Synonymy of the Fish Families

Cheilobranchidae (=Alabetidae) and

Gobiesocidae, with Descriptions

of Two New Species of Alabes

VICTOR G. SPRINGER

and

THOMAS H. FRASER

I

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 234
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 23 4

Synonymy of the Fish Families

Cheilobranchidae (mAlabetidae) and

Gobiesocidae, with Descriptions

of Two New Species of Alabes

Victor G. Springer

and Thomas H. Fraser

SMITHSONIAN INSTITUTION PRESS

City of Washington

1976
ABSTRAC T

Springer, Victor G., and Thomas H. Fraser. Synonymy of the Fish Families

Cheilobranchidae ( = Alabetidae) and Gobiesocidae, with Descriptions of Two

New Species of Alabes. Smithsonian Contributions to Zoology, number 234, 23

pages, 14 figures, 3 tables, 1976.—The genus Alabes, monotypic representative

of the Cheilobranchidae, shares a large number of specializations with the members of the Gobiesocidae, including one not known to occur in any other fishes:

the supracleithrum bears a shallow, concave process at its distal end that articulates with a convex condyle on the anterior surface of the cleithrum. A

vestigial, papillae-bearing "sucking disc" occurs in some specimens of Alabes dorsalis. On the basis of shared specializations, we consider the Cheilobranchidae

to be highly specialized gobiesocids. The osteologies of Alabes dorsalis and the

gobiesocid Trachelochismus pinnulatus are illustrated. Four species are recognized in Alabes: dorsalis (widespread in southern Australia and Tasmania),

parvulits (widespread in southern Australia, Tasmania, and Norfolk Island),

brevis (new species from Western Australia), and hoesei (new species from New

South Wales, South Australia, and Tasmania).

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

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Springer, Victor Gruschka, 1928-

Synonymy of the fish families Cheilobranchidae ( = Alabetidae) and Gobiesocidae.

(Smithsonian contributions to zoology ; no. 234)

Supt. of Docs, no.: SI 1.27:234

1. Alabes. 2. Gobiesocidae. 3. Fishes—Classification. I. Fraser, Thomas H., joint author. II.

Title. III. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no. 234.

QL1.S54 no. 234 [QL638.G6J ">91'.08s [.r>97'..r>8] 76-608046
Contents

Page

Introduction 1

Methods 2

Osteological Material Examined 2

Acknowledgments 2

Osteology of Alabes and the Gobiesocidae $

Cranium 3

Superficial Bones, Jaws, and Suspensorium 3

Gill Arches and Hyoid Apparatus 7

Vertebral Column 9

Pectoral and Pelvic Fins, Girdles, and Supports n

Caudal Fin 13

Miscellaneous Remarks 14

Discussion 14

Conclusions 15

Intrafamilial Relationships of Alabes 15

The Species of Alabes 16

Alabes dorsalis (Richardson) 16

Alabes brevis, new species 19

Alabes parvulus (McCulloch) 19

Alabes hoesei, new species 21

Literature Cited 23

Synonymy of the Fish Families

Cheilobranchidae (=Alabetidae) and

Gobiesocidae, with Descriptions

of Two New Species of Alabes

Victor G. Springer

and Thomas H. Fraser

Introduction

The relationships of the genus Alabes ( = Cheilobranchus Richardson, 1835) have been a source of

confusion to ichthyologists since Alabes was described by Cloquet (1816).1

 Alabes was originally

placed next to the synbranchid eels under the

Anguilliformes (Cuvier, 1817). Swainson (1838),

somewhat presciently, considered Alabes to be intermediate between the eels and the Cyclopteridae,

which at that time embraced the Gobiesocidae, but

he included Alabes in the Muraenidae, even

though his observations should have caused him

to place it in his Synbranchidae. Swainson's

cyclopterid lead was not pursued by later authors.

Richardson (1845) considered his Cheilobranchus

(he was apparently unaware of Alabes) to be a

synbranchiform, and all authors for the next 60

1

 The authorship of Alabes has most often been attributed to

Cuvier (1817), who is actually responsible for the name.

Cuvier, however, furnished Cloquet with the name and description, and Cloquet's publication anticipated Cuvier (who

used the term alabes only in the vernacular sense) by just

a few months. The type-species of the genus is Alahes cwieri

Vaillant, 190">a, 1905b, by subsequent monotypy.

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

Washington, D.C. 20560. Thomas H. Fraser, Eiwiromnental

Quality Laboratory, 590-D Olean Boulevard, Port Charlotte,

Florida 33952.

years treated it, and Alabes, as a synbranchiform

or a genus of questionable relationships. Vaillant

(1905a, 1905b), in the most detailed study of Alabes

to that date, assigned Alabes to the Blenniidae, but

Gill (1906) disagreed, stating that Alabes was

neither a blenniid nor an eel and that its true

relationships would have to await a comparative

study of its osteology. Regan (1912) performed

such a study, albeit superficially, and returned

Alabes to the synbranchiforms, and there it rested

until Gosline (1968) stated that Alabes was possibly closely related to the congrogadoids, which

he included in his suborder Blennioidei. Gosline,

however, perferred to leave Alabes provisionally in

the Synbranchiformes, assuming that it was "so

specialized (degenerate) as to have obscured any

real evidence of relationship." Rosen and Greenwood (in press), in a study of the Synbranchiformes,

again excluded Alabes from close relationship with

that group and suggested that it was, perhaps,

more similar to the blennioids, based on certain

gill-arch specializations. This last study was probably the basis for Greenwood's (1975) inclusion of

the Alabetidae (junior synonym of Cheilobranchidae) in the suborder Blennioidei.

It is the purpose of our study to elucidate the

relationships of Alabes and to assign it to what we

believe is its proper family, the Gobiesocidae. In

addition, we revise the species of Alabes.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

METHODS.—Specimens used for the osteological

study were prepared by the enzyme method (Taylor, 1967). Springer prepared the base drawings

for the osteological illustrations using a Wild M-5

microscope and camera lucida. The final drawings

(Figures 1-10) were inked and labelled by P. K.

Hollingsworth. The osteological illustrations of

Alabes dorsalis and Trachelochismus pinnulatus

are based on a single specimen of each species (see

osteological material list). Three specimens of

Alabes dorsalis and one of A. parvulus were examined to determine osteological consistency. A

large number of Alabes specimens were examined

radiographically. Information on vertebrae, pectoral and pelvic-fin girdles, and certain skull features are easily discerned on radiographs. Only one

osteological preparation of Trachelochismus was

examined.

Vertebral counts, including those of the posteriormost vertebra bearing an epipleural rib, on

non-osteologically prepared specimens were taken

from radiographs. Vertebral counts were not recorded for specimens exhibiting apparent fusions

among vertebrae, with the exception of the penultimate vertebra, which in Alabes appears to be

aberrant more often than not.

The synonymies include only references to original descriptions.

The following institutional abbreviations are

used: AMNH = American Museum of Natural

History; AM = Australian Museum, Sydney;

BMNH = British Museum (Natural History);

MNHN = Museum National d'Histoire Naturelle,

Paris; NMNH = National Museum of Natural

History, Smithsonian Institution; NMNZ = National Museum of New Zealand, Wellington;

QM = Queensland Museum, Brisbane; USNM =

former United States National Museum (Specimens

in NMNH); WAM = Western Australian Museum, Perth.

