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Fish communities on tropical deep reefs are dominated by species that belong to families primarily composed of shallow-water species. Collections of deep-reef fishes via submersibles have allowed us to include these deep-reef species in molecular phylogenies, providing insights into the timing and frequency of invasions from shallow to deep reefs. Here we provide evidence of a new deep-reef invasion in the tribe Gobiosomatini in the family Gobiidae (gobies). We describe two new species, one of which belongs to a new genus, and incorporate these taxa into a time-calibrated molecular phylogeny of Gobiosomatini to show that, collectively, these two genera represent a previously unreported independent invasion on to deep reefs that occurred approximately 20–30 million years ago. These new taxa are readily distinguished from related genera and species by a combination of live coloration, pelvic-fin morphology, meristic characters, head-pore patterns and other osteological characters. We discuss the relevance of these two new species to the systematics of the tribe Gobiosomatini and include a comparison to all known genera in the tribe.

BIRDSONGICHTHYS GEN. NOV. Type species: Birdsongichthys rectus sp. nov. Zoobank registration: urn: lsid: zoobank. org:act: 287F2C89-6296-4FCB-B7B3-ED7B4B4EE73C Diagnosis: Possesses all taxonomic characters present in most members of the Gobiosomatini and the Gobiosoma group, including the following: first dorsal-fin spines VII; dorsal pterygiophore insertion pattern 3-221110; vertebrae 27–11 precaudal and 16 caudal; hypurals 1 and 2 fused to some extent with hypurals 3 and 4 and the terminal vertebral element; one epural; ventral postcleithrum absent. Pelvic fins separate, anterior frenum connecting spines absent, membrane connecting innermost rays absent or weakly developed; pelvic-fin rays 1–5 unbranched and without fleshy or flattened tips; pelvic-fin rays long, extending posteriorly to a point between middle or end of anal fin; side of body covered with ctenoid scales, including two modified basicaudal scales; two anal-fin pterygiophores inserted before first haemal spine; cephalic sensory canals present; second dorsal-fin rays I,9; anal-fin rays I,8; body with three horizontal series of bright yellow spots, one along dorsal midline, one along lateral midline and one along ventral midline; anterior profile of head with gradual slope, not blunt or nearly vertical; eye large (~8.3–10.6% SL). Etymology: The name honours Dr Ray S. Birdsong, who contributed substantially to the knowledge on the systematics of the family Gobiidae, especially taxa from the tropical western Atlantic and eastern Pacific Oceans. The root ‘ichthys’ is derived from the Greek ‘ ikhthús ’ (ιχθύς), fish. Comparisons (Table 1): The presence of VII dorsal spines, 11 + 16 vertebrae and a 3-221110 dorsal pterygiophore pattern (Fig. 4A) all support placement of Birdsongichthys in Gobiosomatini. Gobiosomatini include two main clades, the Microgobius group and the Gobiosoma group. Birdsongichthys shares several characters with genera of the Microgobius group (Antilligobius, Akko Birdsong & Robins, 1995, Bollmannia Jordan, 1890, Microgobius Poey, 1876, Palatogobius and Parrella Ginsburg, 1938), including a laterally compressed body and no substantial fusion between hypurals 3–4 and hypurals 1–2 and the terminal vertebral element. It differs from all Microgobius group genera in having modified basicaudal scales (absent in the Microgobius group) and in having a truncate or emarginate caudal fin (vs. lanceolate in the Microgobius group). The Gobiosoma group contains four lineages: the Barbulifer subgroup, the Nes subgroup, Aboma Jordan & Starks, 1895 and Robinsichthys. Birdsongichthys can be distinguished from all members of the Barbulifer subgroup (Aruma Ginsburg, 1933, Barbulifer, Elacatinus Jordan, 1904, Evermannichthys Metzelaar, 1919, Ginsburgellus Böhlke & Robins, 1968, Gobiosoma, Ophiogobius Hoese, 1976, Pariah Böhlke, 1969, Risor Ginsburg, 1933 and Tigrigobius Fowler, 1931) and Aboma by its completely separate pelvic fins that lack an anterior frenum and have unbranched rays; the Barbulifer subgroup and Aboma have pelvic fins united to form a complete disc with branched rays. Completely separate pelvic fins also characterize species in the Nes subgroup, except Nes and Gobulus. Birdsongichthys differs from most genera of the Nes subgroup and from Robinsichthys in having cephalic lateralis canals and associated head pores. Among Nes subgroup genera, only Eleotrica Ginsburg, 1933, Gymneleotris Bleeker, 1874, and two species of Chriolepis Gilbert, 1892 possess head pores. Birdsongichthys has unbranched pelvic-fin rays, whereas these three genera all have branched pelvicfin rays. The only Gobiosomatini genera containing species with completely unbranched pelvic-fin rays are the monotypic Paedovaricus Van Tassell et al., 2016 [P. imswe (Greenfield, 1981)] and Varicus (unbranched in specimens of five of ten known species in the genus). Birdsongichthys differs from these two genera in having two (vs. one) anal-fin pterygiophores inserted anterior to the first haemal spine. Finally, Birdsongichthys can be further distinguished from Robinsichthys in having 11 + 16 vertebrae (vs. 11 + 17).

BIRDSONGICHTHYS RECTUS SP. NOV. (FIGS 4A, 5–7) Straightfinned goby (English), gobio de aleta recta (Spanish). Gobiidae genus 2, species 1 Robertson et al., 2020: table 1, supplemental plate S13. Zoobank registration: urn: lsid: zoobank. org:act: 016EDDA3-572B-45CB-8B05-676076400186 Holotype: USNM 442874, 14.1 mm SL female, tissue EUS17221, field number CURASUB 17-21, 151 m depth, rubble patch, south and south-east of R / V Chapman mooring, SW of island, Kay Bay, Sint Eustatius, 17.4566, –62.9791, quinaldine, HOV Curasub and R / V Chapman, Carole Baldwin, Luke Tornabene, Bruce Brandt, 19 April 2017. Table 1 . Continued * genus is not monophyletic. ** See Birdsong et al. (1988) for wide range of vertebral counts and pterygiophore insertion patterns. Abbreviations : D 1, first dorsal fin ; D 2, second dorsal fin; A, anal fin; AP , anal fin pterygiophores. Paratypes: MUVS-V-439, 21.0 mm SL male, tissue ROA18021, field number IDABEL18-01, 213 m depth, sand and limestone rubble, outside Halfmoon Bay, Roatan, Honduras, 16.304384°, –86.598204°, quinaldine sulphate, HOV Idabel, D. Ross Robertson, Rachel Manning, Karl Stanley, 4 June 2018; UW 158074, 18.3 mm female, tissue ROA17004, cleared and stained, field number IDABEL17-01, 192 m depth, outside Halfmoon Bay, Roatan, Honduras, 16.305557°, –86.597669°, quinaldine sulphate, HOV Idabel, Luke Tornabene, Ross Robertson, Karl Stanley, 24 July 2017; UW 158128, 12.0 mm female, tissue ROA18044, field number IDABEL18-03, 241 m depth, sand rubble in rock hole, 16.3043°, –86.5982°, quinaldine sulphate, HOV Idabel, Luke Tornabene, Rachel Manning, Karl Stanley, 6 June 2018; UW 158200, 11.5 mm male, tissue ROA18212, field number IDABEL18-08, 213 m depth, sand at base of limestone, 16.3043°, –86.5982°, quinaldine sulphate, HOV Idabel, Luke Tornabene, Katherine Maslenikov, Karl Stanley, 1 June 2018; USNM 442726, 14.4 mm male, tissue EUS17073, field number CURASUB 17-18, 110 m depth, rubble and sand on wall, dropoff west of R / V Chapman mooring, Gallows Bay, Sint Eustatius, 17.4608°, –63.0179°, quinaldine, HOV Curasub and R / V Chapman, Carole Baldwin, Luke Tornabene, Barbara van Bebber, Jordan Casey, 16 April 2017. Diagnosis: Birdsongichthys rectus is the only known member of the genus, so the species diagnosis is that of the diagnosis of the genus. Description: Morphometric data are presented in Table 2. Median and paired fins: first dorsal fin VII(6), first spine extremely elongate in some specimens, extending posteriorly to point between middle of soft dorsal fin and base of anal fin when flat against body; second dorsal fin I,8*(1) or I,9(5); anal fin I,8*(6); pectoral-fin rays 16*(3), 17 (2), 18(1), extending posteriorly to a vertical through second dorsal-fin ray 2 or 3; pelvic fin I,5* (5), all rays unbranched; pelvic fins well separated, lacking anterior frenum, membrane connecting fifth rays absent or reduced; fourth and fifth pelvic-fin rays approximately equal in length, extending posteriorly to base of anal-fin rays 6–7; tips of pelvic-fin rays not ending in fleshy pads; caudal fin truncate or emarginate; branched caudal-fin rays 12 (4)*, segmented caudal-fin rays 17(5)*, caudal fin broken in many specimens. Scales: Scales on head and nape absent; ctenoid scales on side of body present starting above pectoral fin and continuing to caudal peduncle; lateral scale rows 22–23, transverse scale rows 5, scales counted from three specimens including holotype, scales lost in other specimens; pair of modified ctenoid scales with enlarged cteni present at base of caudal fin, one each at dorsal and ventral margins of fin (Fig. 5); abdomen and chest each with small patch of ctenoid scales. Head: Jaw extending posteriorly to a vertical through anterior margin of eye or midpoint of pupil; anterior naris a short tube, posterior naris an opening flush with snout; cephalic lateralis pores small, pores B′, D(s), E′ present in four specimens, pores B′ extremely small and pore D missing in holotype, pores B′ not visible in paratype UW 158200; eyes large, 30–36% HL, positioned dorsolaterally. Upper jaw with two to three rows of teeth anteriorly, outermost row enlarged widely spaced canines continuing along entire upper jaw to the end of the premaxilla, innermost two rows tightly packed, shorter, and conical, becoming single row posteriorly; lower jaw with three to five rows of teeth anteriorly, outermost row restricted to tip of dentary and slightly larger than innermost rows, innermost rows becoming two rows posteriorly. Sensory papillae not visible on most specimens due to small size of specimens and damage to the face from collection. Genitalia: Male urogenital papilla elongate and pointed; female urogenital papilla short, bulbous or slightly conical. Colour in fresh specimens (Fig. 6): Side of body translucent with iridescent scale margins; three horizontal rows of small, bright yellow spots along flank, spots largest in the bottom row; first row with 12–15 spots starting on nape and continuing along dorsal midline, ending just anterior to caudal-fin base; second row with 13–15 spots starting above pectoral fin and continuing along lateral midline, ending just anterior to caudal-fin base; third row with eight to nine spots starting just anterior to anal-fin origin and continuing along ventral midline, ending just anterior to caudal-fin base; tips of snout and upper and lower jaws with small patch of yellow; eye with upperhalf of iris black, lower-half yellow; a broad, oblique yellow to orange stripe extending from lower-rear part of iris down on to preopercle; remainder of head translucent; pink coloration of red gill arches visible through operculum; pectoral-fin base with yellow spot on yellow upper margin of some specimens; dorsal fins with uniform translucent yellow wash with scattered iridophores, and with a distinct horizontal series of yellow spots along midpoints of spines and rays in some specimens; pectoral fins pale or uniformly translucent yellow; caudal fin faintly translucent yellow with two dark yellow, vertical bars at base; anal and pelvic fins uniformly translucent yellow. Colour in preservation (Fig. 7): Body uniformly pale yellowish, no dark pigmentation remains in any preserved specimens, including those fixed either in 95% ethanol or in formalin followed by storage in 70–75% ethanol. Habitat: Known from 110-241 m depth on sand and rubble substrates situated on or around the base of walls or steep rocky slopes on deep reefs. Distribution: Known from Roatan, Honduras and Sint Eustatius, Caribbean Netherlands. Etymology: The epithet is from the Latin adjective rectus, straight, which refers to the straight, unbranched rays of the pelvic fin.

BIRDSONGICHTHYS GEN. NOV. Type species: Birdsongichthys rectus sp. nov. Zoobank registration: urn: lsid: zoobank. org:act: 287F2C89-6296-4FCB-B7B3-ED7B4B4EE73C Diagnosis: Possesses all taxonomic characters present in most members of the Gobiosomatini and the Gobiosoma group, including the following: first dorsal-fin spines VII; dorsal pterygiophore insertion pattern 3-221110; vertebrae 27–11 precaudal and 16 caudal; hypurals 1 and 2 fused to some extent with hypurals 3 and 4 and the terminal vertebral element; one epural; ventral postcleithrum absent. Pelvic fins separate, anterior frenum connecting spines absent, membrane connecting innermost rays absent or weakly developed; pelvic-fin rays 1–5 unbranched and without fleshy or flattened tips; pelvic-fin rays long, extending posteriorly to a point between middle or end of anal fin; side of body covered with ctenoid scales, including two modified basicaudal scales; two anal-fin pterygiophores inserted before first haemal spine; cephalic sensory canals present; second dorsal-fin rays I,9; anal-fin rays I,8; body with three horizontal series of bright yellow spots, one along dorsal midline, one along lateral midline and one along ventral midline; anterior profile of head with gradual slope, not blunt or nearly vertical; eye large (~8.3–10.6% SL). Etymology: The name honours Dr Ray S. Birdsong, who contributed substantially to the knowledge on the systematics of the family Gobiidae, especially taxa from the tropical western Atlantic and eastern Pacific Oceans. The root ‘ichthys’ is derived from the Greek ‘ ikhthús ’ (ιχθύς), fish. Comparisons (Table 1): The presence of VII dorsal spines, 11 + 16 vertebrae and a 3-221110 dorsal pterygiophore pattern (Fig. 4A) all support placement of Birdsongichthys in Gobiosomatini. Gobiosomatini include two main clades, the Microgobius group and the Gobiosoma group. Birdsongichthys shares several characters with genera of the Microgobius group (Antilligobius, Akko Birdsong & Robins, 1995, Bollmannia Jordan, 1890, Microgobius Poey, 1876, Palatogobius and Parrella Ginsburg, 1938), including a laterally compressed body and no substantial fusion between hypurals 3–4 and hypurals 1–2 and the terminal vertebral element. It differs from all Microgobius group genera in having modified basicaudal scales (absent in the Microgobius group) and in having a truncate or emarginate caudal fin (vs. lanceolate in the Microgobius group). The Gobiosoma group contains four lineages: the Barbulifer subgroup, the Nes subgroup, Aboma Jordan & Starks, 1895 and Robinsichthys. Birdsongichthys can be distinguished from all members of the Barbulifer subgroup (Aruma Ginsburg, 1933, Barbulifer, Elacatinus Jordan, 1904, Evermannichthys Metzelaar, 1919, Ginsburgellus Böhlke & Robins, 1968, Gobiosoma, Ophiogobius Hoese, 1976, Pariah Böhlke, 1969, Risor Ginsburg, 1933 and Tigrigobius Fowler, 1931) and Aboma by its completely separate pelvic fins that lack an anterior frenum and have unbranched rays; the Barbulifer subgroup and Aboma have pelvic fins united to form a complete disc with branched rays. Completely separate pelvic fins also characterize species in the Nes subgroup, except Nes and Gobulus. Birdsongichthys differs from most genera of the Nes subgroup and from Robinsichthys in having cephalic lateralis canals and associated head pores. Among Nes subgroup genera, only Eleotrica Ginsburg, 1933, Gymneleotris Bleeker, 1874, and two species of Chriolepis Gilbert, 1892 possess head pores. Birdsongichthys has unbranched pelvic-fin rays, whereas these three genera all have branched pelvicfin rays. The only Gobiosomatini genera containing species with completely unbranched pelvic-fin rays are the monotypic Paedovaricus Van Tassell et al., 2016 [P. imswe (Greenfield, 1981)] and Varicus (unbranched in specimens of five of ten known species in the genus). Birdsongichthys differs from these two genera in having two (vs. one) anal-fin pterygiophores inserted anterior to the first haemal spine. Finally, Birdsongichthys can be further distinguished from Robinsichthys in having 11 + 16 vertebrae (vs. 11 + 17)., Published as part of Tornabene, Luke, Manning, Rachel, Robertson, D. Ross, Van Tassell, James L. & Baldwin, Carole C., 2023, A new lineage of deep-reef gobies from the Caribbean, including two new species and one new genus (Teleostei: Gobiidae: Gobiosomatini), pp. 322-343 in Zoological Journal of the Linnean Society 197 (2) on pages 327-328, DOI: 10.1093/zoolinnean/zlac013, http://zenodo.org/record/7622658, {"references":["Bohlke JE, Robins CR. 1968. Western Atlantic seven-spined gobies, with descriptions of ten new species and a new genus, and comments on Pacific relatives. Proceedings of the Academy of Natural Sciences of Philadelphia 120: 45 - 174."]}

ROBINSICHTHYS NIGRIMARGINATUS SP. NOV. (FIGS 4B, 8–11) Black-margined goby (English), gobio de margen negro (Spanish). Zoobank registration: urn: lsid: zoobank. org:act: 9D9D60B1-2777-47FC-8CB0-6A7E0F70641C Holotype: USNM 446953, 21.0 mm female, tissue CUR19007, field number CURASUB19-02, 229 m depth, sandy bottom, East of Substation downline, Substation Curaçao, Bapor Kibra, Curaçao, 12.0838°, -68.8991°, quinaldine, HOV Curasub, Carole Baldwin, Luke Tornabene, Tico Christiaan, Sarah Yerrace, 7 May 2019. Diagnosis: First dorsal-fin pterygiophore pattern 3-22111*0 (last pterygiophore lacking an associated dorsal-fin spine; Fig. 4B); VI spines in first dorsal fin; 11 + 17 vertebrae; two modified basicaudal scales; head pores absent; second dorsal fin I,9; anal fin I,9; pelvic fins completely united by membrane to form a round disc with well-developed anterior frenum; pectoral-fin rays 19. Description: Morphometric data are presented in Table 2. Median and paired fins: First dorsal fin VI, no spines elongate; second dorsal fin I,9; anal fin I,9; pectoral-fin rays 19, extending posteriorly to a vertical through second dorsal-fin ray 2 or 3; pelvic fin I,5, all rays branched; pelvic fins connected via membrane forming a complete disk with well-developed anterior frenum connecting pelvic spines (Fig. 8); pelvic fin extending posteriorly to just before anus; tips of pelvicfin rays not ending in fleshy pads; caudal fin truncate; branched caudal-fin rays 14, segmented caudal-fin rays 17. Scales: Scales on head and nape absent; ctenoid scales on side of the body present starting above pectoral fin base and ending at base of caudal fin rays; lateral scale rows 24, transverse scale rows six; pair of modified ctenoid scales with enlarged cteni present at base of caudal fin, one each at dorsal and ventral margins of fin; ctenoid scales present on abdomen, scales slightly larger than those on side of body. Head: Jaw extending posteriorly to a vertical through midpoint of the pupil; anterior naris a short tube, posterior naris an opening with raised rim; cephalic lateralis pores absent; eyes large, 41% HL, dorsolateral. Both jaws with two rows of teeth, teeth thin, straight, pointed and evenly spaces posteriorly, becoming less regularly spaced, slightly more recurved anteriorly, with an occasional tooth in outer row slightly larger than others in row. Sensory papillae arranged as in Figure 9, with four prominent transverse rows, including rows 5i/5s present as a continuous single row. Colour in life (Fig. 10): Body translucent with a silvery sheen, an internal white area around gut; body with five dark bars formed by aggregations of black chromatophores overlaying translucent yellow areas, the first bar strongest and widest, extending from the centre of the base of the first dorsal fin to the lower belly, the second to fifth only on the lower-half of the body, thin, formed by less dense aggregations of tiny black chromatophores, the last bar partly fused to the bar on the base of the caudal fin; upper-body profile with six small, translucent yellow blotches, each with a central area of tiny, black chromatophores, the first blotch between the two dorsal fins, the second under the front of the second dorsal fin, the third under the centre of that fin, the fourth to sixth on the caudal peduncle; two elongate internal areas of dark pigment with yellow tones along the spinal column, a short one under the second body bar, a longer one from third body bar to just behind the fourth body bar, with a suggestion of a third patch above the last body bar; when the fish is laterally oriented (Fig. 10A), the silver sheen of the body is present in irregularly dispersed patches in the translucent areas, when the fish is oriented partly towards the viewer (Fig. 10B) that iridescent colour is greenish silver, distributed in patches between the body bars and between the yellow dorsal-profile blotches, and as a distinct vertical greenish silver bar just before the upper-half of the first dark body bar. Head translucent with silver sheen on cheeks; top rear corner of opercle with a translucent orange patch about half pupil size and a larger patch of tiny, welldispersed, black chromatophores over area of pink gills, an indistinct, irregularly shaped translucent orange patch behind centre of eye extending on to nape; mouth translucent, top jaw with silver sheen and a transverse line of black chromatophores across its centre, an indistinct yellow blotch on the snout immediately behind its centre; iris with most of its anterior and rear part overlaid by a dense silvery yellow sheen, top edge of the eyeball silvery yellow; a curved black bar formed by densely aggregated black chromatophores, passing from upper-rear of iris through the pupil and down to rear corner of the mouth, section of bar under eye with relatively large chromatophores overlaying a translucent yellow area, that bar bounded anteriorly and posteriorly by patches of silver iridescence. Base of caudal fin with long, thin, vertical yellow bar with an aggregation of black chromatophores down its centre that partly fuses with last body bar, fin itself translucent with patches of tiny black chromatophores along interradial membranes – two small patches along top of fin, a subterminal band down upper-half of posterior of the fin; a small, vertical patch of black-dotted yellow pigment just behind upper-half of the basal dark bar; first dorsal fin translucent with a large, horizontally oval, solid black blotch covering the outer third of fin between spines 1 and 5, and extending partly towards spine 6, base of fin with short extension of first yellow and black body bar; base of second dorsal fin with two yellow, black dotted extensions of third and fourth yellow body blotches, a black spot immediately above fin base between first two rays, a small, black-dotted, yellow spot on lower third of the fifth ray and another between bases of last two rays, outer edge of fin with a narrow band of dense black chromatophores on interradial membranes; anal fin translucent with scattered, tiny, black chromatophores on interradial membranes, particularly on anterior twothirds of fin; pectoral fin base translucent with a large greenish silver blotch that extends on to basal one-fifth of fin, base of the dorsal ~8 rays of pectoral fin covered with a yellow, black-dotted blotch; remainder of the fin translucent; pelvic fins translucent with small black dots on interradial membranes. Colour in preservation (Fig. 11): Body pale yellowish; side of head below eye with curved, black, vertical bar formed by densely aggregated black chromatophores ending at rear corner of the mouth; operculum covered with scattered chromatophores; chromatophores more densely concentrated on posterior half, forming vertical bar of pigment that extends dorsally on to and across nape; side of body with four concentrations of chromatophores that form vertical bars, first bar complete and positioned beneath centre of first dorsal fin; second bar positioned below origin of second dorsal fin, not extending to dorsal midline, third and fourth bars positioned over end of anal fin and centre of caudal peduncle, respectively, both bars beginning at lateral midline and extending ventrally to ventral midline; a series four or five small patches of chromatophores along dorsal midline, beginning at origin of second dorsal fin and extending to end of caudal peduncle; a singular vertical row of chromatophores over base of caudal fin rays, followed by a small patch of chromatophores on upper-half of caudal fin on anterior one-third of rays; first dorsal fin with distinct black oval on outer third of fin, and smaller dark black spot on middle of fin that is an extension of vertical bar on body; second dorsal fin with three or four small, isolated dark patches of chromatophores on lower-half of fin, distal margin of fin bordered with dark chromatophores; anal fin uniformly covered with scattering of chromatophores; pelvic and pectoral fin without pigment. Genitalia: Female papillae short and conical; male papilla unknown. Habitat: Collected on open sand bottom at 229 m depth. Distribution: Known only from type locality off southwest coast of Curaçao, Caribbean Netherlands. Etymology: The specific epithetic nigrimarginatus is an adjective (black-margined) formed from the Latin niger, glossy black, and marginatus, bordered, in reference to the black margins on the dorsal and caudal fins. Remarks: The new species is placed in the genus Robinsichthys based on the following characters: first dorsal-fin pterygiophore pattern 3-22111*0, with the last pterygiophore lacking an associated dorsal-fin spine (Fig. 4B); VI spines in first dorsal fin; 11 + 17 vertebrae; two modified basicaudal scales; head pores absent; side of body with ctenoid scales; lack of fusion between hypurals 1–2 and hypurals 3–4 and the terminal vertebral element; body with several narrow, dark bands or saddles, head with dark triangular marking below eye. The new species differs from R. arrowsmithensis, and thus the original description of Robinsichthys, most notably in the following ways: (1) having pelvic fins fused to form a complete disk with well-developed anterior frenum vs. having pelvic fins completely separate, lacking frenum; (2) having a large dark blotch on the margin on the first dorsal fin vs. dorsal fin without blotch; (3) having 19 vs. 21–24 (usually 22–23) pectoralfin rays; and (4) having I,9 elements in both the second dorsal fin and anal fin vs. predominately I,10 (I, 9 in one specimen) in both fins of R. arrowsmithensis. Because R. arrowsmithensis was not included in our phylogenetic analysis, we cannot confirm with molecular data that Robinsichthys is monophyletic with regards to Birdsongichthys, and thus whether a new genus is required for B. rectus. However, there are two characters that we regard as apomorphic for Robinsichthy s that are not present in Birdsongichthys. A count of VI dorsal spines (in most specimens) is unique to Robinsichthys among other Gobiosomatini genera (a reduction from the plesiomorphic state of VII spines). Counts of 11 + 17 vertebrae are also considered apomorphic for Robinsichthys, with these counts also being independently derived in members of the Barbulifer subgroup (Elacatinus and allies). Birdsongichthys possesses neither of these synapomorphies. Birdsongichthys has head pores, while both Robinsichthys do not, but head pores and many other morphological characters that may be useful for defining genera of Gobiosomatini have more complex evolutionary histories and may be of limited phylogenetic utility (see Discussion below)., Published as part of Tornabene, Luke, Manning, Rachel, Robertson, D. Ross, Van Tassell, James L. & Baldwin, Carole C., 2023, A new lineage of deep-reef gobies from the Caribbean, including two new species and one new genus (Teleostei: Gobiidae: Gobiosomatini), pp. 322-343 in Zoological Journal of the Linnean Society 197 (2) on pages 337-340, DOI: 10.1093/zoolinnean/zlac013, http://zenodo.org/record/7622658

