PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3440, 20 pp., 8 ®gures, 2 tables May 14, 2004

Review of the Malagasy Sicydiine Gobies (Teleostei: ), with Description of a New Species and Comments on the Taxonomic Status of Gobius lagocephalus Pallas, 1770

JOHN S. SPARKS1 AND DOUGLAS W. NELSON2

ABSTRACT Malagasy sicydiine gobies are reviewed, compared with other members of the subfamily in the Mascarene region, and a new species belonging to the Sicyopterus Gill, 1860 is described on the basis of material collected in northeastern . The new species differs from Sicyopterus franouxi (Pellegrin, 1935), the only other known species of sicydiine goby inhabiting the freshwaters of Madagascar, in the number of branched rays in the second dorsal ®n, the color pattern of the body and ®ns, the number and shape of the premaxillary teeth, and the size and number of scales on the nape and abdomen. Sicyopterus franouxi,a taxon described on the basis of a single juvenile specimen, is redescribed based on adult material. The taxonomic status of Gobius lagocephalus Pallas, 1770, a nominal sicydiine species incorrectly ascribed to the Mascarene islands of the western Indian Ocean, is discussed. This species is herein concluded to be a nomen dubium of uncertain placement beyond the subfamilial level.

INTRODUCTION tropical and subtropical swift-¯owing streams and rivers in the Indo-Paci®c region, Freshwater gobies of the genus Sicyopte- islands of the southwestern Paci®c Ocean, rus Gill, 1860 (subfamily ) inhabit and islands of the Mascarene region in the

1 Department of Ichthyology, Division of Vertebrate Zoology, American Museum of Natural History; reprint re- quests to [email protected] 2 Division of Fishes, Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48109; e-mail: dwnelson@ umich.edu

Copyright ᭧ American Museum of Natural History 2004 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3440 western Indian Ocean. These small ®shes are (®g. 1), and compare the new taxon with S. often referred to as ``rock-climbing gobies'' franouxi (Pellegrin, 1935), the only other si- due to their ability to ascend steep, torrential cydiine goby known to inhabit the freshwa- streams and waterfalls (Fukui, 1979; Parenti ters of Madagascar. Pellegrin described S. and Maciolek, 1993; Balon and Bruton, franouxi on the basis of a single juvenile 1994). Throughout their range, sicydiine go- specimen. Herein, we redescribe this taxon bies exhibit a high degree of island-group en- on the basis of adult material. We also dis- demism (Parenti and Maciolek, 1993, 1996; cuss the of Gobius lagocephalus this study). Two species of Sicyopterus are Pallas, 1770, a nominal sicydiine taxon his- presently known to inhabit the freshwaters of torically and incorrectly ascribed to Mada- Madagascar (Sparks and Stiassny, 2003): gascar and other islands of the Mascarene re- One of these is herein described as new to gion. A summary of the taxonomic conclu- science. sions of this study is presented in table 1. Monophyly of sicydiine gobies is well supported by morphological evidence MATERIALS AND METHODS (Hoese, 1984; Harrison, 1989; Parenti and Maciolek, 1993; Birdsong et al., 1998; Par- Representatives of the new species are de- enti and Thomas, 1998). Parenti and Maci- posited at the University of Michigan Mu- olek (1993) and Parenti and Thomas (1998) seum of Zoology, Ann Arbor (UMMZ), the recognized ®ve genera within the subfamily Museum National d'Histoire Naturelle, Paris Sicydiinae: Sicydium Valenciennes, in Cuvier (MNHN), and at the American Museum of and Valenciennes, 1837, Sicyopterus Gill, Natural History, New York (AMNH). Insti- 1860, Lentipes GuÈnther, 1861, Sicyopus Gill, tutional abbreviations for material examined 1863, and Stiphodon Weber, 1895. On the ba- follow Leviton et al. (1985). Materials ex- sis of a low number of premaxillary teeth (7± amined are listed in appendix 1. 23 vs. Ͼ25 in other Sicydiinae), Watson Osteological characters of the new species (1995a) recognized a sixth sicydiine genus and related taxa were examined using from ReÂunion and Mauritius, Cotylopus Gui- cleared-and-stained individuals (CS), radio- chenot, 1863. The subfamily comprises graphs, or from scanning electron micro- about 100 nominal species; approximately 40 scope (SEM) images. SEM images of dried of these species are presently placed in the and coated premaxillary bones were pro- genus Sicyopterus (Parenti and Maciolek, duced using a Hitachi S4700 Field Emission 1996; Eschmeyer, 1998). Sicyopterus and Si- Scanning Electron Microscope (FE-SEM). cydium are hypothesized to be sister genera Premaxillary tooth counts were taken from based on the following derived features: an either the left or right element; counts are uninterrupted oculoscapular canal extending approximations due to the uncertainty asso- posteriorly from the eye to the posterior mar- ciated with estimating missing teeth. Gaps gin of the opercle, with fusion of oculosca- were included in the tooth count if the gap pular-canal pores H and K (Akihito et al., width was judged to exceed tooth width in 1984; Pezold, 1993), and a blunt ascending that region and it was apparent from the in- process of the premaxilla (Parenti and Ma- sertion point that a tooth was missing. Spec- ciolek, 1993). Akihito and Meguro (1979) imens were cleared and stained using a mod- presented and discussed a number of features i®ed protocol based on Taylor and Van Dyke useful for distinguishing between members (1985). Morphometric measurements were of Sicydium and Sicyopterus. Sicyopterus has recorded to the nearest 0.1 mm using digital been hypothesized to be monophyletic on the or dial calipers. Standard length (SL) is used basis of a marked medial gap between the throughout. Body depth A was measured as left and right premaxillary tooth rows (Par- a vertical transect at the origin of the pelvic enti and Maciolek, 1993, 1996). ®n, body depth B as a vertical transect at the In this paper we review the species of si- origin of the second dorsal ®n, and body cydiine gobies inhabiting Madagascar and depth C at the least depth of the caudal pe- the Mascarene region, describe a new species duncle. The distance between the last spinous of Sicyopterus from northeastern Madagascar ray of the ®rst dorsal ®n and the ®rst ray of 2004 SPARKS AND NELSON: MALAGASY SICYOPTERUS 3

Fig. 1. Map of northern Madagascar illustrating drainages from which the new species has been collected. Solid circles may represent more than a single collecting locality. the second dorsal ®n, as well as the lengths the cephalic seismosensory system follows of the bases of the dorsal and anal ®ns, were that of Akihito et al. (1984). measured from radiographs. Vertebral counts included the ural centrum (ϭ last half-cen- trum). Vertebral and ®n spine/ray counts INSTITUTIONAL ABBREVIATIONS were obtained from radiographs. The termi- nal dorsal- and anal-®n rays, which are AMNH American Museum of Natural History, branched to the base of the ®n, were counted New York as a single element. Transverse scale rows MNHN Museum National d'Histoire Naturelle, were counted from the dorsal margin of the Paris gill opening to the caudal ¯exure (Parenti NHRM Naturhistoriska Riksmuseet, Stockholm and Maciolek, 1993). Scale counts are ap- RUSI J.L.B. Smith Institute of Ichthyology, Grahamstown proximations, due to high intra- and inter- UMMZ University of Michigan, Museum of speci®c variability, irregular arrangement, Zoology, Ann Arbor and because small scale size and the degree USNM National Museum of Natural History, to which scales are embedded makes accu- Smithsonian Institution,Washington, rate counts problematic. Nomenclature for D.C. 4 AMERICAN MUSEUM NOVITATES NO. 3440

