VICTOR G. SPRINGE Revision of the

: Fish Genus Ecsenius

(Blenniidae, Blenniinae,

Salariini)

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 72

victor G. springer Revision of the

Fish Genus Ecsenius

(Blenniidae, Blenniinae,

Salariini)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1971
ABSTRACT

Springer, Victor G. Revision of the Fish Genus Ecsenius (Blenniidae, Blenniinae,

Salariini). Smithsonian Contributions to Zoology, 72:1-74, I971.-The genus

Ecsenius comprises 18 species of Indo-Pacific reef-dwelling fishes. The species are

generally characterized by a high degree of sexual dimorphism and geographic

variation as indicated by statistically significant differences in average numbers

of various meristic and proportional characters and color pattern types.

Keys, illustrations and diagnoses, and nomenclatural and zoogeographical discussions are provided. Four new species are described: E. aroni from the Gulf of

Aqaba, E. oculus from the western Pacific, E. bimaculatus from Borneo, and

E. bandanus from Banda Island.

A discussion is given of a particularly complex taxonomic problem involving

three nominal species occurring in the Red Sea and Gulf of Aqaba. E. nigrovittatus

is known only from the southern Red Sea, E. frontalis is known from the northern

Gulf of Aqaba to the Bay of Tadjoura (Gulf of Aden), and E. albicaudatus is

known from the northern Gulf of Aqaba to the southern Red Sea. Except for

color pattern all three forms are very similar morphologically, have been collected

together from one small rock, and are very different from any other species of

Ecsenius. The problem of whether the three forms constitute one or more species

is not solvable presently, but the three nominal species are treated here as color

pattern forms of single species, E. frontalis.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

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

Washington, D.C. 20402 - Price tl.25
Victor G. Springer Revision of the

Fish Genus Ecsenius

(Blenniidae, Blenniinae,

Salariini)

Introduction

Chapman and Schultz (1952), with seventy specimens for study, last reviewed the Indo-Pacific genus

Ecsenius. They recognized eleven species, including

two species for which they had examined no specimens. Since their review five additional species of

Ecsenius have been described (not all properly allocated to genus). Intensive collecting in recent years

has made available well over one thousand specimens of Ecsenius, including material of all the

described species and additional material of undescribed species. The new material enables the revision of Ecsenius presented here.

I recognize eighteen species of Ecsenius, of which

four are new. Since some species of Ecsenius are restricted to poorly collected, relatively deep water,

and because several of the species appear to be

restricted in their geographic distribution, I believe

that there will be a large increase in the number of

species over that which I report when more thorough collecting has taken place. The faunistically

rich Indo-Malayan region is one of the most poorly

sampled for fishes, and it is therefore probable that

this region will provide most of the new Ecsenius

species collected in the future.

The species of Ecsenius are relatively small, not

exceeding 83 mm standard length, and are known,

with rare exceptions, only from coraliferous areas.

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

Washington, D.C. 20560.

They live at depths from about one meter down

to at least 39 meters, although most species seem

to have a relatively restricted depth range. The

species may occur in the surge zone, tide pools, or

the relatively quiet lagoons of atolls.

The genus is a highly specialized one, exhibiting

anatomical peculiarities not found elsewhere among

the blenniids (see generic discussion). Some of the

species apparently are mimics, many are very colorful, while others are very somber in tone. One species, Ecsenius bicolor, has been maintained and bred

in the laboratory (Wickler, 1965a), although it was

not possible to rear the young.

Chapman and Schultz (1952) believed that the

species of Ecsenius formed a close-knit group. I cannot agree with this. While their generic affiliation is

more clear-cut than it is for the species of many

blenny genera, there seems to have been an unusually high degree of adaptive radiation among the

species of Ecsenius. Although I can recognize small

groups of species within the genus, I am unable to

relate the groups.

Methods

MEASUREMENTS.—When possible, I made all measurements on the left side of each specimen, using a

set of needlepoint dial calipers reading directly to

the nearest tenth of a millimeter. Measurements

were converted to percent of standard length.

Standard length (SL): This measurement was

1
2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

taken from the midtip of the upper lip to the midlateral posterior margin of the hypural vertebra as

indicated externally on the specimen. Variably, the

anteriormost point of the head is in advance of the

upper lip, but this portion of the head was ignored

in measurements.

Nasal cirrus length (Table 10): Most species have

a single, simple cirrus on the posterior rim of the

anterior nostril; two species, bicolor and namiyei,

also have a shorter, simple cirrus on the anterior rim

of the anterior nostril. On occasion any of these cirri

may bear a short secondary branch. The length of

the posterior cirrus was measured as the greatest distance from the point where the cirrus joined the

nostril to the distal tip of the cirrus.

Dorsal fin spine and ray lengths (Tables 5, 7, 8):

These measurements were taken from the proximalmost point of the posterior axil formed by the element with the dorsal body contour to the distal tip

of the element, disregarding the normal curvature

of the element. In a few instances (specimens of E.

namiyei and E. midas and some specimens of E.

frontalis) the lengths of the dorsal rays and spines

were taken from radiographs. This was necessary

because the dark skin overlying the fin elements

obscured them from view.

Longest caudal ray (Table 9): This measurement

was taken from the proximalmost point of the ray,

which overlaps the upper hypural plate, to the

distal tip of the ray.

Dorsal fin notch (Table 5): The notch, if present,

was measured as the greatest distance from the

distal tip of the first dorsal ray to the distal edge of

the fin membrane between the last dorsal spine

and first dorsal ray. This distance was divided by

the length of the first dorsal ray and the resulting

proportion recorded as the nearest one-ninth of the

length of the first dorsal ray. Occasionally the distal

tip of the penultimate dorsal spine extended over

the ultimate spine and attached to the first dorsal

ray. In such instances the notch depth was measured from the point of attachment of the penultimate spine to the tip of the first ray.

Lateral line: The dorsal fin element under which

the lateral line terminated was recorded (Table 6).

Fin elements were numbered consecutively from the

anteriormost to the posteriormost without distinguishing spines from rays. If the lateral line terminated below the space between two elements, the

nearest even-numbered element was recorded. With

few exceptions the lateral line terminates below the

spinous portion of the dorsal fin.

MERISTIC AND OTHER COUNTABLE CHARACTERS

(Tables 1-4, 6, 11, 12, 15-18).—All specimens were

radiographed and counts of the dorsal, anal and

caudal fin elements, caudal and precaudal vertebrae,

and epipleurals were taken from radiographs. Occasionally a dorsal or anal fin ray was absent but its

proximal pterygiophore was present. In such instances the ray was counted as if present. No counts

of caudal vertebrae were recorded for specimens

with apparent vertebral fusions. Pectoral and pelvic

ray counts were made on the specimens. Counts for

bilaterally paired characters are for one side only

(the left, when possible, unless otherwise noted.

Dorsal spines: In some species the terminal dorsal

spine is so reduced that it can be seen only on radiographs or after osteological preparation.

Dorsal rays: Each segmented element was counted

separately (the last two counted as two).

Anal spines: There are invariably two, but the

first in females is frequently not visible externally.

Anal rays: Each segmented element was counted

separately (the last two counted as two); the terminal pterygeophore always supports a single segmented ray (unlike some other blenniids; see

Springer, 1967).

Segmented caudal rays: Except for certain small

specimens (see below) each segmented ray was

counted, including, where they occur, the single

dorsal and ventral rays with a single segmental

joint. I noted that occasional specimens less than

about 25 mm had one or two more segmented caudal

rays than larger specimens. These additional segmented rays were usually the ventralmost and, secondarily, the dorsalmost of the segmented rays, and

they never had more than two segments; they were

either paired structures with right and left halves

or were paired for most of their length and fused

at their tips. I believe that such rays lose their

segmentations with growth and become fused for

much of their length. Counts for specimens under

25 mm SL are not included in Table 3 or the

species descriptions.

Procurrent caudal rays: These are the unsegmented elements described by Springer (1968). As

they are not bilaterally paired structures, they could

be called spines. As they appear to be serially homologous with the segmented rays discussed above and

usually have a bilaterally forked base, they give
NUMBER 72

evidence of having gone through a raylike stage in

ontogeny. In the posterior-most procurrent ray the

base may be cleft for almost half the length of the

element, but the distal portion is always fused and

spinelike with no evidence of a joint line extending

distally from the cleft. Procurrent ray counts for

specimens under 25 mm are not included in the

species descriptions. In Table 3, however, total

caudal element counts do include data from these

specimens.

Total caudal elements: This is the sum of the

dorsal and ventral procurrent caudal rays and the

segmented caudal rays.

Pelvic fins: A spine and three segmented rays are

always present, but the spine is imbedded, closely

applied to the first pelvic ray, and not visible externally. In addition, the innermost pelvic ray is

reduced and often impossible to see externally.

Vertebrae: Precaudal vertebrae are those lacking

a hemal spine; the caudal vertebrae all bear a hemal

spine, although that of the hypural (last) vertebra

is fused with the ventral hypurals.

Epipleural ribs: The anterior epipleural ribs are

not possible to count on radiographs, but from examination of cleared and stained specimens are

known to begin invariably on the first vertebra (the

pleural ribs begin invariably on the third vertebra).

The epipleural rib count is actually a count of the

number of vertebrae from the anteriormost to the

posteriormost bearing an epipleural rib, regardless

of whether it bears a pair or only one, and that on

the right or left side. Thus, occasional vertebrae

lacking epipleural ribs, that occur between vertebrae bearing epipleural ribs, are counted as if they

had these ribs.

Dentary teeth (Table 4): Ecsenius is unique

among blenniids in having two sets of caninesanterior and posterior. The anterior canines, usually a single tooth on each side, occur in line with

the incisoritorm, comblike teeth of the dental plate

(Springer, 1968). They are usually only slightly

larger than the adjacent incisor teeth and are frequently blunted and difficult to distinguish from the

incisors, but in E. midas, and occasional specimens

of other species, they are quite large, pointed, and

distinct. The posterior canines (usually one or,

rarely, two, in all but one species) are posterior,

slightly medial to, and well separated from the anterior teeth. They are small and frequently blunted.

In one species, E. mandibularis, there is a series of

up to eight of these posterior canines on each side.

In making the dentary tooth counts the anterior

canines and all the incisors are included. The posterior canines were counted separately. The posterior canines frequently are not developed in specimens below 25 mm SL.

Premaxillary teeth: All the teeth in the premaxillary dental plate were counted. These counts are

difficult and were made on only a relatively few

specimens.

Gill-rakers and pseudobranchial filaments: All the

gill-rakers on the first arch and the pseudobranchial

filaments of one side were counted.

Color pattern: Unless otherwise stated, color pattern descriptions are based on preserved specimens.

Statistical tests: Student's Mest, two sided (Simp

son, Roe and Lewontin, 1960), was employed to test

the difference between means of some samples. Differences were considered to be significant when

p = .01 or less.

Synonymies: The synonymies under each species

include only references to original species descriptions.

Much of the data concerning each species is to be

found under the general accounts of meristics, sexual dimorphism, etc., and is not always repeated

under the individual species accounts.

Institutional abbreviations: In the material lists

and acknowledgments the following abbreviations

are used:

AMS—Australian Museum, Sydney; ANSP—Academy of Natural Sciences of Philadelphia; BMNH—

British Museum (Natural History); BPBM—Bernice

P. Bishop Museum, Honolulu; CAS—California

Academy of Sciences, San Francisco (also GVF for

those specimens in the George Vanderbilt Foundation collections at CAS); FMRI—Central Fisheries

Marine Research Institute, Mandapam Camp, India; HUI—Hebrew University, Israel (some HUI

numbers include "HUJ" as part of the catalogue

number); ISZZ—Institut fur Spezielle Zoologie und

Zoologisches Museum, Berlin; MNHN—Museum

National d'Histoire Naturelle, Paris; NFIS—NaturMuseum und Forschungs-Institut Senckenberg,

Frankfurt; RU—Rhodes University, Department of

Ichthyology, South Africa; UMML—University of

Miami Rosenstiel School of Marine and Atmospheric Sciences; USNM—United States National

Museum of Natural History; UTAI—University of

Tel Aviv, Israel; UW—University of Washington,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Seattle; ZMA—Zoologisch Museum, Amsterdam;

ZITH—Zoological Institute, University of Tokyo.

Full locality citations have usually not been included in the material lists.

Ecsenius McCulloch

Ecsenius McCulloch, 1923, p. 121 [type-species: Ecsenius mandibularis McCulloch, 1923, by original designation].

Pescadorichthys Tomiyama, 1955, p. 8 [type-species: Salarias

namiyei Jordan and Evermann, 1902, by original designation].

DIAGNOSIS.—Fishes of the tribe Salariini (Springer,

1968), subfamily Blenniinae, (1) lacking an ossified median ethmoidl

, (2) having the lateral extrascapulars fused with the pterotics, (3) four circumorbital bones, (4) both anterior and posterior

canines in the lower jaw (the anterior frequently

difficult to distinguish from the adjacent incisors

except in osteological preparations), (5) dorsomesially extending processes arising from the proximal

ends of at least the third to fifth epipleural ribs, (6)

all fin rays unbranched, (7) upper and/or lower

caudal lobes of most mature males and many females

elongated.

Characters (1), (4), and (5) distinguish Ecsenius

from all other Blenniidae. In addition character (2)

distinguishes Ecsenius from all blenniids except a

few species in the subfamily Nemophidinae, and

character (7) distinguishes Ecsenius from all other

Salariini and Blenniini.

Further characterization: Dorsal spines 10 to 14

(rarely 10 or 11); dorsal rays 12 to 21 (rarely 12);

anal elements II, 13 to 23; pectoral rays 12 to 16;

pelvic elements I, 3 (but obvious pelvic rays 2 or 3);

segmented caudal rays 13 or 14 (rarely 15); vertebrae 10 or 11 (9 and 12 in one specimen each) -f19 to 29 = 29 to 40; dentary incisor teeth (including

anterior canines) 13 to 64; premaxillary incisor

teeth 26 to 148, (usually 2 to 3 times as many upper

incisors as lower incisors).

Lateral line extending posteriorly to beneath level

of seventh dorsal spine to first dorsal ray. Dorsal

1

1 (Springer, 1968) erroneously reported that Ecsenius had

a median ethmoid that was encapsulated by the lateral ethmoids and that projected into the orbital region of the skull.

I have reexamined my material and additional material and

find only a small amount of cartilage in the area where an

ossified median ethmoid would be expected. The structure

labelled as the median ethmoid in Springer (1968, plate 10,

center) is actually a process on the parasphenoid.

fin varying from entire to deeply notched between

dorsal spines and rays. Last dorsal and anal fin rays

attached by membrane to caudal peduncle. Cirri

present on anterior nostril; none on eye or nape.

Distinct occipital fleshy crest absent. Circumorbital

and preoperculomandibular pores in simple series

(no paired pores); one mid-predorsal, supra temporal

pore. Lips entire (no crenulae).

Eighteen species occurring in coraliferous areas

of the Indian and western Pacific oceans and their

associated seas and gulfs.

Subgenera: Ecsenius comprising 17 species, and

Anthiiblennius, monotypic, differentiated in the key.

NOMENCLATURE.—Tomiyama compared Pescadorichthys only to the type-species of Ecsenius, E.

mandibularis. All the characters, except one, that

he used to differentiate the two genera vanish when

all the species treated here are considered. The one

character, presence of a series of canines posteriorly

in each jaw, is distinctive of E. mandibularis. I do

not believe that Pescadorichthys should be maintained, however, in view of the great number of

pecularities exhibited by E. mandibularis in common with the other species here included in

Ecsenius. Indeed, a few specimens of E. mandibularis have only two posterior canines on one

side of the jaw, as do a few specimens of the other

species, and so there is a degree of overlap in the

character.

RELATIONSHIPS.—Ecsenius is such a highly specialized genus that it is not possible to select another

genus of blenniids as its nearest relative. It is only

with reluctance that I do not recognize Ecsenius

as constituting a separate tribe of blenniids. The

excavation of the premaxillaries and dentaries, the

high number of premaxillary teeth, and the floating

nature (Springer, 1968) of the incisor teeth of

Ecsenius place it with the Salariini.

SEXUAL DIMORPHISM.—The species of Ecsenius exhibit a number of diverse morphological features

that are associated with sex and, in males, with

presumed degree of maturity. The primary sexual

dimorphism of Ecsenius is that typical of all

blenniids: the first anal spine in females is usually

greatly reduced, not visible externally and completely included in a somewhat triangular fleshy

lobe embodying a large urogenital opening and a

papilla. In males the anal spines are distinct and

the urogenital opening is a minute orifice at the

tip of a short, slender, or truncate tube.
NUMBER 72

In at least several species of Ecsenius there is a

tendency for males to exceed females in standard

length attained. In E. gravieri the largest male

was 61 mm and the largest female was 46 mm (12

of 29 males over 24 mm exceeded 46 mm, but only

1 of 24 females over 24 mm attained that size). In

E. bicolor the largest male was 77 mm and the

largest female was 59 mm (40 of 128 males over

24 mm exceeded 55 mm, but only 2 of 110 females

over 24 mm exceeded 55 mm). In E. mandibularis

the largest male was 51 mm and the largest female

was 39 mm (23 of 45 males over 24 mm exceeded

39 mm; there were 40 females over 24 mm). In

E. yaeyamaensis the largest male was 51 mm and

the largest female was 42 mm (15 of 84 males over

24 mm exceeded 42 mm; there were 71 females

over 24 mm). In E. nalolo the largest male was

51 mm and the largest female was 43 mm (15 of

81 males over 24 mm exceeded 43 mm; there were

68 females over 24 mm). In E. oculus, E. namiyei,

E. midas, and E. lividinalis, of which there were

relatively few specimens, the largest specimen was

a male. In E. lineatus, of which I had only 13

specimens (6 were males), the largest specimen was

a female. In E. frontalis large males and females

were relatively common in all three color pattern

forms (see discussion under species account), but

the largest female was larger than the largest male

in the nigrovittatus form. The sexes attained about

equal size in E. aroni, E. pulcher, and E. opsifrontalis. The list on this page gives the sizes of the

largest specimen examined for each sex of each

species.

These sexual differences are possibly associated

with territoriality. It has been noted for freshwater

fishes (Collette and Yerger, 1962, p. 220, for significant reference citations) that in those species

with a well-developed territory, the males are larger.

During field observations on nine different species

of Ecsenius I noted that all appeared to have a

very limited range of movement. Individuals were

occasionally seen to drive off fishes (nonblenniids)

that approached them, although most often the

Ecsenius retreated. An individual Ecsenius occupying a hole in a rock in an aquarium would successfully drive off another attacking Ecsenius whether

or not of the same species or of larger size. If the

occupant left the hole and another individual

midas

frontalis

frontalis form

albicaudatus form

nigrovittatus form

pulcher

gravieri

aroni

bicolor

namiyei

lineatus

stigmatura

lividinalis

yaeyamaensis

nalolo

oculus

opsifrontalis

prooculis

bimaculatus

bandanus

mandibularis

female

56.0

505

50.5

48.3

58.0

465

40.8

59.6

665

65.9

44.9

305

42.6

43.2

493

54.8

—

_

—

39.0

male

83.1

58.6

51.6

44.0

58.4

60.8

40.8

77.6

81.8

60.1

37.7

33.0

51.9

51.3

53.8

35.8

395

31.7

31.4

51.1

entered it, however, the new occupant could not

be driven out by the previous occupant.

In E. bicolor (Table 15) and Taiwan specimens

of E. namiyei (Table 16) males have significantly

(p = less than .001) higher average numbers of

dorsal and anal rays and caudal vertebrae than

females. Ethiopian males of E. frontaris (frontalis

form color pattern) had significantly higher average

numbers of anal rays (p = less than .001) and

vertebrae (p = between .02 and .01) than Ethiopian

females (frontalis form color pattern). The other

E. frontalis color pattern forms from Ethiopia and

the frontalis and albicaudatus color pattern forms

from the Gulf of Aqaba did not exhibit sexual

dimorphism in these two characters. In those other

species for which there were large numbers of

specimens, there did not appear to be sexual

dimorphism in these meristics except for numbers

of dorsal rays in E. pulcher, where males had a

significantly higher average number than females

(Table 16; p = .001).

Mandibular (dentary) tooth counts were examined

for sexual dimorphism. Most of the species exhibited none for this character, but in three species,

E. bicolor, E. yaeyamaensis (both Table 4), and

E. nalolo (Table 17), females had significantly

higher average numbers of teeth (p = less than

.001). Gulf of Aqaba and northern Red Sea speci-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mens of E. nalolo were considered in one test,

southern Red Sea and Indian Ocean specimens in

another test. Both tests were significant.

In E. mandibularis the third dorsal spine (Table

7) is considerably longer in males than females

(only the southern Queensland specimens were

checked). Though few specimens were available, it

seems that the same type sexual dimorphism may

also occur in E. opsifrontalis, E. nalolo, and E.

lineatus.

In E. mandibularis, E. yaeyamaensis, E. nalolo,

non-Australian E. bicolor, and E. argus the males

tend to have a higher average fifth dorsal ray length

than females (Table 8). There did not appear to

be much difference between males and females of

Australian E. bicolor in regard to fifth dorsal ray

lengths.

Either the upper and/or lower lobes of the

caudal fin in males (less so in females) of Ecsenius

species generally become increasingly longer with

increase in SL, and males of most species tend to

have longer caudal fins than females (Table 9);

but the two sexes are about the same in E. gravieri

(Figure 11). In E. mandibularis and E. bicolor males

and E. aroni females the caudal fins may reach a

peak relative length well below the maximum SL

and then begin to decrease in relative size with

increase in SL.

Nasal cirrus lengths (Table 10) tend to be longer

in males of E. oculus, E. opsifrontalis, E. yaeyamaensis, E. nalolo, and E. mandibularis than in

females.

In addition to the above features the males of

most, if not all, species of Ecsenius tend to develop

thick fleshy tips on most of the anal rays (Figures

19, 33, 35, 36). Such developments may accompany

sexual maturity as males of similar size within a

species may or may not show such development.

The fleshy ray tips appear to be a parallel of the

condition found in many blenniids where the anal

spines and anterior anal rays may be enveloped in

rugose folds in presumably mature males.

Finally, the color pattern of preserved males is

in general darker or more contrasty than that of

females.

Many of the features discussed above are similarly

and variously exhibited by other species of the

Blenniidae. I am unable to explain why there

should be meristic sexual dimorphism, especially

so as there is usually a broad overlap in meristics

between males and females and the average differences are frequently one element or less in a particular character. On the other hand, the longer

fin elements and cirri of males are probably of

importance in sex recognition and courtship.

MIMICRY.—At least two species of Ecsenius bear

a strong, but superficial, resemblance to species of

the nemophidinine genus Meiacanthus Norman.

