Herpetologica, 49(2), 1993, 229-237 0 1993 by The Herpetologists’ League, Inc
PHYLOGENETIC AND TAXONOMIC ISSUES RELATING TO SALAMANDERS OF THE FAMILY PLETHODONTIDAE
DAVIDB. WAKE Museum of Vertebrate Zoology and Department of Integrative Biology, University of California, Berkeley, CA 94720, USA
THE occasion of the third decennial spects, but he chose not to recognize any Conference on the Biology of Plethodontid taxa between genus and family. Dunn en- Salamanders, and the first publication of visioned two “main groups” of genera the proceedings of the conference, is a pro- along the lines of Cope’s groups-a Pleth- pitious time to take stock concerning di- odon group with attached tongues and a verse phylogenetic and taxonomic issues. Eurycea group with free tongues, “con- As background, the monograph of Cope nected by three intermediate genera which (1889)is used as a point of departure. Cope hardly belong to either group” (Dunn, recognized a family Desmognathidae (for 1926:22)-Stereochilus, Typhlotriton, and Desmognathus) and a family Thoriidae Typhlomolge. Apart from the recognition (for Thorius),both distinguished from the of many more species and a few novel new Plethodontidae by having opisthocoelous genera, the largest difference between the vertebrae. Within the Plethodontidae he taxonomy of today and that of Dunn is his recognized two groups of genera: Pletho- treatment of tropical salamanders. He rec- dontae [Plethodon, Hemidactylium, Bat- ognized only 31 tropical species (about 44% rachoseps, Stereochilus, and Autodax (= of the total number of species of pletho- Aneides)], and Spelerpes [Geotriton (= dontids; tropical species constitute more Hydromantes), G yrinophilus, Manculus than 65% today), all placed in Oedipus. (now included in Eurycea),Spelerpes (Eu- While workers such as Noble (1927, rycea and Pseudotriton), Oedipina, and 1931) quibbled with some of Dunn’s ideas, Oedipus (supergenus Rolitoglossa, minus the monograph remained authoritative for Oedipina and Thorius)].The fundamental several decades. Taylor (1944) described distinction was that the Plethodontae ha5 many tropical species and sorted them into a tongue attached anteriorly, while Spe- a number of genera. Several new genera lerpes has a free tongue. Cope thought that (e.g., Phaeognathus, Haideotriton) and “The generic relationships of the above- many new species were named in North named groups are exceedingly simple, and America as well, but it was not until my the ease with which the animals can be comparative osteological study (Wake, analyzed renders the case free from the 1966) that there was a major change in doubts which constantly arise in discus- taxonomy and phylogenetic perspective. sions of generic relationships as to the That work was published on the eve of the probable omission of characters from the cladistic revolution, and while most taxa argument” (Cope, 1889:121-1 22). are monophyletic and based on shared de- The famous monograph of Dunn (1926) rived character states, there are some in- remains useful today. Dunn documented consistencies with respect to modern cla- the unique features of the family, dis- distic methodology (e.g., in the brief cussed relationships to other families, and discussion of familial relationships) The included a lengthy treatment of relation- main results of that study have remained ships of species within some of the genera surprisingly robust and find wide accep- (e.g., Desmognathus), and of the genera tance to this day. to each other. Only 16 genera and 72 spe- Wake (1966)thought that plethodontids cies were recognized. Dunn noted that were derived from an ambystomatid an- Desmognathus and Leurognathus dif- cestral stock (in 1966 the Ambystomatidae fered from the other genera in many re- included the three subfamilies Ambysto- 229 230 HERPETOLOGICA [Vol. 49, No. 2 matinae, Dicamptodontinae, and Rhyaco- with the hyobranchial apparatus and the tritoninae, all currently recognized as fam- nervous system, and there are unique fea- ilies), that the genera of plethodontids could tures of courtship. Recently Larson and be placed in two subfamilies, Desmogna- Wilson (1989) and Larson (1991) have pro- thinae and Plethodontinae, both with an- vided characters from rRNA sequences that cestral and derived characters, and that further support the monophyly of the fam- the Plethodontinae could be segregated ily, and Sever (1991) has presented some into three tribes, the Hemidactyliini, the characters from the morphology of the clo- Plethodontini, and the Bolitoglossini. A acal region. major departure from prior work was the When I argued for a phylogenetic re- grouping of Hydromantes, Batrachoseps, lationship between the plethodontids and and all of the tropical salamanders (su- the ambystomatids (Wake, 1966), I had pergenus Bolitoglossa) as a monophyletic Rhyacotriton very much in mind. With tribe Bolitoglossini. Another