Evidence for a Monophyletic Gobiinae Author(s): Frank Pezold Source: Copeia, Vol. 1993, No. 3 (Aug. 18, 1993), pp. 634-643 Published by: American Society of Ichthyologists and Herpetologists Stable URL: http://www.jstor.org/stable/1447224 Accessed: 19/03/2009 17:13 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=asih. 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Supported by operating LEE, Freezing- induced in the heart rate of wood grant #GM43796 from the National Institutes changes frogs, of Health to KBS. Rana sylvatica. Ibid. 257:R1046-R1049. LOWRY,O. H., ANDJ. V. PASSONNEAU.1972. A flex- ible system of enzymatic analysis. Academic Press, LITERATURE CITED New York, New York. STOREY,K. B. 1990. Life in a frozen state: adaptive BERGMEYER,H. U. 1974. Methods of enzymatic anal- strategies for natural freeze tolerance in amphibi- ysis. Academic Press, New York, New York. ans and reptiles. Amer.J. Physiol. 258:R559-R568. CHURCHILL,T. A., AND K. B. STOREY.1991. Meta- , ANDJ. M. STOREY. 1984. Biochemical ad- bolic responses to freezing by organs of hatchling aptation for freezing tolerance in the wood frog, painted turtles, Chrysemyspicta marginata and C. p. Rana sylvatica. J. Comp. Physiol. 155:29-36. bellii. Can. J. Zool. 69:2978-2984. , AND . 1988. Freeze tolerance in ani- , AND . 1992a. Natural freezing survival mals. Physiol. Rev. 68:27-84. by painted turtles Chrysemyspicta marginata and C. , AND . 1992. Natural freeze tolerance p. bellii. Amer. J. Physiol. 262:R530-R537. in ectothermic vertebrates. Ann. Rev. Physiol. 54: , AND . 1992b. Responses to freezing 619-637. exposure by hatchling turtles Trachemysscripta ele- , S. P. J. BROOKS, T. A. CHURCHILL, gans: factors influencing the development of freeze ANDR. J. BROOKS.1988. Hatchling turtles survive tolerance by reptiles. J. Exp. Biol. 167:221-233. freezing during winter hibernation. Proc. Natl. , AND . 1992c. Freezing survival of the Acad. Sci. 85:8350-8354. garter snake Thamnophis sirtalis parietalis. Can. J. Zool. 70:99-105. (KBS, TAC, JMS) DEPARTMENTSOF BIOLOGY CLAUSSEN, D. L., P. M. DANIEL, S. AND N. A. JIANG, AND CHEMISTRY, CARLETON UNIVERSITY, ADAMS.1991. Hibernation in the eastern box turtle, OTTAWA, ONTARIO, K1S 5B6 CANADA; AND Terrapene c. carolina. J. Herpetol. 25:334-341. (JRL, MMC) DEPARTMENT OF BIOLOGY, COSTANZO,J.P., ANDD. L. CLAUSSEN.1990. Natural SLIPPERY ROCK UNIVERSITY, SLIPPERY freeze tolerance in the terrestrial turtle, Terrapene ROCK, PENNSYLVANIA carolina. J. Exp. Zool. 254:228-232. 16057. Send reprint requests LAYNE,J. R., AND M. C. FIRST. 1991. Resumption to KBS. Submitted 10 Jan. 1992. Accepted of physiological functions in the wood frog (Rana 16 Aug. 1992. Section editor: G. R. Ultsch. Copeia, 1993(3), pp. 634-643 Evidence for a Monophyletic Gobiinae FRANK PEZOLD Sixty-nine different oculoscapular canal pore configurations are described for 129 gobioid genera. A modified oculoscapular canal structure consisting of a single median anterior interorbital pore and a single terminal pair of nasal pores located near the posterior nares is recognized as synapomorphic for the gobiid subfamily Gobiinae. Some member genera and species have lost the anterior interorbital pore or the oculoscapular canal itself but are included based on relationships determined by other characters. The apomorphic feature is con- cordant with other diagnostic nonpolarized characters. Gobiines have one epural and most species have a 3-22110 first dorsal-fin pterygiophore insertion pattern, two prehemal pterygiophores, and 26 or 27 vertebrae. A new classification of gobioid fishes is proposed to accommodateresults presented here and to integrate information obtained from the competing two-family and six-family classifica- tions now in use. THE limits of family-level taxa within the problem of using different, and often conflict- Gobioidei are poorly defined and a matter ing, character suites on which to base gobioid of much debate. Harrison (1989) recently sum- classifications. I will not review specific argu- marized the major arguments, including the ments of the conflict here, readers are referred ? 