E. bicolor resembles M. atrodorsalis (Giinther) in

both color and morphology. G. Losey (in litteris)

has observed the two species together at Eniwetok,

where it appeared to him that E. bicolor was the

mimic as it was the rarer of the two species. He

described both species as being dark gray-blue

anteriorly and bright yellow-orange posteriorly. My

observations of E. bicolor offer some contrast to

those of Losey's. At One Tree Island on the

southern end of the Great Barrier Reef, I collected

over 100 E. bicolor and only one M. atrodorsalis,

and that one was taken in a collection where no

E. bicolor were seen or collected. The color patterns

were somewhat different also. The E. bicolor were

bright blue anteriorly and red-orange posteriorly.

The M. atrodorsalis was gray-blue anteriorly and

yellow posteriorly. It is possible that there is population variation or that different species of Meiacanthus were involved.

Both E. gravieri and M. nigrolineatus Smith-Vaniz

have been collected together often and are very

similar in morphology and color pattern, both in

life and in preserved material. There is good circumstantial evidence supporting the idea of a

mimic-model relationship between these two species.

In the Gulf of Aqaba the two species are represented only by pale forms; that is, with relatively

little black pigment. In the southern Red Sea both

species are represented only by melanistic forms.

Further considerations of possible mimicry involving these two species will be discussed elsewhere

(Springer and Smith-Vaniz, ms).

Starck (1969) discussed mimicry between E. midas

and the anthiine Anthias squamipinnis Peters. He

believed that E. midas was the mimic because it was

much less abundant than the anthiine. He was not

able to establish the basis for the mimicry but believed that the blenny, when occurring in aggregations of the anthiid, probably was protected from

predation because of the difficulty a predator has in

singling out a prey from a large group. My observations in the Red Sea also indicate that E. midas was
NUMBER 72

exceedingly rare and that A. squamipinnis was one

of the dominant species, but I saw no predation of

or interaction between these two species.

Among the blennies, the only other species reported to be involved in mimicry are the aggressive

nemophidinines Aspidontus taeniatus Quoy and

Gaimard, which mimics the cleaner wrasse Labroides

dimidiatus (Cuvier and Valenciennes)—see Randall

and Randall (I960)—and Runula azalea Jordan

and Bollman, which aggregates with the wrasse

Thalassoma lucasanum (Gill) where mimicry as

such was not specified (see Hobson, 1969).

Zoogeography

Ecsenius occurs from the Red Sea and east African

coast as far west as the Tonga Islands, as far north

as the Ryukyu Islands, and as far south as the

southern end of the Great Barrier Reef. It has

been reported questionably from the Hawaiian

Islands (I doubt its occurrence there; see discussion

under E. bicolor species account). The genus is

unreported from the central and eastern Pacific

and Atlantic oceans. While most of the species are

inhabitants of shallow waters, they are not taken

along shore. With only exceptional individuals, the

species are known only from coraliferous areas.

Over half the species of Ecsenius are relatively

restricted geographically, or form identifiable populations in different areas. Of the five species occurring in the Red Sea, E. aroni (Figure 2), E. gravieri

(Figure 1), and E. frontalis (Figures 1 and 5) occur

only there (the latter two species also occur in the

contiguous Gulf of Aden). Of the other two Red

Sea species, the E. nalolo (Figure 2) population

from the Gulf of Aqaba is recognizable from the

southern Red Sea and Indian Ocean populations

in having more teeth, and the four Red Sea specimens of E. midas (Figure 2), which species is

known from only 12 specimens altogether, have

slightly lower vertebral counts than the six Indian

Ocean specimens. However, the two specimens of

E. midas from the Great Barrier Reef are intermediate in these counts.

Only one species of Ecsenius, E. pulcher (Figure

1), is known from the Persian Gulf and Arabian

Sea, and it is not found elsewhere. Of the four

species known from Australia (all from the Great

Barrier Reef), E. mandibularis (Figure 3) is endemic,

E. yaeyamaensis (Figure 2) is represented by a

unique color pattern population, and the population of E. bicolor (Figure 2) has higher average

numbers of dorsal and anal rays and caudal vertebrae than populations of the species from any other

"W bicolor

A frwrtali*

• flrovitri

• namiyai

O pulch«r

n

FIGURE 1 .—Distribution of certain species of Ecsenius. Irregular broken line indicates approximate

position of andesite line (see zoogeography section).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

O lineatus

A midat

nalolo

opsifrontalis

yaeyamaensis

FICURE 2.—Distribution of certain species of Ecsenius. Irregular hroken line indicates approximate

position of andesite line (see zoogeography section).

locality except the South China Sea. E. prooculis,

E. bimaculatus, and E. bandanus (all Figure 4)

appear to be highly restricted in their ranges, and

E. oculus (Figure 3) comprises several recognizable

color pattern populations in the Pacific.

Only three species occur in both the Indian and

Pacific oceans. E. bicolor (Figure 1), which is widespread in the Pacific, reaches as far west as the

Maldives in the central Indian Ocean. E. oculus

(Figure 3), also moderately widespread in the Pacific, occurs along the Indian Ocean side of Sumatra,

and E. lineatus occurs from northern Taiwan and

the South China Sea to the Maldives. One might

place among these species a fourth, which I have

recognized as two species: E. yaeyamaensis (Figure

2), Pacific Ocean, and E. nalolo (Figure 2), Indian

Ocean, which at the least are completely distinguishable based on color pattern. There is a considerable distance between the nearest known populations of these last two species.

While it is difficult to explain and may only be

an artifact of collecting, only 3 of the 12 species

occurring in the Pacific, E. bicolor (Figure 1), E.

yaeyamaensis (Figure 2) and E. opsifrontalis (Figure

2) have been collected east of the andesite line

(Macdonald, 1949), which separates the continental

from the noncontinental rocks. The approximate

position of the line, based on Menard's (1964, fig.

4.21) delineation, is indicated in Figures 1-4. Of

these three species, E. opsifrontalis occurs only from

the line eastward and E. yaeyamaensis has its eastern

limits at or just beyond the line. The nearest relative of E. opsifrontalis, E. oculus (Figure 3), has its

eastern limit on the line. The last two species have

not been collected together.

It would appear that in the Pacific, at least,

Ecsenius had its evolutionary origin in the older

continental areas and has not been too successful

in invading the more recent oceanic islands. Thus,

E. opsifrontalis is probably a recently evolved

species, perhaps a derivative of E. oculus, and the

presence of E. bicolor east of the andesite line

indicates a relatively recent invasion.

It is interesting to compare the distributions of

the species of Ecsenius with those of the only other

recently revised large genus of salariinine blennies,
NUMBER 72

._ MARSHALL IS

• .Wot/o

• ' ".Maturo ;. ..v.; .? • -./w»

CALEPONIA _4j

lividinalis

O mandibularis

A oculus

stigmatura

|OO

FIGURE 3.—Distribution of certain species of Ecsenius. Irregular broken line indicates approximate

position of andesite line (see zoogeography section).

l8Oe

Entomacrodus (Springer, 1967). Entomacrodus occurs in waters close to rocky shores (right up to

the shore line). The species are most often taken

in areas where corals are sparsely or not at all

present (although the corals may abound in areas

adjacent to where the species occur). Species of the

two genera are not often collected together, and

where they are, one genus or the other usually predominates. Presumably where they are collected

together, by poisoning, the poison has entered

different habitats. Collectors rarely restrict themselves to a single habitat when collecting, but pick

up everything killed, thus mixing the specimens

so taken.

In general, the species of Entomacrodus are

more abundant in collections and more widely

distributed geographically, occurring circumtropically, than are the species of Ecsenius, but no species

of Entomacrodus has been taken in the Red Sea

or Persian Gulf. I collected only one specimen of

Entomacrodus (E. thalassinus) at One Tree Island,

Great Barrier Reef, whereas I collected hundreds
10

of specimens of the three species of Ecsenius occurring there. I collected seven species of Entromacrodus and four species of Ecsenius in Taiwan, and

none were very abundant.

The overall geographic picture presented by the

two genera has two parallels and several differences.

Indian Ocean-Pacific Ocean species pairs are present

in both genera: Ecsenius nalolo-E. yaeyamaensis,

Entomacrodus epalzeocheilus-E. niuafoouensis, Entomacrodus vermiculatus-E. decussatus. Two species

of Entomacrodus, E. sealei and E. cymatobiotus, do

not occur west of the andesite line and one species,

E. stellifer, does not occur east of it. The widespread species of Entomacrodus are more widespread than those of Ecsenius, and Entomacrodus

is circumtropical. This may be the result of the

pelagic ophioblennius larva characteristic of Entomacrodus. The larvae of Ecsenius have not been

collected in the field and nothing is known of their

behavior. Entomacrodus has eight island endemic

species (22 species in the genus), whereas Ecsenius

has only one or two (18 species in the genus), and

these may be found to be more widespread when

more collections are available. Two species of

Entomacrodus have recognizable South China Sea

subspecies, but none of the species of Ecsenius are

so represented (there are few specimens of Ecsenius

from that area). Springer (1967) noted that several

species of Entomacrodus were peripherally distributed in the tropical western and central Pacific;

only Ecsenius mandibularis is so limited in its

distribution. Entomacrodus occurs in Hawaii where

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ecsenius has been reported only once, under peculiar circumstances.

It would seem that the distributional pattern

differences shown by the two genera are probably

the result of their different ecological preferences

and early life history. But these facts alone are

insufficient to explain, on the surface, at least, why

Entomacrodus does not occur in the Red Sea or

Arabian Sea and Persian Gulf. Its absence from the

latter two areas could be an artifact of collecting,

but seems unlikely in view of the fact that many

collections have been made in those areas and that

Entomacrodus is an easier genus to obtain than

Ecsenius where the two co-occur generally. The

absences of Entomacrodus from the Red Sea and

Ecsenius from Hawaii seem real as there has been

a great collecting effort made in those areas in

recent years.

Because they are more difficult to obtain, I feel

certain that the ranges of the species and the

number of species of Ecsenius reported on here

will increase significantly in the course of a few

years. I believe this, in part, because Ecsenius

species frequently occur at greater depths than

have usually been collected in the past; there has

been relatively little collecting at depths over three

meters along the coasts of the Indo-Pacific. On the

other hand, I do not feel that there are many, if

any, species of the shallow-dwelling genus Entomacrodus yet undiscovered, and the ranges of only

a few of those known will increase to any significant

degree with more collecting.

Key to the Subgenera and Species of Ecsenius

1. Anterior canines in lower jaw obvious, more than twice size of adjacent incisoriform teeth;

total incisoriform teeth -)- anterior canines in lower jaw 13-16; total teeth in upper jaw

26-34; caudal forked (Figure 12); mid-predorsal pore of supratemporal series in advance

of level of posterior orbital margin; pseudobranchial filaments 6-12; dorsal -f ventral

procurrent caudal rays 18-20; epipleural ribs 22-25; postcleithra consisting of two wellseparated ossifications, dorsalmost much reduced (subgenus Anthiiblennius)

tnidas (Red Sea, Indian Ocean, eastern Australia)

Anterior canines in lower jaw rarely obvious, when obvious only slightly larger than adjacent

incisoriform teeth; total incisoriform teeth + anterior canines in lower jaw 30-64; total

teeth in upper jaw 97-148; caudal not forked, but often with produced rays on dorsal and

ventral lobes (for instance, Figures 15, 19, 20); mid-predorsal pore of supratemporal series

well posterior to level of posterior orbital margin; pseudobranchial filaments 5-8 (rarely

8); dorsal -f. ventral procurrent rays 10-18; epipleural ribs 11-17; postcleithra consisting

of two subequal articulating ossifications (subgenus Ecsenius) 2

2. Total posterior canines in lower jaw 7-15 (2-8 on each side)

mandibularis (eastern Australia)

Total posterior canines in lower jaw 0-3 (usually 1 on each side) 3
NUMBER 72

3. Segmented dorsal rays 12-15; segmented anal rays 13-17; vertebrae 29-33 4

Segmented dorsal rays 15-20 (only 2 of 252 specimens of bicolor with 15); segmented anal

rays 17-23 (only 1 of 258 specimens of bicolor with 17); vertebrae 33-39 (2 of 4 specimens

of stigmatura and 9 of 234 specimens of bicolor with 33) 11

4. Dark spot extending well anteriorly from anus (Figure 26); dorsal fin notched three-ninths

to five-ninths length of first dorsal ray; last dorsal spine 6 to 10 percent SL; incisoriform

teeth in lower jaw 30-37; segmented anal rays 13-14; caudal vertebrae 19-20; total vertebrae 29-30 lividinalis (Pacific)

No dark spot extending anteriorly from anus (occasionally black lining of gut protrudes

slightly through anus giving appearance of dark ring); dorsal fin notched seven-ninths

to nine-ninths length of first dorsal ray; last dorsal spine 0.4-3.1 percent SL (one specimen

of 35 of E. oculus had length 4.6 percent); incisoriform teeth in lower jaw 41-64; segmented anal rays 14-17 (1 of 42 specimens of opsifrontalis with 14); caudal vertebrae

21-23; total vertebrae 31-33 5

5. Two large, distinct dark spots at caudal base extending posteriorly well out on caudal fin

(Figure 31) oculus (Pacific)

Dark spots, if present at caudal base, small, scarcely encroaching on caudal fin 6

6. Distinct dark stripe, stripes, or spots present on fleshy pectoral base (Figures 27-30); pseudobranchial filaments 5-8 (rarely 5) 7

Fleshy pectoral base not noticeably marked; pseudobranchial filaments 5-6 8

7. Fleshy pectoral base with at least a forked dark stripe, the arms of which extend posteriorly yaeyamaensis (Pacific)

Fleshy pectoral base with an unforked dark stripe (and usually other dark marks)

nalolo (Indian Ocean; Red Sea)

8. Lateral line extending posteriorly to below level of llth-12th (usually 12th) dorsal spine;

segmented dorsal rays 13-14 (14 in only 6 of 42 specimens); segmented anal rays 14 to

16 (usually 15); nasal cirrus 2.3-5.9 percent SL opsifrontalis (Pacific)

Lateral line extending posteriorly to below level of 8th-9th dorsal spine; segmented dorsal

rays 14; segmented anal rays 15-16 (usually 16); nasal cirrus 1.1-2.3 percent SL 9

9. Two large dark spots on side of body in area covered by appressed pectoral fin (Figure 34)

bimaculatus (Pacific)

Body without large dark spots 10

10. Side of body with alternating dark and pale longitudinal stripes (Figure 33)

prooculis (Pacific)

Side of body without distinct marks (Figure 35) bandanus (Pacific)

11. Dark spot extending anteriorly from anus (Figures 24-25); dark stripe originating at ventral

margin of orbit extending posteriorly across head atigmatura (Pacific)

No dark spot in region of anus; dark stripe extending posteriorly across head, if present,

originating at mid-postorbital level 12

12. Anterior nostril with cirri on both anterior and posterior margins; segmented caudal rays

14 (rarely 13); lateral line usually with several vertical pairs of pores beginning anteriorly IS

Anterior nostril with cirrus on posterior margin only; segmented caudal rays 13 (rarely 14

in E. frontalis and in some juveniles of other species under 25 mm SL); lateral line of

simple pores only (rarely with a vertical pair at origin) 14

13. Dorsal rays 15-18; dorsal fin notched two-ninths to seven-ninths length of first dorsal ray

(usually four-ninths or more); last dorsal spine 2.5-6.9 percent SL; lateral line with vertical pairs of pores for one-third to all its length (rarely less than one-half); a dark spot

on paler background on anterior dorsal fin; pseudobranchial filaments usually 6; upper

nasal cirrus length usually more than 4.9 percent SL. bicolor (Pacific and Indian Oceans)

Dorsal rays 18-20 (rarely 18); dorsal fin unnotched; last dorsal spine 11.0-15.9 percent SL;

lateral line with vertical pairs of pores, if present, rarely occupying more than anterior

two-fifths its length; dorsal fin entirely black, occasionally with some obscure longitudinal

stripes anteriorly; pseudobranchial filaments usually 7; upper nasal cirrus length not

more than 4.8 percent SL namiyei (Pacific)

14. Dorsal fin without a notch; last dorsal spine 10.7-20.8 percent SL; pectoral rays 14-16 (14

in 5 or 89 specimens) frontalit (Red Sea;

Gulf of Tadjourah; see discussion under species account for distinct color pattern forms.)
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Dorsal fin with a notch; last dorsal spine 2.2-7.0 percent SL; pectoral rays 13-15 (15 in 5

of 87 specimens) ^

15. Dorsal spines 12; pectoral rays 13 (14 in 1 of 12 specimens); body at about midlevel with a

longitudinal dark stripe, either continuous or as a series of dark bars, extending length

of body and entering caudal fin basally lineatus (Pacific and Indian Oceans)

Dorsal spines 12-14 (12 in 9 of 201 specimens); pectoral rays 13-15 (13 in 2 of 176 specimens) ; body without a broad dark stripe (occasionally with a slender stripe on dorsal

fourth of body, ceasing well anterior to caudal base) 16

16. Mid-distal half of at least spinous dorsal fin black or with black spots in interradial membrane; dorsal and ventral lobes of caudal fin darker than remainder; usually several small

dark spots on posterior third of body; usually a narrow, black longitudinal stripe on

dorsoanterior portion of body gravieri (Red Sea; Bay of Tadjourah)

Mid-distal half of dorsal fin not noticeably marked; dorsal and ventral lobes of caudal not

darker than remainder of caudal; no black spots or longitudinal stripes on body (body

uniformly dark, with a dark spot at caudal base, or with posterior half pale with irregular,

vertical dark bars) 17

17. Dorsal rays 17 or 18; anal rays 19 or 20; nasal cirrus 1.0-3.9 percent SL; dorsal fin notched

seven-ninths to nine-ninths (usually eight-ninths) length of first dorsal ray, body uniformly dark with a darker spot at caudal base E. aroni (Red Sea)

Dorsal rays 18-20; anal rays 19-23 (rarely 19 or 20); nasal cirrus 3.0-7.9 percent SL; dorsal

fin notched five-ninths to eight-ninths (rarely eight-ninths) length of first dorsal ray;

body uniformly dark or posterior half pale with vertical dark bars

E. pulcher (Persian Gulf and northern Arabian Sea)

The Species Groups of Ecsenius

The 18 species of Ecsenius fall into eight groups.

Four of these groups are monotypic, E. midas, E.

frontalis, E. lineatus, and E. mandibularis, and with

the exception of E. lineatus, I hesitate to relate

them closely to each other or to any other group.

E. lineatus may be an offshoot of the yaeyamaensis

group.

Ecsenius midas, as the monotypic member of the

subgenus Anthiiblennius, is well differentiated (see

key, couplet 1). It is also the most specialized of

the species of the genus as evidenced by its extreme

reduction in number of teeth, the increased size of

the anterior canines, the reduced nature of the

postcleithra, and the great anterior displacement

of its dorsal fin origin. E. frontalis has an unnotched

dorsal fin and the highest pectoral ray counts in

the genus. The albicaudatus form of E. frontalis

(see discussion of color pattern under species account) is reminiscent of that found in E. bicolor

and E. namiyei. E. frontalis is separated from the

latter two species, however, in having only a single

nasal cirrus, 13 segmented caudal rays, and no

paired lateral line pores. E. lineatus has no noteworthy peculiarities as found in many of the other

species. Its general robustness, relatively high fin

ray and vertebral counts, large size attained, and

moderately notched dorsal fin separate it from the

yaeyamaensis species group, to which it otherwise

might belong. E. mandibularis is unique in its large

numbers of posterior canine teeth and long third

dorsal spine and fifth dorsal ray lengths. In addition, males of E. mandibularis develop prolongations of all the caudal rays, which in the other

species are primarily restricted to the upper and

lower extremities of the fin.

The pulcher species group comprises E. pulcher,

E. gravieri, and E. aroni. These three species are

morphologically quite similar and differ from each

other primarily in color pattern, depth of the

dorsal fin notch, development of caudal fin lobes,

and the slightly higher meristics of E. pulcher.

They share the pecularity of having typically 14

pectoral rays (otherwise found typically only in

E. stigmatura), and with E. midas and E. frontalis

are the only species with typically 11 precaudal

vertebrae.

The bicolor species group comprises E. bicolor

and E. namiyei, which are distinct from all other

Ecsenius species in having nasal cirri on both the

anterior and posterior margins of the anterior

nostril and paired pores in the lateral line. They

both have typically 14 segmented caudal rays.

The stigmatura species group comprises E. stigmatura and E. lividinalis. The Bornean specimen

allocated to E. stigmatura may be recognizable as
NUMBER 72

13

a separate species when more specimens are known.

The two species are united here in having a black

spot in the anal region (found otherwise only in

E. midas) and typically 14 segmented caudal rays,

but are considerably different in other characters.

E. lividinalis has the lowest tooth, dorsal and anal

fin and vertebral counts, and the highest fin ray

proportions (as percent SL) in the subgenus

Ecsenius. The relationship of E. lividinalis with

E. pulcher is questionable and possibly each should

be placed in its own group.

The yaeyamaensis species group comprises E.

yaeyamaensis, E. nalolo, E. oculus, E. opsifrontalis,

E. prooculis, E. bimaculatus, and E. bandanus. This

group comprises the more or less nondescript, small

species with relatively deep dorsal fin notches and,

for the most part, low vertebral and vertical fin

ray counts. Within this group, there are three subgroups comprising the first two, second two, and

last three species listed. The subgroups are distinguished from each other on color pattern and

slight meristic or proportional differences. Each

group presents certain problems.

The two species of the yaeyamaensis subgroup

might, by some systematists, be considered to comprise a single species. The two species I recognize

are completely differentiated by a color marking

(key couplet 7) and occur in different oceans (Figure

2). In the Pacific Ocean there are also easily recognizable color pattern populations of one species. If

the two species were combined the problem of what

to do with these recognizable populations would

remain, as the situation would then exist of having

two recognizable taxa differing in color pattern

with at least one of these taxa also having populations completely recognizable on the basis of color

pattern (see color pattern description under E.

yaeyamaensis species account).

The two species of the oculus subgroup are distinguished from each other on the basis of color

pattern and the slightly lower meristics of one

species. The two species occupy essentially allopatric

ranges (Figures 2 and 3). There are indications that

some recognizable populations of E. oculus may

occur (see under species account).

The three species of the prooculis subgroup are

known from 5, 3, and 1 specimens, all males. These

species are distinguished from each other only by

their strikingly different color patterns. They have

not been collected from the same locality (Figure 4).

Whether the three species comprise one highly

variable species will require much more material

than is available.

Species Accounts

Ecsenius (Anthiihlennius) midas Starck

FlCURE 12

Ecsenius (Anthiiblennius) midas Starck, 1969, p. 1 [D'Arros

Island, Amirante Islands].

DESCRIPTION.—Dorsal spines 13-14 (usually 13);

dorsal rays 19-21; segmented anal rays 20-23; pectoral rays 13 (12 on one side only of one specimen);

segmented caudal rays 13-15 (usually 14); dorsal

procurrent caudal rays 9-10; ventral procurrent

caudal rays 9-10; total caudal elements 31-34; gillrakers 10-15; pseudobranchial filaments 6-12; lower

incisor teeth 13 to 16; lower jaw posterior canines 1;

total lower jaw posterior canines 2; upper incisor

teeth 26-34; precaudal vertebrae 11; caudal vertebrae 27-29; total vertebrae 38-40; epipleural ribs

22-25. Lateral line with no paired pores, extending

posteriorly to beneath level of 7th-9th dorsal spine.