A list of the osteological preparations examined

is given in the next section; non-prepared specimens of Alabes are listed under the general systematic accounts of the various species. For the

gobiesocids, we attempted to examine the osteology

of one species in at least one genus in each of the

eight subfamilies recognized by Briggs (1955). We

had no osteological material of Briggs' Diplocrepinae; however, our specimen of Gastrocymba,

which genus Briggs placed in the Trachelochisminae, has only three gills and would fit Briggs'

definition of the Diplocrepinae. Briggs reported

that the Trachelochisminae had three and one-half

gills. He had but one specimen of Gastrocymba,

which he was unwilling to dissect, and possibly he

erred in his belief that it had three and one-half

gills. In addition to the osteologically prepared

specimens, we examined many other gobiesocid

species externally and by radiographs; these species

are not listed.

OSTEOLOGICAL MATERIAL EXAMINED (Alabes and the Gobiesocidae).—Alabes dorsalis, USNM 201415, Kangaroo Island,

South Australia, about 60 mm TL (osteology illustrated in

Figures 1, 3-7, and 9); USNM 214100, Green's Beach, Tasmania, 28 mm TL; AMNH 31306, North Point, Western

Australia, about 100 mm TL; Alabes parvulus, AM IA.1253,

Long Reef, New South Wales, about 45 mm TL; Apletodon

pellegrini, USNM uncatalogued, Algoa Bay, South Africa,

2: 19.8-20.8 mm SL; Aspasma minima, USNM uncatalogued,

Japan, 36.2 mm SL; Chorisochismus dentex, USNM uncatalogued, Skoenmarkers Wreck, South Africa, 37.1 mm SL;

Conidens laticephalus, USNM 213838, Taiwan, 3: 15.5-25.3

mm SL; Diademichthys lineatus, USNM 213595, New Caledonia, 3: 26.5-41.0 mm SL; Diplecogaster bimaculata, USNM

213839, Tunisia, 24.5 mm SL; Eckloniaichthys scylliorhiniceps, USNM uncatalogued, Salt Vlei, South Africa, 25.2 mm

SL; Gastrocymba quadriradiata, NMNZ 3330, Auckland

Island, 24.5 mm SL; Gobiesox nigripinnis, USNM 213594,

Dominica, 3: 17.0-40.5; Gouania wildenowi, USNM uncatalogued, Genoa, Italy, about 32 mm SL; Lepadichthys

erythraeus, USNM 213593, Gulf of Aqaba, 3: 20.0-30.6;

Lepadogaster lepadogaster, USNM uncatalogued, Genoa,

Italy, 44.1 mm SL; Rimicola muscarum, USNM uncatalogued,

San Simeon, California, about 30 mm SL; Sicyases sanguineus, USNM 213846, Monte Mar, Chile, 3: 22.9-30.6 mm

SL; Tomicodon fasciatus, USNM 192157, Havana, Cuba, 3:

16.5-33.5 mm SL; Trachelochismus pinnulatus, USNM uncatalogued, Chatham, New Zealand, about 50 mm SL (osteology illustrated in Figures 2-6, 8, and 9).

ACKNOWLEDGMENTS.—We wish to express our appreciation to the following individuals: for loan or

donation of specimens—D. E. Rosen (AMNH),

D. F. Hoese (AM), P. J. Whitehead (BMNH),

M. L. Bauchot (MNHN), G. Allen (WAM), J.

Moreland (NMNZ), R. H. Green and E. O. G. Scott

(Queen Victoria Museum and Art Gallery of Tasmania); for curatorial assistance—E. N. Gramblin

and J. F. McKinney (NMNH); for reviewing the

manuscript—R. H. Gibbs, Jr., and S. H. Weitzman

(both NMNH), and D. E. Rosen (AMNH).
NUMBER 234

Osteology of Alabes and the Gobiesocidae

Unless indicated otherwise, wherever the terms

"gobiesocids" or "Gobiesocidae" are mentioned in

the osteological discussions, they refer only to those

genera and species in the osteological material list.

The osteology of Alabes was treated superficially

and often erroneously by Regan (1912). Among the

gobiesocids, Guitel (1888) treated most of the gross

anatomy of Lepadogaster gouani ( = L. lepadogaster) in considerable detail; Starks (1905) described

the osteology of Caularchus meandricus (=Gobiesox meandricus); Briggs (1955) gave limited osteological information on almost all the genera;

Runyan (1961) imperfectly illustrated the osteology

of Gobiesox strumosus but gave useful information

on variation in meristic osteological characters;

Leray (1961) described and illustrated the osteology

of the skull of Gouania wildenowi; and Rosen and

Patterson (1969) illustrated much of the osteology

of Gobiesox funebris (but we believe that they

have introduced several errors in their labels and

drawings).2

CRANIUM (Figures 1 and 2).—The cranium of

Alabes and the gobiesocids is broad and depressed.

The toothless vomer3 is relatively broad and concave anteriorly. The parasphenoid lacks ascending

processes; hence, the prootics enter the postorbital

area unimpeded. The following bones are not

present: basisphenoid, orbitosphenoid, pterosphenoids, and intercalars. The socket for articulation

with the dorsoanterior condyle of the hyomandibular is formed entirely by the sphenotic (rather

2

 In their figure 4F, Rosen and Patterson indicated that

the parhypural is fused to hypurals 1 and 2, that the uroneural is autogenous, and that there is no cpural. In our

specimens of Gobiesox nigripinnis the parhypural and

epural are present, autogenous, and cartilaginous (Rosen,

pers. comm., believes that the structure we call a parhypural

is not that but a "procurrent cartilage"; we do not agree).

We have not found an autogenous uroneural in any gobiesocid nor are we able to determine if the uroneurals have

been lost or fused to the urostylar complex in gobiesocids.

In their figure 9F they labelled the ectoptcrygoid as part ol

the quadrate. In their figure 53E the frontals, which are not

labelled, appear to be continuous (fused) with the sphenotics.

In their figure 58B they did not indicate whether cartilaginous basibranchials were present because they were only interested in structures that were ossified. The basibranchials

are cartilaginous in our specimens of G. nigripinnis.

3

 Scott (1974) reported that teeth were present on the

vomer, but we have not found this to be true of any ol

our specimens.

than being formed by the sphenotic dorsally and

prootic ventrally, as in many fishes). The supraoccipital lacks a dorsal crest. There is a sensory

canal with three branches in the frontal: an anterior branch that connects with the canal of the

nasal bone; a medial branch (commissure) above

the orbit that is continuous with the same branch

from the opposite frontal (commissure incomplete

in one of three specimens of Diademichthys); and

a posterior or posterolateral branch that, in all but

one of the gobiesocids and Alabes, is continuous

with the canal passing through the sphenotic and

pterotic. In Alabes dorsalis, A. parvulus, and

Gouania there are no sphenotic and pterotic sensory canals; however, based on the presence of a

postocular pore (Figure 10) in the skin lateral to

the sphenotic or pterotic, A. brevis and A. hoesei

probably have a sphenotic or pterotic canal. Neither Alabes nor the gobiesocids possess a supratemporal sensory canal.

The otoliths were not observed in Alabes or most

of the gobiesocids. In Trachelochismus the sagitta

and lapillus are of about the same size; the

asteriscus was not seen. The sagitta was fragile and

hollow, as was about one-third of the volume of

the asteriscus.

