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TAXONOMIC STATUS OF GOMPHOBATES MARMORATUS REINHARDTAND LUumlTKEN 1862 ldquo1861rdquo AND EUPEMPHIX FUSCOMACULATUSSTEINDACHNER 1864 (AMPHIBIA ANURA LEPTODACTYLIDAE)Author(s) Luciana Barreto Nascimento Bruno Vergueiro Silva Pimenta Carlos Alberto GonccedilalvesCruz and Ulisses CaramaschiSource South American Journal of Herpetology 1(3)166-174 2006Published By Brazilian Society of HerpetologyDOI httpdxdoiorg1029941808-9798(2006)1[166TSOGMR]20CO2URL httpwwwbiooneorgdoifull1029941808-97982820062915B1663ATSOGMR5D20CO3B2

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South American Journal of Herpetology 1(3) 2006 166-174copy 2006 Brazilian Society of Herpetology

TAXONOMIC STATUS OF GOMPHOBATES MARMORATUS REINHARDT ANDLUumlTKEN 1862 ldquo1861rdquo AND EUPEMPHIX FUSCOMACULATUS

STEINDACHNER 1864 (AMPHIBIA ANURA LEPTODACTYLIDAE)

LUCIANA BARRETO NASCIMENTO13 BRUNO VERGUEIRO SILVA PIMENTA2CARLOS ALBERTO GONCcedilALVES CRUZ2 AND ULISSES CARAMASCHI2

2 Museu de Ciecircncias Naturais Departamento de Ciecircncias Bioloacutegicas e Programa de Poacutes-graduaccedilatildeo em Zoologia de VertebradosPUC Minas Av Dom Joseacute Gaspar 290 30535-610 Belo Horizonte Minas Gerais Brazil

3 Departamento de Vertebrados Museu NacionalUFRJ Quinta da Boa Vista 20940-040 Rio de Janeiro Rio de Janeiro Brazil3 Corresponding author lunapucminasbr

ABSTRACT An analysis of specimens deposited in herpetological collections and identified as Physalaemus fuscomaculatus demonstratedthat these differ from the types of this taxon Eupemphix fuscomaculatus Consequently the taxonomic status of Gomphobatesmarmoratus and Eupemphix fuscomaculatus was revised based on the types and on data in the literature Gomphobates marmoratusis associated to the genus Physalaemus as Physalaemus marmoratus and refers to specimens previously identified as Physalaemusfuscomaculatus Additionally Eupemphix fuscomaculatus is associated to the genus Pleurodema in the combination Pleurodemafuscomaculata

KEYWORDS Amphibia Anura Taxonomy Physalaemus Pleurodema

INTRODUCTION

Reinhardt and Luumltken (1862 ldquo1861rdquo) describedGomphobates marmoratus based on three syntypescollected in Lagoa Santa (19deg37rsquoS 43deg53rsquoW) State ofMinas Gerais Brazil Currently G marmoratus is ajunior synonym of Physalaemus fuscomaculatus(Steindachner 1864) (Frost 2006)

Steindachner (1864) described Eupemphix fusco-maculatus from one female from ldquoCaiccedilaraacuterdquo [17deg15rsquoS57deg10rsquoW according to Cei (1990)] State of Mato Gros-so Brazil Currently E fuscomaculatus is a juniorsynonym of Physalaemus fuscomaculatus (Stein-dachner 1863) (Frost 2006) This taxon was betterdefined and illustrated by Cei (1990) based on the ho-lotype and on additional specimens from Rio Apa(22deg30rsquoS 57deg00rsquoW) a tributary of Rio Paraguay Heconsidered this species ldquoa relatively uncommon lepto-dactylidrdquo ranging from the type locality to the lowerbasin of Rio Paraguay probably reaching the marginalarea of northern Argentinean provinces southwards

Physalaemus fuscomaculatus is presently allocatedin the P albifrons group (Nascimento et al 2005)based on several morphological characters such asdorsal and ventral color patterns rounded snout largeinguinal glands not associated with a dark ocellus andshovel-like external and internal metatarsal tubercleswith horned distal margins

Several specimens identified as Physalaemusfuscomaculatus collected in localities outside thedistribution range given by Cei (1990) are depositedin herpetological collections (Nascimento et al2005) Analysis of this material demonstrated thatsuch specimens differ from the taxon studied by Cei(1990) but agree perfectly with the types of Gom-phobates marmoratus Reinhardt and Luumltken 1862ldquo1861rdquo

Based on the examination of the types we proposethe revalidation of G marmoratus under a new com-bination and designate a lectotype and paralectotypesWe also assess the taxonomic status ofE fuscomaculatus Steindachner 1863 and propose theallocation of this species in the genus PleurodemaTschudi 1838

Historical background

Reinhardt and Luumltken (1862 ldquo1861rdquo) erected thegenus Gomphobates and described the speciesG marmoratus from Lagoa Santa State of MinasGerais Brazil

Steindachner (1864) described Eupemphix fusco-maculatus from ldquoCaiccedilaraacute in Brasilienrdquo and pointed thepresence of maxillary and vomerine teeth He associ-ated Hiobates fuscomaculatus Fitz Tschudi as a syn-onym of this species

Nascimento LB et al 167

Steindachner (1867) established part of Eupemphixnattereri Steindachner 1863 as a junior synonym ofGomphobates marmoratus associated Eupemphixfuscomaculatus to this genus under the new combi-nation Gomphobates fuscomaculatus and indicatedIliobates fuscomaculatus (assigning it to ldquoFitz Ts-chudirdquo) as a synonym of the latter

Cope (1869 ldquo1868rdquo) described Lystris brachyops(= Pleurodema brachyops) and distinguished this ge-nus from Gomphobates by the presence of fontanelaand vomerine teeth and from Eupemphix by the pres-ence of well-developed teeth on young and adultsEupemphix fuscomaculatus Steindachner was thenincluded in the genus Lystris under the combinationLystris fuscomaculatus (Steindachner)

Peters (1872) included Gomphobates marmora-tus Reinhardt and Luumltken Liuperus marmoratus Bur-meister 1861 and part of Eupemphix nattereri Stein-dachner in the synonymy of Bufo albifrons Spix 1824

Boulenger (1882) allocated Pleurodema TschudiCystignathus Dumeacuteril and Bibron Leiuperus Dumeacuteriland Bibron Pleurodema Guumlnther Gomphobates Re-inhardt and Luumltken Eupemphix Steindachner andLystris Cope in the genus Paludicola Wagler basedon the presence of maxillary teeth Consequently thenew combinations Paludicola fuscomaculata (Stein-dachner) and Paludicola albifrons (Spix) were es-tablished for Lystris fuscomaculatus and Bufo albi-frons respectively Boulenger cited the absence ofvomerine teeth in P fuscomaculata (Steindachner) andestablished its distribution range to Brazil Uruguay andBuenos Aires (Argentina) Paludicola biligonigera(Cope) was associated to Reinhardtrsquos specimens fromLagoa Santa (State of Minas Gerais) and from theStates of Bahia and Paraacute Brazil

Boulenger (1886) indicated the occurrence of Palu-dicola fuscomaculata (Steindachner) and Paludico-la albifrons (Spix) in the State of Rio Grande do SulBrazil Boulenger (1887) presented a diagnosis of Palu-dicola albifrons from Porto Alegre Rio Grande doSul Brazil He indicated that Leiuperus marmoratusBurmeister was not identical to Paludicola albifrons(Spix) as stated by Peters (1872) but was a junior syn-onym of P fuscomaculata (Steindachner) In realityLeiuperus marmoratus DrsquoOrbigny 1847 (referred byBurmeister 1861) Leiuperus marmoratus Burmeis-ter 1861 and Paludicola fuscomaculata Boulenger1886 1887 refers to the currently recognized Phys-alaemus biligonigerus (Cope 1861)

Meacutehely (1904) after examining the type designat-ed by Spix distinguished Paludicola albifrons (Spix)from P fuscomaculata (Steindachner) by the pres-ence of a second tarsal tubercle in the articulation be-tween tibia and tarsus on the former larger than thedistal tarsal tubercle and by the absence of externallyvisible lumbar glands He also indicated thatP fuscomaculata (Steindachner) presents no vomer-ine teeth

Miranda-Ribeiro (1926) included Eupemphix En-gystomops Pleurodema and Paludicola in the fam-ily Paludicolidae Paludicola was characterized by thepresence of ldquomore or less indistinct vomerine teethrdquoHe compared specimens of Paludicola albifrons(Spix) and P fuscomaculata (Steindachner) to the typeof Eupemphix fuscomaculatus Steindachner andagreed with Meacutehely (1904) on the difference amongthese species He also mentioned thatP fuscomaculata (Steindachner) presents ldquovomerineteeth indistinct but present on a line anterior to thechoanaerdquo He indicated the distribution of P albifrons(Spix) for the State of Bahia Brazil and that ofP fuscomaculata (Steindachner) for the states of MatoGrosso Satildeo Paulo Paranaacute and Rio Grande do Sul inBrazil and also in Argentina and Uruguay

