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Redescription of Callogobius clarki (Goren) (Teleostei: Gobiidae), Not a
Synonym of C. bifasciatus (Smith)
Naomi R. Delventhal
1
and Randall D. Mooi
1,2
Redescription of Callogobius clarki (Goren) (Teleostei: Gobiidae), Not a
Synonym of C. bifasciatus (Smith)
Naomi R. Delventhal
1
and Randall D. Mooi
1,2
Callogobius clarki (Goren) is redescribed based on 21 specimens from the Red Sea. This species was originally described
from a single specimen collected from the Red Sea, but subsequent workers have placed it in synonymy with the
Western Indian Ocean species C. bifasciatus (Smith). However, it differs from C. bifasciatus in lacking all head pores, and
having a lower range in several meristic values (pectoral-fin rays usually 15–17 vs. 16–18, lateral scale rows usually 33–39
vs. 39–46, predorsal scale rows 12–15 vs. 16–20, transverse scale rows usually 17–18 vs. 18–22, and transverse
mandibular papillae rows usually 11 vs. 12). Within the genus, C. clarki is a member of the sclateri group, exhibiting a
female urogenital papilla with small distal flaps of skin, elongate ctenii on some caudal peduncle scales, and
preopercular sensory papillae rows not continuous with the transverse opercular rows.
C
ALLOGOBIUS clarki (Goren, 1978) was originally
described as a species of Drombus from a single
specimen collected from the Red Sea. In that
description, Goren (1978) compared C. clarki with two other
Western Indian Ocean and Red Sea gobies then assigned to
Drombus,D. irrasus Smith and D. plumatus Smith (later
placed in Callogobius [Goren, 1980]). Goren did not initially
compare C. clarki with other species of Callogobius.
Although Smith (1958) had described C. bifasciatus (Smith)
and C. flavobrunneus (Smith) in the Western Indian Ocean
one year previous to the description of C. irrasus and C.
plumatus, he had placed them in a different genus,
Mucogobius, and a different family, Eleotridae, presumably
because of the separate pelvic fins in the former two species
(Goren, 1980). The condition of the sensory pores on the
head of C. clarki was not specifically noted by Goren (1978,
1979, 1980), although a key to Red Sea Callogobius used this
feature to differentiate C. clarki from all other local species
(Goren et al., 1991). Randall et al. (1994) and Randall (1995)
synonymized C. clarki with C. bifasciatus without comment.
During the first author’s study of taxonomy of Callogobius,
a striking trend was noted. All specimens of C. bifasciatus
(following Randall et al., 1994) from the Red Sea lacked
sensory pores, but pores were present on every specimen
from outside the Red Sea. Examination of the types of C.
clarki (HUJ 10065) and C. bifasciatus (SAIAB 235) confirmed
the absence and presence of sensory pores in these two
specimens, respectively. This suggested that C. clarki was a
distinct and valid species endemic to the Red Sea as
suggested by Goren et al. (1991), and led to a search for
other possible distinguishing characters between C. clarki
and C. bifasciatus. In this paper, we redescribe C. clarki
(Goren, 1978), with particular attention given to a compar-
ison with C. bifasciatus (Smith, 1958). In addition to sensory
pore distribution, we provide evidence that C. clarki and
C. bifasciatus exhibit different (but overlapping) ranges for
several meristic values; morphometric characteristics exam-
ined did not differ between the two species.
MATERIALS AND METHODS
Institutional abbreviations are as listed at http://www.asih.
org/node/204 (Sabaj Perez, 2010) and in Fricke and
Eschmeyer (2012). Methods for morphometric and meristic
data collection follow Delventhal and Mooi (2013). Mor-
phometric data were limited to specimens that had reached
sexual maturity and that had not become compressed,
folded, or otherwise distorted during preservation and
storage. Data reported in the text are followed by the
number of specimens exhibiting that feature; the holotype
condition is indicated by an asterisk. Bilateral observations
were used when possible. Terminology for sensory papillae
rows follows Delventhal and Mooi (2013). In the redescrip-
tion we have commented only on papillae rows whose
length and orientation are variable among species of
Callogobius and generally consistent within a species.
