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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 234

Synonymy of the Fish Families Cheilobranchidae (=Alabetidae) and Gobiesocidae, with Descriptions of Two New Species of Alabes

Victor G. Springer
and Thomas H. Fraser

SMITHSONIAN INSTITUTION PRESS

City of Washington

1976

A B S T R A C 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), parvulus (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.G6] 591′.08s [597′.58] 76–608046

Synonymy of the Fish Families Cheilobranchidae (=Alabetidae) and Gobiesocidae, with Descriptions of Two New Species of Alabes

Victor G. Springer
and Thomas H. Fraser

____________

Victor G. Springer, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Thomas H. Fraser, Environmental Quality Laboratory, 590-D Olean Boulevard, Port Charlotte, Florida 33952.

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 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.

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).

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 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 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.

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

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

FIGURE 3.—Superficial bones, jaws, and suspensorium: a, Alabes dorsalis (cranium also indicated); b, Trachelochismus pinnulatus. (Articular and angular bones are synonymous with anguloarticular and retroarticular bones of recent authors.)

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 AND HYOID APPARATUS (Figure 4) (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.

FIGURE 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, Alabes dorsalis; b, Trachelochismus pinnulatus.

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 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.

FIGURE 6.—Right hyoid arch (cartilage indicated by pattern of small, open circles): a, Alabes dorsalis; b, Trachelochismus pinnulatus (branchiostegals truncated).

The gobiesocids have 11–20 precaudal, 13–33 caudal, and 25–54 total vertebrae. There are 0–3 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.

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).

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 Bühlke and Robins (1970), Gymnoscyphus ascitus lack, dorsal postcleithra; but all the other genera (entire Gobiesocidae considered) apparently have both dorsal and ventral postcleithra.

FIGURE 8.—Trachelochismus pinnulatus: a, four anteriormost precaudal vertebrae; b, posteriormost two precaudal and anteriormost three caudal vertebrae; c, posteriormost two caudal vertebrae and caudal-fin rays (segmented rays truncated; cartilage indicated by pattern of small, open circles).

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 Böhlke 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 11d; 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 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 Rimicola, 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; metapterygoid; 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; lateral-line 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*.

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.)

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; postcleithra (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 gobiesocids-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; =AIabetidae 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 Böhlke 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 scanty, we propose that Gastrocymba is the genus most closely related to Alabes (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

Cheilobranrhus dorsalis Richardson, 1845 [NW 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 I.16264–001, 14 syntypes, and Tasmania, AM I.16265–001, 17 syntypes].

Alabes cuvieri Vaillant, 1905a:149; 1905b:1714 [Nouvelle-Hollande = 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 on the basis of some trivial, proportional differences (body less compressed, back and belly more obtuse, gill opening more distant from snout). Günther (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.

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

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.)

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. 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.

TABLE 2.—Frequency distributions for number of vertebrae and posterior most vertebra bearing an epipleural rib in Alabes dorsalis

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 I.8951–55 (5). New South Wales: Pittwater, AM IA.5447 (2); Bradley’s Head (Sydney), AM IB.4507 (3); Ulladulla, AM I.18519–001 (6). Victoria: Altona, AM IA.5930 (4), IA.5910 (3); Bell’s Beach, AM I.16981–007 (1), I.16982–012 (1); Port Phillip Bay, AM 1.7618 (1), I.9003 (4), BMNH 1884.11.12.1 (1), 1887.10.10.12–15 (2), 1890.9.16.32–33 (2), WAM P.4579 (1). South Australia: Robe, AM I.13730 (1); Kangaroo Island, AM IA.7 (4). IA.8 (4), IA.10 (4), IA.11 (4), IA.12 (4), USNM 87361 (6), 201415 (2, including one cleared and stained); Moonta, AM IA.7090 (4); Fiddler’s Bay, AM I.17615–005 (2). Western Australia: North Point, Boulder Hill, AMNH 31306 (1, cleared and stained). Tasmania: AM I.16265–001 (17 syntypes of Chilobranchus rufus); Cole’s Bay, AM I.17553–019 (1); Greens Beach, USNM 214100 (2, including one cleared and stained), 214497 (11), 214498 (2), 214499 (10), 214500 (1); Ulverstone, AM I.6260–63 (4); Davenport, AM IB.4564 (1); Oyster Bay, AM I.1103 (8), I.10620–2 (6); Maria Island, AM IA.8065 (1); Eaglehawk Neck, AM I.17545–010 (1); Port Arthur, AM IB.2902 (I); 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 I.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 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).

TABLE 3.—Frequency distributions for number of vertebrae and posteriormost vertebra bearing an epipleural rib in Alabes brevis, A. parvulus, and A. hoesei

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, f, dorsal view. (WAM P.25365–001, Seal Island, Western Australia; drawn by Jack R. Schroeder.)

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 “observations” 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 I.9955.

FIGURE 13.—Alabes parvulus (male, 32 mm TL): a, lateral view; b, ventral view. (AM I.18470–002, Emily Bay, Norfolk Island; drawn by Jack R. Schroeder.)

