UiNIVtRSITY OF
ILLINOIS LIBRARY

*T URBANA CHAMPAIGN
BIOLOGY

;. • ;

t

73

FIELDIANA
Zoology

Published by Field Museum of Natural History

Volume 73, No. 2 _ February 28, 1979

A Review of the
Western Atlantic Starksia ocellata-Complex

(Pisces: Clinidae)

with the Description of Two New Species
and Proposal of Superspecies Status

DAVID W. GREENFIELD

DEPARTMENT OF BIOLOGICAL SCIENCES
NORTHERN ILLINOIS UNIVERSITY

RESEARCH ASSOCIATE, DIVISION OF FISHES
FIELD MUSEUM OF NATURAL HISTORY

ABSTRACT

Starksia ocellata (sensu Bohlke and Springer, 1961) represents a species complex
composed of five species, in addition to S. guttata. S. ocellata occurs on both coasts
of Florida north to North Carolina, and questionably in the Bahama Islands; S.
culebrae ranges from Haiti through Puerto Rico and down the Lesser Antilles to and
including St. Vincent; S. guttata is found in the Tobago Cays and the Grenadines
south to Trinidad and west to Curacao; S. brasiliensis is known from southern
Brazil; S. variabilis occurs at Santa Marta, Colombia; and S. occidentalis ranges
from Panam6 north to Yucatan, Mexico, including certain offshore islands. Of the
above species, two (S. variabilis and S. occidentalis) are described as new, and two
others (S. culebrae and S. brasiliensis) are resurrected from the synonymy of S.
ocellata.

The importance of head coloration in this complex is discussed and examples of
character displacement in meristic and morphometric characters are presented. It is
proposed that the S. oce//ata-complex represents a superspecies composed of six
allospecies. Zoogeographic implications of the superspecies are discussed in relation
to formal provinces.

ABSTRACTO

Starksia ocellata (sensu Bohlke and Springer, 1961) representa un complejo de
especies compuesto de cinco especies ademas de S. guttata: S. ocellata ocurre desde

Library of Congress Catalog Card No.: 78-66776
ISSN 0015-0754

Publication 1294 9

BIOLOGY UBKftRY

APR 17 1979 101BURmaHm

10 FIELDIANA: ZOOLOGY, VOLUME 73

ambas costas de Florida, norte por la costa Atlantica a Norte Carolina, y dudosa-
mente desde las Islas Bahama; S. culebrae ocurre desde Haiti a Puerto Rico y por los
Aritillas menor a St. Vincent; S. guttata ocurre desde los Cayos Tobago y los Grena-
dines hasta Trinidad al sur y hasta Curacao al oeste; S. brasiliensis ocurre en el sur
de Brasil; S. variabilis ocurre en Santa Marta, Colombia; S. occidentalis-ocurre desde
Panama norte a Yucatan, Mexico. Las especies S. occidentalis y S. variabilis son
nuevas, describes por la primera vez en este estudio. Las especies S. culebrae y S.
brasiliensis, anteriormente consideradas como sin6nimas de S. ocellata, son tratadas
como especies distinctas.

La importancia de la coloracibn de la cabeza se discute con respecto a las especies
de este complejo, y se presentan ejemplos de la divergencia de caracteristicos meris-
ticos y morfometricos entre las populaciones continguas de unas especies. Se pro-
pone que este complejo de especies representa una "superespecie" compuesto de
seis "allo-especies," y se discuten las implicaciones zoogeograficas.

INTRODUCTION

Recent collections in Belize and Honduras resulted in the capture
of a series of clinids most closely resembling Starksia ocellata, a
species not yet recorded from the area. Comparison of these speci-
mens with material of S. ocellata from throughout its reported range
demonstrated that not only did the material from Central America
represent an undescribed species, but that Starksia ocellata (sensu
Bohlke and Springer, 1961) represents a species complex.

The first Atlantic species of Starksia (Clinus ocellatus) was de-
scribed by Steindachner in 1876 from the Bahama Islands (possibly
Florida: see S. ocellata). Two additional species of Starksia were
described in 1900: Brannerella brasiliensis Gilbert from near
Macei6, Brazil, and Malacoctenus culebrae Evermann and Marsh
from Puerto Rico. Longley and Hildebrand (1941, p. 258) synony-
mized both S. brasiliensis and S. culebrae with S. ocellata. Bohlke
and Springer (1961, p. 51) listed S. culebrae in the synonymy of S.
ocellata without comment and also retained S. brasiliensis in the
synonymy of S. ocellata stating (p. 53), "... it (S. brasiliensis) may
eventually be shown separable at the subspecific level." Fowler
(1931) described Brannerella guttata, a species closely related to S.
ocellata, from Trinidad. Bohlke and Springer (1961, p. 50) stated,
"S. guttata is like S. ocellata in most regards and it is with some
misgivings that we tentatively retain the two as distinct." Gilbert
(1971) described two new species of Starksia, including S. elongata
(which he believed to be most closely related to S. ocellata) and
followed Bohlke and Springer (1961) in considering S. culebrae and
S. brasiliensis to be synonyms of S. ocellata. Gilbert (1971, p. 204)
recorded the distribution of S. ocellata as follows: "B & S, North

GREENFIELD: STARKSIA OCELLATA COMPLEX 1 1

Carolina, South Carolina, Florida, Cuba, Haiti, Puerto Rico, Virgin
Is., Grenadines (Lesser Antilles), Old Providence Is., Brazil, ques-
tionably from Bahamas; C, Venezuela. New record: Panama."

A comparison of the populations of S. ocellata (sensu Bohlke and
Springer) from throughout its range with the material from Central
America shows striking differences in the color pattern on the side
of the head. The populations fall into six obviously different pat-
terns: Pattern I— North and South Carolina, Atlantic Florida, Gulf
of Mexico, and Bahama Islands?; Pattern II— Haiti, south through
the Lesser Antilles to and including St. Vincent; Pattern III—
Togabo Cays, Grenadines, Tobago Island, Trinidad, and Curacao;
Pattern IV— north coast of South America at Santa Marta, Colom-
bia; Pattern V— coast of Central America from Yucaten to Panama
including Providencia; Pattern VI— southern Brazil.

METHODS

Counts and measurements follow Bohlke and Springer (1961) and
Gilbert (1971), with the exceptions of the method of counting the
number of scales in the lateral line, and in measuring upper jaw
length. All scales are included in the count whether pored or un-
pored, and the straight portion of the lateral line begins with the
first scale whose posterior end is in line with the remainder of the

FIG. 1. Method of counting arched and straight portions of the lateral line. Arrow
indicates location of first scale in straight portion.

lateral line, even if the anterior portion of the pore curves upward
(fig. 1). Upper jaw length is measured by placing one point on the
posterior end of the maxillary and the other on the midline of the
premaxillary. All measurements were made with dial calipers to the
nearest 0.1 mm. and presented as thousandths of standard length
(SL). Vertebral and fin ray counts were taken from radiographs.

12 FIELDIANA: ZOOLOGY, VOLUME 73

The following abbreviations of collections are used in listing
material examined: ANSP, Academy of Natural Sciences of Phila-
delphia; CAS and CAS-SU, California Academy of Sciences; FMNH,
Field Museum of Natural History; GCRL, Gulf Coast Research
Laboratory Museum; LACM, Los Angeles County Museum of
Natural History; MPM, Milwaukee Public Museum; RMNH, Rijks-
museum van Natuurlijke Historic; SIO, Scripps Institution of
Oceanography; UF, University of Florida (Florida State Museum);
UMMZ, University of Michigan Museum of Zoology; USNM,
United States National Museum.

Starksia ocellata (Steindachner). Figures 2-4. Tables 1-3.

Clinus ocellatus Steindachner, 1876, p. 230, pi. 12, fig. 5. Type locality: Bahama
Islands.

Diagnosis.— A species of Starksia with essentially naked belly;
simple orbital cirrus; genital papilla and first anal-fin spine in adult
male united along entire length (fig. 3), papilla projecting beyond tip
of spine a distance equal to a little less than one-fifth length of spine;
first anal-fin spine longer than second anal-fin spine; obvious pelvic-
fin rays 1,2; pectoral-fin rays 13-15 (usually 14); dorsal-fin elements
XX-XXII,6-9 (usually XXI,8); anal-fin elements 11,17-20 (usually
11,18); lateral-line scales, 16-20 in arch (usually 17) and 20-22 in
straight (usually 21), total 37-41 (usually 38); vertebrae 33-35 (usu-
ally 34); no dark diagonal bar on lower part of pectoral-fin base; body
color pattern not consisting of well-defined, dark bands. May be
distinguished from its closest congeners S. culebrae, S. occidentalis,
and S. variabilis by lacking distinct black vertical bars on the lips,
from S. brasiliensis in having two rows of infraorbital pores rather
than a single row, and from S. guttata by the presence of small,
dark, ring-like markings with light centers on the cheek and opercle

FIG. 2. Starksia ocellata, UF 10875, female, 31.3 mm. S.L., Florida.

