Evidence for a Monophyletic 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

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http://www.jstor.org 634 COPEIA, 1993, NO. 3 of Ottawa for performing osmolality measure- sylvatica)after freezing. Amer.J. Physiol. 261 :R134- R137. ments, andJ. A. Duncan of Carleton University ,R. E. AND T. L. HEIL. 1989. for technical support. 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 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 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 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* Eleotrididae Rhyacichthyidae Subfamily Family* Superfamily* Eleotrinae Kraemeriidae Gobioidea Subfamily Family* Family Pirskeninae Microdesmidae Eleotrididae Subfamily* Family* Family* Xenisthminae Xenisthmidae Kraemeriidae Subfamily Family* Family* 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 ,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 ,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 pores Taxa A B C D E F G H I J K L 1 x x 2 2 x x x x x x x x 2 x x 2 1 x x x x x x x x 3 x x 2 1 x x 0 x x x x x 4 x x 0 1 0 x 0 x x x x x 5 x x 0 1 0 x 0 x 0 x x x 6 x x 2 1 0 x 0 x 0 x x x 7 x x 2 1 0 x 0 x 0 0 x x 8 x x 0 1 0 x 0 x 0 0 x x 9 x x 2 1 0 x 0 f 0 0 f x 10 x x 2 1 x x 0 x x x x I11 x x 2 1 0 x x x x x 12 x x 2 1 x x 0 x * * ~~x x 13 x x 2 1 0 x 0 x * * ~~x x 14 x x 2 1 0 x 0 x 15 0 x 0 x 1 16 x x x x x x 1 17 x x 0 x x x 1 18 0 x 0 x x x 1 19 x x 0 x 1 20 x x x 21 0 0 x 0 x x x 22 2 0 x 0 x 23 1 0 x 0 x 24 1 0 x x 25 2 x x x 26 2 0 x 0 x * 27 1 x x 28 1 0 x 29 0 0 x 30 1 x x x f 0 0 f x 31 1 x x x f 0 0 f x 32 1 x x x x 0 x x x 33 1 x x x x x x 34 1 0 x x x * * ~x x 35 1 x x 0 x * x x 36 1 0 x 0 x * x x 37 1 x x x x 38 1 0 x x x 39 1 x x 0 x 40 0 x x 0 x 41 1 0 x 0 x 42 1 x x x 43 1 x x 44 0 x x 45 1 0 x 46 1 47 1 x x 48 0 x x 49 1 50 1 x x x x * * ~~~x x 51 1 0 x x x x x PEZOLD-MONOPHYLETIC GOBIINAE 637

TABLE 2. CONTINUED.

Oculoscapularpores Taxa A B C D E F G H I J K L 52 x 0 1 x x O x 53 x 0 1 0 x 0 x * x 54 x 0 0 0 x 0 x x x 55 x 0 1 0 x x x 56 x 0 0 x x 57 x 0 1 0 x x x 58 x 0 1 x x 0 x 59 x 0 1 0 x 0 x 60 x 0 0 0 x 0 x 61 x 0 1 x x 62 x 0 1 0 x . 63 x 0 0 x 64 0 1 . .. 65 x* 66 2 x 67 x x 0 x 68 * x x 69 0x . x x

associated genera. The remaining members of 1965); Egglestonichthys (Miller and Wongrat, the Amblyopinae, the Trypauchenand Taenioides 1979); (Lachner and Karnella, 1978, groups of Birdsong et al. (1988), have a derived 1980); Gobiopsis(Lachner and McKinney, 1978, fin element to vertebra ratio of 2:1. Transfer- 1979); Gobius (Miller, 1974; Miller and El-Ta- ring Gobioidesto the Gobiinae results in three wil, 1974); Gorogobius(Miller, 1978b); Knipow- gobiid subfamilies out of four hypothesized as itschia (Miller, 1972b); Monishia (Miller, 1978a); monophyletic. The Gobiinae still contains the Odondebuenia (Miller and Tortonese, 1968); bulk of gobioid fishes, however; and its mono- Thorogobius(Miller, 1969); Zebrus(Miller, 1977); phyly is unconfirmed. and gobiosominines (Bohlke and Robins, 1968). Analysis of the oculoscapular canal system in Descriptions of gobies from works on the fol- gobioid fishes revealed a number of distinctive lowing regions were also used: Chagos Archi- structures. In this paper, I propose that a unique pelago (Winterbottom and Emery, 1986); east modification of the anterior oculoscapular canal central Atlantic (Miller, 1981); Japan is synapomorphic for most of the species in- (Akihito et al., 1984); Macaronesia (Miller, cluded in Hoese's Gobiinae. This delimitation 1984). Early papers by Sanzo (1911) and Iljin of the Gobiinae results in the remnant group (1930) on gobioid relationships also provided of genera being placed in an additional subfam- descriptions of the lateralis canals of European ily, the Gobionellinae, for which monophyly is gobies. not established. Incorporation of this assem- Canal and canal pore terminology follows blage into a version of Hoese's classification Takagi (1957) and Akihito et al. (1984). Pore modified to reflect phylogenetic information names (Takagi, 1957) are used in discussions, presented by Springer (1983) obtains a further whereas the lettering system of Akihito et al. refinement of gobioid relationships. Accep- (1984) is used to report pore configurations for tance of the Gobionellinae as a subfamily has a a taxon. All drawings were made using a Wild heuristic value similar to that