ROBINSICHTHYS NIGRIMARGINATUS SP. NOV. (FIGS 4B, 8–11) Black-margined goby (English), gobio de margen negro (Spanish). Zoobank registration: urn: lsid: zoobank. org:act: 9D9D60B1-2777-47FC-8CB0-6A7E0F70641C Holotype: USNM 446953, 21.0 mm female, tissue CUR19007, field number CURASUB19-02, 229 m depth, sandy bottom, East of Substation downline, Substation Curaçao, Bapor Kibra, Curaçao, 12.0838°, -68.8991°, quinaldine, HOV Curasub, Carole Baldwin, Luke Tornabene, Tico Christiaan, Sarah Yerrace, 7 May 2019. Diagnosis: First dorsal-fin pterygiophore pattern 3-22111*0 (last pterygiophore lacking an associated dorsal-fin spine; Fig. 4B); VI spines in first dorsal fin; 11 + 17 vertebrae; two modified basicaudal scales; head pores absent; second dorsal fin I,9; anal fin I,9; pelvic fins completely united by membrane to form a round disc with well-developed anterior frenum; pectoral-fin rays 19. Description: Morphometric data are presented in Table 2. Median and paired fins: First dorsal fin VI, no spines elongate; second dorsal fin I,9; anal fin I,9; pectoral-fin rays 19, extending posteriorly to a vertical through second dorsal-fin ray 2 or 3; pelvic fin I,5, all rays branched; pelvic fins connected via membrane forming a complete disk with well-developed anterior frenum connecting pelvic spines (Fig. 8); pelvic fin extending posteriorly to just before anus; tips of pelvicfin rays not ending in fleshy pads; caudal fin truncate; branched caudal-fin rays 14, segmented caudal-fin rays 17. Scales: Scales on head and nape absent; ctenoid scales on side of the body present starting above pectoral fin base and ending at base of caudal fin rays; lateral scale rows 24, transverse scale rows six; pair of modified ctenoid scales with enlarged cteni present at base of caudal fin, one each at dorsal and ventral margins of fin; ctenoid scales present on abdomen, scales slightly larger than those on side of body. Head: Jaw extending posteriorly to a vertical through midpoint of the pupil; anterior naris a short tube, posterior naris an opening with raised rim; cephalic lateralis pores absent; eyes large, 41% HL, dorsolateral. Both jaws with two rows of teeth, teeth thin, straight, pointed and evenly spaces posteriorly, becoming less regularly spaced, slightly more recurved anteriorly, with an occasional tooth in outer row slightly larger than others in row. Sensory papillae arranged as in Figure 9, with four prominent transverse rows, including rows 5i/5s present as a continuous single row. Colour in life (Fig. 10): Body translucent with a silvery sheen, an internal white area around gut; body with five dark bars formed by aggregations of black chromatophores overlaying translucent yellow areas, the first bar strongest and widest, extending from the centre of the base of the first dorsal fin to the lower belly, the second to fifth only on the lower-half of the body, thin, formed by less dense aggregations of tiny black chromatophores, the last bar partly fused to the bar on the base of the caudal fin; upper-body profile with six small, translucent yellow blotches, each with a central area of tiny, black chromatophores, the first blotch between the two dorsal fins, the second under the front of the second dorsal fin, the third under the centre of that fin, the fourth to sixth on the caudal peduncle; two elongate internal areas of dark pigment with yellow tones along the spinal column, a short one under the second body bar, a longer one from third body bar to just behind the fourth body bar, with a suggestion of a third patch above the last body bar; when the fish is laterally oriented (Fig. 10A), the silver sheen of the body is present in irregularly dispersed patches in the translucent areas, when the fish is oriented partly towards the viewer (Fig. 10B) that iridescent colour is greenish silver, distributed in patches between the body bars and between the yellow dorsal-profile blotches, and as a distinct vertical greenish silver bar just before the upper-half of the first dark body bar. Head translucent with silver sheen on cheeks; top rear corner of opercle with a translucent orange patch about half pupil size and a larger patch of tiny, welldispersed, black chromatophores over area of pink gills, an indistinct, irregularly shaped translucent orange patch behind centre of eye extending on to nape; mouth translucent, top jaw with silver sheen and a transverse line of black chromatophores across its centre, an indistinct yellow blotch on the snout immediately behind its centre; iris with most of its anterior and rear part overlaid by a dense silvery yellow sheen, top edge of the eyeball silvery yellow; a curved black bar formed by densely aggregated black chromatophores, passing from upper-rear of iris through the pupil and down to rear corner of the mouth, section of bar under eye with relatively large chromatophores overlaying a translucent yellow area, that bar bounded anteriorly and posteriorly by patches of silver iridescence. Base of caudal fin with long, thin, vertical yellow bar with an aggregation of black chromatophores down its centre that partly fuses with last body bar, fin itself translucent with patches of tiny black chromatophores along interradial membranes – two small patches along top of fin, a subterminal band down upper-half of posterior of the fin; a small, vertical patch of black-dotted yellow pigment just behind upper-half of the basal dark bar; first dorsal fin translucent with a large, horizontally oval, solid black blotch covering the outer third of fin between spines 1 and 5, and extending partly towards spine 6, base of fin with short extension of first yellow and black body bar; base of second dorsal fin with two yellow, black dotted extensions of third and fourth yellow body blotches, a black spot immediately above fin base between first two rays, a small, black-dotted, yellow spot on lower third of the fifth ray and another between bases of last two rays, outer edge of fin with a narrow band of dense black chromatophores on interradial membranes; anal fin translucent with scattered, tiny, black chromatophores on interradial membranes, particularly on anterior twothirds of fin; pectoral fin base translucent with a large greenish silver blotch that extends on to basal one-fifth of fin, base of the dorsal ~8 rays of pectoral fin covered with a yellow, black-dotted blotch; remainder of the fin translucent; pelvic fins translucent with small black dots on interradial membranes. Colour in preservation (Fig. 11): Body pale yellowish; side of head below eye with curved, black, vertical bar formed by densely aggregated black chromatophores ending at rear corner of the mouth; operculum covered with scattered chromatophores; chromatophores more densely concentrated on posterior half, forming vertical bar of pigment that extends dorsally on to and across nape; side of body with four concentrations of chromatophores that form vertical bars, first bar complete and positioned beneath centre of first dorsal fin; second bar positioned below origin of second dorsal fin, not extending to dorsal midline, third and fourth bars positioned over end of anal fin and centre of caudal peduncle, respectively, both bars beginning at lateral midline and extending ventrally to ventral midline; a series four or five small patches of chromatophores along dorsal midline, beginning at origin of second dorsal fin and extending to end of caudal peduncle; a singular vertical row of chromatophores over base of caudal fin rays, followed by a small patch of chromatophores on upper-half of caudal fin on anterior one-third of rays; first dorsal fin with distinct black oval on outer third of fin, and smaller dark black spot on middle of fin that is an extension of vertical bar on body; second dorsal fin with three or four small, isolated dark patches of chromatophores on lower-half of fin, distal margin of fin bordered with dark chromatophores; anal fin uniformly covered with scattering of chromatophores; pelvic and pectoral fin without pigment. Genitalia: Female papillae short and conical; male papilla unknown. Habitat: Collected on open sand bottom at 229 m depth. Distribution: Known only from type locality off southwest coast of Curaçao, Caribbean Netherlands. Etymology: The specific epithetic nigrimarginatus is an adjective (black-margined) formed from the Latin niger, glossy black, and marginatus, bordered, in reference to the black margins on the dorsal and caudal fins. Remarks: The new species is placed in the genus Robinsichthys based on the following characters: first dorsal-fin pterygiophore pattern 3-22111*0, with the last pterygiophore lacking an associated dorsal-fin spine (Fig. 4B); VI spines in first dorsal fin; 11 + 17 vertebrae; two modified basicaudal scales; head pores absent; side of body with ctenoid scales; lack of fusion between hypurals 1–2 and hypurals 3–4 and the terminal vertebral element; body with several narrow, dark bands or saddles, head with dark triangular marking below eye. The new species differs from R. arrowsmithensis, and thus the original description of Robinsichthys, most notably in the following ways: (1) having pelvic fins fused to form a complete disk with well-developed anterior frenum vs. having pelvic fins completely separate, lacking frenum; (2) having a large dark blotch on the margin on the first dorsal fin vs. dorsal fin without blotch; (3) having 19 vs. 21–24 (usually 22–23) pectoralfin rays; and (4) having I,9 elements in both the second dorsal fin and anal fin vs. predominately I,10 (I, 9 in one specimen) in both fins of R. arrowsmithensis. Because R. arrowsmithensis was not included in our phylogenetic analysis, we cannot confirm with molecular data that Robinsichthys is monophyletic with regards to Birdsongichthys, and thus whether a new genus is required for B. rectus. However, there are two characters that we regard as apomorphic for Robinsichthy s that are not present in Birdsongichthys. A count of VI dorsal spines (in most specimens) is unique to Robinsichthys among other Gobiosomatini genera (a reduction from the plesiomorphic state of VII spines). Counts of 11 + 17 vertebrae are also considered apomorphic for Robinsichthys, with these counts also being independently derived in members of the Barbulifer subgroup (Elacatinus and allies). Birdsongichthys possesses neither of these synapomorphies. Birdsongichthys has head pores, while both Robinsichthys do not, but head pores and many other morphological characters that may be useful for defining genera of Gobiosomatini have more complex evolutionary histories and may be of limited phylogenetic utility (see Discussion below).

BIRDSONGICHTHYS RECTUS SP. NOV. (FIGS 4A, 5–7) Straightfinned goby (English), gobio de aleta recta (Spanish). Gobiidae genus 2, species 1 Robertson et al., 2020: table 1, supplemental plate S13. Zoobank registration: urn: lsid: zoobank. org:act: 016EDDA3-572B-45CB-8B05-676076400186 Holotype: USNM 442874, 14.1 mm SL female, tissue EUS17221, field number CURASUB 17-21, 151 m depth, rubble patch, south and south-east of R / V Chapman mooring, SW of island, Kay Bay, Sint Eustatius, 17.4566, –62.9791, quinaldine, HOV Curasub and R / V Chapman, Carole Baldwin, Luke Tornabene, Bruce Brandt, 19 April 2017. Table 1 . Continued * genus is not monophyletic. ** See Birdsong et al. (1988) for wide range of vertebral counts and pterygiophore insertion patterns. Abbreviations : D 1, first dorsal fin ; D 2, second dorsal fin; A, anal fin; AP , anal fin pterygiophores. Paratypes: MUVS-V-439, 21.0 mm SL male, tissue ROA18021, field number IDABEL18-01, 213 m depth, sand and limestone rubble, outside Halfmoon Bay, Roatan, Honduras, 16.304384°, –86.598204°, quinaldine sulphate, HOV Idabel, D. Ross Robertson, Rachel Manning, Karl Stanley, 4 June 2018; UW 158074, 18.3 mm female, tissue ROA17004, cleared and stained, field number IDABEL17-01, 192 m depth, outside Halfmoon Bay, Roatan, Honduras, 16.305557°, –86.597669°, quinaldine sulphate, HOV Idabel, Luke Tornabene, Ross Robertson, Karl Stanley, 24 July 2017; UW 158128, 12.0 mm female, tissue ROA18044, field number IDABEL18-03, 241 m depth, sand rubble in rock hole, 16.3043°, –86.5982°, quinaldine sulphate, HOV Idabel, Luke Tornabene, Rachel Manning, Karl Stanley, 6 June 2018; UW 158200, 11.5 mm male, tissue ROA18212, field number IDABEL18-08, 213 m depth, sand at base of limestone, 16.3043°, –86.5982°, quinaldine sulphate, HOV Idabel, Luke Tornabene, Katherine Maslenikov, Karl Stanley, 1 June 2018; USNM 442726, 14.4 mm male, tissue EUS17073, field number CURASUB 17-18, 110 m depth, rubble and sand on wall, dropoff west of R / V Chapman mooring, Gallows Bay, Sint Eustatius, 17.4608°, –63.0179°, quinaldine, HOV Curasub and R / V Chapman, Carole Baldwin, Luke Tornabene, Barbara van Bebber, Jordan Casey, 16 April 2017. Diagnosis: Birdsongichthys rectus is the only known member of the genus, so the species diagnosis is that of the diagnosis of the genus. Description: Morphometric data are presented in Table 2. Median and paired fins: first dorsal fin VII(6), first spine extremely elongate in some specimens, extending posteriorly to point between middle of soft dorsal fin and base of anal fin when flat against body; second dorsal fin I,8*(1) or I,9(5); anal fin I,8*(6); pectoral-fin rays 16*(3), 17 (2), 18(1), extending posteriorly to a vertical through second dorsal-fin ray 2 or 3; pelvic fin I,5* (5), all rays unbranched; pelvic fins well separated, lacking anterior frenum, membrane connecting fifth rays absent or reduced; fourth and fifth pelvic-fin rays approximately equal in length, extending posteriorly to base of anal-fin rays 6–7; tips of pelvic-fin rays not ending in fleshy pads; caudal fin truncate or emarginate; branched caudal-fin rays 12 (4)*, segmented caudal-fin rays 17(5)*, caudal fin broken in many specimens. Scales: Scales on head and nape absent; ctenoid scales on side of body present starting above pectoral fin and continuing to caudal peduncle; lateral scale rows 22–23, transverse scale rows 5, scales counted from three specimens including holotype, scales lost in other specimens; pair of modified ctenoid scales with enlarged cteni present at base of caudal fin, one each at dorsal and ventral margins of fin (Fig. 5); abdomen and chest each with small patch of ctenoid scales. Head: Jaw extending posteriorly to a vertical through anterior margin of eye or midpoint of pupil; anterior naris a short tube, posterior naris an opening flush with snout; cephalic lateralis pores small, pores B′, D(s), E′ present in four specimens, pores B′ extremely small and pore D missing in holotype, pores B′ not visible in paratype UW 158200; eyes large, 30–36% HL, positioned dorsolaterally. Upper jaw with two to three rows of teeth anteriorly, outermost row enlarged widely spaced canines continuing along entire upper jaw to the end of the premaxilla, innermost two rows tightly packed, shorter, and conical, becoming single row posteriorly; lower jaw with three to five rows of teeth anteriorly, outermost row restricted to tip of dentary and slightly larger than innermost rows, innermost rows becoming two rows posteriorly. Sensory papillae not visible on most specimens due to small size of specimens and damage to the face from collection. Genitalia: Male urogenital papilla elongate and pointed; female urogenital papilla short, bulbous or slightly conical. Colour in fresh specimens (Fig. 6): Side of body translucent with iridescent scale margins; three horizontal rows of small, bright yellow spots along flank, spots largest in the bottom row; first row with 12–15 spots starting on nape and continuing along dorsal midline, ending just anterior to caudal-fin base; second row with 13–15 spots starting above pectoral fin and continuing along lateral midline, ending just anterior to caudal-fin base; third row with eight to nine spots starting just anterior to anal-fin origin and continuing along ventral midline, ending just anterior to caudal-fin base; tips of snout and upper and lower jaws with small patch of yellow; eye with upperhalf of iris black, lower-half yellow; a broad, oblique yellow to orange stripe extending from lower-rear part of iris down on to preopercle; remainder of head translucent; pink coloration of red gill arches visible through operculum; pectoral-fin base with yellow spot on yellow upper margin of some specimens; dorsal fins with uniform translucent yellow wash with scattered iridophores, and with a distinct horizontal series of yellow spots along midpoints of spines and rays in some specimens; pectoral fins pale or uniformly translucent yellow; caudal fin faintly translucent yellow with two dark yellow, vertical bars at base; anal and pelvic fins uniformly translucent yellow. Colour in preservation (Fig. 7): Body uniformly pale yellowish, no dark pigmentation remains in any preserved specimens, including those fixed either in 95% ethanol or in formalin followed by storage in 70–75% ethanol. Habitat: Known from 110-241 m depth on sand and rubble substrates situated on or around the base of walls or steep rocky slopes on deep reefs. Distribution: Known from Roatan, Honduras and Sint Eustatius, Caribbean Netherlands. Etymology: The epithet is from the Latin adjective rectus, straight, which refers to the straight, unbranched rays of the pelvic fin., Published as part of Tornabene, Luke, Manning, Rachel, Robertson, D. Ross, Van Tassell, James L. & Baldwin, Carole C., 2023, A new lineage of deep-reef gobies from the Caribbean, including two new species and one new genus (Teleostei: Gobiidae: Gobiosomatini), pp. 322-343 in Zoological Journal of the Linnean Society 197 (2) on pages 333-337, DOI: 10.1093/zoolinnean/zlac013, http://zenodo.org/record/7622658, {"references":["Robertson DR, Estape CJ, Estape AM, Pena E, Tornabene L, Baldwin CC. 2020. The marine fishes of St Eustatius Island, northeastern Caribbean: an annotated, photographic catalog. ZooKeys 1007: 145 - 180."]}

Leptoderma ospesca n. sp. (Fig. 1) Holotype. USNM 421478, 197 mm SL; El Salvador, Central America, Eastern Pacific Ocean, research vessel B/ O Miguel Oliver station MOP 11���94, trawl on soft-bottom from 12 �� 53 ' 15.72 " N, 90 �� 9 ' 30.6 " W to 12 �� 53 ' 22.9194 " N, 90 �� 7 ' 56.28 " W, at 1368���1406 m; December 11, 2010; D. Ross Robertson. Diagnosis. Leptoderma ospesca differs from L. affinis, L. macrops and L. retropinna by having a distinct gap between the posterior ends of the dorsal and anal fins and the origins of the upper and lower procurrent caudal-fin rays. From L. lubricum, the new species differs by lacking dermal papillae along the lateral line, by having pectoral fins low on body (vs. high) with 6 fin rays (vs. 7���8), and by having fewer caudal-fin rays (16, vs. 17���18). From L. macrophthalmum, the new species differs by having a longer pre-dorsal length (54.9 % SL, vs. 49.3 %), fewer pectoral-fin rays (6, vs. 8), fewer pelvic-fin rays (5, vs. 8) and longer dorsal- and anal-fin rays (vs. shorter). Leptoderma ospesca also differs from all congeneric species by having fewer pre-ural vertebrae (60, vs. 64���84). Description. Measurements and counts of L. ospesca and comparative data are presented in Table 1. Body naked, relatively long, subcylindrical anteriorly and laterally compressed posteriorly, with greatest body depth at a distance of 24.3 % SL from snout; dermal papillae absent from head, along lateral line and dorsum; lateral line pores inconspicuous. Head length (HL) almost twice greatest body depth. Nostrils with large longitudinal openings, partly covered by a dermal flap, located nearer to eye than to snout tip. Mandibular pores 4, supraorbital pores 3, suborbital pores 4 and preopercular pores 3. Snout blunt, relatively large, its length 30.3 % of HL, shorter than eye. Mouth subterminal. Teeth minute to small, conical and well separated, present in one row on premaxilla and one on dentary; no teeth on palate. Supramaxilla slender, curved. Upper jaw relatively short, barely reaching front border of eye, its length 33.0% of HL. Lower jaw ending under orbit. Eyes large, 37.5 % of HL, directed outward and slightly upward, with upper margin protruding above dorsal profile of head. Interorbital space relatively large, 17.2 % of HL. Postorbital length 32.2 % of HL. Maximal head width (behind eyes) 34.5 % of HL. Pseudobranchiae small. Pectoral fins inserted relatively low on body, uppermost fin ray at level of dorsal corner of gill opening, slightly below midbody. Pelvic fins abdominal, length 71.1 % of pectoral-fin length. Pelvic-fin base located just in front of anus. Genital papilla short, located between anus and anal-fin origin. Anal-fin origin well in advance of dorsal-fin origin; length of dorsal-fin base 60.4 % of length of anal-fin base. Dorsal- and anal-fin rays long, longer rays greater than half body depth at level of anus. Both dorsal and anal fins separated from caudal fin. Procurrent caudal-fin rays numerous, extending far forward on caudal peduncle. Dorsal and anal procurrent caudal sections about equal in length. Caudal fin small, forked, with 8 principal rays in each lobe. Color in life. Head, eye, and anterior part of body bluish black, turning gradually lighter brownish towards posterior part of body (Fig. 1 A). TABLE. 1. M��rph��metric and meristic characters ��f six Leptoderma species. Measurements and c��unts n��t attainable (i.e. structure absent ��r inc��nspicu��us) ��r unavailable are represented an en���dash (���). When the number ��f examined specimens used t�� determine the respective measurement ��r c��unt differs fr��m the t��tal, the number ��f examined specimens is indicated in parenthesis. Character L. affinis L. lubricum L. macrops L. macrophthalmum L. ospesca L.retropinna n= 1 2, * n= 21 3, * n= 52 4 n= 1 5, * n= 1 1, * n= 27 6 (mm) 222.5 29.7 ��� 210 141 ��� 210 151 197 28.4 ��� 152 Morphometric characters as % SL ���d��rsal length ��� 55.1 ��� 61.1 47.3 ��� 56.4 (49) 49.3 59.4 55.0��� 58.7 (6) ���anal length ��� 41.0��� 46.5 (20) 35.5 ��� 44.2 (49) 45.7 46.4 40.9 ��� 45.6 (6) ���pelvic length ��� 36.8 ��� 41.8 (20) 31.0��� 43.8 (49) 39.5 38.0 34.4 ��� 38.7 (6) D��rsal-fin base, length ��� 25.3 ��� 28.9 (5) 44.7 ��� 58.1 (14) 31.1 25.5 41.0��� 47.6 (6) D��rsal pr��current caudal-fin secti��n, length ��� 15.0��� 16.6 (5) 3.2-3.6 (3) 15.5 14.1 4.5-4.7 (2) Anal-fin base, length ��� 36.6 ���41.0 (5) 58.6 ��� 73.6 (14) 40.7 42.2 50.0���59.0 (6) ventral pr��current caudal-fin secti��n, length ��� 14.0��� 16.9 (5) 3.4-4.2 (3) 15.1 14.0 4.1-5.3 (3) Pect��ral-fin length ��� 5.0��� 11.4 (5) 6.0��� 8.4 (14) 7.3 7.0 4.9 ��� 5.8 (5) Pelvic-fin length ��� 1.8 ��� 6.9 (5) 3.9 ��� 5.8 (14) 4.8 5.0 4.0��� 4.9 (5) Greatest b��dy depth c. 11.1 11.1 ��� 14.7 (5) 9.8 ��� 12.3 (14) 12.8 13.4 7.8 ��� 9.2 (6) Caudal-peduncle depth ��� 2.0��� 2.8 (5) 0.9 ��� 1.5 (14) 2.1 1.8 1.1 ��� 1.3 (6) Maximal head width (behind eyes) ��� 8.6 ��� 14.9 (20) 7.8 ��� 9.2 (14) 11.3 8.4 6.6 ��� 8.6 (6) length c. 22.2 22.8 ��� 25.7 (5) 16.7 ��� 24.4 (49) 23.2 24.3 18.7 ��� 21.7 (6) Sn��ut length ��� 7.0��� 8.3 (5) 3.8 ���7.0 (49) 7 7.4 5.3 ���7.0 (6) H��riz��ntal ��rbital diameter ��� 7.5 ���9.0 (5) 5.7 ��� 9.4 (49) 9.3 9.1 3.3 ��� 7.2 (6) P��st��rbital length ��� ��� ��� 6.9 7.8 ��� Inter��rbital distance ��� 3.1 ��� 5.1 (5) 1.6 ��� 5.6 (49) 3.4 4.2 3.3 ��� 5.7 (6) Upper-��aw length ��� 5.6 ��� 8.9 (5) 4.6 ��� 7.3 (49) 7.9 8.0 5.3 ��� 7.3 (6) ������continued on the next page vertebrae, hemal (excl. ur��style) ��� 49 ��� 55 (4) ��� 51 ��� ��� vertebrae, t��tal (excl. ur��style) ��� 64 ��� 77 (20) 79 ��� 84 (35) 66 60 77 (1) Dermal papillae ��n lateral line:es:es (5):es;��;��;�� (5) US;M 421478, net al. (1965), nakami? @ukui (2010), net al. (2011), nakami? @ukui (2010), nFIGURE. 1. Leptoderma ospesca n. sp., holotype (USNM 421478), A: preserved entire specimen; B: newly caught entire specimen, arrows indicate position of gaps between dorsal and anal fin rays and upper and lower procurrent caudal rays, respectively; C close-up of head region of newly caught specimen. Note: The structure near the lateral midline above the anus that is visible in Fig. 1 A (but not obvious in Figs. 1 B or C), is a piece of loose skin adhered to the body. Color in alcohol. Head and anterior part of body black, turning gradually lighter grayish towards posterior part of body (Fig. 1 B). Distribution. Known only from the holotype collected off El Salvador, between 12 �� 53 ' 15.72 " N, 90 �� 9 ' 30.6 " W and 12 �� 53 ' 22.9194 " N, 90 �� 7 ' 56.28 " W, Central America, eastern Pacific Ocean, at a depth of 1368���1406 m. Etymology. The specific name ��� ospesca ��� recognizes the contribution of OSPESCA (Organizaci��n del Sector Pesquero y Ac��icola de Centroamerica) to the discovery of deepwater marine fishes in Central America by sponsoring research cruises of the Spanish research vessel B/ O Miguel Oliver on both the Pacific (2010) and Atlantic (2011) coasts of Central America. Suggested common names. Eastern eel-slickhead; Cabezalisa oriental., Published as part of Angulo, Arturo, Baldwin, Carole C. & Robertson, D. Ross, 2016, A new species of Leptoderma Vaillant, 1886 (Osmeriformes: Alepocephalidae) from the Pacific coast of Central America, pp. 493-500 in Zootaxa 4066 (4) on pages 494-498, DOI: 10.11646/zootaxa.4066.4.10, http://zenodo.org/record/266387, {"references":["Abe, T., Marumo, R. & Kawaguchi, K. (1965) Description of a new alepocephalid fish from Suruga Bay. Japanese Journal of Ichthyology, 13, 69 - 72.","Markle, D. F. (1976) Preliminary studies on the systematics of deep-sea Alocephaoidea (Pisces: Salmoniformes). Ph. D. Dissertation, the Faculty of the School of Marine Science, the College of William and Mary in Virginia, 265 pp."]}

