Revision of the Blenniid

Fish Genus Omobranchus

with Descriptions of Three

New Species and Notes on Other

Species of the Tribe Omobranchini

VICTOR G. SPRINGER

and

MARTIN F. GOMON

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 177
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 177

Revision of the Blenniid

Fish Genus Omobranchus

with Descriptions of Three

New Species and Notes on Other

Species of the Tribe Omobranchini

Victor G. Springer

and Martin F. Gomon

SMITHSONIAN INSTITUTION PRESS

City of Washington

1975
ABSTRACT

Springer, Victor G., and Martin F. Gomon. Revision of the Blenniid Fish Genus

Omobranchus with Descriptions of Three New Species and Notes on Other

Species of the Tribe Omobranchini. Smithsonian Contributions to Zoology,

number 177, 135 pages, 52 figures, 17 tables, 1975.—Omobranchus comprises 19

species, one of which includes two subspecies. Three species are described as

new: O. meniscus, from Thailand, O. steinitzi, from the southern Red Sea, and

O. verticalis, from Queensland. A key is given to all taxa of Omobranchus, and

sexual dimorphism, geographic distribution and variation, habitat, and nomenclature are discussed. All taxa are illustrated and their distributions plotted on

maps. Neotypes are designated for the following species: Blennechis fasciolatus

Valenciennes (1836), Petroscirtes lineolatus Kner (1868a,b), Petroscirtes ferox

Herre (1927).

Omobranchus is native to the Indo-west Pacific, and all taxa, except one subspecies, occur only in areas west of the Andesite Line. One species, O. punctatus,

has been introduced into the Caribbean. Circumstantial evidence indicates that

O. punctatus was probably introduced by ships transporting East Indian coolies

from Madras and Calcutta around southern Africa to the Caribbean between

1845 and the opening of the Panama Canal (1914).

New information (range extensions, taxonomy, errata), since Springer's 1972

Synopsis of the Tribe Omobranchini, is given for the genera Enchelyurus,

Haptogenys, Omox, and Parenchelyurus. Haptogenys quadripora Springer is

synonymized with H. bipunctata (Day).

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5111. SERIES COVER

DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Springer, Victor Gruschka, 1928-

Revision of the blenniid fish genus Omobranchus.

(Smithsonian contributions to zoology, no. 177)

Supt. of Docs, no.: SI 1.27: 177.

1. Omobranchus. 2. Blenniidae. I. Gomon, Martin F., joint author. II. Title. III. Series:

Smithsonian Institution. Smithsonian contributions to zoology, no 177

QL1.S54 no. 177. [QL638.B6] 59I'.08s [597'.58] 74-4249

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402 • Price $2.65 (paper cover)
Contents

Page

Introduction 1

Methods 2

Acknowledgments . 8

Omobranchus Ehrenberg 9

Key to the Species and Subspecies of Omobranchus 17

O. anolius (Valenciennes) 20

O. aurosplendidus (Richardson) 22

O. banditus J. L. B. Smith 24

O. elegans (Steindachner) 26

O. elongatus (Peters) 28

O. fasciolatoceps (Richardson) . 33

O. fasciolatus (Valenciennes) 34

O. ferox (Herre) . 38

O. germaini (Sauvage) 41

O. lineolatus (Kner) 45

O. loxozonus (Jordan and Starks) 48

O. mekranensis (Regan) 50

O. meniscus, new species 52

O. punctatus (Valenciennes) 53

How O. punctatus Became Established in the Caribbean Sea 59

O. rotundiceps (Macleay) 63

O. r. rotundiceps (Macleay) . . 63

O. r. obliquus (Garman) 68

O. steinitzi, new species 70

O. verticalis, new species 71

O. woodi (Gilchrist and Thompson) 72

O. zebra (Bleeker) 74

O. species 76

Questionable Species 76

List of Nominal Taxa of Omobranchus and Their Current Identification 77

Notes on Other Species of the Tribe Omobranchini 78

Enchelyurus ater (Gunther) 78

Enchelyurus flavipes Peters 78

Haptogenys bipunctata (Day) 78

Omox biporos Springer 79

Parenchelyurus hepburni (Snyder) 79

Literature Cited

 8 0

Figures 8-35: The Species of Omobranchus 85-99

Figures 36-52: Graphs 101-135

in

Revision of the Blenniid

Fish Genus Omobranchus

with Descriptions of Three

New Species and Notes on Other

Species of the Tribe Omobranchini

Victor G. Springer

and Martin F. Gomon

Introduction

Springer (1972a) revised the genera of the

blenniid tribe Omobranchini with the exception of

Omobranchus. It is the purpose of our study to

revise Omobranchus and to provide information on

the other species of Omobranchini that has come

to our attention since the 1972 study.

Omobranchus, with 19 species, is the largest of

the six genera in the Omobranchini, which comprises only 29 species altogether. The species of

Omobranchus are small, carnivorous, benthic inhabitants of the shallow, close-shore, marine and

estuarine waters of the Indo-west Pacific (one

species has been introduced into the Caribbean).

At least one species is reported to occur in freshwater. The species most commonly inhabit rocky

and oyster reefs, often near the edge of mangrove

swamps; one species has been observed living in

holes in mangrove roots. Unlike the species of some

Victor G. Springer, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution,

Washington, D.C. 20560. Martin F. Gomon, University of

Miami, Rosenstiel School of Marine and Atmospheric Sciences,

Miami, Florida 33149.

blenniid genera (e.g., Alloblennius, Ecsenius, Cirripectes), the species of Omobranchus are not

definitely known to occur on living corals, although

they may occur on hard substrate near corals. Only

one specimen is definitely known to have been taken

at a depth greater than four meters, while many

specimens have come from depths of less than half

a meter. The species are secretive and few are

brightly colored in life.

The question arises as to how many species of

Omobranchus are yet to be described. Inasmuch

as Omobranchus occurs almost exclusively in shallow, close-shore habitats, which are generally well

collected, one might surmise that few species remain undescribed. Entomacrodus, which also occurs

predominantly in shallow, close-shore waters, was

revised several years ago (Springer, 1967), and considerable collecting in Entomacrodus-type habitats

since 1967 has failed to produce any new species of

Entomacrodus. In Ecsenius, whose species may

occur at greater depths and distances from shore

than those of either Omobranchus or Entomacrodus, new species are continually appearing in

collections (Springer, 1971, 1972b; more undescribed species are at hand). With the possible

exception of the species we treat here as "Omobran1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

chus species," we expect that few new, primarily

marine, species of Omobranchus will be found.

Omobranchus, however, also occurs in estuarine and

freshwater habitats (where Entomacrodus and Ecsenius do not occur). These habitats have not been

well collected in the Indo-west Pacific, and it would

not be surprising to encounter a few undescribed

species of Omobranchus in them.

METHODS

COUNTS.—Dorsal-, anal-, and caudal-fin, vertebral,

and epipleural rib counts were taken from radiographs.

In the dorsal and anal fins each spine and segmented ray (external element) is typically supported by its own pterygiophore (internal element).

Some specimens exhibited a pterygiophore, occasionally two or three, that supported no external

element. The external element count was increased

by one for each such pterygiophore. Usually it was

the segmented-ray portion of the dorsal fin that

lacked external elements, and these anomalies were

most often associated with the posteriormost pterygiophore. Missing external elements anterior to

the posteriormost pterygiophore in either the dorsal

or anal fin can usually be discerned without a

radiograph by the presence of an unusually large

gap between two adjacent external elements.

Total dorsal-fin elements is the sum of the unsegmented (spines) and segmented rays.

With rare exceptions the anal fin has two spines.

In males of some species and in all females, one or

both spines are discernible only on skeletal preparations or from radiographs.

The caudal-fin elements include spinelike dorsal

and ventral procurrent rays and segmented rays

(see Springer, 1968, fig. 10). All rays are typically

simple, but in rare specimens some segmented rays

may be branched once. For most specimens the

count of the segmented rays is identical with that

of the principal caudal-fin rays (those rays that

articulate with a hypural element); however, in

many specimens one or more of the unsegmented,

spinelike rays may articulate with a hypural element or one of the segmented rays may not articulate with a hypural element. The number of segmented rays is the most consistent and useful count

for any species.

Vertebral counts distinguish precaudal and caudal vertebrae. The first caudal vertebra includes the

first centrum with a recognizable hemal spine, and

the last includes the hypural-bearing "centrum"

(actually a complex element). Occasional specimens exhibit fusions among the caudal vertebrae;

caudal vertebral counts for these specimens were

not recorded, with the following exception: the

penultimate centrum is often enlarged and may

bear bifurcate neural and/or hemal spines, but

except for size, the centrum appears normal on

radiographs and was counted as one vertebra.

The pelvic fins invariably consist of a spine (not

visible externally) and two simple, segmented rays.

The pectoral fins consist of 12-14 simple, segmented rays with a strong mode of 13 in all species.

TEETH.—With one partial exception, all species

of Omobranchus normally bear a large, recurved

canine tooth (rarely two) posteriorly on each side

of each jaw. The lower canines are larger than the

upper canines. Mature females of O. fasciolatoceps

lack canine teeth. The remaining teeth in each

jaw, although often slightly pointed, are here considered to be incisors and are much smaller than

the canine teeth. Tooth counts comprise only the

total number of upper jaw (premaxillary) or lower

jaw (dentary) incisor teeth in specimens 25.0 mm

standard length or larger.

All but three of the species of Omobranchus are

characterized by having more incisor teeth in the

lower jaw than in the upper jaw (Table 1). Two

species are characterized by having more than half

their specimens with the same number of teeth in

each jaw, and one species has equal numbers of

specimens with either more teeth in the lower jaw

or an equal number in each jaw. Ten species failed

to exhibit the condition of having more teeth in

the upper jaw than in the lower jaw. Populations

within a taxon were not noticeably different with

regard to the three conditions for numbers of teeth,

except that the Guam population of O. r. obliquus

approached significance (here set at p = .01) in its

difference from the other populations of that subspecies (Chi-square = 6.2; .05 > p > .02; 2 df).

When a difference existed between the number of

teeth in the lower jaw and the number in the upper

jaw, the difference was usually in the range of one

to four teeth. It is not presently possible to explain

the meaning of these differences.

Incisor teeth in either jaw increase in number
NUMBER 177

TABLE 1.—Percentage of specimens of species or populations of Omobranchus exhibiting each of three conditions regarding number of

teeth in upper and lower jaws

Speci«s or population

anolius

aurosplendidus

banditus

elegans

elongatus

fasciolatoceps

fasciolatus

ferox

germaini

lineolatus

loxozonus

mekranensis

meniscus

punctatus

r. rotundiceps

r. obliquuB

Guam

other

steinitzi

verticalis

woodi

zebra

N

34

9

42

154

215

17

50

52

159

5

67

13

2

273

99

45

87

4

3

30

25

Percentage of

specimens with

more lower jaw

teeth than

upper .iaw teeth

47.1

44.4

85.7

88.3

74.0

94.1

76.0

52.0

91.3

100

76.0

100

100

87.2

89.9

55.6

78.1

25.0

66.7

83.3

20.0

Conditions

Percentage of

specimens with

equal numbers

of teeth in

each .law

41.1

44.4

14.3

11.7

22.3

0.0

24.0

46.2

8.2

0.0

24.0

0.0

0.0

10.3

10.1

44.4

21.8

75.0

33.3

6.7

56.0

Percentage of

specimens with

more upper jaw

teeth than

lower .law teeth

11.8

11.1

0.0

0.0

3.3

5.9

0.0

1.9

0.0

0.0

0.0

0.0

0.0

1.8

0.0

0.0

0.0

0.0

0.0

10.0

24.0

with increase in standard length. Linear regression

equations and associated correlation coefficients

(Snedecor and Cochran, 1971) were obtained for

number of teeth in each jaw for each sex of selected populations of each species for which sufficient data were available (Table 2; where only a

limited number of specimens was available for a

species, all specimens were grouped together for

the analysis). Regressions were all positive and

almost all had significant correlation coefficients

(p. < .01; Table 2). Covariance comparisons (Snedecor and Cochran, 1971) were made between males

and females within each population and between

males or females of different populations of the

same species. Covariance comparisons were considered to indicate valid differences when the F values

for either slopes or heights were at the p. ** .001 level

(Table 3). All statistical tests were two-sided and

were computer performed (IBM 1130; based on a

program constructed by P. Poole, Gulf Coast Research Laboratory). Any statistical differences between sexes or populations are discussed in the text.

Selected XY plots (Figures 36-52), mostly of upper

jaw teeth, are presented as examples of species,

population, and sexual differences. These plots are

printed on translucent paper on one side only and
TABLE 2.—Regression equations and associated correlation coefficients for number of upper ber ui or

lower jaw teeth (Y) versus standard length (X) in certain species of Omobranchus (**= significant at p = .01; NS = not significant)

Species, population, sex

anolius

males

females

banditus

males

females

elegans

males

females

elongatus

Madagascar

males

females

Chagos Islands

males

females

Mozambique

males

females

Gulf of Thailand

males

females

fasciolatus

males

females

ferox

males

females

germaini

Hong Kong

males

females

Queensland

males

females

Western Australia

Montebello to Maud

males

females

Shark Bay to Perth

males

females

loxozonus

Japan

males

females

Okinawa

males

females

punctatus

Ceylon

males

females

Gulf of Thailand (east coast)

males

females

China

males

females

Western Australia

males

females

Trinidad

males

females

rotundiceps

Northern Territory

males

females

obliouus

Palau Islands

males

females

Guam

males

females

Palmyra

females

wood!

males

females

»ebra

males

females

Regression equation

lower jaw teeth

Y - 13.6428 + 0.2110X

Y - 14.4446 + 0.2396X

Y = 16.7/74 + 0.1622X

Y = 12.2709 + 0.2849X

Y = 12.6607 + 0.1827X

Y = 13.2074 + 0.1727X

Y • 16.0950 + 0.154U

Y = 14.3674 + 0.1969X

Y = 13.7319 + O.2595X

Y = 7.5892 + 0.4742X

Y = 14.0564 + 0.171 OX

Y - 11.3508 + 0.2701X

Y = 14.2449 + 0.2270X

Y = 9.6895 + 0.4214X

Y = 12.4387 + 0.2811X

Y = 13.9678 + 0.2658X

Y = 12.8756 + 0.1473X

Y « 11.0126 + O.196OX

Y - 14.3719 +0.1841X

Y = 9.5122 + O.3571X

Y = 13.5619 + 0.1986X

Y • 14.3954 + 0.2138X

Y = 12.7926 + 0.2098X

Y - 9.C513 + O.3325X

Y - 8.9368 + 0.3118X

Y = 10.1621 + 0.2837X

Y - 13.5425 + 0.1921X

Y - 10.5804 + O.2758X

Y « 17.9076 + 0.0812X

Y = 13.5538 + 0.2214X

Y = 20.7766 + 0.1706X

Y = 16.7333 + O.3355X

Y - 19.2111 + 0.1754X

Y = 15.7239 + 0.29O7X

Y - 19.4997 + 0.1493X

Y - 18.5295 +0.1740X

Y - 21.9070 + 0.0823X

Y - 18.3451 + 0.1801X

Y - 17.8032 + 0.2526X

Y -20.2373 +0.1969X

Y - 13.7942 + 0.2541X

Y - 11.8393 + O.3376X

Y - 16.3638 + 0.2325X

Y - 10.0620 + 0.4584X

Y = 11.6324 + 0.3471X

Y - 10.2077 + 0.4183X

Y - 10.4757 • 0.4066X

Y - 21.0588 + 0.1094X

Y » 16.8481 + 0.2144X

Y - 8.7708 + 0.2498X

Y - 11.4259 + 0.1845X

N

22

11

28

14

41

42

14

8

7

8

26

27

29

27

27

23

29

16

10

6

35

22

17

20

13

21

39

28

8

7

23

20

35

21

20

15

6

10

18

8

45

38

18

16

23

22

16

15

14

7

13

Correlation

coefficient

0.772 **

C.665 NS

0.703 **

0.860 «

0.842 •*

0.829

 tt£

0.788 «

0.680 NS

0.811 NS

0.932 **

0.692 «

0.751 •*

0.771 **

0.776 **

0.792 «*

0.826 »*

0.748 «*

0.510 NS

0.803 **

0.910 NS1

0.804 **

0.858 «

0.864 **

0.828 *»

0.782 «*

0.787 **

0.910 «*

0.874 **

0.380 NS

0.796 NS

0.834 **

0.845 •*

0.788 **

0.906 **

0.878 **

0.553 NS

0.465 NS

0.645 NS

0.874 **

0.850 **

0.901 «t

0.923 «*

0.830 **

0.661 «*

0.887 **

0.857 •»

0.807 «*

0.515 NS

0.682 M

0.958 •*

0.852 **

Regression equation

upper jaw teeth

Y = 13.9621

Y = 11.9842

Y = 15.828i

Y = 12.5010

Y = 11.4375

Y * 11.0033

Y = 13.9847

Y « 13.8062

Y - 13-2404

Y • 10.6959

Y - 11.7330

Y = 9.6017

Y * 13.0449

Y - 11.2567

Y - 14.2373

Y - 15.7450

Y - 14.5143

Y - 12.2687

Y - 12.1805

Y = 12.6322

Y = 12.0392

Y • 13.4897

Y = 13.1400

Y = 5.5424

Y * 8.0765

Y = 10.1635

Y = 13.1271

Y = 11.7180

Y = 18.0726

Y - 10.5157

Y = 22.9006

Y - 19.0626

Y - 18.3961

Y - 14.4955

Y - 19.1633

Y = 17.9137

Y * 19.7653

Y - 9.6733

Y = 16.9280

Y - 18.7840

Y - 13.5515

Y - 10.1819

Y = 16.1124

Y - 9.6517

Y = 9.5000

Y = 10.7593

Y - 9.2942

Y • 2C.433O

Y - 15.4604

Y = 6.8382

Y - 10.0231

+ 0.1988X

+ 0.2857X

+ 0.H3 U

• 0.2312X

+ O.1635X

+ 0.1770X

+ 0.174'X

+ 0.1914X

+ 0.2224X

• 0.3226X

•f 0.2123X

+ 0.2981 X

• 0.2236X

+ O.3313X

+ O.185OX

+ 0.16J8X

+ 0.C941X

+ O.154OX

+ 0.1838X

+ 0.2212X

+ 0.1810X

+ O.1777X

+ 0.1548X

+ O.3483X

+ 0.2681X

+ 0.2287X

+ 0.1748X

+ 0.2189X

+ 0.0443X

+ 0.2719X

+ O.O526X

+ O.1939X

+ O.1564X

+ O.2756X

+ 0.1140X

+ O.1517X

+ 0.0927X

+ O.4O39X

+ 0.2362X

+ 0.2026X

+ 0.2186X

+ 0.3317X

+ 0.1926X

> 0.4182X

+ 0.3931X

+ 0.3715X

+ 0.4236X

•t- 0.0967X

+ 0.2041x

+ O.3O63X

+ 0.2241X

K

24

11

21

'4

40

42

14

8

7

8

25

27

28

27

29

21

30

16

10

7

35

22

18

20

13

20

40

28

8

7

23

20

34

21

20

14

6

10

18

8

45

37

16

16

24

22

16

17

14

7

13

Correlation

coefficient

0.791 «*

0.576 NS

0.721 «*

0.721 "»

0.866 **

0.902 **

0.821 **

0.696 NS

0.784 NS

0.708 NS

0.733 **

0.775 «

0.798 **

0.710 «

0.692 ••

0.732 • •

0.488 **

0.512 NS

0.774 »*

0.888 **

0.704 *«

0.621 «

0.672 »•

0.865 *«

0.740 •*

0.722 «

0.886 »•

0.830 «

0.173 NS

0.893 •«

0.337 NS

0.629 «•

0.758 **

0.921 «*

0.871 *•

0.521 NS

0.854 *•

0.854 **

O.915 «*

0.774 «

0.846 «

0.920 •»

0.787 **

0.541 «

0.968 «

0.793 «

0.841 **

0.495 NS

0.671 «

0.946 0*

0.865 •»

To be significant, this value should b« 0.917; the regression data, however, have been used in covariance analyses.
NUMBER 177

TABLE 3.—F values for covariance comparisons of regression equations {Table 2)

of lower and upper jaw teeth between males and females within certain species and

populations of Omobranchus (**= significant at p — .001; NS = not significant)

Comparisons

banditus

lower teeth

upper teeth

elegans

lower teeth

upper teeth

elongatus

Mozambique

lower teeth

upper teeth

Gulf of Thailand

lower teeth

upper teeth

fasciolatus

lower teeth

upper teeth

germaini

Hong Kong

lower teeth

upper teeth

Queensland

lower teeth

upper teeth

Western Australia

Montabello to Maud

lower teeth

upper teeth

Shark Bay to Perth

lower teeth

upper teeth

F values

slopes

3.41 NS

1.89 NS

0.14 NS

0.43 NS

2.50 NS

1.67 NS

5.48 NS

2.05 NS

0.06 NS

0.16 NS

2.88 NS

0.20 NS

0.09 NS

0.00 NS

3.75 NS

7.27 NS

0.10 NS

0.20 NS

Degrees of

freedom

1/38

1/37

1/79

1/78

1/49

1/48

1/52

1/51

1/46

1/46

1/12

1/13

1/53

1/53

1/33

1/34

1/30

1/29

F values

heights

0.05 NS

0.22 NS

0.18 NS

0.43 NS

9.47 NS

10.17 NS

22.76 «*

29.18 «*

6.04 NS

4.50 NS

1.42 NS

10.14 NS

22.07 **

9.91 NS

9.94 NS

0.04 NS

0.21 NS

2.48 NS

Degrees of

freedom

1/39

1/38

1/80

1/79

1/50

1/49

1/53

1/52

1/47

1/47

1/13

1/14

1/54

1/54

1/34

1/35

1/31

1/30

Comparisons

loxosonus

Japan

lower teeth

upper teeth

punctatus

Ceylon

lower teeth

Gulf of Thailand

lower teeth

upper teeth

Western Australia

upper teeth

Trinidad

lower teeth

upper teeth

rotundiceps

rotundiceps

Northern Territory

lower teeth

upper teeth

obliquus

Palau Islands

lower teeth

Guam

lower teeth

upper teeth

aebra

lower teeth

upper teeth

F values

slopes

6.48 NS

1.92 NS

7.09 NS

6.71 NS

7.66 NS

6.60 NS

0.52 NS

0.26 NS

6.75 US

9.78 NS

2.42 NS

0.92 NS

0.12 NS

1.33 NS

1.48 NS

Degrees of

freedom

1/63

1/64

1/39

1/52

1/51

1/12

1/22

1/22

1/79

1/78

1/30

1/41

1/42

1/16

1/16

F values

heights

7.16 NS

3.83 NS

28.74 **

8.44 NS

5.63 NS

2.44 NS

0.37 NS

1.31 NS

14.57 *•

4.56 NS

1.93 NS

5.27 MS

6.10 NS

0.03 NS

0.00 NS

Degrees of

freedom

1/64

1/65

1/40

1/53

1/52

1/13

1/23

1/23

1/80

1/79

1/31

1/42

1/43

1/17

1/17

are all drawn to the same scale. The pages containing the plots can be removed and any page superimposed on another to enable comparison of the

plots for different species.

The number of circumorbital bones (Springer,

1968) is reasonably consistent for any given blenniid species (often for a genus). In Omobranchus

the species have either 4 or 5 circumorbital bones

on each side. In those species normally with 5

bones, 4 may occur unilaterally. The dorsalmost

circumorbital bone (dermosphenotic of Springer,

1968) is greatly reduced and often obscured from

view by the next to dorsalmost bone. From two to

seven specimens of each species or subspecies of

Omobranchus were examined for number of circumorbital bones.

MEASUREMENTS.—All measurements were made

with needlepoint dial calipers and recorded to the

nearest 0.1 millimeter. Standard length (SL) was

measured from the midtip of the snout (not necessarily the midtip of the upper lip) to the lateral

midbase of the caudal fin. Head length was measured from the midtip of the snout to the posterodorsal junction of the membranous margin of the

gill opening with the body (dorsal end of gill opening). Dorsal-fin element lengths were measured from

the corner of the posterior angle formed by the

element and the dorsal body contour to the tip of

the element. Caudal-fin ray lengths were measured

from the base to the tip of the ray. The height of

the bladelike, fleshy crest on top of the head was

measured as a perpendicular from the apparent

head contour at the base of the crest to the highest

point on the crest.

OTHER CHARACTERS.—The gill opening is restricted in Omobranchus. The ventral point where the

membrane bordering the gill opening joins the

body relative to the insertion of the pectoral-fin

rays is an important specific character. The gill

opening may loop farther ventrally than the point

of junction between the membrane and the body,

but it is the junction point that we record. When

the gill opening is restricted to the area dorsal to

the pectoral-fin insertion, this condition is recorded

as 0. Otherwise the number of the fin ray (counting

dorsally to ventrally) is recorded. Because the posi-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tion of the opening relative to a particular fin ray

is often a matter of judgment, an allowance of plus

or minus one ray should be assumed. In specimens

smaller than 25 mm SL of any species, the size of

the gill opening may vary considerably and may

extend much farther ventrally than in larger specimens. For that reason we report data only on specimens larger than 24.9 mm SL.

The epipleural rib count is actually a count of

the vertebrae bearing an epipleural rib on either

one or both sides. Occasional caudal vertebrae that

lack epipleural ribs but occur between vertebrae

bearing epipleural ribs are included in the count.

In all cleared and stained specimens, the first epipleural rib is on the first vertebra, and our counts

commenced with the first vertebra even though the

anteriormost epipleural ribs are not distinguishable

on radiographs.

Cephalic sensory pore series and lateral-line tubes

are illustrated in Figure 1. Interorbital pores, all

opening into the frontal bones, are situated in the

interorbital region and number modally 2 or 3 for

a given species. Where the count is 3, there is

10

CO

MP

FIGURE 1.—Diagrammatic illustration of cephalic sensory

pores in Omobranchus. (AN = anterior nostril; CO = circumorbital pores; IO = interorbital pores (both sides included in count); LL = lateral-line tubes (one complete tube

and portion of second illustrated); MP = mandibular pores;

MS = median supratemporal commissural pore; NP = prenasal pores; PN = posterior nostril; PP = preopercular

pores; ST = supratemporal pores.)

typically a pair of anterior pores and a single posterior (apical) pore arranged in a triangular pattern. Where the count is 2, the apical pore is most

often absent. In species possessing a fleshy, bladelike crest on top of the head the apical pore, if

present, is well up on the crest, either on the midline of the crest or displaced to one side. The

circumorbital pores ring the posterior half of the

orbit, beginning dorsally at about the one o'clock

position and extending clockwise to the eight

o'clock position. The two anteriormost circumorbital pores are just posterior or lateral to the nostrils.

There are two prenasal pores on each side, one just

anterior or ventral to each of the nostrils. The only

exception is O. steinitzi, which lacks prenasal pores.

The mandibular, preopercular, and supratemporal

pore series were not used as characters. The total

number of these pores on one side is modally 14

(including the median supratemporal commissural

pore) in all species. Springer (1972a) noted 13

pores in these series of Omobranchus, failing to

include the posteriormost pore of the supratemporal

series in his count.

The lateral-line tubes, when present, are raised,

typically bipored tubes on the body in the area

below the spinous dorsal fin. The tubes are usually

not continuous with each other or with the posterior branch of the supratemporal pore series, although two or three tubes may be connected

occasionally. The tubes are usually close together

but may be widely separated from each other. Pitlike depressions are sometimes present along the

route followed by the lateral-line tubes but were

not included in the lateral-line tube counts. The

anteriormost lateral-line tube rarely originates anterior to the level of the gill opening. In ascertaining

the position of the posteriormost lateral-line tube

relative to a dorsal-fin spine, those tubes that terminated at a level between two spines were recorded

as reaching to below the level of the posteriormost

of the two spines. Where lateral-line tubes were

absent, they were recorded as reaching to below

the level of dorsal-fin spine 0.

All species of Omobranchus except O. banditus

possess a ventrally free fold of skin on the lower

jaw near the corner of the mouth (Figure 2), which

we term the lower-lip flap.

Color pattern descriptions are usually composites

based on a number of specimens. For this reason no

single specimen is likely to exhibit all features of
NUMBER 177

LOWER-LIP

FLAP

a

FIGURE 2.—Diagrammatic illustration of lower-lip flap in

Omobranchus: a, lower-lip flap absent (only O. banditus);

b, lower-lip flap present (all species except O. banditus).

the color pattern given for a species. We use "bars"

and "bands" to refer to vertically oriented markings; "stripes" refer to longitudinally oriented

markings.

Larvae are here defined as specimens in which

there is a posteriorly directed projection at the

angle of the preopercle (Figure 3). The projection

apparently originates as a bony spine, which later

becomes fleshy and flexible before being lost (absorbed?) entirely. Most larvae also possess a large

area of dark melanophores on the pectoral fin and

a relatively large gill opening. All collections, except one, that included larvae also included adults.

The exception, which includes the smallest larvae,

was a night collection made at the surface near a

light, and may indicate that pelagic stages of Omobranchus are more common than the available evidence indicates.

The synonymies include only references to original species descriptions.

In the lists under "Material Examined" abbreviated locality data are given, followed by institutional abbreviations, catalog numbers (if cataloged),

and, in parentheses, the number of specimens (if

more than one) and size range in mm SL.

ABBREVIATIONS.—The following institutional abbreviations are used to denote the location of specimens studied.

AMNH American Museum of Natural History, New York

City

AMS Australian Museum, Sydney (catalog numbers

usually with the prefixes I, IA, and IB)

ANSP Academy of Natural Sciences of Philadelphia

BMNH British Museum (Natural History), London

BPBM Bernice P. Bishop Museum, Honolulu

FIGURE 3.—Morphology of Omobranchus larvae: a, presumable larva of O. lineolatus (after

Weber, 1913); b, presumable larva of O. r. obllquus (after Seale, 1935).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

CAS California Academy of Sciences, San Francisco (catalog or register numbers frequently with the prefixes SU or GVF)

DASF Fisheries Research, Department of Agriculture.

Stock and Fisheries, Kanedobu, Papua

FMNH Field Museum of Natural History, Chicago

GCRL Gulf Coast Research Laboratory, Ocean Springs,

Mississippi

HUJ Hebrew University, Department of Zoology, Jerusalem (catalog numbers frequently with the prefixes Fish, F, and E)

MCSN Museo Civicio di Storia Naturale, Genova

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts

MNHN Museum National d'Histoire Naturelle, Paris (catalog numbers frequently with a prefix A)

NFIS Natur-Museum und Forschungs-Institut Senckenberg, Frankfurt

NMV Naturhistorisches Museum, Vienna

QM Queensland Museum, Brisbane (catalog numbers

frequently with the prefix I)

RMNH Rijksmuseum van Natuurlijke Historic Leiden

RUSI JX.B. Smith Institute of Ichthyology, Rhodes University, Grahamstown, South Africa

SAM South African Museum, Capetown

SIO Scripps Institution of Oceanography, La Jolla,

California (catalog numbers with the prefix JEM)

UG University of Guam, Agana

UMML School of Marine and Atmospheric Sciences, University of Miami, Florida

USNM Division of Fishes, National Museum of Natural

History, Smithsonian Institution, Washington,

D.C. (catalog numbers of the former United

States National Museum)

UZMR Universitetets Zoologiske Museum, Copenhagen

(catalog numbers with the prefixes CN and P)

WAM Western Australian Museum, Perth (catalog numbers with the prefix P)

ZSIC Zoological Survey of India, Calcutta (catalog numbers with the prefix F)

ZITU Zoological Institute, University of Tokyo

ZMA Zoologisch Museum, Universiteit van Amsterdam

ZMB Institut fur Spezielle Zoologie und Zoologisches

Museum, Berlin (East)

ZSZM Zoologisches Staatsinstitut und Zoologisches Museum, Hamburg

ACKNOWLEDGMENTS

A large number of individuals aided our work.

For the loan or exchange of specimens or for providing information on material in the collections

under their supervision, we wish to thank the following people: Dr. D. E. Rosen (AMNH); Drs.

J. R. Paxton and D. Hoese (AMS); Dr. J. E. Bohlke

and Mr. W. F. Smith-Vaniz (ANSP); Messrs. P. J.

Whitehead and G. Palmer (BMNH); Dr. J. E.

Randall (BPBM); Dr. W. N. Eschmeyer and Ms.

P. Sonoda (CAS); Miss P. Kailola (DASF); Mr.

L. P. Woods (FMNH); Mr. C. E. Dawson (GCRL);

Drs. H. Steinitz (deceased) and A. Ben-Tuvia

(HUJ); Dr. E. Tortonese (MCSN); Dr. T. R. Roberts and Mrs. M. Dick (MCZ); Dr. M. L. Bauchot

(MNHN); Dr. W. Klausewitz (NFIS); Dr. P. Kahsbauer (NMV); Mr. R. J. McKay (QM and WAM);

Dr. M. Boeseman (RMNH); Mrs.. M. M. Smith and

Dr. T. H. Fraser (RUSI); Dr. P. A. Hulley (SAM);

Dr. J. E. McKosker (SIO); Dr. R. S. Jones (UG);

Dr. C. R. Robins (UMML); Dr. J. Nielsen (UZMK);

Drs. A. G. K. Menon and P. K. Talwar (ZSIC); Dr.

T. Abe (ZITU); Dr. H. Nijssen (ZMA); Drs. K.

Deckert and C. Karrer (ZMB); Dr. W. Ladiges

(ZSZM); Dr. I. Rubinoff, Smithsonian Tropical Research Institute; Dr. W. R. Courtenay, Jr., Florida

Atlantic University; Dr. I. Tomiyama, Amakusa

Marine Biological Laboratory.

Other colleagues furnished important information, advice, stimulating discussions, or critical

comments on portions of the manuscript: Dr. C.

Sabrowsky, Systematic Entomology Laboratory, Department of Agriculture; Dr. M. Jackson, National

Museum of History and Technology, Smithsonian

Institution; Dr. S. H. Weitzman and Mr. R. H.

Goodyear (USNM); Dr. C. R. Robins, University

of Miami, School of Marine and Atmospheric

Sciences; Dr. B. B. Collette, Systematics Laboratory,

and Dr. W. I. Aron, Ecology and Conservation

Office, National Oceanic and Atmospheric Administration.

The entire manuscript was read and criticized by

Dr. R. H. Gibbs, Jr. (USNM), Dr. L. W. Knapp,

Smithsonian Oceanographic Sorting Center, and

Mr. W. F. Smith-Vaniz (ANSP).

We are indebted to the following individuals for

considerable routine curatorial assistance: Messrs.

E. N. Gramblin, B. B. Hoddess, and F. J. McKinney; Mrs. S. J. Karnella and Ms. J. Ryle, technicians

(USNM). Ms. C. Feely, Arlington, Virginia, generously gave of her time voluntarily. Mesdames D.

Dunn and J. Williams provided arduous secretarial

assistance.

We are also grateful for the extensive efforts of

Mrs. M. F. Bell, Editor, and Mr. V. Shaffer, Series

Production Manager, Smithsonian Institution Press,

who edited and prepared our manuscript for the

printer and shepherded it through to publication.
NUMBER 177

Some of the specimens used in our study were

obtained through support provided by a Smithsonian Foreign Currency grant (Biota of the Red

Sea) to the Hebrew University and Smithsonian

Institution and a grant from the National Geographic Society to V. G. Springer as principal

investigator.

Omobranchus Ehrenberg

Omobranchus Ehrenberg, in Valenciennes in Cuvier and

Valenciennes, 1836:287 [first appearance in synonymy, subsequently made available by Swainson, 1839:274; typespecies: O. fasciolatus Valenciennes, in Cuvier and Valenciennes, 1836, by monotypy].

Graviceps Fowler, 1903:170 [type-species: Petroscirtes elegans

Steindachner, 1876, by original designation].

Cyneichthys Ogilby, 1910: 55 [type-species: Blennechis anolius

Valenciennes, in Cuvier and Valenciennes, 1836, by original

designation, in parentheses1, and monotypy].

Poroalticus Fowler, 1931:403 [type-species: Poroalticus sewalli

Fowler, 1931 ( = Blennechis punctatus Valenciennes, in

Cuvier and Valenciennes, 1836), by monotypy].

Pauloscirtes Whitley, 1935:351 [type-species: Petroscirtes

obliquus Garman, 1903, by original designation].

Cruantus J. L. B. Smith, 1959: 234 [type-species: Omobranchus dealmeida J. L. B. Smith (=Petroscirtes ferox Herre,

1927), by original designation].

NOMENCLATURAL DISCUSSION.—The synonymy of

Omobranchus was discussed by Springer (1972a)

and will not be repeated here. However, some comment is required on the date and authorship of the

name Omobranchus.

The genus Omobranchus and the species Omobranchus fasciolatus first appeared in Valenciennes

(in Cuvier and Valenciennes, 1836) as follows:

Le Blennechis fascie

(Blennechis fasciolatus, nob.; Omobranchus fasciolatus,

Ehremb.).

Valenciennes based his description on a plate

that he erroneously assumed Ehrenberg had published; however, the actual appearance of the plate

(and Ehrenberg's name, Omobranchus fasciolatus)

did not appear until much later (Hemprich and

Ehrenberg, 1899).

Thus, the first appearance of Omobranchus can

be construed to have occurred in synonymy, and the

same is true for O. fasciolatus Ehrenberg.

1

 For a discussion of type-spedes designations in parentheses, see Hubbs, et al., (1963).

According to the International Code of Zoological

Nomenclature, Article 11 (d), "A name first published as a synonym is not thereby made available

unless prior to 1961 it has been treated as an available name with its original date and authorship,

and either adopted as the name of a taxon or used

as a senior homonym."

Swainson (1839:79) next mentioned Omobranchus, as follows:

"Lastly comes Dr. Riippel's sub-genus Petroscertes, joined indeed to Blennichus by M. Valenciennes, but at once distinguished by a single series

of small setaceous teeth, without the large canines

of Omobranchus Ehr. (Blennichus Val.)." (Note

misspelling of Blennechis and Petroscirtes).

On page 182, Swainson gives the genus Blennius

under which he recognizes and characterizes several

genus-group taxa. Among these is Omobranchus,

but not Blennechis. On page 274, Swainson gives

and characterizes "Omobranchus Ehrenberg" as a

genus-group taxon under Blennius. He included

several species in Omobranchus, among which are

"filamentosus. Cuv. xi. 280." and "fasciolatus.

Ehrenb. Ib. 287."

Although Swainson referred to "Cuv." for the

last-mentioned names, it is obvious from the volume

and page citations that he was referring to Valenciennes in Cuvier and Valenciennes (1836).

On the basis of Swainson's treatment of Omobranchus and O. fasciolatus, these names were made

available with their original date and authorship.

There is no need to consider Omobranchus fasciolatus Ehrenberg further as it is a junior objective

homonym of Blennechis fasciolatus Valenciennes.

However, the authorship of Omobranchus is a problem as the International Rules of Zoological Nomenclature furnish no guide as to how to assign

authorship to names first appearing in synonymy.

Omobranchus has been attributed to various authors: De Beaufort (in de Beaufort and Chapman,

1951) attributed it to Cuvier and Valenciennes;

Springer (1963), to Swainson. We choose to recognize Ehrenberg as the author of Omobranchus,

which action is in keeping with the senior author's

recent treatments of the genus (Springer, 1968,

1972a).

DIAGNOSIS.—Dorsal-fin spines X-XIV (rarely X);

segmented dorsal-fin rays 16-26; total dorsal-fin elements 28-37; anal fin II, 18-27; pectoral fin 12-14

(rarely 12 or 14 in any species); pelvic fin 1,2; seg-
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mented caudal-fin rays 12-15 (modally 13 in all

species); dorsal + ventral procurrent caudal-fin

rays 8-18 (rarely 8, 17, or 18), about equally divided

between dorsal and ventral portions; all fin elements unbranched except for an occasional caudalfin ray; precaudal vertebrae 10-12; caudal vertebrae

26-33; total vertebrae 36-43; epipleural ribs 10-25

(rarely 10 or 25); ventral hypural plate (Springer,

1968) autogenous; circumorbital bones 4 or 5; postcleithra 2, not fragmented; nasal bones separate in

juveniles and adults; dentaries connected by suturing joint (except in mature females of O. fasciolatoceps, where the dentaries merely abut); enlarged

posterior canine posteriorly on each side of each

jaw (except in mature females of O. fasciolatoceps,

which lack canines); total premaxillary incisoriform

teeth in specimens 25 mm SL and larger 14-38;

total dentary incisoriform teeth in specimens 25

mm SL and larger 16^8; kinethmoid (Springer,

1968) absent; basisphenoid well developed; interopercle with posteriorly projecting spur (Springer,

1968, fig. 16); pterosphenoid reduced, excluded

from external surface of skull; frontal bones not

fused in adults; interorbital sensory pores 1-4 (modally 2 or 3 in all species); circumorbital sensory

pores 6-10 (modally 7-9); prenasal sensory pores

present (except absent in O. steinitzi); anterior and

posterior nostrils present; bipored lateral-line tubes

on body 0-9; gill opening in specimens 25 mm and

larger varying from restricted entirely above level

of dorsalmost pectoral-fin ray to extending ventrally

to level of 8th pectoral-fin ray; raised, bladelike,

fleshy crest on top of head present or absent; no

cirri on head (the anterior nostrils of Omobranchus

exit from short, fleshy tubes that might be confused

with cirri; only Laiphognathus in the Omobranchini has head cirri, and these arise from the anterior and posterior rims of both the anterior and

posterior nostrils).

COMPARISONS.—Omobranchus is most similar to

Omox and Parenchelyurus within the Omobranchini. It differs from Omox in lacking a kinethmoid

bone, in having strongly modally 2-3 interorbital

pores (versus 4 in Omox), in having a median predorsal supratemporal commissural pore (usually

absent in Omox), and in having the gill opening

rarely extending ventrally below the 6th pectoralfin ray (opposite 8th—11th ray in Omox). Omobranchus differs from Parenchelyurus in having 3, as

opposed to 2, mandibular sensory pores (although,

exceptional specimens of Omobranchus may have

2 on one side). Inasmuch as pores are often difficult

to see in the Omobranchini, it is possible that

specimens of the two species of Parenchelyurus may

be erroneously assigned to Omobranchus. The color

patterns of the two Parenchelyurus species, however,

are quite unlike those of any Omobranchus species

(see Springer, 1972a, figs. 15-16).

SEXUAL DIMORPHISM.—The species of Omobranchus exhibit a variety of different types of sexual

dimorphism. In some species the sexes are so different that they might appear, on superficial examination, to belong to separate species.

PRIMARY SEXUAL DIMORPHISM.—The primary

sexual dimorphism of Omobranchus is that typical

of most blenniids. The first (and occasionally the

second) anal-fin spine of females is greatly reduced,

not visible externally, and completely included in

a somewhat triangular, bilobed, fleshy swelling embodying a relatively large, pocketlike urogenital

opening and a papilla, which Tomiyama (1952)

found to indicate the tip of the first anal-fin spine.

The papilla is greatly reduced and frequently not

noticeable in some species (O. punctatus, mekranensis, banditus). In males (except O. punctatus,

mekranensis, banditus and steinitzi) the anal-fin

spines are usually distinct and the urogenital opening is a minute orifice at the tip of a short, usually

truncate, tube. In the excepted species one or both

anal-fin spines are reduced, closely applied, imbedded in thick skin, and not ordinarily visible

externally. The primary sexual dimorphism of O.

elegans has been figured by Tomiyama (1950), of

O. loxozonus by Tomiyama (1951), of O. fasciolatoceps (as O. uekii) by Tomiyama and Abe (1953)

and of O. punctatus by Tomiyama (1952).

SECONDARY SEXUAL DIMORPHISM.—Most of the

secondary sexual modifications found in Omobranchus appear to confer a higher degree of visibility

on males (larger size, larger structures, more striking color pattern). Higher visibility may function

in sex recognition and/or in territory maintenance.

Modifications in females (more teeth, and other

structural changes in the jaws of one species) serve

an unknown function. Following are the main secondary sexual modifications noted.

1. Standard Length: In 18 of the 20 taxa of Omobranchus the largest male is larger than the largest

female (Table 4; a female O. verticalis arrived too

late to be included in the table; see species account).
NUMBER 177 11

TABLE 4.—Comparisons of sizes (standard length in mm) of species and

subspecies of Omobranchus

Species or subspecies

anolius

aurosplendidus

banditus

elegans

elonfcat.ua

fasciolatoceps

fasciolatus

ferox

germaini

lineolatus

loxozonus

mekranensis

meniscus

punctatua

r. rotundiceps

r. obliguus

steinitzi

verticalis

woodi

zebra

Largest male

60.6

112.2

55.6

64.3

53.8

63.4

49.5

59.9

65.4

36.3

70.7

50.7

60.1

95.0

74.2

56.2

32.0

49.5

81.1

48.1

Largest female

47.0

97.7

46.2

58.0

45.4

61.6

48.4

ca. 48

ca. 49

66.8

57.0

43.5

54.8

77.7

51.9

39.7

25.3

—

62.1

56.5

Largest larva

13.0

—

16.2

13. 0*

18.1

13.9

12.63

ca. 13.5^

Smallest nonlarva

17.2

75.8

21.2

16.6

13.2

3^.5

23.6

30.4

17.0

31.1

19.4

33.3

54.8

17.2

16.9

12.7

25.3

33.3

23.7

25.2

'See 0. lineolatua, discussion under geographical distribution.

2Based on types of Petroscirtes samoensis. See 0. r. obliquus nomenclatural discussion.

on information in Bhattacharya (1916).

Of the remaining two species, O. lineolatus is known

from only four specimens; and O. zebra is represented in our material by 20 specimens. Of the other

18 species, 11 are known from over 50 specimens

each. Generally, in any single, large collection of a

species of Omobranchus the largest specimens are

males. Larger size for males is, generally, a typical

condition for blenniids, and probably indicates territoriality (Collette and Yerger, 1962).

2. Crest: A fleshy, bladelike crest on top of the

head is found in at least some specimens of eight

species of Omobranchus (Table 5). The standard

length at which the crest first becomes apparent is

variable in these species and may be attributable as

much to state of sexual maturity as to size attained.

Inadequate data are available for most of these

species; for instance, no female smaller than 75 mm

and no male smaller than 99 mm are known for

O. aurosplendidus. All specimens of O. aurosplendidus have a crest; the vertical height of the crest

ranges from 5.8-7.7% SL in males and from 2.6-

4.8% in females, and the smallest member of each

sex has the proportionally highest crest for its sex.

In O. fasciolatus the crest is not, or scarcely, apparent in many males less than 33 mm (but always

present, although irregularly variable in size, in

larger males); it is absent, or at best poorly developed, in females. In O. banditus a crest is apparent

in males over 30 mm but never present in females.

All specimens of O. woodi (23.7-81.1 mm), of O.

fasciolatoceps (32.5-63.4 mm), and of O. mekranensis (33.3-50.7 mm) have well-developed crests. The
TABLE 5.—Frequency distributions and character states for certain characters in

Omobranchus species (P = present; A = absent)

Segnented dorsal-fin rays Total dorsal-fin els

16 17 18 19 20 21 22 23 2/. 25 26 28 29 30 31 32 33 34 35 36 37

(see

(see

(sue

(see

(see

Tabla

Table

Table

5

Table

1

Table

8)

9)

10)

12)

12

2

13)

Table 15)

> Table 8)

> Tabla 9)

i Table 10)

(see

(see

(see

(see

(see

(see

Table

Tabla

Table

Table

Table

Table

8 )

9)

10)

3

12)

13)

15)

Segmented anal-fin rays Caudal vertebra

19 20 21 12 23 24 25 26 27 10 " 12 26 27 28 29 30 31 32 36 37 3« 39 40

aaollus

2 46 105 11

> Table 8)

' Table 9)

I Table 10)

i Table 8)

i Table 9)

i Table 10)

see Table 8)

see Table 9)

see Table 10)

(see

(see

(see

2

(see

1

(see

(see

Table

Table

Table

2

Table

1

Table

Table

8 )

9)

10)

_

12)

13)

15)

Lateral-line tubes Last lateral-line tube below dorsal-fin spine

i Tabl<

i Tabli

i Tabli

8)

9)

10)

12 100 65 16 3

8 37

8 I

i Table

i Table

i Table

111 1

Species

tnollue

mroe plena i(lu»

bandltut

elagan*

• i,OTUf tu»

ftntclolatocepe

fxciolatue

ferox

tereujni

UneoUtut

lq»cnor.u»

•wkrtnensia

—nit) cue

puncu,tu»

rotund! ope

• Wlnltml

TerticAU

woodj

SSbr.