Dorsal fin without notch. Third (innermost) pelvic

ray obvious. One cirrus on each anterior nostril.

Color pattern: The Red Sea and Indian Ocean

specimens of this species are generally dusky, darkest anteriorly, and paling on the caudal peduncle

of some specimens. The lower side of the head is

darker anteriorly and on some specimens a dusky

stripe occurs on each side proceeding dorsoposteriorly from the side of the lower jaw and curving

upward along the membranous margin of the

branch iostegal membrane. A few specimens exhibit

a diffusely dusky stripe beginning just behind the

ventroposterior border of the eye and extending

back across the opercle. One specimen had some

diffuse vertical bands on the body. The most conspicuous mark is a dark spot enveloping the anus

and extending anteriorly for a short distance. The

dorsal fin bears a narrow, dark distal stripe beginning anteriorly on the spinous portion and extending to the anterior dorsal rays. This fin is usually

dark anteriorly, making it difficult or impossible

to count the spines externally. The anal fin is pale

dusky, slightly darker on the spines and, in some

specimens, along its distal edge. The caudal fin

varies from immaculate to evenly dusky with its

dorsal and ventral margins narrowly, darkly pig-
14

\ZO°

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

»4O° 160*

bandanus

A bimaculatus

# prooculjs

zo

120 140* 160*

FIGURE 4.—Distribution of certain species of Ecsenius. Irregular broken line indicates approximate

position of andesite line (see zoogeography section).

mented. The pectorals are generally pale with the

rays sometimes narrowly and darkly pigmented.

The pelvics are dusky. The two small Great Barrier

Reef specimens differed from the others most noticeably in having the dorsal fin dusky, but with

the distal portion of the segmented ray part of the

fin unmarked and the edge of the dusky area much

darker than the remainder in the region below the

unmarked part.

Starck (1969) reported the life color to be golden

yellow, darker dorsally except for a fine black

marginal line on the dorsal fin and a black blotch

around the anus. The color of a freshly killed

female (USNM 204550) I collected in the Red Sea

was somewhat different. The head was pale orangedusky with a narrow, pale yellow-green stripe

extending posteriorly from the orbit to the opercle

margin. The underside of the head was pale white

except for the dark dusky lower jaw. The sides of

the body were pale purple-dusky overlying orange.

The venter was chartreuse. The dorsal, anal, and

caudal fins were orange-yellow. The pectoral fins

were pale yellow-green.

GEOGRAPHIC VARIATION.—There are too few specimens of E. mi das to draw firm conclusions, but it

appears from the fact that the four Red Sea specimens have 27 caudal vertebrae and the six Indian

Ocean specimens have 28 or 29 caudal vertebrae,

that population differences occur. The two Great

Barrier Reef specimens have 27 and 28 caudal

vertebrae and are thus intermediate between the

specimens from the other two areas.

MATERIAL.—Gulf of Aqaba, Sinai coast, Dahab,

UTAI 4221 (71.2) and USNM 204707 (77.1); Ras

Muhammad, southern tip of Sinai Peninsula,

USNM 204550 (45.1) and 204551 (68.9); Amirantes

Islands, D'Arros Island, ANSP 111148 (74.4, holotype of E. midas), 111149 (48.7), 111854 (33.5,

cleared and stained), and USNM 202422 (56.0);

Nicobar Islands, NFIS 9579 (83.1); Nicobar Islands,

Tillanchong, NFIS 9577 (44.4); Australia, Queensland, Escape Reef, ANSP 109686 (2:28.5-29.0).

Ecsenius (Ecsenius) frontalis Ehrenberg

FIGURES 13-15

Salarias frontalis Ehrenberg in Cuvier and Valenciennes, 1836,

p. 328 [Massawa, Red Sea].
NUMBER 72 15

Salarias nigrovittatus Ruppell, 1838, p. 136 [Massawa, Red

Sea].

Ecsenius albicaudatus Lotan, 1969, p. 372 [Marsa Murach,

Gulf of Aqaba].

DESCRIPTION.—Dorsal spines 10-13 (rarely 10, 11,

or 13); dorsal rays 17-21 (rarely 17 or 21); segmented anal rays 17-22 (rarely 17, 18, or 22); pectoral rays 14-16 (usually 15); segmented caudal

rays 13 (rarely 14); dorsal procurrent caudal rays

6-8; ventral procurrent caudal rays 6-9 (rarely 9);

total caudal elements 25-30 (rarely 25 or 30); gillrakers 11-17 (rarely 11, 12, or 17); pseudobranchial

filaments 5-7 (usually 6); lower incisor teeth 40-53;

lower jaw posterior canines 0-1; total lower jaw

posterior canines 0-2 (usually 2); upper incisor

teeth 124-130 (4 counts); precaudal vertebrae 10-12

(rarely 10 or 12); caudal vertebrae 23-26 (rarely

23); total vertebrae 34-37 (rarely 34); epipleural

ribs 11-16 (usually 12-14). Lateral line without

paired pores, extending to below level of 8th-12th

dorsal element (usually 9th-llth). Dorsal fin without

notch (one of several hundred specimens had dorsal

fin notched two-ninths length of first dorsal ray).

Third (innermost) pelvic ray varying from obvious

to not obvious (usually not obvious). One cirrus

on each anterior nostril, each cirrus occasionally

forked.

Color pattern: There are three basic color patterns found in E. frontalis. See discussion following,

concerning the systematic status of these patterns.

The smallest specimen examined was a postlarva,

15.0 mm SL, from the Gulf of Aqaba. It was not

possible to assign this specimen to one of the three

color pattern types. The specimen was pale with

several irregular vertical bands composed of spots.

A few small bright yellow individuals of E. frontalis

were also seen in the Gulf of Aqaba. It may be that

the banded and yellow forms represent the preadult

color of the two color pattern forms found in the

Gulf of Aqaba (more below).

The Problem of Ecsenius frontalis, E. albicaudatus, and E. nigrovittatus

Although I have placed E. albicaudatus and E.

nigrovittatus in the synonymy of E. frontalis, I am

uncertain that my action is correct. In the discussion

that follows I treat the three nominal species as

color pattern forms of a single species, E. frontalis,

and present the evidence on which my action is

based.

Ecsenius frontalis and E. nigrovittatus were described from Massawa, Ethiopia, and E. albicaudatus

was described from the Gulf of Aqaba, each from

a single specimen. While the types of E. frontalis

and E. nigrovittatus are in poor condition, data

from these types, together with the distinctive color

patterns given in the original descriptions, make

it possible to identify recently collected specimens

from Massawa with both these nominal species.

The holotype of E. albicaudatus is in excellent

condition. Its distinctive color pattern has been

found in numerous recently collected specimens

from near the type locality.

The morphology and meristics of the three nominal species are very similar, but the species differ

consistently in color pattern. (In Figures 13-15 the

apparent morphological differences exhibited in

these illustrations are individual variations; when

series of each color form are examined complete

morphological overlap is encountered.) I have observed all three color pattern forms alive in their

natural habitat. The frontalis form is generally

brown, darkest anteriorly, grading into yelloworange posteriorly, with a bright yellow-orange

caudal fin. The head and body of the albicaudatus

form are black with a narrow area of variable width

at the caudal peduncle completely unmarked and

pale (sometimes the pale area extends onto the

dorsal fin); the caudal fin is immaculate except

for a faint pink tinge observable only in freshly

preserved specimens. The nigrovittatus form is usually brown with a black stripe on the body bordered

by pale dorsal and ventral margins; the caudal fin

is slightly tinged with orange. Occasional individuals of the nigrovittatus form have yellow-orange

heads.

Two individuals, I believed at the time of capture

to be the frontalis form, were brought to the laboratory alive. When the individuals were placed in

an aquarium there appeared one frontalis form and

one albicaudatus form, which was dark brown with

a yellow-orange caudal—the only albicaudatus form

I have collected with these colors. These and other

individuals of both color pattern forms held in the

aquarium paled considerably after several days,

and it was not possible to tell one form from the

other.

383-026 O - 71 - 2
16

The geographical distributions of the albicaudatus and frontalis forms are nearly identical

(Figure 5). The frontalis form has been taken

widely in the Gulf of Aqaba, in the southern Gulf

of Suez, widely in the Red Sea and in the Bay of

Tadjoura (Gulf of Aden, just outside the southern

entrance to the Red Sea). The albicaudatus form

has a similar distribution to that of the frontalis

form, but it has not been taken outside of the

Red Sea or in the Gulf of Suez. It was observed by

me in the Gulf of Suez, however. The nigrovittatus

form has been taken only in the southern Red Sea,

on and off the coast of Ethiopia.

FIGURE 5.—Distribution of the three color pattern forms of

Ecsenius frontalis: •% — frontalis form, -ft = albicaudatus

form, % — nigrovittatus form.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

I obtained all three forms in the same collection

(poison) from a coral rock outcrop estimated to be

about 3 meters in height and 8-10 meters in circumference; water depth was about 4 meters. The

specimens in this collection included 55 frontalis

forms, 25 nigrovittatus forms, and 6 albicaudatus

forms. My six collections (all made at depths of

7 meters or less) in Ethiopian waters, where the

frontalis and nigrovittatus forms were always taken

together, yielded three times as many frontalis

forms as nigrovittatus forms, and the frontalis forms

were always more abundant. The six albicaudatus

forms mentioned are the only ones known from

Ethiopian waters.

In the Gulf of Aqaba and Gulf of Suez collections, where only the frontalis and albicaudatus

forms were seen or taken, the frontalis form was

generally dominant in the shallower waters and

the albicaudatus form was generally dominant in

the deeper waters. The depth of each collection and

the number of specimens of each form taken in the

poison collections that I made in Israeli waters are

listed below. Because the poison spread, it was not

possible to limit collections to a particular depth.

The collections were all made during the day,

July-September 1969.

Frontalis

form

37

3

9

2

0

1

31

8

1

0

9 (seen commonly)

I observed both the frontalis and albicaudatus

forms together and common at depths of about 3-12

meters on the coral reef preserve at Eilat, Israel.

I cannot say whether one form was much more

dominant than the other; my impressions varied

from day to day.

The frontalis and albicaudatus forms were commonly observed together in the Gulf of Aqaba,

and the frontalis and nigrovittatus forms were

Approximate

depth

(meters)

0-3

0-18

0-15

0-16

0-7.5

0-105

0-10

0-3.5

9-12

21-27

0-9

Albicaudatus

form

10

11

5

14

3

4

1

1

18

3

one seen
NUMBER 72

17

commonly observed together at Massawa, Ethiopia.

I have not seen all three forms together, although

as noted above, I have collected all three from the

same rock. The three forms inhabit live and dead

coral and are not found over the sandy bottom.

They are frequently out from under cover, on top

of the coral. They dart, but only rarely swim

freely, and then for only a few centimeters. Their

movements are only moderately jerky. They feed

by nipping at the rocky substrate. Gut contents

consist of unidentifiable calcareous granules less

than 0.3 mm in diameter, paniculate organic

matter, ostracods, and foraminifera. I noted no

differences in behavior of the three forms, which

appeared to be identical and different from the

behavior of the other species of Ecsenius that occur

together with them in the same general area. For

instance, E. gravieri spends much of its time swimming freely over sandy bottom, where it feeds by

nipping into the sand; E. nalolo was never seen out

from under cover or swimming freely, and its

22

20 -

movements are very jerky; E. aroni is somewhat

similar in behavior (but not morphology or color

pattern) to E. frontalis, but it usually occurs at

greater depths and spends more time under cover;

E. midas, which was not seen alive, was reported

by Starck (1969) to swim with other fishes and to

feed on plankton.

Certain morphological and meristic characters

are given with frequency distributions in Tables

11 and 12 for each of the three forms for the

different localities where they occur. Examination

of the data indicates the possibility of sexual

dimorphism and also that Israeli specimens of the

color forms have higher numbers of certain meristic

characters than Ethiopian specimens of the same

forms. To determine if the differences noted are

statistically significant it is, therefore, necessary to

test the data for the same forms, sexes separated,

from the two areas, and for different forms from

the same area, sexes separated.

15 -

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Israeli frontalis form

Israeli albicaudatus form

Ethiopian frontalis form

Ethiopian albicaudatus form

Ethiopian nigrovittatus form

D

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i

female male

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1

2 2 30 40

Standard Length mm

50 6 0

FIGURE 6.—Graph showing relationship of last dorsal spine length to standard length in the three

color pattern forms of Ecsenius frontalis.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Data for specimens of the frontalis form from

Sanganib Reef (Sudan) and the Gulf of Aden and

for specimens of the albicaudatus form from Ethiopian waters were not included in the tests because

the numbers of specimens were small and the

localities were well removed from Israel or Ethiopia.

The following significant differences were found:

Israeli frontalis form males and females have

higher average numbers of teeth, segmented dorsal

rays, segmented anal rays, and caudal vertebrae

than Ethiopian frontalis form males and females.

Ethiopian frontalis form males have higher average numbers of segmented anal rays than Ethiopian

frontalis form females.

Ethiopian frontalis form males have a close to

significantly (p = between .02 and .01) higher average number of caudal vertebrae than Ethiopian

frontalis form females.

Grouped data (Tables 13 and 14) of proportional

characters also indicate differences among the three

forms. These data were also graphed and the following differences noted:

Israeli frontalis form males have a relatively

longer caudal fin than Israeli frontalis form females

(Figure 10).

Israeli albicaudatus form males have a relatively

longer caudal fin than Israeli albicaudatus form

females (Figure 10).

Israeli frontalis form males have a relatively

longer caudal fin than Ethiopian frontalis form

males (Figure 10).

Ethiopian nigrovittatus form males have a relatively longer caudal fin than Ethiopian nigrovittatus

form females (Figure 9).

Ethiopian nigrovittatus form males have a relatively longer caudal fin than Ethiopian frontalis

form males or females (Figure 9).

Specimens of less than 30 mm SL of the Israeli

frontalis and albicaudatus forms have relatively

longer nasal cirri than specimens of any of the

three Ethiopian forms (Figures 7 and 8).

Specimens of the Israeli frontalis and albicaudatus forms have a relatively longer last dorsal

spine than any of the three Ethiopian forms

5

10

5

-

-

o

-

•

•

D

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o

•

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Ethiopian

Ethiopian

Ethiopian

0

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frontalis form

albicaudatus form

nigrovittatus form

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20 30 40

Standard Length mm

50 60

FIGURE 7.—Graph showing relationship of nasal cirrus length to standard length in the threecolor pattern forms of Ecsenius frontalis from Ethiopian coastal waters.
NUMBER 72 19

(Figure 6). The same relationship exists between

the relative lengths of the third dorsal spine and

fifth dorsal ray of the Israeli frontalis and albicaudatus forms compared with the lengths of these

elements for the three Ethiopian forms (study

graphs).

There is some indication that Israeli frontalis

form males have a relatively longer caudal fin than

Israeli albicaudatus males (Figure 10).

Aside from the fact that there are differences

between the Israeli and Ethiopian populations of

the frontalis form, the above data provide only

limited support for the concept that the three color

pattern forms are different species: The nigrovittatus form exhibits sexual dimorphism in the

length of the caudal fin, which the sympatric

frontalis form does not (but the allopatric Israeli

frontalis form does); nigrovittatus form males have

a longer caudal fin than sympatric males of the

Ethiopian frontalis form (but not longer than the

allopatric Israeli frontalis and albicaudatus form

males), and possibly the Israeli frontalis form males

have longer caudal fins than Israeli albicaudatus

form males. (This does not seem to be true of

Ethiopian frontalis form males versus Ethiopian

albicaudatus form males even though there are

only a few specimens of the latter.)

The fact that the Gulf of Aqaba populations of

the frontalis and albicaudatus color pattern forms

have higher average numbers of each meristic character than their Ethiopian counterparts is probably

due to the cooler water temperatures of the Gulf

of Aqaba relative to those of the Ethiopian coast

(Oren, 1962). The same type variation has been

reported for other Gulf of Aqaba-Red Sea blenniids

(Smith-Vaniz and Springer, in press).

There is, then, no consistent basis other than

color pattern on which to recognize E. frontalis and

E. albicaudatus as separate species. One might

recognize E. nigrovittatus as a separate species,

based on the small differences noted and the fact

that it does not occur at the northern end of the

Red Sea (Gulf of Aqaba).

There are other data that influence the recognition of the three forms as one species. There are

no other species of Ecsenius that appear to be

closely related to the three forms and speciation,

if such occurred, would seem of necessity to have

taken place in the Red Sea. The Red Sea is a

relatively small body of water with a limited amount

of coral reef habitat. It is difficult to postulate how

three species so similar as the three color pattern

forms could have evolved sympatrically, especially

when there does not appear to be a difference in

feeding habits or microhabitat preference.

There are other species of Ecsenius that have

apparently evolved different color pattern forms

that occur sympatrically. These forms are, however,

more obviously derivative of the same basic form

than are the Red Sea color pattern forms of E.

frontalis. Ecsenius pulcher has a solid color form

and one in which the melanophores have been

suppressed in portions of the posterior region of

the body, thus forming vertical dark and pale

stripes (Figure 16). This form is somewhat similar

to the albicaudatus form in which the melanophores

on the caudal peduncle have been suppressed.

Ecsenius bicolor has three basic color pattern forms:

one that is completely dark, one that is paler

(either gradually or abruptly, Figures 20 and 21),

and one that has a diffuse, dark stripe on its side.

Although these three forms duplicate the forms

found in Ecsenius frontalis, they generally are not

so strikingly different from each other, and the

striped form is very rare. The life colors of the

three E. bicolor forms are very much the same as

opposed to the differences noted in the frontalis

forms. It seems, therefore, that the tendency for

species of Ecsenius to exhibit different color pattern

forms is a trait of the genus.

Information on breeding behavior and the color

patterns of the offspring are necessary for reaching

a final solution to the problem of the taxonomic

level that should be accorded the three color pattern

forms of E. frontalis.

I know of no situation involving fishes that is

exactly comparable with that reported here for E.

frontalis. Although only slightly similar, the circumstances Stephens (1963) reported for Acanthemblemaria crockeri Beebe and Tee-Van (Chaenopsidae) is of interest. A. crockeri is represented by

two color pattern forms in the Gulf of California,

the only area where the species occurs. Only one

form was found in the northern end of the Gulf,

and its relative abundance gradually decreased

toward the south until it was not found to occur

in the southern end of the Gulf. The other form

was the only one at the southern end of the Gulf

and it decreased in relative abundance toward the

northern end of the Gulf. The two forms occurred
20

together in the midportion of the Gulf. Stephens

was unable to explain the basis for these circumstances. The situation is somewhat paralleled by

two of the E. frontalis color forms. The albicaudatus

form is abundant at the northern end of the Red

Sea (Gulf of Aqaba) and rare at the southern end.

The nigrovittatus form is common at the southern

end of the Red Sea but absent from the northern

end. I hesitate to place much emphasis on these

similarities.

MATERIAL (localities abbreviated).—Gulf of

Aqaba: USNM 204535 (3 albicaudatus, 25.4-30.5),

USNM 204536 (3 albicaudatus, 19.8-24.2; 1 post

larva, 15.5), USNM 204685 (1 albicaudatus, 51.6;

8 frontalis, 29.2-44.8), USNM 204684 (9 albicaudatus, 18.0-44.4; 37 frontalis, 16.8-43.7), USNM

204534 (18 albicaudatus, 16.6-36.1; 1 frontalis, 36.7),

USNM 204650 (11 albicaudatus, 18.0-37.7; 3 frontalis, 18.7-48.4), USNM 204533 (14 albicaudatus,

19.5-37.5; 2 frontalis, 19.5-21.7), HUI HUJ-F-3540

(holotype of E. albicaudatus, 43.0), HUI E-60/96.16

(albicaudatus, 46.9), HUI HUJ-F-4375 (2 albicaudatus, 19.1-21.4), HUI HUJ-F-3544 (2 frontalis,

40.2-42.1), HUI E-64/34 (4 frontalis, 32.0-41.0),

HUI E-64/3013 (frontalis, 40.0), UTAI NS-3582

(albicaudatus, 43.8); Gulf of Suez, Et Tor: USNM

204651 (9 frontalis, 29.5-44.7); Egypt, Strait of

Jubal: USNM 200613 (1 albicaudatus, 50.5; 6 frontalis, including one cleared and stained, 23.9-58.6);

Egypt, Al Ghardaqa: NFIS 5118 (frontalis, 43.9);

Sudan, Sanganeb Reef: NFIS 8236 (3 frontalis,

40.6-47.4); Ethiopia, Massawa: ISZZ 1947 (5 damaged syntypes of Salarias frontalis), NFIS 1680

(holotype of Salarias nigrovittatus, 29.0), USNM

204517 (4 nigrovittatus, 24.2-36.0; 8 frontalis, 21.2-

34.6); Ethiopia, Harat Island: USNM 204490 (6

albicaudatus, 29.2-46.5; 25 nigrovittatus, 20.4-48.3;

55 frontalis, 21.9-61.4); Ethiopia, Dahlak Islands:

HUI E-62/424A (21 frontalis, 16.3-47.3), HUI

E-62/1280 (2 nigrovittatus, 32.5-34.2), HUI E-62/

428 (2 nigrovittatus, 18.9-40.2), HUI E62/604

(nigrovittatus, 39.2), HUI E-62/3678A (nigrovittatus, 39.5), HUI E-62/1280A (8 frontalis, 30.8-

48.3), HUI E-62/417H (18 nigrovittatus, 16.7-38.5),

NFIS 8757 (7 nigrovittatus, 23.8-44.0); Ethiopia,

Zubair Island: HUI HUJ-F-4628 (4 frontalis 29.4-

35.0); Gulf of Aden, Bay of Djibouti, French Somaliland: MNHN 04-318 (3 frontalis, 37.8-47.3).

Numerous other uncataloged specimens from the

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Gulf of Aqaba and Ethiopia in the USNM collections were examined in order to obtain data on

frequencies of occurrence of the color pattern forms.

In Ethiopian waters, the following localities for

this material are as follows: Isola Delemme (35°54'E,

15°30.5'N), Sciumma Island (40° 00 W E; 15°32'

31"N); Melita Bay (39°49'E; 15°15'N); Difnein

Island (39°20'E, 16°36'N). The frontalis and nigrovittatus forms were present in all these collections.

Ecsenius (Ecsenius) pulcher (Murray)

FIGURE 16

Salarias pulcher Murray, 1887, p. 23 [Manora Rocks, Karachi].

Salarias phntasticus Boulenger, 1897, p. 422 [Mekran Coast,

Persia].

Salarias anomalus Regan, 1905, p. 327 [Mekran Coast, Persian

Gulf].