Leptoderma ospesca n. sp. (Fig. 1) Holotype. USNM 421478, 197 mm SL; El Salvador, Central America, Eastern Pacific Ocean, research vessel B/ O Miguel Oliver station MOP 11–94, trawl on soft-bottom from 12 º 53 ' 15.72 " N, 90 º 9 ' 30.6 " W to 12 º 53 ' 22.9194 " N, 90 º 7 ' 56.28 " W, at 1368–1406 m; December 11, 2010; D. Ross Robertson. Diagnosis. Leptoderma ospesca differs from L. affinis, L. macrops and L. retropinna by having a distinct gap between the posterior ends of the dorsal and anal fins and the origins of the upper and lower procurrent caudal-fin rays. From L. lubricum, the new species differs by lacking dermal papillae along the lateral line, by having pectoral fins low on body (vs. high) with 6 fin rays (vs. 7–8), and by having fewer caudal-fin rays (16, vs. 17–18). From L. macrophthalmum, the new species differs by having a longer pre-dorsal length (54.9 % SL, vs. 49.3 %), fewer pectoral-fin rays (6, vs. 8), fewer pelvic-fin rays (5, vs. 8) and longer dorsal- and anal-fin rays (vs. shorter). Leptoderma ospesca also differs from all congeneric species by having fewer pre-ural vertebrae (60, vs. 64–84). Description. Measurements and counts of L. ospesca and comparative data are presented in Table 1. Body naked, relatively long, subcylindrical anteriorly and laterally compressed posteriorly, with greatest body depth at a distance of 24.3 % SL from snout; dermal papillae absent from head, along lateral line and dorsum; lateral line pores inconspicuous. Head length (HL) almost twice greatest body depth. Nostrils with large longitudinal openings, partly covered by a dermal flap, located nearer to eye than to snout tip. Mandibular pores 4, supraorbital pores 3, suborbital pores 4 and preopercular pores 3. Snout blunt, relatively large, its length 30.3 % of HL, shorter than eye. Mouth subterminal. Teeth minute to small, conical and well separated, present in one row on premaxilla and one on dentary; no teeth on palate. Supramaxilla slender, curved. Upper jaw relatively short, barely reaching front border of eye, its length 33.0% of HL. Lower jaw ending under orbit. Eyes large, 37.5 % of HL, directed outward and slightly upward, with upper margin protruding above dorsal profile of head. Interorbital space relatively large, 17.2 % of HL. Postorbital length 32.2 % of HL. Maximal head width (behind eyes) 34.5 % of HL. Pseudobranchiae small. Pectoral fins inserted relatively low on body, uppermost fin ray at level of dorsal corner of gill opening, slightly below midbody. Pelvic fins abdominal, length 71.1 % of pectoral-fin length. Pelvic-fin base located just in front of anus. Genital papilla short, located between anus and anal-fin origin. Anal-fin origin well in advance of dorsal-fin origin; length of dorsal-fin base 60.4 % of length of anal-fin base. Dorsal- and anal-fin rays long, longer rays greater than half body depth at level of anus. Both dorsal and anal fins separated from caudal fin. Procurrent caudal-fin rays numerous, extending far forward on caudal peduncle. Dorsal and anal procurrent caudal sections about equal in length. Caudal fin small, forked, with 8 principal rays in each lobe. Color in life. Head, eye, and anterior part of body bluish black, turning gradually lighter brownish towards posterior part of body (Fig. 1 A). TABLE. 1. Mοrphοmetric and meristic characters οf six Leptoderma species. Measurements and cοunts nοt attainable (i.e. structure absent οr incοnspicuοus) οr unavailable are represented an en‾dash (‾). When the number οf examined specimens used tο determine the respective measurement οr cοunt differs frοm the tοtal, the number οf examined specimens is indicated in parenthesis. Character L. affinis L. lubricum L. macrops L. macrophthalmum L. ospesca L.retropinna n= 1 2, * n= 21 3, * n= 52 4 n= 1 5, * n= 1 1, * n= 27 6 (mm) 222.5 29.7 ‾ 210 141 ‾ 210 151 197 28.4 ‾ 152 Morphometric characters as % SL ‾dοrsal length ‾ 55.1 ‾ 61.1 47.3 ‾ 56.4 (49) 49.3 59.4 55.0‾ 58.7 (6) ‾anal length ‾ 41.0‾ 46.5 (20) 35.5 ‾ 44.2 (49) 45.7 46.4 40.9 ‾ 45.6 (6) ‾pelvic length ‾ 36.8 ‾ 41.8 (20) 31.0‾ 43.8 (49) 39.5 38.0 34.4 ‾ 38.7 (6) Dοrsal-fin base, length ‾ 25.3 ‾ 28.9 (5) 44.7 ‾ 58.1 (14) 31.1 25.5 41.0‾ 47.6 (6) Dοrsal prοcurrent caudal-fin sectiοn, length ‾ 15.0‾ 16.6 (5) 3.2-3.6 (3) 15.5 14.1 4.5-4.7 (2) Anal-fin base, length ‾ 36.6 ‾41.0 (5) 58.6 ‾ 73.6 (14) 40.7 42.2 50.0‾59.0 (6) ventral prοcurrent caudal-fin sectiοn, length ‾ 14.0‾ 16.9 (5) 3.4-4.2 (3) 15.1 14.0 4.1-5.3 (3) Pectοral-fin length ‾ 5.0‾ 11.4 (5) 6.0‾ 8.4 (14) 7.3 7.0 4.9 ‾ 5.8 (5) Pelvic-fin length ‾ 1.8 ‾ 6.9 (5) 3.9 ‾ 5.8 (14) 4.8 5.0 4.0‾ 4.9 (5) Greatest bοdy depth c. 11.1 11.1 ‾ 14.7 (5) 9.8 ‾ 12.3 (14) 12.8 13.4 7.8 ‾ 9.2 (6) Caudal-peduncle depth ‾ 2.0‾ 2.8 (5) 0.9 ‾ 1.5 (14) 2.1 1.8 1.1 ‾ 1.3 (6) Maximal head width (behind eyes) ‾ 8.6 ‾ 14.9 (20) 7.8 ‾ 9.2 (14) 11.3 8.4 6.6 ‾ 8.6 (6) length c. 22.2 22.8 ‾ 25.7 (5) 16.7 ‾ 24.4 (49) 23.2 24.3 18.7 ‾ 21.7 (6) Snοut length ‾ 7.0‾ 8.3 (5) 3.8 ‾7.0 (49) 7 7.4 5.3 ‾7.0 (6) Hοrizοntal οrbital diameter ‾ 7.5 ‾9.0 (5) 5.7 ‾ 9.4 (49) 9.3 9.1 3.3 ‾ 7.2 (6) Pοstοrbital length ‾ ‾ ‾ 6.9 7.8 ‾ Interοrbital distance ‾ 3.1 ‾ 5.1 (5) 1.6 ‾ 5.6 (49) 3.4 4.2 3.3 ‾ 5.7 (6) Upper-ϳaw length ‾ 5.6 ‾ 8.9 (5) 4.6 ‾ 7.3 (49) 7.9 8.0 5.3 ‾ 7.3 (6) ……continued on the next page vertebrae, hemal (excl. urοstyle) ‾ 49 ‾ 55 (4) ‾ 51 ‾ ‾ vertebrae, tοtal (excl. urοstyle) ‾ 64 ‾ 77 (20) 79 ‾ 84 (35) 66 60 77 (1) Dermal papillae οn lateral line:es:es (5):es;ο;ο;ο (5) US;M 421478, net al. (1965), nakami? @ukui (2010), net al. (2011), nakami? @ukui (2010), n FIGURE. 1. Leptoderma ospesca n. sp., holotype (USNM 421478), A: preserved entire specimen; B: newly caught entire specimen, arrows indicate position of gaps between dorsal and anal fin rays and upper and lower procurrent caudal rays, respectively; C close-up of head region of newly caught specimen. Note: The structure near the lateral midline above the anus that is visible in Fig. 1 A (but not obvious in Figs. 1 B or C), is a piece of loose skin adhered to the body. Color in alcohol. Head and anterior part of body black, turning gradually lighter grayish towards posterior part of body (Fig. 1 B). Distribution. Known only from the holotype collected off El Salvador, between 12 º 53 ' 15.72 " N, 90 º 9 ' 30.6 " W and 12 º 53 ' 22.9194 " N, 90 º 7 ' 56.28 " W, Central America, eastern Pacific Ocean, at a depth of 1368–1406 m. Etymology. The specific name “ ospesca ” recognizes the contribution of OSPESCA (Organización del Sector Pesquero y Acúicola de Centroamerica) to the discovery of deepwater marine fishes in Central America by sponsoring research cruises of the Spanish research vessel B/ O Miguel Oliver on both the Pacific (2010) and Atlantic (2011) coasts of Central America. Suggested common names. Eastern eel-slickhead; Cabezalisa oriental.

Key to species of Leptoderma 1 Both dorsal and anal fins confluent with caudal fin; procurrent caudal-fin rays few (dorsal 7–9; ventral 6–8) not extending far forward on caudal peduncle; dorsal procurrent caudal-fin section about 3.2–4.2 % of SL; ventral procurrent caudal-fin section about 3.4–5.3 % of SL................................................................................. 2 - Both dorsal and anal fins separated from caudal fin; procurrent caudal-fin rays numerous (dorsal 21–29; ventral 19–26) extending far forward on caudal peduncle; dorsal procurrent caudal-fin section 14.1–16.6 % of SL; ventral procurrent caudalfin section 14.0– 16.9 % of SL........................................................................... 4 2 Dermal papillae absent along lateral line (Indo-West Pacific Ocean)................................... L. retropinna - Dermal papillae present along lateral line.................................................................. 3 3 Dorsal-fin rays about 66; anal-fin rays about 85 (Indo-West Pacific)...................................... L. affinis - Dorsal-fin rays 57–67, usually less than 64; anal-fin rays 74–85, usually less than 82 (Atlantic)............... L. macrops 4 Dermal papillae present along lateral line; lateral line pores 55–66; pectoral fin inserted high on body, central fin ray level with dorsal corner of gill opening, slightly above midline of body; length of anal-fin base 36.6 –41.0% of SL (Western Pacific).............................................................................................. L. lubricum - Dermal papillae absent along lateral line; lateral line pores, if conspicuous, about 48; pectoral fin inserted low on body, uppermost fin ray level with dorsal corner of gill opening, slightly below midline of body; length of anal-fin base 40.7–42.2 % of SL.................................................................................................... 5 5 Pre-dorsal region short, length about 49.3 % SL; pectoral-fin rays 8; pelvic-fin rays 8; dorsal- and anal-fin rays short, longer rays less than half body depth at anus; pre-ural vertebrae about 66 (North Atlantic).................. L. macrophthalmum - Pre-dorsal region long, length about 54.9 % SL; pectoral-fin rays 6; pelvic-fin rays 5; dorsal- and anal-fin rays long, longer rays greater than half body depth at anus; pre-ural vertebrae about 60 (Eastern Pacific).......................... L. ospesca

GYMNELEOTRIS BLEEKER, 1874 TYPE SPECIES: GYMNELEOTRIS SEMINUDUS (G UNTHER €, 1864: 304, DESCRIBED AS ELEOTRIS SEMINUDUS G UNTHER €, 1864, BY ORIGINAL DESIGNATION) Diagnosis Possesses all taxonomic characters present in most members of Gobiosomatini and the Gobiosoma group (first dorsal-fin spines VII, pterygiophore insertion pattern of 3 – 221110, 27 vertebrae – 11 precaudal and 16 caudal, hypurals 1 and 2 fused to some extent with hypurals 3 and 4 and the terminal vertebral element, one epural); pelvic fins well separated, lacking both anterior frenum and well-developed membrane connecting innermost rays; pelvic-fin rays 1 – 5 branched and never with flattened or fleshy tips; pelvic-fin rays extending posteriorly to anus, never reaching origin of anterior anal-fin rays; side of body with scales (modified basicaudal scales absent), scales embedded anteriorly; two anal-fin pterygiophores inserted before first haemal spine; papillae rows 5i and 5s separate; cephalic sensory canals present, pores B 0 , F, and H 0 present; second dorsal-fin rays I,10; anal-fin rays I,9; body dark brown to black with prominent narrow white vertical or diagonal bars on head, body, and dorsal fins. The one known species occurs in the eastern Pacific Ocean. Remarks The monotypic Gymneleotris was recovered in a clade with the genera Chriolepis, Pycnomma (now as Chriolepis), and Varicus imswe (now as Paedovaricus gen. nov.). Its position within this clade is unclear, as the sister relationship between Gymneleotris and Paedovaricus is weakly supported (Bayesian posterior probability of 0.69). Morphologically there is little evidence for a sister relationship between the two genera. At present, the only character distinguishing Gymneleotris from Chriolepis (as circumscribed herein) is the absence of modified basicaudal scales in Gymneleotris. The inclusion of additional eastern Pacific taxa and or phylogenetically informative characters may help resolve the position of this taxon and ultimately support a closer relationship (or nested relationship) with Chriolepis. Pending further investigation Gymneleotris is maintained here as a valid genus., Published as part of Tornabene, Luke, Van Tassell, James L., Gilmore, Richard G., Robertson, David Ross, Young, Forrest & Baldwin, Carole C., 2016, Molecular phylogeny, analysis of character evolution, and submersible collections enable a new classification of a diverse group of gobies (Teleostei: Gobiidae: Nes subgroup), including nine new species and four new genera, pp. 764-812 in Zoological Journal of the Linnean Society 177 (4) on page 779, DOI: 10.1111/zoj.12394, http://zenodo.org/record/5460757

Tornabene, Luke, Van Tassell, James L., Gilmore, Richard G., Robertson, David Ross, Young, Forrest, Baldwin, Carole C. (2016): Molecular phylogeny, analysis of character evolution, and submersible collections enable a new classification of a diverse group of gobies (Teleostei: Gobiidae: Nes subgroup), including nine new species and four new genera. Zoological Journal of the Linnean Society 177 (4): 764-812, DOI: 10.1111/zoj.12394, URL: http://doi.wiley.com/10.1111/zoj.12394

A new species of Leptoderma Vaillant, 1886 is described from a single specimen trawled at 1368–1406 m depth off El Salvador, Central America, tropical eastern Pacific. Leptoderma ospesca n. sp. can be readily distinguished from its congeners by the following combination of characters: dermal papillae absent along the lateral line, pectoral-fin rays 6, pelvic-fin rays 5, pre-dorsal length 54.9% of SL, both dorsal and anal fins separated from the caudal fin, dorsal- and anal-fin rays long, procurrent caudal-fin rays numerous and extending far forward on caudal peduncle, caudal-fin rays 16, and total pre-ural vertebrae 60. A key to the species of the genus is presented. UCR::Vicerrectoría de Investigación::Unidades de Investigación::Ciencias Básicas::Centro de Investigación en Ciencias del Mar y Limnología (CIMAR)

A new species of western Atlantic Coryphopterus is described from mesophotic depths off Curaçao, southern Caribbean. Coryphopterus curasub sp. n., is similar to C. dicrus in, among other features, having two prominent pigment spots of roughly equal intensity on the pectoral-fin base, the pelvic fins fused to form a disk, and no pelvic frenum. The two species can be differentiated by body depth (shallower in C. curasub at origin of dorsal fin and caudal peduncle); differences in the pigmentation on the head, trunk, and basicaudal region; and usually by total number of rays (spinous plus soft) in the second dorsal fin (10–11, usually 11, in C. curasub, 10 in C. dicrus). Coryphopterus curasub differs from other Coryphopterus species that have a prominent pigment spot on the lower portion of the pectoral-fin base (C. punctipectophorus and C. venezuelae) in, among other features, lacking a pelvic frenum. Coryphopterus curasub was collected between 70 and 80 m, the deepest depth range known for the genus. Collections of C. venezuelae at depths of 65–69 m extend the depth range of that species by approximately 50 m. Mitochondrial cytochrome c oxidase subunit I (COI) data corroborate the recognition of C. curasub as a distinct species but do not rigorously resolve its relationships within the genus. A revised key to the western Atlantic species of Coryphopterus is presented.

Bythaelurus giddingsi sp. nov. (Figures 1���4, Plates 1���2, Tables 1���2) Proposed common name. Gal��pagos Catshark. Bythaelurus sp.: Compagno (2005 b: 20), McCosker & Rosenblatt (2010: 172, 187), Ruiz et al., (2011: 12). Bythaelurus sp. B: Compagno et al. (2005: 215, pl. 35), Kyne & Simpendorfer (2007: 53), Hearn et al. (2009: 51). Material examined. All specimens are from the Gal��pagos Islands, collected aboard the submersible Johnson Sea- Link during 1995 and 1998. Holotype. CAS 210091, 402 mm TL, female, Darwin Island, 0 1 o 42.0' N, 92 o00.0' W, (station number JSL 3103, field number JM 145), 428 m depth, collected by J.E. McCosker on 18 July 1998. Paratypes. MNHN 2010 -0004 (originally CAS 210092), 324 mm TL, immature male, JM 146; SIO 99-99, 217 mm TL, immature male, JM 147; and CAS 210093, 300 mm TL (partially dissected), immature female, JM 148; Darwin Island, all collected with the holotype. CAS 86559, 261 mm, immature male, Darwin Island, 0 1 o 42.05 ' N, 92 o00.02' W (JSL 3967), 511 m depth, collected by J.E. McCosker on 21 Nov. 1995. USNM 364283, 301 mm TL, immature female, Marchena Island, N. shore, 0 0o 24.0' N, 90 o 26.5 ' W (JSL 3106, JM 175 & CB 98 - 49), 562 m depth, collected by C.C. Baldwin on 19 July 1998. SAM- 35042, 453 mm TL, male, Marchena Island, N. shore, 0 0o 24.0' N, 90 o 26.5 ' W (JSL 3109, JM 199), 454 m depth, collected by J.E. McCosker on 21 July 1998. Diagnosis. A moderate-sized species of Bythaelurus with the following characteristics: snout bluntly rounded; preoral length 4.7���6.8 % TL; head short, its length about equal to pectoral-pelvic space, 21���24 % TL; precaudal length 73���80 % TL; dorsal and anal fins subequal; length of anal-fin base about equal to length of interdorsal space; precaudal vertebrae 81���85 (mean = 82.6); color chocolate brown dorsally, paler on ventral surface from snout to anus; body, flanks, caudal and median fins overlain with pale spots about equal in size to eye, becoming smaller below lateral midline, and showing a bilaterally asymmetrical pattern. Description. Measurements are in millimeters, parenthetically followed by their proportions as percentages of total length and precaudal length, respectively. Proportions and counts of the holotype and paratypes are listed in Tables 1���2. continued next page Total length 402 (100, 124); precaudal length 323 (80.3, 100). Tip of snout to: upper symphysis 41 (10.2, 12.7); nostrils 18.3 (4.6, 5.7); orbits 24 (6.0, 7.4); spiracles 48 (11.9, 14.9); 1 st gill openings 70 (17.4, 21.7); 2 nd gill openings 82 (20.3, 25.4); 3 rd gill openings 89 (22.1, 27.6); 4 th gill openings 92 (22.9, 28.5); 5 th gill openings (= head length) 96.5 (24.0, 29.9); pectoral origins 90 (22.4, 27.9); pelvic origins 190 (47.3, 58.9); 1 st dorsal origin 187 (46.5, 57.9); 2 nd dorsal origin 259 (64.4, 80.2); anal origin 239 (59.5, 74.0); vent 194 (48.3, 60.1). Distance between: vent and caudal-fin tip 208 (51.7, 64.4); 1 st and 2 nd dorsal origins 72 (17.9, 22.3); 1 st and 2 nd dorsal bases 43 (10.7, 13.3); 2 nd dorsal and upper caudal origins 53.5 (13.3, 16.7); 2 nd dorsal base and upper caudal origin 24.5 (60.9, 75.9); pectoral and pelvic origins 100 (24.9, 31.0); pelvic and anal bases 23.5 (58.6, 72.8); anal and lower caudal origins 70 (17.4, 21.7); anal base and lower caudal origin 32 (80.0, 99.1). Eyes (palpebral apertures or fleshy orbits): length 20 (5.0, 6.2); height 7 (1.7, 2.2); width across anterior corners (interorbital) 34 (8.5, 10.5); eyeball diameter 8.5 (2.1, 2.6). Nostril width 12 (3.0, 3.7); internarial space 10 (2.5, 3.1). Spiracles: diameter 4 (1.0, 1.2); space between spiracles and eyes 3.3 (0.8, 1.0). Mouth (jaws in retracted position): length 21.5 (5.4, 6.6); width 46 (11.4, 14.2); width across outer edges of jaws 52 (12.9, 16.1). Gill-opening heights: 1 st 12 (3.0, 3.7); 2 nd 11 (2.7, 3.4); 3 rd 11 (2.7, 3.4); 4 th 11 (2.7, 3.4); 5 th 10 (2.5, 3.1). Head height: at spiracles 35 (8.7, 10.8); at 1 st gill openings 43 (10.7, 13.3); at 5 th gill openings 53 (13.2, 16.4). Trunk height: at 1 st dorsal origin 51 (12.7, 15.8); at pelvic origins 41.5 (10.3, 12.8); at pelvic insertions 24 (6.0, 7.4). Girth: at 1 st dorsal origin 125 (31.1, 38.7); at 2 nd dorsal origin 57 (14.2, 17.6). Caudal peduncle height: at 2 nd dorsal insertion 17 (4.2, 5.3); at upper caudal origin 13 (3.2, 4.0). Caudal peduncle width: at 2 nd dorsal insertion 10.5 (2.6, 3.3); at upper caudal origin 7 (1.7, 2.2). Pectoral fin (right side - left is damaged), length of: anterior margin 50 (12.4, 15.5); posterior margin 45 (11.2, 13.9); origin to free rear tip 35 (8.5, 10.8); inner margin 30.5 (7.6, 9.4). Pelvic fins, length of: anterior margin 26.5 (6.6, 8.3); posterior margin 28.5 (7.1, 8.8); base 29 (7.2, 9.0); origin to free rear tip 40 (10.0, 12.4); inner margin 19.5 (4.9, 6.0); height 18 (4.5, 5.6). 1 st dorsal fin, length of: anterior margin 37.5 (9.3, 11.6); posterior margin 24 (6.0, 7.4); base 28 (7.0, 8.7); inner margin 10 (2.5, 3.1); height 27 (6.7, 8.4). 2 nd dorsal fin, length of: anterior margin 41 (10.2, 12.7); posterior margin 24 (6.0, 7.4); base 29 (7.2, 9.0); inner margin 10 (2.5, 3.1); height 26 (6.5, 8.0). Anal fin, length of: anterior margin 31 (7.7, 9.6); posterior margin 31 (7.7, 9.6); base 32 (8.8, 9.0); inner margin 8.5 (2.1, 2.6); height 21 (5.2, 6.5). Caudal fin, length of: dorsal margin 93 (23.1, 28.8); preventral margin 28 (7.0, 8.7); upper and lower postventral margins (combined) 47 (11.7, 14.6); subterminal margin 18.5 (4.6, 5.7); terminal margin 23 (5.7, 7.1); terminal lobe or sector 26 (6.5, 8.0); width of dorsal lobe at postventral notch 8 (2.0, 2.5); width of ventral lobe at postventral notch 2.5 (0.6, 0.8). Vertebral counts are given in Table 2. Snout bluntly rounded, without a pointed knoblike tip (Figure 2, Plate 1). Head short, its length about equal to pectoral-pelvic space. Head depressed, roughly trapezoidal in cross-section. Outline of head in lateral view convex dorsally. Preoral snout short, nearly 1 / 2 mouth width, broadly rounded in dorsolateral view, minimally indented anterior to nostrils. External eye openings with prominent anterior and posterior eye notches; eyes moderate and spindle-shaped; eye length 4.0��� 5.5 in head length and 3.0��� 4.8 times eye height. Eyes dorsolaterally on head, slightly above lateral midline. Subocular ledge broad and strong. Nictitating lower eyelids rudimentary, with shallow subocular pouches. Spiracles minute, length 3.8���7.6 in eye length, 0.2���0.3 eye lengths behind and below the posterior eye notch. First four gill openings longer than the fifth. Upper edge of each gill opening below eye and spiracle. Gill openings nearly straight, gill filaments not visible from outside. A few minute gill-raker papillae present on gill arches. Nostrils with broad, angular nasal flaps with rounded tips and moderate mesonarial flaps lateral on anterior nasal flaps, moderate excurrent apertures, no posterior nasal flaps. Nostrils nearly reaching level of mouth. Mouth broadly angular, short, its width 1.6���2.5 in head length. Mouth length 2.0��� 3.2 in mouth width. Lower symphysis nearly reaching upper symphysis. Maxillary teeth visible in ventral view (refer to Plate 1 for above characters). Tongue moderate, flat and rounded, filling most of floor of mouth. Many small buccal papillae on tongue and palate. Teeth in 20���23 / 23���26 rows; 3���5 / 4���6 series functional, with more series functional in larger specimens and in posterior tooth rows than anterolaterals. Posterolateral teeth not arranged in diagonal files. No toothless spaces at symphysis. Teeth not strongly differentiated within jaws, but tooth-row groups along jaws demonstrating weakly defined monognathic heterodonty including symphyseals (S), anterior (A), anterolaterals (AL), and posteriors (P) in both jaws. No apparent difference in dentition between genders. Anterolateral teeth (Figure 3) consist of single principal cusp with two pairs of lateral cusplets. Labial face of crown foot moderately concave, lingual face moderately convex. Principal cusp usually erect or showing slight distal inclination; this cusp mildly constricted at base at main crown, expanding slightly apically, tapering to an acute apex. Labial and lingual faces of cusp and cusplets mildly convex with short apicobasal ridges extending from crown foot to about 10 % or less of height of principal cusp and cusplets. Primary cusplets, those proximal to principal cusp, erect or curving slightly toward central cusp, primary cusplets approximately 50 % of height of central cusp; secondary cusplets, those distal to principal cusp, approximately 10���20 % of height of primary cusp. A thin, unserrated cutting edge present on medial and distal edges of primary cusp and lateral cusplets, beginning at crown foot and terminating near crown apices. In posterior teeth, relative height of principal cusp reduced, cusp becoming comparatively wider, and relative height of cusplets increasing but never exceeding height of principal cusp. Symphyseal teeth with relatively narrow and usually containing only one pair of lateral cusplets. Lower teeth similar overall to anterolateral teeth except primary cusplets 60���90 % length of principal cusp in anterolateral teeth. Body robust in head and trunk region, nearly circular in cross section at mid-trunk, laterally compressed and tapering posterior to anus. No predorsal, interdorsal, or postdorsal ridges on midline of back; no postanal ridge between anal-fin base and lower caudal-fin origin. Lateral ridges absent from body. Caudal peduncle moderately elongate, laterally compressed, without lateral keels. Caudal-peduncle height 1.7���2.4 its width at posterior insertion of second dorsal fin, 1.3���2.3 in dorsocaudal space. Dermal denticles, based on an examination of the holotype, vary widely in morphology depending on location on body (Plate 2). Dorsal dermal denticles, from dorsal surface of head and on trunk above lateral line, embedded in skin, spaced apart from (and not overlapping with) one another; these denticles blunt and single-cusped, widest at anterior base, often with two anterolateral lobes, and with 4���8 short, blunt longitudinal keels at base, often with keels extending posteriorly along most of length of denticles and becoming weaker near posterior apex. Lateral sides of the denticles usually weakly concave or occasionally straight; posterior apex acutely pointed, often rising well above skin giving a bristly appearance to dorsal surface of the head and body. Denticles longer than wide, those on dorsal trunk 2���3 times larger, more elongated, and with a more acute posterior apex than those on head. Dorsal dermal denticles, the only pigmented denticles on body, with pigment embedded in their anterior bases. Dermal denticles on lateral flanks colorless and semi-transparent, widely convex centrally and abruptly tapering to a thin, acute posterior apex; denticles longer than wide, with widest point at middle, usually smooth or showing a very weak central keel in middle, that keel absent at anterior base and posterior apex. Denticles are closely spaced and usually not overlapping, but sometimes barely so. Ventral dermal denticles colorless and semi-transparent, very thin without longitudinal keels; denticles wider than long, widest at base, quickly tapering to a blunt posterior apex; some denticles with a very short groove at medial base. Ventral dermal denticles closely-packed, usually with lateral edges barely overlapping neighboring denticles. Dorsal edge of caudal fin, from caudal pit to posterior fin edge and from lateral line to the dorsal edge, with clumped comb-like dermal denticles that together exhibit the file-like ���caudal crest��� characteristic seen in several other Bythaelurus and Parmaturus catshark species (Springer 1979). These denticles transparent, elevated well above skin, and usually overlap 10���50 % atop neighboring denticles. Each denticle consists of a single sharplypointed primary cusp with two (rarely one) large lateral secondary cusplets only nominally shorter than primary cusp; each cusp with a single longitudinal keel extending from anterior base to posterior apex. Denticles somewhat longer than wide, with widely convex lateral margins, the widest point at middle of denticle where lateral cusplets diverge from main body. Denticles on caudal fin below lateral line transparent, lanceolate and basally widest, tapering posteriorly, with straight or weakly convex lateral edges and a bluntly pointed posterior apex. These denticles flat and thin, without lateral cusplets and longitudinal keels, but exhibiting a short central grove at anterior base. Denticles on caudal fin below lateral line slightly elevated above skin and are closely spaced, but not touching or overlapping. In comparing denticle morphology and arrangement between holotype and paratypes, denticles vary little between largest and smallest specimens and between males and females. Relative size of dorsal denticles on head and body of smaller specimens longer, giving them more of a bristly appearance (Plate 1, Figure 4), an ontogenetic trait noted by Shouthall & Sims (2003) in the catshark Scyliorhinus canicula. Pectoral fins triangular, broad and rounded, not falcate, with broadly convex anterior margins, narrowly rounded apices, broadly rounded posterior margins, and free rear tips, inner margins and narrow bases. Pectoral-fin area subequal to area of first dorsal fin. Origins of pectoral fins beneath interspace between third and fourth gill openings. Apex of pectoral fin slightly anterior to its free rear tip when fin is elevated and appressed to body. Pelvic fins broadly triangular. Pelvic anterior margins 1.2���2.4 in pectoral fin anterior margins. Pelvic area subequal to anal-fin area. Pelvic-fin anterior margins slightly curved, apices narrowly rounded, posterior margins nearly straight, free rear tips narrowly rounded, inner margins straight and not fused together over claspers of juvenile males. First dorsal fin high, apically narrow and not falcate, with nearly straight anterior margin, narrowly rounded apex, slightly convex posterior margin, angular free rear tip, and straight inner margin. First dorsal-fin origin over middle of pelvic-fin base, insertion of fin just above a line behind insertion of pelvic fin. First dorsal-fin base 1.5��� 2.5 in interdorsal space, first dorsal-fin height 0.7���1.2 in first dorsal-fin base. Second dorsal fin high, apically narrow and not falcate, subequal to first dorsal-fin in area and about subequal to first dorsal-fin height and base. Second dorsal fin with nearly straight anterior margin, narrowly rounded apex, slightly convex posterior margin, angular free rear tip, and straight inner margin. Second dorsal-fin origin in a line above anal-fin midbase, its insertion above anal-fin free rear tip, and free rear tip in front of upper caudal-fin origin by the length of its base. Second dorsal-fin base 0.6���0.9 in dorsocaudal space, second dorsal-fin height 1.1���1.5 in second dorsal-fin base, second dorsal fin inner margin 1.5���2.6 in second dorsal-fin height and 1.9���2.9 in second dorsal-fin base. Anal fin low, apically broad, not falcate, about equal in size, height and base to that of second dorsal fin. Anterior margin slightly concave, apex broadly rounded, free rear tip bluntly pointed, and inner margin nearly straight. Anal-fin base without preanal ridges, anal-fin origin about 0.7���1.1 times anal-fin base length behind pelvic-fin insertions. Anal-fin posterior margin slanting posterodorsally, anal-fin insertion posterior to apex. Anal-fin base 0.8���1.4 in anal-caudal space, anal-fin height 1.5 ���2.0 in anal-fin base. PLATE 1. Dorsal and ventral views of three specimens of Bythaelurus giddingsi sp. nov. showing relative variation in the shape of the nasal flaps, the mouth, upper dentition, and dermal denticles on dorsum of head. Left: Holotype CAS 210091, adult female, 402 mm TL. Center: Paratype USNM 364283, immature female, 301 mm TL. Right: Paratype CAS 86559, immature male, 261 mm TL. Caudal fin narrow and asymmetrical, with large terminal lobe but ventral lobe not developed. Caudal fin short, dorsal margin 0.2���0.3 in precaudal length. Preventral caudal-fin margin 1.9���3.4 in dorsal caudal-fin margin, terminal lobe 3.0��� 3.7 in dorsal caudal-fin margin. Dorsal caudal-fin margin without lateral undulations but proximally and distally convex with a shallow concavity between the convexities. Preventral caudal-fin margin basally concave and apically straight, tip of ventral caudal-fin lobe bluntly rounded. Postventral margin not differentiated into upper and lower parts, its margin straight to concave. Subterminal notch a narrow slot, subterminal margin straight to concave and terminal margin straight and sometimes notched. Lobe formed by these margins slightly angular, tip of tail broadly rounded. Vertebral counts and statistics are given in Table 2. The precise number of caudal centra difficult to identify due to their small size and abundance of large caudal denticles; therefore counts of both the number of caudal centra and total number of centra are approximate. Transition between MP and DP centra about 4���6 centra behind front of pelvic girdle. Last few MP centra before MP-DP transition hardly enlarged, not forming a "stutter zone" of alternating long and short centra. Dimensions (in mm) of selected centra of holotype: length of penultimate MP 4.02; width of penultimate MP 3.02; midcentral width of penultimate centrum 0.94; length of first PD 2.75; width of first PD 3.08; midcentral width of first PD 1.14. "A ratio" (Springer & Garrick, 1964) of holotype 146. "B ratio" (Springer & Garrick, 1964) of holotype 133. Ratio of DP/MP of holotype 1.0. Color. In ethanol, chocolate brown dorsally, becoming pale brown on ventral surface from snout to anus. Body and flanks overlain with pale spots the largest of which about equal in size to eye, becoming smaller below lateral midline. A pale spot at anterior base of each dorsal fin. Pale spots extending onto caudal and median fins. Caudal region and all fins chocolate brown. The posterior margin of paired and dorsal fins pale. Tongue pale, anterior region of palate darker. Coloration in life can be seen in Figure 1. Size. Relative to other species of Bythaelurus, adult B. giddingsi are moderate in size. Our capture technique, using a suction device, was biased toward smaller individuals, and from the submersible we observed but did not capture specimens that seemed to be slightly larger. Two of our specimens (the 40.2 cm female and the 45.3 cm male) appear to be approaching sexual maturity. Our largest specimen (45.3 cm) is about midway in length between the small species of Bythaelurus ��� B. hispidus (to 29 cm), B. dawsoni (to 35 cm, a subadult male), and B. lutarius (to 39 cm) ��� and the large species, B. canescens (to 70 cm), and B. immaculatus (to 70 cm) (comparative lengths from Compagno, 1984). PLATE 2. Photos of dermal denticles from selected regions of the body of the Holotype of Bythaelurus giddingsi sp. nov., CAS 210091, each at 100 x magnification. Upper left: dorsal surface of head above eye; upp, Published as part of E, John, Long, Douglas J. & Baldwin, Carole C., 2012, Description of a new species of deepwater catshark, Bythaelurus giddingsi sp. nov., from the Gal��pagos Islands (Chondrichthyes: Carcharhiniformes: Scyliorhinidae), pp. 48-59 in Zootaxa 3221 on pages 49-58, DOI: 10.5281/zenodo.280250, {"references":["Compagno, L. J. V. (2005 b) Checklist of Living Chondrichthyes. In W. C. Hamlett (ed.) Reproductive Biology and Phylogeny of Chondrichthyes: Sharks, Batoids and Chimaeras. Reproductive Biology and Phylogeny, Vol. 3. Science Publishers, Inc, Enfield, USA, 503 - 548.","McCosker, J. E. & Rosenblatt, R. H. (2010) The fishes of the Galapagos Archipelago: an update. Proceedings of the California Academy of Sciences, Series 4, 61, Supplement II, No. 11, 167 - 195.","Ruiz, D., Chiriboga, A., & Banks, S. (2011) CDF Checklist of Galapagos Fish - FCD Lista de especies de Peces de Galapagos. In: F. Bungartz, H. Herrera, P. Jaramillo, N. Tirado, G. Jimenez-Uzcategui, D. Ruiz, A. Guezou, & F. Ziemmeck (eds.). Charles Darwin Foundation Galapagos Species Checklist - Lista de Especies de Galapagos de la Fundacion Charles Darwin. Charles Darwin Foundation / Fundacion Charles Darwin, Puerto Ayora, Galapagos, Ecuador.","Kyne, P. M., & Simpfendorfer, C. A. (2007) A collation and summarization of data on deepwater Chondrichthyans: biodiversity, life history, and fisheries. A report prepared by the IUCN SSC Shark Specialist Group for the Marine Conservation Biology Institute, Gland, Switzerland, 137 pp.","Hearn, A., Klimley, A. P. & Ketchum, J. (2009) Vulnerability of top predators to climate change and adaptation options for their coastal and pelagic ecosystems: sharks case study. In J. R. Hoffman, A. Franseca, & C. Drews (eds.) Cetaceans and Other Marine Biodiversity of the Eastern Tropical Pacific: Options for Adapting to Climate Change. International Whaling Commission Full Document 61 - 18, Agenda Item 11.1. 2, 50 - 54.","Springer, S. (1979) A revision of the catsharks, family Scyliorhinidae. NOAA Technical Report NMFS Circular 422, 152 pp.","Southall, E. J. & Sims, D. W. (2003) Shark skin: a function in feeding. Proceedings of the Royal Society of London, Supplemental Biology Letters, 270, S 47 - S 49.","Springer, V. G. & Garrick, J. A. F. (1964) A survey of vertebral numbers in sharks. Proceedings of the United States National Museum, 116,73 - 96.","Compagno, L. J. V. (1984) FAO Species Catalogue. Vol. 4, Sharks of the World. An annotated and illustrated catalogue of shark species known to date. FAO Fisheries Synopsis No. 125. vol. 4, pt. 2 (Carcharhinoformes). United Nations Development Programme / Food and Agriculture Organization of the United Nations, Rome, 251 - 655.","Springer, S. (1966) A review of western Atlantic cat sharks, Sycliorhinidae, with descriptions of a new genus and five new species. Fishery Bulletin, 65,581 - 624.","Bass, A. J., D'Aubrey, J. D. & Kistnasamy, N. (1975) Sharks of the east coast of southern Africa. II. The families Scyliorhinidae and Pseudotriakidae. Investigational Report of the Oceanographic Research Institute, 37, 1 - 64.","Nakaya, K. (1975) Taxonomy, comparative anatomy, and phylogeny of Japanese catsharks, Scyliorhinidae. Memoirs of the Faculty of Fisheries, Hokkaido University, 23, 1 - 94.","Long, D. J. (1994) Historical Biogeography of Sharks from the Northeastern Pacific Ocean. Ph. D. Diss., Department of Integrative Biology, University of California, Berkeley, 371 pp.","Chu, Y. - T., C. - W. Meng, C. - W., Hu, A. & Li, S. (1982) Five new species of elasmobranchiate fishes from the deep waters of South China Sea. Oceanologia et Limnologia Sinica, 13, 301 - 311.","Last, P. R., & Stevens, J. D. (2008) Bythaelurus incanus sp. nov., a new deepwater catshark (Carcharhiniformes: Scyliorhinidae) from northwestern Australia. In P. R. Last, W. T. White, & J. J. Pogonoski (eds.) Descriptions of New Australian Chondrichthyans. CSIRO Marine and Atmospheric Research Paper no. 0 22, Hobart, Australia, 123 - 128.","Gunther, A. (1878) Preliminary notices of deep-sea fishes collected during the voyage of H. M. S. ' Challenger. ' Annals and Magazine of Natural History (series 5), 17 - 28, 179 - 187, 248 - 251.","Alcock, A. W. (1891) Class Pisces. In II-Natural history notes from H. M. Indian marine survey steamer ' Investigator, ' Commander R. F. Hoskyn, R. N. commanding. - Series II., No. 1. On the results of deep-sea dredging during the season 1890 - 91. Annals and Magazine Natural History, (series 6), 16 - 34, 119 - 138.","Springer, S. (1971) A new cat shark (Scyliorhinidae) from New Zealand. Records of the Dominion Museum (Wellington), 7, 235 - 241.","Garman, S. (1913) The Plagiostomia (sharks, skates, and rays). Memoires of the Museum of Comparative Zoology of Harvard College, 515 pp."]}