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FIELDIANA: ZOOLOGY, VOLUME 73

FIG. 3. Gonopodium of
Starksia ocellata, UF 10875,
24.1 mm. S.L. Line equals 1 mm.

falling mostly within a horizontal pale area running from the orbit
posteriorly to the edge of the preopercle. Starksia ocellata has a
longer snout (mean 56) than either S. occidentalis (mean 49) or S.
variabilis (mean 48).

Description.— Meristics are presented in the diagnosis. Measure-
ments are presented in Table 2 and are based on 10 specimens,
17.2-33.4 mm., USNM 116831 and UF 16187.

Color pattern of head.— Starksia ocellata is clearly separable from
other members of the species complex by the presence of small ring-
like markings on the cheek and opercle. These rings are dark with
light centers and fall mostly within a horizontal pale area running
from the orbit posteriorly to the edge of the preopercle (figs. 2, 4).
The anterior portion of the lips have a scattering of melanophores;
however, there are no distinct black vertical bars on the lips. This
color pattern is clearly illustrated in Bohlke and Springer (1961, fig.
15) and in Bohlke and Chaplin (1968, p. 525).

Remarks. —As restricted herein, S. ocellata occurs only along the
Gulf and Atlantic coasts of Florida, the Florida Keys including the
Dry Tortugas and north along the Atlantic coast of the United
States to N. Carolina. Bohlke and Springer (1961) have discussed
the problem of the type locality of S. ocellata. Although listed as
"Bahama-Inseln," this species has not since been collected in the

FIG. 4. Typical head color pat-
tern of Starksia ocellata.

GREENFIELD: STARKSIA OCELLATA COMPLEX 17

Bahamas and it is likely that the type locality may actually be
Florida.

Material examined— N. Carolina-USNM 120131(1). S. Carolina-
UF 7270(2); GCRL 189(2). Gulf of Mexico-UF 7858(2); UF 7859(9).
Florida (Monroe Co.)-UF 10875(9); UF 11876(9); UF 16187(34);
USNM 116831(6); SIO-67-86(3). Florida (Indian R. Co.) UF 12040(1).
Florida (Palm Beach Co.) UF 16018(1). Standard length of
specimens examined 17.2-33.8 mm.

FlG. 5. Starksia culebrae, ANSP 124681, male, 27.9 mm. S.L., St. Lucia.

Starksia culebrae (Evermann and Marsh). Figures 5-7. Tables 1, 2, 4.

Malacoctenus culebrae Evermann and Marsh, 1900a, p. 357. Type locality: reefs out-
side the harbor of Culebra, Puerto Rico.

Diagnosis.— A species of Starksia with essentially naked belly;
simple orbital cirrus; genital papilla and first anal-fin spine in adult
male united along entire length (fig. 6), papilla projecting beyond tip
of spine a distance equal to one-fifth length of spine; first anal-fin
spine longer than second anal-fin spine; obvious pelvic-fin rays 1,2;
pectoral-fin rays 13-14 (usually 14); dorsal-fin elements XX-
XXII,7-9 (usually XXI.8); anal-fin elements 11,17-19 (usually 11,18);
lateral-line scales 16-19 in arch (usually 17) and 20-22 in straight
(usually 21), total 37-41 (usually 38); vertebrae 34-35 (usually 34); no
dark diagonal bar on lower part of pectoral-fin base; body color pat-
tern not consisting of well-defined, dark bands. May be distinguish-
ed from its closest congeners S. ocellata, S. guttata, and S. brasilien-
sis by the presence of distinct black vertical bars on the lips and
from S. occidentalis and S. variabilis by possessing a pale horizontal
bar running from the orbit posteriorly past the edge of the preoper-
cle onto the opercle. The horizontal bar does not branch into a Y, and

18

FIELDIANA: ZOOLOGY, VOLUME 73

FlG. 6. Gonopodium of
Starksia culebrae, ANSP
124679, 25.8 mm. S.L. Line
equals 1 mm.

dark ring-like markings are almost always absent. It may further be
distinguished from S. brasiliensis in having two rows of infraorbital
pores rather than a single row. Starksia culebrae has a longer snout
(mean 58) than S. occidentalis (mean 49) or S. variabilis (mean 48).

Description.— Meristics are presented in the diagnosis. Measure-
ments are presented in Table 2 and are based on 20 specimens,
15.9-27.3 mm., ANSP 134946, ANSP 134945, ANSP 113248, ANSP
124674, ANSP 124679, ANSP 112981, ANSP 124687, and ANSP
124685.

Color pattern of head.— Starksia culebrae is clearly separable from
other members of the species complex by the combination of
distinct black vertical bars on the lips and a pale horizontal bar run-
ning from the orbit posteriorly past the edge of the preopercle onto
the opercle (fig. 7). The horizontal bar does not branch into a Y and
dark ring-like markings are almost always absent. Specimens from
Martinique occasionally have one or two small ring-like markings,
which are located ventral to the horizontal bar. The only other
species with bars on the lips are S. occidentalis from the coast of
Central America and S. variabilis from Colombia, but the pale pat-
terns on the side of the cheek are different. This color pattern is
clearly illustrated in Evermann and Marsh (1900b, fig. 96) and in
Beebe and Tee- Van (1928, p. 236).

Remarks.— Starksia culebrae ranges from Haiti through Puerto
Rico and down the Lesser Antilles to and including St. Vincent.

FIG. 7. Typical head color pat-
tern of Starksia culebrae.

TABLE 4. Meristic characters for Starksia culebrae from various localities.

Dorsal-fin Dorsal-fin Anal-fin
spines soft rays soft rays

Pectoral-
fin rays Verte

brai

XX XXI XXII 7 8 9 17 18 19

13 14 34

35

Haiti

182 10 1 10 1

11 9

2

Puerto Rico

1 1 1

1

St. Barthelemy

45 162162

1 4 5

5

Antigua

42 33 24

6 3

3

Dominica

1 1 1

1

1

Martinique

192 381192

10 8

1

St. Lucia

282 183 93

7 9

3

St. Vincent

10 2 48 1 11

10 3

9

Total

4 45 15 5 39 20 2 37 25

2 49 37

24

Lateral-line scales

Arched Straight
16 17 18 19 20 21 22

Total
36 37 38 39 40

41

Haiti

9 54

5 4

Puerto Rico

1 1

1

St. Barthelemy

14 14

2 3

Antigua

6 42

4 2

Dominica

1 1

1

Martinique

441 441

2331

St. Lucia

32 14

1 2 2

St. Vincent

721 46

351

1

Total

8 33 4 1 12 25 9

3 13 19 9 1

1

19

20 FIELDIANA: ZOOLOGY, VOLUME 73

Whether this species occurs in Cuba is not known. The specimen,
USNM 82548, listed by Bohlke and Springer (1961) from Cuba as S.
ocellata is S. fasciata (Longley). Although S. culebrae is 100 per cent
separable from S. ocellata on the basis of coloration, counts and
measurements cannot be used for separation inasmuch as the only
noticeable difference is an upward shift in the number of vertebrae
in S. culebrae. To the south of St. Vincent S. culebrae is replaced by
S. guttata, which lacks bars on the lips.

The only noticeable instance of geographic variation in popula-
tions of S. culebrae is found in the St. Vincent population, which,
when compared to populations to the north of St. Lucia and Mar-
tinique, shows a modal increase in dorsal-fin soft rays, anal-fin soft
rays, vertebrae, and lateral-line scales. The St. Vincent population is
located at the southernmost limit of the range of S. culebrae, and
thus is geographically closest to populations of S. guttata. These
shifts in certain modal values in S. culebrae result in modal differ-
ences between the two species that are not observed between their
geographically more distant populations, and thus possibly repre-
sent a case of character displacement, a phenomenon supporting
genetic differences between these two species.

Material examined. -Haiti-USNM 178297(5); ANSP 134946(1);
ANSP 134945(5). Puerto Rico-USNM 125973(1), (paratype of M.
culebrae). St. Barthelemy-ANSP 113248(1); ANSP 124674(4);
ANSP 124677(4). Antigua-ANSP 117907(5); UF 11404(1). Domi-
nica-USNM 198272(1). Martinique-ANSP 112981(1); ANSP
113012(2); ANSP 113060(5); ANSP 124679(2); ANSP 124687(2). St.
Lucia-ANSP 124661(1); ANSP 124666(6); ANSP 124681(5). St.
Vincent-ANSP 124623(1); ANSP 124685(12). Standard length of
specimens examined 13.4-31.2 mm.

