WILLIAM F. SMITH-VAN

and VICTOR G. SPRINGF

Synopsis of the

Tribe Salariini,

with Description of

Five New Genera and

Three New Species

Pisces: Blenniidae)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1971 NUMBER 73
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 73

William F. Smith-Vaniz SynopSIS of the

and Victor G. Springer r-pi .1 ^ -, •• .

lnbe oalanini,

with Description of

Five New Genera and

Three New Species

(Pisces: Blenniidae)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON
ABSTRACT

Smith-Vaniz, William F., and Victor G. Springer. Synopsis of the Tribe Salariini,

with Description of Five New Genera and Three New Species (Pisces: Blenniidae). Smithsonian Contributions to Zoology, 73:1-72, 1971.—The blenniid

tribe Salariini is characterized and a key is given to twenty-four genera we

recognized in the tribe. Five of these genera (Alloblennius, Crossosalarias,

Litobranchus, Mimoblennius, Nannosalarias) are here described as new. All

genera are diagnosed and comments are given on their relationships, nomenclature, distribution, and estimated number of species. One genus, Medusablennius, previously placed in the tribe Blenniini is transferred to the Salariini. The

approximately 350 nominal species of the Salariini are listed and each name is

provided with a generic allocation. The institutional depository of primary

type material, if known, is cited for each species. Some annotation to this list

is made where there are involved nomenclatural problems. Three new species

are described: Alticus simplicirrus from the Marquesas Islands; Crossosalarias

macrospilus from the Solomon, Tonga, and Palau Islands, Great Barrier Reef,

and South China Sea; and Mimoblennius cirrosus from the Gulf of Aqaba and

Red Sea. Alloblennius jugularis (Klunzinger), A. pictus (Lotan), Litobranchus

fowleri (Herre), Mimoblennius atrocinctus (Regan), and Nannosalarias nativitatus (Regan) are redescribed. All described and redescribed species and genera

are figured, and illustrations of characteristic cephalic pore patterns are given

for all genera and subgenera.

Alloblennius pictus, one species treated, exhibited statistically significant differences in mean numbers of dorsal and anal fin rays and caudal vertebrae between

specimens from the Gulf of Aqaba and the southern Red Sea. Mean numbers

of these meristic characters were significantly different for males and females of

the southern Red Sea specimens, but were not significantly different for the

Gulf of Aqaba specimens.

Contribution number 1254, University of Miami, Institute of Marine and

.,, Atmospheric Sciences

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

in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1971

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

Washington, D.C. 20402 - Price $1.25 (paper cover)
Contents

Page

Introduction 1

Acknowledgments 1

Methods and Materials 2

Tribe Salariini 4

Nominal Genera and Subgenera in Salariini 6

Key to Genera 8

Synopsis of Genera

A lloblennius 10

Alticus 14

Andamia 17

Antennablennius 18

A trosalarias 18

Cirripectes 19

Crossosalarias 20

Ecsenius 22

Entomacrodus 23

Exallias 23

Glyptoparus 24

Hirculops 24

Istiblennius 25

Litobranchus 26

Medusablennius 28

Mimoblennius 29

Nannosalarias 32

Ophioblennius 35

Pereulixia 36

Praealticus 36

Rhabdoblennius 37

Salarias 38

Scartichthys 39

Stanulus 40

Comparison of Cephalic Sensory Pores 40

Recognition of Genera 49

Annotated List of Nominal Species 52

Tables of Generic Characters 53

Literature Cited 61

William F. Smith-Vaniz

and Victor G. Springer

Synopsis of the

Tribe Salariini,

with Description of

Five New Genera and

Three New Species

(Pisces: Blenniidae)

Introduction

The purpose of this study is to lay the foundation

for a revision of the blenniid fish tribe Salariini.

This study evolved as a result of our attempts to

classify some apparently undescribed genera and

species that belonged in the Salariini (as defined

by Springer, 1968b). Two of the new genera we

describe were mentioned by Springer (1968b), but

were not described in that paper. Medusablennius,

one genus included by Springer in the tribe Blenniini, is here transferred to the Salariini. Description of the five new genera and three species required that we examine the entire nomenclature

of the Blenniidae and a large portion of the blenniid specimens (especially types) available in museums. As a result, we present an annotated list

of the approximately 350 nominal species of the

Salariini and a reinterpretation of and key to the

included genera. While some minor extensions and

corrections are necessary, our findings accord with

our recent revisions of genera of the Salariini

(Springer, 1962, 1967, 1968a, 1971; Springer and

William F. Smith-Vaniz, University of Miami, Rosenstiel School

of Marine and Atmospheric Sciences, Miami, Florida 33149.

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

Washington, D.C. 20560.

Smith-Vaniz, 1968) and Springer's (1968b) osteology and classification of the Blenniidae. Because

13 of the 24 genera we recognize in the Salariini

have not been recently revised, some changes in

the number of genera may be expected.

Acknowledgments

We wish to express our appreciation to the large

number of colleagues who assisted us in obtaining

data for our study, either through hospitality at

their institutions or by providing data on, or loans

of, specimens (abbreviations refer to institutions

and are explained on p. 3) : Drs. D. E. Rosen and

C. L. Smith, AMNH; Drs. F. Talbot and J. Paxton,

AMS; Dr. J. E. Bohlke, ANSP; Drs. P. H. Greenwood and N. B. Marshall and Mr. A. C. Wheeler,

BMNH; Dr. J. E. Randall, BPBM; Dr. W. N. Eschmeyer and Miss P. Sonoda, CAS; Dr. H. Steinitz,

HUI; Drs. C. Karrer and K. Deckert, ISZZ; Dr. G.

Mead and Mrs. M. Dick, MCZ; Dr. M. L. Bauchot,

MNHN; Dr. W. Klausewitz, NFIS; Dr. P. Kahsbauer, NMV; Dr. J. T. Woods, QMB; Dr. M. Boeseman, RMNH; Mrs. M. M. Smith, RU; Dr. M. L.

Penrith, SAMN; Dr. W. Freihofer, SU; Mr. T.

Iwamoto, UMML; Dr. J. Nielsen, UZMK; Dr. A.

N. Svetovidov, ZIAS; Dr. I. Tomiyama, Amakusa

Marine Biological Laboratory; Dr. H. Nijssen,

ZMA; Dr. A. G. K. Menon, ZSIC; Dr. L. Fishelson,

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

University of Tel Aviv, Israel; Dr. R. M. Bailey,

University of Michigan.

Mr. E. N. Gramblin, USNM, was of great assistance in expediting the heavy demands for curatorial work arising during our study.

Drs. B. B. Collette, United States Bureau of Commercial Fisheries, and C. R. Robins, UMML, read

the manuscript and offered valuable suggestions

for its improvement.

This study was made possible by a summer graduate student research appointment to Smith-Vaniz

in 1968 by the Office of Academic Programs, Smithsonian Institution, and a research award to Springer

by the Smithsonian Research Foundation. Important material of some cf the new genera and species

described below was obtained by Springer during

field work in the Gulf of Aqaba and Red Sea under

Smithsonian Foreign Currency Grant 7-0062 (2),

Drs. W. Aron and H. Steinitz, principal investigators, and by Smith-Vaniz in Ceylon under Smithsonian Foreign Currency Grant 3818, Dr. C. R.

Robins and Mr. F. H. Berry, principal investigators.

Methods and Material

COUNTS.—In determining the range of variation

for meristic characters we have drawn on information in our previous generic revisions and the literature for counts on species of which we had little

or no material. When possible, counts were made

on at least ten specimens of each species we examined, in addition to the type material examined.

Usually we counted the dorsal, anal, and caudal

fin rays, the vertebrae, and pleural and epipleural

ribs from radiographs. Cleared and stained material, mostly that reported by Springer (1968b) , was

also used in making counts (or obtaining other

osteological data).

Dorsal-fin elements: The unsegmented rays are

all flexible and are treated as spines. Frequently,

the terminal spine of many species is reduced

greatly in size and is not visible except in osteological preparations or on radiographs. When the

dorsal fin is deeply notched between the spinous

and rayed portions, the last spine is usually difficult

to distinguish externally. As the posteriormost

dorsal pterygiophore always supports a single ray,

each dorsal fin ray was counted and considered a

separate element.

Anal-fin elements: All Salariini have two flexible

anal spines except for aberrant specimens. In sexually undifferentiated specimens and males these

spines are recognizable. In mature males of some

genera they are enveloped in rugose fleshy processes

(to a lesser degree, the skin of some of the anterior

rays of these species may also be plicated). In females of most species the first anal spine is greatly

reduced and completely embedded in the flesh of

the urogenital papilla. The first anal spine of females usually is visible only in osteological preparations or on radiographs. In the Salariini the last

anal-fin pterygiophore may support one or two rays,

in which case the two rays are quite close together

and appear as one ray split to the base. In reporting our anal ray counts we counted two such rays

as one element, but when the last two anal pterygiophores each supported a single ray, the last two

rays were recorded as two elements (see Springer,

1967, for a discussion of the last anal ray in Entomacrodus).

Pectoral-fin ray counts were made on one side

(one fin).

Pectoral-fin radials are reported as a three-part

formula, for example 1-1-2, indicating that one

radial articulates only with the scapula, one radial

articulates with both the scapula and coracoid in

the area where the latter two bones meet, and two

radials articulate only with the coracoid.

Pelvic-fin rays (Figures la and b) : The pelvic

fin always has a single spine, which is greatly reduced, closely applied to the first segmented ray

and is visible only in osteological preparations. In

addition, there are always three or four segmented

pelvic rays in the Salariini, but the intermost ray

may be much reduced and extremely difficult to

find. In some species the innermost ray is completely imbedded and closely applied to the adjacent ray, or it is reduced to a few fragments of bone

visible only in cleared and stained material (for

example Glyptopanis). Unless otherwise indicated,

we report apparent segmented pelvic rays in our

counts for the pelvic fin. Springer (1968b) reported

that the pelvic-ray counts for Salariini ranged from

1,2 to 1,4 neglecting to note that the 1,2 count was

an apparent count.

Vertebral counts: These counts are reported as

total vertebrae (includes hypural vertebra), precaudal vertebrae (those vertebrae anterior to the
NUMBER 73

FIGURE 1.—Ventral view of left pelvic fin showing relative

development of innermost segmented ray: a, Antennablennius velifer, USNM 200205, male, 44.3 mm, Astola Island, W.

Pakistan, b, Alloblennius pictus, USNM 203776, male, 27.7

mm, Strait of Jubal, Red Sea.

first vertebra with a well-developed hemal spine,

which articulates with at least the first anal pterygiophore), and caudal vertebrae (all vertebrae, including the hypural, posterior to the precaudal

vertebrae). Sometimes the last precaudal vertebra

bears a moderate hemal spine, but in these cases

it will be apparent that the first anal pterygiophore is closer to the well-developed hemal spine

of the next vertebra (first caudal).

Epipleural ribs are considered to begin on the

first vertebra and the posteriormost rib is counted

whether on the right or left side. Pleural ribs are

considered to begin on the third vertebra and are

counted similarly.

Teeth: Counts given are total counts for either

both dentaries or both premaxillaries, exclusive of

the posterior dentary canines. In Ecsenins the anterior set of canines, which in most species are

difficult to distinguish from the incisoriform teeth,

are included in the dentary count. Ecsenins is the

only genus with both anterior and posterior canines (Figure 5).

Typically in the Salariini, the number of teeth

increases with increase in size (see Springer and

Smith-Vaniz, 1968, for Atrosnlarias) . In the diagnoses and Table 3, the tooth counts given are

indications of the range for adults, but the actual

range will probably vary from the given range when

all the species of the various genera are surveyed.

Circumorbital bones: The lachrymal and dermosphenotic were included in the counts of circumorbital bones.

MEASUREMENTS.—Standard length (SL) was measured from the midtip of the upper jaw to the middistal end of the hypural vertebra (taken externally) .

HEAD PORES.—All illustrations of cephalic pore

patterns were made using a camera lucida. Pore

diameters are slightly exaggerated for clarity. To

facilitate orientation, small arrows that mark pores

in corresponding positions have been included in

both dorsal and lateral views of most species illustrated. All lateral view pore pattern figures are

presented as if the left side of the specimens were

illustrated. In some instances the right side was

actually drawn but was presented as the left side

for the sake of consistency. In such cases the dorsal

views were adjusted to accord with the mirrorimage figures.

OCCIPITAL CREST.—We define this structure as a

thin, bladelike, longitudinal, median flap on top

of the head. Mature males in many species of the

Salariini (see Springer, 1967, pi. bd) have a raised,

fleshy, swollen area on top of the head that should

not be confused with an occipital crest.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ABBREVIATIONS OF INSTITUTIONS CITED.—

AMNH American Museum of Natural History,

New York City

AMS Australian Museum, Sydney

ANSP Academy of Natural Sciences of Philadelphia

BMNH British Museum (Natural History),

London

BOC Bingham Oceanographic Collection (now

Peabody Museum), New Haven, Connecticut

BPBM Bernice P. Bishop Museum, Honolulu,

Hawaii

CAS California Academy of Sciences, San

Francisco

FMNH Field Museum of Natural History, Chicago

HUI Hebrew University, Jerusalem, Israel

ISZZ Institut fiir Spezielle Zoologie und

Zoologisches Museum, Berlin

KUMF Kasetart University Museum of Fisheries, Bangkok, Thailand

MCZ Museum of Comparative Zoology, Cambridge, Massachusetts

MMS Macleay Museum, University of Sydney,

Australia

MNHN Museum National d'Histoire Naturelle,

Paris

MNRJ Museu Nacional, Rio de Janeiro

NFIS Natur-Museum und Forschungs-Institut

Senckenberg, Frankfurt

NMV Naturhistorisches Museum, Vienna

QMB Queensland Museum, Brisbane

RMNH Rijksmuseum van Natuurlijke Historie,

Leiden

RU Rhodes University, J. L. B. Smith Institute of Ichthyology, Grahamstown,

South Africa

SAMN South African Museum, Capetown

SMNS Staatliches Museum fiir Naturkunde,

Stuttgart

SU Stanford University, Division of Systematic Biology, Palo Alto, California

UMML University of Miami, Rosenstiel School

of Marine and Atmospheric Sciences,

Florida

USNM National Museum of Natural History,

Division of Fishes, Washington, D.C.

UZMK Universitetets Zoologiske Museum, Copenhagen

VMN Vanderbilt Marine Museum, Centerport,

New York

ZIAS Zoological Institute, Academy of Sciences, Leningrad

ZITU Zoological Institute, University of Tokyo

ZMA Zoologisch Museum, Universiteit van

Amsterdam

ZSIC Zoological Survey of India, Calcutta

MATERIAL.—The material upon which our study

is based falls into three categories:

1. Primary type material, mostly examined over

a period of several years prior to this study.

2. Material examined and reported in our previous published papers (see introduction), not necessarily reexamined for the present study.

3. Material in the extensive collections at the

National Museum of Natural History and a few

other museums that was examined during the course

of this study.

The material in the first category is cited in our

annotated list of nominal species. The material in

the second category can be found cited in the relevant publications. The material in the third category is only partially cited in the present paper.

We cited this material in our descriptions of new

genera (even when a new species was not involved)

because we know of no material of these genera

other than that which we list. We also cited some

material that significantly extended the known

geographic range of one genus we previously revised, Atrosalarias. Illustrated material is all cited.

The remaining material was not cited because: we

could not specifically identify the specimens even

though we knew the genus; data on genera were

not reported by species, but were lumped; there is

a possibility that some lots of specimens we examined actually contained more than one species (ignored when we were only interested in generic

ranges of characters) ; and we did not always examine every specimen in a particular lot of specimens (usually only a series of ten).

Tribe Salariini

Type-genus: Salarias Cuvier, 1817

The Salariini constitute the largest tribe of the
NUMBER 73

four tribes of the Blenniidae. We recognize 24

genera with an estimated 150 species in the Salariini.

The tribe is predominantly restricted to the Indian

and western and central Pacific oceans, with only

nine species occurring in the Atlantic and eastern

Pacific oceans. All species lack a swim bladder, are

essentially benthic, and most are rarely seen swimming off the bottom. Some genera have become

secondarily adapted for a semiterrestrial existence

and spend much of their time out of water. An

excellent account of aerial respiration in the genus

Andamia was given by Rao and Hora (1938).

The Salariini have been variously accorded family or subfamily rank, primarily as a result of Norman's (1943) classification of the Blenniidae. Norman used the nature of the jaws and dentition as

primary characters in recognizing the Salariinae.

Springer (1968b) reexamined the nature of the

jaws and dentition of blenniids and accorded the

Salariinae only tribal rank because there were a

few species of blennies that had the jaws and dentition apparently intermediate in structure between

those in the typical Salariini and their closest relatives, the Blenniini, which Norman accorded the

rank of subfamily, Blenniinae. The Salariini and

Blenniini could be combined, but because there

are few intermediate forms, Springer chose to keep

the tribes separate. In the present paper we maintain the separation for much the same reason, as

well as for the fact that all but one genus of the

intermediate forms appears to us to be most similar

to the typical Salariini in general physiognamy.

Although Springer (1968b, p. 44) adequately

discussed the nature of the dentition and jaws of

the Salariini, he (1968b, p. 65) may have been

misleading when he diagnosed the tribe using only

the characters of the jaws and dentition of the

typical members. Because of the importance of the

nature of the jaws and dentition in the recognition

of the Salariini, we present an expanded discussion

of these structures.

In the other tribes of the Blenniidae the premaxillaries and dentaries are essentially closed bony

capsules supporting the functional teeth. The replacement teeth develop inside these capsules and

exit through small foramina in the wall of each

capsule when proceeding into position in the functional row.

In most Salariini (all genera except those in

group 21 of Figure 51) the anteroventral wall of

each premaxillary capsule and the anterior wall of

each dentary capsule are absent, and these bones

are relatively thin, concave (arched) structures

(Figures 2, 3, and 5). In those species with relatively large numbers of incisoriform teeth, the

functional teeth (except the canines, which always

develop in a separate capsular compartment of the

dentaries and are ankylosed to the bone) are

loosely held in a thick band of connective tissue

that fills the concavity of each premaxillary and

dentary. The bases of the functional teeth are not

ankylosed to the bone of the jaws. In those species

with relatively few teeth, the band of connective

tissue is much reduced and the bases of the functional teeth (except the canines) are weakly ankylosed to the jaw bones. The replacement teeth in

both these types develop in the same bands of

connective tissue that hold the functional teeth.

The replacement teeth are dorsal and anterior to

the bases of the functional premaxillary teeth and

ventral and anterior to the bases of the functional

dentary teeth. Inasmuch as the walls of the premaxillary and dentary bones are open, the replacement teeth merely move through the band of connective tissue to take position in the functional row.

The nature of the dentition and jaws in the remaining genera (group 21 of Figure 51), all with

relatively few teeth, is variable. In all these genera

except Hircuiops, the premaxillary is typical of the

other Salariini except that the bone is heavier

(Figure 3). In Hirculops (Figure 4) the premaxillary is open mesially, with normal salariinine replacement of teeth, but laterally the premaxillary

is closed, with the replacement teeth entering the

functional row through foramina in the bone. The

dentaries of the other genera (group 21, Figure 51

and Figure 7), except one species of Rhabdoblennius, (R. snowi), are all blenniinine-like: closed

capsules with the replacement teeth entering the

functional row through foramina in the bone. The

foramina are relatively much larger than those

found in the Blenniini, however, and give the impression that the wall of the dentaries is eroding

or rebuilding. This is further evidenced by Rhabdoblennius snowi (Figure 6), wherein the bone (capsule) of the dentaries is completely open mesially,

but with large gaps and foramina in the wall

laterally.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The structure of the jaws and dentition is obviously specialized in most Salariini, and is less specialized in the Blenniini. Whether the forms with

apparently intermediate dentition are phylogenetically intermediate, or represent developments that

parallel the less specialized blenniinine dentition,

is not known.

Other diagnostic characters of the tribe Salariini

can be found in Springer (1968b, pp. 63—64). Some

extension in the ranges of meristic characters will

be found in our Tables 2 to 6. Any blenniid with

more than 80 teeth in either the upper or lower

jaws belongs to the Salariini, while blenniids with

less than that number may belong to this tribe.

Aside from the nature of the jaws and dentition,

there is no other single character we know of that

sets all the Salariini apart from their closest relatives, the Blenniini.

Nominal Genera and Subgenera in Salariini

The following list of generic group names in the

Salariini is provided to facilitate location of discussion in the text. Each name we synonymize, or

recognize as a subgenus, is followed by the genus

under which we include it:

Alloblennius

Alticops = Istiblennius

Alticus

Andamia

Antennablennius

Anthiiblennius — Ecsenius

Atrosalarias

Basilisciscartes = Alticus

Blenniella = Istiblennius

liletitwphis = Ophioblennius

Cirripectes

Crenalticus — Salarias

Croaltus = Antennablennius

Crossosalarias

Cynoscnrtes = Ophioblennius

Dtimnnia = Alticus

Ecsenius

Entomacrodus

Erpiclitliys — Salarias

Exallias

Fallacirripectes = Stanulus

Giffordella = Entomacrodus

Gloriella = Exallias

Glyptoparus

Halmablennius — Istiblennius

Hepatoscartes= Ophioblennius

Hirculops

Istiblennius

Labroblennius = Ophioblennius

Leoblennius = Exallias

Litanclius = Antennablennius

Litobranchus

Loplialticus = Alticus

Medusablennius

Miinoblennius

Nannosalarias

Negoscartes = Salarias

Nixiblennius — Rhabdoblennius

Opliioblennius

Pereulixia

Pescadorichthys — Ecsenius

Praealticus

Rhabdoblennius

Rupiscartes = Alticus

Salarias

Salarichthys — Entomacrodus

Scartes = Scartichthys

Scartichthys

Scartoblennius = Rhabdoblennius

Stanulus

FIGURES 2-4.—Lateral view of left premaxillary showing functional and replacement teeth: 2, Ecsenius yaeyamaensis,

USNM 200428, male, 38.7 mm, New Britain (dark area on

proximal third of teeth represents band of conective tissue

beneath translucent teeth); 3, Rhabdoblennitis snowi, USNM

142135, female, 38.0 mm, Marshall Islands; 4, Hirculops

conifer, USNM 203123, male, 42.4 mm, Gulf of Suez, Red Sea.

FIGURES 5-7.—Anterolateral view of left dentary showing relative excavation of bone (membranous connective tissue and

functional incisoriform teeth removed): 5, Ecsenius yaeyamaensis, USNM 200428, male, 38.7 mm. New Britain (see

Springer, 1968b, plate 9, for illustration of intact dentary);

6, Rhabdablennius snowi, USNM 198715, female, 44.2 mm,

Howland Island; 7, Rhabdoblennius rhabdotrachelus, USNM

115509, male, 35.3 mm, Phoenix Island.
NUMBER 73
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Genera of the Tribe Salariini

1. Elongate, transverse series of cirri continuous across nape or interrupted at midline of nape

by a narrow hiatus no greater than 25 percent of length of base of either patch of cirri;

lateral line complete, extending to or almost to caudal base 2

Nape cirri, when present, restricted to a single cirrus or a patch of cirri on each side of nape

separated by a wide hiatus nearly equal to or greater than length of base of either patch

of cirri; lateral line complete or incomplete 4

2. Segmented dorsal rays 14 to 16; segmented anal rays 14 to 17 (rarely 14); both upper and

lower jaw teeth freely movable, subequal in breadth; about one-half as many teeth in

lower jaw as in upper; teeth in lower jaw 85 to 135; vomerine teeth absent; one or two

canines present posteriorly on each dentary Cirripectes Swainson, 18S9

Segmented dorsal rays 11 to 13; segmented anal rays 12 to 14; upper jaw teeth freely

movable, lower jaw teeth scarcely movable, nearly twice as broad as upper jaw teeth;

about one-third as many teeth in lower jaw as in upper; teeth in lower jaw less than

65; vomerine teeth present or absent; canines present or absent posteriorly on each

dentary 3

3. Upper lip with 18 to 24 well-developed crenulae; supraorbital cirrus multind, with 5 to 10

branches; lateral line with numerous short side branches anteriorly (Figure 30); no

imbricate scalelike flaps covering anterior lateral-line pores; pair of barbels present on

each side of chin just behind lower lip; dentary canines absent; vomerine teeth

absent Exallias Jordan and Evermann, 1905

Upper lip entire; supraorbital cirrus simple, long and tapering; lateral line consists of short,

separate, longitudinally bipored tube anteriorly; imbricate scalelike flaps covering

at least anterior lateral-line pores (Figure 29) ; no barbels on chin; dentary canines

present; vomerine teeth present Pereulixia Smith, 1959

4. Nuchal and supraorbital cirri absent; all caudal rays unbranched; latcralmost tooth on

each dentary occasionally canine-like, slightly broader (in frontal view) and more robust

than adjacent teeth; lateral extrascapulars fused indistinguishably to pterotic; median

ethmoid unossified; circumorbital bones 4 Ecsenitu McCulloch, 1923

Either nuchal or supraorbital cirri may be absent, but never both (both may be present);

caudal rays branched or unbranched; latcralmost tooth on each dentary similar to

adjacent teeth; lateral extrascapular distinct and separate from pterotic; median ethmoid

ossified; circumorbital bones 5 (except Nannosalarias and Alloblennius, which have 4,

and Medusablennius, which has 2) 5

5. Dorsal spines 9 to 11; all caudal rays unbranched; pectoral rays 15 to 18 (usually 16);

segmented pelvic rays 2; color of head and body almost uniformly dark

Atrosalarias Whitley, 1933

Dorsal spines 12 to 17 (rarely 11); caudal rays branched or unbranched; pectoral rays 13

to 16 (rarely 16); segmented pelvic rays 2 to 4; color of head and body variable 6

6. Total dorsal fin elements 21 to 23; segmented dorsal rays 9 to 12; segmented anal rays

10 to 13; minute, imbricate, scalelike flaps covering anterior lateral-line pores

Stimulus Smith, 1959

Total dorsal fin elements 26 to 38; segmented dorsal rays 13 to 24; segmented anal rays

14 to 28; no minute, imbricate, scalelike flaps covering lateral-line pores 7

7. Teeth in upper jaw freely movable, in adults usually exceeding 120 (110 to 380) 8

Teeth in upper jaw immovable or nearly so, 18 to 80 17

8. All caudal rays unbranched; segmented anal rays 23 to 28 9

Some caudal rays branched near tips, at least in adults; segmented anal rays 17 to 25 10

9. A prominent cup-shaped fleshy disk or appendage behind lower lip (Figure 10)

Andamia Blyth, 1858

No fleshy disk or appendage behind lower lip Alticus Lacdpede, 1880

10. Following combination of characters present: occipital crest, no nuchal cirri, nasal cirri

simple (rarely with single, short side branch), supraorbital cirri pinnately branched

(at least in adults), and single pore present in mid-dorsal area of supratemporal canal

(Figure 21) Praealticus Schultz and Chapman, 1960

Above characters never present in combination H
NUMBER 73

11. Segmented pelvic rays 2 or 3 (each fin) 12

Segmented pelvic rays 4 14

12. Gill membranes bound to isthmus at about, or slightly below, level of ventralmost pectoral

ray; gill membranes occasionally forming a narrow free fold across isthmus; cirri associated with 2 to 4 preoperculomandibular pores on each side; a fleshy flap at base of

first dorsal spine Crossosalarias Smith-Vaniz and Springer

Gill membranes unrestricted laterally and posteriorly; gill membranes forming a deep, free

fold across isthmus; no cirri associated with preoperculomandibular pores; no fleshy flap

at base of first dorsal spine 13

13. Terminal anal ray completely free from caudal peduncle; dorsal spines 13 or 14; segmented pelvic rays 3; anterior anal rays not elongated Istiblermius Whitley, 1943

Terminal anal ray partially bound to caudal peduncle by membrane; dorsal spines 12 or

13, typically 12; segmented pelvic rays 2; anterior anal rays frequently elongated in

mature males Salarias Cuvier, 1817

14. Dorsal fin without notch between spinous and segmented rays (but notched somewhat in

larvae); segmented dorsal rays 19 to 24; segmented anal rays 20 to 22; pectoral rays

typically 15 Ophioblennius Gill, 1860

Dorsal fin with notch between spinous and segmented rays; segmented dorsal rays 13 to 18;

segmented anal rays 14 to 19; pectoral rays typically 14 15

15. Dorsal spines typically 13 (last spine often greatly reduced); vomerine teeth present or

absent, frequently broken off; cirrus on each side of nape usually simple, varying from

no cirri to a small patch (circumtropical, but eastern Pacific species with a single

cirrus) 16

Dorsal spines typically 12; vomerine teeth absent; a patch of cirri on each side of nape

(restricted to the eastern Pacific) Scartichthys Jordan and Evermann, 1898

16. High, thin, fleshy occipital crest present (at least in males); canine tooth absent on dentary

posteriorly; vomerine teeth absent Istiblermiusl Whitley, 1943

Occipital crest absent or present only as a low fold; canine tooth present on dentary posteriorly; vomerine teeth present in most specimens Entomacrodus Gill, 1859

17. Segmented dorsal rays 14 to 16; upper jaw teeth 74 to 80; lower jaw teeth 42 to 58 18

Segmented dorsal rays 17 to 21; upper jaw teeth 18 to 50; lower jaw teeth 16 to 38 19

18. Fleshy crest on midline of nape (low and poorly developed in females); pectoral rays 12

to 14; segmented pelvic rays 2; no vomerine teeth; circumorbital bones 5.