Apogon pillionatus B��hlke and Randall Identification. Adult Apogon pillionatus (Fig. 5) is diagnosed by the combination of eight segmented anal-fin rays, the body and lateral-line scales of similar size, a dark bar just behind the second dorsal fin that does not reach the ventral midline, and a very broad bar on the posterior part of the caudal peduncle (also does not reach the ventral midline). The distance between the two body bars is considerably less than the width of the posterior bar (B��hlke & Chaplin 1993; Gon 2002). No adult specimens of A. pillionatus were collected in this study, and no COl sequences for the species were found in GenBank. The specimen featured in Figure 5 was collected on Saba Bank Atoll prior to our study and is not a DNA voucher specimen. Juveniles identified in this study as A. pillionatus (Appendix 1) have a relatively narrower bar of dark pigment on the posterior part of caudal peduncle than adult A. pillionatus (see ���Juveniles,��� below), and the anterior dark bar is situated behind the posterior base of the second dorsal fin. The identification of those juveniles as A. pillionatus was accomplished by process of elimination and comparative morphological examination. Six western Atlantic Apogon species, Apogon planifrons, A. phenax, A. robinsi, A. townsendi, A. gouldi, and A. pillionatus, have two dark bars on the posterior part of the body: one in the area of the posterior portion of the second dorsal fin and the other on the caudal peduncle (B��hlke & Randall, 1993). In A. townsendi, A. planifrons, and A. gouldi the anterior bar is situated entirely beneath the second dorsal-fin base. Apogon phenax has a wedge-shaped bar situated below and just behind the second dorsal-fin base. Only two Apogon species have the anterior bar well behind the end of the second dorsal fin: A. robinsi and A. pillionatus. Apogon robinsi is easily recognized by the lateral extensions of the premaxillary tooth patches (see ���Identification��� under ��� A. robinsi ���), such that a portion of the dentition lies outside the confines of the mouth (B��hlke & Randall, 1968; B��hlke & Chaplin 1993; Gon 2002). Furthermore, the anterior body bar in A. robinsi extends to the ventral midline, whereas in A. pillionatus it falls short of the ventral midline. The juveniles identified herein as A. pillionatus (Fig. 1) have the anterior bar well behind the second dorsal-fin base, that bar terminating well short of the ventral midline, and no dentition outside of the mouth. Juveniles (Fig. 6). Five juveniles identified as described above range from 15.0 to 17.0 mm SL. The body is pale orange with darker orange coloration on the head and anterior rays of the first dorsal fin. There are melanophores on top of head, behind the eye on the cheek and temporal regions, and on the gut. There are melanophores on the posterior part of the second dorsal- and anal-fin bases, as well as on the distal tips of the middle rays of the first dorsal and anal fins. There are numerous melanophores on the outer rays of the caudal fin. The anterior bar of the body is entirely behind the posterior end of the second dorsal-fin base and does not reach the ventral midline of the body. The bar on the caudal peduncle is broad. As noted above, this bar is narrower in juveniles than in adults, and the space between the anterior and posterior body bars in juveniles is equal to or greater than the width of the posterior bar (this space smaller in adults). Comparisons Among Juveniles. Characters used to separate juvenile A. pillionatus from other Apogon species having two body bars are discussed above (see ���Identification���)., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on pages 8-9, DOI: 10.5281/zenodo.279426, {"references":["Bohlke, J. E. & Chaplin, C. C. G. (1993) Fishes of the Bahamas and Adjacent Tropical Waters. Second Edition. University of Texas Press, Austin, 771 pp.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.) The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391.","Bohlke, J. E. & Randall, J. E. (1968) A key to the shallow-water Atlantic cardinalfishes (Apogonidae), with descriptions of five new species. Proceedings of the Academy of Natural Sciences of Philadelphia, 120, 175 - 206."]}

Apogon affinis (Poey) Identification. Adult A. affinis can be distinguished from other Apogo n species by the combination of cycloid to weakly ctenoid scales, median predorsal scales present, pectoral-fin soft rays 11 or 12 (rarely 13), nine segmented anal-fin rays, and both jaws with a single series of small conical teeth interspersed with several enlarged caniniform teeth (B��hlke & Chaplin 1993, Gon 2002). Two adult specimens identified as A. affinis based on those features were analyzed genetically (Appendix 1). The specimens were taken in trawls and are not in good shape, and we selected a photograph of an adult not analyzed genetically in this study to illustrate the species (Fig. 20). No larvae or juveniles analyzed in this study genetically match A. affinis., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on page 19, DOI: 10.5281/zenodo.279426, {"references":["Bohlke, J. E. & Chaplin, C. C. G. (1993) Fishes of the Bahamas and Adjacent Tropical Waters. Second Edition. University of Texas Press, Austin, 771 pp.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.) The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391."]}

Apogon quadrisquamatus Longley Lineage A Identification. A single adult specimen from Belize (Fig. 28) constitutes the genetic lineage herein referred to as Apogon quadrisquamatus Lineage A. Adult A. quadrisquamatus can be distinguished from other Apogon by the combination of eight segmented anal-fin rays; lateral-line and body scales of similar size; no dark marking or bar beneath the second dorsal fin; caudal-peduncle spot small, circular, of varying intensity, and usually restricted to middle of caudal peduncle; and 12���14, modally 13, gill rakers on the lower limb of the first arch, (B��hlke & Chaplin 1993, Dale 1977, Gon 2002). The adult specimen in this lineage keys to Apogon quadrisquamatus and is distinct from A. mosavi and Apogon sp. 2 in having a circular blotch of melanophores in the center of the peduncle vs. a rectangular bar (A. mosavi) or very diffuse oval (Apogon sp. 2) of melanophores. Apogon robbyi has a similar circular, basicaudal blotch, but that species is distinctive in having dusky stripes on the trunk. Apogon quadrisquamatus Lineage A is further distinguished from A. mosavi in having 12 gill rakers on the lower limb of the first arch (possibly a rudiment is forming), vs. 14���15 in A. mosavi. The body is mostly orange, and the median fins are yellow. Additional material and further study are needed to determine if this lineage and A. quadrisquamatus Lineage B (see next section) are morphologically distinct, and, if so, which one represents A. quadrisquamatus Longley 1934. No larvae or juveniles match the single adult specimen of this lineage in our genetic analysis., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on page 23, DOI: 10.5281/zenodo.279426, {"references":["Bohlke, J. E. & Chaplin, C. C. G. (1993) Fishes of the Bahamas and Adjacent Tropical Waters. Second Edition. University of Texas Press, Austin, 771 pp.","Dale, G. (1977) Apogon mosavi, a new Western Atlantic cardinalfish, and a note on the occurrence of Apogon leptocaulus in the Bahamas. Proceedings of the Biological Society of Washington, 90, 19 - 29.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.) The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391.","Longley, W. H. (1934) Studies on West Indian fishes: description of six new species. Carnegie Institution of Washington Year Book, 33, 257 - 260."]}

Apogon binotatus (Poey) Identification. Eleven adult specimens of A. binotatus provided the basis for genetic identification of larvae and juveniles (Appendix 1, one adult is shown in Fig. 2). Adult A. binotatus can be distinguished from other Apogon by the combination of eight segmented anal-fin rays, body and lateral-line scales of similar size, and body with two distinct dark markings posteriorly ��� a bar below the second dorsal fin and another on the posterior part of the caudal peduncle, both bars narrow, much deeper than wide (B��hlke & Chaplin 1993, Gon 2002). Table 1. Average Kimura 2 -parameter distance summary for Apogon species. Intraspecific averages in bold font. na = not applicable (one specimen). aurolineatus 0.01 = 12) maculatus 0.20 0.01 N = 18) phenax 0.19 0.13 0.00 = 32) binotatus 0.21 0.14 0.10 0.00 = 28) townsendi 0.24 0.15 0.10 0.13 0.00 = 27) pseudomaculatus 0.22 0.12 0.10 0.14 0.14 0.00 = 3) pillionatus 0.23 0.14 0.10 0.12 0.15 0.16 0.01 = 5) lachneri 0.23 0.16 0.10 0.12 0.13 0.14 0.14 0.00 = 11) planifrons 0.22 0.18 0.10 0.10 0.17 0.17 0.11 0.15 0.00 = 6) mosavi 0.23 0.20 0.20 0.22 0.21 0.21 0.24 0.20 0.244 0.00 = 36) robbyi 0.23 0.20 0.20 0.23 0.20 0.19 0.22 0.21 0.22 0.05 0.00 = 3) quadrisquamatus 0.25 0.22 0.20 0.24 0.22 0.18 0.23 0.20 0.24 0.09 0.10 0.00 Lineage A = 1) Apogon sp 0.23 0.14 0.10 0.12 0.14 0.11 0.11 0.14 0.08 0.23 0.20 0.23 na = 1) affinis 0.24 0.14 0.10 0.17 0.15 0.14 0.16 0.16 0.18 0.18 0.20 0.21 0.16 0.00 = 2) robinsi 0.21 0.17 0.20 0.17 0.18 0.18 0.17 0.14 0.18 0.19 0.20 0.20 0.17 0.20 na = 1) Apogon sp. 2 0.24 0.21 0.20 0.24 0.21 0.18 0.23 0.22 0.22 0.07 0.10 0.09 0.23 0.20 0.21 0.00 = 5) quadrisquamatus 0.23 0.17 0.20 0.20 0.19 0.17 0.20 0.22 0.21 0.12 0.10 0.14 0.20 0.20 0.18 0.09 0.00 Lineage B = 5) Juveniles (Fig. 3). Juveniles identified genetically range from 13.5 to 18.0 mm SL. The body is pale. The upper part of the head and the gut are pink. The fins are mostly clear, but there are melanophores on the tips of the anterior rays of the second dorsal and anal fins and on the outer rays of the caudal fin. There are numerous melanophores on top of the head and over the gut. There is a slender bar beneath the end of second dorsal-fin base and a wider bar on the caudal peduncle. There are 12���13 gill rakers on the lower limb of the first gill arch, counts consistent with values for adults (Table 2). Comparisons Among Juveniles. Although adult A. binotatus are easily distinguished from other Apogon species by having both body bars slender (much deeper than wide), the posterior bar in juvenile A. binotatus is broader than the anterior bar. It is still narrow relative to the broad posterior bar in A. townsendi, A. phenax, and A. pillionatus. Juvenile A. binotatus is most similar to juvenile A. phenax, but it can be separated from that species by having the anterior bar slender (vs. at least slightly wedge-shaped in A. phenax) and positioned entirely beneath the second dorsal fin (vs. half or more of the bar behind the fin in A. phenax). The configuration of the two bars in juveniles of A. binotatus is sufficient to separate them from known juveniles of other Apogon. Apogon 14.0 mm SL, DNA # BLZ J. Mounts C. Larvae (Fig. 4). Apogon binotatus larvae analyzed genetically range from 8.5 to 11.0 mm SL. The body is orange to pale orange, and there may be clear (or paler orange) areas on the snout just anterior to the eye, on the caudal peduncle, and near the middle of the trunk. In one 9.0���mm SL specimen there is a wide pale area posterior to the second dorsal fin and anterior to the caudal-fin base. The fins are clear except for a few orange spots along the posterior base of the anal fin and a few spots in the center of the caudal-fin base. There is a line of bright orange pigment on ventral side of the body from the anal fin to the caudal peduncle. There are melanophores on top of the head, in the temporal region, and over the swimbladder. Some specimens have a few melanophores on the lateral surface of the gut. The caudal-peduncle length ranges from 33���37 % SL. Comparisons Among Larvae. Fresh specimens of A. binotatus larvae are extremely similar to A. phenax larvae, but they can often be separated by snout pigment (no or pale orange spot anteriorly in A. binotatus vs. usually a prominent orange spot on snout in A. phenax). From A. planifrons, larval A. binotatus differs in lacking yellow pigment on the anterior portion of the body; from A. maculatus in lacking prominent orange pigment on the first dorsal fin; from A. aurolineatus larvae in lacking orange/yellow dorsal, anal, and pelvic fins; and from A. mosavi larvae in lacking a distinctive pattern of chromatophores on the median fins. We were not able to reliably separate Apogon binotatus larvae from those of A. townsendi and Apogon sp. 1, although our A. binotatus larvae have more orange color on the body than A. townsendi larvae. However, the extent of orange coloration on the body of some Apogon larvae, and whether or not it is disrupted by pale areas, may change ontogenetically (see ��� Apogon phenax,��� below). Preserved larval specimens of A. binotatus have more melanophores on top of the head than larval A. aurolineatus and a longer caudal peduncle (caudal-peduncle length 33���37 % SL in A. binotatus vs. 27���29 % SL in A. aurolineatus). Caudal-peduncle length also is sometimes useful in separating preserved A. binotatus from preserved A. planifrons (caudal-peduncle length 33���37 % SL in A. binotatus, 35���40 % SL in A. planifrons), and it may be useful in separating larval A. binotatus from A. maculatus (caudal-peduncle length 30 % SL in the single larval specimen of A. maculatus in our study material). We have identified no other features to separate preserved A. binotatus larvae from other known Apogon larvae., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on pages 3-8, DOI: 10.5281/zenodo.279426, {"references":["Bohlke, J. E. & Chaplin, C. C. G. (1993) Fishes of the Bahamas and Adjacent Tropical Waters. Second Edition. University of Texas Press, Austin, 771 pp.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.) The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391."]}

Apogon sp. 1 Identification. No adults match the single larval specimen of this lineage in our genetic analysis. This unidentified species could be A. evermanni, A. leptocaulus, or A. gouldi, and specimens of those species are needed for comparative genetic analysis. Alternatively, the unidentified larval specimen may represent an undescribed species. Larvae (Fig. 7). Apogon sp. 1 is represented by one 10.0 mm SL specimen. The body is mostly orange with some transparent areas. The snout is mostly transparent, but there is a small, pale orange spot at the anterior tip. The jaws are pale orange. The rest of the head and anterior portion of the trunk are orange, the area immediately behind the eye the most intense orange. The posterior region of the body is mostly transparent, with one band of pale orange pigment below the second dorsal fin and one spot of bright orange pigment on caudal peduncle. There is a line of bright orange pigment on the ventral midline of the body from the anal fin to the caudal peduncle. All fins are clear except the proximal portions of the anal-fin rays have pale orange coloration. There are melanophores on top of the head, behind the eye on the temporal region, over the swimbladder, and on the lateral surface of the gut. The caudal-peduncle length is 35 % SL. Comparisons Among Larvae. Larval Apogon sp. 1 most closely resemble A. binotatus, A. phenax, A. planifrons, and A. townsendi larvae in having orange pigmentation concentrated mainly on the head and anterior portion of the trunk and transparent areas on the snout and usually also on posterior portions of the trunk. Apogon sp. 1 differs from A. planifrons in lacking yellow pigment on the anterior portion of the body and from larval A. phenax in lacking a prominent orange spot on the snout. From A. maculatus, larval Apogon sp. 1 differs in lacking orange pigment on the first dorsal fin and in having a longer caudal peduncle (35 % SL in Apogon sp. 1, 30% SL in A. maculatus). From A. aurolineatus, larval Apogon sp. 1 differs in lacking orange/yellow pigment on the dorsal, pelvic, and anal fins; in having transparent areas on the trunk (vs. trunk completely orange in A. aurolineatus); and in having a longer caudal peduncle (35 % SL in Apogon sp. 1 vs. 27–29 % SL in A. aurolineatus). Apogon sp. 1 lacks the distinctive pattern of chromatophores on the median fins typical of A. mosavi and the single specimen examined has a longer caudal peduncle (35 % SL in Apogon sp. 1 vs. 31–34 % SL in A. mosavi). Additional larval specimens of Apogon sp. 1 are needed to confirm the differences noted above and to determine if it can be separated from larval A. binotatus and A. townsendi.

Apogon phenax B��hlke and Randall Identification. Sixteen adult specimens of A. phenax provided the basis for genetic identification of larvae and juveniles (Appendix 1, one adult is shown in Fig. 10). Adult A. phenax can be distinguished from other Apogon by the combination of eight segmented anal-fin rays, body and lateral-line scales of similar size, body with two distinct dark markings (one wedge-shaped bar below and just behind second dorsal fin and a bar on the posterior part of caudal peduncle���the distance between the two bars larger than the width of the posterior bar), and 11 to 14 (usually 13���14) gill rakers (B��hlke & Chaplin 1993, Gon 2002). Juveniles (Fig. 11). Eight juveniles, 16.0���22.0 mm SL, are present in our material. The body is pale orange. The head, abdomen, first dorsal fin, bases of second dorsal and anal fins, and posterior portion of the caudal peduncle are darker pink/orange. There are melanophores on the head, gut, outer rays of the caudal fin and on the distal portions of the second dorsal and anal fins. Two dark bars are present on the trunk in all juveniles. The anterior bar is somewhat wedge shaped (slightly broader dorsally than ventrally) and extends ventrally to a point slightly below mid body or slightly above the anal-fin base. This bar is confluent with a stripe of melanophores along the second dorsal-fin base and separated by a gap from a similar stripe along the anal-fin base. The bar on the caudal peduncle extends from the dorsal to the ventral margins of the body. As in adults, the distance between the two bars is greater than the width of the posterior bar. There are 13���14 lower-limb gill rakers on the first arch in the juveniles, which is consistent with values for adults (Table 2). Comparisons Among Juveniles. Juvenile A. phenax can be distinguished from other juvenile Apogon by the same characters that separate adults. Larvae (Fig. 12). Apogon phenax larvae genetically analyzed in this study range from 9.5 to 11.0 mm SL. The smaller specimens are bright orange in life, and the fins are clear (Fig. 12 a). Much of the snout is transparent, but there is a conspicuous orange spot above the upper lip. The upper and lower jaws have scattered orange chromatophores. A nearly solid line of orange pigment extends along the base of the anal fin and ventral midline of the caudal peduncle. A 10.0���mm SL specimen (Fig. 12 b) is paler orange in general, has more pale areas on the snout and jaws, and has a large pale area on the dorsal portion of the caudal peduncle. There are a few orange chromatophores at the bases of the central and ventral rays of the caudal fin. In one 11.0 mm SL specimen (Fig. 12 c), the dark bars characteristic of juveniles and adults are beginning to form posteriorly, and the area between the bars is pale. In all larvae there are melanophores on top of the head, in the temporal region, and over the swim bladder and gut. The caudal-peduncle length ranges from 32���36 % SL. Comparisons Among Larvae. The smallest larvae of A. phenax resemble larvae of A. aurolineatus and A. maculatus in usually having a bright orange body color, but they differ in lacking orange/yellow (A. aurolineatus) or orange (A. maculatus) pigment on the first dorsal fin; A. aurolineatus also has orange second dorsal, anal, and pelvic fins, which are clear in A. phenax larvae. Additionally, larvae of A. phenax have more melanophores on top of the head than A. aurolineatus and a longer caudal peduncle than that species and A. maculatus (caudal-peduncle length 32 % SL or larger in A. phenax, 27���29 % SL in A. aurolineatus, 30 % in A. maculatus). Larger Apogon phenax larvae are very similar to those of A. binotatus, and somewhat similar to those of A. townsendi, but they differ from both in having a conspicuous orange spot above the upper lip (vs. small and pale, if present). Larval A. phenax can be separated from larval A. planifrons by the absence of bright yellow pigment on the head and from larval A. mosavi by the absence of a distinctive pattern of chromatophores on the median fins. We have identified no morphological features to separate preserved A. phenax, A. binotatus, A. townsendi, and A. planifrons larvae. Apogon lachneri B��hlke Identification. Four adult specimens of A.lachneri provided the basis for genetic identification of seven juveniles (Appendix 1; one adult is shown in Fig. 13). The combination of characters that distinguishes A. lachneri adults from other Apogon species is eight segmented anal-fin rays; 16���17 gill rakers on the lower limb of the first arch; lateral-line and body scales of similar size; a small dark saddle behind the second dorsal fin, followed by a white spot (white spot may not be apparent in preserved specimens); a large dark area on the first dorsal fin posterior to the second spine; and anterior portions of the second dorsal and anal fins dark to dusky distally (B��hlke & Chaplin 1993, Gon 2002). Juveniles (Fig. 14). Juveniles examined range from 18.0���22.0 mm SL. In all specimens, the body is pale orange, and the dark pattern of pigment on the dorsal and anal fins typical of adults is conspicuous. The caudal-fin rays are densely covered with melanophores. There are scattered melanophores on the head in the smallest juveniles. The posterior margins of the scales on the dorsal portion of the trunk and caudal peduncle are covered with melanophores, forming roughly diamond-shaped patterns of pigment on the body. The dark saddle behind the second dorsal fin characteristic of adults is beginning to develop or fully present in all specimens. There are 16 gill rakers on the lower limb of the first gill arch. Comparisons Among Juveniles. Juvenile A. lachneri can be distinguished from all Apogon juveniles by the conspicuous dark pigment on the dorsal and anal fins. It resembles A. aurolineatus in lacking dark markings on the caudal peduncle, but it is easily separated from that species by the dark saddle behind the second dorsal fin (vs. no distinguishing marks on the body in juvenile A. aurolineatus). Apogon lachneri juveniles most closely resemble those of A. maculatus and A. pseudomaculatus in having a dark spot of pigment associated with the base of the second dorsal fin, but the position of the spot distinguishes them (behind the last ray of the second dorsal fin in A. lachneri; on the posterior base of that fin in A. maculatus, and well below the posterior base of the second dorsal fin in A. pseudomaculatus). Juvenile A. lachneri can further be distinguished from A.maculatus and A. pseudomaculatus by lacking a dark blotch of pigment on the caudal peduncle., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on pages 12-15, DOI: 10.5281/zenodo.279426, {"references":["Bohlke, J. E. & Chaplin, C. C. G. (1993) Fishes of the Bahamas and Adjacent Tropical Waters. Second Edition. University of Texas Press, Austin, 771 pp.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.) The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391."]}

