Wrasses of the Galapagos Islands, with the description of a new deepwater species of Halichoeres (Perciformes : Labridae)

Abstract

Halichoeres raisneri, new species, is described from three specimens captured by the Johnson Sea Link submersible at 114-125 m off Wolf Island, Galápagos. Distinctive features of the new species include a dorsally projecting fleshy flap along the posterior three-quarters of the upper lip and the absence of a canine tooth at the corner of the upper jaw. The body color of freshly caught females is pale pink with two yellow stripes and five prominent pink spots above the uppermost stripe. Similarities between the new species and other labrids are discussed, and a key to the 16 nominal species of Labridae known from the Galápagos Islands is provided. Decodon melasma is recorded from the Archipelago for the first time.

Full text

Rev. Biol. Trop. 49 Supl. 1: 89-100, 2001
Wrasses of the Galápagos Islands, with the description of a new
deepwater species of Halichoeres (Perciformes: Labridae)
Carole C. Baldwin
1
and John E. McCosker
2
1 Department of Vertebrate Zoology, MRC 159, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A. Fax: 202-357-2986; E-mail: baldwin.carole@nmnh.si.edu
2 California Academy of Sciences, San Francisco, California 94118, U.S.A. Fax: 415-750-7148;
E-mail: jmccosker@calacademy.org
Abstract: Halichoeres raisneri, new species, is described from three specimens captured by the Johnson Sea Link sub-
mersible at 114-125 m off Wolf Island, Galápagos. Distinctive features of the new species include a dorsally projecting
fleshy flap along the posterior three-quarters of the upper lip and the absence of a canine tooth at the corner of the upper
jaw. The body color of freshly caught females is pale pink with two yellow stripes and five prominent pink spots above
the uppermost stripe. Similarities between the new species and other labrids are discussed, and a key to the 16 nominal
species of Labridae known from the Galápagos Islands is provided. Decodon melasma is recorded from the Archipelago
for the first time.
Key words: Labridae, Galápagos Islands, Submersible, New Species, New Records
The nearshore fish fauna of the Galápa-
gos Islands has intrigued ichthyologists and
zoogeographers since Darwin returned to
England with the first specimens for study.
His collection of 15 specimens included a
large wrasse that was later named Cossyphus
d a rw i n i (now Semicossyphus darw i n i
[Jenyns]) in his honor. The 16 labrids known
from the Archipelago represent the fourth
largest family of shorefishes, after 25 ser-
ranids, 23 carangids, and 19 muraenids. Most
of the Galápagos labrid species are variously
distributed from Baja, California, to central
Chile, and occur at one or more of the other
tropical eastern Pacific Islands (Clipperton,
Cocos, Malpelo, Revillagigedos); several
also are known from Indo-Pacific localities.
Only the new species described herein and an
undescribed species of Xyrichtys (Victor and
Wellington 2000) are currently known only
from the Galápagos, and one species,
Xyrichtys victori, is an insular endemic
(Cocos and Galápagos). Based on three
months of scuba and submersible diving
using the Harbor Branch Oceanographic
Institute’s Johnson Sea-Link (JSL) at Galápa-
gos (during November, 1995, a “normal”
period in the eastern Pacific, and June and
July, 1998, a severe El Niño Southern Oscil-
lation period), we are able to add additional
records and observations about Galápagos
labrids.
Grove and Lavenberg (1997) published
the first comprehensive listing and analysis of
Galápagos labrids, along with black and
white drawings and color photographs of
many species. McCosker (1998) corrected
several errors in their work and analysis,
Received: 18-X-2000 Corrected: 23-XI-2000 Accepted: 8-XII-2000