DESCRIPTION.—Dorsal spines 12-14 (usually 13);

dorsal rays 18-20; segmented anal rays 19-23 (usually 21-22); pectoral rays 14-15 (usually 14); segmented caudal rays 13 (rarely 14 in specimens over

25 mm SL; when 14, lowermost ray has only one

segmental line); dorsal procurrent caudal rays 7-8;

ventral procurrent caudal rays 5-8 (usually 7-8);

total caudal elements 26-29 (rarely 26); gill-rakers

13-17 (usually 14-15); pseudobranchial filaments

6-7; lower incisor teeth 44-53; lower jaw posterior

canines 0-1 (rarely 0); total lower jaw posterior

canines 0-2 (usually 2); upper incisor teeth 107-116

(three counts); precaudal vertebrae 11, caudal vertebrae 26-28; total vertebrae 37-39; epipleural ribs

13-14. Lateral line with no paired pores, extending

to below level of 8th to 12th dorsal spine (usually

9th or 10th). Dorsal fin notched five-ninths to eightninths (rarely eight-ninths) length first dorsal ray.

Third (innermost) pelvic ray varying from not

obvious to obvious (usually not obvious). One cirrus

on each anterior nostril, rarely forked.

Color pattern: There are two basic color patterns

found in adult and young adult specimens. These

two patterns are so strikingly different as to have

been the basis for species descriptions, and the

uninformed worker sorts collections containing both

types into different lots. Chapman and Schultz (1952)

believed that the two color patterns were sex linked,

but I find that both sexes exhibit both types of

color pattern.
NUMBER 72 21

The most common color pattern, based on museum specimens, is that in which the head and

body are uniformly dark or dusky brown. Occasional

specimens of this form are paler posteriorly on the

body. A marking slightly darker than the ground

color frequently occurs behind the eye. The underside of the head may also be slightly darker

anteriorly.

In the other color pattern (Figure 8) the head

and anterior half of the body are similar to that

of the form described above, but the posterior half

of the body bears separated pigmentless areas setting

off dark vertical bars of different sizes and intensities. The number of dark bars ranges from 5 to 8

on each side of the body. The smallest specimen

exhibiting this pattern was 32.9 mm SL.

Below about 20 mm SL, specimens may be uniformly dark or bear 7 or 8 pairs of diffuse vertical

bands on each side and irregular dark marks on

the head. Both uniform and banded small specimens occurred in the same collections.

In both adult type color patterns the spinous

dorsal bears a dusky stripe basally and an immaculate area distally. The dusky stripe extends onto

the soft dorsal where it becomes separated into a

darker area suprabasally and a paler area basally.

The distal portion of the rayed dorsal is mostly

unmarked. The anal fin is uniformly pale dusky.

The other fins are variably marked with immaculate

and pale dusky areas.

A color photograph of a living specimen of E.

pulcher, barred form, appears in Axelrod and

Emmens (1969).

NOMENCLATURE.—Salarias phntasticus was not

compared with any other species. I have examined

the two syntypes and find them to be the striped

form of E. pulcher. (An obvious typographical

error is involved in the specific epithet and the

name is usually spelled phantasticus.)

Salarias anomalus was not compared with any

other species. I have examined the syntypes and

find them to be the uniformly dark form of E.

pulcher.

MATERIAL (localities abbreviated).—Persian Gulf:

USNM 201572 (36.5, cleared and stained), 201571

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Ethiopian

Israeli

Israeli

D

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frontalis

frontalis

form

form

albicaudatus form

0

o

o

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o

1

female

•

•

*

o

o

o

o

0

 0

•

male

o

D

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20 30 40

Standard Length mm

50 60

FIGURE 8.—Graph showing relationship of nasal cirrus length to standard length in the two

color pattern forms of Ecsenius frontalis from Israeli coastal waters and one of these forms from

Ethiopian coastal waters.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(32.9), 196505 (2: 42.2-52.1), 196506 (52.3), BMNH

1900.5.9.47-56 (16: 31.4-46.3, syntypes of Salaries

anomalus), 1898.6.29.163 (57.4), 1932.2.18.43 (39.7);

Gulf of Oman: BMNH 1912.11.26.1 (51.2), 1899.5.

8.94 (2: 28.7-34.0); Mekran Coast, Persia: BMNH

1897.9.22.20-21 (2: 54.5-57.9, syntypes of Salarias

phntasticus); Karachi, Pakistan: BMNH 1887.9.22.

59-60 (2: 44.3-44.5, syntypes of Salarias pulcher),

USNM 201867 (3: 38.9-58.0), 201816 (48.3), NFIS

8239 (37.3); Rupan Coast, India; FMRI uncataloged (4: 18.4-30.6, in two lots); Gulf of Kutch,

India: USNM 201863 (16.6); India: NFIS 5120 (9:

21.7-54.6).

Ecsenius (Ecsenius) gravieri (Pellegrin)

FIGURES 17, 18

Salarias gravieri Pellegrin, 1906, p. 5 [Bay of Tadjourah].

Ecsenius klausewitzi Lotan, 1969, p. 371 [Entedebir, Dahlak

Archipelago].

DESCRIPTION.—Dorsal spines 12-14 (usually 13);

dorsal rays 16-19 (rarely 16 or 19); segmented anal

rays 18-21 (rarely 18 or 21); pectoral rays 13-15

(rarely 13); segmented caudal rays 13; dorsal procurrent caudal rays 7-9 (usually 8); ventral procurrent caudal rays 7-9 (usually 8); total caudal

elements 27-31; gill rakers 12-16; pseudobranchial

filaments 6-8 (rarely 8); lower incisor teeth 44-54;

3 8

3 5

3 0

25

20 -

female

Ethiopian frontoli* form

Ethiopian albicaudatus form

Ethiopian nigrovittotus form

male

O

a

* •

a

° • o ^ o

°

o •

D

20 30 4 0

Standard Length mm

50 60

FIGURE 9.-Graph showing relationship of longest caudal ray length to standard length in the three

color pattern forms of Ecsenius frontalis from Ethiopian coastal waters.
NUMBER 72 23

lower jaw posterior canines 0-1; total lower jaw

posterior canines 0-2 (usually 2); precaudal vertebrae 10-11 (rarely 10); caudal vertebrae 24-26

(rarely 24); total vertebrae 35-37 (rarely 35); epipleural ribs 12-15. Lateral line with no paired

pores, extending posteriorly to below level of 8th

to 10th (rarely 8th) dorsal spine. Dorsal fin notched

two-ninths to six-ninths length of first dorsal ray.

Third (innermost) pelvic ray not obvious in specimens over 25 mm SL. One cirrus on each anterior

nostril.

Color pattern: This is a distinctively marked

species. In Gulf of Aqaba, Gulf of Suez, and northern Red Sea specimens, 17.5-60.8 mm SL, the head

is relatively dark with the intensity of the pigment

decreasing gradually on the body and becoming

quite pale posteriorly. At about the midposterior

margin of the orbit of most specimens there is a

dark stripe, or diffuse evidence of one, that extends

posteriorly across the head, darkens greatly on the

body, and usually extends to below at least the

middle of the spinous dorsal fin, but terminates

anterior to the caudal peduncle. At its termination

the stripe continues as a row of variably spaced

dark spots to the base of the dorsal caudal rays.

There are also a variable number of dark spots

posteriorly on the body. In a few specimens the

dark stripe or the spots (but not both) may be

absent.

The spinous dorsal fin bears a jet black row of

dark spots, restricted mostly to the interspinous fin

membranes. These spots decrease in size and intensity posteriorly and may extend onto the interradial membrane of the rayed dorsal for a varying

distance. There is no pigment distally on the rayed

dorsal or anal fins. The upper and lower lobes of

the caudal are dusky.

Specimens from the southern Red Sea and the

Bay of Tadjourah, outside the entrance to the Red

Sea, differ from those described above in having

the dark pigment generally blacker and covering

more area. The stripe on the body may be almost

completely obscured anteriorly by the surrounding

body pigment, but the stripe is absent in the holotype (see Smith, 1959, fig. 12, for an illustration of

the holotype). In addition, some specimens exhibit

dark pigment at the tips of some of the dorsal and

anal rays. Specimens from the northern and southern Red Sea (including the Bay of Tadjourah, type

locality) are easily distinguishable (Figures 17 and

18). I do not believe that the northern populations

merit nomenclatural distinction, however, as there

is a good possibility that the two forms gradually

merge somewhere in the midportion of the Red Sea.

There are no specimens available from the approximately 1500 km between the southernmost collections of the northern form and the northernmost

collections of the southern form.

The two types of color pattern described here

are paralleled in the unrelated blenniid genus

Meiacanthus, which co-occurs with E. gravieri (see

discussion of mimicry on page 6).

In life, E. gravieri is generally slate blue anteriorly

grading into pale yellow posteriorly. The black

markings are black.

REMARKS (see also discussion under E. aroni).—In

the northern Red Sea and Gulf of Aqaba, E. gravieri

occurs commonly in depths of 2 to 10 meters, but

is rarely seen at greater depths. The species is frequently seen swimming freely, but such activity is

punctuated by long periods of rest on the bottom.

E. gravieri is apparently territorial and during

several periods of observation, individuals were

noticed to have a few favorite perching spots. An

individual might leave one of these spots and swim

around, up to about 5 meters distance from such

a spot, but it always returned to one of the favored

spots to perch.

Klausewitz (1960) illustrated E. gravieri but identified the specimen as E. nigrovittatus, a synonym

of £. frontalis.

Lotan (1969) compared E. klausewitzi with E.

gravieri, differentiating E. klausewitzi by its having

a black line on the body, which the holotype of

E. gravieri lacked. The presence or absence of such

a dark line is variable, as noted in the color pattern

description above. The holotype of E. klausewitzi

is a small specimen of the southern Red Sea dark

form of E. gravieri.

MATERIAL (localities abbreviated).—(Pale form)

Gulf of Aqaba: UTAI NS-3580 (3: 20.4-48.1), HUI

HUJ-F-4382 (38.6), HUJ-F-4377 (2: 20.4-21.0),

E-64/30A (2: 20.0-32.4), USNM 204542 (9: 32.1-

55.9), 204555 (15: 19.3-33.2), 204546 (9: 17.5-23.3),

204544 (22: 20.2-52.7), 204543 (3: 19.5-60.8); Ras

Muhammad, southern tip of Sinai Peninsula, Red

Sea: USNM 204545 (9: 27.8-44.1), 204547 (2: 27.8-

41.3); Et Tur, Sinai Peninsula, Gulf of Suez: USNM

204548 (3: 50.1-54.7); Al Ghardaqa, Egypt, northern

Red Sea: NFIS 5122-23 (2: 29.6-34.6).
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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FIGURE 10.—Graph showing relationship of longest caudal ray length to standard length in the

two color pattern forms of Ecsenius frontalis from Israeli coastal waters and one of these forms

from Ethiopian coastal waters.

(Dark form) Dahlak Archipelago, Red Sea: HUI

E-62/3708 15.1, holotype of Ecsenius klausewitzi);

Isola Delemme (15°30.5' N, 39°54' E), Red Sea:

USNM 204479 (56.5); Sheikh el Abu (16°06" N,

39°36.5' E), Red Sea: USNM 204480 (3: 30.5-40.1);

Bay of Tadjourah: MNHN 04-319 (54.2, holotype

of Salarias gravieri).

In addition to the above, a large number of

uncataloged USNM specimens I collected in the

Gulf of Aqaba were examined for color pattern

and ecological data.

Ecsenius (Ecsenius) aroni, new species

FIGURE 19

DESCRIPTION.—Dorsal spines 12-14 (usually 13);

dorsal rays 17-18; segmented anal rays 19-20; pectoral rays 13-15 (usually 14); segmented caudal rays

13; dorsal procurrent caudal rays 6-9 (usually 7 or

8); ventral procurrent caudal rays 6-9 (usually 7

or 8); total caudal elements 25-30; gill-rakers 12-15;

pseudobranchial filaments 6-7; lower incisor teeth

46-57; lower jaw posterior canines 0-1; total lower
NUMBER 72 25

jaw posterior canines 0-2 (usually 2); upper incisor

teeth 130-132 (three counts); precaudal vertebrae

10 or 11 (rarely 10); caudal vertebrae 23-26 (rarely

23); total vertebrae 34-37 (rarely 34); epipleural

ribs 11-14. Lateral line with no paired pores, extending to below level of 8th to 10th dorsal spine

(usually 9th or 10th). Dorsal fin notched six-ninths

to nine-ninths length first dorsal ray. Third (innermost) pelvic ray barely or not obvious. One cirrus

on each anterior nostril.

Color pattern: The color pattern of preserved

specimens is not very variable. The head is dark

brown, but pale underneath. The body is generally

dusky brown with a dark spot on the caudal

peduncle that extends well onto the caudal fin.

For most of its length, the dorsal fin bears a narrow

suprabasal dusky stripe, but is otherwise scarcely

marked. The anal fin is generally dusky and in

mature males the tips of the rays are pale. The

other fins are pale or unmarked.

In life the body is generally dark tan with a

black spot with blue overtones at the base of and

on the caudal fin. Some individuals exhibited a

bright orange stripe along the dorsal body contour.

The head and anal fin are dark brown. The other

fins are pale or unmarked.

HABITAT AND DISTRIBUTION.— Ecsenius aroni is

known only from the Red Sea and Gulf of Aqaba

coasts of the Sinai Peninsula. It occurs in coral

areas only and is commonest at depths greater than

10 meters. It was only rarely seen or taken at depths

as shallow as 3 meters. The species was seen (but

not collected) at the nature preserve in Eilat at

depths as great as 36.6 meters, but was not found

at greater depths, down to 45.7 meters (two dives

planned primarily to search for the species). It

prefers the undersides of ledges and pockets in

the reefs and was not seen in the open areas above

or about the reefs.

RELATIONSHIPS.—Ecsenius aroni is closely related

to E. gravieri, with which it generally occurs in

the Gulf of Aqaba, and E. pulcher, which occurs in

the Persian Gulf and northern Arabian Sea. With

these two species it shares the following unique

combination of characters: typically 13 dorsal spines,

a notched dorsal fin, 11 precaudal vertebrae, typically 14 pectoral rays, and 13 segmented caudal

rays. It differs from these two species primarily in

color pattern, proportions and meristics as indicated

in key couplets 16 and 17. In addition it differs

from the sympatric E. gravieri in that caudal filaments are primarily restricted to males (well developed in both sexes in E. gravieri), and in males the

longest caudal ray tends to increase in relative

length regularly with increase in standard length

(Figure 11). In general, for either sex, E. aroni has

a relatively shorter caudal than E. gravieri (Figure

11). E. aroni attains a much smaller maximum size

than either E. gravieri or E. pulcher (Figure 11).

E. aroni occurs more commonly in greater depths

than the sympatric E. gravieri, which is usually

found in less than 10 meters depth. The two species

behave quite differently. E. gravieri is an obvious

species on the reef and is frequently seen swimming

above the bottom over open sandy areas closely

adjacent to the reef. E. aroni is secretive and was

seen (rarely) free swimming only within small holes

in the reef.

ETYMOLOGY.—This species is named for Dr. William Aron of the Smithsonian Institution in appreciation for his making possible my field work in the

Red Sea. The results of that work form an invaluable contribution to this and other of my studies.

HOLOTYPE.-USNM 204468, male, 36.8 mm SL,

bay at El Himeira, Sinai Peninsula, Gulf of Aqaba.

Depth of collection, to 18 meters. Collected by

V. G. Springer et al., 16 July 1969. Original field

number, VGS 69-2.

PARATYPES (all Sinai Peninsula).-USNM 204690

(12:18.5-40.8), collected with the holotype; USNM

204560 (21:19.4-31.5), 204550 (2:21.4-27.0), 204558

(24.7), all collected in same vicinity as holotype

but on different dates and at different depths;

USNM 204557 (2:21.3-35.4), Marsa Muqabila, Gulf

of Aqaba; USNM 204556 (33.1), bay between Marsa

Mokrakh and El Himeira; USNM 204561 (40.8)

and 204562 (7:25.8-40.5), Ras Muhammad, Red Sea.

Some paratypes will be deposited with the Hebrew

University after publication of this paper.

Ecsenius (Ecsenius) bicolor (Day)

FIGURES 20, 21

Salarias bicolor Day, 1888, p. 798 [Saddle Island, off Kyoukphyoo, Arakan, Burma].

Salarias furcatus Johnstone [not of De Vis], 1904, p. 213

[Chilaw Paar, Ceylon].

Salarias burmanicus Hora and Mukerji, 1936, p. 34 [Maungmagan, Tavoy District, Lower Burma].

Salarias melanosoma Regan, 1909, p. 406 [Christmas Island,

Indian Ocean].
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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FIGURE 11 .—Graph showing relationship of longest caudal ray length to standard length in spedes

of the E. pulcher species group.

Ecsenius hawaiiensis Chapman and Schultz, 1952, p. 526

[Pearl Harbor, Oahu, Hawaii; a possible introduction from

Guam].

DESCRIPTION.—Dorsal spines 11-12 (rarely 11);

dorsal rays 15-18 (rarely 15); segmented anal rays

17-21 (rarely 17 or 21); pectoral rays 12-14 (rarely

12 or 14); segmented caudal rays 13-14 (rarely 13;

15 in one specimen 20.7 mm. SL); dorsal procurrent

caudal rays 6-9 (rarely 6 or 9); ventral procurrent

caudal rays 6-8; total caudal elements 26-31 (rarely

26 or 31); gill rakers 12-18 (rarely 12 or 18); pseudobranchial filaments 5-8 (rarely 5 or 8); lower incisor

teeth 36-46 (one of 243 specimens with 32); lower

jaw posterior canines 0-1; total lower jaw posterior

canines 0-2 (usually 2); upper incisor teeth 114-129

(11 counts); precaudal vertebrae 10 (one specimen

of 233 with 11); caudal vertebrae 23-25; total vertebrae 33-35; epipleural ribs 12-17 (rarely 12 or 17).

Lateral line with vertically paired pores for onequarter to all its length (rarely less than one-half

length), extending posteriorly to beneath level of

10th-13th dorsal fin element (rarely to 13th). Dorsal

fin notched two-ninths to seven-ninths (rarely less

than four-ninths) length first dorsal ray. Third

(innermost) pelvic ray varying from obvious to not

obvious (usually obvious). Two cirri on anterior

nostril, one each arising from anterior and posterior

margins.
NUMBER 72 27

Color pattern: Ecenius bicolor is one of the more

variably pigmented species of Ecsenius. Three of

the nominal species referable to the synonymy of

E. bicolor are merely color pattern variants. There

are, however, two basic color patterns found in

preserved specimens, which, with modifications,

grade into each other. On the basis of life color

variations exhibited by a single individual reported

by Wickler (1965A) and in a group of specimens

reported by Strasburg (1967), it appears that life

color variations may be fixed at the time of preservation. Of the two basic patterns, either one or the

other greatly predominates in individual collections

from any locality. While either type may occur as

the predominant form in individual collections

from the same general locality, collections indicate

that one form or the other is much more often

collected.

In one of the basic patterns the head and body

are, for the most part, completely dark dusky, occasionally slightly less intensely so on the posterior

half of the body. There may be a dark spot just

posterior to the eye. The spinous dorsal fin is dusky

or bears a dusky band on its ventral half with a

black spot between the first and third spines. The

black spot is rarely absent. The rayed dorsal is

more faintly dusky than the spinous dorsal and

slightly darker basally. The anal is dusky, frequently

paler basally and on the tips of the rays. The caudal

bears a faint to dark dusky triangular mark centrally, the base of the triangle vertically aligned

with the caudal base. The uniformly dark color

pattern is the type exhibited by the holotype of

S. melanosoma. A variation of this pattern, wherein

there are several vertical, pale streaks tm the body

and the corners of the mouth are pale, was the

basis for the description of E. hawaiiensis. In some

specimens there is a broad, dark dusky longitudinal

stripe extending from the level of the pectoral axil

posteriorly to some point anterior to the caudal

peduncle. The stripe may be broken into as many

as five portions.

The other basic color pattern is similar to the

first in that the head and body anteriorly are more

or less uniformly dark, but at some point no further

anteriorly than below the level of the third or

fourth dorsal ray, the body gradually or abruptly

becomes paler or completely lacks melanophores.

A colored photograph of a freshly dead specimen

of this color pattern type occurs in Axelrod and

Emmens (1969). In many specimens there are one

to three narrow, vertical, diffusely dusky bars in

the pale area on the body. (One of these is slightly

apparent in figure 21.) A number of specimens

examined also exhibited the longitudinal dark stripe

on the body as described above.

Wickler (1965A) illustrated and discussed life

color variations in E. bicolor, based on aquarium

specimens presumably from Ceylon. He ascertained

that these patterns reflected emotional states. He

described life color generally as dark gray-brown

anteriorly and yellow or orange-red posteriorly.

Strasburg (1967) reported on life coloration and

noted that the lengthwise stripe (described above)

was two toned, black and white, with the white

color turning black in preservative. Strasburg reported that E. bicolor from Eniwetok was bright

ultramarine, sometimes tinged with violet anteriorly,

and orange-yellow posteriorly. G. Losey (in litteris)

described E. bicolor, also from Eniwetok, as bluegray anteriorly and yellow-orange posteriorly. At

One Tree Island, Great Barrier Reef, specimens I

saw in life were electric blue anteriorly and yelloworange posteriorly. One individual was seen with

a rose-colored area posterior to the eye. (Such rosecolored marks are found in excited specimens, according to Wickler.) The illustration in the Tickell

manuscript of the Zoological Society of London,

upon which Day based his description of S. bicolor,

is rich blue anteriorly and red-orange posteriorly.

GEOGRAPHIC VARIATION.—Average numbers of dorsal and anal rays and caudal vertebrae (usually

correlated characters) tend to vary from one locality

to another (Table 15). In general, the highest

averages occur in the Australian specimens. Australian specimens also appeared to have average

longer posterior nasal cirri and shorter fifth dorsal

rays (Tables 8 and 10) than specimens from other

localities. The nasal cirri were sexually dimorphic

in size in the Australian specimens but not in the

others. The length of the fifth dorsal ray was

sexually dimorphic in non-Australian specimens but

was either not dimorphic or only slightly so in

Australian specimens.

REMARKS.—In Figure 1 the acceptable literature

records for E. bicolor entered on the map were

those reported in Day (1888), Hora and Mukerji

(1936), and Kamohara and Yamakawa (1965).

Wickler (1965A, B, C, D) has given a detailed

discussion on the behavior and biology of E. bicolor,
28

including a list of moving picture films deposited

with the Encyclopedia Cinematographies.

Strasburg (1967) reported that E. bicolor occurred

in the lagoon at Eniwetok at depths from about

1 to 7.5 meters, being more common at the latter

depth. In contrast, I collected the species at One

Tree Island only on the ocean side of the island

at depths of about 4 and 12 meters and found the

species common at both depths. Most collections

of the species, however, appear to have come from

the lagoon side of atolls in shallow depths, although

Randall collected a specimen in Guam from a

depth of 21.3 meters.

Chapman and Schultz (1952) mentioned that E.

bicolor (as E. hawaiiensis) might have been introduced into Hawaiian waters from Guam, where

the species is known to occur, as its collection in

Hawaii was made under peculiar circumstances.

They discarded the possibility, however, as they

believed the species to be endemic to Hawaii.

