 and ZMH 7153, which actually contain 51 specimens. As there is no way of determining which 25 specimens was the basis for the original description, all 51 specimens are here considered paralectotypes. Material examined. SULAWESI SELATAN: 641 specimens from nine collections (26.8–80.5 mm SL). ZMH 7151 (radiograph of lectotype of Dermogenys megarrhamphus, female); Lake Towoeti [Towuti]; Vogt, 1978. ZMH 7152 (radiograph of paralectotype, male); collected with lectotype; Vogt, 1978. ZMH 7153 (paralectotypes, 26 females, 47.1–56.0, 24 males, 42.9–47.2); collected with lectotype; Vogt, 1978. CMK 6234 (14 females, 35.4–80.5, 8 males, 28.9–38.9, 8 undet., of which 2 males have been cleared and counterstained);

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Figure 51. Nomorhamphus liemi. A, male, paratype, ZMH 7618, 40.5 mm SL; Maros, Sulawesi Selatan; B, female, USNM 33851, 78.3 mm SL; Maros, Sulawesi Selatan. Table 26. Morphometrics of Nomorhamphus liemi. (A) three female paratypes of N. liemi liemi; (B) 10 female paratypes of N. liemi snijdersi and (C) 10 male paratypes of N. snijdersi. Standard length is expressed in mm; measurement 1 is a proportion; measurements 2–9 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. UJL/UJW 2. SN-P1 3. SN-P2 4. P2-C 5. HDL 6. BDP1 7. BDP2 8. ORBL 9. INTORBL

A

B

C

67.5–73.0 0.9 31.8–33.0 62.7–64.6 37.8–40.0 30.4–31.1 12.9–14.6 12.9–14.6 5.2–5.6 8.4–8.6

54.7–70.2 0.8–1.0 31.5–34.2 60.3–66.5 38.2–41.9 28.9–31.1 14.5–17.4 14.5–18.0 4.7–5.6 8.4–9.3

36.5–45.9 0.9–1.0 34.5–36.6 60.3–82.5 38.1–43.6 31.6–34.4 15.9–18.1 15.9–18.1 5.7–6.7 7.5–9.2

Lake Towuti, about 3 km south of Timampu, estuary of Sungei Tempampu; Kottelat; 30 Jun 1988. USNM 338505 (415, females, 30.0–47.2, males, 28.7–45.2); Towuti, Sungei Lingkoburanga, approx. 8 km SW of Timampu; Parenti, Louie et al.; 9 Aug 1995. USNM

338502 (12 females, 27.5–44.4, 4 males, 32.0–36.6, 27 undet.); Towuti, first stream just S of Sungei Ling Koburanga, where it enters Lake Towuti; Parenti, Louie, et al., 9 Aug 1995. USNM 338501 (14 females, 30.1–68.6, 14 males, 26.8–43.3, 10 undet., of which 2

258

A. D. MEISNER

Figure 52. Nomorhamphus megarrhamphus comb. nov. A, male, USNM 338501, 43.3 mm SL; Lake Towuti, Sulawesi Selatan; B, female, USNM 338501, 61.9 mm SL.

females, 2 males have been cleared and counterstained); Towuti, Sungei Tombala where it empties Lake Mahalona on SW shore; Parenti, Louie et al.; 8 Aug 1995. USNM 338503 (1 female, 76.3, 2 males, 40.3–44.3); Lake Towuti at Kampung Baru, 3 km E of Timampu; Parenti, Louie et al.; 7 Aug 1995. USNM 338500 (60, females, 30.1–56.1, males, 31.4–45.7); Towuti, Sungei Lamapu where it enters western shore of Lake Towuti, 3–4 km SW of Timampu; Parenti, Louie et al., 9 Aug 1995.

NOMORHAMPHUS PECTORALIS (FOWLER, 1934b) COMB. NOV.

(Figures 54, 55; Tables 18–21, 28)

Dermogenys pectoralis Fowler, 1934b: 326–327 (original description, Bubbucan, Philippines). Kottelat, 1992: 269 (taxonomy). Herre, 1953: 160 (listed as a synonym of Dermogenys viviparus). Dermogenys viviparus not of Peters, 1865. Herre, 1944: 47 (collection report). Meisner & Burns, 1997b: 298 (identified as population from Luzon with the Type III form of halfbeak viviparity). Differential diagnosis. The anal fin of N. pectoralis is most similar to N. manifesta, Nomorhamphus pinnamacula, N. vivipara and N. philippina in that the spiculus is elongate. It is distinguished from these other Philippine species by the dorsally offset spiculus comprised of 5–6 segments (vs. 7–10), terminal segment of spiculus short, rarely curved (Fig. 55).

Description. (Table 28) Deep-bodied (BDP1 >13% SL); total vertebrae 39–41 [39]; precaudal vertebrae 21–23 [22] (Table 18); caudal vertebrae 17–19 [17] (Table 19); predorsal scales 35–40; lower jaw short (6.9–18.6 times in SL); upper jaw longer than wide (UJL/UJW 1.2–1.9); biserial or triserial conical teeth on upper and lower jaws; gill rakers tear-drop shaped, one to three teeth; branchiostegal rays 10; pectoral fin pointed, reaching pelvic fin in males; 11–12 pectoralfin rays; second anal-fin ray in males with 8–10 segments proximal to paired spines; spiculus offset dorsally, 5–6 segments, terminal segment short, rarely curved (Fig. 53); anal-fin rays 16–19 [17] (Table 20); dorsal-fin origin over anal-fin ray 6 [6]; dorsalfin rays 11–12 [12] (Table 21); genital papilla in females enlarged, unpigmented; caudal fin truncate; fifth hypural partially fused to the dorsal hypural plate; type III form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 54) Background colour tan, diffuse spot anterior to pectoral fin; scattered melanophores along lateral surfaces; in females, black pigment on middle portion of middle anal-fin rays and in some large females, black pigment on posterior dorsal-fin rays. In males, black pigment on distal portion of dorsal- and caudal-fin rays. Distribution. Luzon, Philippines. Material examined. LUZON, PHILIPPINES: 476 specimens from 14 collections (22.8–88.8 mm SL).

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

259

Oriwong Toll Gate; Herre; 17 Sep 1940. CAS 137630 (22 males, 28.6–46.2, of which 2 males have been cleared and counterstained); Nueva Vizcaya Prov., Oriwong Toll Gate; Herre; 2 Jun 1940. CAS 137636 (6 females, 22.8–29.2, 9 males, 23.2–31.0, 12 undet.); Laguna Prov., Molawin Creek, College of Agriculture; Herre; 17 Oct 1940. CAS 128480 (12 females, 31.4– 59.2, 4 males, 33.2–45.2); Nueva Ecija Prov, Talavera River; Herre; 25 May 1940. CAS 137627 (112, females, 37.7–88.8); Nueva Vizcaya Prov., Sante Fe; Herre, 3 Jun 1931. USNM 138675 (7 females, 38.5–49.3, 10 males, 31.6–41.6, of which 1 male has been cleared and counterstained); Mariquina river, Uana; Albatross; 1 Jan 1908. USNM 197727 (1 female, 27.1, 1 male, 41.3); central Luzon; Johnsen; 1963. USNM 138666 (1 female, 82.6, 2 males, 46.7–52.0); Mato river, Lagondy Gulf; Albatross; 18 Jun 1909. NOMORHAMPHUS PHILIPPINA (LADIGES, 1972) COMB. NOV.

(Figures 56, 57; Tables 18–21, 29)

Dermogenys philippinus Ladiges, 1972: 210–212 (original description, Kulamen Plateau, Cebu, Philippines). Brembach, 1991: 163–164 (anatomy, relationships, distribution). Dermogenys philippina. Kottelat, 1992: 269 (emended spelling).

Figure 53. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus megarrhamphus comb. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. USNM 338501 (45.0 mm).

USNM 93068 (holotype of Dermogenys pectoralis, male, 33.5); Bubbucan; Albatross; 19 Dec 1907. USNM 93069 (paratypes of D. pectoralis, 1 male, 30.8, 1 undet.); collected with holotype. CAS 128479 (94, females, 31.5– 60.3, males 26.7–38.0, of which 2 females, 2 males have been cleared and counterstained); Laguna Prov., Molawin Creek, College of Agriculture; Herre; Apr– May 1931. CAS 137634 (21 females, 25.7–51.4, 13 males, 25.2–37.0, 13 undet.); Rizal Prov., Sala River, Barrio Malaya, Jalajala; Herre; 27 Dec 1940. CAS 128478 (9 females, 31.9–56.7, 16 males, 28.9–47.2, 5 undet.); Nueva Vizcaya Prov., North of Sante Fe; Herre; 27 May 1931. CAS 137628 (52 males, 31.6–44.0); Nueva Vizcaya Prov., Sante Fe; Herre; 3 Jun 1940. CAS 137629 (51 females, 41.3–76.4); Nueva Vizcaya Prov.,

Differential diagnosis. The anal fin of N. philippina is most similar to that of N. pectoralis, N. vivipara, N. pinnimaculata, N. manifesta in that the spiculus is elongate. It is distinguished from other Philippine Nomorhamphus species by the combination of: segmentation of second anal-fin ray in which all segments are approximately equal in length; 9–10 (vs. 5–8) segments in the spiculus, terminal segment turned ventrally, shorter than segmented region, terminal segment shorter than in N. vivipara (Fig. 57); and presence of black pigment on distal tips of dorsal- and anal-fin rays. Description. (Table 29) Deep-bodied (BDP1 >13% SL); vertebrae 36–37 [37]; precaudal vertebrae 21 [21] (Table 18); caudal vertebrae 16 [16] (Table 19); 33–39 predorsal scales; lower jaw short (10.4–18.8 times in SL); upper jaw longer than wide (UJL/UJW 1.3–1.8); biserial conical teeth on upper and lower jaws; pectoral fin pointed not reaching pelvic fin in males; segments in second anal-fin ray in males approximately equal in length; 9–10 segments in spiculus (Fig. 57); analfin rays 14–15 [15] (Table 20); dorsal-fin origin over anal-fin ray 6 or 7 [7]; dorsal-fin rays 12 [12] (Table 21); caudal fin truncate; fifth hypural separate from dorsal hypural plate along most of its length. Colour in alcohol. Background colour tan; thin midlateral stripe extending from opercle to caudal fin;