1993 by the American Society of Ichthyologists and Herpetologists PEZOLD-MONOPHYLETIC GOBIINAE 635 TABLE 1. CONTEMPORARY GOBIOID CLASSIFICATIONS AND PROPOSED MODIFICATIONS. (*) signifies a mono- phyletic taxon. Miller (1973) Hoese (1984) Proposed Family* Family* Superfamily* Rhyacichthyidae Rhyacichthyidae Rhyacichthyoidea Family* Family Family* Gobiidae Eleotrididae Rhyacichthyidae Subfamily Family* Superfamily* Eleotrinae Kraemeriidae Gobioidea Subfamily Family* Family Pirskeninae Microdesmidae Eleotrididae Subfamily* Family* Family* Xenisthminae Xenisthmidae Kraemeriidae Subfamily Family* Family* Gobionellinae Gobiidae Microdesmidae Subfamily Subfamily Family* Tridentigerinae Amblyopinae Xenisthmidae Subfamily Subfamily Family* Gobiinae Gobiinae Gobiidae Subfamily* Subfamily* Subfamily* Kraemeriinae Oxudercinae Amblyopinae Subfamily* Subfamily* Sicydiinae Gobiinae Subfamily Gobionellinae Subfamily* Oxudercinae Subfamily* Sicydiinae to Harrison's work for further information. Two all gobioid fishes. By Millers's classification, sev- classifications of gobioid fishes are currently in en subfamilies encompass the diversity obtained use: a classification that recognizes two families, within the Gobiidae. Other than the Xenisth- with the Gobiidae comprising seven subfami- minae, however, only the Kraemeriinae could lies; and one that recognizes six families, with be delimited by apomorphic features (Table 1). the Gobiidae comprising four subfamilies. Both Hoese (1984) divided gobioid fishes into six classifications accept poly- or paraphyletic fam- families: Rhyacichthyidae, Eleotrididae, Gobi- ily-level taxa; nonmonophyletic subfamilies are idae (including four subfamilies), Xenisthmi- allowed explicitly (Miller, 1973) or implicitly dae, Microdesmidae, and Kraemeriidae (Table (Springer, 1983) in the two-family classification, 1). Of these, all but the Eleotrididae were char- and nonmonophyletic families are proposed in acterized by at least one derived character state. the six-family concept (Hoese, 1984). Subsequently, Springer (1988) followed Hoese Miller (1973) recognized a Gobiidae encom- and accepted the Xenisthmidae as a valid fam- passing all gobioid species but one (about 2000 ily. Within the Gobiidae, monophyly has been species) and a Rhyacichthyidae containing that proposed for the sicydiines (Hoese, 1984; Har- one species, Rhyacichthysaspro (Table 1). The rison, 1989) and the oxudercines (Murdy, 1989). Gobiidae as proposed by Miller share two de- Species of the subfamily Amblyopinae as pre- rived features: one or two epurals and a reduced sented by Hoese (1984) and Birdsong et al. cephalic lateralis system. Rhyacichthysaspro forms (1988) do not share any derived character states. the primitive sister group to all other gobioids. I have noted in an unpublished manuscript (cit- It has three epurals and a well-developed later- ed in Birdsong et al., 1988) that Gobioides,pre- alis on the head. Springer (1983) strengthened viously placed in the Amblyopinae, shares sev- Miller's classification with additional gobiid syn- eral derived features of the cephalic lateralis apomorphies, supported recognition of the xe- with Gobionellus,a member of Hoese's Gobiinae. nisthmines as a distinct subfamily within the Go- Harrison (1989) also noted that Gobioidesshares biidae, and provided four synapomorphies for a derived palatine condition with Gobionellusand 636 COPEIA, 1993, NO. 3 TABLE 2. OCULOSCAPULAR CANAL STRUCTURES AND PORE PATTERNS OBSERVED OR REPORTED IN LITERATURE CITED. Pore terminology follows Akihito et al. (1984). Dots signify canal absent in that region. "0" indicates canal present, but pore is absent from that position. "f" refers to a single pore resulting from the fusion of two separate portions of the canal and their terminal pores. "x" indicates a pore is present. For pores C and D, a "2" indicates paired, separate interorbital canals are present with a pair of pores, whereas " 1" indicates a connection between the canals with a single median pore exhibited. Oculoscapular