Starksia guttata (Fowler). Figures 8, 9a, b, lOa-c. Tables 1, 2, 5.

Brannerella guttata Fowler, 1931, p. 401, text-fig. 3. Type locality: Monas Island,
Trinidad.

Diagnosis.— A species of Starksia with essentially naked belly;
simple orbital cirrus; genital papilla and first anal-fin spine in adult
male united along entire length (fig. 9a, b), papilla projecting beyond
tip of spine a distance equal to one-sixth to one-third length of spine;
first anal-fin spine longer than second anal-fin spine; obvious pelvic-
fin rays 1,2; pectoral-fin rays 13-14 (almost always 14); dorsal-fin
elements XX-XXI.8-9 (usually XXI,8); anal-fin elements 11,17-18
(usually 11,18); lateral-line scales, 15-18 in arch (usually 17) and

GREENFIELD: STARKSIA OCELLATA COMPLEX

21

FIG. 8. Starksia guttata, ANSP 53327-30 (paratype), male, 37.8 mm. S.L., Trini-
dad.

20-22 in straight (usually 21), total 37-39 (usually 38); vertebrae
33-34 (almost always 34); no dark diagonal bar on lower part of
pectoral-fin base; body color pattern not consisting of well-defined,
dark bands. May be distinguished from its closest congeners S.
culebrae, S. occidentalis, and S. variabilis by lacking distinct black
vertical bars on the lips; from S. brasiliensis in having two rows of
infraorbital pores rather than a single row; and from S. ocellata by
usually having a pale horizontal bar which branches into a definite
Y posteriorly, and often lacking dark ring-like markings, or having
solid dark spots overlaying the pale horizontal bar. Starksia guttata
has a longer snout (mean 58) than either S. occidentalis (mean 49) or
S. variabilis (mean 48).

Description.— Meristics are presented in the diagnosis. Measure-
ments are presented in Table 2 and are based on all material listed
except for a bent specimen from LACM 22682 and one from USNM
170202.

Color pattern of head.— Lips dusky anteriorly or evenly peppered
with melanophores, without distinct black vertical bars; cheek
dusky with a pale horizontal bar, equal to or greater than half the

B

FlG. 9. Gonopodium of Starksia guttata. a, ANSP 53325, 36 mm. S.L., Trinidad; b,
ANSP 124624, 23.3 mm. S.L., Union Is. Line equals 1 mm.

22

FIELDIANA: ZOOLOGY, VOLUME 73

TABLE 5. Meristic characters for Starksia guttata from various localities.

Dorsal-fin Dorsal-fin Anal-fin Pectoral-
spines soft rays soft rays fin rays Vertebrae

Tobago Cays

Grenadines

Tobago Is.

Trinidad

Curacao

Total

:x

XXI

8 9

17

18

13 14

33 34

4

1 3

3

1

4

2

2

3

5

2

3

1 4

1 4

3

3

1

2

3

2

6

5 1

6

6

6

1

1

1

1

1

6

13

15 4

6

13

1 18

1 15

Lateral-line scales

Arched
15 16 17 18

Straight
20 21 22

Total
37 38

39

Tobago Cays

121

3

2

1

Grenadines

4 1

1 4

1

3

1

Tobago Is.

1 1

1 1

1

1

Trinidad

1 5 1

1 5 1

2

3

1

Curacao

1

1

1

Total

1 2 13 3

3 13 2

5

9

3

pupil diameter above the maxillary, running from the orbit
posteriorly, usually branching into a definite Y posteriorly and
often lacking dark ring-like markings (fig. lOa, b). Several of the
paratypes have dark spots overlaying the pale horizontal bar, but
this pattern has not been observed outside of Trinidad (fig. lOc). The
Y-shaped pale bar on the side of the head is clearly illustrated in
Randall (1968, fig. 273) (a specimen from Tobago).

Remarks.— The determination of the species limits for S. guttata
has posed a difficult problem. When Bohlke and Springer (1961)
reviewed the genus they had available only the type material of S.

GREENFIELD: STARKSIA OCELLATA COMPLEX

23

B

FlG. 10. Three head color patterns of Starksia guttata: a, b, typical patterns; c,
paratype from Trinidad exhibiting spotted pattern.

guttata. On the basis of the distinctive color pattern of the body and
supposed slight differences in two body proportions they recognized
this species as valid. Bohlke and Springer (1961) and later Gilbert
(1971) used the following characters to separate S. guttata from S.
ocellata (sensu Bohlke and Springer): head length 280-314 SL and
upper jaw length 133-144 SL in S. ocellata vs. head length 328-351
SL and upper jaw length 147-168 SL in S. guttata. Allometric
growth is a factor in the supposed differences in upper jaw length
(fig. 11) and no differences in head length are evident (fig. 12). Thus,
part of the problem in locating additional specimens of S. guttata
was related to the validity of these characters. Further, additional
specimens having the distinctive round black spots on the body
were not located. The color pattern of the body is variable and may
show some clinal variation. Specimens from Trinidad (paratype, fig.
8) and Tobago (fig. 13) have the body covered with distinct round
black spots on a light background. One small specimen (16.1 mm.)
from Tobago is barred and lacks spots, which is perhaps a juvenile
color phase. To the north this spotting pattern becomes less distinct
and blotches more typical of other species in the S. ocellata-complex
begin to appear. At Union Island the pattern of blotches is in-
distinct and the background color dark; however, the round black
spots are still evident (fig. 14). At Little St. Vincent Island the

24

FIELDIANA: ZOOLOGY, VOLUME 73

165
160

-C

_ = 155
z"l 150

•2 145
£ 05

^ H-

' ° 140
o: «

u 9
oil 135

130
125
120 h

0 •

•+

-»• +•

• oi o

15

20

25

30

35

40

STANDARD LENGTH, mm

FIG. 11. Length of upper jaw in thousandths of standard length us. standard
length in millimeters for Starksia guttata (solid circles), S. ocellata (open circles), and
S. culebrae (crosses). Arrows indicate paratypes of S. guttata.

background color is cream with distinct blotches; however, large
black spots are present around the edges of the blotches and the cen-
tral portions of the blotches are pale, thus approaching a spotted
pattern (fig. 15). This clinal explanation is confounded, however, by
the pattern illustrated by Randall (1968, fig. 273) of a specimen from
Tobago that is very similar to the Little St. Vincent Island pattern.
No significant differences in counts or measurements could be
found between the spotted individuals from Trinidad and Tobago
and other specimens here considered to be S. guttata, and thus they
are treated as a single species.

The integrity of this species is demonstrated by comparing it to
the geographically adjacent species. Starksia guttata occurs as far
north as the Tobago Cays in the Grenadines. A single specimen
from Bequia Island, just south of St. Vincent Island (ANSP 124663)
is totally bleached and no color characters are available. Starksia

UJ

.c

Of \J
360

+

c

0)

350

+° 0

c
a

340

• + / /

•6

330

" • + o

1

320

+ ° +(ip* + ^ //

lousanc

310

-hD + x

+ o ^ ^^
V

300

4- •

290

15

20 25 30 35

STANDARD LENGTH, mm

40

FIG. 12. Head length in thousandths of standard length vs. standard length in
millimeters for Starksia guttata (solid circles), S. ocellata (open circles), and S.
culebrae (crosses). Arrows indicate paratypes of S. guttata.

FIG. 13. Starksia guttata, LACM 22682, female, 33.4 mm. S.L., Tobago Is.

25

26

FIELDIANA: ZOOLOGY, VOLUME 73

FIG. 14. Starksia guttata, ANSP 124624, male, 23.3 mm. S.L., Union Is.

culebrae is present at St. Vincent Island where it can be easily
distinguished from S. guttata by the combination of distinct black
vertical bars on the lips and the pale horizontal bar running from the
orbit posteriorly past the edge of the preopercle onto the opercle. To
the south of Trinidad along the southern coast of Brazil, S. brasilien-
sis is present. This species differs from S. guttata in having one, in-
stead of two, rows of infraorbital pores, a lower total number of
lateral-line scales, fewer vertebrae, fewer anal-fin and dorsal-fin
rays, and a distinctive color pattern on the side of the head.

A single specimen from Curacao appears to be S. guttata. This
specimen lacks black vertical bars on the lips and has a typical
Y-shaped bar with light spots on the side of the head (similar to
ANSP 117893 from Little St. Vincent Island). The body coloration
is faded (it was collected in 1917), but some distinct black spots are
evident on the body. To the west, two other species of the S. ocellata-
complex occur: S. variabilis and S. occidentalis. Both of these
species have distinct black vertical bars on the lips, different color

FIG. 15. Starksia guttata, ANSP 117893, male, 30.5 mm. S.L., Little St. Vincent

Is.