SUPERFICIAL BONES, JAWS, AND SUSPENSORIUM

(Figure 3).—Alabes and all the gobiesocids lack the

following bones: mesopterygoids, metapterygoids,

sesamoid articulars, extrascapulars (supratemporals),

and infraorbitals (except the lacrymal, which is

always present; however, the lacrymal is greatly

reduced in Diademichthys). Both Alabes and the

gobiesocids also lack a rostral cartilage (a sphere

of cartilage attached to the dorsoposterior surfaces

of the premaxillary ascending processes). The nasal

bone is relatively long, and in the gobiesocids

varies widely in shape; it lies parallel to the long

ascending process of the premaxillary. A lacrymal

sensory canal is present except in Gouania and

some specimens of Alabes parvulus. The lacrymal

lies lateral to the maxillary and is attached to the

lateralmost process of the lateral ethmoid. It may

be remote anteriorly from the lateral ethmoid

{Alabes and some gobiesocids) or attached closely

to that bone (other gobiesocids). The toothless

palatine is well developed and bears a maxillary

process in all the gobiesocids, but in Alabes the

palatine is reduced to a splint of bone without a

maxillary process. The palatine is not in bony
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

SUPRAOCCIPITAL

BASIOCCIPITAL

EXOCCIPITAL

EPIOTIC

PARASPHENOID

PROOTIC

EXOCCIPITAL

BASIOCCIPITAL

EXOCCIPITAL

BASIOCCIPITAL

LATERAL

ETHMOID

FRONTAL

FIGURE 1.—Alabes dorsalis, cranium: a, dorsal view; b, lateral view; c, ventral view.
NUMBER 234

MEDIAN

ETHMOID

SUPRAOCCIPITAL

EXOCCIPITAL

EPIOTIC

LATERAL

VOMER-i ETHMOID

EPIOTIC

^EXOCCIPITAL

BASIOCCIPITAL

EXOCCIPITAL

BASIOCCIPITAL

FRONTAL

PARASPHENOID

FIGURE 2.—Trachelochismus pinnulatus, cranium: a, dorsal view; b, lateral view;

c, ventral view.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PREMAXILLARY

NASAL

' \ LACRIMAL

ECTOPTERYGOID

HYOMANDIBULAR

OPERCLE

PREMAXILLARY

PALATINE

DENTARY

MAXILLARY

SYMPLECTIC

•QUADRATE

INTEROPERCLE

SUBOPERCLEINTERHYAL

PREOPERCLE

PREMAXILLARY ECTOPTERYGOIDMAXILLARY ^LACRIMAL / r H

NASAL-, / rPALATINE rlNTE OPERCLE

ANGULAR

ARTICULAR

SYMPLECTIC

INTEROPERCLE

 J

PREOPERCLE

SUBOPERCLE

FIGURE 3.—Superficial bones, jaws, and suspcnsoriuni: a, Alabes dorsalis (cranium also indicated) ; b, Trachelochismus pitmulatus. (Articular and angular bones are synonymous with

anguloarticular and retroarticular bones of recent authors.)
NUMBER 234

contact with the ectopterygoid in Alabes or the

gobiesocids, and is well separated from the ectopterygoid in all but Diademichthys, Conidens, and

Gastrocymba. The ectopterygoid is reduced to a

relatively small blade of bone in Alabes and all the

gobiesocids except Diademichthys, where it is quite

large, and usually lacks an anteriorly extending

process (a short process is present in Conidens,

Gastrocymba, and Diplecogaster). The interopercle

is entirely or predominantly medial to the ventral

portion of the preopercle and is well separated

from, though connected by a cordlike ligament to,

the subopercle in Alabes and all the gobiesocids

except Lepadichthys, Aspasma, and Chorisochismus, in which the interopercle and subopercle almost meet.

A large foramen perforates the posterior end of

the articular (anguloarticular) in Trachelochismus,

but is not present in Alabes or the other gobiesocids.

In Alabes dorsalis and the gobiesocids there is a

bioconcave disc of cartilage between the anterolateral head of the vomer and the anteromedial

surface of the maxillary. This cartilage occurs in

a wide variety of perciform fishes; however, the

cartilage is ossified on both sides of the single specimen of A. parvulus examined.

Neither the gobiesocids nor Alabes have scleral

bones in the eyes.

There is no preoperculomandibular sensory canal in Alabes. Among the gobiesocids, only Gastrocymba lacks this canal completely, but Aspasma,

Diademichthys, Diplecogaster, and Lepadichthys

lack the mandibular portion of the canal.

GILL ARCHES (Figure 4) AND HYOID APPARATUS

(Figures 5 and 6).—Alabes and the gobiesocids appear to lack the following structures, either as

bone or cartilage: basibranchial 1 (inasmuch as the

basihyal has a posterior process extending posterior

to the hypohyal—but anterior to hypobranchial 1

—and to which the urohyal attaches, it is possible

that basibranchial 1 is fused with the basihyal);

basibranchial 2 (it is possible that basibranchial 2

is fused with basibranchial 3 because in Alabes and

some of the gobiesocids the element herein termed

basibranchial 3 often has a short anterior process

that extends anterior to the medial end of hypobranchial 2); infrapharygobranchial 1; an uncinate process on epibranchial 1. In addition, there

is only one infrapharyngobranchial (which is

toothed) and one hypohyal on each side. It is

not known whether the infrapharyngobranchial is

infrapharyngobranchial 3 or a complex element

representing a fusion of some or all of infrapharyngobranchials 2-4 (toothplate); nor is it

known whether the hypohyal represents a fusion

of the dorsal and ventral hypohyals, or a loss of

one hypohyal.

In Alabes, basibranchial 3 is cartilaginous;

among the gobiesocids it may be cartilaginous or

ossified (one or the other in any taxon for which

more than one specimen was examined), but is possibly absent in Gastrocymba. In Alabes and all the

gobiesocids except Diademichthys, basibranchial 4,

where present, is cartilaginous. In Diademichthys

basibranchial 4 is ossified. Basibranchial 4 is possibly fused with basibranchial 3 in Apletodon and

appears to be absent in Gastrocymba, Tomicodon,

Gobiesox, Rimicola, and Sicyases.

Hypobranchial 1 is absent in Alabes and ossified in all the gobiesocids except Gastrocymba

and Eckloniaichthys, where it is absent. Hypobranchial 2 is partially ossified in Alabes dorsalis,

cartilaginous in A. parvulus, and, when present in

the gobiesocids, is usually at least partially ossified,

but is cartilaginous in Eckloniaichthys and possibly

absent in Gastrocymba and Diademichthys. Hypobranchial 3 is partially ossified in Alabes dorsalis,

unossified in A. parvulus, and is usually ossified

when present in the gobiesocids, but is cartilaginous

in Eckloniaichthys and Diademichthys, absent in

Lepadichthys, and possibly absent in Gastrocymba.

Ceratobranchials 1-5 and epibranchials 1-4 are

present and ossified in Alabes and all the gobiesocids.

The basihyal is present and ossified in Alabes

and all gobiesocids, but its relative development

is variable. In Alabes and nine of the gobiesocids

the basihyal has an ossified process that extends

well anterior to the anterior level of the hypohyals;

in two other gobiesocids the process is entirely

cartilaginous, and in five of the gobiesocids, no

portion of the basihyal is anterior to the hypohyals.