Dorsal «

9

1

(se

to

3

1

ventral

11 12

12 16

1 6

3

29 50

« Table 6)

(see Tabl

1

(ss

(se

1

3

1

a Tabl

e Tabl

2

2 12

< l )

- 24

2 9

1 1

13 3'

1

e 13)

e 15)

2

9

3 8

procu

13

9

8

37

4

16

34

43

1

9

4

Trent

• 4

5

18

12

21

99

3

15

9

2

16

5

caudal-fin rajs

15 16 17 18

12 12

1

1

62 32 1 3

7

1

2

0

35

9

42

1

(se

20

51

(se

5

11

346

(se

3

1

31

19

Lover end of gil l opening

opposite pectoral-fin ray

5

6 37 52 18 3

> Tabla 6)

2 1

3 25 25 8

> Table 10)

7 40 36 6

2

26

I Tabla 15)

2

1

1

Species

aoollna

aaroapianaldua

bandltus

elemaa

alOM»tm

faeclolaMlcspa

faaclolatta*

faun

xmnmial

llaaola&M

lwoawae

•atamanaia

•sulscva

waarta.taa

rotwBdlcaBw

eflnltaA

*articalla

•abo

10

(se.

(s«i

1

(ss<

(aa.

(aa.

11 12

14 13

i 7

2 3

• Table 6)

4 15

> Tabla 8)

17 38

• Tabla 10)

2

2

1 1

I Tabla 13)

> T*bla 15)

2

13

13

1

39

I

I

3

24

3

14

14

56

45

3

1

4

Bptplaural ribs

15

1

42

34

1

2

16

4

1

6

10

17

2

_

1

11

18

16

1

4

9

19 20 21 22 23

22 5 3

2 3 - - 1

4

Interorbital

po

t 2

2

4

4

1 2

18

55

2

2

2

10

7

res

3

35

9

50

154

243

1

59

242

5

186

2

5OA

(aaa Tabla 15)

1

4

2

33

20

4

3

2

4

8

7

16

1

1

Circuaorbital pores

6

2

(a

2

( •

7

2

7

1

142

4

3

1

8

35

48

16

241

17

49

57

ae Table 10)

1

24

5

186

10

2

603

aa Table 15]

1

3

4

2

33

16

9 10

2 1

5

5

4

2

8

19 2

1

1

Fleshy craat on

head of

•ale feamle

1

P

P

I

' P or A

ar A

I

s r l P

I

P

»

1

>r A
NUMBER 177 13

crests of males of these three species are always

relatively larger than those of comparably sized

females; a few 0. fasciolatoceps females have crests

as high as those of males of comparable size, but

the lateral area of the female crests was obviously

smaller (extreme development of the crest in an

O. fasciolatoceps male is illustrated in Tomiyama

and Abe, 1953). In 0. anolius all the males available (34-60 mm) have very well-developed crests,

whereas females (28-46 mm) either have no crest

or a very poorly developed one, usually represented

by a ridge less than 0.5 mm high. The single available male, 60 mm, of O. meniscus has a crest,

whereas the single available female, 54 mm, has

none.

3. Enlarged Head: In O. elegans, rotundiceps,

loxozonus, and possible other species, the heads of

many of the larger, or presumably mature, males

appear to be enlarged or inflated, particularly dorsally, giving the impression of a slight hump. After

long preservation, the skin on top of the head becomes flaccid, and in the case of shrinkage is drawn

into a low crestlike fold.

4. Anal-fin Tabs: These are small, lappetlike

structures of apparently noncellular composition

that form subdistally to the tips of the segmented

rays of the anal fin of presumably mature males

(see Figure 336 and Springer, 1972a, fig. 86). The

tabs are easily scraped off and are usually not found

on old or poorly preserved specimens. Tabs develop

in most species of Omobranchus and may possibly

occur even in those species for which we have no

records of tabs. The function of the tabs is not

known.

5. Flattened Tips of Dorsal- and Anal-fin Segmented Rays: The tips of the more anterior analfin rays and/or more posterior dorsal-fin rays of

presumably mature males of several species become

expanded or spatulate. The function of these structures is not known.

6. Color Pattern: The males of Omobranchus are

generally darker than females, and the bands on

the body more completely developed than on females. In O. elongatus, rotundiceps, loxozonus, germaini, and fasciolatus, most males exhibit a large,

dark spot or group of spots covering the midportions of the membranes of the central segmented

dorsal-fin rays. Females of these species lack this

marking, although a few females of O. germaini

exhibited indications of the spot. Males of O. ferox

usually have a noticeable dusky spot that females

lack on the posteriormost segmented dorsal-fin rays.

On the other hand, females of O. aurosplendidus

have an intense dark spot, which males lack, at the

anterior end of the dorsal fin. The three males

available of O. verticalis have a dark spot at the

anterior end of the dorsal fin. In the single available female of this species the spot is present but

not quite as dark as in the males.

7. Fin Spine and Ray Lengths: In most species

of Omobranchus the larger, presumably mature,

males have proportionately longer segmented dorsalfin rays (particularly the more posterior rays) than

do females. In the males the tips of the rays tend

to become free of the interradial membranes. This

condition reaches an extreme in some large males

of O. anolius, where the rays become long and

filamentous, attaining a length equal to at least

one-third the standard length. All males of O. aurosplendidus (smallest available, 99 mm) are distinguished from all females in having most of the

dorsal-fin spines prolonged into filaments, the longest equaling more than 40 percent of the standard

length. Males of O. elongatus, rotundiceps, germaini, and loxozonus typically have one or more

caudal-fin rays prolonged and filamentous, a condition rarely present in females.

8. Jaw Modifications: Females of one species of

Omobranchus, O. fasciolatoceps, with spent gonads

exhibit modifications of the jaws that are not approximated in any other species of blenniid. These

females have lost the posterior canines in both jaws.

The hingelike, interdigitating dentary processes

that connect the dentaries anteriorly are also lost

and replaced by a smooth joint at the symphysis.

Along with these modifications, the gape is enlarged, a large flap of skin from the posterior end

of each side of each jaw is extended partway across

the floor and roof of the mouth, both jaws become

broadly rounded anteriorly, the incisoriform teeth

become strongly recurved, and the number of teeth

in the upper jaw is greatly increased. The function

of these modifications is unknown, but they may

indicate mouth brooding, a habit not yet reported

for blennioids. Females of O. fasciolatoceps with

ripe, or near ripe, ovaries exhibit no modifications

of the jaws.

9. Number of Teeth: Four of the nine species of

Omobranchus (elongatus, germaini, punctatus, rotundiceps) that were statistically tested for sexual
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

I So-po- ^ ^

(.^PHILIPPINE ./»». Cm ~*» . •*. . ^MARSHALL IS.

0. ANOLIUS

0. ELE6ANS

. FASCIOLATUS

0. FEROX

O 0- MEKRANENSIS

0- MENISCUS

FIGURE 4. Distribution of certain species of Omobranchus. Dashed line represents approximate

position of Andesite Line.

0. AUROSPLENOIDUS

0- ELONGATUS

• 0. LOXOZONUS

O 0- ROTUNDICEPS OBLIQUUS

0. ROTUNDICEPS ROTUNDICEPS

FIGURE 5.—Distribution of certain species and subspecies of Omobranchus. Dashed line represents

approximate position of Andesite Line. Questionable locality record indicated (see "Geographic

Distribution" under O. r. obliquus).
NUMBER 177 15

O 0. BANDITUS

0. FASCIOLATOCEPS

# 0. GERMAINI

• 0. LINEOLATUS

C> 0. VERTICALIS

• 0. ZEBRA

FIGURE 6.—Distribution of certain species of Omobranchus. Dashed line represents approximate

position of Andesite Line. Questionable locality record indicated (see "Geographic Distribution"

under O. lineolatus).

0. PUNCTATUS

STEINITZI

A 0. W00DI

FIGURE 7.—Distribution of certain species of Omobranchus. Dashed line represents approximate

position of Andesite Line. Questionable locality record and extralimital distribution indicated

for O. punctatus (see "Geographic Distribution" under O. punctatus).
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

dimorphism in number of incisoriform teeth exhibited such dimorphism, although the dimorphism

was not present in all populations of a species, nor

did it necessarily involve the teeth in both jaws.

The results of the tests are given in Table 3. A

fifth species, O. fasciolatoceps, which was not tested,

also snowed differences between males and females

in number of lower jaw teeth (Figure 41), but not

in number of upper jaw teeth (Figure 40). In

species where there was sexual dimorphism, females

had more teeth than males. Springer (1971) found

significant sexual differences in number of lower

jaw teeth in three of 18 species of the blenniid

genus Ecsenius (upper jaw teeth were not examined

for dimorphism). In all three species females had

more teeth than males (in contrast to Omobranchus, tooth numbers in Ecsenius are not related to

SL). We are unable to explain why females should

have more teeth, or why this type of sexual dimorphism is not exhibited consistently in all populations within a species.

ZOOGEOGRAPHY.—The distribution of Omobranchus (Figures 4-7) extends from the northern Red

Sea and southern Africa in the western Indian

Ocean, to the Samoan, Line, and Hawaiian islands

in the central Pacific Ocean (the introduced populations in the Caribbean Sea are ignored here;

see geographic distribution discussion under O.

punctatus).

It is of interest to compare the distributions of

Omobranchus species with those of another blenniid genus, Ecsenius, which has about the same

number of species (Springer 1971, 19726) as Omobranchus but appears to have different ecological

requirements and behavior. (Ecsenius is not as secretive and often occurs in deeper offshore waters

than does Omobranchus. The blenniid/genus Entomacrodus is perhaps more similar to Omobranchus

in ecological requirements and behavior than is

Ecsenius, but there are few parallels in the distributions of Omobranchus and Entomacrodus species;

see Springer, 1967, for Entomacrodus) In general,

Omobranchus occurs farther north and south than

does Ecsenius.

Five species of Omobranchus occur only in the

western Indian Ocean and Red Sea. Of these, two

(woodi, bandittis) occur only in southern Africa,

where Ecsenius has not been reported, and one

(steinitzi) is known only from the Red Sea, where

there are three endemic species of Ecsenius. Omobranchus mekranensis occurs only in the Persian

Gulf and Gulf of Oman, which distribution is similar to that of Ecsenius pulcher, and O. fasciolatus,

which is widespread in the western Indian Ocean,

closely parallels the distribution of Ecsenius nalolo.

There are two endemic species and one endemic

subspecies of Omobranchus (verticalis, anolius, r.

rotundiceps) in Australia, as well as two or three

more broadly distributed species (punctatus, germaini, lineolatusf). These compare well with Ecsenius, which has one endemic species, one endemic

population of another species, and two more broadly

distributed species in Australia; however, Ecsenius

has not been reported from Western Australia

where Omobranchus is common. With regard to

the remaining species of Omobranchus and Ecsenius, there does not appear to be much similarity

in individual species distributions.

With the exception of one subspecies, O. r. obliquus, Omobranchus is limited in distribution to

the area west of the Andesite Line (MacDonald,

1949; modified by later authors). Of the other ten

species in the tribe Omobranchini (Springer, 1972a),

seven occur only west of the Andesite Line. Of the

three species that occur east of the line, one, Parenchelyurus hepburni, barely transcends the Line eastward; one, Enchelyurus ater, barely transcends the

Line westward, and one, Enchelyurus brunneolus,

is found only well east (Hawaii) of the Line.

The Andesite Line marks the separation of the

continental from the oceanic rocks. The oceanic

rocks are of more recent age than the continental

rocks, and apparently there has been insufficient

time for Omobranchus to have spread fully into

Oceania. Although the continental areas may offer

a greater variety of niches that are suitable for

Omobranchus (and other fish genera), and thus

have enhanced speciation of the genus in these

areas, it seems improbable that the oceanic areas

provide niches enough for only one species of Omobranchus. The evidence, therefore, favors a continental area as the area of origin of Omobranchus,

and also indicates that the oceanic fish fauna generally is probably derivative of the continental

fauna. Ladd (1960), however, has presented evidence favoring the dispersal of marine faunas (primarily molluscan) westward from Oceania to the

andesite areas.
NUMBER 177 17

Key to the Species and Subspecies of Omobranchus

1. Total dorsal-fin ray elements 28; segmented anal-fin rays 18; prenasal pores absent

steinitzi, new species (Red Sea)

Total dorsal-fin ray elements more than 28 (more than 29 except in anolius from Australia);

segmented anal-fin rays more than 18 (more than 19 except in anolius, which rarely has

19); prenasal pores present 2

2. Segmented dorsal-fin rays 25-26; total dorsal-fin elements 36-37; bladelike crest present on

top of head; mature males with anterior dorsal-fin spines filamentous (Figure 9)

aurosplendidus (South China Sea)

Segmented dorsal-fin rays 17-24 (rarely 24); total dorsal-fin elements 29-36, but if more than

34 no bladelike crest on top of head; mature males without filamentous anterior

dorsal-fin spines 3

3. Body covered with dense, randomly distributed sprinkling of fine black spots (Figure 12);

circumorbital pores 6-8 (strong mode of 7); lateral-line tubes 0-1

elegans (Japan, China, Korea)

Body not covered with fine sprinkling of numerous black spots (anolius, from Australia,

with up to 5 more or less longitudinal rows of dark spots); circumorbital pores 7-10

(strong mode of 8 or 9 pores; 7 pores common only in Guam population of rotundiceps);

lateral-line tubes 0-9 4

4. Lower-lip flap absent (Figure 2a); fleshy bladelike crest on top of head present on males

over 30 mm SL (absent on females); gill opening restricted to area above level of dorsalmost pectoral-fin ray; lateral-line tubes 5-9; posteriormost tube reaching to below level

of 8th-llth dorsal-fin spine banditus (southeastern Africa)

Lower-lip flap present (Figure 2b); fleshy bladelike crest on top of head present or absent

on males over 30 mm (present or absent on females); gill opening varying from restricted

to area above level of dorsalmost pectoral-fin ray to extending ventrally to opposite 6th

ray; lateral-line tubes 0-9; posteriormost tube, when tubes present, reaching to below

level of 1st—12th dorsal-fin spine 5

5. Conspicuous, slender, black, crescentic mark extending dorsally from dorsoposterior orbital

margin (Figure 25); no lateral-line tubes; gill opening extending ventrally to opposite 3rd

pectoral-fin ray; slender fleshy crest present on top of head of males only

meniscus, new species (Gulf of Thailand)

No black, crescentic mark on head; lateral-line tubes present or absent; gill opening

varying from restricted to area above level of dorsalmost pectoral-fin ray to extending

ventrally to opposite 6th ray; slender fleshy crest present or absent on top of head

of males and females 6

6. Two to five more or less longitudinal rows of small black spots on body, overlying other

features of color pattern; well-developed bladelike fleshy crest on top of head of males,

poorly developed or absent on females; no lateral-line tubes; posterior segmented dorsal-fin

rays of mature males becoming long and filamentous; gill opening restricted to area dorsal

to dorsalmost pectoral-fin ray anolius (eastern Australia)

No rows of black spots on body overlying color pattern; fleshy crest present or absent on

top of head of males and females; lateral-line tubes present or absent; posterior segmented

dorsal-fin rays never filamentous; gill opening varying from restricted to area dorsal to

dorsalmost pectoral-fin ray to extending ventrally to opposite 6th ray 7

7. Fleshy bladelike* crest present on top of head (possibly absent in specimens under 33 mm

SL); gill opening usually restricted to area dorsal to level of dorsalmost pectoral-fin ray

(occasionally reaching opposite 1st ray) 8

No fleshy bladelike* crest on top of head; gill opening varying from restricted to area dorsal

to level of dorsalmost pectoral-fin ray to extending ventrally to opposite 6th ray 11

• In large or mature males of O. rotundiceps and O. loxozonus the head may be fleshy and

dorsally swollen. In preservative the head often shrinks, causing the skin in the dorsal region

of the head to form a loose, fleshy fold that might be confused with a crest. Males with swollen

heads usually also have the dorsal and/or ventral lobes of the caudal fin produced as filaments,

a condition not present in any of the species that fall into key couplets 8-10.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

8. Interorbital pores 3; precaudal vertebrae 11-12; epipleural ribs 14-19 (rarely 14)

tvoodi (southeastern Africa)

Interorbital pores 2; precaudal vertebrae 10-11; epipleural ribs 10-23 9

9. Segmented dorsal-fin rays 18-19; segmented anal-fin rays 20-22; precaudal vertebrae 10;

males frequently with noticeable dark or dusky spot at midlevel of central segmented

dorsal-fin rays; epipleural ribs 10-13

fasciolatus (western half of Indian Ocean, including Red Sea)

Segmented dorsal-fin rays 19-22 (rarely 19); segmented anal-fin rays 22-24; precaudal

vertebrae 10-11; males without noticeable dark or dusky spot on segmented ray portion

of dorsal fin; epipleural ribs 11-23 10

10. Height of crest on top of head 3.1-8.2% SL; mature females without posterior canines in

either jaw and with more than 30 incisoriform teeth in lower jaw; both anal-fin spines of

males discernible externally; precaudal vertebrae 10; epipleural ribs 11-13

fasciolatoceps (Japan, China, South China Sea)

Height of crest on top of head 1.1-4.5% SL; mature females with posterior canines in both

jaws and fewer than 30 incisoriform teeth in lower jaw; 1st, and frequently 2nd, anal-fin

spines of males reduced, not discernible externally; precaudal vertebrae 11; epipleural

ribs 17-23 mekranensis (Persian Gulf; West Pakistan)

11. Interorbital pores 2 12

Interorbital pores 2-4 (2 in fewer than 0.5% of specimens) 13

12. Gill opening restricted to area above level of dorsalmost pectoral-fin ray; lateral-line tubes

3-8 (rarely 3 or 4), extending posteriorly to below level of 7-1 Oth dorsal-fin spine

fasciolatus (western half of Indian Ocean including Red Sea)

Gill opening extending ventrally to level of 3rd-5th pectoral-fin ray; lateral-line tubes 0-4,

extending posteriorly no farther than to below level of 6th dorsal-fin spine

rotundiceps obliquus (Pacific Islands)

13. Head more than 24.0% SL in specimens 25-48 mm SL, and more than 23.5% SL in

specimens 53-56 mm SL; gill opening restricted to area dorsal to level of dorsalmost

pectoral-fin ray (reaching to opposite first ray in only one 25 mm specimen); caudal-fin

rays never filamentous 14

Head less than 24.0% SL in specimens 25-48 mm SL, and less than 23.5% SL in specimens

over 42 mm SL; gill opening varying from restricted to area dorsal to level of dorsalmost

pectoral-fin ray to extending ventrally to opposite 6th ray; caudal-fin rays filamentous in

males of some species (Figures 13, 15, 19, 23, 28) 15

14. Dorsal-fin spines 12; lateral-line tubes 1-4; pale interspaces separating broad, dusky bands

on side of head without small, dark spots; dark spot on side of head posterior to eye an

intensification of dark head band (Figures 33a, 34) zebra (India, South China Sea)

Dorsal-fin spines 11; lateral-line tubes 6-8; pale interspaces separating broad, dusky bands

on head with small, dark spots; large, dark, oval spot with pale margin on side of head

posterior to eye distinct from dark head bands

lineolatus (Australia, New Guinea, Candavu)

15. Broad, dusky to dark band, with pale margin, extending dorsally from one side of body to

other in region just anterior to dorsal fin (band occasionally broken into 3 blotches:

1 on each side and 1 medial); several slender, dusky or dark stripes on side of body in

region covered by appressed pectoral fin; gill opening restricted to area dorsal to level

of dorsalmost pectoral-fin ray (occasionally extending ventrally to opposite dorsalmost

ray); caudal-fin rays never filamentous; 1st, and frequently 2nd, anal-fin spine of males

reduced, not discernible externally; epipleural ribs 15-25 (rarely 15-17)

punctatus (Indian Ocean, Pacific Ocean, Caribbean Sea)

No dark band on body just anterior to dorsal fin; no stripes on body in region covered by

appressed pectoral fin; gill opening varying from restricted to area above level of

dorsalmost pectoral-fin ray to extending ventrally to opposite 6th ray; caudal-fin rays of

males filamentous or not; both anal-fin spines of males discernible externally; epipleural

ribs 10-16 (rarely 16) 16

16. Prominent black or dusky spot covering most of distal portion of membrane between first

2 dorsal-fin spines; prominent dark spot posterior to eye; up to 16 vertical, dusky bands

on body, several partially outlined with slender dark lines; gill opening restricted to

area dorsal to level of 2nd pectoral-fin ray; fewer than 20 incisor teeth in either jaw in
NUMBER 177 19

specimens 30-49 mm SL verticalis, new species (Queensland)

No prominent spot on distal portion of membrane between first 2 dorsal-fin spines; prominent dark spot on head posterior to eye present or absent; dark bands present or absent

on body (bands, when present, usually strongly inclined anteriorly or posteriorly except

occasionally in germaini); bands not outlined with darker lines; gill opening varying from

restricted to area dorsal to level of dorsalmost pectoral-fin ray to extending ventrally to

opposite 6th ray; most species with 20 or more teeth in each jaw in specimens over 30 mm

SL (number increasing with increase in SL) 17

17. Dorsal-fin spines 11 and gill opening restricted to area dorsal to level of dorsalmost

pectoral-fin ray 18

Dorsal-fin spines 11-14 (usually 12-13), but if 11, gill opening extends ventrally at least to

opposite 2nd from dorsalmost pectoral-fin ray 19

18. Lateral-line tubes 6-8, extending posteriorly to below level of 6th-llth dorsal-fin spine;

caudal-fin rays not filamentous lineolatus (Australia, New Guinea, Candavu)

Lateral-line tubes 0-5 (rarely more than 3), extending posteriorly (when present) to below

level of lst-6th dorsal-fin spine (rarely posterior to level of 4th dorsal-fin spine); some

caudal-fin rays of some males filamentous rotundiceps rotundiceps (Australia)

19. 15th segmented dorsal-fin ray 14.7-22.4% SL in males (rarely less than 18.0%), 12.1-16.0%

SL in females (rarely less than 13.5%); gill opening extending ventrally to opposite

3rd-6th pectoral-fin ray (rarely only to 3rd ray); dorsal-fin spines 11-13 (strong mode of

12); lateral-line tubes 1-4; males frequently with dusky spot over last 1-4 segmented

dorsal-fin rays as most prominent mark in dorsal fin (Figure 18); slender, dusky to black

band adjacent to postorbital margin (but separated from margin by slender, pale area);

no dusky pinstripes in segmented ray portion of dorsal fin; circumorbital bones 4

ferox (eastern Africa to Philippine Islands)

15th segmented dorsal-fin ray 9.1-16.0% SL in males (rarely more than 14.0%), 8.4-13.6%

in females (rarely more than 12.9%); gill opening varying from restricted to area above

level of dorsalmost pectoral-fin ray to extending ventrally to opposite 6th ray; dorsal-fin

spines 12-14 (modally 12 or 13); lateral-line tubes 0-9; males frequently with dark spot,

or short series of dark spots, commencing about midway along segmented ray portion of

dorsal fin as most prominent mark in fin (Figures 13, 15, 19, 23, 28); slender, dusky to

black band adjacent to postorbital margin present or absent; dusky pinstripes present

or absent in segmented ray portion of dorsal fin; circumorbital bones 5 20

20. Darkest marking on side of head a prominent spot, which is noticeably closer to preopercular

series of sensory pores than to circumorbital series of pores (spot occupies dorsoposterior

corner of dusky domeshaped area; Figures 28, 29a); dorsal-fin spines 11-13 (rarely 13)

rotundiceps rotundiceps (Australia)

Darkest marking on head a faint to dark band, spot, or blotch, which is either noticeably

closer to circumorbital series of sensory pores than to peropercular series of pores or

only slightly closer to preopercular series (Figures 2ie, 29c and e); dorsal-fine spines

11-14 (rarely 11) 21

21. Dorsal-fin spines 12-13 (rarely 13); underside of head varying from plain dusky to bearing

conspicuous dark chevrons, never with scattered dusky spots or mosaic of chevrons and

vermiculations; prominent dark spot on head posterior to eye approximately midway

between circumorbital and preopercular series of pores or slightly closer to preopercular

series (Figure 29c) rotundiceps obliquus (Pacific Islands)

Dorsal-fin spines 12-14 (strongly modally 13); underside of head variable: plain dusky, with

diffuse dusky chevrons, with scattered dusky to dark spots, or with a mosaic of dark

chevrons and vermiculations; prominent dark spot on head posterior to eye present or

absent, when present closer to circumorbital series of pores than to preopercular series .2 2

22. Segmented dorsal-fin rays 17-20 (modally 18-19); underside of head uniformly dusky or

with scattered dusky to dark spots, which may extend onto prepelvic and prepectoral

areas; side of head with or without prominent dark markings (markings, when present,

variable in form but usually not present as distinct dark spot posterior to eye; occasionally

present as dark band or smudge)

elongatus (eastern Africa to Philippine Islands, excluding Australia)

Segmented dorsal-fin rays 18-23 (modally 19-22); underside of head dusky, bearing dusky
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

chevrons or mosaic of dusky to dark chevrons and vermiculations; side of head with

distinct dark spot posterior to eye as most prominent marking 23

23. Lateral-line tubes 0-8 (usually more than 3); circumorbital pores 7-10 (modally 8-9);

usually 6-8 pairs of more or less vertical, narrow, dark bands or spots on body; underside

of head frequently marked with dark mosaic of chevrons and vermiculations; precaudal

vertebrae 10, rarely 11, in all populations except those from Western Australia, which

may have modally 11 precaudal vertebrae

germaini (Australia north to Taiwan; New Caledonia)

Lateral-line tubes 0-4 (usually less than 3); circumorbital pores 7-9 (rarely 7 or 9); usually

11 or fewer broad, dusky, inclined bands on body; underside of head varying from

uniformly dusky to bearing faint, dusky chevrons; precaudal vertebrae 10-12 (rarely 10) ....

loxozonus (Japan to Okinawa)

Omobranchus anolius (Valenciennes)

FIGURE 8

Blennechis anolius Valenciennes in Cuvier and Valenciennes,

1836:288 [Port Jackson].

Petroscirtes altivelis Steindachner, 1863:1191 [Bombay; locality

probably erroneous].

Blennius unicornis Castelnau, 1879:384 [enters oysters about

Sydney].

Petroscirtes guttatus Macleay, 1881:9 [Port Jackson].

Petroscirtes cristiceps Macleay, 1881:9 [Port Jackson].

Petroscirtes wilsoni Macleay, 1884:171 [Port Jackson].

Salarius galeatus DeVis, 1884a: 147 [Moreton Bay]; 1886; 59

[St. Helena, Moreton Bay; twice described as new; impossible to determine if the same or different specimens

formed basis of descriptions].

DESCRIPTION (see also Table 5).—Dorsal fin XIXIII (XII in 79.2% of specimens), 17-19 = 29-31;

anal fin II, 19-22 (20-21 in 94.3% of specimens);

both anal-fin spines of males discernible externally;

segmented caudal-fin rays 12-13 (13 in 94.3% of

specimens); dorsal + ventral procurrent caudal-fin

rays 11-14; vertebrae 10-11 (11 in only 1 of 42 specimens) + 26-28 (28 in only 1 of 41 specimens) =

36-38 (38 in only 2 of 41 specimens); epipleural

ribs 11-13; prenasal pores present; interorbital

pores 2-4 (3 in 87.5% of specimens); circumorbital

pores 7-10 (8 in 87.5% of specimens); lateral-line

tubes absent; gill opening varying from restricted

to area dorsal to level of dorsalmost pectoral-fin ray

(87.5% of specimens) to extending ventrally to

opposite dorsalmost ray; lower-lip flap present; circumorbital bones 5; lower jaw teeth 20-27; upper

jaw teeth 19-25 (Figure 36); males with fleshy

bladelike crest on top of head (at least in specimens 34-60 mm SL), females with crest poorly developed or absent (at least in specimens 28-46 mm

SL).

COLOR PATTERN.—Males: Head: Three dusky to

dark dusky bands present. First band moderately

wide, extending from anteroventral margin of orbit

across mouth, just anterior to angle of jaws, to chin

where it joins corresponding band of opposite side.

Second band almost twice as broad as first, extending from ventral margin of orbit to ventral midline

of head where it joins corresponding band of opposite side; band, interrupted by eye, continues from

dorsoposterior margin of orbit across anterior nape

to base of fleshy crest. Narrow, pale or pearly lines

border anterior and posterior margins of both portions of second band. Third head band encircles

head posterior to eye; band often obscured by dark

background color of head. Narrow, pale or pearly

line borders posterior margin of third band below

level of ventral margin of orbit. Intensely dark,

short line, blotch, or spot present on third band

just above lateral midline of head. Narrow, pale or

pearly line extends posteroventrally from preopercular head pores near dorsal end of operculum to

anterior edge of fleshy pectoral-fin base. Another

narrow, pale or pearly line occasionally extending

from middle of nape and base of crest, along dorsoposterior edge of operculum and posteroventral

edge of gill opening to upper end of fleshy

pectoral-fin base; line broadens as it passes along

edge of gill opening. Snout, interorbital, nape, and

prepelvic areas dark dusky except where crossed by

pale lines.

Trunk: Body crossed with about 17 to 19 evenly

spaced, narrow, dusky, chevronlike bands flexed

posteriorly; anterior and posterior bands indistinct;

bands originating on dorsal body contour, becoming

faint, then absent on ventral fifth of body; posterior

3 or 4 bands broken, forming 3 dusky stripes: 1 on

lateral midline, 1 each midway between lateral

midline and dorsal and ventral body contours;

dorsal ends of bands occasionally expanded and
NUMBER J77 21

intensified. Horizontal series of small, intensely

dark spots present on lateral midline of body, each

spot lying on body band at angle of flexion (not all

bands with spot). Second horizontal series of small,

dark spots present midway between lateral midline

of body and dorsal body contour, each spot located

on body band. Third horizontal series of small,

dark spots usually present midway between lateral

midline and ventral body contour, at least posteriorly; each spot located on body band. Area covered by appressed pectoral fin with posteroventrally

directed, diagonal row of intensely dark marks or

spots, more or less connected by dark dusky line on

body midlaterally; second parallel line of dark

marks occasionally present just posteriorly on dorsal

half of body.

Pectoral Fin: Pale to transparent with fine dark

specks of pigment scattered over fin; pigment heavy

proximally and distally. Fleshy pectoral-fin base

with large, intensely dark spot, occasionally broken

into several intensely dark marks anteroventrally.

Vertical, narrow, pale line present between large

dark spot and base of pectoral fin; line confluent

dorsally with narrow, pale line on head.

Pelvic Fin: Distal half pale, proximal half dark

dusky, line of demarcation between contrastingly

pigmented areas angled so inner rays darkly pigmented more distally than outer rays.

Dorsal Fin: Pale dusky with dark markings. Membranes between first 5 dorsal-fin spines with 2 or 3

evenly spaced, narrow, horizontal, dark lines; lines

becoming dorsoposteriorly angled on more posterior

spines; parallel series of similar dorsoposteriorly

oriented, narrow, dark lines along rest of fin; lines

becoming faint to absent all but proximally on

posterior half of fin (particularly in large specimens).

Anal Fin: Dark dusky proximally, tips of rays

abruptly pale. Several, often interrupted, posteroventrally directed, narrow, pale lines on proximal

two-thirds of fin; lines faint posteriorly.

Caudal Fin: Proximal end of fin with pale membranes; rays outlined with dark dusky pigment,

occasionally extending posteriorly at midlevel of

fin; some specimens with vertical, narrow, dark

dusky margin on fin at posterior end of fleshy caudal base.

Females: Similar to males except: head background usually not as dark; dark bands and markings on head more apparent; pale spaces between

dark bands slightly broader; body background

paler; body bands less distinct; usually 4th and

occasionally 5th horizontal series of intense, dark

spots present on body; spots in somewhat less distinctly arranged rows; fins less distinctly pigmented;

dorsal fin often only with dark dusky blotches

along base above dorsal ends of body bands.

McCulloch (1917, pi. 10: fig. 2) presented a useful

illustration of O. anolius, which is reproduced in

Scott (1962:247).

GEOGRAPHIC DISTRIBUTION (Figure 4).—Spencer

Gulf, South Australia, to the Queensland coast of

the Gulf of Carpentaria. Scott (1962) reported that

O. anolius occurs in Victoria, but we have seen no

specimens from that state. See "Nomenclatural Discussion" for erroneous locality records.

HABITAT.—Shallow, usually estuarine waters in

and around oyster and other mollusc beds on tidal

mud flats (Thomson and Bennett, 1953).

COMPARISONS.—Omobranchus anolius is unique

in Omobranchus in having the posterior segmented

dorsal-fin rays of larger males long and filamentous.

The 15th ray, for instance, may attain a length of

up to 31.9% SL, more than in any other species of

Omobranchus. In O. ferox and aurosplendidus the

15th ray may attain a length of 22-24% SL, but

none of the rays become filamentous or much exserted beyond the margin of the interradial membrane (the dorsal-fin spines of male aurosplendidus

do, however, become long and filamentous). In O.

anolius there are several series of small, dark spots

that appear to be superimposed on the color pattern of the body. Omobranchus elegans, aurosplendidus and, to a much lesser extent, meniscus also

have a similar spotting, but the spots are smaller,

more numerous, and more randomly distributed in

these three species. Some specimens of O. fasciolatus

appear to have a similar spotting, but the spots are

not so intensely dark and do not appear to overlie

other features of the color pattern.

Omobranchus anolius can be differentiated externally from the Australian populations of those

species of Omobranchus with which it appears to be

broadly sympatric (O. lineolatus, germaini, punctatus, r. rotundiceps, verticalis) most readily by its

lack of lateral-line tubes. The majority of the specimens of O. r. rotundiceps from New South Wales

also lack lateral-line tubes, but these specimens
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

have 33-35 total dorsal-fin elements (versus 29-31

for anolius). O. anolius is also the only Australian

species that may have a fleshy crest on the head.

REMARKS.—Thomson and Bennett (1953) described the nests, development of eggs, and behavior

at hatching of adults and larvae, and Ogilby (1910)

reported on the nests, habitat, and reproductive

behavior.

NOMENCLATURAL DISCUSSION.—None of the authors of the synonyms of O. anolius distinguished

their new species from any other species.

The type-material of P. altivelis comprises three

syntypes, all identifiable as male anolius. Steindachner (1863) gave the type-locality as Bombay, but

otherwise the species has been recorded only from

Australia, except for a specimen cataloged into the

British Museum collections in 1864 and reported

as coming from New Zealand (the only record of

an omobranchine from that country). The fact that

no specimens of this common, oyster-shell inhabiting species have been taken outside Australian

waters in over 100 years seems to indicate that the

species is probably limited to Australia and that

the Bombay and New Zealand records are erroneous.

No type-material is known for B. unicornis. The

description is brief, but the mention of the lack of

head cirri, the presence of an "arched fleshy horn

directed upwards" on the head, and a series of

black spots on the body identifies B. unicornis with

anolius, the only Australian blenniid with this

combination of characters.

The syntypic material of P. guttatus comprises

two female anolius, without obvious fleshy crests

on the head. The only known type-material of

P. cristiceps, described in the same paper with P.

guttatus, comprises four male anolius (listed as cotypes) with fleshy, conspicuous crests on their heads.

This sexual difference probably accounts for Macleay's (1881) description of the species twice in the

same paper.

The holotype of P. wilsoni is a male anolius.

Macleay (1884) noted the similarity of his wilsoni

with his cristiceps, but mentioned no differences.

DeVis (1884a, 1886) twice described Salarias

galeatus. The two descriptions could have been

taken from the same specimen, but this is not certain. Both depict a male anolius, with the typelocalities as Moreton Bay and St. Helena, Morton

Bay. The only possible type-material known for

either species is a male specimen registered as a

"cotype" in the Australian Museum. Although this

specimen is identifiable as anolius, its dorsal-fin

count fits neither of DeVis' descriptions of S.

galeatus.

MATERIAL EXAMINED.—AUSTRALIA: NEW SOUTH

WALES: Sydney, USNM 197621 (26: 17.2-46.0, including 2 cleared and stained), MNHN A.2160 (ca.

48), BMNH 1862.8.1.16 (52.8); Port Jackson, MNHN

A. 1832 (ca. 46, holotype of Blennechis anolius),

AMS 1.16409-001 (2: ca. 44-45, syntypes of Petroscirtes guttatus), I 16410-001 (ca. 49, holotype of

Petroscirtes wilsoni), 1.16411-001 (4: 31.5-52.8,

"cotypes" of Petroscirtes cristiceps); Shoalhaven

River, BMNH 1929.12.3.31.2 (60.6); Clarence River,

BMNH 1914.8.20.238 (53.4). SOUTH AUSTRALIA:

Backy Point (Spencer Gulf), AMS IB.5142 (ca. 47).

QUEENSLAND: Moreton Bay, AMS IA.4596 (41.8),

1.470 (47.0, "cotype" of Salarias galeatus), 1.383 (ca.

28, originally among syntypes of Salarias furtivus =

O. r. rotundiceps); Manly, QM 1.7828 (52.2); Caloundra, AMS 1.6206 (2: 30.5-33.5); Norman River,

QM 1.2200 (ca. 47), 1.2210 (45.2), 1.2211 (47.6),

1.2213 (40.7). INDIA: Bombay (probably erroneous), NMV 71774 (3: 43.9-50.3, syntypes of Petroscirtes altivelis). NEW ZEALAND (probably

erroneous): BMNH 1864.1.17.4 (56.0).

Omobranchus aurosplendidus (Richardson)

FIGURES 9 and 10

Blennius? aurosplendidus Richardson, 1846:265 [Macao].

Petroscirtes lini Herre, 1934:292 [oyster beds, Heungchow,

Kwangtung Province, China].

DESCRIPTION (see also Table 5).—Dorsal fin XIXII (XI in 8 of 10 specimens), 25-26 = 36-37; anal

fin II, 25-27; both anal-fin spines of males discernible externally; segmented caudal-fin rays 13-15

(14 in 3 and 15 in 1 of 10 specimens); vertebrae

10-11 (11 in 1 of 10 specimens) + 31-32 = 41-42;

epipleural ribs 11-13; prenasal pores present; interorbital pores 3; circumorbital pores 7-8 (7 in 7 of 9

specimens); lateral-line tubes 0-2; lateral-line tubes,

when present, extending posteriorly to below level

of 2nd or 3rd dorsal-fin spine; gill opening restricted

to area dorsal to level of dorsalmost pectoral-fin

ray; lower-lip flap present; circumorbital bones 5;

lower jaw teeth 22-28; upper jaw teeth 21-26 (Figure 47); fleshy bladelike crest on top of head of

both sexes.
NUMBER 177 23

COLOR PATTERN.—Males: Head: Broad, dusky to

dark dusky band extending from anteroventral

margin of orbit across corner of mouth to chin,

where it joins corresponding band of opposite side;

band interrupted by eye, continuing on dorsoposterior margin or orbit as rounded dark blotch; background very pale along anterior and posterior

margins of band; pale area broadest posterior to

dorsal segment of band. Fleshy head crest pale with

dark dusky vermiculations. Snout dusky; lower lip

pale. One specimen with broad, pale area extending from just below 1st dorsal-fin spine to upper

end of operculum, then posteroventrally to gill slit,

and ventrally along anterior fleshy pectoral-fin base

to pelvic-fin base. Single (occasionally as many as 4)

small, intensely dark spot posterior to eye midway

between circumorbital and preopercular sensory

pores. Remainder of head indistinctly marked with

dusky to dark dusky pigment.

Trunk: Background dusky, palest ventrally, darkest dorsoanteriorly, with fairly even spread of small,

intensely dark spots and moderately small, pale

spots and blotches.

Pectoral Fin: Pale.

Pelvic Fin: Pale with even spread of fine, dark

specks of pigment.

Dorsal Fin: Dusky to dark dusky anteriorly and

anteroventrally, becoming pale to transparent dorsoposteriorly. Anterior portion of fin with moderately narrow, pale stripe extending from 1st to

about 9th spine midway up fin; several dorsoposteriorly angled, narrow, pale lines present distally;

posteriorly, dark dusky background differentiates

into numerous dorsoposteriorly directed, narrow

lines with pale to transparent interspaces; interspaces broader than adjacent dark dusty lines. Several small, pale spots present on midbase of fin.

Anal Fin: Dusky with 5 or 6 narrow, pale stripes

running length of fin; fin darker anteriorly.

Caudal Fin: Pale dusky, slightly paler proximally.

Females: Similar to males except: background

color of head and body generally paler; dark band

on head slightly narrower, more distinct; segment

of band above eye extending unbroken, or as a

series of irregular dark markings, nearly to dorsal

edge of crest; other markings on crest larger, darker,

often forming an almost complete dark border to

free margin of crest; markings on dorsal and anal

fins much fainter except for large, dark, oblong spot

near outer edge of fin between 1st and 4th or 5th

dorsal-fin spines (spot absent in males).

GEOGRAPHIC DISTRIBUTION (Figure 5).—Hong

Kong, Macao, and vicinity.

HABITAT.—Shallow, brackish water in oyster beds.

COMPARISONS.—Omobranchus aurosplendidus is

one of the most distinctive species of Omobranchus;

however, only large adult specimens are available,

and it is questionable whether all the characters

that set aurosplendidus off from its congeners will

be true of smaller specimens. The long filamentous

dorsal-fin spines of male aurosplendidus are unique

in Omobranchus, and the large, black spot on the

anterior dorsal-fin spines of females may also be

unique, although the single (30 mm SL) female

O. verticalis available has a prominent spot between

the first two dorsal-fin spines. Omobranchus aurosplendidus has more segmented dorsal-fin rays than

any other Omobranchus species, but there is overlap

with other species in total number of dorsal-fin

elements. Although several other species approach

or overlap aurosplendidus in having a large number of segmented anal-fin rays and vertebrae, such

species lack the fleshy crest on top of the head that

is present in both sexes of aurosplendidus. Only

aurosplendidus and lineolatus have modally 11

dorsal-fin spines, and only aurosplendidus and elegans have modally 7 circumorbital pores. Of the

ten specimens of aurosplendidus available, seven

have 13, two have 14, and one 15, segmented caudalfin rays. No other species of Omobranchus has such

a large percentage of its specimens with more than

13 segmented caudal-fin rays (0.0-7.9% in the other

taxa), but perhaps this is due to the small sample

of aurosplendidus. The large numbers of segmented

caudal-fin rays in aurosplendidus are associated

with relatively small total numbers of procurrent

caudal-fin rays, indicating the probability that some

of the procurrent rays became segmented.

NOMENCLATURAL DISCUSSION.—Richardson (1846)

described B. aurosplendidus from an illustration,

now apparently lost (Whitehead, 1969). His mention that the species had a crest on top of the head,

no cirri (this fact mentioned under the description

of the next species, B. fasciolatoceps, he described

in the same study), the first seven dorsal-fin rays

(spines) elongated and numerous black dots on the

body could only refer to a male of the species here

recognized under his name. The type-locality,
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Macao, is close to that (vicinity of Hong Kong)

from which all other specimens of the species have

come.

Herre (1934) did not compare P. lini with any

other species, but his specimens include males that

agree closely with Richardson's description. Herre

did not segregate his holotype from his paratypes,

but merely mentioned that the holotype was a male,

111 mm, with the dorsal-fin elements 37 and the

anal-fin elements 27. The type-material comprises

four males (99.5, 100.8, 110.1, 112.2 mm SL) and

five females (75.8-97.7 mm SL). Herre's measurements must have been SL as the length of the

caudal fin (20.3) of the smallest male added to its

SL would make the total length over 119 mm. The

two largest males are thus candidates for holotype.

Of these, the 110-mm specimen has 37 dorsal-fin

elements (anal fin II, 29), and the largest specimen

has 36 dorsal-fin elements (anal fin II, 26). We

therefore consider the 110-mm specimen to be the

holotype.

MATERIAL EXAMINED.—CHINA: KWANGTUNG PROVINCE: Heungchow, CAS SU29088 (9: 75.8-112.2, including holotype of Petroscirtes lini and 1 cleared

and stained). HONG KONG: USNM 201464 (93.6).

Omobranchus bandittis J. L. B. Smith

FIGURES 2a and 11

Omobranchus bandkus J. L. B. Smith, 1959:232 [Umgazi].

DESCRIPTION (see also Table 5).—Dorsal fin XIXIII (XII in 90.7% of specimens), 19-21 = 31-33;

anal fin II, 21-23; one or both anal-fin spines of

males not discernible externally; segmented caudalfin rays 13 or 15 (15 in only 1 of 55 specimens);

dorsal + ventral procurrent caudal-fin rays 12-16;

vertebrae 10-11 + 27-30 (rarely 27) = 38-40; epipleural ribs 15-21 (rarely 15); prenasal pores present; interorbital pores 2-3 (3 in 92.5% of specimens);

circumorbital pores 7-9 (8 in 88.9% of specimens);

lateral-line tubes 5-9 (rarely 9); lateral-line tubes

extending posteriorly to below level of dorsal-fin

spine 8-11; gill opening restricted to area dorsal to

level of dorsalmost pectoral-fin ray; lower-lip flap

absent (Figure 2a); circumorbital bones 5; lower

jaw teeth 19-27; upper jaw teeth 17-24 (Figure 37);

males over 30 mm SL with bladelike fleshy crest on

top of head; crest absent in males under 30 mm SL

and in all females.

COLOR PATTERN.—Males: Head: Three variable,

dark bands radiating from eye on lower two-thirds

of head; anterior band moderately broad at origin

on anteroventral edge of orbit, flaring broadly as it

crosses mouth just anterior to angle of jaws and

continuing onto chin; 2nd band narrow to moderately broad, extending from midventral margin of

orbit to ventral midline of head, sometimes deflected slightly posteriorly on ventral side of head;

3rd band broad, angled posteroventrally, then ventrally, from posterior margin of orbit to ventral

midline of head; broad (4th) band arising approximately at midside of head in preopercular area,

extending downward across branchiostegal membranes onto prepelvic area; each band meeting or

confluent with corresponding band of opposite side

at ventral midline of head; bands almost evenly

spaced; posterior 3 bands often broken, wavy, or

with branches of pigment occasionally forming irregular vermiculations on cheek. Parallel pair of

narrow, irregular bands, usually angled posteroventrally from near upper end of 4 th head band,

extending across branchiostegal membranes to dark,

vertical band on fleshy pectoral-fin base; lower band

of pair may be quite short; both bands sometimes

incorporated in vermiculations on cheek. Vertically

oriented, moderately broad, wavy, dark line present

just posterior to upper half of eye, continuing onto

nape. Snout with pair of dark, often slivered, bands

along anterodorsal contour of head; bands separated at dorsal midline by pale space; large specimens with snout entirely dusky. Interorbital area

with 2 short, moderately broad, dark lines angled

dorsoposteriorly from dorsal edge of orbit; anterior

line continuing onto fleshy crest, posterior terminating on nape; posterior line appears as disconnected dorsal extension of anterior band on lower

side of head. Fleshy crest with approximately 3

moderately broad, dark bands; posteriormost band

usually continuous with wavy, dark line posterior

to upper half of eye. Large, intensely dark spot

approximately one-third to one-half size of eye on

side of nape just anterior to level of 1st dorsal-fin

spine; spots on both sides loosely joined across

nape by narrow, dusky to dark line; anterodorsally

inclined, short, narrow, dark line parallels anteroventral border of dark spot, separated from spot by

slightly smaller, very pale line.