Glyptoparus Smith, 1959

No fleshy crest on midline of nape; pectoral rays 15; segmented pelvic rays 3; vomerine

teeth present; circumorbital bones 4 Nannosalarias Smith-Vaniz and Springer

19. All caudal rays unbranched 20

Central caudal rays branched near tips 21

20. Cirri on interorbital region, rims of anterior and posterior nostrils, and over eye; nuchal

cirri absent; upper jaw teeth 18 or 19; circumorbital bones 2.

Medusablennius Springer, 1966

No cirri on interorbital region, simple cirrus on anterior nostril and over eye; minute

nuchal cirri present; upper jaw teeth 36 to 42; circumorbital bones 5.

Utobranchus Smith-Vaniz and Springer

21. Dorsal spines typically 13; nasal and nuchal cirri palmate, multifid; supraorbital cirri over

each eye 2 or multifid Mimoblemnus Smith-Vaniz and Springer

Dorsal spines 11 to 13, usually 12; nasal and nuchal cirri, if present, slender and simple;

supraorbital cirri, if present, simple 22

22. Posterior dentary canines well developed; vomer with small conical teeth; last POP position

consisting of a pair of vertical pores (Figures 43, 45, and 46); epipleural ribs 19 to 27. .23

Posterior dentary canines absent or minute; vomerine teeth absent; last POP position

consisting of a single pore (Figures 47-50); epipleural ribs 11 to 16 24

23. Nuchal cirri present; diameter of eye into length of supraorbital cirrus 1.7 to 2.5; precaudal

vertebrae 11 (western Indian Ocean and Red Sea) Hirculops Smith, 1959

Nuchal cirri absent; diameter of eye into length of supraorbital cirrus 1.0 to 1.5; precaudal

vertebrae 10 (western Pacific and eastern Indian oceans) .. Rhabdoblennius Whitley, 1930
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

24. Supraorbital cirri present; nuchal cirri present (minute) or absent; circumorbital bones

4 (Red Sea) Alloblennius Smith-Vaniz and Springer

Supraorbital cirri absent; nuchal cirri present; circumorbital bones 5 (Indian Ocean and

Red Sea) Antennablennius Fowler, 1931

1

 Istiblennius will key out in two places in the key. While most species of Istiblennius have

1,3 pelvic rays, /. unicolor has 1,4 and it alone keys out here.

Alloblennius, new genus

DIAGNOSIS.—Dentary a closed capsule with replacement teeth entering functional series through foramina in jawbone. Anterior and posterior dentary

canines absent. Premaxillary and dentary teeth immovable or nearly so, numbering 30 to 38 in upper

jaw and 30 to 36 in lower jaw. Vomer toothless.

Dorsal rays XI to XIII, usually XII, 16 to 20; anal

rays II, 19 to 22; segmented caudal rays 13, middle

9 branched; pectoral rays 13 to 15 (usually 14);

pelvic rays 1,3 (the innermost pelvic ray in one

species considerably reduced, closely applied to

base of second ray, and visible only in cleared and

stained specimens or upon careful dissection). Terminal anal ray bound by membrane to caudal

peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps

covering lateral-line pores. Preoperculomandibular

pores without cirri. Mid-dorsal supratemporal pores

2-8. Upper lip without free dorsal margin. No cupshaped fleshy disk or appendage behind lower lip.

Gill membranes free. Occipital crest absent. Nuchal

cirri present (minute) or absent; supraorbital and

nasal cirri simple. Postcleithra consisting of two

elongate bones, head of ventral element overlapping ventral end of dorsal element. Lateral extrascapular not fused with pterotic. Median ethmoid

ossified. Circumorbital bones 4. Type-species:

Rhabdoblennius pictus Lotan.

Relationships are discussed under section titled,

"Recognition of Genera."

ETYMOLOGY.—A combination of the Greek allos,

different or strange, and blennos, the name of a

fish, alluding to the unusual discovery of a genus

of blenniid fishes known only from the Red Sea.

From a zoogeographic standpoint Alloblennius

is of interest because it is the only blenniid genus

restricted to the Red Sea. This may not be surprising as a large number of Red Sea fish species

are apparently endemic (Marshall, 1952). Alloblennius appears to be derived from an Antennablennius-like ancestor that invaded the Red Sea,

became isolated, and evolved to its present level

of differentiation.

The genus comprises two species, A. pictus and

A. jugular is. A. pictus differs most notably from

A. jugularis in having the innermost pelvic ray not

obvious, the ventral hypural plate fused proximally

with the urostylar centrum and dorsal hypural

plate, and in lacking nuchal cirri (minute nuchal

cirri are usually present in jugularis). Both species

are described in detail below.

Alloblennius jugularis (Klunzinger)

FIGURES 8a, 50

Iilennius jugularis Klunzinger, 1871, p. 493 [original description; type-locality: Red Sea, Kosseir].

MATERIAL EXAMINED.-ISZZ 10496 (male, 29.5

mm SL), lectotype and ZIAS 2617 (female, 22.7

mm SL), paralectotype, both from Kosseir, Red

Sea; USNM 204695 (6, 30.5-42.8) Kosseir, lat.

26°08'N, long. 34°16'E; USNM 204528 (3, 30.7-

36.2), USNM 204531 (2, 23.3-29.9) and USNM

204532 (1, 27.8), Gulf of Aqaba near Ras Burqa;

USNM 204529 (1, 26.6), Gulf of Aqaba near Marsa

Muqabila; USNM 204530 (5, 21.3-31.5), S end of

Sinai Peninsula at Ras Muhammad; USNM 204612

(1, 19.1), Ethiopia, SW shore of Sciumma Island,

lat. 15°32'31"N, long. 40°00/E; USNM 204488 (26,

16.4-31.0), Ethiopia, Difnein Island, lat. 16°36'N,

long. 39°20'E; HUI E62/417g, (6, 27.6-33.3), Ethiopia, Dahlak Archipelago, Cundabilu.

DESCRIPTION (characters for lectotype in parentheses) .—Nature of dentary and replacement teeth

not investigated. Dentary canines absent. Premaxillary and dentary teeth relatively immovable; premaxillary teeth 30 to 36 (35); dentary teeth 30 to

34 (32). Vomerine teeth absent. Terminal vertebra

with two epurals and autogenous hypural 5 and
NUMBER 73 11

FIGURE 8.—Alloblennius (lateral and ventral views): a, A. jugularis, HUI E62/417g, male, 33.4 mm.

Cundabilu, Dahlak Arquipelago; b, A. pictus, USNM 204524, male, 25.8 mm, Gulf of Aqaba.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ventral hypural plate. Vertebrae 10 + 27 to 29 = 37

to 39 (38); epipleural ribs 12 to 16 (14); last

pleural rib on vertebra 10. Circumorbital bones 4.

Dorsal spines 11 to 13, usually (12); last spine

noticeably reduced; segmented rays 17 to 20 (19);

at least basal two-thirds of terminal dorsal ray

bound by membrane to caudal peduncle and dorsal

procurrent rays; dorsal fin moderately incised between spinous and rayed portions. Anal spines 2;

segmented anal rays 20 to 22 (21); at least basal

two-thirds of terminal anal ray bound by a membrane to caudal peduncle. Caudal fin with 13 segmented rays, middle 8 (9) each branched once;

dorsal procurrent rays 6 or 7; ventral rays 5 to 7

(7). Pelvic rays 1,3, all rays well developed. Pectoral

rays 14. Gill-rakers 10 (lectotype only). Pseudobranchial filaments 5. Nuchal cirri present or absent

(absent); supraorbital and nasal cirri short and

simple. Upper and lower lips entire. Lateral line

of simple pores, considerably reduced, ending at

level of fourth dorsal spine.

COLOR PATTERN.—Mature males: The snout is

dusky. There is a broad band of melanophores behind the eye that becomes diffuse as it extends on

top of the head, but is distinct as it continues ventrally around the eye, and is continuous with a

black area on the underside of the head. The black

area covers the ventral sides of the head but is interrupted by two large, pale spots, one on each

side. Other pale spots may occur on each side lateral to the ventral spots, just anteroventral to the

ventral end of the opercle, or at the ventral end

of the preopercular area. The ventral end of the

opercle is covered by melanophores; the dorsal end

is dusky. The prepelvic area is dark dusky to black

above the level of the pelvic fin base; the dark area

extends onto the ventral half of the fleshy pectoral

base. The venter is black. The body bears six to

eight dusky, bandlike spots at its midlevel. There

may also be a fine stippling of spots over the body.

The spinous dorsal bears a black spot between the

first two dorsal spines; the remainder of the fin is

irregularly marked with dark spots. The rayed portion of the dorsal fin bears a regular pattern of

dark melanophores and pale areas for its entire

length. The tips of the rays and the base of the

anal fin are pale to pale dusky with or without fine

dark spots basally. The remainder of the fin is

dark dusky. The caudal fin bears a pattern of up

to six irregularly vertical rows of dark spots. The

pectoral fins are faintly dusky, and the pelvic fins

are unmarked.

Immature males have very little black pigment,

but may show the same pattern as mature males,

only much less intense. Females are very pale. They

exhibit some pale dusky marks on the sides of the

head and body and a regular pattern of dusky

spots on the dorsal and caudal fins.

REMARKS.—The key and description given by

Klunzinger (1871, p. 493) to distinguish his new

species, Blennius jugularis, contain features indicative of two different genera. In the Berlin Museum

(Klunzinger, whose collections originally resided in

Stuttgart, sold all the specimens reported in his

1871 paper to other museums, see Klausewitz, 1964,

p. xii) there are two specimens, catalog numbers

8025, a 28 mm female, and 10496, a 29.5 mm male,

and in the Leningrad Museum there is a single

specimen, catalog number 2617, a 22.7 mm female,

that were identified as Blennius jugularis by Klunzinger and which were purchased from him. As

all three specimens are from Kosseir, they apparently were included in his 1871 study. While the

fin-ray counts given by Klunzinger agree with both

females, the description of the cirri excludes the

Berlin female, 8025. The description of the cirri

in the key agrees with none of the specimens, but

appears to be a composite from all. The Berlin

female is a species of Antennablennius, possibly

undescribed, and the Berlin male and Leningrad

female represent the species treated here as Alloblennius jugularis. We consider the three specimens

as syntypes of B. jugularis and here designate the

Berlin male, 10496, as lectotype.

Alloblennius pictus (Lotan)

FIGURES \b, 86, and 49

Rhabdoblennius pictus Lotan, 1970, p. 376 [original description; type-locality; Eilat, Gulf of Aqaba].

MATERIAL EXAMINED.—HUI E63/37,l (male, 25.6

mm SL), holotype, Eilat, Gulf of Aqaba, Red Sea,

4 July 1963, A. Ben-Tuvia; USNM 204523 (3, 14.0-

35.4) and USNM 204525 (35, 13.5-27.9), Gulf of

Aqaba near Marsa Muqabila; USNM 204524 (10,

20.2-31.3) and USNM 204527 (2, 14.8-19.8), Gulf

of Aqaba just N of Ras Burqa; USNM 204522 (1,
NUMBER 73 13

19.5), Gulf of Aqaba, bay between Marsa Mokrakh

and El Himeira; USNM 204526 (8, 13.6-25.2), Gulf

of Aqaba between Marset Mahash el Ala and Marset Abu Samra; USNM 203766 (19, including 2

cleared and stained, 16.8-32.6), Red Sea, Strait of

Jubal, lat. 27°14/34"N, long. 33°53'55"E; USNM

204508 (8, 16.8-21.5), Ethiopia, Sciumma Island,

lat. 15°32'31"N, long. 40°40'E; USNM 204498 (9,

18.0-26.1), Ethiopia, Massawa; USNM 204501 (83,

13.8-28.5), Ethiopia, Difnein Island, lat. 16°36'N,

long. 39°20'E; USNM 204510 (1, 17.9), Ethiopia,

Melita Bay, lat. 15°15'N, long. 39°49'E; USNM

204504 (4, 18.6-22.6), Ethiopia, near Harat Island,

lat. 16°08'N, long. 39°26.5'E; USNM 204502 (1,

18.6), Ethiopia, north end Isola Delemme, lat.

15°30.5'N, long. 39°54'E.

DESCRIPTION (characters for holotype in parentheses) .—Dentary a closed capsule with replacement

teeth entering functional series through foramina

in jawbone. Dentary canines absent. Premaxillary

and dentary teeth relatively immovable; premaxillary teeth 34 to 38 (34); dentary teeth 32 to 36

(32). Vomerine teeth absent. Terminal vertebra

with two epurals and autogenous hypural 5; ventral hypural plate fused proximally with urostylar

centrum and dorsal hypural plate. Vertebrae 10 +

26 to 28=36 to 38 (10+27); epipleural ribs 11 to

14 (12); last pleural rib on vertebra 10. Circumorbital bones 4.

Dorsal spines 11 to 13, usually 12 (12); last spine

noticeably reduced; segmented rays 16 to 20 (19);

basal two-thirds of terminal dorsal ray bound by

membrane to caudal peduncle and upper procurrent rays; dorsal fin moderately incised between

spinous and rayed portions. Anal spines 2; segmented rays 19 to 22 (22); basal two-thirds of terminal anal ray bound by a membrane to caudal

peduncle. Caudal fin with 13 segmented rays, middle 9 each branched once; dorsal procurrent rays 6

or 7; ventral rays 5 or 6. Pelvic fin 1,3 (innermost

pelvic ray considerably reduced, closely applied to

base of second ray, and visible only in cleared and

stained specimens or upon careful dissection, Figure 16). Pectoral rays 13 to 15, usually 14 (14). Pectoral radial formula 2-0-2. Gill-rakers 8 to 11.

Pseudobranchial filaments 5. Nuchal cirri absent;

supraorbital and nasal cirri short and simple. Upper and lower lips entire. Lateral line of simple

pores, considerably reduced, ending at level of second to fourth dorsal spine.

PIGMENTATION.—In most specimens the head and

body are pale with loose concentrations of melanophores variously distributed. Groups of melanophores may occur in the mid-predorsal region of

the head, on the snout, and around the orbit, with

an extension from the orbit across the top of the

cheek and another diagonally downward past the

corner of the mouth. The latter extension may be

continuous with the generally dusky to heavily pigmented underside of the head in males. In females

the underside of the head bears only a sparse

sprinkling of melanophores. The breast and a variable portion of the ventral half of the pectoral base

are dusky with melanophores in males, almost unmarked in females. The venter is unmarked in all

specimens but the dark peritoneum shows through

the skin in most specimens. The sides of the body

may bear a midlateral row of somewhat vertical

groups of melanophores and/or scattered ocelli as

shown by the specimen in Figure 8fr. Occasionally

there is a large, concentrated group of melanophores

in the humeral region. There is usually a row of

clusters of melanophores along the dorsal body

contour at the base of approximately every other

dorsal fin element. Usually there is an indication

of a dark spot between the first two dorsal spines

(both sexes) and various other groups of melanophores overlying the dorsal spines. There are about

four clusters of melanophores overlying each dorsal

ray, forming diagonal rows of spots across the length

of the fin. The anal fin usually bears spots at the

base of approximately every other ray, and most of

the distal one-fourth to one-half of the fin is evenly

dusky, except for the pale tips of the rays. The

caudal bears scattered spots, sometimes arranged

in up to five vertical rows. The pectoral and pelvic

fins are unmarked.

REMARKS.—Segmented dorsal and anal ray and

caudal vertebral counts, probably correlated meristic characters, indicate significant population differences between Ethiopian and northern Gulf of

Aqaba Israeli specimens, and sexual differences

between Ethiopian males and females. Israeli specimens do not exhibit significant meristic sexual

dimorphism:
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Israel

males

females

Ethiopia

males

females

16

_

_

_

1

Segmented

dorsal rays

17

_

_

2

5

18

13

4

39

30

19

21

15

23

3

20

3

_

_

_

Av

18.7

18.8

18.3

17.9

19

_

_

1

3

Segmented

anal rays

20

2

2

16

23

21

26

12

46

14

22

10

5

1

_

Av

21.2

21.2

20.7

20.3

Caudal

26

_

I

5

6

27

13

4

41

24

vertebrae

28

19

12

16

2

Av

27.6

27.6

27.2

26.9

The data were tested for significance, using Student's Mest, and the following ^-values and p-values

were obtained. (The values are based on a twosided test, Simpson, Roe, and Lewontin, 1960.) It

was decided beforehand that a p-value of .01 or less

would be considered significant:

Israeli males vs Israeli females for dorsal rays: / = —383, p := .7-.8

Ethiopian males vs Ethiopian females for dorsal rays: t z= 3.909, p zz <.001

Israeli males vs Ethiopian males for dorsal rays: t = 3.452, p — <.001

Israeli females vs Ethiopian females for dorsal rays: t — 6.213, p = <.O01

Israeli males vs Israeli females for anal rays: t zz .338, p — .7-.8

Ethiopian males vs Ethiopian females for anal rays: t zz 4.170, p — <.O01

Israeli males vs Ethiopian males for anal rays: * zz 4.491, p = <.001

Israeli females vs Ethiopian females for anal rays: t zz 5.284, p — <.001

Israeli males vs Israeli females for vertebrae: / zz —.330, p — .7-.8

Ethiopian males vs Ethiopian females for vertebrae: ( zz 2.586, p = .01

Israeli males vs Ethiopian males for vertebrae: t = 3.546, p zz <00 1

Israeli females vs Ethiopian females for vertebrae: t =z 4.821, p zz <.001

Israeli specimens have higher mean numbers of

dorsal and anal rays and caudal vertebrae than

Ethiopian specimens, and Ethiopian males have

higher mean numbers of dorsal and anal rays and

caudal vertebrae than Ethiopian females. Sexual

dimorphism similar to that found in Ethiopian

specimens was noted by Springer (1967) for some

species of Entomacrodus and by Springer (1971)

for some species of Ecsenius.

We have no explanation of why one population

should exhibit meristic sexual dimorphism and the

other not. The higher numbers of fin rays and

vertebrae of the Israeli specimens, however, may

well represent the effect of the cooler surface water

temperature of the Gulf of Aqaba as compared with

those of the Red Sea coast of Ethiopia. Oren (1962)

reported monthly average surface temperatures of

1.8-5.5°C less for the Gulf of Aqaba as compared

with those of the Red Sea coast of Ethiopia.

Seventeen additional males (and two females)

from the Strait of Jubal, Egypt, about 300 km

south of the Israeli localities from which we had

specimens, were not included in the above analyses. These males had means of 18.9 dorsal rays, 21.2

anal rays, and 27.6 caudal vertebrae, indicating no

differences from the Israeli males. Temperature

data for the Red Sea coast of Egypt, where these

specimens were collected, are not available, but

water temperatures generally decrease toward the

north in the Red Sea.

While a large sample of specimens of the related

Alloblennius jugularis was available from the coast

of Ethiopia, only relatively few specimens were

available from the coast of Israel. We noted no sexual dimorphism or geographic variation in meristic

characters such as occurs in A. pictus.

Alticus Commerson

FICURES 9, 15c, 19

Alticus Commerson in Lacepede, 1800, p. 479 [type-species:

A. saltatorius Commerson in Lacepede, zz Blennius saliens

Forster, 1788, by monotypy].

Rupiscartes Swainson, 1839, pp. 79, 182 275 [type-spedes:

Salarias alticus Valenciennes in Cuvier and Valenciennes,

1836, — Blennius saliens Forster, 1788, by monotypy].

Basilisciscartes Fowler, 1939, p. 2 [type-species: Blennius

saliens Forster, 1788, by original designation].

Lophalticus Smith, 1957, p. 889 [type-species: Salarias kirki

Gunther, 1868, by original designation].

Damania Smith, 1959, p. 239 [type-species: Andamia anjouanae Fourmanoir, 1954, by original designation].
NUMBER 73 15

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Anterior dentary canines

absent; posterior dentary canines present or absent, only in males when present. Premaxillary and

dentary teeth freely movable, numbering 215 to

340 in upper jaw and 170 to 290 in lower jaw.

Vomer with or without teeth. Dorsal rays XIII to

XVII, 21 to 23; anal rays II, 24 to 28; segmented

caudal rays 12 or 13, none branched; pectoral rays

14 or 15; pelvic rays 1,3 or 1,4. Terminal anal ray

not bound to caudal peduncle by membrane. Lateral line not consisting of two overlapping, disconnected portions; no scalelike flaps covering lateralline pores. Peroperculomandibular pores without

cirri. A single mid-dorsal supratemporal pore. Upper lip without free dorsal margin. No cup-shaped

fleshy disk or appendage behind lower lip. Gill

membranes free. Occipital crest present. Nuchal

cirri absent; supraorbital and nasal cirri simple or

multifid. Postcleithra consisting of 1—3 small, fragile, well-separated bones on each side. Lateral extrascapular not fused with pterotic. Median ethmoid

ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera." See also discussion and

Table 1 under Praealticus.

Smith (1957, p. 887) proposed the monotypic

genus Lophalticus for Alticus kirki (Giinther),

which was said to differ from other species of Alticus in having vomerine teeth, 14 pectoral rays, and

3 segmented pelvic rays. In the initial stages of our

study we were undecided on whether to recognize

Lophalticus as a genus or a subgenus of Alticus. It

was apparent that the two taxa were closely related,

yet they appeared to differ in a surprisingly large

number of characters. Later, we examined specimens of an Alticus species, which we here describe

as new, that led us to believe that Lophalticus

should be afforded only subgeneric status. A. kirki

differs from the other species of Alticus in having

16 or 17 dorsal spines, as opposed to 13 to 15; 11

precaudal vertebrae, as opposed to 10; the last pair

of pleural ribs on the twelfth vertebra, as opposed

to the tenth or eleventh; 25 to 32 epipleural ribs,

as opposed to 13 to 22; the upper lip entire, as

opposed to crenulate; and teeth on the vomer, as

opposed to absent. Several of the meristic characters

used to distinguish the species of Alticus are correlated and appear to show a trend with kirki merely

the extreme. The presence or absence of vomerine

teeth and lip crenulae are only of specific significance in several genera of Salariini, including the

closely related Praealticus.

The genus Damania was proposed by Smith

(1959, p. 239) for Alticus anjouanae (Fourmanoir), which was said to differ from Alticus in having the dorsal spines filiform and a fleshy knob on

each side of the chin. The "fleshy knob" mentioned

by Smith is only slightly, if at all, less well developed in A. saliens (Forster). The presence of filiform dorsal spines in males of anjouanae will separate that species from other members of the genus,

but we consider the character of only specific importance. Species of the closely related Andamia

also possess filiform dorsal spines and the degree of

elongation varies greatly between species. Chapman

(1951, p. 257) commented on the close relationship

of the two genera, and considered Andamia merely

a specialized extension of Alticus both in structure

and habits. To erect another genus for species of

Alticus with filamentous spines serves no useful

purpose and only obscures the close relationship of

Alticus and Andamia.

As noted by Smith (1959, p. 240) A. anjouanae

may eventually prove identical with A. monochrus

Bleeker, a species that he did not examine. According to Smith, anjouanae differed from monochrus

in having the first, instead of the second, dorsal

spine of males longest, and in lacking canines in

the lower jaw. The presence or absence of canines,

while a valid character, must be used with caution.

The literature is unreliable because most workers

have failed to realize that in those species that have

canines, they are present only in the males. In a

single large collection of Alticus saliens from Samoa

(USNM 115446), dentary canines were present

in all males (27) and absent in all females (75).

DISTRIBUTION.—Red Sea; Indian Ocean; central

and western Pacific Ocean.

Alticus simplicirrus, new species

FlCURE 9

HOLOTYPE.-BMNH 1926.7.12.40 (adult male,

46.8 mm SL), Marquesas Islands, Fatu Hiva, lat.

10°26'S, long. 138°39'W, (presented to the British
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Museum by the Scientific Expeditionary Research

Association, S. Y. "St. George").

PARATYPES.-BMNH 1912.7.12.41 (male, 29.2)

and USNM 203829 (female, 31.8), collected with

the holotype.

DESCRIPTION.—(Meristic and proportional characters for holotype are given first, followed in parentheses by those of the male and then female paratypes when at least one specimen differed from the

others). Dentary an open capsule with replacement

teeth entering functional series through adjacent

excavated area in jaw bone. Dentary canines absent. Premaxillary and dentary teeth freely movable; premaxillary teeth ca. 300; dentary teeth ca.

250. Vomerine teeth absent. Terminal vertebra

with two epurals and autogenous hypural 5. Vertebrae 10 + 31; epipleural ribs ca. 22; last pleural

rib on vertebra 11. Circumorbital bones 5.

Dorsal spines 13; last spine moderately reduced

(see proportional measurements); segmented rays

23 (23;24); basal three-fourths of terminal dorsal

ray bound by membrane to caudal peduncle; dorsal fin strongly incised between spinous and rayed

portions. Anal spines 2; segmented rays 26; terminal anal ray not bound by membrane to caudal

peduncle; interradial membranes of anal fin deeply

incised. Caudal fin with 13 segmented rays, none

branched; dorsal procurrent rays 7, ventral procurrent rays 6 (6;5). Pelvic rays 1,3. Pectoral rays

14. Gill-rakers 18 (-;16). Pseudobranchial filaments 8 (5;6). Nuchal cirri absent; supraorbital

cirri shorter than one-third diameter of orbit, simple (except on left side of holotype, where cirrus

bears a minute branch at its tip); nasal cirri palmate, irregularly branched, left with 6 (4;2) and

right with 5 (2;1) branches. Fleshy occipital crest

moderately well developed in males. Upper lip

crenulae 30 (28;29); crenulae present on lower lip,

except laterally (difficult to count).

FIGURE 9.—Alticus simplicirrus, para type, USNM 203829, female, 31.8 mm, Marquesas Islands.

PROPORTIONAL MEASUREMENTS (as percent SL) .—

Greatest distance from snout tip to posterior margin of opercle 18.8 (21.6;18.8); horizontal fleshy

orbital diameter 6.6 (6.8;6.6); fleshy interorbital

width 1.5(2.6;2.7); orbital cirrus length 1.7(1.7;2.2);

nasal cirrus length 1.3 (1.0; 1.3); insertion of last

dorsal spine to mid-caudal base 7.3 (7.8;7.5); first

dorsal spine length 11.0(7.5;6.6); second 13.3

(8.5;7.8); third 13.9(9.2;8.1); fifth 12.0 (8.9;7.5);

tenth 8.3 (5.8;5.0); thirteenth 3.2 (2.0;2.2); none

filamentous; first dorsal ray length 9.0 (6.8;6.0);

fifth 10.9 (7.5;7.2); tenth 11.8 (8.2;7.8); last 6.8

(4.1;3.5); longest caudal ray length 25.9 (23.6;22.6).