Callogobius clarki (Goren, 1978)
Clark’s Flapheaded Goby
Figures 1–3; Tables 1–5
Drombus clarki Goren, 1978:200–201, fig. 6.—Goren,
1979:36–37, fig. 19, table 1 [description]; Golani,
2006:34 [type list].
Callogobius new species A.—McKinney, 1980:2, fig. 1 (draw-
ing reproduced here as Fig. 3).
Callogobius clarki (Goren).—Goren, 1980:213, 216, tables 1,2;
Dor, 1984:241 [list]; Goren et al., 1991:300 [key]; Goren
and Dor, 1994:63 [list]; Delventhal and Mooi, 2013:162–
163 [species comparison, relationships].
Callogobius bifasciatus (non Smith, 1959). —Randall et al.,
1994:240–241, fig. 8 [in part, description, synonymy of
C. clarki with C. bifasciatus]; Randall, 1995:300, fig. 937 [in
part, description, synonymy of C. clarki with C. bifascia-
tus]; Golani and Bogorodsky, 2010:46 [list].
Holotype.—HUJ 10065 (ex HUJ 56976), 1, 36.5 mm SL,
female, Red Sea, Egypt (formerly Israel), Et Tur, Gulf of Suez,
HUJ team, 1972.
Non-type material.—20 specimens. BMNH 1978.9.8.6, 1,
38.8 mm SL, male, Red Sea, Sudan, Mersa Ar-Rakiyai,
‘‘Manihine’’, 1951; ROM 50227, 1, 34.9 mm SL, male, Red
Sea, Egypt (formerly Israel), Tiran Island, Foul Bay, M.
Goren, 1981; USNM 220031, 11, 14.1–35.1 mm SL, 4 male, 3
female, 1 juvenile, 3 specimens cleared and stained (CS), Red
1
Department of Biological Sciences, 212B Biological Sciences Bldg., University of Manitoba, Winnipeg MB, R3T 2N2 Canada; E-mail:
naomi.delventhal@gmail.com.
2
The Manitoba Museum, 190 Rupert Ave., Winnipeg MB, R3B 0N2 Canada; E-mail: rmooi@manitobamuseum.ca. Send reprint requests to
NRD at this address.
Submitted: 11 July 2013. Accepted: 6 November 2013. Associate Editor: T. Grande.
F2014 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CI-13-078
Copeia 2014, No. 1, 143–148
Sea, Egypt, Gulf of Aqaba, bay at El Himeira, V. Springer
et al., 1969; USNM 220038, 2, 34.6–48.6 mm SL, males, Red
Sea, Eritrea (formerly Ethiopia), Melita Bay (mouth of Melita
Bay), V. Springer et al., 1969; USNM 220090, 1, 43.7 mm SL,
male, Red Sea, Egypt, Reef near road at Marsa Muqabla
[5Marsa el Muqabila], approx. 29u229N, 34u479E, NW of
coast Gulf of Aqaba, V. Springer et al., 1969; USNM 296954,
2, 16.8–24.2 mm SL, 1 male, 1 juvenile, Red Sea, Egypt, Strait
of Jubal, NW edge of Sha’b al Fanadir reef, 27u179230N,
33u489520E, H. Fehlmann et al., 1965; USNM 341181, 2,
31.0–47.9 mm SL, 1 male, 1 female, Red Sea, Egypt, Strait of
Jubal, small reef ca. 300 yds. north of pier at Institute of
Oceanography & Fisheries, Ghardaqa, 27u169380N,
33u479010E, H. Fehlmann et al., 1965.
Diagnosis.—Callogobius clarki is distinguished from all other
described species of Callogobius by the following combina-
tion of characters: head pores absent, 33–41 scales in lateral
series, and preopercular papillae row (Row 20) not contin-
uous with transverse opercular papillae row (Row 21).