Strasburg (1956) believed that the species probably

was brought to Hawaii from Guam in the fouling

on the bottom of a barge, and thus was not a

Hawaiian endemic, but possibly had become established in Hawaiian waters as a consequence of its

introduction. Strasburg did not recognize the species

as E. bicolor, but accepted the name E. hawaiiensis.

Considerable collecting in Hawaiian waters in

recent years has produced no additional material

of E. bicolor other than that reported by Chapman

and Schultz. In view of this and the fact that

Hawaii is considerably removed from the nearest

known locality where E. bicolor occurs, I believe

that the Hawaiian specimens were introductions

and that the species has not become established in

Hawaii.

NOMENCLATURE.—The description of Salarias bicolor was based on a short description and colored

illustration in a handwritten manuscript by Colonel

S. R. Tickell, dated 1875, and titled: "Fishes collected in the seas and fresh waters of British Burma

from 1851 to 1864, vol. 1." The manuscript is

housed in the library of the Zoological Society of

London. Many of the descriptions contained in

the manuscript are the sole bases of many of F.

Day's new species descriptions. The specimens on

which Tickell based his descriptions were evidently

not retained as can be inferred from comments in

the manuscript.

The colored illustration of S. bicolor in the

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

manuscript is of a fish 1% (35 mm) inches long.

The specimen is rich blue anteriorly below the

rayed portion of the dorsal fin; thence it is redorange. The species is portrayed as having a deep

notch in the dorsal fin between the spinous and

rayed portions, somewhat deeper than for any

specimen I have seen. The species is described as

having no tentacles, but a statement was included

to the effect that if there were any tentacles they

"were very likely to be overlooked." The dorsal fin

formula was given as 11 + 18, the anal as 18.

The distinctive life color in combination with

the notched dorsal fin and a dorsal spine count of

at least 11 leaves little doubt of the identification

of Tickell's figure with the species treated here as

E. bicolor. The only blenny that approximates the

appearance of E. bicolor is the nemophidinine

Meiacanthus atrodorsalis (Giinther), which does not

have a notched dorsal and has fewer than 11 dorsal

spines (Tickell's count is probably in error and

the reduced last—12th—dorsal spine was probably

missed). From my observations M. atrodorsalis is

more yellow than orange posteriorly. It would probably be well to designate a neotype for S. bicolor,

but as I have no material from the same geographic

area from which S. bicolor was described, it is not

possible.

Salarias furcatus was not compared with any other

blenny and the description and illustration leave

no doubt that it is a synonym of S. bicolor. I have

been unable to locate the holotype.

Salarias burmanicus was compared only with S.

anomalus (= S. pulcher). The illustration and description clearly indicate bicolor as the species.

W. F. Smith-Vaniz examined the holotype in the

Indian Museum in Calcutta. It bears the number

F 11872/1, type 429.

Salarias melanosoma was compared only with 5.

anomalus. I have examined the syntypic material

and find all specimens to be the same as S. bicolor.

Ecsenius hawaiiensis was compared with E. bicolor, but it was thought that the pale bars on the

sides were distinctive. Wickler (1965A) has described

and illustrated the various color patterns exhibited

by E. bicolor, of which the pale barred form is but

one of several. He concluded that E. bicolor and

E. hawaiiensis were the same species. My examination of the type material of the latter indicated no

morphological differences between that material

and other specimens I assign to E. bicolor.
NUMBER 72

MATERIAL (localities abbreviated).—Christmas Island, Indian Ocean: BMNH 1909.3.4.52-57 (5: 30.0-

48.8, syntypes of Salarias melanosoma), USNM

157373 (40.7, sytitype of Salarias melanosoma); Ceylon: USNM 204623 (43.3), NFIS 5256 (6: 35.4-59.0,

specimens reported by Wickler, 1965A, presumably

from Ceylon); Maldive Islands: NFIS 9578 (49.1);

Nicobar Islands: NFIS 9576 (41.4); Pulo Penju,

Indonesia: USNM 201864 (3: 23.4-52.1); Patong,

Phuket, Thailand: USNM 201755 (5: 25.0-45.0);

Hot du Sud, South China Sea: CAS GVF-2113 (9:

48.3-61.3); Ryukyus: BPBM 7467 (46.4); Guam:

BPBM 7851 (2: 26.0-35.1); Timor: ZMA 109.064

(36.9); Banda Sea: USNM 195718 (2: 36.8-42.0);

Palau Islands: CAS GVF-740 (5: 20.7-48.5), GVF1450 (59.5); Kapingamarangi: CAS GVF-424 (5:

22.0-46.9), GVF-452 (38: 18.5-56.7); Mortlock:

ZSZM 5152 (41.5); Marshall Islands: BPBM 5895

(20: 32.3-67.6), USNM 201472 (8: 16.1-56.6), 201473

(43.9), 201475 (23.4), 201476 (5: 38.0-59.3), 201477

(3: 26.1-48.3), UW 12008 (3: 34.0-0-54.0); Ponape

Island: CAS GVF-496 (20: 24.5-61.0), GVF-497

(50.7); New Britain: USNM 201813 (3; 47.7-49.8);

Neiafu, Tonga Islands: USNM 203334 (2: 39.6-

44.1); Fiji Islands: USNM 201827 (41.9); Espiritu

Santo, New Hebrides: USNM 144293 (2: 29.5-47.5),

144294 (2: 52.2-62.0), 144716 (7: 27.1-45.0), CAS

GVF-1826 (24.5); One Tree Island, Queensland,

Australia: USNM 201368 (74: 38.5-69.3, including

two cleared and stained); Endeavour Reef, Queensland, Australia: ANSP 109699 (2: 31.4-37.7); Oahu,

Hawaii: USNM 112293 (66.5, holotype of Ecsenius

hawaiiensis), 112294 (12: 48.0-77.0).

Ecsenius (Ecsenius) namiyei

(Jordan and Evermann)

FIGURE 22

Salarias namiyei Jordan and Evermann, 1902, p. 362 [Pescadores Islands].

DESCRIPTION.—Dorsal spines 11-12; dorsal rays

18-20; segmented anal rays 20-22; pectoral rays 13

(one specimen each with 11 and 12, and two with

14, on one side only); segmented caudal rays 13-14

(usually 14); dorsal procurrent caudal rays 7-8;

ventral procurrent caudal rays 6-8; total caudal

elements 26-30 (usually 28-29); gill-rakers 12-17;

pseudobranchial filaments 6-8 (usually 7); lower

incisor teeth 35-45; lower jaw posterior canines

29

0-1 (usually 1); total lower jaw posterior canines

1-2 (usually 2); upper incisor teeth 129 (one count);

precaudal vertebrae 10-11; caudal vertebrae 25-27;

total vertebrae 36-37; epipleural ribs 12-14. Lateral

line with vertically paired pores for none to onehalf its length, usually some pairs present, extending

posteriorly to beneath level of 10th-13th dorsal fin

element (usually 11th—12th). Dorsal fin without

notch. Third (innermost) pelvic ray obvious. Two

cirri on each anterior nostril, one each arising from

anterior and posterior margins.

Color pattern: (Taiwan specimens) The body,

head, and dorsal and anal fins are dark black,

with the exception that in the caudal peduncular

region the melanophores are less densely distributed,

and that area is paler. The bodies of some specimens

exhibit three or four short, narrow, pale, vertical,

widely spaced marks on each side, similar to marks

that are found in some specimens of E. bicolor.

Most of the specimens exhibit a row of microscopic

pale spots along the midportion of the body in the

region where the lateral line might be expected to

occur. Still others have a few small oblong intensely

dark spots that underlie the dark surface pigment

on the sides. The spinous dorsal fin bears irregular

deeply pigmented lines below the dark surface

pigment. The anal fin bears an indistinct darker

stripe along its midlength. The caudal, pelvic, and

pectoral fins are dusky with a greater concentration

of pigment along the rays.

The life coloration of the female illustrated in

Figure 13 (preserved color pattern), which had been

in an aquarium for an indefinite period not exceeding a few weeks, was as follows: head and body dark

brown; caudal peduncle grading into yellow-amber

and then orange on proximal portion of caudal fin.

Body with eight small, midlateral, blue-white spots

beginning in the pectoral fin axil and extending no

farther than the level of the last dorsal rays. Similar

spots on pectoral base and on dorsal body contour,

where they were less obvious than the midlateral

spots. Upper lip with blue-white bands separated

by dark interspaces. Pale stripe from behind eye

to lip. Some pale blue dashes on cheek. There is

no evidence of the blue-white markings on the

preserved specimen.

REMARKS.—Tomiyama (1955) has given a thorough redescription of the holotype of E. namiyei.

On the north end of Taiwan I collected this
so

species from depths of about one to three meters

very close to shore.

The specimens, one each, from New Britain and

nearby Kerward Island, are very similar to those

from Taiwan, but the color of their caudal peduncles is of about the same intensity as the rest of

the body. An anal ray count was possible for only

one of these specimens. The count was 20 and is

lower than for any of the Taiwan specimens. The

damaged specimen had a dorsal ray count of 18,

lower than for any Taiwan specimen. These facts

probably indicate racial variation.

MATERiAL.-Pescadores Islands: ZITU 5726 (49.5,

radiograph of holotype of Salarias namiyei); Taiwan

(north end): USNM 203125 (63.6, cleared and

stained), 203126 (2: 58.0-63.2), 203127 (5: 62.0-

80.4), 203128 (5: 55.4-81.8), 204478 (9: 59.0-76.3);

New Britain: USNM 201865 (47.7); Kerward Island:

USNM 201866 (49.0).

Ecsenius (Ecsenius) stigmatura Fowler

FIGURE 23

Ecsenius lineatus Klausewitz, 1962, p. 145 [Madewaru Island,

Fadiffulu Atoll, Maldive Islands].

DESCRIPTION.—Dorsal spines 12; dorsal rays 17-18;

segmented anal rays 19-20; pectoral rays 13 (one

specimen each with 12 and 14 on one side only);

segmented caudal rays 13; dorsal procurrent caudal

rays 5-8 (one specimen each with 5 and 8; all others

had 7); ventral procurrent caudal rays 6-8; total

caudal elements 25-28 (one had 25, none had 26);

gill-rakers 13-16 (one had 16, none had 15); pseudobranchial filaments 6-7 (one had 7); lower incisor

teeth 45-49; lower jaw posterior canines 1; total

lower jaw posterior canines 2; upper incisor teeth

125 (one count); precaudal vertebrae 10-11 (two

had 11); caudal vertebrae 24-26; total vertebrae

34-36 (usually 35); epipleural ribs 13-15. Lateral

line with no paired pores, extending posteriorly to

below 10th dorsal spine. Dorsal fin notched sixninths to seven-ninths length first dorsal ray. Third

(innermost) pelvic ray obvious. One cirrus on each

anterior nostril.

Color pattern: The commonest color pattern,

except as noted below, is that illustrated in Figure

23. The dark midlateral spots on the side of the

body are always nine in number (the anteriormost

spot is actually part of a continuous stripe extending

to the eye). In the holotype and one small specimen,

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

20.5 mm SL, the spots are not evident, but instead

there is a single, broad, uninterrupted, dark midlateral stripe on the side. In another specimen the

spots are connected and are distinguishable only

as slight constrictions along the length of the stripe.

Usually the area of the body below the stripe, or

row of midlateral spots, is abruptly and uniformly

pale, and there is no evidence of a thin pale stripe

below the row of dark spots as in Figure 23. The

thin pale stripe above the row of spots is also

frequently obscured. The presence of the tearshaped spot on the caudal peduncle with the tip

extending onto the central base of the caudal fin

is distinctive of this species.

There is some variation in the markings on the

head. The narrow pale stripes paralleling the dark

postorbital stripe are not always distinguishable.

The underside of the head is most noticeable for

a pale area covering the lower lip and the region

just behind the lip. The remainder of the underside

of the head is variably dusky, sometimes with

indistinct evidence of diffuse dark spots on both

sides. The upper lip may be uniformly or irregularly

dusky, sometimes with three bands anteriorly separated by narrower, paler interspaces.

In life the dark areas are various shades of brown

and black and the pale areas are faintly yellow.

A field assistant described one fresh-caught specimen

as having much bright yellow color, but fading

rapidly after preservation.

REMARKS.—E. lineatus co-occurs with E. namiyei

in northern Taiwan. It was collected at depths of

about 1 to 9 meters, the latter depth somewhat

deeper than that from which specimens of E.

namiyei were obtained.

MATERIAL (localities abbreviated).—Maldive Islands: NFIS 5532 (65.9, holotype of Ecsenius

lineatus); north end of Taiwan: USNM 203129

(4: 42.1-48.2, including one cleared and stained),

203130 (2: 48.6-55.4), 203131 (54.9), 203132 (2:

44.9-49.5), 204477 (60.1); Pratas Reef, South China

Sea: CAS GVF-1749 (20.5); Vietnam: CAS GVF2116 (46.6).

Ecsenius (Ecsenius) stigmatura Fowler

FIGURES 24, 25

Ecsenius stigmatura Fowler in Chapman and Schultz, 1952,

p. 514 [Dammi Island, but see remarks section below].

DESCRipnoN.-Dorsal spines 12; dorsal rays 16-17;

segmented anal rays 18-19; pectoral rays 13-15 (13
NUMBER 72

on only one side of one of the four specimens);

segmented caudal rays 14; dorsal procurrent caudal

rays 7-8; ventral procurrent caudal rays 6-7; total

caudal elements 27-28; gill-rakers 15-17; pseudobranchial filaments 6; lower incisor teeth 37-40;

lower jaw posterior canines 1; total lower jaw posterior canines 2; upper incisor teeth 107 (one count);

precaudal vertebrae 10; caudal vertebrae 23-24;

total vertebrae 33-34; epipleural ribs 12 (one count).

Lateral line with no paired pores, extending posteriorly to beneath level of 9th dorsal spine. Dorsal

fin notched six-ninths to seven-ninths length of

first dorsal ray. Third (innermost) pelvic ray varying from obvious to not obvious. One cirrus on

each anterior nostril.

Color pattern: The ground color of the head and

body is evenly dusky brown. There is a narrow dark

stripe that begins at the posteroventral margin of

the eye and extends posteriorly across the opercle.

Below the opercular portion of the stripe is a

narrow pale stripe that is in line with a similar

area margining the upper fleshy pectoral base. A

large dark spot extends anteriorly from the anus.

Another large dark spot occurs on the middle of

the caudal peduncle and extends onto the bases

of the caudal rays. The stripe has a pale margin

anteriorly. The fins are variably dusky.

Three of the four specimens available are females.

One female is from Masbate, Philippine Islands,

and the others are from Tomahu Island in the

Moluccan Sea. The fourth specimen, a male from

Borneo, may not be the same species. The male

differs from the females in lacking the pale stripe

below the dark stripe on the head and in lacking

the large spot on the caudal peduncle. Whether

these differences are an indication of sexual dimorphism, individual variation, or population differences cannot be decided at present.

A color sketch of the holotype made during the

Albatross expeditions indicates that the specimen

was generally reddish brown in life. The dark streak

behind the eye was deep blue and the pale streak

below this was orange. The pupil was surrounded

by a narrow orange border and this surrounded by

a reddish-brown ring. The anal spot was black with

deep blue overtones. The caudal spot was black

with a pale lavender margin. The spinous dorsal

was reddish brown basally and orange-brown distally. The rayed dorsal and most of the caudal

were pale slate gray. There is a narrow orangebrown stripe along the dorsal body contour extending from the anterior end of the dorsal to the

caudal base.

REMARKS.—In the original published description

accredited by Chapman and Schultz (1952) to

Fowler, the holotype and one paratype of E. stigmatura are reported to have been taken at Dammi

Island, between Jolo and Tawi Tawi straits, at a

depth of 244 fathoms (ca. 443 meters) on 21 September 1909. In an unpublished manuscript kindly

lent me by J. E. Bohlke, however, Fowler stated that

he was unable to ascertain the type locality. How

the published locality was arrived at is not now

known. I became suspicious of the published locality because no salariinine blenniids are known

from depths nearly as great as 244 fathoms.

The original watercolor illustration upon which

the published illustration of E. stigmatura was based

is in the files of the Division of Fishes, National

Museum of Natural History. This illustration bears

a number, 2131, that corresponds with a number

written on a silk tag that was tied to the holotype.

Also written on the illustration was the following:

Tomahu Id, 12/11-09 see color notes. I have been

unable to locate the color notes or any other record

of the tag number 2131, but the locality and date

(interpreted as 11 December 1909) agree with the

published information (Anonymous, 1910, p. 92)

for a collection made by the steamer Albatross

(depth given as 2-15 feet) at Tomahu Island. A

handwritten ledger in the files of the Division of

Fishes, recorded during the Albatross collections,

indicates that blennies were collected at this station.

The published locality and date for the holotype

of E. stigmatura are also in agreement with a published Albatross station (Anonymous, 1910, p. 82),

but the ledger does not record that any blennies

were collected at that station and they would not

be expected from such a depth (ca. 443 meters).

In view of the above information I prefer to

ascribe the Tomahu Island locality to the holotype

and one paratype of E. stigmatura. In addition, as

the second paratype, according to the locality records, came from either beach seining or dynamiting

at a depth up to about 4.5 meters, and no salariinine

is known to have a very extensive depth range, it

seems much more probable that the species is restricted to shallow depths.

MATERiAL.-Tomahu Island: USNM 99379 (45.9,

holotype of Ecsenius stigmatura), 111878 (36.6);

383-026 O - 70 - 3
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Cataingan Bay, east of Masbate Island, Philippines:

USNM 122444 (36.5); Pulav Gaya, Darvel Bay,

Borneo: USNM 201815 (37.7).

Ecsenius (Ecsenius) lividinalis Chapman and

Schultz

FIGURE 26

Ecsenius lividinalis Chapman and Schultz, 1952, p. 517

[Munda, New Georgia, Solomon Islands].

DESCRIPTION.—Dorsal spines 12; dorsal rays 12-14

(one each with 12 and 14); segmented anal rays

13-14; pectoral rays 13-14 (one of 11 with 14 on

one side only); segmented caudal rays 13-15 (usually 14); dorsal procurrent caudal rays 5-6; ventral

procurrent caudal rays 4—6; total caudal elements

23-26; gill-rakers 12-14; pseudobranchial filaments

5-7 (usually 6); lower incisor teeth 30-37; lower jaw

posterior canines 1 (2 on one side in one specimen);

total lower jaw posterior canines 2-3 (3 in one

specimen); upper incisor teeth 98-111 (four counts);

precaudal vertebrae 10-11 (one of 10 with 11);

caudal vertebrae 19-20; total vertebrae 29 or 30;

epipleural ribs 11-13. Lateral line with no paired

pores, extending posteriorly to below level of 9th11th dorsal spine. Dorsal fin notched three-ninths

to five-ninths length of first dorsal ray. Third (innermost) pelvic ray obvious. One cirrus on each anterior

nostril.

Color pattern: The available specimens of E.

lividinalis are all old and to some degree faded.

They appear to have been uniformly dark when

fresh, with the fins perhaps paler than the body.

The most obvious mark is a dark spot that envelops

the anus and extends a short distance anterior from

the anus.

MATERIAL (localities abbreviated).—New Georgia:

USNM 144723 (33.0, holotype of Ecsenius lividinalis), 144292 (6:22.1-26.8, including one cleared

and stained), 144291 (3:22.1-30.5); Philippine Islands: ANSP 109211 (26.0).

Ecsenius (Ecsenius) yaeyamaensis (Aoyagi)

FIGURES 27-29

Salarias yaeyamaensis Aoyagi, 1954, p. 213 [in part, Iriomote

Island, Riukius].

DESCRIPTION.—Dorsal spines 12-13 (rarely 13); dorsal rays 13-15 (usually 15); segmented anal rays

15-17 (usually 16); pectoral rays 12-14 (rarely 12

or 14); segmented caudal rays 13 (14 in one of 120

specimens); dorsal procurrent caudal rays 6-9 (usually 7-8); ventral procurrent caudal rays 6-8 (usually 7); total caudal elements 25-30 (rarely 25 or 30);

gill-rakers 11-16 (rarely 11 or 16); pseudobranchial

filaments 5-8 (rarely 5 or 8); lower incisor teeth

43-56; lower jaw posterior canines 0-1 (usually 1);

total lower jaw posterior canines 0-2 (usually 2);

upper incisor teeth 97-127 (6 counts); precaudal

vertebrae 10; caudal vertebrae 21-23 (usually 22);

total vertebrae 31-33 (usually 32); epipleural ribs

11-14 (rarely 11); lateral line without paired pores,

extending posteriorly to beneath level of 10th-12th

dorsal spine (one specimen each, of 110, to 8th and

12th spine). Dorsal fin notched seven-ninths to

eight-ninths length first dorsal ray (usually eightninths). Third (innermost) pelvic ray varying from

obvious to not obvious (usually not obvious). One

cirrus on each anterior nostril.

Color pattern: Aside from a complex and variable

color pattern, E. yaeyamaensis comprises several geographic populations that are distinguishable only

on color pattern. Although treated here as a separate species, E. nalolo may more properly be placed

with E. yaeyamaensis. In contrast with the color

pattern populations of E. yaeyamaensis, which are

generally of limited geographic occurrence, E. nalolo

occurs over most of the Indian Ocean and Red Sea

with little noticeable geographic variation in color

pattern.

The major feature of the color pattern that links

all the populations of E. yaeyamaensis is a dark,

forked stripe on the fleshy pectoral base. The arms

of the stripe extend onto the bases of the fin rays;

no other species of Ecsenius has such a mark. (E.

nalolo has a single, unforked, stripe on the fleshy

pectoral base.)

The commonest color pattern is best typified by

that illustrated in Figure 27. This color pattern is

found in specimens from the South China Sea,

Taiwan, the Ryukyus, Palaus and adjacent islands,

New Britain, and New Georgia. The illustrated

specimen has a fully developed pattern. Other

specimens from the localities just cited are quite

variable and are allied by lacking specific features

that are found in specimens from other localities.

Banda Sea: A single specimen is available (Figure

29), which has a pattern unlike that from any other

locality. The distinctive features are two long, narrow, dark stripes on the body below the anterior
NUMBER 72

dorsal rays, continuous as dark dashes and ultimately as four dark spots on the caudal peduncle.

(A single, badly faded specimen of E. yaeyamaensis

was available from Flores. Only faint remnants of

the forked stripe on the pectoral base were visible.

In view of the color pattern exhibited by the

specimen from the Banda Sea, which is relatively

close to Flores, it would be important to know

what color pattern is exhibited by specimens of

E. yaeyamaensis from the various islands surrounding the Banda Sea.)

New Hebrides: Three specimens (none illustrated), with two aligned, lengthwise rows of short,

dark dashes on the body grading into spots below

the posterior dorsal rays. If the dashes were to

fuse anteriorly, they would form stripes as in the

preceding form. The fact that the Banda Sea and

the New Hebrides are quite distant from each other

and that almost midway between these two localities,

at New Britain and New Georgia, the common

color pattern occurs, leads me to conclude that the

populations are probably distinct.

