Coryphopterus kuna, a new goby (Perciformes: Gobiidae: Gobiinae) from the western Caribbean, with the identification of the late larval stage and an estimate …

Abstract

A new goby, Coryphopterus kuna, is described from the Atlantic coasts of Panama and Mexico. The species is distinguished from other Coryphopterus spp. by the low median fin and pectoral fin ray counts and the morphology of the pelvic fin. The pelvic fins are fully joined with a rounded outline and have branched and longer innermost pelvic fin rays. There is no frenum connecting the two pelvic fin spines and the fin is heavily speckled with black spots in the male holotype. The late larval stage of C. kuna is identified by DNA sequence matching and is morphologically similar to other larval Coryphopterus spp. but has a distinct melanophore pattern. Examination of the otolith microstructure reveals a relatively long pelagic larval duration of 63 days with a narrowing of the later daily increments suggesting delayed metamorphosis. The species is the first vertebrate to include gene sequence barcoding under the Barcode of Life Data System (BOLD) in the species description.

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Accepted by L. Rocha: 13 Jun. 2007; published: 12 Jul. 2007 51
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2007 · Magnolia Press
Zootaxa 1526: 51–61 (2007)
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Coryphopterus kuna, a new goby (Perciformes: Gobiidae: Gobiinae) from the
western Caribbean, with the identification of the late larval stage and
an estimate of the pelagic larval duration
BENJAMIN C. VICTOR
Ocean Science Foundation, 4051 Glenwood, Irvine, CA 92604 and Guy Harvey Research Institute, Oceanographic Center, Nova
Southeastern University, 8000 North Ocean Drive, Dania Beach, FL 33004. E-mail: ben@coralreeffish.com
Abstract
A new goby, Coryphopterus kuna, is described from the Atlantic coasts of Panama and Mexico. The species is distin-
guished from other Coryphopterus spp. by the low median fin and pectoral fin ray counts and the morphology of the pel-
vic fin. The pelvic fins are fully joined with a rounded outline and have branched and longer innermost pelvic fin rays.
There is no frenum connecting the two pelvic fin spines and the fin is heavily speckled with black spots in the male holo-
type. The late larval stage of C. kuna is identified by DNA sequence matching and is morphologically similar to other
larval Coryphopterus spp. but has a distinct melanophore pattern. Examination of the otolith microstructure reveals a rel-
atively long pelagic larval duration of 63 days with a narrowing of the later daily increments suggesting delayed meta-
morphosis. The species is the first vertebrate to include gene sequence barcoding under the Barcode of Life Data System
(BOLD) in the species description.
Key words: Gobiidae, Goby, New Species, BARCODE, BOLD, Fish, Informatics, Larvae, Larval Identification, DNA,
Larval Stage, Pelagic Larval Duration, Otolith, Panama, Mexico, Caribbean, Western Atlantic
Introduction
Although a number of gobioid species have been recently described from both coasts of the Americas, the
genus Coryphopterus in the New World has seen few changes since the original treatment by Bohlke and Rob-
ins in 1960 and 1962. They listed nine Atlantic species and two eastern Pacific species. Since their treatise,
Coryphopterus venezuelae (Cervig n) has been described from Venezuela (Cervig n 1966, 1994) and Cory-
phopterus tortugae (Jordan) has been redescribed and is widespread in the Caribbean (Garzon-Ferreira and
Acero 1990, Greenfield & Johnson 1999). A number of new Indo-Pacific Coryphopterus spp. have been
described by Randall (2001) who provided a key to the species for that region. However, the validity of
including the Indo-Pacific species in this genus has been questioned recently by Thacker and Cole (2002) and
they suggest that those species be returned to Fusigobius spp. Furthermore, they found the one temperate spe-
cies (from the eastern Pacific) to be unrelated and returned it to Rhinogobiops nicholsii (Bean). In this paper I
describe a new Atlantic species from the western Caribbean, Coryphopterus kuna. Individuals of this species
have been found as an adult in Panama and as larvae northward to Banco Chinchorro, off of the coast of
Yucatan, Mexico. The new species is remarkable for having the lowest fin ray counts of the benthic sand-
perching group and a united, rounded, and darkly-pigmented pelvic fin without a frenum. Despite the morpho-
logical similarities, the mtDNA sequence (COI) for the new species is more than 25% divergent from other
Coryphopterus spp.

