Number of Pages: 3Integrative BiologyGeological Science

Highton, Richard

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AMPHIBIA: CAUDA TA: PLETHODONTIDAE402.1PLETHODON WEHRLEICatalogueof AmericanAmphibiansand Reptiles.HICHTON,RICHARD. 1987. Plethadanwehrlei.Plethodon wehrlei Fowler and DunnAllegheny Plateau woodland salamanderPlethadanglutinasusGreen: Cope, 1889:141 (part). Referencetospecimenswith red spots from caves in Montgomery County,Virginia probably representP. wehrlei.PlethadanwehrleiFowler and Dunn, 1917:23. Type.locality,"TwoLick Hills, Indiana County, Pennsylvania." Holotype,Academyof Natural Sciences of Philadelphia (ANSP) 19123, an adultfemale collected in September 1911 by R. W. Wehrle (ex.amined by author).Plethadandixi Pope and Fowler, 1949:1. Type-locality, "DixieCaverns, Roanoke County, Virginia." Holotype, Field Museumof Natural History (FMNH) 56510, an adult male collected11 July 1948, by John W. Funkhouser, Sarah H. Pope, Clif-ford H. Pope, Hallowell Pope and Whitney Pope (examinedbyauthor).PlethadanjacksaniNewman, 1954:9. Type-locality,"Trillium Vale,elevation 2100 feet, approximately one mile east of Blacks-burg, Montgomery County, Virginia." Holotype, National Mu·seum of Natural History (USNM) 134498, an adult male col-lectedon 11 February 1950, by Walter B. Newman(examinedby author).Plethadanwehrleiwehrlei:Conant, 1958:234.Plethadanwehrleidixi: Conant, 1958:234.• CONTENT. No subspeciesare recognized.• DEFINITION. Plethadanwehrleiis a memberof theP. wehrleigroup of easternPlethadanas definedmorphologicallyby Highton(1962) and biochemicallyby Highton and Larson (1979). Adultsrange from 45-70 mm (body) and 100-160 mm (total)length. P.wehrleihas much smaller dorsal white spots than the only otherspecies in the group, P. punctatus,except in newly discoveredTennessee,Kentucky and Summers County, West Virginia popu-lations of P. wehrleithat possesslarger yellow dorsal spots. In allbut the latter and the northern Pennsylvania and New York pop-ulations, there also may be dorsal red spots, particularly in the0 50100I,II,01 0,,MAP. The solid circle showsthe type locality. Hollow circles in-dicate other records.young.There is oftena smallamountof brassyfleckingon the graydorsumand there is lateralwhitespotting.The chin is usuallylighterthan the belly. The modal number of trunk vertebrae is 18 com-pared to 17 in all membersof the P. glutinasusgroup, the onlyother group in the east comprisinglarge speciesof Plethadan.TheP. wehrleigroup also has more webbingbetweenthe toes than thespeciesof the P. glutinasusgroup.• DESCRIPTIONS.Descriptions are in Fowler and Dunn (1917),Dunn (1926), Bishop(1927,1941,1943), Netting (1936 a, 1936b), Netting et al. (1946), Brooks (1945), Pope and Fowler (1949),Newman (1954), Highton (1962, 1971), Hoffman (1967), Conant(1975), Smith (1978), Behlerand King (1979), Martof et al. (1980)and Cupp and Towles (1983).• ILLUSTRATIONS.Black and whitephotographsof dorsal, lateraland ventral views of specimens from northern Pennsylvania andNew York are in Bishop (1941,1943) and the Maryland specimenin Thompson and Chapman (1978). Pennsylvania specimenswereillustrated by Harrison (1949, 1957) and Fegely (1973). Conant(1958, 1975) illustratedboth the northen and Dixie Caverns types,and Cupp and Towles (1983) the yellow-spottedwestern variant.Color photographsare in Behler and King (1979) and Martof et al.