Apogon mosavi Dale Identification. Seventeen adult specimens of A. mosavi provided the basis for genetic identification of larvae and juveniles (Appendix 1, one adult is shown in Fig. 34). Adult A. mosavi can be distinguished from other Apogo n species by the combination of eight segmented anal-fin rays, body and lateral-line scales of similar size, no dark marking or bar beneath the second dorsal fin, a rectangular to oval bar present on the caudal peduncle that nearly reaches the dorsal and ventral body margins, and 14���15 (rarely 13 or 16) gill rakers on the lower limb of the first gill arch, (Dale 1977, Gon 2002). Our samples include 20 young specimens of A. mosavi, seven of them (14.0���17.0 mm SL) collected in the plankton net and 13 (14.0���20.0 mm SL) with fish anesthetics and dip nets (Appendix 1). As noted in the ���Methods��� section, we describe those collected in the plankton net as larvae and the others as juveniles. However, until melanophores appear on the caudal peduncle in large juveniles, there are no clear morphological differences between the two stages. Juveniles (Fig. 35). In the 15.0 to 20.0 mm SL juvenile specimens of A. mosavi (Fig. 35 a) the body is pale orange and there are no distinctive symmetrical markings on the fins. There is a large blotch of pigment on the caudal peduncle comprising orange chromatophores with sometimes a few melanophores mixed in. There are 14 or 15 gill rakers on the lower limb of the first arch. Fresh specimens of small juveniles (14.0���17.0 mm SL, Fig. 35 b) are mostly pale orange, with paler areas on the snout and jaws, beneath the anterior portion of the second dorsal fin, and on the anterior portion of the caudal peduncle. The dorsal, caudal, anal, and pelvic fins have distinctive blotches of orange pigment. There is a large orange blotch covering the entire first dorsal fin except the bases of the rays. Three orange blotches are present on the second dorsal fin ��� two on the anterior portion of the fin and the third on the posterior base of the fin. There are six orange blotches on the caudal fin: four on the outer caudal-fin rays, two dorsally and two ventrally, and two on the caudal-fin base. Three orange are present blotches on the anal fin ��� two on the anterior portion of the fin and the third on the posterior base of the fin. There is one orange blotch on the pelvic fin. There is symmetry in the position of the orange fin blotches such that those on the dorsal fins and dorsal lobe of the caudal fin mirror those on the pelvic fin, anal fin, and ventral lobe of the caudal fin. There are melanophores on top of the head and internally above the gut. Comparisons Among Juveniles. Young juveniles of A. mosavi most closely resemble those of A. quadrisquamatus and A. robbyi in having distinctive patterns of chromatophores on the fins. See ���Comparisons��� under A. quadrisquamatus and A. robbyi juveniles for characters that distinguish them. Larvae (Fig. 36). The seven larvae (14.0���17.0 mm SL) collected in the plankton have the same patterns of chromatophores and melanophores as those described above for small juveniles. The caudal-peduncle length ranges from 31���34 % SL. Comparisons Among Larvae. Adult A. mosavi are most similar to A. quadrisquamatus and Apogon sp. 2. Larvae of A. quadrisquamatus are unknown, but juvenile A. quadrisquamatus and larval and juvenile A. mosavi have a similar pattern of fin pigment, suggesting that the larvae of A. quadrisquamatus may as well. Assuming the color of fin pigment of larval A. quadrisquamatus is the same as it is in juveniles, as is the case in A. mosavi, larval A. mosavi will differ from larval A. quadrisquamatus in having orange pigment on the fins (vs. orange and yellow). Likewise, larvae of A. robbyi are unknown, but they should differ from A. mosavi larvae in having yellow vs. orange fin pigment. The pattern of fin pigment in larval A. mosavi is sufficient to separate that species from larvae of other known Apogon. We know of no morphological features of preserved A. mosavi larvae that separate them from other Apogon larvae except the numerous melanophores on top of the head and long caudal peduncle (31��� 34 % SL) will distinguish them from larval A. aurolineatus (few or no melanlophores on top of the head and caudalpeduncle length 27���29 % SL)., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on page 27, DOI: 10.5281/zenodo.279426, {"references":["Dale, G. (1977) Apogon mosavi, a new Western Atlantic cardinalfish, and a note on the occurrence of Apogon leptocaulus in the Bahamas. Proceedings of the Biological Society of Washington, 90, 19 - 29.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.) The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391."]}

Apogon planifrons Longley and Hildebrand Identification. One adult specimen of A. planifrons (Fig. 8) provided the basis for genetic identification of five larva (Appendix 1). No juveniles are present in our material. Adult A. planifrons has eight segmented anal-fin rays, the body and lateral-line scales of similar size, the anterior dark bar positioned entirely below the posterior end of the second dorsal fin and distinctly narrower than the dark bar on the caudal peduncle, 15 or 16 circum-caudalpeduncle scales, and usually 15 (14���16) gill rakers on the lower limb of the first arch (B��hlke & Chaplin 1993, B��hlke & Randall 1968, Gon 2002). Apogon planifrons most closely resembles A. townsendi and A. gouldi in having the anterior bar entirely beneath the second dorsal-fin base. It can be separated from those species by lowerlimb rakers (Table 2) and circum-caudal-peduncle scales (15���16 in A. planifrons vs. 12 in A. townsendi and A. gouldi). Apogon planifrons can be further distinguished from A. townsendi in lacking black lateral margins on the caudal-peduncular bar (B��hlke & Chaplin 1993, Smith-Vaniz 1977, Gon 2002). Larvae (Fig. 9). Apogon planifrons larvae genetically analyzed in this study range from 9.0 to 10.0 mm SL. The snout is mostly transparent, and there is pale orange coloration on the jaws. The central and posterior portions of the head and the belly have prominent yellow pigmentation. The posterior region of the body is mostly pale orange to orange in 9.0 and 9.5mm SL specimens, with some pale areas on the caudal peduncle. In 10 ���mm SL specimens, much of the posterior portion of the body is pale, usually with an orange bar beneath the posterior end of the second dorsal fin and an orange blotch on the posterior end of the caudal peduncle. The dorsal fins are clear. There are orange chromatophores at the bases of the pelvic and anal fins, as well as on the proximal portion of the ventral lobe of the caudal fin. There is a line of orange chromatophores along the anal-fin base that extends onto the ventral midline of the caudal peduncle. There are melanophores on top of the head, behind eye in the temporal region, over the swimbladder, and on the lateral surface of the gut. The caudal peduncle is long, 35���40 % SL. Comparisons Among Larvae. Apogon planifrons larvae are easily distinguished from other known Apogon larvae by the bright yellow chromatophores on most of the head and abdominal region and usually by a long caudal peduncle (35���40 % SL in A. planifrons vs. 27���37 % SL in other Apogon species). Caudal-peduncle length alone is useful for separating preserved larval specimens of A. planifrons from larvae of some Apogon species, but A. phenax, A. binotatus, and A. townsendi also have a long caudal peduncle (32���37 % SL). We know of no features that distinguish preserved larvae of A. planifrons from those species., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on page 11, DOI: 10.5281/zenodo.279426, {"references":["Bohlke, J. E. & Chaplin, C. C. G. (1993) Fishes of the Bahamas and Adjacent Tropical Waters. Second Edition. University of Texas Press, Austin, 771 pp.","Bohlke, J. E. & Randall, J. E. (1968) A key to the shallow-water Atlantic cardinalfishes (Apogonidae), with descriptions of five new species. Proceedings of the Academy of Natural Sciences of Philadelphia, 120, 175 - 206.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.) The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391.","Smith-Vaniz, W. F. (1977) Apogon gouldi n. sp., a new cardinalfish from Bermuda (Perciformes: Apogonidae). Notulae Naturae, 452, 1 - 8."]}

Apogon aurolineatus (Mowbray) Identification. Four adult specimens of A. aurolineatus provided the basis for genetic identification of seven larvae and one juvenile (Appendix 1, one adult is shown in Fig. 24). Adult A. aurolineatus can be distinguished from other Apogon by the combination of eight segmented anal-fin rays, 10���11 gill rakers on the lower limb of the first gill arch, 16���18 circum-caudal-peduncle scales, no dark markings or saddles on the posterior portion of the body, and two to four short dark lines radiating from the eye (B��hlke & Chaplin 1993; Gon 2002). Juveniles (Fig. 25). The single juvenile, a reared specimen of 12 mm SL, has a pale salmon body color, and the opercular and abdominal regions are silvery. In preservative, there are no distinctive markings except a few melanophores on top of the head. There are 11 gill rakers on the lower limb of the first gill arch. Comparisons Among Juveniles. Of the Apogon species for which juveniles are known, A. aurolineatus most closely resembles A. quadrisquamatus in lacking dark blotches or markings on the body. The reared juvenile of A. aurolineatus can be separated from juvenile A. quadrisquamatus in having a pale body color (vs. orange in A. quadrisquamatus), in lacking yellow on the dorsal and caudal fins, and in lacking an orange spot on the center of the caudal peduncle. Preserved juveniles of the two species are very similar, but eye diameter may be useful in separating the species (diameter of bony orbit approximately 13 % SL in the 12.0-mm SL juvenile of A. aurolineatus vs. 15 % SL in 14.0���16.0-mm SL juveniles of A. quadrisquamatus). Preserved juveniles of A. aurolineatus also are similar to those of A. robbyi and A. mosavi in having a pale body, but A. aurolineatus lacks the blotch of melanophores on the caudal peduncle present in those species. Larvae (Fig. 26). Apogon aurolineatus larvae genetically analyzed in this study are all approximately 8 mm SL. They are bright orange in life and have orange pelvic, anal, and second dorsal fins. The first dorsal fin is orange at the base, but most of the fin is bright yellow. The pectoral and caudal fins are clear. There are some pale areas on the head���below the anterior portion of the eye and above the tip of the snout. The top of the head has yellow pigment in some specimens. There are no dark markings on the body except sometimes a few melanophores on the top of the head. There are barely observable melanophores scattered on the jaws. The caudal-peduncle length ranges from 27 to 29 % SL. Comparisons Among Larvae. Fresh specimens of A. aurolineatus larvae are easily distinguished from other known Apogon larvae by the combination of bright orange body coloration and yellow pigment on the first dorsal fin. Preserved specimens usually have fewer melanophores on top of the head than larvae of other Apogon (zero to several vs. many) and a shorter caudal peduncle (27���29 % SL vs. 30���40 % SL in other species). Apogon robinsi B��hlke and Randall Identification. One adult specimen of A. robinsi was collected and analyzed genetically (Appendix 1). Because the photograph of that specimen is not of good quality, we selected a photograph of a specimen not included in the genetic analysis to represent the species (Fig. 27). Adult A. robinsi can be distinguished from other Apogo n species by the combination of eight segmented anal-fin rays, body and lateral-line scales of similar size, body with two distinct dark markings (one bar below and just behind second-dorsal fin and a bar on the posterior part of caudal peduncle���the distance between the two bars larger than the width of the posterior bar), and premaxillary dentition extending outside the mouth laterally on the bone (B��hlke & Chaplin 1993, Gon 2002). No larvae or juveniles analyzed in this study genetically match A. robinsi., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on pages 21-23, DOI: 10.5281/zenodo.279426, {"references":["Bohlke, J. E. & Chaplin, C. C. G. (1993) Fishes of the Bahamas and Adjacent Tropical Waters. Second Edition. University of Texas Press, Austin, 771 pp.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.) The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391."]}

Apogon sp. 2 Identification. No larvae or juveniles match the five unidentified adult specimens of this lineage in our genetic analysis (Appendix 1, one adult is shown in Fig. 31). The lineage clusters phenetically with A. quadrisquamatus, A. mosavi, and A. robbyi (Fig. 1), and all of those species lack dark markings on the body. There is a concentration of pigment on the central area of the caudal peduncle that is primarily orange and contains few if any melanophores. In preserved specimens this may appear as a diffuse oval blotch of melanophores or no marking at all. In the other species, there is a distinct basicaudal bar (A. mosavi) or circular spot (A. quadrisquamatus and A. robbyi) of melanophores that is retained in preserved specimens. Specimens in this lineage usually have 13 gill rakers on the lower limb of the first arch vs. usually 14 or 15 in A. mosavi, but additional material is needed to determine if there are modal differences in any counts. This lineage likely represents an undescribed species., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on page 25, DOI: 10.5281/zenodo.279426

Apogon pseudomaculatus Longley Identification. One wild-caught adult specimen of A. pseudomaculatus from Florida provided the basis for genetic identification of one juvenile reared from a wild-caught larva from Belize and one juvenile specimen from Cura��ao (Appendix 1). An adult collected off Cura��ao but not yet analyzed genetically is shown in Figure 18. The combination of characters that distinguishes A. pseudomaculatus adults from other Apogon species is the presence of eight segmented anal-fin rays, body and lateral-line scales of similar size, dark pupil-size spot below posterior end of second dorsal fin, 14���16 circum-caudal-peduncle scales, and a dark pupil-size spot on the caudal peduncle (B��hlke & Chaplin 1993; Gon 2002). A color image of FWRI 20646, the genetically analyzed adult (quality of specimen and image too poor to reproduce here), shows the dark spot beneath the second dorsal fin and another on the caudal peduncle. The spot beneath the second dorsal fin is well below the base of the fin, a diagnostic feature of A. pseudomaculatus. However, there is black pigment on the dorsal, caudal, and anal fins on the FWRI specimen that is not present in the adult specimen from Cura��ao (Fig. 18). Further comparative study, including genetic analysis of the Cura��ao specimen, is needed. Juveniles (Fig. 19). The two juveniles are pale to bright orange. Most fins have some orange coloration, and the first dorsal is predominantly orange. There are melanophores on the anterior rays of the first dorsal, second dorsal, and anal fins, as well as on the anterior base of the second dorsal fin. The outer rays of the caudal fin are densely pigmented. There is a dark spot behind the eye on the opercle and two white stripes in the eye, one above and one below the pupil. There are two dark spots on the body, one on the trunk well below the posterior base of the second dorsal fin and one on the caudal peduncle. The latter is mostly situated above the lateral line and tapers ventrally. There are 13���14 gill rakers on the lower limb of the first gill arch. Comparisons Among Juveniles. Juveniles of A. pseudomaculatus most closely resemble juvenile A. maculatus and A. lachneri in having a spot or blotch of pigment beneath the second dorsal fin (vs. bars of pigment in juveniles of A. binotatus, A. pillionatus, A. phenax and A. townsendi, and no pigment beneath the second dorsal fin in A. aurolineatus). Juvenile A. pseudomaculatus differs from juvenile A. lachneri in having the trunk blotch positioned well below the second dorsal-fin base (vs. just behind the second dorsal-fin base) and in having a dark blotch on the caudal peduncle (lacking in A. lachneri). Apogon pseudomaculatus juveniles can be distinguished from A. maculatus juveniles by the position of the spot beneath the second dorsal fin (well below it in A. pseudomaculatus, on the fin base in A. maculatus), and by the shape of the caudal-peduncle mark (mostly concentrated above the lateral line in A. pseudomaculatus, extending well below the lateral midline in A. maculatus)., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on page 18, DOI: 10.5281/zenodo.279426, {"references":["Bohlke, J. E. & Chaplin, C. C. G. (1993) Fishes of the Bahamas and Adjacent Tropical Waters. Second Edition. University of Texas Press, Austin, 771 pp.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.) The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391."]}

Apogon sp. 1 Identification. No adults match the single larval specimen of this lineage in our genetic analysis. This unidentified species could be A. evermanni, A. leptocaulus, or A. gouldi, and specimens of those species are needed for comparative genetic analysis. Alternatively, the unidentified larval specimen may represent an undescribed species. Larvae (Fig. 7). Apogon sp. 1 is represented by one 10.0 mm SL specimen. The body is mostly orange with some transparent areas. The snout is mostly transparent, but there is a small, pale orange spot at the anterior tip. The jaws are pale orange. The rest of the head and anterior portion of the trunk are orange, the area immediately behind the eye the most intense orange. The posterior region of the body is mostly transparent, with one band of pale orange pigment below the second dorsal fin and one spot of bright orange pigment on caudal peduncle. There is a line of bright orange pigment on the ventral midline of the body from the anal fin to the caudal peduncle. All fins are clear except the proximal portions of the anal-fin rays have pale orange coloration. There are melanophores on top of the head, behind the eye on the temporal region, over the swimbladder, and on the lateral surface of the gut. The caudal-peduncle length is 35 % SL. Comparisons Among Larvae. Larval Apogon sp. 1 most closely resemble A. binotatus, A. phenax, A. planifrons, and A. townsendi larvae in having orange pigmentation concentrated mainly on the head and anterior portion of the trunk and transparent areas on the snout and usually also on posterior portions of the trunk. Apogon sp. 1 differs from A. planifrons in lacking yellow pigment on the anterior portion of the body and from larval A. phenax in lacking a prominent orange spot on the snout. From A. maculatus, larval Apogon sp. 1 differs in lacking orange pigment on the first dorsal fin and in having a longer caudal peduncle (35 % SL in Apogon sp. 1, 30% SL in A. maculatus). From A. aurolineatus, larval Apogon sp. 1 differs in lacking orange/yellow pigment on the dorsal, pelvic, and anal fins; in having transparent areas on the trunk (vs. trunk completely orange in A. aurolineatus); and in having a longer caudal peduncle (35 % SL in Apogon sp. 1 vs. 27���29 % SL in A. aurolineatus). Apogon sp. 1 lacks the distinctive pattern of chromatophores on the median fins typical of A. mosavi and the single specimen examined has a longer caudal peduncle (35 % SL in Apogon sp. 1 vs. 31���34 % SL in A. mosavi). Additional larval specimens of Apogon sp. 1 are needed to confirm the differences noted above and to determine if it can be separated from larval A. binotatus and A. townsendi., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on page 10, DOI: 10.5281/zenodo.279426

Apogon sp. 2 Identification. No larvae or juveniles match the five unidentified adult specimens of this lineage in our genetic analysis (Appendix 1, one adult is shown in Fig. 31). The lineage clusters phenetically with A. quadrisquamatus, A. mosavi, and A. robbyi (Fig. 1), and all of those species lack dark markings on the body. There is a concentration of pigment on the central area of the caudal peduncle that is primarily orange and contains few if any melanophores. In preserved specimens this may appear as a diffuse oval blotch of melanophores or no marking at all. In the other species, there is a distinct basicaudal bar (A. mosavi) or circular spot (A. quadrisquamatus and A. robbyi) of melanophores that is retained in preserved specimens. Specimens in this lineage usually have 13 gill rakers on the lower limb of the first arch vs. usually 14 or 15 in A. mosavi, but additional material is needed to determine if there are modal differences in any counts. This lineage likely represents an undescribed species.

Apogon robbyi Gilbert and Tyler Identification. One adult specimen (Fig. 32), identified based on the presence of seven dusky stripes on the body (Gilbert &Tyler, 1997; Gon 2002), served as the basis for genetic identification of two juveniles (Appendix 1). Juveniles (Fig. 33). The juveniles (17.0 and 22.0 mm SL) have the distinctive body stripes of adults, but the stripes are orange and paler than the dusky stripes in adults and not apparent in preserved specimens. There is yellow pigment on the first and second dorsal fins and upper lobe of the caudal fin that roughly mirrors that on the pelvic, anal, and lower lobe of the caudal fin, respectively. There is a round basicaudal spot of melanophores and orange chromatophores that persists in preserved specimens as a well-defined, medially situated, dark blotch. The larger juvenile has 12 gill rakers on the lower limb. Comparisons Among Juveniles. Juvenile A. robbyi can be separated from other known Apogon juveniles by the seven orange-colored stripes on body. It resembles young A. quadrisquamatus and A. mosavi in having chromatophores on the vertical and pelvic fins, but in A. robbyi this pigment is yellow vs. yellow and orange or all orange. As in A. quadrisquamatus and A. mosavi, the pigment on the dorsal fins and upper caudal lobe appears to mirror that on the pelvic and anal fins and lower lobe of the caudal fin. Modal numbers of gill rakers on the lower limb of the first arch are useful in separating juvenile A. robbyi (12���13) from A. mosavi (14���15) but not from A. quadrisquamatus and Apogon sp. 2 (13)., Published as part of Baldwin, Carole C., Brito, Balam J., Smith, David G., Weigt, Lee A. & Escobar-Briones, Elva, 2011, Identification of early life-history stages of Caribbean Apogon (Perciformes: Apogonidae) through DNA Barcoding, pp. 1-36 in Zootaxa 3133 on page 26, DOI: 10.5281/zenodo.279426, {"references":["Gilbert, C. R. & Tyler, J. C. (1997) Apogon robbyi a new cardinalfish (Perciformes: Apogonidae) from the Caribbean Sea. Bulletin of Marine Science, 60, 764 - 781.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.) The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391."]}