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This species is the first vertebrate to include in the description the DNA barcode developed for the infor-
matics system of the Barcode of Life Data System (BOLD) (Ratnasingham and Hebert 2007, Hanner, pers.
com.). The DNA sequence used for barcoding (COI) was obtained from both the holotype as well as the laval
stage and the close match confirms the larval identification. This new species description is also notable in
that the late larval stage is identified along with the adult. I include a comparison to other regional larval Cory-
phopterus spp. as well as to other gobioid larvae with similar meristics (Victor 2006, 2007). Furthermore, the
daily increments in the otolith microstructure of the transitional stage permits a direct estimation of the
pelagic larval duration, which is 63 days, relatively long for the large group of gobies that settle at a small size
of less than 10 mm SL (Victor 2006, 2007). The pattern of increasingly narrow increments during the latter
part of the pelagic larval phase is similar to that found to represent delayed metamorphosis in the larvae of
some other reef fishes (Victor 1986; Cowen 1991; McCormick 1999).
Materials and methods
Counts, measurements, and techniques follow Randall (2001). All fish lengths are standard length (SL). SIO
is the institutional abbreviation for the Marine Vertebrate Collection of the Scripps Institution of Oceanogra-
phy. Otoliths were extracted, cleaned, dried, and placed in immersion oil and examined under a compound
microscope with transmitted light and polarizing filters. Digital photographs of the otolith increment
sequences were taken at 400X magnification. The mitochondrial DNA sequences were obtained, processed,
and archived following the BOLD procedure outlined in Ratnasingham and Hebert (2007).
Coryphopterus kuna, new species
Fig. 1
Holotype. SIO-07-5, 17.1 mm SL, male, Panama, San Blas Islands, South side of Taintupo reef (9°32'44''N
78°57'26''W), 15 meter depth, sand, dipnet, 30 December 1982, B. Victor. Genbank Accession No. EF550211.
Diagnosis. Dorsal elements VI, I,8; anal fin elements I,8; pectoral fin rays 15; longitudinal scale series 25;
head naked except for scales on the side of the nape reaching near the level of the preopercular margin. Pelvic
fins fully joined medially by membrane, the innermost rays are branched and the longest and there is no fre-
num between the two pelvic fin spines. The pelvic fin is notably darkly-pigmented in the male holotype (the
only known adult specimen). Translucent with moderate black speckling on the head and body, two lines of
black spots along the lowest branchiostegal membranes forming an X across the isthmus, heavy black speck-
ling over the pelvic fin membranes, a faint black spot smaller than the pupil on the proximal upper pectoral fin
rays (not on the base of the fin), a black stripe across the mid-spinous dorsal fin, and prominent black spots in
two rows across the top of the eyeball. Apparently a small species, despite the small size of the holotype it is a
mature male since it has a long pointed urogenital papilla.
Description. Dorsal elements VI + I,8; anal elements I,8; all dorsal and anal soft rays branched, the last to
base; pectoral rays 15, the upper and lowermost unbranched; pelvic elements I,5 with the rays all branched,
united as a disk with a rounded edge, the innermost ray clearly longest, no frenum, a small fold of skin from
each pelvic spine to the body not connected to the fold on the other side; branched caudal rays 12, upper
unbranched caudal rays 9, posterior 3 segmented; lower unbranched caudal rays 8, posterior 2 segmented;
longitudinal scale series 25; transverse scale series 7; circumpeduncular scales 12; gill rakers 2+6; bran-
chiostegal rays 5; vertebrae 10+16; spinous dorsal-fin pterygiophore formula 3-22110.

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NEW SPECIES OF GOBY FROM THE CARIBBEAN
FIGURE 1A–C. Holotype of Coryphopterus kuna, 17.1 mm SL male (SIO-07-5)(A); head and fin markings (B); pelvic
fins united along full-length (C).

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FIGURE 1D, E.
Holotype of
Coryphopterus kuna
, pelvic fin frenum absent (D); prominent melanophores over eyeball (E).

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NEW SPECIES OF GOBY FROM THE CARIBBEAN
FIGURE 2. Transitional larva of Coryphopterus kuna, 7.3 mm SL (SIO-07-55).