Trunk: Pale with approximately 13 evenly

spaced, broad, dark bands, usually reaching from
NUMBER 177 25

dorsal to ventral body contours, sometimes fainter

ventrally and not quite reaching ventral contour;

posterior 2 or 3 bands often represented only as

large midlateral spots or short, broad, midlateral

stripe; spaces separating bands varying from slightly

broader than bands to one and one-half times as

broad as bands; dorsal ends of bands sometimes

expanded slightly, often with pale spot near center

of expansion; dorsal ends of anterior bands sometimes deflected slightly anteriorly, ends of posterior

bands deflected posteriorly; dorsal ends of bands

occasionally separate from rest of bands (more often

true of posterior bands); dorsal ends of posterior

bands occasionally angled and connected irregularly to one another; bands often taper slightly ventrally. Some specimens with a faint, vertical, thin to

moderately broad, dusky line midway in each pale

interspace between body bands, most noticeable

dorsally.

Pectoral Fin: Pale with fairly even spread of fine,

dark specks of pigment on rays; dusky band near

fin base, sometimes confined to dorsal half of fin;

distal portion of fin dusky ventrally. Fleshy pectoralfin base with prominent, broad, dark band extending from near lower end of gill opening to abdomen

just behind pelvic-fin base; bands of opposite sides

with dusky connection across ventral body midline.

Second, narrow, dusky band sometimes present posteriorly on fleshy base of fin.

Pelvic Fin: Pale; dusky proximally where fin

overlies ventral extension of prominent, dark band

on fleshy pectoral-fin base.

Dorsal Fin: Transparent with dusky and dark

markings; base of fin with large, dark blotch at

dorsal end of each body band; blotches extend dorsoposteriorly on fin as broad, dusky bands; longitudinal row of approximately five dark spots present

one-half to two-thirds distance up from fin base;

distal area of fin usually dusky.

Anal Fin: Largely dusky, or pale with dusky

bands; dark spot in membrane posterior to basal

end of every third ray, usually with narrow, dusky

band directed posteroventrally from each spot; distal area of fin dusky; tips of rays abruptly pale.

Caudal Fin: Pale dusky; central portion of fin

base pale with 2 vertically oriented, dark spots,

each with streak of dusky pigment extending slightly posteriorly; dorsal and ventral portions of fin

base dusky.

Females: Similar to males except: dark markings

on head, body, and fins, other than dark spot on

side of nape, usually fainter; body bands noticeably

fainter, particularly posteriorly; pale spots in dorsal

expansions of body bands more apparent.

GEOGRAPHIC DISTRIBUTION (Figure 6).—Port

Maone, Mozambique, south to Port St. Johns, South

Africa. Smith (1959) and Penrith and Penrith (1972)

reported that O. banditus occurred as far north as

Bazaruto Island, Mozambique, but we have seen no

specimens from north of Port Maone.

HABITAT.—Smith (1959) stated that O. banditus

occurred in weedy and muddy areas; Penrith and

Penrith (1972) reported it from rocky intertidal

pools; Millard and Broekhuysen (1970) reported it

from dead oyster shells in the St. Lucia estuary

basin where salinities ranged from 34.4-35.2 o/oo.

COMPARISON.—Omobranchus banditus is unique

in Omobranchus in lacking a lower-lip flap (Figure

2a). Its bold pattern of body bands also readily

distinguishes it. Omobranchus banditus together

with punctatus and mekranensis appear to form a

closely related group that share the following specializations: one or both anal-fin spines of males

reduced and not discernible externally, a large

number of epipleural ribs, a gill opening predominantly restricted to the area dorsal to the level of

the dorsalmost pectoral-fin ray, a dark spot or band

present dorsally on the side just ventroanterior to

the dorsal-fin origin, and a strong trend toward

increased number of precaudal vertebrae to more

than 10. Omobranchus woodi is also close to this

group but lacks the anal-fin spine modification;

however, woodi, banditus, and mekranensis share

in having a fleshy crest on the head of males; the

crest is lacking in punctatus. The only other species

of Omobranchus in which the anal-fin spines of

males are not discernible externally is steinitzi.

Omobranchus steinitzi, however, has the lowest

number of dorsal- and anal-fin elements and vertebrae in the genus. It also has only 14-15 epipleural

ribs and is unique in the genus in lacking prenasal

pores.

Aside from the lack of a lower-lip flap, presence

of modified anal-fin spines, and features of its color

pattern, O. banditus can be distinguished from the

South African populations of those species of Omobranchus with which it appears to be broadly

sympatric as follows: xvoodi has 0-6 (modally 5)

lateral-line tubes (versus 5-9, modally 7, for banditus) and a fleshy crest on the head of females
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(females of banditus lack crests). Omobranchus

ferox has 0-4 lateral-line tubes, which extend posteriorly no farther than to below dorsal-fin spine 7

(versus 5-9 tubes extending posteriorly to below

dorsal-fin spines 8-11); O. ferox has the gill opening

extending ventrally to opposite the 3rd to 6th

pectoral-fin ray (versus restricted to area dorsal to

dorsalmost ray); O. ferox also has 4 circumorbital

bones, 10-13 epipleural ribs, and no fleshy crest on

the head of either sex (versus 5 circumorbital

bones, 15-21 epipleural ribs, and a fleshy crest on

many males); O. punctatus lacks a fleshy crest in

both sexes. Omobranchus elongatus has 0-2 lateralline tubes, which extend posteriorly no farther than

to below the level of secondary dorsal-fin spine, 11-

14 epipleural ribs, gill-opening rarely restricted to

area dorsal to level of dorsalmost pectoral-fin ray

(usually extending ventral to first ray), and no

fleshy crest on the head of either sex. Omobranchus

fasciolatus has 2 interorbital pores (versus 3, rarely

2, for banditus) and usually has fewer vertebrae,

dorsal-fin elements, and epipleural ribs than banditus (see Tables 5 and 8).

NOMENCLATURAL DISCUSSION.—The holotype

(RUDI 232) of O. banditus was not seen by us;

however, the illustration of the holotype (Smith,

1959, pi. 14d) and the description, which calls attention to the diagnostic lack of a flap on the lower

lip, are sufficient for identification of the specimens

here assigned to O. banditus. In the description

Smith states that the type is 50 mm long; in the

legend for the figure of the holotype, he states that

the type is 51 mm long.

MATERIAL EXAMINED.—MOZAMBIQUE: Port

Maone, USNM 197640 (31.5); Inhaca. RUSI 1099

(3: 31.2-42.7), SAM 26406 (ca. 31); Delagoa Bay,

SAM 15547 (7: 22.1-39.3). SOUTH AFRICA: St.

Lucia coast, ANSP 54998-99 (2: 32.5-36.8); Ballito

Bay, RUSI 1095 (18: 24.1-42.2); Durban, ANSP

51282 (39.1), 55121-23 (3: 34.8-53.3), BMNH

1919.4.1.30 (49.3), 1920.7.23.65 (44.5), FMNH 47025

(7: 31.4-48.6), 47223 (6: 25.8-55.8); Port St. Johns.

SAM 23330 (44.6), 23850 (2: 32.1-46.2).

Omobranchus elegans (Steindachner)

FIGURE 12

Petroscirtes elegans Steindachner, 1876:217 [Nangasaki,

Japan].

Petroscirtes lineo-punctatu.s Sauvage, 1880:216 ["Japon"].

DESCRIPTION (see also Table 5).—Dorsal fin XIIXIII (XII in 96.4% of specimens), 20-23 (21-22 in

93.1% of specimens) = 32-35 (33-34 in 92.1% of

specimens); anal fin II, 21-24 (22-23 in 92.0% of

specimens); both anal-fin spines of males discernible

externally; segmented caudal-fin rays 12-14 (13 in

95.5% of specimens); dorsal + ventral procurrent

caudal-fin rays 9-15 (9, 10, and 15 in one specimen

each of 131 specimens); vertebrae 10-12 (11 in

97.5% of specimens) + 28-31 (29-30 in 93.1% of

specimens) = 39-42 (40-41 in 92.2% of specimens);

epipleural ribs 11-18 (11-15 in 98.6% of specimens); prenasal pores present; interorbital pores

2-4 (3 in 96.2% of specimens); circumorbital pores

6-8 (7 in 88.8% of specimens); lateral-line tubes 0

or 1 (1 in 95.5% of specimens); lateral-line tubes,

when present, extending posteriorly to below level

of dorsal-fin spine 1-3; gill opening varying from

restricted to area dorsal to level of dorsalmost

pectoral-fin ray (1 of 117 specimens) to extending

ventrally to opposite 5th ray (opposite rays 2-4 in

91.5% of specimens); lower-lip flap present; circumorbital bones 5; lower jaw teeth 16-27; upper jaw

teeth 15-24 (Figure 36); fleshy bladelike crest on

top of head absent in both sexes (head of some

presumably mature males swollen in appearance).

COLOR PATTERN.—Males: Head: Two moderately

narrow, dark dusky bands, separated by pale interspace, extending from ventral margin of eye; anteriormost band crosses mouth just anterior to angle

of jaws, bends slightly posteriorly and continues to

ventral midline of head, where il joins corresponding band of opposite side; second band extends vertically from ventral margin of orbit, bends slightly

posteriorly on ventral side of head and continues

to ventral midline where it joins corresponding

band of opposite side. Third dark dusky band

usually encircling head posterior to eye; band broad

dorsally and dorsolaterally, tapering below level of

eye of just below preopercular pores and continuing posteroventrally to ventral midline as broader,

often fainter, segment; ventral portion of band

usually broken into mottled pattern; some specimens with short, broad, dark dusky line connecting

posterior margin of orbit to encircling band midlaterally. Dark dusky freckles or mottling present

between and posterior to head bands on ventral

side of head, branchiostegal membranes, and prepelvic area. Snout and interorbital dusky. Broad,

dark dusky band extending from dorsal body con-
NUMBER 177 27

tour at bases of first 2 dorsal-fin spines (and posterior nape in some cpecimens) to just above dorsal

end of gill opening; anteroventral corner of band

connected across dorsal end of operculum by broad,

irregular mark to short, broad, dark dusky bar or

blotch on midanterior edge of operculum. As many

as 12 small, intensely dark spots present on cheek

and area posterior to eye; several spots also usually

present along posteroventral edge of operculum.

Trunk: Up to about 10 broad bands on body,

anterior 4 or 5 bands darkest and broadest; posterior bands very faint, sometimes barely noticeable;

bands originate on dorsal body contour, become

faint to absent on belly or on ventral half of body

posterior to belly; 2nd and 3rd bands often bifurcate ventrally; sometimes narrow, dark bar present

ventrolaterally between 1st and 2nd band. Body

covered by small, intensely dark spots; spots largest

anteriorly, becoming progressively smaller posteriorly and along dorsal and ventral body contours;

occasionally spots tend to be aligned along anterior

and posterior margins of darkest body bands.

Pectoral Fins: Pale with even spread of very fine,

dark spots on rays, spots slightly larger proximally.

Fleshy base with about 10 crowded, moderately

large, intensely dark spots.

Pelvic Fin: Pale.

Dorsal Fin: Pale to transparent with dark to

dusky markings. Base of fin with dark blotchlike

continuations of body bands anteriorly. Dorsoposteriorly slanted dark line arising from blotch above

band at posterior end of nape; 2nd parallel line

arising from dorsal end of first body band present

on anterior portion of fin; lines becoming fainter

distally on fin; 1st and occasionally 2nd line with

several intensely dark spots along its length; distal

half of fin with series of very faint, narrow, dorsoposteriorly directed, dusky lines parallel to distal

ends of first 2 dark lines. Some specimens with

additional small, intensely dark spots mostly on

anterior half of fin, but occasionally on posterior

portion also.

Anal Fin: Pale to transparent with dark area

anterodistally (tips of rays abruptly pale in this

area). Irregular, longitudinal row of moderately

small, dark spots just ventral to ventral body contour, each spot on membrane between bases of 2

adjacent fin rays; spots becoming smaller and

fainter posteriorly.

Caudal Fin: Pale to transparent with several

irregularly placed, horizontal, dusky blotches. Some

specimens with pair of vertically oriented, faint,

dusky blotches at basal edge of fin, each giving rise

to very faint, posteriorly directed, dusky streak.

Females: Similar to males except: head generally

paler; bands on head fainter ventrally, sometimes

fading out before reaching ventral midline; freckles

and mottling on underside of head almost totally

absent; body slightly paler.

GEOGRAPHIC DISTRIBUTION (Figure 4).—Southern

Japan, southern Korea, and Shantung Province,

China. Omobranchus elegans (as Dassori) was reported by Chyung (1954) from Pusan, Masan, and

Cheju-do, Korea. Chyung (1961) identifiably illustrated the species (as Dassori) but we were unable

to ascertain localities from the Korean ideographs.

We have included the Korean localities on Figure 4

on the basis of the 1954 report, together with the

1961 illustration. There is a specimen (USNM

143553) questionably labeled as having come from

Mauritius. This locality is improbable as the species

is otherwise known only from north of about 35°

north latitude; Mauritius is at 20° south latitude.

HABITAT.—Chyung (1954, 1961) reports this species from shallow waters and dead clams.

COMPARISONS.—Omobranchus elegans is instantly

recognizable by its unique color pattern: a heavy

sprinkling of fine, dark spots covering the entire

body, together with prominent dark bands anteriorly on the body. Omobranchus elegans and aurosplendidus, which is allopatric to elegans, are the

only species in the genus with modally 7 circumorbital pores; the other species have modally 8 or 9

pores. Omobranchus aurosplendidus has a fleshy

head crest, which is lacking in elegans. Aside from

color pattern, elegans can be distinguished from the

populations of the three other species of Omobranchus with which it appears to be broadly sympatric

as follows: punctatus has 5-8 lateral-line tubes,

which extend posteriorly at least to below the level

of dorsal-fin spine 7 (versus 0-1 tube, which extends

no farther than to below spine 3); O. punctatus

also has more than 19 epipleural ribs (versus 11-18,

rarely more than 15, in elegans); loxozonus has 0—4

lateral-line tubes, which may extend posteriorly to

below the level of dorsal-fin spine 6, and 12—14

(modally 13) dorsal-fin spines (versus 12-13, rarely

13 in elegans); fasciolatoceps has 3-8 lateral-line

tubes, which extend posteriorly at least to below

the level of dorsal-fin spine 5, 2-3 (modally 2) in-
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

terorbital pores (versus 2-4, modally 3 pores in

elegans), and a fleshy crest on the head (absent in

both sexes of elegans).

NOMENCLATURAL DISCUSSION.—The holotype of

P. elegans is apparently lost; however, the original

description, especially in the details of the color

pattern, clearly refers to the species here accorded

Steindachner's name.

Sauvage (1880) did not compare P. lineopunctatus with any other species. The holotype is a

typical example of O. elegans.

MATERIAL EXAMINED.—CHINA: SHANTUNG PROVINCE: Tsingtao, USNM 130378 (16.6), 130366 (9: ca.

16-42). JAPAN: MNHN 5120 (68.5, holotype of

Petroscirtes lineopunctatus). KUMAMOTO PREFECTURE: Tomioka, USNM 208471 (44.4). YAMAGUCHI

PREFECTURE: Shimonoseki, USNM 71503 (28: 23.2-

58.9); Mitajiri, USNM 70774 (8: 17.6-48.2). KAGOSHIMA PREFCTURE: Akune, Satsuma, USNM 71492

(2: 51.5-53.2). KANAGAWA PREFECTURE: Misaki, Sagami, USNM 71530 (90: 21.2-64.3, including 3

cleared and stained), 208472 (4: 47.7-50.8), 208473

(4: ca. 54-58.0); Hayamamachi, USNM 199522 (4:

43.0-53.3); Yokohama, USNM 44442 (ca. 44). WAKAYAMA PREFECTURE: Wakanoura, USNM 50270 (15:

18.0-58.6). CHIBA PREFECTURE: Pacific coast of

Boshyu, USNM 195811 (47.6).

Omobranchus elongatus (Peters)

FIGURES 13-15

Petroscirtes elongatus Peters, 1855a:249; 1855&:440 [Mossambique].

Petroskirtes kallosoma Bleeker, 1858:227 [Biliton occidentalis,

in mari].

Petroscirtes dispar Fowler, 1937:258 [Bangkok, Siam; junior

primary homonym of Petroscirtes dispar Gunther, 1861,

= Omobranchus punctatus (Valenciennes)].

DESCRIPTION (see also Tables 5 and 6).—Dorsalfin XII-XIV (XIII in 88.9% of specimens), 17-20

(rarely 17) = 30-33 (rarely 30 or 33); anal fin II,

20-23 (rarely 20); both anal-fin spines of males

discernible externally; segmented caudal-fin rays

12-14 (13 in 98.2% of specimens); dorsal + ventral

procurrent caudal-fin rays 10-16 (rarely 10 or 16);

vertebrae 10-11 (10 in 96.5% of specimens) + 27-

30 (rarely 30) = 37-40 (rarely 40); epipleural ribs

11-15 (rarely 11 or 15); prenasal pores present;

interorbital pores 1-4 (3 in 97.6% of specimens);

circumorbital pores 7-9 (8 in 96.5% of specimens);

lateral-line tubes 0-9 (rarely 8 or 9); lateral-line

tubes, when present, extending posteriorly to below

level of dorsal-fin spine 1-10 (rarely 10); gill opening varying from restricted to area dorsal to level

of dorsalmost pectoral-fin ray to extending ventrally to opposite 6th ray (rarely restricted dorsal

to dorsalmost ray or extending ventral to 5th ray);

lower-lip flap present; circumorbital bones 5; lower

jaw teeth 18-26 (Figure 39); upper jaw teeth 15-24

(Figure 38); no fleshy bladelike crest on top of head

of either sex.

COLOR PATTERN.—Males: Head: Two diffusely

dusky to dark bands on side of head below eye;

anterior band extends from anteroventral margin

of eye across corner of mouth to ventral surface of

chin, where it joins corresponding band of opposite

side; second band (frequently absent) extends from

posteroventral margin of eye to ventral surface of

head, where it may join corresponding band of

opposite side. Ventral third of head posterior to

bands, and prepelvic area, with numerous dark

spots, smaller than pupil of eye. Large, dark

smudge present between circumorbital and preopercular series of cephalic sensory pores at level

of lower two-thirds of eye; smudge domeshaped in

some specimens, occasionally with anterior margin

slightly intensified as spot. Operculum occasionally

with anterodorsally directed, pale dusky, oval spot,

partially to completely encircled by distinct, narrow, pale margin; vertical, darkened area present

between spot and posterior preopercular margin;

opercular membrane dusky posterior to spot. Snout

with middorsal dark patch extending posteriorly to

top of head. Ill-defined, horizontal marks dorsolaterally on head posterior to eye in freshly preserved

specimens.

Trunk: About 11 or 12 dark bands laterally; 2nd

through 7th bands arranged somewhat in pairs;

lower half of bands vertical or slanted slightly posteriorly, reaching nearly to ventral midline of body

anteriorly; posteriorly lower half of bands shorter,

often faint or absent, occasionally ventral portions

of all bands poorly developed; upper half of anterior 5 bands inclined dorsoanteriorly; upper half of

succeeding bands decreasingly inclined dorsoposteriorly; upper end of posterior bands often elongate;

bands not reaching onto upper one-fifth of body;

posteriormost 1 or 2 bands usually represented only

as narrow, horizontal, dark spot or spots. Body
TABLE 6.—Frequency distributions for certain characters in populations of

Omobranchus elongatus

Population

Zanzibar

Mozambique

Madagascar

Seychelles Islands

Chagos Islands

Ceylon

Andaman Islands

Nicobar Islands

Sumatra

Billton

Gulf of Thailand

Philippine Islands

Celebes

Ambon

Population

Zanzibar

Mozambique

Madagascar

Seychelles Islands

Chagos Islands

Ceylon

Andaman Islands

Nicobar Islands

Sumatra

Billto n

Gulf of Thailand

Philippine Islands

Celebes

Ambon

Population

Zanzibar

Mozambique

Madagascar

Seychelles Islands

Chagos Islands

Ceylon

Andaman Islands

Hicobar Islands

Sumatra

Biliton

Gulf of Thailand

Philippine Islands

Celebes

Ambon

Population

Zanzibar

Mozambique

Madagascar

Seychelles Islands

Chagos Islands

Ceylon

Andaman Islands

Nicobar Islands

Sumatra

Biliton

Gulf of Thailand

Philippine Islands

Celebes

Ambon

12

1

5

I*

1

1

1

11

1

3

Dorsal-fin

spines

13

1

39

23

6

18

15

1

It

1

1

132

11

10

Ik

1

-

-

-

7

-

1

-

Precaudal

vertebrae

10

2

1*2

26

6

19

Ik

1

3

1

1

11*9

10

13

11

1

_

_

2

.

1

2

1

-

0

~\ O

10

3

X

10.0

10.0

10.0

10.0

10.0

10.1

10.0

10.0

10.2

10.0

1 2

65 3

18 l

2 3

1

3 -

1

X

12.5

12.9

12.8

12.8

12.9

12.9

13-0

13.0

12.9

12.8

27

1

1

2

1

LI ]

6

1

1

Segmented

dorsal-fin rays

17 18

16

2 9

3

1 1

3

1

3

1

9 Ik

1 6

1

9

19

1

25

15

3

15

11

1

1

65

5

k

Caudal

vertebrae

28 29

2

11 28

12 13

k 2

8 10

6 8

3 -

1

1

L06 32

It 2

10 2

lateral-line tubes

3

2

6

1

1

7

1

1

k

Dorsal

11

3

1

3

12

1

28

9

3

1

1

1

25

3

k

2

2

k

1

19

5

k

+

5

3

3

k

2

32

k

2

6 7

1 -

2

2

1 -

1

39 10

1

3 -

30

_

3

1

_

1

_

1

-

-

>

8

-

3

-

•

20

1

k

1

1

1

2

-

2

-

-

X

29-0

28.8

28.5

28.3

28.6

28. k

28.0

28.2

27-7

28.1

9

X

19.5

18.7

18.6

18.7

18.9

18.9

18.2

l8.lt

18.3

18.3

Z

0.

 Q

- 2

- 3.

- k.

- 5.

1 5.

- 4.

- 4

ventral procurrent

caudal-fin rays

13

8

7

7

3

_

1

63

6

1

Ik

-

9

6

9

2

U8

5

5

15

-

-

3

1

1

1

9

-

1

16

-

-

-

-

-

l

l

~

7

0

3

k

3

2

2

3

S

12.

12.

lit.

13-

13-

13-

13.

13-

0

9

2

7

8

2

7

3

37

1

1

1

1

11

it

1

Total

30

3

1

1

6

1

1

dorsal-fin

elements

31

18

11

3

2

2

1

3

1

83

7

1

11

32

2

25

12

It

lit

11

1

1

6o

it

l

Total

vertebrae

38

11

11

it

8

6

3

1

105

6

1

10

39 ko

2

27

13

2

10

9

_

1

33

2

2

0 1

jo n

1

0

2

it

l

.

l

-

_

l

-

-

33

_

2

1

_

1

2

_

1

-

-

X

39.0

38.8

38.5

38.3

38.6

38.5

38.0

*

38.2

37.8

38.1

s

32.

3 1 .

3 1 .

3 1 .

3 1 .

3 1 .

3 1 .

3 1 .

32.

3 1 .

0

6

it

6

8

9

2

It

2

0

11

1

1

1

Segmented anal-fin

rays

20 21 22

1

3 37

9 15

1 5

1 15

2 9

1

2 2

1

1

3 52 87

6 6

1

io 3

Epipleural ribt

12

1

16

2

9

1

1

18

2

2

13 Ik 15

1 - -

18 5 -

21 2 -

1

It - -

8 2 -

1

3 - -

1

97 29 2

8 2 -

1

9 1 -

lest lateral-line tube below

dorsal-fin spine

c

2 3

;*r _ y i ~

15 1

3 1

1

1

3 1

1

it

2

6

-

6

-

1

Lower

5

1

1

it

1

17

3

1

it

end

opposite

1

9

6

2

2

32

5

1

1

15

1

6

it

3

It

-

19

2

1

7

-

1

1

2

1

1

33

3

6

of gill

8 9 10

l - -

3 - -

1 1 -

_ .

25 7 3

3 - -

1 -

opening

iectoral-fin ray

3

28

10

5

1

1

25

It

it

5

2

9

It

2

8

it

1

5

5 6

-

1 3

5

3

1

1

2

3

2 2

?3 X

1 22.

5 22.

3 21.

- 21.

3 22.

It 22.

- 21.

9 21.

- 21.

- 21.

X

12.5

12.7

12.9

12.2

13.1

12.8

13.1

13.0

12.9

X

1.6

3.2

it.8

5.9

6.3

6.8

6.2

6.2

X

2 . 3

2 . 8

lt.0

lt.0

k.3

2 . 9

3.8

!t.7

5

0

8

8

1

1

5

7

5

2
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

bands narrow, broadest medially, tapering dorsally

and ventrally, often surrounded by poorly defined

dusky area; bands usually indistinctly separated

from dusky areas. Pale background spaces between

bands with dorsoanteriorly slanted, very narrow,

paler band extending from just below dorsal fin to

near ventral body midline; posterior paler bands

with dorsal half inclined dorsoposteriorly; paler

bands not evident in faded specimens. Two longitudinal series of paired, elongate, dark spots dorsally on body; dorsal series at junction of dorsal fin

and body; ventral series just ventral to dorsal series,

with each spot directly under corresponding spot

of dorsal series; dorsal series united into narrow

stripe in some specimens. Trunk portion of gill

opening with dusky to dark bar at posterior border

of opercular membrane.

Pectoral Fins: Transparent with uniform spread

of dark specks of pigment. Fleshy base with 0 to 4

small, dark spots, 1 or 2 occasionally extending onto

basal portion of fin membrane.

Pelvic Fins: Pale with uniform spread of fine,

dark specks of pigment.

Dorsal Fin: Generally transparent with series of

dark stripes; stripes on spinous portion moderately broad, curving ventroposteriorly; stripes on

segmented-ray portion narrow, more numerous,

curving dorsoposteriorly. Large, intensely dark spot

(ocellus), encircled by transparent or pale margin,

located slightly above center of fin between 7th and

14th segmented rays (usually extending from 9th to

11 th rays). Series of dusky marks basally on fin associated with adjacent series of body spots. Some

specimens with fin more or less uniformly dusky to

dark, except for narrow, pale, distal margin.

Anal Fin: Dusky to dark, darkest distally, with

narrow, pale, distal margin. Freshly preserved specimens with narrow, curved, oblique, pale lines directed ventroposteriorly from lower ends of paler

bands on body; pale lines terminate proximal to

distal third of fin, become faint to absent posteriorly on fin. Dark spot just posterior to proximal

end of each oblique pale line (after approximately

every 3rd ray base), obscured in specimens with

dark fins.

Caudal Fin: Pale with uniform spread of dark

specks of pigment; dorsal and ventral margins dark.

Caudal peduncle with moderately large, dark spot

just above and below lateral midline at caudal-fin

base: pigment streak from each spot continued posteriorly onto fin.

Females: Color pattern as in males except: overall

background paler; dark markings on trunk and

posterior body better defined; dusky patches surrounding body bands poorly developed or absent;

bands often undeveloped ventrally; narrow, paler

bands in pale spaces between dark bands often

inapparent; dorsal and anal fins largely transparent,

unmarked; ocellus in dorsal fin absent; anal fin

dusky only along proximal edge of pale, distal

border.

Geographic Variation: Although the color pattern variations of specimens from any particular

area appear to encompass the variations found in

the species as a whole, there is a prevalence of specimens with reduced head and fin markings from

southern Africa and Madagascar and specimens

with the same markings intensified from the Gulf

of Thailand and Philippine Islands.

Gulf of Thailand: Both dark bands on head well

developed; spots on underside of head small, distinct, numerous; dusky smudge posterior to eye

usually intensified anteriorly; narrow, paler body

bands (in spaces between dark body bands) not

apparent in many specimens; fleshy pectoral-fin

base usually with 4 distinct spots; dorsal fin with

distinct, dark stripes in males; dorsal-fin ocellus of

males located between 7th and 10th segmented rays,

occasionally reaching posteriorly to 11th.

Madagascar: Anterior dark band on head faint;

2nd band usually faint or absent; spots on underside of head large, indistinct, faint, few in number;

dusky smudge posterior to eye usually without anteriorly intensified area; narrow, paler body bands

usually distinct; fleshy pectoral-fin base often without spots, occasionally with narrow, pale bar at

proximal edge of fin; dorsal fin of males with

stripes obliterated by even, dusky pigmentation at

anterior and posterior ends of fin; dorsal-fin ocellus

of males located between 9th and 14th segmented

rays, occasionally reaching anteriorly to 8th ray.

GEOGRAPHIC VARIATION (see also "Color Pattern").—Variation in meristic characters (Table 6)

occurs in some populations of O. elongatus. The

Gulf of Thailand population has a higher average

number of lateral-line tubes and lower average

numbers of total dorsal- and anal-fin elements and

vertebrae than most of the other populations. The

Mozambique population was notable for having

the lowest numbers of total procurrent caudal-fin
NUMBER 177 31

TABLE 7.—F values for covariance comparisons of regression equations

(Table 2) for number of lower or upper jaw teeth between males or

females from different populations of Omobranchus elongatus (** =

significant at p = .001; NS = not significant)

Comparisons

Lower teeth

males

Madagascar

Madagascar

Mozambique

females

Chagos Is.

Chagos Is.

Mozambique

Upper teeth

males

Madagascar

Madagascar

Mozambique

females

Mozambique

X

X

X

X

X

X

X

X

X

X

Mozambique

Gulf of Thailand

Gulf of Thailand

Mozambique

Gulf of Thailand

Gulf of Thailand

Mozambique

Gulf of Thailand

Gulf of Thailand

Gulf of Thailand

F values

slopes

0.11 NS

1.76 NS

1.07 NS

1.24 NS

0.11 NS

2.87 NS

0.50 NS

0.94 NS

0.04 NS

0.14 NS

Degrees of

freedom

1/36

1/39

1/51

1/31

1/31

1/50

1/35

1/38

1/49

1/50

F values

heights

24.59 **

3.90 NS

67.18 «*

26.70 **

1.84 NS

80.46 *»

7.34 NS

6.10 NS

40.58 «•

64.36 **

Degrees of

freedom

1/37

1/40

1/52

1/32

1/32

1/51

1/36

1/39

1/50

1/51

rays and lateral-line tubes. It was the only population in which some specimens lacked lateral-line

tubes altogether.

Although covariance comparisons of numbers of

jaw teeth were not feasible for all populations of

O. elongatus, significant differences do occur among

those populations that were compared (Table 7).

GEOGRAPHIC DISTRIBUTION (Figure 5).—Tropical

Indian Ocean, excluding the Red Sea and Persian

Gulf, east to the Philippine Islands and Ambon in

the Pacific Ocean.

A completely faded specimen from Sumbawa

Island, Indonesia, questionably referred to O. rotundiceps obliquus could possibly be O. elongatus

(see "Geographic Distribution" under O. r. obliquus).

Menon and Rama Rao (1963) recorded O. kallosoma from Madras and Mandapam, India, and the

Andaman Islands (actually Nicobar Islands). Their

specimens from India are O. elongatus, whereas

their specimen from the Nicobar Islands is O. rotundiceps obliquus. Of the Indian specimens, we

were able to examine only the Mandapam specimen

(ZSIC F6916/2), which arrived too late to be included in Table 6. We have entered the Madras

record for O. elongatus on Figure 5 based on the

color pattern description given by Menon and

Rama Rao. Meristics for the Mandapam specimen

are as follows: dorsal fin XIII,20; anal fin 11,23;

total procurrent caudal-fin rays 12; vertebrae 11 +

29; epipleural ribs 13; lateral-line tubes 2, extending posteriorly to below fourth dorsal-fin spine;

interorbital pores 3; circumorbital pores 8; gill

opening extending ventrally to opposite second

pectoral-fin ray.

Chu, et al. (1962) reported and illustrated O.

elongatus (as O. kallosoma) from Hainan, and Jones

and Kumaran (1965) described and illustrated O.

elongatus from Minicoy, Laccadive Islands. We

have also included these records on the distribution

map.

Munro (1958) mistakenly included O. elongatus

in his checklist of New Guinea fishes. His error was

the result of his reporting the type-locality of Petroscirtes germaini Sauvage, which species he wrongly

considered to be a synonym of O. elongatus, as New

Guinea, instead of New Caledonia. Munro (1967)

continued the error, twice illustrating O. elongatus,

based on published illustrations of extralimital

records.

HABITAT.—Shallow (0-4 m) coastal waters and

tide pools with rocks and little coral. Occasionally

recorded from brackish water and stream mouths

and from rocks with oysters.

COMPARISONS.—Omobranchus elongatus belongs

to a group of species also comprising germaini, rotundiceps, and loxozonus. This group of species is

united by the following unique specializations:

Males may have one or more caudal-fin rays strongly exserted and/or filamentous and a dark ocellus

or group of dark spots usually present near the

middle of the segmented-ray portion of the dorsal
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fin. In addition, all the species have a distinctive

pattern of bands on the body: the anterior bands are

inclined anteriorly from their ventral origin and the

posterior bands are inclined posteriorly. Some males

of O. fasciolatus may have a similar dark blotch in

the dorsal fin, but the pattern of bands on the body is

different and the caudal-fin rays are never exserted or

filamentous. Most male O. fasciolatus have a fleshy

crest on the head, a structure not present in the

elongatus species group. Omobranchus ferox males

may also have a dark blotch in the dorsal fin but it

is positioned at the posterior end of the fin. The

pattern of bars on the body of O. ferox is usually

very faint or absent, but when present it adumbrates

that of the elongatus group; however, the narrow

pale and dark bars adjacent to the posterior margin

of the orbit (Figure 29 f) readily separate O. ferox

from the elongatus group.

With the exception of O. germaini, which overlaps the distribution of O. elongatus and O. rotundiceps, the species of the elongatus group are

predominantly allopatric; however, O. elongatus,

O. germaini, and O. rotundiceps have all been collected at the small island of Ambon in Indonesia,

where each occurred at the mouth of a different

stream. Where two or more of these species occur

together, all or most of the specimens of each

species are readily separable. Overlaps in the overall

ranges for characters of the species are the result of

combining individual population variations within

a species. Problems in identification occur with

specimens that have poorly developed color patterns and counts in the overlap portions of the

ranges for the characters of the population under

consideration.

A pattern of small, dark spots on the underside

of the head (Figures 13b, 146) found in many male

and some female specimens of O. elongatus will

distinguish those specimens from specimens of the

other species in the elongatus group. The spotted

pattern is adumbrated by some specimens of O. r.

rotundiceps from eastern Australia (Figure 296),

where O. elongatus does not occur. Within the

elongatus group, O. elongatus is distinctive in lacking a well-defined dark spot posterior to the eye.

NOMENCLATURAL DISCUSSION.—Bleeker (1858) did

not compare P. kallosoma with any nominal species,

although he mentioned that the color patterns of

the fins distinguished it from all other species of

Petroscirtes, a conglomerate genus at that time.

Bleeker was referring primarily to the stripes and

ocellus found on the dorsal-fin rays (of males),

characters which are also present in O. rotundiceps,

germaini, and loxozonus. The holotype, now faded,

is a typical male O. elongatus.

In the files of RMNH are a number of unpublished colored illustrations that Bleeker intended

to use in his "Atlas." Among these is one labeled

Petroscirtes kallosoma, which was probably drawn

from the holotype. This illustration shows one

more dorsal-fin spine and ray than is present in the

holotype. The distinctive spotting under the head is

not indicated, although Bleeker noted it in his

description.

Fowler (1937) compared his P. dispar only with

P. masyae (= O. punctatus). The holotype is a

small female O. elongatus.

MATERIAL EXAMINED.—ZANZIBAR: USNM 12678

(2: 36.8-ca. 39). MOZAMBIQUE: ZMB 1940 (2: ca.

48-50 syntypes of Petroscirtes elongatus). INHACA

ISLAND: RUSI 1101 (65: 27.8-47.6), 1098 (13: 16.2-

53.8), SAM 25169 (4: 28.0-43.2), USNM 197642

(32.6). MADAGASCAR: Nossi Be, USNM 201876

(6: 37.3-48.6), 201877 (2: 25.2-27.1), 201878 (6:

28.5-35.4), 201879 (25.7), 201891 (4: 27.9-34.3),

201892 (3: ca. 25-31), 209235 (3: 29.8-39.3), 209231

(2: 24.2-27.3). SEYCHELLES ISLANDS: MAHE:

ANSP uncataloged, Sta. F-l (18.8), Sta. F-116

(13.8), Sta. F-l 17 (4: 13.0-14.6). ANONYME: ANSP

uncataloged, Sta. F-21 (2: 24.9-25.7). CHAGOS

ARCHIPELAGO: DIEGO GARCIA ATOLL: USNM

209232 (18: 22.5-43.8), 209230 (33.2). INDIA: Mandapam Camp, ZSIC F6916/2 (44.1). CEYLON:

USNM 209228 (11: 16.0-38.0), 209233 (4: 16.4-

40.4), NFIS 11621 (27.4). ANDAMAN ISLANDS:

USNM 144418 (ca. 33). NICOBAR ISLANDS:

NFIS 11859 (2: 21.8-27.6), 11860 (2: 30.8-34.1).

INDONESIA: SUMATRA: 00° 41' N, 97° 53' 30' E,

USNM 201558 (26.9). BILITON: RMNH 4452 (48.2,

holotype of Petroskirtes kallosoma). CELEBES: Kwandung Bay, ZMA 113014 (28.8). BUTON (also Butung):

USNM 211872 (25.7). AMBON: USNM 211878 (13:

19.5-43.6). GULF OF THAILAND: southeast portion, USNM 119664 (2: 27.4-29.3). GOH KRAM

ISLAND: USNM 209425 (123: 13.2-39.8). GOH SAK

ISLAND: CAS GVF reg. no. 1459 (71: 20.3-46.4). GOH

SAMED ISLAND: CAS GVF reg. no. 2198 (43: 23.1-

47.0). KOH KUT ISLAND: USNM 119661 (4: 28.1-

41.7). THAILAND: Bangkok, ANSP 68255 (24.0,

holotype of Petroscirtes dispar Fowler). PHILIP-
NUMBER 177 33

PINE ISLANDS: NEGROS: CAS GVF reg. no. 1597

(35.9), GVF reg. no. 1598 (3: 31.1-41.6). MINDANAO:

AMNH 15559 (6: 32.3-48.6), FMNH 40581-83 (3:

32.4-35.2). A large number of CAS specimens from

the northern Gulf of Thailand were examined, in

addition to those listed above, but only associated

ecological data were recorded.

Omobranchus fasciolatoceps (Richardson)

FIGURE 16

Blennius fasciolatoceps Richardson, 1846:265 [Macao, based

on a then-unpublished illustration].

Petroscirtes dispar Gunther, 1861:232 [in part; Amoy, China;

name restricted under Omobranchus punctatus].

Chasmodes herklotsi Herre, 1935:288 [southern coast of Hong

Kong].

Petroscirtes uekii Katayama, 1941:591 [Higashiiwase of

Toyama Bay].

DESCRIPTION (see also Table 5).—Dorsal fin XII,

19-22 = 31-34; anal fin II, 22-24; (Katayama and

Ikeda, 1973, reported an anal-fin count of II, 21 for

a specimen from Yamaguchi Prefecture, Japan);

both anal-fin spines of males discernible externally;

segmented caudal-fin rays 12-13 (12 in 1 of 21

specimens); dorsal + ventral procurrent caudal-fin

rays 11-13; vertebrae 10 + 27-30 = 37-40; epipleural ribs 11-13 (13 in 1 of 20 specimens); prenasal pores present; interorbital pores 2-3 (2 in 18

of 19 specimens); circumorbital pores 7-8 (8 in 17

of 20 specimens); lateral-line tubes 3-8; lateral-line

tubes extending posteriorly to below level of dorsalfin spine 5-10; gill opening restricted to area dorsal

to level of dorsalmost pectoral-fin ray; lower-lip flap

present; circumorbital bones 5; lower jaw teeth 26-

48 (Figure 41); upper jaw teeth 24-35 (Figure 40);

fleshy bladelike crest present on top of head of both

sexes.

COLOR PATTERN.—Males: Head: Four moderately

narrow to broad, pale to dusky bands on head.

Anterior band extending from anteroventral margin of orbit across mouth, just anterior to corner

of jaws, onto chin. Second band extending ventrally

from orbit, angled slightly posteriorly on ventral

side of head. Broad, dark dusky continuation of 1st

or 2nd band directed dorsally from dorsal margin

of orbit to anterodorsal edge of fleshy crest, covering anterior half of crest. Third band encircling

head posterior to eye; above level of mouth band

broad, extending onto posterior end of fleshy crest;

below level of mouth band narrow, slanted posteroventrally; large, dark dusky to dark oval spot present on band posterior to level of eye; spot well

defined all but ventrally, with dark anterior and

posterior margins continued dorsally and ventrally

on band in some specimens. All 3 bands may be

confluent ventrally with corresponding bands of

opposite side. Fourth band encircling head just anterior to dorsal-fin origin; ventral end of band terminating just dorsal to pelvic-fin origin (illustration

by Tomiyama and Abe, 1953, shows band splitting

just below level of eye, and posterior arm of band

directed onto fleshy pectoral-fin base; our specimens do not exhibit this pattern but they are

faded).

Trunk: Pale to pale dusky with approximately

10 evenly spaced, faint, broad, dusky bands. Bands

reaching from dorsal to ventral body contour; dorsal half of bands directed slightly dorsoposteriorly,

ventral half slightly ventroposteriorly, bands becoming very faint to absent posteriorly; posterior onefifth of body apparently without bands.

Pectoral Fin: Pale to transparent with dark specks

of pigment scattered basally and concentrated in

single threadlike lines along dorsal and ventral

edges of each ray.

Pelvic Fin: Pale with dark specks of pigment

scattered basally.

Dorsal Fin: Pale with narrow, dark dusky to

dark stripes. Two moderately narrow stripes present

anteriorly on fin; 3rd stripe appearing distally at

about 6th dorsal-fin spine; stripes becoming slightly

narrower and turning dorsoposteriorly near last

dorsal-fin spine; posterior half of fin with series of

narrow, dorsoposteriorly slanted stripes, becoming

faint distally. Large, but not prominent, dusky spot

present distally between about first 3 segmented

dorsal-fin rays in some specimens.

Anal Fin: Dark dusky to dark, tips of rays

abruptly pale.

Caudal Fin: Pale with evenly scattered dark

specks of pigment; duskier basally.

Females (specimens examined greatly faded):

Similar to males except: overall pigment pattern

apparently paler, bands fainter; dorsal-fin markings

less distinct; dusky spot on dorsal fin apparently

absent.

Useful illustrations of O. fasciolatoceps are contained in Tomiyama and Abe (1953, pi. 193) and

Whitehead (1969, pi. 25c).
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

GEOGRAPHIC DISTRIBUTION (Figure 6).—Coast of

mainland China from Macao and Hong Kong north

to Toyama Bay, Japan. The Macao record is based

on the type-locality of Blennius fasciolatoceps.

Tomiyama and Abe (1953) described and illustrated

a specimen (as O. uekii) from Hamana Lake, Honshu, Japan, which is the northernmost record for

the Pacific coast of Japan. Kamohara (1956) reported a specimen (as O. uekii) from the mouth of

the Kogami River in Kochi City, Shikoku; we have

not recorded this locality on the distribution map

because we are unable to verify the identification.

Katayama and Ikeda (1973) described and illustrated

a specimen (as O. uekii) from the Atsu Higashigawa River in Yamaguchi Prefecture, Japan. We

have entered this locality record on the distribution

map.

HABITAT.—Tomiyama and Abe (1953) reported

O. fasciolatoceps (as O. uekii) from brackish water,

and Herre (1934) reported it (as Petroscirtes dispar)

from oyster beds.

COMPARISONS.—Omobranchus fasciolatoceps is

unique in the Blenniidae in the changes in jaw

structure that occur in mature females (see "Sexual

Dimorphism" under the generic account). It is

otherwise separable from most of the other species

of Omobranchus only by a combination of characters. The possession of 2 interorbital pores and a

fleshy crest on the head separates O. fasciolatoceps

from all the Omobranchus species with which it

appears to be broadly sympatric; O. aurosplendidus,

elongatus, ferox, germaini, loxozonus, elegans, punctatus. Omobranchus aurosplendidus also has a crest,

but it has 3 interorbital pores, more dorsal- and

anal-fin elements, more vertebrae, and fewer lateralline tubes than O. fasciolatoceps (Table 5). The

other species have 3, rarely 2 or 4, introrbital pores

and no crest.

NOMENCLATURAL DISCUSSION.—Richardson (1846)

based his description of B. fasciolatoceps on an unpublished colored illustration, later published by

Whitehead (1969, pi. 25c) in black and white. The

illustration depicts an Omobranchus with a crested

head bearing five dark bands and no other distinctive color pattern on the body. The dorsal fin is

without filamentous anterior dorsal-fin spines or a

black spot covering the base of the anteriormost

spines. Except for the 42 dorsal-fin elements depicted (five more than known for any species of

Omobranchus), the illustration reasonably represents a male of the species here treated as O. fasciolatoceps. (Note: Our illustrations of O. fasciolatoceps, Figure 16, were based on faded specimens,

which did not exhibit the dark bands on the head.)

Giinther (1861) based his description of P. dispar

on two specimens representing two species, O. punctatus and O. fasciolatoceps, which were not considered in his discussion. We have designated his specimen of O. punctatus as lectotype of P. dispar and

in doing so have restricted the name.

Herre (1935) noted the similarity of his C. herklotsi to Giinther's P. dispar, but distinguished it

from that species by stating only that its anterior

profile was markedly different. Herre's holotype and

paratypes were all females that do have a unique

snout physiognomy, while Giinther's specimen of

O. fasciolatoceps was a male, thus accounting for

the difference noted by Herre.

Katayama (1941) also compared his P. uekii only

with P. dispar, noting that his species had a higher

crest and shorter snout. The height of the crest and

length of the snout are highly variable characters.

Although the crest of the larger Japanese male

specimens may be more strongly developed than

that of other specimens, we find no reason to recognize the Japanese specimens nomenclaturally.

MATERIAL EXAMINED.—HONG KONG: CAS

SU62067 (2: 50.0-53.0), SU66811 (4: 42.8-63.4),

SU62063 (3: 40.0-45.2), SU61121 (3: 51.0-61.7),

SU30970 (3: 51.1-57.4, holotype and paratypes of

Chasmodes herklotsi; holotype unsegregated, the

largest specimen), USNM 209264 (61.6). CHINA:

Futschau (Foochow), ZSZM 11386 (32.5), ZSZM uncataloged (46.7); Amoy, BMNH 1860.7.20.100 (ca

59, syntype of Petroscirtes dispar Giinther = O.

punctatus). JAPAN: Higashiiwase, near Toyama

City, private collection of Masao Katayama, Yamaguchi University (59.5); Hamamatsu. ZITU 35257

(47.7).

Omobranchus fasciolatus (Valenciennes)

FIGURE 17

Blennechis fasciolatus Valenciennes in Cuvier and Valenciennes, 1836:287 [based on a then-unpublished figure;

neotype designated below].

Petroscirtes striatus Jatzow and Lenz, 1898:512 [junior primary homonym of Petroscirtes striatus Day, 1888, tribe

Nemophini, Bawi, Zanzibar].

Petroscirtes vinciguerrae Borsieri, 1904: 211 [Massaua].
NUMBER 177 35

Omobranchus cristatus Fraser-Brunner, 1951:214 [tide-pools

at Sirah Island, Aden; junior homonym of Petroscrites

cristatus Zugmayer, 1913 (= Omobranchus mekranensis)].

