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Phylogenetic Relationships and Systematic Revision of Central Texas Hemidactyliine Plethodontid Salamanders

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Phylogenetic Relationships and Systematic Revision of Central Texas Hemidactyliine Plethodontid Salamanders HerpetologicalMonographs, 14, 2000, 1-80 ? 2000 by The Herpetologists'League, Inc. PHYLOGENETIC RELATIONSHIPS AND SYSTEMATIC REVISION OF CENTRAL TEXAS HEMIDACTYLIINE PLETHODONTID SALAMANDERS PAUL T. CHIPPINDALE,15 ANDREW H. PRICE,2 JOHN J. WIENS,3 AND DAVID M. HILLIS4 'Dept. of Biology, The University of Texas at Arlington, Arlington, TX 76019, U.S.A. 2Texas Parks and Wildlife Dept., 4200 Smith School Road, Austin, TX 78744, U.S.A. 3Carnegie Museum of Natural History, 4400 Forbes Ave., Pittsburgh, PA 15213-4080, U.S.A. 4Section of Integrative Biology and Institute for Cellular and Molecular Biology, The University of Texas at Austin, Austin, TX 78712, U.S.A. ABSTRACT: Genetic variation and phylogenetic relationships of central Texas nontransforming spring and cave salamanders, genera Eurycea and Typhlomolge (Plethodontidae: Plethodontinae: Hemidactyliini), were examined using 25 allozyme loci and DNA sequence data for a maximum of 356 bp of the mitochondrial cytochrome b gene. Monophyly of the central Texas hemidactyliines is well supported. High levels of divergence occur among many populations and groups of populations, and there clearly are many more species in the group than previously recognized. Many have extremely restricted distributions in isolated islands of aquatic habitat. Several major monophyletic groups were identified that correspond to geographically circumscribed areas of the Edwards Plateau region. The deepest phylogenetic split in the group occurs between populations northeast versus southwest of the Colorado River. Species that have been assigned to the genus Typhlomolge are phylogenetically nested within the central Texas Eurycea; therefore, the genus Typhlomolge is placed in the synonymy of Eurycea. Continued recognition of the species E. latitans, E. nana, E. neotenes, E. pterophila, E. sosorum, E. tridentifera, and E. troglodytes is recommended, but E. neotenes appears to be restricted in range to a small geographic area, and is not widespread in the region as previously thought. The E. latitans and E. troglodytes species complexes are recognized; each con- sists of spring and cave populations that include those at the type localities of the latter two species, plus other populations to which they appear most closely related. Three new species from northeast of the Colorado River are described. Key words: Caudata; Plethodontidae; Eurycea; Typhlomolge; Eurycea chisholmensis new spe- cies; Eurycea naufragia new species; Eurycea tonkawae new species; Phylogeny; Speciation; Central Texas; Allozymes; Cytochrome b THE EDWARDS PLATEAU region of cen- other larval morphological features tral Texas is characterized by Cretaceous throughout their lives). Members of this limestones uplifted since at least mid-Ter- group exhibit a wide range of morpholo- tiary times, dissected and eroded to form gies that vary primarily according to numerous springs and caves (Sweet, whether they occupy surface or subterra- 1978a; Potter and Sweet, 1981; Abbot and nean habitats. Typhlomolge rathbuni, the Woodruff, 1986; Woodruff and Abbott, first member of the group to be described, 1986; and Veni, 1994 review the geologic was discovered after the drilling of an ar- history of the area). These habitat islands tesian well at San Marcos, Hays Co. in are inhabited by a variety of endemic 1895 (Stejneger, 1896). This large cave sal- aquatic organisms, many with extremely amander immediately captured the atten- restricted distributions. Predominant tion of the scientific community due to its among the aquatic vertebrate fauna of the bizarre morphology, including tiny non- region are plethodontid salamanders of the functional vestiges of eyes, loss of pigmen- genera Eurycea and Typhlomolge (tribe tation, long slender legs, and broad flat- almost all of which Hemidactyliini), are tened head; this species and its presumed perennibranchiate (i.e., retain gills and sister species T robusta exhibit some- of the most extreme cave-associated mor- 5 Author to whom correspondence should be ad- phologies known among vertebrates (Pot- dressed (email: [email protected]). ter and Sweet, 1981; Sweet, 1986). Rec- 2 HERPETOLOGICALMONOGRAPHS [Ala[No. 14 ognized as a plethodontid by Emerson ciated morphology (Mitchell and Reddell, (1905) and later placed in the tribe Hem- 1965; Mitchell and Smith, 1972; Sweet, idactyliini by Wake (1966), the relation- 1978a, 1984; Potter and Sweet, 1981). ships of Typhlomolge to other hemidacty- Based on biogeographic and geologic con- liines have remained controversial siderations, multiple invasions of subter- (Mitchell and Reddell, 1965; Mitchell and ranean habitat are likely