NAMERIC
N
MUSEUM
Novia!esD
PUBLISHED BY
THE
AMERICAN
MUSEUM
OF
NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET NEWYORK, N.Y.10024 US.A.
NUMIBER
2616
APRIL13, 1977
C.
LAVETT SMITH AND JAMES C. TYLER
Redescription
of the Gobiid Fish
Coryphopterus
iperne
Bhe
ad
Robins,
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AMERICAN MUSEUM
No iltates
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024
Number 2616,pp. 1-10,figs. 1-3, table 1 April 13, 1977
Redescription
of the
Gobiid Fish
Coryphopterus lipernes
Bohlke
and
Robins,
With
Notes on Its
Habits and
Relationships
C. LAVETT SMITH1 AND JAMES C. TYLER2
ABSTRACT The bluenose goby, Coryphopterus lipernes
Bohike and Robins, was described in 1962 from three specimens collected in the Florida Keys; it has remained poorly known since that time. Recent collections and observations indicate that this species is widespread in the Caribbean and Bahamas but at low population densities wherever it occurs. It is also one of the few
species of western Atlantic reef fishes that lives
in relatively continuous close physical contact
with live corals. All the individuals, observed at
night and during the day, spentmostof the time resting on livecorals, withonlyafewbriefforays
onto nearby algal mats, or off the coral to feed.
In this respect, the bluenosegobyisanecological
counterpart of the Indo-Pacific clownfishes
(Amphiprion). The mechanism by which the bluenose goby avoids being stung by the nemato-cysts of the coelenterates may not be the same as that of the clownfishes because clownfishes
become acclimated to individual anemones, whereas the bluenose goby canmovefreelyback and forth among coral coloniesof both thesame and different species. Other species of fishes
as-sociated with live corals in the West Indies share with the bluenose goby certain features that we
interpret to be specializations for this way oflife.
Within the genus Coryphopterus, two divergent
lineages show progressive specialization toward coral-dwelling, on the one hand, and toward sand-dwelling, on theother.
INTRODUCTION
In the course of our work with Bahamian coral reef fish communities, we repeatedly have
encountered asmall, predominantlyyellowgoby, with conspicuousblue markingsonitshead, that lives in close associationwith the polyps of
mas-sive corals. Although we initially believedthisto
beanundescribed species, itisnowapparentthat
it is Coryphopterus lipernes Bohlke and Robins,
1962. The purpose of the present paper is to supplement the original description ofthis
little-known species with color notes, measurements, observations on ontogenetic changes and sexual
dimorphism, and general information on its
habitsandhabitat.
Within the coral reef cosmos there is a spec-trum ofassociations offishes withinvertebrates. In the West Indian fauna thereareobvious
oblig-atory relationships between the cardinal fish
Apogonstellatus(Cope)and thegastropodconch Strombus gigas Linnaeus, the pearlfish Carapus
'ChairmanandCurator,DepartmentofIchthyology,theAmerican Museum of NaturalHistory.
2ResearchSpecialist,NationalMarineFisheriesService.
bermudensis (Jones) and theholothurian Actin-opyga agassizi (Selenka), and between certain highly specialized gobies such as Risor ruber
(Rosen), Evermannichthys spp. and Pariah sco-tius Bohlke and Robins and massive sponges.
These highly evolved associations usually entail
apparent morphological specializations on the part of the fish as well as behavioral adapta-tions. Other, apparently less rigorous, associa-tions also exist. For example some wrasses and pomacentrids use vase-shaped sponges as
noc-turnal resting places and blennies of the genus
Acanthemblemaria utilize abandoned worm
tubes and the shells of vermetiform gastropods assheltersites.
Coryphopterus lipernes is one of arelatively few fishes that normally lives in close contact
with live coelenterates. Other members of its genus also show specializations that are cor-related with habitat preference. Some arehyaline sand-dwellers, some have more pigment and are
associated with rocky bottoms, and somehover
overthe bottom and nearverticalsurfaces.Since these specializations are derived character states we have tested their usefulness in defining re-lationshipsamongtheWesternAtlantic species of Coryphopterus.