R E V I S TA DE BIOLOGIA T R O P I C A L90
length, “HL,” to head length. Measurements
and most counts follow those of Randall and
Smith (1982). Counts of scale rows between
lateral line and fin origins do not include
small scales at base of fins or the lateral-line
scale. Pored lateral-line scale counts include
the terminal scale on the caudal-fin base pos-
terior to the hypural plate. Counts of verte-
brae, caudal-, dorsal- and anal-fin rays, and
lengths of first anal- and first dorsal-fin
spines were taken from or verified with radi-
ographs. Other measurements were made
with dial calipers to the nearest 0.1 mm. The
branchial skeleton of one paratype was
removed and cleared and stained to facilitate
examination of pharyngeal dentition.
Comparative labrid material examined:
Halichoeres bicolor, USNM 277167, 5 speci-
mens; H. bimaculatus, USNM 358694, 1; H.
biocellatus, USNM 336930, 1; H. bivittatus,
USNM 318887, 2; H. caudalis, U S N M
315469, 1; H. dispilus, USNM 321497, 2;
USNM 321501, 5; H. garnoti, U S N M
318877, 18; H. hortulanus, USNM 345935,
2; H. kallochroma, USNM 327861, 2; H.
margaritaceus, USNM 112976, 9; H. mar -
ginatus, USNM 348651, 1; USNM 332183,
10; H. nicholsi, USNM 321467, 4; H. noto -
spilus, USNM 111466, 111471, 5; H. ornatis -
simus, USNM 333212, 1; H. papilionaceus,
USNM 309333, 2; H. pelicieri, U S N M
222296, 1 paratype; H. podostigma, USNM
277149, 3; H. prosopeion, USNM 333213, 1;
H. scapularis, USNM 332263, 3; Pseudoju -
loides cerasinus, USNM 323918, 1.
Geographical distributions of species pro-
vided in the key to Galápagos labrids include
information from Allen and Robertson
(1994), Bearez (1996), Robertson and Allen
(1996), Grove and Lavenberg (1997), and D.
Ross Robertson (pers. comm.). References
for original descriptions of all labrid species
mentioned herein can be found in Eschmeyer
(1998) or online at the following address:
http://www.calacademy.org/research/ichthy-
ology/catalog/index.html.
based primarily on Randall’s (1995) analysis
of eastern Pacific Thalassoma. Useful keys
to the species of eastern Pacific labrids and
supplementary information may be found in
Bussing (1985, for Costa Rican species) and
Gomon (1995, for eastern Pacific species,
excluding Galápagos). Excellent color pho-
tographs of growth stages of most Galápagos
species are available in Humann (1993) and
Allen and Robertson (1994).
During 1995, the junior author and R.
Grant Gilmore observed and captured from
the JSL the first known Galápagos specimens
of the blackspot wrasse, Decodon melasma
Gomon. They were first observed (7 Nov.
1995, JSL Dive 3941) over sand, rock, and
rock ridge bottoms at 104 m at Española
(=Hood Island) and subsequently collected
(15 Nov. 1995, JSL Dive 3954) from similar
habitats off Cabo Hammond, Isla Fernandina
(=Narborough Island), at 82-105 m. The two
specimens (CAS 210090, 64-95 mm SL) are
typical of the species, which is also known
from the Gulf of California to Peru and Cocos
Island (Allen and Robertson 1994, Gomon,
1995). Another wrasse, resembling Halicho -
eres chierchiae Di Caporiacco, was observed
on 25 July 1998 at North Seymour Island in
Galápagos, but it was not collected or pho-
tographed. The individual, with a prominent
red and blue spot above the posterior tip of
the pectoral fin typical of an adult male (and
from which the common name “wounded
wrasse” is derived), was spotted over a sand
and rock field. We include H. chierchiae,
which is otherwise known from Baja Califor-
nia to Panama (Gomon 1995, pers. obs.), in
the key to Galápagos labrids herein, but fur-
ther investigation is needed to confirm its
presence in Galápagos. Finally, as noted
above, Victor and Wellington (2000) report
on an undescribed species of Xyrichtys from
Galápagos.
MATERIALS AND METHODS
Institutional abbreviations follow Levi-
ton et al. (1985). “SL” refers to standard

Key to the Labridae of the Galápagos Islands
(Modified from Gomon 1995)
1a. Lateral line interrupted (Fig. 1a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1b. Lateral line continuous (Fig. 1b, c), abruptly curved downward in some species . . . . . . . . . . . . . . . . . . . . . . . .4
2a. Dorsal profile of snout not very steep, forming an angle of about 45° with the longitudinal axis of the body (Fig.
2a); top of head and snout somewhat compressed, but not forming a knife-like edge; adults greenish brown with
white spots on each body scale; juveniles green, red, or brown, with white spots on body and head, and 3-4 thin
brown body bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Novaculichthys taeniourus (Lacepède)
(Indo-Pacific; eastern Pacific, from Gulf of California to Panama and the Galápagos and Clipperton Islands)
2b. Dorsal profile of snout steep, forming an angle with longitudinal axis of the body that is noticeably more than 45°
and sometimes almost vertical (Fig. 2b); top of head and snout compressed into a knife-like edge (Xyrichtys
1
) .3
3a. First 2 dorsal-fin spines elongate and well separated from remainder of dorsal fin; body with 4 indistinct broad
brown bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xyrichtys pavo (Valenciennes)
(Indo-Pacific, including Hawaii and Red Sea; Gulf of California to Panama and Galápagos)
3b. First dorsal spines not elongate, not separated from remainder of dorsal fin; terminal phase iridescent blue-green
with scattered black blotches on flanks; initial phase mainly pinkish orange, without blotches or spots; juveniles
white to ye-llowish brown with dark stripe on upper flank . . . . . . . . . . . . . . . . . . . . .Xyrichtys victori Wellington
(Cocos and Galápagos Islands)
4a. Lateral line smoothly curved (Fig. 1b); dorsal-fin spines XI or XII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
4b. Lateral line curved abruptly downward beneath posterior end of dorsal fin (Fig. 1c);
dorsal-fin spines VIII or IX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
5a. Dorsal-fin spines XI; anal fin with 10 segmented rays; predorsal scales reaching a vertical anterior to orbit; col-
oration reddish above, pale below, 3 bright yellow stripes on head and a black blotch above the pectoral fin . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Decodon melasma Gomon
(Gulf of California to Peru, Cocos and Galápagos Islands)
I N T E R N AT I O N A L J O U R N A L OF T R O P I C A L B I O L O G Y AND CONSERVAT I O N 91
Fig. 1. Variation in the shape of the lateral line among eastern Pacific Labridae (from Gomon, 1995).
Fig. 2. Shape of the dorsal profile of the head in (a) Novaculichthys and (b) Xyrichtys (from Gomon, 1995).