PIGMENTATION OF HOLOTYPE.—Lateral and anterior portions of head with numerous fine, pale

spots against dusky background. Indication of large

pale spot on side of head posterior and ventral to

eye with less distinct pale area extending dorso-
NUMBER 73 17

posteriorly from anterior end of large pale spot to

origin of dorsal fin. Dorsal surface of head and eyes

dusky with pale spots in interorbital region. Ventral surface of head pale. Sides of body with numerous irregularly vertical, dusky bands separated

by narrow, white interspaces. Bands becoming indistinct on caudal peduncle. A row of indistinct

white spots along mid-portion of body. Venter pale.

Dorsal fin evenly dusky with narrow, clear areas

basally outlining posterior margins of some spines

and rays; distal tips of most fin elements pale, giving appearance of narrow, pale margin to fin. Anal

fin similarly marked as dorsal, but with distal third

of rays pale. Caudal fin dusky basally, membrane

otherwise mostly clear with dusky areas extending

length of each ray. Pectoral fin similarly marked as

caudal, but fleshy base dusky with pale spots extending onto upper fin rays. Pelvic fins pale dusky.

The female paratype differs most noticeably from

the holotype in having the anal fin completely

clear. In the male paratype the anal fin is marked

with irregular dusky spots.

REMARKS.—Alticus simplicirrus differs from all

previously described species of Alticus in having

simple orbital cirri, as opposed to pinnately

branched cirri; in having 13 segmented caudal

rays, as opposed to 12; and in having the last pair

of pleural ribs on the eleventh vertebra, as opposed

to the tenth or twelfth. It differs from Alticus kirki

most noticeably in having 13 dorsal spines, as opposed to 16 or 17, and in having the lower lip

crenulate as opposed to entire. From all other species of Alticus it differs in having three segmented

pelvic rays, as opposed to four, and in having 14

pectoral rays, as opposed to 15.

ETYMOLOGY.—From the Latin meaning simple

cirrus, referring to the simple nature of the orbital

cirri.

Andamia Blyth

FIGURES 10, 20

Andamia Blyth, 1858, p. 270 [type-species: A. expansa Blyth,

1858, r= Salarias heteropterus Bleeker, 1857a, by monotypy].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Anterior and posterior

dentary canines absent. Premaxillary and dentary

teeth freely movable, numbering ca. 350 to 380 in

upper jaw and ca. 275 to 300 in lower jaw. Vomer

toothless. Dorsal rays XIV to XVII, 17 to 21; anal

rays II, 23 to 25; segmented caudal rays 11 or 12,

none branched; pectoral rays 15; pelvic rays 1,4.

Terminal anal ray bound to caudal peduncle by

membrane (except in A. reyi). Lateral line not

consisting of two overlapping, disconnected portions; no scalelike flaps covering lateral-line pores.

Preoperculomandibular pores without cirri. A

single mid-dorsal supratemporal pore. Upper lip

without free dorsal margin. A cup-shaped fleshy

disk or appendage behind lower lip. Gill membranes free. Occipital crest present or absent.

Nuchal cirri absent; supraorbital cirri pinnately

branched; nasal cirri simple. Postdeithra consisting

of 1—3 small, fragile, well-separated bones on each

side. Lateral extrascapular not fused with pterotic.

Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

Alticus and Andamia are quite closely related.

One might unite these two genera and recognize

them as only subgenera. We maintain the separation because of the unique cup-shaped labial disk

present in Andamia (Figure 10) and because both

genera are well entrenched in the literature.

disk

FIGURE 10.—Ventral view of head showing labial disk of

Andamia heteroptera, USNM 137867, male, 53.5 mm, Porongpong Island, Philippines (see also Figure 20).
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Andamia rcyi, the least specialized member of

the genus, differs from its congeners and agrees with

the closely related Alticus in having the last anal

ray not bound to the caudal peduncle and in having an occipital crest in males (less well developed

than in Alticus, however). As noted by Roa and

Hora (1938, p. 400), Andamia roai (= reyi) is

more aquatic in its habits than A. heteroptera.

DISTRIBUTION.—Eastern Indian Ocean; central

Pacific Ocean.

Antennablennius Fowler

FIGURES la, 47, 48

Antennablennius Fowler, 1931, p. 248 [type-species: Blennius

hypenetes Klunzinger, 1871, by original designation].

Croaltus Smith, 1959, p. 247 [type-species: Blennius bifilum

Ciinther, 1861, by original designation].

Litanchus Smith, 1959, p. 248 [as a subgenus of Antennablennius; type-species: Antennablennius velifer Smith, 1959,

by original designation].

DIAGNOSIS.—Dentary a closed capsule with replacement teeth entering functional series through

foramina in jawbone. Anterior dentary canines

absent; posterior dentary canines present (minute) or absent. Premaxillary and dentary teeth

immovable or nearly so, numbering 32 to 42 in

upper jaw and 26 to 34 in lower jaw. Vomer

toothless. Dorsal rays XII or XIII, 17 to 21;

anal rays II, 19 to 23; segmented caudal rays 13,

middle 9 branched; pectoral rays 14; pelvic rays

I, 3. Terminal anal ray bound to caudal peduncle

by membrane. Lateral line not consisting of two

overlapping disconnected portions; no scalelike

flaps covering lateral-line pores. Preoperculomandibular pores without cirri. Mid-dorsal supratemporal pores 1—4 (typically 3 in subgenus Antennablennius and usually 2 in subgenus Croaltus).

Upper lip without free dorsal margin. No cupshaped fleshy disk or appendage behind lower lip.

Gill membranes free. Occipital crest present or absent. Nuchal and nasal cirri simple; supraorbital

cirri absent. Postcleithra consisting of two elongate

bones, head of ventral element overlapping ventral

end of dorsal element. Lateral extrascapular not

fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

Fraser-Brunner (1951) described several new species of Antennablennius and included a key to the

species. Subsequently, Smith (1959) described one

additional new species and revised the work of

Fraser-Brunner.

Antennablennius bifilum (Giinther) differs from

other members of the genus in possessing nuchal

cirri that are set close together, their bases nearly

touching, and in usually having 2 instead of 3 pores

at the mid-dorsal point of the supratemporal sensory canal (Figure 48). In 54 specimens examined

1 had one, 34 had two, 18 had three, and 1 had

four pores in this position. The nuchal cirri are

greatly elongated in males of A. bifilum. Smith

(1959, p. 247) proposed the monotypic genus Croaltus for bifilum, based only on the relative lengths

of the nuchal cirri. In view of the close relationship

of Antennablennius to Alloblcnnius, we feel that

Croaltus should be afforded only subgeneric status.

At least in the males of some species of Antennablennius a cutaneous flap is present anteriorly on

the first dorsal spine. For those species Smith (1959)

erected the subgenus Litanchus. A. bifilum also

possesses a cutaneous flap on the first dorsal spine.

The degree of development is variable in those

species that have the flap, being well developed in

an apparently undescribed species (USNM 201-

868) that we have examined, and only slightly

developed in A. velifer Smith. In at least the males

of some species a well-developed fleshy occipital

crest is present, while in other species both sexes

are without a crest. We question the significance

attributed to the cutaneous flap on the first dorsal

spine, while ignoring the presence or absence of an

occipital crest as a generic group character.

DISTRIBUTION.—Restricted to the Red Sea and Indian Ocean.

Atrosalarias Whitley

FIGURE 27

Atrosalarias Whitley, 1933, p. 93 [type-spedes: Salarias phaiosotna Bleeker, 1855b, = Salarias fuscus Ruppell, 1838, by

original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through

excavated area in jaw bone. Anterior dentary canines absent; posterior dentary canines present.

Premaxillary and dentary teeth freely movable,
NUMBER 73 19

numbering 185 to 226 in upper jaw and 120 to 147

in lower jaw. Vomer toothless. Dorsal rays IX to

XI, 18 to 22; anal rays II, 18 to 21; segmented

caudal rays 10 to 14 (usually 12 or 13), none

branched; pectoral rays 15 to 18 (usually 16);

pelvic rays 1,3 (innermost pelvic ray minute, visible only in cleared and stained material). Terminal anal ray bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping

disconnected portions; no scalelike flaps covering

lateral-line pores. Preoperculomandibular pores

without cirri. A single mid-dorsal supratemporal

pore. Upper lip without free dorsal margin. No cupshaped fleshy disk or appendage behind lower lip.

Gill membranes free. Occipital crest absent. Nuchal,

supraorbital, and nasal cirri simple. Postcleithra

consisting of two elongate bones, head of ventral

element overlapping ventral end of dorsal element.

Lateral extrascapular not fused with pterotic.

Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section title

"Recognition of Genera."

The systematics and distribution of this monotypic genus have recently been reported by Springer

and Smith-Vaniz (1968). We used Salarias fuscus

Riippell, 1838, as the senior specific synonym for

the genus, but gave the date as 1835 (the same as

that given on the title page of the article and the

same to which it has almost always been referred).

We now note that Riippell referred to Cuvier and

Valenciennes (1836) in the paper in which he described S. fuscus. Thus, Salarias ruficaudus Valenciennes, in Cuvier and Valenciennes, 1836, which we

tentatively placed in junior synonymy under S. fuscus, predates S. fuscus. We have since examined the

holotype of S. ruficaudus and can verify that it and

S. fuscus are the same species (and subspecies, both

described from the Red Sea). Sawyer (1952) established the date of publication of that portion of

Ruppell's work referring to blenniids as 1838.

(Note: the Ruppell paper that we refer to as 1830

is, as far as we know, usually referred to as 1828.

In the Division of Fishes, USNM, however, there

is a copy of this study in the original dated wrappers. These wrappers show that this study appeared

in parts with dates of 1828, 1829, and 1830. All the

blenniid accounts appeared in a part dated 1830.)

In view of this, the two subspecies we (1968)

recognized in Atrosalarias might be referred to as

A. r. ruficaudus and A. r. holomelas. We had called

the latter subspecies A. fuscus holomelas, but because S. fuscus has always been used as a synonym

senior to S. ruficaudus in all studies where both

species names have been included, because we know

of no study using S. ruficaudus as other than a listed

name referring back to Cuvier and Valenciennes

(1836)—and the most recent of these is Giinther

(1861)—because the species is a common one that

for many years has been recognized as fuscus, because the genus has been revised recently, and

because we believe only confusion would result if

the name we accepted as a senior synonym were

changed, we retain the name Salarias fuscus Riippell for the monotypic genus Atrosalarias. In doing

this we comply with the spirit of Rule 23b of the

International Code of Zoological Nomenclature.

Since publication of our revision we have examined specimens of A. fuscus holomelas in the California Academy of Sciences that extend the known

distribution of the subspecies. These specimens are

reported here under their George Vanderbilt Foundation register numbers (localities abbreviated):

South Viet Nam, numbers 2116, 2789, 2790; Jokaj

Island, off Ponape Island, number 497; Palaus,

Koror Island, number 1412. The last two localities

refute our postulation that the specimens from

Boston Island, Marshall Islands, might be an

erroneous locality record.

In addition to the distinguishing characters listed

in the Key and Tables 2 to 6, mature males of

Atrosalarias differ from Salarias in having the spines

and anterior rays of the anal fin frequently rugose,

and the anterior anal rays never elongated.

DISTRIBUTION.—Red Sea; Indian Ocean; western

and central Pacific Ocean.

Cirripectes Swainson

FIGURES 31, 32

Cirripectes Swainson, 1839, pp. 79, 80, 182, 275 [type-spedes:

Salarias variolosus Valenciennes in Cuvier and Valendennes,

1836, by monotypy].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Anterior dentary canines

absent; posterior dentary canines present. Premaxillary and dentary teeth freely movable, numbering

180 to 270 in upper jaw and 85 to 134 in lower jaw.
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Vomer toothless. Dorsal rays XII, 14 to 16; anal

rays II, 14 to 16 (rarely 14); segmented caudal rays

13, middle 9 branched; pectoral rays 14 to 16 (usually 15); pelvic rays 1,3 or 1,4. Terminal aflal ray

not bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps covering lateralline pores. Preoperculomandibular pores without

cirri. Mid-dorsal supratemporal pores numerous,

relatively small, and difficult to count. Upper lip

without free dorsal margin. No cup-shaped fleshy

disk or appendage behind lower lip. Gill membranes free. Occipital crest absent. Nuchal cirri

consisting of an elongate, transverse series of cirri

continuous across nape or interrupted at midline

of nape by a narrow hiatus no greater than 25

percent length of base of either patch of cirri;

supraorbital cirri simple or multifid; nasal cirri

multifid. Postcleithra consisting of two elongate

bones, head of ventral element overlapping ventral

end of dorsal element. Lateral extrascapulaf not

fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

Schultz and Chapman (1960, p. 307) discussed

the problem of the valid spelling of this genuS and

showed that the correct spelling is Cirripectes, not

Cirripectus.

Schultz and Strasburg (1953) discussed the JndoPacific species of Cirripectes, Strasburg (1956) reported on the Hawaiian species, and Smith (1959)

treated the western Indian Ocean species. Our examination of recent collections of Cirripectes

housed in the National Museum of Natural History

indicates that this genus needs additional studyThe only characters that we know of that will

distinguish the eastern Pacific genus Scartichthys

from Cirripectes are the number of rays in the median fins (see Table 4) and the position and development of the nuchal cirri. In Cirripectes the

elongate, transverse series of cirri is continuous

across the nape or is interrupted at the midline of

the nape by a narrow hiatus no greater thart 25

percent of the length of the base of either patch

of cirri, and the individual cirri are usually unbranched and arranged in a linear series. In Scartichthys the patch of cirri on each side of the nape

is separated by a wide hiatus equal to or greater

than the length of the base of either patch of cirri

and the individual cirri are frequently branched,

and their arrangement is often irregular.

The data at hand suggest that Scartichthys and

Cirripectes may be combined when the species are

better known. We prefer to maintain the status quo

until the group can be studied in greater detail.

DISTRIBUTION.—Red Sea; Indian Ocean; western

and central Pacific Ocean to Easter Island.

Crossosalarias, new genus

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Anterior dentary canines

absent; posterior dentary canines present. Premaxillary and dentary teeth freely movable, numbering

about 150 in upper jaw and 85 to 100 in lower jaw.

Vomer toothless. Dorsal rays XII, 16 to 18; anal

rays II, 18 to 20; segmented caudal rays 13, middle

9 branched; pectoral rays 15; pelvic rays I, 3. Terminal anal ray bound by membrane to caudal

peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps

covering lateral-line pores. Cirri associated with 2

to 4 preoperculomandibular pores on each side.

Mid-dorsal supratemporal pores 3. Upper lip without free dorsal margin. No cup-shaped fleshy disk

or appendage behind lower lip. Gill membranes

joined to isthmus at about, or slightly below, level

of ventralmost pectoral ray (posterior edge of gill

membrane occasionally with a narrow free fold,

depth no greater than diameter of pupil, extending

across isthmus). Occipital crest absent. Nuchal cirri

palmate, with short side branches; supraorbital and

nasal cirri multifid. Postcleithra consisting of two

elongate bones, head of ventral element overlapping ventral end of dorsal element. Lateral extrascapular not fused with pterotic. Median ethmoid

ossified. Circumorbital bones 5. Type-species: Crossosalarias macrospilus, new species.

Relationships are discussed under section titled

"Recognition of Genera."

ETYMOLOGY.—A combination of the Greek krosso,

fringe or tassel, referring to the cirri associated with

the preoperculomandibular pores, and Salarias, the

genus that Crossosalarias most closely resembles.
NUMBER 73 21

FIGURE 11.—Crossosalarias macrQspilus, holotype, USNM 203096, male, 51.0 mm, Solomon Islands; lower left insert is ventral view of head, lower right insert, dorsal view of head.

Crossosalarias macrospilus, new species

FIGURES 11, 26

HOLOTYPE.-USNM 203096 (adult male, 51.0 mm

SL), Te Vega Cruise 6, sta. 247, Solomon Islands,

Bougainville, Tautsina Island, east of Kieta Peninsula, north end of island, lat. 6°12'S, long. 155°40'E,

depth ca. 1-8 meters, 11 March 1965, Daniel

M. Cohen.

PARATYPES.-USNM 203764 (2, 54.0-61.1), Te

Vega Cruise 7, sta. 301, Tonga Islands, Nivatobutabu Island, lat. 16°00'S, long. 175°53'W, depth

0-7 meters, 31 May 1965, Rolf L. Bolin; CAS 24154

(1, 55.1), original no. GVF 1750 sta. HK 73, South

China Sea, Pratas Reef, south channel, lat. 20°38'N,

long. 116°45'E, 25 May 1958, Rolf L. Bolin; BPBM

7460 (1, 53.9), Ryukyu Islands, Taketomi, Ishigaki,

coral reef, depth ca. 6 meters, 24 May 1968, John

E. Randall; ANSP 109689 (1, 341), Australia,

Queensland, Endeavor Reef, depth 7.6-18.3 meters,

4 January 1969, J. C. Tyler and C. L. Smith,

TS,A-3.

DESCRIPTION.— (Meristic characters for holotype

in parentheses.) Dentary an open capsule with replacement teeth entering functional series through

excavated area of jaw bone. Posterior dentary canines present. Premaxillary and dentary teeth freely

movable; premaxillary teeth 140 to 158 (158);

dentary teeth 85 to 97 (91). Vomerine teeth absent.

Terminal vertebra with two epurals and autogenous

hypural 5 and ventral hypural plate. Vertebrae 10

or 11 (10) + 24 to 26 (24) = 34 to 37 (34); epipleural

ribs 13 to 15; last pleural rib on vertebra 11. Circumorbital bones 5.

Dorsal spines 12; last spine noticeably reduced;

fleshy flap on anterior border of first spine; segmented rays 16 to 19 (16); basal three-fourths of

terminal dorsal ray bound by membrane to procurrent caudal rays; dorsal fin deeply incised between spinous and rayed portions. Anal spines 2;

segmented rays 18 to 20 (18) ; basal two-thirds of

terminal anal ray bound by membrane to caudal

peduncle; interradial membranes of anal fin deeply

incised. Caudal fin with 13 segmented rays, middle

9 each branched; dorsal procurrent rays 7, ventral

procurrent rays 6 or 7. Pelvic rays 1,3, innermost

closely bound for entire length to middle ray. Pectoral rays 15. Pectoral radial formula 2—0-2 (as

determined from radiographs). Gill-rakers 19 to

22 (19). Pseudobranchial filaments 7 to 9 (7). Gill

membranes joined to isthmus at about, or slightly

below, level of ventralmost pectoral rays (posterior
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

edge of gill membrane occasionally with a narrow

free fold, depth no greater than diameter of pupil,

extending across isthmus). Nuchal cirri palmate,

with 4 to 13 short branches; supraorbital cirri

conspicuous, with 4 to 8 short branches; nasal cirri

short, with 2 to 5 branches. Upper and lower lips

entire; corner of lower lip with a fleshy barbel

about one-half diameter of pupil. Some pores of

infraorbital series in horizontal pairs; cirri associated with 2 to 4 preoperculomandibular pores on

each side. Lateral line of simple pores, continuous

to point below penultimate dorsal spine.

PROPORTIONAL MEASUREMENTS.— (Expressed as percent of SL; figures in parentheses are for the paratypes, except smallest, in order of size, the largest

paratype first). Longest caudal ray 23.7 (20.8;25.0;

25.0;23.0); head length 23.7 (21.8;22.7;21.5;23.3);

horizontal bony orbital diameter 7.1(6.5;6.8;6.3;7.4);

bony interorbital width 1.9 (1.3;1.5;1.2;1.6); supraorbital cirrus length 2.9 (3.9;2.7;2.6;3.3); penultimate dorsal spine length 11.0 (9.7;9.6;11.5;9.7); ultimate spine 3.9(2.8;3.4;4.6;4.1); first dorsal ray

length 16.9(12.1;14.2;13.9;12.1); height of interradial membrane between ultimate spine and first

dorsal ray 3.1 (3.0;3.1;3.5;2.6).

PIGMENTATION.—The following description is

based on the holotype. Body and head peppered

with fine melanophores, becoming concentrated

anteriorly as dark reticulations, especially beneath

appressed pectoral fin, and on opercle and interorbital region. Sides posteriorly with pale spots

(ocelli ?) approximately equal to diameter of pupil. Nape with a conspicuous black spot, approximately equal to eye diameter, extending from

anterior margin of fleshy flap on first dorsal spine

to base of second dorsal spine. Base of each pectoral

fin with heavy concentration of melanophores.

Background color of throat and venter white. Posterior half of throat with two conspicuous black

gular patches united at their bases. Band of melanophores behind median half of lower lip with pale

area at symphysis of lower jaw. All fin rays with

dark, somewhat regularly spaced, cross bands; dorsal and caudal fins with melanophores extending

onto interradial membranes adjacent to markings

on rays. Interradial membranes of anal fin almost

uniformly dusky.

In most features of pigmentation the paratypes of

Crossosalarias macrospilus agree with the holotype.

In the two Tonga specimens the gular markings are

less distinct and the predorsal spot is smaller. The

latter difference is perhaps correlated with the

less developed fleshy flap on the first dorsal spine

in these specimens. In the male Tonga specimen

the distal margin of the anal fin is noticeably more

heavily pigmented than in the female.

RELATIONSHIPS.—Relationships are discussed under "Recognition of Genera" section. Superficially

C. macrospilus most closely resembles Salarias fasciatus. In addition to the distinguishing characters

given in the diagnosis and key, S. fasciatus differs in

lacking a conspicuous predorsal spot and in having

a poorly developed notch in the dorsal fin.

ETYMOLOGY.—The specific name, macrospilus, is

derived from the Greek makros, large, and spilos,

a spot, referring to the conspicuous, black, predorsal

spot characteristic of the species. It is to be treated

as an appositional noun.

Ecsenius McCulloch

FIGURES 17, 18

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

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

namiyei Jordan and Evermann, 1903, by original designation].

Anthiiblennius Starck, 1969, p. 1 [as a subgenus of Ecsenius;

type-species; Ecsenius (Anthiiblennius) midas Starck, 1969,

by original designation].

DIAGONSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Anterior and posterior

dentary canines present (anterior canines slightly

broader and more robust than adjacent incisors).

Premaxillary and dentary teeth freely movable to

nearly immovable, numbering 26 to 148 in upper

jaw and 13 to 64 in lower jaw. Vomer toothless.

Dorsal rays XI to XIV, 13 to 21; anal rays II, 13 to

23; segmented caudal rays 13 to 15, none branched;

pectoral rays 12 to 15 (rarely 12); pelvic rays 1,3

(innermost ray well developed to minute, usually

visible only in cleared and stained material). Terminal anal ray bound by membrane to caudal

peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps

covering lateral-line pores. Preoperculomandibular

pores without cirri. A single mid-dorsal supratem-
NUMBER 73 23

poral pore. Upper lip without free dorsal margin.

No cup-shaped fleshy disk or appendage behind

lower lip. Gill membranes free. Occipital crest absent. Nuchal and supraorbital cirri absent; nasal

cirri simple or paired. Postcleithra consisting of two

elongate bones, head of ventral element overlapping

ventral end of dorsal element (subgenus Ecsenius)

or postcleithra consisting of 2 small, fragile, wellseparated bones on each side (subgenus Anthiiblennins). Lateral extrascapular fused with pterotic.

Median ethmoid unossified. Circumorbital bones 4.

Relationships are discussed under section titled

"Recognition of Genera."

The most recent revision of Ecsenius is that of

Springer (1971). An interesting new subgenus and

species, Ecsenius (Anthiiblennins) midas, was recently described by Starck (1969). Anthiiblennius

differs from all other species of Ecsenius primarily

in having fewer incisoriform teeth, 26 to 34 in the

upper jaw, as opposed to 105 to 148; and 13 to 18

in the lower jaw, as opposed to 30 to 64.

DISTRIBUTION.—Red Sea; Indian Ocean; central

and western Pacific Ocean.

Entotnacrodus Gill

FIGURES 35, 36

Entomacrodus Gill, 1859, p. 168 [type-species: E. nigricam

Gill, 1859, by monotypy].

Salarichthys Guichcnot, 1867, p. % [type-spedes: Salarias

vomerinus Valenciennes in Cuvier and Valenciennes, 1836,

by original designation].

Giffordella Fowler, 1932a, p. 14 [type-species: G. corneliae

Fowler, 1932, by original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Anterior dentary canines

absent; posterior dentary canines present. Premaxillary and dentary canines freely movable, numbering

116 to 224 in upper jaw and 92 to 170 in lower

jaw. Vomer with small conical teeth. Dorsal rays

XII to XIV, 13 to 18; anal rays II, 14 to 19; segmented caudal rays 13, middle 9 branched. Pectoral

rays 12 to 15 (usually 14); pelvic rays 1,4. Terminal

anal ray not bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps covering lateral-line pores. Preoperculomandibular

pores without cirri. Mid-dorsal supratemporal pores

3 to more than 50. Upper lip with free dorsal margin. No cup-shaped fleshy disk or appendage behind

lower lip. Gill membranes free. Occipital crest absent. Nuchal cirri present or absent; supraorbital

and nasal cirri simple to multifid. Postcleithra consisting of two elongate bones, head of ventral element overlapping ventral end of dorsal element.

Lateral extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

The genus Entomacrodus has recently been revised by Springer (1967). This genus appears to

be most closely related to Stanulus, but is easily

distinguished by the characters given in the key.

Two nominal forms, missed by Springer, and their

proper allocation are given in the list of nominal

species of the Salariini.

DISTRIBUTION.—Circumtropical.

Exallias Jordan and Evermann

FlCURE 30

Exallias Jordan and Evermann, 1905, p. 503 [type-spedes:

Salarias brevis Kner, 1868, by original designation].

Gloriella Schultz, 1941, p. 17 [type-spedes: Cirripectes caninus

Herre, 1936 = Salarias brevis Kner, 1868a and b, by original designation].

Leoblennius Reid, 1943, p. 282 [type-spedes: L. schultzi Reid,

1943 = Salarias brevis Kner, 1868, by original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Anterior and posterior

dentary canines absent. Premaxillary teeth freely

movable, numbering 194 to 210; dentary teeth fairly

rigid numbering 52 to 62. Vomer toothless. Dorsal

rays XII, 12 or 13; anal rays II, 14; segmented

caudal rays 13; middle 9 branched; pectoral rays

15; pelvic rays 1,4. Terminal anal ray not bound by

membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions;

no scalelike flaps covering lateral-line pores. Preoperculomandibular pores without cirri. Mid-dorsal

supratemporal pores numerous, relatively small and

difficult to count. Upper lip without free dorsal

margin. No cup-shaped fleshy disk or appendage

behind lower lip. Gill membranes free. Occipital

crest absent. Nuchal cirri consisting of an elongate,

transverse series of cirri interrupted at midline of
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

nape by a narrow hiatus; supraorbital and nasal

cirri multifid. Postcleithra consisting of two elongate bones, head of ventral element overlapping

ventral end of dorsal element. Lateral extrascapular

not fused with pterotic. Median ethmoid ossified.

Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

This apparently monotypic genus appears to be

most closely related to Cirripectes.

DISTRIBUTION.—Red Sea; Indian Ocean; central

and western Pacific Ocean.

Glyptoparus Smith

FIGURE 41

Glyptoparus Smith, 1959, p. 249 [type-species: G. delicatulus

Smith, 1959, by original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jawbone. Anterior dentary canines

absent; posterior dentary canines present, usually

2 or 3 posterior canines on each dentary. Premaxillary and dentary teeth fairly rigid, 74 to 80 in upper jaw and 50 to 58 in lower jaw. Vomer toothless.