260

A. D. MEISNER Table 27. Morphometrics of Nomorhamphus megarrhamphus comb. nov. (A) 10 female paralectotypes (ZMH 7153); (B) seven male paralectotypes; (C) 10 females; (D) nine male. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

D

47.1–56.0 4.5–5.2 1.6–2.3 28.5–32.1 55.6–60.6 40.4–42.4 25.9–28.2 10.8–12.9 10.8–12.9 4.9–5.5 6.6–7.3

42.9–47.2 4.0–4.8 1.6–1.8 29.4–32.3 56.4–60.8 39.2–43.2 26.0–29.2 9.8–11.4 9.8–11.4 5.4–6.0 .6–6.8

38.9–68.3 4.5–5.7 1.5–2.0 32.3–35.1 59.2–63.3 39.2–40.9 28.1–30.8 9.7–13.4 9.7–13.4 4.3–5.7 4.3–6.7

39.0–47.1 4.2–5.4 1.3–1.6 33.3–35.3 59.3–63.5 38.4–41.8 30.3–31.3 11.0–12.1 11.0–12.1 5.6–6.1 5.5–6.4

Figure 54. Nomorhamphus pectoralis comb. nov. A, holotype, USNM 93068, male, 33.5 mm SL; Bubbucan, Philippines; B, female, CAS 128479, 60.5 mm SL; Laguna Province, Luzon, Philippines.

black pigment on distal tips of dorsal- and anal-fin rays; diffuse black spot anterior to pectoral fin. Distribution. Cebu and Mindanao, Philippines.

ZMH 4534 (radiograph of holotype of Dermogenys philippinus, male); Kulamen Plateau. ZMH 4535 (paratypes of Dermogenys philippinus, 1 female, 34.4, 2 males, 40.3–43.0); collected with holotype.

Material examined. Cebu. Four specimens from two collections (34.4–43.0 mm SL).

MINDANAO. 11 specimens from two collections (21.1–61.5 mm SL).

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

261

Nomorhamphus towoeti. Kottelat et al., 1993: 122 (distribution). Meisner & Burns, 1997b: 298 (reproductive biology, viviparity). Diagnosis. Three species of Nomorhamphus (N. celebensis, N. kolonodalensis and N. towoetii ) that live in the Malili lakes region of Sulawesi Selatan and the Lake Poso region of Sulawesi Tengah are characterized by anal-fin morphology in which the second anal-fin ray in males has 10 segments proximal to paired spines, segments of this ray approximately equal in length, and a distinctly segmented, straight spiculus that is not noticeably thickened (Fig. 48). Nomorhamphus towoetii is distinguished by the combination of: pigmentation pattern in which females are dusky, and most large males are completely melanistic (Fig. 58); and upper jaw slightly longer than wide, UJL/ UJW 1.0–1.3 (versus <1.0 in N. celebensis and 1.1–1.8 in N. kolonodalensis).

Figure 55. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus pectoralis comb. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. CAS 137630 (44.0 mm).

FMNH 50934 (2 females, 47.5–61.5); Cotobato, Burungkot, Upi; Anonevo; 3 Jan 1947. FMNH 50933 (6 females, 21.1–45.1, 3 males, 23.0–28.8); Cotobato, Burungkot, Upi; Anonevo; 10 Jan 1947.

NOMORHAMPHUS TOWOETII LADIGES, 1972 (Figures 47, 58; Tables 18–21, 30)

Nomorhamphus towoetii Ladiges, 1972: 207–209 (original description, Lake Towoeti [Towuti], Celebes [Sulawesi]). Meyer, 1985: 414 (characters, distribution). Brembach, 1991: 189 (anatomy, relationships, distribution). Downing & Burns, 1995: 330 (testis study).

Description. (Table 30) Deep-bodied (BDP1 >14% SL); vertebrae 39–40 [40]; precaudal vertebrae 22–23 [22] (Table 18); caudal vertebrae 17–18 [18] (Table 19); 31–39 predorsal scales; length of lower jaw reduced, upper and lower jaw approximately equal in length; upper jaw longer than wide (UJL/UJW 1.0–1.3); biserial or triserial conical teeth on upper and lower jaws; gill rakers short, base expanded, many teeth; branchiostegal rays 8; small, pigmented mandibular appendage in males and females; pectoral fin pointed, not reaching base of pelvic fin; pectoral-fin rays 11–12; genital papilla in females wide, flap-like, lightly pigmented; second anal-fin ray of males with 10 segments proximal to paired spines, length of segments approximately equal; thin, straight spiculus, distinctly segmented; anal-fin rays 13–14 [13] (Table 20); dorsalfin origin over anal-fin ray 4 [4]; dorsal-fin rays 12–13 [13] (Table 21); caudal fin truncate; fifth hypural fused to dorsal hypural plate; type IV form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 58) Background colour olive; females with a diffuse midlateral stripe, more prominent posteriorly; small black spot anterior to pectoral fin; large concentration of melanophores over most of lateral body surfaces; large males melanistic; females with black pigment on distal tips of anal- and pelvicfin rays; other fins dusky. Distribution. Lake Towuti, Sulawesi Selatan and Lake Poso, Sulawesi Tenggara, Indonesia. Material examined. SULAWESI TENGAH: 110 specimens from five collections (23.5–63.5 mm SL). ZMH 4532 (radiograph of holotype of Nomorhamphus towoetii, female); Lake Towoeti [Towuti]. ZMH 4533

262

A. D. MEISNER Table 28. Morphometrics of Nomorhamphus pectoralis comb. nov. (A) male holotype (USNM 93068); (B) male paratype (USNM 93069); (C) 18 females; and (D) 20 males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

D

33.5 6.9 1.5 37.3 62.4 41.5 33.4 17.3 17.3 7.5 8.7

30.8 — 1.3 36.7 63.3 40.9 32.1 17.9 17.9 7.5 8.4

42.6–88.8 8.3–18.6 1.3–1.7 32.9–37.3 59.4–64.4 37.8–41.6 28.3–33.1 13.8–19.0 14.3–19.0 5.0–6.1 7.2–8.8

31.1–47.2 7.2–12.8 1.2–1.9 34.1–37.7 57.3–63.2 38.9–43.7 30.4–33.5 14.6–20.0 15.4–21.2 5.7–6.8 6.9–8.3

Figure 56. Nomorhamphus philippina comb. nov. A, male, paratype, ZMH 4535, 40.6 mm SL; Cebu, Philippines; B, female, FMNH 50934, 61.8 mm SL; Cotobato, Mindanao, Philippines.

(paratypes of Nomorhamphus towoetii, 2 females, 48.2– 63.5); collected with holotype. USNM 338353 (21 females, 29.7–62.4, 25 males, 31.1–40.2); Tengah, Segele,

western shore of Lake Poso where it empties at Tentena; Parenti, Louie, Beta, Young; 11 Aug 1995. USNM 338352 (27, 14 females, 23.5–44.9, 13 males, 26.1–36.9,

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

263

of which 2 males, 2 females have been cleared and counterstained); Tengah, Segele, west bank of Lake Poso at Tentena; Parenti, Louie, Beta, Young; 13 Aug 1995. CMK 6187 (7 females, 36.7–59.3, 14 males, 30.0– 47.6, of which 1 male, 1 female have been cleared and counterstained); Sungei Balambano (road from Malili to Soroako); Kottelat and Kottelat-Kloetzli; 19 Jun 1988.

NOMORHAMPHUS VIVIPARA (PETERS, 1865) COMB. NOV.

(Figures 59, 60; Tables 18–21, 31)

Hemiramphus viviparus Peters, 1865: 132–133 (original description, Bassey River, Samar Island, Philippines). Hemirhamphus (Dermatogenys) viviparus. Peters, 1868: 273 (distribution). Dermogenys viviparus. Herre, 1931: 25 (collection report). Mohr, 1936a: 51–53 (characters, distribution). Herre, 1944: 45–48 (characters, distribution). Herre, 1953: 159 (synonymy). Brembach, 1991: 168–169 (characters, relationships, distribution). Downing & Burns, 1995: 330 (testis study). Meisner & Burns, 1997b: 298 (reproductive biology, viviparity). Dermogenys viviparus mindanensis Herre, 1944: 48 (original description, Agusan Province, Mindanao). Herre, 1953: 160 (collections report). Brembach, 1991: 168–169 (characters, distribution).

Figure 57. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus philippina comb. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. ZMH 4535 (40.6 mm).

Differential diagnosis. The anal fin of Nomorhamphus vivipara is most similar to that of N. pectoralis, N. philippina, N. pinnimaculata and N. manifesta in that

Table 29. Morphometrics of Nomorhamphus philippina comb. nov. (A) female paratype (ZMH 4535); (B) two male paratypes; (C) four females; (D) two males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

45.5 10.5 1.7 38.4 64.2 36.6 35.8 18.5 20.0 7.5 7.3

40.3–43.0 15.5–18.8 1.8 37.0–38.2 63.5–64.2 38.6–39.5 33.0–33.7 14.9–15.1 14.9–15.1 6.7 7.2

∗ Proportion based on two specimens.

C 35.5–61.5 10.4–11.6∗ 1.3–1.7 33.2–37.7 62.1–63.8 39.4–40.8 30.4–34.3 13.5–15.3 13.5–15.3 6.0–7.1 6.9–8.2

D 25.9–28.8 — — 33.6–34.4 60.0–64.2 52.7–42.9 30.5–31.3 15.3–15.8 15.3–15.8 7.3–7.7 7.6–7.7

264

A. D. MEISNER

Figure 58. Nomorhamphus towoetii. A, male, USNM 338352, 38.2 mm SL; Lake Poso, Sulawesi Tengah; B, female, USNM 338352, 59.6 mm SL.