R E V I S TA DE BIOLOGIA T R O P I C A L92
5b. Dorsal-fin spines XII; anal fin with 12 segmented rays; predorsal scales reaching or nearly reaching a vertical
through posterior margin of orbit; coloration not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Scaly basal sheath absent on dorsal and anal fins; 53-56 pored lateral-line scales; initial phase red with a pale chin
and a yellow blotch above the pectoral-fin base, terminal phase dark gray with a pale chin and an obvious large yel-
low spot above the pectoral-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Semicossyphus darwini (Jenyns)
(mainland Ecuador to central Peru, and Galápagos)
6b. Scaly basal sheath present on dorsal and anal fins; 31-33 pored lateral-line scales; coloration not as above
(Bodianus ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
7a. Terminal phase with a pronounced bump on forehead and elongate rays in median fins; body coloration of juve-
niles and initial phase yellow, the anterior 2/3 becoming pinkish in adult females, with two black stripes on upper
half of flank, beginning behind eye and extending to caudal base; body coloration of terminal male gray to green
with a faint yellow bar at mid-body, head pinkish, chin white . . . . . . . . . . . . . . . . . . .Bodianus diplotaenia (Gill)
(Baja California to northern Chile, Clipperton, Cocos, Revillagigedos, and Gálapagos Islands)
7b. Terminal phase without an exaggerated forehead bump, median fins not greatly elongated; body coloration of juve-
niles pale yellow to white, with three black stripes on head and body, the middle stripe beginning on the snout and
extending to the caudal fin; body coloration of adults extremely variable, from brown to dark gray to bright orange
and white, overlain with black blotches . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Bodianus eclancheri (Valenciennes)
(mainland Ecuador to central Chile, and Galápagos)
8a. Dorsal-fin spines VIII (Thalassoma) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8b. Dorsal-fin spines IX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
9a. Branched pectoral-fin rays 13; caudal fin truncate; juveniles, females, and primary males with lengthwise bands of
yellowish brown and red on head and body; secondary males with a greenish purple head, a broad yellow vertical
band in the thoracic region, and the remainder of the body greenish blue; maximum size 15 cm SL . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thalassoma lucasanum (Gill)
(Gulf of California to mainland Ecuador, Cocos, Galápagos, and Malpelo)
9b. Branched pectoral-fin rays 14; caudal fin of terminal phase with elongate lobes; coloration green or blue-green with
2-3 radiating lines from eye; adults larger, may attain 43 cm SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Coloration green or blue-green with a slender red streak on each scale; head pink with green lines behind and
beneath eye; may attain 24 cm SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thalassoma grammaticum Gilbert
(Gulf of California to Panama, Clipperton, Cocos, Galápagos, and Revillagigedos Islands)
10b. Coloration green, with red stripes and vertical lines along body, and red lines behind eye and on snout and fore-
head; may attain 43 cm SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thalassoma purpureum (Forsskål)
(Indo-Pacific eastward to Clipperton, Cocos, Galápagos, and Panama)
11a. Anterior jaw teeth not enlarged, most teeth incisor-like (Fig. 3a); initial phase gray, with fine white speckling on
upper half and 2 black spots over caudal-fin base; terminal phase greenish dorsally, pale ventrally, with thin blue
stripes before and behind eye, and a red-orange patch above pectoral base . . . . Stethojulis bandanensis (Bleeker)
(western Pacific to Costa Rica, Panama, Clipperton, Cocos, and Galápagos Islands)
Fig. 3. Dentition in (a) Stethojulis and (b) Halichoeres (from Gomon, 1995).