GREENFIELD: STARKSIA OCELLATA COMPLEX 27

patterns on the sides of the heads, shorter snouts, and fewer total
lateral-line scales, vertebrae, and anal-fin rays.

Material examined.— TRINIDAD (paratypes) ANSP 53325(1);
ANSP 53326(1); ANSP 53327-30(4). TOBAGO ISLAND-LACM
22682(2); ANSP 98488(1). GRENADINES (B.W.I.)-Tobago Cays
USNM 170202(2); Little St. Vincent Island ANSP 114377(2), ANSP
117893(1), ANSP 124658(1); Union Island ANSP 124624(1).
CURA£AO-RMNH 9822(1). Standard length of specimens examin-
ed 16.1-38.2 mm.

FlG. 16. Starksia occidentalis , FMNH 83716 (holotype), female, 29.6 mm. S.L.,
Belize.

Starksia occidentalis new species. Figures 16-18. Tables 1, 2, 6.

Holotype.— FMNH 83716, adult female 29.6 mm. SL., Belize, Tar-
pon Cay (about 16° 10' N. lat., 88° 40' W. long.), depth 0-1 m., 19 Ju-
ly 1974, D. W. Greenfield, T. Greenfield, R. L. Woods, R. Williamson
(field no. G-74-15).

Paratypes.— BELIZE-FMNH 84388 (3 females, 16.0-26.3 mm.),
taken with holotype; FMNH 84386 (1 female, 27.8 mm.), Bugle
Cays, depth 0-1.2 m., 16 July 1974, D. W. Greenfield, T. Greenfield,
R. L. Woods, R. Williamson; UF 23348 (1 female, 28.5 mm.), French-
man's Cay, depth 0-1. 2m., 19 July 1974, D. W. Greenfield, T.
Greenfield; FMNH 84385 (1 male, 26.9 mm.), Barrier Reef, Gallows
Point, depth .9 m., 15 April 1973, D. W. Greenfield, T. Greenfield,
A. Drew, M. Drew, J. Russo, D. Wildrick, R. Woods; FMNH 84389
(1 female, 25.0 mm.), Middle Snake Cay, depth 1.5 m., 20 July 1974,
D. W. Greenfield.

HONDURAS-FMNH 84377 (1 male 25.8, 4 females 17.0-28.0

28 FIELDIANA: ZOOLOGY, VOLUME 73

mm.), Hog Islands, Big Hog Island, depth .9 m., 20 May 1974, D.
W. Greenfield, T. Greenfield, R. K. Johnson; UMMZ 200204 (1 male
20.5, 5 females 17.5-20.3 mm.), Hog Islands, Big Hog Island, depth
0-.9 m., 21 May 1975, R. K. Johnson, R. R. Miller, F. Miller, G.
Glodek; FMNH 84379 (1 male, 22.0, 1 female 20.2 mm), Hog
Islands, Little Hog Island, depth .6 m., 19 May 1975, R. K.
Johnson; FMNH 84380 (1 female 23.3 mm.), Hog Islands, Little
Hog Island, depth 4.6 m., 18 May 1975, D. W. Greenfield, T. Green-
field, R. K. Johnson, G. Glodek, N. Hylton; FMNH 84381 (1 male,

17.1, 5 females 9.4-16.2 mm.), Hog Islands, N.W. Cay, depth 15.2
m., 20 May 1975, D. W. Greenfield, T. Greenfield, R. K. Johnson, R.
R. Miller, F. Miller, G. Glodek, N. Hylton; FMNH 84382 (1 female

20.2, 4 males, 16.6 - 23.0 mm.), Roatan, Cow Island in Bay of Port
Royal, depth 1.5 m., 2 May 1975, D. W. Greenfield, T. Greenfield, R.
K. Johnson, R. R. Miller, F. Miller, G. Glodek; FMNH 84383 (1 male
24.2, 6 females 16.7-25.2 mm.), Hog Islands, Little Hog Island,
depth 4.6 m., 19 May 1975, D. W. Greenfield, T. Greenfield, R. K.
Johnson, G. Glodek; FMNH 84384 (2 males both 21.4, 7 females
15.9-25.6 mm.), Hog Islands, Big Hog Island, depth 1.8 m., 21 May
1975, D. W. Greenfield, T. Greenfield, R. K. Johnson, R. R. Miller,
F. Miller, G. Glodek.

OLD PROVIDENCE -USNM 107110 (2 males 17.6-24.6 mm.),
shore, 6 August 1938, W. L. Schmitt and presidential party; UF
24303 (1 female 25.6 mm.), S.W. Coast, Santa Catalina, 26 August
1968, Tyler, Tyler, Faunce, Perdew, Londono, Freidenberg, Ander-
son.

MEXICO-MPM 11369 (1 female 24.1 mm.), Yucatan, Quintana
Roo, Ascension Bay, 16 June 1976, Spieler, Yeo, Noeske; MPM
11577 (1 female 10.8 mm.), Yucatan, Quintana Roo, Ascension Bay,
Cayo Culebra, 17 June 1976, Spieler, Noeske, Yeo; USNM 192388 (1
female 20.5 mm.), Yucatan, Quintana Roo, Ascension Bay,
Nichaabin Reef, 14 April 1960, Smithsonian-Bredin Caribbean Ex-
pedition IV; USNM 192399 (1 male 19.4 mm.), Yucatan, 1960,
Smithsonian-Bredin Caribbean Expedition IV; USNM 192401 (1
male 27.8 mm.), Yucatan, Quintana Roo, Ascension Bay, 16 April
1960, Smithsonian-Bredin Caribbean Expedition IV; USNM
192416 (1 male 17.0 mm.), Yucatan, 1960, Smithsonian-Bredin Car-
ribean Expedition IV.

PANAMA-S10 67-45 (1 female 24.5 mm.), Toro Point, depth 0-4.6
m., 23 March 1967, R. Rosenblatt, I. and R. Rubinoff; CAS 31610 (2

GREENFIELD: STARKSIA OCELLATA COMPLEX

29

TABLE 6. Meristic characters for Starksia occidentalis from various localities.

Dorsal-fin Dorsal-fin Anal-fin

spines soft rays soft rays

Yucatan

2

4

Belize

2

6

Honduras

9

28

Providencia

3

Panama

5

5

Total

18

46

Pectoral-
fin rays Vertebrae

XX XXI 7 8 9 16 17 18 19 13 14 32 33 34 35

42114

341 152

5 27 5 22 14 1

21 12

19 82

18 46 9 46 9 2 37 24 1

6

1

4

1

7

3

5

2

17

22

15

1

3

1

2

7

1 7

3

5

40

1 33

28

1

Lateral-line scales

Arched
16 17

Straight
19 20 21

Total
35 36 37

38

Yucatan

3 2

2

3

2 1

2

Belize

3 5

2

4 2

4

3

1

Honduras

4 8

7

6

1 7

4

1

Providencia

2

1

1

1

1

Panama

6

4

2

4

2

Total

10 23

16

16 2

3 17

12

2

males 21.2-26.3, 3 females 16.2-26.5 mm.), San Bias Archipelago,
Morbetupo, depth 10 m., J 3 May 1974, J. E. McCosker, D. Diener;
CAS 31661 (1 male 23.7 mm.), San Bias Archipelago, Rio Tigre,
depth 10 m., 14 May 1974, J. E. McCosker, D. Diener, R. Warner;
CAS 31727 (1 male 16.8, \ female 13.7 mm.), San Bias Archipelago,
Cocos-Banderas Cays, depth 5 m., 15 May 1974, J. E. McCosker, S.
McCosker, D. Diener; GCRL 3611 (1 immature 13.1 mm.), Devils
Beach, Fort Sherman, depth to 9.1 m., 28 July 1968, C. E. Dawson.