Table 30. Morphometrics of Nomorhamphus towoetii. (A) two female paratypes (ZMH 4533); (B) 16 females; and (C) 14 males. Standard length is expressed in mm; measurement 1 is a proportion; measurements 2–9 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1.UJL/UJW 2. SN-P1 3. SN-P2 4. P2-C 5. HDL 6. BDP1 7. BDP2 8. ORBL 9. INTORBL

A

B

C

48.2–63.5 1.3 34.2–35.7 63.9 40.2–40.9 30.9–34.2 14.1–14.2 14.1–14.2 5.4–6.4 7.9–8.2

47.1–65.5 1.0–1.1 33.0–35.5 59.4–62.6 39.1–42.0 30.1–32.1 13.9–16.0 13.9–16.0 5.2–6.1 7.5–8.2

32.5–40.2 1.0–1.2 34.2–37.6 60.0–64.6 39.1–42.5 30.1–34.3 14.5–16.5 14.5–16.5 5.6–8.0 5.4–8.5

the spiculus is elongate. The anal fin of Nomorhamphus vivipara is distinguished from other Philippine Nomorhamphus by: a spiculus with 7 segments, terminal segment elongate, slender, and curved ventrally; segments 4–5–6 in anal-fin ray two noticeably elongate; distinguished from N. bakeri which also occurs on

Mindanao by segmentation of the second anal-fin ray in males (Fig. 60). Description. (Table 31) Slender-bodied (BDP1 <14% SL); vertebrae 39–41 [40]; precaudal vertebrae 21–24 [23] (Table 18); caudal vertebrae 15–18 [17] (Table 19);

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS

265

Figure 59. Nomorhamphus vivipara comb. nov. A, male, CAS 137829, 34.0 mm SL; Sulu Province, Jolo Island, Philippines; B, female, CAS 137829, 46.1 mm SL.

34–41 predorsal scales; lower jaw elongate (approximately 5.3–7.9 times in SL); upper jaw longer than wide (UJL/UJW 1.3–1.7 ); biserial conical teeth on upper and lower jaws; gill rakers tear-drop shaped, 1–3 teeth; branchiostegal rays 9; pectoral fin pointed reaching pelvic fin in males; pectoral-fin rays 11–12; second anal-fin ray in males with 8 segments proximal to paired spines, segments 4–5–6 noticeably elongate, segments 5 and 6 approximately equal in length; spiculus with 7 segments, terminal segment elongate, slender, curved ventrally (Fig. 60); anal-fin rays 14–17 [14] (Table 20); dorsal-fin origin over anal-fin ray 4 or 5 [4]; dorsal-fin rays 9–12 [12] (Table 19); caudal fin truncate; fifth hypural separate from dorsal hypural plate along most of its length; type IV form of halfbeak viviparity (Meisner and Burns, 1997b). Colour in alcohol. (Fig. 60) Background colour tan; distinct midlateral stripe from pectoral fin to base of caudal fin; small distinct spot anterior to pectoral fin; black pigment on distal tips of posterior anal-fin rays, base and distal tips of pelvic-fin rays, distal tips of unmodified anal-fin rays, and posterior dorsal-fin rays in males; in females, black pigment on distal tips of anal-fin rays, and base of pectoral fins. Distribution. Luzon, Samar, Mindanao, and Jolo, Philippines.

Remarks. A lectotype is not designated because diagnostic characters are found in the andropodium and the remaining syntype is a female in poor condition. Dermogenys is a feminine noun and the specific name should be emended to D. vivipara. Material examined. SAMAR. One specimen from one collection (49.0 mm SL). ZMB 6267 (syntype of Hemiramphus viviparus female, 49.0); Basseyfluss; Jagor. LUZON. Six specimens (34.3–71.4 mm SL).

from

one

collection

ZMB 6721 (6 of 12 examined; 2 females, 58.2–71.4, 4 males, 34.3–45.4); Bach Yassot; Jagor. MINDANAO: 85 specimens from three collections (21.2–61.0 mm SL). CAS 137631 (syntypes of Dermogenys viviparus mindanensis, 74, females, 35.1–46.6, males, 29.7–31.6, of which 2 males, 2 females have been cleared and counterstained); Lake Mainit, Jabonga, Agusan Province; Herre; 1940. FMNH 50933 (4 females, 21.2–45.5, 5 males, 24.5–29.2); Burunghot, Upi, Cotabato; Anonuevo; 1947. FMNH 50934 (2 females, 47.7–61.0); Burunghot, Upi, Cotabato; Anonuevo; 1947.

266

A. D. MEISNER

Hemiramphus weberi. Weber, 1913: 202 (collection report). Dermogenys weberi. Weber & de Beaufort, 1922: 138 (characters, distribution). Mohr, 1936a: 53–55 (characters, distribution). Brembach, 1991: 171 (characters, relationships, distribution). Kottelat et al., 1993: 118–119 (distribution). Downing and Burns, 1995: 330 (testis study). Meisner & Burns, 1997b: 298 (reproductive biology, viviparity). Differential diagnosis. Nomorhamphus weberi is distinguished from other Nomorhamphus by unsegmented, sickle-shaped spiculus (Fig. 62); and distinguished from N. weberi by absence of a black spot on posterior dorsal-fin rays in females (Fig. 61).

Figure 60. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus vivipara comb. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. CAS 137829 (33.0 mm).

JOLO: 133 specimens (26.9–57.5 mm SL).

from

one

collection

Description. (Table 32) Slender-bodied (BDP1 <14% SL); vertebrae 42–46 [42]; precaudal vertebrae 24–27 [24] (Table 18); caudal vertebrae 16–19 [18] (Table 19); predorsal scales 41–44; lower jaw elongate (3.7–4.9 times in SL); upper jaw longer than wide (UJL/UJW 1.1–1.6); up to three rows of conical teeth on upper jaw; gill rakers tear-drop shaped, many teeth; branchiostegal rays 10–12; pectoral fin pointed not reaching base of pelvic fin; pectoral-fin rays 12; second anal-fin ray in males with 9–10 segments proximal to paired spines, length of segments approximately equal, segments four and five of second anal-fin ray in males thickened anteriorly; unsegmented, sickle-shaped spiculus (Fig. 62); first anal pterygiophore thin, angled anteriorly; anal-fin rays 16–18 [18] (Table 20); in females, genital papilla slightly elongate, pigmented; dorsal-fin origin over anal-fin ray 6–8 [8]; dorsal-fin rays 10–12 [11] (Table 21); caudal fin truncate; fifth hypural fused to dorsal hypural plate; type IV form of halfbeak viviparity (Meisner & Burns, 1997b). Colour in alcohol. (Fig. 61) Background colour olive or whitish ventrally; concentration of melanophores on dorsal surface; diffuse midlateral stripe, most prominent posteriorly; black pigment at base and distal tips of pelvic-fin rays; all other fins dusky.

CAS 137829 (133, females, 27.8–57.5, males, 26.9– 34.7, of which 2 males, 2 females have been cleared and counterstained); Sulu Province; Herre; 17 Sep 1940.

Distribution. Lake Matano and Lake Mahalona, Sulawesi Selatan, Indonesia.

NOMORHAMPHUS WEBERI (BOULENGER, 1897)

NMBA 1065 (holotype of Hemirhamphus weberi, female, 63.0); Lake Matana [Matano].

COMB. NOV.

(Figures 61, 62; Tables 18–21, 32)

Hemirhamphus weberi Boulenger, 1897: 429 (original description, Lake Mantana [Matano], Celebes).

Material examined. SULAWESI SELATAN: 278 specimens from seven collections (22.1–72.0 mm SL).

USNM 338506 (156, females, 63.3–71.1, males, 40.0– 41.7, of which 2 females, 2 males have been cleared and counterstained); Luwu, Nuha, Soroako; Louie, Parenti,

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267

Table 31. Morphometrics of Nomorhamphus vivipara comb. nov. (A) female syntype (ZMB 6267); (B) 10 females; and (C) 10 males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed in text

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

49.0 — 1.5 32.7 66.3 40.6 29.6 10.8 10.8 6.5 7.8

38.9–50.7 6.3–7.9∗ 1.3–1.6 31.9–33.8 58.4–63.1 39.6–41.5 28.8–31.0 10.5–13.4 10.9–13.4 4.9–6.2 6.6–7.5

31.2–35.0 5.3–6.0∗ 1.4–1.7 34.6–37.1 61.1–62.2 39.7–42.1 32.2–34.2 12.2–14.0 12.2–14.6 6.0–6.9 6.9–7.6

∗ Proportion based on four specimens.

Figure 61. Nomorhamphus weberi comb. nov. A, USNM 338506, male, 42.6 mm from Lake Matano, Sulawesi Selatan. B, USNM 338506, female, 66.1 mm.

Amos; 6 Aug 1995. USNM 338504 (5 undet.); Luwu, Nuha, stream entering small inlet approximately 6–7 km N of Soroako, western shore of Lake Matano; Louie and Parenti; 1995. AMNH 9578 (1 female, 57.7); Soroako, Lake Matano; Abendanon. CMK 6196A (47 females, 32.6–66.7, 16 males, 39.2–43.8); Lake Matano,

east of Soroako; Kottelat; 19 Jun 1988. CMK 6497 (4 females, 32.3–72.0, 15 undet.); Lake Mahalona, south coast; Kottelat and Werner; 16 Mar 1989. CMK 6275 (16 females, 28.1–39.9, 17 males, 22.1–29.4); Lake Matano, south coast, 13 km west of Soroako; Kottelat; 5 Jul 1988.

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the second anal-fin ray of males has 10 approximately equal-sized segments proximal to paired spines; distinctly segmented, straight spiculus, not noticeably thickened (Fig. 46). Among these species, N. kolonodalensis is distinguished by the combination of: short lower jaw (8.8–22.2 times in SL vs. upper and lower jaws approximately equal in length); longer upper jaw (UJL/UJW 1.1–1.8 vs. 1.2 or less); gill rakers teardrop shaped usually without teeth versus teardropshaped with many teeth; fifth hypural separate from dorsal hypural plate along most of its length versus completely fused. Description. (Table 34) Slender-bodied (BDP1 <15% SL); vertebrae 37–39 [37]; precaudal vertebrae 21–23 [22] (Table 18); caudal vertebrae 15–17 [15] (Table 19); predorsal scales 47–53; lower jaw short (8.8–22.2 times in SL); upper jaw longer than wide (UJL/UJW 1.1–1.8); biserial or triserial conical teeth on upper and lower jaws; gill rakers tear-drop shaped, 1–3 teeth; branchiostegal rays 8–9; pectoral fin pointed, nearly reaching base of pelvic fin in males; 10–11 pectoral-fin rays [11]; second anal-fin ray of males with 10 segments proximal to paired spines, length of segments approximately equal; spiculus straight, distinctly segmented (Fig. 76); anal-fin rays 13–14 [15] (Table 20); dorsal-fin origin over anal-fin ray 4 or 5; dorsal-fin rays 10–11 [12] (Table 21); fifth hypural separate from dorsal hypural plate along most of its length; caudal fin truncate; small spermatozeugmata; type IV form of halfbeak viviparity (Meisner & Burns, 1997b).