novel feature the breakup of the Ambystomatidae, the was the placement of Hemidactylium with possibility of a sister taxon relationship with the Eurycea group of genera of Dunn the Rhyacotritonidae (fide Good and Wake, (1926). I envisioned the Hemidactyliini as 1992) must be seriously considered. Larson the central evolving stock, giving rise first and Wilson (1989) and Larson (1991) have to the desmognathines, next to the boli- shown that plethodontids are very distinct toglossines, then to the plethodontines, and from other families, and occupy a rather finally to Hemidactylium on the one hand basal position. Rhyacotritionids also are and the remaining hemidactyliines on the rather basal, and remote from both am- other, but I was uncertain on the place- bystomatids and dicamptodontids (Good ment of Hemidactylium, which ended up and Wake, 1992; Larson, 1991; Sever, 1991, with the Eurycea group mainly by default. 1992). The rRNA sequence data have sup- 1 use the 1966 taxonomy and phylogenetic ported some traditional groupings (notably interpretation as a point of departure for the monophyly of the Hynobiidae + Cryp- my re-evaluation. tobranchidae), while they have challenged others (the widely accepted grouping of FAMILYPLETHODONTIDAE the Plethodontidae with the Ambystomati- There has been no proposal to include dae, or alternatively with the Salamandri- plethodontids as members of any other dae). Further resolution of relationships of currently recognized family of salaman- the families of salamanders is likely to come ders since the admission of Typhlomolge with additional sequence data, and by to the Plethodontidae (Fowler and Dunn, combining sequence data with traditional 1917). Dunn (1926) showed that Typhlo- characters (such work is in progress, Lar- molge was a plethodontid; his conclusions son and Dimmick, personal communica- were foreshadowed by Emerson’s (1905) tion). demonstration that the genus was not a proteid. There have been no shifts of gen- SUBFAMILYDESMOGNATHINAE era from other families into the Pletho- This is a well supported, monophyletic dontidae since that time. While Soler (1950) group (Schwenk and Wake, 1993; Soler, proposed recognition of a family Desmog- 1950; Wake, 1966). Phaeognathus has nathidae, he acknowledged close relation- many autapomorphies, and it seems to be ship with the Plethodontidae. a basal derivative within the subfamily. All The Plethodontidae is well supported by remaining species fit well within Desmog- character data-lunglessness is universal, nathus, although Leurognathus, which all metamorphosed individuals have a na- contains a single species with a few autapo- solabial groove, only adult plethodontids morphies, is universally recognized. This lack an ossified pterygoid bone, pletho- may well render Desmognathus paraphy- dontids have a unique arrangement of vo- letic. This problem may be solved by DNA merine and postvomerine teeth, there are sequence data (Titus, 1992).A special phy- a number of unique features associated logenetic puzzle of the subfamily is the June 19931 HERPETOLOGICA 23 1 number of times that direct development called global) heterochrony could affect has evolved. Direct development is found many characters at once, leading to an in- in Phaeognathus, D. aeneus, and D. flated impression of the extent of homo- wrighti. It seems unlikely that aquatic lar- plasy, but so far it has not proven possible vae have re-evolved in the subfamily (for to sort coevolving complexes of characters one thing, a more ancestral hyobranchial that might be unconnected functionally system is present in desmognathine larvae from independently evolving characters. I than in the remaining plethodontids: Wake, personally like to use new data sets to test 1966). However, Phaeognathus is a sister hypotheses based on old data, and so I have taxon of Desmognathus, and if one or both long advocated the use of molecular and of the species of Desmognathus that have other kinds of data. Linda Maxson and I direct development should prove to be bas- started a collaboration with the goal of test- al within the genus, that possibility must ing alternative morphologically based be considered. phylogenetic hypotheses, but the taxa are too differentiated for microcomplement SUBFAMILYPLETHODONTINAE fixation of albumin to be effective. Protein Wake (1966) recognized three tribes of electrophoresis is useful for comparing genera: Plethodontini (for Plethodon, closely related species and even genera, Aneides, and Ensatina),Bolitoglossini (for but the higher taxa are beyond the limit Bolitoglossa, Chiropterotriton, Lineatri- of effectiveness of the technique. On the ton, Oedipina, Parvimolge, Pseudoeury- other hand, the taxa are not sufficiently cea, Thorius, Batrachoseps, and Hydro- distinct to be in the range of resolution mantes), and a catch-all Hemidactyliini using the ribosomal RNA sequences stud- (all other