ACKNOWLEDGMENTS
Assistance with our extensive scuba
observa-tions of Coryphopterus lipernes and associated
fishes dwelling on live coral surfaces was
help-fully provided atBimini,Bahamas, byMrs.Helga
andMr.MilanTyler,andMr.PedroRomerof the LernerMarine Laboratory staff made our
boating
operations as efficient as possible. Mrs. Norma
Feinberg of the Department of Ichthyology of the American Museum assisted with the
identifi-cations of species of Coryphopterus that were
compared with C. lipernes. Drs. James W. Atz and Gareth J. Nelson helpfully criticized the
manuscript and figures 1 and 2 were drawn by
Ms.CarolGeneSchleifer.
REDESCRIPTION OF
Coryphopteruslipernes Bohlke andRobins, 1962
Figure 1
Material Examined. (All specimens are
cata-logued atthe American Museum ofNatural
His-tory; all lengths are given as standard length; all specimens were collected by J. C. Tyler, C. L. Smith and associates.)
VIRGIN ISLANDS: St. John. AMNH 34948, one probable 9, 11.5 mm., from TEKTITE study reef, Beehive Cove, Lameshur Bay, depth 11Im.,
October 8, 1973. (Listed in table 4, Smith and
Tyler, 1975, as Coryphopterus sp.). AMNH 31118, seven additional specimens, one probable
9, two probable d and fourjuveniles, 8.1-15.4 mm., collected on coral head near the above
lo-cality atsimilar depth,October 11, 1973. BAHAMAS: Turtle Rocks, south of Bimini.
AMNH 33931, 19 21.0 mm.,depth7m., May 5, 1975; AMNH 33925, 1d 24.5 mm., depth 7 m.,
June 26, 1975; AMNH 33908, one juvenile 10.7 mm., 19 21.5 mm., 3d 18.0, 20.5, 24.4 mm.
depth 7 m., June 27, 1975; AMNH 33946, 1d
23.2 mm., depth 7 m., June 29, 1975; AMNH 33703, two juveniles 11.8 and 14.4 mm., 1d 23.7 mm., depth 7m., August 2, 1975;AMNH
33944, 1d 20.3 mm., depth 7 m., August 6, 1975. Berry Islands, Mamma Rhoda Rock. AMNH 34743, 19 22.7 mm.,depth7m.,
Febru-ary 5, 1968.
Meristic and morphometric features of nine specimens, eight from St. John, Virgin Islands,
and one from TurtleRocks, Bimini,Bahamas,are
presentedintable 1.All specimens agreewith the characters of the genus given by Bohlke and Robins (1960), exceptthat thefleshy padonthe shoulder girdle is not well developed. Although the specimens examined by Bohlke and Robins
apparently did not have enlarged neural and hemal spines on the penultimate vertebra, we
find these to be present on a 21 mm. specimen, as seeninX-rays.
The following color and morphological notes
supplement the original description by Bohlke
and Robins (1962, pp. 186-187). Living
speci-mens photographed during daylight hours are predominantlyhyaline yellow with light electric-blue markings onthe head. These blue markings include a round spot at the tip ofthesnout,the
dorsal surface of the eyeball, and three
longi-tudinallines: one inthemidlinefrom behind the
eyes to the nape, and one frombehind eacheye
to above the operculum. Two bluishwhitelines,
one on each side of themidline and apparently
TABLE 1
Proportional Measurements and Meristics of Nine Specimens of
Coryphopterus lipernesa
(Standard lengthinMillimeters;allproportions in thousandths of the standardlength.)