Figure 62. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus weberi comb. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. USNM 338506 (43.0 mm).

Colour in alcohol. (Fig. 63) Background colour olive or whitish; concentration of melanophores along dorsal and lateral surfaces; diffuse black spot anterior to pectoral fin; black pigment along ventral surface of lower jaw; males and females with black pigment on distal tips of posterior dorsal-, anal-, and pelvic-fin rays. Distribution. Kolonodale, Sulawesi Tengah and sites near Lake Towuti, Sulawesi Selatan, Indonesia. Material examined. SULAWESI TENGAH: 68 specimens from six collections (26.3–64.4 mm SL).

NOMORHAMPHUS KOLONODALENSIS MEISNER & LOUIE, 2000 (Figures 46, 63; Tables 18–21, 34)

Nomorhamphus sp. Meisner & Burns, 1997b: 298 (identified as population from Sulawesi with the Type IV form of halfbeak viviparity). Differential diagnosis. Three species of Nomorhamphus, N. kolonodalensis, N. towoetii and N. celebensis, that live in the Malili lakes region of Sulawesi Selatan and the Lake Poso region of Sulawesi Tengah are characterized by an anal-fin morphology in which

Holotype: MZB 8638 (male, 43.5); Poso, Kolonodale, Mondowe along road to Tiu, near bridge at home no. 53 in Mondowe; Louie and Soerto; 1995. Paratypes: USNM 338494 (8 females, 35.5–64.4, 10 males, 35.1–40.7, 1 undet., of which 2 males, 1 female have been cleared and counterstained); Poso, Kolonodale, Mondowe along Rd to Tiu, near bridge at home No. 53 in Mondowe; Louie and Soerto; 1995. USNM 338490 (7 females, 27.7–61.6, 3 males, 36.0– 39.7, of which 1 male, 1 female have been cleared and counterstained); Kolonodale, Koroloama 5 km SE of

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Table 32. Morphometrics of Nomorhamphus weberi comb nov. (A) female holotype (ZMB 1065); (B) 20 females; and (C) 14 males. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentage of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/ UJW SN-P1 SN-P2 P22-C HDL BDP1 BDP2 ORBL INTORBL

A

B

63.0 — 1.5 32.3 63.3 41.3 27.6 113.2 13.2 5.9 5.7

55.0–72.1 3.9–4.9 1.2–1.4 31.7–33.3 59.2–62.6 39.1–42.0 25.6–29.6 12.0–14.0 12.0–14.0 5.2–6.2 5.3–7.4

C 40.0–44.2 3.7–4.5 1.1–1.6 31.5–34.1 58.1–61.9 40.3–43.6 215.8–29.5 10.1–12.3 10.1–12.3 5.4–6.1 4.6–6.4

Figure 63. Nomorhamphus kolonodalensis. A, paratype, male, USNM 338490, 39.7 mm SL; Poso, Sulawesi Tengah; B, paratype, female, USNM 338490, 53.7 mm SL.

Kolonodale; Louie and Soerto; 1995. USNM 338495 (1 female, 42.4, 3 males, 26.3–38.5, 5 undet.); Poso, Beteleme, Lawonke from streams between Mandula,

Palawi Kecil and Buli; Louie and Soerto; 1995. USNM 338491 (7 females, 34.0–56.8, 3 males, 27.7–44.0, 2 undet.); Poso, near Poona (SW 4 km), Palawi River;

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Table 33. Morphometrics of Nomorhamphus kolonodalensis. (A) male holotype (MZB 8638); (B) 16 female paratypes; and (C) 15 male paratypes. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. SL/LJL 2. UJL/UJW 3. P2-C 4. HDL 5. BDP1 6. ORBL 7. INTORBL

A

B

C

43.5 10.1 2.1 41.4 37.0 18.9 7.1 7.1

46.1–67.9 10.4–22.2 1.2–1.6 38.3–41.0 29.7–35.8 11.8–14.9 4.9–6.5 6.5–8.4

35.7–47.7 8.8–16.5 1.1–1.8 36.9–41.1 28.8–34.8 11.8–15.0 5.2–7.7 6.2–7.8

NOMORHAMPHUS MANIFESTA SP. NOV. (Figures 64, 65; Tables 18–21, 34)

Dermogenys viviparus not of Peters, 1865. Herre, 1944: 47 (collection report). Herre, 1953: 160 (collection report). Differential diagnosis. The anal fin of N. manifesta is most similar to N. pectoralis, N. pinnimaculata, N. philippina and N. vivipara in that the spiculus is elongate (Fig. 65). Nomorhamphus manifesta is distinguished from other Philippine Nomorhamphus by: fin pigmentation in which there is black pigment on the distal half of dorsal-fin rays, distal tips of pelvicfin rays, distal tip of caudal-fin rays, and in females distal tips of anterior 1–2 anal-fin rays; spiculus typically with 5–6 segments, rarely 7 and terminal segment longer than in N. pectoralis. Description. (Table 34) Slender-bodied (BDP1 <14% SL); vertebrae 38–40 [40]; precaudal vertebrae 21–24 [23] (Table 18); caudal vertebrae 17–18 [17] (Table 19); predorsal scales 47–52; lower jaw elongate (6.1–11.5 times in SL); upper jaw longer than wide (UJL/UJW 1.2–1.6 ); biserial conical teeth on upper and lower jaws; gill rakers tear-drop shaped, 1–3 teeth; branchiostegal rays 9; pectoral-fin rays 10–13; dorsal-fin origin over anal-fin ray 4 or 5 [5]; second anal-fin ray in males with 8–9 segments proximal to paired spines, length of segments approximately equal; spiculus with 5–6 segments; terminal segment elongate with distal tip curved ventrally (Fig. 65); anal-fin rays 15 [15] (Table 20); dorsal-fin rays 10–12 [10] (Table 21); caudal fin

Louie and Soeroto, 1995. USNM 338492 (5 females, 37.7–50.4, 10 males, 34.1–44.7, 2 undet.); Poso, Kolonodale, Tiu, Lake Tiu, headwater input, mountain streams, 1–2 km from Desa Tiu; Louie, Soeroto, Tamanampo, Desatu; 1995. SULAWESI SELATAN: 11 specimens from one collection (44.6–67.9 mm SL). CMK 7576 (7 females, 63.4–67.9, 3 males, 44.6–47.7, 1 undet.); Sungei Masiluk, a stream about 10 km SE of Lingkona (E. shore of Lake Towuti) apparently draining to Banda Sea; D.B. Susanto; 1991.

Table 34. Morphometrics of Nomorhamphus manifesta sp. nov. (A) male holotype (CAS 129706); (B) allotype (=paratype) (CAS 169841); (C) 10 female paratypes; and (D) 10 males paratypes. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. SL/LJL 2 UJL/UJW 3. SN-P1 4. SN-P2 5. P2-C 6. HDL 7. BDP1 8. BDP2 9. ORBL 10. INTORBL

A

B

37.2 7.1 1.5 34.4 64.0 41.9 30.1 13.7 13.7 5.4 6.5

66.3 8.1 1.6 31.7 64.0 38.8 28.2 13.3 13.3 4.8 7.4

∗ Proportion based on eight specimens.

C 34.4–57.2 8.0–11.5∗ 1.2–1.5 30.7–32.4 59.6–65.1 38.2–42.5 27.9–30.1 10.2–12.2 9.9–12.6 5.3–6.1 5.9–7.0

D 31.0–38.4 6.1–7.7 1.2–1.5 32.0–36.1 60.1–64.1 37.8–42.9 29.8–33.1 10.2–13.6 10.8–14.0 5.7–6.6 5.7–8.1

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Figure 64. Nomorhamphus manifesta sp. nov. A, Holotype, male, CAS 129706, 37.2 mm SL; Ilokos Norte Province, Philippines; B, allotype (=paratype), CAS 169840, 66.3 mm SL.

truncate; fifth hypural separate from dorsal hypural plate along most of its length.

NOMORHAMPHUS PINNIMACULATA SP. NOV. (Figures 66, 67; Table 18–21, 35)

Colour in alcohol. (Fig. 64) Background colour tan, scattered melanophores on lateral surfaces, faint midlateral stripe on caudal peduncle; black pigment along ventral surface of lower jaw; males and females with black pigment on distal half of pelvic-, dorsal-, and caudal-fin rays; females with black pigment on distal tips of anteriormost anal-fin rays.

Differential diagnosis. The anal fin of N. pinnimaculata is most similar to that of N. pectoralis, N. vivipara, N. philippina and N. manifesta in that the spiculus is elongate. It is distinguished from other Philippine Nomorhamphus by: segmentation of second anal-fin ray, segments four and five slightly elongate, approximately equal in length; spiculus with 7–8 segments, terminal segment turned ventrally but shorter than segmented region (Fig. 95); terminal segment shorter than in N. vivipara; black pigmentation on anterior anal-fin rays of females.

Etymology. Manifesta from Latin manifesta (unadorned) in reference to the lack of distinctive body pigmentation. Distribution. Ilokos Norte Province, Luzon, Philippines. Material examined. One hundred and ninety-eight specimens from one collection (32.9–66.7 mm SL). Holotype: CAS 129706 (male, 37.2) Ilokos Norte Province, Solsona; Lopez; 12 Apr 1934. Measurements listed in Table 34. Paratypes: Allotype (=Paratype): CAS 169840 female (66.3); collected with holotype. CAS 169841 (196, females 42.8–66.7, males 32.9–38.2, of which 2 males and 1 female have been cleared and counterstained).