The basihyal extends posterior to the level of the

joint between the basihyal and urohyal in nine of

the gobiesocids, but in four of these the extension

is only slight; it does not extend posterior to the

joint in Alabes and the other gobiesocids.
BASIBRANCHIAL 3

HYPOBRANCHIAL 2

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

-HYPOBRANCHIAL 3

BASIBRANCHIAL 4

BASIHYAL

CERATOBRANCHIALS 1-5

EPIBRANCHIALS 1-4

INFRAPHARYNGOBRANCHIAL

BASIBRANCHIAL 3

L HYPOBRANCHIALS 1-3

BASIBRANCHIAL 4

BASIHYAL

CERATOBRANCHIALS 1-5

EPIBRANCHIALS 1-4

INFRAPHARYNGOBRANCHIAL

Fio.i'RE 4.—Dorsal view of left gill arches and unpaired median elements (cartilage indicated by pattern of small, open circles; epi- and infrapharyngobranchials folded back,

relative positions especially distorted in b; gill rakers and teeth indicated but not labelled; position of basihyal relative to more posterior median elements indicated): a,

A tabes donalis; b, Traclielochismus pitmulatus.
NUMBER 234

BASIHYAL ^HYPOHYAL

CERATOHYAL

OSSEOUS

INCLUSIONS

(SESAMOID)

UROHYAL

FIGURE 5.—Lateral view of basihyal and urohyal and dorsal view of urohyal

(cartilage indicated by pattern of small, open circles) : a, Alabes dorsalis (relationship of adjacent bones on right side indicated; extraneous osseous inclusions indicated); b, Trachelochismus pinnulatus.

The joint between the epihyal (posterior ceratohyal) and ceratohyal (anterior ceratohyal) is nonsutured in Alabes and all the gobiesocids.

The number of branchiostegals is always three in

all species of Alabes (all articulate with the ceratohyal) and normally five or six in the gobiesocids

(most species in the Gobiesocidae—entire family

considered—have six, with seven in variant specimens), but the articulations of the branchiostegals

are quite variable. For instance, all six articulate

with the ceratohyal in Aspasma, five with the ceratohyal and one with the epihyal in Diplecogaster,

and four with the ceratohyal and two with the

epihyal in Gobiesox. Where five branchiostegals

are present (Gastrocymba, Eckloniaichthys), it is the

anteriormost branchiostegal that is absent.

Osseous inclusions (sesamoid bones) in various

ligaments in the anterior region of the hyoid

apparatus were noted in Alabes (Figure 5) and

several of the gobiesocids.

VERTEBRAL COLUMN (Figures 7 and 8).—Alabes

has 18-25 precaudal, 39-55 caudal, and 60-78 total
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

HYPOHYAL

INTERHYAL

EPIHYAL

CERATOHYAL

BRANCHIOSTEGALS

INTERHYAL'

EPIHYAL

L

HYPOHYAL

BRANCHIOSTEGALS

FIGURE 6.—Right hyoid arch (cartilage indicated by pattern of small, open

circles): a, Alabes dorsalis; b, Trachelochismus pinnulatus (branchiostegals

truncated).

vertebrae (see Tables 2 and 3). The number of

precaudal or caudal (but not total) vertebrae is

difficult to determine, especially from radiographs,

and our figures should be considered only as approximations. The last one or two precaudal vertebrae have complete hemal arches but no hemal

spines. The epipleural ribs begin on the third

from anteriormost vertebra and extend no farther

posteriorly than the third caudal vertebra. All but

the posteriormost epipleural ribs attach directly

to the constricted portions of their respective

centra; there are no pleural ribs.

The gobiesocids have 11-20 precaudal, 13-33

caudal, and 25-54 total vertebrae. There are 0-3
NUMBER 234 11

FIRST COMPLETE

HEMAL ARCH

FIRST HEMAL SPINE

rEPURAL

UROSTYLAR COMPLEX

PARHYPURAL CAUDAL-FIN RAYSJ

FIGURE 7.—Alabes dorsalis: a, five anteriormost precaudal vertebrae; b, posteriormost precaudal and anteriormost three

caudal vertebrae; c, posteriormost three caudal vertebrae and

caudal-fin rays (rays truncated, parhypural cartilaginous).

precaudal vertebrae with complete hemal arches.

The epipleural ribs begin on the second or third

from anteriormost precaudal vertebrae and extend

as far posteriorly as the last precaudal to the tenth

caudal vertebra. The articulation of the epipleural

ribs varies from all ribs attaching to the constricted

portions of their respective centra to only the precaudal ribs attaching to the constricted portions

of the centra, with the caudal ribs gradually being

displaced ventrally and attaching to the hemal

arches. All gobiesocids have pleural ribs that attach to the epipleural ribs well proximal to the

distal ends of the epipleural ribs (an uncommon

condition in fishes, where the epipleural ribs, except for the anteriormost one or two, ordinarily

attach to the pleural ribs). The pleural ribs begin

on the epipleural ribs of the third from anteriormost precaudal vertebra and extend no farther

posteriorly than the third from posteriormost to

the posteriormost precaudal vertebra.

Vertebral fusions and anomalies are common in

Alabes, but not in the gobiesocids.

PECTORAL AND PELVIC FINS, GIRDLES, AND SUPPORTS (including sucking disc) (Figures 9 and 11).—

In Alabes and all the gobiesocids the posttemporal

lacks a ventral arm, but has in its place a strong

ligament that attaches to the exoccipital. The

posttemporal is somewhat sigmoid in shape and

its longitudinal axis is nearly perpendicular to the

longitudinal axis of the skull (in most, if not all,

other acanthopterygiian fishes the longitudinal axis

of the posttemporal is nearly parallel to the longitudinal axis of the skull).

In Alabes and all the gobiesocids, the supracleithrum bears a shallow, concave process on its

distal end that articulates with a convex condyle

on the anterior surface of the cleithrum. The joint

between these two bones is not duplicated in other

fishes, where the supracleithrum is a compressed,

bladelike bone whose medial surface is joined

(appressed) to the flattened lateral surface of the

cleithrum. Furthermore, the longitudinal axis of

the supracleithrum is roughly perpendicular to the

longitudinal axis of the posttemporal, whereas in

other acanthopterygiian fishes the axes of these

two bones are parallel.

Alabes lacks a scapula and pectoral-fin radials

and rays. The coracoid of A. dorsalis is reduced

to a small, rodlike bone with a relatively large

cartilage on its posterior surface; all the other

species of Alabes lack coracoids. The gobiesocids

have essentially normal scapulae and coracoids,

four radials, and 16-31 simple pectoral-fin rays.

Alabes lacks postcleithra; Gastrocymba quadriradiata, Gastroscyphus hectoris, and, according to

Bohlke and Robins (1970), Gymnoscyphus ascitus

lack,dorsal postcleithra; but all the other genera
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

UROSTYLAR

COMPLEX-.

FIRST COMPLETE

HEMAL ARCH

PLEURAL RIB CAUDAL-FIN RAYS

PARHYPURAL

FIGURE 8.—Trachelochismus pinnulatus: a, four auuriormost prccaudal vertebrae; b, postcrionnost two prccaudal and antcriormost three caudal vertebrae; c, postcriormost two caudal

vertebrae and caudal-fin rays (segmented rays truncated; cartilage indicated by pattern of

small, open circles).