The first revision of the genus Paludicola waspresented by Parker (1927) who distributed its spe-cies among the genera Physalaemus Fitzinger 1826Pseudopaludicola Miranda-Ribeiro 1926 and Pleu-rodema Paludicola biligonigera (Cope)P fuscomaculata (Steindachner) and P albifrons(Spix) were included in the genus Physalaemus Healso discussed the differences indicated by Meheacutely(1904) between Physalaemus albifrons (Spix) andP fuscomaculatus (Steindachner) Parker (1927) re-moved Gomphobates marmoratus Reinhardt andLuumltken from the synonymy with P albifrons (Spix)and considered it a junior synonym ofP fuscomaculatus (Steindachner) after requestingthe analysis of the type-specimens of P albifrons(Spix)

Cochran (1955) mentioned the absence of vomer-ine teeth in Physalaemus fuscomaculatus (Stein-dachner) ldquoalthough the bony projection on which theywould have arisen can be felt and in a few cases seenbetween the choanaerdquo The description presented wasbased on specimens from the States of Minas GeraisRio Grande do Sul and Satildeo Paulo (Brazil) ArgentinaBolivia and Paraguay

168 Taxonomic status of Gomphobates marmoratus

Milstead (1960) characterized Physalaemus bili-gonigerus (Cope) and P fuscomaculatus (Stein-dachner) species occurring in the State of Rio Grandedo Sul Brazil and pointed that the first differs from allcongeneric species by the presence of vomerine teethHe observed differences on snout-vent length betweenspecimens of P fuscomaculatus (Steindachner) fromthe States of Satildeo Paulo and Rio Grande do Sul sug-gesting that this taxon might include two distinct spe-cies or geographic races

Milstead (1963) after analyzing the holotype ofP biligonigerus (Cope) verified the absence ofvomerine teeth stating that it was a ldquotypical specimenof what is now called P fuscomaculatusrdquo (Stein-dachner) Therefore he considered that the speciestreated by Parker (1927) Cochran (1955) and Mil-stead (1960) as P fuscomaculatus (Steindachner)should be in fact called P biligonigerus (Cope) basedon the precedence of this name He suggested theaplication of the name ldquobiligonigerusrdquo to the smallerspecimens from the south whereas ldquofuscomaculatusrdquoshould be applied to the more widespread northernform

Barrio (1965) agreed with Milstead (1963) that theforms previously identified as Physalaemus fuscomac-ulatus (Steindachner) in Argentina corresponded toP biligonigerus (Cope) but disagreed with the dis-tinction between P biligonigerus (Cope) related tothe southern smaller specimens andP fuscomaculatus (Steindachner) associated to thenorthern larger specimens

Lynch (1970) indicated the absence of vomerineteeth (referred as prevomerine teeth) for all species ofpaludicoline leptodactylids except for Physalaemusbiligonigerus (Cope) which sometimes has them

Frost (1985) mentioned the occurrence of Phys-alaemus fuscomaculatus (Steindachner) in Argenti-na Cei (1987) considered it ldquoquite improbablerdquo andpresented a summarized account on the differencesbetween P biligonigerus (Cope) andP fuscomaculatus (Steindachner) He also indicatedthe absence of vomerine teeth on both forms

Cei (1990) defined and illustrated the taxon Phys-alaemus fuscomaculatus considering that ldquoCaiccedilaraacuterdquohas been misspelled and should be written ldquoCaissaraacuterdquoHe stated that P fuscomaculatus (Steindachner) andP biligonigerus (Cope) were considered as a largecomplex of species of Physalaemus biligonigerus(Cope) since the arrangement of Boulenger (1882)

He pointed out that the morphology of the former spe-cies is inaccurately determined because Stein-dachnerrsquos description of the type clearly indicated thepresence of maxillary and vomerine teeth Physalae-mus fuscomaculatus (Steindachner) was character-ized morphologically based on the holotype and on spec-imens collected at Rio Apa a tributary of Rio Para-guay not far from the type-locality adding informationto Steindachnerrsquos (1863) diagnosis The presence ofvomerine teeth is emphasized as well as the rarity ofthis species in herpetological collections

Nascimento et al (2005) diagnosed the genusPhysalaemus by the presence of quadratojugal bonesand absence of vomerine teeth among other charac-teristics These characters distinguish Physalaemusfrom Pleurodema They also associatedP fuscomaculatus (Steindachner) to the P albifronsspecies group

MATERIAL AND METHODS

Acronyms used in the text are as follow MCNAM(Museu de Ciecircncias Naturais Pontifiacutecia UniversidadeCatoacutelica de Minas Gerais Belo Horizonte Brazil)MNRJ (Museu Nacional Rio de Janeiro Brazil)MZUSP (Museu de Zoologia Universidade de SatildeoPaulo Satildeo Paulo Brazil) NHMW (NaturhistorischesMuseum of Wien Vienna Austria) UFRGS (Univer-sidade Federal do Rio Grande do Sul Porto AlegreBrazil) R and ZMUC (Zoological Museum Universi-ty of Copenhagen Denmark)

Morphometric data follow Cei (1980) and the abre-viations used for are SVL (snout-vent length) HL (headlength) HW (head width) ED (eye diameter) IOD(interorbital distance) UEW (upper eyelid width) END(eye-nostril distance) NSD (nostril-snout distance)IND (internarial distance) UL (upper arm length) AL(arm length) HAL (hand length) TL (thigh length) SL(shank length) FL (foot length from the inner meta-tarsal tubercle to the distal point of fourth toe)

RESULTS AND DISCUSSION

We examined two of the three syntypes of Gom-phobates marmoratus Reinhardt and Luumltken R 1193(formerly ZMUC 22) is in very good conditions all thedistinctive characters of the species are easily ob-served The coloration is quite faded due to the time ithas been preserved Skin on flanks is extensively fold-

Nascimento LB et al 169

ed a feature not observed on living and recently pre-served specimens An incision has been made in thepectoral girdle region but it did not cause too muchdamage Fingers and toes are slightly curved A smalllabel written ldquoType 22rdquo is tied on the right leg R 1194(formerly ZMUC 23) is not well preserved as R 1193It is flaccid and because of that it is not possible toobserve the glands and the coloration is more fadedThe third syntype R 11125 (formerly ZMUC 26) is acleaned skeleton and was not analyzed

We found verified that vomerine teeth and darkocellus on lumbar glands are not present on the typesof Gomphobates marmoratus Reinhardt and LuumltkenSteindachner (1864) diagnosed and Cei (1990) veri-fied the presence of these characters in the holotypeof Eupemphix fuscomaculatus and in additional spec-imens from Rio Apa Paraguay Additionally the pres-ence of vomerine teeth distinguishes Pleurodema fromPhysalaemus (Nascimento et al 2005) We there-fore conclude that Gomphobates marmoratus Rein-hardt and Luumltken should be removed from the synony-my with Physalaemus fuscomaculatus (Steindachner)and recognized as a full species under a new combi-nation Moreover due to the presence of vomerineteeth Eupemphix fuscomaculatus Steindachnershould be allocated to the genus Pleurodema

Species Accounts

Physalaemus marmoratus (Reinhardt and Luumltken1862 ldquo1861rdquo) n comb

Gomphobates marmoratus Reinhardt and Luumltken1862 ldquo1861rdquo

Eupemphix nattereri Steindachner 1863 (part)Paludicola albifrons ndash (not of Spix 1824) Peters

1872 (part)Paludicola biligonigera ndash Boulenger 1882 (part)Paludicola fuscomaculata ndash Meheacutely 1904Physalaemus fuscomaculatus ndash Parker 1927 (part)

Lectotype ndash R 1193 (formerly ZMUC 22) male col-lected in Lagoa Santa State of Minas Gerais Brazil inNovember 1854 by J B Reinhardt

Paralectotype ndash R 1194 (formerly ZMUC 23) malecollected in Lagoa Santa State of Minas Gerais Bra-zil in 04 August 1841 by PW Lund R 11125 (former-ly ZMUC 26) cleaned skeleton collected in Lagoa

Santa State of Minas Gerais Brazil no date by PWLund

Original description of Gomphobates marmoratus ndashldquoThis species seems not only to be much larger thanthe preceding (G notatus) but is plumper about likethe plumper of the Cystignathi The body is thickerand wider the head rounded Although the skin is looseand has many irregular folds the preceding lateral folddoes not seem to be missing besides there are the foldaround the belly the crossfold between the arms andthe inflated throat skin as in the preceding the threeavailable specimens also have openings next to thetongue which lead down to the vocal sacs What con-tributes to giving the species a different make-up isthat there are not so few large warts on the back (oneof them shows that these can disappear almost com-pletely on soft specimens) The differences are reducedto the following the small points on the outer posteriorborder of the tarsus are missing the blade of the footis wider and there is a trace of a fringe around thelarger toes The markings are really to a great extentthe same as on G notatus but much stronger and moredistinct the warts on the back are bordered by darkrings The dark lines characteristic of G notatus andthe 2 round spots on the hind part of the back are miss-ing here A white line which occupies the middle of thehind part of the back is more distinct than the preced-ing species In all other important respects they corre-spond despite the difference in their make-uprdquo (Trans-lation from old Danish by Astrid Schmidt-Nielsen)