Description.—See Table 1 for selected morphometric values,
with comparisons to values for C. bifasciatus. See Tables 2–4
for meristic values. Starred values indicate those of the
holotype. Dorsal-fin elements VI+I,10 (19*), VI+I,11 (1);
anal-fin elements I,9 (20*); pectoral-fin elements 12 (1), 15
(3), 16 (27*), 17 (9); pelvic-fin elements I,5 (40*); un-
branched and branched principal caudal-fin rays i8+7i (18*),
i8+8 (2); procurrent rays 4+4 (5), 4+5 (1), 5+4 (5), 5+5 (8*),
6+6 (1); scales in lateral series 33 (2), 34 (2), 35 (8), 36 (5), 37
(7*), 38 (6), 39 (2), 41 (1); scales in predorsal series 12 (1), 13
(3), 14 (7), 15 (6*); scales in transverse series 17 (9), 18 (7*),
19 (1).
Teeth and tongue based on three cleared-and-stained
specimens from USNM 220031. Teeth of upper jaw conical,
slender and slightly curved inward; outer row enlarged and
spaced at a distance of about 2/3 to 1/3 the height of the
tooth; about one to four tightly spaced inner rows of smaller
teeth (in any single specimen, one posterior row gradually
increasing to about four inner rows anteriorly). Teeth in
lower jaw conical, slender and slightly curved inward; an
anterior outer row with about 3–5 somewhat enlarged teeth
on each side; a mid-posterior innermost row of enlarged
teeth (some of these greatly enlarged); one to four rows of
smaller teeth extending between and posterior to the
Fig. 1. Callogobius clarki, holotype, HUJ 10065, 36.3 mm SL female: (A) left lateral view; (B) right lateral view.
Fig. 2. Female urogenital papilla of Callogobius clarki, USNM 341181,
31.0 mm SL, ventral view. Note the broad lateral flaps of skin. An 5
anus; AS 5anal-fin spine; LF 5lateral flap; UP 5urogenital papilla.
Scale bar 1 mm.
144 Copeia 2014, No. 1
enlarged outer and inner rows. Tongue rather slender, tip
rounded.
Anterior nostril a slender tube, may reach outer edge of
upper lip in some specimens; posterior nostril a short,
upright tube.
Pelvic fins partially joined, membrane probably reaching
between 2/3–5/6 length of fin (all available specimens with
torn pelvic fins); fifth pelvic-fin ray shortened, about nine-
tenths length of fourth pelvic-fin ray on holotype, frequent-
ly shorter on other specimens; all pelvic-fin rays branched.
Frenum present, relatively thin and fragile (weak).
Urogenital papilla with little or no pigment, long and
tapering in males; broad with distal flaps of skin in females
(Fig. 2).
Pores absent. Individual pore replacement papillae (or
occasionally short ridges of up to 4 papillae) present in
interorbital, temporal, and preopercular regions.
Head papillae row configuration (Fig. 3): postnasal rows
(Row 2) long and joined across the midline (17*), or
occasionally separate (2); anterior suborbital row (Row 9)
moderately long to long, approaching eye (38*); mid-
suborbital row (Row 10) nearly reaches or reaches eye
(38*); posterior suborbital rows (Row 11) short to moderate
length, well separated (26), nearly touching (5*), or slightly
overlapping (7*); longitudinal maxillary row (Row 14)
unbroken, extends posteriorly below (9) or below and
beyond (29*) the transverse cheek row (Row 13); longitudi-
nal mandibular row (Row 15) unbroken (38*); approximate-
ly 10 (2), 11 (30*), or 12 (6*) transverse mandibular rows
(Row 16) on each lower jaw; postorbital rows (Row 17)
medium length (between 2/3 and 4/5 distance from dorsal
midline to bony edge of cranium) (5) or long (more than 4/5
distance from dorsal midline to bony edge of the cranium)
(14*), well separated (11), nearly touch or touch (6*), or
continuous across the midline (2); preopercular row (Row
20) anterior to and not continuous with transverse opercular
row (Row 21) (38*).