(entire Gobiesocidae considered) apparently have

both dorsal and ventral postcleithra.

The pelvises (pelvic bones) of Alabcs dorsalis are

greatly reduced with relatively large cartilaginous

anterior ends, and each pelvis supports three

greatly reduced fin rays; it was not possible to determine if any of the three represent a spine. All

the other species of Alabes lack pelvises and pelvicfin rays. The pelvises of the gobiesocids are well

developed, but highly modified for support of the

sucking disc; each pelvis supports a modified spine

and four simple, segmented rays.
NUMBER 234 13

O PELVISi

PELVIC-FIN RAYS

CORACOID

CLEITHRA

PECTORAL-FIN

RADIALS

SUPRACLEITHRUMPOSTTEMPORAL

SCAPULA

SUPRACLEITHRUM

POSTTEMPORAL

CLEITHRUM

CORACOID

VENTRAL POSTCLEITHRUM

DORSAL POSTCLEITHRUM

PELVIC-FIN RAYS

PELVIC-FIN SPINE

*- PELVIS

FIGURE 9.—a, Alabes dorsalis, ventral view of pelvic and

pectoral-fin girdles; b, Trachelochismus pinnulatus, lateral

view of pectoral-fin girdle (fin rays omitted); c, Trachelochismus pinnulatus, ventral view of pelvises and right pelvic

fin and postcleithra (cartilage indicated by pattern of small,

open circles).

The postcleithra, pelvises, and pelvic-fin rays of

gobiesocids function in support of the ventral sucking disc, which characterizes the Gobiesocidae.

Externally, the sucking disc bears fleshy papillae

(one exception is Gymnoscyphus ascitus, according

to Bohlke and Robins, 1970). Alabes dorsalis has,

at most, a vestigial sucking disc, which was present

in only nine of more than 150 specimens examined.

In these nine specimens the skin between the

pelvic fins forms a fold, which is covered with tiny

papillae that are extremely difficult to see. The

fold may be the result of shrinkage during preservation. The vestigial sucking disc is illustrated in

Figure lid; the usual appearance of the region

around the pelvic fins is illustrated in lie. None

of the other species of Alabes showed evidence of

a sucking disc.

CAUDAL FIN.—In Alabes the entire caudal fin is

greatly reduced in size relative to the rest of the

body and relative to its development in the gobiesocids. All of the hypurals are fused into a fan

(which is, in turn, fused to the urostylar complex)

and are indistinguishable individually. In A.

dorsalis there is no diastema separating the dorsal

and ventral halves of the hypural fan, although a

tiny foramen (see Figure 7c) representing a vestigial

diastema may be present in the hypural plate. The

parhypural is autogenous and cartilaginous; there

is a single epural, which is ossified; the uroneurals

are either absent or fused indistinguishably with

the urostylar complex. All of the fin rays are segmented and there are a total of 8-11 rays, of which

7 or 8 articulate with the hypural fan; there may

be 0-2 dorsal procurrent rays and 0-1 ventral procurrent rays. The neural spine of preural centrum

2 is long. The caudal fin of A. brevis appears to be

similar to that of A. dorsalis, but there is a diastema

present. The caudal fins of the other species are

quite variable, but usually are not well developed

and often appear to be aberrant.

In the gobiesocids the caudal fin is better developed than in Alabes, but does vary from greatly

reduced (Gastrocymba) to not much reduced

(Trachelochismus) in size relative to the rest of

the body. The condition of the hypurals and uroneurals is similar to that of Alabes, but a diastema

always separates the dorsal and ventral halves of

the hypural fan. The parhypural, when present,

is reduced to a distal plate that is autogenous and

either ossified or cartilaginous (both conditions occur in a single species of Tomicodon). In Gastrocymba, Aspasma, and Rimicola it was not possible
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

to determine if the parhypural is present, but if

present it is cartilaginous and greatly reduced or

fused to the hypural fan. There is a single epural,

which may be ossified or cartilaginous (both conditions occur in a single species of Conidens). It

was not possible to determine if an epural is present in Gastrocymba, Eckloniaichthys, and Rimxcola, but if present it is cartilaginous and greatly

reduced. The fin rays comprise both segmented

(principal and procurrent) and non-segmented

(procurrent) simple rays totaling 16-27 rays, of

which 8-14 articulate with the hypural fan. There

may be 3-7 dorsal procurrent rays and 3-8 ventral

procurrent rays. The neural spine of preural centrum 2 is long.

Miscellaneous Remarks

Alabes has three gills; the gobiesocids have three

or three and one-half gills. The gill opening of

Alabes consists of a common opening to both gill

chambers. The opening is situated on the midventral

side of the head. In A. dorsalis it extends dorsally

on either side to, or slightly dorsal to, the dorsal

level of attachment of the pelvic-fin membrane to

the body. In the other species of Alabes the gill

opening is restricted entirely to the midventral

area.

In the gobiesocids there are two types of gill

opening. One type consists of a common opening

similar to that of Alabes except that the opening

extends much farther dorsally on either side (as

far as the dorsal margin of the pectoral-fin base).

In the other type, there is a separate opening to

the gill chamber on each side of the fish.

Neither Alabes nor the gobiesocids have scales

or lateral-line canals on the body. Both Alabes and

the gobiesocids have a genital (urogenital?) papilla

just posterior to the anus.

Waite (1923) noted that Alabes dorsalis is green

with darker spots when alive, but that the color

changes to red in certain preservatives (the same

change takes place in Alabes parvulus). We have

noted that certain gobiesocids (Diademichthys,

Lepadichthys) also turn red or pink when placed

in formalin.

Discussion

Alabes shares at least one specialized osteological

character with gobiesocids that is not found in other

fishes: the structure of the joint between the supracleithrum and cleithrum. The angle at which the

posttemporal articulates relative to the longitudinal axis of the skull may also be unique to Alabes

and the gobiesocids.

There are many additional, specialized characters (mostly reductional and osteological) that

Alabes shares with some or all the gobiesocids.

None of these characters is unique to Alabes and

the gobiesocids, but the number and combination

of these characters is not duplicated in any other

group of fishes. Included among these specializations are the following (* indicates conditions in

Alabes and some gobiesocids). Absence of: scales;

basisphenoid; orbitosphenoid; pterosphenoid; intercalar; ascending processes on parasphenoid;

supraoccipital crest; dermosphenotic; metapteryANP

AN DLP PN POP

ANP

FIGURE 10.—Diagrammatic illustration of sensory canal pores

in Alabes. (Abbreviations: AN •» anterior nostril; ANP =

anterior nasal canal pore; DLP — dorsal lacrymal canal

pore; PN — posterior nostril; PNP — posterior nasal canal

pore; POP — postocular pores; VLP — ventral lacrymal

canal pore.)
NUMBER 234 15

goid; mesopterygoid; sesamoid articular (cornomeckelian); infraorbitals (except lacrymal); lateral

or median extrascapulars; scleral bones; vomerine

and palatine teeth; Baudelot's ligament; rostral cartilage; infrapharyngobranchial 1; uncinate process

on epibranchial 1; hypobranchial 1*; basibranchials 1 and 2 (except that basibranchial 1 is possibly

fused to the basihyal); ventral process on posttemporal; ribs on first vertebra; ribs on second vertebra*; autogenous hypurals or uroneurals; lateralline canals on body; and supratemporal, infraorbital (except lacrymal), and preoperculomandibular*

sensory canals. Presence of: only one hypohyal; no

more than one epural; cartilaginous parhypural*;

eight principal caudal-fin rays*; and a long neural

spine on preural centrum 2. In addition: the socket

for articulation with the dorsoanterior condyle of

the hyomandibular is formed entirely by the sphenotic; most of the epipleural ribs attach directly

to the constricted portion of their respective centra;

infrapharyngobranchials 2, 3, and the toothplate of

4 are represented by a single toothed bone (the

composition of which is unknown); the interopercle is mostly medial to the preopercle and is

not in bony contact with the subopercle; the palatine is not in bony contact with the ectopterygoid;

and there are only three gills*.