DESCRIPTION (see also Tables 5 and 8).—Dorsal

fin XI-XII (XII in 95.0% of specimens), 18-19 =

30-31; anal fin II, 20-22; both anal-fin spines of

males discernible externally; segmented caudal-fin

rays 13; dorsal + ventral procurrent caudal-fin rays

12-15; vertebrae 10 + 26-28 (26 in only 1 of 51

specimens) = 36-38; epipleural ribs 12-16 (16 in

only 1 of 53 specimens); prenasal pores present;

interorbital pores 2; circumorbital pores 7-9 (8 in

88.9% of specimens); lateral-line tubes 3-8 (5-7 in

90.5% of specimens); lateral-line tubes extending

posteriorly to below level of dorsal-fin spine 7-10;

gill opening varying from resricted to area dorsal

to level of dorsalmost pectoral-fin ray to extending

ventrally to opposite 2nd ray (restricted to area

dorsal to dorsalmost ray in 95.0% of specimens);

lower-lip flap present; circumorbital bones 5; lower

jaw teeth 18-26; upper jaw teeth 18-25 (Figure

42); fleshy bladelike crest present on top of head of

males over 33 mm SL, present or absent on smaller

males, poorly developed or absent on females.

COLOR PATTERN.—Males: Head: Two narrow,

dark bands present between anterior margin of

orbit and upper lip; row of intense, small, dark

spots occasionally present along each band. Three

broad, mottled, dark bands on lower half of head;

anterior band extending from anteroventral margin

of orbit across mouth just anterior to angle of jaws

to chin, usually joining corresponding band of opposite side on chin; 2nd band extending ventrally

from posteroventral margin of orbit, directed slightly posteriorly at level of anterior preopercular

pores; 3rd band extending ventroposteriorly from

posterior margin of orbit; cheek portion of 3rd band

faint or absent in most specimens; posterior 2 bands

TABLE 8.—Frequency distributions for certain characters in populations of

Omobranchus fasciolatus

Population

Northern Red

Southern Red

Gulf of Aden

Persian Gulf

Arabian Sea

Mombasa

Mozambique

Madagascar

Population

Northern Red

Southern Red

Gulf of Aden

Persian Gulf

Arabian Sea

Mombasa

Mozambique

Madagascar

Sea

Sea

Sea

Sea

Dorsal-fin

spines

11

1

2

12

5

5

6

12

12

1

1

15

X

12.

11.

12.

1 1 .

12.

12.

0

8

0

9

0

0

Segmented

anal-fin rays

20

1

2

1

4

1

3

21

4

3

5

9

5

1

1

12

22

1

1

6

X

20.8

20.8

20.8

20.8

21.4

20.8

Segmented

dorsal-fin

rays

18

3

3

2

6

3

12

1

<

12

1

6

1

19

2

3

4

8

9

1

1

3

X

18.4

18.5

18.7

18.6

18.8

18.2 ]

Dorsal + ventral

procurrent

;audal-fin rays

13

1

3

-

4

2

14 15

4

3

11 2

2

1

9 1

X

13

13

14

12

13

Total

dorsal-fin

elements

30

3

4

2

8

3

L2

8

5

0

7

8

31

2

2

4

6

9

1

1

3

12

2

X

30.4

30.3

30.7

30.4

30.8

30.2

Epipleural

13 Ik

1 2

1 3

4 7

2 7

3 6

15

2

2

3

1

1

5

Precaudal

vertebrae

10

5

6

14

12

1

1

15

ribs

16

1

X 3

14.2

14.2

13.4 1

14.1

14.3

26

1

Caudal

vertebrae

27

2

5

7

4

14

28

3

1

6

8

1

1

Lateral-line

4

1

1

5 6

2 2

5

2 8

3

1

2 8

7

1

1

2

7

4

X

27.6

27.2

27.5

27.7

26.9

tubes

8 X

5.8

6.2

5.6

2 6.9

6.0

Total

vertebrae

36 37

2

5

7

4

1 14

38

3

1

6

8

l

l

X

37.6

37.2

37-5

37-7

36.9

tube below

dorsal-fin spine

7

2

1

2

2

8 9

1 2

3 2

8 3

2 6

1

8 5

10

1

4

X

8.0

8.2

8.2

9.2

8.2
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

usually terminating before reaching ventral midline

of head. Underside of head with dusky spots or

freckles in spaces between bands and in area posterior to 3rd band, particularly well developed posteriorly; narrow, vertical, dusky streaks or lines of

small, dark spots located more dorsally between

bands. Large, dusky, oval area directly behind eye,

extending from level of upper end of orbit to ventral extent of preopercular pores; intensely dark

semicircular spot approximately half size of eye

covering dorsal end of oval area; oval area dusky

anteriorly and medially, usually paler ventrally

and ventroposteriorly; narrow, pale margin along

curved dorsal edge of semicircular spot; some specimens with narrow, posteroventrally inclined, pale

slash across center of oval area. Operculum generally indistinctly mottled except for narrow, pale

slash originating just above pale, dorsal margin of

semicircular spot and running diagonally downward

across upper portion of operculum; posteroventral

portion of operculum sometimes with several tiny,

dark spots; posterior opercular membrane pale.

Interorbital area occasionally with several broad,

dark bands, spots, or blotches continuing dorsally

onto fleshy crest.

Trunk: Approximately 17 vertical, broad, dusky

to dark bands on body; tips of bands reaching to

or near dorsal and ventral body contours, usually

becoming fainter ventrally; occasionally 1 or 2 middle bands not reaching as high as adjacent bands:

posterior bands often faint and blotchlike; dorsal

ends of anterior bands directed slightly anteriorly,

posterior bands slightly posteriorly; dorsal ends of

bands wavy, giving dorsal one-fourth of body color

pattern less regular appearance; some specimens

with bands paired, bands in a band pair only

slightly closer together than band pairs; pigment

along vertical axes of bands slightly intensified;

intensification greatest at 3 points, at lateral midline of body, halfway up dorsal half of body, and

halfway down ventral half of body, giving appearance of 3 longitudinal rows of dark spots in faded

specimens; rows usually not well developed in

males; when evident, midlateral row usually best

developed.

Pectoral Fin: Fin and fleshy base pale with even

spread of dark specks of pigment. Some specimens

with about 4 tiny, dark spots evenly spaced along

basal edge of fin.

Pelvic Fin: Pale with even spread of dark specks

of pigment.

Dorsal Fin: Mostly dusky proximally; membrane

pale to transparent distally, usually with slightly

dusky edge; spinous portion occasionally with dusky

mottling or several dorsoanteriorly slanted, broad,

dusky bands proximally, otherwise evenly dusky,

except more transparent proximally. Large, intensely dark, oval spot usually present somewhat

distally between 9th and 14th segmented rays.

Anal Fin: Pale dusky proximally, dusky to dark

distally, tips of rays abruptly pale.

Caudal Fin: Pale with even spread of dark

specks of pigment. Vertically oriented pair of diffuse, dusky blotches at base of fin giving rise to 2

diffuse, dusky stripes; stripes may converge into 1

posteriorly.

Females: Similar to males except: body bands

largely reduced or fainter; intensifications of pigment in bands usually forming 3 rather distinct

rows of dark spots and often a 4th row above anterior portion of dorsal row; other spots often present

along vertical axes of bands and dorsally; dorsal

and anal fins weakly pigmented, dorsal fin uniformly very pale dusky proximally, transparent distally, dark oval spot in dorsal fin absent.

GEOGRAPHIC VARIATION.—There are slight shifts

in the averages for meristic characters in the various

populations of O. fasciolatus that may prove to be

descriptive of the populations when more specimens are available (Table 8).

GEOGRAPHIC DISTRIBUTION (Figure 4).—Bitter

Lakes of the Suez Canal south to Bazaruto Island,

Mozambique, and east from Persian Gulf to northwest India; Madagascar. Smith (1959) reported O.

fasciolatus (as O. mekranensis) from Malindi and

Shimoni, Kenya, Zanzibar (based on Jatzow and

Lenz, 1898), and Ibo and Bazaruto Island, Mozambique. We have not seen Smith's specimens, but

based on his description and illustration of a specimen from Ibo, we accept his records and have

recorded them on the distribution map.

HABITAT.—Shallow coastal waters and tidepools

in rocky areas and under nonestuarine conditions.

In the "Material Examined" section, the specimen

recorded as having come from Kabret, which is in

the Bitter Lakes, is of interest. The specimen was

collected by the Cambridge Suez Canal Expedition

of 1924, at which time the salinity of the Bitter
NUMBER 177 37

Lakes was 48.5-53.5 o/oo (Fox, 1926, tables 5 and

11). In view of the ability of O. fasciolatus to exist

in this high salinity barrier, it is conceivable that it

might penetrate to, and become established in, the

Mediterranean Sea.

COMPARISONS.—Omobranchus fasciolatus is separable from most of the other species of Omobranchus only by a combination of characters. It is most

easily confused with O. mekranensis, with which it

is sympatric (both species have been taken in the

same collection). It differs from O. mekranensis

most obviously in having 30-31 total dorsal-fin elements (versus 32-33), 10 + 26-28 = 36-38 vertebrae (versus 11 + 28-29 = 39-40), and both anal-fin

spines of males discernible externally (versus one

or both spines not discernible). Male O. fasciolatus

may have a dark spot, which O. mekranensis lacks,

near the center of the segmented dorsal-fin rays.

Omobranchus fasciolatus may be most easily separated from the other Omobranchus species (banditus, elongatus, ferox, punctatus, and woodi) with

which it appears to be broadly sympatric as follows:

from all in having typically 2 interorbital pores

(rarely fewer than 3 pores in the other species);

from O. banditus in having a lower-lip flap and in

lacking prominent, discrete dark bands on the body;

from O. elongatus in frequently having a fleshy

crest, 12 dorsal-fin spines (usually more than 12 in

O. elongatus), the gill opening usually restricted to

the area dorsal to the level of the dorsalmost

pectoral-fin ray (versus usually extending ventrally

to or below 2nd from dorsalmost ray in O. elongatus), and in never having filamentous caudal-fin rays

in males; from O. ferox in having 30-31 total

dorsal-fin elements (versus 32-35) and in having

the gill opening restricted dorsal to the level of the

3rd pectoral-fin ray (versus extending ventral to

the second ray in O. ferox), and in frequently having a fleshy crest; from O. punctatus in frequently

having a fleshy crest, both anal-fin spines of males

discernible externally, 12-16 epipleural ribs (versus

15-25, rarely less than 18 in O. punctatus), and 10

precaudal vertebrae (versus 10-12, rarely 10); from

O. woodi in having 10 precaudal vertebrae (versus

11-12), and usually fewer total dorsal-fin elements

and vertebrae (Tables 5 and 8).

NOMENCLATURAL DISCUSSION.— (For discussion of

the date and authorship of Blennechis fasciolatus

see "Nomenclatural Discussion" under generic account). Valenciennes' (1836) description of B. fasciolatus was based on an unpublished illustration,

which Valenciennes erroneously believed to have

been published, but that was first published by

Hemprich and Ehrenberg (1889). Hemprich and

Ehrenberg noted that the specimen on which the

figure was based came from the Red Sea and was

lost. The general appearance of the figure, presence

of canine teeth posteriorly in both jaws, and the

banded color pattern definitely portray a species of

the generic taxon that we recognize as comprising

all the species in the present study, but it is not

possible to absolutely determine which, if any, of

these species the figure represents. Only two species

of Omobranchus are known from the Red Sea:

fasciolatus and steinitzi. Of these two only the specimens we treat here as fasciolatus exhibit a color

pattern similar to the type shown in the illustration.

Inasmuch as fasciolatus is the type-species of Omobranchus, it is important to tie the generic and

specific names to a specimen in order to stabilize

the nomenclature. We, therefore, designate USNM

204487, male, 44.5 mm SL, from the Ethiopian

Naval Base, Massawa Ethiopia (collected by V. G.

Springer and P. Anastos, 10 August 1969) as neotype of Blennechis fasciolatus. A female specimen

(USNM 209265) was collected with the neotype.

Jatzow and Lenz (1898) described P. striatus

from two specimens from Bawi, Zanzibar. They did

not compare their species with any other; their

specimens are no longer extant. The species was

illustrated, however, and based on the crested head,

color pattern, total number of dorsal-fin elements,

and locality of origin, it is obviously a synonym of

O. fasciolatus.

Of the species of Omobranchus, Borsieri (1904)

compared his P. vinciguerrae only with O. lineolatus

(Kner). We have examined the holotype of P. vinciguerrae and find it to be a male specimen of O.

fasciolatus. The crest on the head is represented

only by a ridge less than 0.3 mm high, and is rather

low for a specimen its size (39.1 mm SL); however,

all other characters of the specimen are in good

agreement with those of other specmens of O.

fasciolatus.

Fraser-Brunner (1951) stated that his O. cristatus

differed from all other Omobranchus species in having a crest on the head. He did mention, however,

that male O. striatus has such a crest (it is not

clear how he knew that Jatzow and Lenz's specimens included a male as they did not mention sex
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and their illustration is inadequate for determining

sex). He stated that O. cristatus differed from O.

striatus in proportions, fin-ray counts, and color

pattern. The fin-ray counts reported by Jatzow and

Lentz for O. striatus, however, are within the range

of those reported by Fraser-Brunner for O. cristatus.

Jatzow and Lenz reported the two proportions they

gave in terms of total length, and Fraser-Brunner

reported his proportions for the same characters in

terms of standard length. When this difference in

method is considered, the two sets of proportions

would be similar, if not identical. Finally, color

pattern in O. fasciolatus is quite variable, and we

have examined specimens that are similar in color

pattern to that described and illustrated by both

Jatzow and Lenz and Fraser-Brunner.

MATERIAL EXAMINED.—NORTHERN RED SEA:

SUEZ CANAL: Kabret, BMNH 1925.9.19.139 (47.1).

GULF OF SUEZ: Ras Misalla, HUJ Fish 303 (31.1);

Abu Rodees, HUJ F-4379 (39.6); Et Tur, HUJ

Fish 44 (33.9). SOUTHERN RED SEA: ETHIOPIA:

USNM 130834 (33.8); Massawa, USNM 204487

(44.5, neotype of Blennechis fasciolatus), 209265

(35.8), MCSN 32051 (39.1, holotype of Petroscirtes

vinciguerrae). DAHLAK ARCHIPELAGO: Cundabilu,

HUJ E62/44 (34.1); Entedebir, HUJ E62/4037

(40.2). GULF OF ADEN: SIRAH ISLAND: BMNH

1954.4.26.68 (38.0, holotype of Omobranchus cristatus Fraser-Brunner), 1954.4.26.69-74 (6: 26.5-

41.7). PERSIAN GULF: SAUDI ARABIA: Ras Tanura, USNM 147984 (9: 23.6-47.3). ZAAL ISLAND,

USNM 147980 (5: 27.4-34.0). ARABIAN SEA:

WEST PAKISTAN: Astola Island, USNM 201869 (11:

38.6-48.4). INDIA: Okha Point, USNM 201889

(38.1). KENYA: Mombasa, ANSP 53260 (40.0).

MOZAMBIQUE: Cape Delgado, USNM 197641

(44.7). MADAGASCAR: Nossi Be, USNM 201880

(11: 23.7-49.5), 209267 (32.2), 201890 (38.3), 201894

(2: 29.6-32.1). NO LOCALITY: BMNH 1878.9.26.7

(36.0).

Omobranchus ferox (Herre)

FIGURES 18, 29/

Petroscirtes ferox Herre, 1927:277 [60 syntypes from vicinity

of Ambulong, Talisay, and around Volcano Island, all in

Lake Taal; neotype designated below].

Petroscirtes kranjiensis Herre, 1940:25 [mangrove swamp

drained by Kranji River, Singapore Island].

Petroscirtes waterousi Herre, 1942:112 [swamp on Waterous

Hacienda, Mangarin, Mindoro, Philippine Islands].

Petroscirtes feliciana Herre, 1942:112 [mangrove swamp beside

Fishery Station at Cagayan, Oriental Misamis, Mindanao,

Philippine Islands].

Omobranchus dealmeida J. L. B. Smith, 1949:104 [rock pool,

Ponte Maone, Delagoa Bay].

DESCRIPTION (see also Tables 5 and 9).—Dorsal

fin XI-XIII (XII in 85.2% of specimens), 20-23 =

32-35; anal fin II, 22-26; both anal-fin spines of

males discernible externally; segmented caudal-fin

rays 13-15 (13 in 95.4% of specimens); dorsal +

ventral procurrent caudal-fin rays 10-15 (12-14 in

96.7% of specimens); vertebrae 10-11 (11 in only 1

of 75 specimens) + 28-30 = 38-40; epipleural ribs

10-13 (11-12 in 96.5% of specimens); prenasal

pores present; interorbital pores 2-3 (3 in 96.8% of

specimens); circumorbital pores 8-9 (8 in 96.8% of

specimens); lateral-line tubes 0-4; lateral-line tubes,

when present, extending posteriorly to below level

of dorsal-fin spine 2-7; gill opening extending ventrally to level of pectoral-fin ray 3-6; lower-lip flap

present; circumorbital bones 4; lower jaw teeth

16-22; upper jaw teeth 14-21 (Figure 37); no fleshy

bladelike crest on top of head of either sex.

COLOR PATTERN (available specimens all greatly

faded).—Males. Head: Two very faint, dusky bands

present below eye; distinctive, almost vertical, narrow, pale line, approximately two-thirds eye diameter in length, bordered posteriorly by intense dark

line of similar length, present along posteroventral

margin of orbit; lower dorsoanterior edge of pale

line also with very narrow, intensely dark, marginal

line. Dark dusky blotch apparent posteriorly on

operculum in some specimens.

Trunk: Up to 12 or 13 evenly spaced, moderately

narrow, dark dusky bands reaching almost from

dorsal to ventral body contour present in some

specimens (most specimens with color pattern almost totally faded); bands nearly vertical, though

apparently with slight anteriorly directed angle at

lateral midline of body; bands only about half as

broad as adjacent pale dusky interspaces; some

bands incomplete dorsally or dorsal ends separate

from rest of band. Row of paired dark blotches

present on dorsal body contour, each pair at dorsal

end of corresponding body band.

Pectoral Fin: Pale with even spread of dark

specks of pigment; very fine, threadlike, dark line

running length of each ray. Dusky blotch present

on dorsal half of fleshy pectoral-fin base.
NUMBER 177 39

TABLE 9.—Frequency distributions for certain characters in populations of

Omobranchus ferox

Population

Mozambique

Ceylon

India

Nias Island

Singapore

Gulf of Thailand

Hong Kong

Philippine Islands

Population

Mozambique

Ceylon

Nias Island

Singapore

Gulf of Thailand

Hong Kong

Philippine Islands

11

1

2

1

Dorsal-fin

spines

12 13

5 -

4

3 -

2

22 1

1

27 6

Pre caudal

vertebrae

10

1

5

3

2

25

1

33

11

-

_

-

-

1

X

10.0

10 0

10.0

10.0

10.0

10.0

X

12.0

12.0

12.0

12.0

12.1

Caudal

28 29

1 3

3

16

2 18

Segmented

dorsal-fin

20 21

1 4

2 1

1 18

9 19

vertebrae

30

1

1

_

2

9

1

14

X

29.0

30.0

29.0

30.0

29.4

29.4

22

1

-

3

-

2

6

1

6

rays

23 X

- 20.8

1 22.2

- 20.3

- 22.0

- 21.2

- 20.9

Total vertebrae

38

1

2

39 40

1

3 1

4

3 -

2

16 9

1

17 15

X

39.0

40 0

39.0

40.0

39.4

39.4

Total

32

1

2

1

7

dorsal-fin

elements

33

1

4

1

19

18

34 35

-

3 l

-

2 -

5 -

1

9 -

Lateral-line

0

•I

1

4•3

1

11

1

6

2 3 h

I ' -

l l -

1 - -

7 5 2

9 6 3

X

32.8

34.2

32.3

34.0

33.2

33-0

tubes

X

1.2

0 8

2-5

1.5

1.9

1.8

Segmented

anal-fin rays

22 23 24 25 26 5c

1

4 1 -

2 1

3 - -

2

1 13 11 -

1

2 22 10 -

Last lateral-line

below dorsal-fin

0 1 2 3 4 5 6

4 - - - 1

1 - •?___ _

1 1 - -

1 1 - - -

10 6 5 3 -

l

5 7 3 9 -

- 23.2

1 24.8

- 23.0

- 24.0

- 23-4

- 23.2

tube

spine

7 x

- 2.8

_ \ 5

- 3-5

- 2.5

1 3-2

- 3-7

Pelvic Fin: Pale with even spread of dark specks

of pigment proximally.

Dorsal Fin: Pale with scattered fine, dark spots.

Indication of some color pattern in the dorsal fin,

possibly 1 or 2 longitudinal stripes as described by

Herre for O. waterousi; however, not apparent on

best marked specimen available. Large, dark spot,

diffuse in some specimens, present distally between

last 3 or 4 segmented rays.

Anal Fin: Dark dusky, ray tips abruptly pale.

Caudal Fin: Pale, membranes transparent distally, dorsal and ventral edges dusky. Two broad,

longitudinal, dark dusky streaks present on fin, 1

just above and 1 just below lateral midline of fin.

Female: Apparently similar to males except for

absence of dark blotch on posterior distal portion

of dorsal fin.

GEOGRAPHIC VARIATION.—There is some evidence

(Table 9) that the Indian population of O. ferox

has greater numbers of meristic elements than do

the other populations. Rama Rao (1968) also reported a high number of segmented dorsal- and

anal-fin rays for his specimens from India.

GEOGRAPHIC DISTRIBUTION (Figure 4).—Philippine

Islands, South China Sea, west to Delagoa Bay,

Mozambique. The two specimens known from Mozambique (Smith, 1959) may possibly be the result

of artificial introductions similar to that we propose for O. punctatus (p. 63) from the same area.

The wide gap in distribution between the Mozambique locality and the nearest locality of occurrence

to the east (India), together with the estuarine

habitat of the species, furnishes circumstantial

evidence favoring a hypothesis of artificial introduction.

We have examined specimens of O. ferox from

India only from Vizgapatnam (about 17° 3C N) on

the Bengal coast. Rama Rao (1968) has reliably

reported and illustrated the species, as Cruantus

dealmeida, from Nizampatnam (above 16°00'N) on

the same coast.

HABITAT.—Shallow waters, most frequently collected in mangrove swamps and river estuaries; also

from a freshwater lake. One collection was recorded

as having come from oysters in a river.

COMPARISONS.—Omobranchus ferox differs from

all other species of Omobranchus except O. zebra,

with which it is broadly sympatric, in having 4 circumorbital bones (4 bones is a unilateral variant

in the other species). It differs from O. zebra most

obviously in having the gill opening extending ventrally to opposite the 3rd to 6th pectoral-fin ray
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(versus never extending ventral to level of dorsalmost ray), in lacking well-defined bands on the

head, and in having a shorter head (see key couplet

13).

Omobranchus ferox may be most readily separated from the other species of Omobranchus that

occupy the same general geographic area as follows:

from O. aurosplendidus and fasciolatoceps in lacking a fleshy crest on the head and in having the gill

opening extending much farther ventrally (restricted to area dorsal to dorsalmost pectoral-fin ray

in the other species); from O. meniscus in having

lateral-line tubes (versus lacking tubes), 20-23 segmented dorsal-fin rays (versus 19 rays), males without a fleshy crest, and no dark, crescentic marking

extending dorsally from the postorbital margin;

from O. punctatus in having the gill opening extending farther ventrally (never below level of

dorsalmost pectoral-fin ray in punctatus), and no

more than 4 lateral-line tubes (versus rarely less

than 4); from O. elongatus and germaini in not

having any distinct, dark markings on the underside of the head and on the body, and in the nature

of the dark spot behind the eye (compare Figure

29/ with Figures 29e and 13-15).

Males of O. ferox are unique in Omobranchus in

having a dark spot distally at the posterior end of

the dorsal fin (Figure 18).

NOMENCLATURAL DISCUSSION.—Herre (1927) described Petroscirtes ferox based on 60 syntypes from

Lake Taal (= Lake Bombon), Luzon, Philippine

Islands. We have been unable to locate any of the

specimens upon which his description was based.

Herre did not compare his species with any other,

and while the description and illustration best fit

the species here called O. ferox, the determination

is not unequivocal. For this reason, and in order

to stabilize the name of the taxon in question, we

here designate CAS SU67264, male, 38.1 mm SL,

Lake Bombon, collected by Herre, May 1931, as

neotype of Petroscirtes ferox. The neotype is one

of 19 specimens (14.5-41.7 mm SL) originally cataloged as CAS SU28446 (the remaining 18 specimens retain this number) under the name Petroscirtes ferox, and were presumably identified as such

by Herre. In the description of P. ferox, Herre

stated that the total number of dorsal-fin elements

ranged from 32-36, which is one element more than

we have found for the species and two more than

we found among the 34 specimens available from

the Philippine Islands.

Herre (1940) did not compare his Petroscirtes

kranjiensis with any other species. We have examined the holotype and paratype and find them to

be specimens of O. ferox.

Herre (1942) described two more species, P.

waterousi and P. feliciana, also synonymous with

P. ferox. His comparison of these two species is

contained in a key to the Philippine species of

Petroscirtes, which included several species in the

blenniid tribe Nemophini and the following nominal species of Omobranchus: ferox, waterousi,

kallosoma (= elongatus), feliciana and loxias

(= rotundiceps obliquus). His first major separation

in the key was only ecologically based: freshwater

species versus marine species. His P. ferox keyed

out as the only freshwater species. All the remaining Omobranchus species and two nemophinine

species, P. bankanensis and P. eretes, fall under

couplets H and HH of his key. Petroscirtes waterousi is segregated in couplet H, which reads "no

cross bands or stripes on body," but in couplet I

leading to waterousi, he stated contradictorily:

". . . with faint indications of a dozen narrow crossbars. . . ." Later, in couplet LL leading to P. feliciana, he stated: ".. . 12 angulate or vertical dusky

brown crossbars." He, therefore, did not differentiate his species, and we can find no reason to

recognize them.

Smith (1949) did not compare his O. dealmeida

with any other species. We have examined the holotype and find it to be a specimen of O. ferox.

MATERIAL EXAMINED.—MOZAMBIQUE: DELAGOA BAY: Ponte Maone, RUSI 233 (44.1, holotype

of Omobranchus dealmeida). CEYLON: Trincomalee, USNM 209268 (5: 39.0-49.8). INDIA: Vizagapatnam, CAS SU67265 (4: 29.2-36.5). INDONESIA:

NIAS ISLAND: USNM 201559 (3: 30.4-34.5, including one cleared and stained). SINGAPORE: CAS

SU33007 (42.4, holotype of Petroscirtes kranjiensis),

SU33008 (40.5). THAILAND: northeast coast Gulf

of Thailand, USNM 119663 (ca. 45), CAS GVF reg.

no. 1486 (23: 35.7-ca. 49), GVF reg. no. 2176 (38.8).

HONG KONG: CAS SU62066 (42.6). PHILIPPINE

ISLANDS: Manila fish market, USNM 99372 (4:

53.1-55.7), 122383 (54.6). PALAWAN: USNM 122382

(37.9). LUZON: Pangasinan Province, USNM 209345

(34.4), 209346 (8: 36.8-58.3); Batangas Province,
NUMBER 177 41

CAS SU28446 (18: 14.8-41.7), SU67264 (38.1, neotype of Petroscirtes ferox). MINDANAO: CAS SU36671

(54.7, holotype of Petroscirtes feliciana), SU36672

(46.1). MINDORO: CAS SU36673 (38.4, holotype of

Petroscirtes waterousi).

Omobranchus germaini (Sauvage)

FIGURES 19, 20, 29c

Petroscirtes germaini Sauvage, 1883:158 [Nouvelle-Caledonie].

Graviceps alexanderi Whitley, 1945:33 [Fremantle, Western

Australia, living in holes in wharf piles].

DESCRIPTION (see also Tables 5 and 10).—Dorsal

fin XII-XIV (XIII in 84.6% of specimens), 18-23 =

31-36; anal fin II, 21-26; both anal-fin spines of

males discernible externally; segmented caudal-fin

rays 13-14 (13 in 98.8% of specimens); dorsal +

ventral procurrent caudal-fin rays 11-18 (12-16 in

97.5% of specimens); vertebrae 10-11 + 28-32 =

38-43; epipleural ribs 11-15 (12-14 in 97.4% of

specimens); prenasal pores present; interorbital

pores 2-5 (3 in 96.7% of specimens); circumorbital

pores 7-10 (modally 8 or 9); lateral-line tubes 0-8

(0 in only 1 of 243 specimens); lateral-line tubes,

when present, extending posteriorly to below level

of dorsal-fin spine 1-10; gill opening varying from

restricted to area dorsal to level of dorsalmost

pectoral-fin ray to extending ventrally to opposite

6th ray (extending below level of 4th ray in only 3

of 230 specimens); lower-lip flap present; circumorbital bones 4-5 (of 7 specimens examined all had

5 except 1 specimen with 4 on one side and 5 on

the other; the 4 bones resulted from a fusion of the

posteriormost 2 circumorbials); lower jaw teeth 16-

28 (Figure 48); upper jaw teeth 14-25; no fleshy

bladelike crest on top of head of either sex.

COLOR PATTERN.—Males: Head: Ventral half to

one-third with about 4 irregular, moderate to

broad, evenly spaced bands; anteriormost band

joining corresponding band from opposite side

across ventral surface; remaining 3 bands variable,

with none to all bands joining corresponding band

of opposite side across ventral surface. Anteriormost

band extends from chin to anterior edge of orbit;

remaining 3 bands originate ventral and posterior

to ventral extremity of orbit. Pale spaces between

bands slightly broader than adjacent bands; spaces

divided by vertical series of narrow to moderately

broad dashes or broken lines. Ventral surface with

irregular transverse series of uninterrupted and

interrupted dark bands. One or more irregularly

positioned, short, broad, dark lines posteriorly in

branchiostegal region. Dorsoanteriorly darkened,

dome-shaped area posterior to eye between circumorbital and preopercular series of cephalic sensory

pores; vertically elongate, intense, black spot, about

as broad as pupil diameter, at level of pupil just

inside anterior margin of dome-shaped area. Operculum with large, dorsoanteriorly inclined, dusky

area enclosed by pale border; border sometimes

incomplete posteroventrally; dark blotch sometimes

present above pale border. Short, dark band just

anterior to gill opening. Trunk portion of gill

opening darkly pigmented. Head and nape uniformly dusky dorsally.

Trunk: Unpaired dark band in area covered by

appressed pectoral fin followed posteriorly by 7 or

8 pairs of dark, equally broad bands. Bands slant

posteroventrally below lateral midline; unpaired

band and first 2 band pairs slant anterodorsally

above midline; remaining bands slant posterodorsally above midline. Bands usually well developed

ventrally, occasionally faint to absent; bands broken

into dark mottling or spotting on dorsal quarter of

body. Posterior bands often poorly developed with

posteriormost band usually represented as elongate

midlateral spot. Each band with intensification of

pigment midlaterally, sometimes continuing ventrally as medial sliver of dark pigment (intensifications appear as series of paired spots in faded specimens). Unpaired band and band pairs encircled by

narrow, pale margins, paler than interspaces separating band pairs; interspaces between band pairs

broader than individual bands. Narrow, dark stripe

on body along dorsal-fin base, sometimes broken

into series of spots in faded specimens.

Pectoral Fin: Pale with uniform spread of dark

specks of pigment. Fleshy pectoral-fin base crossed

by 1-3 dark or dusky bars; anteriormost bar darkest, occasionally extending onto head.

Pelvic Fin: Pale to slightly dusky.

Dorsal Fin: Pale with series of dark, irregular

stripes; stripes broad, inclined anterodorsally on

anteror portion of fin, less defined, narrower, inclined dorsoposteriorly on posterior portion. Posterior 3rd of dorsal fin uniformly dusky. Some specimens with a series of more-or-less joined, intensified,

black blotches, usually extending from 8th to 11th

segmented ray (variable in range between 7th and
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 10.—Frequency distributions for certain characters in populations of

Omobranchus gennaini

Population

Taiwan

Hong Kong

Philippine Islands

Singapore

Ambon

A istralia

Queensland

Capricorn Islands

Swains Islands

Little Hope Island

Arnhemland

Western Australia

Montebello

Northwest Cape

Point Cloates

Maud Landing

Shark Bay

Port Denison

Rottnest Island

New Caledonia

Population

Taiwan

Hong Kong

Philippine Islands

Singapore

Ambon

Australia

Queensland

Capricorn Islands

Swains Islands

Little Hope Island

Arnhe mland

Western Australia

Monte bello

Northwest Cape

Point Cloates

Maud Landing

Shark Bay

Port Denison

Rottnest Island

New Caledonia

Population

Taiwan

Hong Kong

Philippine Islands

Singapore

Ambon

Australia

Queensland

Dorsal-fin

spines

12 13 14

1 7 -

2 15 1

3 -

1

3 -

4 49 3

2 -

12 1

6 -

1 1 -

2

7 12 -

7 18 -

4 40 4

1 13 1

29 -

2 1

Precaudal

vertebrae

10

8

16

3

1

3

55

2

1C

5

2

1

12

14

9

6

12

2

Capricorn Islands

Swains Islands

Little Hope Island

Amhemland

Western Australia

Monte bello

Northwest Cape

Point Cloates

Maud Landing

Shark Bay

Port Denison

Rottnest Island

New Caledonia

11 X

- 10.0

2 10.1

- 10.0

- 10.0

1 10.0

- 10.0

2 10.2

1 10.2

- 10.0

1 10.5

7 10.4

11 10.4

39 10.8

9 10.6

17 10.6

- 10.0

2

12.9

12.9

13.0

13.0

13.0

13.0

13-1

13.0

12.5

13.0

12.6

12.7

13-0

13.0

13.0

13.3

28

7

2

2

h

18

4

2

Segmented Dorsal-fin

rays

19

1

14

3

3

2

4

2

2

20

7

-

-

1

-

30

5

-

1

4

5

19

21 22

-

-

-

24 1

2

8 -

-

2

1

12 3

15 3

21 6

3 7

9 20

3

Caudal vertebrae

29

7

10

1

1

1

8

2

2

4

3

10

Lateral-line

0 1 2

1

1

2

6 15

1

1 4

1

2

1

1 2 4

1 1

3

2

4

-

-

8

1

1

1

4

3

2

7

4

1

6

-

-

14

3

2

2

7

5

15

4

6

5

4

2

-

1

1

9

3

5

_

_

7

7

14

4

5

30

_

-

-

-

41

7

-

1

1

8

17

25

1

4

31

_

-

-

-

6

2

3

-

1

1

6

5

10

7

20

1

tubes

6

1

3

2

-

2

1

_

6

12

3

3

7

1

-

-

-

-

_

2

3

32

_

-

-

-

-

-

-

-

-

-

1

-

2

7

5

1

8

-

-

-

-

•

m

1

X

29.0

28.6

28.3

28.3

30.0

3l.o

30.1

28.3

30.5

30.5

30.2

30.1

30.1

31.4

31.0

31.5

X

4.5

4.3

4.7

3-0

3.2

2.5

3.5

4.7

4 . 0

4 . 0

4.1

4.7

5.0

4.6

3-5

1.5

23

-

-

-

_

-

-

-

-

-

-

-

5

-

38

6

2

2

3

0 1

1

1 -

X

19.

18.

19.

19.

2 0 .

2 1 .

2 0 .

9

8

0

0

4

0

6

18.7

2 1 .

2 1 .

2 0 .

2 0 .

2 0 .

2 2.

2 1 .

2 2 .

0

0

9

8

6

1

7

0

Total

39

7

10

1

1

1

9

1

3

l

40

_

1

-

-

40

7

-

1

1

10

11

15

31

6

2

Total Dorsal-fin

elements

32

2

11

3

l

3

4

4

2

2

vertebrae

41

_

-

-

-

8

2

4

-

1

1

8

12

23

4

13

1

1*2

_

-

-

-

-

_

-

-

-

-

-

1

8

7

14

1

33

6

1

-

-

27

4

-

1

1

9

9

16

43

_

-

-

-

_

_

-

_

_

1

_

1

4

2

-

Last lateral-line

2

4

-

1

2

1

3

1

1

1

12

1

4

1

34

-

-

-

24

2

9

-

1

.

9

14

27

4

9

2

X

39.0

38.7

38.3

38.3

40.0

41.0

40.2

38.5

40.5

40.5

40.6

40.5

40.9

42.0

41.6

41.5

tube

dorsal-fin spine

4

1

3

1

7

1

2

1

1

5

5

1

4

_

-

5

1

1

i.

3

4

4

6

1

6

4

3

-

6

1

1

2

3

1

3

3

3

7

1

2

_

1

1

13

2

1

4

2

10

2

4

1

8

1

3

2

-

6

4

2

"

6

7

21

7

7

35

-

-

-

1

-

-

-

l

1

.

3

5

20

1

36 X

- 32

- 31

- 32

- 32

- 33

- 34

- 33

- 31

- 33

- 33

- 33

- 33

6 35

- 34

- 34

8

7

0

0

4

0

7

7

5

0

6

5

6

l

7

3

Circumorbital

7 8

6

14

3

1

3

1 53

2

13

1

9

2

4

-

_

2

_

5

2

1

5 14

4 19

8 39

2 3

1 13 1

1

Delow

9

1

_

-

1

-

1

6

10 X

_ 6.C

- 5-9

- 6.3

- 4.7

- 5-2

- 4.5

- 5.4

- 6.5

5«5

- 8.5

- 6.6

4 7.5

- 7-3

- 7.3

- 5-6

- 4.7

I

1

10

-

_

_

_

1

_

_

_

2

1

1

-

21

2

Segmented Anal-fin

rays

22

1

12

3

1

3

2

5

1

pores

X

8.2

8 . 2

8 . 0

8 . 0

8 . 0

8 . 0

8 . 0

9.2

9.0

8.5

8.7

8.9

8.8

8.6

£5

8.5

Lower

23

7

4

-

-

27

4

1

6

7

16

11

1

1

1

1

end

24

-

-

-

26

2

9

-

2

1

11

14

30

Jl

2

25

-

-

-

3

_

-

_

1

2

3

l

8

12

1

26

-

-

_

_

_

_

_

_

_

_

1

4

3

Bpipleural

12

7

1

1

13

2

3

1

1

7

8

3

of

13

8

6

2

1

2

36

1

8

3

_

2

13

12

33

ll

1

14

1

_

7

1

_

_

2

5

6

4

14

1

X

22^9

22.1

22.0

22.0

23-5

24.0

23-7

22.2

24.0

24.5

23.8

23.8

23.7

2S. 1

24.6

24.3

ribs

15

_

_

_

1

_

_

_

_

-

ill l opening

opposite pectoral-fin

C

5

5

4

3

2

1

1

11

5

1

1

2

2

5

9

3

9

1

2

5

5

24

1

9

9

18

2

7

3

3

6

2

1

1

1

7

6

11

2

7

4 5

- 1

5 _

1 _

2 -

- -

- -

1 1 1 rH 1 1 1 11 i-H CVJ lf\ CVI rH 1 1

6

.

l

-

-

ray

X

2 . 7

2.9

3.3

3-7

1.6

0.6

0.7

O ft

2»U

2.4

2.0

2.3

1.9

2.0

X

13.0

12.5

12.T

12.7

12.9

13.5

12.8

12.5

11.5

13.0

13.1

12.8

12.9

13.1

13.*

13.5
NUMBER 177 43

13th rays); others with single large, dusky to intense

dark spot in same area. Distal margin of fin with

narrow white edge.

Anal Fin: Dusky, slightly darker distally with

narrow, white, distal margin. Small, poorly denned,

dark spot on membrane at fin base posterior to

every 2nd ray.

Caudal Fin: Uniformly pale. Caudal peduncle

with 2 moderately sized spots at fin base, 1 just

above and 1 just below lateral midline of body;

pigment streak from each spot continued posteriorly

onto fin.

Females: Color pattern as in males but considerably less intense; body bands very narrow, ventral

portions faint to absent. Fin pigment usually faint;

spot or spots between 7th and 13th segmented

dorsal-fin rays faint or absent.

Geographic Variation: Taiwan: Dark pigment

on dorsal portion of body generally better developed; lines and marks in pale interspaces above

lateral midline of body intensified, producing mottled appearance on trunk and posterior body. Dark

blotches present on top of head and nape. Females

with better development of spots (normally characteristic of males) in region of 7th to 13th dorsal-fin

rays. Ventral edge of caudal fin occasionally dusky.

Hong Kong and Northern and Western Australia:

Trunk and posterior body mottled dorsally, similar

to Taiwan specimens. Anterior end of dorsal fin

with narrow, anterodorsally inclined, dark stripes.

GEOGRAPHIC VARIATION (see also color pattern).—

Population variation was noted in most meristic

characters of O. germaini (Table 10). In general,

the Australian populations have the highest average numbers of elements for each character, but

within Australia most of the Arhnemland population's averages are noticeably low and closer to

those of the extra-Australian populations.

The Western Australian populations exhibit increases in numbers of precaudal, caudal, and total

vertebrae. Similar increases are exhibited by the

Western Australian populations of O. rotundiceps,

whereas the Western Australian populations of O.

punctatus exhibit an increase in the number of

precaudal vertebrae, but not caudal or total vertebrae. Increase in number of precaudal vertebrae

was not encountered in any other species of Omobranchus. It is noteworthy that three species of

Omobranchns should each exhibit increases in

number of precaudal vertebrae in about the same

geographic area, even if presently inexplicable.

Although covariance comparisons of numbers of

jaw teeth (Table 11) were not feasible for all populations of O. germaini, it was found that significant

differences in numbers of lower teeth occurred between females of the Hong Kong and southwestern

Australian (Shark Bay to Perth) populations and

between the southwestern Australian and Queensland populations. For upper jaw teeth, significant

differences occurred between females of the Hong

Kong and northwestern Australian (Montebello to

Maud), between Hong Kong and southwestern Australian, and between the southwestern Australian

and Queensland populations; the comparisons between the northwestern Australian and Queensland

populations were almost significant.

GEOGRAPHIC DISTRIBUTION (Figure 6).—Western

Pacific Ocean from northern Taiwan south to the

Tropic of Capricorn on the Australian east coast

and Perth on the Australian west coast; New

Caledonia.

HABITAT.—Found in shallow coastal waters and

tide pools around rocks and coral, usually at normal seawater salinities. At least 1 collection from

rocks at the mouth of a small stream. Usually in

calm waters but common on the open-ocean (highenergy) side of One Tree Island in areas inundated

at high tide and subjected to the action of breakers.

COMPARISONS.—Omobranchus germaini is a member of the elongatus species group (see "Comparisons" under O. elongatus). Males and sometimes

females of O. germaini are distinguishable from all

other Omobranchus species by the mosaiclike pattern of dark spots and bars that is often present

on the underside of the head (Figure 29e). Within

the elongatus group, O. germaini is most similar to

O. loxozonus, which is allopatric. The color pattern on the head and body of O. germaini is usually

more contrasty and the number of body bands is

usually larger than in O. loxozonus. Where populations of the two species are geographically closest,

there are marked differences in numbers of fin rays,

vertebrae, and lateral-line tubes (see Tables 5, 10,

12), although overlaps in the ranges for these characters exist.

NOMENCLATURAL DISCUSSION.—Whitley (1945) did

not compare his G. alexanderi with O. germaini.

We have not seen his holotype (WAM P.671), but

based on his report of the number of dorsal-fin

spines (14), total number of dorsal-fin elements
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 11.—F values for covariance comparisons of regression equations

(Table 2) for number of lower or upper jaw teeth between males or

females from different populations of Omobranchus germaini (*• —

significant at p = .001; NS = not significant)

Comparisons

Lower teeth

males

hong Kong X Queensland

Western Australia

Montebello-Maud X Hong Kong

Shark Bay-Perth X Hong Kone

Montebello-Maud X Queensland

Shark Bay-Perth X Queensland

Montebello-Maud X Shark Bay-Perth

females

Hong Kong X Queensland

Western Australia

Montebello-Maud X Hong Kong

Shark Bay-Perth X Hong Kong

Montebello-Maud X Queensland

Shark Bay-Perth X Queensland

Montebello-Maud X Shark Bay-Perth

Upper teeth

males

Hong Kong X Queensland

Western Australia

Montebello-Maud X Hon^ Kong

bhark Bay-Perth X Hong Kong

Kontebello-Maud X Queensland

Shark Bay-Perth X Queensland

Montebello-Maud X Shark Bay-Perth

females

hong Kong X Queensland

Western Australia

Montebello-Maud X Hong Kong

Shark Bay-Perth X Hong Kong

Montebello-Maud X Queensland

Snark Bay-Perth X wueerjsland

Montebello-Maud X Shark Bay-Perth

F values

slopas

0.06 NS

0.23 NS

1.95 NS

0.05 NS

2.33 NS

1.89 NS

4.18 NS

0.06 NS

0.75 NS

3.90 NS

1.51 NS

0.42 NS

0.00 NS

0.19 NS

0.86 NS

C.I8 NS

1.04 NS

1.89 NS

0.23 NS

2.43 NS

0.01 NS

5.18 NS

0.43 NS

2.65 NS

Degrees of

freedom

1/41

1/23

1/19

1/48

1/44

1/26

1/24

1/22

1/23

1/38

1/39

1/37

1/41

1/24

1/19

1/49

1/44

1/27

1/25

1/23

1/23

1/38

1/38

1/36

F values

heights

0.27 NS

2.22 NS

1.23 NS

0.69 NS

0.10 NS

0.15 NS

0.03 NS

4.62 NS

14.89 **

5.07 NS

38.03 «*

5.10 NS

0.17 NS

0.11 NS

2.77 NS

0.00 NS

0.91 NS

1.47 NS

1.31 NS

30.58 **

31.89 **

10.97 NS

15.54 **

0.30 NS

Degrees of

freedom

1/42

1/24

1/20

1/49

1/45

1/27

1/25

1/23

1/24

1/39

1/40

1/38

1/42

1/25

1/20

1/50

1/45

1/28

1/26

1/24

1/24

1/39

1/39

1/37

(33), segmented anal-fin rays (25), the presence of a

crescent-shaped blue spot close behind the eye, and

the type-locality (Freemantle, Western Australia),

we believe that the holotype could only be O. germaini. In addition, all specimens of O. germaini

sent to us from the Western Australian Museum

were identified as O. alexanderi, and were presumably so identified based on knowledge of Whitley's

holotype. (These specimens were part of the material treated in an unpublished master's thesis by

N. Milward, University of Western Australia, on

the systematics and biology of Western Australian

Blenniidae, Tripterygiidae, and Clinidae, 1962.)

MATERIAL EXAMINED.—TAIWAN: USNM 204027

(5: 44.7-50.4), 208479 (2: 40.1-42.8), 208480 (58.3).

HONG KONG: CAS GVF reg. no. 1730 (2: 37.0-

38.4), GVF reg. no. 1723 (5: 25.1-36.7), SU62062

(8: 23.8-47.5), USNM 197977 (2: ca. 25-33), 197983

(38.0). PHILIPPINE ISLANDS: CEBU: USNM

207270 (3: 25.7-35.3). SINGAPORE: CAS SU32887

(29.7). INDONESIA: AMBON: USNM 209531 (3:

30.3-37.1). AUSTRALIA: GULF OF CARPENTARIA

(Arnhemland): USNM 174338 (34.2), 174342 (2:

41.7-44.0), 174343 (2: 29.3-32.9), 207959 (31.4).

GREAT BARRIER REEF: Little Hope Island, ANSP

109700 (12: 16.1-51.3), 109701 (29.4); Swain Reefs,

AMS IB.6220-1 (2: 42.6-ca. 45); One Tree Island,

USNM 208475 (29: 32.2-54.0), 208476 (ca. 40),

208478 (2: ca. 34-37.4), CAS 13780 (20: 37.8-65.4);

Heron Island, BPBM 15129 (4: 30.1-50.2), USNM

208477 (2: 29.4-36.1). WESTERN AUSTRALIA: Montebello Island, BMNH 1961.8.16.76 (31.7), 1961.8.