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 I.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), I.17019–026 (1); Manly, Freshwater Bay, AM I.9266, in part (20 syntypes, others at AM not examined), I.9954 (lectotype), I.9955 (3 syntypes). South Australia: Fiddler’s Bay, AM I.17615–004 (I). Tasmania: Cole’s Bay, AM I.17553–005 (1). Western Australia. Safety Harbor, WAM P.25465-001 (4). Norfolk Island: Emily Bay, AM I.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 I.17615–003: 30 mm TL, Fiddler’s Bay. South Australia. AM 1.18471–001: 25 mm TL, Freshwater Bay. Manly, New South Wales (formerly a syntype of Cheilobranchus parvulus). AM I.17553-017: 44 mm TL, Cole’s Bay. Tasmania. AM 1.17553–018: 34 and 38 mm TL, Cole’s Bay, Tasmania.

FIGURE 14.—Alabes hoesei (a–c, holotype, 34 mm TL; d–f, paratype, 28 mm TL): a, lateral view; b, dorsal view; c, ventral view; d, lateral view; e, dorsal view; f, ventral view. (Robe, South Australia; a–c, AM I.18470-001; d–f, AM I.18470-002; drawn by Jack R. Schroeder.)

Literature Cited

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Böhlke, J. E. and C. R. Robins

1970. A New Genus and Species of Deep-dwelling Clingfish from the Lesser Antilles. Notulae Naturae, 434:1–12.

Briggs, J. C.

1955. A Monograph of the Clingfishes (Order Xenopterygii). Stanford Ichthyological Bulletin, 6:1–224.

Cloquet, H.

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Cuvier, G. L. F. C. D.

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Gill, T.

1872. Arrangement of the Families of Fishes or Classes Pisces, Marsipobranchii, and Leptocardii. Smithsonian Miscellaneous Collections, 247: xlvi + 49 pages.

1906. The Fish Genus Alabes or Cheilobranchus. Science, new series, 23(589):584–585.

Gosline, W. A.

1968. The Suborders of Perciform Fishes. Proceedings of the United States National Museum, 124(3647): 1–78.

Greenwood, P. H.

1975. J. R. Norman’s A History of Fishes. Third edition, xxv + 467 pages. New York: John Wiley and Sons.

Guitel, F.

1888. Recherches sur les lepadogasters. Archives de Zoologie Experiméntal et Générale, series 2, 6:423–647, plates 24–37.

Günther, A.

1870. Catalogue of the Fishes in the British Museum. Volume 8, xxv + 549 pages. London.

Leray, C.

1961. Contribution a l’etude osteologique de Gouania wildenowi Risso (teleosteens) (squelette œphalique et ceintures). Cahiers de Biologie Marine, 2(1):41–52.

Macleay, W.

1882. Descriptive Catalogue of the Fishes of Australia, part 3. Proceedings of the Linnean Society of New South Wales, 6(2):202–386.

McCulloch, A. R.

1909. Studies in Australian Fishes, number 2. Records of the Australian Museum, 8(4):315-321, plates 90-91.

Regan, C. T.

1912. The Anatomy and Classification of the Symbranchoid Eels. Annals and Magazine of Natural History, series 8, 9:387–390, plate 9.

Richardson, J.

1845. Ichthyology of the Voyage of H. M. S. Erebus and Terror. Pages 17–52 in number [2] in volume II of J. Richardson and J. E. Gray, editors, The Zoology of the Voyage of H. M. S. Erebus and Terror... 1839–1813. London.

Rosen, D. E., and P. H. Greenwood

In press. [A study of the Synbranchiformes—title indefinite].

Rosen, D. E., and C. Patterson

1969. The Structure and Relationships of the Paracanthopterygian Fishes. Bulletin of the American Museum of Natural History, 141(3):357–474, plates 52–78.

Runyan, S.

1961. Early Development of the Clingfish. Gobiesox strumosus Cope. Chesapeake Science, 2(3–4):113–141.

Scott, E. O. G.

1974. Observations on Some Tasmanian Fishes, part XX. Papers and Proceedings of the Royal Society of Tasmania, 108:171–197.

Swainson, W.

1838. The Natural History of Fishes, Amphibians, and Reptiles, or Monocardian Animals. Volume 1: vi + 368 pages; volume 2: vi + 452 pages. London.

Starks, E. C.

1905. The Osteology of Caularchus maeandricus (Girard). Biological Bulletin, 9(5):292–303.

Taylor, W. R.

1967. An Enzyme Method of Clearing and Staining Small Vertebrates. Proceedings of the United States National Museum, 122(3596):1–17.

Vaillant, L.

1905a. Description de poissons nouveaux ou imparfaitement connus de la collection du Muséum D’Histoire Naturelle: Le Genre Alabes de Cuvier. Nouvelles Archives du Museum D’Histoire Naturelle, series 4, 7(1):145–152; and (2):153–158.

1905b. Le genre Alabes de Cuvier. Comptes Rendus Hebdomadaires des Séances de l’Academic des Sciences, 140:1713–1715.

Waite, E. R.

1923. The Fishes of South Australia. 243 pages. Adelaide: Government Printer.

____________

¹The 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 Cloquct’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 Alabes cuvieri Vaillant, 1905a, 1905b, by subsequent monotypy.

²In 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 epural. 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 ectopterygoid as part of 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 basihranchials were present because they were only interested in structures that were ossified. The basibranchials are cartilaginous in our specimens of G. nigripinnis.

³Scott (1974) reported that teeth were present on the vomer, but we have not found this to be true of any of our specimens.

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