Astrapogon Identification. Astrapogon specimens in our samples form 3 distinct genetic lineages corresponding to the three known species (Fig. 7). Intraspecific variation ranges from 0.00– 1.30 % nucleotide substitutions per site as opposed to 15.62–20.06 % for interspecific variation (Table 6). We identified adult A. puncticulatus and A. stellatus through counts of pectoral-fin rays and gill rakers in adults (A. puncticulatus typically with 16 pectoral-fin rays and 12–13 lower-limb gillrakers; A. stellatus usually with 15 pectoral-fin rays and 11 lowerlimb gill rakers—Böhlke & Randall 1968). By process of elimination, the third clade, comprising only larvae, is A. alutus. A. alutus N= 3 A. puncticulatus N= 18 A. stellatus N= 7 A. alutus 0.72 % (0.62–0.93) A. puncticulatus 19.11 % (18.44–20.06) 0.39 % (0.00– 1.30) A. stellatus 16.06 % (15.62–16.62) 17.13 % (16.60–18.23) 0.29 % (0.00– 0.62) Larvae and Juveniles (Figs. 8, 9). Astrapogon larvae can be distinguished from larvae of other apogonids by the heavily pigmented (often black) first dorsal and pelvic fins, and usually by the trunk melanophores, which, when expanded, cover most of the body and variously obscure the pale orange chromatophores beneath. (Note: reared juvenile specimens are available for A. puncticulatus and A. stellatus but not for A. alutus; we incorporate notes on the juveniles within the appropriate sections below.) Astrapogon alutus: Three larvae, 5–6 mm SL. First dorsal and pelvic fins heavily pigmented, with large dark spots (interradial membranes of first dorsal fin almost black in one 5 -mm SL specimen) and yellow chromatophores. Second dorsal and anal fins mostly clear, one or two melanophores extending from trunk onto bases of fins. Caudal fin with no pigment in one 5 mm specimen, with one orange chromatophore at base of one ray in dorsal lobe in the other 5 mm specimen, and with several orange chromatophores at bases of central and ventral rays in 6 mm specimen. Posterior end of caudal peduncle clear, trunk pigment ending abruptly on anterior part of peduncle. Background color of head and body yellow and orange. Large melanophores present on head and body. Snout, jaws, cheek, temporal region, and lateral surface of gut with melanophores. A dense cap of melanophores visible over swimbladder. Astrapogon puncticulatus: Size range of larvae 7.5–12 mm SL; size range of juveniles 10–14.5 mm SL. First dorsal and pelvic fins heavily pigmented in larvae, almost black but with some yellow pigment on or between fin elements. Second dorsal fin mostly clear, one or two melanophores usually extending from trunk onto base of fin. Anal fin primarily clear, a melanophore usually extending from trunk onto base of fin and one melanophore usually present at base of anterior part of fin—sometimes on one side of body only. Caudal fin usually with a few orange chromatophores at bases of central rays. In reared specimens 12 mm SL and larger, heavy pigment present on second dorsal and anal fins, but both fins with distinctive clear areas; second dorsal usually without pigment along distal edges of all rays, posteriormost one or two rays, and near proximal base of posterior fin rays; these clear areas roughly forming a semicircle around pigmented portion; anal fin with similar clear areas in most specimens. Caudal fin in reared specimens mostly dark; distal tips of all rays and proximal portions of central (and sometimes additional) rays clear. In larvae, posterior end of caudal peduncle with three melanophores in vertical bar. In one 9 -mm SL larva this bar of melanophores separated by a small but distinct gap from the pigment immediately anterior to it; in another 9 mm specimen this small gap present on left side of body only. Background color of head and body mostly orange, some yellow pigment above swimbladder. Large melanophores present on head, usually including top of head, snout, jaws, cheek, and temporal region; head pigment sometimes including four conspicuous lines of pigment radiating from eye—one dorsally, two posteriorly, and one ventrally. Large melanophores present on body but conspicuously absent on dorsal trunk just anterior to first dorsal fin, just behind second dorsal fin, and posterior to gut; when trunk melanophores in expanded state, these areas conspicuously paler than remainder of trunk. In one of smallest larvae available, 8.5 mm, trunk lacking melanophores except two beneath first dorsal fin and one on caudal peduncle. In a 7.5 -mm specimen, trunk melanophores contracted but present in typical number and pattern for A. puncticulatus. Reared juveniles usually darker in overall body color than larvae and exhibiting less orange coloration; pale areas as described above still evident in most juveniles. Lines of pigment radiating from eye observed in larvae usually visible in juveniles. Cap of melanophores usually visible over gut and swimbladder as in larvae. Astrapogon stellatus: One larva, 7 mm SL; four juveniles reared from wildcaught larvae, 10–13 mm SL. In the larva, first dorsal and pelvic fins heavily pigmented, almost black but with some yellow pigment on or between fin elements. Second dorsal and anal fins mostly clear, trunk melanophores extending onto base of fin. In reared specimens, second dorsal and anal fins each with a blotch of melanophores set within and usually entirely surrounded by pale remainder of fin. In specimens with greatly expanded melanophores, basal fin pigment sometimes merging slightly with the blotch of melanophores on fin. Caudal fin without melanophores in larva but with a few scattered orange chromatophores at bases of central and ventral rays. In reared specimens caudal fin with two large, rounded melanophores at base of fin and one or two smaller melanophores above upper spot and below lower; fin rays clear centrally; two concentrations of melanophores present posteriorly (appearing either as two patches of small spots or as two large pigmented blotches), one on upper caudal lobe and one on lower. Tips of fin rays clear. Background color of head and body mostly orange in larval specimen, some yellow anteriorly; no chromatophores on body in reared specimens. Large, greatly expanded melanophores covering head and body in larva. Trunk nearly chocolate brown in reared specimens, with expanded but distinct melanophores in other two reared specimens. Posterior end of caudal peduncle in larva with at least two melanophores in vertical bar and with a clear area anterior to those spots. Caudal peduncle uniformly dark in chocolate brown reared specimens, with small clear area anterior to bar of melanophores on posterior end of caudal peduncle in other reared specimens. Snout, jaws, cheek, temporal region, and lateral surface of gut with melanophores. A dense cap of melanophores visible over gut and swimbladder in larva. Comparisons: Larvae of A. puncticulatus differ from the other species in usually having three areas on the trunk that conspicuously lack melanophores: just anterior to origin of first dorsal fin, just posterior to second dorsal fin, and just above anterior end of anal fin (just posterior to the gut). These gaps can be seen in specimens in which the trunk melanophores are contracted (Fig. 8 E) or expanded (Fig. 8 D). The gap behind the second dorsal fin is sometimes small (Fig. 8 F) because of an additional melanophore on the dorsal portion of the caudal peduncle, and in one unusual specimen (Fig. 8 G), so few melanophores are present that there is no conspicuous pattern of gaps. Larvae of A. stellatus sometimes exhibit less distinctive pale areas in similar but slightly different locations than those in A. puncticulatus (Fig. 8 C): the gap anterior to origin of first dorsal fin is a bit more forward, the gap at posterior end of second dorsal fin is below the fin (vs. behind it), and the gap above anal fin is above the anterior third of the fin (vs. above the most anterior point of the fin). Larvae of A. puncticulatus have an overall paler appearance than larvae of A. alutus and A. stellatus; this is, in part, because the pale areas described above reveal more of the pale orange background body color but also because the trunk melanophores, even when expanded, are generally smaller or lighter than those in the other species. Most A. puncticulatus larvae lack a clear area on the caudal peduncle anterior to the posterior bar of melanophores, a feature present in the larval A. stellatus; the entire peduncle is clear in 5 - and 6 -mm SL specimens of A. alutus. A small clear gap is present on the caudal peduncle in two 9 -mm SL specimens of A. puncticulatus, however, suggesting that it could be present in early ontogenetic stages of all three species. It is not possible with our limited larval Astrapogon material to definitively diagnose larvae of A. alutus and A. stellatus. Astrapogon alutus differs from the other species in having yellow and orange body coloration vs. primarily orange. However, the largest (and only) larvae of A. alutus we have collected are 5 and 6 mm SL, and the smallest larvae of A. puncticulatus and A. stellatus are 8.5 and 7 mm SL, respectively. Accordingly, the differences in body coloration could be attributable to differences in size or developmental stage. Astrapogon alutus is unique in having no melanophores on the caudal peduncle, and A. stellatus has a bar of melanophores posteriorly on the caudal peduncle preceded by a clear area. Those differences may reflect variation typical of early larval stages: in a series of Astrapogon larvae from a single collection (all approximately 6 mm SL) not identified using DNA but representing either A. alutus or A. stellatus based on trunk pigmentation, the pigment on the caudal peduncle is highly variable: pigment is absent on the caudal peduncle in one specimen, bears a single spot on one side and several on the other in another specimen, has a posterior bar of pigment and a clear area preceding it in several specimens, and has the peduncle entirely pigmented in another. Juveniles of A. puncticulatus differ from those of A. stellatus in the patterns of pigment on the second dorsal, anal, and caudal fins: in A. puncticulatus, the second dorsal and anal fins have a large blotch of pigment with clear areas forming roughly a semicircle around the posterior end of the blotch; in A. stellatus, the pigment blotch on the second dorsal and anal fins is smaller and completely surrounded by clear areas. Caudal-fin pigment of juvenile A. puncticulatus comprises a horizontally elongate oval centered on the proximal portion of the fin, whereas in A. stellatus the caudal fin has two large, rounded melanophores at the base of the fin and two concentrations of melanophores further out on the fin. Identification of juvenile A. alutus is needed to determine if the fin pigment in A. puncticulatus or A. stellatus, or both, is diagnostic. Comments on previous identifications of Astrapogon larvae . Lara (2006: 1391) provided illustrations of two specimens of A. stellatus, 5.7 and 7.8 mm SL. The 5.7 mm specimen looks very similar to both our 6 mm specimen of A. alutus and 7 mm specimen of A. stellatus except that it lacks a clear area on the caudal peduncle. Lara’s 7.8 mm specimen of A. stellatus bears the distinctive pale areas in front of the first dorsal fin, beneath the posterior portion of the second dorsal fin, and above the anterior third of the anal fin characteristic of some of our larval A. stellatus. As noted above, A. puncticulatus has similar pale areas, but it is never as uniformly covered with melanophores as in Lara’s specimen and our A. stellatus. Lara (2006: 1389) illustrated five larval and juvenile specimens of A. puncticulatus. The pale areas characteristic of A. puncticulatus appear to be present in the 5.6, 8.6, and 14.3 mm specimens (her figures A, B, D). The 10.5 mm larva (her figure C) appears to have small melanophores anterior to the first dorsal fin and several melanophores behind the second dorsal fin. The absence in that specimen of pigment on the first dorsal and pelvic fins is odd and unlike that of any larval Astrapogon specimen we have examined. Possibly there is geographical variation in pigment patterns of Astrapogon between Florida (where Lara’s specimens were obtained) and Belize. The 16.8 mm juvenile (her figure E) appears to have the clear area on the proximal part of the posterior half of the second dorsal fin typical of A. puncticulatus and a similar band on the anal fin. Lara (2006) did not indicate how she identified the larval stages of Astrapogon prior to their developing the full complement of gill rakers and pectoral-fin rays, and her descriptions contain some errors. In her description of A. stellatus larvae, Lara (2006: 1390) noted that, by 5mm, most of the body is heavily pigmented except for the dorsal third, but her illustrations of 5.7 and 7.8 mm specimens depict heavy pigmentation on the dorsal third—consistent with our specimens of A. stellatus. She also indicated (2006: 1390) that “D 1 & P 1 fins darkly pigmented,” but the pectoral fin is clear in her illustrations and presumably she meant the pelvic fins: as she correctly noted later in the description, the first dorsal and pelvic fins are heavily pigmented. Likewise, in the description of A. puncticulatus, she noted that small larvae have large melanophores on the head and anterior half of the body, but her illustration of a 5.6 mm specimen reveals large melanophores on the posterior half of the body as well. Large melanophores are present posteriorly in our specimens. She mistakenly indicated the presence of heavy pigment on the pectoral fin as in A. stellatus. One image of A. puncticulatus was located on the internet (Victor 2008). The 9.2 -mm SL larva from San Blas, Panama, agrees well in pigmentation with our A. puncticulatus.

Phaeoptyx Identification. Our Phaeoptyx specimens form three distinct genetic lineages, corresponding to the three known species (Fig. 1). Intraspecific variation ranges from 0.00– 2.19 % nucleotide substitutions per site as opposed to 15.20–20.80 % for interspecific variation (Table 2). See “Comments on Identification of Adult Phaeoptyx,” below, for a discussion of morphological identification of adult Phaeoptyx. P. p i g m e n t a r i a N= 44 P. conklini N= 26 P. xenus N= 6 P. p i g m e n t a r i a 0.56 % (0.00– 2.19) P. conklini 16.79 % (15.65–19.02) 0.78 % (0.00– 5.81) P. xenus 16.03 % (15.20–20.80) 18.10 % (16.71–20.80) 0.43 % (0.00– 0.93) Larvae (Fig. 2). Phaeoptyx larvae are easily distinguished from other apogonids by the presence of yellow chromatophores on the first dorsal and pelvic fins and usually melanophores on some portion of the pelvic fin. The yellow fin pigment typically disappears about the same time as the blotch of melanophores appears on the caudal peduncle, and here we separate larval and juvenile stages based on this transition in pigment. This also corresponds well with settlement from the plankton, as most of our net-collected larvae that have caudalpeduncle pigment do not have the pigment well defined as it is in juveniles and adults. Diagnostic features of the species are given below. Phaeoptyx conklini: Size range 10–13.5 mm SL. First dorsal fin yellow; no melanophores. Second dorsal fin clear. Pelvic fin yellow, usually more strongly so proximally, with a prominent melanophore at the base and linear series of melanophores along the fin rays to the distal tip of the fin. Anal fin mostly clear; orange, if present, restricted to base. Posterior end of caudal peduncle with two dense, vertical crescent-shaped bands of orange chromatophores overlying posterior margin of hypural plate, the dorsal one only partially formed in small specimens. Caudal fin with orange chromatophores on bases of central rays; remainder of fin without pigment. Head mostly orange, some specimens with yellow chromatophores over posterior portion of brain; scattered melanophores present on top of head, a couple sometimes present on cheek, and heavy pigment developing in temporal region by approximately 12 mm. Scattered melanophores present on gut and a cap of melanophores usually visible over swimbladder in all larvae, and a series of internal melanophores usually developing along anterior portion of vertebral column by 12 mm. Phaeoptyx pigmentaria: Size range 13–18 mm SL. First dorsal fin usually yellow, last two elements sometimes orange; melanophores, if present, restricted to the distal half of anterior spines. Second dorsal fin with orange stripe along proximal one fifth to one third of fin rays. Pelvic fin yellow; melanophores, if present, restricted to distal half of fin (or distal tips of fin rays). Anal fin mostly orange. Posterior end of caudal peduncle with two dense, vertical crescent-shaped bands of orange chromatophores overlying posterior margin of hypural plate; caudal fin mostly orange, melanophores usually present at distal ends of outer branched rays of dorsal and ventral lobes. Head mostly orange, some specimens with yellow chromatophores over posterior portion of brain; scattered melanophores present on top of head and a few present on cheek/ temporal region by 13 mm; larger larvae usually also with melanophores on distal margins of premaxilla and dentary. Scattered melanophores present on gut, and a cap of melanophores visible over swimbladder in all larvae; a series of internal melanophores developing along anterior portion of vertebral column by 15 mm. Phaeoptyx xenus: Size range 8–10 mm SL. First dorsal fin primarily yellow, last two fin elements clear; no melanophores. Second dorsal-fin spine elongate, extending when depressed beyond posterior base of second dorsal fin in 8 -mm SL specimen, to middle of second dorsal fin in 10 -mm SL specimen. Second dorsal fin clear. Pelvic fin orange at base; the remainder of fin yellow with scattered melanophores on the distal three quarters. Anal fin primarily clear. Posterior end of caudal peduncle with two vertical crescent-shaped bands of orange chromatophores overlying posterior margin of hypural plate, the dorsal one only partially formed in 10 mm specimen, absent in 8 mm specimen. Caudal fin with a few scattered chromatophores on anterior portion of central rays; remainder of fin clear. Head mostly orange, with bright yellow chromatophores over posterior portion of brain; a few melanophores present on top of head and two to several present on temporal region. Lateral surface of gut yellow in 8 -mm specimen, pale yellow and with a few melanophores in 10 mm specimen. A cap of melanophores developing over swimbladder in 8 mm specimen, well developed and prominent at 10 mm. The 10 mm specimen also with a well-developed series of internal melanophores on anterior portion of vertebral column. Comparisons (Table 3): The patterns of chromatophores and melanophores on the fins usually distinguish larvae of the three species. Phaeoptyx pigmentaria is unique in typically having melanophores on the first dorsal and caudal fins, a wide orange stripe at the base of the second dorsal fin and a mostly orange caudal fin. Phaeoptyx conklini is distinctive in having a prominent melanophore at the base of (and usually spots along the entire length of) the pelvic fin; in the other species, melanophores are restricted to the distal half (P. p i g m e n t a r i a, if present) to three quarters (P. x e n u s) of the fin. Phaeoptyx xenus is distinctive in having the second dorsal-fin spine elongate. Some of the differences noted above may be related to developmental stage and size: although P. pigmentaria is the most common larval Phaeoptyx species in our samples, we have not obtained larvae smaller than 13 mm, whereas the largest larvae of P. xenus and P. conklini we have collected are 10 and 13.5 mm, respectively. Species Second D 1 Melanophores: D 1 Melanophores: P 2 Melanophores: Caudal Fin Element Elongate Species Chromatophores: D 1 Chromatophores: D 2 Chromatophores: Caudal Fin Juveniles (Fig. 3). Phaeoptyx juveniles can be distinguished from other juvenile apogonids by the pattern of pigment – from Astrapogon in lacking heavily pigmented first dorsal and pelvic fins, and from Apogon in always having a blotch of pigment on the caudal peduncle (vs. caudal-peduncle spot present only in some species), trunk beginning to develop uniform covering of melanophores (vs. melanophores only in discrete spots or bars), and in having a pale pink color (vs. having bright orange chromatophores covering most of trunk). Phaeoptyx conklini: Size range 14–22 mm SL. Snout and upper and lower jaws with orange chromatophores, never yellow. Tip of snout blunt, forming almost a 90 o angle with dorsal profile. Second dorsal and anal fins lacking distinct stripes of orange pigment but entire fin may be orange. Specimens 15 mm SL and smaller lacking melanophores on second dorsal and anal fins. Specimens 17 mm SL and larger with distinct stripes of melanophores forming at bases or slightly apart from bases of second dorsal and anal fins. Posterior end of caudal peduncle with vertical, dark bar spanning most of peduncle; anterior edge of bar irregular, the central portion usually extending further anteriorly than rest of bar. Posterior end of peduncle retaining the two vertical crescent-shaped bands of orange chromatophores of larvae. Caudal fin orange and with numerous melanophores in wildcaught juveniles, usually pale and with fewer melanophores in reared specimens. Teeth, when developed, small (see Fig. 5). Phaeoptyx pigmentaria: Size range 17–21 mm SL. Snout and upper and lower jaws usually with yellow chromatophores. Tip of snout rounded. No stripe of melanophores along bases of second dorsal and anal fins. Second dorsal fin with orange stripe along proximal one fifth to one third of fin rays. Anal fin with similar orange stripe in wildcaught specimens (lacking in some reared juveniles). Caudal peduncle with dark bar usually spanning entire peduncle; anterior edge of bar relatively straight and uniform. Posterior end of peduncle retaining the two vertical crescent-shaped bands of orange chromatophores of larvae. Caudal fin mostly pale orange, with numerous scattered melanophores. Teeth, when developed, enlarged (see Fig. 5). Phaeoptyx xenus: Presumably juveniles are typically larger than our largest larva, 10 mm SL, and smaller than our smallest adult, 28 mm SL; our description here is based on one reared larva measuring 11 mm SL and a wild-caught juvenile of 21 mm SL. Snout and upper and lower jaws pale yellow; tip of snout rounded. Second dorsal and anal fins lacking distinct stripes of pigment in 11 mm specimen, with well-developed stripes of pigment in 21 mm specimen. First dorsal and caudal fins with scattered melanophores in 21 mm specimen. Center of caudal peduncle with vertical oval of melanophores, oval not reaching dorsal and ventral body margins. Posterior end of caudal peduncle with two crescent-shaped lines of orange chromatophores as in larvae. Teeth small. Species Yellow Snout Tip of Snout Enlarged Teeth in Melanophores: Stripes at Bases of D 2 and A and Jaws Jaws Species Melanophores: Caudal Peduncle Chromatophores: D 2 P 1 Rays Comparisons (Table 4): Juveniles typically can be separated by the following: size and shape of the blotch of pigment on the caudal peduncle (spanning entire depth of peduncle and with a relatively uniform anterior margin in P. pigmentaria, usually spanning entire depth of peduncle and with irregular anterior margin in P. conklini, more oval shaped and not reaching dorsal and ventral body margins in P. xenus); presence or absence of yellow pigment on the snout and tip of lower jaw (usually present in P. pigmentaria and P. xenus, absent in P. conklini); shape of the tip of the snout (blunt in P. conklini, rounded in P. pigmentaria and P. x e n u s); presence or absence of basal stripes of melanophores on the second dorsal and anal fins (absent in P. pigmentaria, present and set slightly apart from bases of fins in P. conklini, present and covering bases of fins in P. xenus); and presence or absence of orange pigment proximally on second dorsal (and usually anal) fin (present in P. pigmentaria, absent in P. c o n k l i n i and P. xenus). Larger juveniles of P. pigmentaria differ from the others in having enlarged teeth. Additionally, Phaeoptyx xenus, which is not common in our samples at any life-history stage, has 10 pectoral-fin rays vs. 11–13 in the other two species. Comments on Identification of Adult Phaeoptyx (Fig. 4, Table 5) . Adult P. p i g m e n t a r i a is easily identified by the absence of a well-developed pigment stripe along the bases of the second dorsal and anal fins (although there may be a small stripe comprising one or two rows of melanophores as well as numerous scattered spots on both fins – Fig. 4 D); presence of enlarged teeth in the upper and lower jaws (Fig. 5); and by having 11–13 (rarely 13) lower-limb gill rakers (vs. 13–15, usually 14, in P. x e n u s, 14–16, usually 15, in P. conklini—Böhlke and Chaplin 1968). Phaeoptyx conklini has traditionally been separated from P. xenus by the presence of wide, dark stripes of pigment along the bases of the second dorsal and anal fins, a larger eye, and modally one more gill raker (15 vs. 14) on the lower limb (Böhlke & Chaplin 1968; Gon 2002). There is overlap in extremes of eye diameter (Fig. 6), the stripe of fin pigment may be wider in P. xenus than in P. conklini, and accurate gill-raker counts are sometimes difficult to make in small adults, especially in the field. As noted by Böhlke and Chaplin (1968), the fin stripes in P. conklini are usually darker and more prominent than in P. xenus and set slightly apart from the bases vs. directly along the bases. However, the intensity of the stripes is most informative when both species are available for comparison, and several of our P. conklini adults lack the pale streaks or gaps between the stripes of fin pigment and fin bases. The stripe of pigment on the anal fin does not extend anteriorly to the first and second anal-fin elements in P. xenus, whereas it does extend to the first two elements in P. conklini. Additionally, adult P. conklini retain the very blunt snout of juveniles, and it is not yellow as it is in adult P. pigmentaria and P. x e n u s. If the mouth is closed, the morphology of the snout is often the most distinctive diagnostic character of P. conklini. Phaeoptyx conklini usually has the pigment spot on the caudal peduncle extending to or near the dorsal and ventral body margins, whereas this spot is typically restricted to the center of the caudal peduncle in P. xenus. Finally, adults of all three species can usually be separated by the size, number, and distribution of melanophores on the trunk: in P. pigmentaria, there is usually one melanophore per scale, and the pigment forms uniform rows of spots from anterior to posterior (Fig. 4 C); in P. conklini, there are usually two or more melanophores per scale posteriorly, and the pigment spots are uniformly spaced on the trunk (Fig. 4 B); in P. xenus, the trunk pigment is irregular, with larger melanophores scattered along the center and smaller, more densely clustered melanophores dorsally and ventrally (Fig. 4 E, F). As with most other characters of Phaeoptyx, however, there is variation within species, and one specimen of P. pigmentaria, for example, has a pattern of trunk melanophores more like that of P. xenus than other P. pigmentaria (Fig. 4 D). Comments on Previous Identifications of Early Life Stages of Phaeoptyx. Lara (2006: 1393) provided illustrations of a 12 mm larva and a 16.2 mm late larval or juvenile specimen of P. conklini. The larva has the hallmark P. conklini pelvic-fin pigment—melanophores along the entire length of the fin including a spot at the base. In the description of larvae, however, Lara (2006: 1392) did not mention pelvic-fin pigment. The 16.2 -mm specimen exhibits the blunt snout, small teeth, and caudal-peduncle pigment spanning entire depth of peduncle and with irregular anterior edge typical of juvenile P. conklini. Lara (2006: 1395) provided an illustration of a 17.2 -mm SL specimen of P. pigmentaria that resembles our juvenile specimens of that species in having the blotch of melanophores on the caudal peduncle extending the entire depth of the peduncle and lacking a blunt snout, although the snout in that illustration is considerably more pointed than the snout is in any of our larval, juvenile, and adult specimens. Lara (2006: 1394) noted that larval P. pigmentaria has little or no pigment on the fins, but most of our specimens have diagnostic melanophores at the distal tips of the first dorsal, pelvic, and caudal fins. Lara (2006: 1397) illustrated 13.9 - and 22.1 -mm SL specimens of P. x e n u s. The configuration of the blotch of pigment on the caudal peduncle, which, in both specimens does not extend to the dorsal and ventral margins of the body, agrees with our juveniles of P. xenus. The pelvic fin of the 13.9 mm specimen lacks the pigment characteristic of our smaller P. x e n u s larvae. One additional illustration of Phaeoptyx provided by Lara (2006: 1399: Figure 3 C) could be P. pigmentaria based on the absence of pelvicfin pigment. 40 35 30 HL 25 % as Phaeoptyx conklini diameter 20 Phaeoptyx Phaeoptyx xenus pigmentaria Eye 15 10 5 0 0 5 10 15 20 25 30 35 40 45 50 Standard length (mm)

Astrapogon Identification. Astrapogon specimens in our samples form 3 distinct genetic lineages corresponding to the three known species (Fig. 7). Intraspecific variation ranges from 0.00��� 1.30 % nucleotide substitutions per site as opposed to 15.62���20.06 % for interspecific variation (Table 6). We identified adult A. puncticulatus and A. stellatus through counts of pectoral-fin rays and gill rakers in adults (A. puncticulatus typically with 16 pectoral-fin rays and 12���13 lower-limb gillrakers; A. stellatus usually with 15 pectoral-fin rays and 11 lowerlimb gill rakers���B��hlke & Randall 1968). By process of elimination, the third clade, comprising only larvae, is A. alutus. A. alutus N= 3 A. puncticulatus N= 18 A. stellatus N= 7 A. alutus 0.72 % (0.62���0.93) A. puncticulatus 19.11 % (18.44���20.06) 0.39 % (0.00��� 1.30) A. stellatus 16.06 % (15.62���16.62) 17.13 % (16.60���18.23) 0.29 % (0.00��� 0.62) Larvae and Juveniles (Figs. 8, 9). Astrapogon larvae can be distinguished from larvae of other apogonids by the heavily pigmented (often black) first dorsal and pelvic fins, and usually by the trunk melanophores, which, when expanded, cover most of the body and variously obscure the pale orange chromatophores beneath. (Note: reared juvenile specimens are available for A. puncticulatus and A. stellatus but not for A. alutus; we incorporate notes on the juveniles within the appropriate sections below.) Astrapogon alutus: Three larvae, 5���6 mm SL. First dorsal and pelvic fins heavily pigmented, with large dark spots (interradial membranes of first dorsal fin almost black in one 5 -mm SL specimen) and yellow chromatophores. Second dorsal and anal fins mostly clear, one or two melanophores extending from trunk onto bases of fins. Caudal fin with no pigment in one 5 mm specimen, with one orange chromatophore at base of one ray in dorsal lobe in the other 5 mm specimen, and with several orange chromatophores at bases of central and ventral rays in 6 mm specimen. Posterior end of caudal peduncle clear, trunk pigment ending abruptly on anterior part of peduncle. Background color of head and body yellow and orange. Large melanophores present on head and body. Snout, jaws, cheek, temporal region, and lateral surface of gut with melanophores. A dense cap of melanophores visible over swimbladder. Astrapogon puncticulatus: Size range of larvae 7.5���12 mm SL; size range of juveniles 10���14.5 mm SL. First dorsal and pelvic fins heavily pigmented in larvae, almost black but with some yellow pigment on or between fin elements. Second dorsal fin mostly clear, one or two melanophores usually extending from trunk onto base of fin. Anal fin primarily clear, a melanophore usually extending from trunk onto base of fin and one melanophore usually present at base of anterior part of fin���sometimes on one side of body only. Caudal fin usually with a few orange chromatophores at bases of central rays. In reared specimens 12 mm SL and larger, heavy pigment present on second dorsal and anal fins, but both fins with distinctive clear areas; second dorsal usually without pigment along distal edges of all rays, posteriormost one or two rays, and near proximal base of posterior fin rays; these clear areas roughly forming a semicircle around pigmented portion; anal fin with similar clear areas in most specimens. Caudal fin in reared specimens mostly dark; distal tips of all rays and proximal portions of central (and sometimes additional) rays clear. In larvae, posterior end of caudal peduncle with three melanophores in vertical bar. In one 9 -mm SL larva this bar of melanophores separated by a small but distinct gap from the pigment immediately anterior to it; in another 9 mm specimen this small gap present on left side of body only. Background color of head and body mostly orange, some yellow pigment above swimbladder. Large melanophores present on head, usually including top of head, snout, jaws, cheek, and temporal region; head pigment sometimes including four conspicuous lines of pigment radiating from eye���one dorsally, two posteriorly, and one ventrally. Large melanophores present on body but conspicuously absent on dorsal trunk just anterior to first dorsal fin, just behind second dorsal fin, and posterior to gut; when trunk melanophores in expanded state, these areas conspicuously paler than remainder of trunk. In one of smallest larvae available, 8.5 mm, trunk lacking melanophores except two beneath first dorsal fin and one on caudal peduncle. In a 7.5 -mm specimen, trunk melanophores contracted but present in typical number and pattern for A. puncticulatus. Reared juveniles usually darker in overall body color than larvae and exhibiting less orange coloration; pale areas as described above still evident in most juveniles. Lines of pigment radiating from eye observed in larvae usually visible in juveniles. Cap of melanophores usually visible over gut and swimbladder as in larvae. Astrapogon stellatus: One larva, 7 mm SL; four juveniles reared from wildcaught larvae, 10���13 mm SL. In the larva, first dorsal and pelvic fins heavily pigmented, almost black but with some yellow pigment on or between fin elements. Second dorsal and anal fins mostly clear, trunk melanophores extending onto base of fin. In reared specimens, second dorsal and anal fins each with a blotch of melanophores set within and usually entirely surrounded by pale remainder of fin. In specimens with greatly expanded melanophores, basal fin pigment sometimes merging slightly with the blotch of melanophores on fin. Caudal fin without melanophores in larva but with a few scattered orange chromatophores at bases of central and ventral rays. In reared specimens caudal fin with two large, rounded melanophores at base of fin and one or two smaller melanophores above upper spot and below lower; fin rays clear centrally; two concentrations of melanophores present posteriorly (appearing either as two patches of small spots or as two large pigmented blotches), one on upper caudal lobe and one on lower. Tips of fin rays clear. Background color of head and body mostly orange in larval specimen, some yellow anteriorly; no chromatophores on body in reared specimens. Large, greatly expanded melanophores covering head and body in larva. Trunk nearly chocolate brown in reared specimens, with expanded but distinct melanophores in other two reared specimens. Posterior end of caudal peduncle in larva with at least two melanophores in vertical bar and with a clear area anterior to those spots. Caudal peduncle uniformly dark in chocolate brown reared specimens, with small clear area anterior to bar of melanophores on posterior end of caudal peduncle in other reared specimens. Snout, jaws, cheek, temporal region, and lateral surface of gut with melanophores. A dense cap of melanophores visible over gut and swimbladder in larva. Comparisons: Larvae of A. puncticulatus differ from the other species in usually having three areas on the trunk that conspicuously lack melanophores: just anterior to origin of first dorsal fin, just posterior to second dorsal fin, and just above anterior end of anal fin (just posterior to the gut). These gaps can be seen in specimens in which the trunk melanophores are contracted (Fig. 8 E) or expanded (Fig. 8 D). The gap behind the second dorsal fin is sometimes small (Fig. 8 F) because of an additional melanophore on the dorsal portion of the caudal peduncle, and in one unusual specimen (Fig. 8 G), so few melanophores are present that there is no conspicuous pattern of gaps. Larvae of A. stellatus sometimes exhibit less distinctive pale areas in similar but slightly different locations than those in A. puncticulatus (Fig. 8 C): the gap anterior to origin of first dorsal fin is a bit more forward, the gap at posterior end of second dorsal fin is below the fin (vs. behind it), and the gap above anal fin is above the anterior third of the fin (vs. above the most anterior point of the fin). Larvae of A. puncticulatus have an overall paler appearance than larvae of A. alutus and A. stellatus; this is, in part, because the pale areas described above reveal more of the pale orange background body color but also because the trunk melanophores, even when expanded, are generally smaller or lighter than those in the other species. Most A. puncticulatus larvae lack a clear area on the caudal peduncle anterior to the posterior bar of melanophores, a feature present in the larval A. stellatus; the entire peduncle is clear in 5 - and 6 -mm SL specimens of A. alutus. A small clear gap is present on the caudal peduncle in two 9 -mm SL specimens of A. puncticulatus, however, suggesting that it could be present in early ontogenetic stages of all three species. It is not possible with our limited larval Astrapogon material to definitively diagnose larvae of A. alutus and A. stellatus. Astrapogon alutus differs from the other species in having yellow and orange body coloration vs. primarily orange. However, the largest (and only) larvae of A. alutus we have collected are 5 and 6 mm SL, and the smallest larvae of A. puncticulatus and A. stellatus are 8.5 and 7 mm SL, respectively. Accordingly, the differences in body coloration could be attributable to differences in size or developmental stage. Astrapogon alutus is unique in having no melanophores on the caudal peduncle, and A. stellatus has a bar of melanophores posteriorly on the caudal peduncle preceded by a clear area. Those differences may reflect variation typical of early larval stages: in a series of Astrapogon larvae from a single collection (all approximately 6 mm SL) not identified using DNA but representing either A. alutus or A. stellatus based on trunk pigmentation, the pigment on the caudal peduncle is highly variable: pigment is absent on the caudal peduncle in one specimen, bears a single spot on one side and several on the other in another specimen, has a posterior bar of pigment and a clear area preceding it in several specimens, and has the peduncle entirely pigmented in another. Juveniles of A. puncticulatus differ from those of A. stellatus in the patterns of pigment on the second dorsal, anal, and caudal fins: in A. puncticulatus, the second dorsal and anal fins have a large blotch of pigment with clear areas forming roughly a semicircle around the posterior end of the blotch; in A. stellatus, the pigment blotch on the second dorsal and anal fins is smaller and completely surrounded by clear areas. Caudal-fin pigment of juvenile A. puncticulatus comprises a horizontally elongate oval centered on the proximal portion of the fin, whereas in A. stellatus the caudal fin has two large, rounded melanophores at the base of the fin and two concentrations of melanophores further out on the fin. Identification of juvenile A. alutus is needed to determine if the fin pigment in A. puncticulatus or A. stellatus, or both, is diagnostic. Comments on previous identifications of Astrapogon larvae . Lara (2006: 1391) provided illustrations of two specimens of A. stellatus, 5.7 and 7.8 mm SL. The 5.7 mm specimen looks very similar to both our 6 mm specimen of A. alutus and 7 mm specimen of A. stellatus except that it lacks a clear area on the caudal peduncle. Lara���s 7.8 mm specimen of A. stellatus bears the distinctive pale areas in front of the first dorsal fin, beneath the posterior portion of the second dorsal fin, and above the anterior third of the anal fin characteristic of some of our larval A. stellatus. As noted above, A. puncticulatus has similar pale areas, but it is never as uniformly covered with melanophores as in Lara���s specimen and our A. stellatus. Lara (2006: 1389) illustrated five larval and juvenile specimens of A. puncticulatus. The pale areas characteristic of A. puncticulatus appear to be present in the 5.6, 8.6, and 14.3 mm specimens (her figures A, B, D). The 10.5 mm larva (her figure C) appears to have small melanophores anterior to the first dorsal fin and several melanophores behind the second dorsal fin. The absence in that specimen of pigment on the first dorsal and pelvic fins is odd and unlike that of any larval Astrapogon specimen we have examined. Possibly there is geographical variation in pigment patterns of Astrapogon between Florida (where Lara���s specimens were obtained) and Belize. The 16.8 mm juvenile (her figure E) appears to have the clear area on the proximal part of the posterior half of the second dorsal fin typical of A. puncticulatus and a similar band on the anal fin. Lara (2006) did not indicate how she identified the larval stages of Astrapogon prior to their developing the full complement of gill rakers and pectoral-fin rays, and her descriptions contain some errors. In her description of A. stellatus larvae, Lara (2006: 1390) noted that, by 5mm, most of the body is heavily pigmented except for the dorsal third, but her illustrations of 5.7 and 7.8 mm specimens depict heavy pigmentation on the dorsal third���consistent with our specimens of A. stellatus. She also indicated (2006: 1390) that ���D 1 & P 1 fins darkly pigmented,��� but the pectoral fin is clear in her illustrations and presumably she meant the pelvic fins: as she correctly noted later in the description, the first dorsal and pelvic fins are heavily pigmented. Likewise, in the description of A. puncticulatus, she noted that small larvae have large melanophores on the head and anterior half of the body, but her illustration of a 5.6 mm specimen reveals large melanophores on the posterior half of the body as well. Large melanophores are present posteriorly in our specimens. She mistakenly indicated the presence of heavy pigment on the pectoral fin as in A. stellatus. One image of A. puncticulatus was located on the internet (Victor 2008). The 9.2 -mm SL larva from San Blas, Panama, agrees well in pigmentation with our A. puncticulatus., Published as part of Baldwin, Carole C., Mounts, Julie H., Smith, David G. & Weigt, Lee A., 2009, Genetic identification and color descriptions of early life-history stages of Belizean Phaeoptyx and Astrapogon (Teleostei: Apogonidae) with Comments on identification of adult Phaeoptyx, pp. 1-22 in Zootaxa 2008 on pages 15-21, DOI: 10.5281/zenodo.185742, {"references":["Bohlke, J. E. & Randall, J. E. (1968) A key to the shallow-water Atlantic cardinalfishes (Apogonidae), with descriptions of five new species. Proceedings of the Academy of Natural Sciences of Philadelphia, 120, 175 - 206.","Lara, M. R. (2006) Apogonidae: Cardinalfishes. In: Richards, W. J. (Ed.), Early stages of Atlantic fishes: an identification guide for western central North Atlantic. CRC Press, Boca Raton, pp. 1363 - 1399.","Victor, B. (2008) A photographic guide to the larvae of coral reef fishes. Available from: http: // www. coralreeffish. com / larvae. html / (2 / 14 / 08)."]}