Australia: All of a considerable number of specimens (not all cited in the material list) from the

Capricorn group of islands at the southern end of

the Great Barrier Reef and Endeavour Reef, off

northern Queensland, exhibit numerous fine, dark

spots on the posterior half of the body (Figure 28).

These spots occur in at least two horizontally

aligned rows, but there is usually a sprinkling of

unaligned spots both above and between the rows.

All of the specimens are larger than those from

the New Hebrides and it may be that Australian

specimens of sizes equal to those from the New

Hebrides will be similar in color pattern.

REMARKS.—In Figure 2, a literature record for

E. yaeyamaensis is included, based on Aoyagi's (1954)

record from Iriomote Island, Ryukyus. For geographic variation see color pattern description

above.

NOMENCLATURE.—Aoyagi (1954) described Salarias yaeyamaensis on the basis of three specimens:

two males, 49.4 and 46 mm total length from

Iriomote Island, and a female, 49 mm total length

from Ishigaki Island. Both Islands are in the

southern Ryukyus. Aoyagi wrote that the description was based on the 49.4 mm male, but he also

described the female. A male, presumably the 49.4

mm specimen, was illustrated as was also the female.

No holotype was specifically designated. No indication of where the type material was deposited

was given, and I have been unable to locate Aoyagi's

specimens. The male, well illustrated, is clearly the

species that I treat here as E. yaeyamaensis. The

female, also well illustrated, is clearly the species

I describe below as Ecsenius oculus. (Aoyagi believed that color pattern differences between the

males and female were indicative of sexual dimorphism.) In order to fix the name of Aoyagi's

species, I here designate the larger male, 49.9 mm

total length, as lectotype of Salarias yaeyamaensis

Aoyagi.

MATERIAL (localities abbreviated).—South end of

Taiwan: USNM 203133 (35.2), 203134 (2: 38.5-

39.0), 203135 (42.0), 203136 (2: 39.7-42.6), 203137

(10: 26.5-49.0), 203138 (4: 37.7-41.8); Vietnam:

CAS GVF-2072 (38.8); Flores: ZMA 109.100 (33.4);

Banda Sea: USNM 202477 (26.6); Yap: CAS GVF894 (11: 21.9-34.3), GVF-794 (9: 26.1-51.9), GVF1912 (4: 28.7-34.8), GVF-1913 (25.0), GVF-1915

(36.7), GVF-1924 (31.7), GVF-1946 (37.7), GVF1923 (2: 34.5-39.9); Palaus: CAS GVF-735 (46.4),

GVF-740 (11: 31.9-43.0), GVF-802 (2: 24.7-29.8),

GVF-807 (39.9), GVF-843 (4: 30.0-39.0), GVF-1450

(6: 31.6-43.6), GVF-1868 (46.2), GVF-1869 (41.5);

Western Caroline Islands, Sorol Atoll: CAS GVF993 (31.6); Bougainville: USNM 201824 (2: 30.4-

39.8); New Britain: USNM 200428 (3: 38.2-38.7,

including one cleared and stained), 201812 (32.3);

New Hebrides: USNM 195787 (3: 20.6-26.0); One

Tree Island, Great Barrier Reef: USNM 201818

(29: 32.1-46.8), 201822 (15: 34.1-45.4), 201836 (4:

32.0-36.0); Endeavour Reef, Great Barrier Reef:

ANSP 109691 (31: 20.0-46.6), 109690 (52: 13.5-

37.8).

Ecsenius (Ecsenius) nalolo Smith

FIGURE SO

Ecsenius nalolo Smith, 1959, p. 245 [Pinda, Mozambique].

Ecsenius minutus Klausewitz, 1963, p. 357 [Addu Atoll, Maldive Islands].

DESCRIPTION.—Dorsal spines 11-13 (rarely 11 or

13); dorsal rays 12-15 (usually 13 or 14); segmented

anal rays 14-17 (rarely 14 or 17); pectoral rays 12-13

(rarely 12); segmented caudal rays 12-14 (rarely 12

or 14); dorsal procurrent caudal rays 6-9; ventral

procurrent caudal rays 5-9 (rarely 5 or 9); total

caudal elements 25-31; gill rakers 12-17 (rarely 12
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

or 17); pseudobranchial filaments 6-8; lower incisor

teeth 42-63; lower jaw posterior canines 0-1 (usually 1); total lower jaw posterior canines 0-2 (usually 2); upper incisor teeth 132 (one count); precaudal vertebrae 10; caudal vertebrae 21-23; total

vertebrae 31-33; epipleural ribs 11-14 (usually 12-

13). Lateral line without paired pores, extending

posteriorly to below level of 8th-11th dorsal spine

(rarely 8th). Dorsal fin notched seven-ninths to

eight-ninths length of first dorsal ray. Third (innermost) pelvic ray varying from obvious to not

obvious. One cirrus on each anterior nostril.

Color pattern (see also description under Ecsenius

yaeyamaensis): Description of the color pattern of

E. nalolo is difficult because of individual variation

and the complexity of the pattern itself. The most

consistent feature is the presence of a dark stripe

on the fleshy pectoral base that extends onto the

fin rays. Occasionally there are dark spots above

and/or below the stripe on the fleshly base and,

frequently, on the bases of the rays. The stripe is

never forked as in E. yaeyamaensis.

The head typically bears a dark stripe, varying

in intensity along its length, and extending from

the midposterior margin of the orbit across the

top of the opercle to the posterior bony margin

of the opercle. Above this stripe is a parallel narrow

pale stripe, then a stripe paler than the first described stripe, then a broad pale stripe and next a

thin, narrow, dusky stripe just lateral to the midline

of the top of the head. The last listed stripe may

intensify as a dark spot just anterior to the level

of the dorsal fin origin. Below the midorbital stripe

the cheek may be pale dusky or, in some non-Red

Sea males, peppered with tiny dark spots (Figure

30). The snout region is variably pale to dark dusky.

The underside of the head in males may be almost

black with a sharply defined, pale, triangular area,

apex directed posteriorly, just behind the lower lip.

In other males and females the underside of the

head is dusky with a similar pale triangular area

and a dorsally curving dark line on each side that

extends from behind the corner of the lower lip

up along the posterior margin of the opercle.

The dark stripes on the head may continue onto

the body anteriorly where they usually break up

into short, broad, dark dashes. Below these dashes

in the region above the venter, there are a variable

number of dark lines and splotches. The remainder

of the body is variably dusky with pale spots.

The pectoral fin and base were described above.

There is usually a dark or dusky spot in the pectoral

axil (also found in E. yaeyamaensis and to lesser

extent in E. oculus). The pelvics are pale to dark

dusky. The caudal is not noticeably marked, but

may be dusky basally. The dorsal fin is dusky

basally, mostly unmarked otherwise. The anal is

pale dusky in males, with the tips of the rays

noticeably paler in presumably mature males.

Females are paler overall than males and the

body of the female may be almost completely

unmarked.

GEOGRAPHIC VARIATION.—Mandibular tooth counts

(Table 17) of specimens from the northern Red

Sea (including the Gulf of Aqaba) are usually much

higher than those for specimens from the southern

Red Sea and Indian Ocean. (See also discussion of

this under E. oculus, geographic variation.)

REMARKS.—Specimens as small as 13.1 mm SL

were examined; they were not ophioblennius type

larvae, but were typical juveniles.

NOMENCLATURE.—Ecsenius minutus was not compared with any other species of fish. I have examined

the holotype and find it to be a young specimen of

E. nalolo.

MATERIAL (localities abbreviated).—Gulf of

Aqaba: USNM 204571 (39: 13.1-46.4), 204568 (20:

15.3-38.4), 204569 (61: 16.1-42.5), HUI E-63/36 (3:

18.8-25.1), HUJ-F-3531 (25.1), HUJ-F-4376 (6:

14.5-32.0), HUJ-F-3542 (31.6), E-63/33 (2: 26.2-

28.2), HUJ-F-3553 (3: 22.0-33.8), HUJ-F-4384

(35.9), HUJ-F-3551 (3: 17.5-34.8). Red Sea: southern tip of Sinai Peninsula, USNM 204570 (36: 21.8-

44.9); Egypt, NFIS 5119 (38.0), USNM 200602 (4:

30.3-45.0, including one cleared and stained),

201825 (28.1), 201754 (2: 32.3-43.1); Sudan, NFIS

(36.6); Zubair Island, HUI HUJ-F-4622 (25.2);

Ethiopia, USNM 204481 (9: 16.6-31.4). Aden:

USNM 203954 (16.8). Indian Ocean: Amirantes

Islands, USNM 201845 (7: 28.8-47.6), ANSP uncataloged (35.8); Farquhar Island, USNM 203955

(48.7); Aldabra Atoll, USNM 203072 (15: 29.9-

46.0), 203075 (11: 27.7-38.7), 201828 (6: 22.9-51.3);

Comoro Islands, USNM 201826 (2: 30.1-31.9),

201521 (31.4); Mozambique, RU BP2127C (39.2,

holotype of Ecsenius nalolo), BP2127B (34.3),

BP2127A (33.4), BP1924 (29.7), RU uncataloged

(5: 25.2-39.2, Pinda; 31.0, Ilha); Maldive Islands,

NFIS 6363 (20.0, holotype of Ecsenius minutus);
NUMBER 72

Chagos Archipelago, USNM 203071 (43.2), 203073

(2: 30.9-49.2).

Ecsenius (Ecsenius) oculus, new species

FIGURE 31

Salarias yaeyamaensis Aoyagi, 1954, p. 213 [in part, Ishigaki

Island].

DESCRipnoN.-Dorsal spines 12; dorsal rays 12-15

(usually 13-14); segmented anal rays 14-17 (usually

15-16); pectoral rays 13-14 (rarely 14); segmented

caudal rays 12-14 (rarely 12 or 14); dorsal procurrent caudal rays 7-10 (usually 8-9); ventral procurrent caudal rays 6-9 (usually 7-8); total caudal

elements 27-32 (usually 28-30); gill-rakers 12-18

(usually less than 16); pseudobranchial filaments

5-6; lower incisor teeth 47-64; lower jaw posterior

canines 0-1; total lower jaw posterior canines 1-2

(rarely 1); upper incisor teeth 138-148 (two counts);

precaudal vertebrae 10; caudal vertebrae 21-24

(usually 22-23); total vertebrae 31-34; (usually 32

or 33); epipleural ribs 11-14 (rarely 11). Lateral

line with no paired pores, extending to below level

of 10th spine to first dorsal ray (usually beyond 11th

spine). Dorsal fin notched nine-ninths length first

dorsal ray (one specimen of 46 had notch sevenninths). Third (innermost) pelvic ray obvious (not

obvious in one specimen only). One cirrus on each

anterior nostril, occasionally branched.

Color pattern: The color pattern of this species

is variable, but most specimens are characterized

by having at least three pairs of large, dark, pale

margined spots on each side of the body. The most

common pairs of spots are on or just above the

midline of the body. These spots frequently give

the appearance of sunglasses. Occasionally there

are similar pairs of smaller dark spots with pale

margins along the dorsal body contour in line

with the pairs lower on the sides; sometimes only

the dorsal pairs of spots are present. The general

body color is less dark than the spots and is punctuated with pale areas of various sizes and shapes.

There is invariably a vertical pair of dark marks

(sometimes fusing, but always recognizable) on the

caudal peduncle, extending onto the caudal fin

and there decreasing in intensity. The dorsal spot

of the pair extends diagonally downward on the

caudal fin, tapering posteriorly, and the ventral

spot extends horizontally, also tapering posteriorly.

35

The extensions frequently meet posteriorly. These

two spots are characteristic of the species. The

caudal is otherwise clear or variably pale dusky.

The head pattern is variable, but frequently

there is a middorsal pale stripe extending from

just anterior to the dorsal fin to between the eyes.

There is a broader dusky stripe (sometimes represented posteriorly only, as a dark spot) on either

side of the median pale stripe, followed laterally by

another pale stripe, then a broader dusky stripe,

another narrower pale stripe, and then a broader

dark stripe. This last stripe begins at about the

posterior midorbital level and extends posteriorly

onto the body for a short distance. Sometimes there

is another thin, pale stripe ventral to this last dark

stripe. The cheeks are irregularly dusky with pale

marks. The corners of the upper lip may be darker

than the surrounding color. Occasionally the ventral

margin of the opercle is dark, and this darkness

extends as a stripe onto the lower surface of the

head, meeting a similar mark from the other side.

The underside of the head is otherwise dusky

except for a pale unmarked area around the lower

lip.

The fleshy pectoral base is dusky with indications

of one, rarely two, darker dusky stripes, the more

dorsal of which may be bordered by pale areas.

The pectoral fin is mostly unmarked except for a

dusky area basally on the rays. The pelvic fins are

pale to dark dusky. The dorsal may bear a basal

or subbasal dusky stripe with the color mostly

restricted to the membrane or to the fin elements.

The anal fin is mostly uniformly dusky, paling at

the distal tips of the rays.

There are some striking variations from the

commonest type color pattern described above. It is

probable that some of these represent population

differences as they are in some degree paralleled

by meristic and tooth count differences (Table 18).

Three specimens (USNM 195716) collected by

Longley and presumably from the Banda Sea area

(see appendix) are somewhat faded, but the two

spots at the caudal peduncle extending onto the

caudal fin are still apparent, as are pairs of large

dark spots on the dorsal half of the body. These

specimens differ in having four dark spots in a

longitudinal series posteriorly on the ventral half

of the body. These spots are of about equal size

and intensity as those of the upper half of the body.
36

Four specimens from the New Hebrides (AMS

1-6521-24) are similar to those from the Banda Sea,

but exhibit about 6 to 8 dark spots on the ventral

half of the body. The dark spots on the dorsal half

of the body are not segregated as pairs but appear

as a somewhat irregularly linear series.

The single specimen available from the Palau

Islands (CAS GVF-880) differs from all others of

the species in having three large, dark, saddle-like

spots with pale ventral margins posteriorly on the

dorsal body contour. Each spot is separated from

the next by a slightly paler dark spot, which is

darker than the ground color of the body. The two

spots at the caudal base are relatively larger than

those for any other specimen.

A specimen from Hot du Sud, South China Sea

(CAS GVF-2113), is unique in having only a single

dark spot on the body, on the dorsal body contour

at the end of the dorsal fin. The body color is

generally dark, but paler than the spot, and is

punctuated with rows of pale, round or oblong

spots. The spots at the caudal base are normal.

The fleshy pectoral base bears a dark bar with a

shallow fork at its posterior end. The anterior end

grades into the dusky color of the pectoral base.

Above and below the bar are pale spots. The color

pattern is, in general, somewhat intermediate between that of typical E. oculus and E. yaeyamaensis.

Other specimens from the South China Sea are more

normal.

GEOGRAPHIC VARIATION.—In addition to color pattern variation (see above), there is some evidence

that populations from various localities differ in

meristics and tooth numbers (Table 18). Until more

specimens from more localities are available, it is

not possible to say definitely what the cause of this

variation is. There is a tendency for the meristic

characters to gradually decrease in numbers toward

the south to the equator and then to increase south

of the equator. Tooth numbers seem to exhibit a

sharp break near the northern end of the range of

the species.

A possible explanation for the observed meristic

variation may be the fact that water temperatures

tend to increase in a north-south direction to the

equator and decrease southward from the equator.

It has been demonstrated often in fishes that meristics decrease in numbers with increase in water

temperatures, but tooth numbers have not been

examined for such a correlation. In this respect

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the tooth numbers of Ecsenius nalolo specimens

from the northern Red Sea including the Gulf of

Aqaba) are higher than those of the southern Red

Sea and Indian Ocean specimens (Table 12). Gulf

of Aqaba water temperatures average lower than

those of the southern Red Sea (and Indian Ocean1)

during every month (Oren, 1962).

REMARKS.-Aoyagi (1954) illustrated a female E.

oculus from the Ryukyus that he identified as E.

yaeyamaensis, and Kamohara (1954) illustrated a

male, from Takarajima, northern Ryukyus. Kamohara's record is included on Figure 3 although the

specimen was not seen.

RELATIONSHIPS.—Ecsenius oculus is most closely

related to E. opsifrontalis, from which it is distinguished in usually having more dorsal and anal

rays (Table 1), usually shorter third and last dorsal

spines (Tables 5 and 7), and in overall color pattern

(compare Figures 31 and 32).

ETYMOLOGY.—The specific name oculus is Latin

for eye, and refers to the eyelike spots on the body

of the species.

HOLOTYPE.-USNM 203140, male, 53.8 mm SL,

immediately south of cut between large outstanding

rock and Ch'uan-fan-shih, south end of Taiwan.

Collector V. G. Springer, et al., 24 April 1968. Depth

approximately 6 meters. Original field number VGS

68-3.

PARATYPES.-USNM 203923 (7: 38.0-52.4), collected with the holotype. USNM 203142 (4: 43.9-

50.5,) just north of type locality. USNM 203141

(45.5), northwest of swimming beach of Sha Toa,

south end of Taiwan. USNM 203139 (3: 40.8-42.7),

just south of Chin-chiao-wan, south end of Taiwan.

BPBM 7464 (19.5), Taketoni Island, Ishigaki, Ryukyus. BPBM 7468 (13: 19.7-47.6), Ishigaki City

Harbor, Ishigaki, Ryukyus. CAS GVF-2113 (41.3),

west shore Hot du Sud, South China Sea. USNM

201811 (4: 26.2-32.5), Goh Huyong, Similand Island,

Thailand. USNM 201560 (2: 30.4), Pulo Jara, Strait

of Malacca, Indonesia. USNM 201522 (3: 31.8-

39.9), Pulo Mega, Mentawei Islands, Indonesia.

CAS GVF-880 (40.7), vicinity of Garudowaishi

Point, Nardueis Island, Palau Islands. USNM

195716 (3: 31.0-37.8), Banda Sea. AMS 1.6521-24

(4: 34.7-41.8), New Hebrides.

Much additional field data is available for the

specimens listed above from Taiwan and those

bearing California Academy of Sciences numbers.

The data are on my field data sheets for the year
NUMBER 72 37

1968 (in the files of the Division of Fishes, National

Museum of Natural History) and the George Vanderbilt Foundation register sheets at the California

Academy of Sciences.

Ecsenius (Ecsenius) opsifrontalis Chapman

and Schultz

FIGURE 32

Ecsenius opsifrontalis Chapman and Schultz, 1952, p. 521

[Rongelap Island, Marshall Islands].

DESCRIPTION.—Dorsal spines 12; dorsal rays 13-14

(usually 13); segmented anal rays 14-16 (usually 15);

pectoral rays 12-13 (rarely 12); segmented caudal

rays 13; dorsal procurrent caudal rays 8-9; ventral

procurrent caudal rays 7-8; total caudal elements

28-30; gill rakers 12-15; pseudobranchial filaments

5-6; lower incisor teeth 43-50; lower jaw posterior

canines 0-1; total lower jaw posterior canines 1-2;

upper incisor teeth 127 (one count); precaudal

vertebrae 9-10 (rarely 9); caudal vertebrae 21-22

(usually 22); total vertebrae 31-32 (usually 32);

epipleural ribs 12 (one count). Lateral line with

no paired pores, extending to below level of 11th12th dorsal spine (usually 12th). Dorsal fin notched

eight-ninths to nine-ninths length of first dorsal

ray. Third (innermost) pelvic ray not obvious. One

cirrus on each anterior nostril.

Color pattern: This is usually a faintly marked

species. The most noticeable and consistent marking

is a narrow stripe extending posteriorly from the

orbit across the top of the opercle and, broadening

and becoming fainter, along the upper portion of

the body, sometimes onto the bases of the caudal

rays. The body portion of the stripe is frequently

absent in part or completely. The second most

frequent and obvious marking is a dark edging of

the anterior margin of the lower lip. The head is

otherwise variably dusky, sometimes with one or

two dusky stripes connecting the orbits across the

anterior interorbital region. A second longitudinal

stripe occasionally occurs along the ventral half of

the body, extending onto the bases of the caudal

rays. Characteristic of many specimens are up to

10 faint to dark dusky, narrow, vertical stripes on

the upper two thirds of the body. Except for the

anal fin, which is dusky, and the body stripes

extending onto the base of the caudal, the fins are

essentially unmarked. In some specimens the black

lining of the intestine everts at the anus and gives

the appearance of a black ring in this region.

MATERiAL.-Guam: USNM 203742 (25.5); Kapingamarangi: CAS GVF-440 (23: 15.6-32.4), GVF-468

(5: 17.8-33.8), GVF-446 (3: 18.6-28.7), GVF-452 (6:

21.9-35.8); Marshall Islands: USNM 142065 (29.7,

holotype of Ecsenius opsifrontalis), 142066 (26.1),

152978 (34.8), 202548 (32.3).

Ecsenius (Ecsenius) prooculis Chapman and

Schultz

FIGURE 33

Ecsenius prooculis Chapman and Schultz, 1952, p. 519

[Munda Lagoon near Sassavelle, New Gorgia, Solomon

Islands].

DESCRIPTION.—Dorsal spines 12; dorsal rays 14;

segmented anal rays 15-16; pectoral rays 13; segmented caudal rays 13; dorsal procurrent caudal

rays 7; ventral procurrent caudal rays 6-8; total

caudal elements 26-28; gill-rakers 11-13; pseudobranchial filaments 6; lower incisor teeth 41-49;

lower jaw posterior canines 1; total lower jaw posterior canines 2; precaudal vertebrae 10; caudal

vertebrae 21-22; total vertebrae 31-32; epipleural

ribs 12 (one count). Lateral line with no paired

pores, extending posteriorly to below level of 8th9th dorsal spine. Dorsal fin notched eight-ninths

length of first dorsal ray. Third (innermost) pelvic

ray obvious. One cirrus on each anterior nostril.

Color pattern: This species is characterized by a

pattern of three dark stripes, separated by pale

dusky interspaces, extending the length of the body.

The dorsalmost stripe extends along the dorsal

body contour. The second and darkest stripe extends along the midlength of the body, and the

ventralmost, and least dark of the three, extends

along the ventral body contour. The dorsalmost

stripe gradually diminishes in intensity anteriorly

and fades into the head. The middle stripe extends

anteriorly onto the head (entering just below the

dorsal level of the opercle) as a narrow dark line

that reaches the posterior margin on the eye. The

ventralmost stripe broadens anteriorly and may

spread over much of the venter. All three stripes

extend onto the bases of the caudal rays where

the stripes fuse and form a crescentic dusky area.

Just above the head portion of the middle stripe

there may be an indication of a dark spot, which
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in one specimen is faintly continuous with a spot

on the dorsal margin of the opercle. The under

surface of the head is generally paler than the sides

of the head and there is faint indication in all

specimens of a dusky stripe extending a short distance posteriorly from the anteriormost mandibular

sensory pore on each side. The stripe on each side

shows indications of two concentrations of pigment,

which make the stripe appear as two diffusely

connected spots.

The holotype exhibits some dark spots on the

head not found in the other specimens. There is

a spot behind each anterior nostril and five spots

in the interorbital region: two anteriorly, two posteriorly, and one in the midline slightly closer to

the anterior pair.