TABLE 1 Taxonomic Conclusions of the Present Study

SYSTEMATIC ACCOUNTS 1933: 153±154, pl. 2, ®g. 5; Pellegrin, 1935: 72; Arnoult, 1959: 99; Catala, 1982: 60. Sicyopterus franouxi (Pellegrin, 1935) Sicyopterus fasciatus (misidenti®cations): Kiener, Figures 2±3, 5A, 6A, B 1963: 64±65, pl. 33; MaugeÂ, 1986: 383; Stiass- Sicydium lagocephalum (nomen dubium): Pollen, ny and Raminosoa, 1994: 139; Stiassny and 1875: 6; Sauvage, 1891: 520; Catala, 1982: 60. Harrison, 2000: 151±153. (nomen dubium): Kie- Sicydium franouxi Pellegrin, 1935: Catala, 1982: ner, 1963: 64, pl. 33; MaugeÂ, 1986: 383±384; 60. Reinthal and Stiassny, 1991: 234; Stiassny and TYPE MATERIAL EXAMINED: MNHN 1935± Raminosoa, 1994: 139; Stiassny and Harrison, 0017, holotype, juvenile, 44 mm SL, sex un- 2000: 151±153. determined; southeastern Madagascar, region Sicydium laticeps (misidenti®cations): Pollen, 1875: 6; Sauvage, 1891: 378, 520, 531, pl. 40A, of Ranopitso (Fort-Dauphin), Ankondro Riv- ®g. 2, 2a, pl. 47, ®g. 5. er, near to Tsimelahy; R. Catala. Sicyopterus laticeps (misidenti®cation): Kiener, ADDITIONAL NONTYPE MATERIAL EXAM- 1963: pl. 33. INED: Total of 33 specimens, all from Mad- Sicydium fasciatum (misidenti®cations): Pellegrin, agascar. AMNH 97068 (1, 39.0 mm SL), Ta- 2004 SPARKS AND NELSON: MALAGASY SICYOPTERUS 5

Fig. 2. Sicyopterus franouxi, holotype, MNHN 1935-0017, 44.4 mm SL, juvenile, Madagascar: region of Ranopitso (Fort-Dauphin), Akondro River, near Tsimelahy. matave Province, drainage, 97149 (4, 76.2±110.1 mm SL), Tamatave stream by Ambinanindra village, M.L.J. Province, Mangoro River drainage, Nosivolo Stiassny, P.N. Reinthal, and G.J.P. Naylor, 19 River below Ampasimaniona village, 16 km Sept. 1990; AMNH 97071 (3, 53.4±84.5 mm ENE of Marolambo, M.L.J. Stiassny, P.N. SL), Tamatave Province, Mangoro River Reinthal, and G.J.P. Naylor, 21 Sept. 1990; drainage, Sahala River near Andranovolo, AMNH 215495 (2, 113.9±131.0 mm SL), M.L.J. Stiassny, P.N. Reinthal, and G.J.P. Andapa Region, , downstream Naylor, 19 Sept. 1990; AMNH 97080 (7, of Belaoko, P. de Rham, 21 Oct. 1993; 62.9±95.3 mm SL), Tamatave Province, MNHN 1891±0731 (1, 95.0 mm SL, adult Mangoro River drainage, be- male), Tamatave Province, Sahamandrevo; low Ampasimaniona village, 26 km ENE of MNHN 1891±0732 (1, 94.0 mm SL, adult Marolambo, M.L.J. Stiassny, P.N. Reinthal, female), Tamatave Province, Sahamandrevo; and G.J.P. Naylor, 20 Sept. 1990; AMNH MNHN 1891±0733 (1, 80.5 mm SL, likely