I N T E R N AT I O N A L J O U R N A L OF T R O P I C A L B I O L O G Y AND CONSERVAT I O N 93
to ventrally projecting flap on lower lip; no
canine tooth at corner of upper jaw; body
color (of freshly caught female) pale pink
with two prominent yellow stripes, one from
tip of snout to upper caudal-fin base, the other
from pectoral-fin base to lower caudal-fin
base; five prominent pink spots above upper
yellow stripe, dispersed along lateral line; no
dark blotch on caudal-fin base; lateral-line
complete, abruptly deflected downward
below posterior end of dorsal fin; 27 tubed
scales in series, tubes simple and with single
opening; ultimate pored scale located on cau-
dal-fin base; dorsal-fin rays IX,12; anal-fin
rays III,12, first spine visible without dissec-
tion; head naked; scales on chest region
smaller than other body scales; upper and
lower jaws each with a pair of canine teeth
near symphysis and with a series of conical
Halichoeres raisneri, new species
Figs. 4-5, Table 1
Holotype: USNM 357795, 90.0 mm SL,
female, SE Wolf Island, Galápagos,
01°23.2´N, 91°48.6´W, 115 m, JSL II Sta.
3087, C. Baldwin and J. Gomezjurado, 22
June 1998.
Paratypes: CAS 209810, 78.8 mm SL,
female, and 39.8 mm SL, sex undetermined,
SE Wolf Island, Galápagos, 01°23.2´N,
91°48.6´W, 114-125 m, JSL II Sta. 3086, C.
Baldwin and J. McCosker, 22 June 1998.
Diagnosis: Halichoeres raisneri is dis-
tinguishable from other species of the genus
on the basis of the following combination of
characters: posterior three-quarters of upper
lip with dorsally projecting fleshy flap similar
11b. Anterior jaw teeth enlarged and canine-like (Fig. 3b); color not as above (Halichoeres) . . . . . . . . . . . . . . . .….12
12a. Upper jaw with one or more prominent canines at corner of mouth; dorsal fin with 11 segmented rays . . . . . .13
12b. Upper jaw without prominent canines at corner of mouth; dorsal fin with 11 or 12 segmented rays . . . . . . . . 15
13a. Each lateral-line scale usually with only a single pore; body coloration pink to orange (live individuals capable of
turning blue) with distinct black spot above middle of pectoral fin. . . . . . . . . . . .Halichoeres dispilus (Günther)
(Gulf of California to Peru, Galápagos, and Cocos)
13b. Each anterior lateral-line scale usually with 2-3 pores; coloration not as above . . . . . . . . . . . . . . . . . . . . . . . .14
14a. Terminal phase greenish blue with green bars above midbody, yellow to greenish yellow below, purple midlater-
ally with a prominent red and blue spot above pectoral fin tip; initial phase greenish with purple bars above, yel-
low below, an orange reticulated pattern midlaterally and a black spot on segmented portion of dorsal fin . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halichoeres chierchiae Caporiacco
(Baja California to Panama and Galápagos)
14b. Terminal phase greenish blue with a diffuse black bar on upper body near distal tip of pectoral fin, a prominent yel-
low blotch preceding it; initial phase pale yellow with irregular dark blotches on sides and a large dark green ocel-
lus on dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Halichoeres nicholsi (Jordan & Gilbert)
(Gulf of California to mainland Ecuador, Revillagigedos, and Galápagos)
15a. Segmented dorsal rays 12; a prominent fleshy skin flap at posterior end of upper lip; coloration of living females
pale pink with 2 prominent yellow body stripes and 5 prominent pink spots above uppermost yellow stripe . . . ..
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Halichoeres raisneri n. sp.
(Wolf Island, Galápagos)
15b. Segmented dorsal rays 11; upper lip may be creased, but prominent posterior skin flap absent; coloration of initial
phase dark green above, light green below, with 5-6 alternating yellow and black patches at base of dorsal fin; ter-
minal phase with 7-8 blackish bars on upper half of body separated by narrow yellow bars, a blackish patch behind
pectoral fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Halichoeres notospilus (Günther)
(Gulf of California to Peru, Galápagos and Revillagigedos Islands)
1
not included is an undescribed species of Xyrichtys from Galápagos (Victor and Wellington 2000)

R E V I S TA DE BIOLOGIA T R O P I C A L94
Fig. 4. Top: Holotype of Halichoeres raisneri, new species, USNM 357795, 90.0 mm SL, Wolf Island, Galápagos
Islands. Middle: Paratype of H. raisneri (prior to conservation), CAS 209810, 78.8 mm SL. Bottom: In situ photograph
of H raisneri, JSLII Sta. 3086.