Body elongate, depth 6.13 in SL, and compressed, width 1.12 in depth; ventral part of head and chest
broad and nearly flat; head triangular when viewed from above, its length 3.8 in SL, no head crest; snout
pointed and short, its length 4.14 in head; orbit diameter 3.05 in head, the eye extending above dorsal profile

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of head; interorbital space extremely narrow, 33.5 in head; caudal-peduncle depth 2.42 in head; caudal-pedun-
cle long, its length 1.31 in head. Mouth large, the maxilla ending below anterior 1/3 of pupil, the upper-jaw
length 2.23 in head; lower jaw projecting; mouth oblique, the gape forming an angle of 35° to horizontal axis
of head and body; upper jaw and lower jaws with multiple irregular rows of well spaced-apart slender curved
conical teeth. Tongue truncate with rounded corners.
Gill opening extending forward to below middle of opercle. Gill rakers nubs in holotype. Head naked
except for scales on side of nape extending forward nearly to eye; no scales on fins except a few on base of
caudal fin that are smaller than largest scales on body. Scales ctenoid except those on side of nape, thorax,
prepectoral area, and a few just above base of pelvic fins that are cycloid. Anterior nostril a short membranous
tube at level of middle of eye. Head pores prominent, as follows: a nasal pore, an anterior and a posterior
interorbital pore, a postorbital pore, an infraorbital pore below the postorbital, a pore at each end of a lateral
sensory canal; a short posterior lateral canal with a pore at each end, and 3 preopercular pores (i.e. Birdsong’s
B’, C, D, E, F, G, H’, K’, L’, M’, N, O’). Head papillae in rows vertically along the lower opercle and along
the lower rim of the preopercle extending forward along the line of the lower jaw.
Origin of dorsal fin behind upper base of pectoral fin, predorsal distance 2.79 in SL; spines of fins slender
and flexible; 1st dorsal fin lower than 2nd; 1st dorsal spine through the fifth about equal in length, 2.2 in head,
sixth spine shorter; spine of 2nd dorsal fin 4.33 in head; 1st dorsal soft ray longest, 1.93 in head; origin of anal
fin below base of 1st soft ray of 2nd dorsal fin, preanal distance 1.84 in SL; anal spine 4.0 in head; caudal fin
moderately rounded, 3.58 in SL; pectoral fins pointed, the middle rays longest, 3.45 in SL; origin of pelvic
fins directly beneath base of pectoral fins, prepelvic distance 3.39 in SL; pelvic fins fully joined by mem-
brane; pelvic frenum absent; pelvic spine 5.27 in head; 5th pelvic soft ray longest, nearly reaching origin of
anal fin, its length 4.22 in SL (1.29 in head); 4th pelvic soft ray about 94% length of 5th ray. Length of genital
papilla of male holotype almost equal to pupil diameter and narrow, the length-to-width ratio is 3.1.
Barcode sequence. A 652-nucleotide sequence of the section of COI gene used for barcoding by the
BOLD informatics database (Ratnasingham and Hebert 2007) was obtained for both the holotype and the lar-
val stage specimen (Genbank accession numbers EF550211 and EF550210 and protein sequences ABQ22956
and ABQ22955 respectively). The two sequences were a close match with 0.62% sequence divergence. Fol-
lowing the database management recommendation of the BOLD the 652 bp sequence (5' - 3') of the holotype
is presented here as well:
CCTTTACCTAGTATTCGGGGCCTGAGCCGGGATGGTAGGCACTTCCCTTAGCCTCCTTATCCGGGC
CGAACTAAGCCAACCTGGCGCCCTTCTGGGTGACGATCAGATCTATAACGTAATTGTCACCGCCC
ACGCATTCGTAATAATTTTCTTTATAGTGATGCCACTCATGATTGGAGGGTTCGGAAACTGACTCGT
CCCCCTAATGATCGGGGCCCCCGATATGGCATTCCCACGTATGAATAATATAAGCTTTTGACTCCTG
CCTCCTTCTTTTCTGCTTCTCCTAGCATCTTCGGGGGTAGAGGCTGGAGCTGGGACAGGTTGAAC
TGTCTACCCTCCGTTATCAGGCAACCTTGCTCATGCTGGAGCATCAGTCGATTTAACAATTTTTTCT
CTTCACCTAGCAGGTATTTCATCAATTCTGGGGGCGATCAATTTTATTACAACAATCCTTAACATGA
AACCTCCCGCCACTTCCCAGCACCAGACACCTCTGTTTGTTTGATCCGTGTTAATTACGGCAGTAC