Fig. 3. Representative adult Sicyopterus franouxi. A, UMMZ 234878, 87.0 mm SL, male, north- eastern Madagascar: Masoala Peninsula, River. B, UMMZ 234878, 83.1 mm SL, female, northeastern Madagascar: Masoala Peninsula, Ankavanana River. 6 AMERICAN MUSEUM NOVITATES NO. 3440 male), Tamatave Province, Sahamandrevo; posteriorly to midpoint of second dorsal ®n. MNHN 1891±0734 (1, 71.0 mm SL, male), Anal ®n with one weak spine and 10 Tamatave Province, Sahamandrevo; MNHN branched rays. Pectoral ®n large and fan- 1966±1081 (2, 78.8±89.0 mm SL), Majunga shaped, tips of ®n rays slightly exserted from Province, Sahatelo River, Kiener and There- edge of ®n membrane, but not elongate or zien, Oct. 1962; UMMZ 234874 (2, 78.8± ®lamentous. Caudal ®n rounded. Pelvic ®ns 78.9 mm SL), Masoala Peninsula, Lohanto- fused into strong, cuplike suction disc. Total zona River, JSS 94-32, J.S. Sparks, K.J. Ris- vertebral count 25±26: 10 precaudal ϩ 15 eng, and guides, 9 Sept. 1994; UMMZ caudal, 11 precaudal ϩ 14 caudal, 10 pre- 234876 (1, 74.6 mm SL), Antalaha Province, caudal ϩ 16 caudal. Three preopercular canal Masoala Peninsula, Ankavanana River (15Њ pores (MЈ, N, and OЈ) present. Oculoscapu- 18Ј 14Љ S, 50Њ 13Ј 20Љ E), JSS 94-42, J.S. lar-canal pore pattern comprised of AЈ,B,C, Sparks, K.J. Riseng, and guides, 27 Sept. D (single), F, HK, and LЈ (®g. 4). Pore AЈ 1994; UMMZ 234878 (3, 83.1-87.0 mm SL), located just anterior and medial to anterior Antalaha Province, Masoala Peninsula, An- nasal tube; pore B is just anterior and medial kavanana River (15Њ 18Ј 35Љ S, 50Њ 14Ј 08Љ to posterior nasal opening. Pattern of free E), JSS 94-40, J.S. Sparks, K.J. Riseng, and neuromasts (pit organs) on head and anterior guides, 25 Sept. 1994; UMMZ 234880 (3, part of body similar to that of the new spe- 72.3-86.5 mm SL, 1 ex. CS), Antalaha Prov- cies (see description below; ®g. 4). ince, Masoala Peninsula, large unnamed trib- MOUTH AND TEETH: Mouth subterminal utary of Ankavanana River, JSS 94-45, J.S. and wide. Upper jaw short, extends posteri- Sparks, K.J. Riseng, and guides, Sept. 1994; orly at most to level of vertical through an- UMMZ 236538 (1, 97.0 mm SL), Fianarant- terior margin of orbit (®g. 5A). Upper lip soa Province, Parc National de Ranomafana, broad, lower edge weakly crenulate. Sym- above waterfall (21Њ 15Ј 32Љ physeal cleft in upper lip shallow and poorly S, 47Њ 25Ј 17Љ E), J.S. Sparks and K.J. Ris- developed. Lateral cleft wide throughout, not eng, Aug. 1994. expanded dorsally (®g. 5A). Single row of DIAGNOSIS: A species of Sicyopterus dis- ¯eshy, lightly pigmented papillae present be- tinguished from congeners by the following tween upper lip and premaxilla; papillae at combination of characters: a broad, darkly symphyseal and lateral cleft larger than oth- pigmented, midlateral band extending the ers in row. Enlarged lateral papilla expanded length of the body and onto the caudal ®n; laterally into area of cleft. Premaxillae sep- an absence of spotting on the posterior ¯ank arated by median gap. Premaxillary teeth tri- and caudal-®n base; second dorsal ®n with cuspid; cusps short and broad, including me- one weak spine and 10 branched rays; broad dian cusp (®g. 6A, B). Tooth shaft expanded lateral cleft in upper lip; and scales on the at distal end, near articulation with tooth nape, abdomen, and behind the pectoral-®n crown, and proximal to insertion on premax- axil markedly reduced in size compared to illa; shaft tapered medially (®g. 6B). Artic- scales on the sides of the body. ulation of tooth shaft and crown forms prom- DESCRIPTION:BODY AND FINS: Selected inent angled ridge (®g. 6B). Premaxillary proportional measurements and meristic data tooth count (approximately) 53 to 59, ar- are presented in table 2. Body cylindrical an- rayed in single functional row. Numerous teriorly; becoming progressively laterally rows of replacement teeth (Mochizuki and compressed posterior to second dorsal-®n or- Fukui, 1983) present on upper jaw. Dentary igin. Head large, blunt, and rounded in lateral teeth composed of outer row of setiform (la- pro®le. First dorsal ®n with six weakly de- bial) teeth forming horizontal plate and an veloped spines; second dorsal ®n with one inner row of caniniform, slightly curved weak spine and 10 branched rays. Sixth spine teeth; anteriormost one to two and posteri- of ®rst dorsal ®n widely separated from pre- ormost two to three of these caniniform teeth ceding spine. First dorsal ®n more elongate largest. Gill rakers weakly developed. than second; third spine longest. This spine SQUAMATION: Body covered with strongly markedly longer in males than in females. In ctenoid scales; frequently irregularly ar- adult males, depressed third spine extends ranged. Head and underside of body anterior 2004 SPARKS AND NELSON: MALAGASY SICYOPTERUS 7 S. franouxi , n.sp., and TABLE 2 Sicyopterus punctissimus Morphometric and Meristic Data of Measurements (mm) in percent standard length (SL) or percent head length (HL), unless noted otherwise. 8 AMERICAN MUSEUM NOVITATES NO. 3440

Fig. 4. Diagrammatic illustration of oculoscapular-canal pore pattern and pattern of free neuromasts (ϭ pit organs) on head and anterior part of body of S. franouxi and S. punctissimus. A, left lateral view. B, dorsal view. Abbreviations: an, anterior nasal tube; pn, posterior nasal opening; AЈ,B,C,D,F,HK, LЈ,MЈ,N,OЈ, cephalic seismosensory pores (see text).

to pelvic disk naked. Scales of nape, abdo- posterior edge of pelvic disk and anus. Pre- men, and immediately behind pectoral-®n dorsal scale count 26±31. axil markedly reduced in size compared to SEXUAL DIMORPHISM: Third spine of ®rst those on ¯anks. Scales in axil of pectoral ®n dorsal ®n in males markedly elongate. Pig- and under free edges of pelvic disk embed- mentation and coloration of females less pro- ded in integument. Caudal ®n scaled over nounced. Males much darker overall, nearly one-third of length; scales markedly reduced solid black in life, with several lighter gold- in size posteriorly on scaled part of ®n. Ap- en-brown bars on ¯anks. Size and shape of proximately 65±71 rows of scales in lateral urogenital papilla differs between sexes, as series. Approximately 27±37 scales between has been reported for other sicydiine gobies 2004 SPARKS AND NELSON: MALAGASY SICYOPTERUS 9

brown to iridescent golden bars on ¯anks. Dark, wide midlateral stripe discernible only in females and more lightly pigmented males. Midlateral stripe extends length of body. Females brownish and considerably lighter overall in coloration. Females light brown to whitish on throat, belly, and dorsal to anal-®n base. Rays of second dorsal ®n, in both males and females, with dark brown spots. No spotting evident on ¯anks or near caudal base of either sex. PIGMENTATION AND COLORATION IN PRES- ERVATION (FIGS. 2, 3): Upper part of head and body dark brown in both sexes. Males nearly uniform brown to grayish-brown. Lower part of head, beginning at level of upper lip, whit- ish to very light brown. Thin, curved, sub- orbital bar extends anteroventrally from orbit to upper lip. Wide, brown, midlateral stripe present and more evident in females, which are lighter overall. Midlateral stripe begins on head and terminates on anterior region of caudal ®n. Faint brownish vertical bars dis- cernible on ¯anks in females, especially above midline. Abdomen and pelvic disk light brown in females, medium brown to grayish in males. Pectoral ®ns and caudal ®n of males nearly uniform dark brown or gray, margins somewhat lighter. Pectoral ®ns of fe- males light to medium brown with dark brown rays. Rays of ®rst dorsal and caudal ®n dark brown or gray in males, dark brown Fig. 5. Left lateral view of head of (A) Si- in females. Anal ®n of males dark grayish- cyopterus franouxi, UMMZ 234878, 87.0 mm SL brown; distal margin of ®n lighter brown. and (B) Sicyopterus punctissimus, AMNH Anal ®n of females brown to grayish proxi- 226656, paratype, 108.7 mm SL. Upper lip stip- mal to base, followed by black submarginal pled to facilitate visualization of lateral cleft. band, and light brown along margin. Rays of second dorsal with dark brown spots in both males and females. No spotting on ¯anks or (e.g., Parenti and Maciolek, 1993; Watson, near caudal-®n base. 1995a, 1995b). Urogenital papilla of males DISTRIBUTION AND HABITAT: Although S. bulbous and rounded distally; in females pa- franouxi exhibits a somewhat patchy distri- pilla bilobed and tapered distally. Gap pre- bution throughout its range, the species has sent between distal tip of genital papilla and been collected from drainages spanning near- anal-®n origin in females; in males, distal tip ly the entire eastern coast of Madagascar. We of genital papilla extends to anal-®n origin. have examined a single lot of S. franouxi PIGMENTATION AND COLORATION IN LIFE: (MNHN 1966-1081) that was reportedly col- Males black on head, dorsum, and ¯anks, lected in northwestern Madagascar, in the re- whitish on throat and belly. Some males, pre- gion of Mahajunga (ϭ Majunga). We know sumably exhibiting breeding coloration, of no other records of S. franouxi from west- nearly solid black, with only small whitish or ern Madagascar, and the species has not been silvery region on throat and belly. Solid collected in a number of recent surveys fo- black males characterized by several golden- cusing on that region. Sicyopterus franouxi 10 AMERICAN MUSEUM NOVITATES NO. 3440