Specimen Number 1 2 3 4 5 6 7 8 9 x + s.d. Standardlength 8.1 8.7 11.5 13.0 13.1 13.2 13.9 15.4 21.0 13.1 3.8 Headlength 309 333 296 315 298 303 302 305 281 304.6 14.2 Head width 160 172 139 162 176 174 165 169 167 164.9 11.1 Eyelength 111 115 104 108 107 114 115 110 105 109.0 4.2 Maxillarylength 86 92 96 100 107 98 86 91 90 94.0 6.9 Snout length 62 57 52 69 53 61 58 58 48 57.6 6.2 Postorbital head length 136 149 130 146 153 144 144 149 133 142.7 7.9 Interorbital width 12 11 9 15 15 11 14 13 12 12.4 2.0 Bodydepth 160 184 148 185 176 182 173 169 200 175.2 15.2 Caudal peduncle depth 74 92 96 97 99 106 101 84 105 94.9 10.3 Caudalpeduncle length 259 310 252 279 260 242 252 247 252 261.4 21.0 Upper caudalrays - - - 246 - - -
-Midcaudalrays - - - 237 - - - -Pectoral fin length - 287 243 300 298 295 273 286 281 282.9 18.5 Pelvic fm length 210 230 226 254 229 258 259 240 233 237.7 16.6 Snouttopelvic finorigin 321 368 304 369 336 341 360 338 305 338.0 24.7 Snouttodorsal finorigin 395 379 357 354 359 379 374 364 348 367.7 15.1 Length of first
dorsal fin base 136 138 130 138 122 129 137 130 129 132.1 5.5 Length of second
dorsal fin base 198 207 174 192 198 212 187 201 171 193.3 13.9 Longestdorsal finspine - 218 - 177 176 189 187 182 186 187.9 14.2 Longest dorsal fin ray 173 - - 169 137 167 180 169 143 162.6 16.1 Snout to anal finorigin 543 575 557 608 565 598 597 578 567 576.4 21.2 Lengthofanal fin base 160 172 148 162 176 189 173 169 167 168.4 11.4 Longestanalfin
ray - 149 - 131 115 144 158 156 143 142.3 15.0 Dorsal fin VI-lo VI-lo VI-lo VI-10 VI-10 VI-10 VI-lo VI-lo VI-lo -
-Analfin 10 10 10 10 10 10 10 10 10 - -Pectoralfin 17-17 16-17 17-17 17-17 16-17 17-17 17-17 17-17 - -Lateralscales - 25 28 25 26 26 26 26 27 - -Circumpeduncular scales 12 12 11 12 12 12 11 12 12 - -Gill rakers 11 10 11 11 10 - 9 11 10 -
-aColumn
1,2;4-8.AMNH31118. St.John,VirginIslands. Column3. AMNH 34948. St.John, VirginIslands.behind the head to below the first dorsal fin. There are two or three other small bluishwhite
spots, apparently in the spinal meninges, the last ofwhich is at the level of the base of the caudal fin. The iris is blue. The areas behind each eye andthe lower parts of the opercula are light blue. One specimen from Turtle Rocks obtained May 5, 1975 was examined after it had been in formalin for 10 days. The yellow pigment that remained was concentrated along the edges of the scales onthe dorsal surface ofthe body and
in the mid-dorsalline. There was an intense yel-low line behind the middle of the eye, ending over the pectoral fin. Another bright yellow line ran on the lower sides from behind the middle of the pectoral fin base to over the end of the anal fin base. This lower line was broken posteriorly into short segments.
A second specimen from Turtle Rocks, col-lected June 26, 1975, was examined after about 15 hours in formalin. Body and head were uni-form yellow above; belly and underside of head pure white below level of lower rim of orbit. Cheeks behind each eye with three dark lines; interspaces intensely yellow. Deep yellow bar
acrossupper part ofoperculum.Prepectoral area with two longitudinal bands of melanophores:
one extending forward from base of uppermost rays, another forward from base of middle rays. Scattered melanophores forming faint third line
forward from base of lowerpectoralrays. Between these lines, prepectoral area intensely yellow.
Top ofhead and upper part of body uniformly
pale yellow; sides yellow, shading to white
ven-trally. Peritoneum as seen through body wall bright silvery. A narrow, intense yellowline ex-tends behind the pectoral fin alongsideof body, ending at lower half of caudalbase. Anotherless intense and less regular line parallel and slightly above it. Pectorals,both dorsals, and caudal fins with pale reddish rays; pelvic rays pure white.
Blue spots on head generally faded but intense on upper cheek and operculum and on topsof
eyes.
Fresh specimens collected June 27, 1975 had orange-yellow streaks along bases of soft dorsal and anal fins. Each ray ofdorsal, pectoral, and caudal fins was suffused with orange-red. In living fish, spinalmeninges are pigmentedandare seen as dark line with three light interruptions wherepigment is less intense.
At night, with the fish resting quiescent on
live coral surfaces, the color fadesto an overall dull gray, and the unpigmented areas of the meninges stand out as chalkwhite spots. Neither the overall yellow coloration nor the blue head
markingscan beseen in this nocturnal pattern.