I N T E R N AT I O N A L J O U R N A L OF T R O P I C A L B I O L O G Y AND CONSERVAT I O N 95
large, forward-projecting canine teeth near
symphysis, lower pair sliding between upper
pair when mouth closed. A single row of con-
ical teeth in both jaws posterior to canine
teeth, about 9 in upper jaw, 10 in lower jaw –
teeth of 78.8-mm SL paratype damaged and
counts not included; single row of very small
nodular teeth in single row posterior to coni-
cal teeth, about 6 in upper jaw in holotype, 8
in lower jaw – nodular teeth in paratypes, if
present, small and difficult to discern. A few
tiny teeth present in both jaws behind canine
teeth and medial to anteriormost conical
teeth.
Pharyngeal dentition of 78.8-mm SL
paratype comprising paired upper third-pha-
ryngeal tooth plates and median lower tooth-
plate formed by fused fifth ceratobranchials.
Upper tooth plates roughly triangular, with 17
teeth in five transverse rows; first (anterior-
most) row with a single tooth, second with
three, third with four, fourth with five, fifth
(posteriormost) with four; size of teeth
decreasing posteriorly; all teeth roughly con-
ical but with broad bases and slender, curved
tips. Lower pharyngeal plate a shallow “Y”
shape in dorsal view with teeth covering all
but the distal upper arms of the “Y.” Posteri-
or end of stem with a single row of three
teeth, broadening to two rows of ca. four teeth
anteriorly on stem; curved part of plate with
teeth posterior to canines; free margin of ver-
tical preopercular limb smooth, reaching dor-
sally to horizontal through ventral edge of
eye, free margin of horizontal preopercular
limb smooth, reaching anteriorly to vertical
through front of eye; snout long, length 26-
29% HL; body slender, depth 20-22% SL;
and caudal peduncle narrow, depth less than
or approximately equal to length.
Description: Data for morphometric
features are given in Table 1. Dorsal-fin rays
IX,12; anal-fin rays III,12; pectoral-fin rays
12; principal caudal-fin rays 8+7 (8+8 seg-
mented rays), procurrent caudal-fin rays 5+4
(5+5 unsegmented rays); vertebrae 10+15;
lateral-line scales 27; scale rows between lat-
eral line and dorsal-fin origin 2; scale rows
between lateral line and anal-fin origin 7;
suborbital pores (from posterior point of orbit
to and including lacrimal) 9; gill rakers 16-18
(18 in holotype, 16 in 78.8-mm SL paratype,
too small to accurately count in 39.8-mm
paratype).
Mouth terminal and of moderate size,
upper jaw reaching vertical through anterior
nostril. Lips fleshy, lower lip with a broad
flap projecting ventrally, posterior three-quar-
ters of upper lip with more slender flap pro-
jecting dorsally (Fig. 5A). Inner surface of
lips near base of teeth strongly papillose.
Upper and lower jaws each with a pair of
Fig. 5. Right lateral view of head. (a) Halichoeres raisneri, holotype, USNM 357795, 90.0 mm SL; (b) Halichoeres
dispilus, USNM 321501, 106 mm SL.

R E V I S TA DE BIOLOGIA T R O P I C A L96
Lateral line complete, running closely
along dorsal-fin base anteriorly, abruptly dip-
ping ventrally beneath posterior end of soft
dorsal fin and then leveling out to a straight
line mid-laterally on caudal peduncle. Scales
20-23 in the series containing the abrupt cur-
vature of the lateral line. Lateral-line scales
each with a single pore.
Head naked, body scales terminating on
predorsal region just posterior to a vertical
through upper limb of preopercle. Ventrally,
scales extending anteriorly to posterior end of
isthmus. Scales cycloid, posterior margin
bluntly pointed. Scales on most of body
large, greatest height (in dorsoventral plane)
ca. 15-17% HL; scales on abdomen smaller
and decreasing in size anteriorly, the most
anterior ones ca. 4-5% HL in height. Dorsal,
anal, and pectoral fins naked. A single large
about four rows of teeth, those in posterior-
most row largest; all teeth roughly conical,
similar in shape to those of upper plates (i.e.,
somewhat broad at base, more slender distal-
ly).
Nostrils anterior to upper quarter of
orbit. Anterior nostril in short tube; posterior
nostril partially covered by small membra-
nous flap from anterior edge. Distance
between anterior and posterior nostrils about
half the distance between posterior nostril
and orbit. Distance between anterior nostril
and tip of snout approximately equal to diam-
eter of orbit.
Margins of all bones of opercular series
smooth; opercle terminating posteriorly in
bluntly pointed flap, the posterior tip reaching
a vertical through base of third dorsal-fin
spine; outer edge of opercular flap membra-
nous.
TABLE 1
Data on morphometric features of type specimens of Halichoeres raisneri. Standard length is in mm; other measure -
ments, in percentage of standard length.
Holotype Paratypes
USNM 357795 CAS 209810
Standard length (mm) 90.0 78.8 39.8
Depth of body 22 21 20
Width of body 11 10 8.8
Head length 32 31 32
Snout length 9.4 8.8 8.3
Orbit diameter 6.1 6.0 7.0
Body interorbital width 4.7 4.4 4.5
Length of upper jaw 7.4 6.6 7.0
Least depth of caudal peduncle 10 10 11
Length of caudal peduncle 12 10 12
Predorsal length 29 28 31
Length of dorsal-fin base 65 62 60
Length of depressed dorsal fin 75 71 72
Length of first dorsal-fin spine 5.6 6.6 7.6
Length of ninth dorsal-fin spine 10 11 10
Length of anal-fin base 34 37 36
Length of depressed anal fin 41 43 43
Length of first anal-fin spine 2.8 3.8 3.3
Length of third anal-fin spine 7.0 7.9 8.3
Length of upper segmented
caudal-fin rays 21 20 23
Length of central segmented
caudal-fin rays 20 19 23
Length of pectoral fin 18 16 19
Length of pelvic fin 15 13 13