(1980). Drawingsof lateral and ventral aspectsof the holotypearein Fowler and Dunn (1917). Color drawings of both northern andsouthern(red-spotted)types are in Smith (1978). Drawings of ven-tral and dorsal viewsof the holotypeof P. jacksani are in Newman(1954). Drawings of the inside of the mouth are in Fowler andDunn (1917) and Bishop (1941) and the webbingof the toes inHighton (1962). Photographsof sectionsof the testis,vas deferens,cloacaand spermathecaare in Pauley and England (1969). A scan-ning electronmicrographof a maxillary tooth is in Coss (1974).• DISTRIBUTION.This speciesis known from Monroe and Wash-ington counties, Ohio, and Cattaraugus County, New York, southin the Allegheny Plateau through western Pennsylvania, westernMaryland and West Virginia to southwesternVirginia (where it alsooccurs in parts of the Valley and Ridge, Blue Ridge and Piedmontprovinces)to StokesCounty, North Carolina. The newly discoveredyellow-spottedmorph is known from Summers County, West Vir-ginia; Letcher County, Kentucky; and CampbellCounty, Tennessee.Erroneous records are Fowler and Dunn (1917), a paratype fromJuniata Co., Pennsylvania (Netting, 1936b); Lachner (1942) forMercer County, Pennsylvania; Netting et al. (1946) for RaleighCo., West Virginia; and Grobman (1949) for Nelson County, Vir-ginia. The record from Hancock County, in the northern panhandleof West Virginia, needsconfirmation.• FOSSILRECORD. None.• PERTINENTLITERATURE. Summariesof the known distributionof P. wehrleiin Ohio were given by Walker (1933) and Thomas(1951); in New York by Bishop(1927,1941) and Johnson (1961);in Pennsylvaniaby Highton (1971) and McCoy (1982); in Marylandby Thompson and Chapman (1978); in North Carolina, West Vir-ginia and Virginia by Hoffman (1967) and Highton (1971); in Ken-tucky by Cupp and Towles (1983); and in Tennesseeby Redmondand Jones (1985). Distributionalinteractionswith otherspecieswerediscussedby Grobman (1944) and Highton (1971). The relation-ships of P. wehrleiwere discussedby Fowler and Dunn (1917),Dunn (1926), Grobman(1944), Newman(1954) and Highton (1962,1971). Highton and Larson (1979) compared electrophoreticvari-ation in 29 proteinswith that in all 25 other speciesof Plethadonrecognizedat that time. Its albuminwas comparedimmunologicallyto severalother speciesof Plethadanin Maxson et al. (1979) andMaxson et al. (1984). Chromosomalmorphology,genomesize andDNA sequencehomologiesto easternand westernspeciesof Pleth-adanwerestudiedby Mizuno and Macgregor (1974). Larson (1984)reviewedpatternsof evolutionin the salamanderfamily Plethodon-tidae and made a number of comparisonsof P. wehrleito otherspecies.Wake (1981) discussedthe advisabilityof recognizingiso-latedallopatricspeciessuch as P. wehrleiand P. punctatuson thebasisof allozymedata.Wake (1963, 1966) includedone P. wehrleifrom Virginia inhis osteologicalstudiesof the family'Plethodontidae.Thurow (1968)studied morphometrics.Vertebral variation was given by Highton(1962, 1971). Coss (1974) did a comparative scanning electronmicroscopic study of maxillary and premaxillary dentition. Hilton(1951) studiedthe nasal gland. Dodd and Brodie (1976) described402.2the mentalgland. Habitat distributionwas given by Brooks (1945,1948), Bishop(1927,1941,1943), Netting (1936a, 1936b), Net-ting et a!. (1946), Newman (1954), and Pauley (1980). Netting(1933), Reese (1933, 1934), Jackson (1944), Fowler (1951) andCooper (1961) recorded its occurrence in caves in Virginia andWest Virginia. Life historydataare in Bishop(1927, 1941), Fowler(1951), Johnson (1961), Pauley and England (1969) and Hall andStafford(1972). Food recordsare in Bishop(1941), Johnson (1961),Hall (1976) and Pauley (1977). Pauley (1977, 1978a, 1978b,1978c, 1978d) studiedthe ecologicalinteractionof P. wehrleiandP. cinereusin West Virginia. Pauley (1980) studied its altitudinaldistributionin the mountainsof West Virginia. Thurow (1976) stud-ied interspecificaggression.• ETYMOLOGY. P. wehrleiis a patronym honoring the personwho was probably the first to collect the speciesin Pennsylvania,Richard W. Wehrle, a jeweler and naturalist from Indiana, Penn-sylvania.COMMENTDorsal red spottingis rare or absentin northern populationsof P. wehrlei, but in somesouthernpopulationsit is common andthis wasthebasisfor thedescriptionof P. jacksoniNewman(1954).Recently Cupp and Towles (1983) reported a new yellow spottedvariant from SummersCounty, West Virginia, and Letcher County,Kentucky. It has also been found in Campbell County, Tennesseeby Redmondand Jones (1985).The ranges of P. wehrleiand its close relative P. punctatuswere believed by Highton (1971) to be separated by 45 km ofuninhabited territory. P. wehrleihas recently been taken aboutmidwaybetweenthe closestrecordsof the two species.It shouldbesought at lower elevations on the western slopes of ShenandoahMountain, West Virginia to determineif the rangesof the two formscontact or overlap.I thank the National Science Foundation for financial supportof my researchon P. wehrlei(Grant DEB-81-17983).LITERATURECITEDBehler, John 1., and F. Wayne King. 1979. The AudubonSocietyfield guide to North American reptiles and amphibians.AlfredA. Knopf, New York. 720 p.Bishop, Sherman C. 1927. The amphibiansand reptiles of Alle-gany State Park. New York State Mus. Handb. 3:1-134.1941. The salamandersof New York. New York State Mus.Bull. 324:1-365.1943. Handbookof salamanders.ComstockPub. Co., Ithaca,New York, xiv + 555 p.Brooks, Maurice. 1945. Notes on amphibiansfrom Bickle's Knob,West Virginia. Copeia 1945(4):231.1948. Notes on the Cheat Mountain salamander. Ibid.1948(4):239-244.Conant, Roger. 1958. A field guide to reptilesand amphibiansofthe United States and Canada east of the 100th meridian.Houghton Mifflin Co., Boston. xv +366 p.1975. A field guideto reptilesand amphibiansof easternandcentral North America. Secondedition.Houghton Mifflin Co.,Boston. xviii +429 p.Cooper, John E. 1961. Cave records for the salamanderPleth-odonr. richmondiPope, with notes on additionalcave-asso-ciated species.Herpetologica 17(4):250-255.Cope, Edward D. 1889. The Batrachia of North America. Bull.U.s. Nat. Mus. (34):1-525.Coss, H. T. 1974. Maxillary and premaxillary dentition of sala-manders of the tribe Plethodontini (family Plethodontidae).Ph.D. Dissertation,ClemsonUniversity, ix +70 p.Cupp, Paul V., Jr., and Donald T. Towles. 1983. A new variantof Plethodonwehrleiin Kentucky and West Virginia. Trans.Kentucky Acad. Sci. 44:157-158.Dodd, C. Kenneth, Jr., and Edmund D. Brodie, Jr. 1976. Obser-vations on the mental hedonic gland-clusterof eastern sala-mandersof the genusPlethodon.ChesapeakeSci. 