Phaeoptyx Identification. Our Phaeoptyx specimens form three distinct genetic lineages, corresponding to the three known species (Fig. 1). Intraspecific variation ranges from 0.00��� 2.19 % nucleotide substitutions per site as opposed to 15.20���20.80 % for interspecific variation (Table 2). See ���Comments on Identification of Adult Phaeoptyx,��� below, for a discussion of morphological identification of adult Phaeoptyx. P. p i g m e n t a r i a N= 44 P. conklini N= 26 P. xenus N= 6 P. p i g m e n t a r i a 0.56 % (0.00��� 2.19) P. conklini 16.79 % (15.65���19.02) 0.78 % (0.00��� 5.81) P. xenus 16.03 % (15.20���20.80) 18.10 % (16.71���20.80) 0.43 % (0.00��� 0.93) Larvae (Fig. 2). Phaeoptyx larvae are easily distinguished from other apogonids by the presence of yellow chromatophores on the first dorsal and pelvic fins and usually melanophores on some portion of the pelvic fin. The yellow fin pigment typically disappears about the same time as the blotch of melanophores appears on the caudal peduncle, and here we separate larval and juvenile stages based on this transition in pigment. This also corresponds well with settlement from the plankton, as most of our net-collected larvae that have caudalpeduncle pigment do not have the pigment well defined as it is in juveniles and adults. Diagnostic features of the species are given below. Phaeoptyx conklini: Size range 10���13.5 mm SL. First dorsal fin yellow; no melanophores. Second dorsal fin clear. Pelvic fin yellow, usually more strongly so proximally, with a prominent melanophore at the base and linear series of melanophores along the fin rays to the distal tip of the fin. Anal fin mostly clear; orange, if present, restricted to base. Posterior end of caudal peduncle with two dense, vertical crescent-shaped bands of orange chromatophores overlying posterior margin of hypural plate, the dorsal one only partially formed in small specimens. Caudal fin with orange chromatophores on bases of central rays; remainder of fin without pigment. Head mostly orange, some specimens with yellow chromatophores over posterior portion of brain; scattered melanophores present on top of head, a couple sometimes present on cheek, and heavy pigment developing in temporal region by approximately 12 mm. Scattered melanophores present on gut and a cap of melanophores usually visible over swimbladder in all larvae, and a series of internal melanophores usually developing along anterior portion of vertebral column by 12 mm. Phaeoptyx pigmentaria: Size range 13���18 mm SL. First dorsal fin usually yellow, last two elements sometimes orange; melanophores, if present, restricted to the distal half of anterior spines. Second dorsal fin with orange stripe along proximal one fifth to one third of fin rays. Pelvic fin yellow; melanophores, if present, restricted to distal half of fin (or distal tips of fin rays). Anal fin mostly orange. Posterior end of caudal peduncle with two dense, vertical crescent-shaped bands of orange chromatophores overlying posterior margin of hypural plate; caudal fin mostly orange, melanophores usually present at distal ends of outer branched rays of dorsal and ventral lobes. Head mostly orange, some specimens with yellow chromatophores over posterior portion of brain; scattered melanophores present on top of head and a few present on cheek/ temporal region by 13 mm; larger larvae usually also with melanophores on distal margins of premaxilla and dentary. Scattered melanophores present on gut, and a cap of melanophores visible over swimbladder in all larvae; a series of internal melanophores developing along anterior portion of vertebral column by 15 mm. Phaeoptyx xenus: Size range 8���10 mm SL. First dorsal fin primarily yellow, last two fin elements clear; no melanophores. Second dorsal-fin spine elongate, extending when depressed beyond posterior base of second dorsal fin in 8 -mm SL specimen, to middle of second dorsal fin in 10 -mm SL specimen. Second dorsal fin clear. Pelvic fin orange at base; the remainder of fin yellow with scattered melanophores on the distal three quarters. Anal fin primarily clear. Posterior end of caudal peduncle with two vertical crescent-shaped bands of orange chromatophores overlying posterior margin of hypural plate, the dorsal one only partially formed in 10 mm specimen, absent in 8 mm specimen. Caudal fin with a few scattered chromatophores on anterior portion of central rays; remainder of fin clear. Head mostly orange, with bright yellow chromatophores over posterior portion of brain; a few melanophores present on top of head and two to several present on temporal region. Lateral surface of gut yellow in 8 -mm specimen, pale yellow and with a few melanophores in 10 mm specimen. A cap of melanophores developing over swimbladder in 8 mm specimen, well developed and prominent at 10 mm. The 10 mm specimen also with a well-developed series of internal melanophores on anterior portion of vertebral column. Comparisons (Table 3): The patterns of chromatophores and melanophores on the fins usually distinguish larvae of the three species. Phaeoptyx pigmentaria is unique in typically having melanophores on the first dorsal and caudal fins, a wide orange stripe at the base of the second dorsal fin and a mostly orange caudal fin. Phaeoptyx conklini is distinctive in having a prominent melanophore at the base of (and usually spots along the entire length of) the pelvic fin; in the other species, melanophores are restricted to the distal half (P. p i g m e n t a r i a, if present) to three quarters (P. x e n u s) of the fin. Phaeoptyx xenus is distinctive in having the second dorsal-fin spine elongate. Some of the differences noted above may be related to developmental stage and size: although P. pigmentaria is the most common larval Phaeoptyx species in our samples, we have not obtained larvae smaller than 13 mm, whereas the largest larvae of P. xenus and P. conklini we have collected are 10 and 13.5 mm, respectively. Species Second D 1 Melanophores: D 1 Melanophores: P 2 Melanophores: Caudal Fin Element Elongate Species Chromatophores: D 1 Chromatophores: D 2 Chromatophores: Caudal Fin Juveniles (Fig. 3). Phaeoptyx juveniles can be distinguished from other juvenile apogonids by the pattern of pigment ��� from Astrapogon in lacking heavily pigmented first dorsal and pelvic fins, and from Apogon in always having a blotch of pigment on the caudal peduncle (vs. caudal-peduncle spot present only in some species), trunk beginning to develop uniform covering of melanophores (vs. melanophores only in discrete spots or bars), and in having a pale pink color (vs. having bright orange chromatophores covering most of trunk). Phaeoptyx conklini: Size range 14���22 mm SL. Snout and upper and lower jaws with orange chromatophores, never yellow. Tip of snout blunt, forming almost a 90 o angle with dorsal profile. Second dorsal and anal fins lacking distinct stripes of orange pigment but entire fin may be orange. Specimens 15 mm SL and smaller lacking melanophores on second dorsal and anal fins. Specimens 17 mm SL and larger with distinct stripes of melanophores forming at bases or slightly apart from bases of second dorsal and anal fins. Posterior end of caudal peduncle with vertical, dark bar spanning most of peduncle; anterior edge of bar irregular, the central portion usually extending further anteriorly than rest of bar. Posterior end of peduncle retaining the two vertical crescent-shaped bands of orange chromatophores of larvae. Caudal fin orange and with numerous melanophores in wildcaught juveniles, usually pale and with fewer melanophores in reared specimens. Teeth, when developed, small (see Fig. 5). Phaeoptyx pigmentaria: Size range 17���21 mm SL. Snout and upper and lower jaws usually with yellow chromatophores. Tip of snout rounded. No stripe of melanophores along bases of second dorsal and anal fins. Second dorsal fin with orange stripe along proximal one fifth to one third of fin rays. Anal fin with similar orange stripe in wildcaught specimens (lacking in some reared juveniles). Caudal peduncle with dark bar usually spanning entire peduncle; anterior edge of bar relatively straight and uniform. Posterior end of peduncle retaining the two vertical crescent-shaped bands of orange chromatophores of larvae. Caudal fin mostly pale orange, with numerous scattered melanophores. Teeth, when developed, enlarged (see Fig. 5). Phaeoptyx xenus: Presumably juveniles are typically larger than our largest larva, 10 mm SL, and smaller than our smallest adult, 28 mm SL; our description here is based on one reared larva measuring 11 mm SL and a wild-caught juvenile of 21 mm SL. Snout and upper and lower jaws pale yellow; tip of snout rounded. Second dorsal and anal fins lacking distinct stripes of pigment in 11 mm specimen, with well-developed stripes of pigment in 21 mm specimen. First dorsal and caudal fins with scattered melanophores in 21 mm specimen. Center of caudal peduncle with vertical oval of melanophores, oval not reaching dorsal and ventral body margins. Posterior end of caudal peduncle with two crescent-shaped lines of orange chromatophores as in larvae. Teeth small. Species Yellow Snout Tip of Snout Enlarged Teeth in Melanophores: Stripes at Bases of D 2 and A and Jaws Jaws Species Melanophores: Caudal Peduncle Chromatophores: D 2 P 1 Rays Comparisons (Table 4): Juveniles typically can be separated by the following: size and shape of the blotch of pigment on the caudal peduncle (spanning entire depth of peduncle and with a relatively uniform anterior margin in P. pigmentaria, usually spanning entire depth of peduncle and with irregular anterior margin in P. conklini, more oval shaped and not reaching dorsal and ventral body margins in P. xenus); presence or absence of yellow pigment on the snout and tip of lower jaw (usually present in P. pigmentaria and P. xenus, absent in P. conklini); shape of the tip of the snout (blunt in P. conklini, rounded in P. pigmentaria and P. x e n u s); presence or absence of basal stripes of melanophores on the second dorsal and anal fins (absent in P. pigmentaria, present and set slightly apart from bases of fins in P. conklini, present and covering bases of fins in P. xenus); and presence or absence of orange pigment proximally on second dorsal (and usually anal) fin (present in P. pigmentaria, absent in P. c o n k l i n i and P. xenus). Larger juveniles of P. pigmentaria differ from the others in having enlarged teeth. Additionally, Phaeoptyx xenus, which is not common in our samples at any life-history stage, has 10 pectoral-fin rays vs. 11���13 in the other two species. Comments on Identification of Adult Phaeoptyx (Fig. 4, Table 5) . Adult P. p i g m e n t a r i a is easily identified by the absence of a well-developed pigment stripe along the bases of the second dorsal and anal fins (although there may be a small stripe comprising one or two rows of melanophores as well as numerous scattered spots on both fins ��� Fig. 4 D); presence of enlarged teeth in the upper and lower jaws (Fig. 5); and by having 11���13 (rarely 13) lower-limb gill rakers (vs. 13���15, usually 14, in P. x e n u s, 14���16, usually 15, in P. conklini���B��hlke and Chaplin 1968). Phaeoptyx conklini has traditionally been separated from P. xenus by the presence of wide, dark stripes of pigment along the bases of the second dorsal and anal fins, a larger eye, and modally one more gill raker (15 vs. 14) on the lower limb (B��hlke & Chaplin 1968; Gon 2002). There is overlap in extremes of eye diameter (Fig. 6), the stripe of fin pigment may be wider in P. xenus than in P. conklini, and accurate gill-raker counts are sometimes difficult to make in small adults, especially in the field. As noted by B��hlke and Chaplin (1968), the fin stripes in P. conklini are usually darker and more prominent than in P. xenus and set slightly apart from the bases vs. directly along the bases. However, the intensity of the stripes is most informative when both species are available for comparison, and several of our P. conklini adults lack the pale streaks or gaps between the stripes of fin pigment and fin bases. The stripe of pigment on the anal fin does not extend anteriorly to the first and second anal-fin elements in P. xenus, whereas it does extend to the first two elements in P. conklini. Additionally, adult P. conklini retain the very blunt snout of juveniles, and it is not yellow as it is in adult P. pigmentaria and P. x e n u s. If the mouth is closed, the morphology of the snout is often the most distinctive diagnostic character of P. conklini. Phaeoptyx conklini usually has the pigment spot on the caudal peduncle extending to or near the dorsal and ventral body margins, whereas this spot is typically restricted to the center of the caudal peduncle in P. xenus. Finally, adults of all three species can usually be separated by the size, number, and distribution of melanophores on the trunk: in P. pigmentaria, there is usually one melanophore per scale, and the pigment forms uniform rows of spots from anterior to posterior (Fig. 4 C); in P. conklini, there are usually two or more melanophores per scale posteriorly, and the pigment spots are uniformly spaced on the trunk (Fig. 4 B); in P. xenus, the trunk pigment is irregular, with larger melanophores scattered along the center and smaller, more densely clustered melanophores dorsally and ventrally (Fig. 4 E, F). As with most other characters of Phaeoptyx, however, there is variation within species, and one specimen of P. pigmentaria, for example, has a pattern of trunk melanophores more like that of P. xenus than other P. pigmentaria (Fig. 4 D). Comments on Previous Identifications of Early Life Stages of Phaeoptyx. Lara (2006: 1393) provided illustrations of a 12 mm larva and a 16.2 mm late larval or juvenile specimen of P. conklini. The larva has the hallmark P. conklini pelvic-fin pigment���melanophores along the entire length of the fin including a spot at the base. In the description of larvae, however, Lara (2006: 1392) did not mention pelvic-fin pigment. The 16.2 -mm specimen exhibits the blunt snout, small teeth, and caudal-peduncle pigment spanning entire depth of peduncle and with irregular anterior edge typical of juvenile P. conklini. Lara (2006: 1395) provided an illustration of a 17.2 -mm SL specimen of P. pigmentaria that resembles our juvenile specimens of that species in having the blotch of melanophores on the caudal peduncle extending the entire depth of the peduncle and lacking a blunt snout, although the snout in that illustration is considerably more pointed than the snout is in any of our larval, juvenile, and adult specimens. Lara (2006: 1394) noted that larval P. pigmentaria has little or no pigment on the fins, but most of our specimens have diagnostic melanophores at the distal tips of the first dorsal, pelvic, and caudal fins. Lara (2006: 1397) illustrated 13.9 - and 22.1 -mm SL specimens of P. x e n u s. The configuration of the blotch of pigment on the caudal peduncle, which, in both specimens does not extend to the dorsal and ventral margins of the body, agrees with our juveniles of P. xenus. The pelvic fin of the 13.9 mm specimen lacks the pigment characteristic of our smaller P. x e n u s larvae. One additional illustration of Phaeoptyx provided by Lara (2006: 1399: Figure 3 C) could be P. pigmentaria based on the absence of pelvicfin pigment. 40 35 30 HL 25 % as Phaeoptyx conklini diameter 20 Phaeoptyx Phaeoptyx xenus pigmentaria Eye 15 10 5 0 0 5 10 15 20 25 30 35 40 45 50 Standard length (mm), Published as part of Baldwin, Carole C., Mounts, Julie H., Smith, David G. & Weigt, Lee A., 2009, Genetic identification and color descriptions of early life-history stages of Belizean Phaeoptyx and Astrapogon (Teleostei: Apogonidae) with Comments on identification of adult Phaeoptyx, pp. 1-22 in Zootaxa 2008 on pages 6-15, DOI: 10.5281/zenodo.185742, {"references":["Bohlke, J. E. & Chaplin, C. C. G. (1968) Fishes of the Bahamas and Adjacent Tropical Waters. Second Edition. University of Texas Press, Austin, 771 pp.","Gon, O. (2002) Apogonidae. In: Carpenter, K. (Ed.), The living marine resources of the Western Central North Atlantic. Vol. 3: Bony fishes part 2 (Opistognathidae to Molidae), sea turtles and marine mammals. FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5, Rome, pp. 1386 - 1391.","Lara, M. R. (2006) Apogonidae: Cardinalfishes. In: Richards, W. J. (Ed.), Early stages of Atlantic fishes: an identification guide for western central North Atlantic. CRC Press, Boca Raton, pp. 1363 - 1399."]}

Phaeoptyx and Astrapogon are represented in the Caribbean by six species (P. conklini, P. pigmentaria, P. xenus, A. alutus, A. stellatus, and A. puncticulatus). Species identification of larvae and juveniles is problematic because characters used to distinguish adults (e.g., patterns of pigmentation and numbers of gill rakers) are absent, incomplete, or difficult to discern in the young stages. Neighbor-joining trees derived from mitochondrial cytochrome oxidase 1 sequences (DNA Barcoding) were used to match early life stages and adults. Subsequent comparative analysis of preserved voucher specimens from which the DNA was extracted or digital color photographs of those specimens taken prior to preservation yielded sufficient information to separate all early life-history stages of Belizean Phaeoptyx and Astrapogon and provided additional information for field identification of adult Phaeoptyx. Patterns of chromatophores in fresh material, combined with patterns of melanophores, provide the easiest means of separating the life-history stages of Phaeoptyx. Larvae of Astrapogon species are morphologically very similar, and some differences in pigmentation detected among them may reflect different stages of development. Continued implementation of the DNA Barcoding methods and field protocol outlined herein should prove valuable in accurately identifying much more of the ichthyoplankton fauna of the Caribbean.

Akko brevis (G��nther 1864) Fig. 3 Amblyopus brevis G��nther 1864: 151, type locality Panama, Pacific coast. Holotype: BMNH 1864.1. 26.416. Gobioides brevis ��� Palmer 1952: 55 Tyntlastes brevis Jordan and Eigenmann 1887: 511 Material examined: El Salvador ��� USNM 371781, 11 specimens, Gulf of Fonseca, shrimp trawler Amancer, Captain Francisco Antonio Lopez, 13 ��06.95���N, 87 �� 52.15 ���W to 13 ��08.79���N, 87 �� 47.79 ���W, 20.0 m. USNM 371782, 28 specimens, same as above except 13 ��09.54���N, 87 �� 48.30 ���W to 13 ��08.42���N, 87 ��48.00���W. USNM 371782, 13 specimens, Gulf of Fonseca, trawl, 13 ��09.14���N, 87 �� 51.89 ���W to 13 ��09.87���N, 87 �� 50.91 ���W, 14.5 ���17.0 m, mud. USNM 371784, 20 specimens, same as above. USNM 371785, 4 specimens, Gulf of Fonseca, trawl, 13 ��09.99���N, 87 �� 50.75 ���W to 13 �� 11.01 ���N, 87 �� 49.56 ���W, 14.0���12.0 m, mud. USNM 371786, 7 specimens, Gulf of Fonseca, trawl, 13 �� 11.97 ���N, 87 �� 48.30 ���W to 13 �� 10.44 ���N, 87 �� 47.79 ���W, 10.4���13.3 m, mud. USNM 371787, 94 specimens, off Las Tunas, Departmento La Union, trawl, 13 ��04.74���N, 88 ��02.82���W to 13 ��05.02���N, 88 ��01.27���W, 21.0���25.0 m, mud. EL��01��017(JVT collection), 2 specimens cleared and stained, shrimp trawler, 13 ��09.54���N, 087�� 48.30 ���W to 13 ��08.42���N, 087��48.00���N, 20.0 m, mud. Panama��� URC 2554.011; 7 specimens, Panama, Darien; 23 Jan 2000 San Miguel, midway between Pta. Garachine and Pta. San Lorenzo, 18 km W Pta. Pati��o. BMNH 1864.1. 26.416 holotype Amblyopus brevis, Pacific coast of Panama, radiograph. USNM 339614; 1 specimen, 1954, field number W 54��326 station 47, Pacific Panama. USNM 43428; 2 specimens, Panama: U.S. Colombia (Equador to Panama) Albatross Expedition, F.C. 5577, 15 Mar 1888. Peru��� USNM 77581; 1 specimen, Paita, Peru, 13 Apr 1907. USNM 128187; 1 specimen, Sechura Bay near Sechura 16 May 1941. Description: Morphometric data are given in Table 1. Frequency distributions of numbers of pectoral�� and caudal��fin rays are given in Table 2. 21 36 3 10 87 23 1 11 90 14 Median fins: Each count is followed parenthetically by the number of specimens recorded with that count; an * indicates value for holotype. The first dorsal fin has seven spines (145 *) or six (1) [specimen missing the posteriormost spine], 15 second dorsal��fin elements (146 *), and 15 anal��fin elements (143 *). The caudal fin is lanceolate, with 17 (133 *) segmented rays, 3���5 procurrent rays in the upper lobe, and 2���5 in the lower lobe (Table 2). Rays of the median fins are serially branched three to four times. Caudal skeleton: Hypurals 1���2 are fused to one another, and hypurals 3, 4, and the terminal half centrum are fused into a single unit. The parhypural supports the lower segmented element, hypural 5 is free, and the single epural supports the uppermost segmented element. Paired fins: The pectoral fins are relatively short, falling just short of the posterior extreme of the pelvic fin when depressed. There are 19���21 pectoral��fin rays, all dichotomously branched except for the uppermost and lowermost rays. The pelvic fins are united, forming a disk with a well��developed anterior frenum. The pelvic base is a muscular pedicel with strong muscle bundles extending from the vicinity of the pelvic process to the vicinity of the pelvic spine, with some of the fibers inserting on the spines. All pelvic��fin rays are branched dichotomously four to five times. The pelvic fin extends posteriorly to a point that is short of the anus by a distance equal to 2���3 eye diameters. Pectoral and pelvic girdle: The osteology is the same as that described by Birdsong & Robins (1995) for A. dionaea. Head: The mouth is large, capable of expanding to a complete circle equal to the body diameter in live specimens. When the mouth is closed, the opening is oriented at an angle of 50 �� to the horizontal. The upper jaw is slightly protrusible, and there is no rostral frenum. An oral velum is present with dorsal and ventral projections. Upper and lower lips possess 10���15 fleshy papillae along the lateral upper edges. The anterior nostril is composed internally of a thickened, U��shaped, fleshy tube (the anterior section of which forms the external nostril, the remainder is visible under the overlying skin). A histological examination of this structure shows it to possess some smooth muscle fibers. The posterior nostril consists of a short tube or raised rim with no muscular development. The gill opening extends from the fourth pectoral ray ventrally to just above the pelvic��fin base, terminating at a membrane supported by the second branchiostegal ray and extending to the ventral��most pectoral ray. No pseudobranch present. The eye is small, covered with skin, slightly recessed, and has a large lens. Head canal pores B��� and G��� are present. No preopercle pores are present. Sensory papillae: The sensory papillae pattern is shown in Fig 2. Horizontal row b has an elongate anterior extension ending under the eye; row d is complete, not divided into two sections, and row ot extends ventrally onto the branchiostegals, ending just past the second branchiostegal ray. The first vertical papillae row is complete, extending from the eye to near row d. Additional vertical rows are divided by row b, with two rows above and four rows below row b. Only the last vertical row below row b extends below the level of row d. Dorsal rows x 1 and x 2 are continuous, or nearly so, as is row n. Jaws: The upper jaw has a single row of enlarged teeth, 10���11 on each premaxilla in males and 16���18 in females. The lower jaw consists of two rows of teeth, an outer row of 6���8 enlarged teeth in males (7���10 in females), and an inner row of 13���15 small teeth in males (18���20 in females). There is an unossified extension of the ventromedial process of the dentary and the ventral margin of the anguloarticular. The lacrimal is large and ossified. Myology: The superficial adductor mandibulae is subdivided into the A 1 beta, A 1 alpha, and A 2 bundles (Fig. 4). The A 1 beta passes under the large lacrimal and inserts directly by a broad non��ligamentous connection onto the posterodorsal margin of the maxilla. Both the A 1 alpha and A 2 combine via a strong tendon, which inserts on the coronoid process of the dentary. The ramus mandibularis V passes between the A 1 and A 2. The origin of the A 1 alpha is along the anterior margin of the dorsal half of the preopercle. Adductor mandibulae 2 originates along the anterior margin of the ventral half of the preopercle and along the quadrate, but the area of origin does not reach the condyle of the quadrate. Braincase: The osteology of the cranial region of Akko brevis and A. dionaea is essentially the same. The primary difference is the lateral expansion of the frontals and parasphenoid in A. dionaea, which is absent in A. brevis. Vertebral column: There are 11 precaudal and 16 caudal vertebrae (144 *). Pleural ribs of vertebrae 3���11 articulate with their respective parapophyses. Epineural bones are found on vertebrae 1���21. Epineurals 1 and 2 articulate with the parapophyses, 3���10 with the pleural ribs, 11 articulates with the parapophysis dorsal to its articulation with the pleural rib, and the remaining epineurals articulate with the centrum of each vertebra. The posterior process of the neural spine present in A. dionaea on each of vertebrae 1���6 is variably present or absent from most of the neural spines in A. brevis. When present it ranges in size from very small to large, and size is not consistent between specimens. The haemal spines of caudal vertebrae 13 and 14 are broad but not bifurcated. There are 2 anal��fin pterygiophores preceding the first haemal spine. Scales: Scales are cycloid, small, embedded, and non��overlapping anteriorly, becoming larger and overlapping on the caudal peduncle. There are 53���60 in the lateral series. Predorsal scales extend forward to the region above the anterior portion of the opercle. The pectoral��fin base, operculum, and breast are naked. No modified basicaudal scales are present. Viscera: A small gas bladder is located in the posterior 1 / 3 of the body cavity and is characterized by a thick tunica interna. The intestine is coiled twice, and its total length is about equal to �� body length. A single gravid female contained about 700 eggs in one of the ovaries, each about 0.5 mm in diameter. Genitalia: In males the genital papilla is short, thin, and pointed. Gravid females possess a short, rounded, bulbous papilla with a rounded opening and a narrow slit along the dorsal margin. Non��gravid females do not have a bulbous papilla. Pigment: The general body color is whitish with a light pink overtone in live specimens. The upper body has numerous densely packed melanophores, the pigment becoming less dense ventrally and ending at the lateral midline. The belly is whitish from the ventral surface to the lateral midline. The head and upper lip have densely packed small melanophores dorsally that become less concentrated ventrally and terminate at the ventral midline. The pectoral, pelvic, and anal fins are transparent. The caudal fin is black except for the midbasal area, which is whitish with a few scattered melanophores. The dorsal fins have scattered melanophores restricted to the membranes between the fin elements that become more concentrated towards the margin of the fin. Neither males nor females possess any genital pigmentation., Published as part of Van, James L., Tassell & Baldwin, Carole C., 2004, A review of the gobiid genus Akko (Teleostei: Gobiidae) with description of a new species, pp. 1-15 in Zootaxa 462 on pages 6-10, DOI: 10.5281/zenodo.157403, {"references":["Gunther, A. (1864) On some new species of Central American fishes. Proceedings of the Zoological Society of London, part 1, 23 - 27.","Palmer, G. (1952) Notes on the fishes of the genus Gobioides, with description of a new species. Annals and Magazine of Natural History, 5, 50 - 57.","Jordan, D. S. & Eigenmann, C. H. (1887) A review of the Gobiidae of North America. Proceedings of the United States National Museum, 9, 477 - 518 [for 1886].","Birdsong, R. S. & Robins, C. R. (1995) New genus and species of seven-spined goby (Gobiidae: Gobiosomini) from the offing of the Amazon River, Brazil. Copeia, 1995, 676 - 683."]}

