Research Article |
Corresponding author: Carole C. Baldwin ( baldwinc@si.edu ) Academic editor: Nina Bogutskaya
© 2015 Carole C. Baldwin, D. Ross Robertson.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Baldwin CC, Robertson DR (2015) A new, mesophotic Coryphopterus goby (Teleostei, Gobiidae) from the southern Caribbean, with comments on relationships and depth distributions within the genus. ZooKeys 513: 123-142. https://doi.org/10.3897/zookeys.513.9998
|
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.
Coryphopterus curasub , Coryphopterus dicrus , submersible, Substation Curaçao, Deep Reef Observation Project (DROP), DNA barcoding, phylogeny
Coryphopterus gobies live in mostly shallow warm waters of the western Atlantic and eastern Pacific Oceans, dwelling on sand around coral and rocky reefs or hovering above or perching on reef structures. Twelve species are known from the western Atlantic and one from the eastern Pacific (
A new western Atlantic species, Coryphopterus kuna Victor, was described in 2007 but not included in the molecular phylogeny of
Four specimens of the new species and three of C. venezuelae were collected using Substation Curaçao’s (http://www.substation-Curacao.com) manned submersible Curasub. The sub has two flexible, hydraulic arms, one of which is equipped with a quinaldine-ejection system and the other with a suction hose. Anesthetized fish specimens were captured with the suction hose, which empties into a vented plexiglass cylinder attached to the outside of the sub. At the surface, the specimens were photographed, tissue sampled, and preserved. Preserved specimens were later photographed to document preserved pigment pattern and X-rayed with a digital radiography system. Counts and measurements follow
Tissue samples for DNA Barcoding were stored in saturated salt-DMSO (dimethyl sulfoxide) buffer (Seutin et al. 1991). DNA extraction, PCR, sequencing cytochrome c oxidase subunit I (COI), and editing COI sequences were performed as outlined by
GenSeq nomenclature (
Curaçao, southern Caribbean
USNM 406373, Smithsonian DNA number CUR 11373, 33.3 mm SL, female, Curasub submersible, sta. 11-05, southern Caribbean, Curaçao, east of downline off Substation Curaçao dock, near 12°05.069'N, 68°53.886'W, 80 m, quinaldine, 30 May 2011, D. R. Robertson, B. Brandt, A Schrier, K. Stewart.
USNM 430037, CUR 13302, 30.0 mm SL, male, Curasub submersible, sta. 13-29, southern Caribbean, Curaçao, east of downline off Substation Curaçao dock, near 12°05.069'N, 68°53.886'W, 70–72 m, quinaldine, 30 October 2013, C. C. Baldwin, D. R. Robertson, B. Brandt, C. Castillo, L. Ybarrondo. USNM 431328, CUR 14003, 31.0 mm SL, male, Curasub submersible, sta. 14-01, southern Caribbean, Curaçao, east of downline off Substation Curaçao dock, near 12°05.069'N, 68°53.886'W, 73 m, quinaldine, 17 March 2014, C. C. Baldwin, D. R. Robertson, B. Brandt, C. Castillo, H. Reichardt. USNM 430019, CUR 13303, 17.5 mm SL, immature (same collection locality as USNM 430037), cleared and stained.
The combination of six spines in the first dorsal fin, fewer than 20 rays in the second dorsal fin, pelvic fin with one spine and five soft rays, head pores present, no free pectoral-fin rays, no scales on top of head, and no prominent crest on top of head anteriorly from first dorsal fin support the placement of C. curasub in the genus Coryphopterus (
A species of Coryphopterus distinguishable from its congeners by the following combination of characters: total second dorsal-fin rays (spinous plus soft) 10-11, usually 11; total anal-fin rays (spinous plus soft) 10; pectoral-fin rays 19-20; pelvic fins united; no pelvic frenum; pectoral-fin base with two prominent dark spots (yellow with dark spotting in life) of roughly equal intensity, one on dorsal portion of fin base and one on ventral part; no distinct black blotch behind orbit above opercle; no dark triangular blotch immediately behind middle of orbit; blotches of pigment on trunk mostly yellow; few melanophores and yellow dots interspersed among yellow blotches of pigment on trunk; no black ring of pigment surrounding anus; dark triangular blotch variously developed beneath anteroventral corner of orbit; basicaudal blotch cross-shaped, with prominent anterior projection; and two yellow/orange blotches on base of caudal fin situated immediately behind basicaudal blotch.
