History of Gobioid Classification

 By: James L. Van Tassell

1998

In 1758, Linnaeus recognized one genus (Gobius) with seven species.  This was increased to seven genera in 1800 by Lacepède, eleven genera in 1801 by Bloch and Schneider, twenty-three genera by Günther in 1861b, and ninety-nine genera by Bleeker in 1874.  Günther (1861b), Bleeker (1874), and Jordan (1885-1923) proposed the earliest classification systems for gobioid interrelationships.  These systems were based primarily on the structure of the fins, but also utilized numbers and type of scales (ctenoid vs. cycloid), teeth, and placement of eyes.

Günther (1861b) recognized four groups of gobioids: the Gobiinae, containing species currently recognized as gobiids, eleotrids, and periophthalmids; the Amblyopina, containing the genus Amblyopus; the Trypauchenina, containing Trypauchen and Trypauchenichthys; and the Callionymina with three genera: Platyptera, Callionymus, and Vulsus.  In 1874 Bleeker diagnosed four subfamilies: Eleotriformes, Gobiiformes, Amblyopodiformes (Amblyopina and Trypauchena of Günther), and Luciogobiiformes (gobioids without a first dorsal fin).

The first classification based largely on osteology, e.g. bones of the cranium and pectoral girdle and number of vertebrae, was by Regan (1911).  He recognized three families: the Eleotridae, Gobiidae, and Psammichthyidae.  The Eleotridae were separated from the Gobiidae on the basis of the shape of the palatine and the greater development of the mesopterygoid and scapula in the eleotrids.  Based on these characteristics, the genus Rhyacichthys was also placed within the eleotrids.  The family Gobiidae was divided into two subfamilies, Gobiinae and Periophthalminae, with the taenioids included within the Gobiinae. Regan established the family Psammichthyidae (=Kraemeriidae according to Maugé 1986 in Eschmeyer, 1990) and provisionally placed it within the Gobioidei.

Sanzo (1911) was the first to study the gobioid lateralis system.  This system of external neuromast organs (so-called ‘sensory papillae’) forms distinct patterns on the head region.  Neuromast patterns can be divided into two basic groupings, one where the neuromasts are primarily in longitudinal rows (fig. 1) and the other with neuromasts in transverse rows (fig. 2). These patterns have been shown to be useful in defining genera and species (Hoese 1971, 1983; Gill et al., 1992) and have been used extensively by Miller (1963, 1973, 1992b, to cite a few) in defining taxa and arranging systems of classification.

Jordan (1923) produced an extensive treatment of the group.  He divided the order Gobioidei into eight families (Rhyacichthyidae, Eleotridae, Gobiidae, Periophthalmidae, Gobioididae, Trypauchenidae, Doliichthyidae, and Psammichthyidae) but gave no diagnoses for these families.  Duncker (1928) recognized four families (Eleotridae, Gobioididae, Periophthalmidae, and Gobiidae), based on the work of others.  Berg (1940) separated the group into two superfamilies (Eleotrioidae and Gobioidae) comprising three families in total for the groups.  The superfamilies separated the eleotrids and Rhyacichthys from all other gobioids.  His three families were the Eleotridae, the Gobiidae (including taeniodids), and the Periophthalmidae (including kraemeriids).

Ginsburg (1933a) and Koumans (1953) both pointed out the inadequacies of the classification system of gobioids, but it was not until 1955 that Gosline made the next major osteological study.  Gosline gave evidence for the placement of the microdesmids and kraemeriids among the gobioids and demonstrated that the shape of the palatine and the presence or absence of the scapula did not adequately separate the Eleotridae and the Gobiidae.  He based his separation of these two families on the presence of six branchiostegal rays in eleotrids and five in gobiids.  In 1960, Takagi made an extensive study of the cephalic sensory canal systems of gobies from Japan.  A portion of the study was published in 1967 and the complete work published in 1988.  Takagi examined 82 species of 54 genera from Japanese waters and amended the terminology of the cephalic sensory canals, head pores, and sensory papillae patterns.  He divided the gobioids into two groups, those possessing both cephalic sensory canals and sensory papillae, which he considered as the plesiomorphic condition, and those possessing only sensory papillae, the derived condition.  Akihito (1963, 1967) conducted an extensive study on the scapula of gobioids, noting that the scapula is well developed in the primitive genera.  Later, in 1969, Akihito produced a detailed study of the higher classification of gobioid fishes based on the osteology of 85 species in 71 genera.