Dorsal rays XII or XIII, 15 or 16; anal rays II, 18;

segmented caudal rays 13, middle 9 branched; pectoral rays 13; pelvic rays 1,3 (the innermost pelvic

ray reduced to a few fragments of bone visible only

in cleared and stained material). Terminal anal

ray bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps covering

lateral-line pores. Preoperculomandibular pores

without cirri. Mid-dorsal supratemporal pores 1 or

2. Upper lip without free dorsal margin. No cupshaped fleshy disk or appendage behind lower lip.

Gill membranes free. Occipital crest present. Nuchal

and nasal cirri simple; supraorbital cirri simple

(minute) or absent. Postcleithra consisting of two

elongate bones, head of ventral element overlapping ventral end of dorsal element. Lateral extrascapular not fused with pterotic. Median ethmoid

ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

The description of Glyptoparus was based only on

specimens from the western Indian Ocean. We find

that Glyptoparus is also generally distributed

throughout the central Pacific.

The characteristic color pattern of Glyptoparus

easily distinguishes it from all other species of

blenniids that we have examined. Especially diagnostic is the narrow, dusky crescent extending across

the throat to the corners of the mouth, and the

dark spot on each side of the gill membranes just

anterior to the base of the pelvic fins.

DISTRIBUTION.—Indian Ocean; central and western Pacific Ocean.

Hirculops Smith

FIGURE 46

Hirculops Smith, 1959, p. 247 [type-spedes: Blennius cornifer

Riippcll, 1829, by original designation].

DIAGNOSIS.—Dentary a closed capsule with replacement teeth entering functional series through

foramina in jaw bone. Premaxillary open mesially,

but closed laterally with replacement teeth entering

functional row through foramina in the bone. Anterior dentary canines absent; posterior dentary

canines present. Premaxillary and dentary teeth

immovable or nearly so, numbering 34 to 44 in

upper jaw and 22 to 26 in lower jaw. Vomer with

small conical teeth. Dorsal rays XII, 20 or 21; anal

rays II, 20 to 23; segmented caudal rays 13, middle

9 branched; pectoral rays 14; pelvic rays I, 3. Terminal anal ray bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps covering lateral-line pores. Preoperculomandibular pores

without cirri. Mid-dorsal supratemporal pores 8—15.

Upper lip without free dorsal margin. No cup

shaped fleshy disk or appendage behind lower lip.

Gill membranes free. Occipital crest absent. Nuchal,

supraorbital, and nasal cirri simple. Postcleithra

of two elongate bones, head of ventral element

overlapping ventral end of dorsal element. Lateral

extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

The genus Hirculops appears to be most closely

related to Rhabdoblennius, but it is easily distinguished by the characters given in the key. Smith

(1959) recognized a single species with two sub-
NUMBER 73 25

species: one in the Red Sea and one in the western

Indian Ocean. He did not specify the differences

between the subspecies except to say that they differed "in certain markings from Klunzinger's detailed notes, and as Red Sea forms are quite often

unique, the African form here kept subspecifically

distinct."

DISTRIBUTION.—Restricted to the Red Sea and

western Indian Ocean.

Istiblennius Whitley

FICURES 15d, 22-25

Istiblennius Whitley, 1943, p. 185 [type-species: Salarias

muelleri Klunzinger, 1880, by original designation].

Blenniella Reid, 1943, p. 383 [type-species: B. rhessodon Reid,

1943 — Salarias gibbifrons Quoy and Gaimard, 1824, by

original designation].

Halmablennius Smith, 1948, p. 340 [type-species: Salarias

unicolor Riippell, 1838, by original designation].

Alticops Smith, 1948, p. 340 [type-species: Salarias periophthalmus Valenciennes in Cuvier and Valenciennes, 1836,

by original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through

excavated area in jaw bone. Anterior dentary canines absent; posterior dentary canines present or

absent. Premaxillary and dentary teeth freely movable, numbering 205 to 225 in upper jaw and 165

to 195 in lower jaw. Vomer toothless. Dorsal rays

XIII or XIV, 18 to 24; anal rays II, 17 to 25; segmented caudal rays 13, middle 9 branched; pectoral

rays 12 to 15 (rarely 15); pelvic rays 1,3 or 1,4. Terminal anal ray not bound by membrane to caudal

peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps

covering lateral-line pores. Preoperculomandibular

pores without cirri. Three (0-2 in only one species)

or more mid-dorsal supratemporal pores. Upper lip

without free dorsal margin. No cup-shaped fleshy

disk or appendage behind lower lip. Gill membranes free. Occipital crest present or absent.

Nuchal cirri present or absent; supraorbital and

nasal cirri simple to multifid. Postcleithra consisting of two elongate bones, head of ventral element

overlapping ventral end of dorsal element. Lateral

extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

In his treatment of the western Indian Ocean

Blenniidae, Smith (1959) distinguished Halmablennius from Istiblennius solely on the type of

supraorbital cirri present in the two groups. In

Halmablennius the supraorbital cirri were described

as being "bushy," while in Istiblennius the cirri

were described as being simple or very weakly

branched. In light of the variation frequently encountered in other genera (for instance Entomacrodus), we do not believe that this difference is sufficient for generic recognition. The type-species of

Halmablennus, I. unicolor, however, does differ

from all other species of Istiblennius (including

those with bushy supraorbital cirri) that we examined in having 1,4 pelvic rays rather than 1,3.

Whether this difference will merit recognition as

a valid generic group character when Istiblennius

is revised is open to question.

In our examination of the material available for

this study we find that Istiblennius gibbifrons is

one of the most distinctive members of the genus.

In the type collection of the National Museum of

Natural History alone this species is represented

by the following junior synonyms: Salarias rutilus,

Istiblennius afilinuchalis, and Istiblennius rodenbaughi. If Istiblennius gibbifrons is indicative of

the amount of synonymy that occurs in Istiblennius

(and we believe it is), then the actual size of the

genus is considerably smaller than the list of 102

nominal species (see list) would suggest.

Istiblennius gibbifrons is readily distinguished

from all other species in the genus by the following

combination of characters: 12 precaudal vertebrae,

no nuchal cirri, no fleshy occipital crest, simple

supraorbital cirri, mid-dorsal area of supratemporal

canal with 6 to 12 or more pores, and the lateral

line with numerous, regularly spaced side branches.

Istiblennius coronatus differs from gibbifrons primarily in having the supraorbital cirri usually

branched, 3 to 8 pores in the corresponding position of the supratemporal canal, and the side

branches of the lateral line less numerous and more

irregularly spaced. Specimens (USNM 142204) that

we tentatively identify as /. paulus, also with 12

precaudal vertebrae (most species have 10 or 11),

have small nuchal cirri, consistently 3 pores in the

mid-dorsal area of the supratemporal canal, and the

lateral line with reduced side branches. In view of

the overlapping combinations of characters shown
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

by species in Istiblenniiis, we hesitate to assess their

importance as generic characters.

Several species of Istiblenniiis, as here defined,

show a remarkable resemblance to species of Praealticus, and there appears to be no single character

(except, perhaps, the nature of the supratemporal

pores) that will distinguish all species of either

genus (see discussion and Table 1 under Praealticus). We urge future workers to consider carefully

all the nominal species in both genera before describing additional species of either Istiblenniiis or

Praealticus.

DISTRIBUTION.—Red Sea; Indian Ocean; central

and western Pacific Ocean.

Litobranchus, new genus

DIAGNOSIS.—Dentary a closed capsule with replacement teeth entering functional series through foramina in jawbone. Anterior dentary canines absent;

posterior dentary canines present (frequently absent in females). Premaxillary and dentary teeth immovable or nearly so, numbering 36 to 42 in upper

jaw and 23 to 30 in lower jaw. Vomer toothless.

Dorsal rays XII, 18 to 20; anal rays II, 20 to 22;

segmented caudal rays 12, none branched; pectoral

rays 14; pelvic rays 1,3. Terminal anal ray bound

by membrane to caudal peduncle. Lateral line not

consisting of two overlapping disconnected portions; no scalelike flaps covering lateral-line pores.

Preoperculomandibular pores without cirri. Middorsal supratemporal pores 2-4, usually 2. Upper

lip without free dorsal margin. No cup-shaped

fleshy disk or appendage behind lower lip. Gill

membranes free. Occipital crest absent. Nuchal,

supraorbital, and nasal cirri simple. Postcleithra

consisting of two elongate bones, head of ventral

element overlapping ventral end of dorsal element.

Lateral extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5. Typespecies: Salarias foxvleri Herre.

Relationships are discussed under section titled

"Recognition of Genera."

ETYMOLOGY.—A combination of the Greek litos,

simple, and branchos, fin, referring to the unbranched rays characteristic of the genus.

An interesting parallel is offered by the relationship of the caudal fin structure and branching of

principal caudal rays in Litobranchus and Medusablcnnius and the unrelated Atrosalarias to their

respective relatives. Litobranchus and Medusablennius agree with their closest allies in having a generalized caudal fin containing an autogenous hypural 5, but differ in having all the caudal rays

unbranched. Atrosalarias also differs from its nearest allies in having all simple caudal rays, but agrees

with most members of the closely related Salarias in

having a specialized caudal fin that lacks an autogenous hypural 5.

FIGURE 12.—Litobranchus fowleri: Top, SU 40246, male, 28.1 mm; bottom SU 30971, female,

28.2 mm. Both specimens from Oriental Negros, Philippine Islands.
NUMBER 73 27

Litobranchus fowleri (Herre)

FIGURES 12, 42

Salaries fowleri Herre, 1936, p. 364, pi. 2, fig. 7 [original

description; type-locality: Philippine Islands, Oriental Negros, tide pool at Dumaqucte].

MATERIAL EXAMINED.—SU 30971 (adult female,

29.9 mm SL), holotype of Salarias fowleri and paratypes (6 [2 not seen], 16.0-28.2) in same bottle,

Philippine Islands, Oriental Negros, tide pool at

Dumaquete, lat. 9°1S"N, long. 123°18'E, 23 April

1936, A. W. C. T. Herre; USNM 203763 (1, cleared

and stained, 27.4), paratype originally with SU

30971; data for following collections same as SU

30971, except for date of collection: SU 29582 (11 [3

not seen], 23.7-34.4), SU 32295 (1, 28.8), SU 40244

(1, 26.4), SU 40246 (1, 28.1), USNM 203768 (1,

30.9), and USNM 203769 (2, including 1 cleared

and stained, 31.0-32.0). ZMA 109.098 (1, 31.0),

original no. Siboga Expedition sta. 33, Indonesia,

Lombok, Pijut Bay, lat. 8°47'S, long. 116°3rE,

depth 9-22 meters, 24/26 March 1899, M. Weber;

ZMA 109.097 (1, 24.8) and 109.099 (2, 25.1-30.2),

New Guinea, Waigeo Island, Saonek Anchorage,

lat. 0°21'S, long. 130°56'E, December 1909, L. F.

de Beaufort; CAS 24687 (3, 18.7-23.6), GVF Reg.

#881, sta. 95, Palau Islands, tide pool on reef off

SW point of Nardueis Island, lat. 7°20'44"N, long.

134°35'08"E, 4 September 1956, H. A. Fehlmann

and A. W. Scott; CAS 24688 (1, 22.0), GVF Reg.

#901, sta. 115, Palau Islands, Peleliu, south side

of channel between Ngurungor Point and Ngusuwal Island, east side of Peleliu, lat. 7°00'36"N, long.

134°16'26"E, depth 0-0.5m, tide pool, rock, and

sand bottom, 19 September 1956, H. A. Fehlmann,

et al.; CAS 24689 (2, 23.9-34.5), GVF Reg. #1600,

sta. 20, Philippine Islands, tide pools at Ayuguiton,

14 km south of Dumaquete City, Oriental Negros,

19 April 1958, W. C. Brown, et al.

DESCRIPTION.— (Meristic characters for holotype

in parentheses). Dentary a closed capsule with replacement teeth entering functional series through

foramina in jawbone. Posterior dentary canines

present (frequently absent in females) . Premaxillary and dentary teeth immovable or nearly so;

premaxillary teeth 36 to 42; dentary teeth 28 to 30.

Vomerine teeth absent. Terminal vertebra with two

epurals and autogenous hypural 5 and ventral hypural plate. Vertebrae 10 + 27 or 28 (usually 28) =

37 or 38; epipleural ribs 13 or 14; last pleural rib

on vertebra 10. Circumorbital bones 5.

Dorsal spines 12; last spine moderately reduced;

segmented rays 18 to 20(19); basal two-thirds of

terminal dorsal ray bound by membrane to caudal

peduncle; dorsal fin moderately incised between

spinous and rayed portions. Anal spines 2; segmented rays 20 to 22 (21); basal two-thirds of terminal anal ray bound by membrane to caudal peduncle. Caudal fin with 12 segmented rays, none

branched; dorsal procurrent rays 5 to 8 (usually 6) ,

ventral rays 4 or 5. Pelvic rays 1,3. Pectoral rays 14.

Pectoral radial formula 2-0-2. Gill-rakers 6 to 8(7).

Pseudobranchial filaments 5. Nuchal, supraorbital,

and nasal cirri simple. Upper and lower lips entire.

Pores of infraorbital series simple. Lateral line

short, continuous to point below 4th to 6th dorsal

spine, pores simple.

PIGMENTATION.—The pigmentation of the sexes

closely approaches that shown in Figure 12. Males

are characterized by 7 dark lateral bands that extend across the dorsum, several rows of pale, horizontally elongated spots on sides, and the fins not

heavily pigmented, except distally on the interradial membranes of the anterior 4 or 5 dorsal spines.

In several of the largest (presumably nuptial)

males the head is very dark. In the females there

are no rows of pale spots and the lateral bands are

less well defined. In contrast to the males, the body

and the dorsal fin are characterized by numerous

dark spots.

GEOGRAPHIC VARIATION.—The three New Guinea

specimens differ in several respects from the other

material and were not used in preparing the above

description. We provisionally assign these specimens to Litobranchus fowleri.

In specimens from the Palaus, Philippines, and

Indonesia, the supraorbital cirri are minute and

there is no apparent sexual dimorphism associated

with these structures. In 7 females (21.8—31.0 mm

SL) the ratio of the cirrus length into the orbital

diameter was 4.3 to 8.5, average 5.6; in 10 males

(22.4-34.0) the ratio was 4.0 to 8.0, average 5.4.

The New Guinean specimens have longer cirri that

clearly reflect sexual dimorphism. In the single

female specimen the cirri/orbital ratio is 1.8; in

both males it is 0.65. In all three specimens the

number of dentary teeth is low, 23 (versus 28 to

30). The number of premaxillary teeth is within
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the range of those shown by the other specimens.

The interradial membranes of the anal fin are

more deeply incised and the anal segmented rays

approximately one-third longer in New Guinean

males than in Philippine males of nearly equal size.

The New Guinean males also differ in having the

body almost completely devoid of pigment, but this

may be the result of fading.

REMARKS.—Chapman (1951, p. 349) reported

Blennius jugularis Klunzinger from the IndoAustralian Archipelago, an area from which we

have no other record of this species. His record was

based on a single female 32 mm long, taken at

Siboga Station 149 (Insel Fau bei Gebe), in the

Amsterdam Museum. The description of the specimen is sufficient to exclude all species of Blenniidae

known to us except Litobranchus fowleri, and the

color description is indicative of females of that

species.

Under the synonymy of B. jugularis, Chapman

recorded "Salarias spec. M. Weber, Siboga Exped.

Fische, 1913, p. 537." Several unidentified species

of Salarias are listed (Weber, 1913, p. 537). No.

22, Salarias spec, of Weber, is a 16 mm specimen

from Siboga sta. 149, and no. 23, Salarias spec, is

recorded as a juvenile [no length given] from sta.

33. The only Siboga specimen of Litobranchus

fowleri presently housed in the Amsterdam Museum

is ZMA 109.098, a 31 mm female from sta. 33 (see

material examined). We suspect that this is the

specimen reported by Chapman and that the discrepancy in locality data was due to a recording

error. Chapman (1951, p. 350) noted that the specimen he identified as B. jugularis was closely related

to fowleri but differed in lacking dentary canines.

Although dentary canines are typically well developed in males of Litobranchus fowleri, they are

frequently absent in females.

Medusablennius Springer

FIGURE 44

Medusablennius Springer, 1966a, p. 56 [type-species: M.

chant Springer, by original designation].

DIAGNOSIS.—Dentary a closed capsule with replacement teeth entering functional series through foramina in jawbone. Anterior and posterior dentary

canines absent. Premaxillary and dentary teeth immovable or nearly so, numbering 17 to 19 in upper

jaw and 16 to 18 in lower jaw. Vomer toothless.

Dorsal rays XI to XIII, usually XII, 17 or 18;

anal rays II, 18 or 19; segmented caudal rays 13,

none branched; pectoral rays 13; pelvic rays 1,3.

Terminal anal ray bound by membrane to caudal

peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps

covering lateral-line pores. Preoperculomandibular

pores without cirri. Mid-dorsal supratemporal pores

2. Upper lip with a free dorsal margin extending

across snout. No cup-shaped fleshy disk or appendage behind lower lip. Gill membranes free. Occipital crest absent. Nuchal cirri absent; supraorbital

cirri simple; complex cirri on rims of anterior and

posterior nostrils and on interorbital region. Postcleithra consisting of two elongate bones, head of

ventral element overlapping ventral end of dorsal

element. Lateral extrascapular not fused with

pterotic. Median ethmoid ossified. Circumorbital

bones 2.

Relationships are discussed under section titled

"Recognition of Genera."

Springer (1968b) included Medusablennius in

his tribe Blenniini. We now find that this action

was erroneous, as even the information on the jaws

contained in the original description was sufficient

to have indicated that Medusablennius is allied to

the Salariini. The removal of Medusablennius from

the Blenniini makes that group more uniform in

anatomical composition: all Blenniini have 5 circumorbital bones and several branched caudal rays.

Medusablennius, unique in the Blenniidae in having

only 2 circumorbital bones, has unbranched caudal

rays, as do several other genera of Salariini (as well

as all genera of Omobranchini and Nemophidinae).

Springer (1968b, p. 56) erroneously stated that

Medusablennius had branched caudal rays and an

autogenous ventral hypural plate. The original

description (Springer, 1966a), which is correct, indicates that the caudal rays are all simple and that

the ventral hypural plate is fused with the urostylar

centrum and upper hypural plate.

Springer (1966a) had only two specimens of

Medusablennius at the time of his description. Two

additional collections, from near the type locality,

of this monotypic genus have since been discovered:

CAS 24741 (2, 12.0-13.5), original no. GVF 80,

sta. 26, Tuamotu Archipelago, Raroia Atoll, outer
NUMBER 73 29

reef channel next to Lithothamnion ridge opposite

Ngarumaoa Village, lat. 16°1'S, long. 142°26/W,

21 July 1952, R. R. Harry (Rofen) et al. and CAS

24742 (6, 12.1-12.9), original no. GVF 113, sta. 59,

collection data same as preceding except date 17

August 1952.

The holotype and largest known specimen of

Medusablennius is a male, 15.4 mm standard length.

CAS 24742 contains a 12.1 mm female with welldeveloped ova. All specimens of Medusablennius

examined have well-developed cirri (Figure 44) at

a size when the cirri of other salariinine species has

scarcely begun to develop. The above leads us to

believe that these specimens of Medusablennius are

mature individuals and that the species easily ranks

as the smallest member of the family Blenniidae.

Mimoblennius, new genus

Undescribcd genus (B) Springer, 1968b, p. 10.

DIAGNOSIS.—Dentary a closed capsule with replacement teeth entering functional series through

foramina in bone. Anterior dentary canines absent;

posterior dentary canines present. Premaxillary and

dentary teeth immovable or nearly so, numbering

30 to 34 in upper jaw and 25 to 30 in lower jaw.

Vomer toothless. Dorsal rays XIII, 16 to 19; anal

rays II, 19 to 22; segmented caudal rays 13, middle

9 branched; pectoral rays 14 to 15 (usually 14);

pelvic rays 1,3 (the innermost pelvic ray is considably reduced, closely applied to the base of the

second ray, and visible only in cleared and stained

specimens or upon careful dissection). Terminal

anal ray bound by membrane to caudal peduncle.

Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps covering

lateral-line pores. Preoperculomandibular pores

without cirri. Mid-dorsal supratemporal pores 3.

Upper lip without free dorsal margin. No cupshaped fleshy disk or appendage behind lower lip.

Gill membranes free. Occipital crest absent. Nuchal

cirri palmate; supraorbital cirri over each eye 2 or

multifid; nasal cirri palmate, multifid. Postdeithra

consisting of two elongate bones, head of ventral

element overlapping ventral end of dorsal element.

Lateral extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5. Typespecies: Blennius atrocinctus Regan.

Relationships are discussed under section titled

"Recognition of Genera."

ETYMOLOGY.—A combination of the Greek mimos,

a mimic, and Blennius, the nominal genus for

which it was originally mistaken and shows the

greatest superficial resemblance.

In general physiognomy Mimoblennius strongly

resembles the genus Blennius (tribe Blenniini),

but differs in the typically salariinine nature of the

premaxillaries. In having two discrete nuchal cirri

the genus differs from all species of Blennius known

to us except B. riodourensis, an eastern Atlantic

species described from Rio d'Ouro, Spanish West

Africa (Metzelaar, 1919, p. 291). In the few other

species of Blennius that have nuchal cirri, the cirri

are numerous and arranged in a median predorsal

row or patch.

The genus is comprised of two species, M. atrocinctus and M. cirrosus. They differ most notably

in the development of their supraorbital and posterior nasal cirri and robustness of head and body

(difficult to define quantitatively but readily a p

parent when specimens of the same size are compared) . In M. atrocinctus each supraorbital cirrus

consists of two cirri that arise from a short base

and the posterior nasal cirri are minute or absent.

In M. cirrosus each supraorbital cirrus typically

consists of 3 or more cirri that arise from a relatively longer common base and the posterior nasal

cirri are prominent palmate flaps with 2 to 5 short

branches. In the former species the head and body

is more robust, tapering posteriorly (Figure 13).

Both species are described in detail below. M. atrocinctus is known only from Hong Kong, Christmas

Island (eastern Indian Ocean), and Ceylon. M. cirrosus is apparently restricted to the Red Sea.

Mimoblennius atrocinctus (Regan)

FIGURES 13a, 40

Blennius atrocinctus Regan, 1909, pp. 404-^405, pi. LXVI,

fig. 1 [original description; type-locality: Christmas Island].

MATERIAL EXAMINED.—BMNH 1909.3.4.70 (juvenile, 21.5 mm SL), holotype of Blennius atrocinctus,

Christmas Island, lat. 10°30'S, long. 105°40'E, 1909,

C. W. Andrews; USNM 203095 (1, 44.2), original

no. GVF 1731, sta. HK 54, Hong Kong, New Territories, north end of Shelter Island, entrance to Port
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 13.—Mimoblennius: a, M. atrocinctus, USNM 204624, male, 36.3 mm, Trincomalee, Ceylon; b, M. cirrosus, holotype, USNM 204491, male, 35.2 mm, Sheikh el Abu, Harat Island,

Ethiopia.

Shelter, lat. 22°20/00"N, long. 114°17'42"E, depth

ca. 1-4 meters, 24 April 1958, Rolph L. Bolin;

USNM 197980 (3, including 1 cleared and stained,

20.1-23.4) and CAS 24155 (2, 24.3-32.4), locality

data same as USNM 203095, 20 August 1963,

W. L. Y. Chan; USNM 204624 (7, 14.5-40.7), Ceylon, Trincomalee, small cove north of harbor, lat.

8°34'N, long. 81°13'E, large rock outcrop devoid

of coral, approx. 50 meters off shore, depth 0-4

meters, 26 June 1969, W. F. Smith-Vaniz, S-V

69-119.

DESCRIPTION.— (Meristic characters for holotype

in parentheses.) Dentary a closed capsule with replacement teeth entering functional series through

foramina in bone. Canine on each dentary posteriorly. Premaxillary and dentary teeth immovable

or nearly so; premaxillary teeth 30 to 34 (32); dentary teeth 25 to 30 (26). Vomerine teeth absent.

Terminal vertebrae with two epurals and autogenous hypural 5 and ventral hypural plate. Vertebrae 10 + 25 to 28 = 35 to 38 (38); epipleural

ribs 15 to 19; last pleural rib on vertebra 10. Circumorbital bones 5.

Dorsal spines 13; last spine noticeably reduced;

segmented rays 16 to 19 (17); basal two-thirds of

terminal dorsal ray bound by membrane to caudal

peduncle; dorsal fin moderately incised between

spinous and rayed portions. Anal spines 2; segmented rays 19 to 21 (20) ; basal two-thirds of terminal anal ray bound by membrane to caudal
NUMBER 73 31

peduncle. Caudal fin with 13 segmented rays, middle 9 each branched; dorsal and ventral procurrent

rays 7. Pelvic rays 1,3. Pectoral rays 14 or 15(14).

Pectoral radial formula 2—0—2 (based on 1 cleared

and stained specimen). Gill-rakers 10 to 11. Pseudobranchial filaments 5. Nuchal cirri palmate, with

4 or 5 short, rounded branches; supraorbital cirri

consisting of 2 long, slender cirri over each eye;

anterior nasal cirri palmate, with 3 to 5 branches,

posterior nasal cirri absent or minute. Upper and

lower lips entire. Lateral line of simple pores ending beneath 4th or 6th dorsal spine or continuous

to point below level of 4th dorsal spine, thence

curving downward and continuous as short, separate, longitudinally bipored tubes to point opposite

tip of median rays of appressed pectoral fin (at

level of 11th dorsal spine).

In most specimens the head and body have a

pale ground color. The pattern of melanophores is

considerably variable. Most specimens bear a dusky

mark along the posterior border of the orbit and

irregular dusky marks over the opercle. The underside of the head and prepelvic region are dusky to

black with occasional evidence of bands under the

head. The side of the body may be irregularly

dusky, darker on the ventral half, with dark spots

at midlevel, or with about 7 vertical dark bands.

The dorsal fin usually bears a dark spot between

the first two rays, usually paler than in M. cirrosus.

There may be dark extensions of the body bands

on to the basal portion of the dorsal fin and sundry other dusky marks over the fin. The anal fin

may be pale dusky, darker at the ray tips, or almost

uniformly black. The caudal fin rays bear dusky

spots aligned in irregular vertical rows. The pectoral fin rays are dusky and the pelvic fin rays are

dusky to almost black.

The life colors were given by Regan (1909, p.

405) as "reddish, with 6 blackish cross-bars continued on the basal part of the dorsal fin; anal

blackish."

GEOGRAPHICAL VARIATION.—See comments under

Mimoblennius cirrosus.

Mimoblennius cirrosus, new species

FIGURES 13ft, 39

HOLOTYPE.-USNM 204491 (adult male, 35.2 mm

SL), Ethiopia, Sheikh el Abu, off lighthouse just

west of south end of Harat Island, lat. 16°08'N,

long. 39°26.5'E, depth to 4 meters, 14 August 1969,

Victor G. Springer, E. Clark, and Z. Zelleke, VGS

69-14.

PARATYPES.-USNM 204645 (14, 21.8-37.0), taken

with the holotype; HUI SLR-270d (4, 25.0-31.7),

Israel, Ophir Bay, 29 August 1967; USNM 204563

(1, 23.9), northwest coast of Gulf of Aqaba, bay

between Marsa Mokrakh and El Himeira, depth

0-3 meters, 15 July 1969, V. G. Springer, et al.,

VGS 69-1; USNM 204564 (1, 29.0), northwest

coast Gulf of Aqaba, bay at El Himeira, depth to

18 meters, 16 July 1969, V. G. Springer, et al., VGS

69-2; USNM 204565 (2, 15.9-19.2), depth to 16

meters, 19 July 1969, V. G. Springer, et al., VGS

69-4; USNM 204566 (I, 23.0), depth 21.3 to 24.4

meters, 9 September 1969, V. G. Springer, et al.,

VGS 69-24; USNM 204569 (6, 26.1-32.5), Red Sea,

Strait of Jubal, southern end of Sinai Peninsula

at Ras Muhammad, depth to 9.2 meters, V. G.