TCCTCCTTTTATCTCTTCCCGTACTAGCTGCGGGCATTACTATACTCCTGACGGACCGAAACCTAA
ACACCACATTTTTTGACCCTGCAGGCGGGGGGGACCCAATCCTCTACCAACACCT
Color of holotype in alcohol. Pale yellowish with sparse black speckling and white and iridescent spot-
ting. Black spots on the body small and few; in irregular rows along the dorsal aspect below the base of the
dorsal fins, along the dorsal midline, lateral midline, and along the posteriormost portion of the ventral mid-
line of the tail. White spots, mostly made up of clusters of tiny single leukophores, sparse over the body and
along the lateral midline and speckling the operculum and jaws and a broad iridescent stripe across the oper-
culum onto the pectoral fin. No distinct spot at the base of the caudal fin but a thin line of melanophores at the

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NEW SPECIES OF GOBY FROM THE CARIBBEAN
base of the first six ventral caudal fin rays. A small patch of melanophores on the proximal portion of the pec-
toral fin rays between the 2nd and 6th rays (not on the fin base), a few melanophores on the inner side of the
pectoral fin base (the axil), and a white patch along the proximal portions of the 5th through 15th pectoral fin
rays. Sparse black spots speckling the top of the head and an indistinct black triangle directly below the orbital
rim down to the end of the maxilla. Prominent black spots over the dorsal aspect of the eyeball in two irregular
rows: two spots in the inner row, about seven in the outer row. Black spots line the lowest branchiostegal rays
in an X shape across the isthmus and then in a patch just forward of the pelvic fin. Pelvic fin is extensively
speckled with black spots concentrated along the fin membranes. Spinous dorsal fin has a distinct black spot
at the base of the first interspinous membrane and an additional black stripe along the mid-portion of the fin
across all of the spines and membranes and a white edging to the fin membrane tips; second dorsal fin is cov-
ered in fine white spotting as are the caudal and anal fins. Anal fin has an additional dusting of fine black
spots.
Distribution. Known from the Western Caribbean at the San Blas Islands of Panama (the adult) and
Banco Chinchorro, Mexico (larval specimen).
Etymology. Named for the Kuna indigenous people of the Kuna Yala, the region of Atlantic Panama in
which the holotype was collected, in recognition of their cooperation in marine biological research. Although
Coryphopterus is masculine, kuna is a noun in apposition and the “a” ending is thus appropriate.
Comparisons. The regional congenerics that share the low median and pectoral fin ray counts are dis-
tinctly different: the masked goby Coryphopterus personatus (Jordan and Thompson) and the glass goby
Coryphopterus hyalinus (Bohlke and Robins) are hovering, not benthic, species with divided and unmarked
pelvic fins, dark masks from the snout through the eye, black rings around the anus, and orange, not brown or
black, body markings (Bohlke and Robins 1960, 1962; Randall 1996).
Among the benthic Coryphopterus spp., all other species have more than 15 pectoral fin rays: 16 or 17 in
Coryphopterus alloides (Bohlke and Robins), 16 to 18 in Coryphopterus lipernes (Bohlke and Robins), and
17 to 20 in Coryphopterus dicrus (Bohlke and Robins), Coryphopterus eidolon (Bohlke and Robins), C. glau-
cofraenum, (along with C. tortugae and C. venezuelae), Coryphopterus punctipectophorus (Springer) and
Coryphopterus thrix (Bohlke and Robins) (Bohlke and Robins 1960, 1962; Springer 1960). Similarly, all
other Coryphopterus species have more than nine second dorsal fin elements: 10 or 11 (except a rare 9
recorded for C. glaucofraenum, C. alloides and C. thrix). The 9 anal fin elements are shared only with C.
alloides and a rare specimen of C. eidolon (Bohlke and Robins 1960).