Diagnosis ndash A species belonging to the P albifronsspecies group diagnosed by the following combinationof characters (1) size large for the group (SVL formales 333-421 mm females 330-475 mm) (2) bodyrobust (3) head slightly wider than long (4) snoutrounded in dorsal and lateral views (5) canthus ros-tralis rounded (6) vocal sac subgular well developedextending to the border of chest with belly (7) fingersand toes robust fringed with horned tips (8) tarsaltubercle present (9) metatarsal tubercles compressedprotruding with horned distal margins (10) toes webbedon base (11) large inguinal glands with the same colorpattern of dorsum (12) dorsal color pattern presentingan ldquoomegardquo-shaped mark and stripes extending toflanks and inguinal region and (13) presence of a darkbrown interorbital bar Physalaemus marmoratus canbe distinguished from all the other members of the

170 Taxonomic status of Gomphobates marmoratus

P albifrons group due to its larger size (SVL com-bined for males of the other species 282-326 mmfemales 284-341 mm) robust fingers (slender longerfingers in the other species) and dorsal color patternPhysalaemus marmoratus differs from P albifronsand P biligonigerus by the fringed fingers (fringesabsent or poorly marked in P albifrons andP biligonigerus) from P albifrons andP santafecinus by the presence of long rounded orirregular glandular ridges on dorsum (dorsal skin smoothin P albifrons and granulated in P santafecinus)from P albifrons by the presence of one tarsal tuber-cle (two in P albifrons) and large inguinal glands (in-distinct in P albifrons) from P biligonigerus by in-guinal glands with same color pattern of dorsum (in-guinal glands with color pattern darker than dorsum inP biligonigerus) from P santafecinus by the ab-sence of granules on forearms (presence of a line ofgranules on the outer margins of forearms inP santafecinus) and absence of a white longitudinalline on sacral region (evident vertebral line inP santafecinus)

Description of lectotype ndash Body robust head slightlywider than long snout rounded in dorsal (Fig 1) andlateral views (Fig 2) nostrils elliptical not protuber-ant located near the tip of snout oriented dorsolateral-ly canthus rostralis rounded loreal region slightly con-cave eyes protuberant eye diameter larger than in-terorbital distance tympanum indistinct supratympanicfold short slightly marked dorsolateral fold absentvocal sac subgular well developed extending to theborder of chest with belly choanae small oval tonguesmall narrow rounded on posterior border which isnot indented maxillary and premaxillary teeth presentvomerine teeth absent Arms short robust upper armsshorter than forearms outer margin of forearms with-out ridges or granules fingers robust fringed fingertips not expanded horned finger lengths IltIVltIIltIII(Fig 3) nuptial asperities on thumbs and on the medialborder of the inner carpal tubercle but not continuouscarpal tubercles large elliptical with the same sizesupernumerary tubercles protruding conical subartic-ular tubercles single large protruding conical Legsmoderately robust tibia slightly longer than thigh tar-sal fold absent toes slender long slightly fringed (fringeabsent on toe I) toes webbed on base tips not ex-panded horned toe lengths IltIIltVltIIIltIV (Fig 4)tarsal tubercle present metatarsal tubercles large shov-

el-shaped protruding with distal margins horned dis-tance between inner and outer metatarsal tuberclesshorter than that between the former and tarsal tuber-cles supernumerary tubercles absent subarticular tu-bercles single large protruding conical outer marginof subarticular tubercle of toe I weakly horned dorsalpattern presenting an ldquoomegardquo-shaped mark and stripes

Figure 2 Lateral view of the lectotype of Gomphobates marmora-tus Reinhardt and Luumltken 1862 ldquo1861rdquo

Figure 1 Dorsal view of the lectotype of Gomphobates marmora-tus Reinhardt and Luumltken 1862 ldquo1861rdquo

Figure 3 Ventral view of the hand of the lectotype of Gompho-bates marmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Nascimento LB et al 171

extending to flanks and inguinal region dorsum andflanks with long rounded or irregular glandular ridgesupper eyelids finely rugose ventral surfaces smoothexcept on cloacal region and thighs which are cov-ered by large granules ventral disc distinct inguinalgland large ovoid General color pattern in preserva-tive cream light brown blotches and sinuous stripesirregular generally wide bordered by brown lines lightbrown stripes continuous or interrupted tending to forman ldquoomegardquo-shaped mark under scapular region in-terorbital bar dark brown white longitudinal line onsacral region vertical bars dark brown and cream al-ternated from the snout to posterior corner of mouthtransversal bars dark brown and cream alternated ondorsum of arms and legs gular region grayish chestbelly ventral surfaces of arms and legs cream palmarand plantar surfaces grayish over cream backgroundnuptial asperities light brown horned margins of fin-gers toes and metatarsal tubercles dark brown in-guinal gland with the same color pattern of dorsum

Measurements ndash SVL 406 HW 124 HL 112 THL162 TL 173 FL 190 ED 51 ID 36 END 24UEW 39 IND 26

Variation ndash Paralectotype R 1194 shows finger tipsless horned inner border of toe I with a small fringeand subarticular tubercles on toes II III IV and Vhorned Granular ridges granules and inguinal glandsare less evident probably due to preservation Thegeneral color pattern follows that of the lectotype butcolors are more faded Measurements of paralecto-type SVL 385 HW 98 HL 106 THL 150 TL 157FL 170 ED 44 ID 43 END 29 UEW 37 IND

26 Some of the additional specimens examinedshowed a narrower head and more rugose dorsumPalmar and plantar tubercles may be less or morehorned This variation is also observed among tips offingers and toes on a single specimen General colorpattern on dorsum and nuptial asperities of recentlypreserved specimens vary from gray to dark brownthe white longitudinal line on sacral region may be fad-ed Gular region of females follows the color patternof chest and belly Morphometric variation is present-ed in Table 1

Tadpole ndash External morphology and internal oral mor-phology were described by Nomura et al (2003) asPhysalaemus fuscomaculatus

Distribution and ecology ndash Physalaemus marmora-tus breeds in temporary shallow ponds of open habi-tats in the States of Bahia Espiacuterito Santo Rio de Jan-eiro Satildeo Paulo Minas Gerais Goiaacutes Mato Grossoand Mato Grosso do Sul (Brazil) and also in Para-guay Males call floating on the water with the largevocal sac inflated During the axillary amplexus a foamnest is constructed on the water surface where theunpigmented eggs are deposited

Remarks ndash Reinhardt and Luumltken (1862 ldquo1861rdquo) des-ignated three syntypes in the original description ofG marmoratus one was collected by Reinhardt theother two were sent by Dr PW Lund one to theUniversity Museum of Copenhagen and the other tothe ldquoRoyal Museum of Natural Historyrdquo DenmarkLater on all natural history collections in Denmark wereunited into one large collection now ldquoZoological Mu-

Figure 4 Ventral view of the foot of the lectotype of Gomphobatesmarmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Table 1 Mean (x) standard deviation (SD) and range of measure-ments (in mm) of males and females of Physalaemus marmoratus(Reinhardt and Luumltken 1862 ldquo1861rdquo)

Males (N=19) Females (N=13)x SD Range x SD Range

SVL 382 21 333-421 418 42 330-475HL 102 06 92-112 106 07 90-117HW 113 08 10-128 125 10 109-138ED 45 03 40-51 46 05 37-53END 26 02 23-28 27 02 24-31IND 26 01 23-28 27 02 24-31IOD 33 04 27-43 34 04 28-41UEW 39 03 33-46 42 04 37-50THL 157 11 138-176 162 15 139-190TL 155 10 138-173 159 12 136-178FL 174 11 159-190 177 12 156-193

172 Taxonomic status of Gomphobates marmoratus

seum of the University of Copenhagenrdquo (ZMUC MAndersen pers comm) When we consulted theZMUC Curator about the loan of the syntypes he lo-cated two specimens R 1193 (here designated as thelectotype) and R 1194 (here designated as one of theparalectotypes) When asked about the third syntypethe Assistant-Curator could not trace it with certaintybecause none of the three other specimens ofG marmoratus deposited in the ZMUC collection car-ried a label indicating which was the third type How-ever he provided all the information available in thejars labels One specimen (R 1170 formerly ZMUC24) was collected by E Warming which did not col-lect any of the types Specimen R 1195 (formerlyZMUC 25) is just labeled ldquoMinas Geraisrdquo with nocollector and date The last specimen (R 11125 for-merly ZMUC 26 a cleaned skeleton loose bones andskin in alcohol) was collected by PW Lund but thelabel does not indicate the locality and date only aLundrsquos handwriting note saying ldquoNew genus of frogs(1)rdquo We considered this latter as the third syntypebecause of the evidences shown in the label