Miller (1963) briefly outlined the major differences between gobiids and eleotrids, commenting that the position of separate pelvic fins was not sufficient to separate eleotrids from gobiids.  He produced a considerably different classification based largely on osteological characters.  The characters he included are the number of epurals, hypural connection, the presence or absence of the endopterygoid, the number of branchiostegal rays, development of the scapula, the number of pectoral radials, presence or absence of the postcleithrum, a metapterygoid bridge to the quadrate, a preoperculum-symplectic bridge, and the extent of development of the oculoscapular and preopercular canals.  Miller (1973) divided the suborder Gobioidei into two families: Rhyacichthyidae, with only one species, and Gobiidae, with close to 2000 species.  Rhyacichthyidae was recognized by two plesiomorphies, the presence of three epurals and a well-developed cephalic lateralis system.  Miller’s Gobiidae shared the derived character states of one or two epurals and a reduced cephalic lateralis system.  Miller (1973) further divided the Gobiidae into seven subfamilies: Eleotrinae, Pirskeninae, Xenisthminae, Gobionellinae, Tridentigerinae, Gobiinae, and Kraemeriinae and included the Pholidichthyidae within the Gobiinae.  Springer (1983) strengthened Miller’s classification of the Gobioidei by incorporating additional synapomorphies.  He removed the Pholidichthyidae because it did not share  any of the synapomorphies that he considered diagnostic for the gobioids.

Conflicting phylogenetic schemes based on synapomorphies in skeletal and sensory papillae systems were discussed by Miller et al. in 1980.  Using electrophoretic techniques to differentiate hemoglobin polymorphisms, they attempted to resolve the conflicting hypotheses, but no close relationship to either of the phylogenetic schemes was apparent from the study.  Springer (1983) studied the Gobioidei cladistically and proposed the four following synapomorphies for all gobioids: parietals absent, pelvic intercleithral cartilage present, dorsal end of interhyal fails to meet the dorsal end of the symplectic, and basibranchial 1 cartilaginous.  To this list Miller (1992a) added the presence of a sperm duct gland.  Subsequently, others have provided a partial resolution of groups within the Eleotridae (Hoese & Gill, 1993), or given evidence for monophyly within the Gobiidae: sicydiines (Hoese, 1984; Harrison, 1989, 1993; Parenti and Maciolek, 1993); oxudercines (Murdy, 1989); Amblyopinae and Gobiinae (Pezold, 1993).

Some resolution to the phylogenetic problems within the lower members of the Gobioidei was presented by Hoese and Gill (1993).  They were able to define three families (Rhyacichthyidae, Odontobutidae, Gobiidae) and divide the Gobiidae into the subfamilies Butinae, Eleotridinae, and Gobiinae based on sixteen characters.  The Gobiidae were diagnosed according to the following synapomorphies: (1) no autogenous middle radial in the first pterygiophore of the second dorsal fin; (2) upper proximal radial of the pectoral fin usually in contact with the cleithrum and extending well above the scapula; (3) anterior elongation of the procurrent caudal cartilage;  and (4) scales without transforming ctenii. 

Pezold (1993) divided Hoese’s (1984) Gobiinae into a monophyletic Gobiinae and a smaller group, the subfamily Gobionellinae, for which monophyly could not be established.  Pezold’s Gobiinae is diagnosed by the presence of a single anterior interorbital pore, a single posterior pair of nasal pores, the interorbital portion of the oculoscapular canal fused, one epural, most species with 3-22110 first dorsal fin pterygiophore formula, 26 or 27 vertebrae, and two prehemal pterygiophores in most species.