Other than fin ray counts, the pelvic fin morphology is distinctive for Coryphopterus kuna, i.e. a united
rounded dark pelvic fin with the innermost rays longer than the next ray and no frenum connecting the pelvic
fin spines into a sucking disk. C. alloides, the species that shares nine anal fin elements with C. kuna (and has
only one or two more pectoral fin rays) has a distinctively divided pelvic fin morphology (and the innermost
ray unbranched) (Bohlke and Chaplin 1993). C. lipernes also has divided pelvic fins (Bohlke and Robins
1962). The remaining united-pelvic-fin group comprises C. dicrus with the innermost pelvic fin rays mark-
edly shorter and no frenum, C. glaucofraenum (and both C. tortugae and C. venezuelae) with the innermost
pelvic fin rays about equal to the next ray and with a frenum, and C. eidolon, C. thrix, and C. punctipectopho-
rus with a distinct frenum and the innermost pelvic fin rays somewhat shorter than the next ray (Bohlke and
Robins 1960; Springer 1960; Bohlke and Chaplin 1993).
In markings Coryphopterus kuna most resembles the benthic species without prominent stripes behind the
eye. It shares the dorsal fin stripe with C. alloides, but the latter species does not have fused pelvic fins. It is
distinguished from C. thrix by having the stripe across the spinous dorsal fin and the absence of the pectoral
fin base spot and extended second dorsal fin spine (but this spot is often indistinct and the fin spine is not
extended in C. thrix less than 20 mm SL). Males of Coryphopterus spp. usually do share the dusky pelvic fins
of the holotype of C. kuna. C. eidolon is a pale goby without dark markings and has orange head stripes some-
times outlined in thin black lines. The prominent black spots on the dorsal aspect of the eyeball is shared with

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several other Coryphopterus spp., however, in most other species the spots are few (usually three or four) and
large and not in two distinct rows.
Remarks. The holotype was perching on the bottom on a fine sandy plain in relatively deep (15-20m)
water and was collected with a dipnet. Along with the holotype, several other sandbed fishes were collected:
Chaenopsis spp., Diplogrammus pauciradiatus (Gill), and Achirus lineatus (Linnaeus). Although Cory-
phopterus spp. are ubiquitous in Caribbean fish collections and abundant in the region, this was the only spec-
imen of this species collected by me in the San Blas Islands of Panama. Explanations for why this species is so
elusive may rest on the fact that it occupies an obscure habitat: it was collected in deeper water than is usually
surveyed, was far from reef substrate, and the area is subject to some siltation from the Panamanian mainland.
In addition, the species is not particularly different-looking from the usual sand-bed gobies and thus would not
attract attention. Nonetheless, one would have expected its occurrence in trawling samples elsewhere and its
meristics would have immediately distinguished the specimen. Its occurrence only in the Western Caribbean
also assures that it was not discovered in the extensive collecting focused on the Bahamas, the Antilles, and in
US waters.
DNA Sequence. The COI sequences of the adult and larval specimens matched closely (0.62% diver-
gence) and confirm the identification of the larval specimen. The DNA sequence for Coryphopterus kuna is
quite divergent from other Coryphopterus spp. and Fusigobius spp., despite the morphological similarities.
Percent sequence divergence from other regional Coryphopterus spp. is 25.2% from C. dicrus, 27.8% from C.
glaucofraenum, 25.9% from C. lipernes, 25.4% from C. personatus, 26.1% from C. thrix, 25.8% from C. tor-
tugae and 26.5% from the eastern Pacific species C. urospilus. The percent sequence divergence is similar
from the Indo-Pacific species: 26.1% from Fusigobius signipinnis (Hoese and Obika) and 24.9% from
Fusigobius neophytus (Gunther) (equivalent to Randall’s (2001) Coryphopterus signipinnis and Cory-
phopterus neophytus). This marked divergence raises the question of the status of the genus Coryphopterus
and the relatedness of the many species that have resided within the genus at one time or another (Randall
2001; Thacker and Cole 2002). Thus the position of this species in the genus is provisional and elucidating the
relationships of these species would await a more complete molecular phylogenetic survey of the numerous
other species and related genera in the Gobiidae.
Early life history. A larval Coryphopterus kuna collected from a light trap at the Banco Chinchorro, off
of Yucatan, in Mexico is illustrated in Fig. 2. The specimen was collected by Dave Jones on March 29, 2006
(SIO-07-55). The larvae of C. kuna are recognized by the fin ray count of D-VI,9 A-9 pectoral 15 and the
identification is confirmed by the almost identical DNA sequence to the adult holotype. The body is relatively
thin, long and narrow with a large eye and a terminal mouth. The pectoral fins are long, reaching to the level
of the vent. The pelvic fins are also long, reaching to the vent, apparently united and with no visible frenum.