Dorsal-fin rays VI + I, 9-10 (9 in one paratype, 10 in other specimens), total second dorsal-fin rays 10 or 11 (10 in one paratype, 11 in other specimens); anal-fin rays I, 9; all soft dorsal- and anal-fin rays branched. Pectoral-fin rays 19, 19 (paratypes) or 20, 20 (holotype); all pectoral rays branched except splint-like uppermost and lowermost rays. Pelvic-fin rays I, 5; all soft rays branched; fins united, no frenum. Total caudal-fin rays (including procurrent rays) 30 (holotype) or 31 (cleared and stained paratype; can’t assess number from radiographs of other paratypes); segmented caudal rays17; branched caudal rays12 (6+6); unbranched caudal rays 18 in holotype (9+9), 19 (10+9) in cleared and stained paratype. Dorsal-fin formula 3-22110. Vertebrae 10 precaudal +16 caudal. Epineurals 10 pairs. Ribs on vertebrae 3-10. Anal-fin pterygiophores anterior to first haemal spine 2. Gill rakers on first arch, including rudiments, 2 + 8 (holotype and adult paratypes), 0 + 8 in cleared and stained juvenile paratype. Branchiostegal rays 5. Numerous scales abraded and missing on all specimens, one paratype with approximately 22 scales in longitudinal series, 6 scales between origin of second dorsal fin obliquely downward to anal fin, and approximately 17 circum-peduncular scales.
Measurements of holotype in parentheses following extremes for holotype plus two adult paratypes. Juvenile paratype bent and not measured prior to clearing and staining. Body elongate, maximum depth from base of dorsal-fin spines 17–19% SL (17); body compressed, greatest width just posterior to gill opening 14–15% SL (14); head length 30–32% SL (32); snout length (to fleshy edge of orbit) 6.0–7.1% SL (6.0); greatest fleshy orbit diameter 9.3–10% SL (9.3); least fleshy interorbital 1.3–3.3% SL (3.3); caudal-peduncle length 23–24% SL (23); least caudal-peduncle depth 11–12% SL (11); length of dorsal-fin base 39–46% SL (46); first dorsal spine 14–16% SL (16); second dorsal spine 15–18% SL (17); third and longest dorsal spine 16–20% SL (18); sixth and shortest dorsal spine 7.0–8.4% SL (8.4); seventh dorsal spine (first element of second dorsal fin) 14–16% SL (16); last dorsal soft ray 16–17% SL (16); anal-fin spine 8.7–11% SL (11); last anal soft ray 20–22% SL (20); pectoral fin 32–36% SL (36), fin reaching vertical through second anal-fin soft ray, longest pectoral rays the 11th-14th rays from top of fin; pelvic fin 24–32% SL (27), fin reaching origin of anal fin or terminating slightly before origin, longest pelvic ray the fourth.
Trunk, belly, and pre-pelvic region scaled, head and predorsal region naked; scales ctenoid except on pre-pelvic region, where they are cycloid. Upper jaw with several rows of small conical teeth, outermost teeth largest but smaller than outermost dentary teeth; dentary with outer row of fairly large conical teeth and several inner rows of smaller teeth; innermost teeth intermediate in size between teeth of outermost row and those adjacent to it. No teeth on vomer, palatines, or pterygoids. Anterior nare opening on short tube; posterior nare a simple opening. Head pores prominent: nasal pore, anterior interorbital pore, posterior interorbital pore, infraorbital pore, postorbital pore, pore at each end of lateral sensory canal, pore at each end of posterior lateral canal, and three preopercular pores (pores B’, C and D [both single], E, F, G, H’, K’, L’, M’, N, O’). A very low, thin ridge of tissue extending from just posterior to interobital region to base of first dorsal fin.
When photographed against a light background (Fig.
Coryphopterus curasub sp. n., type specimens: A, B USNM 406373, holotype, Smithsonian DNA number CUR 11373, 33.3 mm SL, female – after preservation (A) and before preservation (B) C USNM 431328, Paratype, CUR 14003, 31.0 mm SL, male, before preservation D USNM 430019, Paratype, CUR 13303, 17.5 mm SL, immature, before preservation and clearing and staining. Note that the dark color on the posterior portion of the caudal fin is an artifact of flash photography and does not reflect the existence of dark pigment. Photos by Ian Silver-Gorges (A) and D. R. Robertson and C. C. Baldwin (B–D).