Alabes differs most prominently from the gobiesocids in exhibiting the following specializations. Absence of: dorsal, anal, and pectoral-fin

rays, pterygiophores, and radials; maxillary process

on the palatine; pleural ribs; postdeithra (the

dorsal postcleithrum is absent in some gobiesocids);

and scapulae. Reduction in: number of branchiostegals and pelvic-fin rays; size and/or structure of

pelvises, sucking disc, coracoids, and gill opening.

Increase in: number of vertebrae.

No specialization was found that unites all the

gobiesocids to the exclusion of Alabes. This statement is based on our assumption that the most

specialized state for a character must be considered as having passed through (and hence embodying) all the character states leading to it.

For instance whereas all gobiesocids have at least

one pair of specialized postcleithra modified for

support of the sucking disc, Alabes has no postcleithra. We believe that evolution of the postcleithra has gone from two to one to no pairs of

postcleithra in gobiesocid-Alabes phylogeny.

Conclusions

Inasmuch as (1) Alabes exhibits at least one

specialization found otherwise only in the gobiesocids, (2) it shares a large number of other specializations with all or some gobiesocids, (3) all the

other specializations of Alabes (except number of

vertebrae) can be derived by reduction from

gobiesocid character states (many of which show

tendencies toward reduction even among the gobiesocids), and (4) the gobiesocids share no specialization not also found in Alabes, we believe that

Alabes is a gobiesocid. We, therefore, synonymize

the Cheilobranchidae Gill (1872, emended from the

original Chilobranchidae; =Alabetidae Gill, 1906,

occasionally spelled Alabidae in the literature)

with the Gobiesocidae Bleeker (1860, emended

from the original Gobiesocioidei).

Intrafamilial Relationships of Alabes

The relationships of Alabes to the other genera

are difficult to assess because of the extreme state

of reduction in Alabes of most of the characters

ordinarily used to relate gobiesocids. We agree

with Bohlke and Robins (1970) that Briggs' (1955)

subfamilial classification is in need of major revision, but we prefer not to attempt that extensive

task. The brief discussion that follows is a distillation of our impressions based on examination

(varying from superficial to detailed) of numerous

gobiesocid genera and species.

With the exception of number of vertebrae,

Alabes appears to have followed an overall path of

reduction, much more so than any other clingfish

genus. As far as we know, the genus Gastrocymba,

among the other gobiesocids, exhibits the greatest

overall degree of reduction (for example: no

preoperculomandibular sensory canal or pores,

only one pair of postcleithra, only five branchiostegals, epipleural ribs begin on third vertebra).

In addition, Gastrocymba has more vertebrae (total:

53-54) than any other gobiesocid except Alabes

(total: 60-78). A high number of vertebrae appears

to be a specialization in gobiesocids; hence, the

general trend toward reduction in Gastrocymba,

together with an increase in number of vertebrae,

indicates two paths of specialization coupled otherwise only in Alabes. While the data are admittedly
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

scanty, we propose that Gastrocymba is the genus

most closely related to A lobes (that is, the two

genera form a sister group).

The Species of Alabes

We recognize four species in Alabes, two of

which we describe as new. For purposes of identification, we provide a table of diagnostic characters (Table 1) for the species rather than a key.

The table also serves as the primary description

of each species, and many of the characters are not

repeated in the formal descriptions.

Alabes dorsalis (Richardson)

FIGURE 11

Cheilobranchus dorsalis Richardson, 1845 [N'W coast of Australia, BMNH 1848.3.18.37, 4 syntypes].

Cheilobranchus aptenodytum Richardson, 1845:51 [above

highwater mark on Penguin Island, 72°S latitude, BMNH

1848.3.18.75, holotype].

Chilobranchus rufus Macleay, 1881:266 [Port Jackson, AM

1.16264-001, 14 syntypes, and Tasmania, AM 1.16265-001,

17 syntypes].

Alabes cuvieri Vaillant, 19O5a:149; 1905b:1714 [NouvelleHollandc = Australia, MNHN 2181, 3 syntypes].

The diagnostic characters of this species are

given in Tables 1 and 2 and are not repeated here.

REMARKS.—Variation in numbers and number

of posteriormost vertebra bearing an epipleural rib

is reported in Table 2. In eastern Australia there

is a tendency for average numbers of vertebrae to

increase from north to south, with the exception

that specimens from South Australia have lower

averages than might be expected from the longitudinal position of the localities. The trends in the

average number of the posteriormost vertebra

bearing an epipleural rib are not clear, but the

highest averages are found in populations from the

area (Tasmania) also having the highest average

numbers of total vertebrae.

NOMENCLATURAL DISCUSSION.—Richardson differentiated his C. aptenodytum from his C. dorsalis

TABLE 1.—Comparison of the diagnostic characters of the species of Alabes

dorsalis brevis hoesei

Pelvises and pelvic-fin

rays

Coracoids

Gill-opening width

Posterior nasal canal

pore*

Dorsal and ventral

lacryroal canal pores*

Postocular sensory

canal pores*

Posteriormost epipleural

rib on vertebra

Diastema in hypural

plate

Longest specimen (TL)

Present, obvious

Present

Present

^ eye diameter

Absent

Present

1

66-78

21-28

Absent

120 mm

Absent, fin-fold

questionably

present

Absent

Absent

^ eye diameter

Usually absent

(present unilaterally in 1

specimen)

2 (3 unilaterally

in 1 specimen)

60-61

21

Present

23 mm

Absent

Absent

«ey e diameter

Usually absent

(present unilaterally in 3 and

bilaterally in 1

specimen)

Usually present

1

63-74

16-20

Present or absent

47 nrn

Absent

Absent

«ey e diameter

Present (absent

unilaterally in

2 specimens)

Present

2

67-71

14-16

Present or absent

44 mn

Number of specimens

examined over 150

*See Figure 10 for pore terminology
NUMBER 234 17

FIGURE 11.—Alabes dorsalis: a, lateral view; b, lateral view of head region; c, ventral view of

head region; d, ventral view of gill opening, pelvic fins, and vestigial sucking disc; e, ventral

view of gill opening and pelvic fins, (a-d, USNM 201415, Kangaroo Island, South Australia,

ca. 77 mm TL; e, USNM 214499, Green's Beach, Tasmania; drawn by Jack R. Schroeder.)

on the basis of some trivial, proportional differences (body less compressed, back and belly more

obtuse, gill opening more distant from snout).