The family Rhyacichthyidae is generally accepted as the sister group to all other gobioids.  Monophyly of the remaining gobioids is supported by three synapomorphies (Springer, 1983): (1) lateral line canal not extending onto the body; (2) ventral process of the hyomandibula broad with the dorsal tips of the interhyal and symplectic widely separated from each other; and (3) mandibula sensory canal absent.  To this list Hoese and Gill (1993) added (4) change in the position of the penultimate branchiostegal ray; and (5) reduction of head canals with the separation of the preopercular canal from the oculoscapular canal and possibly the reduction in the number of pores.

 

The Tribe Gobiosomini

There are about 100 genera in the Gobiinae worldwide with 29 genera in the New World, 26 of which are restricted to the New World (Birdsong and Robins, 1995).  The New World goby fauna is dominated by the tribe Gobiosomini (Birdsong, 1975) in the Gobiinae (sensu Pezold, 1993).  The Gobiosomini contains 24 of the 26 New World genera and thus, according to Birdsong and Robins (1995), represent 40% of the New World gobioid genera.

The Gobiosomini, as originally proposed by Birdsong (1975) and later revised by Birdsong et al. (1988), unite what was commonly called the American seven-spined gobies and several closely allied genera, all endemic to the New World.  Additional genera were added to the tribe by Hoese (1976) and Birdsong & Robins (1995).  Genera currently included in the tribe are Akko Birdsong and Robins, 1995; Aruma Ginsburg, 1933; Barbulifer Eigenmann & Eigenmann, 1888; Bollmannia Jordan in Jordan and Bollmann, 1890; Chriolepis Gilbert, 1892; Eleotrica Ginsburg, 1933a; Enypnias Jordan & Evermann, 1898; Evermannichthys Metzelaar, 1920; Ginsburgellus Böhlke & Robins, 1968; Gobiosoma Girard, 1858 (including Aboma Jordan and Starks, 1895 and Elacatinus Jordan, 1904); Gobulus Ginsburg, 1933; Gymneleotris Bleeker, 1874; Microgobius Poey, 1876; Nes Ginsburg, 1933;, Ophiogobius Gill, 1863; Palatogobius Gilbert, 1971; Pariah Böhlke, 1969; Parrella Ginsburg, 1938; Psilotris Ginsburg, 1953; Pycnomma Rutter, 1904; Risor Ginsburg, 1933; Robinsichthys Birdsong, 1988; and Varicus Robins & Böhlke, 1961.  Böhlke and Robins (1968) defined the genus Gobiosoma to include the subgenera Austrogobius de Buen, 1950,; Elacatinus Jordan, 1904; Garmannia Jordan and Evermann in Jordan, 1895; Gobiosoma Girard, 1858; and Tigrigobius Fowler, 1931.  While the tribe may not be monophyletic, later work by Birdsong et al. (1988) inferred a subset of the group, known as the ‘Gobiosoma Group’ (all genera except Bollmannia, Microgobius, Palatogobius, Akko, and Parrella) to be monophyletic.

Many of the genera within the tribe are monotypic. They include Aboma, Akko, Aruma, Eleotrica, Ginsburgellus, Gymneleotris,  Nes, Ophiogobius, Palatogobius, Pariah, Risor, and Robinsichthys.  Several contain only a few species (Pycnomma, 2 spp.; Enypnias, 2 spp.; Psilotris, 3 spp.; Gobulus, 4 spp.; Evermannichthys, 4 spp.; Parrella, 5 spp.) and are rather distinctive.  The most speciose genera, other than Gobiosoma (37 spp.), are Microgobius (14 spp.) and Bollmannia (13 spp.).