Description. (Table 35) Deep-bodied (BDP1 >14% SL); vertebrae 37–38 [38]; precaudal vertebrae 20–21 [21]; caudal vertebrae 16–17 [17]; predorsal scales 33–38; lower jaw elongate (5.6–20.6 times in SL); upper jaw longer than wide (UJL/UJW 1.2–1.7); biserial conical teeth on upper and lower jaws; gill rakers tear-drop shaped, 1–3 teeth; second anal-fin ray in males 8–9 segments proximal to paired spines; spiculus with 7–8 segments, terminal segment turned ventrally but shorter than segmented region (Fig. 67); anal-fin rays 13–14 [14]; dorsal-fin rays 10–12 [11]; caudal fin truncate; fifth hypural separate from dorsal hypural plate along most of its length; type III form of halfbeak viviparity.

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Distribution. Leyte, Philippines. Material examined. LEYTE: 125 specimens from three collections (24.6–50.0 mm SL). Holotype: ZRC 46173 (male, 41.0); creek at eastern end of Tunga, riffles; Kottelat; 7 Jul 1993. Morphometric measurements listed in Table 35. Allotype (=paratype): ZRC 46174 female (39.7). Paratypes: CMK 9984 (9 females, 27.3–47.2, 20 males, 28.8–41.0, 6 undet.); Hillosig Creek, 1.3 km north of Mahalplag junction on road from Baybay to Tacloban; Kottelat et al.; 9 Jul 1993. CMK 9980 (22 females, 24.6–50.0, 37 males, 29.2–40.5, 30 undet.); creek at eastern end of Tunga, riffles; Kottelat; 7 Jul 1993. NOMORHAMPHUS ROSSI SP. NOV. (Figures 68, 69; Tables 18–21, 36)

Dermogenys sp. Downing & Burns, 1995: 330 (testis study). Meisner & Burns, 1997b: 298 (identified as population from Luzon with the Type III form of halfbeak viviparity). Diagnosis. Nomorhamphus rossi is distinguished from all other Nomorhamphus by: second anal-fin ray in males in which segment five is greatly elongate, comprising one half of the ray; spiculus thick, unsegmented distally; offset dorsally at about one fourth its length, with distal tip pointed ventrally (Fig. 97).

Colour in alcohol. (Fig. 66) Background colour olive; brown pigment over lateral body surfaces; thin midlateral stripe, prominent posteriorly; black pigment along length of lower jaw; distinct, black oval spot on body anterior to pectoral fin; in males, black pigment along anterior 1–3 dorsal-fin rays and on distal tips of pelvic-fin rays; in females fin pigmentation as in males but with prominent black spot anteriorly on anal fin; all other fins dusky.

Description. (Table 36) Slender-bodied (BDP1 <16% SL); large size, females reaching 100.1 mm, largest male 46.6 mm; vertebrae 39–41[41]; precaudal vertebrae 23–24 [24] (Table 18); caudal vertebrae 17–18 [17] (Table 19); predorsal scales 37–44; lower jaw elongate (4.5–8.3 times in SL); upper jaw longer than wide (UJL/UJW 1.2–1.7); biserial or triserial conical teeth on upper and lower jaws; gill rakers tear-drop shaped, 1–3 teeth; branchiostegal rays 10; pectoral fin pointed extending to base of the pelvic fin in males; pectoral-fin rays 12; second anal-fin ray in males with 7 segments proximal to paired spines; segment five greatly elongate, comprising one half of ray; spiculus thick, unsegmented distally, offset dorsally at about one fourth its length, distal tip oriented ventrally (Fig. 69); analfin rays 14–17 [15] (Table 20); dorsal-fin origin over anal-fin ray 6 or 7 [7]; dorsal-fin rays 10–12 [12] (Table 21); caudal fin truncate; fifth hypural separate from dorsal hypural plate along most of its length; type III form of halfbeak viviparity (Meisner & Burns, 1997b).

Etymology. Pinnimaculata from Latin pinna (fin) and macula (spot) in reference to the diagnostic black spot on the anterior anal-fin rays in the female and dorsalfin rays in males.

Colour in alcohol. (Fig. 68) Large concentration of melanophores dorsally, ventral surface olive with scattered melanophores; diffuse oval spot anterior to pectoral fin; black pigment on lower jaw; females, black

Figure 65. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus manifesta sp. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. CAS 169841 (34.0 mm).

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Figure 66. Nomorhamphus pinnimaculata sp. nov. A, holotype, male, ZRC 46173, 41.0 mm SL; Leyte, Philippines; B, allotype (=paratype) ZRC 46174, female, 39.7 mm SL.

pigment on distal tips of caudal-fin rays, distal tips of pelvic-fin rays; anterior 1–2 anal-fin rays pigmented along their length; black pigment on distal one half of dorsal-fin rays; in some males black pigment on distal tips of caudal-fin rays and on distal tips of posterior anal- and dorsal- and pelvic-fin rays; both sexes with small concentration of melanophores anterior to dorsal fin. Etymology. Named for Charles ‘Andy’ Ross, herpetologist, who collected all the specimens of this species. Distribution. Cagayan Province, Luzon, Philippines. Material examined. LUZON: 264 specimens from one collection (39.3–100.1 mm SL). Holotype: USNM 333262 male (40.3) Baggao Municipality, Barrovia Barangay hot springs, Intel River, pool in river bordered by waterfall (upriver) and rapids (downriver); Ross et al.; 5 May 1989. Measurements listed in Table 36. Paratypes: Allotype (=Paratype): USNM 363186 female (100.1). USNM 363187 (72 females, 62.9–86.2, 47 males, 39.3–

46.6, 143 undet., of which 1 male, 1 female have been cleared and counterstained); Baggao Municipality, Barrovia Barangay hot springs, Intel River, pool in river bordered by waterfall (upriver) and rapids (downriver); Ross et al.; 5 May 1989.

DISCUSSION The taxonomy of the halfbeaks assigned to the genera Dermogenys and Nomorhamphus has been in a state of confusion for over a century. Much of this confusion stems from the lack of a rigorous methodology for proposing species relationships as well as from the use of overlapping meristic and morphometric characters in proposing species limits. The problem with generic limits dates from the original description of Nomorhamphus (Weber & de Beaufort, 1922). Without any diagnostic characters presented in that description, the implication seemed to be that the lack of an elongate lower jaw was characteristic of Nomorhamphus. Following that, when a new species was discovered that had characteristics of both Dermogenys and Nomorhamphus, it was described as a Nomorhamphus if it lacked an elongate lower jaw, or as a Dermogenys if the lower jaw was elongate. In examining these fishes, however, it became apparent

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Figure 67. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus pinnimaculata sp. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled.

that there is a continuum of lower jaw lengths so that classification based on this character is artificial, as evidenced by the arbitrary reclassification of species with short lower jaws such as N. ebrardtii by Meyer (1985). In an attempt to minimize importance of such characters in a phylogenetic analysis, other hypotheses, based the distinction between Dermogenys and Nomorhamphus on the absence of ‘true’ gill rakers on the first arch in Nomorhamphus (Anderson & Collette, 1991; Collette, 1995). This character, too, is uninformative, as it has been found in this study to be more widely distributed among the viviparous halfbeaks (Hemirhamphodon, Dermogenys and Nomorhamphus) and not diagnostic of any one genus. In the absence of defining characters for either genus, the traditional classification is artificial (Table 1, left column).

Despite the recognition of the taxonomic significance of the modified anal fin of Dermogenys, previous researchers continued to use overlapping meristic and morphometric characters to diagnose viviparous halfbeak species (e.g., Brembach, 1991). It is clear from the present study, however, that these characters are not appropriate for addressing this problem; they are largely uninformative at the species level. Here, more specimens from a wider geographic range were examined and characters of the anal fin were used to determine the number of valid species as well as to recognize several new species. Another important finding is the identity of D. pusilla, a species name that has been misused throughout the literature and in museum collections. Dermogenys s.l. (Table 1) is paraphyletic (Downing & Burns, 1995; Meisner & Burns, 1997a,b; Meisner & Collette, 1999), and I propose that the name Dermogenys be restricted to the clade containing D. pusilla, the type species of the genus. Dermogenys s.s. comprises 12 species: D. pusilla, D. burmanica, D. siamensis, D. collettei, D. orientalis, D. sumatrana, D. bispina, D. palawanensis, D. robertsi and D. bruneiensis. Two species D. vogti and D. brachynotopterus are tentatively included in this genus (see systematic accounts). I support retaining the name Nomorhamphus for the clade containing N. celebensis, the type species of Nomorhamphus. However, these findings also support expanding the generic limits to include seven species previously classified as Dermogenys (D. ebrardtii, D. megarrhamphus, D. weberi, D. vivipara, D. pectoralis, D. bakeri, D. philippina) as well as N. celebensis, N. towoetii, N. liemi, N. brembachi, N. kolonodalensis, N. pinnimaculata, N. manifesta and N. rossi. Nomorhamphus hageni is also tentatively included in this clade (see systematics accounts). Two species were not available for phylogenetic analysis. Nomorhamphus bakeri and N. philippina are placed in an unresolved position within Nomorhamphus because they possess two diagnostic characters of that genus: (1) absence of a well-defined geniculus; and (2) an oval lacrimal. In addition to confusion over species limits, there has also been some ambiguity concerning the relationships among the genera of internally fertilized halfbeaks. In previous hypotheses, Dermogenys and Nomorhamphus were placed in an unresolved polytomy with Hemirhamphodon and Tondanichthys (Collette, 1995). Tondanichthys, known only from immature specimens, was inferred to be viviparous from that preliminary phylogeny. In the present phylogenetic analysis, utilizing osteological data as well as reproductive characters, the sister group relationship between Hemirhamphodon and (Dermogenys+Nomorhamphus) is well supported. Tondanichthys is placed in an unresolved position with

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Table 35. Morphometrics of Nomorhamphus pinnimaculata sp. nov. (A) male holotype (ZRC 46173); (B) allotype (=paratype) (ZRC 46174); (C) 13 female paratypes; and (D) 16 male paratypes. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentage of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

D

41.0 7.6 1.4 37.3 64.6 39.3 34.4 16.3 16.3 6.3 8.0

39.7 9.9 1.5 34.3 58.2 41.6 31.7 15.6 15.6 6.0 6.3

35.7–49.6 6.3–20.6 1.4–1.7 32.6–36.7 61.0–63.7 39.1–43.2 30.1–33.9 14.6–16.6 14.6–16.6 5.0–7.0 7.2–8.4

35.6–41.1 5.6–12.4 1.2–1.6 34.1–38.6 60.9–64.7 39.9–42.8 31.3–35.1 15.2–18.0 15.2–18.0 5.5–6.5 7.2–8.8

Figure 68. Nomorhamphus rossi sp. nov. A, holotype, USNM 333262, male, 40.3 mm SL; Intel River, Cagayan Province, Luzon, Philippines; B, allotype (=paratype), USNM 363186, 100.1 mm SL, collected with the holotype.