16.78 (43.2); North West Cape, WAM P4837 (2:

41.0-ca. 45); Point Cloates, WAM P10720-36 (19:

28.5-59.4); Maud Landing, WAM P7735-59 (25:

32.4-47.5); Shark Bay, WAM P7257-7306 (48:

22.9-42.8) Port Denison, WAM P4836 (6: ca. 37-

48), P10403-11 (9: 16.2-38.8); Rottnest Island,

WAM P4779 (2: 42.4-49.7), P6296 (45.7), P10115-

22 (8: 32.6-42.6), P10123-28 (6: ca. 36-40.5).
NUMBER 177 45

P10393-402 (11: 17.9-46.5); near Perth, NFIS 9635

(51.6). NEW CALEDONIA: MNHN A-4891 (45.5,

holotype of Petroscirtes germaini); Noumea, CAS

SU47017 (36.2), AMS IB.2239 (41.6).

Omobranchus lineolatus (Kner)

FIGURES 21, 22

Petroscirtes lineolatus Kner, 1868a:29; 1868b:331 [Candavu].

Graviceps darwini Whitley, 1958:47 [Port Darwin. Northern

Territory of Australia].

Description (see also Table 5).—Dorsal fin XI,

19-21 = 30-32; anal fin II, 20-23; both anal-fin

spines of males discernible externally; segmented

caudal-fin rays 13-14; dorsal + ventral procurrent

caudal-fin rays 11-14; vertebrae 10 + 26-29 = 36-

39; epipleural ribs 11-13; prenasal pores present;

interorbital pores 3; circumorbital pores 8; lateralline tubes 4-8; lateral-line tubes extending posteriorly to below level of dorsal-fin spine 6-11; gill

opening restricted to area dorsal to level of dorsalmost pectoral-fin ray; lower-lip flap present; circumorbital bones 5; lower jaw teeth 22-30; upper jaw

teeth 20-27 (Figure 44); no fleshy bladelike crest

on top of head of either sex.

COLOR PATTERN.—Head: Pale with 5 dark bands

approximately half width of pale interspaces. Anteriormost band extends from anteroventral margin

of orbit across mouth, just anterior to angle of

jaws, onto chin; 2nd band extends ventrally from

eye and is deflected posteriorly on underside of

head; 3rd band extends ventrally from posteroventral edge of eye to ventral portion of head; anterior

3 bands meet corresponding bands of opposite side

at ventral midline of head; 4th band originates

laterally on nape, curves ventroanteriorly, then

posteriorly, and extends to point just above anterior end of pelvic girdle; 5th band extends ventrally

from near upper end of gill opening to uppermost

branchiostegal membrane fold. Each pale interspace contains a medial, vertical series of about 3-5

small, dark spots. A large, dark, ovoid spot, slightly

smaller than eye, present posterior to eye midway

between dorsal origins of 3rd and 4th cephalic dark

bands (illustrated specimen, Figure 21a, has spot

on left side slightly darker along anterior edge,

resembling a posteriorly opening crescent; spot on

right side is uniformly dark); spot sandwiched between small, dark, longitudinal dorsal mark and

ventral dark blotch that is confluent with upper

end of 3rd band. Dark, tear-shaped mark directed

posteriorly from dorsoposterior edge of eye. Series

of irregularly shaped, well-defined, dark spots less

than half pupil diameter along top of head posterior to anterior nostrils. Smaller, less defined, dark

blotches and spots extend posteriorly along nape

and sides of body below anterior end of dorsal fin.

Trunk: Pale with about 9 or 10 pairs of darker,

irregularly vertical bands originating along dorsalfin base and fading out on approximately ventral

quarter of body; anteriorly, banding less defined

and paired nature less apparent.

Pectoral Fin: Pale with broad, dark band along

base (illustrated specimen, Figure 21a, had band

broken into small dorsal spot and large ventral

blotch on left side of body). Several small, dark

freckles present on fleshy pectoral-fin base and basal

part of rays.

Pelvic Fins: Uniformly pale.

Dorsal and Anal Fins: Uniformly dusky with

several irregularly placed pale spots, each slightly

smaller than pupil of eye; pale spots noticeable

only in freshest specimen. Series of dark lines in

dorsal fin; each line extending from base of a

dorsal-fin ray, along posterior edge of ray to fin

margin.

Caudal Fin: Dusky with numerous small, pale

spots. Series of 4 short, dark stripes on basal portion of fin just posterior to caudal peduncle; series

appears superficially as broad bar. Narrow, longitudinal lines of darker pigment present posteriorly.

The largest specimen available (female, Figure

22) differs from the above description in having the

cephalic bands very faint laterally on the head and

the spotting in the cheek region intensified into a

densely speckled pattern. The dark lines in the

dorsal fin are directed slightly more diagonally and

cross several succeeding rays, a condition similar to

that shown in Kner's (1868b) illustration of Petroscirtes lineolatus (Figure 216). The dark lines in the

anal fin illustrated by Kner are not present in any

of our specimens.

There was no noticeable difference in color pattern between males and females.

GEOGRAPHICAL DISTRIBUTION (Figure 6).—Northern and Western Australia and southern New

Guinea. The holotype of O. lineolatus was reported

to have come from Candavu, Fiji Islands. As noted

in the "Nomenclatural Discussion," we question the
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

validity of this locality record (also see p. 63 under

O. punctatus).

Weber (1913, fig. 114; the same illustration appears in de Beaufort and Chapman, 1951, fig. 53,

and we reproduce it in our Figure 3a) illustrated a

planktonic larva that he listed as Petroscirtes spec,

juv. from Laiwui, Obi, Indonesia (Siboga station

142). He mentioned only one specimen, but there

are three, 12.1-13.0 mm SL, in ZMA 109.362 from

the same station and locality. Weber's illustration

is representative of these specimens, particularly in

the preopercular spination. The bony projections

anterior to each dorsal-fin ray element except the

anteriormost, which lacks a projection, that Weber

portrays are actually external projections of the

dorsal-fin pterygiophores. The specimens definitely

belong in the tribe Omobranchini (as evidenced

by their having only two segmented pelvic-fin rays

and hinged dentaries), but it is not possible to

identify them with certainty. Based on the following

meristics obtained from a radiograph, they could be

O. lineolatus or Laiphognathus multimaculatus;

dorsal fin XI, 20-21; anal fin II, 20-21 (or 21-22,

uncertain), vertebrae 10 + 27-28.

HABITAT.—Little known; taken once from rocky

ironstone reef in "salt" water at a depth up to 1

meter; once from rock holes on a reef.

COMPARISON.—Omobranchus lineolatus is separable from most of the other species of Omobranchus only by a combination of characters. It differs

from all the other species of Omobranchus except

O. aurosplendidus in having modally 11 dorsal-fin

spines (all populations of the other species have

modally 12 or 13 spines). It is prominently different

from O. aurosplendidus in lacking a fleshy crest

and in having fewer dorsal- and anal-fin elements

and vertebrae. In small specimens of O. lineolatus,

the color pattern of the head (Figure 21) renders

the species instantly recognizable. It is most likely

to be confused with members of the elongatus

species group, but only O. rotundiceps of that group

ever has as few as 11 dorsal-fin spines. In having

the gill opening restricted to the area dorsal to the

level of the dorsalmost pectoral-fin ray, O. lineolatus could only be confused with the nominal

subspecies, O. r. rotundiceps, which infrequently

has more than 3 lateral-line tubes, whereas O. lineolatus usually has more than 4.

NOMENCLATURAL DISCUSSION.—Kner (1868a)

briefly described P. lineolatus from Candavu (in

the Fiji Islands) without noting the number of

specimens he was treating. He accorded the species

a number (catalog?), 834, and stated that the species resembled Salarias semilineatus Kner in color

pattern. Kner (1868b) again described P. lineolatus

as a new species from Candavu based on a single

specimen, which he accorded the number 834d (we

have been unable to determine the allocation of

834a-c). Kner (1868b) was merely expanding on

Kner (1868a), as all the new species in 1868a were

redescribed in more detail and as new in 1868b.

In the second description all the characters of the

first description were repeated except for the color

pattern, which was described entirely differently.

At the end of the second description he noted that

there was a second specimen from Candavu, number 834e, that differed from the first in several

characters, including those repeated in the first and

second descriptions of P. lineolatus, and particularly

in color pattern, in which the second specimen was

said to resemble Salarias semilineatus. Kner concluded his description of the second specimen stating, "Wenn dieses Exemplar als neue Art gel ten

kann, so diirste die Benennung P. semilineatus nicht

unpassend erschienen."

As a result of the information thus far presented,

we believe that Kner's first description of P. lineolatus was based on at least (and possibly only) two

specimens (syntypes): one served as the basis for

the counts and proportions and the other for the

color pattern. In the second description Kner effectively restricted the name P. lineolatus to the specimen that provided the counts and proportions (this

specimen is also the holotype of the second P. lineolatus description, and the second P. lineolatus is a

junior objective primary homonym of the first P.

lineolatus).

Dr. P. Kahsbauer and one of us (VGS) have

searched the collection of the Vienna museum and

have been unable to locate the holotype of the second P. lineolatus. However, the description and

illustration provided in the second account clearly

define a taxon applicable to certain specimens we

have examined, and P. lineolatus is the oldest name

for that taxon. The nomenclatural situation is complicated, however, by other circumstances. First,

we believe that one of the syntypes of P. lineolatus

(1868a) is extant. We have found a specimen labeled as Petroscirtes semilineatus in the Vienna

museum, NMV 12561, from Candavu, that closely
NUMBER 177 47

agrees with Kner's description of P. semilineatus

(but there is no indication of the number 834e with

the specimen). Furthermore, this specimen is identifiable as O. punctatus and is the only specimen

of that species known from Candavu. If we are correct, this specimen also represents the holotype of

P. semilineatus, for Kner's description constituted a

valid species description under Article 17 (8) of the

International Code of Zoological Nomenclature.

Even if our interpretation of the Code is erroneous,

Giinther's (1877) first usage of P. semilineatus as a

senior synonym made the name available.

Secondly, if our interpretation of Kner's second

description of P. lineolatus as constituting a restriction of that name is not nomenclaturally acceptable, the only remaining syntype (the holotype of

P. semilineatus) might be designated as the lectotype of P. lineolatus (1868a). Such an action would

result in P. semilineatus' becoming a junior objective synonym of P. lineolatus, and P. lineolatus

would then be a junior synonym of O. punctatus.

The net result would be the loss of a name, P.

lineolatus, that on other grounds would be the

senior synonym of another taxon. In itself this

would not be bad (there is a junior synonym available, Graviceps darwini, for that taxon), but P.

lineolatus has been used several times as a senior

synonym of a recognizable species (Giinther, 1877;

Whitley, 1927, 1958; Fowler, 1928, 1959; Norman,

1943; Springer, 1972a). The designation of the extant syntype would also subvert Kner's obvious

intensions with regard to P. lineolatus.

We therefore believe that the designation of a

neotype for the first description of P. lineolatus is

desirable. To do so requires us to arbitrarily assume

that the description was based either on only one

specimen that was lost, or on more than one specimen, all of which are lost, because the Code (Article 75) predicates neotype designation on the fact

that no holotype, syntype, or lectotype exists. To circumvent future controversy, a neotype is also

needed for the second description of P. lineolatus,

to replace the missing holotype. To avoid confusion, the same specimen should be designated

neotype for both these descriptions. Before taking

these actions further comment is necessary.

Both P. lineolatus and P. semilineatus were based

on specimens reportedly from Candavu. Neither of

the taxa we allocate to these names has since (or

previously) been recorded from or near Candavu,

albeit there are very few specimens of the first

taxon known altogether (the second taxon is a

commonly collected species). Aside from the lack of

additional specimens from Candavu, we suspect

that the locality is in error because only one species

of Omobranchus, O. rotundiceps, is definitely

known to occur so far east in the western Pacific

Ocean, although numerous collections of fishes

from Fiji and areas to the east have been made,

especially in recent years. Also, there is a possibility

that the collector of the specimens (source not furnished with the descriptions) made an error. According to Kner (1868a, b), the specimens on which

he reported were sent to the Museum Godeffroy by

various people, including a large number of boat

captains. Locality records for specimens obtained by

nonprofessional biologists, especially those of over

a century ago, have often been suspect.

The Code, Article 75 (c) (5), requires that a neotype come as nearly as practicable from the original

type-locality. Of the specimens we treat as O. lineolatus, the one that comes nearest to being from

Candavu is the one contained in USNM 174335,

from Port Bradshaw, Northern Territory, Australia.

This specimen has almost completely lost the distinctive color pattern of the species, and it does

not, therefore, appear to be a good choice for a

neotype. The holotype of Graviceps darwini Whitley, AMS IA.4298, from nearby Port Darwin, Northern Territory, Australia, is in excellent condition,

and the color pattern is still clearly represented.

We therefore designate the holotype of G. darwini

as the neotype of Petroscirtes lineolatus Kner,

1868a, and Petroscirtes lineolatus Kner, 1868b. In

so doing, G. darwini becomes a junior objective

synonym of P. lineolatus.

Whitley (1958) compared his G. darwini with

P. lineolatus, which he believed different based on

slight differences in color pattern shown by the

illustration of P. lineolatus as compared with his

specimen of G. darwini. We are more impressed

with the similarities than the differences. The number of dark spots on the head is variable (see Figures 21 and 22), and the other differences noted by

Whitley are probably due to minor introductions

of the illustrator, but in any event are within the

range of intraspecific variation we have noted in

other species.

MATERIAL EXAMINED.—AUSTRALIA: NORTHWEST AUSTRALIA: BMNH 1933.8.14.7 (34.5). NORTH-
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ERN TERRITORY: Port Darwin, AMS IA.4298 (31.1,

holotype of Graviceps darwini and neotype of Petroscirtes lineolatus); Port Bradshaw, USNM 174335

(37.4). WESTERN AUSTRALIA: Broome, WAM P10100

(36.3). NEW GUINEA: Merauke, RMNH 26918

(66.8).

Omobranchus loxozonus (Jordan and Starks)

FIGURE 23

Petroscirtes loxozonus Jordan and Starks, 1906:705 [Tanegashima].

Description (see also Tables 5 and 12).—Dorsalfin spines XII-XIV (modally XIII), 19-22 = 32-35;

anal fin II, 21-25; both anal-fin spines of males

discernible externally; segmented caudal-fin rays

13-15 (13 in 92.1% of specimens); dorsal + ventral

procurrent caudal-fin rays 10-15; vertebrae 10-12

(11 in 93.8% of specimens) + 28-31 = 39-42; epipleural ribs 12-16 (13-15 in 97.3% of specimens);

prenasal pores present; interorbital pores 2-4 (3 in

96.0% of specimens); circumorbital pores 7-9 (8 in

95.5% of specimens); lateral-line tubes 0-4 (0-3 in

98.5% of specimens); lateral-line tubes, when present, extending posteriorly to below level of dorsalfin spine 1-6 (1-5 in 99.0% of specimens); gill

opening varying from restricted to area dorsal to

level of dorsalmost pectoral-fin ray to extending

ventrally to opposite 4th ray; lower-lip flap present;

circumorbital bones 5; lower jaw teeth 18-28; upper jaw teeth 16-27 (Figure 43); no fleshy bladelike

crest on top of head of either sex (head of some

presumably mature males swollen in appearance).

COLOR PATTERN.—Males: Head: Dusky band extending from anteroventral margin of orbit across

angle of jaws to chin, where it joins corresponding

band of opposite side; dorsal portion of band with

narrow, pale anterior and posterior margins. Four

evenly spaced, parallel, dusky bands more posteriorly on lower half to third of head (bands greatly

faded or absent in most specimens examined); 1st

band angled posteroventrally from posteroventral

margin of orbit, directed anteriorly for short distance at level of anterior preopercular sensory

pores, then redirected posteroventrally to ventral

TABLE 12.—Frequency distributions for certain meristic characters in populations

of Omobranchus loxozonus

Population

Japan

Wakayama Prefecture

Kagawa Prefecture

Miyazaki Prefecture

Tanega-shima

Okinawa

Population

Japan

Wakayama

12

1

3

11

1

Dorsal-fir

spines

13 14

5 2

1

4

66 l

20 1

Prefecture

Kagawa Prefecture

Miyazaki Prefecture

Tanega-shima

Okinawa

I

13-

12.

12.

13.

10

1

3

l

6

9

0

Segmented

dorsal-fin rays

19 20

1 5

6

2 30

3

Precaudal

vertebrae

11

7

1

7

68

22

12 5

10

11

3 11

11

21

2

1

1

39

13

9

0

0

0

22 X

20.1

20.1

7 20.6

6 21.1

Caudal

28 29

1 5

1 5

5 38

2

Total dorsal-fin

elements

32

1

2

3

33 34 35

4 3

1

5

35 36 4

3 13 6

vertebrae

30 :

2

1

1

30

16

31 X

29.1

29.0

3 29.4

4 30.1

33.

32.

33.

34.

2

7

5

1

Total

39

3

1

4

40

4

5

42

2

21

1

2

Segmented

anal-fin rays

22

4

2

21

1

23 24 25

3

1

4

48 8

15 5 1

vertebrae

41

1

1

1

29

16

42

3

k

X

39-8

40.0

40.4

41.1

X

2 2 .

2 2 .

2 2 .

23-

1*

4

8

3
NUMBER 177 49

midline of head; dorsal half of band with narrow,

pale anterior marginal line continuing dorsally

along posterior edge of orbit, and narrow, pale posterior marginal line continuing dorsally as anterior

margin of dorsoposteriorly lying, dome-shaped,

dusky smudge; succeeding 3 bands each originating

slightly higher on head than preceding band; last 2

bands extending ventrally onto prepelvic and prepectoral areas respectively; bands confluent ventrally with corresponding bands of opposite side.

Large dome-shaped, dusky smudge approximately

size of eye posterior to eye between circumorbital

and preopercular series of cephalic sensory pores;

smudge bordered anteriorly and posteriorly by narrow, pale, marginal lines; intense, dark spot approximately size of pupil in dorsoanterior corner of

smudge. Operculum with vertically elongate, less

dark spot encircled by narrow, pale, marginal line;

slightly narrower, pale line running anterodorsally

from dorsal end of spot, passing above dusky

smudge and curving to join anterior, pale, marginal line of smudge; faint, dusky bar present posterior to spot. Snout and interorbital area darker

than background.

Trunk: Approximately 11 more or less vertical,

broad, dusky to dark bands present on body; pigment along central vertical axis of each band more

intense than along edges; intensification sometimes

occupying almost all of some bands, sometimes restricted to portion of band at lateral midline of

body; bands usually not reaching onto upper and

lower one-fourth to one-fifth of body (bands very

broad, diffuse, and reaching more dorsally in very

large specimens than in small specimens); anterior

5 or 6 bands inclined dorsoanteriorly; succeeding 4

or 5 bands inclined dorsoposteriorly at least above

lateral midline (dorsal portion of approximately

9th band drawn into longitudinal streak above

ventral midline of body), posterior 1 or 2 bands

represented as short, narrow, stripes on lateral midline of body; often 1 or 2 middle bands broken into

side-by-side paired, narrower bands. Elongate, dark

blotch or spot usually present at dorsal tip of each

body band, at least anteriorly, forming longitudinal

row of spots just above body bands; each blotch or

spot lies just below corresponding spot in row of

approximately 8 moderately large pairs of evenly

spaced, dark spots on dorsal body contour at base

of dorsal fin. Dusky to dark blotches sometimes

present between and below ventral tips of posterior

bands. Best preserved specimens with narrow, pale

lines midway between and parallel with body bands;

pale lines extending from above dorsal tips of

body bands to ventral body contour where they

continue onto anal fin.

Pectoral Fin: Fin and fleshy base pale with even

spread of dark specks of pigment.

Pelvic Fin: Pale.

Dorsal Fin: Membranes transparent with dusky

to dark markings. Spinous ray portion of fin with

several broad, dorsoanteriorly slanted to longitudinal dusky stripes; very large specimens with spinous

portion largely dusky, stripes indistinct. Segmented

ray portion with narrow, dusky, often faint stripes;

oval, dusky to dark spot in distal portion of fin

between 10th and 14th segmented rays (usually

reaching from 11th to 13th); spot absent in many

specimens.

Anal Fin: Pale proximally, dusky distally, tips

of rays abruptly pale. Best preserved specimens with

several distinct, narrow, pale lines slanted ventroposteriorly from ventral ends of narrow, pale lines

on body.

Caudal Fin: Largely pale, sprinkled with dark

specks of pigment. Two vertically oriented, dusky

blotches at base of fin; very dark specimens with

diffuse, dusky stripe directed posteriorly from ventral blotch.

Females: Similar to males except: body bands

narrower, intensified portions much darker, usually

occupying most of band; other body markings darker; stripes on dorsal fin usually more distinct; dusky

to dark oval spot on dorsal fin absent.

GEOGRAPHIC VARIATION.—Variation in meristic

characters was noted only in the Okinawa population of O. loxozonus, which tended to have higher

averages for most characters (Table 12).

GEOGRAPHICAL DISTRIBUTION (Figure 5).—Southern Japan and Okinawa. Kamohara (1957) reported O. loxozonus as occurring as far north as Kominato, Chiba Prefecture. We have not been able to

verify this record and have not, therefore, entered

it on the distribution map.

HABITAT.—Little information is available, but

some collections have come from shallow, rocky

tidepools.

COMPARISONS.—Omobranchus loxozonus is a

member of the elongatus species groups (see "Comparisons" under O. elongatus). Within the elongatus

species group, O. loxozonus is most similar to O.
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

germaini, which is allopatric. The color pattern on

the head and body of O. loxozonus is less contrasty

than that usually exhibited by O. germaini, and the

number of body bands is less. Where populations

of the two species are geographically closest there

are marked differences in numbers of fin-rays, vertebrae, and lateral-line tubes (see Tables 5, 10, 12),

although overlaps in the ranges for these characters

exist.

Omobranchus loxozonus can be most readily distinguished from those species of Omobranchus with

which it appears to be sympatric as follows: from

O. fasciolatoceps in lacking a fleshy crest, in having

the gill opening extending ventrally to opposite the

1st to 4th pectoral-fin rays (versus restricted to area

dorsal to dorsalmost ray), and in rarely having as

few as 2 interorbital pores (versus rarely having as

many as 3 pores); from O. elegans in lacking the

dense covering of small, dark spots on the body and

in infrequently having as few as 7 circumorbital

pores (versus strongly modally 7 pores); and from

O. punctatus in having 0-4, rarely more than 3,

lateral-line tubes (versus 2-8, rarely fewer than 4

tubes), and in having the gill opening usually extending ventral to the level of the dorsalmost

pectoral-fin ray (versus usually restricted to area

dorsal to dorsalmost ray and never extending ventrally below dorsalmost ray).

MATERIAL EXAMINED.—JAPAN: WAKAVAMA PREFECTURE: CAS uncataloged (ca. 41); Shirahama,

CAS GVF reg. no. 747 (6: 49.5-64.8), GVF reg. no.

748 (ca. 49). KAGAWA PREFECTURE: Toyohama, CAS

uncataloged (ca. 51). MIYAZAKI PREFECTURE: NObeoka, USNM 199520 (7: ca. 34-59.5). TANEGASHIMA: BMNH 1923.2.26.341-50 (6: 37.4-47.8), CAS

SU9364 (53.5, presumed holotype of Petroscirtes

loxozonus, see Springer and Eschmeyer, 1974),

SU9783 (25: 28.8-58.0), USNM 53275 (16:32.7-54.6,

including putative holotype of Petroscirtes loxozonus), 70761 (142: 19.4-70.7). OKINAWA: USNM

70763 (4: 34.2-ca. 51), 123770 (ca. 45), 132754

(37.9), 132812 (13: 22.2-42.1), 209269 (2: 27.9-42.5),

209715 (ca. 40).

Omobranchus mekranensis (Regan)

FIGURE 24

Petroscirtes mekranensis Regan, 1905:328 [Jask, Mekran

Coast].

Petroscirtes cristatus Zugmayer, 1913:20 [Ormara].

DESCRIPTION (see also Table 5).—Dorsal fin XIXII (XI in one of 13 specimens), 20-21 = 32-33;

anal fin II, 22-23; one or both anal-fin spines of

males not discernible externally; segmented caudalfin rays 13-14 (14 in one of 13 specimens); dorsal

+ ventral procurrent caudal-fin rays 12-14; vertebrae 11 + 28-29 = 39-40; epipleural ribs 17-23;

prenasal pores present; interorbital pores 2; circumorbital pores 8; lateral-line tubes 6-7; lateralline tubes extending posteriorly to below level of

dorsal-fin spine 9-11; gill opening restricted to area

dorsal to level of dorsalmost pectoral-fin ray; lowerlip flap present; circumorbital bones 5; lower jaw

teeth 22-27; upper jaw teeth 18-23 (Figure 44);

fleshy bladelike crest present on top of head of

both sexes.

COLOR PATTERN:—Males: Head: Three narrow,

dark bands radiating from eye on ventral twothirds of head; anterior band flared ventrally, extending from anteroventral edge of orbit across

mouth, just anterior to angle of jaws, to ventral

midline of head, where it joins corresponding band

of opposite side; 2nd band extending ventrally and

slightly posteriorly from ventral margin of orbit,

flaring sharply midventrally on cheek and terminating just before reaching ventral midline of head;

3rd band extending posteroventrally from posterior

or posteroventral margin of orbit, becoming faint

or disjunct ventrally on cheek, recommencing as

broader, darker continuation, which is displaced

slightly posteriorly, and continuing across branchiostegal membranes onto prepelvic area, where it

joins corresponding band of opposite side. Moderately narrow, dark line (4th band) arises from dark

blotch on upper end of operculum and runs posteroventrally across branchiostegal membranes,

where it joins broad, dark band on fleshy pectoralfin base. Series of narrow, dark spots, dashes, or

lines present in each pale space separating dark

head bands; each series parallel to and midway

between adjacent head bands; each series except

last continuous on underside of head with corresponding series of opposite side. Distinct, vertically

oriented, dark, oval spot approximately one-half to

two-thirds size of eye present just posterior to

dorsal two-thirds of eye. Patch of intensely dark

melanophores present on free opercular membrane

margin; anterior edge of margin sometimes darkened to form narrow line running into upper end

of band on fleshy pectoral-fin base. Snout dusky.
NUMBER 177 51

Interorbital and nape areas with irregular, narrow,

dark, vertical lines mostly originating at dorsal

edge of orbit, extending to dorsal midline and onto

fleshy bladelike crest as pattern of vermiculations.

Broad saddlelike, dark band on nape just anterior

to 1st dorsal-fin spine; ventral end of band almost

confluent with dark blotch at upper end of

operculum.

Trunk: Pale to pale dusky with 7 pairs of broad,

dark bands; posterior 2 bands often faint to absent

(paired nature of bands less apparent in very large

specimens); pale spaces between band pairs, one and

one-half to two times as broad as adjacent bands.

Bands extending from dorsal body contour almost

to ventral body contour or becoming faint on ventral 3rd of body. Dorsal ends of body bands often

expanded, occasionally more intense than rest of

band, often with pale spot in center or near dorsal

end of expanded area (particularly in anterior

bands). Expanded dorsal ends of anterior 2 or 3

band pairs often deflected slightly anteriorly, those

of posterior 2 or 3 pairs deflected slightly posteriorly; expanded portions of anterior and posterior

bands sometimes separated from rest of band. Very

faint, narrow, vertical, dusky lines sometimes present midway between bands.

Pectoral Fin: Pale with even spread of dark

specks of pigment. Narrow, faint, dusky bar extending along basal portion of upper half of fin; ventrodistal portion of fin dusky. Fleshy base with nearly

vertical, broad, dark band extending from edge of

branchiostegal membrane to chest, usually faint or

terminating on prepelvic area.

Pelvic Fin: Pale with even spread of dark specks

of pigment.

Dorsal Fin: Transparent with dusky to dark

markings. Dark dusky blotch at fin base just above

each body band; some specimens with each blotch

consisting of 2 closely paired blotches. Spinous portion of fin with moderately broad, dusky to dark

stripe; some specimens with basal dark blotches

extending dorsoposteriorly as faint, moderately

broad, dusky bands; proximal portion of fin often

dusky.

Anal Fin: Largely dusky, especially distally;

proximal portion often slightly transparent. Tips

of rays abruptly pale, particularly anterior rays.

Caudal Fin: Pale with 2 or 3 dusky streaks directed posteriorly from 1 or 2 vertically oriented,

large dusky areas on basal one-fifth of fin. Proximal

edge with narrow, dark dusky bar centrally. One

large specimen had very fine, intensely dark spots

spread over dorsal, anal, and caudal fins.

Females: Similar to males except: dark markings

on head, body, and fins (other than oval spot posterior to eye and saddle-like band on nape) fainter;

body bands noticeably fainter; bands not distinct

as in males, even in darkly pigmented females.

GEOGRAPHIC DISTRIBUTION (Figure 4).—Persian

Gulf, Gulf of Oman, and northern Arabian Sea.

The Persian Gulf record was not accompanied by

a specific locality; the record on the distribution

map for the Persian Gulf is to be considered as

generalized. Reported as occurring at Minicoy

Atoll, Laccadive Archipelago, by Nagabhushanam

and Chandrasekhara Rao (1972), which, if true,

would represent a significant range extension. We

were unable to verify the identification.

HABITAT.—Known only for 1 collection; rocks,

sand, and sparse coral, thus indicating normal

ocean salinites.

COMPARISONS.—Omobranchus mekranensis is a

member of the banditus species group (see "Comparisons" under O. banditus). Superficially it is

most similar to O. fasciolatus (for comparison see

O. fasciolatus). The only species of Omobranchus,

other than O. fasciolatus, with which O. mekranensis appears to be sympatric is O. punctatus, which

lacks a fleshy crest and rarely has as few as 2 interorbital pores (only 2 in O. mekranensis).

NOMENCLATURAL DISCUSSION.—Zugmayer (1913)

compared his P. cristatus only with P. dispar, which

he believed to be the only species of Petroscirtes

{Omobranchus) with a crest on top of the head.

The four syntypes of P. cristatus are no longer extant (Dr. F. Terofal, Munich, in litt.), and one

must depend entirely on the original description to

determine the identity of P. cristatus. The important identifying characters contained in the description are the following (translated from the German): dorsal fin 33, an oval, dark blue spot restricted to the preopercle (Vordeckel), 10-12 spots

along the base of the dorsal fin, which, on the anterior half of the body, send down transverse bandlike continuations toward the lateral line. (Presumably the dark blue color described by Zugmayer is based on fresh material. In blenniids, this

color is lost in preservation and is represented by

dusky gray.)

Among the crested forms of Omobranchus in the
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Indian Ocean, we recognize four species: O. banditus and O. woodi (both restricted to southern

Africa), O. fasciolatus (western half of the Indian

Ocean), and O. mekranensis (Persian Gulf, Gulf of

Oman, and northern Arabian Sea). The last two

species have been taken together in a single collection at Astola Island, close to Ormara, the typelocality of P. cristatus. Of these four species, O.

fasciolatus can be excluded from consideration on

the basis of its lower total number of dorsal-fin ray

elements (30-31). The preopercular spot of O. fasciolatus (Figure 17) is not so distinctly oval as is

the spot of O. mekranensis. Similarly, the transverse

body bands of O. fasciolatus are less distinct than

those of O. mekranensis. Both O, woodi and O.

banditus also lack the preopercular oval spot (although such a spot does occur dorsoanteriorly on

the body in O. banditus). The number of dorsal-fin

ray elements and color pattern, as well as the typelocality, agree well with what is known for O. mekranensis, and for these reasons we synonymize P.

cristatus with O. mekranensis.

MATERIAL EXAMINED.—PERSIAN GULF: BMNH

1900.7.25.41 (43.5). IRAN (Gulf of Oman): Jask,

BMNH 1904.5.25.94 (ca. 46, holotype of Petroscirtes mekranensis). WEST PAKISTAN: Astola

Island: USNM 200604 (11: 33.3-50.7, including

two cleared and stained).

Omobranchus meniscus, new species

FIGURE 25

DESCRIPTION (where variation is indicated, the

holotype is flagged with an asterisk).—Dorsal fin

XII, 19; anal fin II, 21-22*; both anal-fin spines of

male discernible externally; segmented caudal-fin

rays 13; dorsal + ventral procurrent caudal-fin

rays 9-10#; vertebrae 10 + 26-27*; epipleural ribs

11-12*; prenasal pores present; interorbital pores

3; circumorbital pores 8; lateral-line tubes absent;

gill opening extending ventrally to opposite 3rd

pectoral-fin ray; lower-lip flap present; circumorbital bones 5; male with fleshy, bladelike crest on

top of head (2.8 percent SL), female lacks crest.

Teeth (Figure 43): The 60.1 mm SL male holotype has 20 premaxillary and 22 dentary incisoriform teeth. The 54.8 mm SL female paratype has

21 premaxillary and 23 dentary incisoriform teeth.

Both specimens have dorsal and ventral posterior

caniniform teeth.

COLOR PATTERN (specimens faded).—Male: Head:

Moderately narrow, dusky band extending from

anteroventral margin of orbit at least to corner of

mouth. Large, intensely dark, anteriorly opening,

crescentic mark curving dorsally from dorsoposterior margin of orbit, almost reaching to base of

fleshy head crest. Snout dark dusky with sensory

pores and nostrils pale ringed. Dark dusky vermiculations present on crest. Dark, oval blotch

present laterally on nape just anterior to dorsal-fin

origin, narrowly joined to similar blotch on opposite side at dorsal midline; 3 to 5 small, intensely

dark spots present on blotch in dorsoposteriorly

directed line.

Trunk: Several faint, moderately broad, dusky

bands apparently present on side, extending approximately from dorsal to ventral body contours.

Indication of broad, dark dusky stripe just below

dorsal body contour extending from just behind

head to caudal peduncle (possibly artifact of preservation); longitudinal series of dusky blotches or

dusky stripe apparently present at dorsal-fin base.

Number of very small, intensely dark spots scattered along body, probably restricted somewhat to

body bands. Connected horizontal line of very narrow, dark dashes present posteriorly on lateral midline of body.

Pectoral Fin: Pale, transparent.

Pelvic Fin: Pale.

Dorsal Fin: Dusky anteriorly, pale to transparent

posteriorly. Anterior half of fin with several darkish streaks, apparently representing dorsoposteriorly

angled dark lines. Posterior half of fin with streaks

very faint, fin transparent distally.

Anal Fin: Pale with dark membranes between

tips of rays.

Caudal Fin: Pale. Midportion of base dusky, becoming faint distally.

Female: Apparently similar to male, appears to

have slightly less developed body bands and less

distinct dorsal-fin markings (crest absent).

COMPARISONS.—Omobranchus meniscus is easily

distinguishable from all other species of Omobranchus by the dark, crescentic marking extending

dorsally from the posterior margin of the orbit. It

is otherwise distinguishable only on the basis of a

complex of characters. It appears probable that
NUMBER 177 53

when more specimens of 0. meniscus are available,

the species will exhibit a low number of meristic

elements compared with those of the other species

in the genus.

There are eight species of Omobranchus. in which

the males may have a fleshy, bladelike crest on top

of the head. Of these, all but O. meniscus have the

gill opening predominantly restricted to the area

dorsal to the level of the dorsalmost pectoral-fin

ray. Only one of these species with a restricted gill

opening, O. fasciolatus, has the gill opening extending ventrally as far as the level of the second

pectoral-fin ray (and this condition was found in

only one of 54 specimens), whereas both specimens

of O. meniscus have the gill opening extending

ventrally to opposite the third pectoral-fin ray.

Of the eight crested species, only O. anolius, O.

aurosplendidus, and O. meniscus commonly lack

bipored lateral-line tubes.

The two specimens of O. meniscus have nine or

ten total procurrent caudal-fin rays. Among the

other crested species only O. aurosplendidus has as

few as ten total procurrent caudal-fin rays.

Possibly the head of O. meniscus is proportionately larger than that of most of the other Omobranchus species. The head length of the two

specimens of O. meniscus, 54.8 and 60.1 mm SL,

are 24.2 and 23.6% SL, respectively. The head

length of O. zebra is proportionately about the

same as that of O. meniscus at similar sizes (the

largest specimen of O. zebra available is only 56.5

mm SL). Omobranchus aurosplendidus may also

have a head length proportionately the same as

that of O. meniscus, but this is not certain as there

are no specimens available less than 75 mm SL.

Where specimens of comparable size to those of

O. meniscus are available, the head length is about

17-22% SL in the other Omobranchus species.

ETYMOLOGY.—The specific epithet meniscus comes

from Latin, meaning "crescent," and refers to the

dark, crescent-shaped mark on the head that distinguishes the species. Meniscus is here used as a

noun in apposition.

HOLOTYPE.—USNM 119685, male, 60.1 mm SL,

mouth of river at Chantabun, Thailand. Collected

by Mai Mah, 10 April 1933; habitat unknown.

PARATYPE.—USNM 211153, female, 54.8 mm SL,

same data as holotype.

Omobranchus punctatm (Valenciennes)

FIGURES 26 and 27

Blennechis punctatus Valenciennes in Cuvier and Valenciennes, 1836:286 [le canal de Bombay].

Petroscirtes dispar Gunther, 1861:232 [in part; Amoy, China;

lectotype designated below].

Petroscirtes semilineatus Kner, 1868b:333 [Candavu].

tPetroskirtes japonicus Bleeker, 1869:246 [Jedo, Japonia].

Salarias decipiens DeVis, 1884b:694 [Cardwell, Queensland

Coast].

Salarias helenae DeVis, 1884b:697 [St. Helena, Moreton Bay].

Salarias sindensis Day, 1888:263 [Kurrachee].

Aspidontus dasson Jordan and Snyder, 1902:456 [Wakanoura,

Japan].

Petroscirtes kochi Weber, 1908:263 [Merauke-Fluss, Sudkiiste,

New Guinea, Brackwasser].

Poroalticus sewalli Fowler, 1931:403 [tide pools at Brighton

Beach, Trinidad].

Petroscirtes masyae H. M. Smith, 1934:316 [tide pool on Koh

Chula, a rocky islet in the Gulf of Siam off Lem Sing,

Southeast Siam].

Omobranchus japonicus scalaius J. L. B. Smith, 1959:232

[Delagoa].

DESCRIPTION (see also Tables 5 and 13).—Dorsal

fin XI-XIII (XII in 90.6% of specimens), 19-24 =

31-36; anal fin II, 20-26; one or both anal-fin spines

of males not discernible externally; segmented

caudal-fin rays 12-14 (13 in 97.9% of specimens);

dorsal + ventral procurrent caudal-fin rays 8-16

(rarely 8-11); vertebrae 10-12 + 26-31 = 37-41;

epipleural ribs 15-25 (rarely 15-17 or 25); prenasal

pores present; interorbital pores 2-4 (3 in 95.6% of

specimens); circumorbital pores 6-10 (8 in 92.8%

of specimens; 6 or 10 in 0.6% of specimens); lateralline tubes 2-8 (usually more than 4); lateral-line

tubes extending posteriorly to below level of dorsalfin spine 3-12 (usually 7-11); gill opening either

restricted to area dorsal to level of dorsalmost

pectoral-fin ray (93.0% of specimens) or extending

ventrally to opposite dorsalmost ray; lower-lip flap

present; circumorbital bones 4-5 (of 5 specimens

examined all had 5 except one specimen with 5 on

one side and 4 on the other; the 4 bones resulted

from a fusion of the 2nd and 3rd circumorbitals

from the anteriormost); lower jaw teeth 22-38

(Figures 49-52); upper jaw teeth 21-33; no fleshy

bladelike crest on top of head of either sex.

COLOR PATTERN.—Males: Head: Broad, ventrally

flared, dark dusky band running from anteroventral

margin of orbit across mouth, just anterior to angle

of jaws, onto chin where it joins corresponding
TABLE IS.—Frequency distributions for certain characters in populations of Omobranchus punctatus

Population

Mozambique

Persian Gulf

Gulf of Oman

Arabian Sea

India

Gulf of Kutch

Bombay

Vizagapatnam

Maldive Islands

Ceylon

Kikobar Islands

Burma

Thailand

West Coast

Oulf of Thailand

West Coast

North Coast

East Coast

Hong Kong (vicinity )

China

Amoy

Chusan Island

Japan

Philippine Islands

Halrnahera

Hew Guinea

Bougainville

Australia

Shark Bay

Ons low

Horth Coast

East Coast

Candavu (?)

Trinidad

Population

Mozambique

Persian Gulf

Gulf of Oman

Arabian Sea

India

Gulf of Kutch

Bombay

Vizagapatnam

Maldive Islands

Ceylon

Kikobar Islands

Burma

Thailand

nest Coast

Gulf of Thailand

West Coast

north Coast

East Coast

Hong Kong (vicinity)

China

Amoy

Chusan Island

Japan

Philippine Islands

Raima he ra

Hew Guinea

Bougainville

Australia

West Coast

Shark Bay

Onslou

north Coast

East Coast

Candava (?)

Panama

Trinidad

Population

Mozambique

Puraian Oulf

Oulf of Oman

Arabian Sea

Oulf ii Kutch

Bombay

Vizagapatnam

Maldive Islands

Ceylon

Ilkobar Islands

Burma

Thailand

West Coast

Oulf of Thailand

West Coast

North Coast

Fact Coast

Hong Kong (vicinity)

China Amoy

Chusan Island

Japan

Philippine Islands

HaUar.era

lew Guinea

Bougainville

A.strain

Shark toy

Onslo*

lorth Coast

Eait Coast

Candavu (?)

Parana

Trinidad

Do

11

2

3

5

4

2

1

1

1

1

2

rsal-fin

spines

12

6

2

_

6

1

10

5

1

541

1

25

1

29

63

13

1

24

9

3

2

2

1

6

7

6 0

1

21

33

13

-

_

-

-

3

1

1

2

-

1

1

6

I

12.0

12.0

11.0

12.0

12.0

12.0

12.0

12.0

11.2

11.9

12.0

11.9

12.0

11.9

12.2

12.0

12.0

12.0

12.0

12.0

12.0

12.1

Precaudal

vertebrae

1 0

1

2

1

6

14

2

2

1

1

2

2

3

3

1

1

1

1 1

4

2

2

3

7

5

1

53

1

1

25

3

29

55

15

1

24

6

4

1

2

1

8

4

7

61

1

20

32

iate

.

1

5

1

2

12

-

1

2

-

1

1

-

_

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1

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12

2

-

1

-

-

I

10.8

11.0

11.0

11.0

11.2

11.0

11.0

11.0

10.8

10.8

10.8

11.0

11.0

11.1

11.0

10.5

11.0

11.6

n . 3

11.0

11.0

10.9

10.9

al-line tube

2

1

1

.

1

7

7

22

9

1

1

2

2

6

U

2 -

1

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4 1

64 58

1

1

20 18

2 2

3* 25

25 24

12

23 51

2 5

3 1

1

2

1

< 11

3 1

6 -

21 25

1

6 1

14 11

19

1

1

1

1

1

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26

2

1

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1

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4

2

.

3

1

3

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1

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17

19

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5

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1

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1

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11

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13

14

3

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4

5

1

3

4

5.5

5-5

6 . 0

5 . 7

5 . 9

6 . 2

6 . 4

6.3

6 . 5

6 . 0

6 . 2

7 . 2

6 . 8

6 . 7

6 . 2

6 . 0

6 . 0

6.6

5.3

5.8

6 . 4

5.6

5.6

2 8

3

1

2

1

1

7

1

42

1

11

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39

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La

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23

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24

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1

1

1

1

3

2

1

1 7

2

1

1

1

2

1 2

2 4

X

20.3

21.0

21.0

21.2

21.2

20.0

20.4

20.2

21.0

20.6

20.8

22.8

22.7

22.1

20.0

-'*

19.5

20.0

20.3

21.9

20.9

20.7

28.0

27.5

28.0

28.5

28.0

27.2

27.9

27.3

27.8

27-7

28.0

29-5

29.4

28.6

27.2

26.5

86.5

28.2

27.0

27.3

28.8

28.2

28.1

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1-fln

3

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1

2

3

37

17

1

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13

13

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5

3

4

5

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1

1

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50

1

18

2

28

2 0

4

27

.

3

1

2

1

8

.

4

19

1

6

6

31

1

1

1

1

1

To

32

2

1

3

34

1

1

21

3

20

19

3

It

5

3

11

Total

1

2

2

1

ube

-

_

-

6

_

45

1

11

_

30

25

7

38

1

1

.

6

2

22

3

12

1

1

4

9

1

12

2

16

17

3

.

1

1

4

5

3

It

tal

el

33

3

2

5

6

1

22

5

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14

46

1

1

1

2

10

17

25

ver

4

1

2

1

7

1

43

1

12

2

15

46

2

1

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5

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12

25

belou

-

-

1

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3

1

5

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3

8

3

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1

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34

1

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1

1

2

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1

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1

It

5

9

8

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35

:

-

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9

1

16

1

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2-

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-

-

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1

1

2

-

1

3

6

7

17

5

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_

14

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43

1

6

5

7 . 8

6 . 0

8.5

7 . 7

9 . 7

8.4

9 . 0

8 . 8

8.2

8 . 9

9 . 2

10.1

9.4

10.1

9.2

8 . 0

8 . 0

9.2

8.5

8.5

9.3

8.6

8.4

-

-

-

-

-

-

-

-

-

_

7

1

8

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-

-

_

l

_

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6

36

;

-

-

i

-

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38.8

38.5

39.0

39.5

39.2

38.2

38.8

38.4

38.5

38.6

38.8

40.5

40.3

39-7

38.2

37.0

37.5

39.8

38.3

38.3

39.9

39.1

39.0

1

X

32.3

33.0

33.

33.

33.

32.

32.

32.

32.

32.

0

2

2

0

4

2

2

4

32.6

34.

34.

34.

32.

31.

31.

33.

32.

32.

7

6

0

2

5

1,

0

3

33.9

32.8

32.8

1

1

1

Segmer.

20 21

Dsrwl

1

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1

1

1

1

1

3

2

22

i

1

?

1

7

4

26

1

1

20

2

24

24

1

3

5

7

9

12

ted

ray

23

i

1

2

2

3

-

30

5

2

9

41

2

1

12

-

48

1

12

25

• ventral p

caudal-fin r

2

1

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1

2

1

Eplpleu

2

1

1

1

2

4 7

5 8

1

1

2 1

1 1

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1

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12

20

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1

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1

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1

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22

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7

11

3

1

8

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13

5

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22.5

22.5

23.0

22.7

22.3

21.6

22.6

22.2

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22.3

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24.9

24.7

23.6

2S.2

20.5

21.C

22. ,'

21.5

21.7

22.9

22.6

22.8

*

13.8

14.5

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14. u

13.4

14.0

13."

13.£

13-5

14.5

13.6

13-?

13.1

14.0

13-5

13-5

13.0

14.2

13.4

13-7

12.4

12.9

X

20.2

20.5

23.0

23.3

23-1

2O.d

22.0

22.0

20.0

19. fc

20.4

21. J

22. C

21.4

23.0

23.1

22.7

21."1

21.6

22.C

21.1

£1.9
NUMBER 177 55

band of opposite side; band, interrupted by orbit,

continues from dorsoposterior margin of orbit to

dorsal midline of head, decreasing in definition.

Second broad, dark dusky band present posterior

to eye completely encircling head, often faint to

absent midlaterally, being represented only by narrow, intensely dark, anterior marginal line, dusky

to dark blotch, or curved, diagonal, moderately

broad, dark dusky line posterior and ventroposterior to eye. Third broad, dark dusky band extending from dorsal end of operculum to ventral side

of head, sometimes confluent with corresponding

band of opposite side at ventral midline; band

often faint midlaterally on operculum. Snout and

interorbital dark dusky with large, irregular, pale

blotches or spots anteromedially to eyes. Nape dark

dusky with irregular pale spots and blotches anteriorly. Nape posterolaterally with prominent, dark,

anteroventrally intensified blotch just anterior to

dorsal-fin origin; corresponding blotches on opposite sides often continuous as band, sometimes with

dusky blotch present between them middorsally;

dark blotches, or band, usually set off by pale

margin. Short, nearly horizontal, dark line present

at dorsal end of 3rd dark head band.

