A New Species of Mud Turtle, Genus Kinosternon

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    Society for the Study of Amphibians and Reptiles

    A New Species of Mud Turtle, Genus Kinosternon, from Oaxaca, MexicoAuthor(s): James F. Berry and John B. IversonSource: Journal of Herpetology, Vol. 14, No. 4 (Oct. 31, 1980), pp. 313-320Published by: Society for the Study of Amphibians and ReptilesStable URL: http://www.jstor.org/stable/1563684 .

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    JOURNAL O F HERPETOLOGYVolume 14 31 October 1980 Number4

    A New Species of MudTurtle,Genus Kinosternon,fromOaxaca,MexicoJames F. Berry1Departmentf Biology,Universityf Utah,Salt LakeCity,Utah84112

    John B. Iverson2,3Department f Natural cience, Florida tate Museum,Gainesville,Florida 2611

    ABSTRACT-The relationshipsbetween Kinosteron scorpioides and K. integrumare dis-cussed. A new allopatric pecies, Kinosternonaxacae, is described romcoastalOaxaca,Mexico.K.oaxacae is a memberof the scorpioides species complex(withntegrum), utdiffers rom hesetwo species in its reducedplastron,narrower arapace,and axillary nd inguinal cutes in broadcontact. A dichotomouskey forthe identificationf the adultKinosteron of Oaxacais presented.

    INTRODUCTIONSystematic relationshipswithin he mud turtles(genus Kinosternon)f Mexicoare notoriouslyconfused. Several recent contributionsBerryand Legler1980; Conant and Berry1978; Iverson1978, 1979; and Iverson and Berry 1979) have clarifiedmany of the taxonomic problemsassociated withthe genus in northernMexico,butrelationships mongthe taxa insouthernMexicoremainin disorder.Particular isagreementsurrounds he systematicrelationships etween KinosternonntegrumLeconte and K.scorpioides Linnaeus(K.cruentatumDumeril nd Bibronn Dumeril nd Dumeril1851 has been relegatedto a subspecies of K.scorpioides by Legler1965 and Iverson1976).Siebenrock (1906, 1907) was first to consider integrum'a subspecies of K. scorpioides, an

    arrangementfollowedsubsequently by Ahl (1934), Wermuthand Mertens(1961, 1977), CasasAndreu (1967), and Morafka 1977). Mostotherauthors(e.g. Smith and Taylor1950; Duellman1961; Casas Andreu1965;Hardy nd McDiarmid969;Pritchard979;andSmithand Smith1980)have consideredintegruma species distinct rom corpioides.Pritchard1967)referred o integrumas a subspecies of scorpioides in his text (p. 40), but as a distinct pecies in his index(p. 285).We have examined the relationshipsbetween Kinosternonntegrumand K. scorpioides indetail. The two taxa are knownto occupythe same riverbasins only in the Rio Panucoand RioPapaloapan basins in eastern Mexico.Although hey are nowhere knownto occur at the samelocality,we findthe lackof evidenceof intermediacyintergradation)nd the presenceof numerousmorphologicaldifferencesbetweenthe taxa to be ampleevidence thatKinosternonntegrum s a1Present address:Department f Biology,Elmhurst ollege,Elmhurst,llinois 0126.2Present address:Department f Biology,EarlhamCollege,Richmond,ndiana 7374.30rder of authorshipwas determinedby the toss of a coin.1980 JOURNAL OF HtnrtlOLOGY 14(4):313-320 313

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    JAMESF. BERRYANDJOHNB. IVERSONspecies closely related to but specificallydistinct romK. scorpioides. We cannot agree withMorafka's1977) suggestionthat ntegrums a "Mexican ighplateaurace of scorpioides" ince itoccupies lowlandhabitatsvirtuallyo sea levelalongmuchof the PacificCoast of westernMexico(albeit t is restrictedo the MexicanPlateau neasternMexico; n this,cf. Iverson ndBerry1979).The relationships mongall membersof the Kinosternoncorpioidescomplex includingntegrum)are discussed by Berry 1978).In examiningthe distributions f K. scorpioides cruentatumand K. integrum n southernOaxaca, Mexico where theirgeographicranges are adjacent,we discovered he existence of athird,undescribed,allopatric pecies of Kinosternon.