genera). The Plethodontini and ied to date (Larson and Wilson, 1989). I Hemidactyliini were treated as sister taxa, am confident that in time molecules with and this group was the sister taxon of the the appropriate rates of molecular evolu- Bolitoglossini. While there has been gen- tion will be identified, and that compar- eral acceptance of these three groups, there ative sequence analysis will contribute pos- has been a low level of debate about the itively to our understanding of cladistics reality of the Hemidactyliini and about in plethodontids. Episodes of reciprocal il- which two of these three groups are sister lumination obtained by testing phyloge- taxa. Lombard and Wake (1986) accepted netic hypotheses with different data sets the three tribes and the desmognathines as will lead to deeper understanding of the four primary taxa and analyzed relation- nature and degree of independence of ships of them, concluding that Desmog- characters, and at that point a total evi- nathinae was basal and that Bolitoglossini dence approach to understanding phylo- and Plethodontini were terminal sister taxa. genetics of the family will be appropriate. Presch (1989) objected to using these four taxa as OTU’s. He attempted a reanalysis TRIBEHEMIDACTYLIINI of the data set, but recorded some char- There is reason to question if this is a acters in ways that I find to be unaccept- monophyletic taxon. The problem is with able. Presch showed that no hypothesis of Hemidactylium, which does not fit com- relationships is robust, but Lombard and fortably with the other genera, as Wake Wake had made the same point. New neu- (1966) acknowledged. Historically, Hemi- rological evidence adds support for the sis- dactylium was considered to be a close ter group relationship of the Plethodontini relative of Plethodon (Dunn, 1926). The and Bolitoglossini (Wake et al., 1987). genus was grouped with the other hemi- Extensive morphological homoplasy in dactyliines almost by default, for it lacked plethodontids makes it likely that molec- the derived direct development of the Bo- ular characters will be needed to resolve litoglossini and Plethodontini, it lacked the the relationships of higher taxa. Paedo- derived morphological traits of the Des- morphosis is a factor to be taken seriously mognathinae, and I was reluctant to es- in the family, and organismal-wide (so- tablish a fourth tribe that included only a 232 HERPETOLOGICA [Vol. 49, No. 2 single monotypic genus. However, subse- menclature are in a state of transition, so quently Wake and Lombard (1972) did the case is not so simple as Dubois (1984b) suggest that Hemidactylium might best be implies. Recently an appeal has been made placed in its own tribe, but failed to make to the International Commision on Zoo- this move in their later analysis (Lombard logical Nomenclature to suppress the name and Wake, 1986). The remaining genera Mycetoglossina and to conserve the Hemi- of the Hemidactylini appear to form a dactyliini (Smith and Wake, 1993b), and monophyletic group. I recommend maintaining the traditional Stereochilus, Gyrinophilus, Pseudotri- taxonomy until the matter receives formal ton, and Hemidactylium are relatively action. noncontroversial as far as their generic sta- tus is concerned, although there have been TRIBEPLETHODONTINI attempts to combine Gyrinophius and This grouping of three genera is sup- Pseudotriton (Grobman, 1959; countered ported by a variety of morphological and by Martof and Rose, 1962). However, the molecular evidence (Jackman, this confer- remaining genera are difficult. I included ence; Larson et al., 1981; Wake, 1963, Manculus quadridigitata in Eurycea, as 1966), and Ensatina is acknowledged as had Dunn (1926), but argued for separate the sister taxon of Plethodon + Aneides. generic status for Typhlotriton, Haideo- Aneides is a monophyletic group, but mo- triton, and Typhlomolge. The last genus, lecular evidence (Jackman, in progress, in particular, has been controversial (e.g., Larson et al., 1981) increasingly points to Mitchell and Reddell, 1965; Mitchell and Plethodon as being paraphyletic, with Smith, 1972; Potter and Sweet, 1981).The western members of the genus having a two species currently assigned to Typh- sister group relationship to Aneides. One lomolge differ osteologically from peren- solution is to place all species in the tribe nibranchiate species assigned to Eurycea, in a single genus Plethodon, because En- but it may be that these traits have been satina includes only a single species and derived within the framework of a mono- is apparently the sister taxon of the others, phyletic group of species, and if so all of but such a genus would be large and in- these may eventually be combined in a convenient. An alternative is to place just single genus. Molecular studies in progress the species of Plethodon and Aneides in by Chippindale and Hillis (presented