I N T E R N AT I O N A L J O U R N A L OF T R O P I C A L B I O L O G Y AND CONSERVAT I O N 97
yellow patch between lilac stripe and posteri-
or lilac spot; a pale yellow stripe beneath and
behind eye; upper jaw pink. Dorsal and anal
fins mostly transparent — distal one-third of
fins yellow, fin tips transparent; caudal fin
yellow, outermost principal rays notably so;
pectoral fin transparent; pelvic-fin spine yel-
low to orange, rest of fin transparent.
Other Material: After this paper was
accepted for publication, the junior author
learned that William Bussing of the Museo de
Zoologia, Universidad de Costa Rica, had in
his possession an unidentified adult male
wrasse from Cocos Island that resembled our
new species from Galápagos in having a long
pointed snout. Bussing compared his speci-
men (UCR 2127.005, 143 mm SL,
5°34’30”N, 87°2’35”W) to our description
and photographs of H. raisneri and suggested
it is probably the male of the new species.
Subsequent examination of the Cocos speci-
men by the senior author corroborates the
similarity, but in the absence of female speci-
mens from Cocos or male specimens from
Galápagos, we are reluctant to designate the
Cocos specimen as type material for H. rais -
neri. The Cocos specimen has most of the
diagnostic features of H. raisneri, including
the fleshy flap on the upper lip; no canine
tooth at the corner of the upper jaw; a com-
plete lateral line, abruptly deflected down-
ward at ca. the twentieth scale of the series,
the last scale on the caudal-fin base; IX,12
dorsal-fin rays, III,12 anal-fin rays, a long
snout (33% HL), and slender body (depth
23% SL). The Cocos specimen differs from
type specimens of H. raisneri in having fewer
lateral-line scales (scales missing on both
sides, but ca. 25 on left side of specimen vs.
27 in H. raisneri). In alcohol, the Cocos
specimen is light brown with darker areas
along the base of the dorsal fin and along the
lateral midline. There are three postorbital
lines of pigment on the head, a small patch of
pigment in front of the dorsal portion of the
eye, a line of pigment between the anteroven-
tral portion of the eye and the posterior end of
the upper jaw, and a line of pigment along the
scale extending posteriorly between bases of
right and left pelvic fins. Several rows of
scales on caudal-fin base, scales becoming
progressively smaller posteriorly.
Caudal fin slightly rounded, length 1.6
in HLin holotype (1.4-1.6 in paratypes). Ori-
gin of dorsal fin slightly anterior to vertical
through base of pectoral fin (over second lat-
eral-line scale). Dorsal fin long, 1.5 (1.6-1.7)
in SL. Pelvic fin short, falling well short of
anus, 2.2 (2.3-2.5) in HL. Pectoral fin 1.8
(1.7-1.9) in HL, anal-fin base 3.0 (2.7-2.8) in
SL.
Color of holotype in ethyl alcohol pale
with no distinctive markings. Tiny dots of
dark pigment scattered on head and mid- to
upper body, heaviest on lacrimal, opercle, and
occiput, and along base of dorsal fin where
they form ca. 11 faint spots, the last one at
base of last dorsal soft ray. No pigment on
fins. The 78.8-mm SL paratype also with a
poorly demarcated line of melanophores
extending posteriorly from posterior tip of
opercle to base of upper caudal-fin lobe and
with a faint spot on base of fifth dorsal soft
ray. In the 39.8-mm SL paratype, the spot on
fifth dorsal soft ray and the lines of pigment
along dorsal-fin base and between opercle
and upper caudal-fin lobe more prominent,
the latter terminating in a relatively large dif-
fuse spot on caudal fin.
Color of female immediately after cap-
ture: body pink, with two prominent yellow
stripes, upper one from tip of snout to upper
caudal-fin base, lower one from pectoral-fin
base to lower caudal-fin base; between yel-
low stripes, the pink body color forming
another stripe that is edged dorsally and ven-
trally with orange scalloping. Above upper
yellow stripe, five pale pink spots dispersed
along and lying directly on lateral line.
(From the elevated vantage point of observers
in the submersible, darker areas between the
pale spots appeared as a series of saddles on
the back.) Head with a lilac stripe along dor-
sal midline of frontal region and between
orbits, a lilac spot at tip of snout, and another
just anterior to first dorsal-fin spine; a small