Pleurodema fuscomaculata (Steindachner 1864)n comb

Eupemphix fuscomaculatus Steindachner 1864Hiobates fuscomaculatus Steindachner 1864Gomphobates fuscomaculatus ndash Steindachner 1867Iliobates fuscomaculatus ndash Steindachner 1867Lystris fuscomaculatus ndash Cope 1869 ldquo1868rdquoPaludicola fuscomaculata ndash Boulenger 1882Physalaemus fuscomaculatus ndash Parker 1927 (part)

Holotype ndash NHMW 4316 female collected by JNatterer in Caiccedilara Municipality of Caacuteceres State ofMato Grosso Brazil (Fig 5 and Fig 6)

A very good description of the holotype of Eupem-phix fuscomaculatus as well as comparisons withother coespecific specimens and notes on distributionis provided by Cei (1990)

Remarks ndash The presence or absence of vomerine teethseems to be variable among genera of the currentlyLeiuperidae (sensu Frost et al 2006) Consequentlyit is not yet possible to determine with certainty forwhat taxon or group of taxa the presence or absenceof vomerine teeth represent a synapomorphic stateThis would require a complete sampling of all species

and all genera involved which is beyond the scope ofthis paper Until this can be accomplished we tenta-tively use the new combination Pleurodema fusco-maculata

RESUMO

A anaacutelise de exemplares depositados em coleccedilotildeesherpetoloacutegicas e identificados como Physalaemus fus-comaculatus demonstrou que estes diferem do tipodeste taacutexon Eupemphix fuscomaculatus Consequumlen-temente o status taxonocircmico de Gomphobates mar-moratus e Eupemphix fuscomaculatus foi revisadocom base nos tipos e na literatura Gomphobatesmarmoratus eacute associado ao gecircnero Physalaemus

Figure 6 Ventral view of the type of Eupemphix fuscomaculatusSteindachner 1864

Figure 5 Dorsal view of the type of Eupemphix fuscomaculatusSteindachner 1864

Nascimento LB et al 173

como Physalaemus marmoratus e se refere aosexemplares previamente identificados como Physala-emus fuscomaculatus Adicionalmente Eupemphixfuscomaculatus eacute associado ao gecircnero Pleurodemacom a combinaccedilatildeo de Pleurodema fuscomaculata

ACKNOWLEDGMENTS

We thank Giovanni Vinciprova (UFRGS) Jens Boslashdtker Ras-mussen (in memorian) and Mogens Andersen (Curator and Assis-tant-Curator of the Zoological Museum University of Copen-hagen Denmark respectively) for providing specimens under theircare Jose P Langone (Museo de Historia Natural de MontevideoUruguay) for providing photos of the type of Eupemphix fusco-maculatus Miguel A Andrade for providing photos of the lecto-type of Gomphobates marmoratus Ceacutelio FB Haddad for criti-cally reading the manuscript Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for the fellowships grant-ed to BVSP CAGC and UC and financial support

LITERATURE CITED

BARRIO A 1965 El genero Physalaemus (Anura Leptodactylidae)en la Argentina Physis 25(70)421-448

BOULENGER GA 1882 Catalogue of Batrachia Salientia siveEcaudata and Batrachia Apoda in the collection of the BritishMuseum 2nd Ed London British Museum xxx + 503p

BOULENGER GA 1886 A synopsis of the reptiles and batrachiansof the Province Rio Grande do Sul Brazil Annals and Magazineof Natural History 18(5)423-445

BOULENGER GA 1887 Description of a new or little-known SouthAmerican frogs of the genera Paludicola and Hyla Annals andMagazine of Natural History 2(5)295-300

CEI JM 1980 Amphibians of Argentina Monitore ZoologicoItaliano (NS) Firenze Monografia 2609p

CEI JM 1987 Additional notes to ldquoAmphibians of Argentinardquo anupdate 1980-1986 Monitore Zoologico Italiano 21209-272

CEI JM 1990 On a paraguayan sample of a long time confusedspecies Physalaemus fuscomaculatus (Steindachner 1864)(Anura Leptodactylidae) Bolletin di Museo Regionale diScienze Naturali di Torino 8(1)215-231

COCHRAN DM 1955 Frogs of Southeastern Brazil United StatesNatural Museum Bulletin 206 xvi + 423p

COPE ED 1869 ldquo1868rdquo Sixth contribution to the herpetology ofTropical America Proceedings of the Academy Natural Sciencesof Philadelphia 20305-313

FROST DR (ED) 1985 Amphibian species of the world Ataxonomic and geographical reference Lawrence Allen PressInc and The Association of Systematics Collections v + 732p

FROST DR 2006 Amphibian Species of the World an OnlineReference Version 40 (17 August 2006) Electronic Databaseaccessible at httpresearchamnhorgherpetologyamphibia

indexhtml American Museum of Natural History New YorkUSA [captured on 20 August 2006]

LYNCH JD 1970 Systematic status of the American leptodactylidfrog genera Engystomops Eupemphix and PhysalaemusCopeia 1970(3)488-496

MEacuteHELY L VON 1904 Investigations on paraguayan batrachiansAnnales Historico-Naturales Musei Nationalis Hungarici2207-232

MILSTEAD WW 1960 Frogs of the genus Physalaemus in SouthernBrazil with description of a new species Copeia1960(2)83-89

MILSTEAD WW 1963 Notes on brazilian frogs of the generaPseudopaludicola and Physalaemus Copeia 1963 (3)565-566

MIRANDA-RIBEIRO A 1926 Notas para servirem ao estudo dosgymnobatrachios (Anura) brasileiros Arquivos do MuseuNacional Rio de Janeiro 271-227

NASCIMENTO LB U CARAMASCHI AND CAG CRUZ 2005Taxonomic review of the species groups of the genusPhysalaemus Fitzinger 1826 with the revalidation of the generaEngystomops Jimeacutenez-de-la-Espada 1872 and EupemphixSteindachner 1863 (Amphibia Anura Leptodactylidae)Arquivos do Museu Nacional Rio de Janeiro 63(2)297-320

NOMURA F DC ROSSA-FERES AND VHM PRADO 2003 Thetadpole of Physalaemus fuscomaculatus (AnuraLeptodactylidae) with a description of internal oralmorphology Zootaxa 3701-8

PARKER HW 1927 A revision of the frogs of the generaPseudopaludicola Physalaemus and Pleurodema Annals andMagazine of Natural History ser 9(20)450-478

PETERS W 1872 Uumlber eine Sammlung von Batrachiern aus Neu-Freiburg in Brasilien Monatsberichte der Akademie derWissenschaften zu Berlin 1872680-684

REINHARDT J AND CF LUumlTKEN 1862 ldquo1861rdquo Bidgrad til Kundskabom Brasiliens Padder og Krybdyr Videnskabelige Meddelelserfra den Naturhistoriske Forening i Kjoslashbenhavn3(10-15)143-242

SPIX JB 1824 Animalia nova sive species novae Testudinum etRanarum quas it itinere per Brazilian annisMDCCCXVIII-MDCCXX jussu et auspiciis MaximillianiJosephi I Bavarrie Regis Monachii xxii + 53p

STEINDACHNER F 1863 Uumlber einige neue Batrachier aus denSammlungen des Wiener Museums Sitzungsberichte derAkademie der Wissenschaften 48186-192

STEINDACHNER F 1864 Batrachologische MitteilungenVerhandelugen Zoologisch-Botanischen Gesellschaft in Wien14239-288

STEINDACHNER F 1867 Amphibien In Reise der OesterreichischenFregatte Novara um die Erde in den Jahren 1857 1858 1859unter den Befehlen des Commodore B von Wuumlllerstorf-UrbairZoologischer Theil 70p

Received 28 April 2006Accepted 16 November 2006

174 Taxonomic status of Gomphobates marmoratus

APPENDIX I

Specimens Examined

Physalaemus albifrons BRAZIL MARANHAtildeO MNRJ 24216-24226 Barreirinha CEARAacute MNRJ 14940-14942 MNRJ 24059 MNRJ 24062-24072 Brejo Santo MNRJ 6636-6674 Mucuripe Fortaleza MNRJ 1125MNRJ 6636-6676 Fortaleza SERGIPE MNRJ 17976-17981 Santa Luzia do Itanhy BAHIA MNRJ 1094-1096Barreiras MNRJ 114-116 MNRJ 1094-1096 MNRJ 1113 MNRJ 1102-1104 MNRJ 1089-1091 Bom Jesus daLapa MNRJ 1105 Juazeiro MZUSP 76521 MZUSP 82303 MINAS GERAIS MNRJ 27180-27189 LageadoMNRJ 27179 MNRJ 21743-21745 Manga MCNAM 206-209 MCNAM 213-216 Porteirinha