Springer, et al., VGS 69-28.

DESCRIPTION.— (Meristic characters for holotype

in parentheses.) Dentary a closed capsule with replacement teeth entering functional series through

foramina in bone. Canine on each dentary posteriorly. Premaxillary and dentary teeth immovable or

nearly so; premaxillary teeth 30 to 34 (30); dentary

teeth 24 to 30 (28). Vomerine teeth absent. Terminal vertebra with two epurals and autogenous

hypural 5 and ventral hypural plate. Vertebrae

10 + 27 or 28 = 37 or 38(38); epipleural ribs 13

to 16; last pleural rib on vertebra 10. Circumorbital

bones 5.

Dorsal spines 12 or 13 (13); last spine noticeably

reduced; segmented rays 17 to 19 (18); basal twothirds of terminal dorsal ray bound by membrane

to caudal peduncle; dorsal fin moderately incised

between spinous and rayed portions. Anal spines

2; segmented rays 20 to 22 (21); basal two-thirds

of terminal anal ray bound by membrane to caudal

peduncle. Caudal fin with 13 segmented rays, middle 9 each branched; dorsal procurrent rays 5 to

8 (6); ventral procurrent rays 5 to 7 (6). Pelvic rays

I, 3. Pectoral rays 14. Pectoral radial formula 2-0—2

(based on 1 cleared and stained specimen). Gillrakers 10 to 12. Pseudobranchial filaments 5 or 6.

Nuchal cirri palmate, with 3 or 4 broadly rounded

lobes (best developed in large individuals); each

supraorbital cirrus consisting of 1 to 6, typically
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3 or more, well-defined cirri; nasal cirri present on

both anterior and posterior nostrils, posterior cirri

consisting of palmate flaps with 2 to 5 short branches

and anterior cirri consisting of a short raised tube

with 1 or 2 narrow elongate flaps on posterior

margin. Upper and lower lip entire. Lateral line

consisting of simple, short, separate, longitudinally

bipored tubes that end beneath 4th to 6th dorsal

spine.

PIGMENTATION.—Color pattern is extremely variable; most specimens, however, tend to exhibit a

network of dusky melanophores surrounding a few

irregular, pale spots on the head and numerous

small, round, pale spots on the body. The pale spots

in life are red, and on the body form numerous

regular vertical bands. The diameters of the

spots decrease gradually posteriorly on the body.

The bandlike pattern is not always obvious in

preserved specimens. Small specimens may appear

to have the body covered with dusky spots while

large specimens may have some flecklike dark spots

on the sides, and a few specimens, including the

holotype, have dusky spots on the ventral body

contour paralleling the anal fin base. The ventral

side of the head may bear an adumbration of a

dusky transverse band or two bands separated by

a pale area. The prepelvic area and adjacent ventral portion of the head may appear dusky or

unmarked.

The dorsal fin may bear diffuse pale to dark

dusky spots densely to loosely distributed over its

surface. Some specimens exhibit a dark spot between the first two dorsal spines. The caudal and

anal fins are similarly marked to the dorsal, but

never have a distinct dark spot. The anal fin is

frequently darker than the other fins. The pectoral

fins bear fine melanophores along the lengths of

the rays. The pelvic fins are unmarked.

ETYMOLOGY.—The Latin word cirrosus means

bearing cirri.

GEOGRAPHICAL VARIATION.—Frequency distributions for numbers of segmented dorsal and anal

rays and caudal vertebrae were tabulated for specimens of Mimoblennius cirrosus and Af. atrocinctus:

M. cirrosus

Israel

males

females

Ethiopia

males

females

Af. atrocinctus

Ceylon

males

females

Christmas Island

female

Hong Kong

males

females

segmented dorsal

16

1

2

17

2

1

2

3

1

2

rays

18

2

5

6

6

1

1

2

19

3

3

1

segmented anal

19

2

20

2

1

2

5

1

2

rays

21

2

7

5

6

2

1

22

3

1

1

caudal vertebrae

25

1

26

1

1

27

1

5

1

5

3

1

28

4

5

5

3

2

4

These data indicate that there are mean differences

in number of segmented dorsal and anal rays between Israeli and Ethiopian specimens of Af. cirrosus similar to those found between Israeli and Ethiopian specimens of Alloblennius pictus {q-v.).

There is also a suggestion that differences exist

between Hong Kong (South China Sea) and Indian

Ocean specimens of Af. atrocinctus, and that males

tend to have higher meristic values than females.

Springer (1967) reported differences between South

China Sea and non-South China Sea populations of

two species of Entomacrodus.

Nannosalarias, new genus

Undescribed genus (A) Springer, 1968b, p. 10.

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through
NUMBER 73 33

excavated area in jawbone. Anterior dentary canines absent; posterior dentary canines present.

Premaxillary and dentary teeth fairly rigid, 76 to

80 in upper jaw and 42 to 52 in lower jaw. Vomer

with small conical teeth. Dorsal rays XII, 15 or 16;

anal rays II, 16 to 18; segmented caudal rays 13,

middle 9 branched; pectoral rays 14 or 15 (usually

15); pelvic rays I, 3. Terminal anal ray bound by

membrane to caudal peduncle. Lateral line not

consisting of two overlapping disconnected portions; no scalelike flaps covering lateral-line pores.

Preoperculomandibular pores without cirri. Middorsal supratemporal pores typically 3. Upper lip

without free dorsal margin. No cup-shaped fleshy

disk or appendage behind lower lip. Gill membranes free. Occipital crest absent. Nuchal, supraorbital, and nasal cirri simple. Postcleithra consisting of two elongate bones, head of ventral element

overlapping ventral end of dorsal element. Lateral

extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 4. Type-species:

Blennius nativitatus Regan.

Relationships are discussed under section titled

"Recognition of Genera."

Etymology: A combination of the Greek nannos,

a dwarf (mature adults range from 22.5—34.8 mm

SL) , and Salarias, the type-genus of the Salariini.

Nannosalarias nativitatus (Regan)

FIGURES 14, 38

Blennius nativitatus Regan, 1909, pp. 404-405, pi. LXVI, fig.

2 [original description; type-locality: Christmas Island].

MATERIAL EXAMINED.-BNNH 1909.3.4.42 (adult

male, 30.1 mm SL), lectotype of Blennius nativitatus, Christmas Island, lat. 10°30'S, long. 105°40'E,

1909, C. W. Andrews; BMNH 1909.3.4.43-52 (5

of 15 paratopotypes, examined, 25.2-33.7); USNM

157372 (2 paratopotypes, 22.2-33.0); USNM 203098

(10, 24.9—33.0); Australia, Queensland, Capricorn

group, One Tree Island, south reef face about 1.6

km from island, lat. 23°30'S, long. 153°08'E, depth

0-4.6 meters, 22 November 1966, Victor G. Springer,

VGS 66-7; AMS IB 8207 (5, 26.6-30.4), same data

as USNM 203098; USNM 201369 (9, including 1

cleared and stained, 22.5-34.8), One Tree Island,

off reef crest on west side of island just above drop

off, depth 7.1-12.2 meters, 7 December 1966, Victor

G. Springer, VGS 66-16; AMS uncat. (1, 22.7), sta.

FT 96, Australia, One Tree Island, south face,

depth 10 meters, 26 November 1966, F. Talbot, et

al.; AMS uncat. (4, 26.3-34.3), sta. FT 204, Australia, One Tree Island, reef slope on south face,

depth 8 meters, 22 September 1967, F. Talbot and

B. Goldman; ANSP 109688 (2, 23.5-26.3), Australia, Queensland, Endeavour Reef, depth 1.3—2.4

meters, 13 January 1969, J. C. Tyler and C. L.

Smith, TS.A-13; ANSP 109687 (8, 15.7-27.7),

Australia, Queensland, Endeavour Reef, depth

1.5-4.6 meters, 16 January 1969, J. C. Tyler and

C. L. Smith, TS.A-17; USNM 195700 (1, 25.5),

presumably Dutch East Indies; CAS 24156 (5, 25.4-

30.1), same data as USNM 201369; USNM 203097

(2, 28.5-32.3), original no. GVF 1737, sta. HK 60,

South China Sea, Pratas Reef, about 0.6 km west

by north of Southwest Horn Island, lat. 20°36'40"N,

long. 116°43'50"E, 21 May 1958, Rolph L. Bolin;

USNM 203767 (2, 19.3-23.8), Te Vega Cruise 7,

sta. 301, Tonga Islands, Nivatobutabu Island, lat.

16°00'S, long. 175°53'W, depth 0-7 meters, 31 May

1965, Rolph L. Bolin.

DESCRIPTION.— (Characters for holotype in parentheses.) Dentary an open capsule with replacement

teeth entering functional series through excavated

area in jaw bone. Canine on each dentary posteriorly. Premaxillary and dentary teeth movable, but

fairly rigid; premaxillary teeth 76 to 80 (80); dentary teeth 42 to 52 (42). Vomer with 5 or 6 small,

conical teeth. Terminal vertebra with two epurals

and autogenous hypural 5 and ventral hypural

plate. Vertebrae 10 + 24 or 25 = 34 or 35; epipleural ribs 14 to 16; last pleural rib on vertebra

10. Circumorbital bones 4.

Dorsal spines 12 (rarely 13); last spine noticeably reduced; segmented rays 15 to 16 (16) [18 in

one specimen]; basal two-thirds of terminal dorsal

ray bound by membrane to caudal peduncle; dorsal

fin moderately incised between spinous and rayed

portions. Anal spines 2; anal fin rays 16 to 18(17);

basal two-thirds of terminal anal ray bound by

membrane to'caudal peduncle. Caudal fin with 13

segmented rays, middle 9 each branched once;

dorsal procurrent rays 6 to 8 (7), ventral rays 5 to

7 (6). Pelvic rays I, 3. Pectoral rays 15, rarely 14.

Pectoral radial formula 2—1-1 (based on 1 cleared

and stained specimen). Gill-rakers 12 to 14. Pseudobranchial filaments 5. Nuchal, supraorbital, and
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 14.—Nannosalarias nativitatus, from top to bottom: USNM 203097, male, 28.5 mm and

female, 32.3 mm, South China Sea; USNM 203098, male, 29.5 mm and USNM 201369, female,

26.7 mm, Queensland, Australia. Inserts are ventral side of head of males.
NUMBER 78 35

nasal cirri short and simple. Upper and lower lips

entire. Some pores of infraorbital series in pairs.

Lateral line of simple pores, incomplete, continuous

to point below 7th or 8th dorsal spine.

PIGMENTATION.—The following description is a

composite based on Australian specimens. Background color of head and body pale. Sides with

concentrations of melanophores tending to form

bands. Melanophores becoming more numerous on

dorsum and head, especially beneath eye and at

corners of mouth. Throat with two large, dark

gular spots extending posteriorly from lower lip.

(In the Tonga specimens the gular spots have become united as a single large spot.) Gular spots

well defined and intensified in some males, coalesced and diffuse in others, and even less well defined in females. Spinous dorsal fin of some specimens heavily pigmented on basal one-fifth, otherwise unmarked. Soft dorsal with a heavy concentration of pigment on basal one-fifth and on distal

margin of fin, rays otherwise largely unmarked.

Anal fin with interradial membranes heavily pigmented, especially distally. Caudal fin rays faintly

outlined with melanophores; interradial membranes

of unbranched rays heavily pigmented in some

males. Paired fins unmarked. Females are usually

paler than males, and in some specimens the fins

are completely devoid of pigment.

Regan (1909, p. 405) gives the life colors as

follows: "color reddish, with about 8 darker crossbars, sometimes appearing only as a series of oblong

blotches along the middle of the sides; fins pale."

GEOGRAPHIC VARIATION.—The two South China

Sea specimens of Nannosalarias differ in several

respects from the Australian and Tongan material,

especially in pigmentation. In the male the leading

edge of the first dorsal spine is intensely pigmented,

and the basal half of the spinous dorsal fin has a

heavy concentration of melanophores. The head

is uniformly dark and the two gular spots (not

apparent in the photograph) are smaller, located

farther posteriad, and more widely separated. The

female differs in having well-defined concentrations

of pigment in approximately the position of several

of the nreoperculomandibular pores. The gular

spots, although not as well defined, are in the same

position as those of the male.

The two South China Sea specimens also differ

from the other material in having longer supraorbital cirri. In the male (28.5) the ratio of the

cirri length into the orbital diameter is 1.0; in 8

Australian males (22.5—34.8) this ratio is 1.6 to

2.7, average 2.0. In the female (32.3) the cirri/

orbital ratio is 1.8; in 6 Australian females (25.5—

29.5) it is 2.0 to 5.3, average 3.0.

We do not believe that the differences exhibited

by the two South China Sea specimens warrant

description as a taxon separate from that of the

Australian and Tongan specimens. Springer (1967)

did recognize a separate subspecies for the South

China Sea populations of Entomacrodus thalassinus,

based mainly on relative length of the supraorbital

cirrus, and the South China Sea population of

E. stellifer, based primarily on color pattern. In

both instances, however, considerable material was

available.

Ophioblennius Gill

FIGURE 34

Blennophis Valenciennes in Webb and Berthelot, 1843, p. 60

[type-species: B. webbi Valenciennes in Webb and Berthelot, 1843, = Salarias atlanticus Valenciennes in Cuvicr and

Valenciennes, 1836, by original designation; a junior homonym of Blennophis Swainson, 1839, Clinidae].

Ophioblennius Gill, 1860, p. 103 [a substitute name for

Blennophis Valenciennes in Webb and Berthelot, 1843,

taking the same type-species: Blennophis webbi Valen.

ciennes in Webb and Berthelot, 1843].

Labroblennius Borodin, 1928, p. 31 [type-species: L. nicholsi

Borodin, 1928, = Ophioblennius steindachneri Jordan and

Evcrman, 1896, by monotypy].

Cynoscartes Norman, 1943, p. 31 [type-species: Salarias

atlanticus Valenciennes in Cuvier and Valenciennes, 1836,

by original designation].

Hepatoscartes Fowler, 1944, p. 230 [type-species: H. unbrifasciatus Fowler, 1944, = Ophioblennius steindachneri

Jordan and Evermann, 1896, by original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through

excavated area in jaw bone. Anterior dentary canines absent; posterior dentary canines present.

Premaxillary and dentary teeth freely movable,

numbering 184 to 224 in upper jaw and 102 to 108

in lower jaw. Vomer toothless. Dorsal rays XI to

XIII, 19 to 24; anal rays II, 20 to 25; segmented

caudal rays 13, middle 9 branched; pectoral rays 14

or 15 (usually 15); pelvic rays I, 4. Terminal anal

ray not bound by membrane to caudal peduncle.

Lateral line consisting of two overlapping dis-
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

connected portions; no scalelike flaps covering

lateral-line pores. Preoperculomandibular pores

without cirri. Mid-dorsal supratemporal pores 5 to

more than 50. Upper lip without free dorsal margin. No cup-shaped fleshy disk or appendage behind lower lip. Gill membranes free. Occipital

crest absent. Nuchal cirri consisting of a pair of

cirri or a patch of 3 to 15 or more cirri on each

side of nape; supraorbital cirri simple; nasal cirri

multifid. Postcleithra consisting of two elongate

bones, head of ventral element overlapping ventral

end of dorsal element. Lateral extrascapular not

fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

The genus Ophioblennius was revised by

Springer (1962) . Ophioblennius appears to be most

closely related to Scartichthys. In addition to the

characters given in the key to distinguish these two

genera, Ophioblennius differs in having dentary

canines that are equal to or greater than the diameter of the pupil, instead of having canines less

than one-half the pupil diameter (exclusive of

larvae), and the lateral line consisting of two

overlapping disconnected portions instead of being

continuous.

DISTRIBUTION.—Restricted to the Atlantic Ocean

and eastern Pacific Ocean.

Pereulixia Smith

FIGURE 29

Pereulixia Smith, 1959, p. 238 [type-species: Salarias kosiensis Regan, 1908, by original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through

excavated area in jaw bone. Anterior dentary

canines absent; posterior dentary canines present.

Premaxillary teeth freely movable, numbering 155

to 157; dentary teeth fairly rigid, numbering 43 to

53. Vomer with small conical teeth. Dorsal rays

XI or XII, 11 or 12; anal rays II, 12 to 14; segmented caudal rays 13, middle 9 branched; pectoral

rays 15 or 16 (usually 15); pelvic rays I, 4. Terminal

anal ray not bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; imbricate scalelike

flaps covering at least anterior lateral-line pores.

Preoperculomandibular pores without cirri. Middorsal supratemporal pores numerous, relatively

small and difficult to count. Upper lip without free

dorsal margin. No cup-shaped fleshy disk or appendage behind lower lip. Gill membranes free.

Occipital crest absent. Nuchal cirri consisting of a

series of 4 elongate, transverse, contiguous or

slightly overlapping patches of cirri interrupted at

midline by a narrow hiatus; supraorbital cirri simple, long, and tapering; nasal cirri multifid. Postcleithra consisting of two elongate bones, head of

ventral element overlapping ventral end of dorsal

element. Lateral extrascapular not fused with

pterotic. Median ethmoid ossified. Circumorbital

bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

Pereulixia is apparently monotypic and rare in

museum collections; we know of fewer than 10

specimens.

DISTRIBUTION.—Known only from the western

Indian Ocean.

Praealticus Schultz and Chapman

FIGURES 15a, b; 21

Praealticus Schultz and Chapman, 1960, p. 368 [type-species:

Salarias natalis Regan, 1909, by original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through

excavated area in jaw bone. Anterior dentary

canines absent; posterior dentary canines present

or absent. Premaxillary and dentary teeth freely

movable, numbering 190 to 280 in upper jaw and

128 to 214 in lower jaw. Vomer with or without

teeth. Dorsal rays XII to XIV, 18 to 20; anal rays

II, 18 to 23; segmented caudal rays 12 or 13,

branched rays 0 to 9 (usually 4 to 8); pectoral rays

14 or 15; pelvic rays I, 3 or I, 4. Terminal anal ray

not bound by membrane to caudal peduncle.

Lateral line not consisting of two overlapping

disconnected portions; no scalelike flaps covering

lateral-line pores. Preoperculomandibular pores

without cirri. A single mid-dorsal supratemporal

pore. Upper lip without free dorsal margin. No

cup-shaped fleshy disk or appendage behind lower

lip. Gill membranes free. Occipital crest present.
NUMBER 73 37

Nuchal cirri absent; supraorbital cirri pinnately

branched; nasal cirri simple, occasionally with

weak points. Postcleithra consisting of 1—3 small,

fragile, well-separated bones on each side (except

P. schmidti in which postcleithra consist of two

elongate bones, head of ventral element overlapping ventral end of dorsal element). Lateral extrascapular not fused with pterotic. Median ethmoid

ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

The fact that this genus, which is widely distribFICURE 15.—Postcleithra of selected species of Salariini showing relative development (gray bar to left of postcleithra

represents pectoral fin base): a, Praealticus tanegasimae,

USNM 123398, male, 61.0 mm. Okinawa, b, Praealticus margaritatus, USNM 115481, female, 49.7 mm, Samoa Islands;

c, Alticus saliens, USNM 115446, male, 61.9 mm, Samoa Island; d, Istiblennius coronatus, USNM 200611, female, 56.6

mm, Marshall Islands.

uted throughout much of the Indo-Pacific, was not

recognized until 1960 attests to its remarkable resemblance to two other (though not necessarily

closely related) genera. Praealticus is well named,

for it has frequently been confused with species of

Alticus. The close superficial similarity of the two

genera is well illustrated by the drawing of the

holotype of Alticus semicrenatus Chapman, which

was published in the same work as the original

description of Praealticus and was not recognized

by Schultz and Chapman as being a species of their

new genus.

Although we consider Praealticus more closely

related to Alticus than to Istiblennius, species of

Istiblennius are most often confused with Praealticus and are the most difficult to distinguish from

it. A comparison of the characters most useful in

distinguishing these three genera is given in Table 1.

DISTRIBUTION.—Eastern Indian Ocean; central

and western Pacific Ocean.

Rhabdoblennius Whitley

FIGURES 43, 45

Rhabdoblennius Whitley, 1930, p. 20 [type-species: Blennius

rhabdotrachelus Fowler and Ball, 1924, by original designation].

Nixiblennius Whitley, 1930, p. 20 [type-species: Blennius

snowi Fowler, 1928, by original designation].

Scartoblennius Fowler, 1946, p. 174 [type-species: Blennius

ellipes Jordan and Starks, 1906, by original designation].

DIAGNOSIS.—Dentary a closed capsule (open mesially in R. snowi, but with large gaps and foramina

in the wall laterally) with replacement teeth entering functional series through foramina in jaw bone.

Anterior dentary canines absent; posterior dentary

canines present. Premaxillary and dentary teeth

immovable or nearly so, numbering 44 to 50 in

upper jaw and 28 to 38 in lower jaw. Vomer with

small conical teeth. Dorsal rays XII, 18 to 21; anal

rays II, 20 to 22; segmented caudal rays 13, middle

9 branched; pectoral rays 13 or 14 (usually 14) ;

pelvic rays 1,3. Terminal anal ray bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; no

scalelike flaps covering lateral-line pores. Preoperculomandibular pores without cirri. Mid-dorsal

supra temporal pores 3—10. Upper lip without free

dorsal margin. No cup-shaped fleshy disk or ap-
38

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Character

Pores in middorsal area of

supratemporal canal

Nature of postcleithra (figure

15)

Branched caudal rays1

Fleshy occipital crest in adult

males2 ...

Nasal cirri

Nuchal cirri

Total dorsal rays

Segmented anal rays

TABLE 1.—Comparison of

Praealticus

1 (Figure 21)

1-3 small, fragile, wellseparated bones on each

side (figures I5a-b)

[except schmidti Fowler

($) =biorni Fowler ($ ) ]

0-9 (usually 4-8)

always present

simple or weakly, pinnately

fringed

absent

31-33

18-23

three genera of Salariini

Alticus

1 (Figure 19)

1-3 small, fragile, wellseparated bones on each

side (Figure 15c)

0

always present

simple to multifid

absent

35-38

24-28

Istiblennius

0-12 (3 in most species;

0-2 in only one species;

Figures 22-25).

Two elongate bones, head

of ventral element

overlapping ventral end

of dorsal element

(Figure 15d).

8-9 (usually 9)

present or absent

simple to multifid, deeply

split except in very

young (simple in only

two species) .

present or absent

31-37

17-25

1

 The caudal rays are frequently unbranched in very young individuals, even in genera that typically have branched caudal

rays. In Praealticus the caudal rays are frequently branched only at their tips and the branching may be missed easily.

2

 Even in species that typically have a fleshy occipital crest, it is always much better developed in males than in females and

is frequently absent in young of both sexes.

pendage behind lower lip. Gill membranes free.

Occipital crest absent. Nuchal cirri absent; supraorbital and nasal cirri simple. Postcleithra consisting of two elongate bones, head of ventral element

overlapping ventral end of dorsal element. Lateral

extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

Norman (1943) first placed Nixiblennius in

synonymy with Rhabdoblennius. Schultz and Chap

man (1960) first synonymized Scartoblennius.

Whitley (1930) proposed the genus Nixiblennius

for Blennius snozvi on the same page that he erected

the genus Rhabdoblennius for B. rhabdotrachelus,

but gave no reasons for erecting the genera and no

characters to separate them. Later, Fowler (1946)

proposed the genus Scartoblennius for Blennius

ellipes, but did not compare it with either Rhabdoblennius or Nixiblennius. Rhabdoblennius rhabdotrachelus does differ from the previously mentioned

species in possessing a "pseudobasisphenoid"

(Springer, 1968, p. 44), but we do not believe that

this difference warrants rhabdotrachelus being

placed in its own monotypic genus. The species

comprising Rhabdoblennius are few in number

(probably only 6 or 7) and, except for the presence

of a pseudobasisphenoid in rhabdotrachelus, are

quite close osteologically.

DISTRIBUTION.—Central and western Pacific Ocean.

Salarias Cuvier

FIGURES 16, 28

Salarias Cuvier, 1817, p. 251 [type-species: 5. quadripennis

Cuvier, 1817, = Blennius fasciatus Bloch, 1786, by monoErpichthys Swainson, 1839, p. 275 [type-species: Salarias

quadripennis Cuvier, 1817, = Blennius fasciatus Bloch, 1786,

by subsequent designation].

Negoscartes Whitley, 1930, p. 20 [type-species: Salarias irroratus Alleyne and Macleay, 1877, by original designation].

Crenalticus Whitley, 1930, p. 21 [type-species: Salarias crenulatus pallidus Whitley, 1926, = Salarias sinuosus Snyder,

1908, by original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jawbone. Anterior dentary canines

absent; posterior dentary canines present. Premaxillary and dentary teeth freely movable, numbering

110 to 168 in upper jaw and 72 to 134 in lower
NUMBER 73 39

jaw. Vomer toothless. Dorsal rays XII or XIII, 16 to

20; anal rays II, 18 to 21; segmented caudal rays 13,

middle 9 branched; pectoral rays 13 to 15 (rarely 15);

pelvic rays I, 3 (innermost pelvic ray minute, visible

only in cleared and stained material). Terminal anal

ray bound by membrane to caudal peduncle. Lateral

line not consisting of two overlapping disconnected

portions; no scalelike flaps covering lateral-line

pores. Preoperculomandibular pores without cirri.

Mid-dorsal supra temporal pores 1—3. Upper lip

without free dorsal margin. No cup-shaped fleshy

disk or appendage behind lower lip. Gill membranes free. Occipital crest present or absent.

Nuchal cirri simple or palmate; supraorbital and

nasal cirri simple to multifid. Postcleithra consisting of two elongate bones, head of ventral element

overlapping ventral end of dorsal element. Lateral

extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

Whitley (1930, p. 20) erected the genus Negoscartes to include those species of Salarias that have

a well-developed notch in the dorsal fin and large

mandibular canines. Salarias fasciatus was erroneously thought to lack canines. The degree of notching in the dorsal fin varies from well developed to

absent in the species of Salarias and in other genera

of Salariini (for example Ecsenius). Norman (1943)

also recognized Negoscartes, but the defining characters he used are without substance. Salarias fasciatus and S. ceramensis do differ from the other

species of Salarias in attaining a greater size as

adults, having more teeth in the upper jaw, and

nuchal and supraorbital cirri that are broad and

multifid. Several of these characters, however, appear to show a trend, with fasciatus and ceramensis

merely the extremes. We believe that the characters

used in the key and diagnosis to define the genus

are of much greater importance than the differences

noted above.

The genus Crenalticus (Whitley, 1930, p. 21)

was proposed for Salarias crenulatus (Weber) [=

S. sinuosus Snyder], which differs from the other

species of Salarias in possessing a crenulate upper

lip. We consider this character to be of only specific

significance. Springer (1967) found that the degree

or presence of crenulation of the upper lip was

only of specific importance in the salariinine genus

Entomacrodus.

DISTRIBUTION.—Red Sea; Indian Ocean; central

and western Pacific Ocean.

Scartichthys Jordan and Evermann

FlCURE 3

Scartes Jordan and Evermann, 1896, p. 471 [type-species:

Salarias rubropunctatus Valenciennes in Cuvier and Valencinnes, 1836, by original designation; a junior homonym of

Scartes Swainson, 1835, Mammalia].

Scartichthys Jordan and Evermann, 1898, p. 2395 [type-species:

Salarias rubropunctatus Valenciennes in Cuvier and Valenciennes, 1836, by original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jawbone. Anterior dentary canines

absent; posterior dentary canines present. Premaxillary and dentary teeth freely movable, numbering

144 to 256 in upper jaw and 67 to 123 in lower

jaw. Vomer toothless. Dorsal rays XII, 17 or 18;

anal rays II, 18 to 20; segmented caudal rays 13,

middle 9 branched; pectoral rays 14; pelvic rays

1,4. Terminal anal ray not bound by membrane

to caudal peduncle. Lateral line not consisting of

two overlapping disconnected portions; no scalelike

flaps covering lateral-line pores. Preoperculamandibular pores without cirri. Mid-dorsal supratemporal pores 5—12. Upper lip without free dorsal

margin. No cup-shaped fleshy disk or appendage

behind lower lip. Gill membranes free. Occipital

crest absent. Nuchal cirri consisting of a patch of

cirri on each side of nape separated by a wide

hiatus equal to or greater than length of base of

either patch of cirri; supraorbital and nasal cirri

multifid. Postcleithra consisting of two elongate

bones, head of ventral element overlapping ventral

end of dorsal element. Lateral extrascapular not

fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

Specimens housed in the National Museum of

Natural History that we have examined indicate

that the genus consists of at least two well-defined

species (also see discussion under Cirripectes).