    Kinosternonaxacae, sp. nov.Oaxaca MudTurtleFigs. 1 and 2

    Holotype.-UC 48857, whole, preserved,adultmale;collected11.6 km N. of Pochutla SanPedroPochutla),alongMexicanHwy.175 (ca. 235 m),Oaxaca,Mexico 15046'N,96028'W)on 13July1971 byT. Paul Maslinand the UCSummer xpeditiono Oaxaca.Alsobearinga tag "UIMNHField 15892".Allotype.-AMNH88884, whole, preserved,adultfemale;collected11.3 km N. of Pochutla,Oaxaca, Mexicoon 12 August1962 by RogerConant.Paratypes.-A total of six, whole preservedspecimensfromOaxaca,Mexico:UIMNH975,adultmale, Pochutla;KU137680,adultmale,4.8 kmS. of Candelaria;KU87296, adult emale,1km NNWof San GabrielMixtepec 800 m); KU38209-10, juveniles,and KU38211, subadultfemale, Chacalapa(betweenPuertoAngeland Miahuatlan).Diagnosis.-Kinosternonoaxacae is a largespecies (largestspecimena male 159 mm incarapace length,CL) belongingto the scorpioidesspecies complex (with ntegrumand scorpi-oides). It is most similar o K. integrum,but is distinguishableromall otherKinosteronby thefollowingcombination f characteristicsn adults(cf.Table1):(1) a depressed,stronglyricarinatecarapace; (2) a relativelysmall plastron(smallerin males than in females) which does notcompletelyclose the ventralopeningof the shell (maximalwidthof plastralhindlobe57-63% ofmaximalcarapace width,CW);(3) anterior nd posteriorplastralobes bothfreelymoveable; 4)posteriorplastralobe witha distinctanal notchwhich s moreemarginatenmales thaninfemales(Figs. 1 and 2); (5) fixedportion f plastronong,interabdominaleam 24-27%of CL; 6) bridgeofmoderate ength(22-25%of CL),notgroovedposteriorly;7)axillaryndinguinalcutes incontact,inguinal n contact withmarginalcute M5;(8) firstvertebralcute, V1,contactsM2;(9) opposedpatches of hornyscales on the posteriorhighand crus (claspingorgans)lacking n males andfemales; and (10) tail of males and femalesterminatingn a horny pine.Descriptionof species (based on adultsof thetype series):Carapacerelatively epressedandwide (CWand carapace height58-66% and 32-37% of CLrespectively),withthree prominent,parallel, ongitudinal eels. Ground olor of carapacebrown o black,or mottledbrownand black;seams darker nlighter oloredspecimens.Carapacialcutes imbricate;ertebralcutes variablensize (V2-V4 may be longest, V1 or V3-V5may be widest,and V1 or V4-V5may be shortest).Nuchalscute small, longer hanbroadbothdorsallyandventrally.V1usually 83%) ontactingM2.Dorsalmarginsof M1-M9aligned;M10higher,slopingupwardrom ts contactwithM9to seambetweenV5 and fourthpleural,henabruptlyownwardo contactM11;M11notsignificantlyigherthan M9.Plastronrelatively mall, smallerin males than in females (plastronwidthat anteriorhinge65-67% of CWinbothsexes, butmaximalwidthof plastral indlobe 7-59%and62-63%of CW nmales and females respectively).Plastron oncave in males, morenearly latin females. Plastralgroundcoloryellowto brownwithdarker eams, sometimeswithdarkbrown tains,particularlynthe bridgeand ventralsurfacesof the marginals.Plastronwithtwofreelykinetichinges;anterior

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    FIGURE . Photographs f theplastron nd head of males of Kinostemon axacae (UC48857,holotype,ett), . integrumUU78cruentatumUU 7631, right).

    9, 1"t

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    fl ~ ~ ~~ ~ ~ ~ ~ &'~~~~~~~~~~

    ? :?4

    l ~l, ' L

    FIGURE 2. Photographs of the plastron of females of Kinosternon oaxacae (AMNH88884, allotype, left), K. integrum (UU 78cruentatum (UU 5150, right).