Fig. 6. Scanning electron micrographs of right premaxillary teeth in medial view. A, B, Sicyopterus franouxi, UMMZ 234880, 79.7 mm SL, 100ϫ and 250ϫ magni®cation. C, D, Sicyopterus punctissimus, paratype, UMMZ 234879, 73.4 mm SL, 100ϫ and 250ϫ magni®cation.

occurs in clear, swift-¯owing rivers and 1875: 6; Sauvage, 1891: 378, 520, 531, pl. 40A, streams. The species is frequently captured ®g. 2, 2a, pl. 47, ®g. 5. quite far inland and upstream of large falls Sicyopterus laticeps (misidenti®cation): Kiener, (e.g., Namorona River, Ranomafana National 1963: pl. 33. Park, in the southeastern highlands). HOLOTYPE: UMMZ 242048, 77.6 mm SL, ETYMOLOGY: Pellegrin dedicated the spe- adult male; Madagascar, Masoala Peninsula, cies to and named it after Mr. Franoux, a large unnamed cascading tributary of Anka- collaborator of Mr. R. Catala, who collected the holotype. vanana River (15Њ 18Ј S; 50Њ 16Ј E), JSS 94- 45; collected using small seines, J.S. Sparks, Sicyopterus punctissimus, new species K.J. Riseng, and local Malagasy guides, 29 Figures 5B, 6C, D, 7 Sept. 1994. PARATYPES: Total of 10 specimens, all Sicydium lagocephalum (nomen dubium): Pollen, from northeastern Madagascar: AMNH 1875: 6; Sauvage, 1891: 520; Catala, 1982: 60. 215498 (3, 76.1±121.9 mm SL), Andapa Re- Sicyopterus lagocephalus (nomen dubium): Kie- gion, Lokoho River, downstream of Belaoko, ner, 1963: 64, pl. 33; MaugeÂ, 1986: 383±384; Reinthal and Stiassny, 1991: 234; Stiassny and P. de Rham, 21 Oct. 1993; AMNH 226656 Raminosoa, 1994: 139; Stiassny and Harrison, (1, 106.7 mm SL), Diego-Suarez Region, 2000: 151±153. Manantenina River, southeastern boundary Sicydium laticeps (misidenti®cations): Pollen, of Parc National de Marojejy (14Њ 18Ј S, 49Њ 2004 SPARKS AND NELSON: MALAGASY SICYOPTERUS 11

Fig. 7. Sicyopterus punctissimus. A, holotype, UMMZ 242048, 77.6 mm SL, adult male. B, paratype, UMMZ 234879, 73.0 mm SL, adult female. Northeastern Madagascar: Masoala Peninsula, Ankavanana River.