Diagnosis.— A species of Starksia with essentially naked belly;
simple orbital cirrus; genital papilla and first anal-fin spine in adult

30 FIELDIANA: ZOOLOGY, VOLUME 73

male united along entire length (fig. 17), papilla projecting beyond
tip of spine a distance equal to a little less than one-third length of
spine; first anal-fin spine longer than second anal-fin spine; obvious

FlG. 17. Gonopodium of Starksia occidentalis, UF 10875, 24.1 mm. S.L. Line
equals 1 mm.

pelvic-fin rays 1,2; pectoral-fin rays 13-14 (usually 14); dorsal-fin
elements XX-XXI.7-9 (usually XXI,8); anal-fin elements H, 16-19
(usually 11,17); lateral-line scales, 16-17 in arch (usually 17) and
19-21 in straight (usually 19), total 35-38 (usually 36); no dark
diagonal bar on lower part of pectoral-fin base; body color pattern
not consisting of well-defined, dark bands. May be distinguished
from its closest congeners S. guttata, S. brasiliensis, and S. ocellata
by the presence of distinct black vertical bars on the lips, and from
S. culebrae by a Y-shaped pale bar running posteriorly from the or-
bit with distinct, dark, ring-like markings usually present under the
ventralmost portion of the Y. It may further be distinguished from
S. ocellata, S. culebrae, and S. guttata by possessing a lower number
of vertebrae (mode 33 in occidentalis vs. 34 in others), a lower
number of anal-fin soft rays (mode 17 in occidentalis vs. 18 in
others), and a lower number of lateral-line scales in the straight por-
tion (mode 19 or 20 in occidentalis vs. 21 in others). Starksia oc-
cidentalis also has a shorter snout than other species except S.
variabilis (mean 49 in occidentalis vs. 56 jn ocellata, 58 in culebrae,
58 in guttata, and 59 in brasiliensis. It also differs from S. brasilien-
sis in having two rows of infraorbital pores rather than a single row.
Starksia occidentalis differs from its closest congener, S. variabilis,
by having fewer anal-fin soft rays (mode 17 in S. occidentalis vs. 18),
fewer total lateral-line scales (mode 36 in S. occidentalis vs. 37), a
longer head (mean 310 in S. occidentalis vs. 304), and by its post-

GREENFIELD: STARKSIA OCELLATA COMPLEX 31

orbital Y-shaped pale bar bordered ventrally (usually) with distinct
dark ring-like markings.

Description.— Data for the holotype are presented first, followed
in parentheses by the range for the paratypes, plus the mean for
morphometric data and mode for meristics. Dorsal-fin elements
XXI,7 (XX-XXI.7-9, usually XXI.8); anal-fin elements 11,17
(11,16-19, usually 11,17); pectoral-fin rays 14 (13-14, usually 14);
vertebrae 33 (32-35, usually 33); lateral-line scales 17 in arch (16-17,
usually 17) and 21 in straight (19-21, usually 19 or 20), total 38
(35-38, usually 36).

Measurements are based on 20 specimens, FMNH 83716, FMNH
84377, FMNH 84385, FMNH 84386, FMNH 84387, FMNH 84388,
FMNH 84389, USNM 192401, USNM 192399, CAS 31610. Length
of head 290 (286-355, 310); depth of body 209 (165-213, 195);
diameter of eye 81 (74-105, 85); length of snout 37 (37-61, 49); length
of upper jaw 132 (116-147, 135); length of pectoral fin 226 (203-294,
237); length of ventral fin 223 (174-281, 212); length of first dorsal-
fin spine 71 (65-112, 83).

Narrow simple cirri present on nape, top of eyeball and rear
margin of anterior nostril, orbital cirrus the longest, nostril cirrus
three-fourths of orbital cirrus and cirrus on nape slightly shorter
than nostril cirrus; teeth present on vomer and palatine bones; most
or all scales of posterior part of lateral line with tubes and pores,
scales of anterior arched portion with tubes and pores except usual-
ly the last one or two, which curve down to meet the straight por-
tion; third pelvic ray greatly reduced and not obvious; pectoral fin
extending posteriorly to between bases of 1st and 2nd soft anal-fin
rays; belly mostly naked except for four or five scale rows im-
mediately anterior to anus.

Color pattern of Aead— Lips with distinct black vertical bars,
anteriormost bar begins on upper lip just below nostril and runs
ventrally onto lower lip, second bar begins on upper lip slightly pos-
terior to first bar, third bar begins on upper lip under suborbital
pore under eye, and fourth begins on maxillary and narrows ventral-
ly toward lower jaw; cheek dusky with a pale Y-shaped horizontal
marking running posteriorly from the orbit onto the opercle, its ven-
tral margin with distinct, dark, ring-like markings with pale centers;
top of head with brown band running between dorsal edges of oper-
cle. Color pattern of body: background cream overlaid by three
longitudinal rows of brown blotches, each blotch slightly smaller

32 FIELDIANA: ZOOLOGY, VOLUME 73

than eye; dorsalmost row of 10 blotches located along base of dorsal
fin, extending up onto bases of fin rays and membrane, first blotch
located at base of first dorsal-fin spine, last blotch over hypural

FlG. 18. Typical head color
pattern of Starksia occidentalis.

(forming upper hypural marking); second row slightly dorsal to
midline of body, each blotch slightly posterior to blotch above; third
row slightly below midline of body and each blotch slightly posterior
to blotch above and thus midway between blotches in dorsalmost
row; distinctness of blotches variable, but usually well defined;
pelvic fins cream; pectoral fins cream except for two dark, ring-like
markings with cream centers located on basal portion of rays, one
dorsally and one ventrally, dorsal fin with distinct dark brown spot
on membrane between 1st and 2nd dorsal-fin spines in males, re-
mainder of fin with scattered brown spots on fin rays, melanophores
form spots running onto membranes; anal fin similar to dorsal fin;
caudal fin crossed by a series of four or five light brown bars.

Ety mology. — The name occidentalis refers to the fact that this
species has the westernmost distribution of any species in the S.
ocellata-complex.

Remarks.— Starksia occidentalis ranges from the east side of the
Yucatan peninsula, Mexico, south along the coast of Central Amer-
ica to Panama. It is also present on the island of Old Providence off
Nicaragua. With the exception of the southern species, S. brasilien-
sis, S. occidentalis exhibits the greatest divergence of all the Stark-
sia species in the S. ocellata-complex. In addition to the distinctive
color pattern, it has a shorter snout length and lower counts for
lateral-line scales, vertebrae, and anal-fin soft rays. In Belize, where
the most thorough collections have been made, it appears to be
restricted to areas near continental influence. It has been taken on
the Barrier Reef and around cays inside of the reef, but never on the
atolls outside of the Barrier Reef. It appears to be most common in
shallow waters, 1.5 m. or less; however, six individuals (FMNH
84381) were taken at a depth of 15.2 m. off Honduras. A discussion

GREENFIELD: STARKSIA OCELLATA COMPLEX

33

FlG. 19. Starksia variabilis, FMNH 83717 (holotype), male 29.8 mm. S.L., Santa
Marta, Colombia.

of the ecological relationships of S. occidentalis to other species in
the genus is presented by Greenfield and Johnson (MS).

Starksia variabilis new species. Figures 19-21. Tables 1, 2.

Holotype— FMNH 83717, adult male 29.8 mm. SL., Colombia,
northeast of Bahia de Nenquange, Parque Nacional Tayrona, Santa
Marta (about 11°40'N. lat., 74°30'W. long), depth 1 m. (±.2 m.),
August, 1976, Arturo Acero P.

Paratypes.— All collected with the holotype. FMNH 83718 (5
males, 23.7-32.9 mm. and 2 females, 25.1-27.6 mm.); USNM 217832
(1 female, 28.9 mm.); GCRL 15739 (1 male, 24.8 mm.); UF 23349 (1
male, 23.3 mm.); CAS 40229 (1 female, 21.7 mm.).

Diagnosis.— A species of Starksia with essentially naked belly;
simple orbital cirrus; genital papilla and first anal-fin spine in adult
male united along entire length (fig. 20), the papilla projecting a
short distance beyond tip of spine; first anal-fin spine longer than
second anal-fin spine; obvious pelvic-fin rays 1,2; pectoral-fin rays
13-15 (usually 14); dorsal-fin elements XX-XXI.7-9 (usually XXI,8);
anal-fin elements 11,17-18 (usually 11,18); lateral-line scales, 16-18 in
arch (usually 17) and 20-21 in straight (usually 20), total 36-37
(usually 37); no dark diagonal bar on lower part of pectoral-fin base;

FIG. 20. Gonopodium of
Starksia variabilis, FMNH
83718 (paratype), male, 32.9
mm. S.L., Santa Marta, Colom-
bia.