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    NEW KINOSTERNONFROM OAXACATABLE1. Character atiosuseful ndistinguishing inosternonaxacae, K. ntegrum, nd K.scorpioidescruentatum. oreachratio expressedas percent) here s a meanwith herangebeneath.Character bbreviationsot ncludedn hetextare:PWant=widthof anterior lastralobe at hinge;PWpost= maximalwidth fposterior lastralobe;Ab= interabdominaleam length;Br=bridge length;Ht = maximal arapace height;F = interfemoraleam length;Post. Lobe= maximalengthof posterior lastrallobe; P = interpectoraleam length;and Ant. Lobe = lengthof anteriorplastral obe.

    CL PWant PWpost Ab Br Ht F P(mm.) CW CW CL CW CL Post Lobe Ant. Lobe

    MALESK. oaxacae (3) 153 65.0 58.4 25.7 35.0 34.1 27.7 13.2150-159 64.6-65.5 57.2-59.4 24.4-27.3 33.7-36.0 32.1-35.6 23.3-32.3 9.4-17.7K. integrumRio Coyuca- (18) 154 68.7 62.4 25.7 35.0 34.8 27.8 15.5Rio Papagayo 110-118 62.5-76.9 56.8-69.4 22.7-27.5 29.5-38.0 31.5-37.4 22.9-36.4 7.4-26.8

    Upper Rio (9) 151 75.3 68.3 26.1 35.7 36.4 25.9 13.4Balsas 113-183 71.3-78.8 64.4-70.5 25.0-27.7 30.8-39.8 34.5-39.7 20.0-32.8 5.6-19.6Rio Verde (6) 150 73.5 65.6 26.3 36.2 36.2 27.8 17.3133-177 63.7-88.4 63.0-70.8 24.1-28.2 33.0-39.2 33.7-41.8 24.2-31.0 14.0-23.2

    K. scorpioidesRio Tehuantepec (24) 122 73.1 69.5 24.9 43.4 42.9 7.4 16.4219-133 67.5-80.5 63.8-75.0 20.5-27.2 40.4-47.5 34.2-47.6 0.0-16.7 9.5-26.7FEMALESK. oaxacae (2) 128 65.8 61.9 26.3 36.8 36.4 32.5 14.0126-130 64.9-66.7 61.7-62.0 26.2-26.4 35.2-38.3 35.8-36.9 32.4-32.5 10.0-17.9

    K. integrumRio Covuca- (39) 142 69.4 65.0 28.6 37.6 39.9 29.1 13.4Rio Papagayo 126-174 63.2-76.7 60.0-71.1 24.6-33.3 32.2-42.3 34.4-44.4 18.2-36.8 4.4-27.1Upper Rio (23) 140 73.0 70.1 27.8 37.8 38.9 21.1 13.8Balsas 123-157 66.7-76.0 64.4-73.3 26.0-31.0 32.0-41.3 32.8-43.8 13.6-26.9 5.0-21.4Rio Verde (10) 131 71.4 65.5 28.3 37.5 41.4 25.5 12.6107-154 68.0-75.6 62.7-70.2 26.3-30.9 34.5-39.6 37.8-45.8 15.4-32.6 6.5-19.4

    K. scorpioidesRio Tehuantepec (49) 124 71.3 68.4 27.1 44.0 48.6 6.1 10.6105-138 65.9-75.9 63.9-72.9 24.8-29.9 40.0-48.5 42.9-53.2 0.0-15.8 2.4-23.8hinge straight (at right angles to midline), posterior hinge curved (Figs. 1 and 2); posterior lobeslightly constricted at hinge; anal notch distinct, more emarginate in males than in females. Axillaryand inguinal scutes in contact; axillaries extending from mid-M4 to posterior M5 or anterior M6,inguinals from posterior M5 or anterior M6 to anterior or mid-M8 (posterior portion of inguinaltending to be replaced by skin in older, larger individuals). Plastral scutes in order of decreasinglength at the midline: abdominal-anal-gular-humeral--femoral-pectoral (33%), or abdominal-anal-gular-femoral-humeral-pectoral (67%).Rostral shield "V"-shaped or bell-shaped. Maxillarysheath strongly hooked in larger speci-mens, weakly hooked in smaller specimens. Head mottled dark brown to black and cream dorsally,cream to yellow with few, faint spots ventrally (more nearly immaculate in females than males);laterally, those patterns integrated in a reticulated network (Fig. 1). Darkpigment coalesced in someto form indistinct light or dark head stripes, but no well-defined head stripes present. Jaw sheathscream to yellow, heavily streaked with dark brown to black in males, faintlystreaked or immaculatein females. Skin of limbs dark brown or gray with darker spots dorsally, immaculate cream to palegray ventrally; tail in both sexes uniformlybrown or gray dorsally and ventrally.Three to four pairsof barbels on chin between mandibular symphysis and throat at level of mid-tympanum;anteriorpair longest, remaining pairs of subequal length. Four to six poorly defined rows of papillae onlateral and dorsal surfaces of the neck.