17(2):129-131.Dunn, Emmett Reid. 1926. The salamandersof the family Pleth-odontidae.Smith College Ann. Pub. viii +441 p.Fegely, Tom. 1973. The unseenamphibians.Part II. Pa. Angler42(8): inside back cover.Fowler, Henry W., and Emmett Reid Dunn. 1917. Notes onsalamanders.Proc. Acad. Natur. Sci. Philadelphia69:7-28.Fowler, James A. 1951. Preliminary observationson an aggre-gation of Plethodondixi. Herpetologica7(3):147-148.Grobman, Arnold B. 1944. The distribution of the salamandersof the genusPlethodonin easternUnited Statesand Canada.Ann. New York Acad. Sci. 45(7):261-316.1949. Some recent collections of Plethodonfrom Virginiawith the descriptionof a new form. Proc. Bio!' Soc. Washing-ton 62:135-142.Hall, Russell J. 1976. Summer foods of the salamander,Pietho-donwehrlei(Amphibia,Urodela, Plethodontidae).J. Herpeto!.10(2):129-131.- and Duane P. Stafford. 1972. Studies in the life history ofWehrle's salamander,Plethodonwehrlei.Herpetologica28(4):300-309 .Harrison, Hal H. 1949. Pennsylvania reptiles and amphibiansNo.8-salamanders. Pennsylvania Angler 18(11):24.1957. Pennsylvania reptiles and amphibians. PennsylvaniaFish Comm., Harrisburg, third edition, 25 p.Highton, Richard. 1962. Revision of North American salaman-ders of the genus Plethodon.Bull. Florida State Mus. 6(3):235-367.1971 (1972). DistributionalinteractionsamongeasternNorthAmerican salamandersof the genusPlethodon.In The distri-butional history of the biota of the southern Appalachians.Part III: Vertebrates:139-188. Res. Div. Mono. (4), VirginiaPolytechnic Inst., Blacksburg.and Allan Larson. 1979. The genetic relationships of thesalamandersof the genus Plethodon.Syst. Zoo!. 28(4):579-599.Hilton, William A. 1951. A nasalgland in plethodontidsalaman-ders. Copeia 1951(1):75-76.Hoffman, Richard 1. 1967. Distributionalrecordsfor threespeciesof Plethodonin Virginia. Radford Rev. 21(3):201-214.Jackson, Herbert W. 1944. A preliminary check list of the cavefauna of southwestVirginia. Bull. Nat. SpeleologicalSoc. 6:56-57.Johnson, Wilbert W. 1961. A life history study of Plethodonwehrleiin the high plateaus. Ph.D. dissertation,St. Bona-venture University, xi +91 p.Lachner, Ernest A. 1942. An aggregationof snakes and sala-mandersduring hibernation.Copeia 1942(4):262-263.Larson, Allan. 1984. Neontological inferences of evolutionarypattern and processin the salamanderfamily Plethodontidae.Evo!' Bio!' 17:119-217.Martof, Bernard S., William M. Palmer, Joseph R. Bailey, andJulian R. Harrison III. 1980. Amphibians and reptiles ofthe Carolinasand Virginia. University of North Carolina Press,Chapel Hill. 264 p.Maxson, Linda R., Richard Highton, and David B. Wake. 1979.Albumin evolution and its phylogenetic implications in theplethodontidsalamandergeneraPlethodonand Ensatina.Co-peia 1979(3):502-508.-, and David Ondrula. 1984. Immunological evidence ongeneticrelationshipsof Plethodondorsalis.J. Herpeto!. 18(3):341-344.McCoy, C. J. 1982. Amphibians and reptiles in Pennsylvania.Special Pub. Carnegie Mus. Nat. Hist. (6): 91 p.Mizuno, Shigeki, and Herbert C. Macgregor. 1974. Chromo-somes,DNA sequences,and evolution in salamandersof thegenusPlethodon.Chromosoma48:239-296.Netting, M. Graham. 