Gunther (1870) considered the two species to be

synonymous and we find no basis for considering

them distinct. We are dubious of the correctness

of the type-localities of both species. We know of

no other records of Alabes from northwestern

Australia, which would be rather warm relative

to those localities where Alabes is known with

reasonable certainty to occur. Alabes dorsalis appears to be uncommon in Western Australia, from

where only one specimen (from the southern coast)

is definitely known to have been collected. The

type-locality of C. aptenodytum from 72°S latitude

is most suspicious. The specimen supposedly was

taken on land and had presumably passed undamaged through the gut of a penguin. Aside from

these peculiar circumstances, we know of no littoral

species or genus of fish that is common to both

Antarctica and Australia.

Macleay distinguished his Chilobranchus rufus

from Cheilobranchus dorsalis on the basis of the

former's having dark spots on the body, which the

latter lacked. The presence or absence of the spots

in A. dorsalis, as well as their number, is variable.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 2.—Frequency distributions for number of vertebrae and posteriormost vertebra bearing

an epipleural rib in Alabes dorsalis

Localities 66 67 68 69 70 71 72 73 74 75 76 77 78

Posteriormost vertebra

bearing an epipleural rib

21 22 23 24 25 26 27 28

Queensland

New South Wales

Ulladulla

Bradley's Head

Pittwater

Victoria

Port Phillip Bay

Bell's Beach

Altona

South Australia

Moonta

Kangaroo Island

Robe

Fiddler's Bay

Tasmania

Green's Beach

Ulverstone

Davenport

Oyster Bay

Maria Island

Eaglehawk Neck

Port Arthur

Hobart

Kt. Esperance

D'Entrecasteaux Channel

Wedge Bay

Great Taylor Bay

South Port

Western Australia

North Point

Type specimens

dorsalis (northwest coast of Australia).

aptenodytum (Penguin Island, 72°S lat.).

rufus (Taiinania)

cuvieri (Australia)

22.6

1

2

1

1

1

2

6

1

1

-

1

1

10

1

1

1

-

3

1

1

6

4

1

1

2

3

1

2

3

1

1

2

2

2

2

1

1

1

10

1

1

1

2

1

1

8

1

1

3

1

5

1

1

1

1

3

1

5

3

1

.

2

1

2

1

.

1

6

1

3

2

1

1

_

70.3

71.3

69.5

1 72.2

71.0

71.0

68.5

69.0

68.5

72.2

72.5

73.5

73.2

74.0

1 75.4

74.0

74.2

1

1

1

2

2

1

1

7

1

1

1

1

3

1

1

3

1

9

1

1

7

2

1

1

1

1

3

.

2

2

1

4

1

2

2

8

9

1

7

5

1

1

.

1

4

2

1

2

7

1

4

5

1

3

1

3

1

3

1

3

1

22.8

22.0

22.5

24.1

23.5

23.8

24.0

23.1

22.5

24.3

23.0

24.4

25.4

23.7

25.3

24.2

23.0

72.5

73.0 2

1

5

1

1

4

1

3

4

24.8

23.7

23.5

Scott (1974) reported 0-9 spots per side in Tasmanian specimens. There is also a tendency for

spots to be absent in long preserved specimens,

indicating that they fade. Both males and females

exhibit the spots.

Vaillant distinguished A. cuvieri on the basis of

its having the dorsal-fin fold origin well posterior

to the head rather than near the nape (Scott, 1974,

distinguished Chilobranchus rufus from Cheilobranchus dorsalis using the same character). The

origin of the fin folds in A. dorsalis are difficult

to determine, but we have seen no specimen in

which we would consider the dorsal fold as originating close to the head (including one specimen

identified by Scott as A. dorsalis). During preservation, shrinkage appears to cause the formation

of pseudo-folds in the skin, and these folds may

appear to be continuous with the natural fin fold.

The dorsal-fin fold origin in A. dorsalis varies in

position from slightly in advance to slightly posterior to a vertical from the anus.

MATERIAL EXAMINED.—AUSTRALIA: MNHN 2181 (3 syntypes of Alabes cuvieri); NW coast, BMNH 1848.3.18.37 (4

syntypes of Cheilobranchus dorsalis). Queensland: QM

1.8951-55 (5). New South Wales: Pittwater, AM IA.5447 (2);

Bradley's Head (Sydney), AM IB.45O7 (3); Ulladulla, AM

1.18519-001 (6). Victoria: Altona, AM IA.5930 (4), IA.5910

(3); Bell's Beach, AM I.16981-OO7 (1), 1.16982-012 (1); Port

Phillip Bay, AM 1.7618 (1), 1.9003 (4), BMNH 1884.11.12.1

(1), 1887.10.10.12-15 (2), 1890.9.16.32-33 (2), WAM P.4579

(I). South Australia: Robe, AM 1.13730 (1); Kangaroo Island,

AM IA.7 (4), IA.8 (4), IA.10 (4), IA.ll (4), IA.12 (4), USNM

87361 (6), 201415 (2, including one cleared and stained);

Moonta, AM IA.7090 (4); Fiddler's Bay, AM 1.17615-005 (2).

Western Australia: North Point, Boulder Hill, AMNH 31306

(1, cleared and stained). Tasmania: AM 1.16265-001 (17 syntypes of Chilobranchus rufus); Cole's Bay, AM 1.17553-019

(1); Green's Beach, USNM 214100 (2, including one cleared
NUMBER 234 19

and stained), 214497 (11), 214498 (2), 214499 (10), 214500 (1);

Ulverstone, AM 1.6260-63 (4); Davenport, AM IB.4564 (1);

Oyster Bay, AM 1.1103 (8), 1.10620-2 (6); Maria Island, AM

IA.8065 (1); Eaglehawk Neck, AM I.17545-O1O (1); Port

Arthur, AM IB.2902 (1); Hobart, AM 1.14207 (6), IB.2481

(4), QM 1.265 (1), 1.8948-50 (3); Mt. Esperance, AM 1.12284

(3); D'Entrecasteaux Channel, AM IA.4110 (4), IA.4111 (3);

Wedge Bay, AM 1.17193-003 (1); Great Taylor Bay, AM

IB.1291 (5); South Port, AM IA.3617 (4). PENGUIN ISLAND

(supposedly 72°S latitude): BMNH 1848.3.18.75 (holotype of

Cheilobranchus aptenodytum).

Alabes brevis, new species

DESCRIPTION.—The diagnostic characters of this

species are given in Tables 1 and 3 and are not

repeated here. The three known specimens are

quite small, not in good condition, and are probably juveniles. No evidence of color pattern is

present on any of the specimens. The caudal fin

is well developed for an Alabes, and there are 9

or 10 caudal-fin rays, 8 articulating with the hypural fan, which has a diastema separating the

dorsal and ventral portions.

COMPARISONS.—Alabes brevis is distinguishable

from all the other species of Alabes by its low

number of vertebrae, and from all but A. hoesei

in having 2 postocular pores. Aside from number

of vertebrae, it differs from A. hoesei in usually

lacking a posterior nasal canal pore, having more

epipleural ribs, and a larger gill opening.

ETYMOLOGY.—The specific epithet is from the

Latin, meaning "short," and refers to the reduced

number of vertebrae that characterizes the species.