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Figure 69. Diagrammatic representation of the anterior five modified anal-fin rays of the andropodium of Nomorhamphus rossi sp. nov. Middle radials absent and ossified distal radials not illustrated; cartilage not illustrated; bone is stippled. USNM 363187 (47.0 mm).

Zenarchopterus, suggesting that Tondanichthys may not be viviparous, a suggestion made also by Meisner & Collette (1999). It is interesting to note that several characters supporting the monophyly of the viviparous genera involve reductions in both the size and number of skeletal elements which may be correlated with an overall reduction in body size. Within the Atherinomorpha, internal fertilization has arisen independently several times, including once in the halfbeaks (Parenti, 1981). Atherinomorph fishes known to be obligately internally fertilized possess some type of intromittant organ. These organs are often modified anal fins, as they are in the halfbeaks. The role of the modified anal fin in halfbeaks is not clear. If in fact it functions in the transfer of sperm, there would be considerable contact with the aqueous environment, resulting in potential loss of sperm. The

transfer of spermatozoa in the form of some kind of packet may decrease the chance of such losses (Ginzburg, 1968; Burns et al., 1995). Indeed, all other atherinomorph fishes known to be internally fertilizing have evolved mechanisms that result in at least some degree of packaging. Within the halfbeaks, it is most parsimonious to infer that such mechanisms for sperm packaging evolved once at the level of Zenarchopterus. After initial appearance, sperm packaging mechanisms have undergone tremendous diversification. A unique form of spermatogenesis is found in Zenarchopterus, in which free spermatozoa are released into the testicular ducts and sperm are subsequently packaged into either spermatozeugmata or spermatophores. Six different sperm bundle morphologies have been described for this genus where only the anterior portion of the testis is spermatogenic (Grier & Collette, 1987). There is some uncertainty as to whether Zenarchopterus is internally fertilized. The presence of sperm bundles in all species and the observation of sperm within the ovary of one species suggests that these fishes are at least inseminated. Gross dissections and histological observations of the ovaries have not found any fertilized eggs so the timing of fertilization remains in question. Another unique method of sperm packet formation is found in Hemirhamphodon. In this genus, the sperm bundles are formed within the spermatocysts prior to spermiation and five different sperm bundle types have ben described (Downing & Burns, 1995). Yet another unique mechanism of sperm bundle formation is found in Dermogenys+Nomorhamphus. As in Hemirhamphodon, the spermatozeugmata are formed within spermatocysts and two sperm bundle morphologies are known (Downing & Burns, 1995). Although viviparity in these fishes has been documented since the original description of Dermogenys (see Roberts, 1993), details of reproductive biology were scarce. The information available indicates that viviparity is tremendously variable, even within a single genus. Superfetation has been reported in Hemirhamphodon (Brembach, 1976; Roberts, 1989). However, observations made in the aquarium literature suggest that H. tengah may lay fertilized eggs (Bork & Mayland, 1998). In a recent histological survey of the ovaries, viviparity in Dermogenys and Nomorhamphus has been divided into five unique types based on morphological modifications involving both maternal and embryonic tissues (Meisner & Burns, 1997b). To address questions concerning the evolution of viviparity in halfbeaks, it is first necessary to place the group into some historical phylogenetic context. This has been accomplished through the present study. The task of addressing such questions remains difficult, however, due to the lack of comparative knowledge of viviparity in Hemirhamphodon. Despite this,

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Table 36. Morphometrics of Nomorhamphus rossi sp. nov. (A) male holotype (USNM 333262); (B) allotype (=paratype); (C) 15 female paratypes; and (D) 14 male paratypes. Standard length is expressed in mm; measurements 1 and 2 are proportions; measurements 3–10 are expressed as percentages of standard length; abbreviations listed on p. 202

SL 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

SL/LJL UJL/UJW SN-P1 SN-P2 P2-C HDL BDP1 BDP2 ORBL INTORBL

A

B

C

D

40.3 5.3 1.3 36.7 61.3 42.7 33.0 15.4 15.4 7.2 6.7

100.1 7.5 1.3 32.4 61.2 40.3 28.5 13/6 14.5 4.5 7.7

42.3–86.2 6.3–8.3 1.2–1.7 32.2–33.6 58.6–62.3 39.1–42.8 26.5–31.7 11.6–14.2 11.8–15.2 4.4–6.9 7.8–8.6

39.3–46.9 4.5–6.9 1.2–1.7 32.3–37.7 58.3–62.6 40.4–43.1 30.5–34.4 12.2–16.0 15.0–18.0 5.1–18.0 6.8–8.2

some general statements about the evolution of viviparity in halfbeaks are possible. First, it appears that viviparity arose from the oviparous condition once, at the level of Hemirhamphodon. Intraluminal development appears to be the plesiomorphic condition due to its presence in Hemirhamphodon, with intrafollicular gestation being the derived state occurring in Dermogenys. One of the difficulties in discussing the evolution of the five types of viviparity is that these types appear to be unique to Dermogenys and Nomorhamphus. Aside from these very general conclusions, the current hypothesis of relationships does not allow for more precise statements concerning the order and evolution of the five types of viviparity identified in Dermogenys and Nomorhamphus. Characters of the reproductive biology of these fishes have been informative in inferring phylogenetic relationships at different taxonomic levels, including distinguishing between Dermogenys and Nomorhamphus as well as diagnosing more inclusive groups within each genus. This study presents a clear diagnosis of species as well as an hypothesis of the phylogenetic relationships among the species in these genera. Despite this new information, however, resolution of species relationships within several clades was not possible. The taxonomy of these species had been confused in the past partly by the lack of rigorous methodology in proposing species limits and relationships, but also by the reliance on ‘traditional’ meristic and morphometric characters at all taxonomic levels. One of the goals of phylogenetic analysis is to find the level in the hierarchy where characters are most informative. I conclude that characters of the anal fin are informative at the species level, histological characters of the gonads are informative both at diagnosing species groups

within genera and in distinguishing between Dermogenys and Nomorhamphus, and osteological characters are most informative at the generic level.

ACKNOWLEDGEMENTS I thank Lynne R. Parenti, John R. Burns, and Bruce B. Collette for all of their support, patience, and encouragement generously provided throughout this study. I would also like to thank Mary Sangrey and the Research Training Program at the National Museum of Natural History for sparking my initial interest in halfbeak systematics. For lively discussions of many aspects of my research, I thank Thomas A. Munroe. Funding for the collection of halfbeaks in Brunei was provided by the Pew Charitable Trust. For loan of specimens I thank: Maurice Kottelat (CMK); Horst Wilkens (ZMH); Gerald R. Smith, Doug W. Nelson, and William L. Fink (UMMZ); Mark W. Westneat, Barry Chernoff, and Mary Anne Rogers (FMNH); Kelvin Lim and C. M. Young (ZRC); Karsten Hartel (MCZ); David Catania, Tomio Iwamoto, and Carl J. Ferraris (CAS); Isaa¨c J. H. Isbru¨ker (ZMA); Scott A. Schaeffer (formerly ANSP); Melanie L.J. Stiassny and Barbara Brown (AMNH); Hans J. Paepke (ZMB); Tyson R. Roberts (MVT); Martien J. P. van Oijen (RMNH); Jean Claude Hureau (MNHN); and Nigel R. Merret (BMNH). For the gift of live specimens I thank John R. Burns. For technical assistance and loan coordination I thank Lisa Palmer, Jeff Clayton, Ruth Gibbons, Kris Murphy, and Sandra Raerdon. For computer assistance I thank J. Andrew Cooper. Keiko Hiratsuka Moore drew Fig. 31. Ruth Gibbons photographed Fig. 30.

278

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Submitted in partial fulfilment of the requirements for the degree of Doctor of Philosophy, The George Washington University, Washington, D.C.

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Clemen G, Wanninger AC, Greven H. 1997. The development of the dentigerous bones and teeth in the hemiramphid fish Dermogenys pusillus (Atheriniformes, Teleostei). Annals of Anatomy 179: 165–174. Collette BB. 1982. Two new species of freshwater halfbeaks (Pisces: Hemiramphidae) of the genus Zenarchopterus from New Guinea. Copeia 1982: 265–276. Collette BB. 1985. Zenarchopterus ornithocephala, a new species of freshwater halfbeak (Pisces: Hemiramphidae) from Vogelkop Peninsula of New Guinea. Proceedings of the Biological Society of Washington 98: 107–111. Collette BB. 1995. Tondanichthys kottelati, a new genus and species of freshwater halfbeak (Teleostei: Hemiramphidae) from Sulawesi. Ichthyological Exploration of Freshwaters 6: 171–174. Collette BB, McGowen GE, Parin NV, Mito S. 1984. Beloniformes: development and relationships. In: Mosher HG, Richards WJ, Kendall AW, Richardson SL, eds. Ontogeny and systematics of fishes. American Society of Ichthyologists and Herpetologists Special Publication, 1, 335–354. Day F. 1878. The fishes of India being a natural history of the fishes known to inhabit the seas and freshwaters of India, Burma, and Ceylon. New Dehli: 1971 reprint issued by Today and Tomorrow’s Book Agency. Dingerkus G, Uhler LD. 1977. Enzyme clearing of alcian blue stained whole small vertebrates for demonstration of cartilage. Stain Technology 52: 229–232. Downing AL, Burns JR. 1995. Testis morphology and spermatozeugma formation in three genera of viviparous halfbeaks: Nomorhamphus, Dermogenys, and Hemirhamphodon (Teleostei: Hemiramphidae). Journal of Morphology 225: 329–343. Fowler HW. 1934a. Zoological results of the third DeSchauensee Siamese expedition, part I Fishes. Academy of Natural Sciences Philadelphia 86: 67–163. Fowler HW. 1934b. Descriptions of new fishes obtained 1907–1910, chiefly in the Philippine islands and adjacent seas. Academy of Natural Sciences Philadelphia 85: 233– 367. Fowler HW, Bean BA. 1922. Fishes from Formosa and the Philippine islands. Proceedings of the United States National Museum 62: 1–73. Ginzburg AS. 1968. Fertilization in fishes and the problem of polyspermy. Moscow: Academy of Science USSR. Greven H, Wanninger AC, Clemen G. 1997. Dentigerous bones and dentition in the hemiramphid fish Dermogenys pusillus (Atheriniformes, Teleostei). Annals of Anatomy 179: 21–32. Grier HJ, Collette BB. 1987. Unique spermatozeugmata in testes of halfbeaks of the genus Zenarchopterus (Teleostei: Hemiramphidae). Copeia 1987: 300–311. Grier HJ, Fitzsimmons JM, Linton JR. 1978. Structure and ultrastructure of the testis and sperm formation in goodeid teleosts. Journal of Morphology 156: 419–438. Gu ¨ nther A. 1866. Catalogue of fishes of the British Museum VI. London: Taylor and Francis. Hauser DL, Presch W. 1991. The effect of ordered characters on phylogenetic reconstruction. Cladistics 7: 243–265.