Color in preservation.—(Figs. 1, 3) General color pattern
brown dusky mottling and bars on pale yellow-brown head
and body. Head markings poorly defined, mottled even in
small specimens. Ventral region of head with little pigment.
Papillae row coloration continuous with surrounding mark-
ings but with greater contrast, typically alternating light and
dark. Trunk with wide vertical bar extending below first
dorsal fin, reaching to, but not across pale belly. Irregular,
mottled marking may be visible between dorsal fins,
marking often wide ventrally. Wide, slanted bar from
second dorsal fin to posterior edge of the anal fin and
around caudal peduncle. Wide bar sometimes well defined
Fig. 3. Callogobius clarki, USNM 220038, 48.6 mm SL, male (illustration P09712 by Jack R. Schroeder, Smithsonian Institution, NMNH, Division of
Fishes): (A) dorsal view of head; (B) lateral view of head and body; (C) lateral view of head enlarged from (B). Numbers are added to identify papillae
rows used in the redescription (numbers follow Akihito and Meguro, 1977; names follow Delventhal and Mooi, 2013): 2 5postnasal row; 9 5
anterior suborbital row; 10 5mid-suborbital row; 11 5posterior suborbital row(s); 12 5longitudinal cheek row; 13 5transverse cheek row; 14 5
longitudinal maxillary row; 15 5longitudinal mandibular row; 16 5transverse mandibular rows; 17 5postorbital row; 20 5preopercular row; 21 5
transverse opercular row.
Delventhal and Mooi—Redescription of Callogobius clarki 145
on caudal peduncle, edge of hypural plate and proximal
regions of caudal fin. First and second dorsal fins with
irregular rows of pale spots and/or bands. Pectoral fins
mottled, often with small spot medially on pectoral-fin base.
Pelvic fins mostly pale, anal fin darker. Caudal fin with fine
irregular bands. Urogenital papilla pale; in female, lateral
and distal flaps lightly pigmented (Fig. 2).
Etymology.—Goren named this species in honor of Dr.
Eugenie Clark ‘‘for her outstanding contributions to the
knowledge of the fish fauna of the Red Sea.’’ We suggest the
vernacular name Clark’s Flapheaded Goby for this species.
Distribution.—Known only from the Red Sea.
Comparisons.—Four described species of Callogobius are
known to lack all head pores as adults: C. clarki,C. crassus,
C. hastatus, and C. winterbottomi.Callogobius clarki differs
from C. crassus in having higher lateral scale counts (33–41
vs. 19–22), higher second dorsal- and anal-fin element
counts (normally I,10 and I,9 vs. I,7–8 and I,6–7, respec-
tively), preopercular papillae row not continuous with
transverse opercular row (vs. continuous), longitudinal
maxillary papillae row and longitudinal mandibular row
continuous (vs. broken and displaced), and pelvic fins
joined at least 2/3 length (vs. joined only at the base).
Callogobius clarki can be distinguished from C. hastatus by
color pattern (mottled with wide bars vs. finely speckled
with or without narrow bars), body shape (moderately stout
with rounded fins vs. very slender with elongate fins),
preopercular papillae row not continuous with transverse
opercular row (vs. continuous) and fifth pelvic-fin ray
shorter than the fourth, and pelvic fins with a short and
fragile (weak) frenum (vs. rounded pelvic disc with a robust
frenum). Callogobius clarki differs from C. winterbottomi in
having higher lateral scale counts (33–41 vs. 23–26) and
higher second dorsal- and anal-fin ray counts (I,10 and I,9
vs. I,9 and I,7, respectively). All other described species of
Callogobius can be distinguished from C. clarki by the
presence of head pores on all specimens larger than 10–
15 mm. Tiny juveniles of C. sclateri,C. flavobrunneus, or C.
bifasciatus that have not yet developed pores could be
confused with C. clarki; these species share a similar body
shape, color pattern, and configuration of preopercular
papillae row and transverse opercular papillae row. Callogo-
bius sclateri and C. flavobrunneus can be distinguished by
separate pelvic fins (joined at least 2/3 length in C. clarki),
and in addition C. sclateri has a lower lateral scale count (28–
31). Callogobius bifasciatus generally has higher counts for
several meristics (pectoral-fin rays, lateral scale rows,
predorsal scale rows, transverse scale rows, transverse
mandibular papillae rows; Tables 2–5); additionally, it has
not been collected from the Red Sea.