Physalaemus biligonigerus BOLIacuteVIA SANTA CRUZ AMNH 144362-144370 AMNH 144371-144385 AMNH144432-144433 BRAZIL SANTA CATARINA MNRJ 31128-31153 RIO GRANDE DO SUL UFGRS 1924-1929 General Cacircmara UFGRS 1776 Uruguaiana UFGRS 1619 Tramandaiacute UFGRS 1319-20 UFRGS 1342Parque Estadual de Itapoatilde Viamatildeo PARAGUAY AMNH 23807 Paraguayan Chaco AMNH 50669-50671Colocircnia Nueva Itaacutelia Villeta URUGUAY DEPARTAMENTO RIO NEGRO MNRJ 28554

Physalaemus marmoratus BRAZIL Bahia MNRJ 28443 MNRJ 28489 Caravelas DISTRITO FEDERALMNRJ 3415 Ribeiratildeo do Torto GoiaacutesMinas Gerais MCNAM 3915-3918 MCNAM 3964-3965 MCNAM 4796region of Queimados Hydroelectric Dam (AHE Queimados) Minas Gerais MCNAM 1259 Bocaiuacuteva MCNAM3155-3157 Conselheiro Mata MNRJ 1020-1021 Lagoa Santa MNRJ 40876-40877 Pirapora MCNAM 2135-2138 MCNAM 2204-2209 MCNAM 2238 MCNAM 2604 MCNAM 2738 MCNAM 2740-2745 MCNAM2748-2751 MCNAM 3147-3148 Santana do Riacho ESPIacuteRITO SANTO MNRJ 40878-40879 Piuacutema RIO DEJANEIRO MNRJ 1015-1019 Campos dos Goytacazes MNRJ 3366 MNRJ 13872-13914 MNRJ 40587 MNRJ40880 MNRJ 40881 Satildeo Joatildeo da Barra MATO GROSSO DO SUL MNRJ 24875 Miranda MNRJ 3070 MNRJ13263 Salobra SAtildeO PAULO MNRJ 40882-40883 Botucatu MNRJ 34666-34671 Campinas MNRJ 37043-37045 Piraju PARAGUAY MNRJ 12710-12730 Asuncioacuten MNRJ 12696-12709 Brejo de Ipuatilde

Physalaemus santafecinus ARGENTINA CORRIENTES UFGRS 1943-1947 MZUSP 83258-83259 Ituza-ingoacute Estacircncia Santa Tecla

Nascimento LB et al 167

Steindachner (1867) established part of Eupemphixnattereri Steindachner 1863 as a junior synonym ofGomphobates marmoratus associated Eupemphixfuscomaculatus to this genus under the new combi-nation Gomphobates fuscomaculatus and indicatedIliobates fuscomaculatus (assigning it to ldquoFitz Ts-chudirdquo) as a synonym of the latter

Cope (1869 ldquo1868rdquo) described Lystris brachyops(= Pleurodema brachyops) and distinguished this ge-nus from Gomphobates by the presence of fontanelaand vomerine teeth and from Eupemphix by the pres-ence of well-developed teeth on young and adultsEupemphix fuscomaculatus Steindachner was thenincluded in the genus Lystris under the combinationLystris fuscomaculatus (Steindachner)

Peters (1872) included Gomphobates marmora-tus Reinhardt and Luumltken Liuperus marmoratus Bur-meister 1861 and part of Eupemphix nattereri Stein-dachner in the synonymy of Bufo albifrons Spix 1824

Boulenger (1882) allocated Pleurodema TschudiCystignathus Dumeacuteril and Bibron Leiuperus Dumeacuteriland Bibron Pleurodema Guumlnther Gomphobates Re-inhardt and Luumltken Eupemphix Steindachner andLystris Cope in the genus Paludicola Wagler basedon the presence of maxillary teeth Consequently thenew combinations Paludicola fuscomaculata (Stein-dachner) and Paludicola albifrons (Spix) were es-tablished for Lystris fuscomaculatus and Bufo albi-frons respectively Boulenger cited the absence ofvomerine teeth in P fuscomaculata (Steindachner) andestablished its distribution range to Brazil Uruguay andBuenos Aires (Argentina) Paludicola biligonigera(Cope) was associated to Reinhardtrsquos specimens fromLagoa Santa (State of Minas Gerais) and from theStates of Bahia and Paraacute Brazil

Boulenger (1886) indicated the occurrence of Palu-dicola fuscomaculata (Steindachner) and Paludico-la albifrons (Spix) in the State of Rio Grande do SulBrazil Boulenger (1887) presented a diagnosis of Palu-dicola albifrons from Porto Alegre Rio Grande doSul Brazil He indicated that Leiuperus marmoratusBurmeister was not identical to Paludicola albifrons(Spix) as stated by Peters (1872) but was a junior syn-onym of P fuscomaculata (Steindachner) In realityLeiuperus marmoratus DrsquoOrbigny 1847 (referred byBurmeister 1861) Leiuperus marmoratus Burmeis-ter 1861 and Paludicola fuscomaculata Boulenger1886 1887 refers to the currently recognized Phys-alaemus biligonigerus (Cope 1861)

Meacutehely (1904) after examining the type designat-ed by Spix distinguished Paludicola albifrons (Spix)from P fuscomaculata (Steindachner) by the pres-ence of a second tarsal tubercle in the articulation be-tween tibia and tarsus on the former larger than thedistal tarsal tubercle and by the absence of externallyvisible lumbar glands He also indicated thatP fuscomaculata (Steindachner) presents no vomer-ine teeth

Miranda-Ribeiro (1926) included Eupemphix En-gystomops Pleurodema and Paludicola in the fam-ily Paludicolidae Paludicola was characterized by thepresence of ldquomore or less indistinct vomerine teethrdquoHe compared specimens of Paludicola albifrons(Spix) and P fuscomaculata (Steindachner) to the typeof Eupemphix fuscomaculatus Steindachner andagreed with Meacutehely (1904) on the difference amongthese species He also mentioned thatP fuscomaculata (Steindachner) presents ldquovomerineteeth indistinct but present on a line anterior to thechoanaerdquo He indicated the distribution of P albifrons(Spix) for the State of Bahia Brazil and that ofP fuscomaculata (Steindachner) for the states of MatoGrosso Satildeo Paulo Paranaacute and Rio Grande do Sul inBrazil and also in Argentina and Uruguay

The first revision of the genus Paludicola waspresented by Parker (1927) who distributed its spe-cies among the genera Physalaemus Fitzinger 1826Pseudopaludicola Miranda-Ribeiro 1926 and Pleu-rodema Paludicola biligonigera (Cope)P fuscomaculata (Steindachner) and P albifrons(Spix) were included in the genus Physalaemus Healso discussed the differences indicated by Meheacutely(1904) between Physalaemus albifrons (Spix) andP fuscomaculatus (Steindachner) Parker (1927) re-moved Gomphobates marmoratus Reinhardt andLuumltken from the synonymy with P albifrons (Spix)and considered it a junior synonym ofP fuscomaculatus (Steindachner) after requestingthe analysis of the type-specimens of P albifrons(Spix)

Cochran (1955) mentioned the absence of vomer-ine teeth in Physalaemus fuscomaculatus (Stein-dachner) ldquoalthough the bony projection on which theywould have arisen can be felt and in a few cases seenbetween the choanaerdquo The description presented wasbased on specimens from the States of Minas GeraisRio Grande do Sul and Satildeo Paulo (Brazil) ArgentinaBolivia and Paraguay

168 Taxonomic status of Gomphobates marmoratus

Milstead (1960) characterized Physalaemus bili-gonigerus (Cope) and P fuscomaculatus (Stein-dachner) species occurring in the State of Rio Grandedo Sul Brazil and pointed that the first differs from allcongeneric species by the presence of vomerine teethHe observed differences on snout-vent length betweenspecimens of P fuscomaculatus (Steindachner) fromthe States of Satildeo Paulo and Rio Grande do Sul sug-gesting that this taxon might include two distinct spe-cies or geographic races

Milstead (1963) after analyzing the holotype ofP biligonigerus (Cope) verified the absence ofvomerine teeth stating that it was a ldquotypical specimenof what is now called P fuscomaculatusrdquo (Stein-dachner) Therefore he considered that the speciestreated by Parker (1927) Cochran (1955) and Mil-stead (1960) as P fuscomaculatus (Steindachner)should be in fact called P biligonigerus (Cope) basedon the precedence of this name He suggested theaplication of the name ldquobiligonigerusrdquo to the smallerspecimens from the south whereas ldquofuscomaculatusrdquoshould be applied to the more widespread northernform

Barrio (1965) agreed with Milstead (1963) that theforms previously identified as Physalaemus fuscomac-ulatus (Steindachner) in Argentina corresponded toP biligonigerus (Cope) but disagreed with the dis-tinction between P biligonigerus (Cope) related tothe southern smaller specimens andP fuscomaculatus (Steindachner) associated to thenorthern larger specimens

Lynch (1970) indicated the absence of vomerineteeth (referred as prevomerine teeth) for all species ofpaludicoline leptodactylids except for Physalaemusbiligonigerus (Cope) which sometimes has them