Male paratypes (Fig.
Color of holotype in alcohol (Fig.
Known from 70–80 m off Curaçao, southern Caribbean.
Notes recorded during the submersible dive on which the 33.0 mm SL paratype (USNM 431328) was collected indicate that it occurred on sand with rubble patches on a 45°slope.
Named for the manned submersible Curasub, which is owned and operated by Substation Curaçao, in recognition of the contributions of this vehicle to increasing our knowledge of the Caribbean deep-reef fish fauna.
“Yellow-spotted sand goby” refers to the yellow spots on the trunk and the collection habitat.
Coryphopterus curasub is most similar to C. dicrus (Fig.
Comparison of A Coryphopterus curasub sp. n., holotype, USNM 406373, 33.3 mm SL, and its most similar congener B Coryphopterus dicrus, USNM 413296, 30 mm SL. Note the differences in the shape of the basicaudal pigment marking (with distinct anterior projection in C. curasub), body depth (shallower in C. curasub), head pigment (absence of a distinct blotch of black pigment immediately posterior to the orbit and presence of a black triangle of pigment beneath the anteroventral portion of orbit in C. curasub (present and absent, respectively, in C. dicrus), and trunk pigment (blotches predominantly yellow with few melanophores interspersed among them in C. curasub vs. blotches predominantly orange/rust with numerous melanophores interspersed among them in C. dicrus).
Of the 14 apomorphic morphological characters of Coryphopterus species tabulated by
COI sequences derived from tissue samples from the four type specimens of C. curasub and three specimens of C. venezuelae collected by submersible as part of this study (
The strict consensus of a maximum parsimony analysis of the COI region of 42 individuals of Coryphopterus and Lophogobius cyprinoides. Fusigobius duospilus and Rhinogobiops nicholsii were outgroups in the analysis. Numbers above branches represent bootstrap support values > 50. Note: C. punctipectophorus from the Gulf of Mexico was not available for inclusion in this analysis.
Average Kimura two–parameter distance summary for species of western Atlantic Coryphopterus based on cytochrome c oxidase I (COI) sequences of individuals represented in the neighbor–joining tree in Figure
curasub | dicrus | lipernes | hyalinus | personatus | tortugae | glaucofraenum | venezuelae | thrix | eidolon | alloides | kuna | |
---|---|---|---|---|---|---|---|---|---|---|---|---|
C. curasub (n=4) | 0.10 | |||||||||||
C. dicrus (n=24) | 18.00 | 0.60 | ||||||||||
C. lipernes (n=7) | 19.40 | 21.70 | 0.10 | |||||||||
C. hyalinus (n=11) | 18.00 | 19.60 | 14.90 | 0.10 | ||||||||
C. personatus (n=11) | 19.00 | 19.00 | 15.60 | 7.20 | 0.10 | |||||||
C. tortugae (n=21) | 17.40 | 17.50 | 19.60 | 21.10 | 20.10 | 0.20 | ||||||
C. glaucofraenum (n=26) | 17.80 | 20.60 | 20.70 | 21.70 | 21.50 | 12.10 | 0.20 | |||||
C. venezuelae (n=39) | 17.40 | 18.30 | 21.40 | 20.90 | 20.10 | 9.90 | 9.50 | 0.40 | ||||
C. thrix (n=7) | 22.90 | 21.30 | 21.90 | 21.10 | 19.70 | 19.00 | 21.20 | 19.60 | 0.20 | |||
C. eidolon (n=21) | 22.90 | 19.40 | 25.20 | 19.20 | 18.00 | 19.70 | 12.10 | 18.90 | 19.60 | 0.10 | ||
C. alloides (n=4) | 17.90 | 17.90 | 21.90 | 17.90 | 18.40 | 20.00 | 21.30 | 18.90 | 20.20 | 18.50 | 3.70 | |
C. kuna (n=5) | 22.70 | 24.90 | 26.40 | 23.20 | 25.70 | 25.50 | 27.90 | 26.00 | 24.90 | 25.80 | 23.60 | 0.50 |
Depth ranges of Coryphopterus species are shown in Figure
Depth ranges for Coryphopterus species. Data are from
(Modified from
1 | Black ring surrounding anus; pelvic fins separate, frenum absent | 2 |
– | No black ring around anus; pelvic fins separate or fused, frenum present or absent | 4 |
2 | One anterior interorbital pore | 3 |
– | Two anterior interorbital pores | Coryphopterus hyalinus |
3 | Total second dorsal-fin rays (spinous plus soft) typically 11; total anal-fin rays (spinous plus soft) typically 11; in life, head with orange pigment, body translucent with several square- or rectangular-shaped orange blotches internally; preserved specimens lacking conspicuous postorbital stripes of melanophores but with dark “mask” around eye | Coryphopterus personatus |
