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Phylogeny and Evolution of the Gobiid Genus <I>Coryphopterus</I>

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Phylogeny and Evolution of the Gobiid Genus <I>Coryphopterus</I> BULLETIN OF MARINE SCIENCE, 70(3): 837–850, 2002 PHYLOGENY AND EVOLUTION OF THE GOBIID GENUS CORYPHOPTERUS Christine E. Thacker and Kathleen S. Cole ABSTRACT We use morphological and molecular data to resolve the relationships among species of the gobiid genus Coryphopterus, and between Coryphopterus and putative sister taxa Lophogobius and Fusigobius. Characters of the external morphology and the mitochon- drial ND2 gene were combined in a total evidence cladistic analysis. The single most parsimonious topology that resulted indicates that Coryphopterus is not monophyletic, and we advocate the removal of C. nicholsii from Coryphopterus and resurrection of the name Rhinogobiops nicholsii. Our phylogeny agrees in many respects with an earlier hypothesis advanced by Smith and Tyler (1977); interpretations of the evolution of mor- phology and ecology on the two topologies are compared. The phylogeny also indicates that Coryphopterus is not closely related to Fusigobius, and so the synonomy of Randall (1995) should not be used. Instead, Coryphopterus is more closely related to the tran- sisthmian Lophogobius than to any Indo-Pacific species examined, a finding that pro- vides insight into the biogeography of the group. An ancillary result of our study is the suggestion that Fusigobius is not monophyletic, however, a comprehensive revision and phylogenetic analysis of Fusigobius is beyond the scope of this paper. The goby genus Coryphopterus Gill includes nine western Atlantic and two eastern Pacific species. Among gobies, Coryphopterus species are unusual in that they occupy a range of habitats and have a sequential (protogynous) hermaphroditic life history. Spe- cies of Coryphopterus are generally small (11 to 50 mm SL) and vary in their general morphology. Most are pale, with few black, yellow or blue markings; the hover gobies (C. personatus (Jordan and Thompson) and C. hyalinus Böhlke and Robins) and the peppermint goby (C. lipernes Böhlke and Robins) are the smallest Coryphopterus, and are orange to gold with white markings. As is typical for gobies, most Coryphopterus species are benthic, inhabiting rocky or sandy substrates near reefs. The hover gobies and peppermint goby are exceptions to these patterns: hover gobies are found in large schooling aggregations above coral reefs, and the peppermint goby perches on corals; these three species are also found at deeper depths (up to 30 m) than the other Coryphopterus species. Previous work on Coryphopterus has mostly been taxonomic rather than phylogenetic. In the revision of Böhlke and Robins (1960), the two Pacific (Coryphopterus nicholsii (Bean) and C. urospilus Ginsberg) and six western Atlantic (C. alloides, C. dicrus, C. thrix, C. eidolon all new species, C. glaucofrenum Gill and C. punctipectophorus Springer) species are discussed and a key is provided. Böhlke and Robins (1960) provide 21 char- acters that are present in Coryphopterus, including the presence of six dorsal spines, with the sixth widely separated from the fifth; seventeen segmented caudal rays; no teeth on vomer or palatine; jaw teeth in several rows, the inner and outer rows enlarged; second dorsal and anal fins relatively short (13–15 second dorsal and 12–13 anal elements in C. nicholsii; 9–11 second dorsal and 12–13 anal elements in the other species); pelvic fin counts of I,5/I,5 with disc complete or deeply incised, frenum present or absent; enlarged neural and hemal spines on the vertebra preceding the hypural plate; body scaled, but 837 838 BULLETIN OF MARINE SCIENCE, VOL. 70, NO. 3, 2002 cheek and opercle unscaled; and a fleshy ridge, variously developed, from dorsal fin origin to a point between and just behind the eyes. None of these characters is unique to Coryphopterus species, and most are fairly widespread among gobies. A further three western Atlantic species, Coryphopterus personatus (formerly Eviota personata), C. hyalinus and C. lipernes, were described in 1962 (Böhlke and Robins, 1962). Coryphopterus tortugae was discussed as a pallid form of C. glaucofrenum by Böhlke and Robins (1960); based on collections in the Colombian Caribbean, Garzon-Ferreira and Acero (1990) redescribe C. tortugae as a distinct species, and Greenfield and Johnson (1999) also recognize C. tortugae. The primary character used to differentiate the species is the pigment on the caudal peduncle: either two separate blotches (C. glaucofrenum) or a single bar (C. tortugae). Garzon-Ferreira and Acero (1990) also report that their C. tortugae specimens are overall more pallid and less robust than C. glaucofrenum. The coloration differences were attributed by Böhlke and Robins (1960) to differences in habitat where the