at a single genus Plethodon, but such a taxon this conference) suggest that Typhlotriton would still be large Another alternative and Naideotriton also require renewed at- would be to expand Aneides to include tention, and they, too, might fall within western Plethodon, but I know of no mor- the framework of an expanded Eurycea. phological character evidence for such a Dubois (19846) raised a point relating grouping. A final alternative is to name a to taxonomic priority. He argued that an new genus for the western species of Pleth- older and more appropriate name for the odon, but I know of only weak morpho- tribe Hemidactyliini is the Mycetoglossini, logical or molecular character evidence for based on the argument that Bonaparte such a move. Ideally, while I would like (1839) had used the name M ycetoglossa to see genera to be monophyletic, I would as a substitute name (invalid) for Pseu- also like them to be diagnosed by mor- dofriton, and had later used the term My- phological characters, because I think that cetoglossina as a subfamilial category a major goal of taxonomy is convenience. (Bonaparte, 1850).Dunn (1926) and other I would like to comment briefly on new- workers ignored this family-group name, ly discovered information concerning the recognizing the priority of the name Pleth- curious name Ensatina eschscholtzii pla- odontidae (Gray, 1850), listed by Bona- tensis. The status of this taxon, which is parte (1850) as a synonym of his Myce- based on a single specimen purportedly toglossina. The name Hemidactyliini collected near Montevideo, Uruguay (Ji- (based on Hallowell, 1856) has become well mhez de la Espada, 1875) (and hence the established, and the rules on zoological no- name, derived from Rio de la Plata), has June 19931 HERPETOLOGICA 233 been considered by Dunn (1926) and My- sister group relationship of Batrachoseps ers and de Carvalho (1945). The latter au- and Bolitoglossa, with Hydromantes be- thors suggested that a specimen from the ing an earlier derivative (Lombard and Sierra Nevada of California had been car- Wake, 1986), and this hypothesis is fa- ried by a miner back to Uruguay, where vored by cytological evidence as well (Ses- it eventually reached a traveling compan- sions and Kezer, 1991). The problem with ion of Jimknez de la Espada, who in turn this hypothesis is that it requires that com- first noticed the specimen as he was pack- pletely free tongues have evolved twice ing his third shipment in Chile for trans- within the Bolitoglossini, assuming that the portation to Spain. Savage (1978) present- genioglossal muscle of Batrachoseps has ed a brief summary of the “Comisibn not reappeared, which seems highly un- Cientifica del Pacifico”, 1862-1865. He likely (Lombard and Wake, 1977; Roth stated that Jimknez de la Espada visited and Wake, 1985). However, the prevailing the “countries of Central America” on two hypothesis of phylogenetic relationships of different occasions. He also noted that a the plethodontid genera already requires zoologist on the expedition, Fernando that fully free tongues have evolved in- Amor, died in December 1863, in San dependently in the Eurycea group, so this Francisco, California, of an illness con- homoplasy is not so unexpected as it might tracted in the Atacama Desert. Savage did seem (Lombard and Wake, 1986; Wake, not mention that at least one ship from the 1991). expedition went as far north as San Fran- The genus Hydromantes has many aut- cisco while the “Comisih” was in prog- apomorphies (Lombard and Wake, 1977, ress. This fact has been vividly illustrated 1986; Wake, 1966). Lanza and Vanni by the recent publication of a remarkable (1981) separated the European and Amer- set of photographs by Rafael Castro Or- ican species of Hydromantes into two for- dbiiez (Calatayud Arinero and Puig-Sam- mal taxa, using the name Hydromantoides per, 1992), a photographer with the “Co- for the American species. Wake (1966) mision”. What particularly struck me were presented osteological evidence that the three plates (75, 85, 86) taken in present- European and American species could be day Calaveras Big Trees State Park, in the distinguished, and Wake et al. (1978) central Sierra Nevada of California, where showed that the two groups differed in I have been conducting a field study of biochemical characters as well. Conse- Ensatina eschscholtzii platensis for the quently, the only debate is whether it is past eight years. The salamanders are useful to separate a clearly monophyletic abundant in this area, and I suggest that genus into two genera. I find the two groups it was this previously unrecorded visit by to be very similar in terms of morphology, members of the “Comisibn” that accounts ecology, and behavior, and they share an for the specimen that was mislabeled as absolutely unique food-capturing system being from Uruguay and that ultimately (Lombard and Wake, 