provided by rec- M-5 dissecting microscope and camera lucida. ognition of the Eleotrididae. Where preopercular canals are present in go- biids, the canals have no more than three pores. to canal or the total loss MATERIALS AND METHODS Reduction a two-pored of the canal has occurred independently many Collections examined are listed below. Cteno- times (e.g., Takagi, 1989). The preopercular gobius and Gobionellus specimens are listed in canal system was, therefore, considered sepa- Pezold (1984). Information was drawn from the rately from the more complex oculoscapular literature for: Chromogobius(Miller, 1971); Cor- system because (1) preopercular canal modifi- cyrogobius (Miller, 1972a); Didogobius (Miller, cations often appear independently of oculo- 638 COPEIA, 1993, NO. 3

TABLE 3. GENERA EXHIBITING OCULOSCAPULAR TABLE 3. CONTINUED. CANALS AND PORE PATTERNS DESCRIBED IN TABLE 2. 42. Pleurosicya 1. Butis, Bostrychusa 43. Eviota,a,b Gobiopsisa.b 2. Oxyeleotris 44. Eviotaa,b 3. Ophiocara 45. Risor 4. Bostrychusa 46. Stonogobiopsab 5. Parviparma 47. Eviotaab 6. Awaousa 48. Eviotaab 7. Gobionellus 49. Stonogobiopsab 8. Gobioides 50. Barbuligobius,aBathygobiusa 9. Sicydium, Sicyopterus 51. Mahidolia 10. Xenisthmus 52. Gobiopsisab 11. Awaousa 53. ,Gobiomorus, Parachaeturichthys 12. Tridentiger 54. ,Deltentosteus, Gobiopsis,abHetereleo- 13. , , Lentipes, , Sicy- trisa opus, ,Stiphodon 55. Hemieleotris,Leptophilypnus, 14. , 56. Gobitrichinotus,Kraemeriab 15. Tukugobiusa 57. Callogobius,a.b Parrella, Vanderhorstiaa 16. Coryphopterusa 58. Gobiopsis,ab Ptereleotrisa 17. ,aRhinogobius 59. Aruma, Chasmichthys,Didogobius, Eleotrica, Pycnom- 18. Pterogobius,aSagamia, Tukugobiusa ma, Suruga 19. , Nemateleotris 60. Barbulifer, Callogobiusab 20. Bryaninops 61. Gobiopsis,abPtereleotrisa 21. 62. Callogobius,abParkraemeria, 22. ChaenogobiusSab 63. Apocryptesa 23. Knipowitschia 64. Apocryptes,aBoleophthalmus, Parapocryptes 24. ChaenogobiusSab 65. Scartelaos 25. Hypseleotrisa 66. Gobiomorphus,abHypseleotrisa 26. Hyrcanogobius 67. Gobiopsisa.b 27. 68. Percottus 28. Chaenogobiusa"b 69. Gobiomorphusa'b 29. aGenera with more than one pore pattern for the oculoscapular canal. 30. Bathygobius,aGobius,a Mauligobius b Genera in which the oculoscapular canal is completely lost in some 31. Bathygobiusa species. 32. Fusigobius,ano connection between J and K. 33. Acentrogobius,aAmblyeleotris,a Amblygobius, Asterrop- scapular canal changes; (2) there are teryx,Barbuligobius,a Bathygobius,a Bollmannia,a Ca- relatively few, simple states possible for the billus, Caffrogobius, Callogobius,ab Coryphopterus,a preopercular canal; and similar, but con- Cristatogobius, Cryptocentroides, (3) nonhomologous, Cryptocentrus,a ditions have arisen a number of times Ctenogobiops,, Exyrias, Favonigobius, Fu- apparently within the siogobius,aGladiogobius, Glossogobius,aGobius,a Ha- family. zeus, Heteroplopomus,Istigobius,a Macrodontogobius, Mangarinus, Monishia, Myersina,a Oplopomus, Sig- RESULTS nigobius, Silhouettea, Thorogobius, Valenciennea, Vanderhorstia,aYongeichthyes, Zebrus Sixty-nine different oculoscapular canal pore 34. Bollmannia,a Gobiusa patterns were observed and/or derived from 35. Amblyeleotris,aBuenia, Cryptocentrus,aEnypnias,a published accounts (Tables 2-3). Although this Gobiosoma,aGobiusculus, Istigobius,a Pomatoschistusa information does not include all gobioid taxa, 36. Pomatoschistusa 129 genera are represented in the tables. Gen- 37. Acentrogobius,aCallogobius,ab Chromogobius, Discor- era with species lacking oculoscapular canals dipinna, Flabelligobius, Glossogobius,aMillerigobius, were noted only if canals were present in some Myersina,aOdondebuenia, Vanderhorstiaa species. 38. of F. examined Fusigobiusa (specimens neophytus Pore configurations outlined in Tables 2 and here) 3 range from patterns unique to specific genera 39. Elacatinus, Enypnias,a Eviota,a.b Garmannia, Gins- to shared diverse taxa. A Heter- patterns by many large burgellus, Gobiodon, Gobiosoma,aGorogobius, of to oeleotris,a,b group "typical" gobiids usually assigned Tomiyamichthys the Gobiinae 40. subfamily (e.g., Hoese, 1984) is Paragobiodon characterized an canal 41. Gobiosoma,aPomatoschistusa by oculoscapular with a single median anterior interorbital pore (pore PEZOLD-MONOPHYLETIC GOBIINAE 639

/ 0! I0

/ Hi M' N N/ ,,, /X_^K'o/

Fig. 2. The oculoscapularcanal and pores of Evor- thoduslyricus. Labeling as in Figure 1.