The species of Scartichthys constitute the largest

members of the family Blenniidae, although not the
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

longest. (Xiphasia Swainson, of the Nemophidinae,

is eel-like and attains a length over 500 mm SL.)

Adults frequently exceed 175 mm in standard

length.

DISTRIBUTION.—Restricted to the eastern Pacific

Ocean.

Stanulns Smith

FIGURE 7

Stanulus Smith, 1959, p. 246 [type-species: S. seychellensis

Smith, 1959, by original designation].

Fallacirripectes Schultz and Chapman, 1960, p. 362 [typespecies: F. minutus Schultz and Chapman, 1960, = Stanulus

seychellensis Smith, 1959, by original designation].

DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jawbone. Anterior dentary canines

absent; posterior dentary canines present. Premaxillary and dentary canines freely movable, numbering 118 to 138 in upper jaw and 62 to 82 in lower

jaw. Vomer with or without teeth. Dorsal rays XII,

9 to 12; anal rays II, 10 to 13; segmented caudal

rays 13, middle 9 branched; pectoral rays 14 to 16

(usually 15); pelvic rays I, 4. Terminal anal ray

not bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; minute, imbricate, scalelike flaps

covering anterior lateral-line pores. Preoperculomandibular pores without cirri. Mid-dorsal supratemporal pores 3—22. Upper lip without free dorsal

margin. No cupshaped fleshy disk or appendage

behind lower lip. Gill membranes free. Occipital

crest absent. Nuchal, supraorbital, and nasal cirri

simple. Postcleithra consisting of two elongate

bones, head of ventral element overlapping ventral

end of dorsal element. Lateral extrascapular not

fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.

Relationships are discussed under section titled

"Recognition of Genera."

This genus, consisting of two species, has been

revised recently by Springer (1968).

DISTRIBUTION.—Indian Ocean; central and western

Pacific Ocean.

Comparison of Cephalic Sensory Pores

Although the sensory canal system has generally

been neglected in the classification of blenniid

fishes, Springer (1967) used the nature of the sensory pores as a specific character to distinguish

certain species of the salariinine genus Entomacrodus. In our study we find that the cephalic sensory

pores have utility as generic characters and may

indicate phyletic relationships. We have illustrated

the pore patterns of representative species of each

genus and subgenus of Salariini (Figures 17—50)

as an adjunct to our generic discussions and to encourage future students of the Blenniidae to survey

the character more fully.

Correlation between development of cephalic

pores and the ecology and/or behavior of blenniid

fishes is not readily apparent. Blennius ocellaris

(subfamily Blenniinae), the deepest dwelling (to

at least 400 meters) member of the family, has

simple pores, as do species of the salariinine genera

Alticus and Andamia, which spend much of their

existence out of water. Unlike the previously mentioned genera, all species of the subfamily Nemophidinae spend most of their time actively swimming, yet they share in common a reduced sensory

pore system. The only general observation that we

can make regarding sensory pores is that there

seems to be a strong trend for the pores to be reduced in number in the more specialized forms.

The cephalic sensory canals in the tribe Salariini

agree with the typical perciform pattern described

by Branson and Moore (1962). Some of the major

sensory canals and associated pores of a representative salariinine genus are shown in Figure 16. The

sensory pore series of the head that we found of

importance are defined below.

Infraorbital (IFO) and supraorbital (SO) series:

Pores in the positions occupied by these canals.

Supratemporal series (ST) : Only pores in the

region anterior to the first dorsal spine and between

the bases of the nuchal cirri (if present) , extending

anteriorly across the top of the head, are discussed

below. In some genera, for instance Entomacrodus,

pores anterior to the level of the nuchal cirri frequently spread laterally and cover an expanse

broader than the distance between the nuchal cirri;

all pores anterior to the nuchal cirri in this midregion are included in our counts. In some specimens this count may be difficult to make. Springer

(1967, p. 6) termed pores in this series "predorsal

commissural pores."
NUMBER 73

41

IFO

ST

FIGURE 16.—Semidiagramatic illustration of cephalic sensory canals and pores in Salarias sinuosut

based on several specimens from Queensland, Australia, with abbreviations as follows: MDB,

mandibular series; POP, preopercular series; POM, preoperculomandibular series; IFO, infraorbital canal; SO, supraorbital canal; SOC, supraorbital commissural canal; INO, interorbital

canal; PT, pterotic canal; ST, supratemporal canal; LI, lateralis. Positions of preopercle,

operde, and subopercle are shown in dark gray.

Mandibular series (MDB): The number of mandibular pores on each side of the head beginning

with the pore nearest the symphysis of the lower

jaw and terminating posteriorly with the pore immediately behind or slightly dorsal to the membranous corner of the mouth. In some genera it is

difficult to determine by external examination

alone where the break occurs between the MDB

series and the POP series. In the following discussion of pore patterns the reader should experience

little difficulty determining which pores we include

in either series if reference is made to the illustrations.

Preopercular series (POP): In this series we

consider the first pore position (depending on the

genus or species, a position may be occupied by

one or more pores, all connected by a common

canal with a single opening in the preopercular

bone) as the dorsalmost, which occurs slightly

above the level of the greatest distance between the

posterior margins of the preopercle and operde,

excluding the branchiostegal membrane.

In the Salariini there may be either five or six

positions in the POP series, depending on the

genus, describing an arc between the first and last

positions. The last pore position is the ventralmost
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of the series and is posterior to the last MDB pore.

In some genera the last POP position is always

occupied by a vertical pair of pores, in other genera

by a single pore. On the figures we have placed

an "x" below the last POP position.

In the following account of pore patterns the

reader should refer to Figures 17-50. Unless stated

otherwise, the species illustrated is considered typical for all species of the genus. To facilitate comparison of genera the sequence of presentation

follows that of Figure 51 (from top to bottom) . In

many species the pores are difficult to see and are

easily overlooked. Overlying mucus must be carefully removed and drying of the area in which pores

occur greatly facilitates examination.

Ecsenius (Figures 17-18). All pore positions

occupied by simple pores. MDB pores 4. While the

basic pore pattern is the same for both subgenera,

there has been a posterior displacement of the mandibular pores in Anthiiblennius, perhaps correlated

with changes in the structure of the lower jaw.

Anthiiblennius also differs in having a more anterior

placement of the mid-dorsal ST pore. This shift in

position is apparently the result of the more anterior placement of the first dorsal spine and its

pterygiophore.

Andamia (Figure 20). All pore positions occupied

by simple pores. A single MBD pore. Reduction of

MDB pores is probably correlated with the presence

of a prominent cup-shaped fleshy disk behind the

lower lip (Figure 10). A single mid-dorsal ST pore

present.

Alticus (Figure 19). All pore positions occupied

by simple pores. MDB pores 2 or 3. Reduction of

MDB pores in Alticus and Andamia (4 or more in

other salariinine genera except group 21 of Figure

51) is another reason for placing Alticus as we do

in Figure 51. A single mid-dorsal ST pore present.

Praealticus (Figure 21). All pore position occupied by simple pores. MDB pores 4. Distribution of

ST pores is identical to that found in the related

Alticus and Andamia, and is characterized primarily

by the presence of a single mid-dorsal ST pore.

Istiblennius (Figures 22-25). This large assemblage of species, which shows the greatest range of

variation in its other characters, likewise exhibits

much variation in pore development. IFO positions

occupied by multiple or simple pores, simple in

most species. MDB pores 5 or 6, 5 in most species.

POP positions consisting of simple pores in all

species except /. gibbifrons in which the pores at

each position are double. Also, since an ophioblennius larval stage is known only for gibbifrons

among the Istiblennius species, placement of gibbifrons in a separate genus may be indicated (see

discussion under general synoptic account). Middorsal ST pores 0-12 or more, 3 or more in all

except I. steinitzi.

The genera illustrated in Figures 29-34 (group

7 of Figure 51) are very difficult to characterize

individually because of the complex nature of their

sensory pores. In Pereulixia (Figure 29), Exalhas

(Figure 30), and Cirripectes (Figures 31—32) most

or all of the pore positions are occupied by relatively small and numerous pores arranged in clusters. Each pore in a cluster at a given position is

united by a minute branch to a common canal. In

most species of Cirripectes these pores are very

minute and easily overlooked. In Scartichthys (Figure 33) and Ophioblennius (Figure 34) the pores

are larger and less numerous. In all of these genera

there tends to be an increase in the number of

pores with increase in standard length.

Stanulus (Figure 37). IFO typically containing

positions with some multiple pores. POP positions

typically with simple pores. MDB pores 4 or 5. Middorsal ST pores 3-22, the number increases with

increase in standard length (Springer, 1968a).

Entomacrodus (Figures 35-36). IFO typically

containing several positions with multiple pores.

POP positions variable, consisting of simple, paired

or mulitple pores. MDB pore positions (some positions may have multiple pores) 4-6. ST pores in

mid-dorsal region 3 to more than 50. In those species that have a relatively high number of ST pores,

the number increases with increase in standard

length.

Nannosalarias (Figure 38). IFO positions occupied by multiple pores. POP positions with simple

pores. MDB pores usually 5, occasionally 4. Middorsal ST pores typically 3.

Crossosalarias (Figure 26). Some IFO position

typically with multiple pores. POP positions with

simple pores. MDB pores 5. Mid-dorsal ST pores 3.

Atrosalarias (Figure 27). All pore positions occupied by simple pores. MDB pores 4. A single middorsal ST pore present.
NUMBER 73 43

E. (Ecsenius) E. (Anthiblennius)

Alticus

Andamia

Praealticus Istiblennius

FIGURES 17-22.—Comparison of cephalic sensory pores in selected species of Salariini: 17, Ecsenius (E.) bicolor, USNM 201823, male, 61.5 mm, Queensland, Australia; 18, Ecsenius (Anthiiblennius) midas, paratype, USNM 202422, female, 56.0 mm, Amirante Islands; 19, Alticus saliens,

USNM 115447, male, 69.5 mm, Tutuila Island, Samoa; 20, Andamia heteroptera, USNM

137867, male, 53.5 mm, Porongpong Island, Philippines; 21, Praealticus tanegasimae, USNM

123398, male, 58.5 mm, Okinawa; 22, Istiblennius edentulus, USNM 142201, male, 59.2 mm.

Marshall Islands.
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

23 Wiblennius 24 Istiblennius

Istib'tfnnius Crossosalarias

27 Airosalarias 28 Salorias

FIGURES 23-28.—Comparison of cephalic sensory pores in selected species of Salariini: 23, Istiblennius gibbifrons, USNM 200545, male, 65.0 mm, Marshall Islands; 24, Istiblennius coronatus,

USNM 142068, female, 75.5 mm, Marshall Islands; 25, Istiblennius paulm, USNM 142202, male,

66.0 mm, Marshall Islands; 26, Crossosalarias macrospilus, holotype, USNM 203096, male, 51.0

mm, Solomon Islands; 27, Atrosalarias fuscus holomelas, USNM 201255, male, 62.1 mm, Queensland, Australia; 28, Salaries fasciatus, USNM 203105, female, 52.0 mm, Queensland, Australia.
NUMBER 73 45

Pereulixia

Exallias

Cirripectes

Cirripactes

33 Scortichthys Ophioblennius

FIGURES 29-34.—Comparison of cephalic sensory pores in selected spedes of Salariini. In the following genera some or all of the MDB and ST pores are not shown: 29, Pereulixia kosiensis,

USNM 204035, male, 64.9 mm, Inhaca, Mozambique; 30, Exallias brevis, USNM 1999420, male,

85.5 mm, Malacca Straits, Indonesia; 31, Cirripectes obscurus, BPBM (uncat.), male, 106 mm,

Oahu, Hawaii; 32. Cirripectes jenningsi, USNM 200615, female, 69.6 mm, Vostok Island; 33,

Scarttchthys variolatus, USNM 176546, female, 90.3, Juan Fernandez Island, Chile (right supraorbital cirrus not illustrated); 34, Ophioblennius atlanticus macclurei, UMML 6523, female,

75.5 mm, Bahama Islands.
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Entomacrodus

Entomacrodus

Stanulus

- \

Nannosalarias

Mimoblennius 40 Mimoblennius

FIGURES 35-40.—Comparison of cephalic sensory pores in selected species of Salariini: 35, Entomacrodus chiostictus, USNM 256392-F3, male, 62.5 mm, Bahia Pinas, Panama; 36, Entomacrodus

striatus, USNM 198141, male, 68.9 mm, Phoenix Islands; 37, Stanulus seychellensis, USNM

201580, male, 24.8 mm, Queensland, Australia; 38, Nannosalarias nativitatus, USNM 201369,

male, 32.8 mm, Queensland, Australia; 39, Mimoblennius cirrosus, paratype, USNM 204645,

female, 28.2 mm, Sheikh el Abu, Harat Island, Ethiopia; 40, Mimoblennius atrocinctus, USNM

201395, male, 442 mm, Hong Kong.
NUMBER 73 47

Glyptoparus 42 Litobranchus

Rhabdoblennius Medusablennius

Hirculops

FIGURES 41-46.—Comparison of cephalic sensory pores in selected species of Salariini: 41, Glyptoparus delicatulus, USNM 203106, male, 30.4 mm, Aldabra Island; 42, Litobranchus fowleri,

USNM 203769, male, 31.0 mm, Oriental Negros, Philippines; 43, Rhabdoblennius snowi, USNM

204634, male, 40.7 mm, Gilbert Islands; 44, Medusablennius chani, CAS 24741, male, 12.0 mm,

Tuamotu Archipelago; 45, Rhabdoblennius rhabdotrachelus, USNM 200608, male, 34.0 mm,

Solomon Islands; 46, Hirculops cornifer, USNM 200030, female, 46.5 mm, Gulf of Suez, Red

Sea.
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

A. (Antennoblennius)

A. (Croaltus)

Alloblenniiis Alloblennius

FIGURES 47-50.—Comparison of cephalic sensory pores in selected species of Salariini: 47, Antennablennius (A.) velifer, USNM 200205, male, 44.7 mm, Astola Island, West Pakistan; 48, Antennablennius (Croaltus) bifilum, USNM 200207, male, 495 mm, Astola Island, West Pakistan; 49,

Alloblennius pictus, USNM 203766, male, 30.1 mm, Strait of Jubal, Red Sea; 50, Alloblennius

jugularis, HUI £62/417g., male, 30.9 mm, Cundabilu, Dahlak Archipelago.

Salaries (Figures 16 and 28). All pore positions

occupied by simple pores. MDB pores 4. Mid-dorsal

ST pores 1-3.

Glyptoparus (Figure 41). All pore positions

occupied by simple pores. MDB pores 4. Mid-dorsal

ST pores 1 or 2, usually 2.

Medusablennius (Figure 44). All pore positions

occupied by simple pores. MDB pores 3. Mid-dorsal

ST pores 2.

Mimoblennius (Figures 39-40). IFO with some

positions occupied by multiple pores. POP positions with simple pores except for the last position,

which consists of a vertical pair of pores. MDB

pores 3. Mid-dorsal ST pores 3.

Litobranchus (Figure 42). All pore positions

occupied by simple pores. MDB pores 3. Mid-dorsal

ST pores 2-4, usually 2.

Rhabdoblennins (Figures 43 and 45). IFO positions occupied by simple or multiple pores. POP

positions with simple pores except for the last position, which consists of a vertical pair of pores.

MDB pores 3 or 4. Mid-dorsal ST pores 3-10.

Hirculops (Figure 46). IFO with some positions

occupied by multiple pores. POP pores simple except for last position, which consists of a vertical

pair of pores. MDB pores 3. Mid-dorsal ST pores

8-15.

Antennablennius (Figures 47-48). IFO with some

positions occupied by multiple pores. All POP

positions occupied by simple pores. MDB pores 3.
NUMBER 73

49

Mid-dorsal ST pores 1-4, typically 3 in subgenus

Antennablennius and usually 2 in subgenus Croaltus (see general comments under Antennablennius).

Alloblennius (Figures 49-50). IFO positions usually occupied by simple pores or with relatively

few paired pores. All POP positions occupied by

simple pores. MDB pores 3. Mid-dorsal ST pores

3-8.

Recognition of Genera

In any study embracing as many genera and

species as in our study, it behooves the authors to

discuss their generic concept. The only rule that

can be consistently applied to the genus-group

category is that the species of a particular taxon be

more closely related to each other than to species

in another taxon of equal rank. Recognition of

monotypic taxa, above the species level, simply infers a high degree of differentiation in a single

species. Obviously, the level of differentiation that

one chooses to regard as meriting generic (as opposed to subgeneric) recognition will depend upon

the number and stability of characters within each

family of animals considered and the subjective

bias of the taxonomist involved. In the final analysis the genus-subgenus system of nomenclature is

subjective.

For every genus that we recognize we have spent

much time in discussion and debate challenging each

other to defend an action, whether it be to synonymize a genus, recognize one in synonymy, erect a

new genus, or not erect a genus where one might

be recognizable. We have taken special note of the

TRIBE

SALARIINI

1

6

15

16

20

21

2

5

7

13

8

4

•* 11 -

3

9

8

9

17

10

12

1

 1 12

14

r ••

I,,

 |4

18

19

22

23

24

25

27

1 11

ix

 2 8

1 2*

ESTIMATED NUMBER OF

SPECIES IN GENUS

ECSENIUS

ANDAMIA

ALTICUS

PRAEALTICUS

ISTIBLENNIUS

OPHIOBLENNIUS

PEREULIXIA

EXALLIAS

CIRRIPECTES

SCARTICHTHYS

STANULUS

ENTOMACRODUS

NANNOSALARIAS

CROSSOSALARIAS

ATROSALARIAS

SALARIAS

GLYPTOPARUS

MEDUSABLENNIUS

MIMOBLENNIUS

LITOBRANCHUS

RHABDOBLENNIUS

HIRCULOPS

ANTENNABLENNIUS

ALLOBLENNIUS

18-24

4 - 5

8-10

10-12

25-30

2

1

1

10-15

2

2

22-24

1

1

1

8-12

1

1

2

1 - 2

6 - 8

1

8-10

2

138-170

FIGURE 51.—Schematic representation of genera in tribe Salariini showing relationships. Numbered branches of dendrogram are discussed under section titled "Recognition of Genera."
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fact that some of these genera are as yet poorly

known. From these discussions has evolved the

dendrogram shown in Figure 51, which is a schematic representation of our concept of the relationships of the salariinine genera that we recognize.

The dendrogram does not imply that one genus is

more primitive than, or ancestral to, another. The

arrangement simply places those genera that we

consider to be most closely related together in

brackets. In the following discussion we attempt

to explain the reasons for our arrangement of the

genera of Salariini in Figure 51 and comment on

why we recognize some of them. Numbers given

in the following discussion correspond to those

shown on the figure.

1. Ecsenius is the most distinct genus in the

Salariini. Indeed, it occupies a unique position in

the Blenniidae and perhaps merits a separate tribe.

The general nature of its dentition, however, at

least places Ecsenius as an offshoot of the Salariini.

The following information is the result of a revision of the genus by Springer. Ecsenius differs from

all other blenniids in having dorsal processes extending from the proximal portions of the epipleural ribs on at least the third through fifth

vertebrae; in having both anterior and posterior

canines on the dentary (the anterior canines are

difficult to distinguish from the incisoriform teeth

in most species, but their nature is apparent in

osteological preparations); and in lacking an ossified median ethmoid. In addition to the above

characters, Ecsenius also differs from the other

salariinine genera in having the lateral extrascapular fused with the pterotic.

2. Excluding Ecsenius, there appear to be at least

four major phyletic lines within the Salariini. Their

relationships to each other, however, are not clear.

Alticus and its allies comprise a group of slender,

elongate species that closely approach each other

in general physiognomy. They share in common

high numbers of dentary and premaxillary teeth,

a reduced pore system, a fleshy occipital crest (secondarily lost in all except one species of Andamia),

and loss of nuchal cirri. There has been a tendency

for the ventralmost rays of the caudal fin to become

thickened in most species, probably in response to

their habit of frequently leaving the water and

using their tails to hop from one pool to another.

They have either unbranched caudal rays or the

number of branched rays is reduced, usually 4 to 8

(typically 9 in those Salariini with branched rays).

With the exception of one species of the unrelated

genus Ecsenius (E. midas) and one species of Praealticus, this group of genera is unique among the

Salariini in having the postcleithra consisting of

one to three small, fragile, well-separated bones on

each side (Figures 15a, b, c).

3. Alticus and Andamia are clearly more closely

related to each other than to Praealticus. In adults

of Praealticus, unlike Alticus and Andamia, the

tips of some of the caudal rays are always branched

and the terminal anal pterygiophore frequently

bears two rays (one in Alticus and Andamia) . With

the exception of Alticus simplicirrus (see text discussion under Alticus), the general nature of the

supraorbital cirri is different in the two groups.

4. Andamia is unique among the genera of Salariini in having a cup-shaped fleshy disk or appendage behind the lower lip (Figures 10 and 20).

5. The precise placement of Istiblennius is uncertain, but if difficulty in distinguishing the genus

is an indication, then it is closely related to Praealticus (see text discussion under Praealticus). Since

an ophioblennius larval stage is known for only one

species of Istiblennius (gibbifrons), for none of

the presumed closest relatives of Istiblennius (group

2), and for most (if not all) of the unrelated genera

of group 6, Istiblennius as here defined may be

polyphyletic.

6. Included are those genera that share the following combination of characters: ophioblennius

larvae, the anal spines of mature males enveloped

in conspicuous fleshy rugosities, multiple pores on

the head tending to increase in number with increase in standard length, I, 4 pelvic rays (except

for one species of Cirripectes, which has I, 3), and

the terminal anal ray not bound by a membrane

to the caudal peduncle. Most of the species typically

have the last anal pterygiophore bearing two rays.

7. Included are those genera with complete lateral lines (extending almost to the caudal base)

and nuchal cirri consisting of a transverse series

across the nape or a patch of cirri on each side of

the nape. One species of Entomacrodus (E. marmoratus) overlaps this group.

8. Ophioblennius differs from its nearest relatives

in having the lateral line consisting of two overlapping, disconnected portions (instead of being
NUMBER 73 51

continuous to the caudal base), the dorsal fin without a notch between spinous and segmented portions (but notched somewhat in larvae), and in

having more segmented dorsal and anal rays. It is

the only genus of the group that has become established in the Atlantic Ocean.

9. Pereulixia differs from its nearest allies in having small, imbricate, scalelike flaps covering at

least the anterior lateral-line pores, vomerine teeth,

simple supraorbital cirri that are long and tapering

(simple but short in Cirripectes jenningsi), and

the upper lip entire.

10. Exallias differs from both Cirripectes and

Scartichthys in having about one-third as many

teeth in the lower jaw as in the upper (instead of

about one-half as many teeth in the lower jaw as

in the upper), no dentary canines, a pair of barbels

on each side of the chin just behind the lower lip,

and fewer segmented dorsal and anal rays.

11. We know of no single character that alone

will distinguish Cirripectes and Scartichthys from

their respective relatives. This, it seems to us, is

not too surprising when dealing with a relatively

large number of genera and species that have

evolved along similar lines.

12. As indicated by the dendrogram, Cirripectes

and Scartichthys are quite closely related. We considered treating them as subgenera but believe that

it would be premature to do so until the species

of these genera are better known. For the present

we recognize both genera because they are well entrenched in the literature (Strasburg and Schultz,

1953; Cohen, 1956), and we encountered no difficulty in assigning species to one or the other of

these genera (see discussion under Cirripectes).

13. Both Entomacrodus and Stanulus have recently been revised by Springer (1967; 1968).

There is considerable overall similarity between

the two species of Stanulus, especially S. talboti,

and the species of Entomacrodus.

14. Stanulus differs from Entomacrodus primarily

in having minute, imbricate scalelike flaps covering

the lateral-line pores, typically one less dorsal spine,

one more pectoral ray, and fewer segmented dorsal

and anal rays.

15. Nannosalarias does not appear to show especially close affinities to any other genus. We

tentatively place it near the base of those genera

centering around Cirripectes and Entomacrodus

because it shares with that group a distinctive type

of sexual dimorphism in which the anal spines of

mature males become enveloped in conspicuous

fleshy rugosities. Nannosalarias differs from those

genera in the general nature of its head pore distribution, and in having one less circumorbital

bone, one less segmented pelvic ray, and a membranous attachment of the terminal anal ray to the

caudal peduncle.

16. Such a wide range of characters is shown

between the species of Crossosalarias, Atrosalarias,

and Salarias that it is difficult to define them collectively as a group. The various combinations of

characters shared between the three genera, together

with their general physiognomy, however, leads us

to believe that they are more closely related to

each other than to other genera of Salariini.

17. Crossosalarias differs from all other genera

of Salariini in having restricted gill openings (the

gill membranes are bound to the isthmus at about,

or slightly below, the level of the ventralmost pectoral ray) and cirri associated with 3 or 4 preoperculomandibular pores on each side. It differs

from Salarias and Atrosalarias in having the innermost pelvic ray well developed, as opposed to rudimentary, and a fleshy flap at the base of the first

dorsal spine.

18. Atrosalarias differs from both Salarias and

Crossosalarias in having no branched caudal rays,

typically one more pectoral ray in each fin (usually

16), one to three less dorsal spines, and in having

the skin of the anal spines and first several segmented anal rays rugose in mature males (see

illustration in Springer and Smith-Vaniz, 1968).

19. Salarias differs from all other genera of Blenniidae in having the anterior anal segmented rays

elongated in large, mature males.

20. We place Glyptoparus as we do because it

does not appear to show any obvious affinities

with the other groups of salariinine genera. As in

one species of Ecsenius (E. mandibularis), Glyptoparus frequently has more than one (usually two

or three) posterior canines on each dentary, arranged in linear series. It does not, however, have

both anterior and posterior dentary canines or any

of the other numerous distinguishing osteological

characters of Ecsenius.

21. Included here are those genera that have

relatively few premaxillary (18 to 50) and dentary
52

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(16 to 38) teeth, and have the dentary replacement

teeth generally making their appearance through

foramina in the bone (Figure 7, except Rhabdoblennius snowi, Figure 6, in which the dentaries

are moderately excavated). All of the species belonging to this group of genera are relatively small,

have the terminal anal ray bound by a membrane

to the caudal peduncle, and usually 14 pectoral rays

(except Medusablennius, which has 13).

22. Medusablennius differs from its nearest relatives in having considerably fewer teeth, only two

circumorbital bones (no other salariinine has less

than 4), typically 13 pectoral rays, and attaining

maturity at a size when the sex of the other species

is hardly distinguishable. While its complex cirri

are to some extent duplicated in Mimoblennius

cirrosus, no salariinine species has such well-developed cirri at so small a size. Simple caudal rays are

found in group 21 only in Medusablennius and

Litobranchus; Medusablennius, however, has 13

segmented rays and Litobranchus has only 12. Except for Alloblennius pictus, Medusablennius is the

only genus of salariinines in which the ventral

hypural plate is fused to the urostylar centrum.

23. Mimoblennius differs from all other genera

in group 21 in having 13, instead of typically 12

dorsal spines. Although Mimoblennius has no

unique character that will separate it from all other

genera, when compared with individual genera in

group 21 there are numerous characters that set

it apart.

24. Litobranchus differs from all members of

group 21, except Medusablennius, in having simple

caudal rays. It differs from all members of group 21

in having only 12 instead of 13 segmented caudal

rays.