fish were caught: clear, deeper water for C. tortugae and murkier, in- shore water for C. glaucofrenum. Böhlke and Robins (1960) also report that a full range of intermediates was observed. Garzon-Ferreira and Acero (1990) claim that no intergra- dation exists in the caudal pigmentation. The C. glaucofrenum examined for this study were collected from localities ranging from Panama, throughout the Greater and Lesser Antilles, to Venezuela, and all but one lot from Costa Rica (LACM 2250) exhibited two distinct blotches, without intervening pigment, on the caudal peduncle. The four fish in the Costa Rican lot were darker than the other fishes examined, and there was some dark pigment between the caudal peduncle blotches; however, the two blotches were clearly distinguishable and no other notable differences were observed. Because of the variation found in C. glaucofrenum and the presence of intermediates observed by Böhlke and Robins (1960), C. tortugae is not treated as a separate species in this study. Smith (1997) recognizes another species similar to C. glaucofrenum, C. venezulae. Originally described as a subspecies of C. glaucofrenum (Cervigón, 1966), it differs from C. glaucofrenum slightly in pigmentation and has one additional dorsal ray; we did not examine this spe- cies to determine its validity, and do not consider it further. Coryphopterus is currently placed in the Priolepis group of the gobiid subfamily Gobiinae (Birdsong et al., 1988), a large (54 genera) phenetically distinct subdivision of Gobioidei that includes most of the gobiids inhabiting the Indo-Pacific and Pacific Plate, as well as a significant portion of the western Atlantic gobiid fauna. Members of the Priolepis group share a dorsal-fin pterygiophore formula of 3-22110, 10 precaudal and 16 caudal vertebrae, one epural and two anal-fin pterygiophores anterior to the first he- mal spine. More precise interrelationships between the Priolepis group genera, including the relationships of Coryphopterus, are not known. Two noteworthy candidates for the sister taxon to Coryphopterus have been proposed, both members of the Priolepis group. The first, Lophogobius Gill, includes three species, two western Atlantic (L. cyprinoides (Pallas) and L. androsensis Breder) and one eastern Pacific (L. cristulatus Ginsburg). Lophogobius shares several similarities with Coryphopterus, including gonad structure (which varies considerably among gobies) and a protogynous hermaphroditic sexual pat- tern (Cole, 1988). Lophogobius cyprinoides also has a conspicuous fleshy head crest similar to the fleshy ridge seen among Coryphopterus species. The development of this crest varies among species of Lophogobius: in L. cyprinoides the crest is prominent, and in the illustration of the holotype of L. androsensis (Breder, 1932) a large crest is figured. THACKER AND COLE: CORYPHOPTERUS PHYLOGENY 839 Lophogobius cristulatus is described as differing from L. cyprinoides in that the crest is present, but markedly smaller (Ginsburg, 1939). An additional candidate that has been proposed as a close relative of Coryphopterus is Fusigobius Whitley. Fusigobius is an Indo-Pacific genus which also has a gonad struc- ture similar to that of Coryphopterus and also exhibits protogynous hermaphroditism (Cole, 1988). There are six described Fusigobius species: F. neophytus (Günther, 1877), the type species, and F. l ongispinus Goren, 1978; F. duospilus Hoese and Reader, 1985; F. signipinnis Hoese and Obika, 1988; F. inframaculatus (Randall, 1994); and F. aureus Chen and Shao, 1997. Several species of Fusigobius are also similar in coloration and external morphology to those species of Coryphopterus that are associated with benthic reef substrate. This similarity led Randall (1995) to place all Indo-Pacific species of Fusigobius in the genus Coryphopterus, although other authors (Chen and Shao, 1997) advocate recognition of a Fusigobius distinct from Coryphopterus until a revision of Fusigobius is available. Several species of Fusigobius remain undescribed; the described species have only slight meristic differences, but vary in the degree to which their pelvic fins are joined, from separate to a complete pelvic disk. They are distinguished from one another primarily on the basis of color pattern. Determination of which (if either) of these genera is the sister taxon to Coryphopterus will inform hypotheses about the biogeography of the genera. If Fusigobius is the sister genus (or the same genus), the phylogeny would be consistent with the hypothesis that Coryphopterus originated in the Pacific and subsequently radiated into the Caribbean prior to the last closing of the land bridge betweeen South and Central America. Alterna- tively, if Lophogobius is the sister group to Coryphopterus, origin of Coryphopterus in the eastern Pacific or western Atlantic is hypothesized. Additionally,
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