The dorsal and anal fin bases are relatively short, the caudal peduncle long and narrowing rapidly and there
are 5 to 7 procurrent caudal fin rays. Larval C. kuna are lightly marked, mostly along the lower body: melano-
phores are on the ventral midline at the isthmus and the pelvic fin insertion and then in a row along the anal fin
base, paired and one per side between the third and ninth element, then after a space, there is a row of melan-
ophores extending along the ventral midline of the caudal peduncle ending near the start of the lower procur-
rent caudal fin rays. Internal melanophores are present around the sacculus, at the dorsal surface of the swim
bladder, and around the gut near the vent. Transitional larval C. kuna develop a pattern of large discrete mel-
anophores on the dorsal aspect of the body, most notably three or four large melanophores over the eyeball, a
triangle of three with the vertex forward at the anterior midline between the eyes, several around the back of
the braincase, and then a row of sometimes paired melanophores along the dorsal midline of the body at the
base of the soft dorsal fin rays. A large stripe of iridophores extends backward on the head from the upper eye.
Melanophores develop along the first dorsal spine and at the base and tip of the second dorsal fin spine. There
are no melanophores at the angle of the jaw or on the caudal fin rays.

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NEW SPECIES OF GOBY FROM THE CARIBBEAN
Larval Coryphopterus kuna have similar proportions and morphology to larval C. dicrus, C. glaucofrae-
num, and C. tortugae (the latter species are visibly indistinguishable as larvae and are identified by mtDNA
sequences only) and larval C. personatus/hyalinus (Victor 2006, 2007). However, these similar larvae do not
share the low fin ray counts of the C. kuna larval type. Larval C. kuna differ from larval C. dicrus, C. glaucof-
raenum, and C. tortugae in missing the lower caudal fin melanophores and the melanophores at the angle of
the jaw and behind the last dorsal fin ray. At transition, larval C. kuna have a pattern of a few discrete large
melanophores on top of the head vs. stripes going back from the eye of transitional C. dicrus, C. glaucofrae-
num, and C. tortugae larvae and a patch of tiny melanophores on top of the head and a distinctive patch
around the vent in transitional C. personatus/hyalinus larvae.
The melanophores over the dorsal aspect of the eyeball of larval C. kuna are more prominent than is
observed in most other larval Coryphopterus spp. and break up into two stripes of relatively smaller melano-
phores on the eyeballs of adult C. kuna. This description of larval C. kuna agrees with most of the characters
enumerated for an unknown larval type (Coryphopterus B of Baldwin and Smith (2003)), except one of the
larvae of that type that was raised in captivity for a few days had apparently divided pelvic fins and C. kuna
has fused pelvic fins. This is not likely an ontogenetic change, since divided pelvic fins in Coryphopterus spp.
are a derived character and at least two of the species with divided pelvic fins, C. personatus and C. lipernes,
have fused pelvic fins as larvae and at settlement (Victor 2007). Since the collection site was Belize, included
within the range of C. kuna (Yucatan to Panama), further collections should resolve whether there are more
cryptic species of Coryphopterus in the region.
Other gobioid larvae that may rarely overlap the fin ray counts for larval Coryphopterus kuna comprise
the gobies Lythrypnus spp. and Bathgobius curacao (Metzelaar), as well as the eleotrid Eleotris amblyopsis
(Cope). The larvae of these potentially confounding species have been identified and characterized in Victor
(2006, 2007) and they are quite different from the C. kuna larval type. Larval Lythrypnus spp. are shorter and
wider than larval Coryphopterus spp. and have a conspicuous melanophore at the angle of the jaw, not present
on the larval C. kuna. Furthermore, Lythrypnus spp. larvae have no dorsal melanophores before other distinc-
tive transitional melanophore patterns develop on the head. The transitional larvae of Lythrypnus spp. have a
characteristic Òradiating spokesÓ pattern of melanophores around the eye that develops before other melano-
phores along the dorsal aspect (but occasionally a single melanophore at the rear edge of the dorsal fin devel-
ops early).