Frost (1985) mentioned the occurrence of Phys-alaemus fuscomaculatus (Steindachner) in Argenti-na Cei (1987) considered it ldquoquite improbablerdquo andpresented a summarized account on the differencesbetween P biligonigerus (Cope) andP fuscomaculatus (Steindachner) He also indicatedthe absence of vomerine teeth on both forms

Cei (1990) defined and illustrated the taxon Phys-alaemus fuscomaculatus considering that ldquoCaiccedilaraacuterdquohas been misspelled and should be written ldquoCaissaraacuterdquoHe stated that P fuscomaculatus (Steindachner) andP biligonigerus (Cope) were considered as a largecomplex of species of Physalaemus biligonigerus(Cope) since the arrangement of Boulenger (1882)

He pointed out that the morphology of the former spe-cies is inaccurately determined because Stein-dachnerrsquos description of the type clearly indicated thepresence of maxillary and vomerine teeth Physalae-mus fuscomaculatus (Steindachner) was character-ized morphologically based on the holotype and on spec-imens collected at Rio Apa a tributary of Rio Para-guay not far from the type-locality adding informationto Steindachnerrsquos (1863) diagnosis The presence ofvomerine teeth is emphasized as well as the rarity ofthis species in herpetological collections

Nascimento et al (2005) diagnosed the genusPhysalaemus by the presence of quadratojugal bonesand absence of vomerine teeth among other charac-teristics These characters distinguish Physalaemusfrom Pleurodema They also associatedP fuscomaculatus (Steindachner) to the P albifronsspecies group

MATERIAL AND METHODS

Acronyms used in the text are as follow MCNAM(Museu de Ciecircncias Naturais Pontifiacutecia UniversidadeCatoacutelica de Minas Gerais Belo Horizonte Brazil)MNRJ (Museu Nacional Rio de Janeiro Brazil)MZUSP (Museu de Zoologia Universidade de SatildeoPaulo Satildeo Paulo Brazil) NHMW (NaturhistorischesMuseum of Wien Vienna Austria) UFRGS (Univer-sidade Federal do Rio Grande do Sul Porto AlegreBrazil) R and ZMUC (Zoological Museum Universi-ty of Copenhagen Denmark)

Morphometric data follow Cei (1980) and the abre-viations used for are SVL (snout-vent length) HL (headlength) HW (head width) ED (eye diameter) IOD(interorbital distance) UEW (upper eyelid width) END(eye-nostril distance) NSD (nostril-snout distance)IND (internarial distance) UL (upper arm length) AL(arm length) HAL (hand length) TL (thigh length) SL(shank length) FL (foot length from the inner meta-tarsal tubercle to the distal point of fourth toe)

RESULTS AND DISCUSSION

We examined two of the three syntypes of Gom-phobates marmoratus Reinhardt and Luumltken R 1193(formerly ZMUC 22) is in very good conditions all thedistinctive characters of the species are easily ob-served The coloration is quite faded due to the time ithas been preserved Skin on flanks is extensively fold-

Nascimento LB et al 169

ed a feature not observed on living and recently pre-served specimens An incision has been made in thepectoral girdle region but it did not cause too muchdamage Fingers and toes are slightly curved A smalllabel written ldquoType 22rdquo is tied on the right leg R 1194(formerly ZMUC 23) is not well preserved as R 1193It is flaccid and because of that it is not possible toobserve the glands and the coloration is more fadedThe third syntype R 11125 (formerly ZMUC 26) is acleaned skeleton and was not analyzed

We found verified that vomerine teeth and darkocellus on lumbar glands are not present on the typesof Gomphobates marmoratus Reinhardt and LuumltkenSteindachner (1864) diagnosed and Cei (1990) veri-fied the presence of these characters in the holotypeof Eupemphix fuscomaculatus and in additional spec-imens from Rio Apa Paraguay Additionally the pres-ence of vomerine teeth distinguishes Pleurodema fromPhysalaemus (Nascimento et al 2005) We there-fore conclude that Gomphobates marmoratus Rein-hardt and Luumltken should be removed from the synony-my with Physalaemus fuscomaculatus (Steindachner)and recognized as a full species under a new combi-nation Moreover due to the presence of vomerineteeth Eupemphix fuscomaculatus Steindachnershould be allocated to the genus Pleurodema

Species Accounts

Physalaemus marmoratus (Reinhardt and Luumltken1862 ldquo1861rdquo) n comb

Gomphobates marmoratus Reinhardt and Luumltken1862 ldquo1861rdquo

Eupemphix nattereri Steindachner 1863 (part)Paludicola albifrons ndash (not of Spix 1824) Peters

1872 (part)Paludicola biligonigera ndash Boulenger 1882 (part)Paludicola fuscomaculata ndash Meheacutely 1904Physalaemus fuscomaculatus ndash Parker 1927 (part)

Lectotype ndash R 1193 (formerly ZMUC 22) male col-lected in Lagoa Santa State of Minas Gerais Brazil inNovember 1854 by J B Reinhardt

Paralectotype ndash R 1194 (formerly ZMUC 23) malecollected in Lagoa Santa State of Minas Gerais Bra-zil in 04 August 1841 by PW Lund R 11125 (former-ly ZMUC 26) cleaned skeleton collected in Lagoa

Santa State of Minas Gerais Brazil no date by PWLund

Original description of Gomphobates marmoratus ndashldquoThis species seems not only to be much larger thanthe preceding (G notatus) but is plumper about likethe plumper of the Cystignathi The body is thickerand wider the head rounded Although the skin is looseand has many irregular folds the preceding lateral folddoes not seem to be missing besides there are the foldaround the belly the crossfold between the arms andthe inflated throat skin as in the preceding the threeavailable specimens also have openings next to thetongue which lead down to the vocal sacs What con-tributes to giving the species a different make-up isthat there are not so few large warts on the back (oneof them shows that these can disappear almost com-pletely on soft specimens) The differences are reducedto the following the small points on the outer posteriorborder of the tarsus are missing the blade of the footis wider and there is a trace of a fringe around thelarger toes The markings are really to a great extentthe same as on G notatus but much stronger and moredistinct the warts on the back are bordered by darkrings The dark lines characteristic of G notatus andthe 2 round spots on the hind part of the back are miss-ing here A white line which occupies the middle of thehind part of the back is more distinct than the preced-ing species In all other important respects they corre-spond despite the difference in their make-uprdquo (Trans-lation from old Danish by Astrid Schmidt-Nielsen)

Diagnosis ndash A species belonging to the P albifronsspecies group diagnosed by the following combinationof characters (1) size large for the group (SVL formales 333-421 mm females 330-475 mm) (2) bodyrobust (3) head slightly wider than long (4) snoutrounded in dorsal and lateral views (5) canthus ros-tralis rounded (6) vocal sac subgular well developedextending to the border of chest with belly (7) fingersand toes robust fringed with horned tips (8) tarsaltubercle present (9) metatarsal tubercles compressedprotruding with horned distal margins (10) toes webbedon base (11) large inguinal glands with the same colorpattern of dorsum (12) dorsal color pattern presentingan ldquoomegardquo-shaped mark and stripes extending toflanks and inguinal region and (13) presence of a darkbrown interorbital bar Physalaemus marmoratus canbe distinguished from all the other members of the

170 Taxonomic status of Gomphobates marmoratus

P albifrons group due to its larger size (SVL com-bined for males of the other species 282-326 mmfemales 284-341 mm) robust fingers (slender longerfingers in the other species) and dorsal color patternPhysalaemus marmoratus differs from P albifronsand P biligonigerus by the fringed fingers (fringesabsent or poorly marked in P albifrons andP biligonigerus) from P albifrons andP santafecinus by the presence of long rounded orirregular glandular ridges on dorsum (dorsal skin smoothin P albifrons and granulated in P santafecinus)from P albifrons by the presence of one tarsal tuber-cle (two in P albifrons) and large inguinal glands (in-distinct in P albifrons) from P biligonigerus by in-guinal glands with same color pattern of dorsum (in-guinal glands with color pattern darker than dorsum inP biligonigerus) from P santafecinus by the ab-sence of granules on forearms (presence of a line ofgranules on the outer margins of forearms inP santafecinus) and absence of a white longitudinalline on sacral region (evident vertebral line inP santafecinus)

Description of lectotype ndash Body robust head slightlywider than long snout rounded in dorsal (Fig 1) andlateral views (Fig 2) nostrils elliptical not protuber-ant located near the tip of snout oriented dorsolateral-ly canthus rostralis rounded loreal region slightly con-cave eyes protuberant eye diameter larger than in-terorbital distance tympanum indistinct supratympanicfold short slightly marked dorsolateral fold absentvocal sac subgular well developed extending to theborder of chest with belly choanae small oval tonguesmall narrow rounded on posterior border which isnot indented maxillary and premaxillary teeth presentvomerine teeth absent Arms short robust upper armsshorter than forearms outer margin of forearms with-out ridges or granules fingers robust fringed fingertips not expanded horned finger lengths IltIVltIIltIII(Fig 3) nuptial asperities on thumbs and on the medialborder of the inner carpal tubercle but not continuouscarpal tubercles large elliptical with the same sizesupernumerary tubercles protruding conical subartic-ular tubercles single large protruding conical Legsmoderately robust tibia slightly longer than thigh tar-sal fold absent toes slender long slightly fringed (fringeabsent on toe I) toes webbed on base tips not ex-panded horned toe lengths IltIIltVltIIIltIV (Fig 4)tarsal tubercle present metatarsal tubercles large shov-

el-shaped protruding with distal margins horned dis-tance between inner and outer metatarsal tuberclesshorter than that between the former and tarsal tuber-cles supernumerary tubercles absent subarticular tu-bercles single large protruding conical outer marginof subarticular tubercle of toe I weakly horned dorsalpattern presenting an ldquoomegardquo-shaped mark and stripes