Fig. 1. The oculoscapularcanal and pores of Cryp- The Indo-Pacific is tocentroidesinsignis. Canal pores labeled as in Akihito comm.). Bryaninops et al. (1984). Dotted lines indicate canal. Terminal tentatively included in the Gobiinae because of pores indicated by '. an = anterior nares, pn = pos- its relationship to Pleurosicya (Larson, 1985). terior nares. These are small obligate symbionts associated with corals and sea grasses. All the species of Bryaninops and Coryphopterus hyalinus have C). The canal terminates in a single pair of pores shortened snout canals with a single pair of nasal (B') near the posterior nares (Fig. 1). This mod- pores. Because some species of Bryaninops show ification of the lateralis is unique among gobioid variability in the connection of the posterior fishes. The more general condition is illustrated interorbital section of the oculoscapular canal in Fig. 2. ontogenetically (Larson, 1985), the condition Takagi (1989) noted that species with the of the anterior section is probably secondarily modified oculoscapular pore arrangement usu- derived. Several other gobiid genera (Tables 2- ally have a single, unified interorbital canal (his 3: 17-20) have reduced canals on the snout and type III condition). One exception he found was a single pair of nasal pores, but do not have Callogobiusokinawae, in which the canals are sep- united anterior interorbital canals. arate although a single anterior interorbital pore D. Hoese (pers. comm.) has observed a species is present. Other species of Callogobius have ca- of Nesogobius with two pairs of pores on the nals united and a single pore, or no pore. Takagi snout (A and B in Table 3) and a single median presented evidence that a unified interorbital anterior interorbital pore. Because I have not canal is a specialization found only within gobies seen this genus, and have no information on and some blennioid fishes. other species of the genus, I cannot comment The single AIP is a stable character for go- on the derivation of the canal structure. biines, with only two of the 129 genera con- Species with the derived anterior interorbital taining a species showing a reversal from the condition (single AIP and one pair of nasal pores; single-pored to the two-pored state. One species Tables 2-3: 30 and 32-46) usually have a su- of Coryphopterus,C. hyalinus, has two anterior praotic pore (E in Table 2; not in 34, 36, 38, interorbital pores [see Smith and Tyler (1977) 41, 45, and 46) and/or a posterior otic pore (G for cladistic evidence supporting a reversal]. The in Table 2; lacking in 35, 36, 39-41, and 43- interorbital or anterior portion of the right and 46). Takagi (1989) also noted the general as- left oculoscapular canals initially develop sep- sociation of the supraotic and posterior otic arately in gobioid fishes and may fuse later in pores with the interorbital canal modification. development. Consequently, developmental The association of one or both of these pores truncation could lead to reversals to the two- may be diagnostic for some gobiine genera (e.g., pored (unconnected canal) state (D. Hoese, pers. Amblygobius,Asterropteryx, Cabillus, Caffrogobius, 640 COPEIA, 1993, NO. 3

Table 2: 33), but their presence in a number of (1973) also permits greater information retriev- gobioid fishes lacking the modified interorbital, al within the Gobiidae; natural subfamilies and including some eleotridids (Table 2: 1-3), in- possibly tribes may be discerned. To incorpo- dicates they are plesiomorphic for the Gobiinae. rate phylogenetic information presently ob- tained only in the two-family classification pro- Miller as Subfamily Gobiinae posed by (1973), developed by Springer (1983), I propose the use of two superfamilies, Diagnosis.