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS Hennig W. 1966. Phylogenetic systematics. Urbana: University of Illinois Press. Herre AWCT. 1931. The fishes of the Herre Philippine Expedition of 1931. Notes on fishes in the Zoological Museum of Stanford University. Herre AWCT. 1944. A review of the halfbeaks or Hemiramphidae of the Philippines and adjacent waters. Stanford University Publications University Series Biological Sciences 9: 39–86. Herre AWCT. 1953. Check list of Philippine fishes. Research report 20. Washington, D.C. Inger RF. 1955. Ecological notes on the fish fauna of a coastal drainage of North Borneo. Fieldiana: Zoology 37: 47–90. Inger RF, Chin PK. 1962. The freshwater fishes of North Borneo. Fieldiana: Zoology 45. International Commission on Zoological Nomenclature. 1985. International code of zoological nomenclature. International Commission on Zoological Nomenclature. Berkeley: University of California Press. Jayaram KC. 1981. The freshwater fishes of India, Pakistan, Bangladesh, Burma and Sri Lanka. Calcutta: Sri Aurobindo Press. Jordan DS. 1923. The genera of fishes and a classification of fishes. Stanford: Stanford University Press. Jordan DS, Seale A. 1908. List of fishes in the river at Buytenzorg, Java by Dr Douglas Houghton Campbell. Proceedings of the United States National Museum 33: 535– 540. Kottelat M. 1985. Freshwater fishes of Kampuchea. Hydrobiologia 121: 249–279. Kottelat M. 1987. Nomenclature status of the fish names created by J.C. van Hasselt (1823) and some cobitoid genera. Japanese Journal of Ichthyology 4: 368–375. Kottelat M. 1992. Book review: Lebendgeba¨rende Halbschna¨bler – Untersuchungen zur Verbreitung, Morphologie, Systematik und Fortpflanzungsbiologie der lebendgeba¨renden Halbschna¨bler der Gattungen Dermogenys und Nomorhamphus (Hemirhamphidae [sic]: Pisces). Raffles Bulletin of Zoology 40: 267–272. Kottelat M, Whitten AJ, Kartikasari SN, Wirjoatmodjo S. 1993. Freshwater fishes of western Indonesia and Sulawesi. Jakarta: Periplus Editions Limited. Ladiges W. 1972. Zwei neue Hemirhamphiden von Celebes und Cebu (Philippinen). Mitteilungen aus dem Hamburgischen Zoologischen und Institut 68: 207–212. Leviton AE, Gibbs RH Jr, Heal E, Dawson CE. 1985. Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985: 802–832. Meisner AD, Burns JR. 1997a. Testis and andropodial development in the viviparous halfbeak Dermogenys sp. (Teleostei: Hemiramphidae). Copeia 1997: 44–52. Meisner AD, Burns JR. 1997b. Viviparity in the halfbeak genera Dermogenys and Nomorhamphus (Teleostei: Hemiramphidae). Journal of Morphology 234: 295–317. Meisner AD, Collette BB. 1998. A new species of viviparous halfbeak, Dermogenys bispina (Teleostei: Hemiramphidae)

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from Sabah (North Borneo). Raffles Bulletin of Zoology 46: 373–380. Meisner AD, Collette BB. 1999. Generic relationships of the internally fertilized southeast Asian halfbeaks (Hemiramphidae: Zenarchopterinae). In: Se´ret B, Sire J-Y, eds. Proc. 5th Indo–Pac. Fish Conf., Noume´a, 1997. Paris: Soc. Fr. Ichthyol, 69–76. Meisner AD, Louie KD. 2000. Nomorhamphus kolonodalensis, a new species of viviparous halfbeak from Sulawesi (Teleostei: Hemiramphidae). Ichthyological Exploration of Freshwaters 11: 361–368. Meyer MK. 1985. Lebendgeba¨rende Zierfische Arten der Welt. Melle, Germany: Verlag fu¨r Natur und Heimtierkunde. Mohr E. 1936a. Hemirhamphiden Studien IV. Die Gattung Dermogenys van Hasselt. Mitteilungen aus dem Zoologischen Museum in Berlin 21: 34–55. Mohr E. 1936b. Hemirhamphiden Studien V. Die Gattung Nomorhamphus Weber and de Beaufort. Mitteilungen aus dem Zoologischen Museum in Berlin 21: 55–58. Mukerji DD. 1935. Description of a new species of hemirhamphid fish, Dermogenys burmanica from lower Burma, with notes on sexual dimorphism and its taxonomic significance. Records of the Indian Museum 37: 213–218. Nelson G, Platnick N. 1981. Systematics and biogeography: Cladistics and vicariance. New York: Columbia University Press. Parenti LR. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei, Atherinomorpha). Bulletin of the American Museum of Natural History 168: 341–557. Parenti LR, Meisner ALD. 1995. Fishes of the Belait River, Brunei Darussalam, Borneo. Brunei Museum Journal (in press). ¨ ber lebedig geba¨rende Arten der FischPeters UCH. 1865. U gattung Hemiramphus. Auszug aus dem Monatsbericht der Ko¨nigl Akademie der Wissenschaften zu Berlin 1865: 132–133. ¨ ber die van Hrn. Dr. F. Jagor in Peters UCH. 1868. U dem ostindischen Archipel gesammelten und dem Ko¨nigl. Zoologischen Museum u¨bergebenen Fische. Auszug aus dem Monatsbericht der Ko¨nigliche Akademie der Wissenschaften zu Berlin 1868: 254–281. Popta CML. 1912. Fortsetzung der Beschreibung von neuen Fischarten der Sunda Expedition. Notes from the Leyden Museum 34: 185–193. Roberts TR. 1989. The freshwater fishes of Western Borneo (Kalimantan Barat, Indonesia). Memoirs of the California Academy of Sciences 14: 1–210. Roberts TR. 1993. The freshwater fishes of Java, as observed by Kuhl and van Hasselt in 1820–23. Zoologische Verhandelingen (Leiden) 285: 1–94. Rosen DE. 1964. The relationships and taxonomic position of the halfbeaks, killifishes, silversides and their relatives. Bulletin of the American Museum of Natural History 127: 219–267. Rosen DE, Parenti LR. 1981. Relationships of Oryzias, and the groups of atherinomorph fishes. American Museum Novitates 2719: 1–25.

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Swofford DL. 1993. PAUP: Phylogenetic Analysis Using Parsimony, ver. 3.1.1. Program and documentation. Urbana: Illinois Natural History Survey. Talwar PK, Jhingram AG. 1991. Inland fishes of India and adjacent countries. vol. 2. New Delhi: Oxford and IBH Publishing Co. Pvt. Ltd. van Hasselt JC. 1823. Uittreksel uit een’ brief van den Heer J.C. van Hasselt, aan den Heer C.J. Temminck, geschreven uit Tjecande, Residentie Bantam, den 29sten Dec 1822. Algemeene Konst- en Letter Bode voor het jaar 1823, II, Deel 35: 130–133. Vogt D. 1978a. Kennen Sie Nomorhamphus? Die Aquarien und Terrarien Zeitschrift 31: 222–226. Vogt D. 1978b. Der Fisch wird teuer bezahlt im Hochland von Maros mit einer vor la¨ufigen Beschreibung von Nomorhamphus liemi n. sp. Die Aquarien und Terrarien Zeitschrift 1: 5–9. Weber M. 1894. Die Su¨sswasser Fische des indischen Archipels, Nebst Bemerkungen u¨ber den Ursprung der Fauna von Celebes. Zoologische Ergebnisse einer Reise in Niederla¨ndisch Ost–Indien 1894: 405–476. Weber M. 1913. Neue Beitra¨ge zur Kenntnis der Su¨sswasserfische von Celebes. Ergebnisse einer Reise von E.C. Abendanon in Celebes 1913: 197–213. Weber M, de Beaufort LF. 1922. The fishes of the Indo– Australian Archipelago IV. Heteromi, Solenichthyes, Synentognathi, Percesoces, Labyrinthici, Microcyprini. Leiden, E.J. Brill. 4: 1–410. Weed AC. 1933. Notes on the fishes of the family Hemirhamphidae [sic]. Field Museum of Natural History, Zoological Series 20: 41–65.