Akko rossi sp. nov. Fig. 1 Holotype: USNM 371780, 90 mm SL, female, Gulf of Fonseca, El Salvador, trawl, 13��12.89���N, 87��51.29���W to 13��13.74���N, 87��49.81���W, 7.0-9.5 m, mud. Description: Morphometric data for the holotype (the only known specimen) are given in Table 1. Characters given in the Diagnosis are not repeated here. Median fins: The second dorsal and anal fins both have 16 elements, the first element a spine in both. The caudal fin is lanceolate with 17 segmented rays, 4 upper and 5 lower procurrent rays. Median fin rays are serially branched with 4 to 5 branches. Caudal skeleton: Hypurals 1-2 are fused to one another but not to other elements in the caudal skeleton. Hypurals 3-4 are fused to one another and to the terminal half centrum. A small parhypural supports the lowermost segmented ray, hypural 5 is free, and the single epural supports the uppermost segmented ray. Paired fins: The pectoral fin is short, ending posteriorly before the posterior extension of the pelvic fin, well in front of the anus; the pectoral fin has 19 dichotomously branched rays. Pelvic fins are united forming a disk with a well-developed anterior frenum. The pelvic base is formed into a muscular pedicel with strong muscle bundles extending from the vicinity of the pelvic process to the vicinity of the pelvic spine, with some of the fibers inserting on the spines. Rays 1, 2, and 3 are serially branched; rays 4 and 5 dichotomously branched. Head: The mouth is large, the opening at an angle of 50�� to the horizontal. No rostral frenum is present, and the upper jaw is only slightly protrusible. The oral velum has dorsal and ventral projections at the midline. The anterior nostril is a short thickened tube. The posterior nostril is a slit. There is no pseudobranch. The eye is tiny with a large lens occupying most of the eye. The gill opening extends from the level of the sixth pectoral fin ray ventrally to the pelvic-fin base, terminating in a membrane supported by the second branchiostegal ray. The cephalic canal system has a small pore medial and just posterior to the anterior nostril (B���) and a second small pore posterior to the eye (G���). Sensory papillae: The sensory papillae are slightly abraded, and no drawing is included, but the pattern is the same as that of Akko brevis (Fig 2). Horizontal row b has an elongate anterior extension ending under the eye; row d is complete, not divided into two sections, and row ot extends ventrally onto the branchiostegals, ending just past the second branchiostegal ray. The first vertical papillae row is complete, extending from the eye to near row d. Additional vertical rows are divided by row b, with two rows above and four rows below. Only the last vertical row below row b extends below the level of row d. Dorsal rows x1 and x2 are continuous or nearly so as is row n. Jaws: The upper jaw has a single row of large depressible teeth, 16 on each premaxilla in the holotype. The lower jaw has two rows of teeth, the outer row consisting of 11 large evenly spaced teeth, the inner row with smaller widely spaced teeth. Vertebral column: There are 11 precaudal and 17 caudal vertebrae. Two anal-fin pterygiophores precede the anteriormost haemal spine. The dorsal pterygiophore pattern is 3-221110. Scales: Scales are small, cycloid, embedded, and non-overlapping anteriorly, becoming larger and overlapping posteriorly. The body is scaled from the predorsal area over the operculum to the base of the caudal fin, with about 115 scales in the lateral series. Genitalia: The female genital papilla is short and broad, opening through a slit. Pigment: The head and dorsal portion of the trunk are brown in the preserved specimen. The ventral section, below the midline, is pale tan anteriorly, becoming tan to light brown posteriorly. The fins are dark brown, becoming light brown or speckled brown at the base only. The pigment is restricted to the interradial membranes in all fins except the first dorsal, in which the membrane covering the spines is also brown. The genital papilla is brown, the tip and remainder of the surrounding ventral area white. Etymology: The specific epithet is in honor of our friend and colleague, D. Ross Robertson, of the Smithsonian���s Tropical Research Institute in Panama, who has made substantial contributions to our understanding of diversity of tropical eastern Pacific shorefishes. Distribution: The only known specimen of Akko rossi was collected from the Gulf of Fonseca, El Salvador., Published as part of James L. Van Tassell & Carole C. Baldwin, 2004, A review of the gobiid genus Akko (Teleostei: Gobiidae) with description of a new species., pp. 1-15 in Zootaxa 462 on pages 3-6

Akko Birdsong and Robins Diagnosis: Body elongate; vertebrae 11 + 16���17 = 27���28; first dorsal fin with seven spines; dorsal��pterygiophore formula 3�� 221110; second dorsal and anal fins with equal numbers of elements, 15 or 16 present in each; dorsal, caudal, and anal fins continuous, connected by a membrane; upper and lower jaws with enlarged widely spaced teeth which overlap the lips of both jaws; head with very small eyes (2���8 % of head length); mouth large, the opening at an angle of 50 �� to the horizontal; a cephalic lateral��line canal with two pores (B���,G���). Pores very reduced in size and occasionally absent. Sensory papillae pattern in a transverse pattern; horizontal row b elongate, extending to vertical through middle of eye; dorsal rows n elongate, approaching or meeting at midline; rows x 1 and x 2 continuous., Published as part of Van, James L., Tassell & Baldwin, Carole C., 2004, A review of the gobiid genus Akko (Teleostei: Gobiidae) with description of a new species, pp. 1-15 in Zootaxa 462 on page 3, DOI: 10.5281/zenodo.157403

Van, James L., Tassell, Baldwin, Carole C. (2004): A review of the gobiid genus Akko (Teleostei: Gobiidae) with description of a new species. Zootaxa 462: 1-15, DOI: 10.5281/zenodo.157403

[[Akko Birdsong and Robins]] Discussion All species of Akko are easily recognized by their large teeth, elongate body, small eyes, anterior nostril located within a fleshy tube, and by having the dorsal, caudal, and anal fins connected by membrane. The species can be differentiated by numbers of lateral scales, vertebrae, and fin rays, and by color. In A. dionaea the lateral scales do not overlap on the caudal peduncle, and there are approximately 76 in the series; A. brevis and A. rossi both have overlapping scales on the caudal peduncle, with 53-60 and 155 scales, respectively, in the lateral series. Color patterns are distinctive, even in preserved specimens. Akko dionaea has no trunk or head melanophores, and only the distal two-thirds of the caudal fin and posterior portion of the anal fin have some melanophores; A. brevis possesses melanophores on the upper portion of the trunk, on the interradial membranes of the dorsal fin, and on the caudal fin; A. rossi has a heavily pigmented dorsal trunk, and all fins are dark brown. Vertebral counts in A. dionaea and A. brevis are 11 precaudal and 16 caudal, 11+17 in A. rossi; additional specimens of A. rossi are needed to determine if this vertebral count is characteristic for the species. Akko dionaea can be separated from the Pacific species by having fewer pectoral-fin rays: 17 in A. dionaea, 19-21 in A. brevis, and 19 in A. rossi. The second dorsal and anal fins both have 15 elements in A. dionaea and A. brevis but 16 in A. rossi. The pelvic-fin rays are branched in the three species, with A. dionaea and A. brevis having all rays dichotomously branched. In A. rossi, rays 1,2,3 are serially branched, and rays 4,5 are dichotomously branched. Differences that readily separate the three species are given in Table 3. The head pores in all species of Akko are very small and difficult to observe. All species of Akko have a short cephalic head canal with two pores, B��� and G���. Pore G��� was reported as absent by Birdsong and Robins (1995) in A. dionaea, and, although it is absent in the holotype, we have observed it in several of the paratypes. The three species also possess small fleshy flaps on the posterior dorsal edges of the upper and lower lips, a character not mentioned in the original description of A. dionaea. Birdsong and Robins (1995) stated that row ot in A. dionaea has a disjunct extension on the branchiostegals. Our examination of the types of A. dionaea with 0.5mm fiber optic lighting shows this row to be complete, not disjunct, and similar to our Figure 2 of A. brevis. A large ossified lacrimal is present in all species of Akko. In all other Gobiosomatini and most members of the Gobiinae the lacrimal is small. Akko brevis has the ability to open its mouth in a complete circle equal to the body diameter. The large lacrimal, which is attached to the maxilla by a broad tendinous sheath, may play a role in this movement. Akko belongs within the tribe Gobiosomatini based on the presence of seven spines in the first dorsal fin, a dorsal-pterygiophore formula of 3-221110, and a vertebral count of 11 precaudal and 16 caudal vertebrae (17 in rossi). Although Birdsong and Robins (1995) placed Akko within the tribe, they did not comment further on its relationships within the group because of the large number of autapomorphies possessed by A. dionaea. Additional information now available from mtDNA places Akko within the ��� Microgobius ��� group of the Gobiosomatini (Birdsong et al. 1988) and provides support for a sister-group relationship with Microgobius (Ruber et al. 2003). Further supporting the placement of Akko within the ��� Microgobius ��� group are patterns of sensory papillae. There are two general categories of sensory papillae patterns within the Gobiosomatini: In the ��� Gobiosoma ��� group, row n on the dorsal surface of the head is short, the right and left elements never joining at the dorsal midline, and rows x1 and x2 are not connected to form a continuous row (Fig. 5A); in the ��� Microgobius ��� group row n is generally elongate, the left and right elements frequently joining at the dorsal midline, and rows x1 and x2 are united to form a single row (Fig. 5B). This arrangement of sensory papillae has been observed in most species of Microgobius, Bollmannia, and Parrella of the ��� Microgobius ��� group by one of us (JVT). All species of Akko possess a papillae pattern similar to that of the ��� Microgobius ��� group. Myological features, particularly origin and insertion patterns within the adductor mandibulae complex, also support the proposed relationship between Akko and other species of the ��� Microgobius ��� group. In the ��� Gobiosoma ��� group (Fig. 6A) the adductor mandibulae1 complex (A1) originates along the dorsal half of the preopercle and along the lateral edges of the pterotic and sphenotic. The A1 complex separates into A1 beta and A1 alpha sections. An additional separation of the A1, the A1 gamma, occurs in all genera of the ��� Gobiosoma ��� group except Aruma, Barbulifer, Gymneleotris, Chriolepis, Eleotrica, Pycnomma, Gobulus, Nes (Van Tassell, 1998). The A1 gamma inserts on the maxilla and on the A2 gamma, the A1 beta inserts on the maxilla, and the A1 alpha inserts on the coronoid process of the dentary. The adductor mandibulae 2 (A2) originates on the ventral half of the preopercle and on the quadrate. The A2 separates into two sections, one inserting along with the A1 alpha on the coronoid process and the other inserting on the maxilla and A1 gamma, near or at the insertion of the primordial ligament. In the ��� Microgobius ��� group the A1 complex does not originate along the sphenotic, and the A1 gamma is absent. In Bollmannia (Fig. 6B) the A1 originates along the dorsal half of the preopercle and on the pterotic; it inserts via two heads, one on the maxilla, in conjunction with the A1 beta, and the second via a tendon on the coronoid process of the dentary. An A1 beta is present medial to the large A1 alpha, originating on the hyomandibular and inserting with the A1 alpha on the maxilla. In Microgobius there is a partial separation of an A1 beta from A1 alpha anteriorly, just prior to the insertion on the maxilla; otherwise, the A1 remains a single muscle mass. The adductor mandibulae 2 is a single mass originating along the ventral half of the preopercle and on the quadrate. It inserts, along with the A1 alpha, via a tendon onto the dentary. The cheek myology pattern of Akko is most similar to that of the ��� Microgobius ��� group (Fig. 4). Cheek myology of Parrella and Palatogobius in the ��� Microgobius ��� group has not been examined., Published as part of James L. Van Tassell & Carole C. Baldwin, 2004, A review of the gobiid genus Akko (Teleostei: Gobiidae) with description of a new species., pp. 1-15 in Zootaxa 462 on pages 10-14

Akko rossi sp. nov. Fig. 1 Holotype: USNM 371780, 90 mm SL, female, Gulf of Fonseca, El Salvador, trawl, 13 �� 12.89 ���N, 87 �� 51.29 ���W to 13 �� 13.74 ���N, 87 �� 49.81 ���W, 7.0��� 9.5 m, mud. Description: Morphometric data for the holotype (the only known specimen) are given in Table 1. Characters given in the Diagnosis are not repeated here. Median fins: The second dorsal and anal fins both have 16 elements, the first element a spine in both. The caudal fin is lanceolate with 17 segmented rays, 4 upper and 5 lower procurrent rays. Median fin rays are serially branched with 4 to 5 branches. Caudal skeleton: Hypurals 1���2 are fused to one another but not to other elements in the caudal skeleton. Hypurals 3���4 are fused to one another and to the terminal half centrum. A small parhypural supports the lowermost segmented ray, hypural 5 is free, and the single epural supports the uppermost segmented ray. Paired fins: The pectoral fin is short, ending posteriorly before the posterior extension of the pelvic fin, well in front of the anus; the pectoral fin has 19 dichotomously branched rays. Pelvic fins are united forming a disk with a well��developed anterior frenum. The pelvic base is formed into a muscular pedicel with strong muscle bundles extending from the vicinity of the pelvic process to the vicinity of the pelvic spine, with some of the fibers inserting on the spines. Rays 1, 2, and 3 are serially branched; rays 4 and 5 dichotomously branched. A. rossi A. brevis males (n= 12) A. brevis females (n= 9) Head: The mouth is large, the opening at an angle of 50 �� to the horizontal. No rostral frenum is present, and the upper jaw is only slightly protrusible. The oral velum has dorsal and ventral projections at the midline. The anterior nostril is a short thickened tube. The posterior nostril is a slit. There is no pseudobranch. The eye is tiny with a large lens occupying most of the eye. The gill opening extends from the level of the sixth pectoral fin ray ventrally to the pelvic��fin base, terminating in a membrane supported by the second branchiostegal ray. The cephalic canal system has a small pore medial and just posterior to the anterior nostril (B���) and a second small pore posterior to the eye (G���). Sensory papillae: The sensory papillae are slightly abraded, and no drawing is included, but the pattern is the same as that of Akko brevis (Fig 2). Horizontal row b has an elongate anterior extension ending under the eye; row d is complete, not divided into two sections, and row ot extends ventrally onto the branchiostegals, ending just past the second branchiostegal ray. The first vertical papillae row is complete, extending from the eye to near row d. Additional vertical rows are divided by row b, with two rows above and four rows below. Only the last vertical row below row b extends below the level of row d. Dorsal rows x 1 and x 2 are continuous or nearly so as is row n. Jaws: The upper jaw has a single row of large depressible teeth, 16 on each premaxilla in the holotype. The lower jaw has two rows of teeth, the outer row consisting of 11 large evenly spaced teeth, the inner row with smaller widely spaced teeth. Vertebral column: There are 11 precaudal and 17 caudal vertebrae. Two anal��fin pterygiophores precede the anteriormost haemal spine. The dorsal pterygiophore pattern is 3�� 221110. Scales: Scales are small, cycloid, embedded, and non��overlapping anteriorly, becoming larger and overlapping posteriorly. The body is scaled from the predorsal area over the operculum to the base of the caudal fin, with about 115 scales in the lateral series. Genitalia: The female genital papilla is short and broad, opening through a slit. Pigment: The head and dorsal portion of the trunk are brown in the preserved specimen. The ventral section, below the midline, is pale tan anteriorly, becoming tan to light brown posteriorly. The fins are dark brown, becoming light brown or speckled brown at the base only. The pigment is restricted to the interradial membranes in all fins except the first dorsal, in which the membrane covering the spines is also brown. The genital papilla is brown, the tip and remainder of the surrounding ventral area white. Etymology: The specific epithet is in honor of our friend and colleague, D. Ross Robertson, of the Smithsonian���s Tropical Research Institute in Panama, who has made substantial contributions to our understanding of diversity of tropical eastern Pacific shorefishes. Distribution: The only known specimen of Akko rossi was collected from the Gulf of Fonseca, El Salvador., Published as part of Van, James L., Tassell & Baldwin, Carole C., 2004, A review of the gobiid genus Akko (Teleostei: Gobiidae) with description of a new species, pp. 1-15 in Zootaxa 462 on pages 3-6, DOI: 10.5281/zenodo.157403

Pseudogramma xanthum, nomen novum (Figure 1, Table 1) Pseudogramma australis australis Randall and Baldwin, 1997 (type locality Temoe Atoll). Holotype: BPBM 13531, 39.1 mm, Tuamotu Archipelago, Gambier Group, Temoe Atoll (23��20���S, 134��30���W), N side of seaward reef, small cave, 41 m, rotenone, J.E. Randall and D.B. Cannoy, 16 December 1970. Additional material examined: BMNH 1996.6.27.1, 41.0 mm, Pitcairn Island, coral rubble and stone bottom, 99-102 m, station 6, dredge haul 17, crew of ���Pele���, 20 October 1967; BPBM 37393, 4:22.4-35.0 mm, same data as holotype; BPBM 16893, 33.1 mm, Pitcairn Island, N side off Gannet Ridge, coral reef, 40-44.5 m, rotenone, J.E. Randall, D.B. Cannoy, J.R. Haywood, R.R. Costello, J.D. Bryant, and S.R. Christian, 6 January 1971; BPBM 13974, 2: 22.5-26.0 mm, Cook Islands, Rarotonga, off oil tanker buoy near harbor entrance, 23-30.5 m, rotenone, J.E. Randall and D.B. Cannoy, 11 March 1971; USNM 327795, 3: 26-27 mm, Tonga, Ha���apai Group, Ofolanga Island, SW side (19��36���15���S, 174��28���15���W), deep reef slope with sand channel at base and steep dropoff at end, 21-32 m, rotenone, J.T. Williams, B.B Collette, G.D. Johnson, D.G. Smith, C.C. Baldwin, and E.A. Powers, 12 November 1993. Diagnosis: Dorsal rays VII, 19-21; anal rays III, 16-18 (usually 17); pectoral rays 14- 15 (usually 15); one incomplete lateral line, pored scales 28-32; longitudinal scale series 50-52; gill rakers 6 + 10 or 11; a slender triangular dermal flap dorsally on eye of adults; tubular anterior nostril reaching about half way to posterior nostril when laid back; head length 40.3-43.1% SL; snout length 7.5-7.9% SL; orbit diameter 9.0-10.2% SL; upper-jaw length 19.3-21.1% SL; predorsal length 38.2-40.3% SL; head and body yellow in life with rows of large white blotches, a pale-edged brown spot on opercle; dark red dots on head, many in rows, and scattered dark red dots on body; largest specimen, 39.1mm SL. Description: Dorsal rays VII, 20 (19-21); anal rays III, 17 (16-18); pectoral rays 15 (14-15), all rays branched in adults; pelvic rays I, 5; principal caudal rays 17; upper procurrent caudal rays 4; lower procurrent caudal rays 3; one incomplete lateral line, the pored scales 30 (28-32); longitudinal scale series 50 (about 50-52); scale rows directly above lateral line to origin of dorsal fin about 6; scale rows directly below lateral line to origin of anal fin about 17 (rows partly overlapping and difficult to count); gill rakers 6 + 11 (6 + 11 or 12); pseudobranchial filaments 10 (7-9); branchiostegal rays 7; vertebrae 10+16. Body depth 3.55 (3.6-4.25) in SL; body width 1.8 (1.6-2.0) in body depth; head length 2.45 (2.3-2.5) in SL; snout length 5.4 (5.3-5.6) in head length; orbit diameter 4.3 (4.1-4.6) in head length; interorbital narrow and nearly flat, the least fleshy width 11.5 (10.9-12.5) in head length; caudal-peduncle depth 3.2 (3.25-3.5) in head length; caudal-peduncle length 5.35 (5.25-5.45) in head length. Mouth terminal or with lower jaw slightly projecting, oblique, forming an angle of about 35�� to horizontal axis of head, and large, the maxilla extending well beyond a vertical at rear edge of eye, the upper-jaw length 2.1 (1.95-2.15) in head length; a small, triangular, ventrally-projecting, bony process posteriorly on maxilla; a small fixed incurved canine tooth (sometimes a close-set pair) anteriorly on each side of upper jaw; an inner band of small, conical, inwardly depressible teeth at front of upper jaw in about 6 irregular rows, progressively longer medially, the innermost symphyseal teeth longer than canines, though more slender; band of teeth narrowing to 1 or 2 irregular rows posteriorly in jaw; dentition of lower jaw similar to upper but without the fixed canines, with only 4-5 rows of teeth anteriorly, the largest teeth of inner row continuing along side of jaw; vomer with a V-shaped patch of small conical teeth in 3 irregular rows; palatines with a long band of small conical teeth in 1-2 irregular rows. Tongue long and slender, reaching slightly anterior to vomerine teeth. Opercle with 3 flat spines covered by scales; posterior margin of preopercle with a strong spine projecting obliquely downward, its base at level of lower edge of eye. A slender triangular flap longer than pupil extending upward from dorsal surface of eye (small and more triangular on smallest paratypes). Tubular anterior nostril near base of upper lip at level of lower edge of pupil, reaching about half distance to posterior nostril when laid back; posterior nostril at edge of orbit, slightly dorsal to anterior nostril, and vertically oval with a slight rim. A pair of large pores in midinterorbital space with a median pore behind; 12 other large pores encircling orbit; 4 large pores on mandible continuing as 7 pores along edge of preopercle. Scales on body small and adherent, their surface with prominent horizontal ridges, those posterior to about tenth lateral-line scale rhomboid and ctenoid, the middle cteni longest; small cycloid scales on nape extending into posterior interorbital space; small cycloid scales on opercle and cheek; rest of head naked; small scales on about basal half of fins except pelvics where only basally; no scales in axil of pectoral fins; lateral line paralleling dorsal contour of body, ending below middle of base of soft portion of dorsal fin. Origin of dorsal fin above fifth lateral-line scale, the predorsal length 2.6 (2.5-2.6) in SL; first dorsal spine 5.6 (5.4-5.75) in head length; fifth or sixth dorsal spines longest, 4.0 (3.8-3.95) in head length; last dorsal spine 4.95 (4.5-4.85) in head length; longest dorsal soft ray 2.85 (2.6-2.75) in head length; origin of anal fin below base of third dorsal soft ray, the preanal length 1.5 (1.5-1.55) in SL; first anal spine 6.9 (7.l-8.55) in head length; second anal spine clearly largest, 4.7 (4.2-4.8) in head length; longest anal soft ray 2.8 (2.55-2.8) in head length; caudal fin rounded and small, 1.8 (1.6-1.5) in head length; middle pectoral rays longest, 1.3 (1.2-1.3)in head length; origin of pelvic fins anterior to base of pectorals, the prepelvic distance 2.95 (2.8-3.1) in SL; second and third pelvic soft rays longest, 2.65 (2.6-2.7) in head. Color of holotype entirely pale yellowish. Color when fresh light yellow, shading to white on abdomen, chest, and ventral half of head, with two rows of indistinct pale blotches larger than pupil, one positioned midlaterally and one on lower side, scattered dark red dots (red with dark brown centers) on body; opercle with irregular pale-edged brown blotch rimmed with dark red dots; numerous dark red dots on dorsal half of head, those on lower half of head more widely separated but most arranged in lines, four of which pass ventrally from anterior half of eye, the first three crossing lips onto chin; fins light yellowish, the dorsal with a few dark red dots on base. A color photo (by JTW) of a specimen from Tonga is essentially the same as that of the holotype (Fig. 1) from the Tuamotu Archipelago. Etymology: The specific epithet is derived from the Greek adjective xanthos, meaning yellow, in reference to the ground color of the body. Remarks: The species redescribed above was first treated as a subspecies of Pseudogramma australis, but was mistakenly named P. a. australis. The name P. australis has been shown above to be occupied by the Easter Island species and does not apply to the western species. The subspecies P. a. pasquensis is a junior synonym of P. australis. Thus there is no name available for the western species of the P. australis group, and we provide P. xanthum as the new replacement name for P. a. australis. Pseudogramma australis and P. xanthum have the same basic color pattern, but differ in ground color, red for the former and yellow for the latter. However, we have records of the life color of P. xanthum from only Temoe and Tonga. In addition to the meristic differences between P. xanthum and P. australis discussed for the two subspecies of australis in Randall & Baldwin (1997: tables 1-3), and the smaller size of xanthum, we note the following differences in proportional measurements: P. xanthum has a larger head (40.7- 43.1% SL, compared to 37.0-39.4% for P. australis); longer snout (7.5-7.9% SL, compared to 6.7-7.4% for P. australis); longer upper jaw (19.3-21.1% SL, compared to 18.5- 19.7%); and longer predorsal length (38.2-40.3% SL, compared to 36.4-38.2%). The eye size of P. xanthum is larger (9.0-10.2% SL, compared to 8.4-9.1%), but this is probably due to the smaller size of the specimens of P. xanthum. From the tables, P. xanthum seems to have a shorter caudal fin, but only three specimens could be measured due to the poor condition of most specimens., Published as part of John E. Randall, Carole C. Baldwin & Jeffrey T. Williams, 2002, Pseudogramma xanthum, a new replacement name for a serranid fish from the subtropical South Pacific Ocean with description of the species., pp. 1-8 in Zootaxa 40 on pages 3-7