-A single anterior interorbital pore the Gobioidea (=Gobiidae of Miller, 1973; present, one pair of nasal pores (posterior), in- Springer, 1983) and Rhyacichthyoidea (in- terorbital portion of oculoscapular canal fused; cludes the Rhyacichthyidae of Miller, 1973; anterior interorbital pore, posterior nasal pores, Springer, 1983; Hoese, 1984). interorbital section of oculoscapular canal or For those gobies outside of the Gobiinae, Si- entire oculoscapular canal lost in some species. cydiinae, Oxudercinae, and Amblyopinae, the One epural; most species with 3-22110 first dor- name Gobionellinae proposed by Miller (1973) sal-fin pterygiophore insertion pattern or de- may be used. However, the names Brachygo- rivative; most species with 26 or 27 vertebrae biinae, Chaeturichthyinae, Platygobiinae, and and two prehemal pterygiophores. Luciogobiinae may also be available (Bleeker, The Gobiinae as recognized here include the 1874). Birdsong et al. (1988) proposed three Priolepis, Microgobius, Gobius, Bathygobius,Gobio- monophyletic groups placed here with the go- soma, Pomatoschistus, and Kelloggella groups of bionellines, the Acanthogobius,, and Chas- Birdsong et al. (1988). Included genera are list- michthysgroups (Appendix 2). Species contained ed in Appendix 1. Although some members of in all three of these groups show elevated ver- these groups have lost the anterior interorbital tebral numbers (30-42), and members of the pore, their generic or generic group relation- Astrabe and Chasmichthysgroups have posteri- ships may be recognized by other characters. orly displaced insertions of the first dorsal fin. The bulk of the gobiine species I recognize have A posteriorly displaced first dorsal fin is often either a 3-22110 or 3-221110 first dorsal pte- obtained in some eleotridids (Guavina, Odonto- rygiophore insertion pattern and a single ep- butis, Ophieleotris,Gobiomorphus, Mogurnda) and ural, although these osteological characters are a few other gobiids (Evermannichthys,Lethrops, not polarized herein. Most other insertion pat- and ).Lethrops and Synechogobius, terns found in the gobiines are more easily de- both northern Pacific gobiid genera, are prob- rived from the 3-22110 or 3-221110 patterns ably related to members of the Astrabeand Chas- than the 3-1220 pattern. An exception is Po- michthysgroups. The condition reported for Ev- matoschistuswhich has 3-122100 and 3-1221000 ermannichthys (Birdsong et al., 1988) is patterns (Birdsong et al., 1988). Until conflict- undoubtedly homoplastic. Posteriorly displaced ing information is obtained from characters that first dorsal fins in eleotridids insert in the fourth can be polarized, Pomatoschistus is retained in interneural space. All members of the Astrabe the Gobiinae. group and two genera of the Chasmichthysgroup The single anterior interorbital pore delimits (Clevelandia and )have first pteryg- a smaller group than the Gobiinae of Hoese iophores inserting in the fifth through tenth (1984); however, it still includes most gobiids interneural spaces. Insertions beyond the fourth (Birdsong et al., 1988). Recognition of the Go- interneural space are derived conditions, but biinae as defined here also leaves a number of polarity of the fourth interneural insertion is gobiids (though smaller) outside of a monophy- ambiguous. letic subfamily (Appendix 2). Addressing this The number of vertebrae offers less insight problem (and ignoring the problem of the re- as a character than the first dorsal-fin insertion. lationships of the included species for the mo- Relatively high numbers of vertebrae (30 or ment), I follow Hoese (1984) and recognize six more) are also found in eleotridids (Gobiomor- families of gobioids as well as five subfamilies phus, Micropercops,Mogurnda, Odontobutis,Per- of gobiids (Table 1). Distinguishing major go- cottus, Philypnodon, Tateurndina), kraemeriids bioid lineages as families is more informative to (Gobitrichonotus),microdesmids (Cerdale, Clark- individuals not familiar with the intricacies of ichthys, Gunnellichthys, Microdesmus, Paragunne- gobioid systematics than designating the lin- lichthys) and other gobiids (Deltentosteus, Ever- eages subfamilies. A species' family member- mannichthys, Gobioides, Lethrops, Neogobius, ship is more often referenced by nonsystema- Nesogobius, Odontamblyopus,Ophiogobius, Poma- tists than is its membership in other categories toschistus, Synechogobius,Taenioides, Trypauchen, above the genus level. Using families to delimit Trypauchenichthys).Considering the plasticity for lower levels of relationship than those of Miller vertebral number exhibited both in and be- PEZOLD-MONOPHYLETIC GOBIINAE 641 tween gobioid species (Birdsong et al., 1988), 199452 (7); CAS-SU 19339 (2); CAS 51044 (1); ANSP 143106 (1). even transformation