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    JAMESF. BERRYANDJOHNB. IVERSONHands and feet small, fullywebbed,withwelldevelopedclaws. Discrete,opposed patches ofhorny,tuberculate cales (claspingorgans)on posteriorhighandcruslackingnbothsexes. Tailofmales elongate and prehensile,greaterthan half lengthof posteriorplastral obe;tailof femalesshort, less than halflengthof posteriorplastralobe. Tailinboth sexes with ourrows of dorsal and

    lateralpapillae,and terminalhory spines.Ontogeneticvariation.-The threeimmature aratypesof Kinostemon axacae (KU38209-11,withCL= 55, 84, and 100 mm)differ rom he adult ype specimens inhaving: 1) a proportionatelywider, more evenly roundedcarapace (CW = 73-79% of CL); (2) medialcarapacialkeel moreprominentthan lateral keels; (3) groundcolor of carapace olive to brown with darker seams(carapacialscutes of KU 38210 much darkernear theirposteromedialborders,gradually ighteranterolaterally);4) V1 contactingM2in KU38210 only;(5) M10morenearly he same heightasthe other marginals in smaller specimens; (6) plastronproportionatelymaller than in adults(plastronwidthat anteriorhinge and mid-hind obe 55-62%/and 52-57% of CLrespectively); 7)plastronof the smallest specimen (KU38209) witha faint,gray,foliateplattem;and (8) head withadistinct, lateral,cream-colored tripeextending romposteroventraldge of orbit o angle of jaws.Distribution.-Kinostemonoaxacae is knownonlyfrom he basins of the Rios ColotepecandTonameca in coastal Oaxaca between 100 and 800 m elevation(Fig.3).

    KEYTOTHEADULT CL> 90 mm)KINOSTERNONF OAXACA,MEXICO1. Interabdominaleam length< 22% of CL,< 80% of plastralorelobe ength;gular ength< 52%of plastral orelobelength;maximaldorsalwidthof gularscute > maximal entralwidth;headbrown or gray, withtwo broad,cream or yellowpostorbitaltripes,sometimes broken nto aseries of blotches or mottles......................................... K.leucostomum

    (Atlanticdrainages of the Rios Papaloapanand Coat7acoalcos and theirtributaries, orth-westwardand eastward)Interabdominaleam length> 21% of CL,> 80% of plastralorelobe ength;gular cute length> 50% of plastral orelobelength n94% of individuals;maximaldorsalwidthof gularscute -maximalventralwidth;head not colored as above ..................................... 22. Interfemoral eam length < 20% of maximal engthof plastralhindlobe;plastralhindlobenotnotched or only very slightlynotched posteriorly Figs. 1 and 2); bridgelength> 40% ofmaximalCW.................................................................. 3

    FIGURE . Ranges of Kinostemonn souther Oaxaca,Mexico.Triangles,K.integrum; ots, K.oaxacae (star, ype locality);squares,K.scorpioidescruentatum. rincipalivers renumbered: erde 1), Colotepec 2),Tonaneca(3),andTehuantepec4).Elevationsabove 2400 m are shaded.