R E V I S TA DE BIOLOGIA T R O P I C A L98
three-quarters of the upper lip (Fig. 5A) sim-
ilar to the ventrally projecting flap on the
lower lip typical of all Halichoeres (as in H.
raisneri, Fig. 5A, andH. dispilus, Fig. 5B). In
some species of Halichoeres (we did not
examine all species), the upper lip may be
very fleshy and variously creased, but among
those examined only H. raisneri has a dis-
crete flap of skin that can be lifted at its distal
edge and folded back towards the gape. Hali -
choeres raisneri also lacks a feature diagnos-
tic of most species of the genus, a prominent
canine tooth at the posterior end of the upper
jaw at the level of the buccal commissure.
Randall (in litt.) informed us that in some
species of Halichoeres, including the type
species H. bimaculatus, this tooth is not fully
developed until the adult stage, but we doubt
that its absence in H. raisneri is ontogenetic
because the two largest specimens are ripe
females. This canine tooth also is lacking in
several other eastern Pacific species of Hali -
choeres: H. adustus, H. notospilus, H. insu -
laris, H. malpelo, and H. melanotis (Allen
and Robertson 1994, Gomon 1995). Possibly,
the absence of the canine tooth in those
species and H. raisneri warrants recognition
of a separate genus. Randall and Condé
(1989) erected Frontilabrusfor a wrasse from
the Maldives that they noted is very similar to
Halichoeresexcept that it lacks a canine tooth
at the rear of the upper jaw, has a steep dorsal
profile of the head, and has two pairs of
canine teeth (vs. one) at the front of the lower
jaw. Halichoeres raisneri has a relatively
shallow dorsal head profile and a single pair
of canine teeth at the front of the lower jaw.
We note, however, that H. insularis, known
only from the Islas Revillagigedos, lacks the
canine tooth at the rear of the upper jaw and
has two pairs of canine teeth at the front of
the lower jaw (Allen and Robertson 1994).
Among other labrid genera, H. raisneri
appears most similar to P s e u d o j u l o i d e s
F o w l e r, especially in having an elongate
body. Pseudojuloides differs from Halicho -
eres most notably in having small, chisel-like
incisiform teeth along the sides of the jaws
dorsal midline of the head that extends to the
base of the spinous dorsal fin. There are three
wavy lines of pigment on the membrane of
the dorsal fin, each extending from the first
spine to approximately the sixth segmented
ray. The posterior tips of the caudal-fin rays
are pigmented, pigment extending several
millimeters anteriorly on the uppermost and
lowermost rays of the dorsal and ventral
lobes, respectively. Bussing (in lit., 4 July
2000) indicated that the color of the fish was
faded when he received it.
E t y m o l o g y : Named r a i s n e r i f o r
William R. Raisner, Jr., a veteran pilot who
lost his life in a tragic ultralight plane acci-
dent on 26 June 1998 during the expedition
on which the new species was collected. Bill
was as enthusiastic about our underwater
dives in the submersible as about his aerial
endeavors, and it is in the spirit of a shared
passion for venturing beyond the earth’s sur-
face that we name in his honor and memory a
new species collected from submersible
“flights.” We give H. raisneri the common
name “five-spot wrasse” in reference to the
pale pink spots along the lateral line.
Locality: Halichoeres raisneri is known
only from the type locality, off Wolf Island,
Galápagos (but see comments under “Other
Material” above).
Habitat: The three type specimens were
collected at depths of 114-125 m on a sand
bottom characterized by little rock rubble.
Several other individuals were observed,
often in pairs, but we did not observe signifi-
cantly larger (and presumably male) individ-
uals. One individual was seen taking refuge
in a broken sea-urchin test, but most were
observed over sand. When disturbed by
motions of the mechanical collecting arm of
the sub, the fishes would dive head first into
the sand.
R e m a r k s : Although very similar in
most aspects of its morphology to other
species of Halichoeres, and keying out to that
genus with the labrid keys of Randall (1986)
and Gomon (1995), H. raisneri differs in hav-
ing a dorsally projecting flap on the posterior