Sexual Dimorphism. Thegenitalpapilla ofthe
adult male islongandslender,that of thefemale
isshort and broad. Thepapillain both sexes has amidventralband of densemelanophores,butits lateral edgesand dorsal surface are unpigmented
orhaveonlyafew scatteredmelanophores. The second dorsal spine of the male is
strik-ingly elongated, reaching to the posterior end of the base of thesoft dorsalfin in somespecimens.
There is no elongationof the second dorsalspine
in the female, in contrast to the situation in
1
3~~~~~~~~~~17
SMITH ANDTYLER: GOBIID FISH
Coryphopterus personatus (Jordan and Thomp-son) inwhich both sexes havethe second dorsal spineprolonged.
The posterior two rays of the soft dorsal and
anal fins are also slightly elongated in the male,
forming lobelike extensions that reach the base
of the procurrent caudal rays in large specimens. In the female, the ends of the depressed dorsal
and anal fins fall abouthalfwaybetween the end
of the respective fin base and the base of the anterior-most procurrent caudal rays. The pelvic
fins are longerinthemale,reachingtoorslightly beyond the origin of the anal fin; the pelvicfins
end at a level between the anus and anal-fin in
thefemale.
Ontogenetic Changes. Our smallest specimens have theinner rays ofthe pelvic finscompletely connected bymembrane, thus forminga typical
gobiid pelvic disc (fig. 2A). Inlarger specimens, the interpelvic membrane is almost completely absent (as shown in fig. 2B) so that the pelvic finsareessentiallyseparate.
LIFE HISTORY
A 21 mm. female obtained at Turtle Rocks
(250 40'N lat.) on May 5, 1975 hadlarge eggs.
On June 26, 1975, a smallindividual, estimated
to be less than 5 mm., was seen there on a
Col-pophyllia colony but was not taken. This sug-gests alate spring spawningseasonfor the Bimini area. An 8.7 mm. individual was taken in the Virgin Islands
(180
19' N lat.) on October 11, 1973,and this may indicateeitheralater spawn-ing period for the more southerly locality,or that the breeding seasonispossibly prolonged
inthewarmerpartsofthe speciesrange.
DISTRIBUTION
At the time it was originally described,
Cory-phopterus lipemes was known
only
from threespecimens from the FloridaKeys.Ourspecimens
now extend the known range to the Virgin
Is-lands and to the areasencompassing the Bimini
and Berry islands of the Bahamas. We find no
differences between the Bahamian and Virgin
Is-land specimens. We also have aphotograph ofa
specimen from Glovers Reef, British Honduras,
and sight records from off Providencia
(Old
Prov-idence) IslandnearthecoastofNicaragua,Grand
A 2 2 --z 7 ''C -vVW\f P1 .f3
-FIG. 2. Pelvic fins of Coryphopteruslipernes. A. AMNH 33908, a juvenile 10.7 mm. S.l. B. AMNH 33908, an adult male 24.4 mm. S.l. Cayman Island (C. Gilbert, pers. commun.), and
Freeport, Grand Bahama. Recently it has been reported from Providenciales Island, Caicos Bank; Discovery Bay, Jamaica; and Curacao, Netherlands Antilles (Colin, 1975). The species has been taken at depths ranging from 7 m. in the Bahamasto26m.inthe Florida Keys.
HABITS
Coryphopterus lipemes is closely associated with live corals. It is not an especially abundant species, and most individuals we have seenwere solitary or members of small groups oftwo or
threeinhabiting a large coral colony, with only a few colonies in any given region acting as host to these fish. For example, our series of seven
speci-mens from the Virgin Islands was collected alive
in test tubes from two large contiguous colonies
of Diplorialabyrinthifornis (Linnaeus).
Colin (1975, p. 191) provided data on the
abundance of C. lipernes and several species of Gobiosoma in Discovery Bay, Jamaica.
Ap-parently there are considerable fluctuations in abundance. The population density was lower in February, 1972 than in August, 1971 but reached high levels in June, 1972 and February, 1973.
Several species of massive corals serve ashosts;
we have observed C. lipernes on the following: Colpophyllia natans(Houttuyn)
Diploria labyrinthiformis (Linnaeus)
Montastrea annularis(EllisandSolander) Montastreacavernosa (Linnaeus)
Siderastreasp.