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    NEWKINOSTERNONROMOAXACAInterfemoraleam length> 19%of maximalengthof plastralhindlobe;plastralhindlobewithadistinctposteriornotch(Figs. 1 and 2); bridge length< 40%of maximalCW............ 43. Carapace with a single, longitudinal,medialkeel, or lateralkeels considerablyess prominentthan medialkeel; anteriorwidthof plastralhindlobe< 45% of CL; ourth ostal scute usually

    (73%)contacting11thmarginals;maximal hell height< 41% of CL;anterior orderof plastralhindlobeslightlycurved ................................................... K. acutum(Atlanticdrainages of the Rios Papaloapanand Coatzacoalcosand theirtributaries,north-westwardand eastward)Carapace with three longitudinal,parallelkeels of subequal prominence;anteriorwidth ofplastral hindlobe > 42% of CL; fourth costal scutes usually not (83%) contacting 11thmarginals;maximalshell height> 38%of CL;anteriorborderof plastralhindlobea straightline (Figs. 1 and 2) ..........................................K. scorpioidescruentatum(Pacificdrainagesof the RioTehuantepeceastward;Atlantic rainagesof the RioPapaloapannorthwestward)4. Widthof plastral orelobeat anteriorhinge > 67%of CW(88%of individuals);maximalwidthofplastralhindlobe > 59.5% of CW in males (91%)and > 62% of CW in females (93%);interfemoraleam length< 46%of bridge length,and < 12% of maximalplastron ength ...........................................................................K . integ um(Pacific drainages of the Rio Verdenorthand westward)Widthof plastral orelobe< 67% of CW;maximalwidthof plastralhindlobe< 59.5%of CWinmales, < 62% of CWinfemales;interfemoraleam length> 38%of bridge ength,and > 9%of maximalplastron ength................................................. K. oaxacae(Pacific drainages of the Rios Colotepecand Tonameca)

    SPECIMENSEXAMINEDStandard museum acronyms(Duellman,Fritts,and Leviton1978) are used throughouthispaper.

    Kinosternonoaxacae-UC 48857; UIMNH 975; AMNH88884; KU87296, 38209-11, 137680.Kinosternon ntegrum-Rio Coyuca-Rio Papagayo: USNM 46275, 108729-30, 108732; FMNH116514, 116523-24,116540; KU87292-94; UMMZ 0964; UU 7839-40, 7842-44, 7847-84, 7919-20.UpperRio Balsas (RiosMixteco,Acatlan,Atoyac):USNM108590, 108593, 108609-10,108614-15,108617, 108624, 108626, 108628-29, 108631; KU40135-38, 62527; UMMZ 02173-76, 102183,119496, 125364-65; FSM20647; UU 7951, 12086-87, 12095; MVZ76528; CAS 87218.Rio Verde: LACM64498-99; UMMZ119407; KU 87297; UIMNH 966, 9968, 9971-74, 61002,61004-05; UC 48853-54; EAL858.Kinosternon corpioides-Rio Tehuantepec:USNM109105-23, 113278;UMMZ2183-85, 82187-213, 82215-22, 82224-26, 82228, 82230-33, 82235-38, 118633-34;UU 7950.Kinosternon eucostomum-Rio Papaloapan-Rio Jamapa: USNM 109189-92, 109194-96, 109198, 109202, 109205-11;UMMZ89388, 89390, 89392-93, 118290-93; KU 40129-30, 47375-76, 51449; UU 8555-88,10258-61, 11875; FSM32628-29.Rio Coatzacoalcos: AMNH97997; KU24447; UU7953-55.Kinosternonacutum-(see list in Iverson1976).

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    JAMESF. BERRYANDJOHNB. IVERSONRESUMEN

    Las relaciones entre Kinosternoncorpioides y K. integrum e discuten brevemente.Unanueva especie alopatrica,Kinosternonaxacae, se describede la costa de Mexico.K.oaxacae esmiembrodel complejode especies de scorpioides (con integrum),pero se difierede estas dosespecies en el plastronreducido, l carapachomas estrecho,y escudos axillaresy inguinales ncontacto extenso. Una llave dicotomaparalos Kinosternondultosde Oaxacase presenta.