– | Total second dorsal-fin rays (spinous plus soft) typically 10; total anal-fin rays (spinous plus soft) typically 10; in life, head and body predominantly yellow, with blue-white stripes extending posteriorly from dorsal and ventral portions of orbit; a dusky internal stripe along posterior section of vertebral column; preserved specimens with postorbital stripes of melanophores and scattered spots over entire body | Coryphopterus lipernes |
4 | No distinct black blotch behind eye above opercle in adults; pigment mark above opercle, if present, no larger or darker than other marks behind eye; pelvic fins separate or fused, frenum present or absent (see |
5 |
– | Distinct black blotch or triangle behind eye above opercle in adults, blotch usually larger and darker than other pigment in stripe behind eye; pelvic fins fused to form disc, frenum present (see |
11 |
5 | Total anal-fin rays (spinous plus soft) 8–9 (usually 9), pectoral-fin rays 15–17, pelvic fins separate or fused, frenum absent | 6 |
– | Total anal-fin rays (spinous plus soft) 10–11, pectoral-fin rays 17– 20, pelvic fins fused, frenum present or absent | 7 |
6 | Total second dorsal-fin rays (spinous plus soft) 9; total anal-fin rays (spinous plus soft) 9; pectoral-fin rays 15; first dorsal fin with stripe of black pigment; in life, head and body with orange spots and blotches, sometimes a flag of dark pigment on 1st–3rd dorsal spines; pelvic fins fused to form a disc | Coryphopterus kuna |
– | Total second dorsal-fin rays (spinous plus soft) 10; total anal-fin rays (spinous plus soft) 9 (rarely 8); pectoral-fin rays 16–17; black blotch or bar between 2nd and 3rd dorsal spines; in life, head and anterior portion of body mottled orange, posterior portion of body mottled yellow; pelvic fins separate | Coryphopterus alloides |
7 | Pectoral-fin base with two prominent dark spots of equal intensity, one dorsally and one ventrally; upper spot usually with swath of melanophores extending posteriorly onto pectoral-fin rays; sides of body freckled with scattered large and small blotches of melanophores (blotches associated with orange, rust, or yellow pigment in life); pelvic frenum absent | 8 |
– | Pectoral-fin base with or without two prominent dark spots; if two spots present, upper spot more intense; sides of body with few dark markings (with few to many yellow spots in life) or with three rows of light markings (coral pink/orange in life); pelvic frenum present | 9 |
8 | Total second dorsal-fin rays (spinous plus soft) 10; a dark triangle immediately behind orbit; no dark triangle under front of orbit; basicaudal mark a vertical dumbbell that in life incorporates two large orange spots on the base of the caudal fin; maximum body depth beneath spinous dorsal fin 20–26% SL, least depth of caudal peduncle 13–15% SL, depth range 0–56 m | Coryphopterus dicrus |
– | Total second dorsal-fin rays (spinous plus soft) 10 or 11, usually 11; no dark triangle behind orbit; dark triangle variously developed beneath front part of orbit; basicaudal blotch cross-shaped, the anterior horizontal projection prominent; in life, two large yellow spots on base of caudal fin just posterior to basicaudal blotch; maximum body depth beneath spinous dorsal fin 17–19% SL; least depth of caudal peduncle 9–12% SL; depth range 70–80 m | Coryphopterus curasub sp. n. |
9 | Pectoral-fin base without prominent dark markings, but may have scattered melanophores; sides of body with few if any dark markings (with yellow to orange spots and stripes in life) except for several dark streaks internally along spinal cord and a thin dark basicaudal bar | Coryphopterus eidolon |
– | Pectoral-fin base with prominent markings; sides of body with or without numerous dark markings | 10 |