34 FIELDIANA: ZOOLOGY, VOLUME 73

body color pattern not consisting of well-defined, narrow, dark
bands. May be distinguished from its closest congeners S. guttata,
S. brasiliensis, and S. ocellata by the presence of distinct black ver-
tical bars on the lips, from S. culebrae by a reticulated pale pattern
on the side of the head, but never with a single horizontal pale bar.
It may further be distinguished from S. ocellata, S. culebrae, and S.
guttata by possessing a lower number of lateral-line scales in the
straight portion (mode 20 in S. variabilis vs. 21 in others). Starksia
variabilis has a shorter snout than all other species in the complex
except S. occidentalis (mean 48 in S. variabilis vs. 56 in S. ocellata,
58 in S. culebrae, 58 in S. guttata, and 59 in S. brasiliensis). It also
differs from S. brasiliensis in having two rows of infraorbital pores
rather than a single row. Starksia variabilis differs from its most
closely-related species, S. occidentalis, by having more anal-fin soft
rays (mode 18 in S. variabilis vs. 17), more total lateral-line scales
(mode 37 in S. variabilis vs. 36), a shorter head (mean 304 in S.
variabilis vs. 310), and by the distinctive color pattern on the side of
the head. S. occidentalis has a Y-shaped pale bar running posteriorly
from the orbit with distinct, dark ring-like markings usually present
under the ventralmost portion of the Y. In S. variabilis the pale pat-
tern on the side of the head is more complex and variable (fig. 21).
None of the individuals have the simple Y patterns, but rather have
a more reticulated pattern with the pale area enclosing dark blot-
ches. Occasionally dark ring-like markings are present within the
pale area, ventral to it, or both.

Description.— Data for the holotype are presented first, followed
in parentheses by the range for the paratypes, plus the mean for
morphometric data and mode for meristics. Dorsal-fin elements
XXI,8 (XX-XXI.7-9, usually XXI.8); anal-fin elements 11,18 (11,17-
18, usually 11,18); pectoral-fin rays 14 (13-15, usually 14); vertebrae
34 (33-34, usually 33 or 34); lateral-line scales 17 in arch (16-18,
usually 17) and 20 in straight (20-21, usually 20), total 37 (36-38,
usually 37).

Measurements are based on all 12 specimens in the type series.
Length of head 302 (289-316, 304); depth of body 201 (174-210, 192);
diameter of eye 80 (76-90, 83); length of snout 57 (36-58, 48); length
of upper jaw 151 (127-151, 138); length of pectoral fin 228 (212-258,
233); length of ventral fin 178 (145-211, 193); length of first dorsal-
fin spine 74 (62-82, 75).

Narrow simple cirri present on nape, top of eyeball and rear
margin of anterior nostril, nostril cirrus the longest, orbital slightly

GREENFIELD: STARKSIA OCELLATA COMPLEX

35

shorter and nape slightly shorter than orbital cirrus; teeth present
on vomer and palatine bones; most or all scales of posterior part of
lateral line with tubes and pores, scales of anterior arched portion
with tubes and pores except usually the last one or two which curve
down to meet the straight portion; third pelvic ray greatly reduced
and not obvious; pectoral fin extending posteriorly to base of 1st
soft anal-fin ray; belly mostly naked except for five or six scale rows
immediately anterior to anus.

Color pattern of head.— Lips with distinct black vertical bars,
anteriormost bar beginning on upper lip just below nostril and run-
ning ventrally onto lower lip, second bar beginning on upper lip
slightly posterior to first bar, third bar beginning on upper lip under
suborbital pore under eye, fourth bar beginning on anterior portion
of maxillary and running ventrally onto lower jaw either as a solid
bar or as separated blotches, occasionally isolated blotches present
between third and fourth bar, a distinct black blotch on posterior
margin of maxillary; cheek dusky brown with variable pale pat-

FIG. 21. Typical head color patterns of Starksia variabilis.

36 FIELDIANA: ZOOLOGY, VOLUME 73

terns, beginning as a single pale bar at the posterior margin of the
eye and branching into two or more portions posteriorly across the
cheek, the pale areas enclosing dark brown blotches, occasionally
with dark ring-like markings present either within the pale area,
ventral to it or both; interorbital white with a black blotch in the
center; top of head with a dark brown rectangular blotch in area be-
tween posterior margin of eye and posterior margin of preopercle, a
white band posterior to the rectangular blotch, followed posteriorly
by a white chevron anterior to the 1st dorsal-fin spine. Color pattern
of body: background cream or white overlaid by three longitudinal
rows of brown blotches, each blotch slightly smaller than eye; dor-
salmost row of 10 blotches located along base of dorsal fin extend-
ing up onto bases of fin rays and membranes, first blotch located at
base of first dorsal-fin spine, last blotch over hypural (forming upper
hypural marking); second row slightly dorsal to midline of body,
each blotch slightly posterior to blotch above; third row slightly
below midline of body and each blotch slightly posterior to blotch
above and thus midway between blotches in dorsalmost row;
distinctness of blotches variable, but usually well defined, distinct
black or dark brown dots often present around edges of blotches;
pelvic fins cream; pectoral-fin rays dusky, membranes cream, two
dark-brown markings, either solid or ring-like with cream centers
located on basal portion of rays, one dorsally and one ventrally; dor-
sal fin with distinct, dark brown spot on membrane between 1st and
2nd dorsal-fin spines in males, remainder of fin with alternating
dark brown and white areas corresponding to the dorsalmost row of
blotches on body, each spine and soft ray with alternating white and
dark brown bars. Anal fin similar to dorsal fin; caudal fin dusky
with scattered black melanophores.

Etymology. —The name variabilis refers to the unusual variation
of the coloration on the side of the head when compared to other
members of the S. oce//ato-complex.

Remarks.— Starksia variabilis is known only from the type series
collected at Santa Marta, Colombia. This species is clearly most
closely related to S. occidentalis which occurs along the Atlantic
coast of Central America, based on the presence of barred lips in
both species, a shared short snout (mean = 48 in S. variabilis and 49
in S. occidentalis whereas all other species in the complex have
means of 56 or greater), and the variable, somewhat reticulated
color pattern on the side of the head of S. variabilis which could be
derived from the basic Y-shaped pattern present in S. occidentalis.

GREENFIELD: STARKSIA OCELLATA COMPLEX

37

UJ £

I -o

345
340
335

£ 330

e

" 325

TI

| 320
o

^ 315

£ 310

| 305

o

^ 300

295
290
285 t

3

- O

i i

1 - 1

13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33
STANDARD LENGTH, mm

FIG. 22. Head length in thousandths of standard length vs. standard length in
millimeters for Starksia variabilis (solid circles) and S. occidentalis from Panama
(open circles).

It can be argued that S. variabilis is simply a geographic variant
of S. occidentalis; however, I have decided to recognize it as a dis-
tinct species for several reasons. Starksia variabilis exhibits modal
differences in counts for anal-fin soft rays and total number of later-
al-line scales that are greater than the differences exhibited between
other species in the complex (i.e., S. ocellata and S. culebrae) and
also greater than those exhibited by other species pairs in this
morphologically conservative genus (i.e., S. sluiteri, S. fasciata, and
S. y-lineata). In other members of the species complex, coloration on
the side of the head has been considered to be a significant character
and S. variabilis exhibits a pattern that despite its variability is
clearly different from S. occidentalis. This variability in itself may
serve as an additional indication of genetic distinctness, since in S.
occidentalis there is little variation in coloration of the side of the
head. In addition, when the freshly-caught specimens of S. variabilis
were received they still retained some of their live coloration, which
revealed that the ring-like markings under the eye and cheek were
lemon yellow. In contrast, live specimens from Belize have white
markings and no yellow: Starksia variabilis has a shorter head

38 FIELDIANA: ZOOLOGY, VOLUME 73

length (mean = 304) than S. occidentalis (mean = 310). This charac-
ter exhibits allometric growth, with smaller individuals having
longer heads; however, specimens of comparable size are separable.
When values for individuals from the total range of S. occidentalis
are plotted against the values for S. variabilis, considerable overlap
occurs; however, when only the Panam6 population (which is geo-
graphically closest to S. variabilis) is plotted, 100 per cent separa-
tion is possible. The Panam£ population of S. occidentalis exhibits
values of 317 or greater, whereas S. variabilis exhibits values of 316
or less (fig. 22). This shift in values in adjacent populations of S. oc-
cidentalis can be interpreted as character displacement, a feature in-
dicating genetic divergence.

The lack of collections from the area between Santa Marta, Colom-
bia, and Panam6 is indeed unfortunate, since this is a straight-line
distance of at least 500 km. and the coastline distance would be even
greater. It is recognized that clinal variation between these loca-
tions is possible; however, until further collections are obtained I
recognize these two species as distinct.

Starksia brasiliensis (Gilbert). Figures 23-25. Tables 1, 2.

BrannereUa brasiliensis Gilbert 1900, p. 180, pi. 9, fig. 1 Type locality: coral reef near
Macei6, Brazil.