    LITERATUREITEDAhl,E. 1934. Ubereine Sammlung on Reptilien ndAmphibienus Mexiko. ool.Anz.,106:184-186.Berry,J. F. 1978. Variation nd systematicsin the Kinosteron scorpioidesand K. leucostomum omplexes (Reptilia:Testudines:Kinosternidae)f Mexicoand CentralAmerica.Ph.D.dissertation, niv.of Utah,SaltLakeCity,326 pp.andJohnM. Legler.1980. A new turtle genusKinosternon)romSonora,Mexico.Contrib.ci. Nat.Hist.Mus.

    Los Angeles Co. 325:1-12.Casas Andreu,G. 1965. Estudiopreliminarobre las tortugasde agua dulceen Mexico.An. Inst.Nac. Inves.Biol.-Pesq.,1:365-401..1967. Contribucionl conocimiento e las tortugasdulceacuicolas e Mexico.Mexico,D. F.,Univ.Nac.Auton.Mexico,Fac. Ciencias,Dept.Biol.,96 pp.Conant,R. and J. F. Berry.1978. Turtles f the familyKinosternidaenthe southwesternUnitedStatesandadjacentMexico:Identificationnd distribution.mer.Mus.NovitatesNo. 2642:1-18.Duellman,W. E. 1961.The amphibians nd reptilesof Michoacan,Mexico.Univ.KansasPubl.,Mus.Nat. Hist. 15:1-148., T. Fritts, ndA. E. Leviton. 978.Museum cronyms.Herp.Review9: 5-9.Dumeril,A. M. C. and A. H. A. Dumeril. 851. Cataloguemethodique e la collectiondes reptilesde Museumd'HistoireNaturelleGideand Boudry,Paris,224 pp.Hardy,L.M.and R. W.McDiarmid.969. Theamphibiansndreptiles f Sinaloa,Mexico.Univ.KansasPubl.,Mus.Natl.Hist.18:39-252.Iverson,J. B. 1976. The genus Kinostemonn Belize (Testudines:Kinosternidae). erpetologica2:258-262. (See alsoAddendum,Herpetologica2:443).1978. Distributionalroblems f the turtle enus Kinosteron nthe American outhwestandadjacentMexico.Copeia 1978:476-479..1979. A taxonomic eappraisal f the yellowmudturtle,Kinosternonlavescens(Testudines:Kinosternidae).Copeia 1979:212-225.and J. F. Berry.1979.Thegenus Kinosternonn northeasternMexico.Herpetologica5:318-324.LeConte,J. 1854. Descriptions f fournewspecies of Kinosterum.Proc.Acad.Nat. Sci. Philadelphia:180-190.Legler,J. M. 1965. A new species of turtle,genus Kinosternon,romCentralAmerica.Univ.KansasPubl.,Mus. Nat.Hist.15:615-625.Linnaeus,C. 1766. Systemanaturaeperregnatrianaturae, ecundum lasses, ordines,genera,species cumcharacteribus,differentiis,ocis. 12thEd.,Vol. 1. L.Salvius,Stockholm,32 pp.Morafka,D. J. 1977.A biogeographicalnalysisof the Chihuahuan esert hroughtsherpetofauna.iogeographica:1-313.Pritchard, . C. H. 1967. Livingurtlesof the world,T. F. H. Publs.,Jersey City,288 pp.

    .1979. Encyclopediaf turtles.T. F. H. Publs.,Neptune,NewJersey,895 pp.Siebenrock,F. 1906. Schildkrotenus Sudmexiko. ool.Anz.30:94-102..1907. Die Schildkrotenfamilieinosternidaem. Sber. Akad.Wiss.Wien,Math.Nat.KI.116:527-599.Smith,H. M.and E. H.Taylor. 950.Anannotated hecklist ndkeyto thereptiles f Mexico xclusive f thesnakes.Bull.U.S.Nat. Mus. 199:1-253.and R. B. Smith.1980.Synopsisof the herpetofaunaf Mexico.Vol.VI.Guide o Mexicanurtles. ohnJohnson,NorthBennington,Vermont, viii,1044pp.Wermuth,H. and R. Mertens.1961.Schildkroten, rokodile, ruckenechsen. ischerVerlag, ena,422 pp.and . 1977.ListederrezentenAmphibienndReptilien. estudines,Crocodylia, hynchocephalia.Das Tierreich 95:319-352.

    Accepted13 May1980Copyright 980 Societyfor the Studyof Amphibiansnd Reptiles

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