series of this character Evorthodus lyricus: UF 100059 (12); TNHC 10623 (29). Favonigobius may aliciae: CAS 56374 (3). F. reichei: CAS-SU 38598 (1); USNM 258131 offer little information at higher levels of uni- (4). Fusigobius neophytus: CAS-SU 9010 (2). Ginsburgellus sp.: USNM the concordance of first 170262 (1). Glossogobiusbicirrhosus: CAS 51058 (12). G. callidus: ANSP versality. Although G. CAS 56375 cauerensis: CAS dorsal fin and vertebral number character states 55287-9 (3). giuris: (16). Gnatholepis 51548. G. thompsoni:UMMZ 174286 (5). Gobiodonquinquestrigatus: CAS- suggests the existence of a northern Pacific go- SU 26717 (1). Gobioidesbroussoneti: USNM 233612 (11); ANSP 121256 biid clade, its limits are defined. The (1). G. africanus: BMNH 1939.7.12.33 (1). G. ansorgii: BMNH poorly 1968.11.15.77 (1). G. grahamae: BMNH 1959.3.17.161 (1). Gobiomor- synapomorphies proposed by Birdsong et al. phus australis: USNM 265059 (1). G. gobioides: ANSP 122811 (45). Go- for the Astrabe, and Chas- biomorphussp.: USNM 210751 (5). Gobiomorusdormitor: CAS-SU 61392 (1988) Acanthogobius, UF 21835 Gobius CAS-SU 61441 further refinement and (4). Gobiosomalongipala: (1). niger: (2). michthysgroups require G. paganellus: CAS-SU 1688 (2). Gobiopterussp.: USNM 217264 (3). resolution against other character state distri- Guavina guavina: CAS-SU 52357 (3). Hemieleotris latifasciatus: USNM butions. 26500t) (3); ANSP 71084-96 (13). Hetereleotriszonatus: USNM 210413 (1). Hypseleotriscyprinoides: USNM 262027 (39). H. guntheri:ANSP 31675- Although not delimited as a group in Appen- 91 (16). H. tohizonae:USNM 143781 (1). Istigobius ornatus: CAS 46503 dix 2, Harrison (1989) hypothesized a relation- (5). Leptophilypnusfluviatilis: USNM 249722 (16); ANSP 122361 (7). Lophogobiuscyprinoides: CAS 52008 (1); TNHC 10925 (2); TNHC 10862 ship between the genera Ctenogobius,Evorthodus, (2). Mahidolia mystacina:CAS 51063 (1); MNHN 1965-217 (2); RMNH Gnatholepis,Gobioides, Gobionellus, Oligolepis, and 15260 (1); MNHN 2967 (1). Microgobiusthalassinus: UF 30508 (4). Mo- based an gurnda mogurnda: ANSP 89831 (3). abei: CAS-SU 30266 Stenogobius upon apomorphic palatine (2); Mugilogobiussp.: USNM uncat. IIOE, Seychelles, stn. F-70 (2). Odon- structure. The same structure was observed in tanblyopus sp.: USNM 86955 (9). : UMMZ 100537 , but that was as the sis- (25); RMNH 14325 (3). O.jaarmani: USNM 217267 (8); USNM 217266 genus suggested 0. stomias: USNM 257137 USNM 258782 USNM 99296 ter to the because a (2). (13); (9); group sicydiines of syna- (1). Ophiocaraporocephala: CAS-SU 38579 (4). Oplopomusoplopomrus: CAS pomorphic suborbital free neuromast pattern. 51589. Oxyeleotrismarmorata: USNM 230328 (2); ANSP 87352 (3). Oxu- there is difference in detail, Harri- richthysauchenolepis: RMNH 4506 (2). 0. keiensis:RUSI 17043 (8); RUSI Although 16786 (1). 0. lonchotus:ANSP 23350 (1); ANSP 90998 (3); ANSP 28055- son's general hypothesis of a close relationship 56 (2); CAS 23328 (14). 0. microlepis:ANSP 88946 (1). 0. ophalmonema: of these Awaous, is CAS 61040 (1). 0. papuensis: LACM 37382-2 (1). 0. stigmalophius:ANSP genera, including supported tentacularis: ANSP 100179 other derived features of the later- 81233 (1); ANSP 81855 (1). 0. (1). Oxyurich- by cephalic thys sp.: CAS 51047 (75). Parapocryptescantonensis: UMMZ 100616 (4). alis (Pezold, unpubl. cited in Birdsong et al., Parrella ginsburgi: CAS-SU 46827 (19). Panriparma straminea: CAS-SU No formal is 29701 (1). Percottusglehni: USNM 86108 (1). Periophthalmuscantonensis: 1988). group recognition given CAS-SU 61102 (5). Philypnodon grandiceps: USNM 59976 (9). Pomatos- here, however, because the relationship of these chistus microps:ANSP 147148 (20); CAS-SU 32183 (3). Porogobiusschle- genera to other gobionellines, oxudercines, and geli: CAS-SU 63028 (2). Ptereleotris calliurus: USNM 118102 (4). P. is unclear helenae: CAS-SU 63565 (1). Quietula ycauda: CAS 41895 (5). Redigobius sicydiines (Harrison, 1988; unpubl.). balteata: CAS 51061 (9). R. bikolanus: NLU uncat., John Lake Coll.; Acceptance of the Gobiinae as defined by the Australia. R. dispar: CAS-SU 69019 (4). R. macrognathos:CAS-SU 38647 derived canal confers new infor- (3). Rhinogobiuscandidus: CAS-SU 23185 (2). R. kurodai: CAS-SU 23529 oculoscapular (5). R. hadropterus: CAS-SU 23536 (5). R. leavelli: CAS-SU 32832 (3). mation on relationship for a large number of R. similis: CAS-SU 58338 (3); RMNH 12509 (1). R. taiwanus: CAS-SU gobiids. The Gobionellinae simply becomes a 23182 (1); CAS-SU 23104 (2); CAS-SU 23167 (1). Rhyacichthysaspro: smaller of USNM 247300 (5). Sicydiumpittieri: CAS 52012 (3); TNHC 11524 (1); assemblage convenience. TNHC 11507 (1); TNHC 11499 (1); TNHC 11510 (1); TNHC 11521 (1). S. plunieri: USNM uncat. acc. no. 249592, Dominica (3). Sicydium sp.: MCZ 33968 (1). Sicyopterusouwensi: CAS-SU 61629 (2). Stenogobius MATERIAL EXAMINED genivittatus: ANSP 86151 (9); ANSP 28016-19 (4); BPBM 26373 (4); BPBM 26380 (2); CAS 51056 (7); CAS 52011 (3). S. gymnopomus:RMNH 4552 (4). S. laterisquamatus: ZMA 116.477 (2); WAM P27847-007 (3); Institutional abbreviations follow Leviton et al. unless oth- (1985) WAM P28206-002 (1); NLU 62766, Laloki River (4). Stigmatogobius erwise noted. sadanundio: USNM 263326 (4); ANSP 77797 (1). Stiphodon elegans: CAS caninus: CAS Acanthogobiusflavimanus: 52003 (4). Acentrogobius USNM 260937 (29). Taenioides cirratus: USNM 243405 (1). Taenioides 51065 CAS-SU 26209 CAS-SU 33073 (2). Amblyeleotrisfontanesii: (1); sp.: USNM uncat. SOSC ref. no. 134, Madagascar (4); ANSP 63086- USNM 160741 A. randalli: USNM 220085 A. steinitzi:USNM (5); (2). (1). 87 (2). Tridentigerobscurus:CAS-SU 6590 (3); CAS 52013 (3). Trnpauchen 282957 minuta: USNM 265060 bato: UMMZ (2). Aphia (1). Apocryptes vagina: USNM 119644 (1). Tukugobiusbucculentus: CAS-SU 39400 (5); 187890 Awaous CAS-SU 55635 A. stamineus: ANSP (9). guineensis: (2). CAS-SU 26366 (5); CAS-SU 26363 (1). T. carpenteri: CAS-SU 26367 29510-13 A. ANSP 144525 FMNH 93277 UF (4); tajasica: (3); (3); (5). T. philippinus: CAS-SU 38601 (5). Tyntlastes brevis: USNM 77581 30510 of Texas Marine Sciences Institute 334 (1); University (UTMSI) (1). Valenciennea longipinnis: CAS 13762 (2); USNM 259429 (6). V. mur- A. transandeanus: TNHC 11519 TNHC 11506 TNHC (3). (1); (1); alis: CAS 51060 (1). V. sexguttatus: CAS 59878 (2). Vanderhorstiaorna- 11509 TNHC 11511 Awaous ANSP 149484 (1); (1). sp.: (1). Bathygobius tissima:CAS 59865. Yongeichthyescriniger: CAS-SU 9425 (1). Y. nebulosus: soporator:TNHC 10469 (5); TNHC 10809 (6). Boleophthalmusboddarti: CAS-SU 20331 (3). CAS-SU 33141 (1). Bollmannia chlanydes: CAS 42777 (1). B. communis: USNM 195539 (1). Bostrychussinensis: USNM 85907 (6). Brachyambly- opus sp.: USNM 243403 (45). Butis butis: USNM 261350 (13); ANSP 63023-39 (17). Callogobius sp.: CAS 5105 (16). Chaenogobiusurotaenia: ACKNOWLEDGMENTS CAS 52005 (2). Chaeturichthysstigmatias: CAS-SU 61210 (5). Corypho- pterusglaucofrenum: FMNH 48426 (2). C. nicholsii: CAS-SU 48966 (15). This was of a Cryptocentroidesinsignis: CAS 51064 (1). Cryptocentruscryptocentrus: MNHN paper part manuscript previ- A.1166(1); BMNH 1975.5.5.3-4(2). C.filifer: FMNH 3814 (3); UMMZ ously read by R. Birdsong, R. C. Cashner, D. 142629 (1); MNHN 7196 (1). C. strigilliceps: CAS 52009 (1). Cryptocen- Hoese, H. Larson, E. O. Murdy, L. Parenti, and trus sp.: CAS 52010 (1); CAS 52006 (3). Dormitator latifrons: CAS-SU 55012 (1). D. lebretoni: ANSP 38675-78 (4). D. maculatus: MSU 1273 R. Winterbottom. All reviewers provided help- (2); MSU 7692 (4). Dormitator sp.: ANSP 99855 (2). Drombus palackyi: ful comments. It was at R. Birdsong's suggestion USNM 57971 (1). Eleotris pisonis: ANSP 99833 (3); ANSP 104302 (3); that this work was from the TNHC 8115 (2); TNHC 10473 (1). E. picta: ANSP 104165 (2). Eleotris separated larger sp.: CAS-SU 52359 (1); CAS-SU 52360 (2). Erotelis smaragdus: UMMZ manuscript. All conclusions are my own and 642 COPEIA, 1993, NO. 3 may or may not be shared by the reviewers in the genus Gobiopsisand a redescription of Feia nym- their entirety. Specimens and information on pha Smith. Ibid. 299:1-18. materials in their care were provided by W. LARSON, H. K. 1985. A revision of the gobiid genus Eschmeyer, T. Iwamoto, P. Sonoda, M. Hearne, Bryaninops (Pisces), with a description of six new B. Chernoff, W. Smith-Vaniz, E. Bohlke, R. R. species. The Beagle, Occ. Pap. Northern Terr. Mus. Arts and Sci. 2:57-93. Miller, A. S. C. R. Gilbert, Snyder-Creighton, A. R. H. E. ANDC. E. G. H. B. K. E. A. LEVITON, E., GIBBS, JR., HEAL, Burgess, J. Miller, Hartel, DAWSON. 