Diagnosis.— A species of Starksia with essentially naked belly;
simple orbital cirrus; genital papilla and first anal-fin spine in adult
male united along entire length (fig. 24), papilla projecting beyond
tip of spine a distance equal to one-fourth length of spine; first anal-
fin spine longer than second anal-fin spine; obvious pelvic-fin rays
1,2; pectoral-fin rays 14; dorsal-fin elements XX-XXI.7-8 (usually
XXI.7); anal-fin elements 11,16-17 (usually 11,17); lateral-line scales,
15-16 in arch (usually 15) and 19-20 in straight (usually 20), total

FlG. 23. Starksia brasiliensis, GCRL 9476, male, 24.6 mm. S.L., Brazil.

GREENFIELD: STARKSIA OCELLATA COMPLEX

39

FIG. 24. Gonopodium of
Starksia brasiliensis. GCRL
9476, 24.6 mm. S.L. Line equals
1 mm.

34-36 (usually 35); no dark diagonal bar on lower part of pectoral-fin
base; vertebrae 30(1), 33(3); body color pattern not consisting of
well-defined, dark bands. May be distinguished from its closest con-
geners S. ocellata, S. guttata, S. culebrae, S, occidentalis, and S.
variabilis in having a single row of infraorbital pores rather than
two rows. It may be further separated from S. culebrae, S. occiden-
talis, and S. variabilis by lacking distinct black vertical bars on the
lips. Starksia brasiliensis is clearly separable from S. guttata and S.
ocellata in having a very narrow Y-shaped pale bar extending from
the orbit posteriorly to the preopercular margin.

Description.— Meristics are presented in the diagnosis. Measure-
ments are presented in Table 2 and are based on the four specimens
listed on p. 32.

Color pattern of head. — Starksia brasiliensis is clearly separable
from other members of the species complex by the presence of a very
narrow, Y-shaped pale bar that extends from the orbit posteriorly to
the preopercular margin. The diameter of the bar directly above the
maxillary is considerably less than one-half of the pupil diameter,
whereas in S. guttata the diameter of the bar is equal to or greater
than half of the pupil diameter. A few small, distinct, dark ring-like
markings with pale centers are present under the ventralmost por-

FiG. 25. Typical head color
pattern of Starksia brasiliensis.

40 FIELDIANA: ZOOLOGY, VOLUME 73

B

FlG. 26. Head pore patterns for (a) Starksia brasiliensis , GCRL 9476 and (b) S. gut-
tata, ANSP 117893.

tion of the Y. An additional dark ring-like marking may be present
inside the Y, as well as three larger ring-like markings on the oper-
cle. The lips lack distinct black vertical bars, but are peppered with
black pigment that is more heavily concentrated anteriorly (fig. 25).
The lack of distinct vertical bars on the lips separates this species
from S. culebrae, S. variabilis, and S. occidentalis . This color pattern
is clearly illustrated in Gilbert (1900, pi. 9, fig. 1).

Remarks. —Starksia brasiliensis is known only from Brazil, south
of Recife. It is possible that the mouth of the Amazon forms a bar-
rier to its northward dispersal; however, shallow-water marine col-
lections from the mouth of the Amazon north to Trinidad are not
available. In addition to the distinctive color pattern, S. brasiliensis
has fewer scales in the lateral line and a lower number of soft dorsal-
fin rays. In addition, S. brasiliensis may be separated from all other
species in the complex by having a single series of infraorbital pores
rather than two series (fig. 26). This species bears the closest resem-
blance to S. guttata, which is geographically closest. In addition to
the similarities of the Y-shaped pattern on the head, the body of S.
brasiliensis tends to approach a spotted pattern as in S. guttata, but
not distinctly so.

Material examined— Brazil-CAS-SU 7750(1) 25.0 mm. SL,
holotype; CAS-SU 53510(1) 20.8 mm., paratype; GCRL 9476(2) 24.6
and 26.0 mm.; Bahia, Isla Itaparica, Barra do Gil, 13°00'S lat.,
-38°37'W. long.

DISCUSSION
The recognition of five separate species within the currently rec-

GREENFIELD: STARKSIA OCELLATA COMPLEX 41

ognized species limits of S. ocellata (sensu Bohlke and Springer,
1961), in addition to S. guttata, presents another example of the im-
portance of color patterns within the genus Starksia. Gilbert (1965)
discussed the fact that three species, S. y-lineata, S. fasciata, and S.
sluiteri, can only be distinguished by color pattern, exhibiting no
differences in meristics or morphometrics. Meristic and morphomet-
ric differences between other species pairs is often only modal. I
believe that the color pattern on the lips and side of the head are par-
ticularly important as possible mechanisms of species recognition.
Breder (1941) stated that when a gravid female of the clinid Para-
clinus marmoratus approached a nesting site guarded by a male, the
male underwent a striking color change, "... an intense purple col-
oration which was nearly black and dotted with brilliant metallic
blue." In addition, a head-on, open-mouth threat display was ob-
served in the male. In the closely related family Chaenopsidae,
Wickler (1967) has shown that Emblemaria pandionis has a head-on
display that includes opening the mouth and swaying the head from
side to side. Greenfield and Johnson (MS) have reported similar dis-
plays for Acanthemblemaria spinosa. Wickler (1967, p. 541) noted
the following for blennioid fishes: "Nodding in blennies can be
either a courtship or a threat movement, depending on the particu-
lar species. Since these fishes usually lie within a hole, with only the
head and sometimes the forepart of the body visible, colour patterns
underlining the nodding movements are mainly confined to these
parts of the body. Head and face colourations are, therefore, charac-
teristic for the species and/or sex or motivational state of the fish"
(italics mine).

The importance of head coloration in species recognition is further
supported by the fact that within the S. ocellata-complex, species
with and without bars on the lips have alternating ranges (fig. 27).
Starksia ocellata from Florida lacks bars on the lips, whereas the
species to the south, S. culebrae, has distinct black bars on the lips.
Starksia guttata, which occurs to the south of S. culebrae and
reaches west along the coast of South America at least to Curacao,
lacks bars on the lips. Farther west in Colombia S. guttata is re-
placed by S. variabilis, which has very prominent black bars on the
lips. Starksia brasiliensis lacks bars on the lips, as does S. guttata;
however, it occurs (as far as is known) far to the south of S. guttata,
south of Recife in Brazil. The only pair of species for which this cor-
relation does not hold includes S. variabilis from Colombia and S. oc-
cidentalis from the coast of Central America. Both of these species

42

FIELDIANA: ZOOLOGY, VOLUME 73

FIG. 27. Distribution of the Starksia ocellata superspecies: A, S. ocellata; B, S.
culebrae; C, D, S. guttata; E, S. variabilis; F, S. occidentalis; G, S. brasiliensis.

have bars on the lips; however, they differ in the coloration of the
sides of the head, in addition to which S. variabilis has lemon yellow
spots under the eye and on the cheek, whereas S. occidentalis has
white spots.

As far as is known, all of the species within the S. ocellata-com-
plex have allopatric distributions. In fact, in the case of S. culebrae
and S. guttata, there appears to be a rather sharp break between
these two forms at St. Vincent Island. However, the possibility of
past or present parapatry is suggested by the two cases of apparent
character displacement (S. occidentalis and S. variabilis; S. culebrae
and S. guttata), which usually develops as a result of interactions
while in contact. The allopatric distribution of these species could be
considered evidence that the observed variation is a£ the subspecific
level. However, on the basis of coloration, and in some cases meris-
tics and morphometrics, it is possible to identify all individuals
without knowledge of the locality from which they were obtained.

GREENFIELD: STARKSIA OCELLATA COMPLEX 43

Thus, the morphological differentiation is above that normally ob-
served at the subspecific level. This complex of entirely allopatric
species seems to fit well into the superspecies category proposed by
Amadon (1966, p. 245). Amadon defined the superspecies as follows,
"A group of entirely or essentially allopatric taxa that were once
races of a single species but which now have achieved species
status." Mayr (1963, p. 499) has defined the category as "... a
monophyletic group of entirely or essentially allopatric species that
are too distinct to be included in a single species." Following the ter-
minology proposed by Amadon, each of the species within the <S.
ocellata superspecies would be termed an allospecies. As summar-
ized by Amadon (1966, p. 247), the use of the superspecies concept
"... will remove the temptation to treat as subspecies various allo-
patric forms which are so distinct as to make such a procedure ques-
tionable."

The discovery of this superspecies with its six allospecies poses
some interesting zoogeographical questions. Briggs (1974) has
divided the tropical western Atlantic into three zoogeographical
provinces, Caribbean, Brazilian, and West Indian. The recognition
of Starksia braziliensis as a distinct species adds to the level of
endemism for the Brazilian Province. In an example from another
family, investigation of the Eupomacentrus fuscus -complex showed
that E. fuscus is present only off Brazil, whereas E. dorsopunicans
(previously called E. fuscus) replaces it to the north (Greenfield and
Woods, 1974). Gilbert (1977) has shown that Apogon maculatus is
replaced by A. americanus in Brazil. Thus it is likely that further in-
vestigations of widespread species may show that levels of
endemism in the Brazilian Province are greater than has previously
been recognized.