Remarks.—Goren (1978, 1980) reported a collection date of
20 January 1970 for the holotype. This is in contrast to the
specimen label and the HUJ written catalog, both of which
record the date as 20 January 1972. The latter date is
probably correct (and is the date reported in the present
paper) because on this date a collection was made at Et Tur
(Por et al., 1972; D. Golani, pers. comm.).
Our scale counts for the holotype vary slightly from those
reported by Goren (1978), although some variation in
reported values might be expected from worker to worker
given the irregular scale rows in many species of Callogobius.
In the present study, both authors independently completed
scale counts for the majority of specimens; these were
consistent, and used for comparison between C. clarki and
C. bifasciatus.
The larger specimen of USNM 220038 (a 48.6 mm SL
male) was formerly designated as Callogobius simulus
Lachner and McKinney holotype MS, but remained unpub-
lished. The lot USNM 220031 (8 specimens alcohol, 3 CS)
Table 1. Selected morphometric characters for Callogobius clarki and C. bifasciatus. The holotype values of C. clarki are compared with values of a
similarly sized specimen of C. bifasciatus (BMNH 2000.4.19.761) as the holotype of the latter is a small juvenile in poor condition. Pooled values are
presented as averages followed by range in parentheses. Values for specimens smaller than 15 mm SL are not included.
Callogobius clarki Callogobius bifasciatus
Holotype n=16 BMNH 2000.4.19.761 n=16
Gender F 10M, 6F F 6M, 10F
Standard length (SL; mm) 36.5 16.8–48.6 37.8 22.4–46.5
Head length (HL; %of SL) 30.1 31.2 (28.6–32.5) 29.4 29.7 (27.3–32.0)
Head depth (%of HL) 52.7 48.3 (43.6–54.6) 48.6 47.1 (44.0–54.9)
Head width (%of HL) 66.4 67.9 (60.2–79.4) 71.2 70.0 (60.4–78.9)
Interorbital width (%of HL) 7.9 7.7 (6.4–9.7) 9.6 8.7 (7.3–10.1)
Pre-dorsal fin distance (%of SL) 39.5 38.3 (35.5–41.0) 36.5 37.0 (34.6–39.0)
Pre-anal fin distance (%of SL) 62.7 61.5 (58.0–63.6) 61.9 60.4 (58.4–62.2)
Pre-pelvic fin distance (%of SL) 32.1 32.8 (30.2–34.5) 29.9 30.4 (29.1–32.2)
Pectoral fin length (%of SL) 23.8 25.5 (23.5–27.1) 24.3 24.2 (20.6–27.1)
Pelvic fin length (%of SL) 18.6 21.8 (18.6–24.2) 22.0 19.9 (17.8–22.0)
Caudal fin length (%of SL) 23.6 26.9 (22.0–31.0) 25.4 25.2 (21.7–28.2)
Caudal peduncle depth (%of SL) 12.6 12.5 (11.2–13.7) 12.7 12.6 (10.9–13.6)
Table 2. Pectoral-fin ray count frequencies for Callogobius clarki (n=
21) and C. bifasciatus (n=22). Holotype values are marked by *.
Bilateral counts included when available.
Pectoral-fin ray number
15 16 17 18
C. clarki 3 27* 11
C. bifasciatus 1 15 13 13*
146 Copeia 2014, No. 1
was also designated Callogobius simulus. McKinney (1980)
referred to C. clarki as ‘‘Callogobius new species A’’ and
presented two illustrations (a dorsal view of the head and a
lateral view of the head and body) of the larger specimen of
USNM 220038. These images, by Jack R. Schroeder, are
printed here with permission of the Smithsonian Institu-
tion, NMNH, Division of Fishes (Fig. 3).