25. We believe that the genera of group 25 are

more closely related to each other than to any of

the other genera in group 21 because they appear

to be relatively unspecialized when compared with

the other genera. They exhibit none of the obvious

specializations of cirri or caudal fin structure found

in the other three genera. In his review of the

Blenniidae, Norman (1943) regarded Rhabdoblennius and Antennablennius as the most primitive

members of the subfamily Salariinae. He considered these two genera so closely related that he

accorded them only subgeneric rank.

26. Rhabdoblennius and Hirculops differ from

Antennablennius and Alloblennius in having vomerine teeth, well-developed canines in the lower

jaw (instead of lacking canines or having them

only weakly developed), and in having the last

POP pore position with a vertical pair of pores

(instead of with a single pore). Our concept of the

relationship of these four genera to each other has

been influenced by their geographic distributions

(see general accounts). The members of each pair

of genera are closely related and occupy adjacent

geographic areas with little or only slight overlap.

In each pair one genus is relatively speciose and

the other is either monotypic (Hirculops) or known

to comprise only two species (Alloblennius).

27. Hirculops differs from Rhabdoblennius and

all other genera included in group 21 in consistently

having 11 (instead of 10) precaudal vertebrae and

in having the premaxillary closed laterally with

replacement teeth entering the functional row

through foramina in the bone (Figure 4). Hirculops can also be distinguished from Rhabdoblennius by its extremely long supraorbital cirri

(Figure 46) and by its small nuchal cirri (absent

in Rhabdoblennius). The mutually exclusive geographic distributions of these two taxa reinforces

their generic separation.

28. Antennablennius differs from Alloblennius

in having 5, instead of 4, circumorbitals, and in not

having supraorbital cirri. Alloblennius is apparently restricted to the Red Sea and Antennablennius is known from the Red Sea and Indian Ocean.

Annotated List of Nominal Species, Tribe Salariini

The following list gives in order: (1) the scientific name in alphabetical order by species as it

originally appeared (hyphens omitted and capitalized specific names in lower case, however); (2)

the author or authors (Cuvier and Valenciennes,

1836, is abbreviated to C & V, 1836); (3) date of

publication (some species were described as new

by the same author on two occasions); (4) page

reference (see literature cited section for complete

reference); (5) the genus to which we currently

assign the species, if such is possible; (6) one or

two letters of the alphabet that indicate the basis

on which generic allocation was made (or other

pertinent information), as follows: A—holotype,

lectotype, neotype, or syntypes examined; B—para-
NUMBER 73 53

type examined; C—literature or other information

obtained sufficient for determination; D—probably

belongs in this genus, but should be verified (in

some instances this is impossible); E—questionable,

could belong in a different tribe of Blenniidae but

probably not; F—name unavailable by reason of not

having been presented properly, a nomen nudum;

(7) one or more museum abbreviations (see methods section for corresponding museum name) and

catalog numbers indicating holotypes, lectotypes,

neotypes, or syntypes. (Parentheses around information in 7 signifies that either we are not certain

of the information or have not seen the specimens

in question). Occasionally, comments referring to

a particular name have been included immediately

below that name. Noteworthy misspellings have

been included, as well as some names of questionable validity.

Following this list is a list of available names in

the genus Salarias that refer to species belonging

in blenniid tribes other than the Salariini. We

know of no other species originally described in

genera of the Salariini that do not belong in the

Salariini.

Species, author, publication date, and page reference

Antennablennius adenensis Fraser-Brunner, 1951, p. 219

Salarias aequipinnis Gunther, 1861, p. 253

Istiblennius afilinuchalis Schultz & Chapman, 1960, p. 356

Ecsenius albicaudatus Lotan, 1970, p. 372

Salarias alboapicalis Ogilby, 1899, p. 742

Salarias alboguttatus Kner, 1867, p. 16

Salarias aldabraensis Regan, 1912, p. 302

Salarias alticus Valenciennes in C & V, 1836, p. 337

Salarias amboinensis Bleeker, 1857a, p. 67

Praealticus amboinensis litteratus Schultz 8c Chapman, 1960,

p. 370

Blennius amphibius Walbaum, 1792, p. 187

Salarias andamanensis Day, 1878, p. 332

(misspelling of S. andamensis, see below)

Salarias andamensis Day, 1870b, p. 611

Salarias andersoni Jordan 8c Starks, 1906, p. 703

Salarias andersonii Day, 1878, p. 331

Salarias aneitensis Gunther, 1877, p. 205

Andamia anjouanae Founnanoir, 1954, p. 207

In his original description Founnanoir mentioned collecting 12

specimens, but he neither designated a holotype nor indicated

where he deposited his specimens. The specimen we list was

collected in the Comores by Founnanoir in 1954 and was identified by him. The total length of the specimen, 63.2 mm,

agrees closely with the average size mentioned by him in the

description. It seems probable that this specimen is a syntype.

Salarias anomalus Regan, 1905, p. 327

Salarias arenatus Bleeker, 1855a, p. 173

Blennius arnoldorum Curtis, 1938, p. 38

Ecsenius aroni Springer, 1971, p. 24

Alticus aspilus Bleeker, 1869, p. 235

Salarias atkinsoni Jordan Jc Seale, 1906, p. 428

Salarias atlanticus Valenciennes in C & V, 1836, p. 321

Salarias atratus Macleay, 1882, p. 361

The type material was reported as present in the Macleay

Museum by Stanbury (1969:209) , but neither of us was able to

Genus

Antennablennius

Andamia

Istiblennius

Ecsenius

Cirripectes

Salarias

Alticus

Alticus

Praealticus

Praealticus

Alticus

Istiblennius

Istiblennius

Salarias

Alticus

Entomacrodus

Alticus

Generic

allocation

A

A

A

A

A

C

A

A

A

A

C

C

C

A

A

C

A

Museum abbreviations

and

Catalog numbers

BMNH 1954.426.1

BMNH 1858.421.424

USNM 115421

HUI HUJ-F-3540

AMS1-1860-3

BMNH 19125.320-22

MNHN A-1793, A-1794,

A-1795.A-1797,

855

RMNH4781

USNM 124116

USNM 53273

ZSIC274

MNHN 1965-220

Ecsenius

Entomacrodus

Alticus

Ecsenius

Alticus

Entomacrodus

Ophioblennius

Istiblennius

A

C

D

A

A

A

A

D

BMNH 1900.5.9.47-56

18995.8.94-95

USNM 204468

RMNH4481

USNM 51791

MNHN A-2024

(MMS F-1047)
54

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Species, author, publication date, and page reference

find the specimens (searched for in 1966 and .1969, after appearance of Stanbury's publication). Stanbury, personal communication, did not see the specimens. An entry in the Macleay Museum catalog indicates that two specimens in poor

condition were present in 1965. The specimens were reidentified as "rivulatus" in the catalog. Other specimens in the Macleay Museum identified by De Vis as atratus belong in Istiblennius.

Salarias atrimarginatus Fowler, 1946, p. 182

Blennius atrocinctus Regan, 1909, p. 405

Salarias aureopuncticeps Fowler, 1946, p. 179

Salarias auridens Alleyne & Macleay, 1877, p. 338

Antennablennius australis Fraser-Brunner, 1951, p. 217

Salarias azureus Seale, 1906, p. 87

Blennius baliensis Nichols, 1954, p. 1

Ecsenius bandanus Springer, 1971, p. 38

Salarias barbatus Gilchrist Thompson, 1908, p. 108

Scartichthys basiliscus Fowler, 1904, p. 552

Salarias beani Fowler, 1928, p. 441

Salarias belemites De Vis, 1884b, p. 695

Salarias bellus Gunther, 1861, p. 256

Istiblennius bellus impudens Smith, 1959, p. 242

Salarias bicolor Day, 1888, p. 798

Blennius bifilum Gunther, 1861, p. 225

Salarias bilineatus Peters, 1868, p. 269

Salarias bilitonensis Bleeker, 1858b, p. 231

Ecsenius bimaculatus Springer, 1971, p. 38

Salarias biorni Fowler, 1946, p. 181

Salarias biseriatus Valenciennes in C & V, 1836, p. 316

Salarias bleekeri Chapman, 1953, p. 338

Salarias brasiliensis Sauvage, 1880, p. 217

Salarias brevis Kner, 1868a, p. 29; 1868b, p. 334

Salarias brevoorti Fowler, 1946, p. 177

Salarias bryani Jordan & Seale, 1906, p. 430

Salarias burmanicus Hora & Mukerji, 1936, p. 34

Entomacrodus cadenati Springer, 1966b, p. 59

Salarias caesius Seale, 1906, p. 88

Entomacrodus calurus Fowler, 1904, p. 555

Salarias calvus De Vis, 1884b, p. 697

Blennius canescens Carman, 1903, p. 236

Cirripectes caninus Herre, 1936a, p. 284

Ophioblennius capillus Reid, 1943, p. 381

Salarias castaneus Valenciennes in C & V, 1836, p. 324

Salarias caudofasciatus Regan, 1909, p. 405

Salarias caudolineatus Gunther, 1877, p. 209

Salarias celebicus Bleeker, 1854, p. 250

Salarias ceramensis Bleeker, 1852, p. 701

Salarias cervus Cuvier & Valcncinnes in Sauvage, 1880, p. 218

Medusablennius chani Springer, 1966a, p. 56

Entomacrodus chapmani Springer, 1967, p. 95

Salarias cheverti Macleay, 1881, p. 12

Salarias chiostictus Jordan & Gilbert, 1882, p. 363

Salarias chrysospilos Bleeker, 1857a, p. 66

Istiblennius chrysospilos insulinus Smith, 1959, p. 243

Blennius cinereus Castelnau, 1875, p. 26

Genus

Museum abbreviations

Generic and

allocation Catalog numbers

Istiblennius

Mimoblennius

Istiblennius

Istiblennius

Antennablennius

Istiblennius

Salarias

Ecsenius

Istiblennius

Istiblennius

Istiblennius

Istiblennius

Istiblennius

Istiblennius

Ecsenius

Antennablennius

Praealticus

Istiblennius

Ecsenius

Praealticus

Istiblennius

Istiblennius

Salarias

Exallias

Istiblennius

Istiblennius

Ecsenius

Entomacrodus

Praealticus

Istiblennius

Salarias

Cirripectes

Exallias

Cirripectes

Cirripectes

Entomacrodus

Istiblennius

Salarias

Salarias

Istiblennius

Medusablennius

Entomacrodus

Istiblennius

Entomacrodus

Istiblennius

Istiblennius

Istiblennius

A

A

A

A

A

A

A

A

C

C

A

A

A

C

C

A

A

A

A

A

A

A

A

C

C

A

A

A

A

A

A

A

A

A

A

A

C

A

A

A

A

A

A

A

A

C

£

ANSP 72052

BMNH 1909.3.4.70

ANSP 72050

MMS F-1051

BMNH 1920.12.6.24

BPBM 783

AMNH 16987

USNM 195717

(SAMN)

(ANSP 27802)

BPBM 1063

QMB1-224

BMNH 1855.9.19.700

(RU)

based on an illustration

BMNH 1859.5.7.70

ISZZ 6707

RMNH 4792

USNM 201817

ANSP 72051

MNHN A-2150

SU 37136

MNHN A-2148

(ANSP 72048)

USNM 51794

ZSICF-11872/1

MNHN 1965-705

BPBM 785

ANSP 27807

QMB 1-1355

MCZ 28294

SU 29082

USNM 120032

MNHN A-l 799

BMNH 1909.3.4.68

RMNH 4786

RMNH 4783

MNHN 2208

SU 62057

MCZ 29446

MMSF-1051A

USNM 28117

RMNH 4788

(RU)
NUMBER 73 55

Species, author, publication date, and page reference

Mimoblennius cirrosus Smith-Vaniz & Springer

Ophioblennius clarki Reid, 1943, p. 380

Salarias colei Herre, 1934, p. 96

Salarias concolor Philippi, 1896, p. 380

Giffordella corneliae Fowler, 1932a, p. 14

Blennius cornifer Riippell, 1830, p. 112

Hirculops cornifer menos Smith, 1959, p. 247

Salarias coronatus Giinther, 1872b, p. 424

Salarias crenulatus Weber, 1909, p. 144

Salarias crenulatus pallidus Whitley, 1926, p. 235

Salarias cristiceps Alleyne & Macleay, 1877, p. 338

Entomacrodus cruentatus Garman, 1899, p. 123

Salarias cruentipinnis Day, 1888, p. 797

Cirripectes cruentus Smith, 1959, p. 238

Rupiscartes cubensis Mowbray in Breder, 1927, p. 85

Salarias cuvieri Giinther, 1861, p. 248

In the description of S. cuvieri Giinther listed Salarias variolatus C & V as a synonym. He gave no reason for erecting a

new name for C & V's species. Giinther based his description

on material we list here, but he also cited data contained in

C 8c V's description. Thus, there is some question as to what

constitutes the type material of S. cuvieri: Giinther's specimens,

C & V's specimen, or both. Since Giinther's specimens are a

different species from C fc V's specimen, this problem must be

settled when the genus is revised.

Salarias, cyanostigma Bleeker, 1849, p. 18

Andamia cyclocheilus Weber, 1909, p. 143

Blennius cyclops var. guttatus Kossman & Rauber, 1877, p. 21

Blennius cyclops var. punctatus Kossman & Rauber, 1877, p. 21

Entomacrodus cymatobiotus Schultz & Chapman, 1960, p. 335

Salarias cypho Jenkins, 1903, p. 507

Salarias dama Valenciennes in C & V, 1836, p. 336

Blennius dama Hemprich & Ehrenberg, 1899, p. 3

The nomenclatural status of this name is debatable.

Salarias dayi Whitley, 1929, p. 136

A substitute name for "5. alboguttatus Day, 1876," which is

not listed above because Day was using Kner's S. alboguttatus.

Day neglected to mention Kner in the original listing, but

corrected the omission later (1888).

Salarias deani Jordan & Seale, 1905, p. 799

Entomacrodus decoratus Poey, 1868, p. 398

Salarias decussatus Bleeker, 1858b, p. 230

Glyptoparus delicatulus Smith, 1959, p. 249

Salarias diproktopterus Bleeker, 1857b, p. 69

Salarias doliatus Sauvage, 1880, p. 217

Salarias dussumieri Valenciennes in C & V, 1836, p. 310

Blennius edentulus Bloch, 1801, p. 172

Blennius ellipes Jordan & Starks, 1906, p. 702

Scartichthys enosimae Jordan & Snyder, 1902, p. 460

Salarias epalzeocheilus Bleeker, 1859b, p. 344

Salarias epalzeorhynchus Bleeker, 1859c, p. 218

Misspelling of epalzeocheilus often accorded to Giinther (1861).

Salarias eques Steindachner, 1898, p. 307

Genus

Mimoblennius

Cirripectes

Praealticus

Scartichthys

Entomacrodus

Hirculops

Hirculops

Isliblennius

Salarias

Salarias

Istiblennius

Entomacrodus

Cirripectes

Cirripectes

Ophioblennius

Scartichthys

Generic

allocation

A

A

A

C

A

A

C

A

A

A

A

A

C

C

C

A

Museum abbreviations

and

Catalog numbers

USNM 204491

CAS 24686

SU 25520

USNM 91821

NFIS 589

(RU)

BMNH 1871.3.29.42

ZMA 109.062,109.063

AMSIA-2609,

1-7104-7111

MMS F-1050B

MCZ 28695

based on an illustration

(RU)

(BOC 376)

BMNH 1850.6.14.19,

1850.6.14.21

Istiblennius

Andamia

?

?

Entomacrodus

Istiblennius

Istiblennius

Istiblennius and

Salarias (?)

Salarias

A

A

E

E

A

A

A

A

RMNH 4794

ZMA 109.061

USNM 142183

USNM 50697

ISZZ 1957

ISZZ 1957.

1946 (?)

Istiblennius

Entomacrodus

Entomacrodus

Glyptoparus

Istiblennius

Istiblennius

Istiblennius

Istiblennius

Rhabdoblennius

Istiblennius

Entomacrodus

Entomacrodus

Scartichthys

A

C

A

C

A

A

A

C

A

A

C

A

USNM 51950

RMNH 4778

(RU)

RMNH 4468

MNHN 2548

MNHN A-1740

USNM 53272

SU7068

ISZZ 15658
56

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Species, author, publication date, and page reference

Ophioblennius erythraeus Fowler, 1944, p. 251

Alticus evermanni Jordan & Seale, 1906, p. 422

Andamia expansa Blyth, 1858, p. 271

Blennius fasciatus Bloch, 1786, p. 29

Ophioblennius fernandezensis Clark, 1938, p. 184

Ophioblennius ferox Beebe & Tee-Van, 1928, p. 242

Salarias filamentosus Alleyne 8c Madeay, 1877, p. 337

Salarias flaviumbrinus Riippell, 1830, p. 113

Salarias fluctatus Fowler, 1945, p. 70

Salarias forsteri Valenciennes in C & V, 1836, p. 315

Salarias fowleri Herre, 1936b, p. 364

Salarias fraenatus Valenciennes in C &V, 1836, p. 342

Also spelled "froenatus"

Salarias frontalis Ehrenberg in C k V, 1836, p. 328

Salarias fronto Gunther, 1861, p. 255

Salarias furcatus Johnstone, 1904, p. 213

This is a homonym of 5. furcatus De Vis, 1884b, which belongs

in a different blcnniid tribe.

Salarias furvus De Vis, 1884b, p. 696

Cirripectus fuscoguttatus Strasburg & Schultz, 1953, p. 130

Salarias fuscus Riippell, 1838, p. 135

Salarias garmani Jordan & Seale, 1906, p. 429

Salarias geminatus Alleyne & Madeay, 1877, p. 336

Cirripectus gibbifrons Smith, 1947, p. 815

Salarias gibbifrons Quoy & Gaimard in C & V, 1836, p. 312

Istiblennius gibbifrons insolitus Smith, 1959, p. 242

Salarias gigas Steindachner, 1876, p. 220

Salarias gilberti Bryan & Herre, 1903, p. 135

Antennablennius girad Fraser-Brunner, 1951, p. 218

Blennius gobioides Forster, 1844, p. 283

Salarias goesii Bleeker, 1859a, p. 19

Salarias gravieri Pellegrin, 1906, p. 93

Salarias griseus De Vis, 1884a, p. 450

Salarias guttatus Valenciennes in C & V, 1836, p. 308

Salarias harmandi Sauvage, 1880, p. 218

Salarias hasseltii Bleeker, 1851, p. 257

Ecsenius hawaiiensis Chapman & Schultz, 1952, p. 526

Salarias hendriksii Bleeker, 1858b, p. 233

Salarias heteropterus Bleeker, 1857a, p. 65

Salarias histrionicus Kuhl & Van Hasselt in C & V, 1836, p. 320

Salarias holomelas Gunther, 1872, p. 399

Blennius hypenetes Klunzinger, 1871, p. 492

Entomacrodus incisolabiatus Schultz & Chapman, I960, p. 332

Cirripectes indrambaryae Smith, 1934, p. 322

Salarias insulae Ogilby, 1899, p. 741

Salarias interruptus Bleeker, 1857b, p. 68

Salarias irroratus Alleyne 8c Macleay, 1877, p. 337

Cirripectes jenningsi Schultz, 1943, p. 274

Blennius jugularis Klunzinger, 1871, p. 493

Salarias kellersi Fowler, 1932b, p. 6

Salarias kikaiensis Aoyagi, 1954b, p. 239

Springer (1967) omitted this spedes in his revision of Ento.

macrodus. The name is a junior synonym of Entomacrodus

decussatus (Bleeker, 1858b).

Salarias kingii Valenciennes in C 8c V, 1836, p. 334

Genus

Ophioblennius

Istiblennius

Andamia

Salarias

Scartichthys

Ophioblennius

Cirripectes

Istiblennius

Istiblennius

Istiblennius

Litobranchus

Entomacrodus

Ecsenius

Istiblennius

Ecsenius

Salarias

Cirripectes

Atrosalarias

Istiblennius

Istiblennius

Cirripectes

Istiblennius

Istiblennius

Scartichthys

Istiblennius

Antennablennius

Alticus

Praealticus

Ecsenius

Salarias

Salarias

Salarias

Istiblennius

Ecsenius

Istiblennius

Andamia

Salarias

Atrosalarias

Antennablennius

Entomacrodus

Cirripectes

Istiblennius

Istiblennius

Salarias

Cirripectes

Alloblennius

Istiblennius

Entomacrodus

Generic

allocation

A

A

(?)A,C

C

A

A

A

A

A

A

A

A

A

A

C

A

A

A

A

A

C

A

C

C

A

A

D

A

A

A

A

A

C

A

C

A

F

A

A

A

C

D

A

A

A

A

A

C

Museum abbreviations

and

Catalog numbers

ANSP 69997

USNM 51789

(?) BMNH 1924.3.7.3

CAS 5557

USNM 170901

MMS F-1035

NFIS 1893,8053-6

ANSP 71603

ISZZ 1942

SU3097

MNHN 996

ISZZ 1947

BMNH 1845.6.22.196

QMB 1-225

USNM 113634

NFIS 1832

USNM 51792

MMS F-1050

(RU)

MNHN 4002

(RU)

BPBM 2454

BMNH 1932.2.18.41-42

RMNH4659

MNHN 04-319

QMB 1-11/76

MNHN A-2049

MNHN A-2146

based on an illustration

USNM 112293

RMNH4779

BMNH 1872.10.18.108

ISZZ 8024,10498

USNM 142188

(KUMF 0174)

QMB 1-862

RMNH4787

AMS1-1042

USNM 115486

ISZZ 10496

USNM 91944

Istiblennius MNHN A-2048
NUMBER 73 57

Species, author, publication date, and page reference Genus

Salarias kirkii Giinther, 1868, p. 458

Lophalticus kirkii magnusi Klausewitz, 1964a, p. 133

Ecsenius klausewitzii Lotan, 1970, p. 371

Salarias kosiensis Regan, 1908, p. 254

Blennius kuasut Montrouzier, 1856, p. 463

Salarias kuhlii Bleeker, 1851, p. 258

Salarias lacunicola Fowler, 1946, p. 178

Ophioblennius lanieri Seale, 1940, p. 40

Entomacrodus leopardus Fowler, 1904, p. 554

Salarias leopardus Day, 1870a, p. 518

Salarias leopardus Fowler, 1938, p. 82

Salarias lighti Herre, 1938, p. 65

Ecsenius lineatus Klausewitz, 1962, p. 145

Salarias lineatus Valenciennes in C & V, 1836, p. 314

Salarias lineolatus Alleyne & Macleay, 1877, p. 336

Cirripectus lineopunctatus Strasburg, 1956, p. 248

Ecsenius lividinalis Chapman & Shultz, 1952, p. 517

Salarias lividus Thiolliere in Montrouzier, 1856, p. 463

Salarias luctuosus Whitley, 1929, p. 136

This is a replacement name for Salarias andersoni Jordan &

Starks, which is a junior homonym of 5. andersonii Day.

Rupiscartes macclurei Silvester, 1915, p. 217

Alticus macclurei Silvester, 1918, p. 24

The above two nominal species were described as new using

the same specimen as holotype. The second name is, therefore,

an objective synonym of the first.

Salarias macneilli Whitley, 1928, p. 227

Salarias macneilli coloratus Whitley, 1928, p. 229

Crossosalarias macrospilus Smith-Vaniz & Springer

Entomacrodus macrospilus Springer, 1967, p. 147

Ecsenius mandibularis McCulloch, 1923, p. 122

Salarias marcusi Bryan & Herre, 1903, p. 137

Salarias margaritaceus Poey, 1860, p. 289

Alticus margaritarius Snyder, 1908, p. 106

Alticus margaritatus Kendall & Radcliffe, 1912, p. 157

Blennius marmoratus Bennett, 1828, p. 35

Salarias martini Herre, 1942, p. 2

Ophioblennius mazorkae Hildebrand, 1946, p. 384

Salarias mccullochi Fowler it Bean, 1923, p. 25

Salarias melanocephalus Bleeker, 1849, p. 18

Salarias melanosoma Regan, 1909, p. 406

Salarias meleagris Valenciennes in C & V, 1836, p. 332

Ecsenius {Anthiiblennius) midas Starck, 1969, p. 1

Ecsenius minutus Klausewitz, 1963, p. 357

Fallacirripectes minutus Schultz & Chapman, 1960, p. 363

Salarias modestus Philippi, 1896, p. 381

Alticus monochrus Bleeker, 1869, p. 234

Salarias montanoi Sauvage, 1880, p. 219

Salarias mulleri Klunzinger, 1880, p. 388

Salarias multilineatus Fowler, 1945, p. 68

Museum abbreviations

Generic and

allocation Catalog numbers

Alticus

Alticus

Ecsenius

Pereulixia

?

?

Entomacrodus

Ophioblennius

Istiblennius

Exallias

Istiblennius

En tomacrodus

Ecsenius

Istiblennius

Salarias

Cirripectes

Ecsenius

Istiblennius

Salarias

Ophioblennius

Ophioblennius

Praealticus

Praealticus

Crossosalarias

En tomacrodus

Ecsenius

Istiblennius

Entomacrodus

Praealticus

Praealticus

Entomacrodus

Istiblennius

Scartichthys

Praealticus

Istiblennius

Ecsenius

Istiblennius

Ecsenius

Ecsenius

Stanulus

Scartichthys

Alticus

Alticus

Istiblennius

Istiblennius

A

A

A

A

F

£

A

A

A

C

A

A

A

A

A

A

A

D

C

C

A

A

A

A

A

A

A

A

A

C

A

A

A

A

A

A

A

A

A

C

A

A

A

A

BMNH 1868.2.29.48

NFIS6444

HUIE-62/3708

BMNH 1906.11.9.111

based on an illustration

ANSP 72049

CAS 5743

ANSP 27805

ANSP 68305

SU 33001

NFIS5532

MNHN 1396

MMS F-1052A

USNM 164198

USNM 144723

Lost, see

Montrouzier (1856)

Lost, see

Springer (1962)

Lost, see

Springer (1962)

AMS1-14286

AMSIA-2943

USNM 203096

USNM 200279

AMS 1-7112

BPBM 2556

MCZ 12513

USNM 62243

USNM 65409

SU 38253

USNM 128188

USNM 83293

RMNH 6297

USNM 157373,

BMNH 1909.3.4.52-57

MNHNA-1808.A-2041

ANSP 111149

NFIS 6367

USNM 142153

RMNH 4474

MNHN A-2199

NFIS (originally

SMNS 1519)

ANSP 71602
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Species, author, publication date, and page reference

Salarias muscarus Snyder, 1908, p. 109

Alticus musilae Jordan & Seale, 1906, p. 425

Ecsenius nalolo Smith, 1959, p. 245

Salarias namiyei Jordan & Evermann, 1902, p. 362

Salarias tmtulis Regan, 1909, p. 405

Blennius nativitatus Regan, 1909, p. 404

Labroblennius nicholsi Borodin, 1928, p. 31

Salarias niger Ehrenberg in C & V, 1836, p. 327

Bauchot (1967) gives reference to the supposed types of this

species. The specimens in question are the types of S. niger

Kossmann & Rauber.

Salarias niger Kossmann & Rauber, 1877, p. 21

Entomacrodus nigricans Gill, 1859, p. 168

Salarias nigripes Seale, 1901, p. 127

Salarias nigrovittatus Riippell, 1838, p. 136

Salarias nitidus Gunther, 1861, p. 243

Salarias nitidus Fourmanoir, 1954, p. 207

Salarias niuafoouensis Fowler, 1932b, p. 7

Alticus novemmaculosus Snyder, 1908, p. 107

Exallias obscurus Borodin, 1927, p. 1

Ecsenius oculus Springer, 1971, p. 35

Salarias olivaceus Blyth, 1858, p. 271

Salarias oortii Bleeker, 1851, p. 257

Ecsenius opsifrontalis Chapman & Schultz, 1952, p. 521

Alticus orientalis Tomiyama, 1955, p. 10

Salarias omatus Ehrenberg in C & V, 1836, p. 320

Blennius omatus Hemprich & Ehrenberg, 1899, p. 3

The nomenclatural status of this name is debatable.