1977).Furthermore, was sent to Madrid. if one compares degree of molecular di- vergence and accepts a general molecular TRIBEBOLITOGLOSSINI evolutionary clock, the two groups of Hy- I know of no challenges to the mono- dromantes are about as different from each phyly of this taxon, which includes about other (Wake et al., 1978) as are the species one-half of all the species of salamanders of Aneides from each other (Larson et al., of all families. Currently there are three 1981), and in general the degree of genetic recognized supergenera-Hydromantes, differentiation relative to that in other in Europe and western North America, plethodontid genera is low (cf. Larson, Batrachoseps, in western North America, 1984). So this is very much a matter of and Bolitoglossa, which occurs in mainly personal choice, and I choose to recognize tropical America from northeastern Mex- a single genus. A new twist was added ico to Brazil, Bolivia and Peru. Morpho- when Dubois (1984~)published one of a logical evidence favors the hypothesis of a series of papers on the nomenclature of 234 HERPETOLOGICA [Vol. 49, No. 2 amphibians. Dunn (1923) argued that arguments dating back 90 years earlier. Geotriton was preoccupied by a salaman- We have had stability for 70 years, and drid, and he assumed that Hydromantes pending a formal decision on a recent ap- was the next available name. This is a very peal to the International Commission on complicated story, discussed in detail by Zoological Nomenclature (Smith and Dubois, who claims that Dunn erred, on Wake, 1993a), I recommend continued use technical grounds, when he selected the of Hydromantes for the European and name Hydromantes. The name is credited American species traditionally associated to Gistel (1848), who apparently thought with this name. Batrachoseps is another that he was applying the name to the spe- problematic genus that might well be sep- cies known today as Hydromantes itali- arated into subgenera or genera. The two CUS. According to Dubois, the name Hy- major groups in the genus are differenti- dromantes was a substitute name for ated by many biochemical characters, but Geotriton, a name first used by Bonaparte the morphological data are not easily in- (1831) (but lacking a diagnosis or a list of terpreted (Wake, 1989). The name Pleth- included species, it is a nomen nudum). In opsis is available for Batrachoseps wrighti 1832, Bonaparte again used the name and and its relatives (B. campi and an unde- associated it only with a single species, Sal- scribed species). However, the monophyly amandra exigua Laurenti, 1768. This tax- of Batrachoseps, as currently recognized, on is now considered to be a synonym of is unquestioned, so the only reason to di- Triturus vulgaris. Dubois claims that this vide the genus would be for convenience species must be considered to be the type (the genus will soon be larger; there are species of Geotriton, even though Bona- several undescribed species: Wake and co- parte (1837) clearly applied the name to workers, in preparation). a species that is included in present-day There are old records of Batrachoseps Hydromantes. On narrow, technical from Alaska (Cope, 1889; Dunn, 1926) and grounds, which I do not have the space to from Nevado de Colima, Jalisco, Mexico present here, Dubois argued that Hydro- (Gadow, 1905). I have examined all of the mantes was invalid for plethodontid sal- specimens involved and agree with pre- amanders and that the appropriate name vious authors (Hendrickson, 1953; Steb- for the genus was the new name of Lanza bins, 1951; Stebbins and Lowe, 1949) that and Vanni (1981), Hydromantoides. Du- there are reasons to question the records bois briefly considered the biological evi- from Alaska. Ron Crombie (in litt.) has dence and concluded that it might be ap- qtudied letters from Lt. Nichols, who sent propriate to recognize subgenera for the the Alaskan specimen to the National Mu- European and American species. Because seum. The ship “Hassler” moved between Lanza and Vanni had designated Spe- southeastern Alaska and Mare Island (Val- lerpes platycephalous Camp, 1916, as the lejo, San Francisco Bay Region), Califor- type species of Nydromantoides, Dubois nia. Much of Nichols’ material was from proposed a new subgeneric name for the Alaska, but perhaps he inadvertently mixed European species, Hydromantoides (Spe- a specimen from Mare Island or vicinity, leomantes), with Hydromantes italicus where Batrachoseps is abundant, with his Dunn, 1923, as the type species. Thus, if Alaskan material. I think it unlikely that one accepts the nomenclatural argument Batrachoseps is in Alaska, especially be- of Dubois and the taxonomic argument of cause species with similar morphology are Lanza and Vanni, the correct generic name not known to occur north of extreme for the California Hydromantes is Hydro- southeastern Oregon. mantoides, and the correct generic name The Mexican specimen, a juvenile, is less for the European Hydromantes is Speleo- readily dismissed. Gadow (1908) recount- mantes. I reject both arguments. Hydro- ed in detail the circumstances of capture. mantes has been used almost universally Stebbins (1951), who examined the spec- in a large literature since 1923, and I see imen, considered it to be correctly assigned no useful purpose in dredging up technical to Batrachoseps. However, I suspect that June 19931 HERPETOLOGICA 235