Briggs has also recognized the West Indian Province (Bahamas to
Grenada) as being distinct from the Caribbean Province (including
Florida, Central America, and the northern coast of South America),
based mainly on the level of endemism. For the fishes, this decision
was based in part on work on the fishes of the Bahama Islands
(Bohlke and Chaplin, 1968), and work on the seven-spined gobies
(Bohlke and Robins, 1968). Briggs states that for the gobies, of the
24 species occurring in the West Indies, only 11 appear to be shared
with the mainland (Caribbean Province). Our collections from Belize
(of which only a portion of the gobies have been identified) and the
records from Colombia presented by Palacio (1974) reduce the
number of species occurring only at the Bahama Islands, Antilles,

44 FIELDIANA: ZOOLOGY, VOLUME 73

or both to only five. Briggs also states that of the 466 shore species
treated by Bohlke and Chaplin (1968), 87, or about 19 per cent, have
not been taken outside the West Indies. Examining just the blenni-
oid fishes collected from Belize and Honduras shows that 14 of the
species considered to be in the group of 87 also occur along the coast
of Central America (Greenfield and Johnson, MS). In addition, three
more species of blennioids described since Bohlke's and Chaplin's
treatment, and considered to occur only in the West Indies, have
been taken along the Central American coast. If unpublished rec-
ords from other families were considered, the 77 species left in this
category would be further reduced. Thus, additional collecting along
the coast of Central and South America has shown that the distribu-
tions of many reef-fish species in the tropical Western Atlantic are
wider than was once believed, and that the levels of endemism cited
for a particular locality may simply be a function of the intensity of
the collecting.

The separation of the Starksia ocellata superspecies into six allo-
species presents a paradox. Whereas other reef species are being
shown to have widespread distributions throughout the tropical
Western Atlantic, the species of Starksia exhibit distinct, restricted
distributions. Although this pattern is unusual, it is not unique. Col-
lette (1974) reviewed the toadfish genus Sanopus, recognizing two
species from Yucatan, Mexico (S. splendidus and S. johnsoni), one
species from Belize (S. astrifer), and a fourth (S. barbatus) from Hon-
duras south to Panama. The two Yucatan species are sympatric;
however, this pair and all other species appear to have allopatric
distributions. Intensive collections throughout Belize and Hon-
duras have emphasized this separation: the only species of Sanopus
we have collected at Belize is S. astrifer (Greenfield and Greenfield,
1973), and a month of intensive collecting in Honduras yielded only
specimens of S. barbatus. Sanopus astrifer and S. barbatus appear
to occupy very similar habitats, caves or hollows underneath coral
heads, usually with sand bottoms. Collette (1974) indicates similar
habitats for S. splendidus and S. johnsoni.

It would thus appear that attempts to divide the fish fauna of the
tropical Western Atlantic into provinces is premature. Although re-
cent strides have been made in investigating the fish faunas of Cen-
tral America and the northern coast of South America, many areas
lack proper coverage. Until such surveys have been completed, the
use of levels of endemism to designate fish faunal provinces will be
meaningless.

GREENFIELD: STARKSIA OCELLATA COMPLEX 45

A KEY TO THE ALLOSPECIES OF THE SUPERSPECIES Starksia ocellata

1. Infraorbital pores in a single series (fig. 26a); narrow Y-shaped pale bar extending
from orbit posteriorly to preopercular margin, diameter of bar directly above
maxillary considerably less than one-half pupil diameter; lips lacking distinct ver-
tical bars S. brasiliensis (Gilbert)

Brazil

Infraorbital pores in two rows (fig. 26b); no Y-shaped pale bar on side of head or, if
present, equal to or greater than one-half pupil diameter; distinct black vertical
bars on lips either present or absent 2

2. Lips with distinct black vertical bars 3

Lips lacking distinct black vertical bars 4

3. Side of head with a pale horizontal bar running from orbit posteriorly past edge
of preopercle onto opercle, not branching into a Y (fig. 7); snout length 46-72,
mean 58 S. culebrae (Evermann and Marsh)

Greater Antilles and south to St. Vincent

Side of head without a simple pale horizontal bar; pale area either Y-shaped or
reticulated and enclosing dark blotches; snout length 36-61, mean 48 or 49 5

4. Side of head with small, dark ring-like markings with light centers, most included
within a horizontal pale area running from the orbit posteriorly to the edge of the
preopercle (fig. 4) S. ocellata (Steindachner)

Florida north to North Carolina

Side of head usually with a pale horizontal bar originating from the orbit and
branching into a definite Y posteriorly, occasionally with dark ring-like mark-
ings, or overlaid with solid dark spots (fig. 10a,b,c) S. guttata (Fowler)

Curacao to Trinidad and north up Lesser Antilles
to Tobago Cays in the Grenadines.

5. Side of head with a Y-shaped pale bar usually bordered ventrally with distinct
dark ring-like markings (fig. 18); anal-fin soft rays 16-19, usually 17

S. occidentalis n. sp.

Atlantic coast of Central America

Side of head without a simple Y pattern, pale area reticulated and enclosing dark

blotches (fig. 21); anal-fin soft rays 17-18, usually 18 S. variabilis n. sp.

Colombia

ACKNOWLEDGEMENTS

I am indebted to the government of Belize, and especially to
Winston Miller, Fisheries Administrator, for granting permission to
collect fish specimens. I would also like to thank the government of
the Republic of Honduras and especially Lie. Humberto Caballero L.,
Director General de Recursos Naturales Renovables, for granting
permission to collect fish specimens. The following persons assisted
in field work: A. Drew, M. Drew, G. Glodek, T. Greenfield, N.
Hylton, R. K. Johnson, F. Miller, R. R. Miller, J. Russo, D. Wildrick,
R. Williamson, and R. L. Woods. I would also like to thank: Rev.
Leonard E. Dieckman, S.J. for assistance in arranging field work in

46 FIELDIANA: ZOOLOGY, VOLUME 73

Belize; Gilbert and Marsha Jo Lomont for providing accommoda-
tions at Glover's Reef Village; Nick Hylton, Captain of the MS.
Miss Sabrina, for providing invaluable assistance in field work in
Honduras. Ms. J. Glaser provided photographs of the specimens
and drawings and prepared the graphs. The following curators kind-
ly lent material for study: Dr. M. Boeseman, Rijksmuseum van
Natuurlijke Historic; Dr. James E. Boh Ike, The Academy of
Natural Sciences of Philadelphia; Mr. C. E. Dawson, Gulf Coast
Research Laboratory Museum; Dr. William N. Eschmeyer, Califor-
nia Academy of Sciences; Dr. Carter R. Gilbert, University of Flor-
ida; Dr. Robert K. Johnson, Field Museum of Natural History; Dr.
Robert J. Lavenberg, Los Angeles County Museum; Dr. Richard E.
Spieler, Milwaukee Public Museum; Dr. Victor G. Springer, U.S.
National Museum. I am especially grateful to Sr. Arturo Acero P.,
College of Marine Sciences, Universidad Jorge Tadeo Lozano, Santa
Marta, Colombia, for collecting a series of Starksia for me which
proved to be an undescribed species, S. variabilis. Ms. Lillian
Dempster kindly assisted me with a nomenclatural problem. Dr.
Carter R. Gilbert has been of considerable assistance in locating
specimens, has reviewed the manuscript, and made many valuable
suggestions. My wife, Terry Greenfield, has assisted in the prepara-
tion of the manuscript, has served as a sounding board for various
taxonomic decisions, and has provided most of the assistance in the
field work.

The Associated Universities for International Education provided
transportation to Belize in connection with its summer course in
tropical ecology. This research was supported by grants to D. W.
Greenfield from the Johnson Fund of the American Philosophical
Society (no. 982) and from the Council of Academic Deans, Northern
Illinois University. This paper is based in part on the results of the
Miskito Coast Expedition (1975) to Honduras and Nicaragua, joint-
ly sponsored by Field Museum of Natural History, Chicago; Nor-
thern Illinois University, DeKalb; and the University of Michigan,
Ann Arbor. The expedition was supported in part by grants from
the Johnson Fund (no. 1220) of the American Philosophical Society;
from the Wrigley Fund for Marine Biological Research to R. K.
Johnson; and by a grant from the Graduate School, Northern
Illinois University to D. W. Greenfield.

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UNIVERSITY OF ILLINOIS-URBANA