Trunk: Four or five, more or less parallel, evenly

spaced, slightly wavy, dark, horizontal lines present

anteriorly on dorsal three-fourths of body; large

specimens with lines extending posteriorly for half

body length; lines much shorter in small specimens;

midlateralmost line on body reaching forward to

center of gill opening; dorsalmost line usually,

formed from series of 8 or 9 more or less joined

horizontal dashes; midlateral 3 lines followed posteriorly by single midlateral series of dusky to dark

rectangular blotches, becoming faint posteriorly;

all blotches very faint in large specimens; blotches

reach far forward in small specimens; some specimens with blotches continued ventrally as very

faint, broad, dusky streaks. Dorsal body contour

with series of about 11 joined pairs of intensely

dark blotches; anteriorly each pair of blotches positioned above corresponding dash in dorsalmost

horizontal dark body line.

PECTORAL FIN: Pale with even spread of dark

specks of pigment on rays; membranes mostly transparent. Lower half of fleshy pectoral-fin base with

faint to prominent, broad, dark dusky bar immediately posterior to branchiostegal membrane; fleshy

pectoral-fin base sometimes with dark line along

base of fin, particularly dorsally.

Pelvic Fin: Pale with fairly even spread of dark

specks of pigment, somewhat darker proximally.

Dorsal Fin: Transparent to slightly dusky with

narrow, dark dusky stripe one-third distance up

from base; stripe becoming faint to absent posteriorly. Dorsoposterior portion of fin often dusky.

Some specimens with distal edge of fin narrowly

dusky.

Anal Fin: Pale dusky to dark dusky, especially

distally; ray tips abruptly pale, particularly anteriorly on fin.

Caudal Fin: Pale to dusky with vertically oriented

pair of slightly darker blotches at base of fin; dusky

streak directed posteriorly from each blotch in

some specimens.

Females: Similar to males except: horizontal

lines on body less distinct; midlateral, rectangular,

dark blotches well developed, remaining distinct

more posteriorly, often appearing associated with

more dorsal row of dashes and row of blotches on

dorsal body contour giving body banded appearance (banded appearance present in some males

but not usually to extent of that in females).

Geographical Variation: This species is fairly

consistent in color pattern throughout its range. A

great deal of variation occurs in head pattern (particularly midlaterally) largely due to the indistinctness of the markings posterior to the eye and on

the cheek. Specimens in the three lots examined

from Ceylon, however, were somewhat distinct in

the following particulars: head with dark markings

behind eye usually joined to form reticulated pattern; dorsalmost horizontal line on body not usually

clearly broken into dashes, posterior ends of next

two lines confluent with second and first midlateral

blotches respectively; blotches on lateral body midline less angular, extending slightly dorsally and

ventrally, particularly in smaller specimens; series

of paired blotches on dorsal body contour extending onto base of dorsal fin in many specimens.

GEOGRAPHIC VARIATION ( see also "Color Pattern").—Population variation was noted in most

meristic characters of O. punctatus (Table 13).

There is no particular pattern shown by these variations. The Western Australian populations exhibit

an increase in the number of precaudal vertebrae

that is duplicated by no other population of O.
56

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 14.—F values for covariance comparisons of regression equations

(Table 2) for number of lower or upper jaw teeth between males or

females from different populations of Omobranchus elongatus (** =

significant at p = .001; NS — not significant)

Comparisons

Lower teeth

males

Ceylon X Gulf of Thailand

Ceylon X China

Ceylon X Trinidad

Gulf of Thailand X China

Gulf of Thailand X Trinidad

China X Trinidad

females

Ceylon X Gulf of Thailand

Ceylon X Trinidad

Gulf of Thailand X Trinidad

Upper teeth

males

Gulf of Thailand X China

Gulf of Thailand X W. Australia

Gulf of Thailand X Trinidad

China X W. Australia

China X Trinidad

W. Australia X Trinidad

females

Ceylon X Gulf of Thailand

Ceylon X W. Australia

Ceylon X Trinidad

Gulf of Thailand X W. Australia

Gulf of Tr-ailar.d X Trinidad

W. Australia X Trinidad

F values

slopes

0.01 NS

O.46 NS

3.50 NS

0.69 NS

3.44 NS

6.22 NS

0.54 NS

3.49 US

2.23 NS

2.03 NS

1.32 NS

4.43 NS

0.34 NS

12.04 NS

6.20 NS

1.81 NS

2.99 NS

0.01 NS

1.74 NS

1.42 NS

2.37 NS

Degrees of

freedom

1/54

1/39

1/37

1/51

1/49

1/34

1/37

1/24

1/25

1/50

1/36

1/48

1/22

1/34

1/20

1/37

1/26

1/24

1/27

1/25

1/14

F values

heights

13.72 **

38.75 «*

0.42 NS

5.35 NS

18.95 **

27.73 **

59.23 **

9.95 NS

5.38 NS

9.75 NS

7.13 NS

24.13 **

0.77 NS

45.89 *»

32.65 *•

22.12 •»

22.40 **

0.00 NS

0.74 NS

12.46 us

13.07 NS

Degrees of

freedom

1/55

1/40

1/38

1/52

1/50

V35

1/38

1/25

1/26

1/51

1/37

1/49

1/23

1/35

1/21

1/38

1/27

1/25

1/28

1/26

1/15

punctatus. Similar increases in number of precaudal vertebrae are also exhibited by the Western

Australian populations of O. germaini and O. r.

rotundiceps, but no others. The factors causing this

peculiar type of variation are unknown.

Although covariance comparisons of numbers of

jaw teeth (Table 14) were not feasible for all populations of O. punctatus, several of the populations

exhibited significant differences in numbers of

teeth. In general, if one sex of one population

exhibited a significant difference in the number of

teeth in one jaw from that exhibited by the same

sex of another population, then both sexes of each

population differed significantly in numbers of

teeth in both jaws; where the differences in the

other jaw or other sex were not significant they

were nearly so.

GEOGRAPHIC DISTRIBUTION (Figure 7).—Recorded

from one restricted locality (Delagoa Bay) in the

western Indian Ocean, otherwise common from the

Persian Gulf to Japan and eastern Australia. One

questionable record from the Fiji Islands (discussion below). Common in Trinidad and in Panama

around the Atlantic entrance to the Panama Canal

and in lower Gatun Locks. (For discussion of

Atlantic distribution see p. 59.)

Yazdani (1963) reported Petroscirtes kochi

(= O. punctatus) from Phoenix Bay, Port Blair,

Andaman Islands. We are unable to verify this

identification and have not entered the record on

the distribution map for O. punctatus. Chyung

(1954) reported O. punctatus (as Dasson) from

Masan, Korea, and (1961) identifiably illustrated it.

We are unable to translate the Korean localities in

the 1961 publication but based on the 1954 locality,

together with the 1961 illustration, we have included the 1954 Korean locality on Figure 7.

HABITAT.—Shallow, often murky, marine and

occasionally brackish waters at or close to the shoreline or near river mouths or mangroves. In tidepools and areas with rocks (often encrusted with

barnacles or oysters), sparse corals, and sargassum.

Once taken at a salinity of 28 0/00 in Gatun Locks,

Panama. Reported (Lachner, Robins, Courtenay,

1970) common around docks in Trinidad.

COMPARISONS.—Omobranchus punctatus is a

member of the banditus species group (see "Comparisons" under O. banditus). It differs from the
NUMBER 177 57

other members of its group as follows: from all in

overall color pattern and in never having a fleshy

crest on the head, and from O. banditus in having

a lower-lip flap. It is otherwise distinguishable from

the other species of Omobranchus only by a combination of characters, although when its color

pattern is strongly manifested the species is recognizable at sight. The narrow, dark longitudinal

lines on the anterior quarter of the body occur in

no other species of Omobranchus.

NOMENCLATURAL DISCUSSION.—Giinther (1861)

described P. dispar on the basis of two specimens

representing different species, both with available

older names (O. punctatus and O. fasciolatoceps).

No comparison was made with any other species as

he erroneously thought the fleshy head crest, exhibited by one of his specimens, to be a unique

character among the species of Petroscirtes (which

in Gunther's day included Omobranchus). In order

to stabilize the name, we here designate the larger

of Gunther's two syntypes (BMNH 1860.7.20.99),

identifiable as O. punctatus, as the lectotype of

P. dispar. The other (crested) syntype bears the

catalog number BMNH 1860.7.20.100.

Kner (1868b) did not compare his P. semilineatus

with O. punctatus or O. dispar. We have examined the presumable holotype of P. semilineatus

and find it to be identifiable with O. punctatus.

(For a discussion of the complexities surrounding

the description of P. semilineatus see "Nomenclatural Discussion" under O. lineolatus).

The most commonly applied name to the species

we treat as O. punctatus is O. japonicus (Bleeker).

Bleeker (1869) described P. japonicus from Tokyo

on the basis of a single specimen, now lost. He

compared his species only with (the lectotype of)

P. dispar and noted that it differed from P. dispar

in having its head contained six and one-third

times in the total length (as opposed to seven

times) and in having three fewer dorsal-fin elements. Bleeker noted that his specimen was remarkable for its numerous jaw teeth, 36-40 in each

jaw.

Bleeker's description and illustration of P. japonicus clearly apply to a species of the genus Omobranchus, but it is impossible for us to assign it

with certainty to any species we know in that genus.

The only species of Omobranchus that has the jaw

teeth so numerous is O. fasciolatoceps (known from

China and Japan), and in that species only mature

females may have as many teeth as Bleeker reported. However, mature females of O. fasciolatoceps lack canine teeth, whereas P. japonicus was

described as having canine teeth. Furthermore, O.

fasciolatoceps has an obvious, high, fleshy crest on

top of the head in both sexes. Bleeker reported that

his specimen lacked a crest and no indication of a

crest is present in his figure.

There are three other species of Omobranchus

known from Japan. Of these, O. elegans can probably be excluded from consideration as identifiable

with O. japonicus because the black-spotted color

pattern of O. elegans is so distinctive and persistent

that Bleeker would hardly have overlooked it. In

the unlikely event that P. japonicus should prove

to be synonymous with O. elegans, O. japonicus

would have priority as it is the older name.

Of the remaining two Japanese species of Omobranchus—O. loxozonus and O. punctatus—neither

is a more likely candidate solely on the basis of

Bleeker's description. However, P. japonicus was

described from Tokyo, and of the two species, only

O. punctatus has been recorded from this far north

on the Japanese coast. We have therefore questionably synonymized P. japonicus with O. punctatus.

DeVis (1884b), who worked at the Queensland

Museum, briefly described S. decipiens without comparing it with any other species. We have examined

the holotype (only one specimen mentioned in the

description) and find it to be a female O. punctatus. A few pages later, in the same publication,

DeVis described S. helenae (number of specimens

not indicated), also without comparison. The description of S. helenae is slightly less brief than that

of S. decipiens, but almost identical with that of

S. decipiens, where the same characters were described. In the Queensland Museum there are three

specimens (QM 1.1361) that are cataloged as "types"

of S. helenae. In addition to these, the Australian

Museum has two lots of specimens received from

the Queensland Museum (AMS 1.12694, two specimens, and 1.445, five specimens) that are cataloged

as "cotypes" of S. helenae. There is some question

concerning the validity of the type-status accorded

to the specimens in AMS 1.445, but for the sake of

argument we assume that they are types. All the

QM and AMS specimens mentioned, except one

contained in AMS 1.445 that is identifiable as O. r.

rotundiceps, are conspecific with O. punctatus. To

stabilize the name of S. helenae we here designate
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

one specimen from QM 1.1361 as the lectotype of

S. helenae. The lectotype can be distinguished from

the other syntypes by the followng combination of

characters: 65.8 mm SL; 6 lateral-line tubes on the

left side; last epipleural rib on vertebra 21; dorsal

fin XII, 22; anal fin II, 23; 7 upper and 8 lower

procurrent caudal-fin rays, 29 upper and 30 lower

incisoriform teeth. The two syntypes formerly with

the lectotype are now cataloged as QM 1.10330.

Day (1888) described S. sindensis without comparison with any other species. The syntypic material comprises three specimens, all of which are

identifiable as O. punctatus.

Jordan and Snyder (1902) described A. dasson,

which they compared only with P. japonicus, noting that P. japonicus had more teeth than their

specimens. They reported that they had two specimens, which they called the type (sic), from Wakanoura. These two specimens, which must be

accorded syntypic status, are both identifiable as

O. punctatus.

Jordan and Snyder (1902) reported that the paratype (cotype) of Aspidontus dasson came from "Agu

in Shima." We are unable to locate this supposed

Japanese locality and suspect that it was originally

intended to be Aguni Shima, an island off Okinawa.

However, as Jordan and Starks (1907) gave no

records of fishes from Aguni Shima in their list of

fishes from the Ryukyus, we have not included their

record on the distribution map for O. punctatus.

Weber (1908) compared his P. kochi only with

O. lineolatus, and then only to state that it was

different without specifying how. We have examined the two syntypes of P. kochi and find them to

be identifiable as male O. punctatus.

Fowler (1931) did not compare his P. sewalli

with any other species. Inasmuch as his specimens

came from Trinidad, it is understandable that he

would not have looked for an applicable IndoPacific species name for his specimens. Although he

designated a catalog number for the holotype

(ANSP 53318) and a series of numbers for the paratypes (ANSP 53319-21, indicating 3 specimens), all

four types are presently included in a single jar

labeled with all the catalog numbers. Fowler did

note that the holotype was 57 mm TL and that the

paratypes were 50-55 mm TL; thus, the largest

specimen (which we find to be 47.6 mm SL, 55.5

mm TL) is probably the holotype. The next largest

specimen is 45.0 mm SL, but the caudal fin is

damaged and the TL cannot be measured. It is not

important to decide which specimen is the holotype

as all are identifiable with O. punctatus.

H. M. Smith (1934) did not compare P. masyae

with any other species. We have not seen the holotype (reported by Monkolprasit, 1969, to be in the

collections of Kasetsart University, Bangkok). The

description of the color pattern clearly applies only

to O. punctatus. Furthermore, there is a photograph of a drawing by L. Masya (for whom the

species was named), published as a postcard, in the

illustration files of the Division of Fishes, USNM,

that carries the legend, "Petroscirtes masyae." The

drawing was apparently based on the holotype,

which was reported to be 5.9 cm TL. The reproduced drawing, which is 9.4 cm TL, is stated on

the postcard to be X 1.6 natural size, or 9.4 cm.

This illustration clearly indicates the unique color

pattern of O. punctatus.

J. L. B. Smith (1959) described his subspecies,

O. japonicus scalatus, claiming that the stripes on

the head and pectoral-fin base differentiated his

form from the "typical form." The color pattern

described and illustrated by Smith is found in

specimens from throughout the range of O. punctatus. Although some populations tend to diverge

from others in color pattern, the differences are

minor and unworthy of nomenclatural recognition.

MATERIAL EXAMINED.—MOZAMBIQUE: Delagoa

Bay, RUSI 1096 (2: 11.9-39.9), 1102 (56.2), SAM

25385 (38.3), USNM 197417 (2: ca. 43-55). IRAN:

Bushire, (Persian Gulf), UZMK CN6-7 (38.8),

CN6 (ca. 54); Jask (Gulf of Oman), BMNH 1904.

5.25.92-3 (2: ca. 44-46). PAKISTAN (Arabian

Sea): Karachi (= Kurrachee), BMNH 1889.2.1.

3616-18 (3: 43.6-53.5, syntypes of Salarias sindensis), 1898.6. 29.148-9 (2: 30.5-45.1), UZMK P.75390

(34.9). INDIA: Okha Point (Gulf of Kutch), USNM

201882 (58.6); Bombay, ANSP 100170 (54.6), 122231

(68.8), BMNH 1887.9.22.67-73 (7: 30.9-57.7),

MNHN 716 (40.2, holotype of Blennechis punctatus); Vizagapatnam, CAS SU41332 (5: 27.2-54.4).

MALDIVE ISLANDS: Seenicappa, FMNH 71357

(44.3). CEYLON: Trincomalee, USNM 205449 (31:

21.2-37.7), 206389 (109: 16.3-55.5), 208275 (23.3).

NICOBAR ISLANDS: GREAT NICOBAR ISLAND:

NFIS 11861 (39.0). BURMA: TAVOY COAST: USNM

89499 (23.6). THAILAND: WEST COAST: Phuket,

USNM 201871 (ca. 42); mouth of Pakchan River,

CAS GVF reg. no. 2208 (48: 21.5-54.3). GULF OF
NUMBER 177 59

THAILAND: west coast (Chumphon Province), Matapora, CAS GVF reg. no. 2184 (3: 57.6-66.4), GVF

reg. no. 2191 (40.7); north coast, Bangkok, CAS

GVF reg. no. 2067 (45.0); vicinity of Choi Buri

(= Chon Buri), CAS GVF reg. no. 1542 (200: 23.0-

71.6), GVF reg. no. 1543 (9: 44.9-64.2); east coast,

vicinity of Chantaburi (= Chantabun), CAS GVF

reg. no. 1586 (3: 22.3-60.2), USNM 119660 (26.0);

Goh Proet (= Kaw Praed?), CAS SU62090 (110: 27.4-

63.7); vicinity of Trat Bay, CAS GVF reg. no. 1487

(5: 38.5-50.1). HONG KONG and vicinity (including Kwangtung Province, China); CAS SU29465

(14: 70.2-95.0), SU62061 (67.4). CHINA (other

than Kwangtung Province); Amoy, BMNH 1860.7.

20.99 (ca. 90, lectotype of Petroscirtes dispar Giinther). CHUSAN ISLAND: Pu-tau, CAS SU32420 (88:

21.8-70.2). JAPAN: NAGASAKI PREFECTURE: Takegaki, USNM 199521 (3: 26.4-52.0). MIYAZAKI PREFECTURE: Nobeoka, USNM 199519 (ca. 76). WAKAYAMA PREFECTURE: Wakanoura, CAS SU7O7O (2:

44.0-49.0, syntypes of Aspidontus dassori). HIROSHIMA

PREFECTURE: Hiroshima, ZITU 24762 (74.8). TOKYO

PREFECTURE: Tokyo, ZITU 31779 (34.8). INLAND SEA

OF JAPAN: BMNH 1905.6.6.347 (52.0). PHILIPPINE ISLANDS: LUZON: Manat (Pangasinan Province), USNM 209347 (2: 47.7-54.5). PALAWAN:

Nakoda Bay, USNM 99388 (36.0). MINDANAO:

AMNH 40019 (32.8). INDONESIA: HALMAHERA:

Teluk Kan, USNM 208278 (2: 24.6-39.5). NEW

GUINEA: Merauke River, ZMA 109.102 (2: 32.8-

42.3). SOLOMON ISLANDS: BOUGAINVILLE: Torokina, DASF FO1929 (39.6). AUSTRALIA: WESTERN AUSTRALIA: Onslow, WAM P4841 (5: 31.4-

48.8), PI 1245 (25.9); Shark Bay, WAM P7257-7306,

in part (2: 31.6-41.7), P7361-32 (2: 38.4-38.7),

P7342 (16: 29.6-61.1). NORTHERN TERRITORY: Yirrkala, USNM 174331 (2: 20.9-33.1); Groote Eylandt,

USNM 174332 (26.2); north of Port Darwin, USNM

174330 (5: 37.8-52.7, including one cleared and

stained). QUEENSLAND: Weipa, AMS IB.5307 (39.7);

Townsville, BPBM 14952 (26: 16.1-71.5); Bowen

(including Port Denison), RMNH 11424 (43.6),

AMS IA.1796 (3: 40.2-55.8); Brisbane and vicinity,

AMS 1.445 (4: ca. 47-78.4), IA.3670 (6: 37.5-65.2),

IA.4597 (ca. 54), IA.4948 (5: 32.5-ca. 67), 1.9871

(76.5), 1.9872 (61.8), 1.9873 (59.1), 1.9874 (3: 34.0-

39.8), 1.9875 (52.1), 1.11365 (2: 45.3-67.9), 1.12616

(2: 64.3-66.9), 1.12694 (2: 52.4-55.0), QM 1.1352

(ca. 48, holotype of Salarias decipiens), 1.1361 (65.8,

lectotype of Salarias helenae), 1.10330 (2: 49.9-64.6),

USNM 209348 (2: 44.0-45.2). FIJI ISLANDS: CANDAVU: NMV 12561 (41.5, holotype of Petroscirtes

semilineatus). PANAMA (Atlantic): Bahia Limon,

SI0 JEM70-63 (3: 39.2-52.1), JEM71-60 (5: 17.7-

27.5), USNM 201439 (44.4), 208276 (10: 37.4-69.4);

Gatun Locks (lowest chamber), GCRL 8784 (2:

43.9-54.2), 9095 (40.3). TRINIDAD: Brighton

Beach, ANSP 53318-21 (4. 42.8-47.6, including unsegregated holotype of Poroalticus sewalli); Mayaro

Point, USNM 204059 (24: 22.0-58.2) Chaguarmus

Bay, UMML 24718 (3: 23.0-55.4); Saline Bay,

USNM 204060 (11: 24.9-36.9). GULF OF PARIA:

BMNH 1933.11.20.12-13 (2: 35.7-63.8).

How O. punctatus BECAME ESTABLISHED IN THE

CARIBBEAN SEA

Omobranchus punctatus is the only Indo-Pacific

marine fish species, other than circumtropical

pelagic forms, known to be established in the western Atlantic. Its known occurrence in the western

Atlantic dates from 30 April 1930, when four specimens were collected at Brighton Beach, Trinidad

(Fowler, 1931, reported as Poroalticus sewalli). Since

1930, O. punctatus has been collected repeatedly

and in large numbers in Trinidad, including points

near the extreme corners of the island. Omobranchus punctatus (as O. dasson) was reported to be

the most common fish around the docks north of

Port of Spain, Trinidad (Lachner, Robins, and

Courtenay, 1970).

Cervigon (1966) reported that one specimen of

O. punctatus (as O. dasson) was collected 23 May

1961 on the shore at Guiria, Gulf of Paria, Venezuela, which is about 75 km west of Trinidad. The

only other known occurrences of O. punctatus in

the Atlantic are in Panama: at Limon Bay, which

is at the entrance to the Panama Canal, and in

the adjacent lower Gatun Locks of the Canal. The

first Panamanian specimen was collected on 3 December 1966 in Limon Bay. Since that date several

more specimens have been taken in Panama, all

from Limon Bay, but the species does not appear

to be nearly so common in Panama as in Trinidad.

The wide area between the Caribbean occurrences of O. punctatus appears to represent a real

distributional void, as there has been continuous

and extensive collecting of Caribbean shore fishes

for many years, and no specimens of O. punctatus,

other than from the Trinidad-Gulf of Paria area
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and Panama, have been obtained. C. E. Dawson,

who obtained the first Panamanian specimen, has

since made numerous marine and estuarine fish

collections in Panama from points approximately

5 km west of the entrance to Limon Bay eastward

to Portobello and some of the San Bias Islands. He

has obtained O. punctatus only from Limon Bay

and the lower Gatun Locks. Omobranchus punctatus also has not been obtained in the eastern Pacific. Aside from extensive fish collecting throughout the tropical eastern Pacific by others, Dawson,

since 1967, has been making fish collections within

an arc of 8 km surrounding the Pacific entrance

to the Panama Canal, into the Canal as far north

as Miraflores locks, and along the Pacific coasts of

Costa Rica and Colombia, and has not obtained

O. punctatus from any of these areas.

The first question that arises concerning the distribution of O. punctatus is whether the Caribbean

population (s) is a relic population of a once

circumtropically distributed species or whether it is

the result of an introduction. We believe that it is

the result of an introduction. The circumstantial

evidence supporting our belief follows.

Omobranchus punctatus is the only tropical Indowest Pacific benthic shore fish that also occurs in

the western Atlantic.2 The blenniids are a relatively speciose group that includes many endemic

populations and species of relatively restricted distribution, indicating a relatively rapid rate of

speciation for the family as a whole. Such rapid

speciation is indicated, for instance, by the fact

that no blenniid species is native to both the

western Atlantic and eastern Pacific oceans, which

were connected most recently two or three million

years ago (Woodring, 1966).3 Nor is there a blen-

'Sauvage (1880) described Salarias brasiliensis from Santa

Catarina, Brazil. Springer (1963) noted that Sauvage's species

was a synonym of the common Indo-west Pacific species,

Salarias fasciatus, and that there was good reason to question

the locality data given by Sauvage. S. fasciatus has not otherwise been reported as occurring outside the Indo-west Pacific.

* Dawson (1970) reported the western Atlantic blenniid

Lupinoblennius dispar from the unused Miraflores Third

Lock on the Pacific side of the Panama Canal, and Hildebrand (1939) reported it, as Blennius sp., from Gatun Locks

on the Atlantic side. Dawson believed, as do we, that its

presence in these locks was the result of immigration through

the Canal from the Caribbean. J. C. McCosker informs us

that he has taken the Atlantic blenniid Hypleurochilus

aequipinnis in the Miraflores Third Lock. We postulate a

niid species common to the eastern and western

Pacific Ocean. Furthermore, all species of the

blenniid tribe Omobranchini, other than O. punctatus, are restricted to the Indo-west Pacific. If a

species of Indo-west Pacific blenniid was successful

in maintaining a naturally disjunct population in

the western Atlantic, one should reasonably expect

similar distributions for other Indo-west Pacific

benthic species, yet there are no other similar distributions. Therefore, the presence of O. punctatus

in the western Atlantic must be the result of an

artificial introduction.

The presence of O. punctatus in Panama is probably the result of an artificial introduction from

Trinidad, but first we must discuss the possible

method by which O. punctatus was introduced into

Trinidad.

Dawson (1973) presented a good case for the

method of introduction of the Indo-west Pacific

small, benthic, gobioid fish Prionobutis koilomatodon into the eastern Pacific (Panama Canal): in

ship's ballast water, which was emptied into the

Canal. Ballast water is preferable to bilge water as

a means of hypothesized transport by motorized

vessels because bilge water is mainly derived from

avenues that would permit entry into the ship of

few, if any, marine organisms, and bilge water is

usually too foul and oily to support live fishes.

Ballast water, in contrast, is drawn through a

strainer, with perforations about one-half inch in

diameter (although the diameter often greatly increases through corrosion) directly from the sea

into a ship's relatively clean ballast tanks (Melvin

Jackson, personal communications; however, Rubinoff, 1970, noted that modern nonstainless steel

ballast tanks may be coated with highly toxic anticorrosion paints). Each ship usually has two ballast

water intakes: one located on the side below the

waterline and the other on the bottom of the hull.

Both bilge and ballast water would offer possibilities for fish transport on sailing ships, the bilges

of which are generally cleaner than those of motorized vessels. Sailing ships rarely used water for ballasting (Melvin Jackson, personal communication),

similar immigration for this species. Rubinoff and Rubinoff

(1968) believed that the presence of the western Atlantic

gobioid, Lophogobius cyprinoides, in the Miraflores Third

Lock was the result of colonization from the Atlantic through

the Canal.
NUMBER 177 61

but as early as 1901, at least, water ballast was used

on some sailing vessels (Lubbock, 1935).

The ballast-bilge water method of introduction

probably applies also to the Trinidad population

of O. punctatus and to the two other New World

(eastern Pacific) introductions of Indo-Pacific marine fishes (Brittan, Albrecht, and Hopkirk, 1963;

Lachner, Robins, and Courtenay, 1970), which are

both gobioids (Acanthogobius flavimanus, Tridentiger trigonocephalus) and relatively small benthic

fishes. Small size is favored for introduction by ballast water (and probably bilge water also) because

of the small size, at least initially, of the holes in

the ballast strainer. Omobranchus punctatus would

be a good candidate for being sucked into a ship's

ballast tanks because of its small size, abundance

around docks (at least in Trinidad; Lachner, Robins, and Courtenay, 1970), and because of the general inclination of blenniids, including Omobranchus, to inhabit tubelike holes. Omobranchus might

well be expected to enter the holes in the ballast

strainer of a ship that had been resting in port,

and then be sucked into the ballast tanks when the

pumps were started.

Inasmuch as O. punctatus has been taken in the

Caribbean and not in the eastern Pacific it seems

unlikely that its introduction into the Caribbean

was effected through the Panama Canal via a transPacific crossing. Ships intending to clear their bilges

and ballast tanks must do so before entering the

Canal (Rubinoff, 1970), and although some ships

illegally discharge bilge and ballast after entering

the locks (Dawson, 1973), it is most unlikely that

they would wait until the end of the traverse to do

so. More likely they would discharge at the beginning of the traverse. This being the case, one would

expect to find O. punctatus at the Pacific side of the

Canal had it arrived via a Pacific crossing. On the

other hand, one would expect a ship entering the

Canal from the Caribbean side to discharge its

bilge or ballast in Limon Bay (just prior to making

its traverse) or in Gatun Locks (just after entering

the Canal). Omobranchus punctatus in Panama is

most common in Limon Bay, where it has been

collected repeatedly. It has been taken in only one

collection in Gatun Locks and only three specimens

were obtained (indicating that the lock environment is not particularly suitable, that periodic

emptying of the lock for cleaning decimates the

population, or that there has been insufficient time

or introductions for the population to have developed large numbers). Another consideration that

would argue against O. punctatus' having entered

the Caribbean through the Panama Canal is the

probability that it was introduced into the Caribbean prior to 16 August 1914, the date that the

first ship passed through the Panama Canal; our

reasons for this assumption will become apparent

below.

Hildebrand (1939) reported on collections of

fishes obtained from Gatun Locks (Atlantic side)

in 1935 and Pedro Miguel and Miraflores locks (Pacific side) in 1937, when these locks had been dewatered for their periodic cleaning. Omobranchus

punctatus was not present in his collections. The

evidence thus favors the introduction of O. punctatus in Trinidad (first appearance in collections

in 1930) before its presence in Panama (first appearance in collections in 1966) and to some extent

supports our belief that the Panama population is

a derivative of the Trinidad population. The very

close averages for various meristic characters (Table

13) and the uniquely low averages in common for

one of these characters, total number of procurrent

caudal-fin rays, exhibited by the Trinidad and

Panama populations of O. punctatus also point to

a common origin for the two populations. (The

relationship of these data to those of other populations of O. punctatus is by no means clear, but

certainly seems to preclude derivation from some

populations: China, Hong Kong, Australia. In general, the averages are close to those of specimens

from western Thailand and, perhaps, Ceylon,

which are relatively close geographically to the area

we will propose below as the source of the Caribbean populations.)

If one accepts the proposition that O. punctatus

did not arrive in the Caribbean via the Panama

Canal, then it must have arrived by passage across

the Atlantic Ocean. Such passage could have been

made either around the southern tip of Africa or

via the Red Sea, Suez Canal, and Straits of Gibraltar. In the latter case its arrival could not have

been earlier than 1869, the year the Suez Canal was

opened. In either case a locality in the Indian

Ocean portion of the distribution of O. punctatus

appears to be the most probable source of the

parental population of the Trinidad introduction.

There are two main reasons for such an assumption. The first is proximity: survival during trans-
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

port is dependent on, among other things, the

length of time spent in transport, and Indian Ocean

localities are closer to Trinidad than are Pacific

Ocean localities (considering the direction of transport proposed above). The second reason concerns

the commerce in East Indian coolie labor that occurred between 1838 and 1917 (Erickson, 1934). It

is for the second reason also that we believe the

Trinidad population of O. punctatus originated at

Madras or, more probably, Calcutta.

With the 1833 abolition of slavery in the British

Empire, there developed a pressing demand by

many of the British colonies for emigrant laborers.

This demand was met by the importation of indentured east Indian coolies from Calcutta and, to a

lesser extent, Madras on the east coast of India.

The first Indian coolies sent to the New World

arrived in British Guiana in 1838. Because of cruel

practices the importation of coolies was suspended

after 1838, but recommenced in 1844 after new

safeguards for the coolies had been established.

From 1845 until 1917, when immigration was

suspended, there was a great flow of Indian coolies

into British Guiana, Trinidad, Jamaica, and, to

much lesser extent, other Caribbean Islands. Of

these areas, British Guiana received the most coolies, approximately 240,000 (Gangulee, 1947); Trinidad received 143,900 (Klass, 1961); and Jamaica

received 36,400 (Roberts, 1957). These numbers

reflect the total number of arrivals only and not

the greater number of coolies that initially embarked on the ships, because many died along the

way (for instance, see Swinton and Swinton, 1859,

for the account of one voyage from Calcutta to

Trinidad during which 120 of 324 coolies died

before reaching Trinidad; this may be an extreme

case; for other, lesser, mortalities see Lubbock,

1935).

As far as we can determine, a sailing vessel

(which was the type predominantly used) carried

an average of about 380 coolies (range 269-515;

Lubbock, 1935), but at the end of the period of

immigration the steamers were carrying about 1000

coolies (Lubbock, 1935; although Klass, 1961, reported that the last steamship to carry coolies to

Trinidad arrived in 1917 with only 394 coolies).

Using the average figures and allowing for a most

conservative 5 percent mortality, it would have

required 400 shipments to bring the coolies to

Trinidad. The means for transport of O. punctatus

to Trinidad were, thus, abundantly available.

Coolie immigration into Trinidad (and elsewhere in the Caribbean) was essentially terminated

in 1914 (because of World War I) and ended in

April 1917 (Klass, 1961; Roberts, 1957), thus accounting for our belief that the introduction of O.

punctatus into Trinidad occurred prior to the 1914

opening of the Panama Canal. In view of the probable introduction of O. punctatus into Trinidad

prior to the opening of the Canal, the similarity in

meristic characters shared by the Trinidad and

Panama populations, the fact that O. punctatus is

much more common at Trinidad than at Panama,

the gap in distribution between Trinidad and Panama, and the probable means of ship transport of

O. punctatus, we believe that the occurrence of O.

punctatus in Panama is the result of a ship's acquiring the species in Trinidad, proceeding to the

Canal, and discharging its acquisition at the entrance to the Canal just before making the Canal

traverse. (We should interject here that although

O. punctatus has not been reported from British

Guiana we would not be surprised to find it there.

Few, if any, shore fish collections have been made

in British Guiana. Jamaica has, however, been well

collected; see Caldwell, 1966.)

From the initiation of the Indian coolie emigration until about 1910, only sailing ships were used

to transport the coolies (the 1910 date is inferred

from data in Lubbock, 1935). Their trip to the

New World was made only during the warm months

and followed a route around the southern tip of

Africa with one stop, St. Helena (Swinton and

Swinton, 1859; Lubbock, 1935). The Suez Canal

route was not used. When steamers replaced the

sailing ships they followed the same route (Lubbock, 1935). As far as we can determine, the sailing

ships required 69-101 days to make the journey

from Calcutta to British Guiana or Trinidad (Lubbock, 1935; Swinton and Swinton, 1859). The steamers were probably able to halve the transit time.

While speed and preferred method of ballasting

would favor a steamer as the vessel for introducing

O. punctatus to Trinidad, the much larger number

of sailing vessels making the crossing might favor a

sailer.

(It is perhaps relevant to note here that a specimen of a blennid, Petroscirtes breviceps [DASF

FO213O], was taken from the bilge of a ship that

traveled from northwest Australia to Port Moresby
NUMBER 177 63

in September 1972. The data with the specimens

indicates that there was a continuous flow of seawater in the ship's hold.)

In postulating that the origin of the Trinidad

population of O. punctatus came from Madras or

Calcutta, we are faced with the difficulty of a lack

of records of O. punctatus from either of these two

localities. This difficulty may be more apparent

than real, as O. punctatus has been collected from

Ceylon, which is south of Madras, from Vizagapatnam, which is between Madras and Calcutta, and

from the Tavoy coast of Burma, which is east and

south of Calcutta. The east and northeast coasts of

India have not been well collected for fishes, and

there is no reason to believe that O. punctatus does

not occur at one or both, Calcutta or Madras.

Mauritus, Ceylon, southern Africa, and Fiji also

received large numbers of Indian coolies from

Madras and Calcutta (Gangulee, 1947) during the

same period that coolies were arriving in the New

World. For this reason alone, one must generally

hold in suspect the naturalness of the apparent

distributions of small, benthic shore fishes in the

Indo-west Pacific. It is possible, for instance, that

the isolated populations of O. punctatus and O.

ferox at Delagoa Bay (southeast African coast) are

the result of man-induced introductions. J. L. B.

Smith (1959) reported that O. japonicus (= punctatus) was taken "nowhere else in the whole W. Ind.

Ocean" despite careful search, and neither O. punctatus nor O. ferox have since been taken elsewhere

in the western Indian Ocean.

The occurrence of O. punctatus at Candavu (Fiji

Islands) must also be questioned. This record is

based on Kner's (1868a) description of Petroscirtes

semilineatus, from one specimen, which was given

at the same time as his description of P. lineolatus,

also based on one specimen from Candavu. Candavu is well removed from the verifiable distribution of both these species, and neither species has

been retaken at Candavu or Fiji, in spite of considerable recent collecting. Furthermore, Kner received many of his specimens from boat captains,

and it is quite possible that the localities were

erroneously recorded. Perhaps the specimens came

from the boats themselves, which had acquired

them in another port only to be noticed after arrival in Candavu, but we shall never know (see

also "Nomenclatural Discussion" under O. lineolatus).

Omobranchus rotundiceps (Macleay)

(For synonymy see subspecies)

COMPARISONS.—Omobranchus rotundiceps is a

member of the elongatus species group (see "Comparisons" under O. elongatus). Omobranchus rotundiceps is the only species in the group with

modally 12 dorsal-fin spines; the others have modally 13 spines. It further differs from the other

species in the group in having the dark spot posterior to the eye either closer to the preopercular

series of pores than to the circumorbital series or

about midway between the two series. In the other

species the dark spot is either absent or closer to

the circumorbital series of pores. Aside from its

species group characters, O. rotundiceps is separaable from most of the other species of Omobranchus only by a combination of characters.

Omobranchus rotundiceps rotundiceps (Macleay)

FIGURES 28, 29a,b

Petroscirtes fasciolatus Macleay, 1881:8 [Port Jackson; secondary junior homonym of Blennechis fasciolatus Ehrenberg in Cuvier and Valenciennes, 1836, which also belongs

in Omobranchus].

Petroscirtes rotundiceps Macleay, 1881:9 [Port Jackson; lectotype designated below].

Salarias furcatus DeVis, 1884b:696 [St. Helena, Moreton Bay].

Salarias furtivus DeVis, 1886:60 [Moreton Bay].

Petroscirtes macleayi Ogilby, 1887:38 [replacement name for

Petroscirtes fasciolatus Macleay].

Graviceps angelus Whitley, 1959:320 [Heron Island, Capricorn Group, Queensland].

DESCRIPTION (see also Table 15).—Dorsal fin XIXIII (XII in 85.2% of specimens), 18-24 = 30-36

(rarely 30); anal-fin II, 20-26 (rarely 20); both analfin spines of males discernible externally; segmented caudal-fin rays 11-14 (13 in 95.2% of specimens;

11 in only 1 of 230 specimens); dorsal + ventral

procurrent caudal-fin rays 10-16; vertebrae 10-11 +

27-33 (rarely 27 or 33) = 37-43 (rarely 37); epipleural ribs 11-14 (rarely 11); prenasal pores present; interorbital pores 1-4 (3 in 96.8% of specimens);

circumorbital pores 6-9 (8 in 94.7% of specimens;

6 in only 1 of 251 specimens); lateral-line tubes 0-5

(rarely 4 or 5); lateral-line tubes, when present, extending posteriorly to below level of dorsal-fin

spine 1-6 (rarely 5 or 6); gill opening varying from

restricted to area dorsal to level of dorsalmost
TABLE 15.—Frequency distributions for certain characters in subspecies and

populations of Omobranchus rotundiceps

Subspecies •

populatio!

Dorsal-fin

spines

Segnented dorsal-fin

Epipleural ribs

18 19 20 21 22 23 2k 30 31 32 33 34 35 36 1 11 12 13 11 15

Philippine Islands

Guam

Yap

Falau Islands

Ambon

Timor

Hew Guinea

lev Caledonia

Vanikoro

Aneityum

FIJI Islands

Samoa Islands

Palmyra

Hawaiian Islands

Harthern Territory

•estern Australia

Montebello

Exmouth Gulf

Shark Bay

Cottesloe

Que ens 1 **TT1

Townsville

Port Bowen

Cumberland Islands

Beron Island

Gladstone

Moreton Bay

Hew South Wales

15 .

1.7

16

6k

2

1

k

1

It

2 0

k

26

- 11.9

3 12.0

- 11.9

- 12.0

- 12.0

- 12.0

- 12.0

- 12.0

- 12.0

- 12.0

1 12.0

- - 19.0

- - 19.0

- - - 18.6

- - - 18.8

- - 19.0

- - - 19.5

5

3

1

9

2

1

3

11

12

12

.9

. 2

. 2

12 63 75 16

It 12 - 11.8

20.0

19.9

19-5

19.2

19.2

- 20.5

- 21.8

1 22.9

- 22.2

It Ik 2 -

4 12 72 77 5 -

30.9

31-0

30.5

30.7

31.0

30.8

32.0

31.9

31.5

31.2

31.2

- 32.k

- 3k.O

3 35.2

- 33.9

19 57 3 2.8

3 Ik - 2.6

53 - 3.0

2 - 3.0

5 3.0

- 3.0

2 31 110 2k

1 3

8 3

2 12 3

- 13.2

- 12.8

- 13.0

- 12.7

- lk.O

- 13.1

- 12.7

12.9

13.8

13.3

13.0

13.5

Subspecies and

population

0. r. obliquus

Philippine Islands

Guam

Tap

Palau Islands

Ambon

Timor

Hew Guinea

Sew Caledonia

Vanikoro

Aneityum

Fiji Islands

Samoa Islands

Palmyra

Hawaiian Islands

0. r. rotundicep.

Horthern Territory

Western Australia

Montebello

Fxmouth Gulf

Shirk Bay

Cottesloe

Queensland

Townsville

Port Bowen

Cumberland Islands

Heron Island

Oladstone

Moreton Bay

Hew South Wales

Subspecies and

population

0. r. obliquua

Philippine Islands

Guam

Tap

Palau Islands

Timor

Sew Guinea

He. Caledonia

Vanikoro

Fiji Islands

Samoa Islands

Palmyra

Hawaiian Islands

Northern ^rritory

Western Australia

•tante bello

Sxsouth Gulf

Shark Bay

Cottesloe

Queensland

Townavllle

Port Bowen

Cumberland Island

Heron Island

Gladstone

Moretjn Bay

«Vw South Wales

Preca

1 0

16

52

17

65

2

k

1

k

2

2 9

2

26

k

16k

1

1

2 0

1

2

1

1

8

16

11

-

1

.

_

1

-

1

-

k

3

1

11

1

-

2

2

idal

crae

5

3

1

3

S

10.0

10.0

10.0

10.0

10.0

10.0

10.0

10.0

10.0

10.0

10.0

10.0

10.0

10.8

10.9

10.0

10.0

10.0

10.1

Se

Ik

38

10

k9

2

1

It

1

11

It

10

2 7

18

12

2 1

2

1

1

1

5

gmsn

22

3

15

2

13

3

2

Ik

1

15

1

60

28

15

33

k2

2

3

1

6

1

18

k

9

Caudal v

29

1

1

2

2

1

.

1

23

1

6

1

77

2

30

-

-

-

1

.

1

-

68

5

3

k

ted anal-fin

ray

23

*

-

1

.

1

k

-

91

1

2

3

3

2k

-

-

-

-

6

3

1

8

12

1

2

2

5

9

ertebrme

31 32

-

-

-

-

8

3

1

6

12

1

2

1

1

5

6

-

-

-

-

6

1

3

-

5

_

-

k

1

2

.

1

.

5

- 21

- 21

- 20

- 21

- 21

2

3

8

2

8

- 21.0

• 22

• 21

2

5

- 22.5

- 23.8

- 2k

- 2k

- 2k

- 2k

- 23

1 2k

3

0

0

0

.6

. 2

33

_

-

-

_

-

1

X

28.1

27.7

27.3

27.7

28.5

27.8

28.5

28.8

28.5

28.3

28.2

29. k

31.0

31.5

3O.o

31.0

30.6

30.9

Dorsal

1 0

13

2

6

k

37

18

12

21

1

k

• ve

38

15

33

k2

1

2

3

1

5

1

19

3

8

Total V.

39 kO

1

1

2

2

1

.

1

2k

1

6

1

_

I

.

_

-1

.

1

-

77 69

1

fttral proc

caudal-fl

17

k

12

7

18

1

1

-

5

16

7

2k

1

1

2

k

33

1

3

1

6

5

k

k

22

1

3

1

1

2

kk

2

1

k

2

7

1 rays

5 2

1

1

3 -

1

59 12

2

10 -

1

5 1

1

1

3 -

1

5

3

k

ir

rtebrae

kl k2

- -

-

-

-

_

- -

10 -

1 2

1

7

12 . 3

1

2

1

1

5 -

7 5

•ent

- 13-2

- 11.3

- 12.1

- 12.2

- 12.8

- 12.8

- U.5

- 13.0

7 13.k

- 13.5

- 13.9

- 13.3

1 13-7

• 12.1

k3

„

-

I

_

_

_

-

-

5

1

.

1

k

2

1

10

1 0

X

38.1

37.7

37.3

37.7

37.0

38.5

37.8

38.5

38.9

38.5

38.3

38.2

39.k

111.7

k2.k

k0.9

k0.6

kl.O

3

3 0

k

Ik

2

1

1

1

117

3

8

1

7

1

7

8

0

k6

I

6

Ik

7

25

5

13

1

-

3

1

53

3

3

1

-

Lower end of gil l oper

opposite pectoral-fin

1

37

I- 1

a

16

3

2 0

1

-

8

12

1

1

_

1

-

7

3

1

k

5

k

3

m

-

3

3

1

-

2

2

3

k

1

2

2

2

32

k

1

_

1

1

2

1

-

-

3

6

9

3

10

2

5

9

2

7

1

1

1

_

-

2

-

-

k

3

17

2k

1

1

2

1

12

2

I

1

_

-

2

8

a.k

2

1

2

1.2

3

3

1

1

1

1

1

0

k

k

5

k

3

. 2

o.k

5 6

. .

19 -

12 2

_ _

1

1

1 -

2 -

- -

1 -

_ _

- "

Las

I

k -

2 -

1 -

10 1

10 -

ing

ray

7 8

- 1

. _

- -

X

3-1

k . 2

3-5

k.O

3.3

3.2

3.6

2.8

1. 0

1.5

O.k

0 . 2

lateral-li

do

k

33

3

16

2

1

1

2

2

1

121

3

7

1

7

2

1

7

7

•sal-fit

6

21

5

12

1

-

3

1

3

k9

1

k

3

1

2

Ik

1

18

1

-

7

13

.

1

„

1

1

Clrcumorbital

pores

6

1

K t

7

1

L3

1

2

1

"

lbe

spine

c C

J

.

7

2

1

_

-

2

1

.

1

_

-

1

1

5

k

1

3

1

-

8

13

67

15

52

1

k

2

2

18

1

25

5

187

1

11

1

2

1

8

be

7

-

2

-

-

3

-

1

1

_

-

k

1

.

3

ow

1

2 .

2 .

k.

3.

2

"

2 .

k.

2.

k.

2 .

2 .

2 .

2 .

2 .

2.

2 .

2 .

9

6

0

0

5

0

0

o

k

3

2

3

2

5

3

c

2

0 . 9

X

7.

7.

3.

3.

8.

• >

8

0

0

8 . 0

8.

8.

8.

8.

8.

g

8.

8.

8.