Figure 2 Lateral view of the lectotype of Gomphobates marmora-tus Reinhardt and Luumltken 1862 ldquo1861rdquo

Figure 1 Dorsal view of the lectotype of Gomphobates marmora-tus Reinhardt and Luumltken 1862 ldquo1861rdquo

Figure 3 Ventral view of the hand of the lectotype of Gompho-bates marmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Nascimento LB et al 171

extending to flanks and inguinal region dorsum andflanks with long rounded or irregular glandular ridgesupper eyelids finely rugose ventral surfaces smoothexcept on cloacal region and thighs which are cov-ered by large granules ventral disc distinct inguinalgland large ovoid General color pattern in preserva-tive cream light brown blotches and sinuous stripesirregular generally wide bordered by brown lines lightbrown stripes continuous or interrupted tending to forman ldquoomegardquo-shaped mark under scapular region in-terorbital bar dark brown white longitudinal line onsacral region vertical bars dark brown and cream al-ternated from the snout to posterior corner of mouthtransversal bars dark brown and cream alternated ondorsum of arms and legs gular region grayish chestbelly ventral surfaces of arms and legs cream palmarand plantar surfaces grayish over cream backgroundnuptial asperities light brown horned margins of fin-gers toes and metatarsal tubercles dark brown in-guinal gland with the same color pattern of dorsum

Measurements ndash SVL 406 HW 124 HL 112 THL162 TL 173 FL 190 ED 51 ID 36 END 24UEW 39 IND 26

Variation ndash Paralectotype R 1194 shows finger tipsless horned inner border of toe I with a small fringeand subarticular tubercles on toes II III IV and Vhorned Granular ridges granules and inguinal glandsare less evident probably due to preservation Thegeneral color pattern follows that of the lectotype butcolors are more faded Measurements of paralecto-type SVL 385 HW 98 HL 106 THL 150 TL 157FL 170 ED 44 ID 43 END 29 UEW 37 IND

26 Some of the additional specimens examinedshowed a narrower head and more rugose dorsumPalmar and plantar tubercles may be less or morehorned This variation is also observed among tips offingers and toes on a single specimen General colorpattern on dorsum and nuptial asperities of recentlypreserved specimens vary from gray to dark brownthe white longitudinal line on sacral region may be fad-ed Gular region of females follows the color patternof chest and belly Morphometric variation is present-ed in Table 1

Tadpole ndash External morphology and internal oral mor-phology were described by Nomura et al (2003) asPhysalaemus fuscomaculatus

Distribution and ecology ndash Physalaemus marmora-tus breeds in temporary shallow ponds of open habi-tats in the States of Bahia Espiacuterito Santo Rio de Jan-eiro Satildeo Paulo Minas Gerais Goiaacutes Mato Grossoand Mato Grosso do Sul (Brazil) and also in Para-guay Males call floating on the water with the largevocal sac inflated During the axillary amplexus a foamnest is constructed on the water surface where theunpigmented eggs are deposited

Remarks ndash Reinhardt and Luumltken (1862 ldquo1861rdquo) des-ignated three syntypes in the original description ofG marmoratus one was collected by Reinhardt theother two were sent by Dr PW Lund one to theUniversity Museum of Copenhagen and the other tothe ldquoRoyal Museum of Natural Historyrdquo DenmarkLater on all natural history collections in Denmark wereunited into one large collection now ldquoZoological Mu-

Figure 4 Ventral view of the foot of the lectotype of Gomphobatesmarmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Table 1 Mean (x) standard deviation (SD) and range of measure-ments (in mm) of males and females of Physalaemus marmoratus(Reinhardt and Luumltken 1862 ldquo1861rdquo)

Males (N=19) Females (N=13)x SD Range x SD Range

SVL 382 21 333-421 418 42 330-475HL 102 06 92-112 106 07 90-117HW 113 08 10-128 125 10 109-138ED 45 03 40-51 46 05 37-53END 26 02 23-28 27 02 24-31IND 26 01 23-28 27 02 24-31IOD 33 04 27-43 34 04 28-41UEW 39 03 33-46 42 04 37-50THL 157 11 138-176 162 15 139-190TL 155 10 138-173 159 12 136-178FL 174 11 159-190 177 12 156-193

172 Taxonomic status of Gomphobates marmoratus

seum of the University of Copenhagenrdquo (ZMUC MAndersen pers comm) When we consulted theZMUC Curator about the loan of the syntypes he lo-cated two specimens R 1193 (here designated as thelectotype) and R 1194 (here designated as one of theparalectotypes) When asked about the third syntypethe Assistant-Curator could not trace it with certaintybecause none of the three other specimens ofG marmoratus deposited in the ZMUC collection car-ried a label indicating which was the third type How-ever he provided all the information available in thejars labels One specimen (R 1170 formerly ZMUC24) was collected by E Warming which did not col-lect any of the types Specimen R 1195 (formerlyZMUC 25) is just labeled ldquoMinas Geraisrdquo with nocollector and date The last specimen (R 11125 for-merly ZMUC 26 a cleaned skeleton loose bones andskin in alcohol) was collected by PW Lund but thelabel does not indicate the locality and date only aLundrsquos handwriting note saying ldquoNew genus of frogs(1)rdquo We considered this latter as the third syntypebecause of the evidences shown in the label

Pleurodema fuscomaculata (Steindachner 1864)n comb

Eupemphix fuscomaculatus Steindachner 1864Hiobates fuscomaculatus Steindachner 1864Gomphobates fuscomaculatus ndash Steindachner 1867Iliobates fuscomaculatus ndash Steindachner 1867Lystris fuscomaculatus ndash Cope 1869 ldquo1868rdquoPaludicola fuscomaculata ndash Boulenger 1882Physalaemus fuscomaculatus ndash Parker 1927 (part)

Holotype ndash NHMW 4316 female collected by JNatterer in Caiccedilara Municipality of Caacuteceres State ofMato Grosso Brazil (Fig 5 and Fig 6)

A very good description of the holotype of Eupem-phix fuscomaculatus as well as comparisons withother coespecific specimens and notes on distributionis provided by Cei (1990)

Remarks ndash The presence or absence of vomerine teethseems to be variable among genera of the currentlyLeiuperidae (sensu Frost et al 2006) Consequentlyit is not yet possible to determine with certainty forwhat taxon or group of taxa the presence or absenceof vomerine teeth represent a synapomorphic stateThis would require a complete sampling of all species

and all genera involved which is beyond the scope ofthis paper Until this can be accomplished we tenta-tively use the new combination Pleurodema fusco-maculata

RESUMO

A anaacutelise de exemplares depositados em coleccedilotildeesherpetoloacutegicas e identificados como Physalaemus fus-comaculatus demonstrou que estes diferem do tipodeste taacutexon Eupemphix fuscomaculatus Consequumlen-temente o status taxonocircmico de Gomphobates mar-moratus e Eupemphix fuscomaculatus foi revisadocom base nos tipos e na literatura Gomphobatesmarmoratus eacute associado ao gecircnero Physalaemus

Figure 6 Ventral view of the type of Eupemphix fuscomaculatusSteindachner 1864

Figure 5 Dorsal view of the type of Eupemphix fuscomaculatusSteindachner 1864

Nascimento LB et al 173

como Physalaemus marmoratus e se refere aosexemplares previamente identificados como Physala-emus fuscomaculatus Adicionalmente Eupemphixfuscomaculatus eacute associado ao gecircnero Pleurodemacom a combinaccedilatildeo de Pleurodema fuscomaculata

ACKNOWLEDGMENTS

We thank Giovanni Vinciprova (UFRGS) Jens Boslashdtker Ras-mussen (in memorian) and Mogens Andersen (Curator and Assis-tant-Curator of the Zoological Museum University of Copen-hagen Denmark respectively) for providing specimens under theircare Jose P Langone (Museo de Historia Natural de MontevideoUruguay) for providing photos of the type of Eupemphix fusco-maculatus Miguel A Andrade for providing photos of the lecto-type of Gomphobates marmoratus Ceacutelio FB Haddad for criti-cally reading the manuscript Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for the fellowships grant-ed to BVSP CAGC and UC and financial support