1985. Standards in and ich- Wheeler, M. Chambers, M-L. herpetology Holloway, J. thyology: Part I. Standard codes for in- V. G. S. T. Wil- symbolic Bauchot, Springer, Jewett, J. stitutional resource collections in herpetology and liams, R. K. Johnson, D. J. Stewart, T. Grande, . Copeia 1985:802-832. C. E. Dawson, S. Poss, M. J. P. van Oijen, J. MILLER, P.J. 1965. A new genus and species of gobiid Randall, R. Winterbottom, J. Nielsen, D. Mo- from the eastern Mediterranean. Amer. Mus. sier, T. van den Audenaerde, M. Louette, H. Nat. Hist. Ser. 13, 8:161-172, 2 pls. Nijssen, P. C. Heemstra, R. G. Gilmore, and G. .1969. Systematics and biology of the leopard- Gobius Gobi- Allen. Portions of this study were supported by spotted goby, ephippiatus (Teleostei: with of a new and notes on travel from Xi, the Smithsonian idae), description genus grants Sigma the of G. Kolombatovic. the of Natural Sciences of identity macrolepis J. Mar. Institution, Academy Biol. Ass. U.K. 49:831-855. and the California of Philadelphia, Academy 1971. A revision of the Mediterranean gobiid Sciences. genus Chromogobius(Teleostei-Perciformes).J. Zool., London 164:305-334. LITERATURE CITED . 1972a. Generic status and redescription of the Mediterranean fish Gobius liechtensteiniKolom- PRINCEAKIHITO, M. HAYASHI, AND T. YOSHINO. batovic, 1891 (Teleostei: Gobioidea), and its affin- 1984. Suborder Gobioidei, p. 236-289. In: The ities with certain American and Indo-Pacific gobies. fishes of the Japanese Archipelago. H. Masuda, K. J. Nat. Hist. 6:395-407. Amaoka,C. Araga,and T. Uyeno (eds.).Tokai Univ. . 1972b. Gobiid fishes of the Caspian genus Press, Tokyo, Japan. Knipowitschia from the Adriatic Sea. J. Mar. Biol. BIRDSONG,R. S., E. O. MURDY,AND F. L. PEZOLD. Ass. U.K. 52:145-160. 1988. A study of the vertebral column and median . 1973. The osteology and adaptive features fin osteology in gobioid fishes with comments on of Rhyacichthysaspro (Teleostei: Gobioidei) and the gobioid relationships. Bull. Mar. Sci. 42:174-214. classification of gobioid fishes. J. Zool. London 171: BLEEKER,P. 1874. Esquisse d'un systeme naturel des 397-434. Gobioides. Arch. Neerl. Sci. Nat. 9:289-331. 1974. New species of Gobius (Teleostei: Go- BOHLKE, J. E., AND C. R. ROBINS. 1968. Western biidae) from the western English Channel, with a Atlantic seven-spined gobies, with descriptions of key to related species in the British and Irish fauna. ten new species and a new genus, and comments Ibid. 174:467-480. on Pacific relatives. Proc. Acad. Nat. Sci. Philadel- . 1977. Gobies from Rhodes and the system- phia 120:45-174. atic features of Zebrus zebrus (Teleostei: Gobiidae). HARRISON, I. J. 1989. Specialization of the gobioid Zool. J. Linn. Soc. 60:339-362. palatopterygoquadrate complex and its relevance . 1978a. The systematic position and origin of to gobioid systematics. J. Nat. Hist. 23:325-353. Gobius ocheticus Norman, 1927, from the Suez Ca- HOESE,D. F. 1984. Gobioidei: relationships, p. 588- nal. Ibid. 62:39-58. 591. In: Ontogeny and systematics of fishes. H. G. . 1978b. The status of the west African fish Moser, W.J. Richards, D. M. Cohen, M. P. Fahay, Gobiusnigricinctus with reference to New World au- A. W. Kendall, Jr., and S. L. Richardson (eds.). tochthones and an Old World colour-analogue. Ibid. Amer. Soc. Ich. Herp. Spec. Publ. 1, Gainesville, 64:27-39. Florida. 1981. Gobiidae. FAO Species Identification ILJIN,B. S. 1930. Le systeme des gobiides. Trab. Inst. Sheets. East Central Atlantic 2:1-8. Esp. Oceanogr. 2:1-63. . 1984. The gobiid fishes of temperate Ma- LACHNER,E. A., AND S. J. KARNELLA.1978. Fishes caronesia (eastern Atlantic). Zool. Soc., London 204: of the genus Eviota of the Red Sea with descriptions 363-412. of three new species (Teleostei: Gobiidae). Smith- , AND M. Y. EL-TAWIL. 1974. A multidisci- sonian Contr. Zool. 286:1-23. plinary approach to a new species of Gobius(Teleos- ,AND . 1980. Fishes of the Indo-Pacific tei: Gobioidea) from southern Cornwall. Ibid. 174: genus Eviota with descriptions of eight new species 539-574. (Teleostei: Gobiidae). Ibid. 315:1-127. , AND E. TORTONESE.1968. Distribution and ,ANDJ. F. MCKINNEY.1978. A revision of the systematics of the gobiid fish Odondebueniabalearica Indo-Pacific Fish genus Gobiopsiswith descriptions (Pellegrin Fage). Estr. Dag. Ann. Mus. St. Nat. Ge- of four new species (Pisces: Gobiidae). Ibid. 262:1- nova 77:342-359. 52. ,AND P. WONGRAT.1979. A new goby (Teleos- ,AND . 1979. Two new gobiid fishes of tei: Gobiidae) from the South China Sea and its PEZOLD-MONOPHYLETIC GOBIINAE 643