33Ј E), M.L.J. Stiassny and ®sherwomen, 10 ®ns with light-colored (whitish to yellow) Nov. 1996; MNHN 1968-174 (2, 44.5±66.5 distal margins; second dorsal ®n with one mm SL), no locality data (incorrect coordi- weak spine and 11 branched rays; lateral nates reported in MNHN database), Kiener; cleft in the upper lip markedly expanded dor- UMMZ 234877 (1, 74.9 mm SL), Masoala sally; and scales on the nape and abdomen Peninsula, Ankavanana River (15Њ 18Ј 14Љ S, similar in size to scales on the sides of the 50Њ 13Ј 20Љ E), JSS 94-42, small seine, J.S. body. Sparks, K.J. Riseng, and local Malagasy DESCRIPTION:BODY AND FINS: Selected guides, 27 Sept. 1994; UMMZ 234879 (2, proportional measurements and meristic data 73±75.3 mm SL, 1 ex. CS), Masoala Penin- presented in table 2. Body robust, cylindrical sula, Ankavanana River, near Projet Masoala anteriorly, becoming somewhat laterally site 1005 (15Њ 18Ј 35Љ S, 50Њ 14Ј 08Љ E), JSS compressed posterior to origin of second dor- 94-40, small seine, J.S. Sparks, K.J. Riseng, sal ®n. Body depth greatest approximately at and local Malagasy guides, 25±26 Sept. level of ®rst dorsal ®n, but not varying great- 1994; UMMZ 234881 (1, 93.2 mm SL), data ly along entire body length. Head large, as for holotype. blunt, and rounded in lateral pro®le. First DIAGNOSIS: The new species differs from dorsal ®n with six weakly developed spines; all congeners in possessing the following second dorsal ®n with one weak spine and combination of characters: a broad, darkly 11 branched rays. Anal ®n with one weak pigmented, midlateral band on the posterior spine and 10 branched rays. First dorsal ®n half of the body extending onto the base of slightly more elongate than second; third the caudal ®n; numerous small dark (brown- spine longest. This spine markedly longer in ish) spots on the posterior half of the body, males than in females. In males, depressed dorsal and ventral to this midlateral band; nu- third spine extends well posterior of second merous small dark spots on the second dorsal dorsal-®n origin. Sixth spine of ®rst dorsal ®n; second dorsal, anal, caudal, and pectoral ®n very small; widely separated from pre- 12 AMERICAN MUSEUM NOVITATES NO. 3440 ceding spine. Second dorsal ®n and anal ®n OЈ present (®g. 4). Pore AЈ located just an- approximately same height; all rays of each terior and medial to anterior nasal tube; pore ®n branched near tips. Pectoral ®n large and B is just anterior and medial to posterior na- fan-shaped; length of longest ray approxi- sal opening. Based on material examined and mately equal to head length. All pectoral ®n descriptions and illustrations of other gobiid rays, except uppermost 4±6, branched; tips taxa in the literature (e.g., Akihito et al., of ®n rays slightly exserted from edge of ®n 1984; Pezold, 1993) this pore pattern appears membrane, but not elongated or ``silklike'' to be unique within the subfamily to the gen- (i.e., ®lamentous). Posterior margin of caudal era Sicyopterus and Sicydium. All examined ®n rounded; length of longest rays approxi- species of Sicyopterus exhibited these char- mately equal to head length. Pelvic ®ns fused acter states, although in one species of the into robust, cuplike suction disc. Total ver- latter genus (Sicydium crenilabrum) pore AЈ tebral count 25: 10 precaudal ϩ 15 caudal. is located posterior to the anterior nasal tube MOUTH AND TEETH: Mouth subterminal, (Harrison, 1993: ®g. 5). A survey of these large, and wide. Upper jaw long, extends features throughout the subfamily is not posteriorly to level of vertical through mid- complete. dle of orbit (®g. 5B). Upper lip broad, its Free neuromasts (ϭ pit organs) on the lower edge weakly crenulate. Symphyseal head and body are present as follows (®g. 4). cleft in upper lip simple although relatively There is an almost straight line of 3±5 organs deep; lateral cleft extremely narrow ventrally starting at the tip of the snout and terminat- (i.e., slitlike) and markedly expanded and ing anterior and medial to the anterior nasal rounded dorsally (®g. 5B). Large, ¯eshy pa- tube, followed by a brief interruption, then pilla just inside symphyseal cleft of upper continuing posteriorly in a line of 6±8 organs lip. Single row of ¯eshy, lightly pigmented to just medial to the posterior nasal opening. papillae present between upper lip and pre- A row of pit organs traverses the occipital maxilla; papillae at symphyseal and lateral region, beginning just posterior to the eye cleft larger than others in row. Lateral papilla and passing just posterior to cephalic pore D. expanded laterally (into area of cleft). Pre- Another transverse row just posterior to, and maxillae separated by median gap. Premax- essentially continuous with, this occipital illary teeth tricuspid (®g. 6C, D). Lateral row extends about one-third of the distance cusps relatively broad and median cusp slen- to the dorsal midline (®g. 4B). A series of der (®g. 6D). Tooth shaft not expanded dis- irregularly spaced pit organs, variable in tally, but tapered from base to articulation number, is located medially and just posterior with tooth crown. Premaxillary tooth count to the anterior transverse row. An anteropos- (approximately) 67 to 80, arrayed in single teriorly oriented row beginning at the upper, functional row. Numerous rows of replace- posterior margin of the orbit abuts the lower ment teeth (Mochizuki and Fukui, 1983) pre- portion of the posterior, transverse occipital sent on upper jaw. Dentary teeth composed row. Several short rows of organs radiate of outer row of setiform (labial) teeth form- across the suborbital region and cheek. A ing horizontal plate and an inner row of ca- short row of pit organs extends dorsoven- niniform, slightly curved teeth; anteriormost trally about midway between the anterior and one to two and posteriormost two of these posterior nasal openings, and another short, caniniform teeth largest. In largest specimens anteroposteriorly directed row abuts this ver- of type series (e.g. AMNH 215498: 121.9 tical row at about its midpoint. A curved row and 106.7 mm SL) labial teeth almost entire- of about 12±15 organs, beginning at the an- ly buried in ¯esh of lower lip and all cani- teriormost internarial free neuromast, abuts niform teeth of inner tooth row comparative- the lowermost portion of the ®rst (anterior- ly larger than in smaller individuals. Gill rak- most) suborbital row. Three rows of organs ers weakly developed. are present in the opercular region (®g. 4A): SEISMOSENSORY SYSTEM: Cephalic seismo- a nearly straight dorsoventral row of 11±15 sensory poresÐOculoscapular canal contin- organs beginning just posterior and dorsal to uous; pores AЈ, B, C, D (single), F, HK, and preopercular cephalic pore MЈ, at the anterior LЈ present; preopercular canal pores MЈ,N, portion of the opercle, and terminating near 2004 SPARKS AND NELSON: MALAGASY SICYOPTERUS 13 the ventral margin of the opercle; an approx- spotting present on caudal half of body and imately dorsoventral row of about 6±10 or- on second dorsal ®n. Spotting particularly gans near the posterior margin of the opercle; prevalent on caudal peduncle and near caudal and a generally shorter, anteroposteriorly di- base. Pectoral ®ns and caudal ®n yellow or rected row of 5±8 organs located slightly be- golden at margins of upper and lower lobes, low the lowermost portion of the posterior dark brown or blackish elsewhere. Pelvic opercular row. In the preopercular-mandibu- disk brown anteriorly, cream to pale yellow lar region there is a double row of 10±12 elsewhere. Anal ®n dark brown or gray, hy- pairs of pit organs between pore OЈ of the aline along distal margin. First dorsal ®n preopercle and the posterior corner of the brown to blackish proximal to body and ligh- mouth. A few other free neuromasts, scat- ter brown to yellowish along dorsal margin. tered about the head, are not arranged in any Second dorsal ®n dark brown to blackish discernible pattern. proximal to body. Margin of second dorsal SQUAMATION: Body covered with regularly ®n olive to yellow distally. imbricate, strongly ctenoid scales. Head and Females similar in coloration to males, but underside of body anterior to pelvic disk de- more muted overall. Females also lighter void of scales. Scales of nape and abdomen ventrally, and over more expansive area that of approximately same size as those of sides extends further dorsally onto ¯anks and of body. In axil of pectoral ®n and under free above anal-®n base. Margins of pectoral ®ns edges of pelvic disk, scales embedded in in- and caudal ®n whitish to pale yellow. tegument. Caudal ®n scaled over approxi- PIGMENTATION AND COLORATION IN PRES- mately one-®fth to one-fourth of length; ERVATION (FIG. 7): Upper part of head and scales become progressively reduced in size body dark brown. Lower part of head, begin- posteriorly on scaled portion of ®n. Approx- ning at level of upper lip, whitish to very imately 50±55 rows of scales in lateral series. light brown. Dark brown to black suborbital Approximately 13±18 scales present between bar extends ventrally or anteroventrally from posterior edge of pelvic disk and anus. Pre- orbit to upper lip. Brown pigmentation of dorsal scale count 12±14. head and anterior portion of body continues SEXUAL DIMORPHISM: In males, third spine onto posterior portion of body as wide mid- of ®rst dorsal ®n elongate, usually greater lateral stripe. In males, this midlateral stripe than twice the length of ®rst spine. Pigmen- begins under anterior rays of second dorsal tation and coloration of females differs ®n and terminates on anterior portion of cau- somewhat from males, mainly in being more dal ®n. In females, midlateral stripe begins muted, but not as strongly dimorphic as in on head, extends length of body, and termi- some sicydiine gobies (see color descriptions nates on anterior portion of caudal ®n, as in below). Size and shape of urogenital papilla males. Compared to males, midlateral stripe differs between sexes. This variation has of females not as deep on caudal peduncle. been discussed and/or ®gured for various si- Above and particularly below lateral stripe, cydiine genera by other authors (e.g., Parenti body more lightly pigmented due to lighter and Maciolek, 1993; Watson, 1995a, 1995b). pigmentation on edges of scales. This results In males, urogenital papilla bulbous and in distinct, small brown spots, which cover rounded distally, whereas in females papilla 75±90% of each scale on this part of body. bilobed and more tapered distally. Gap pre- These small brown spots extend slightly pos- sent in females between distal tip of genital terior to terminal scale rows and onto ante- papilla and anal-®n origin, whereas in males, rior part of caudal ®n, forming indistinct distal tip of genital papilla extends to anal- crescent-shaped pattern. Caudal ®n a uniform ®n origin. light brown, with narrow, whitish border at PIGMENTATION AND COLORATION IN LIFE: margins of upper and lower lobes. A few in- Males dark brown to black on head, dorsum, distinct darker spots present on upper part of and ¯anks, whitish to silvery on throat and caudal ®n. First dorsal ®n uniformly dark belly (lighter areas restricted ventrally in brown. Brown pigmentation of second dorsal males). Dark, brown to black, wide midla- ®n rays interrupted by areas of lighter color, teral stripe present. Dark brown to blackish resulting in an irregularly spotted pattern. 14 AMERICAN MUSEUM NOVITATES NO. 3440