Larval Bathgobius curacao are very different and have rows of melanophores along the dorsal midline,
the ventral midline, and internally along the spine and are smaller at the same stage of development than larval
Coryphopterus kuna. Larval Eleotris amblyopsis have a completely different appearance: they are larger with
a stout thick body and the eleotrid larval melanophore pattern of long streaks or rows of melanophores along
the ventral midline along with a conspicuous patch of melanophores covering the caudal peduncle. In addi-
tion, most Eleotris amblyopsis larvae have narrowed eyes and distinctive melanophores covering the surface
of the iris (Victor 2006, 2007).
Otolith microstructure. The sagittal otoliths of the transitional stage of larval Coryphopterus kuna reveal
a clear increment array (Fig. 3). These arrays have been interpreted as daily in numerous otolith studies for
reef fishes and experimentally validated for a variety of fishes, although not for this species in particular
(Thorrold and Hare 2002). At the center of the core there is a rod-like or oblong primordium about 7 microns
long (typical of gobiid otoliths). The core is an area with no distinct increments extending about 20 microns
from the center and outlined by an oval ring that can be brightly demarkated at some focal planes. This core
region is formed before hatching in gobies. Surrounding the core there is an increment array of relatively nar-
row but clearly delineated dark and light lines easily visible in all quadrants of the otolith when illuminated
with transmitted light from the lateral aspect of the otolith. The optimal array for counting and measuring
increments is along the longest axis of the sagitta. The lapillus shows a similar increment pattern, but since
this pair of otoliths in gobies is often much smaller than the sagittae, the array is compressed and the narrower

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bands merge together. The asterisci, the third pair of otoliths in bony fishes, usually have indistinct increments
and are not used in otolith aging.
The count of 63 days from the edge of the core region to the edge of the otolith indicates a pelagic larval
life of 63 days. In species with broadcast eggs, some number of days need to be added to the increment count
for the hatching and embryonic developmental stages (Victor 1991). However, these gobies have brooded
eggs that hatch with the core already developed and the hatchlings begin their pelagic life at that point. The
capture of transitionally-marked larvae at light traps off the reef edge indicate that the larvae were about to
settle, probably that same night, onto the reef. Thus otolith counts from transitional larvae caught at the reef
edge are one of the most direct estimates of pelagic larval duration. In later life stages, a settlement mark
needs to be interpreted and validated, and the counts subdivided to infer a pelagic larval duration (Victor
1991; Thorrold and Hare 2002).
The array of increment widths on the otoliths of larval Coryphopterus kuna show a distinct narrowing at
day 30-35 from increments as wide as 5 microns down to increments as narrow as one micron that continue
over the next 25 days. Near the very edge of the otolith there are a few indistinct slightly wider increments that
could represent faster growth associated with approach to shore waters or transitional changes in head mor-
phology commonly found in transitional larvae of reef fishes. A similar pattern of narrowing otolith incre-
ments during the latter portion of the pelagic larval life has been shown to correlate with slower growth in
another reef fish (Victor 1986) and indicates delayed metamorphosis, a phenomenon common in marine
invertebrate larvae (Pechenik 1990), and perhaps an important part of the pelagic stage of many reef fishes.
FIGURE 3. Sagittal otolith of transitional larva of Coryphopterus kuna, lateral view, photographed at 400x. The array of
daily otolith increments extends along the longest radius of the sagitta from the center of the otolith (at left) to the edge of
the otolith (at right). The primordium is the horizontal black oblong at the left edge of the figure and is about 7 microns
long, the core (pre-hatching) is the area without clear increments about 20 microns out from the center.
Acknowledgements
I appreciate the cooperation of the Smithsonian Tropical Research Institute, the government of Panama, and
the Kuna people of the Kuna Yala of the Comarca of San Blas. K. Clifton, L. Clifton, S. Floeter, J. Gauvain,
D. R. Robertson, B. Ruttenberg, M. Schildhauer, G. Walsh, and R. Warner provided invaluable assistance in
Panama and elsewhere. D. Jones graciously provided the larval Coryphopterus kuna and J. Randall contrib-
uted the Indo-Pacific tissues. The cooperation of H.J. Walker and P. Hastings at the Marine Vertebrate Collec-
tion of the Scripps Institution of Oceanography is appreciated and comments by A. Acero, D. Greenfield, and
L. Rocha were very helpful. In particular, I thank M. Shivji and R. Hanner and the BOLD team, C. Maitland,

Zootaxa 1526 © 2007 Magnolia Press · 61
NEW SPECIES OF GOBY FROM THE CARIBBEAN
A. Borisenko, R. Breese, and G. Downs, for their help with my introduction to barcoding and efficiently man-
aging the informatics side of the project.