The rayed portion of the dorsal fin bears a diffuse

dusky stripe in its lower third. Otherwise the fin

bears only some melanophores basally. The anal

fin is uniformly dusky, except for the tips of the

rays, which are pale. The pelvics are pale dusky and

the pectorals are clear except that the rays are

faintly overlain with melanophores.

REMARKS.—see under Ecsenius bandanus.

MATERiAL.-New Georgia: USNM 144722 (39.5,

holotype of Ecsenius prooculis); New Britain:

USNM 201819 (4:28.2-34.0).

Ecsenius (Ecsenius) bimaculatus, new species

FIGURE 34

DESCRIPTION (data for holotype followed in parentheses by data for largest and smallest paratypes, if

different).—Dorsal spines 12; dorsal rays 14; segmented anal rays 16 (16, 15); pectoral rays 13 (13,

12—but 13 on opposite side); segmented caudal rays

13; dorsal procurrent caudal rays 7 (7, -); ventral

procurrent rays 6 (6, -); total caudal elements 26

(26, -); gill-rakers 12 (14, 14); pseudobranchial filaments 6; lower incisor teeth 42 (42, 41); lower jaw

posterior canines 1; total lower jaw posterior canines

2; precaudal vertebrae 10; caudal vertebrae 22;

total vertebrae 32; epipleural ribs - (-, 12). Lateral

line with no paired pores, extending to below level

of 10th dorsal spine (9th, 10th). Dorsal fin notched

eight-ninths length of first dorsal ray. Third (innermost) pelvic ray obvious. One cirrus on each anterior

nostril.

Color pattern: The head is dusky above the level

of the upper third of the eye. Immediately below

this dusky area is a dark stripe that encompasses

the two circumorbital pores at the midposterior

orbital margin. This stripe extends posteriorly

across the head along the dorsal opercular margin

and enters the body, where it narrows and becomes

diffuse, ending below the third or fourth dorsal

spine. The head is less dark below the stripe than

above. One specimen exhibited a dark spot on

either side of the head at the dorsoposterior margin

of the eye just above the dark stripe. There are two

dusky spots on each side of the ventral surface of

the head. The anterior spot of each pair is just

behind the anteriormost pore of the mandibular

series, and the posterior is a little medial to the

fifth pore of this series.

The body is generally dusky, slightly darker

anteriorly above the level where the head stripe

enters the body. Just below the body portion of

the stripe, the body pales abruptly or gradually.

There are two distinct, dark, oblong spots on each

side just below the midlevel of the body under

the third or fourth through ninth dorsal spines.

There is a narrow dark line at the dorsal body

contour that continues on the base of the spinous

dorsal only. The rayed portion of the dorsal is

clear basally and has a diffuse stripe extending

along its length just above the clear area. The anal

fin is evenly dusky except for a paling at the distal

ends of the rays. The caudal is faintly sprinkled

with melanophores except for a faint crescentric

concentration of melanophores at its base. The

pectorals are marked similarly to the caudal; the

pelvics are unmarked.

RELATIONSHIPS.—See under Ecsenius bandanus.

MATERiAL.-Holotype, USNM 201817, male, 31.0

mm SL, Pulav Bohidulong, Darvel Bay, east end

of Borneo. Fringing coral reef on eastern (outer)

side of island; depth ca. 1.5-3.6 meters; 2 February

1965, Te Vega Cruise 6, station 216.

PARATYPES.-USNM 203922, two males, 31.7 and

30.6 mm SL collected with the holotype.

ETYMOLOGY.—The name bimaculatus refers to the

characteristic two dark spots on the side of the body.

Ecsenius (Ecsenius) bandanus, new species

FIGURE 35

DESCRIPTION (holotype and only known sped-
NUMBER 72 39

men).—Dorsal spines 12; dorsal rays 14; segmented

anal rays 16; pectoral rays 13; segmented caudal

rays 13; dorsal procurrent caudal rays 8; ventral

procurrent caudal rays 7; total caudal elements 28;

gill-rakers 13; pseudobranchial filaments 6; lower

incisor teeth 43; lower jaw posterior canines 1;

total lower jaw posterior canines 2; precaudal vertebrae 10; caudal vertebrae 21; total vertebrae 31.

Lateral line with no paired pores, extending posteriorly to below level of 9th dorsal spine. Dorsal

fin notched eight-ninths length of first dorsal ray.

Third (innermost) pelvic ray obvious. One cirrus

on each anterior nostril.

Color pattern: Distinctive markings appear only

on the head of this species, which has as its most

noticeable marking a broad stripe extending posteriorly from just behind the eye across the top of

the opercle. On the right side of the specimen there

is faint indication of a continuation of this stripe

on the body to just beyond the level of the pectoral

base. There is also faint evidence anteriorly that

the stripe continues dorsally following the margin

of the eye and meeting in the interorbital region

the stripe from the other side. There is a faint,

narrow band in the anterior interorbital region

connecting the two orbits, and a dark sprinkling

of melanophores around the anterior nostrils. On

each side of the undersurface of the head there is

a dusky spot encompassing the anteriormost two

pores of the mandibular series and a similar spot

just medial to the fifth pore of this series. The

rayed portion of the dorsal fin has a faintly dusky

stripe extending the length of the fin on its basal

half. The anal is dusky except for the unmarked

basal half. The specimen is otherwise evenly dusky.

RELATIONSHIPS AND REMARKS.—Ecsenius bandanus

is most and very closely related to E. bimaculatus

and E. prooculis: the prooculis species subgroup of

the yaeyamaensis species group. The species of this

subgroup differ from one another only in matters

of color pattern, and in this they are strikingly

different from one another (Figures 33-35). The

species are rare in collections and all known specimens are males. The species occupy different geographic areas (Figure 4).

Divergence of the species probably occurred as a

result of isolation in the different island groups,

either by differentiation of an ancestral founding

species that invaded new island areas, or after the

breaking up of the geographic range of a widespread

ancestral species. The distribution of the three

species correlates somewhat with biogeographic

areas recognized for terrestrial organisms. Thorne

(1963) prepared a biogeographical analysis of the

islands of the Pacific based on terrestrial organisms.

The tropical portion of the Pacific was called the

Oriental Region, which was divided into five subregions. Each subregion was divided into provinces

and some provinces were divided into districts. E.

bimaculatus occupies the Bornean District of the

Malayan Province of the Indo-Malayan Subregion.

E. bandanus occupies the Moluccan Province of the

Papuan Subregion, and E. prooculis occupies the

Bismarckian and Solomonian Districts of the Bismarckian Province of the Papuan Subregion. If

these distributions are indicative of distributional

patterns generally, then additional species in the

prooculis subgroup can be expected to occur in

other provinces and subregions.

HOLOTYPE.-USNM 195717, male, 31.4 mm SL,

Banda Sea, Indonesia (see discussion of Banda Sea

locality in Appendix).

ETYMOLOGY.—Named for the area from where the

holotype is believed to have come.

Ecsenius (Ecsenius) mandibularis McCulloch

FIGURE 36

Ecsenius mandibularis McCulloch, 1923, p. 122 [Masthead

Island, Australia].

DESCRIPTION.—Dorsal spines 12 (rarely 11 or 13);

dorsal rays 14-16 (usually 15); segmented anal rays

16-18 (rarely 18); pectoral rays 13 (rarely 12); segmented caudal rays 13; dorsal procurrent caudal

rays 6-7 (one of 53 with 8); ventral procurrent

caudal rays 5-7; total caudal elements 24-27; gillrakers 12-16 (usually 14); pseudobranchial filaments

6 (rarely 7); lower incisor teeth 45-53; lower jaw

posterior canines 2-8 (usually 5-7); total lower jaw

posterior canines 7-15; upper incisor teeth 120-121

(two counts); precaudal vertebrae 10 (one of 91

specimens with 11); caudal vertebrae 22-23 (rarely

21 or 24); total vertebrae 31-34 (usually 32-33);

epipleural ribs 12 or 13 (rarely 11 or 14). Lateral

line with no paired pores, extending posteriorly to

below 10th-11th dorsal spine. Dorsal fin notched

seven-ninths to eight-ninths length of first dorsal

ray (usually eight-ninths). Third (innermost) pelvic

ray obvious to not obvious (45 of 51 specimens with
40

ray obvious). One cirrus, rarely forked, on each

anterior nostril.

Color pattern: Specimens may range from almost

uniformly pale, with no distinct marks, to quite

dark anteriorly, grading into moderately dusky posteriorly with two longitudinal rows of dark spots

on the body. When the spots are present the upper

row usually begins between the dorsal fin origin

and the lateral line and terminates at some point

below the dorsal fin rays. The ventral row of spots

begins at about the midlateral line of the body no

farther anteriorly than the level of the anus and

extends to the caudal base. The upper row of spots

may be obscured by the dark color of the body.

I have seen collections from One Tree Island taken

within a week of each other in which almost all

the specimens in one collection exhibited spotting

and almost none of the specimens in the other

exhibited spotting. While not so restricted, the

spotting seems to be found predominantly in males.

In many specimens there is a narrow, dark stripe

extending from the posterior margin of the orbit

across the top of the opercle. There may also be a

narrow, dark stripe on each side of the underside

of the head, beginning somewhat behind the tip

of the lower jaw and extending to the margin of

the branchiostegal membrane, thence continuous

along the ventral margin of the fleshy pectoral base.

The anterior margin of the lower jaw may be dark

in pale as well as dark specimens. Some nonspotted

specimens exhibit adumbrations of several vertical

dusky bars on the upper half of the body.

The fins are not distinctively marked, usually

having an overall dusky appearance. Melanophores

usually concentrate along the fin elements. The

caudal is darker along its midlength; the tips of

the anal rays may be pale, especially in mature

males; there may be a dark crescentic area at the

base of the pectoral rays.

REMARKS.—At One Tree Island E. mandibularis

was collected with E. yaeyamaensis (also rarely with

E. bicolor). Superficially the two species are very

similar and difficult to tell apart in the field. They

can be separated readily, however, by the fact that

E. yaeyamaensis always has a dark forked stripe on

the fleshy pectoral base. This mark is not present

in E. mandibularis.

MATERIAL.—Australia, Queensland: Endeavour

Reef, ANSP 109694 (2:28.1-35.2), 109692 (30.3),

109696 (14:21.4-37.2), 109698 (20.6); Little Hope

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Island, ANSP 109693 (4:27.3-37.1), 109695 (18.6);

Big Hope Island, ANSP 109697 (29.8); Gillett Cay,

Swains Reef, AMS IB-6222 (4:28.5-35.7), IB-6237

(2:15.6-17.9); Masthead Island, Capricorn Group,

AMS 1-7112 (48.4, holotype of Ecsenius mandibularis, 1-7114-6 (6: 33.6-50.6); One Tree Island,

Capricorn Group, USNM 201367 (2 of 56: 31.5-

44.3, cleared and stained), 201821 (57:29.7-49.1);

Hoskyn Island, Bunker Group, AMS IA-3585 (35.0).

In addition, several series of specimens at USNM

from One Tree Island were cursorily examined in

preparing the color pattern description, and one

specimen from USNM 201820, also One Tree Island,

was used for preparation of figure 36).

ACKNOWLEDGMENTS.—For loans of specimens and

other favors aiding in the completion of this study

I wish to thank: F. H. Talbot and J. R. Paxton,

AMS; J. E. Bohlke and J. C. Tyler, ANSP; P. H.

Greenwood, A. C. Wheeler, N. B. Marshall, and

G. Palmer, BMNH; J. E. Randall, BPBM; W. N.

Eschmeyer and P. Sonoda, CAS; S. Rajagopalan

and R. S. Lai Mohan, FMRI; H. Steinitz, HUI;

K. Deckert and C. Karrer, ISZZ; E. Tortonese,

MCSN; M. L. Bauchot, MNHN: W. Klausewitz,

NFIS: M. Smith, RU; H. Nijssen, ZMA; I. Tomiyama, Amakusa Marine Biological Laboratory,

Japan; L. Fishelson, UTAI; W. Freihofer, Stanford

University.

Figures 12, 17, 20, 25, 28, and 29 were drawn by

Sharon L. Chambers; Figures 13, 18, 24, 26, 27, 32,

33, 35, and 36 were drawn by Martha H. Lester;

Figures 22 and 23 were drawn by Michelle R. Davis;

Figure 21 was drawn by Sandra Collum; Figure 31

was drawn by Ann Schreitz; Figure 30 was drawn

by Barbara Holden; Figures 14, 15, 18, 19, and 34

were drawn by Jack R. Schroeder. The graphs were

prepared by Richard Goodyear.

J. A. Peters, National Museum of Natural History,

was extremely helpful in advising me on computer

use and programing. W. F. Smith-Vaniz, University

of Miami Rosenstiel School of Marine and Atmospheric Sciences, and B. B. Collette, United States

Fish and Wildlife Service, called important references to my attention. E. A. Lachner, R. H. Gibbs,

Jr., S. H. Weitzman, and R. Goodyear, Division of

Fishes, National Museum of Natural History, offered critical comment on portions of the manuscript and useful advice during discussions of some

of the problems raised during my study.
NUMBER 72 41

Considerable financial and field support was provided by the staffs of the Department of Zoology

and Marine Laboratory of Hebrew University, under

a foreign currency grant, SFC-7-0062 (2), Drs. W.

Aron and H. Steinitz, principal investigators. Much

field support was provided by the staffs of the

United States Navy Medical Research Unit Number

2, Captain R. H. Watten, commanding, and the

Imperial Ethiopian Navy, H. I. H. Commodore

Alexander Desta, commanding.

This study was aided by a grant to me from the

Smithsonian Research Foundation (SG0661084).

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1949. Hawaiian Petrographic Province. Bulletin of the

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McCulloch, A. R.

1923. Fishes from Australia and Lord Howe Island. No. 2.

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Menard, H. W.

1964. Marine Geology of the Pacific. 271 pages.

Murray, J. A.

1887. New Species of Fish from Rurrachee and the Persian Gulf. Journal of the Bombay Natural History

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Oren, O. H.

1962. A Note on the Hydrography of the Gulf of Eylath.

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Randall, J. E. and H. A. Randall

1960. Examples of Mimicry and Protective Resemblance
42

in Tropical Marine Fishes. Bulletin of Marine

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Regan, C. T.

1905. On Fishes from the Persian Gulf, the Sea of Oman,

and Karachi, Collected by Mr. F. W. Townsend.

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1909. A Collection of Fishes Made by Dr. C. W. Andrews,

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Schultz, L. P.

1966. Pseudorhegma diagramma, a New Genus and Species

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Simpson, G. S., A. Roe and R. C. Lewontin

1960. Quantitative Zoology. Revised edition, Harcourt,

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Smith, J. L. B.

1959. Fishes of the Families Blenniidae and Salariidae of

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Smith-Vaniz, W. F. and V. G. Springer

In Synopsis of the Tribe Salariini, with Descriptions

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Starck, W. A., II

1969. Ecsenius (Anthiiblennius) midas, a New Subgenus

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Strasburg, D. W.

1956. Notes on the Blennioid Fishes of Hawaii with Descriptions of Two New Species. Pacific Science, 10(3):

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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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Appendix

NOTE ON BANDA SEA LOCALITIES.—After W. H.

Longley's death in 1937, the Carnegie Institution

of Washington presented his fish collections to the

Smithsonian Institution. The only localities associated with Longley's specimens were general ones

accompanying the containers holding his fishes

(Schultz, 1966, p. 194). In general, these localities

seem applicable, but it is known that some mixing

of specimens from different localities did occur.

During his lifetime Longley collected marine

fishes from only two general areas: Dry Tortugas,

Florida, and the western tropical Pacific Ocean.

Many of Longley's unlabeled specimens were obviously tropical western Atlantic species and a

number of these were saved and cataloged into the

collections of the Division of Fishes, National Museum of Natural History, as having come from

Tortugas. Some of this group of specimens, however, were discarded when they were found mixed

with obviously Pacific Ocean species.

The remainder of Longley's collections were

obviously tropical western Pacific in origin, and

all but a small lot were accorded the same general

locality by L. P. Schultz, formerly curator, Division

of Fishes, when they were entered into the collections: Dutch East Indies. The small lot (catalog

numbers 160665-160717) was catalogued with the

locality datum only of Hawaii. At least one species

in this lot, Enchelyurus brunneolus (Jenkins), is a

Hawaiian endemic

Prior to 1926, Longley (1918, 1920) made one or

two trips to the tropical western Pacific where he

spent time observing fishes at Pukoo, Molokai,

Hawaiian Islands, and Pago Pago, Samoa. There

is no information that he did any collecting on these
NUMBER 72 43

trips. During 1926-27 Longley (1927) made another

trip to the tropical western Pacific. On this trip he

spent two weeks at Pukoo, nine weeks at Pago Pago,

one week at Amboina, and eight weeks at Banda

Island (the last two localities are in the Banda Sea

and are about 250 km apart) collecting and making

field observations on the habits of fishes. He also

stopped in Sydney, but mentioned nothing about

his work there.

I have been unable to locate any of Longley's

specimens that bear Samoan locality data, although

on the basis of his handwritten field notes in the

files of the Division of Fishes, National Museum

of Natural History, he definitely made collections

of fishes in Samoa. Because some of the containers

holding Longley's fishes did have specimens from

obviously mixed localities, one cannot exclude the

possibility that the Dutch East Indies locality may

be erroneous for any particular specimen.

Longley appears to have made field notes on

almost every specimen he collected during his

1926-27 trip. On the basis of his color pattern

description of a blenny from Banda Island, I feel

fairly confident that the specimen I list in the

material of E. yaeyamaensis as having come from

the Banda Sea in fact came from Banda Island..

With regard to the specimens of E. bicolor, E.

oculus, and E. bandanus that I list as having come

from the Banda Sea, I am less certain, but I

believe that none of these came from the Hawaiian

Islands or Samoa.

Ecsenius bicolor has been reported from the

Hawaiian Islands only under unusual circumstances

(see remarks under E. bicolor species account) and

has never been found in Hawaii subsequent to the

original report despite considerable collecting in

that area in recent years. Samoa has been fairly

well collected since Longley was there and no

specimens of Ecsenius have been taken there.

Ecsenius generally does not occur much farther west

than 170° east longitude, and Samoa is located at

approximately 170° west longitude. It is possible

that the wide-ranging E. bicolor may turn up in

Samoa as it is the only species that comes close to

that island group. The Banda Sea, however, is well

within the range of E. bicolor, and E. oculus and

E. bandanus are most typical of species that seem

to be limited to west of 160° west longitude.

As Longley collected only at Amboina and Banda

in the Dutch East Indies, his Ecsenius specimens

from this area could have come from either island.

The two islands are close together and I have

elected to use the locality "Banda Sea" for his

Ecsenius specimens.

It is probable that proof of a Banda Sea (Amboina

or Banda) locality for Longley's Ecsenius specimens

will never be possible. Strong evidence favoring the

Banda Sea would be the re-collection of the species

described herein as Ecsenius bandanus, and also

the re-collection of the peculiar color pattern forms

of E. oculus and E. yaeyamaensis reported here as

occurring in the Banda Sea. Longley's specimens are

much too valuable to discard because of the locality

data problem. Aside from their intrinsic value as

specimens, I wish to use the present circumstances

as a test of the predictive value of systematic work.

TABLE I.—Frequency distributions for certain meristic characters in species of Ecsenius

Species Dorsal spines

10 11

1 10

2

2

1

2

12

261

3

4

2

251

23

13

4

11

208

127

48

41

5

3

1

on

13

11

5

58

77

45

4

4

14

1

5

6

1

Dorsal rays

12 13 14 15 16 17 18 19 20 21

Anal rays

13 14 15 16 17 18 19 20 21 22 23

midas

frontalis....

pulcher

grdvieri. . ..

aroni

bicolor

namiyei. . . .

lineatus . . . .

stigmatura. .

lividinalis...

yaeyamaensis

nalolo

oculus

opsifrontalis.

prooculis

bimaculatus.

bandanus. . .

mandibularis

10

11

1 26

2 12

36

1

186

103

31

6

5

3

1

21

2

24

1

3

65

4

54

2

3

3

45

22

164

10

2

108

11

38

26

32

1

3

6

96

38

1

9

5

59

16

15

1

3

n

8

3

1

1

17

30

12

37

1

1

182

93

32

4

4

2

1

1

1

12

2

3

9

3

33

2

97

1

44

11

163

4

2

1

118

3

36

36

59

1

8

5

45

21

3

2

12

4

1

37

11

35 56

(For sexual dimorphism and geographic variation see individual species accounts and section on sexual dimorphism.)
TABLE 2.—Frequency distributions for number of vertebrae in species of Ecsenius

Species Precaudal vertebrae

9 10 11 12

Caudal vertebrae

19 20 21 22 23 24 25 26 27 28 29

Total vertebrae

29 30 31 32 33 34 35 36 37 38 39 40

midas

frontalis... .

pulcfur

gravieri. . ..

aroni

bicolor

namiyei

lineatus.

stigmatura. .

lividinalis

yaeyamaensis.

nalolo

oculus

opsifrontalis..

prooculus

bimaculatus. .

bandanus . . . .

mandibularis.

12

1 282

44

2 83

1 47

237

21

11

4

8

202

124

37

34

5

3

1

90

10

17

4

3

2

1

1

179

94

23

33

3

3

34

2

1

9

2

15

10

19

49

80

5

9

145

3

2

1

6

158

49

34

82

4

9

34

4

31

4

13

1

30 8

10

17

4

4

2

179

94

23

32

3

3

9

2

15

10

19

1

1

145

1

2

1

94

6

9

82

1

11

169

49

35

15

1

32

4

29

3

9

30 8

33 50

(For sexual dimorphitm and geographic variation see individual species accounts and section on sexual dimorphism.)
TABLE $.—Frequency distributions for certain characters in species of Ecsenius

Species

midas

Jrontalis

pulcher

gravieri

oront

bicolor

namiyei

lineatus

stigmatura

lividinalis

yaeyamaensis

nalolo

oculus

opsifrontalis

prooculis

bimaculatus

bandanas

mandibularis

Pectoral rays

12 13 14 15 16

1 11

15 236 27

60 4

1 62 6

1 39 3

4 231 4

1 23 1

11 1

1 2 1

10 1

4 119 3

2 67

44 1

2 39

5

1 2

1

2 65

Segmented caudal rays in

specimens over 26 mm SL

12 13 14 15

1 7 4

190 6

31 2

51

29

2 268

3 22

12

4

2 5 1

113 1 1

1 84 1

1 40 4

25

5

3

1

87

Total caudal rays

23 24 25 26 27 28 29 30 31 32 33 34

1 1 7 3

7 21 116 37 26 2

2 7 9 7

5 13 47 5 4

1 5 12 13 10 3

2 21 29 100 13 4

1 4 8 9 2

1-9 2

1 2

2-5 2

2 8 36 18 13 1

4 4 9 9 24 5 1

2 12 8 11 2 1

3 4 3

1 2 1

2

1

6 23 26 11
TABLE 4.—Frequency distributions for number of dentary incisor teeth (including anterior canines) in species of Ecsenius

Species Number of teeth

13 14 15 16 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64

midas

frontalis

(see also table 12)

pulcher

gravieri

aroni

bicolor

male

female

namiyei

lineatus

stigmatura

lividinalis

yaeyamaensis

male

female

nalolo

(see also table 17)

oculus

(see also table 18)

opsifrontalis

prooculis

bimaculatus

bandanas

mandibularis

2-2 8

1 1 1 4 9 22 40 39 51 28 32 9 12 3

237 9 10 3 8 11 2 1

2 - 3 8 10 11 14 13 3 4 1

1-- 1 1289844 1

1 -

1 2 4 13 53 32 26 3 3 1

1 _____ _ 2 13 22 36 15 14 1 1

1111 2 3 5 4 3 3 1

4 1 2 5 1

12- 1

1 1 4 3- 1

1 2 13 10 12 18 12 2 3 1

1--2--6 6 14 8 11642 1

1 1 2 4 4 9 15 6 8 10 11 14 16 16 12 13 12 11 7 5 3 2

1113-234333544231 1

1 2

1 3 6 3 15 9 - 3

2 - - 1 - - 1

4 4 7 17 18 10 6 2 1
G

1

TABLE 5.—Frequency distributions for depth of dorsal fin notch (to nearest ninth of length of first dorsal ray) and length of last dorsal

spine as percent standard length in species of Ecsenius

Species

Notch depth

0123456 7 8 9

Last dorsal spine length (lower class limits)

0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0 10.0 11.0 12.0 13.0 14.0 15.0 16.0 17.0 18.0 19.0 20.0

midas

frontalis...

pulcker... .

gravieri. . .

aroni

bicolor. . . .

namiyei. . .