Distal margin of second dorsal ®n more DISCUSSION faintly pigmented. Anal ®n brown, with MALAGASY AND MASCARENE SICYOPTERUS, darker brown submarginal streak and whitish COMPARISONS AND COMMENTS distal margin. Pectoral ®ns uniformly brown, with much lighter brown streak extending Preserved specimens of the new species along medial 2±3 rays, and with distinct are easily differentiated from S. franouxi, the whitish distal margin. Underside of head and only other sicydiine goby presently known to gular region light brown. Abdomen and pel- occur in the freshwaters of Madagascar, by a vic disk whitish. number of features. The number of branched Little sexual dimorphism evident with re- rays in the second dorsal ®n is invariably 11, spect to pigmentation or color pattern in pre- whereas in S. franouxi the number of served material, particularly in comparison branched rays is 10. In S. punctissimus, the with some other species of sicydiine gobies. scales of the nape and abdomen are approx- In females, however, anal ®n light brown in imately the same size as those on the ¯anks. color, making brown submarginal streak In S. franouxi the scales of the nape and ab- more prominent in females than in males. In domen (and behind the pectoral-®n axil) are addition, lower half of caudal ®n more darkly markedly reduced in size in comparison to pigmented than upper half, and midlateral the scales of the sides of the body. Compared stripe on body extends, as more faint stripe, to S. franouxi, S. punctissimus has a higher to distal portions of caudal rays. Midlateral number of premaxillary teeth (67±80 vs. 53± stripe more prominent in females due to ligh- 59), fewer scales in lateral series (50±55 vs. ter overall coloration, especially anteriorly. 65±71), fewer predorsal scales (12±14 vs. Flank region in females, between midlateral 26±31), and fewer scale rows between the stripe and anal ®n, more weakly pigmented pelvic disk and anus (13±18 vs. 27±37). Pre- and with fewer spots than in males. maxillary teeth in S. punctissimus are char- In largest specimens of type series acterized by a slender median cusp (®g. 6D; (AMNH 215498, 106.7±121.9 mm SL, and vs. broad in S. franouxi; ®g. 6B), and a tooth AMNH 226656, 108.7 mm SL), midlateral shaft that is tapered over its length (®g. 6C; stripe and spotting pattern on posterior re- vs. tooth shaft expanded both proximal to gion of body and second dorsal ®n much less base and distally, near articulation with pronounced, due to darker overall coloration crown; ®g. 6A). In S. punctissimus the upper of these individuals. In largest female spec- jaw extends posteriorly to about the level of imen, anal ®n more uniformly dark brown. mid orbit, whereas in S. franouxi, the upper Prominent whitish distal margins of second jaw extends at most to the level of the an- dorsal, anal, caudal, and pectoral ®ns remain terior margin of the orbit (®g. 5). The lateral readily apparent regardless of size. cleft in the upper lip is a narrow slit ventrally DISTRIBUTION AND HABITAT: The new spe- and is markedly expanded and rounded dor- cies is known from only a few isolated col- sally in S. punctissimus (®g. 5B); it is wide lection localities in northeastern Madagascar and about of equal width throughout in S. (®g. 1). All specimens collected by the ®rst franouxi (®g. 5A). author (catalogued as UMMZ lots) were cap- Comparison of morphometric measure- tured in clear, swift-¯owing streams and riv- ments taken on the two species (table 2) re- ers on the eastern slope of the Masoala Pen- veals few differences between them. Body insula, northeastern Madagascar. Sicyopterus depth data, at all three points at which mea- franouxi is sympatric with the new species, surements were taken, were slightly greater but unlike S. punctissimus, the range of S. in S. punctissimus than in S. franouxi. The franouxi extends much farther south (to the length of the base of the second dorsal ®n is southeastern highlands). greater in the former species, and is probably ETYMOLOGY: The speci®c name, punctis- attributable to the greater number of ®n rays simus, is a Latin adjective, superlative form, (11 vs. 10). The length of the anal-®n base and refers to the presence of numerous small in S. punctissimus also appears to be slightly spots on the sides of the body and second greater than that in S. franouxi. The basic dorsal ®n in the new species. color pattern (in preservation) of the two spe- 2004 SPARKS AND NELSON: MALAGASY SICYOPTERUS 15 cies is similar: brownish anteriorly, light-col- on the body wider than the ground color''. ored or whitish ventrally, with a broad, None of the specimens of Sicyopterus from brown, midlateral stripe along the side of the Madagascar (or the Mascarene region for that body. However, in S. franouxi the midlateral matter) that we have examined exhibit a sim- stripe is well de®ned at, and even anterior to, ilar color pattern, although certain specimens a vertical from the origin of the ®rst dorsal of S. franouxi do possess some faint, vertical (®gs. 2, 3), whereas in the new species the bars on the sides of the body. However, in S. overall ground coloration of the body is franouxi the wide, dark midlateral band darker much farther posteriorly and the mid- largely obscures these bars. Moreover, other lateral stripe is frequently not evident until character states observed in this Myanmar about the origin of the second dorsal ®n (®g. specimen (e.g., the thickened upper lip with 7). In S. franouxi there are also several faint numerous transverse plicae and the extreme- vertical bars on the ¯anks, which are slightly ly high number of premaxillary teeth) are darker than the ground color. These bars signi®cantly different from those observed in were not readily discernable in all specimens the Mascarene material. We therefore con- examined; however, similar bars are not pre- clude that attributions of S. fasciatus to Mad- sent in any specimen of the new species. The agascar are incorrect, and that specimens so body, including the caudal peduncle and cau- identi®ed in the past are simply adults of S. dal-®n base, of S. franouxi is devoid of dis- franouxi. tinctive brown spots (®gs. 2, 3), and the ®ns Sicyopterus laticeps, a species inhabiting lack the whitish margins, as well as the light the Mascarene islands of ReÂunion and Maur- brown streak extending along the medial 2± itius, but reported to occur in Madagascar ac- 3 rays of the pectoral ®n, characteristic of the cording to Valenciennes, in Cuvier and Va- new species (®g. 7). lenciennes (1837), and Pollen (1875), lacks In the course of this investigation, we have a dark, midlateral stripe, and the caudal ®n also been able to clarify the identity of spec- of this species has distinctive, black submar- imens from Madagascar that have been er- ginal bars on both the upper and lower lobes roneously attributed to Sicyopterus fasciatus. (®g. 8). We have not examined any material Pellegrin (1933) ®rst reported S. fasciatus assignable to S. laticeps from Madagascar, (Day, 1874) from Madagascar: four speci- and conclude that the species does not occur mens from ``la rivieÁre Sahembendrana (reÂ- on the island. gion de Tamatave)''. Examination of these There is potentially yet another Mascarene specimens indicates that they are adult spec- species of Sicyopterus, which inhabits the imens of Sicyopterus franouxi. Pellegrin freshwaters of the Comoros Islands. Balon (1935) described S. franouxi on the basis of and Bruton (1994), referring to this popula- a single juvenile specimen, measuring only tion as S. lagocephalus, provided an excel- 44.4 mm SL (MNHN 1935-0017). He ap- lent description of the ®sh, as well as a de- parently misidenti®ed the four adults he ex- tailed account of its habitat and biology. amined (MNHN 1891-0731 to 0734) as S. Specimens of this species (which we refer fasciatus, and considered them to be distinct herein to as Sicyopterus sp. ``Comoros'') are from S. franouxi. Similar to S. franouxi, S. similar to S. laticeps in the presence of dark fasciatus possesses scales of reduced size on (blackish) submarginal bars on the upper and the nape and abdomen, which may at least lower lobes of the caudal ®n, but they differ partly explain the numerous historical mis- from other Mascarene sicydiine gobies in the identi®cations of S. franouxi. overall coloration of the body. In the Com- The type material of S. fasciatus has been oros specimens, the body is uniformly dark lost, although we have been able to examine brown on the head, dorsum, and ¯anks, and a recently collected specimen from the Rak- is whitish on the underside. A few vague ver- hine region of Myanmar (ϭ Burma). This tical, darker brown bars on the ¯anks are dis- specimen compares very well with Day's cernible in some specimens. (1874, 1876) description and specimen he Finally, Gobius caeruleus (Commerson, in ®guredÐparticularly the color pattern com- LacepeÁde, 1800), ``le Gobie Bleu'', also a si- posed of prominent ``vertical darker bands cydiine goby described from material col- 16 AMERICAN MUSEUM NOVITATES NO. 3440