References
Baldwin, C. C. & Smith. D. G. (2003) Larval Gobiidae (Teleostei: Perciformes) of Carrie Bow Cay, Belize, Central
America. Bulletin of Marine Science, 72, 639–74.
Bohlke, J. E. & Chaplin C.G. (1993) Fishes of the Bahamas and Adjacent Tropical Waters. University of Texas Press,
Austin, 544 pp.
Bohlke, J. E. & Robins C. R. (1960) A revision of the gobioid fish genus Coryphopterus. Proceedings of the National
Academy of Sciences, Philadelphia,112 (5), 103–128.
Bohlke, J. E. & Robins C. R. (1962) The taxonomic position of the West Atlantic goby, Eviota personata, with descrip-
tions of two new related species. Proceedings of the National Academy of Sciences, Philadelphia, 114, 175–189.
Cervig n, F. (1966) Los peces marinos de Venezuela. Estacion de Investigaciones Marinas de Margarita, Caracas,
Venezuela, 951 pp.
Cervig n, F. (1994) Los peces marinos de Venezuela. Volume 3. Fundaci n Cientifica Los Roques, Caracas,Venezuela,
295 pp.
Cowen, R. K. (1991) Variation in the planktonic larval duration of the temperate wrasse Semicossyphus pulcher. Marine
Ecology Progress Series, 69, 9–15.
Garzon-Ferreira, J. & Acero, P. (1990) Redescription of Coryphopterus tortugae (Jordan) (Osteichthyes: Gobiidae), a
valid species of goby from the western Atlantic. Northeast Gulf Science, 11(2), 105–112.
Greenfield D.W. & Johnson R.K. (1999) Assemblage structure and habitat associations of western Caribbean gobies
(Teleostei: Gobiidae). Copeia, 1999, 251–266.
McCormick, M.I. (1999). Delayed metamorphosis of a tropical reef fish (Acanthurus triostegus): a field experiment.
Marine Ecology Progress Series, 176, 25–38.
Pechenik, J.A. (1990) Delayed metamorphosis by larvae of benthic marine invertebrates: Does it occur? Is there a price
to pay? Ophelia, 32, 63–94.
Ratnasingham, S. & Hebert, P. D. N. (2007) BOLD: The Barcode of Life Data System (www.barcodinglife.org). Molec-
ular Ecology Notes (in press).
Randall, J. E. (1996) Caribbean reef fishes. 3rd edition. T.F.H. Publications, Hong Kong, 368 pp.
Randall, J. E. (2001) Five new Indo-Pacific gobiid fishes of the genus Coryphopterus. Zoological Studies, 40, 206–225.
Springer, V. (1960) A new gobiid fish from the eastern Gulf of Mexico. Bulletin of Marine Sciences of the Gulf and
Caribbean, 237–240.
Thacker, C.E. & Cole, K.S. (2002) Phylogeny and evolution of the gobiid genus Coryphopterus. Bulletin of Marine Sci-
ence 70 (3), 837–850.
Thorrold, S. R. & Hare, J. A. (2002) Otolith applications in reef fish ecology. In: Sale, P. F. [ed] Ecology of Fishes on
Coral Reefs. Academic Press, San Diego, pp. 243–264.
Victor, B.C. (1986) Delayed metamorphosis with reduced larval growth in a coral reef fish, Thalassoma bifasciatum.
Canadian Journal of Fisheries and Aquatic Sciences, 43, 1208–13.
Victor, B.C. (1991) Settlement strategies and biogeography of reef fishes. In: Sale, P. (Ed), The Ecology of Fishes on
Coral Reefs. Academic Press, Orlando, Florida, pp. 231–60.
Victor, B.C. (2006) The late-stage larvae of Caribbean gobies, eleotrids, and microdesmids: identification guide and pat-
terns of size and age at settlement. In: Program book and abstracts, 2006 joint meeting of the American Society of
Ichthyologists and Herpetologists July 12–17, 2006. Allen Press, Lawrence, Kansas, pp.128–129.
Victor, B.C. (2007) A photographic guide to the larvae of coral reef fishes. B. C. Victor. Available from http://www.cor-
alreeffish.com (2/26/2007).

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