DISCUSSION
With reference only to the original descriptions and
apparently with only one Red Sea specimen at hand, Randall
et al. (1994) synonymized Callogobius clarki and C. bifascia-
tus without comment. Randall (1995) and Golani and
Bogorodsky (2010) followed this decision. Our study
indicates that C. clarki is a valid Red Sea endemic species.
Not only does C. clarki differ strikingly in lacking sensory
pores, it differs in count frequencies of pectoral-fin rays
(Table 2), lateral scale rows (Table 3), predorsal and trans-
verse scale rows (Table 4), and transverse mandibular
papillae rows (Table 5). Moreover, the anterior suborbital
papillae row (Row 9) is relatively longer in most specimens
of C. bifasciatus, nearly reaching the eye in 13 out of 14
specimens examined for this character. All specimens of C.
bifasciatus examined as comparative material in this study
had dorsal-fin element counts of VI+I,10 and anal-fin
element counts of I,9. We observed a lateral scale count
range of 37–48 (40 for the holotype) and predorsal scale
range of 16–20 (17 on the holotype). In contrast, Randall et
al. (1994) reported C. bifasciatus having ‘‘dorsal rays VI–I, 9–
10 (usually 9); anal rays I, 8–9 (usually 8); . . . longitudinal
scale series 47–53; . . . predorsal scales 21–23.’’ Of their
material, all but one specimen (from the Red Sea), is from
the Arabian Gulf or Oman, as is most of our material of C.
bifasciatus. Randall et al. (1994) reported about 47 lateral
series scales on the holotype of C. bifasciatus, but our counts
agree with those of Smith (1958), who recorded 40.
Both C. clarki and C. bifasciatus belong to the sclateri group
defined by Delventhal and Mooi (2013) as a subgroup of
Callogobius having a modified female urogenital papilla with
lateral distal flaps, and elongate ctenii on caudal peduncle
scales. The modified female urogenital papilla (Fig. 2) may
be unique to the sclateri group among gobies, although we
have yet to examine representatives of all goby genera for
this character.
Of the five species of Callogobius reported from the Red
Sea, two are endemic (C. clarki,C. dori); C. amikami is known
from one Red Sea specimen and a photo record from Oman
(Debelius, 1993:283; Randall, 1995:329), whereas C. flavo-
brunneus and C. maculipinnis are widespread throughout the
Indo-West Pacific. Randall (1998) estimated Red Sea ende-
mism in fishes to be about 14%, although he included
species that extended their ranges into the Gulf of Aden; for
example, no chaetodontids are restricted to the Red Sea, but
Randall suggested 7 of 17 were endemic. However, some
families exhibit extremely high endemism. Of 12 species of
pseudochromids listed for the Red Sea (Golani and Bogor-
odsky, 2010), nine are found only there. This supports
Randall’s (1998) expectation that smaller fishes will exhibit
higher endemism due to more rapid generation times,
amplified by life history traits such as demersal eggs and
parental care. However, the Gobiidae (sensu Gill and Mooi,
2012) does not support this generalization. Despite being
small with short generation times and having demersal eggs
with parental care, gobiids exhibit less than 16%Red Sea
endemism (19 of 120 presumed native species as listed in
Golani and Bogorodsky, 2010), only marginally higher than
Red Sea fishes as a whole. However, our knowledge of small,
cryptic fishes is far from complete; better understanding of
taxonomy is likely to reduce the number of apparent
widespread species and increase the number of recognized
endemics.
MATERIAL EXAMINED
Callogobius bifasciatus (30 specimens): BMNH 1994.1.18.163–
166, 4, 26.9–40.1 mm SL, 2 male, 2 female, Kuwait, Arabian
Gulf, Qaru Island, N. Downing, 1985; BMNH 2000.4.19.755,
1, 40.2 mm SL, male, United Arab Emirates, Abu Dhabi,
Jazirat Dagalah, 24u129180N, 52u56960E, A. C. Gill et al.,
1999; BMNH 2000.4.19.756, 1, 44.5 mm SL, female, United
Arab Emirates, Abu Dhabi, Ushsh Island, 24u189150N,
52u529180E, A. Gill et al., 1999; BMNH 2000.4.19.758–759,
2, 24.5-39.3 mm SL, 1 male, 1 female, United Arab Emirates,
Abu Dhabi, Ghasha Island, 24u259270N, 52u389440E, A. C.