HOLOTYPE.—WAM 24385: 20 mm TL; Green

Island, Rottnest, Western Australia; among weed

and rock, limestone reef platform; collected by

L. M. Joll, 25 April 1972.

PARATYPES.—WAM 24386-7: 2 specimens, 16-23

mm TL; same data as holotype.

Alabes parvulus (McCulloch)

FIGURES 12 AND 13

Cheilobranchus parvulus McCulloch, 1909:316 [rock pools

near Sydney; AM 1.9954, lectotype, designated in account

below].

The diagnostic characters of this species are given

in Tables 1 and 3 and are not repeated here.

REMARKS.—The disposition of the sensory canal

pores in A. parvulus is variable and to some extent

related to geography. Specimens from New South

Wales and South Australia have lacrymal canal

TABLE 3.—Frequency distributions for number of vertebrae and posteriormost vertebra bearing

an epipleural rib in Alabes brevis, A. parvulus, and A. hoesei

Species 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74

Posteriormost vertebra

bearing an epipleural rib

7 1

ng an epipl

17 18 19

Alabes brevis

Western Austra1ia

Rottnest

Alabes parvulus

New South Wales

Ulladulla

Manly

Long Reef

South Australia

Fiddler's Bay

Tasmania

Cole's Bay

Western Australia

Seal Island

Norfolk Island

Emily Bay

Alabes hoesei

New South Wales

Manly

South Australia

Robe

Fiddler's Bay

Tasmania

Cole's Bay

1

1

1

4*

1

1

4

1

1

2

2

1

72.0

70.2

71.6

66.7

1

1

2

1

2

1

1

2

11*

2

1

7 1

18.3

18.7

17.5

16.3

1113 *

1

70.0 1 3* 2

1

15.2

15.7

•Denotes count for holotvoe or lectotvpe
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 12.—Alabes parvulus (a-c, female, 42 mm TL; d-f, male, 38 mm TL): a, lateral view;

b, dorsal view; c, ventral view; d, lateral view; e, ventral view, /, dorsal view. (WAM P.25365-001,

Seal Island, Western Australia; drawn by Jack R. Schroeder.)

pores and occasionally have posterior nasal canal

pores. Specimens from Tasmania, Western Australia, and Norfolk Island lack lacrymal canal

pores and none have posterior nasal canal pores

(some specimens from Tasmania and Western Australia exhibit blind porelike depressions unilaterally in the positions where the dorsal or ventral

lacrymal canal pores ordinarily occur).

There are indications that numbers of vertebrae

and epipleural ribs vary geographically (Table 3).

The one available Norfolk Island specimen

(male) has a slightly different color pattern (Figure

13) from that of either the male or female Western

Australian specimens (Figure 12), which agree

closely with McCulloch's (1909) description of the

color pattern of New South Wales specimens. (All

we saw were faded; McCulloch's description appears to be based on both sexes.) The dorsal-fin

fold origin of the Norfolk Island specimen is much

farther posteriorly than that of any other Alabes

specimen we have seen, and it is possible that the

Norfolk Island species will merit description if

additional specimens exhibit the same differences

shown by our single specimen. The caudal region

of the Norfolk Island specimen is malformed.

LECTOTYPE DESIGNATION.—McCulloch did not

designate a holotype or specify the number of

specimens he had examined when he described

C. parvulus, but he did indicate that he had

"many." He gave, however, the total length of

only one specimen (47 mm) in the descriptive account, although he mentioned under "observa-
NUMBER 234 21

FIGURE 13.—Alabes parvulus (male, 32 mm TL): a, lateral view; b, ventral view. (AM 1.18470-002,

Emily Bay, Norfolk Island; drawn by Jack R. Schroeder.)

tions" that females had large eggs at 37 mm. The

only specimen among the syntypes that is close to

47 mm is contained in a lot of four specimens

collected at Freshwater Bay, Manly, by Basset Hull,

during May 1909, and bearing the catalog number

AM 1.9954-5. This specimen, which is 47 mm TL,

is here designated the lectotype of C. parvulus and

assigned the catalog number 1.9954. The three

other syntypes in the lot now bear the catalog

number 1.9955.

There are a considerable number of other syntypes (of which we have seen only 20) at AM bearing the catalog number 1.9266, also collected by

Hull at Manly, but dated January 1908. One

syntype formerly in 1.9266 has been recataloged

as 1.18471-001, as it represents an undescribed species; it is designated a paratype of Alabes hoesei

under the description of that species.

MATERIALS EXAMINED.—New South Wales: Ulladulla, AM

IB.447 (3); Long Reef, AM IA.1253 (2, including one cleared

and stained), IA.5892 (2), 1.17019-026 (1); Manly, Freshwater

Bay, AM 1.9266, in part (20 syntypes, others at AM not

examined), 1.9954 (lectotype), 1.9955 (3 syntypes). South Australia: Fiddler's Bay, AM 1.17615-004 (1). Tasmania: Cole's

Bay, AM 1.17553-005 (1). Western Australia: Safety Harbor,

WAM P.25465-001 (4). Norfolk Island: Emily Bay, AM

1.18470-002 (1).

Alabes hoesei, new species

FIGURE 14

DESCRIPTION.—The diagnostic characters of this

species are given in Tables 1 and 3 and are not

repeated here. Only two specimens showed evidence of a color pattern (Figure 14); the other

specimens were faded. The color pattern closely

resembles that of the female A. parvulus in

Figure 12a.

REMARKS.—Alabes hoesei, A. parvulus, and A.

dorsalis have all been obtained in the same collection from Cole's Bay, Tasmania.

COMPARISONS.—Alabes hoesei resembles A. brevis

in having two postocular pores (the other two

species have only one), but differs from A. brevis

in having a posterior nasal canal pore (unilaterally

present in one of the three specimens of A. brevis),

more vertebrae, fewer epipleural ribs, and a smaller gill opening. A. hoesei is probably most closely

related to A. parvulus; both have highly restricted

gill openings. A. hoesei differs from A. parvulus in

having two postocular pores, posterior nasal canal

pores (rarely present in A. parvulus), and modally

fewer epipleural ribs.

ETYMOLOGY.—Named for Dr. D. F. Hoese (AM)

who provided much of the material upon which

our study is based.

HOLOTYPE.—AM 1.18470-001: 34 mm TL; Robe,

South Australia; rock platform; collected by D. F.

Hoese, K. and R. Meguro, and R. Kuronuma,

3 October 1975 (original number DFH 75-212).

PARATYPES.—AM 1.18470-002: 5 specimens, 26-28 mm TL;

same data as holotype. AM 1.17615-003: 30 mm TL, Fiddler's

Bay, South Australia. AM I.18471-OO!: 25 mm TL, Freshwater Bay, Manly, New South Wales (formerly a syntype

of Cheilobranchus parvulus). AM 1.17553-017: 44 mm TL,

Coles Bay, Tasmania. AM 1.17553-018: 34 and 38 mm TL,

Cole's Bay, Tasmania.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 14.—Alabes hoesei (a-c, holotype, 34 mm TL; d-f, para type, 28 mm TL): a, lateral view;

/>, dorsal view; c, ventral view; d, lateral view; e, dorsal view; /, ventral view. (Robe, South

Australia; a-c, AM 1.18470-001; d-f, AM 1.18470-002; drawn by Jack R. Schroeder.)
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U.S. GOVERNMENT PRINTING OFFICE: 1976 211-559/27 23

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