to have occurred Smith, 1972; Potter and Sweet, 1981; (Mitchell and Smith, 1972; Sweet, 1978a, Lombard and Wake, 1986). For the re- 1982, 1984; Potter and Sweet, 1981), but mainder of this paper, we will use the ge- in at least some cases (especially in the ar- nus name Eurycea for these two species, eas inhabited by E. tridentifera, E. latitans, a taxonomic change proposed by Mitchell and perhaps E. troglodytes) there is the and Reddell (1965) and Mitchell and potential for subterranean gene flow Smith (1972). Molecular phylogenetic ev- among cave populations (Sweet, 1978a, idence presented here and by Chippindale 1984). (1995) supports this taxonomic shift. A key issue is whether the central Texas Additional central Texas hemidactyliines hemidactyliines are monophyletic. Wake were not recognized until 1937, when (1966) and Potter and Sweet (1981) sug- Bishop and Wright described Eurycea gested that members of what they consid- neotenes from a spring near Helotes in ered the genus Typhlomolge may be de- Bexar Co. In subsequent decades, several rived from an early Tertiary invasion of the more spring and cave species were de- area whereas Eurycea arrived later (per- scribed: E. nana Bishop 1941 from San haps in the late Miocene or Plio-Pleisto- Marcos Springs in Hays Co., E. latitans cene). According to this scenario, the ex- Smith and Potter 1946 from Cascade Cav- treme cave-associated morphologies of E. erns in Kendall Co., E. pterophila Burger, rathbuni and E. robusta (the two species Smith and Potter 1950 from Fern Bank formerly placed in Typhlomolge) may re- Springs in Hays Co., E. troglodytes Baker flect the long period of time spent under- 1957 from Valdina Farms Sinkhole in Me- ground. The range of (less extreme) cave- dina Co., E. tridentifera Mitchell and Red- associated morphologies seen in other dell 1965 from Honey Creek Cave in Co- central Texas subterranean Eurycea could mal Co., and E. sosorum Chippindale, then represent convergences with these Price and Hillis 1993 from Barton Springs species, varying in degree according to the in Travis Co. The status and relationships length of time spent underground and of these taxa have been problematic; most constrained by the putative common an- recently Sweet (1978a,b, 1984) questioned cestry of members of the Edwards Plateau the status of E. pterophila (which he syn- Eurycea group exclusive of E. rathbuni onymized under E. neotenes) as well as E. and E. robusta. Potter and Sweet (1981) latitans and E. troglodytes, which he con- found that the taxa they considered Ty- sidered hybrids between other species. phlomolge and some central Texas cave With the exception of Bogart's (1967) Eurycea exhibit the same general evolu- chromosomal studies, all inferences of re- tionary trends in head form (broadening lationships among the central Texas hem- and flattening of the skull). However, Pot- idactyliines have been based on morphol- ter and Sweet (1981) demonstrated that ogy, and no truly phylogenetic analysis of the osteological basis of these changes dif- the group has been attempted. Morpho- fers in the two groups. Given this result, logical variation in the group has proven Potter and Sweet (1981) recommended confusing; many surface dwellers from that the genus Typhlomolge continue to be throughout the region appear similar to recognized, a reasonable move given the one another based on external morphology information then available. (e.g., Mitchell and Smith, 1972; Hamilton, Extreme cave-associated morphologies 1973; Sweet, 1978a, 1982; Chippindale et are seen in some other non-Texan hemi- al., 1993), while subterranean dwellers dis- dactyliine plethodontids, including Typhlo- play a spectrum of degrees of cave-asso- triton spelaeus from the Ozark region and 2000] HERPETOLOGICAL MONOGRAPHS 3 Haideotriton wallacei from northern Flor- tyliine genera that are suspected to be ida and Georgia. We included these taxa closely related to or nested within Eury- in the present study; both are thought to cea. We based the choice of outgroup have close affinities to Eurycea and/or members on published morphological what has been considered Typhlomolge work (Wake, 1966; Sweet, 1977b; Lom- (Wake, 1966; Lombard and Wake, 1986). bard and Wake, 1986) and molecular data We also included representatives of all from a study in progress of higher-level species groups currently recognized within hemidactyliine relationships (Chippindale, Eurycea. Use of multiple outgroups is par- unpublished). The outgroup consisted of ticularly important because hemidactyliine E. bislineata (Renfrew Co. Ont.), E. wild- relationships are poorly understood, and erae (Watauga Co. NC), E. quadridigitata the closest relatives of the central Texas (yellow-bellied form, Tyler Co. TX), E. group are uncertain (but see Sweet, 1977b quadridigitata (silver-bellied form, for additional discussion). Charleston Co. SC), E. 1. longicauda (Bal-
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