Characters used to unite the Gobiosomini are a dorsal fin pterygiophore formula of 3-221110 ( in all genera except Evermannichthys, Pariah, and Risor), a  vertebral count of 11+16-17 (in all but Evermannichthys, and Pariah), and the fusion of hypurals 1+2 with 3+4 (in all genera except Aboma, Akko, Bollmannia, Microgobius, Palatogobius, and Parrella) (Birdsong, 1975).

The largest genus, Gobiosoma, has been divided into as few as four or as many as seven subgenera (Ginsburg, 1933b, 1944; Böhlke and Robins, 1968; Hoese, 1971) based on squamation, cephalic pore patterns, or sensory papillae patterns.  The characters used to define the genus and to separate the subgenera phenetically include (1) the number of pores and the extent of development of the head lateral line canal system, (2) elongation of dorsal fin spines, (3) precaudal and caudal vertebral numbers, (4) the shape of the ‘tongue’,  (5) the extent of squamation on the body and (6) the presence or absence of basicaudal scales.  While the species within Gobiosoma are, in general, well-defined, their phylogenetic relationships have not been investigated cladistically.  Previous authors agree little on the arrangement of the species within the subgenera or which subgenera to include within the genus Gobiosoma.

The species of Gobiosoma sensu (Böhlke and Robins, 1968) were first studied by Isaac Ginsburg in a series of papers from 1933 to 1953.  He recognized three genera  (Aboma, Gobiosoma, Garmannia) based on the extent of squamation: Aboma (monotypic), completely scaled; Garmannia, scaled at least on the posterior half of the trunk and possessing four transverse scales on the caudal peduncle; and Gobiosoma, either completely naked or possessing only two modified basicaudal scales.  Gobiosoma and Garmannia were divided into numerous subgenera: Gobiosoma into the subgenera Elacatinus, Nes, Gobiosoma, Aruma, Dilepidion Ginsburg, 1933, Gerhardinus Meek & Hildebrand, 1928, and Garmannia into the subgenera Tigrigobius, Gobicula, Ginsburg 1944, Gobiolepis Ginsburg, 1944, Garmannia, Gobiohelpis Ginsburg, 1944, Gobiculina Ginsburg, 1944, and Risor, again based on the extent of squamation.  Ginsburg erected the subgenera  as  temporary holding areas for the species until he could obtain sufficient data on each species to change the arrangement.  In his final published papers, he alluded to sensory papillae patterns as a character that could be used to rearrange the species.  His later work on this subject, however, was never published.

Böhlke and Robins reviewed the Atlantic species of Gobiosoma and closely related genera in 1968 and included nominal Pacific species of Gobiosoma in discussing general relationships.  The genera included in their study were Gobiosoma, Risor, Ginsburgellus, Nes, Aruma, Enypnias, Barbulifer, Eleotrica, Gymneleotris, and Pycnomma.  Ten new species and one new genus were described. They elevated Ginsburg's subgenera Nes and Aruma to genera and made Garmannia a subgenus of Gobiosoma.  Their classification was based on the presence or absence of oculoscapular and preopercular canal pores and numbers of vertebrae rather than on scale patterns as adopted by Ginsburg.  According to Böhlke and Robins, Gobiosoma was composed of five subgenera (Elacatinus, Gobiosoma, Austrogobius, Tigrigobius, Garmannia).

Hoese (1971) revised the eastern Pacific species of Gobiosoma in his doctoral dissertation.  The nine Pacific species, three of which were new, are separated by color pattern, extent of squamation, presence or absence of head barbels, cephalic lateral line pore patterns, fin ray counts, sensory papillae patterns, vertebral counts, elongation of the sphenotic, and changes in the length to width ratio of the skull.  He included the genus Aboma as a subgenus of Gobiosoma and commented on the type species, Aboma etheostoma, as perhaps the most primitive member of Gobiosoma.  Several new species were described and the subgenus Gobiolepis (Ginsburg 1944) was resurrected.  However, the new species described in his thesis still remain unpublished.

 

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