APPENDIX 1 CHARACTERS AND STATES 1. Premaxilla. 0: right and left bones form triangular plate, approximating an equilateral triangle; 1: right and left bones form narrow-based triangular plate (UJL/ UJW >1.0); 2: right and left bones form a broad plate, roughly trapezoidal, upper jaw approximately as long as wide. 2. Uniserial conical teeth extending medially on premaxillae. 0: oral teeth form up to four rows along outer edge of upper jaws; 1: row of teeth extends medially in a concave row from outer rows of teeth to a point at about one half the length of premaxilla. 3. Five or six rows of conical teeth present anteriorly on premaxillae. 0: follows description for character 2; 1: large patch of conical teeth present anteriorly on upper jaw. 4. Maxilla. 0: short, distal half of bone oriented vertically, proximal half oriented slightly anterior; 1: elongate, thin, distal half angled anteriorly. 5. Lower jaw. 0: lower jaw extends past upper jaw forming a well-defined ‘beak’; 1: lower jaw not elongate. 6. Nasal fossa. 0: large; 1: small. 7. Nasal papilla. 0: short, spatulate; 1: elongate, extending out of nasal fossa. 8. Epibranchial one. 0: epibranchial one largest of four epibranchials, extends anteriorly and ventral, passes

anterior extent of pharyngobranchial three, base only slightly expanded; 1: reduction in length of epibranchial one to approximately length of epibranchial four; base expanded. 9. Epibranchial four. 0: ramus on posterior surface near point of articulation with pharyngobranchial three; 1: ramus absent. 10. Epibranchial four. 0: epibranchial four straight with surface articulating with pharyngobranchial three expanded; 1: curved slightly at mid length, proximal articular surface just slightly expanded; 2: proximal and distal articular surfaces reduced, each oriented posteriorly. 11. Pharyngobranchial three. 0: pharyngobranchial three elements fused forming a broad plate that narrows posteriorly, elongate along anterior-posterior axis; 1: fused bones form a broad plate, expanded posteriorly. 12. Anterior processes of pharyngobranchial three. 0: processes in line with anterior–posterior axis; 1: processes offset laterally. 13. Gill rakers. 0: elongate, many teeth along entire length; 1: reduced, roughly tear-drop shaped, one to three teeth; 2: reduced, base wide, many teeth on dorsal surface; 3: reduced, more or less tear-drop shaped, many teeth on dorsal surface. 14. Lacrimal. 0: round, about as long as wide; 1: oval, much longer than wide. 15. Opercle. 0: large, extends dorsally to contact skull; 1: reduced, does not extend dorsally past midpoint of hyomandibula. 16. Endopterygoid. 0: slender; 1: expanded, flange along anterior surface. 17. Autopalatine. 0: slender; without flange; 1: short, flange along anterior edge; 2: elongate, flange along anterior surface. 18. Post-temporal process. 0: absent, appears as flange or nub near point of articulation with cleithrum; 1: elongate process on anterior surface, near point of articulation with cleithrum. 19. Pectoral fin position. 0: pectoral girdle vertical, radials oriented dorsally, with high set pectoral fins; 1: pectoral girdle more ventrally oriented with the radials oriented caudad. 20. Pelvic girdle. 0: anterior extension of pelvic bones angled toward midline, anterior tips in close proximity; 1: pelvic bones widely separated, anterior processes lie directly along axis of body. 21. Ventral extension of the pelvic girdle. 0: thin, bifurcated; 1: ventral extension with an entire, expanded flange. 22. Anterior lateral extension of pelvic girdle. 0: absent; 1: present. 23. First dorsal pterygiophore. 0: ventral process present, dorsoanterior extension slight; 1: ventral process greatly reduced or absent, dorsoanterior extension expanded. 24. Shape of the caudal fin. 0: forked; 1: round or slightly notched. 25. Caudal skeleton. 0: expanded uroneural, neural plates, and epurals; 1: reduction in size of bony elements. 26. Parhypural. 0: autogenous; 1: fused to ventral hypural plate along its length. 27. Fifth hypural. 0: fused only proximally to dorsal hypural plate; 1: fused to dorsal hypural plate along its length.

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS 28. Origin of the anal fin. 0: posterior to dorsal fin origin. 1: anterior to dorsal fin origin. 29. Anal-fin rays one through seven in males. 0: unmodified; 1: modified into an andropodium. 30. First anal pterygiophore in males. 0: unmodified, not differing from more posterior radials; 1: thick, lateral flanges present along most of length; 2: thin, angled sharply anterior. 31. Geniculus. 0: absent or only very weakly developed; 1: present as a distinct, thinning of second ray at segment three or four. 32. Cap of bone on paired spines, second anal-fin ray in males. 0: absent; 1: present. 33. Mode of spermatogenesis. 0: spermatids unorganized within spermatocysts during spermiogenesis; 1: spermatids distributed unevenly around periphery of spermatocysts; 2: spermatids evenly distributed around periphery of spermatocysts. 34. Sperm bundles in which the spermatid nuclei are

281

oriented toward one side and the flagella towards the other. 0: absent; 1: large; 2: small. 35. Thread-like sperm nuclei. 0: absent; 1: present. 36. Insemination. 0: absent; 1: present. 37. Genital papilla in females. 0: inconspicuous or only slightly enlarged, unpigmented; 1: elongate, pigmented. 38. Melanophores anterior to the anal fin in females. 0: diffuse; 1: arranged into a distinct spot. 39. Site of gestation. 0: oviparous; 1: development intrafollicular until parturition; 2: significant period of intraluminal gestation; 3: embryos spend an intermediate period of time developing within ovarian lumen. 40. Type I form of viviparity. 0: absent; 1: present. 41. Type II form of viviparity. 0: absent; 1: present. 42. Type III form of viviparity. 0: absent; 1: present. 43. Type IV form of viviparity. 0: absent; 1: present. 44. Type V form of viviparity. 0: absent; 1: present. 45. Ovigerous ridge. 0: absent or only weakly developed; 1: well-developed.

1–5

00000 00000 00000 00000 10010 11010 11010 11010 11010 11010 11010 11010 11010 11010 11010 20111 20011 10010 20111 10010 10010 10011 10010 10010 10010 10010 10010 10010

Character

Hemiramphus Hyporhamphus Zenarchopterus Tondanichthys Hemirhamphodon Dermogenys burmanica D. bispina D. bruneiensis D. pusilla D. orientalis D. siamensis D. sumatrana D. robertsi D. palawanensis D. collettei Nomorhamphus brembachi N. celebensis N. ebrardtii N. liemi N. megarrhamphus N. pectoralis N. towoetii N. vivipara N. weberi N. manifesta N. pinnimaculata N. rossi N. kolonodalensis 00000 00000 11000 11000 11112 11111 11112 11112 11111 11112 11111 11112 11112 11112 11111 11112 11112 11112 11112 11112 11112 11112 11112 11112 11112 11112 11112 11112

6–10 00000 00000 10000 10000 11101 11101 11101 11101 11101 11101 11101 11101 11101 11101 11101 11211 11311 11311 11211 11311 11111 11311 11111 11311 11111 11111 11111 11111

11–15 00000 00000 00110 00110 00111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 12111 12111 12111 12111 12111 12111 12111 12111 12111 12111 12111 12111 12111

16–20 00000 00000 00010 00010 10011 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111 11111

21–25 00000 00000 00000 00000 10000 10111 10111 10111 10111 10111 10111 10111 10111 10111 10111 11111 11111 11111 11111 11112 10111 11111 10111 11112 10111 10111 10111 10111

26–30 00000 00000 00100 00??? 00100 10210 11211 11211 10210 10211 10210 11211 11211 11211 10210 00220 00220 00220 00220 00220 00220 00220 00220 00220 00220 00220 00220 00220

31–35

0=plesimorphic condition; 2=apomorphic condition; ?=unknown character state.

APPENDIX 2: Character Matrix

00000 00000 10000 ????? 10030 10111 11110 11110 10111 11110 10111 11110 10110 11110 10111 100?? 100?? 10020 10020 10020 10020 10020 10020 10020 10020 10020 10020 10020

36–40

00000 00000 00000 ????? 00000 00000 10000 10000 00000 10000 00000 10000 10000 10000 00000 ????? ????? 00011 00101 00101 01001 00101 00101 00101 01001 00101 01001 00101

41–45

282 A. D. MEISNER

SYSTEMATIC REVISION OF DERMOGENYS AND NOMORHAMPHUS APPENDIX 3 INDEX OF SPECIES (NEW COMBINATIONS AND SPECIES IN BOLD)

Genus Dermogenys Kuhl & van Hasselt in van Hasselt, 1823 ............................................................. Dermogenys brachynotopterus (Bleeker, 1853a) ..... Dermogenys burmanica Mukerji, 1935 ................... Dermogenys orientalis (Weber, 1894) ...................... Dermogenys pusilla Kuhl & van Hasselt in van Hasselt, 1823 ............................................................. Dermogenys siamensis Fowler, 1934a .................... Dermogenys sumatrana (Bleeker, 1853b) ............... Dermogenys vogti Brembach, 1982 .......................... Dermogenys bispina Meisner & Collette, 1998 ...... Dermogenys bruneiensis sp. nov. ....................... Dermogenys robertsi sp. nov. .............................. Dermogenys palawanensis sp. nov. ................... Dermogenys collettei sp. nov. .............................. Genus Nomorhamphus Weber & de Beaufort, 1922 Nomorhamphus bakeri (Fowler & Bean, 1922) comb. nov. ................................................................

215 218 219 222 224 227 229 231 231 235 236 237 238 242 244

Nomorhamphus brembachi Vogt, 1978a ................. Nomorhamphus celebensis Weber & de Beaufort, 1922 ........................................................................... Nomorhamphus ebrardtii (Popta, 1912) comb. nov. ............................................................................ Nomorhamphus hageni (Popta, 1912) ..................... Nomorhamphus liemi Vogt, 1978 ............................ Nomorhamphus megarrhamphus (Brembach, 1982) comb. nov. ..................................................... Nomorhamphus pectoralis (Fowler, 1934b) comb. nov. ................................................................ Nomorhamphus philippina (Ladiges, 1972) comb. nov. ................................................................ Nomorhamphus towoetti Ladiger, 1972 .................. Nomorhamphus vivipara (Peters, 1865) comb. nov. ............................................................................ Nomorhamphus weberi (Boulenger, 1897) comb. nov. ............................................................................ Nomorhamphus kolonodalensis Meisner & Louie, 2000 ........................................................................... Nomorhamphus manifesta sp. nov. ................... Nomorhamphus pinnimaculata sp. nov. ......... Nomorhamphus rossi sp. nov. ............................

283 245 249 251 252 253 255 258 259 261 263 266 268 270 271 272