Mycetophyllia lamarckiana (Edwards and Haime)
In addition, Colin (1975, p. 253) presented a photograph of C. lipemes on the coral Stephan-ocoenia michelini and listed (p. 202)it as occur-ring onAgaricia spp., Poritesastreoides, Diploria clivosa, D. strigosa, and Manicina areolata. Indi-viduals sometimes enter holes, disappear under
the free edges of the colonies, or rest onnearby algal mats.
We observed one extraordinarily large coral mound off Turtle Rocks in the Bimini chain, Bahamas, on five consecutive daylight periods from June 27, to July 1, 1975. This hemi-spherical mound (approx. 5 m. high and 5 m. in
diameter) was at a depth of 9 m. It was dominated by Montastrea annularis, which made up perhaps 95 percent ofitssurface. Most of the M. annularis was of the helmet form, but a few
colonieswerepartially ramose.
On the south side of the main mound, a
colony of Colpophyllia natans measuring ap-proximately 1 m. in diameter had four
Cory-phopterus lipernes onJune28,thedayonwhich we cropped one to serveas a voucher specimen. On June 29, four individualswere present again when we first approached the colony, but one
left as soon as we arrived. It moved to a nearby colony of Montastrea cavernosa, where it
re-mained until July 1 when it was again seen on
the Colpophyllia. Of the three C. lipernes that remained on theColpophyllia colony, two were
large individuals and one was small. These
ap-peared to have partitioned the coral surfaceinto
definite home ranges, although one of -the large
individuals apparently ranged over more of the surface than did the othertwo.OnAugust2, this large plate ofColpophyllia had twolarge andtwo
small C. lipernes during daylight (18:30 to
19:00). Later the same day, at 20:30 to 21:30,
one of the larger and one of the smaller indi-viduals were found quiescent and in their drab nighttime coloration. Further observations of C. lipernes onthisplateofColpophyllia during day-light and nighttime on August 6 and 12 con-firmed that this goby is brightly colored and
active by daylight, but drab-colored and
quies-cent ontheexposed coral surfacesatnight. On the southeast side of thedome, therewere
two adjacent colonies ofMontastrea cavernosa, one grayishin color andonegreenish. One
Cory-phopterus lipernes was seen in the gray
Mon-tastrea on June 28, and it moved freely back and forth between Montastrea and Mycetophyllia lamarckiana. On June 29, the C. lipernes wason the greenish colony of Montastrea. We did not observe this individual on June 30orJuly 1, but
itor aspecimenof similar sizewas presentduring
most observations made from August 2 to 12 during daylight andatnight.
On the north side of the same dome, a flat colony ofMycetophyllia lamarckiana about 30 cm.in diameter had oneC. lipernesfrom June27
to 30.Thisindividual, apparentlyafemale,spent
much ofits daylight time on thealgalmats
adja-cent to the colony and occasionallymade brief foraysonto the nearbyMontastreaannularis.We
didnot observethis coralplateduringtheAugust 2to 12 surveillanceperiods.
Although the C. lipemes of this huge coral mound occasionallymoved onto theMontastrea annularis that dominated it, these gobies spent
almost all of the time on the other species of corals that made up only about 5 percentofthe surface.
Feeding consisted ofthe fish actively chasing
particles in the water, sometimes leaving and hovering over the coral for several seconds at a
SMITHANDTYLER: GOBIIDFISH the coral surface. Detrital particles were taken in
and sometimes spit out. Clouds of small mysid shrimps were seen nearby during daylight, but they were never pursued by the gobies. Colin (1975, p. 207) also noted C. lipernes "hovering above coral heads apparently eating plankton." Occasionally, other fishes such as Poma-centrusplanifrons and Holocentrus rufus, passed close to the corals, sometimes remaining almost in contact with the corals for a few moments. At such times, a Coryphopterus lipernes would
move away to avoid the larger fish but did not seekany shelter nor leave the coral surface.