1933. The amphibiansof West Virginia.West Virginia Wild!. 11:5-6, 15.1936a. Wehrle's salamander,PlethodonwehrleiFowler andDunn, in West Virginia. Proc. West Virginia Acad. Sci. 10:89-93.1936b. Wehrle's salamander,PlethodonwehrleiFowler andDunn, in Pennsylvania.Proc. PennsylvaniaAcad. Sci. 10:28-30.-, N. Bayard Green, and Neil D. Richmond. 1946. The occur-rence of Wehrle's salamander,PlethodonwehrleiFowler andDunn, in Virginia. Proc. Bio!' Soc. Washington 59:157-160.Newman, Walter B. 1954. A new plethodontidsalamanderfromsouthwesternVirginia. Herpetologica 1O(1):9-14.Pauley, Thomas K. 1977. Microdistribution of two sympatricspeciesof plethodontidsalamanders.Ph.D. Dissertation,WestVirginia University. ix + 110 p.1978a. Plants as indicators of occurence of two sympatricPlethodonspecies.Bull. Maryland Herpeto!. Soc. 14(1):29-35.1978b. Food types and distributionas a Plethodonhabitatpartitioningfactor. Ibid. 14(2):79-82.1978c. Moisture as a factor regulating habitat partitioningbetweentwo sympatricPlethodon(Amphibia, Urodela, Pleth-odontidae)species.1. Herpeto!. 12(4):491-493.1978d. Temperature and insolationas factors regulatingthepartitioning of habitats of two sympatric Plethodonspecies.Proc. W. Va. Acad. Sci. 50(2-4):77-84.1980. Field noteson thedistributionof terrestrialamphibiansand reptilesof the West Virginia mountainsabove975 meters.Proc. W. Va. Acad. Sci. 52(2):84-92.and Wayne H. England. 1969. Time of mating and eggdepositionin the salamanderPlethodonwehrleiFowler andDunn, in West Virginia Proc. West Virginia Acad. Sci. 41:156-160.Pope, Clifford H., and James A. Fowler. 1949. A new speciesofsalamander(Plethodon)from southwesternVirginia. Natur.Hist. Misc. (47):1-4.Redmond, William H., and Robert 1. Jones. 1985. Geographicdistribution.Plethodonwehrlei.Herpeto!. Rev. 16(1):31.Reese, Albert M. 1933. The fauna of West Virginia caves.Proc.W. Va. Acad. Sci. 7:39-53.1934. The fauna of West Virginia caves.Bull. Soc. de Cluj.Roumanie 7:571-573.Smith, Hobart M. 1978. A guide to the field identification.Am-phibians of North America. Golden Press, New York. 160 p.402.3Thomas, Edward C. 1951. Distribution of Ohio animals. Ohio 1.Sci.51(4):153-167.Thompson, Edward 1., and Joseph A. Chapman. 1978. The firstrecord of Wehrle's salamanderfrom Maryland. Proc. Penn-sylvania Acad. Sci. 52:103.Thurow, Gordon R. 1968. On thesmallblack Plethodonproblem.Western Illinois Univ. Ser. Bio!' Sci. (6):1-48.1976. Aggressionand competitionin easternPlethodon(Am-phibia, Urodela, Plethodontidae).J. Herpeto!. 10(4):277-291.Wake, David B. 1963. Comparativeosteologyof the plethodontidsalamandergenusAneides.J. Morpho!' Il3(1):77-Il8.1966. Comparativeosteologyand evolution of the lunglesssalamanders,family Plethodontidae.Mem. So. California Acad.Sci. 4: I-Ill.1981. The application of allozyme evidenceto problems inthe evolutionof morphology.Proc. Second International Con-gress of Syst. and Evo!' Bio!.:257-270.Walker, Charles. F. 1933. Somenew amphibianrecords for Ohio.Copeia 1933(4):224.RICHARD HIGHTON, UNIVERSITY OF MARYLAND, COLLEGE PARK,MARYLAND20742.Primary editor for this account, Jaime D. Villa.Published27 March 1987. Copyright 1987 by the SOCIETYFORTHESTUDYOF AMPHIBIANSANDREPTILES.

Plethodon wehrlei

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Plethodon wehrlei

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