Fig. 8. Sicyopterus laticeps. A, MNHN 918, syntype, adult male, 97.6 mm SL, ReÂunion. B, MNHN 1984-806, adult male, 96.7 mm SL, ReÂunion.

lected in ReÂunion, is determined to be an descripsit, ignoravit autem patriam.'' We available name. Based on Commerson's de- translate this as: ``I myself regard the small scription, however, we are unable to deter- ®sh to be from America. Koelreuter de- mine if the species is conspeci®c with S. la- scribed [the species] from a specimen in the ticeps. No type material exists for this spe- St. Petersburg Museum, however he was ig- cies, and we defer resolution of the nomen- norant of [its] native land.'' clatural problem of the ReÂunion Sicyopterus The description and ®gures of the speci- to a later study. men presented by Pallas (1770) leave little doubt that the ®sh was a sicydiine goby. TAXONOMIC STATUS OF GOBIUS However, identi®cation of the specimen to LAGOCEPHALUS PALLAS, 1770 the speci®c (or even generic) level is not pos- Gobius lagocephalus was described by sible from the description or ®gures. Char- Pallas on the basis of a single specimen ex- acter states (e.g., number of rays in the sec- amined by him in the St. Petersburg Muse- ond dorsal ®n [10] and pectoral ®n [15]) re- um. The specimen had earlier been described ported by Pallas, are inconclusive in estab- as a Gobius by Joseph Koelreuter (1764). lishing taxonomic identity. The specimen Neither Pallas nor Koelreuter knew where upon which Pallas based his description has the specimen had been collected. Pallas been lost. wrote: ``Pisculum ipse ex America habui. Valenciennes, in Cuvier and Valenciennes Koelreuterus e specimen Musei Petropolitani (1837), placing the species into the newly de- 2004 SPARKS AND NELSON: MALAGASY SICYOPTERUS 17 scribed genus Sicydium, was the ®rst to at- study, the specimens recently designated as tribute Gobius lagocephalus to the Mascar- neotypes of S. lagocephalus are valid as S. ene region: ``. . . des iles de France et de laticeps. The neotype of S. lagocephalus des- Bourbon ...''.Valenciennes, in this same ignated by Fricke (1999: 523) is a syntype paper, described Sicydium laticeps from ``les of S. laticeps (MNHN 841). eaux douces de l'IÃle de Bourbon ...''(ϭ The numerous synonyms for New Cale- freshwaters of the island of Bourbon). This donian sicydiine gobies listed by Watson et latter name is the correct one for the species al. (2000; n ϭ 12) under Sicyopterus lago- of Sicyopterus inhabiting the island of ReÂ- cephalus are also problematic. As we have union (ϭ l'IÃle de Bourbon). The syntypes of stated above, translation of Pallas (1770) re- S. laticeps (which we have examined) are in veals that neither he nor Koelreuter knew the good condition (®g. 8A), and pigmentation collecting site of the described specimen. pattern and coloration are well preserved. Pe- Pallas regarded it ``. . . to be from America'' ters (1868) ascribed S. lagocephalus to the (see above). Attempting to establish this no- Comoros Islands, and it appears that Pollen men dubium as senior synonym to numerous (1875) was the ®rst to attribute this species other nominal sicydiine taxa only further (along with S. laticeps) to Madagascar, al- confuses nomenclatural issues in this specio- though he did not provide speci®c localities se and wide-ranging group of gobies. Watson (e.g., ``Le PeÃches aà Madagascar et ses DeÂ- et al. (2000) neither compared their diagnosis pendances''). Likewise, several more recent for Sicyopterus lagocephalus with the de- ichthyofaunal inventory studies have also re- scription presented by Pallas nor did they ported S. lagocephalus from the freshwaters justify selection of a neotype locality remote of Madagascar (e.g., Kiener, 1963; Catala, from the conjectured ``native land'' stated in 1982; MaugeÂ, 1986; Reinthal and Stiassny, the original description of the species. As we 1991; Stiassny and Raminosoa, 1994; Stiass- have demonstrated, although the description ny and Harrison, 2000). and ®gures of Gobius lagocephalus present- We consider Gobius lagocephalus to be a ed by Pallas (1770) leave little doubt that the nomen dubium of uncertain placement (in- ®sh described was a sicydiine goby, accurate certae sedis) beyond the subfamily level. identi®cation of the specimen to even the ge- Therefore, the name is not available for any neric level is certainly not possible. It is en- of the presently recognized Mascarene sicy- diine gobies (likewise, it is not available for tirely feasible that the specimen described by any species of Sicyopterus). However, due to Pallas (1770) is a member of Sicydium, a ge- historical misapplication of the name to sev- nus that occurs in the tropical eastern Paci®c eral species of Mascarene sicydiine gobies, and Atlantic Oceans, including the Caribbean we have included the name in the synonymy Sea (i.e., American Seas). Hence, although of both known Malagasy species. the research presented by these authors pro- Recently, both Fricke (1999: 523) and G. vides some interesting descriptions of habitat Marquet in Watson et al. (2000: 13) have and ecology, as well as many excellent color designated neotypes of Sicyopterus lagoce- illustrations of New Caledonian Sicyopterus phalus from specimens collected in the Mas- species, the taxonomic conclusions reached carene region (ReÂunion). Gobius lagoce- in the article are not supported by the avail- phalus is not an available name for the rea- able evidence. Based on our studies of Mal- sons detailed above. In addition, Eschmeyer agasy and Mascarene sicydiine gobies, we (1998; online version updated March 13, are of the opinion that, with further study of 2003) points out that neither of these desig- Sicyopterus interrelationships, including pro- nations is valid because they fail to meet posed molecular analyses, claims of a high conditions set out in Article 75.3 of the In- degree of island-group endemism for these ternational Code of Zoological Nomenclature ®shes (Parenti and Maciolek, 1993) will be (1999). Fricke later rescinded his neotype corroborated and additional geographically designation for S. lagocephalus, along with distinct populations that have historically numerous others (see Fricke, 2000: 639). In been placed in S. lagocephalus will be re- accordance with the conclusions of this vealed. 18 AMERICAN MUSEUM NOVITATES NO. 3440