it is a member of the supergenus Bolito- Museum Godeff roy, associated with an an- glossa, some members of which occasion- imal dealer in Hamburg).However, I have ally have four rather than five toes (Wake, examined the type specimen and on the 1991), because the specimen appears to basis of its having a subocular groove that have a free tongue (it is possible that the intercepts the lip, an autapomorphy of specimen has been damaged, however). Thorius, I assign it to that genus, possibly The supergenus Bolitoglossa has been as a synonym of T. narisovalis of Oaxaca, very difficult to analyze phylogenetically which has nostrils of similar size and shape. because the group as a whole is highly derived morphologically, and there are CONCLUSIONS relatively few characters (Wake and Elias, Many opportunities exist for developing 1983). Some large, well defined genera and testing phylogenetic hypotheses for (Thorius and Oedipina) have been widely plethodontid salamanders. Robust phylo- but not universally recognized for over 100 genetic hypotheses have been difficult to years. Homoplasy is so great in the super- obtain, mainly because of the extent of genus that Dunn (1926) reacted by simply morphological homoplasy. As we come to recognizing a single genus (Oedipus, later understand the nature of homoplasy and shown to be preoccupied by an insect). the biological basis for homoplasy in par- Taylor (1944) accomplished a significant ticular characters (Wake, 1991; Wake and advance by breaking this genus into seven, Larson, 1987), I expect more progress in and Tanner (1950) added an eighth. Wake solving the phylogenetic problems that I and Brame (1963) showed that Taylor’s have outlined. Cytological (e.g., Sessions genus Magnadigita was not valid, and Eli- and Kezer, 1991) and molecular (e.g., Lar- as and Wake (1983), Wake and Elias son, 1991) data not only will add to the (1983), and Wake and Johnson (1989) add- data base, but should also help to sort ho- ed four genera, for a present-day total of moplasy from synapomorphy, and help US eleven. Major problems remain, for five of to determine how independent the cur- these (Bradytriton, Ixalotriton, Lineatri- rently recognized morphological charac- ton, Nyctanolis, and Parvimolge) are ters are from one another. In another de- monotypic, one (Pseudoeurycea) is sus- cade, I hope that we will have achieved pected to be paraphyletic, and one (Boli- robust phylogenetic hypotheses for the toglossa) is very large (containing about a genera and suprageneric taxa, and a stable, quarter of all species of salamanders). phylogenetic taxonomy. There has been recent progress in devel- Acknowledgments. --I thank P. Alberch, R. Crom- oping Phylogenetic hypotheses for the bie, E. Jockusch, and G. Peters for information and group (Sessions and Kezer, 1991; Wake and assistance, and P. Chippindale, A. Larson, S. Marks, Elias, 1983; Wake and Johnson, 1989), but H. Smith, and M. Wake for discussion and comments the effort is hampered by the vast amount on the manuscript. My research has been supported of homoplasy and the fact that many of by NSF (BSR 8619360). the species are poorly known. Doubtless LITERATURECITED molecular data will help in testing alter- BONAPARTE,C. L. 1831. Saggio d’una distribuzione native hypotheses based on morphological metodica degli animali vertebrati a sangue freddo. and karyological data, and such work is Giorn. Arcad. 52:129-209. underway in my laboratory. . 1832. Triton exiguus. Triton picciolino. In There is a worrisome old record of a Iconografia della fauna italica per le quattro classi tropical salamander, Spelerpes infuscatus degli animali vertebrati, tumi 11, fasc. I:243-244, pl. 83. Salviucci, Rome. (Peters, 1879), from Haiti. Dunn (1924) , 1837. Geotriton fuscus. Geotritone del Savi. did not examine the unique holotype, but In Iconografia della fauna italica per le quattro he nonetheless considered the taxon to be classi degli animali vertebrati, tome 11, fax. XIX: a synonym of present-day Lineatriton lin- 255-256, pl. 84. . 1839. Euproctus platycephalus. Euprotto eola. I agree with Dunn in considering the del Rusconi. In Iconografia della fauna italica per locality to be in error (the specimen, in the le quattro classi degli animali vertebrati, tome 11, Berlin Museum, was obtained from the fasc. XXVI:60-68, pl. 85. 236 HERPETOLOGICA [Vol. 49, No. 2
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