Uneatus. . .

stigmatura.

lividinalis.

12

277

24

- 1

14 18 5 1

2 2 17 33 7

1 24 1

2 1 9 42 45 13

6 26

2 17 9

6 15 20

1

1 1

9 4 1

18 25 21 9 15 10

2 15 37 12 2

2 1 9 8

4 3 1

yaeyamaensxs. •

nalolo

oculus

opsifrontalis

prooculis

bimaculatus

bandanus....

mandibularis. . ..

3 10

1 3

7 101

3 14 40 19

1 - 45

4 18

5

1 2

1

7 55

7

2

2 27 64

1 26 42

21 13 -

4 16 3

1 3 1

3

1

34 22

1-1- 1
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 6.—Frequency distributions for certain characters in species of Ecsenius

Species

Lateral line extends posteriorly to below

level of dorsal Jin element

7 8 9 10 11 12 13

Pseudobranchial filaments

5 6 7 9 10 11 12

midas

frontalis... .

pulcher

gravieri. . ..

aroni

bicolor

namiyei. . ..

liruatus. . . .

stigmatura. .

lividinalis...

yaeyamaensis

nalolo

oculus

opsifrontalis.

prooculis....

bimaculatus.

bandanas. . .

mandibularis

8

3

4

3

7

1

1

1

2

2

36

15

41

13

3

3

-

9

3

1

161

35

30

25

51

1

13

5

74

66

2

3

25

4

64

5

2

34

27

3

3

1

1

46

16

1

1

37

24

1

1

5

1 28 12

5

4

1

2

11

5

1

81

28

10

7

163

3

10

4

7

77

21

17

16

5

3

1

45

-

3

8

4

3

26

18

1

1

38

24

1

2

1

1

2

3

1

3
NUMBER 72 51

TABLE 1—Frequency distributions of third dorsal spine length as percent standard length in species of

Ecsenius

Species

Percent SL (lower class limits)

7.0 8.0 9.0 10.0 11.0 12.0 13.0 14.0 15.0 16.0 17.0 18.0 19.0 20.0 21.0 22.0

midas

Males...

Females.

frontalis

Males...

Females.

pulcher

Males...

Females.

gratneri

Males...

Females.

aroni

Males...

Females.

bicolor

Males...

Females.

namiyei

Males...

Females.

lineatus

Males...

Females.

stigmatura

Females.

lividinalis

Males...

Females.

yaeyamaensts

Males...

Females.

nalolo

Males...

Females.

ocutus

Males...

Females.

opsifrontalis

Males...

Females.

prooculis

Males...

bimaculatus

Males...

bandanas

Males...

mandibularis

Males...

Females.

2

1

1

2

1

-

1

2

2

1

1

1

1

1

-

-

1

3

1

1

1

14

8

4

-

1

4

2

_

3

-

1

13

10

7

6

5

8

2

23

8

3

3

_

3

17

10

2

1

3

3

1

9

9

3

2

12

7

5

5

16

in 5

2

5

2

25

22

5

4

7

5

2

13

7

-

11

3

7

8

1

1

1

1

1

11

5

10

16

6

2

5

1

3

6

9

-

4

2

1

1

2

1

2

13

3

2

2

4

3

1

4

5

3

1

_

1

2

3

1

2

1

6

3

3

_

2

2

2

1

1

1

1

2

1
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 8.—Frequency distributions of fifth dorsal ray length as percent standard length in species of Ecsenius

Species

Percent SL (lower class limits)

10.0 11.0 12.0 13.0 14.0 15.0 16.0 17.0 18.0 19.0 20.0 21.0 22.0

midas

Males

Females

frontalis

Males

Females

pulcher

Males

Females

gravteri

Males

Females

aroni

Males

Females

bicolor non-Australian

Males

Females

bicolor Australian

Males

Females

namiyei

Males

Females

stigmatura

Males

Females

lividinalis

Males

Females

yaeyamaensis

Males

Females

nalolo

Males

Females

oculus

Males

Females

opsifrontalis

Males

Females

prooculis

Males

bimaculatus

Males

bandanas

Males

mandibularis

Males

Females

10

4

9

19

13

6

1

3

5

2

26

14

1

10

4

1

1

3

17

14

1

13

8

10

7

2

2 1

5 5

2 1
NUMBER 72 53

TABLE 9.—Frequency distributions of length of longest caudal ray as percent standard length in species of

Ecsenius

Species

midas

Males

Females

frontalis

Males

Females

ptdcher

Males

Females

gravieri

Males .. .

Females

aroni

Males... .

Females

bicolor

Males

Females

namiyei

Males

Females

lineatus

Males

Females..

stigmatura

Males

Females..

lividinalis

Males

Females..

yaeyamaensis

Males

Females

nalolo

Males

Females

oculus

Males

Females

opsifrontalis

Males

Females

prooculis

Males

bimaculatus

Males

bandanas

Males

mandibularis

Males

Females

18.0 20.0

4

10

1

1 4

1

1

4

3

2 3

2

1

1

22.0

13

49

2

3

2

4

2

2

17

2

2

1

8

29

2

11

?,

12

1

5

8

24.0

1

26

32

2

1

9

8

5

11

4

3

1

1

26

11

14

n

s

3

•5

1

1

6

26.0

2

2

13

5

1

4

1

2

6

1

4

1

2

4

4

6

7

2

1

2

1

28.0

2

12

2

2

4

2

1

7

-\

1

1

3

2

2

1

4

Percent SL (lower class limits)

30.0 32.0 34.0 36.0 38.0 40.0

11

1

2

?

1

4

?

?

1

4

7.

1

3

3

1

7

4

3

7

3

3

3

1

1

6

1

5

4

2

6

1

1

1

1

3

1

3

3

10

_

2

1

2

2

4

4

?.

6

1

1

1

6

3

-

8

2

42.0

4

3

1

2

2

44.0 46.0 48.0 50.0 52.0

4 - 1

2

1

2 1 3 3 3
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 10.-Frequency distributions of nasal cirrus length as percent standard length in species of Ecsenius

Species

midas

Males

Females.. .

frontalis

Males

FrmaVs,, ,

(ndcktr

Males

Females

gravtert

Males

Females

ofont

Males

Females... .

bicolor non-Australian

Males

Females

bicolor Australian

Males

Females

namiyei

Males

Females.. .

lineatus

Males

Females

sttgmatura

Males

Females... .

lividinalis

Males

Females.

yaeyamaensis

Males

Females

nalolo

Males

Females

oculus

Males

Females...

opsifrontalis

Males

Females.

prooculis

Males

bimaculatus

Males

bandanus

Males

mandibularis

Males...

Females

0.0

1

1

1

3

1.0

1

1

1

2

1

27

29

2

5

5

2

3

13

2.0

1

6

7

9

10

1

2

1

32

17

17

24

4

8

2

1

1

16

1

3.0

1

2

1

1

1

3

7

7

4

3

2

9

4

1

to to

17

2

4

6

1

6

6

1

4.0

1

1

7

2

4

3

14

7

7

5

1

6

3

1

2

4

8

1

4

3

Percent SL {lower class limits)

5.0 6.0 7.0 8.0 9.0 10.0 11.0 12.0 13.0

3

1

15

17

4

3

3

20

9

to to

3

4

1

2

2

1

1

1

15

29

1

2

9

5

9

2

1

1

2

1

21

14

1

3

1

5

5

10 3 6 2 2 1

11 6 6 5 1

1

1

1

4 1

(In E. fttcofor and E. namiyei, which have two cirri on each nostril, the posterior cirrus was measured.)
TABLE 11.—Frequency distributions for certain characters of the three color pattern forms of Ecsenius frontalis

Locality and form

Dorsal spines

11 12 13

Dorsal rays

17 18 19 20 21

Anal rays

18 19 20 21 22

Precaudal vertebrae

10 11 12

Caudal vertebrae

23 24 25 26

Pectoral rays

14 15 16

Israel

frontalis form

males

females

albicaudatus form

males

females

Sudan

frontalis form

males

females... .

Ethiopia

frontalis form

males

females

albicaudatus form

males

females

nigrovittatus form

males

females

Gulf of Aden

frontalis form

males

females....

4 36

1 28

2 26

2 25

40

1 45

2

4

25

1 24

4 18 17 2

15 14

11 16

3 15 11 1

1

1 1

2

1

22

36

1

4

18

19

16

10

1

6

8

1 1

1

3 21 14

18 10

14 10

1 19 10

2

3

1

1

2

22

32

1

3

18

17

16

11

1

5

7

1 37

29

27

31

40

1 45

2

4

25

26

4 24 10

23 6

21

1 23

2 30

2 24

21

26

1

1

16

30

3

12

14

24

16

2

1

12

10 2

3

1

1

2

4

35

45

1

3

22

23

2

1

Dau from Egyptian and Gulf of Suez localities are combined with dau from Israel; dau from all southern Red Sea localities are combined with dau from

Ethiopia.
TABLE

Locality and form

Israel

frontalis form

males

females

albicaudatus form

males

females

Sudan

frontalis form

males .

females . .. .

Ethiopia

frontalis form

males

females

albicaudatus form

males

females

nigrovittatus form

males

females

Gulf of Aden

frontalis form

males

females

12.—Frequency distributions for

25

1

1

1

1

1

1

1

Total caudal fin rays

(procurrent and principal)

26 27 28 29

4

1

1

3

6

2

1

10

10

7

10

1

25

23

1

1

12

13

1

1

1

3

2

1

7

9

1

8

3

2

2

2

3

3

6

1

1

3

3

30

1

1

certain characters o F the threecolor pattern forms

Dentary incisor teeth (including anterior canines)

40 41 42 43 44 45 46 47 48 49 50 51 52

1

1-11 3

1 4

1-1 1

1

2

1

1

6 12

8 12

1

3 5

4 5

1 1

?

5

1

6

4

7

7

S

1

4

3

4

8

1

7

10

1

1

9

ft

7

1

?

?

s

9 3 5

6 4 3

8

7 2 4

1

1

of Ecsenius frontalu

53

1

2

Lateral line terminates

below level of dorsal spine

8 9 10 11 12

2 6

7

3

6

1

2

1 1

1

7

2

77

18

7

17.

1

27

34

1

4

17

15

2

1

1

1

2

1

6

8

1 1

5

Data from Egyptian and Gulf of Suez localities are combined with data from Israel; data from all southern Red Sea localities are combined with data from

Ethiopia.
TABLE 13.—Frequency distributions for measurements of certain fin elements of the three color pattern forms of Ecsenius frontalis

(see also figure 6)

•z

c

I

Locality and form

Third dorsal spine length as percent SL

Lower class limits

10.0 11.0 12.0 13.0 14.0 15.0 16.0 17.0

Last dorsal spine length as percent SL

Lower class limits

10.0 11.0 12.0 13.0 14.0 15.0 16.0 17.0 18.0 19.0 20.0 12.0 13.0 14.0

Fifth dorsal ray length as percent SL

Lower class limits

\.O 15.0 16.0 17.0 18.0 19.0 20.0 21.0 22.0

Israel

frontalis form

males

females

albicaudatus form

males

females

4 1

2 -

2 1

1 4 1

1

2

_

5

3

1

5

5

5

4

1

3

1

-

1

1

1

1

1

1

1

2

1

2

1

3

3

1

6

5

3

1

3

-

1

6

1

1

_

_

1

_

_

2

Sudan

frontalis form

males

females... .

Ethiopia

frontalis form

males

females

albicaudatus form

males

females

nigrovittatus form

males

females

3 3 4

3 2 2

1 1

1

3 5

7 7

6

3 8 1

3 1 2

1

- 1

5 4

3

1 - 1

1 3

2 2 4 6

6 4 1

1 - - 1

2 3 3 4

3-3 1

Data from Egyptian and Gulf of Suez localities are combined with data from Israel; data from all southern Red Sea localities are combined with data from

Ethiopia.
TABLE 14.—Frequency distributions for measurements of nasal cirrus and longest caudal ray of the three color pattern forms of Ecsenius

frontalis (see also figures 7-10)

Locality and form

Nasal cirrus length as percent SL

Lower class limits

1 . 0 2. 0 3. 0 4. 0 5. 0 6. 0 7. 0 8. 0 9. 0 10. 0 11. 0 12. 0 13. 0

Longest caudal ray length as percent SL

Lower class limits

20.0 22.0 24.0 26.0 28.0 30.0 32.0 34.0 36.0

Israel

frontalis form

males

females

albicaudatus form

males

females

Sudan

frontalis form

males

females....

Ethiopia

frontalis form

males

females

albicaudatus form

males

females

nigrovittatus form

males

females

1

5

_

6

5

1

1

3

2

1

1

3

3

1

6

-

2

1

3

3

1

3

4

2

1

_

4

1

1

1

1 1

2

10

1

9

1 1

4

2

2

7

5

7

7

7

14

2

1

7

10

6

1

7

4

4

1

1

1

1

1

1

1

3

5

1

1

8

15

3

2

11

13

17

1

3

2

6

2

_

1

_

Data from Egyptian and Gulf of Suez localities are combined with data from Israel; data from all southern Red Sea localities are combined with data from

Ethiopia.
NUMBER 72 59

TABLE 15.—Sexual dimorphism and geographic variation in frequencies of some meristic characters as indicated by specimens of Ecsenius bicolor from selected localities

Locality

Australia

Males

Females ... .

New Hebrides

Males

Females.. . .. .

Kapingamarangi

Males

Females

Ponape

Males

Females

Marshall Islands

Males

Females

South China Sea

Males

Females.

Malacca Straits

Males

Females

Christmas Island (Indian

Ocean)

Females

15

1

1

1

1

16

2

1

4

4

9

0

?

9

1

?

Dorsal rays

17 18

21

31

4

1

21

8

8

4

24

2

1

2

5

20

2

1

1

5

1

Av.

17.5

17.0

17.0

16 2

16.9

16.5

17.0

16.3

16.8

16.1

17.8

17.3

15.5

15.7

17.0

17 18

1

4

11

1 1

6

1

Anal rays

19 20 21

12

28

4

5

23

13

6

2

7?

6

7

7.

7

4

28 1

6

2

3

2

2

5

3

1

Av.

19.7

19.1

19.3

19.0

19.1

18.8

19.2

18.2

19.0

18.5

19.7

20.0

19.0

18.7

19.2

Caudal vertebrae

23 24 25 Av.

7

20

2

1 4

16

16

7

6 7

15

10

2

1

2

3

3

28

13

4

8

1

10

1

4

2

1

24.8

24.4

24.7

23 8

24.3

24.0

24.1

23.5

24.4

24.1

24.7

24.7

24.0

24.0

24.2
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 16.—Sexual dimorphism in frequencies of some meristic characters as indicated by specimens of

Ecsenius namiyei from Taiwan and Escenius pulcher

Species

namiyei

Males.. .

pulcher

Males

Females

18

3

7

Dorsal rays

19 20

1

7

17

17

13

1

14

2

Av.

19.9

19.1

19.3

18 S

19

1

20

1

Anal

21

4

8

11

9

rays

22

10

?1

14

23

1

Av.

21.7

21.0

21.6

21.5

25

1

1

Caudal vertebrae

26 27 28

5

7

1

3

8

21

9

S

3

Av.

26.5

25.9

27.1

27.0

TABLE 17.—Frequency distributions for numbers of dentary incisor teeth (including anterior canines) in

Ecsenius nalolo

Locality

Northern Red Sea and

Gulf of Aqaba

Males

Females

Middle Red Sea

Females

Southern Red Sea

Males

Females

Indian Ocean

Males

Females

Number of teeth

42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63

6 8 14 15 13 4 2 2

3 1 14979753 2

1-221- 2

1 1

1122367343 1

111 4 1 4 4 4 3 1
TABLE 18.—Variation in meristic characters and number of dentary incisor teeth (including anterior canines) in populations oi Ecsenius

oculus

Locality

Ryukyu s

Taiwa n

H o t d u Sud. .

Thailand... .

Palau

Mentawe i Island... .

Strait of Malacca... .

Band a Sea. .

N e w Hebride s ... .

12

1

1

Dorsal rays

13 14

3

3

1

3

1

1

12

12

1

1

1

4

15

2

1

Anal rays

14 15 16 17

3

3

1

3

1 1

1

11 3

13

1

1

2

4

Caudal vertebrae

21 22 23 24

1

1

1

1

4

11

3

1

2

1

1

9 1

3

1

2

4

47 48 49 50 51

1 - 1 -

1 -

1 - 1

1 -

52

1

1

-

53

-

1

2

Dentary incisor teeth

54 55 56 57

1

1

1

1

1 3 3

2

58

CO CM

59

CM CM <-

60

1

3

61

1

1

62 63 64

2 1 1

1

Localities arranged from north to south. Dashed line separates localities north of equator from those south of equator.

FIGURES 12-36

383-026 O - 70 - 5

NUMBER 72 65

FIGURE 12.—Ecsenius midas, ANSP 111148, holotype, male, 74.4 mm SL,

D'Arros Island, Amirante Islands, Seychelles. Insert presents ventral

aspect of region around anus.

FIGURE 13.—Ecsenius frontalis, USNM 200613, male, 58.6 mm SL,

frontalis color pattern form, Strait of Jubal, Egypt, Red Sea.

FIGURE H.-Ecsenius frontalis, USNM 204650. female. 37.3 mm SL, albicaudatus

color pattern form, El Himeira, Sinai Peninsula, Gulf of Aqaba.
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

10

FIGURE 15.—Ecsenius frontalis, USNM 204490, male, 38.7 mm SL,

nigrovittatus color pattern form. Sheikh el Abu, Ethiopia, Red Sea.

l cm

FIGURE 16.—Ecsenius pulcher, USNM 201571, male, 32.9 mm SL,

banded form, Tarut Bay, Ras Tanura, Persian Gulf.

cm FIGURE 17.—Ecsenius gravieri, HUI HUJ-F-4382, male, 38.6 mm SL,

pale form, Dahab, Sinai Peninsula, Gulf of Aqaba.
NUMBER 72 67

10

FIGURE 18.—Ecsenius gravieri, USNM 204480, female, 40.1 mm SL,

dark form, Sheikh el Abu, Ethiopia, Red Sea.

FIGURE 19.—Ecsenius aroni, USNM 204468, male. 36.8 mm SL, El Himeira,

Sinai Peninsula, Gulf of Aqaba.

FIGURE 20— Ecsenius bicolor, USNM 201368, male, 66.3 mm SL,

One Tree Island, Queensland, Australia.
68

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

l c m FIGURE 21.—Ecsenius bicolor, NFIS 9578, female, 49.1 mm SL,

Madewaru Island, Fadiffulu Atoll, Maldives.

icm ,

FIGURE 22.—Ecsenius namiyei, USNM 203125, female,

63.6 mm SL, north end of Taiwan.

icm FIGURE 23.—Ecsenius lineatus, USNM 203129, male, 48.2 mm SL,

just SW of Yeh-Liu, north end of Taiwan.
NUMBER 72

69

1 cm i i

5 mm

FIGURE 24.—Ecsemus stigmatura, USNM 111878, female, 36.6 mm SL, Tomahu Island, Indonesia.

Insert presents ventral aspect of region about anus.

icm T

FIGURE 2S.-Ecsenius stigmatura, USNM 201815, male, 37.7 mm SL, Pulav Gaya, Darvel Bay,

Borneo. Insert presents ventral aspect of region about anus.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 26.—Ecsenius lividinalis, USNM 144292, male, 25.8 mm SL, Munda Lagoon, New Georgia,

Solomon Islands. Insert presents ventral aspect of region about anus.

l cm

FIGURE 27.—Ecsenius yaeyamaensis, USNM 203136, female, 42.6 mm SL,

typical color form Ch'uan-fan-shih, south end of Taiwan.

FIGURE 28.—Ecsenius yaeyamaensis, USNM 201818, male, 38.1 mm

SL, spotted form, One Tree Island, Queensland, Australia.
NUMBER 72

71

cm

FIGURE 29.—Ecsenius yaeyamaensis, USNM 202477, male, 26.6 mm SL,

striped form, Banda Sea, Indonesia.

FIGURE SO.—Ecsenius nalolo, USNM 203072, male 43.3 mm SL, Aldabra Atoll, Indian Ocean

(penultimate dorsal spine abnormally reduced). Insert presents ventral aspect of head.

-
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

i cm

FIGURE 31.—Ecsenius oculus, USNM 201522, male, 39.9 mm SL, Pulo Mega, Mentawei Islands,

Indonesia (forked nasal cirrus is atypical).

5 mm

FIGURE 32.—Ecsenius opsifrontalis, CAS GVF-468, male, 325 mm SL, Kapingamarangi Atoll.

Insert presents anterior view of head.
NUMBER 72 73

l cm

FIGURE S3.—Ecsenius prooculis, USNM 201819, male, 305 mm SL, Simpson Harbour, Rabaul, New

Britain. Insert presents enlarged view of two of the anal rays exhibiting fleshy envelopment of

the ray tips, a common characteristic of presumably mature males of most species of Ecsenius.

\0

FIGURE 34.—Ecsenius bimaculatus, USNM 203922, male, 31.7 mm

SL, Pulav Bohidulong, Darvel Bay, Borneo. Insert presents region

of body hidden by pectoral fin.
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

/ /

cm

FIGURE 35.—Ecsenius bandanus, USNM 195717, holotype, male, 31.4 mm SL,

Banda Sea, Indonesia.

L_^^_J

FIGURE 36.—Ecsenius mandibularis, USNM 201820, male, 51.1 mm SL, One Tree Island,

Queensland, Australia.

U.S. GOVERNMENT PRINTING OFFICE : 1970 O - 383-026
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