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APPENDIX 1

MATERIALS EXAMINED type, 97.6 mm SL), ReÂunion; MNHN 1984± 806 (2, 51.0±96.7 mm SL), ReÂunion. Representative sicydiine taxa examined in the Sicyopterus lividus: USNM 322470 (4, paratypes, present study are listed below. Type specimens 39.3±42.7 mm SL), Caroline Islands, Ponape; are listed ®rst. Institutional catalog number, num- USNM (1, 62.6 mm SL), Caroline Islands, ber of specimens examined, and size range follow Pohnpei. the species name. CS denotes material cleared and Sicyopterus longi®lis: USNM 346647 (1, 64.6 mm stained for bone and/or cartilage. SL), Philippine Islands, Luzon Island, Quezon Province, Gen. Nakar, Agos River, Brgy. Mai- Sicydium multipunctatum: UMMZ 189598 (1 CS, gang; USNM 346653 (1, 71.2 mm SL), Phil- 90 mm SL), Mexico, Colima, Rio de Comala, ippines, Quezon, Gen. Nakar, Brgy. Banglos, tributary of Rio Armeria drainage, south of Agos River. Comala and ca. 5 mi north of Colima. Sicyopterus marquesensis: USNM 109370 (2, Sicydium plumieri: USNM 313724 (42, 29.6± paratypes, 42.1±48.1 mm SL), Marquesas Is- 102.2 mm SL), Caribbean, Dominica B.W.I., lands, Tohetaivau, Oomoa Valley, Fatuhiva. foot of Tra-Falgar Falls and portions of Trois Sicyopterus micrurus: UMMZ 189873 (10, 25.0± Pitons River behind hydroelectric station. 26.0 mm SL), Society Islands, Tahiti-iti, Vavi Sicyopterus brevis: USNM 313863 (5, 24.3±27.1 Valley; USNM 348156 (2, 29.1±35.5 mm SL), mm SL), American Samoa, Soonapule Stream American Samoa, Tau Island, National Park of (Seetaga) near mouth. American Samoa, Laufuti Stream. Sicyopterus cynocephalus: USNM 340134 (1, Sicyopterus ouwensi: USNM 313867 (3, 24.0± 63.5 mm SL), Philippine Islands, Palawan, 26.0 mm SL), American Samoa, Soonapule Plaridel River. Stream (Seetala) near mouth. Sicyopterus eudentatus: USNM 322473 (1, para- Sicyopterus pugnans: USNM 348154 (3, 32.3± type, 100.0 mm SL), Caroline Islands, Ponape, 42.1 mm SL), American Samoa, Tau Island, Nanpil River, transect above reservoir. National Park of American Samoa, Laufuti Sicyopterus extraneus: USNM 135671 (2, 103.3± Stream. 122.5 mm SL), Taiwan, Koroton; USNM Sicyopterus rapa: USNM 330077 (1, paratype, 135740 (2, 21.5±62.9 mm SL), Philippine Is- 64.8 mm SL), French Polynesia, Rapa, shallow lands, Camiguin Id (between Leyte and Min- water in Pania River at head of Haurei Bay. danao). : UMMZ 196862 (2, 71.0± Sicyopterus fasciatus: NHRM (NRM) 40847 (1, 72.6 mm SL, 2 CS), Hawaiian Islands, Kauai, 87.4 mm SL), Myanmar, Rakhine State, Taung- Wainiha River, 2 km above mouth; USNM gok Thade River Drainage: Yan Khaw Chaung, 214001 (5, 25.3±82.0 mm SL), Hawaiian Is- ca. 4 km from Gwetauk village (18Њ 47Ј 48ЉN, lands, Maui Island, east Maui, Hanawi Stream; 94Њ 21Ј 46ЉE). USNM 214005 (1, 113.2 mm SL), Hawaiian Sicyopterus fuliag: USNM 135738 (1, 101.0 mm Islands, Maui Island, west Maui, Waihee SL), Philippines, Mindanao, Nonucan River, Stream. Iligan Branch, Camp Overton. Sicyopterus taeniurus: USNM 66030 (2, 84.5± Sicyopterus inana: USNM 109379 (1, paratype, 91.3 mm SL), Society Islands, Tahiti, Tatana 26.5 mm SL), Society Islands, Tahiti, Papenoo River; USNM 82967 (1, 75.1 mm SL), Society Valley; USNM 109371 (1, ϳ25 mm SL), Mar- Islands, Tahiti. quesas Islands, Oomoa Valley, Fatuhiva. Sicyopterus tauae: USNM 51787 (1, holotype, Sicyopterus japonicus: UMMZ 194566 (2, 75.9± 27.5 mm SL), Western Samoa, Ahia, Upolu Is- 80.6 mm SL, 2 CS); UMMZ 194573 (14, 60.0± land, Vaisigano River. 68.0 mm SL), Taiwan, I-Ian, Tau Chang, Lan Sicyopterus sp. ``Comoros'': MNHN 1931-257 (2, Yan Chi (Nomonhan), Ta-tung Hsiang; USNM 23.5±25.4 mm SL), Comoros, Anjouan Island; 191283 (1, 104.0 mm SL), Taiwan, Chu-Tung, RUSI 27713 (10, 66.5±91.5 mm SL), Comoros Hsin-Chu Hsien, small stream in coastal plain; Islands, Anjouan Island, Tatinga River; RUSI USNM 336718 (1, 58.6 mm SL), Taiwan, Tai- 036381 (12, 23.7±110.6 mm SL), Comoros Is- dong, Xin-Gong XI (ϭ New Port Stream). lands, Anjouan Island, Tatinga River; UMMZ Sicyopterus lachrymosus: USNM 135739 (12, 237080 (6, 79.0±94.0 mm SL, 1 CS), Comoros 46.3±67.6 mm SL), Philippine islands, Luzon, Islands, Anjouan Island, Tatinga River; UMMZ Dumaca River (branch of Tayabes); USNM 237081 (4, 33.0±52.0 mm SL), Comoros Is- 313864 (1, 50.0 mm SL), Philippine Islands, lands, Anjouan Island, Tatinga River. northern Luzon, Ilocos Province, Tagudin. Sicyopterus sp. ``Pohnpei'': USNM 362310 (1, Sicyopterus laticeps: MNHN 841 (1, syntype, 95.0 mm SL), Pohnpei, Lehn Mesi River, 3.2 91.1 mm SL), ReÂunion; MNHN 918 (1, syn- km NE of Salapwuk.