10 | Pectoral-fin base with distinct, often large, pigment spot dorsally, spot usually dark above, diffuse below, often with dots trailing ventrally; ventral dots coalescing into a separate spot in some specimens (ventral spot, if present, less intense than dorsal spot); total second dorsal-fin rays (spinous plus soft) 9–10; second dorsal spine filamentous; in life, first dorsal fin without orange stripes | Coryphopterus thrix |
– | Pectoral-fin base with prominent dark spot or blotch ventrally; total second dorsal-fin rays (spinous plus soft) 11; second dorsal spine not filamentous; in life, first dorsal fin with two broad orange stripes | Coryphopterus punctipectophorus |
11 | Body usually pale, pigment primarily comprising three rows of markings on side of body; lower row comprising small, mostly vertically elongate markings, some of which may be crescent shaped or some part of an X-shape but rarely well-defined X’s; height of any X-shaped markings considerably less than eye diameter; pigment mark above opercle usually a triangle, and basicaudal pigment usually a central bar | Coryphopterus tortugae |
– | Body heavily pigmented or pale but no vertically elongate or crescent-shaped markings in ventral row of pigment on side of body; height of any X-shaped markings three-quarters of or equal to eye diameter; pigment mark above opercle triangular, rounded, or with two peaks; basicaudal pigment variable: two separate spots, a vertical dumbbell, a central bar, or a C-shaped marking | 12 |
12 | Pectoral-fin base with dark spot or rectangle ventrally (may be associated with bright yellow pigment in life); one or two bars or blotches sometimes present dorsally; three rows of dark markings on side of body, some in lower row large, X-shaped markings in heavily pigmented specimens, small, circular blotches in paler specimens; pigment mark above opercle triangular or round | Coryphopterus venezuelae |
– | Pectoral-fin base rarely with prominent dark marking ventrally (may have one to three light to moderate concentrations of melanophores); body with three rows of dark marks, most of those in the lower row large and distinctively X-shaped; pigment mark above opercle usually with two well-defined peaks | Coryphopterus glaucofraenum |
Exploratory submersible diving to 300 m off Curaçao is resulting in the discovery of numerous new fish species, only a few of which have been described to date (
For contributing in various ways to this project, we thank the following (in alphabetical order): Bruce Brandt, Barry Brown, Cristina Castillo, Amy Driskell, Tico Christiaan, Rob Loendersloot, Dan Mulcahy, Diane Pitassy, Sandra Raredon, Laureen Schenk, Adriaan Schrier, Ian Silver-Gorges, Jennifer Strotman, Laura Tancredi, Barbara van Bebber, and Lee Weigt. Funding for the Smithsonian Institution’s Deep Reef Observation Project was provided internally by the Consortium for Understanding and Sustaining a Biodiverse Planet to CCB, the Competitive Grants for the Promotion of Science program to CCB and DRR, and the Herbert R. and Evelyn Axelrod Endowment Fund for systematic ichthyology to CCB; and externally by National Geographic Society’s Committee for Research and Exploration to CCB (Grant #9102-12). This is Ocean Heritage Foundation/Curacao Sea Aquarium/Substation Curacao (OHF/SCA/SC) contribution number 17.
Links between DNA voucher specimens, GenBank accession numbers, and cytochrome c oxidase subunit I (COI) sequences of Coryphopterus curasub sp. n. and C. venezuelae.
Catalog Number/DNA Number | GenBank No. | GenSeq Designation |
---|---|---|
Coryphopterus curasub sp. n. | ||
USNM 406373, CUR 11373, Holotype | KT020955 | Geneseq-1 COI |
USNM 430037, CUR 13302, Paratype | KT020957 | Genseq-2 COI |
USNM 431328, CUR 14003, Paratype | KT020961 | Genseq-2 COI |
USNM 430019, CUR 13303, Paratype | KT020958 | Genseq-2 COI |
Coryphopterus venezuelae | ||
USNM 413804, CUR 12273 | KT020959 | Genseq-4 COI |
USNM 413992, CUR 13008 | KT020956 | Genseq-4 COI |
USNM 430016, CUR 13328 | KT020960 | Genseq-4 COI |