ACTIVITY CYCLES
Coryphopterus lipernes is a day-active species. During our observation periods, sunrise occurred approximately at 06:45 to 07:00, and sunset at 20:00 to 20:15, with the moon at about one-quarter. Feeding behavior was observed from 07:40 to 08:15 and from 12:20 to 19:00 hours from June 28toJuly 1, and August 2, 6, and 12, 1975, at Turtle Rocks, Bimini. At night, C.
lipernes assumes itsnocturnal,drab color pattern and rests motionless in valleys of meandriform corals. Quiescent fish can beapproached closely
and are especially easy to collect at night by prodding them into a glass test tube. Our
ob-servations of quiescent nighttime behavior were
made at 20:30 to 22:25 on August2,6, and 12. DISCUSSION
Coryphopterus lipernes is associated with several different live corals and individual fish move freely from ohe to the other. This indicates that the mechanisin by which it avoids being stung by coral nematocysts is quite different from that of the Indo-Pacific clownfishes
(Amphiprion), which develop ad hoc immunity after a period ofacclimation, during which time the fishes' slime is altered so that it prevents the anemone's nematocysts from discharging
(Mariscal, 1970). It remains a question whether other fishes that live in close association with coelenterates utilize the same or different
mech-anisms for protection against the host's
nemato-cysts.
The habit of resting onandhunting overlive
corals is shared by several West Indian fishes. Along with C. lipernes, themostcommonspecies found on live coralsisPseudemblemariasignifera
(Ginsberg), which darts rapidly over the surface ofthe colonyasit searches for foodorwhenit is
disturbed. Unlike C. lipernes,Pseudemblemaria spends much of the time hiddenin plants orin
cavities on the dead surface of the reef (Smith
and Tyler, 1972, p. 156).Pseudemblemaria does
not venture ashigh above the coral surfaceasC. lipernes, nor does it hover like C. personatus
(Jordan andThompson). Pseudemblemaria is
pre-dominantly colorless with reddish brown blotches and flecks. Its spinal meninges are
pig-mentedwith alternating bands of red andwhite,
a pattern not unlike that of the meninges of C. lipernes. Webelieve that P. signifera oftenretires atnight into crevices orholes in the reef (Smith
and Tyler, 1972, p. 156) even though we ob-served it on thesurface of coral platesduringall
three of our nighttime surveillance periods of August 2, 6, and 12 from 20:30 to 22:25 at Turtle Rocks, Bimini, when it remained
quies-cent in valleys of the meandriform corals Col-pophyllianatans andMycetophyllialamarckiana.
The cryptic and semitransparent colorationofP. signiferaismuch the sameby dayandnight.
Atdepths of 16m.offFreeport,Grand
Baha-mas, we observed that Emblemaria bahamensis
(Stevens) has daylight and nighttime habits
sim-ilartothoseof Pseudemblemaria.
In the Virgin Islands, Smith and Tyler(1972, p. 164) reported that the goby Gobiosoma
saucrum (Robins) is closelyassociated with coral
heads of Siderastrea spp. and Montastrea
an-nularis. It, too, ishyaline with internal pigment and exhibits a double yellow ocellus in the
peri-toneum that forms afigure 8 pattem within the body wall. The cleaner gobies Gobiosoma
eve-lynae Bohlke and Robins and G. genie Bohike and Robins also utilize the surface of livecorals,
frequently Montastrea sp. and Agaricia sp., as cleaning stations. These scaleless gobies have
heavy skin pigmentation and a thick mucous
coating. They are boldly striped duringtheday
but fade to apale, nearly uniform gray atnight
as they rest motionless between coral polyps.
Colin (1975, p. 207) listed nine forms of the
subgenus Elacatinus (genus Gobiosoma) that live oncorals.
Another species with nighttime habits similar
to those of Coryphopterus lipernes and Pseu-demblemaria signifera is Coryphopterus per-sonatus. Conspicuous during the day, C.
perso-natus hovers and feeds a few centimeters above soft andhard corals, and feeds bypecking inthe watercolumn.Collette andTalbot (1972,p. 122)
and Smith andTyler (1972, p. 158) reportedthat it settles down on coral surfaces atnightfall and
disappears from sight until the morning
change-over. However, at night between 20:30 and
22:25, on August 2, 6, and 12 at Turtle Rocks,
Bimini, we observed numerous individuals of C. personatus resting quietly on the surface of Col-pophyllia natans andMycetophyllia lamarckiana, usually in valleys on the coral face. These all showed coloration and color pattern similar to those ofdaytime. Coryphopteruspersonatus
be-came active by daylight, or atleastby07:40,40
minutes after sunrise, hovering in the water
column. We suspect that some individuals ofC.
personatus do retire into holesorcrevicesonthe
reef at night, while others rest motionless but
exposed onlivecoralsurfaces.