I N T E R N AT I O N A L J O U R N A L OF T R O P I C A L B I O L O G Y AND CONSERVAT I O N 99
Halichoeres raisneri also is similar in
color pattern to another deepwater labrid,
Polylepion russelli from the Hawaiian and
Ryukyu Islands (Gomon and Randall 1978).
The latter differs from H. raisneri in having
three (vs. two) yellow stripes, the dorsalmost
running along the base of the dorsal fin. The
two species are markedly different in other
morphological aspects (e.g., counts of fin
rays, scales, and vertebrae - Gomon and Ran-
dall 1978).
ACKNOWLEDGMENTS
Many individuals have assisted us in this
project as well as with the operation of the
Harbor Branch Vessel R/V Seward Johnson
and its submersible, the Johnson Sea-Link. In
particular we wish to thank Don Liberatore,
ace sub pilot, and the staff of Harbor Branch
Oceanographic Institution. Special thanks
are due Julie Piraino of the Smithsonian Insti-
tution’s Marine Station at Link Port, FL, and
David Pawson of the Smithsonian’s National
Museum of Natural History for helping us
prepare for collecting on a ship docked in Ft.
Pierce, FL, while we were in Washington,
D.C., and San Francisco, CA. Martin Gomon
allowed us to modify his labrid key, and
Grant Gilmore provided information on
Decodon. William Bussing allowed us to
examine and report on the unidentified UCR
labrid specimen from Isla del Coco, and made
counts, measurements, and illustrations of
that specimen available to us. Bill also pro-
vided helpful comments on the manuscript.
Julie Mount and Sara Pringle improved our
digitized color images in Figure 4. For all of
the above, we are grateful. For assistance and
permission to study in Ecuador, we thank:
Eliecer Cruz, Arturo Izurieta Valery and
Eduardo Amador, Parque Nacional Galápa-
gos; and Robert Bensted-Smith and Chantal
Blanton, Estacíon Cientifica Charles Darwin.
We also thank Al Giddings of Images Unlim-
ited, David Clark, Inc., the David and Lucile
Packard Foundation, the Discovery Channel,
Ocean Films, the Smithsonian Institution’s
vs. conical ones in H a l i c h o e res ( R a n d a l l
1986). Although the elongate body of H.
raisneri is within the extremes for both gen-
era (Randall 1986), it has conical teeth in the
jaws. Further study of the limits of Halicho -
eres and related genera is needed, and pend-
ing a cladistic revision of labrid genera that
includes all species of Halichoeres, we assign
raisneri to Halichoeres.
Among Halichoeres known from Galá-
pagos, H. raisneri shares with H. notospilus
the absence of a canine tooth at the posterior
end of the upper jaw, but H. raisneri has 12
segmented dorsal rays (vs. 11 in H. noto -
spilus), and the two species have very differ-
ent body coloration. Although somewhat
similar in coloration to H. dispilus, H. rais -
neri differs in having 12 segmented dorsal
rays (vs. 11 in H. dispilus), no dark spot on
the lateral line below the anterior part of the
dorsal fin, no canine tooth at the rear of the
upper jaw, fewer infraorbital sensory pores,
and a fleshy flap on the upper lip. Halicho -
eres nicholsi and H. chierchiae are distinct
from H. raisneri in, among many other fea-
tures, having two or three (vs. one) pores in
each lateral-line scale.
The color pattern of the 90-mm SL
female holotype of H. raisneri is similar to
that of the 86.4-mm SL female holotype and
50-mm SL female paratype of H. pelicieri
from Mauritius (Randall and Smith 1982:
Plate 5C-D). Halichoeres raisneri differs
most notably in lacking a large black spot at
the posterior end of the yellow stripe extend-
ing from the snout to the caudal-fin base, and
in having five prominent pink spots along the
lateral line (Randall and Smith’s photographs
of H. pelicierishow several faint yellow spots
along the lateral line in the paratype). The
two species also have similar or identical
numbers of fin rays, gill rakers, and lateral-
line scales, but they are separable on the basis
of numbers of dorsal- and anal-fin soft rays
(12 in both fins in H. raisneri, 11 in H.
pelicieri), and the absence of a canine tooth at
the buccal commissure and presence of a
fleshy flap on the upper lip in H. raisneri.

R E V I S TA DE BIOLOGIA T R O P I C A L100
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Hawaiian fishes of the labrid tribe Bodianini. Bull.
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Galápagos Islands. Stanford University, Stanford.
863 p.
Humann, P. 1993. Reef fish identification Galápagos.
New World Pub., Jacksonville-Libri Mundi, Quito.
192 p.
Leviton, A. E., R. H. Gibbs, Jr., E. Heal, & C. E. Daw-
son. 1985. Standards in herpetology and ichthyol-
ogy: part I. Standard symbolic codes for institu-
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McCosker, J. E. 1998. (Review of) The fishes of the
Galápagos Islands by J. S. Grove & R. J. Laven-
berg, 1997, Stanford University. Copeia 1998:809-
812.
Randall, J. E. 1986. Family No. 220: Labridae, p. 683-
706 In: Smith, M. M.& P. C. Heemstra (eds.).
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Randall, J. E. 1995. On the validity of the eastern Pacif-
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Discovery Center Project, and the IMAX cor-
poration, Inc., for grants and other assistance.
We are grateful to Mandalay Media Arts,
LLC, Smithsonian Institution’s Discovery
Center Project, and IMAX Corporation for
contributing funds to publish the color
images (Fig. 4).
RESUMEN
Se describe una nueva especie, Halichoeres rais -
neri, capturada a 114-125 m de profundidad en el archip-
iélago de Galápagos. Se presenta una clave para las 16
especies nominales de Labridae de Galápagos y se reg-
istra por vez primera Decodon melasma.
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