Salarias oryx Ehrenberg in C & V, 1836, p. 335

Andamia pacifica Tomiyama, 1955, p. 13

Blennius pardalis Castelnau, 1875, p. 26

Salarias paulus Bryan & Herre, 1903, p. 136

Salarias pauper De Vis, 1884b, p. 695

Salarias penotus Sauvage, 1891, p. 389

Misspelling of Salarias fraenatus Valenciennes

Salarias percophthalmus Swainson, 1839, p. 274

Misspelling of Salarias periophthalmus Valenciennes

Blennius periophthalmoides Macleay, 1883, p. 269

Salarias periophthalmus Valenciennes in C & V, 1836, p. 311

Salarias periophthalmus visayanus Herre, 1937, p. 97

Blennius persicus Regan, 1905, p. 327

Salarias personatus Fowler, 1945, p. 71

Cirripectes perustus Smith, 1959, p. 238

Salarias petersoni Fowler, 1940, p. 189

Salarias phaiosoma Bleeker, 1855b, p. 317

Ophioblennius phalacrus Clark, 1938, p. 184

Salarias phntasticus Boulenger, 1897, p. 422

An obvious typographical error is involved and the species is

usually referred to as S. phantasticus.

Rhabdoblennius pictus Lotan, 1970, p. 376

Ophioblennius pinchoti Fowler, 1932a, p. 13

Genus

Istiblennius

Entomacrodus

Ecsenius

Ecsenius

Praealticus

Nannosalarias

Ophioblennius

?

Generic

allocation

A

A

A

C

A

A

A

E

Museum abbreviations

and

Catalog numbers

USNM 62245

USNM 51790

RU BP-2127C

ZITU 5726

BMNH 1909.3.4.58-67,

USNM 157374

BMNH 1909.3.4.42

VMM 408

Atrosalarias

Entomacrodus

Cirripectes

Ecsenius

Rhabdoblennius

Salarias

Entomacrodus

Istiblennius

Cirripectes

Ecsenius

Istiblennius

Istiblennius

Ecsenius

Alticus

Salarias

Salarias

Istiblennius

Andamia

Istiblennius

Istiblennius

Salarias

Entomacrodus

Istiblennius

?

Istiblennius

Istiblennius

Antennablennius

Istiblennius

Cirripectes

Scartichthys

Atrosalarias

Exallias

Ecsenius

Alloblennius

Ophioblennius

A

A

A

A

A

C

A

A

A

A

D

C

A

C

F

C

A

C

E

A

A

E

A

A

A

A

C

A

C

A

A

A

A

ISZZ 9854

USNM 34625

BPBM271

NFIS 1680

BMNH 1848.3.16.124-

125

USNM 91932

USNM 62244

AMNH 9363

USNM 203140

based on an illustration

USNM 142065

(ZITU 48353)

MNHN A-1803, A-1804

ISZZ 1954,1955

(ZITU 37924)

BPBM 2455

QMB 1-13/1344

(MMS)

MNHN 857

SU 28435

BMNH 1900.7.25.42-44

ANSP 71604

(RU)

ANSP 69150

CAS 5566

BMNH 1897.9.22.20-21

HUI HUJ-E63/37.1

USNM 91819
NUMBER 73 59

Species, author, publication date, and page reference

Entomacrodus plurifilis marshallensis Schultz & Chapman, 1960,

p. 341

Entomacrodus plurifilis plurifilis Schultz & Chapman, 1960, p. 338

Salarias (Cirripectes) polyzona Bleeker, 1868, p. 392

Salarias poptae Fowler, 1925, p. 19

Salarias priamensis Bleeker, 1853, p. 268

Ecsenius prooculis Chapman & Schultz, 1952, p. 519

Salarias pulcher Murray, 1887, p. 47

Salarias punctillatus Klunzniger, 1880, p. 389

Salarias quadricornis Valenciennes in C & V, 1836, p. 329

Salarias quadricornis status coloratus Klunzinger, 1871, p. 488

Salarias quadricornis status hyalinus Klunzinger, 1871, p. 487

Salarias quadricornis status transiens Klunzinger, 1871, p. 488

Salarias quadricornis status unitus Klunzinger, 1871, p. 488

We do not believe that the preceding four nominal forms are

available nomcnclaturally, but include them here for the sake

of completeness.

Salarias quadripennis Cuvier, 1817, p. 251

Spelled quadripinnis by Cuvier (1829) and subsequent authors.

Rupiscartes quagga Fowler & Ball, 1924, p. 273

Salarias raaltenii Bleeker, 1851, p. 257

Entomacrodus randalli Springer, 1967, p. 71

Andamia raoi Hora in Rao & Hora, 1938, p. 398

Salarias rarotongenis Whitley, 1965, p. 118

Salarias rechingeri Steindachner, 1906, p. 1411

Jstiblennius rennellensis Rofen, 1958, p. 199

Cirripectes reticulatus Fowler, 1946, p. 173

Salarias reuteri Lenz, 1881, p. 506

Salarias reyi Sauvage, 1880, p. 219

Blennius rhabdotrachelus Fowler & Ball, 1924, p. 272

Blenniella rhessodon Reid, 1943, p. 383

Salarias rivulatus Ruppell, 1830, p. 114

Istiblennius rodenbaughi Schultz & Chapman, 1960, p. 358

Entomacrodus rofeni Springer, 1967, p. 86

Salarias rotundifrons Thiolliere in Montrouzier, 1856, p. 463

Salarias rubropunctatus Valenciennes in C & V, 1836, p. 348

Salarias ruficaudus Ehrenberg in C & V, 1836, p. 328

Salarias rutilus Jenkins, 1903, p. 509

Blennius saliens Lacepede, 1800, p. 479

Salarias saltans Jenkins, 1903, p. 508

Alticus saltatorius Commerson in Lacepede, 1800, p. 479

Salarias sanna Whitley, 1929, p. 137

Replacement name for Salarias macneilli coloratus, Whitley,

which Whitley believed was a homonym of Salarias quadricornis coloratus Klunzinger (see above). We do not believe

Klunzinger's name is available nomenclaturally; therefore,

Salarias sanna is a junior objective snyonym of Salarias macneilli coloratus.

Salarias scandens Ehrenberg in C & V, 1836, p. 341

Salarias schmidti Fowler, 1946, p. 176

Genus

Entomacrodus

Entomacrodus

Cirripectes

Praealticus

Salarias

Ecsenius

Ecsenius

Salarias

Istiblennius

Istiblennius

Gobiidae

Istiblennius

Istiblennius

Generic

allocation

A

A

A

A

C

A

A

A

A

A

A

A

A

Museum abbreviations

and

Catalog numbers

USNM 142155

USNM 144720

RMNH 4797

BPBM 3413

USNM 144722

BMNH 1887.9.22.59-60

NFIS (orig. nos. SMNS

3660,4035)

MNHN 767, A-2002,

A-2003, A-2004, A-2005

A-2007

ISZZ 8085

ISZZ 8084

ISZZ 8086

ISZZ 8087

Salarias (UZMK)

Cirripectes

?

Entomacrodus

Andamia

Entomacrodus

Istiblennius

Praealticus

Cirripectes

Entomacrodus

Andamia

Rhabdoblennius

Istiblennius

Istiblennius

Istiblennius

Entomacrodus

Salarias

Scartichthys

A trosalarias

Istiblennius

A Iticus

Istiblennius

Alticus

Praealticus

A

E

A

A

A

C

B,C

C

E

A

A

A

A

A

A

D

A

A

A

C

A

C

BPBM 3420

Based on an illustration

USNM 199985

ZSICF-12933/1

AMSIZ-5358

(UZMK)

(ANSP 72043)

Destroyed; see

Springer (1967)

MNHN A-2201

BPBM 3419

USNM 118029

NFIS 1843

USNM 142067

SU 62035

lost, see Montrouzier

(1856)

MNHN A-2038

ISZZ 1948

USNM 50695

USNM 50696

Alticus

Praealticus ANSP 72046
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Species, author, publication date, and page reference

Salarias schultzei Bleeker, 1859b, p. 345

Leoblennius schultzi Reid, 1943, p. 382

Entomacrodus sealei Bryan & Herre, 1903, p. 138

Salarias sebae Valenciennes in C & V, 1836, p. 323

Alticus semicrenatus Chapman, 1953, p. 270

Blennophis semifasciatus Kner & Steindachner, 1866, p. 369

Salarias semilineatus Kner, 1867, p. 14

Salarias sertatus Garman, 1903, p. 236

Salarias sexfasciatus Von Bonde, 1924, p. 34

Salarias sexfilum Gunther, 1861, p. 241

Stanulus seychellensis Smith, 1959, p. 246

Salarias siamensis Smith, 1934, p. 320

Blennius sitnus Svief, 1779, p. 198

Salarias sindonis Jordan & Seale, 1906, p. 427

Salarias simonyi Steindachner, 1902, p. 148

Alticus simplicirrus Smith-Vaniz & Springer

Salarias sinuosus Snyder, 1908, p. 109

Salarias sinuosus indicus Smith, 1959, p. 241

Blennius snowi Fowler, 1928, p. 431

Salarias spaldingi Macleay, 1878, p. 358

Ophioblennius steindachneri Jordan & Evermann, 1898, p. 2401

Ophioblennius steindachneri clipper tonensis Springer, 1962,

p. 431

Salarias steindachnerii Pfeffer, 1893a, p. 143; 1893b, p. 15

Istiblennius steinitzi Lotan, 1970, p. 367

Scartichthys stellifer Jordan & Snyder, 1902, p. 461

Scartichthys stigmatopterus Fowler, 1904, p. 553

Cirripectes stigmaticus Strasburg & Schultz, 1953, p. 132

Ecsenius stigmatura Fowler in Chapman & Schultz, 1952, p. 514

Entomacrodus strasburgi Springer, 1967, p. 88

Salarias striatomaculatus Kner & Steindachner, 1866, p. 368

Salarias striatus Quoy & Gaimard in C & V, 1836, p. 309

Salarias strhiatus Day, 1878, p. 333

Salarias sublineatus De Vis, 1884b, p. 695

Salarias sumatranus Bleeker, 1851, p. 256

Stanulus talboti Springer, 1968a, p. 119

Salarias tanegasimae Jordan & Starks, 1906, p. 704

Salarias tetradactylus Bleeker, 1858b, p. 228

Salarias textilis Quoy & Gaimard in C & V, 1836, p. 307

Alticus thalassinus Jordan & Seale, 1906, p. 425

Entomacrodus thalassinus longicirrus Springer, 1967, p. 145

Alticus triangulus Chapman, 1953, p. 269

Blennius tridactylus Bloch, 1801, p. 176

This is a junior primary homonym of Blennius tridactylus

Lacepede (1800, p. 484), which belongs in another tribe. While

Bloch's description could apply to a number of different blenniids, it has consistently been applied to a species of Alticus.

Ophioblennius trinitatis Ribeiro, 1919, p. 177

Blennius truncatus Forster, 1844, p. 231

Blennius truncatus Poey, 1861, p. 424

Salarias tubuensis Seale, 1906, p. 87

Hepatoscartes umbrifasciatus Fowler, 1944, p. 230

Salarias undecimalis Jordan & Seale, 1905, p. 801

Salarias unicolor Ruppell, 1838, p. 136

Salarias unimaculatus Aoyagi, 1954a, p. 215

Genus

Istiblennius

Exallias

Entomacrodus

Cirripectes

Praealticus

Scartichthys

Salarias

Alticus

Antennablennius

Salarias

Stanulus

Istiblennius

Istiblennius

Antennablennius

A Iticus

Salarias

Salarias

Rhabdoblennius

Salarias

Ophioblennius

Ophioblennius

Istiblennius

Istiblennius

Entomacrodus

Istiblennius

Cirripectes

Ecsenius

Entomacrodus

Istiblennius

Entomacrodus

Istiblennius

Salarias

Istiblennius

Stanulus

Praealticus

Andamia

Entomacrodus

Entomacrodus

Entomacrodus

Praealticus

Alticus

Generic

allocation

C

A

A

A

A

E

C

C

C

A

A

C

p

A

A

A

A

A

C

C

A(?)

A

A

A

A

A

D

A

A

A

C

A

A(?)

A

C

A

A

A

A

A

A

A

E

Museum abbreviations

and

Catalog numbers

USNM 118037

BPBM 2458

MNHN A-2059

SU 17779

(MCZ)

BMNH 1855.9.19.835

RU (uncataloged)

(KUMFO177)

USNM 51793

NMV (uncataloged)

BMNH 1926.7.12.40

USNM 62246

(RU)

(MCZ 3634)

MMS F1045 (?)

NMV (uncataloged)

USNM 196030

ZSZM 6791

HUI E62/4040b

SU7069

(ANSP 27803)

USNM 164962

USNM 99379

USNM 179202

MNHN A-1796

ZSIC2011(?)

QMB1-13/1367

USNM 202421

USNM 53274

RMNH 4780

MNHN A-2025

USNM 51795

SU 62089

SU 17780

Ophioblennius

Istiblennius

Ophioblennius

Istiblennius

Ophioblennius

Salarias

Istiblennius

Istiblennius

C

E

C

A

A

A

A

C

(MNRJ 5423)

BPBM 784

ANSP 70211

USNM 51942

NFIS 1866
NUMBER 73 61

Species, author, publication date, and page reference

Ophioblennius vanderbilti Fowler, 1938, p. 242

Salarichthys vanderbilti Fowler, 1944, p. 234

Salarias variolatus Valenciennes in C & V, 1836, p. 346

Salarias variolosa Dclfin, 1899, p. 96

Salarias variolosus, Valendennes in C & V, 1836, p. 317

Cirripectes variolosus patuki De Buen, 1963, p. 58

Blennius variopunctatus Jatzow & Lenz, 1898, p. 511

Antennablennius (Litanchus) velifer Smith, 1959, p. 249

Salarias vermicularis Swainson, 1839, p. 274

A misspelling, which Springer (1967) omitted, of vermiculatus

Valenciennes.

Salarias vermiculatoides Bleeker, 1858a, p. 40

Salarias vermiculatus, Valenciennes in C & V, 1836, p. 301

Salarias viridis Valenciennes in C & V, 1836, p. 344

Blennius vitipinnis Castelnau, 1875, p. 25

Salarias vomerinus Valendennes in C 8c V, 1836, p. 344

Salarias walaensis Herre, 1931, p. 14

Salarias walensis Herre, 1935, p. 437

Ophioblennius watsoni Reid, 1943, p. 376

Blennophis webbii Valendennes in Webb & Berthelot, 1843, p. 61

Fallacirripectes wellsi Schultz & Chapman, 1960, p. 365

Entomacrodus wolffi Rofen, 1958, p. 202

Ophioblennius xiphiodon Clark, 1938, p. 183

Salarias yaeyamaensis Aoyagi, 1954a, p. 213

Salarias zamboangae Evermann & Seale, 1906, p. 512

Salarias zebra Varillant & Sauvage, 1875, p. 281

The following names available in the genus

Salarias pertain either to species in blenniid tribes

other than the Salariini or to unidentifiable species:

Salarias cephalotes Thiolierre in Montrouzier—unidentifiable

Salarias chilensis Clark—Blenniini

Salarias curtus Boulenger—Blenniini

Salarias cyclops Riippell—Blenniini

Salarias decipiens De Vis— Omobranchini

Salarias furcatus De Vis—Omobranchini

Genus

Museum abbreviations

Generic and

allocation Catalog numbers

Cirripectes C

Entomacrodus A

Scartichthys A

? F

Cirripectes A

Cirripectes C

Antennablennius C

Antennablennius C

En tomacrodus

Entomacrodus A

Entomacrodus A

Scartichthys A

Istiblennius A

Entomacrodus A

Rhabdoblennius F

Rhabdoblennius C

Ophioblennius A

Ophioblennius A

Stanulus A

Entomacrodus A

Scartichthys A

Ecsenius C

Istiblennius A

Istiblennius A

(ANSP 68432)

ANSP 70070

MNHN 4410

MNHN4411

(NFIS, lost)

(RU)

RMNH4777

MNHNA-1809

MNHNA-2128

BMNH 1883.7.4.56,

MNHNA-2120.A-2121

MNHN 3082

(FMNH 17393)

USNM 89614

MNHNA-2181

USNM 142154

UZMK P.75354

CAS 5543

USNM 55623

MNHN 8917,8918

Salarias furtivus De Vis—Omobranchini

Salarias galeatus De Vis—Omobranchini

Salarias helenae De Vis—Omobranchini

Salarius [sic] lupus De Vis—Nemophidini

Salarias neilli Day—Blenniini

Salarias opercularis Murray—Blenniini

Salarias sindensis Day—Omobranchini

Salarias symplocos Hilgendorf—Blenniini

Salarias varus Risso—Blenniini

Salarias vitoeridens De Vis—Nemo>hidin i i

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TABLE 2.—Summary of certain characters used to define genera of Salariini

(horizontal lines enclose major groups of genera—see section on recognition of genera)

Genus

Ecsenius

Andamia

Alticus

Praealticus

Istiblennius

Circumorbital

Species1 Incisoriform teeth2 Dentary canines Vomerine teeth bones

Premax- Denillary tary

24 (18) 148 64 present absent 4

26 13

5 (3) 382 300 absent absent 5

358 276

10 (5) 340 288 present present or absent 5

215 172 (males only)

or absent

(both sexes)

12 (8) 280 214 present or present or absent 5

190 128 absent

30 (18-f) 224 196 present or absent 5

208 164 absent
68

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 2.—Continued

Genus

Ophioblennius

Pereulixia

Exallias

Cirripectes

Scartichthys

Stanulus

Entomacrodus

Nannosalarias

Crossosalarias

Atrosalarias

Salarias

Glyptoparus

Medusablennius

Mimoblennius

Litobranchus

Rhabdoblennius

Hirculops

Antennablennius

Alloblennius

Circumorbital

Species1 Incisoriform teeth2 Dentary canines Vomerine teeth bones

Premax- DenMary tary

2 (2) 224 108 present absent 5

184 102

1 (1) 155 53 present present 5

157 43

1 (1) 210 62 absent absent 5

194 52

15 (8) 270 134 present absent 5

180 85

2 (2) 256 123 present absent 5

144 67

2 (2) 138 82 present present or absent 5

118 62

24 (22) 224 170 present present 5

116 92

1 (1) 80 52 present present 4

76 46

1 (1) 157 97 present absent 5

140 85

1 (1) 226 147 present absent 5

185 120

12 (7) 168 134 present absent 5

110 72

1 (1) 80 58 present absent 5

74 50

1 (1) 19 18 absent absent 2

17 16

2 (2) 34 30 present absent 5

32 24

2 (1) 42 30 present absent 5

40 23 (frequently

absent in

females)

8 (6) 50 38 present present 5

44 28

1 (1) 44 26 present present 5

34 22

10 (5) 42 34 present or absent 5

31 26 absent (minute

when present)

2 (2) 38 36 absent absent 4

34 30

1

 Estimated maximum number of species in genus followed, in parentheses, by number examined in our study.

2

 Counts are of adults; when more than one spedes was available, counts are of spedes with the most (upper pair of figures)

and least (lower pair of figures) number of teeth. When only one spedes was available, the upper pair of figures represents

the maximum counts recorded and the lower pair of figures Tepresents the minimum counts recorded for adults.
NUMBER 73 69

TABLE 3.—Summary of certain characters used to define genera of Salariini

(horizontal lines enclose major groups of genera—see section on recognition of genera)

Genus

Ecsenius

Andamia1

Alticus

Praealticus

Istiblennius

Ophioblennius

Pereulixia

Exallias

Cirripectes

Scartichthys

Stanulus

Entomacrodus

Nannosalarias

Crossosalarias

Atrosalarias

Salarias

Glyptoparus

Medusablennius

Mimoblennius

Litobranchus

Rhabdoblennius

Hirculops

Antennablennius

Alloblennius

Terminal

pleural ribs

Epipleural on vertebra Precaudal Caudal Total

ribs number vertebrae vertebrae vertebrae

11 to 25 lOorll 10tol2 19to29 29to40

14 to 19 10 10 28 to 30 38to40

13 to 32 10 to 12 lOorll 31 to 33 41to44

(11 in

A. kirki only)

11 to IS 10 10 26to28 36to38

12to25 lOtolS 10tol2 24to31 35to42

17tol9 11 10 22to26 32to36

13 or 14 10 10 20 30

19 to 21 11 10 20 30

17 to22 10 to 12 10 19 to 22 29to32

15 to 18 11 10 24 34

11 to 14 10 10 18 to 21 28 to 31

(rarely 11)

13 to 16 11 10 23 to 26 33 to 36

14tol6 10 10 24to25 34or35

13 to 15 11 lOorll 24 to 26 34 to 37

(11 in 4 of 5)

13 to 15 lOorll 10 22 to 26 32 to 36

12 to 16 lOorll 10 24 to 27 34 to 37

12orl3 10 10 24or25 34or35

? ? 10 26or27 36or37

13 to 19 10 10 25 to 28 35 to 38

13 to 15 10 10 27 or 28 37 or 38

19to22 10 10 27to29 37to39

25 to27 11 11 27 to29 38to40

12 to 15 10 10 27 to 29 37 to 39

11 to 16 10 10 26 to 29 36 to 39

1

 Fin ray counts for Andamia based in part on those reported in Chapman (1951).
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 4.—Summary of certain characters used to define genera of Salariini

(horizontal lines enclose major groups of genera—see section on recognition of genera)

Genus

Ecsenius

Andamia

Alticus

Praealticus

Istiblennius

Ophioblennius

Pereulixia

Exallias

Cirripectes

Scartichthys

Stanulus

Entomacrodus

Nannosalarias

Crossosalarias

Atrosalarias

Salarias

Glyptoparus

Mcdusablennius

Mimoblennius

Litobranchus

Rhabdoblennius

Hirculops

AntennabUnnius

Alloblennius

Dorsal rays Anal rays Caudal rays

spinous segmented total segmented branched segmented

11 to 14 13 to 21 25 to 34 13 to 23 0 13 to 15

(rarely 11) (rarely 15)

14 to 17 17 to 21 33 or 34 23 to 25 0 11 or 12

13 to 17 21 to 23 35 to 38 24 to 28 0 12 or 13

(13 in ,4.

simplicirrus

only)

12 to 14 18 to 20 31 to 33 18 to 23 0to9 12 or 13

(usually

4 to 8)

13 or 14 18 to 24 31 to 37 17 to 25 9 13

11 to 13 19 to 24 31 to 34 20 to 25 9 13

(12 in over

99% of specimens)

11 or 12 11 or 12 23 or 24 12 to 14 9 13

(usually 12)

12 12 or 13 24 or 25 14 9 13

12 14 to 16 26 to 28 14 to 16 9 13

(rarely 14)

12 17 or 18 29 or 30 18 to 20 9 13

12 9 to 12 21 to 23 10 to 13 9 13

(rarely 9)

12 to 14 13 to 18 26 to 31 14 to 19 9 13

(usually 13)

12 or 13 14 to 16 26 to 28 16 to 18 9 13

(usually 12)

12 16 to 19 28 to 31 18 to 20 9 13

9 to 11 18 to 22 28 to 32 18 to 21 0 10 to 14

(rarely 9) (usually

12 or 13)

12orl3 16to20 29to32 18to21 9 13

(usually 12)

12 or 13 15 or 16 27 or 28 18 9 13

(usually 12)

11 to 13 17 or 18 28 to 30 19 or 19 0 13

(usually 12)

12 to 14 16 to 19 29 to 32 19 to 22 9 13

(typically 13)

12 18to20 30to32 20to22 0 12

12 18to21 30to33 20to22 9 13

12 20 or 21 32 or 33 20 to 23 9 13

12orl3 17to21 29to33 19to23 9 13

(usually 12)

11 to 13 16 to 20 28 to 32 19 to 22 9 13

(usually 12)
NUMBER 73 71

TABLE 5.—Summary of certain characters used to define genera of Salariini

(horizontal lines enclose major groups of genera—see section on recognition of genera)

Genus

Ecsenius

Andamia

Alticus

Praealticus

Istiblennius

Ophioblennius

Pereulixia

Exallias

Cirripectes

Scartichthys

Stimulus

Entomacrodus

Nannosalarias

Crossosalarias

A trosalarias

Salarias

Glyptoparus

Medusablennius

Mimoblennius

Litobranchus

Rhabdoblennius

Hirculops

Antennablennius

Alloblennius

Terminal anal

Apparent ray bound by

segmented membrane Terminal anal Fleshy

Pectoral pelvic to caudal ray divided occipital

rays rays

1

 peduncle to base crest2

12 to 15 2 or 3 yes no absent

(rarely 12)

15 4 yes (except no present or absent

A. reyi)

14 or 15 3 or 4 no no present

14 or 15 3 or 4 no frequently present

12 to 15 3 or 4 no frequently present or absent

(rarely 15) (3 in most

species)

14 or 15 4 no frequently absent

(usaully 15)

15 or 16 4 no frequently absent

(usually 15)

15 4 no frequently absent

14 to 16 3 or 4 no frequently absent

(usually 15) (4 in most

species)

14 4 no frequently absent

14 to 16 4 no frequently absent

(usually 15)

12 to 15 4 no frequently absent

(usually 14)

14 or 15 3 yes no absent

(usually 15)

15 3 yes no absent

15 to 18 2 yes no absent

(usually 16)

13 to 15 2 yes no present or absent

(rarely 13)

13 2 yes no present

13 3 yes no absent

14 or 15 2 yes no absent

(usually 14)

14 3 yes no absent

13 or 14 3 yes no absent

(usually 14)

14 3 yes no absent

14 3 yes no present or absent

13 to 15 2 or 3 yes no absent

(usually 14)

1

 Refer to methods and material section for discussion of pelvic-fin rays.

2

 In species that have a fleshy occipital crest, it is always much better developed in males than in females and is frequently

absent in young of both sexes.
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 6.—Summary of certain characters used to define genera of Salariini

(horizontal lines enclose major groups of genera—see section on recognition of genera)

Genus

Ecsenius

Andamia

Alticus

Praealticus

Istiblennius

Ophioblennius

Pereulixia

Exallias

Cirripectes

Scartichthys

Stanulus

Entomacrodus

Nannosalarias

Crossosalarias

Atrosalarias

Salarias

Glyptoparus

Medusablennius

Mimoblennius

Litobranchus

Rhabdoblennius

Hirculops

Antennablennius

Alloblcnnius

Nuchal cirri Supraorbital cirri Nasal cirri

absent absent simple or paired

absent present (multifid) simple

absent present (simple or multifid) simple or multifid

absent present (pinnated branched) simple

(occasionally with

weak lateral points)

present or absent present simple to multifid

(highly variable) (usually multifid)

present present (simple) multifid

present present (long and simple) multifid

present present (multifid) multifid

present present (simple or multifid) multifid

present present (multifid) multifid

present (small) present (simple) simple

present or absent present (simple to multifid) simple to multifid

(present in most species)

present present (simple) simple

present present (multifid) multifid

present present (simple) simple

present present (simple or multifid) simple to multifid

present present or absent simple

(simple; minute)

absent present (variable) complex, multifid

present present (pair over each eye simple to multifid

or multifid)

present (minute) present (simple; minute to simple

well developed)

absent present (simple) simple

present present simple

(simple; well developed)

present absent simple

present (minute) present (ample; minute to simple

or absent well developed)

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In the bibliography (usually called "Literature Cited"), spell out book, journal, and article

titles, using initial caps with all words except minor terms such as "and, of, the." (For capitalization of titles in foreign languages, follow the national practice of each language.) Underscore

(for italics) book and journal titles. Use the colon-parentheses system for volume, number, and

page citations: "10(2) :5-9." Spell out such words as "figures" and "plates" (or "pages" when

used alone).

For free copies of his own paper, a Smithsonian author should indicate his requirements on

"Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will receive

50 free copies; order forms for quantities above this amount with instructions for payment will be

supplied when page proof is forwarded.