Gill et al., 1999; BMNH 2000.4.19.760–766, 7, 22.4–39.9 mm
Table 3. Lateral scale count frequencies for Callogobius clarki (n=18) and C. bifasciatus (n=21). Holotype values are marked by *. Bilateral counts
included when available.
Lateral series
33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48
C. clarki 2 2 8 5 7* 6 2 1
C. bifasciatus 1 1 2 7* 7 7 3 4 3 1
Table 4. Predorsal and transverse scale count frequencies for Callogobius clarki (n=17) and C. bifasciatus (n=20 for predorsal and 21 for
transverse). Holotype values are marked by *; that of transverse series for C. bifasciatus from Smith (1958) as specimen condition did not permit
counts at time of our observation.
Predorsal series Transverse series
12 13 14 15 16 17 18 19 20 17 18 19 20 21 22
C. clarki 1 3 7 6* 9 7* 1
C. bifasciatus 1 6* 7 4 2 5* 6 3 3 4
Delventhal and Mooi—Redescription of Callogobius clarki 147
SL, 1 male, 1 male?, 5 females, United Arab Emirates, Abu
Dhabi, 24u399230N, 54u319160E, A. Gill, 1999; ROM 39892, 4,
33.9–45.0 mm SL, 2 male, 1 female, 1 CS, Oman, B. N. G.
Simm, 1981; ROM 39899, 4, 14.8–34.8 mm, 3 female, 1
juvenile, Oman, NE of Sur, B. N. G. Simm, 1982; ROM
39919, 3, 40.6–46.5 mm SL, 3 male, Oman, near Kalhat
(Qalhat), B. N. G. Simm, 1981; SAIAB 235, holotype, 1,
21.0 mm SL, female?, Tanzania, Pemba Island, J. L. B. Smith;
SAIAB 003419, 1, 63.4 mm SL, male, Tanzania, Pemba
Island, J. L. B. Smith, 1952; SAIAB 46417, 1, 24.2 mm SL,
male, South Africa, Aliwal Shoal, Cathedral, C. Buxton et al.,
1994; SAIAB 46477, 1, 28.5 mm SL, male, South Africa,
Natal, Aliwal Shoal, Hospital Reef, off Scottburgh, C. Buxton
et al., 1994.
Type specimens of other species of Callogobius were
examined as listed in Delventhal and Mooi (2013).
ACKNOWLEDGMENTS
We are grateful to D. Golani (HUJ) for loaning us the
holotype of Callogobius clarki; to J. Maclaine (BMNH), J.
Williams (USNM), and R. Winterbottom (ROM) for loans of
specimens of C. clarki and C. bifasciatus. Special thanks to
L. Palmer (USNM) who facilitated use of the J. Schroeder
illustration P09712 of C. simulus (5C. clarki). O. Gon hosted
the first author’s visit to SAIAB. A. Gill provided advice,
encouragement, and humor. A. Matheson and J. Cordova
(University of Manitoba Accessibility Services) assisted in
typing the manuscript. This study was supported in part by a
University of Manitoba Graduate Fellowship and an NSF
Graduate Research Fellowship to NRD, and Natural Sciences
and Engineering Research Council of Canada Discovery
Grant 327844-06 to RDM.
LITERATURE CITED
Akihito, Prince, and K. Meguro. 1977. Five species of the
genus Callogobius found in Japan and their relationships.
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Table 5. Transverse mandibular papillae row (Row 16) count
frequencies for Callogobius clarki (n=19) and C. bifasciatus (n=
14); counts taken bilaterally. Holotype values for C. clarki are marked by
*; counts were not taken from the holotype of C. bifasciatus at time
of observation.
10 11 12
C. clarki 2 30* 6*
C. bifasciatus 1 27
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