The
Tribe Gobiosomatini
by
James Van Tassell
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)[note
the correct name of the tribe is Gobiosomatini, see Smith and
Baldwin 1999] 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 Aboma Jordan and Starks, 1895; Akko
Birdsong and Robins, 1995; Aruma Ginsburg, 1933;
Barbulifer Eigenmann & Eigenmann, 1888; Bollmannia
Jordan in Jordan and Bollmann, 1890; Chriolepis
Gilbert, 1892; Elacatinus Jordan, 1904;Eleotrica
Ginsburg, 1933a; Enypnias Jordan & Evermann,
1898; Evermannichthys Metzelaar, 1920; Ginsburgellus
Böhlke & Robins, 1968; Gobiosoma Girard,
1858; 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.