significancefor gobioid classification.Zool.J. Linn. Caffrogobius,Callogobius, Chromogobius, Coryphopterus, Cristatogobius, Cryp- Soc. 67:239-257. tocentroides,Cryptocentrus, Ctenogobiops, Didogobius, Discordipinna, Drom- bus, Fu- MURDY,E. 0. 1989. A taxonomic revision and cla- Egglestonichthys,Eviota, Exyrias, Favonigobius, Feia, Flabelligobius, sigobius, Garmannia, Gladiogobius,Glossogobius, Gobiodon, Gobiopsis, Gobius, distic analysisof the oxudercine gobies (Gobiidae: Gobiusculus, Gorogobius,Hazeus, Hetereleotris, Heteroplopomus,Istigobius, Oxudercinae). Rec. Austral. Mus. Supplement 11: Kelloggella, Lesueurigobius, Lophogobius, Lotilia, Luposicya, Lythrypnus, 1-93. Macrodontigobius,Mahidolia, Mangarinus, Mauligobius, Millerigobius,Mon- ishia, F. 1984. A revision of the fish Myersina,Nematogobius, Odondebuenia, Oplopomops, Oplopomus, Opua, PEZOLD, gobioid genus Parachaeturichthys,Paragobiodon, Paratrimma, Parkraemeria, Platygobiop- Gobionellus. Unpubl. Ph.D. diss. Univ. of Texas, sis, Pleurosicya, Pomatoschistus,Porogobius, Priolepis, Psammogobius,Psilo- Austin, Texas. gobius, Signigobius, Silhouettea, Stonogobiops,Sueviota, Tenacigobius, Thoro- SANZO,L. 1911. Distribuzione della cutanee gobius, Tomiyamichthys,Trimma, Trimmatom,Valenciennea, Vanderhorstia, papille Vanneaugobius, Yongeichthys,Zebrus, Zonogobius (organi ciatiforme) e suo valore sistematico nei Gobi. Mitt. zool. Stn. Neapel 20:249-328. SMITH, C. L., ANDJ. C. TYLER. 1977. Redescription Tribe Gobiosomini of the gobiid fish Coryphopteruslipernes Bohlke and Robins, with notes on its habits and relationships. Aruma, Barbulifer, Bollmannia, , Elacatinus, Eleotrica, Enyp- Amer. Mus. Novit. 2616:1-10. nias, Evermannichthys,Ginsburgellus, Gobiosoma,Gobulus, Gymneleotris,Mi- Nes, Pariah, G. crogobius, Ophiogobius,Palatogobius, Parrella, Psilotris, Pyc- SPRINGER, V. 1983. Tyson belos, new genus and nomma, Risor, Robinsichthys, species of western Pacific fish (Gobiidae, Xenisth- minae), with discussionsof gobioid osteology and classification.Smithsonian Contr. Zool. 390:1-40. APPENDIX2. GENERA . 1988. Rotuma lewisi, new genus and species ASSIGNED TO THE GOBIONELLINAE of fish from the southwest Pacific (Gobioidei, Xe- Members of the Astrabe and nisthmidae). Proc. Biol. Soc. Washington 101:530- Acanthogobius, Chasmichthysgroups, pro- 539. posed monophyletic by Birdsong et al. (1988), are listed separately. Aphia, Awaous, , Calamiana, Ctenogobius, Deltentosteus, TAKAGI,K. 1957. Descriptionsof some new gobioid Evorthodus,Gnatholepis, Gobioides, Gobionellus, , Hyrcanogobius, fishes of Japan, with a proposition on the sensory Knipowitschia, Lethrops, Mesogobius, Mistichthys,Mugilogobius, Neogobius, line as a taxonomiccharacter. Univ. Nesogobius, Oligolepis, Oxyurichthys,, ,Pseudogo- system J. Tokyo bius, Fish. 43:96-126. Redigobius, , Schismatogobius,Stenogobius, Stigmatogobius, Synechogobius,Tamanka, Tridentiger, Tukugobius,Vitraria, Vomerogobius . 1989. Cephalic sensory canal system of the gobioid fishes of Japan: comparative morphology with special reference to phylogenetic significance. AcanthogobiusGroup Ibid. 75:499-568. WINTERBOTTOM, R., ANDA. R. EMERY.1986. Review Acanthogobius,Amblychaeturichthys, Chaeturichthys, Lophiogobius, Ptero- gobius, Sagamia, Suruga of the gobioid fishes of the Chagos Archipelago, Central Indian Ocean. Royal Ont. Mus. Life Sci. Contr. 142:1-82. Astrabe Group

BIOLOGY DEPARTMENT AND MUSEUM OF Astrabe, , Eutaeniichthys,Leucopsarion, , Typhlogobius ZOOLOGY, NORTHEAST LOUISIANA UNIVER- SITY, MONROE, LOUISIANA 71209. Submitted ChasmichthysGroup 20 March 1992. Accepted 26 June 1992. Sec- tion editor: R. Winterbottom. ,Chasmichthys, Clevelandia, Eucyclogobius,, Ilvp- nus, Lepidogobius,Quietula

APPENDIX 1. GENERA INCLUDED IN THE GOBIINAE

Members of the Tribe Gobiosomini are listed separately. Acentrogobius,Amblyeleotris, Amblygobius, Amoya, Asterropteryx,Aulopa- reia, Austrolethops,Barbuligobius, Bathygobius, Bryaninops, Buenia, Cabillus,