RELATIONSHIPS WITHIN THE GENUS CORYPHOPTER US
Bohlke and Robins (1960) recognized six species ofCoryphopterus from the western At-lantic. In 1962, the same authors demonstrated that the species previously called Eviota per-sonata was also a member of this complex and described two additional species, C. hyalinus and C. lipernes, bringing the total number of western
Atlantic species tonine. Coryphopterushyalinus
was said to differ from C. personatus by having less pigment, the anus not in the center of a black pigment spot, and three interorbital pores instead of two. The differences between C. hyalinus and C. personatus are not absolute,
however; in a large series, we have found some intermediates. We therefore accept the species hyalinus with considerable reservation, especially
since we have not been able to separate C.
hy-alinusfromC. personatusinthefield.
These nine species show well-marked habitat differences. Some occur on sandy or silty bot-toms that arelightin color; such species show a reduction in pigmentation. Others are confined
to rocky surfaces and havecorrespondinglymore
melanin pigment so that they have spotted orat least punctate patterns. The hoverers tend to be
reddishin color withablackperiproct.Also
cor-related with hoveringis areduction ofthe
inter-pelvic membrane. Generalizedand derived condi-tions of morphology and pigment are clearly determinable for these gobies and subject to cladisticanalysis.
Our interpretation of therelationships of the Western Atlantic species of Coryphopterus is presented in figure 3. Derived characters used in
the analysisarediscussed with numbers indicated
onthe figure.
1. The synapomorphies (shared specialized characters) that unite this complex are in-cluded among the generic characters that
were enumerated by Bohlke and Robins
1960, 1962). Some of these are
undoubt-edly plesiomorphic (shared generalized
char-acters), but until all other related gobiid
genera have been adequately studied, it will be difficult to becertain of the status of all these characters. We would be surprised, however, if the head-pore pattern and the presence of a fleshy crest on the predorsal midlineis notshowntobederived.
2. Emarginate (rather than round) pelvic disc. In C. glaucofraenum and C. punctipec-tophorous the disc is of the usualrounded gobiid form. In the C. thrix-C. hyalinus line, the fifth rays are shorter than the
fourth so that thedisc is emarginate.
3. Pelvic frenum lost. In C. glaucofraenum-C. thrix thereisawell-developedfrenum (mem-brane) across thebase of the pelvic rays. In C. dicrus-C. hyalinus the frenum is absent.
A frenum occurs ina wide variety of gobies
and we consider its absence to be the
de-rivedcondition.
4. Separate pelvic fins. In C. alloides-C. hy-alinus the pelvic fins are separate,with only
a trace of the connecting membraneat the
very base. In the young of C. lipernes the
membrane is complete, but it isvestigial in adults. There can be little question thatthe loss of the membrane is the derived condi-tion. Bohlke and Robins (1960) figured
the inner (fifth) pelvic rays of C. alloides
as unbranched but in ourspecimen (AMNH 29352) from the Berry Islands, Bahamas,
it is deeply branched, although both
branches areslender.
5. Pigmented periproct. In C. lipernes-C.
hy-alinus there is a conspicuous black ring
un-SMITH AND TYLER: GOBIID FISH
c
iC
12. hyaline sand dweller / \\2. emarginate pelvics 1. 21 characters
FIG. 3. Inferred relationships of the Western Atlantic species of Coryphopterus. For a discussion of thecharactersseetext.
pigmented periprocts, as do most other
fishes.
6. Filamentous dorsal. The presence of pro-duced dorsal-fin spines occurs in both sexes of C. thrix and C. eidolon, although only
small specimens ofC. eidolon have the rays produced. In C. lipernes only males have produced rays, whereas in C. personatus (and presumably hyalinus) both sexes ex-hibit them. We suggest that this feature has been developed and lost repeatedly within the group.
7. Synapomorphies that unite C. personatus
and C. hyalinus are the habit of hovering,
the reddishcoloration,and the characteristic
head markings (the latter being less well developedinC. hyalinus).
8. An autapomorphic feature of C. hyalinus is the presence of two anterior interorbital poresinstead ofone.
9. Autapomorphies for C. lipernes include
yel-low and blue coloration, coral-dwelling habits, and sexual dimorphism inthe dorsal-finspines.