0

0

0

0

0

o

1

o

0

8 . 0
NUMBER 177 65

pectoral-fin ray to extending ventrally to opposite

5th ray (rarely extending below level of 3rd ray);

lower-lip flap present; circumorbital bones 5; lower

jaw teeth 18-31; upper jaw teeth 16-29 (Figure 45);

no fleshy bladelike crest on top of head of either

sex (head of some presumably mature males swollen

in appearance).

COLOR PATTERN (described first from northern

and western Australian specimens exhibiting most

developed pattern, followed by comparison with

eastern Australian specimens).—Males: Head: Narrow, dark band extending anteroventrally from

anteroventral margin of orbit across mouth, just

anterior to corner of jaws, to chin, confluent across

chin with corresponding band of opposite side.

Four slightly to considerably diffuse, posteroventrally slanted, narrow, dusky bands posteriorly on

lower half to third of head and prepelvic area;

anteriormost band extending from below posteroventral edge of orbit to ventral surface of head,

usually with interruption and slight anterior shift

of ventral part of band at preopercular series of

pores; succeeding bands evenly spaced, paralleling

1st band; each band originating slightly higher on

side of head than preceding band; 3rd and/or 4th

band arising dorsally from diffuse, dusky area on

operculum and continuing onto perpelvic area;

additional (5th) small band originating on operculum and continuing onto prepectoral area; anterior bands usually confluent with corresponding

bands of opposite side at ventral midline of head;

posterior bands frequently interrupted on ventral

surface of head. Large, oval to dome-shaped, dark

area, approximately size of eye, present posterior to

eye between circumorbital and preopercular series

of sensory pores; dome-shaped area encircled by

narrow, pale margin, which is less distinct ventrally; intense dark spot, equal to or smaller than

pupil of eye, at dorsoposterior corner of domeshaped area. Free opercular membrane pale. Predorsal area dark dusky with scattered dark spots in

some specimens.

Trunk: Ten to 12 broad, ventrally tapering, dark

dusky bands present on side of body; anterior 2 or

3 bands merging together to form dark dusky area;

posterior 4 or 5 bands each represented as midlateral spot, or bands (spots) joined together to form

narrow, midlateral stripe (especially in large specimens); anterior 5 or 6 bands slanted posteroventrally, usually not extending onto upper quarter of

body; all bands terminating dorsal to ventral body

contour; pigment of each band often narrowly

intensified along central, vertical axis of band; some

specimens with bands represented only as dark, midlateral blotches (in freshly preserved specimens,

midlateral portion of bands more intense, forming

interrupted midlateral stripe extending onto head);

posterior bands and midlateral spots with slender

dorsoposterior extensions; approximately 9th band

(spot) prolonged dorsally into narrow, posteriorly

extending stripe dorsal to midlateral stripe; both

stripes directed ventrally at caudal-fin base and

continuing onto fin. Adjacent body bands, posterior

to 2nd or 3rd band, each separated by vertical,

well-defined, slender, pale line; pale lines nearly

reaching ventral body contour, terminating dorsally

at level of dorsal ends of body bands, or directed

sharply anteriorly anterior to about 6th body band,

or angled posteriorly posterior to about 7th body

band; free longitudinal pale lines sometimes present above dorsal ends of vertical, pale lines. Series

of approximately 8 pairs of dark, dorsally open

half-rings along base of dorsal fin; rings of each

anterior pair joined. Dark peritoneum visible

through abdominal wall.

Pectoral Fins: Slightly dusky. Fleshy base with

dark band midway between opercular edge and

bases of rays; vertically oriented pair of dark

spots or faint bar at base of fin rays.

Pelvic Fins: Pale.

Dorsal Fin: Transparent with dark markings.

Spinous portion with 3 pairs of broad, dorsoanteriorly directed or vertical bands; each pair above

corresponding pair of dark, dorsally open halfrings. Segmented ray portion with approximately

8 narrow, dark stripes; tips of rays abruptly pale;

large, diffusely dusky to black, oval spot on distal

portion of fin in area of 10th to 16th (usually centering on 14th or 15th) segmented rays. Large specimens with spinous portion dusky except for 2 large,

pale spots near distal edge: 1 near central spines,

1 near posteriormost spines; segmented ray portion

of fin dusky near distal edge, transparent proximally; black, oval spot apparent.

Anal Fin: Dusky to dark distally, transparent

proximally. Tips of rays abruptly pale.

Caudal Fin: Pale with 2 narrow, dark stripes

directed posteriorly and slightly ventrally from

posterior ends of midlateral and more dorsal pos-
66

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

terior body stripes (each stripe sometimes intensified as dark spot at caudal-fin base).

Females: Similar to males except: midlateral intensifications of body bands darker, generally

broader, better defined, reaching farther ventrally;

dusky portion of bands surrounding intensifications

reduced, fainter; areas between posterior bands on

lower half of body with additional intense, dark

bands or spots; dorsal extensions of posterior bands

(spots) broken into spots; posterior midlateral

stripe and more dorsal stripe indicated only by 2

longitudinal series of spots; body more mottled or

spotted dorsally; dorsal fin less pigmented, especially distally; dark bands of spinous dorsal fin, if

present, much narrower; black, oval spot and all

but basal dark stripes of segmented ray position

of dorsal fin absent; stripes on caudal fin better

defined or present as intense, dark streaks.

Eastern Australian Specimens: Head bands narrow, broken into dashes or irregular series of spots,

often completely replaced by large number of fine

spots and speckles (Figure 29a,b); complete head

bands, when present, usually occur only posteriorly.

GEOGRAPHIC VARIATION (see also "Color Pattern").—Population variation was noted in most

meristic characters of O. rotundiceps, both within

a subspecies and between subspecies (Tables 15

and 16). The meristics of O. r. rotundiceps generally ranged and averaged higher in number than

they did in O. r. obliquus. The northern Australian

population of O. r. rotundiceps, which are geographically closest to the population of O. r. obliquus, are intermediate in meristics between the

two subspecies (see also "Color Pattern" under O. r.

rotundiceps).

The Western Australian populations of O. r.

rotundiceps exhibit increases in numbers of precaudal, caudal, and total vertebrae. Similar increases

are exhibited by the Western Australian populations of O. germaini, whereas the Western Australian populations of O. punctatus exhibit an increase

in the number of precaudal, but not caudal and

total, vertebrae. Increase in number of precaudal

vertebrae is not encountered in any other species of

Omobranchus. It is noteworthy, even if presently

inexplicable, that three species of Omobranchus

exhibit increases in number of precaudal vertebrae

in about the same geographic area.

Although covariance comparisons of numbers of

jaw teeth (Table 16) were not feasible for all populations of O. rotundiceps, almost all the tests that

were performed between pairs of populations were

either significant or almost significant.

GEOGRAPHIC DISTRIBUTION (Figure 5).—From

Cottesloe (near Perth), Western Australia, northward around Australia to Sydney, New South Wales.

HABITAT.—Shallow marine and brackish water;

in tide pools, around ironstone reefs, rocks, and

coral. Most often taken in marine habitats.

COMPARISONS.—Omobranchus r. rotundiceps is

completely separable from O. r. obliquus only on

the nature of the dark spot posterior to the orbit.

In O. r. rotundiceps this spot is much closer to the

preopercular series of pores than to the circumorbital series (Figures 28a and 29a) and consists of a

concentration of pigment along the posterodorsal

portion of a dark dusky, dome-shoped area. In O. r.

obliquus the spot is about midway between the

circumorbital and preopercular series of pores and

consists of a concentration of pigment over the

dorsal portion of a dark dusky, dome-shaped area

(Figure 29c). Omobranchus r. rotundiceps may have

the color pattern on the underside of the head consisting of about 3 or 4 narrow, dark bands (Figure

286; northern and Western Australian populations)

or of a group of dark spots and slender bands (Figure 29b; eastern Australian populations). In O. r.

obliquus the pattern under the head consists of

dark, slender bands (Figure 29d). Where populations of the two subspecies are geographically closest, there is broad overlap in all meristic characters

(Table 15).

NOMENCLATURAL DISCUSSION.—Macleay (1881)

did not indicate the number of specimens on which

he based his description of P. rotundiceps, and

there is no indication that he had more than one

specimen as he mentioned no variation in characters. The size was given as 2.5 inches, which might

indicate a single type-specimen. The putative types,

formerly in the Macleay Museum, are now cataloged into the Australian Museum collections.

There are two conspecific specimens, 50.7 and 36.6

mm SL. Only the larger of these two is near enough

in size to be considered as a type. This specimen,

which we here designate the lectotype of P. rotundiceps, is now cataloged as AMS 1.16413-001. The

smaller specimen is now cataloged as AMS 1.16413-

002.

DeVis (1884b) did not compare his Salarias furcatus with any other species. The putative holotype
NUMBER 177 67

TABLE 16.—F values for covariance comparisons of regression equations

(Table 2) for number of lower or upper jaw teeth between males or

females from different populations of Omobranchus rotundiceps (** =

significant at p = .001; NS = not significant)

Comparisons

Lower teet h

males

Northern Territor y X

Northern Territory A

Palau Is . X Guam

Northern Territor y X

Northern Territor y X

Northern Territor y X

Palau In . X Guam

Palau Is . X Palmyra

Guam X Palmyra

Upper tent h

males

Northern Territor y X

Northern Territor y X

Palau Is . X Guam

females

Northern Territor y X

Northern Territor y X

Guam X Paln\yra

Palau Is.

'luam

Palau Is.

Guam

Palnyra

Palau Is.

Guam

Guam

Palmyra

F value s

slopes

0.23 NS

3.92 NS

3.92 NS

0.85 NS

1.38 NS

0.78 NS

0.08 NS

0.10 NS

C.01 NS

0.28 NS

13.63 **

14.86 **

0.32 NS

1.46 NS

0.30 NS

Degrees of

freedom

1/59

1/64

1/37

1/50

1/56

1/50

1/34

1/28

1/34

1/59

1/65

1/36

V55

1/49

1/34

F values

heights

28.69 * *

6.65 NS

7.02 NS

31.22 **

7.31 NS

7.84 NS

10.19 NS

7.12 NS

0.06 NS

20.03 *»

(23.94)

(0.00)

36.79 **

45.74 **

0.17 NS

Degrees of

freedom

1/60

1/65

1/38

1/51

1/57

1/51

1/35

1/29

1/35

1/60

1/66

1/39

1/56

1/50

1/35

(QM 1.11/114) is a female O. r. rotundiceps, and

although its characteristics do not all agree with

those given in DeVis' description, the color pattern

of the head given in the description is particularly

distinctive of O. r. rotundiceps. On the basis of the

description and the identity of the putative holotype, we synonymize S. furcatus with O. rotundiceps.

DeVis (1886) also failed to compare his Salarias

furtivus with any other species. The description

alludes to at least three specimens (from Moreton

Bay), which must be considered as syntypes, and

could pertain to either O. r. rotundiceps or O. germaini. There is no type-material of S. furtivus at

the QM, where DeVis deposited his specimens.

However, DeVis (1887) mentioned that five species

of fishes (comprising 75 specimens) were exchanged

with the Australian Museum in 1886. There is no

correspondence at the AMS concerning the exchange, but AMS 1.383-4 (five specimens) was cataloged in 1886 and listed as "cotypes" of S. furtivus

(see also McCulloch, 1929:341). Considering the

year the specimens were cataloged, the identification given them, and the mention in the report by

DeVis (1887), we consider the specimens to be syntypes of S. furtivus. The five specimens comprise

four identifiable as O. r. rotundiceps and one as

O. anolius. We here designate one of the O. r. rotundiceps specimens as lectotype of Salarias furtivus,

which is now cataloged as AMS 1.16728-001. The

other three specimens of O. r. rotundiceps are now

cataloged as AMS 1.384, and the specimen of O.

anolius as AMS. 1.383. The lectotype is a male,

distinguishable from all the other syntypes in having three lateral-line tubes on the left side, as opposed to none or one tube, and in having the

posteriormost dorsal-fin ray represented only by a

fragment of its base (distinguishable only on a

radiograph), as opposed to the posteriormost ray

being present in its entirety in the other specimens.

Petroscirtes macleayi Ogilby is a replacement

name for Petroscirtes fasciolatus Macleay, which is

preoccupied. Neither Macleay nor Ogilby compared

the species with any other. The type-material,

which was originally at the Macleay Museum but

is now deposited at AMS, comprises 12 syntypes, all

identifiable as O. r. rotundiceps. We here designate

one specimen, a male, as lectotype. The lectotype

is now cataloged as AMS 1.16412-002. It is distinguished from all the other syntypes, catalogued as

AMS 1.16412-001, in having the anal-fin rays II, 26,

as opposed to II, 24-25, and from the other four

males in having the dorsal-fin rays XII, 23, as o p

posed to XII, 21-22.

Whitley (1959) differentiated his Graviceps angelus from "Petroscirtes elegans and congeners" on

the basis of unspecified "fin counts and color pat-
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tern" and from its supposed nearest relative, Petroscirtes kranjiensis, on proportions and mouth position. The holotype of G. angelus is a male O. r.

rotundiceps. Whitley reported the dorsal-fin formula as 13, 21, whereas we find it to be XII, 22, and

the pectoral-fin rays as 14, whereas we find 13 in

each fin.

MATERIAL EXAMINED.—AUSTRALIA: WESTERN

AUSTRALIA: Montebello, BMNH 1961.8.16.71-73

(37.2), 1961.8.16.74-75 (2: 35.0-39.0), 1961.8.16.74-

77 (36.9); Exmouth Gulf, WAM PI 1141 (34.4);

Shark Bay,- WAM 8148-56 (9: 26.2-49.4), P7649

(51.4), WAM uncataloged (2: 33.4-34.6); Cottesloe,

WAM P6300 (31.3). NORTHERN TERRITORY: 7 miles

north of Darwin, USNM 174263 (42: 18.9-48.6);

East Alligator River at Cahill's Landing (Van Diemen Gulf), USNM 174336 (22.7); vicinity of Cape

Arnhem, USNM 174337 (18: 28.1-48.2), 174339

(139: 18.9-48.5), 174344 (5: 28.9-37.5); Groote Eylandt, USNM 174334 (8: 24.2-37.7), 174340 (29:

20.6-43.7), 174341 (107: 21.4-65.6, including three

cleared and stained). QUEENSLAND: Townsville,

BPBM 15020 (20: 13.9-60.5); Pt. Bowen, BMNH

1871.9.13.192 (74.2); Cumberland Islands, AMS

IB.5826 (48.1), IB.6004 (36.3); Heron Island, AMS

IB.3995 (47.5, holotype of Graviceps angelus),

IB.4057 (38.8); Gladstone, AMS IA.6093 (29.3);

Moreton Bay, AMS 1.445 (ca. 50), 1.16728-001 (ca.

42, lectotype of Solaris furtivus), 1.384 (3: ca. 39-

45), ANSP 98718 (68.7), MCZ 38548 (51.9), QM

1.11/114 (35.5, holotype of Salarias furcatus DeVis).

NEW SOUTH WALES: Newcastle, USNM 148633

(55.8); Port Jackson, AMS 1.16412-002 (51.9, lectotype of Petroscirtes fasciolatus Macleay and Petroscirtes macleayx), 1.16412-001 (11:. 35.6-51.8),

1.16413-001 (50.7, lectotype of Petroscirtes rotundiceps), 1.16413-002 (36.6); Sydney Harbor, USNM

205712 (3: 40.1-56.9).

Omobranchus rotundiceps obliquus (Carman)

FIGURES 29c,d

Petroscirtes obliquus Garman, 1903:237 [Suva, Fiji Islands].

Hypleurochilus loxias Jordan and Seale, 1905:802 [Negros,

Philippine Islands].

THypleurochilus samoensis Seale, 1935:374 [Pago Pago. Tutu ila Island, Samoa].

DESCRIPTION (see also Table 15).—Dorsal-fin spines

XI-XIII (XII in 95.0% of specimens); segmented

dorsal-fin rays 18-21 (rarely 21); total dorsal-fin

elements 30-33 (rarely 33); anal fin II, 20-23; both

anal-fin spines of males discernible externally; segmented caudal-fin rays 11-13 (1 specimen each, of

210, with 11 and 12 rays); dorsal + ventral procurrent caudal-fin rays 10-15; vertebrae 10-11 (rarely

11) + 27-30 (rarely 30) = 37-40 (rarely 40); epipleural ribs 11-15 (rarely 11 or 15); prenasal pores

present; interorbital pores 1-4 (rarely 1); circumorbital pores 7-9 (modally 8, rarely 9); lateral-line

tubes 0-6 (rarely 5 or 6); lateral-line tubes, when

present, extending posteriorly to below level of

dorsal-fin spine 1-7 (rarely 1 or 7); gill opening

extending ventrally to opposite 2nd to 8th pectoralfin ray (to 7th in 0 and 8th in 2 of 170 specimens;

of the 2, 1 each had the opening of the opposite

side at the level of the 4th and 5th rays); lower-lip

flap present; circumorbital bones 5; lower jaw teeth

19-29; upper jaw teeth 18-28 (Figure 45); no fleshy

bladelike crest on top of head of either sex (head

of some presumably mature males swollen in appearance).

COLOR PATTERN.—Closely resembles O. r. rotundiceps (q. v.) with the following notable exceptions:

Males: Bands on head better defined; band at

posteroventral margin of eye not interrupted at

preopercular series of sensory pores; it and following band confluent on underside of head with corresponding bands of opposite side; last 2 bands

often separated from corresponding bands of opposite side by narrow, pale space; last band continuous dorsally with broad, dusky area. Dome-shaped,

dark dusky area posterior to eye, reaching farther

ventrally; intense dark spot, larger than pupil of

eye, occupying entire dorsal portion of dome-shaped

area. Vertically elongate, less dark, oval area outlined, except occasionally ventrally, by narrow, pale

margin, present on operculum just posterior to

broad, dusky area continuous with 4th head band;

vertically elongate, dark smudge just posterior to

pale, margined, oval area, confluent with dark area

on top of head. No spots on top of head. Bands on

trunk better defined; dark intensifications of body

bands broad, not extending below midlateral line

of body. Posterior midlateral stripe and more dorsal stripe not developed; midlateral. stripe represented by series of dark spots; tips of anterodorsal

and posterodorsal extensions of body bands disjunct

from bands, forming longitudinal series of dark

spots or dashes. Dark bar on fleshy pectoral-fin base
NUMBER 177 69

well defined, continued anteroventrally onto prepelvic area. Dorsal fin largely transparent with several narrow, dark stripes extending length of fin;

diffuse concentration of pigment on segmented rays

and dorsoposteriorly on interradial membranes;

diffuse, dusky to black, oval spot placed more anteriorly, betweeen 8th and 12th (usually between 9th

and 11th) segmented rays. Anal fin wfth slender,

evenly spaced, posteroventrally inclined dusky

bands; distal edge of fin dusky with abruptly pale

ray tips. Caudal fin with vertically oriented pair of

large, dark spots at base; stripes faint to absent,

often only ventral stripe apparent.

Females: Sexual dimorphism as in O. r. rotundiceps except dark intensifications of body bands not

extending much farther ventrally; additional bands

or spots not developed ventrally between main

body bands; dorsal spotting and vermiculations

intensified and more numerous.

GEOGRAPHIC VARIATION.—See O. r. rotundiceps.

GEOGRAPHIC DISTRIBUTION (Figure 5).—Car Nicobar Island, Indian Ocean, west to Hawaii, but apparently absent from the eastern Caroline, Marshall, and Gilbert islands.

A completely faded specimen ZMA 113.013 (Female, 27.2 mm), from Sumbawa Island, Indonesia,

is identifiable as O. r. obliquus or O. elongatus. We

questionably allocate it to O. r. obliquus based on

its having 12 dorsal-fin spines, a number uncommon

in O. elongatus. Characters for the specimen are:

dorsal fin XII, 19; anal fin II, 21; total procurrent

caudal-fin rays 13; vertebrae 10 + 27; epipleural

ribs 14; 1 lateral-line tube, positioned below 5th

dorsal-fin spine; interorbital pores 3; circumorbital

pores 8.

Menon and Rama Rao (1963) recorded O. kallosoma from Madras and Mandapam, India, and the

Andaman Islands. Their specimens from India are

O. elongatus, whereas their specimen from the

Andaman Islands (actually Car Nicobar Island,

Nicobar Islands), is O. r. obliquus. The Car Nicobar

specimen, which represents the only Indian Ocean

record for the subspecies, arrived too late to be

included in Table 15. Meristics for the Car Nicobar

specimen are as follows: dorsal fin XII, 19; anal fin,

II, 21, vertebrae 10 + 28; epipleural ribs 13; lateralline tubes 1, reaching to below 2nd dorsal-fin spine;

gill opening extending ventrally to opposite

pectoral-fin ray 4.

HABITAT.—Shallow, usually close to shore, marine

waters. Frequently in tidepools, on muddy to sandy

bottoms with eel grass and algae, and near the edge

of mangrove swamps; once taken at the junction of

a small freshwater stream and marine bay. Live and

dead coral often recorded as present. Also taken

from a concrete tank in Hawaii (Strasburg, 1956),

and from around a rusting barge in the western

Caroline Islands.

COMPARISONS.—See O. r. rotundiceps.

NOMENCLATURAL DISCUSSION.—Jordan and Seale

(1905) did not compare their Hypleurochilus loxias

with any other species. It seems strange that they

failed to note the close resemblance of their illustration of the holotype (fig. 20) with Garman's

(1903, pi. 4: fig. 3) illustration of the holotype of

Petroscirtes obliquus. Jordan and Seale reported

that the holotype of H. loxias had D. XII, 20 and

A. 20. Their illustration indicates D. XII, 19 and

A I, 21, whereas we find the counts to be D. XI,

19 and A. II, 21 (the specimen is a female and the

first anal-fin spine can be seen only on a radiograph).

Springer (1972a) believed that the types of Hypleurochilus samoensis most probably represented

the postlarvae of Parenchelyurus hepburni, noting,

however, that the dorsal-fin formula of the types

was more representative of an Omobranchus species

than of P. hepburni. He excluded Omobranchus

from consideration because he erroneously believed

that Omobranchus was not known from Samoa.

Both of the types of H. samoensis have 12 dorsal-fin

spines, an unusual number for Parenchelyurus, but

a common number for Omobranchus. It therefore

appears that the types of H. samoensis are probably identifiable as O. rotundiceps obliquus, the

only taxon of Omobranchus occurring in Oceania.

Seale's (1935) illustration of the "type" of H. samoensis is reproduced as our Figure 36.

MATERIAL EXAMINED.—NICOBAR ISLANDS:

CAR NICOBAR: ZSIC F2764/2 (35.4). PHILIPPINE

ISLANDS: NEGROS: USNM 51952 (28.82, holotype

of Hypleurochils loxias), 137828 (4: 24.6-43.1),

209426 (4: 32.8-37.3), CAS GVF reg. no. 1597 (31.9),

GVF reg. no. 1618 (3: 25.2-37.6), GVF reg. no.

2671 (12.7-39.9). GUAM: CAS GVF reg. no. 1858

(122:14.8-46.2), UG 4125 (3:14.6-33.7). CAROLINE

ISLANDS: YAP: CAS GVF reg. no. 1910 (ca. 40),

GVF reg. no. 1927 (3: 35.9-41.6) GVF reg. no. 1928

(12: 26.9-44.0), GVF reg. no. 1929 (39.0). PALAU

ISLANDS: Koror, CAS SU62058 (3: 30.3-43.9), GVF

reg. no. 814 (21.1), GVF reg. no. 1380 (5: 13.9-
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

29.3), GVF reg. no. 1894 (2: 27.1-52.0); Babalthuap,

CAS GVF reg. no. 1397 (20.6), GVF reg. no. 1992

(32.3); Arakabesan, GVF reg. no. 1995 (31.5); Urukthapel, CAS SU62064 (5: 29.6-37.8); Aluptagel,

CAS SU62068 (42: 22.8-45.5), GVF reg. no. 1414

(4: 24.1-28.1). INDONESIA: AMBON: RMNH 20201

(2: ca. 38-41), USNM 211874 (23.8, arrived too late

to be included in Table 15). BUTON: USNM 211873

(4: 20.5-28.3, arrived too late to be included in

Table 15). TIMOR: RMNH 20340 (ca. 40). NEW

GUINEA: Port Moresby, DASF 1929 (38.0). TROBRIAND ISLANDS: USNM 205697 (3: 25.6-41.0). NEW

CALEDONIA: USNM 210689 (25.7). SANTA

CRUZ ISLANDS: Vanikoro: USNM 201893 (4:

19.5-26.8). NEW HEBRIDES: ANEITYUM: ANSP

91584 (2: ca. 28-37). FIJI ISLANDS: Vm LEVU:

ANSP 93928 (2: 29.0-31.2), MCZ 28297 (25.6, holotype of Petroscirtes obliquus). OVALAU: CAS

SU24926 (18: 22.9-ca. 39), FMNH 24928-37 (10:

ca. 25-48.1). SAMOA: TUTUILA: CAS 5515-5516

(12.3-12.6, holotype and paratype of Hypleurochilus

samoensis), USNM 209427 (ca. 20). UPOLU: USNM

52240 (ca. 30). LINE ISLANDS: PALMYRA: BPBM

15060 (2: 33.3-39.8), 15061 (3: 32.9-33.7), CAS

SU62059 (20: 24.4-42.4), GVF reg. no. 45 (2: 28.5-

32.7). HAWAIIAN ISLANDS: OAHU: BPBM 6205

(32.4), 15062 (50.9), 15063 (2: 35.1-56.2), USNM

198223 (54.8).

Omobranchus steinitzi, new species

FIGURE 30

DESCRIPTION (where variation is indicated, the

holotype is flagged with an asterisk; see also Table

5).—Dorsal fin XI-XII*, 16*-17 = 28; anal fin II,

18; one or both anal-fin spines of males not discernible externally; segmented caudal-fin rays 13; dorsal + ventral procurrent caudal-fin rays 10-12*;

vertebrae 10 + 26 = 36; epipleural ribs 14*-15;

prenasal pores absent; interorbital pores 3; circumorbital pores 8; lateral-line tubes 4-6*; lateral-line

tubes extending posteriorly to below level of dorsalfin spine 6-9 (to 8 on one side and 9 on the other

in the holotype); gill opening restricted to area

dorsal to level of dorsalmost pectoral-fin ray; lowerlip flap present; circumorbital bones 5; lower jaw

teeth 22-26 (24*); upper jaw teeth 22-24* (Figure

43); no fleshy bladelike crest on top of head of

either sex.

COLOR PATTERN.—Males: Head. Up to 8 evenly

spaced, dusky bands present on ventral half of

head; 1st, 3rd, 5th, and 7th bands narrow; 2nd, 4th,

6th, and 8th bands slightly broader; 1st band connecting anterior margin of orbit and ventral edge

of upper lip just posterior to tip of snout; 2nd band

extending anteroventrally from nape to dorsoposterior margin of orbit, interrupted by eye and continuing from anteroventral margin of orbit across

mouth, just anterior to corner of jaws, to chin; 3rd

band extending from ventral margin of orbit to

underside of head; 4th band paralleling 3rd from

posteroventral margin of orbit to underside of

head: 5th band originating at about level of ventral

margin of orbit and extending to ventral side of

head; 6th band extending from level just ventral to

origin of 5th band and splitting near preopercular

sensory pores, with 1 branch continuing to ventral

side of head and other continuing posteroventrally

to prepelvic area; 7th band short, extending posteroventrally from about level of ventral margin of

orbit to near center of branchiostegal membrane;

8th band extending from level of center of eye to

point just dorsoanterior to origin of pelvis fins;

bands may join corresponding bands of opposite

side on ventral side of head (largest specimens

mostly dusky or duskily mottled on head with only

4 anterior bands apparent). Large, dusky, oval spot

slightly smaller than eye present just posterior to

eye; dorsal half of spot dark (sometimes only portion of spot apparent) with narrow, pale margin.

Faint, broad, dusky band extending from dorsal

side of spot to nape. Snout and interorbital dusky;

nape with pale spots and blotches posteriorly.

Trunk: Approximately 7 pairs of narrow, dark

bands present on trunk; posterior 2 pairs each

coalesced into short, broad, irregular dash at lateral midline, 5 anterior band pairs distinct on midlateral half of body, bands in each pair broadly

separated, but faintly joined at lateral midline in

some specimens, giving each band pair an H configuration (banding obscured by mottled dusky pigment in some specimens). Five or 6 pairs of large,

broad, dark circles with pale centers present in

longitudinal series on dorsal quarter of trunk; circles more or less associated with dorsal ends of lateral bands or band pairs. Ventral quarter of trunk

with series of irregular, dark marks usually present

ventrally between tips of lateral bands along base

of anal fin, especially posteriorly.
NUMBER 177 71

Pectoral Fin: Transparent with fine, dark specks

of pigment scattered on rays; ventral rays dusky

distally. Narrow to broad, dusky band present midway along fleshy pectoral-fin base. Large, faint,

dusky blotch present dorsally on fleshy base at base

of fin rays, 2nd spot sometimes present at middle of

fin base.

Pelvic Fin: Pale with fin dark specks of pigment

basally.

Dorsal Fin: Transparent with broad, dusky to

dark, marginal stripe on distal third to half of fin;

anterior half of fin with 2 darker, narrow stripes,

1 on proximal edge and 1 on distal edge of broad

stripe. Tips of some segmented rays pale, others

dark.

Anal Fin: Dark; rays transparent to pale proximally; tips of rays and membranes posterior to ray

tips abruptly pale.

Caudal Fin: Pale, with some dusky pigment. Two

large dusky spots present at base of fin, 1 just above

and 1 just below lateral midline, each spot drawn

into dusky streak posteriorly. Dorsal and ventral

edges of fin dusky.

Female: Similar to males except: markings on

head and trunk more distinct, less obscured by

overall dusky pigment; background paler; bands

on head and trunk well defined; dorsal fin with

only 2 narrow, dark stripes (stripes not as dark)

apparent anteriorly; posteriorly, dorsal fin transparent with irregular dark blotches present just

above dark circles on trunk; anal fin with dark

pigment confined to narrow stripe near distal edge

of fin, pigment reaching slightly more basally on

rays.

COMPARISONS.—Omobranchus steinitzi is unique

among the specie* of Omobranchus in lacking prenasal sensory pores; it also has fewer dorsal- and

anal-fin elements than any of the other species. In

having one or both of the anal-fin spines of males

not discernible externally it resembles the banditus

species group, but it does not have the large number of epipleural ribs or color pattern features that

characterize that group.

ETYMOLOGY.—Named in honor of the late Dr. H.

Steinitz in recognition of his contributions to the

field of marine biology.

HOLOTYPE.—USNM 209433, male, 32.0 mm SL,

Cundabilu, Dahlak Archipelago, Red Sea, collected

20 March 1962, H. Steinitz, E. Clark, A. Ben-Tuvia.

PARATYPES.—USNM 209434, 2 males, 25.6-29.1

mm SL; HUJ E62/399a, female, 25.3 mm SL, all

with same data as holotype.

Omobranchus verticalis, new species

FIGURE 31

DESCRIPTION (see also Table 5; where variation

is indicated the holotype is flagged with an asterisk).—Dorsal fin XII, 19*-20 = 31*-32; anal fin

II, 22; both anal-fin spines of males discernible externally; segmented caudal-fin rays 13; dorsal +

ventral procurrent caudal-fin rays 14-15*; vertebrae

10 + 27*-28 = 37*-38; epipleural ribs 13; prenasal

pores present; interorbital pores 3*-4; circumorbital pores 8; lateral-line tubes 3-5 (4 on both sides

of holotype); lateral-line tubes extending posteriorly to below level of dorsal-fin spine 3-9 (to 6 on

one side and 9 on the other in the holotype); gill

opening either restricted to area dorsal to level of

dorsalmost pectoral-fin ray* or extending ventrally

to opposite dorsalmost ray; lower-lip flap present;

circumorbital bones 5; lower jaw teeth 16*—18; upper

jaw teeth 15-17 (16*; Figure 42); no fleshy bladelike crest on top of head of males.

The above description is based only on the three

available males. The following description is based

on a female that arrived too late to be included

above or in Tables 1, 4, and 5 or Figure 42. Dorsal

fin XIII, 19 = 32; anal fin II, 22; segmented caudalfin rays 13; dorsal + ventral procurrent caudal-fin

rays 14; vertebrae 10 + 28 = 38; epipleural ribs

12; prenasal pores present; interorbital pores 3; circumorbital pores 8; lateral-line tubes 2; lateral-line

tubes extending posteriorly to below level of dorsalfin spine 3; gill opening restricted to area dorsal to

level of dorsalmost pectoral-fin ray; lower-lip flap

present; lower jaw teeth 16; upper jaw teeth 15; no

fleshy bladelike crest on top of head.

COLOR PATTERN.—Males: Head: Four moderately

narrow, dark bands present on head; all bands confluent across ventral surface of head with corresponding band of opposite side; anterior band

extends from anteroventral margin of orbit across

mouth, just anterior to corner, to chin; 2nd band

extends ventrally from posteroventral margin of

orbit; 3rd band originates broadly on nape midway

between eye and dorsal-fin origin, extends first

anteroventrally, then slightly posteriorly just below

posterior margin of orbit, where it narrows slightly
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and continues to ventral surface of head; 3rd band

bears dark intensification (spot) on anterior margin

in area posterior to eye; 4th band extends ventrally,

slightly posteriorly, from upper end of operculum.

Several very fine, intensely dark spots present on

opercular membrane just anterior to gill slit. Snout

and interorbital dusky. Several pale blotches present on nape; dusky blotch present on nape just

anterior to dorsal-fin origin.

Trunk: Approximately 10 to 12 dusky bands on

body; bands on midlateral half of body, bordered

by anterior and posterior narrow, dark margins;

margins may converge slightly dorsally and ventrally, giving bands double convex appearance;

bands generally broader anteriorly (although first

2 or 3 narrow) and progressively narrower posteriorly, the last 5 or 6 may be reduced to single

dark, dusky, vertical lines, becoming faint to absent at posterior end of body; bands evenly spaced,

spaces between bands slightly narrower than broadest band; ends of bands indistinct, dorsal end of

each band connected by diffuse, irregular, dusky

extension of band to corresponding blotch in series

of dark, evenly spaced blotches lying along dorsalfin base.

Pectoral Fin: Pale, slightly dusky area present

midlaterally on base of fin; narrow dark band

directed posteroventrally across fleshy fin base from

ventral end of gill slit to ventral surface of trunk

posterior to pelvic-fin base.

Pelvic Fin: Pale.

Dorsal Fin: Pale, with fairly even spread of fine

dusky specks of pigment, pigment more dense anteriorly. Row of dusky blotches on body at base of

dorsal fin continued slightly onto fin. Large, oval,

dark spot present on distal three-fourths of membrane between first 2 spines.

Anal Fin: Pale dusky, darker anteriorly, ray tips

abruptly pale. Dark dusky blotch present on membrane posterior to every 2nd ray base.

Caudal Fin: Pale with vertically oriented pair of

dusky blotches at fin base, each blotch continued

posteriorly as dusky streak.

Female: Similar to males except: narrow, pale,

median stripe extending from between anterior

nostril to interorbital area; pale spot enclosing

posterior nostril; dark area on nape forming chainlike configuration enclosing pale spots; dark body

b bands not converging dorsally or ventrally, less

bbroad anteriorly, more uniformly pigmented; spinous portion of dorsal fin with distal half dusky,

proximal half pale except for dusky blotches associated with body bands; dark spot at anterior end

of dorsal fin slightly smaller and less intense.

COMPARISONS.—Omobranchus verticalis is separable from all other species of Omobranchus on

features of its color pattern. Males have a dark spot

between the anterior two dorsal-fin spines, a mark

not found in males of the other species. The nature of

the body bands of O. verticalis is also distinctive.

Omobranchus verticalis is otherwise separable from

most of the species of Omobranchus only by a combination of characters. Besides color pattern, it may be

separated most readily from those populations (eastern Australian) of Omobranchus species with which

it appears to be broadly sympatric as follows: from

O. anolius in lacking a fleshy crest on the head and

in having lateral-line tubes; from O. germaini,

O. rotundiceps, and O. punctatus in having 31-32

total dorsal-fin elements (versus 32-36 in the other

species; only a few specimens of O. germaini with

32) and 37-38 total vertebrae (39-43 in the other

species); from O. punctatus in having 13 epipleural

ribs (versus 19-24) and both anal-fin spines of

males discernible externally; from O. rotundiceps

in having 3-5 lateral-line tubes, which extend posteriorly to a level below dorsal fin spines 6-8 (versus 0-3, rarely 3, tubes extending, when present, to

below dorsal-fin spines 1-4), and in having 22 segmented anal-fin rays (versus 23-26); from O. lineolatus in having 12 dorsal-fin spines (versus 11) and

22 segmented anal-fin rays (versus 20).

HOLOTYPE.—AMS IA.4938, male, 38.8 mm SL,

Boat Passage, Brisbane River, Queensland, in hole

in mangrove roots high up in root and almost at

high-water mark, coll. T. Iredale, May 1931.

PARATYPES.—AMS 1.17150-001, male, 49.5 mm

SL, and USNM 210616, 33.3 mm SL, male, collected with the holotype. ANSP 128000, female, 30.4

mm SL, vicinity of Brisbane, probably Serpentine

Creek mangrove area, 1970.

Omobranchus woodi (Gilchrist and Thompson)

FIGURE 32

Aspidontus woodi Gilchrist and Thompson, 1908:105 [Nahoon

River].

DESCRIPTION (see also Table 5).—Dorsal fin XIXIII (XII in 94.6% of specimens), 19-21 = 31-33;
NUMBER 177 73

anal fin II, 20-22 (up to 23 according to J. L. B.

Smith, 1965 and Penrith and Penrith, 1972); both

anal-fin spines of males discernible externally; segmented caudal-fin rays 13; dorsal + ventral procurrent caudal-fin rays 12-15 (rarely 15); vertebrae

11-12 (11 in 92.0% of specimens) + 27-29 = 38-

40; epipleural ribs 14-19 (rarely 14); prenasal pores

present; interorbital pores 2-4 (3 in 97.2% of specimens); circumorbital pores 7-9 (8 in 94.3% of

specimens); lateral-line tubes 0-6 (0 in one, 1 in

none, and 2 in one of 33 specimens); lateral-line

tubes, when present, extending posteriorly to below

level of dorsal-fin spine 2-10 (to 2 in one specimen, to 5-10 in all other specimens); gill opening

either restricted to area dorsal to level of dorsalmost pectoral-fin ray (31 of 32 specimens) or extending ventrally to opposite dorsalmost ray (1

specimen); lower-lip flap present; circumorbital

bones 5; lower jaw teeth 25-31; upper jaw teeth

23-30 (Figure 46); fleshy bladelike crest on top of

head of both sexes.

COLOR PATTERN.—Males (pattern of most large

individuals obscured by dusky to dark dusky shading): Head: Faint, diffuse, dusky band extending

from anteroventral margin of orbit across mouth,

just anterior to corner of jaws, to chin, where it

joins corresponding band of opposite side; band

interrupted by eye dorsally and continued from

dorsoposterior margin of orbit onto fleshy crest.

Two (occasionally 3) vertical, intensely dark marks

separated by narrow, pale interspace on side of

head posterior to eye. Circumorbital sensory pores

along posterior half of eye ringed by intensely dark

pigment. Several narrow, faint, dusky lines radiating from dorsal and ventral margins of orbit. Snout

and anterior inteorbital dusky with several pale

spots or small blotches anterior to eye. Crest and

nape usually paler than rest of head with dorsoposteriorly slanted, dusky band extending onto

crest from orbital margin. Dark ventroanteriorly

slanted saddle extending across nape just anterior

to dorsal-fin origin; ventral end of saddle (separate

spot in some specimens) broad, somewhat darker,

positioned posterior to 2 dark marks on side of head.

Narrow, dark bar present on operculum just ventral to saddle, occasionally continuing across ventral

surface on head to other side. Several scattered,

small, intensely dark spots sometimes present on

operculum.

Trunk: Up to 11 or 12 evenly spaced, faint,

broad, dark dusky bands present on trunk, ventral

half of anterior 5 bands slanted posteroventrally,

posterior bands nearly vertical, dorsal ends of

bands irregular, sometimes separate; bands becoming faint to absent posteriorly. Row of approximately 15 evenly spaced, faint, dusky blotches on

dorsal body contour at dorsal-fin base. Many small,

intensely dark spots occasionally present dorsoposteriorly to pectoral-fin base; approximately half of

spots hidden by appressed pectoral fin. Freshly preserved specimens with trunk evenly covered by

small, pale spots.

Pectoral Fin: Pale dusky with even spread of fine

dark specks of pigment. Fleshy base pale dusky with

dusky blotch ventrally.

Pelvic Fin: Pale dusky with even spread of fine

dark specks of pigment.

Dorsal Fin: Pale dusky to transparent with even

spread of fine dark specks of pigment, slightly

duskier near base.

Anal Fin: Dark dusky to dark, occasionally with

many indistinct, small, pale spots; tips of rays

abruptly pale.

Caudal Fin: Pale dusky, duskier on ventral half

of fin. Ventral edge of fin dark near base.

Females: Similar to males except: overall pigment paler; 1 freshly preserved specimen with 2

additional, very faint, broad, dusky bands slanted

posteroventrally on ventral side of head; anteriormost band ventral to eye, joins corresponding band

of opposite side; 2nd band broadly connected to

dark saddle dorsally, reaching ventrally to prepelvic

area but not to ventral midline; bands on trunk

much fainter dorsally and ventrally, restricted much

more to lateral midline, especially posteriorly; occasionally more small intensely dark spots near

fleshy pectoral-fin base.

Omobranchus woodi has been illustrated in color

in Smith and Smith (1967:112).

GEOGRAPHIC DISTRIBUTION (Figure 7).—Western

coast of South Africa.

HABITAT.—Quiet backwaters in estuaries and

tidal rivers; in burrows under stones in muddy

areas (Smith and Smith, 1967).

COMPARISONS.—Omobranchus woodi is distinguishable from most of the other species of Omobranchus only on the basis of a combination of

characters. It is closely related to the banditus species group (see "Comparisons" under O. banditus),

but differs from that group in particulars of color
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pattern and in having both anal-fin spines of

males discernible externally. A fleshy crest on top

of the head is present in all specimens of both

sexes, and this character alone will distinguish

O. woodi from those species with which it appears

to be broadly sympatric, except O. banditus and

O. fasciolatus. It is readily separable from O. banditus in having a lower-lip flap and in lacking the

prominent dark body bands found in that species.

It can be distinguished from O. fasciolatus in having 3 (versus 2) interorbital pores and 11-12 precaudal vertebrae (versus 10).

While it is not readily apparent to the uninitiated, O. woodi exhibits certain features of color

pattern on the head that distinguish it from all

other species of Omobranchus. The distinguishing

pattern consists of 2 or 3, more or less parallel

short, dark, vertical marks posterodorsal to the

orbit, and the dark ringing of the posterior circumorbital pores (Figure 32).

MATERIAL EXAMINED.—SOUTH AFRICA: Durban Bay, SAM 17400 (ca. 29); Port Elizabeth, SAM

12024 (2: 62.1-ca. 74), BMNH uncataloged (3: ca.

51-81.1), 1900.4.20.9 (ca. 56); Nahoon River, SAM

9898 (77.9, holotype of Aspindontus woodi); Kowie

River, RUSI 2364 (58.3); Kariega River, RUSI 2361

(48.0); Knysna and vicinity, RUSI 2365 (12: 23.7-

68.7), 2357 (12: 45.0-75.7), 2359 (2: 60.8-70.2),

SAM 25384 (ca. 40); Lake Sifungwe, RUSI uncataloged (43.6). NO LOCALITY: SAM 23984 (ca. 37).

Omobranchus zebra (Bleeker)

FIGURES 33, 34

Petroscirtes zebra Bleeker, 1868:279 [Singapore in mari].

Petroscirtes bhattacharyae Chaudhuri, 1916:107 [Barkul Point,

Chilka Lake}.

DESCRIPTION (see also Table 5).—Dorsal fin XII,

18-20 = 30-32: anal fin II, 20-22; both anal-fin

spines of males discernible externally; segmented

caudal-fin rays 13; dorsal + ventral procurrent

caudal-fin rays 11-14; vertebrae 10 + 27-28 = 37-

38; epipleural ribs 12-15; prenasal pores present;

interorbital pores 3; circumorbital pores 7-8 (8 in

16 of 19 specimens); lateral-line tubes 1-4; lateralline tubes extending posteriorly to below level of

dorsal-fin spines 2-7 (2-5 in all but 1 specimen);

gill opening either restricted to area dorsal to level

of dorsalmost pectoral-fin ray (19 of 20 specimens)

or extending ventrally to opposite dorsalmost ray

(1 specimen); lower-lip flap present; circumorbital

bones 4; lower jaw teeth 16-22; upper jaw teeth

15-24 (Figure 46); no fleshy bladelike crest on top

of head of either sex.

COLOR PATTERN.—Males: Head: Four broad, dark

bands present on head; anterior band running from

anteroventral margin of orbit across mouth, midway between corner of jaws and anterior contour

of head, to chin; band interrupted dorsally by eye,

continuing dorsally for short distance from dorsoposterior margin of orbit; 2nd band angled slightly

posteroventrally from posteroventral margin of

orbit to ventral midline of head; 3rd band encircling head with dorsal one-third directed anteroventrally from center of nape to point just posterior

to eye, then slightly posteroventrally to ventral

midline; 3rd band bears dark intensification on

anterior margin just posterior to eye; 4th band

encircling head just anterior to dorsal-fin origin;

dorsal half of 4th band angled slightly anteroventrally, ventral half slightly posteroventrally; 3rd and

4th bands usually fainter ventrally, especially on

cheek and branchiostegal region; all 4 bands join

corresponding bands of opposite side on underside

of head, but are slightly fainter at ventral midline,

particularly 1st band. Short, posteroventrally slanted, diffuse, dusky line usually present anteriorly on

free opercular membrane. Snout and interorbital

area dusky, blending with dorsal end of 1st band,

forming confluent dark dusky area with well-defined

posterior margin across top of head.

Trunk: Background pale to pale dusky, slightly

darker dorsally, with dark markings. Bleeker's

(1868) description and unpublished illustration of

the holotype (our Figure 34) indicate 8 evenly

spaced, broad, dark bands present on the body, each

almost reaching from the dorsal to the ventral body

contour (holotype now completely faded). Our material, however, exhibits only the anterior 2-4 bands,

and even these may be faint. Where the bands are

not prominent our specimens may exhibit a series

of large, circular, dark spots, each appearing where

a band would cross the lateral body midline; the

last two bands are usually not represented in our

specimens; the first two bands are usually developed,

at least dorsally, and slant slightly posteroventrally.

Pectoral Fin: Rays pale with even spread of fine,

dark specks of pigment, membranes transparent.

Posteroventrally slanted, broad, dark band travers-
NUMBER 177 75

ing fleshy pectoral-fin base, anteriorly connected to

faint dusky line on free opercular membrane dorsally and continued slightly onto abdomen ventrally.

Pelvic Fin: Pale with even spread of fine, dark

specks of pigment.

Dorsal Fin: Pale with even spread of fine, dark

specks of pigment and dark dusky to dark markings.

Series of paired dark blotches (diffusely dusky and

unpaired in some specimens) present along fin base

anteriorly above dorsal ends of body bands; blotches

becoming faint, then absent, midway along fin;

each blotch covering membrane between bases of 2

adjacent rays; pairs of blotches separated by single,

unpigmented membrane space. Broad, dark, marginal stripe present on spinous portion of fin; pigment on anterior distal edge of 1st spine and

membrane between distal tips of first 2 spines intensified, often the only prominent dark markings

in dorsal fin of small specimens.

Anal Fin: Dusky, ray tips abruptly pale.

Caudal Fin: Pale with even spread of fine, dusky

specks of pigment on rays and membranes bordering rays. Basal margin of fin dusky; 2 longitudinal,

diffuse, dusky streaks on fin, 1 just above and 1 just

below lateral midline of fin.