10. Dorsal fin coloration. A prominent bar on
the anterior interradial membranes of the
spiny dorsal is autapomorphic for C. al-loides.
11. A generally speckled color pattern,
partic-ularly with two prominent spots on the pectoral base, is apomorphic for C. dicrus. 12. Sand-dwelling habits. The C. dicrus-C.
hy-alinuslineage shows a progressively increased specialization of the pelvic fin, culminating in their complete separation. A specialized pigment pattern has also appeared. We be-lieve that C. dicrus and C. alloides are spe-cialized for rock-dwelling; dicrus on the basis of our own observations and alloides because of its rarity in collections and from the fact that it is never taken in large
num-bers. Hoverers such as personatus and hyalinus are taken in swarms. We report here thatC. lipernesdwellsinlivecoral.
Coryphopterus thrix, C. eidolon, C. glaucofraenum, and, to a lesser extent, C. punctipectophorus are hyaline forms that live around the base of reefs and feed on minute invertebrates at the surface of the sand bottom. We consider this habit and
as-sociated pigment loss to be asingle apomor-phic character complex withinthe C. thrix-C. hyalinus lineage. An alternative hypothe-sis, which we tentatively reject, is that sand-dwelling has developed once and that emarginate pelvics have developed
inde-peAidently atleasttwice.
13. The filamentous dorsal spines unite C. thrix and C. eidolon. We believe that this is an
independent development and does not in-dicate any relationship with lipernes, per-sonatus or hyalinus, all of which also have prolonged dorsal spines.
14. A large spot at the upper pelvic base is autapomorphicforC. thrix.
15. The yellow pigmentaround thehead and
an-terior partof the bodyisautapomorphic.
16. Anautapomorphy for C. punctipectophorus is the presence ofalarge spot on the lower
part of thebaseof thepectoral fin.
17. Autapomorphic features of C.
glaucofrae-num are polytypic coloration (some indi-viduals pale, some with considerable pig-ment) and the presence of a spot over the
operculum.
18. We are unable to find a synapomorphy to unite C. punctipectophorus with C. glau-cofraenum or with any other Cory-phopterus. On the basis ofpelvic disc
struc-ture, C. punctipectophorus seems to be closest to C. glaucofraenum. We believe, however, that this is a primitive condition and therefore consider the relations of C. punctipectophorus and C. glaucofraenum to be unresolved.
LITERATURE CITED
Bohlke, James E., and C.Richard Robins
1960. A revision of the gobiid fish genus Coryphopterus. Proc. Acad. Nat. Sci.
Philadelphia, vol. 112, no. 5, pp. 103-128.
1962. The taxonomic position of the west
Atlantic goby, Eviota personata, with descriptions of two related species. Ibid., vol. 114,no. 5, pp. 175-189. Collette, Bruce B., and Frank H. Talbot
1972. Activity patterns of coral reef fishes with emphasis on nocturnal-diurnal changeover. In Collette, Bruce B., and
Sylvia A. Earle (eds.), Results of the Tektite program: ecology of coral reef fishes. Los Angeles Co. Mus. Sci. Bull., vol. 14, pp. 98-124.
Colin,Patrick
1975. The Neon Gobies: The comparative
biology of the Gobies of the genus Gobiosoma, subgenus Elacatinus,
Pis-ces: Gobidae) in the tropical
West-ern North Atlantic Ocean. Neptune City, N.J., T. F. H. Publications, Inc.,
304pp. Mariscal, Richard N.
1970. Anexperimental analysis of the protec-tion of Amphiprion xanthurus Cuvier and Valenciennes and some other
anemone fishes from sea anemones.
Jour. Exp. Marine Biol. Ecol., vol. 4,
pp. 134-149.
Smith,C. Lavett, andJamesC.Tyler
1972. Space resource sharing in a coral reef
fish community. In Collette, Bruce B.,
and Sylvia A. Earle (eds.), Results of the Tektite program: ecology of coral
reef fishes. Los Angeles County Mus. Sci. Bull.,vol. 14,pp. 125-170, 1 chart.
1975. Succession and stability in fish
com-munities ofdome-shaped parchreefs in the West Indies. American Museum Novitates,no. 2572,pp. 1-18.
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