Females: Similar to males except: series of spots

on lateral midline of body very faint to absent;

dark markings on dorsal fin slightly fainter, anal fin

slightly paler.

GEOGRAPHIC DISTRIBUTION (Figure 6).—Bay of

Bengal coast of India to Singapore.

Menon and Talwar (1972) reported O. zebra

from Galathea Bay, Great Nicobar Island. Their

description appears to apply to O. zebra but we

have not seen their specimen and have not, therefore, included the record on Figure 6.

Dutt and Visveswara Rao (1961) reported O. zebra

(as Petroscirtes bipunctata) nesting in holes in mangrove stems in the Godavari River estuary (16°28'N,

82°03'E) in India. We became aware of this reference

too late to include the record on Figure 6.

HABITAT.—Mangrove areas, sand and mud bottom, brown stained water, low salinity areas. Bhattacharya (1916) reported larval stages, as P. bhattacharyae, 3.2-15.9 mm total length, from Chilka

Lake waters of densities 1.006-1.007, corrected to

15° C. Converted to salinity, these densities equal

a range of about 8.2-10.8 o/oo.

COMPARISONS.—Omobranchus zebra differs from

all other species of Omobranchus except O. ferox,

with which it is broadly sympatric, in having 4 circumorbital bones (4 bones is an uncommon variant

in the other species). It differs from O. ferox most

obviously in having the gill opening almost always

restricted to the area dorsal to the level of the

dorsalmost pectoral-fin ray, rarely extending ventrally to opposite the dorsalmost ray (versus opposite the 3rd to 6th ray in O. ferox), in having welldefined bands on the head, and in having a longer

head (see key couplet 13).

Aside from the number of circumorbital bones,

O. zebra may be most readily separated from the

other species with which it appears to be broadly

sympatric as follows: from all in having a longer

head (key couplet 13); from O. elongatus and O.

germaini in lacking those characters of the elongatus species group (see "Comparisons" under O.

elongatus) and in having 12 dorsal-fin spines (versus

12-14, modally 13 spines); from O. meniscus in

lacking a fleshy crest in males, in having lateral-line

tubes, and in having the gill opening restricted to the

area dorsal to the level of the 2nd pectoral-fin ray

(versus tubes absent and gill-opening extending ventrally to opposite 3rd ray); and from O. punctatus

in having 1-4 (rarely 4) lateral-line tubes, which

extend posteriorly to below the level of dorsal-fin

spines 2-6 (rarely 5-6; versus 3-8 tubes, rarely 3—4,

which extend posteriorly to below dorsal-fin spines

6-11, rarely 6-7).

NOMENCLATURAL DISCUSSION.—Bleeker (1868) did

not illustrate his holotype of P. zebra, but he did

have a colored illustration prepared, which he intended to use in his "Atlas." This illustration is in

the files of the RMNH. As the holotype is now

faded and the illustration indicates a more complete banding on the body than any specimen we

have examined, we have included a copy of the

illustration, much enlarged, as Figure 34.

Chadhuri (1916) did not compare his Petroscirtes

bhattacharyae with any other species. We have not

seen the holotype, but from information contained

in the original description, data on the holotype,

which was examined for us, and our examination

of 2 of the 11 paratypes, we are convinced that P.

bhattacharyae is a synonym of O. zebra.

MATERIAL EXAMINED.—INDIA: Lake Chilka,

ZISC F8764/1 (41.8, holotype of Petroscirtes bhattacharyae, information furnished by W. F. SmithVaniz), 8774 (25.2), 8775 (39.6). THAILAND: Pakchan River mouth (Indian Ocean), CAS SU62065
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(2: 39.9-56.5). GULF OF THAILAND: between Choi

Buri and Bangsaen, CAS GVF reg. no. 1542 (9:

25.2-53.8); Lem Nam Point, Trat Bay, CAS GVF

reg. no. 1486 (35.9), GVF reg. no. 1487 (5: 36.2-

48.1). SINGAPORE: RMNH 4454 (44.4, holotype

of Petroscirtes zebra).

Omobranchus species

FIGURE 35

Among all the specimens of Omobranchus that

we examined, there was only one (USNM 211092)

that we could not identify with reasonable certainty. The specimen is a small (22.7 mm SL) male

from a predominantly "worm rock" habitat about

one-half mile (0.6 km) offshore of Hikkaduwa,

Ceylon. The specimen is noteworthy for several

reasons, among which are the following: it is the

only transformed specimen of Omobranchus that

we know of that has not come from a close-to-shore

or freshwater habitat and it was taken at depth of

35-45 ft (10.7-13.7 m), much greater than that

verifiable for any other collection of Omobranchus,

which genus has rarely been reported from a depth

as great as 4 meters. Furthermore, the color pattern

on the head is unusually well developed for so

small a specimen. Because of its small size and the

possibility that important characteristic structures

may not have had time to develop (for instance, a

fleshy crest, filamentous caudal-fin rays) or because

it might represent an aberrant specimen of a known

species, we are unwilling to describe it as a new

species.

The specimen has the following characteristics:

dorsal fin XII, 19; anal fin II, 21; segmented caudalfin rays 13; dorsal + ventral procurrent caudal-fin

rays 12; vertebrae 10 + 28 = 38; epipleural ribs

14; prenasal pores present; interorbital pores 3;

circumorbital pores 8; lateral-line tubes 7, extending posteriorly to below level of dorsal-fin spine 10;

lower-lip flap present, gill opening extending ventrally to opposite 2nd pectoral-fin ray; 5 circumorbital bones; upper jaw incisor teeth 20; lower jaw

incisor teeth 23.

The specimen appears to be most similar to

O. fasciolatus, but aside from being somewhat outside the geographic range of that species, it has 3

interorbital pores, rather than 2, and a welldeveloped dark banding under its head and on the

fleshy pectoral-fin base that are not found in O.

fasciolatus. Nor does O. fasciolatus have a dark spot

immediately posterior to the orbital margin as does

the specimen. The number of jaw teeth is slightly

higher than what we would expect for so small a

specimen of O. fasciolatus.

In color pattern and other respects the specimen

appears similar to O. zebra, which possibly occurs

in Ceylon. The specimen differs from O. zebra in

general physiognomy, in having more lateral-line

tubes (which extend farther posteriorly than in

O. zebra), more caudal and total vertebrae, more

circumorbital bones, and far more jaw teeth than

would be expected in so small a specimen of O.

zebra.

It is markedly different from O. punctatus and

O. elongatus, the other two species that occur in

the same general geographic area with it.

Questionable Species

Blennius pardalis Castlenau (1875) was questionably allocated to the genus Istiblennius (tribe Salariini) by Smith-Vaniz and Springer (1971). Springer

(1972a) overlooked his previous generic allocation

and questionably assigned B. pardalis to Omobranchus, noting that it might even belong in the tribe

Blenniini. We now believe that B. pardalis should

be referred back to the Salariini and particularly

to Salarias fasciatus (Bloch).

Castlenau described B. pardalis (from Cape York,

Torres Strait) as lacking orbital cirri, having the

head rounded and cut vertically in front, and having the dorsal fin of about uniform height for its

entire length. The color pattern description includes the following information: anterior half of

back with narrow, longitudinal, black lines; body

entirely covered with oblong whitish blotches; dorsal fin brown with transverse whitish lines formed

of oval or rounded blotches and with series of

minute black blotches near edge of fin; caudal fin

rounded, with three transverse series of black dots;

anal fin brown, marked with whitish round blotches.

The color pattern is not found in any species of

the tribes Omobranchini, Nemophini, Phenablenniini, or Blenniini (the last-named tribe is either

rare in, or absent from, the tropical Indo-Australian

archipelago). The color pattern seems most applicable to a member of the tribe Salariini, but not to
NUMBER 177 77

Ecsenius, the only genus of that tribe in the IndoAustralian archipelago normally lacking orbital

cirri, nor to Istiblennius, which has a deeply

notched dorsal fin. If, however, Castlenau was in

error in reporting the absence of orbital cirri, the

color pattern description best applies to Salarias

fasciatus, a common, nearshort inhabitant of the

I ndo-Australian region.

List of Nominal Taxa of Omobranchus and Their Current Identification

NOMINAL TAXA

Graviceps alexanderi Whitley, 1945

Petroscirtes altivelis Steindachner, 1863

Graviceps angelus Whitley, 1959

Blennechis anolius Valenciennes in Cuvier and

Valenciennes, 1836

Blennius auro-splendidus Richardson, 1846

Omobranchus bandittis J.L.B. Smith, 1959

Petroscirtes bhattacharyae Chaudhuri, 1916

Petroscirtes cristatus Zugmayer, 1913

Omobranchus cristatus Fraser-Brunner, 1951

Petroscirtes cristiceps Macleay, 1881

Graviceps darwini Whitley, 1958

Aspidontus dasson Jordan and Snyder, 1902

Omobranchus dealmeida J.L.B. Smith, 1949

Salarias decipiens DeVis, 1884b

Petroscirtes dispar Gunther, 1861

Petroscirtes dispar Fowler, 1937

Petroscirtes elegans Steindachner, 1876

Petroscirtes elongatus Peters, 1855a, 1855b

Blennius fasciolatoceps Richardson, 1846

Blennechis fasciolatus Valenciennes in Cuvier

and Valenciennes, 1836

Petroscirtes fasciolatus Macleay, 1881

Petroscirtes feliciana Herre, 1942

Petroscirtes ferox Herre, 1927

Salarias furcatus DeVis, 1884b

Salarias furtivus DeVis, 1886

Salarias galeatus DeVis, 1884a

Petroscirtes germaini Sauvage, 1883

Petroscirtes guttatus Macleay, 1881

Salarias helenae DeVis, 1884b

Chasmodes Herklotsi Herre, 1935

Petroskirtes japonicus Bleeker, 1869

Omobranchus japonicus scalatus J.L.B. Smith, 1959

Petroskirtes kallosoma Bleeker, 1858

Petroscirtes kochi Weber, 1908

Petroscirtes kranjiensis Herre, 1940

Petroscirtes lineolatus Kner, 1868a, 1868b

Petroscirtes lineo-punctatus Sauvage, 1800

Petroscirtes lini Herre, 1934

Hypleurochilus loxias Jordan and Seale, 1905

Petroscirtes loxozonus Jordan and Starks, 1906

Petroscirtes macleayi Ogilby, 1887

Petroscirtes masyae H.M. Smith, 1934

Petroscirtes mekranensis Regan, 1905

Omobranchus meniscus Springer and Gomon

Petroscirtes obliquus Carman, 1903

CURRENT IDENTIFICATION

Omobranchus germaini

Omobranchus anolius

Omobranchus r. rotundiceps

Omobranchus anolius

Omobranchus aurosplendidus

Omobranchus banditus

Omobranchus zebra

Omobranchus mekranensis

Omobranchus fasciolatus

Omobranchus anolius

Omobranchus lineolatus

Omobranchus punctatus

Omobranchus ferox

Omobranchus punctatus

Omobranchus punctatus

Omobranchus elongatus

Omobranchus elegans

Omobranchus elongatus

Omobranchus fasciolatoceps

Omobranchus fasciolatus

Omobranchus r. rotundiceps

Omobranchus ferox

Omobranchus ferox

Omobranchus r. rotundiceps

Omobranchus r. rotundiceps

Omobranchus anolius

Omobranchus germaini

Omobranchus anolius

Omobranchus punctatus

Omobranchus fasciolatoceps

Omobranchus punctatus

Omobranchus punctatus

Omobranchus elongatus

Omobranchus punctatus

Omobranchus ferox

Omobranchus lineolatus

Omobranchus elegans

Omobranchus aurosplendidus

Omobranchus rotundiceps obliquus

Ombranchus loxozonus

Omobranchus r. rotundiceps

Omobranchus punctatus

Omobranchus mekranensis

Omobranchus meniscus

Omobranchus rotundiceps obliquus
78

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

NOMINAL TAXA

Blennechis punctatus Valenciennes in Cuvier

and Valenciennes, 1836

Petroscirtes rotundiceps Macleay, 1881

Hypleurochilus samoensis Seale, 1935

Petroscirtes semilineatus Kner, 1868b

Poroalticus sewalli Fowler, 1931

Salarias sindensis Day, 1888

Omobranchus steinitzi Springer and Gomon

Petroscirtes striatus Jatzow and Lenz, 1898

Petroscirtes uekii Katayama, 1941

Blennius unicornis Castelnau, 1879

Omobranchus verticalis Springer and Gomori

Petroscirtes vinciguerrae Borsieri, 1904

Petroscirtes wateroust Herre, 1942

Petroscirtes wilsoni Macleay, 1884

Aspidontus woodi Gilchrist and Thompson, 1908

Petroscirtes zebra Bleeker, 1868

CURRENT IDENTIFICATION

Omobranchus punctatus

Omobranchus r. rotundiceps

^Omobranchus rotundiceps obliquus

Omobranchus punctatus

Omobranchus punctatus

Omobranchus punctatus

Omobranchus steinitzi

Omobranchus fasciolatus

Omobranchus fasciolatoceps

Omobranchus anolius

Omobranchus verticalis

Omobranchus fasciolatus

Omobranchus ferox

Omobranchus anolius

Omobranchus woodi

Omobranchus zebra

Notes on Other Species of the Tribe Omobranchini

Enchelyurus ater (Gunther)

The following specimens extend the geographic

range of this species beyond that reported by

Springer (1972a): NEW HEBRIDES: AMS 1.14313

(6: 23.5-28.6); Vila, Sandwich Island (= Efate Island?), AMS 1.6323-4 (2: 20.7-32.7), IB.3609 (18.9);

Santo (= Espiritu Santo?), AMS 1.6513 (26.3).

LORD HOWE ISLAND: AMS IA.969 (6: 19.0-

ca. 47); IB.5995 (32.4).

Important meristic characters for these specimens

are given in Table 17. The high counts of the Lord

Howe Island specimens are similar to those of specimens from New Caledonia and Rapa, whereas the

low counts of the New Hebrides specimens are

similar to those of the other populations (Springer,

1972a, Tables 1-2).

Enchelyurus flavipes Peters

Springer (1972a) incorrectly cited RMNH 20813,

from Gonto Soea, near Makassar, and ZMB 5193,

from Singapore, as the types of Enchelyurus flavipes

var. nigerrima Weber. The correct catalog number

for the two syntypes is ZMA 112.678, from Barang

Island near Makassar. We have examined both syntypes and find no reason to recognize the validity

of Weber's infraspecific name.

Menon and Talwar (1972) reported on a specimen of E. flavipes (as E. flaviceps) from Galathea

Bay, Great Nicobar Island. If their identification

of the specimen is correct, the locality would represent the first record of E. flavipes in the Indian

Ocean.

Haptogenys bipunctata (Day)

On the basis of information in Day (1876) Springer (1972a) assigned Petroscirtes bipunctatus from

Calicut, India, to the genus Omobranchus, and

noted that the holotype was lost. Menon and Yazdani (1968) did not mention the existence of the

holotype in their purportedly complete catalog of

the types in the collections of the Zoological Survey

of India. W. F. Smith-Vaniz, who visited ZSIC in

1969 with the intention of examining all the blenniid types, was not apprised that it was present.

Recently, Dr. P. Talwar (ZSIC) notified us that the

type was available (ZSIC No. 2082). Information

on the holotype, kindly furnished us by Dr. A. G. K.

Menon, convinces us that P. bipunctatus is a species

of Haptogenys and is undoubtedly the same as

H. quadripora Springer, which we here place in

synonymy with H. bipunctata (name emended here

to agree with gender of the genus). Springer had

but a single specimen (USNM) of H. bipunctata (as

H. quadripora), from Koh Tao, Gulf of Thailand.

Another specimen (MCZ 47202, female, 32.0 mm

SL) of H. bipunctata, from Lam Goh Peninsula,

south Thailand (6°57'N, 99°43'E), has also come

to our attention. Both this specimen and the holotype of P. bipunctatus have three mandibular sens-
NUMBER 177 79

TABLE 17.—Frequency distributions for certain meristic characters in two

populations of Enchelyurus ater

Population

Nev Hebrides

Lord Howe

Dorsal-fin

spines

8 9 10

3 7

6 1

Segmented

dorsal-fin rays

20

1

21 22

7 2

2

23 24

3 2

Total

dorsal-fin elements

29

3

30 31

6 1

1

32

4

33

2

Total anal-fin

elements

20 21 22 23

3 7

4 3

Caudal

23

2

24

7

vertebrae

25

1

3

26 27

3 1

Total

33 34

2 7

vertebrae

35

1

3

36

3

37

1

ory pores, whereas the USNM specimen has two

pores, indicating the probability that the USNM

specimen is aberrant. The USNM specimen also

may have been unsual in having the shortest pelvicfin ray less than half the length (42.5%) of the longest. The MCZ specimen has the shortest ray 54.5%

the length of the longest.

The following meristic data from the ZSIC and

MCZ specimens expand the description of the species over that given by Springer; dorsal fin XII, 19

(2) anal fin II, 21 (2); pectoral fin 13 (MCZ); interorbital pores 4 (2); circumorbital pores 8 (MCZ);

gill opening extending ventrally to pectoral-fin ray

13 (2); upper teeth 18 (MCZ); lower teeth 20

(MCZ). Ventral hypural plate autogenous.

Otnox biporos Springer

Springer (1972a) described O. biporos from seven

specimens, two of which were from New Guinea

(Madang Harbor, north coast). Nineteen more specimens from New Guinea (south coast) are now

available: AMS 1.17541-001 (6: 25.2^36.9) and

BPBM 15914 (10: 26.1-40.7), Port Moresby, and

DASF FO1655 (3: 24.7-39.2) Lelea. All the collections were taken in mangrove areas.

The additional specimens modify the characterization of the genus (and its monotypic species) as

given by Springer. All of the specimens have the

characteristic four interorbital pores, except one,

which has two pores. Fourteen specimens agree with

Springer's description in lacking a median supratemporal commissural pore, whereas five of the

specimens have this pore. On the basis of pore

distribution, therefore, some specimens of Omox

could be misidentified as Omobranchus. The much

less restricted gill opening of Omox (extending to

opposite pectoral-fin ray 8-11) will distinguish

Omox from Omobranchus. Male Omox over 37 mm

SL possess a kinethmoid, which is not present in

any specimen of Omobranchus that we examined

for this bone.

The frequencies for total dorsal-fin elements of

Omox biporos from Thailand and New Guinea given

by Springer (1972a, table 1) should each be shifted

one column to the left.

Parenchelyurus hepburni (Snyder)

The following specimens represent new locality

records for the species, which is now known to

extend from the Amirante Islands, in the western

Indian Ocean, to Samoa, in the central Pacific

Ocean: DASF FO3200 (26.8), Port Moresby, New

Guinea; USNM 211875 (2: 22.7-32.4), Ambon Island, Indonesia; USNM 211876 (23.0), Buton (also

Butung), Indonesia; NFIS 11862 (2: 13.0-ca. 22),

Great Nicobar Island, Nicobar Islands; ANSP station no. F-102 (8: 17.1-24.9), St. Joseph Island,

Amirante Islands. The species appears to vary little

throughout its geographic range.

Yamakawa (1971) reported and illustrated P.

hepburni (as Omobranchus hepburni) from Yoronto

and Kikaijima. These localities represent the northernmost records for the species.

Springer (1972a) believed that the types of Hypleurochilus samoensis Seale might represent the

postlarvae of P. hepburni. It now appears more likely

that these types are larval stages of Omobranchus

r. obliquus, based on their dorsal-fin formulae.

Both species of Parenchelyurus have two mandibular sensory pores, as indicated in Springer's

(1972a) key to the species. Due to a typographical

error, the number of mandibular pores was given as

three in Springer's generic diagnosis.
80

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Literature Cited

Bhattacharya, D. R.

1916. Stages in the Life History of Gobius, Petroscirtes,

and Hemirhamphus. Memoirs of the Indian Museum, 5 (4): 383-392.

Bleeker, P.

1858. Vierde bijdrage tot de kennis der vischfauna van

Biliton. Tijdschrift voor Nederlandsch-Indie, 15:

219-240.

1868. Description de deux especes nouvelles de Blenniofdes de l'lnde Archipe"lagique. Verslagen en

Mededeelingen der Koninklihje Akademie van

Wetenschappen, Letterkunde, en Schoone Kunsten

te Amsterdam, series 2, 2:278-280.

1869. Neuvieme notice sur la faune ichthyologique du

Japon. Verslagen en Mededeelingen der Koninklikje

Akademie van Wetenschappen, Letterkunde, en

Schoone Kunsten te Amsterdam, series 2, 3:239-252.

Borsieri, C.

1904. Contribuzione alia conoscenza della fauna ittiologica

della colonia Eritrea. Annali del Museo Civico di

Storia Naturale di Genova, series 3a, 1:187-220.

Brittan, M. R., A. B. Albrecht, and J. B. Hopkirk

1963. An Oriental Goby Collected in the San Joaquin

River Delta Near Stockton, California. California

Fish and Game, 49 (4): 302-304.

Caldwell, D. K.

1966. Marine and Freshwater Fishes of Jamaica. Bulletin

of the Institute of Jamaica, Science Series, 7:1-120.

Castelnau, F., de

1875. Researches on the Fishes of Australia. In Official

Record, Philadelphia Centennial Exhibition of 1876,

Intercolonial Exhibition Essays, 1875-1876, 11:3-52.

1879. Essay on the Ichthyology of Port Jackson. Proceedings of the Linnean Society of New South Wales,

3 (4):347-402.

Cervigon, M. F.

1966. Los peces marinos de Venezuela. Estacion de Investigaciones Marinas de Margarita, Fundacion La

Salle de Ciencias Naturales, monograph 12, volume

2:443-951.

Chaudhuri, B. L.

1916. Descriptions of Two New Fish from the Chilka

Lake. Records of the Indian Museum, 12 (3): 105-108.

Chu, Y. T., et al.

1962. Synopsis of Fishes of Nanhai. xxxvii + 1184 pages.

Peiping. [In Chinese.]

Chyung Moon Ki

1954. Korean Fishes. 574 pages. Seoul.

1961. Fishes. In Illustrated Encyclopedia, The Fauna of

Korea (2). iv + 861 pages. Seoul: Ministry of Education.

Collette, B. B., and R. W. Yerger

1962. The American Percid Fishes of the Subgenus Villora. Tulane Studies in Zoology, 9(4):213-230.

Cuvier, G. L. F. D., and A. Valenciennes

1836. Histoire naturelle des poissons. Volume 11, 506

pages. Paris.

Dawson, C. E.

1970. The Caribbean Atlantic Blenny Lupinoblennius

dispar (Tribe Blenniini), with Observations on a

Pacific Population. Proceedings of the Biological

Society of Washington, 83 (26):273-285.

1973. Occurrence of an Exotic Eleotrid Fish in Panama

with Discussion of Probable Origin and Mode of

Introduction. Copeia, 1973 (1): 141-144.

Day, F.

1876. The Fishes of India . . . Burma and Ceylon. Volume

1, part 2, pages 169-368, plates 41-81. London.

1888. Observations on the Fishes of India, Part 1. Proceedings of the Scientific Meetings of the Zoological

Society of London for the Year 1888, 3:258-265.

de Beaufort, L. F., and W. M. Chapman

1951. The Fishes of the Indo-Australian Archipelago.

Volume 9, xi + 484 pages, Leiden: E. J. Brill.

DeVis, C. W.

1884a. On New Fish from Moreton Bay. Proceedings of

the Royal Society of Queensland, 1 (3): 144-147.

1884b. New Fishes in the Queensland Museum, No. 4.

Proceedings of the Linnean Society of New South

Wales, 9 (3):685-698.

1886. On a Lizard and Three Species of Salarias, &c. Proceedings of the Royal Society of Queensland, 1885,

2 (2): 56-60.

1887. Annual Report to the Trustees of the Queensland

Museum. Pages 1-14 in Queensland Museum' (Report of Board of Trustees for the Year 1886).

Dutt, S., and V. Visveswara Rao

1961. On the Breeding Habits and Early Developmental

Stages of Petroscirtes bipunctatus Day. Journal of

the Zoological Society of India, 12 (2): 125-274.

Erickson, E. L.

1934. The Introduction of East Indian Coolies into the

British West Indies. The Journal of Modern History, 6 (2): 127-146.

Fowler, H. W.

1903. Descriptions of Several New Fishes from Zanzibar

Island, Two of Which Are New. Proceedings of

the Academy of Natural Sciences of Philadelphia,

55:161-176.

1928. The Fishes of Oceania. Occasional Papers of the

Bernice P. Bishop Museum, Memoir 10: 540 pages.

1931. Fishes Obtained by the Barber Asphalt Company

in Trinidad and Venezuela in 1930. Proceedings of

the Academy of Natural Sciences of Philadelphia,

83:391-410.

1937. Zoological Results of the Third De Schauensee

Siamese Expedition, Part 8: Fishes Obtained in 1936.

Proceedings of the Academy of Natural Sciences of

Philadelphia, 89:125-264.

1959. Fishes of Fiji. 670 pages. Suva, Fiji.
NUMBER 177 81

Fox, H. M.

1926. Cambridge Expedition to the Suez Canal, 1924, I:

General Part. Transactions of the Zoological Society

of London, 22 (1): 1-64.

Fraser-Brunner, A.

1951. Some New Blennioid Fishes, with a Key to the

Genus Antennablennius. Annals and Magazine of

Natural History, series 12, 4 (39):213-220.

Gangulee, N.

1947. Indians in the Empire Overseas. 263 pages. London:

New India Publishing House Limited.

Garman, S.

1903. Some Fishes from Australasia. Bulletin of the Museum of Comparative Zoology, 39 (8):229-241.

Gilchrist, J. D. F., and W. W. Thompson

1908. The Blenniidae of South Africa. Annals of the

South African Museum, 6 (2):97-143.

Giinther, A. C. L. G.

1861. Catalogue of the Acanthopterygian Fishes in the

Collection of the British Museum. Volume 3, 586

pages. London.

1877. Andrew Garrett's Fische der Sudsee, VI. Journal des

Museum Godeffroy, 13:169-216.

Hemprich, F. G., and C. G. Ehrenberg

1899. Pisces (F. Hilgendof, editor). Pages 7-9 in Symbolae,

Physicae . . . , Zoologica. Plates 1-10. Berlin.

Herre, A. W. C. T.

1927. Four New Fishes from Lake Taal (Bombon). Philippine Journal of Science, 34 (3):273-28O.

1934. Notes on New or Little Known Fishes from Southeastern China. Lingnan Science Journal, 13 (2):

285-296.

1935. Notes on Fishes in the Zoological Museum of

Stanford University, VI: New and Rare Hong Kong

Fishes Obtained in 1934. Hong Kong Naturalist,

6 (3,4):285-293.

1940. New Species of Fishes from the Malay Peninsula

and Borneo. Bulletin of the Raffles Museum, 16:

5-26.

1942. Contributions from the Zoological Museum of Stanford University, California, IX: Two New Species

of Petroscirtes and a Key to the Philippine Species.

Copeia, 1942 (2): 111-116.

Hildebrand, S. F.

1939. The Panama Canal as a Passageway for Fishes, with

Lists and Remarks on the Fishes and Invertebrates

Observed. Zoologica, 24 (3): 15-45.

Hubbs, C. L., L. P. Schultz, V. G. Springer, and J. E. Randall

1963. Comments on the Proposed Rejection of the Type

Designations of Jordan & Evermann, 1896-1900 and

1896. Z.N. (S.) 1279. Bulletin of Zoological Nomenclature, 20 (4):257-259.

Jatzow, R., and H. Lenz

1898. Fische von Ost-Afrika, Madagaskar und Aldabra.

Abhandlungen der Senckenbergischen naturforschenden Gesellschaft, 21 (3):497-531.

Jones, S., and M. Kumaran

1965. New Records of Fishes from the Seas around India,

Part II. Journal of the Marine Biological Association of India, 7(1): 108-123.

Jordan, D. S., and A. Seale

1905. List of Fishes Collected by Dr. Bashford Dean on

the Island of Negros, Philippines. Proceedings of

the United States National Museum, 28 (1407): 769-

803.

Jordan, D. S., and J. O. Snyder

1902. A Review of the Blennioid Fishes of Japan. Proceedings of the United States National Museum,

25 (1293):441-504.

Jordan, D. S., and E. C. Starks

1906. List of Fishes Collected on Tanega and Yaku, Offshore Islands of Southern Japan, by Robert Van

Vleck Anderson, with Descriptions of Seven New

Species. Proceedings of the United States National

Museum, 30 (1462):695-706.

1907. List of Fishes Recorded from Okinawa or the Riu

Kiu Islands of Japan. Proceedings of the United

States National Museum, 32 (1541):491-504.

Kamohara, T.

1956. Nine Noteworthy Additions to the Marine Fish

Fauna of Prov. Tosa, Including One New to

Science. Reports of the Usa Marine Biological Station, 3 (1): 1-4.

1957. List of Fishes from Amami-Oshima and Adjacent

Regions, Kagoshima Prefecture, Japan. Reports of

the Usa Marine Biological Station, 4(1): 1-65.

Katayama, M.

1941. A New Blennoid Fish from Toyama Bay. The Zoological Magazine [Dobutsugaku Zasshi], 53 (12):591—

592.

Katayama, M., and H. Ikeda

1973. [Omobranchus uekii from the Inland Sea.] Yamaguchi-ken no Shizen, 3 (8)^8:6-7. [In Japanese.]

Klass, M.

1961. East Indians in Trinidad, xxvii + 265 pages. New

York and London: Columbia University Press.

Kner, R.

1868a. t)ber neue Fische aus dem Museum der Herren

Johann Casar Godeffroy und Sohn in Hamburg.

Sitzungsberichte der Kaiserlichen Akademie der

Wissenschaften in Wien, 58 (1),7:26-3I.

1868b. Folge neuer Fische aus dem Museum der Herren

Joh. Cas. Godeffroy und Sohn in Hamburg. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften in Wien, 58(l),8:293-356.

Lachner, E. A., C. R. Robins, and W. R. Courtenay, Jr.

1970. Exotic Fishes and Other Aquatic Organisms Introduced into North America. Smithsonian Contributions of Zoology, 59:1-29.

Ladd, H. S.

1960. Origin of the Pacific Island Molluscan Fauna.

American Journal of Science, Bradly Volume,

258-A: 137-150.
82

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Lubbock, B.

1935. Coolie Ships and Oil Sailers, x + 180 pages. Glasgow:

Brown, Son and Ferguson.

MacDonald, G. A.

1949. Hawaiian Petrographic Province. Bulletin of the

Geological Society of America, 60:1541-1596.

Macleay, W.

1881. Descriptive Catalogue of the Fishes of Australia,

Part 3. Proceedings of the Linnean Society of New

South Wales, 6(1): 1-138.

1884. Notices on New Fishes. Proceedings of the Linnean

Society of New South Wales, 9 (1): 170-172.

McCulloch, A. R.

1917. Ichthyological Notes. Australian Zoologist, 1 (4):

89-93.

1929. A Checklist of the Fishes Recorded from Australia,

Part 3. Australian Museum, Memoir, 5:329—436.

Menon, A. G. K., and K. V. Rama Rao

1963. On a New Record of the Blenniid Fish Omobranchus kallosoma (Bleeker) from India. Science

and Culture, 29:362.

Menon, A. G. K., and P. K. Talwar

1972. Fishes of the Great Nicobar Expedition, 1966, with

Description of a New Gobioid Fish of the Family

Kraemeriidae. Records of the Zoological Survey of

India, 66 (1-4): 35-62.

Menon, A. G. K., and G. M. Yazdani

1968. Catalogue of Type-Specimens in the Zoological Survey of India, Part 2: Fishes. Records of the Zoological Survey of India, 61 (l-2):91-190.

Millard, N. A. H., and G. J. Broekhuysen

1970. The Ecology of South African Estuaries, Part X: St.

Lucia: A Second Report. Zoologica Africana, 5 (2):

277-307.

Monkolprasit, S. P.

1969. Type Specimens of Fishes at the Kasetsart University

Museum of Fisheries. Notes from Faculty of Fisheries, Kasetsart University, Bangkok, 5:3-8.

Munro, I. S. R.

1958. The Fishes of the New Guinea Region. Papua and

New Guinea Agricultural Journal, 10 (4):97-369.

[Reprinted with same pagination as Territory of

Papua and New Guinea, Fisheries Bulletin J].

1967. The Fishes of New Guinea, xxxvii+650 pages. Port

Moresby, New Guinea: Department of Agriculture,

Stock and Fisheries.

Nagabhushanam, A. K., and G. Chandrasekhara Rao

1972. An Ecological Survey of the Marine Fauna of

Minicoy Atoll (Laccadive Archipelago, Arabian Sea).

Mitteilungen aus dem Zoologischen Museum in

Berlin, 48 (2):265-324.

Norman, J. R.

1943. Notes on the Blennioid Fishes. I: A Provisional

Synopsis of the Genera of the Family Blenniidae.

Annals and Magazine of Natural History, series 11,

10 (72):793-812.

Ogilby, J. D.

1887. Catalogue of the Fishes of New South Wales, with

Their Principal Synonyms. Appendix A in Report

of the Commissioners of Fisheries up to 31 December 1886, Department of Fisheries, New South

Wales. 67 pages.

1910. On New or Insufficiently Described Fishes. Proceedings of the Royal Society of Queensland, 23:

1-55.

Penrith, M. J., and M. L. Penrith

1972. The Blenniidae of Western Southern Africa. Cimbebasia, series A, 2 (5):65-90.

Peters, W.

1855a. Uebersicht der in Mossambique beobachten Fische.

Archiv fur Naturgeschichte, 1:234-282.

1855b. Ubersicht der in Mossambique beobachten Seefische.

Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin, 1855:428-466.

Rama Rao, K. V.

1968. On a Record of the Blenniid Fish Cruantus dealmeida (Smith) from Indian Waters. Journal of the

Bombay Natural History Society, 64 (3):570-572.

Regan, C. T.

1905. On Fishes from the Persian Gulf, the Sea of Oman,

and Karachi, Collected by Mr. F. W. Townsend.

Journal of the Bombay Natural History Society, 16:

318-333.

Richardson, J.

1846. Report on the Ichthyology of the Seas of China

and Japan. Pages 187-320 in Report of the Fifteenth

Meeting of the British Association for the Advancement of Science.

Roberts, G. W.

1957. The Population of Jamaica, xxii + 356 pages. Cambridge University Press.

Rubinoff, I.

1970. The Sea-level Canal Controversy. Biological Conservation, 3 (l):33-36.

Rubinoff, R. W., and I. Rubinoff

1968. Interoceanic Colonization of a Marine Goby

through the Panama Canal. Nature, 217 (5127):476-

478.

Sauvage, H. E.

1880. Description de quelques Blenniofdes de la collection

du Museum d'Histoire Naturelle. Bulletin de la

Societe Philomantique de Paris, series 7, 4(1879-

1880):215-220.

1883. Descriptions de quelques poissons de la collection

du Museum d'Histoire Naturelle. Bulletin de la

Societe Philomantique de Paris, series 7, 7:156-161.

Scott, T. D.

1962. The Marine and Fresh Water Fishes of South Australia. 338 pages. Adelaide: Government Printer.

Seale, A.

1935. The Templeton Crocker Expedition to Western

Polynesian and Melanesian Islands, 1933, No. 27:
NUMBER 177 83

Fishes. Proceedings of the California Academy of

Sciences, series 4, 21:337-378.

Smith, H. M.

1934. The Blenniid Fishes of Siam, with Description of

New Species. Journal of the Siam Society, 9 (3):

315-323.

Smith, J. L. B.

1949. Forty-two Fishes New to South Africa with Notes on

Others. Annals and Magazine of Natural History,

series 12, 2:97-111.

1959. Fishes of the Families Blenniidae and Salariidae of

the Western Indian Ocean. Rhodes University, Department of Ichthyology, Ichthyological Bulletin,

14:229-252.

1965. New Records and New Species of Fishes from

South Africa, Chiefly from Natal. Rhodes University, Department of Ichthyology, Occasional Paper,

4:26-42.

Smith, J. L. B., and M. M. Smith

1967. Fishes of the Tsitsikama Coastal National Park. 161

pages. Johannesburg: The Swan Press Limited.

Smith-Vaniz, W. F., and V. G. Springer

1971. Synopsis of the Tribe Salariini, with Description of

Five New Genera and Three New Species (Pisces:

Blenniidae). Smithsonian Contributions to Zoology,

71:1-72.

Snedecor, G. W., and W. G. Cochran

1971. Statistical Methods. 6th edition, xiv+593 pages.

Ames, Iowa: Iowa State University Press.

Springer, V. G.

1963. Two Species of Indo-west Pacific Blenniid Fishes

Erroneously Described from the Western Atlantic

Ocean. Copeia, 1963 (2):452-454.

1967. Revision of the Circumtropical Shorefish Genus

Entomacrodus (Blenniidae: Salariinae). Proceedings of the United States National Museum, 122

(3582): 1-150.

1968. Osteology and Classification of the Fishes of the

Family Blenniidae. United States National Museum

Bulletin 284:1-85.

1971. Revision of the Fish Genus Ecsenius (Blenniidae,

Blenniinae, Salariini). Smithsonian Contributions

to Zoology, 72:1-74.

1972a. Synopsis of the Tribe Omobranchini with Descriptions of Three New Genera and Two New Species

(Pisces: Blenniidae). Smithsonian Contributions to

Zoology, 130:1-31.

1972b. Additions to Revisions of the Blenniid Fish Genera

Ecsenius and Entomacrodus, with Descriptions of

Three New Species of Ecsenius. Smithsonian Contributions to Zoology, 134:1-13.

Springer, V. G., and W. N. Eschmeyer

1974. Location of Holotypes of Japanese Fishes (Jordan

and Starks, 1906) and Types of Hawaiian Fishes

(Jordan and Evermann, 1903). Copeia, 1974(2):566-

569.

Steindachner, F.

1863. Ichthyologische Mittheilungen (VI). Verhandlungen

der Zoologisch-Botanischen Gesellschaft in Wien,

13:1189-1192.

1876. Ichthyologische Beitrage (V). Sitzungsberichte der

Kaiserlichen Akademie der Wissenschaften in Wien,

74(l-2),l:49-240.

Strasburg, D. W.

1956. Notes on the Blennioid Fishes of Hawaii with Descriptions of Two New Species. Pacific Science,

10 (3):214-267.

Swainson, W.

1839. The Natural History of Fishes, Amphibians and

Reptiles. . . . Volume 2, 452 pages. London.

Swinton, E., and J. Swinton

1859. Journal of a Voyage with Coolie Emigrants, from

Calcutta to Trinidad. 16 pages. London: Alfred W.

Bennett.

Thomson, J. M., and A. E. Bennett

1953. The Oyster Blenny, Omobranchus anolius (Valenciennes) (Blenniidae). Australian Journal of Marine

and Freshwater Research, 4 (2):227-233.

Tomiyama, I.

1950. On a Japanese Blennioid Fish, Dasson elegans

(Steindachner). The Zoological Magazine [Dobutsugaku Zasshi], 59 (9): 17. [In Japanese with English

summary.]

1951. On a Japanese Blennioid Fish, Dasson loxozonus

(Jordan and Starks). The Zoological Magazine

[Dobutsugaku Zasshi], 60(ll):227. [In Japanese with

English summary.]

1952. On a Japanese Blennioid Fish, Dasson japonicus

(Bleeker). The Zoological Magazine [Dobutsugaku

Zasshi], 61(1):11-13. [In Japanese with English

summary.]

Tomiyama, I., and T. Abe

1953. Omobranchus uekii (Katayama) (Blenniidae). Pages

975-978 in volume 49 in Figures and Descriptions

of the Fishes of Japan. Tokyo.

Weber, M.

1908. Susswasserfische von Neu-Guinea: ein Beitrag zur

Frage nach dem fiirheren Zusammenhang von NeuGuinea und Australien. Pages 201-267 in Nova

Guinea, part 2 in volume 5 (Zoology) in Resultats

de I'expedition scientifique Neerlandaise a la

Nouvelle-Guinee en 1903 sous les auspices de Arthur

Wichmann.

1913. Die Fische der Siboga-Expedition. Volume 57 in

Siboga Expedition Report. 710 pages.

Whitehead, P. J. P.

1969. The Reeves Collection of Chinese Fish Drawings.

Bulletin of the British Museum (Natural History),

Historical Series, 3 (7): 193-233.

Whitley, G. P.

1927. A Check-list of Fishes Recorded from Fijian Waters.
84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Journal of the Pan-Pacific Research Institution,

2(1):3-S.

1935. Fishes from Princess Charlotte Bay, North Queensland. Records of the South Australian Museum,

5 (3):345-365.

1945. New Sharks and Fishes from Western Australia,

Part 2. Australian Zoologist, 11 (l):l-45.

1958. Descriptions and Records of Fishes. Proceedings of

the Royal Zoological Society of New South Wales,

1956-1957:28-51.

1959. Ichthyological Snippets. Australian Zoologist, 12(4):

310-323.

Woodring, W. P.

1966. The Panama Land Bridge as a Sea Barrier. Proceedings of the American Philosophical Society,

110(6):425-433.

Yamakawa, T.

1971. Additional Records of Marine Fishes from Amami

(iv). Reports of the Usa Marine Biological Station,

18 (2): 1-21.

Yazdani, G. M.

1963. On a New Record of a Blenniid Fish, Petroscirtes

kochi M. Weber, from Andamans. Current Science,

32 (9):413.

Zugmayer, E.

1913. Die Fische von Balutschistan. Abhandlungen der

Koniglich Bayerischen Akademie der Wissenschaften,

Mathematisch-physikalische Klasse, 26 (6): 1-35.
Figures 8-35

The Species of Omobranchus
86

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 8. Omobranchus anolius, USNM 197621, male, 46.9 mm SL Sydney, Australia.

(Drawn by J. R. Schroeder.)

FIGURE 9—Omobranchus aurosplendidus, CAS SU 29088, male, 99.5 mm SL,

Heungchow, Kwangtung Province, China. (Drawn by J. R. Schroeder.)
NUMBER 177 87

tiu

FIGURE 10—Omobranchus aurospendidus, USNM 201464, female, 93.6 mm SL, Hong Kong.

(Drawn by J. R. Schroeder.)

FIGURE 11.—Omobranchus bandittis, BMNH 1919.4.1.30, male, 49.5 mm SL, Durban, South Africa:

a, lateral view; b, ventral side of head. (Drawn by S. L. Chambers.)
88

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 12. Omobranchus elegans, USNM 71530, male, 53.8 mm SL, Misaki, Sagami Bay, Japan:

a, lateral view; b, ventral side of head. (Drawn by J. R. Schroeder.)

a

FIGURE 13—Omobranchus elongatus, USNM 209425, male, 38.9 mm SL, Goh Kram, Gulf of

Thailand: a, lateral view; b, ventral side of head. (Drawn by J. R. Schroeder.)
NUMBER 177 89

FIGURE 14.—Omobranchus elongatus, USNM 209425, female, 34.3 mm SL, Goh Kram, Gulf of

Thailand: a, lateral view; b, ventral side of head. (Drawn by J .R. Schroeder.)

FIGURE 15.—Omobranchus elongatus, USNM 201876, male, 45.7 mm SL,

near Marokibani, Madagascar. (Drawn by J. R. Schroeder.)
90

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 16.—Omobranchus fasciolatoceps, CAS SU61121, Hong Kong: a, female,

51.0 mm SL; b, male, 61.7 mm SL. (Drawn by K. H. Moore.)

a

a

FIGURE 17.—Omobranchus fasciolatus, USNM 201869, Astola Island, West Pakistan, a-b, male,

44.2 mm SL: a, lateral view; b, ventral side of head; c, female, 48.4 mm SL. (Drawn by J. R.

Schroeder.)
>\\\\Vv

FIGURE 18 Omobranchus ferox, USNM 201559, male, 34.5 mm SL, Nias Island, Indonesia.

(Drawn by B. L. Holden.)

a

FIGURE 19.—Omobranchus germaini, CAS 13780, male, 63.7 mm SL, One Tree Island, Great

Barrier Reef: a, lateral view; b, ventral side of head. (Drawn by T. R. Schroeder.)

FIGURE 20.—Omobranchus germaini, USNM 204027, female, 45.1 mm SL,

northwest of Yeh-Liu, Taiwan. (Drawn by J. R. Schroeder.)
92

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 21.—Omobranchus lineolatus: a, WAM P.10100, male, 36.3 mm SL, Riddell Beach,

Broome, Western Australia (Drawn by J. R. Schroeder.); b, holotype of Petroscirtes lineolatus,

after Kner (1868b).

a

FIGURE 22.—Omobranchus lineolatus, RMNH 26918, female, 66.8 mm SL, Merauke, New Guinea:

a, lateral view; b, ventral side of head. (Drawn by J. R. Schroeder.)
NUMBER 177 93

FIGURE 23.—Omobranchus loxozonus, USNM 70761, male, 49.1 mm SL, Tanegashima, Japan:

a, lateral view; b, ventral side of head; c, area of head just posterior to eye. (Drawn by J. R.

Schroeder.)

FIGURE 24.—Omobranchus mekranensis, USNM 200604, male, 46.4 mm SL,

Astola Island, West Pakistan. (Drawn by J. R. Schroeder.)
94

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

i

FIGURE 25 Omobranchus meniscus, Chantabun, Thailand: a, USNM 119685, holotype, male,

60.1 mm SL; b, USNM 211153, female, 54.8 mm SL. (Drawn by J. R. Schroeder.)

FIGURE 26.—Omobranchus punctatus, USNM 206389, male, 49.4 mm SL, Trincomalee, Ceylon.

(Drawn by J. R. Schroeder.)
NUMBER 177 95

FIGURE 27.—Omobranchus punctatus, USNM 174330, male, 52.7 mm SL,

Northern Territory, Australia. (Drawn by M. H. Lester.)

a

FIGURE 28 Omobranchus r. rotundiceps, USNM 174337, male, 48.2 mm SL. Northern Territory,

Australia: a, lateral view; b, ventral side of head. (Drawn by J. R. Schroeder.)
96

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 29.—Head color pattern exhibited by certain species of Omobranchus. a-b, O. r. rotundiceps, USNM 205712, male, 40.1 mm SL, Sydney, Australia: a, lateral view; b, ventral view.

c-d, O. r. obliquus, CAS GVF reg. no. 1858, male, 34.9 mm SL, Guam: c, lateral view;d, ventral

view, e, O. germaini, ANSP 109700, male, 49.6 mm SL, Little Hope Island, Great Barrier Reef.

/, O. ferox, CAS GVF reg. no. 1486, female, 38.9 mm SL, Trat Bay, Gulf of Thailand. (Drawn by

J. R. Schrocder.)
NUMBER 177 97

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FIGURE 30.—Omobranchus steinitzi, USNM 209433, holotype, male, 32.0 ram SL,

Cundabilu, Dahlak Archipelago, Red Sea. (Drawn by J. R. Schroeder.)

FIGURE 31.—Omobranchus verticalis, USNM 210616, male, 33.3 mm SL, Boat Passage,

Brisbane River, Queensland, Australia. (Drawn by J. R. Schroeder.)

FIGURE 32.—Omobranchus woodi, RUSI 2357, male, 69.6 mm SL, Knysna Lagoon, South Africa.

(Drawn by J. R. Schroeder.)
98

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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FIGURE 33.—Omobranchus zebra, USNM 209380, male, 43.0 mm SL, Trat Bay Gulf of Thailand:

a, lateral view; b, tips of some anal-fin rays: two anterior rays with subdistal tabs; two posterior

rays without tabs; all four rays with flattened tips (see "Sexual Dimorphism" under Omobranchus). (Drawn by J. R. Schroeder.)

FIGURE 34.—Omobranchus zebra, photocopy of unpublished Bleeker color plate representing

holotype of Petroscirtes zebra Bleeker. (Courtesy of RMNH.)
NUMBER 177 99

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FIGURE 35.—Omobranchus species, USNM 211092, male, 22.7 mm SL, Hikkaduwa, Ceylon.

(Drawn by J. R. Schroeder.)

Figures 36-52

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