LITERATURE CITED

BARRIO A 1965 El genero Physalaemus (Anura Leptodactylidae)en la Argentina Physis 25(70)421-448

BOULENGER GA 1882 Catalogue of Batrachia Salientia siveEcaudata and Batrachia Apoda in the collection of the BritishMuseum 2nd Ed London British Museum xxx + 503p

BOULENGER GA 1886 A synopsis of the reptiles and batrachiansof the Province Rio Grande do Sul Brazil Annals and Magazineof Natural History 18(5)423-445

BOULENGER GA 1887 Description of a new or little-known SouthAmerican frogs of the genera Paludicola and Hyla Annals andMagazine of Natural History 2(5)295-300

CEI JM 1980 Amphibians of Argentina Monitore ZoologicoItaliano (NS) Firenze Monografia 2609p

CEI JM 1987 Additional notes to ldquoAmphibians of Argentinardquo anupdate 1980-1986 Monitore Zoologico Italiano 21209-272

CEI JM 1990 On a paraguayan sample of a long time confusedspecies Physalaemus fuscomaculatus (Steindachner 1864)(Anura Leptodactylidae) Bolletin di Museo Regionale diScienze Naturali di Torino 8(1)215-231

COCHRAN DM 1955 Frogs of Southeastern Brazil United StatesNatural Museum Bulletin 206 xvi + 423p

COPE ED 1869 ldquo1868rdquo Sixth contribution to the herpetology ofTropical America Proceedings of the Academy Natural Sciencesof Philadelphia 20305-313

FROST DR (ED) 1985 Amphibian species of the world Ataxonomic and geographical reference Lawrence Allen PressInc and The Association of Systematics Collections v + 732p

FROST DR 2006 Amphibian Species of the World an OnlineReference Version 40 (17 August 2006) Electronic Databaseaccessible at httpresearchamnhorgherpetologyamphibia

indexhtml American Museum of Natural History New YorkUSA [captured on 20 August 2006]

LYNCH JD 1970 Systematic status of the American leptodactylidfrog genera Engystomops Eupemphix and PhysalaemusCopeia 1970(3)488-496

MEacuteHELY L VON 1904 Investigations on paraguayan batrachiansAnnales Historico-Naturales Musei Nationalis Hungarici2207-232

MILSTEAD WW 1960 Frogs of the genus Physalaemus in SouthernBrazil with description of a new species Copeia1960(2)83-89

MILSTEAD WW 1963 Notes on brazilian frogs of the generaPseudopaludicola and Physalaemus Copeia 1963 (3)565-566

MIRANDA-RIBEIRO A 1926 Notas para servirem ao estudo dosgymnobatrachios (Anura) brasileiros Arquivos do MuseuNacional Rio de Janeiro 271-227

NASCIMENTO LB U CARAMASCHI AND CAG CRUZ 2005Taxonomic review of the species groups of the genusPhysalaemus Fitzinger 1826 with the revalidation of the generaEngystomops Jimeacutenez-de-la-Espada 1872 and EupemphixSteindachner 1863 (Amphibia Anura Leptodactylidae)Arquivos do Museu Nacional Rio de Janeiro 63(2)297-320

NOMURA F DC ROSSA-FERES AND VHM PRADO 2003 Thetadpole of Physalaemus fuscomaculatus (AnuraLeptodactylidae) with a description of internal oralmorphology Zootaxa 3701-8

PARKER HW 1927 A revision of the frogs of the generaPseudopaludicola Physalaemus and Pleurodema Annals andMagazine of Natural History ser 9(20)450-478

PETERS W 1872 Uumlber eine Sammlung von Batrachiern aus Neu-Freiburg in Brasilien Monatsberichte der Akademie derWissenschaften zu Berlin 1872680-684

REINHARDT J AND CF LUumlTKEN 1862 ldquo1861rdquo Bidgrad til Kundskabom Brasiliens Padder og Krybdyr Videnskabelige Meddelelserfra den Naturhistoriske Forening i Kjoslashbenhavn3(10-15)143-242

SPIX JB 1824 Animalia nova sive species novae Testudinum etRanarum quas it itinere per Brazilian annisMDCCCXVIII-MDCCXX jussu et auspiciis MaximillianiJosephi I Bavarrie Regis Monachii xxii + 53p

STEINDACHNER F 1863 Uumlber einige neue Batrachier aus denSammlungen des Wiener Museums Sitzungsberichte derAkademie der Wissenschaften 48186-192

STEINDACHNER F 1864 Batrachologische MitteilungenVerhandelugen Zoologisch-Botanischen Gesellschaft in Wien14239-288

STEINDACHNER F 1867 Amphibien In Reise der OesterreichischenFregatte Novara um die Erde in den Jahren 1857 1858 1859unter den Befehlen des Commodore B von Wuumlllerstorf-UrbairZoologischer Theil 70p

Received 28 April 2006Accepted 16 November 2006

174 Taxonomic status of Gomphobates marmoratus

APPENDIX I

Specimens Examined

Physalaemus albifrons BRAZIL MARANHAtildeO MNRJ 24216-24226 Barreirinha CEARAacute MNRJ 14940-14942 MNRJ 24059 MNRJ 24062-24072 Brejo Santo MNRJ 6636-6674 Mucuripe Fortaleza MNRJ 1125MNRJ 6636-6676 Fortaleza SERGIPE MNRJ 17976-17981 Santa Luzia do Itanhy BAHIA MNRJ 1094-1096Barreiras MNRJ 114-116 MNRJ 1094-1096 MNRJ 1113 MNRJ 1102-1104 MNRJ 1089-1091 Bom Jesus daLapa MNRJ 1105 Juazeiro MZUSP 76521 MZUSP 82303 MINAS GERAIS MNRJ 27180-27189 LageadoMNRJ 27179 MNRJ 21743-21745 Manga MCNAM 206-209 MCNAM 213-216 Porteirinha

Physalaemus biligonigerus BOLIacuteVIA SANTA CRUZ AMNH 144362-144370 AMNH 144371-144385 AMNH144432-144433 BRAZIL SANTA CATARINA MNRJ 31128-31153 RIO GRANDE DO SUL UFGRS 1924-1929 General Cacircmara UFGRS 1776 Uruguaiana UFGRS 1619 Tramandaiacute UFGRS 1319-20 UFRGS 1342Parque Estadual de Itapoatilde Viamatildeo PARAGUAY AMNH 23807 Paraguayan Chaco AMNH 50669-50671Colocircnia Nueva Itaacutelia Villeta URUGUAY DEPARTAMENTO RIO NEGRO MNRJ 28554

Physalaemus marmoratus BRAZIL Bahia MNRJ 28443 MNRJ 28489 Caravelas DISTRITO FEDERALMNRJ 3415 Ribeiratildeo do Torto GoiaacutesMinas Gerais MCNAM 3915-3918 MCNAM 3964-3965 MCNAM 4796region of Queimados Hydroelectric Dam (AHE Queimados) Minas Gerais MCNAM 1259 Bocaiuacuteva MCNAM3155-3157 Conselheiro Mata MNRJ 1020-1021 Lagoa Santa MNRJ 40876-40877 Pirapora MCNAM 2135-2138 MCNAM 2204-2209 MCNAM 2238 MCNAM 2604 MCNAM 2738 MCNAM 2740-2745 MCNAM2748-2751 MCNAM 3147-3148 Santana do Riacho ESPIacuteRITO SANTO MNRJ 40878-40879 Piuacutema RIO DEJANEIRO MNRJ 1015-1019 Campos dos Goytacazes MNRJ 3366 MNRJ 13872-13914 MNRJ 40587 MNRJ40880 MNRJ 40881 Satildeo Joatildeo da Barra MATO GROSSO DO SUL MNRJ 24875 Miranda MNRJ 3070 MNRJ13263 Salobra SAtildeO PAULO MNRJ 40882-40883 Botucatu MNRJ 34666-34671 Campinas MNRJ 37043-37045 Piraju PARAGUAY MNRJ 12710-12730 Asuncioacuten MNRJ 12696-12709 Brejo de Ipuatilde

Physalaemus santafecinus ARGENTINA CORRIENTES UFGRS 1943-1947 MZUSP 83258-83259 Ituza-ingoacute Estacircncia Santa Tecla

168 Taxonomic status of Gomphobates marmoratus

Milstead (1960) characterized Physalaemus bili-gonigerus (Cope) and P fuscomaculatus (Stein-dachner) species occurring in the State of Rio Grandedo Sul Brazil and pointed that the first differs from allcongeneric species by the presence of vomerine teethHe observed differences on snout-vent length betweenspecimens of P fuscomaculatus (Steindachner) fromthe States of Satildeo Paulo and Rio Grande do Sul sug-gesting that this taxon might include two distinct spe-cies or geographic races

Milstead (1963) after analyzing the holotype ofP biligonigerus (Cope) verified the absence ofvomerine teeth stating that it was a ldquotypical specimenof what is now called P fuscomaculatusrdquo (Stein-dachner) Therefore he considered that the speciestreated by Parker (1927) Cochran (1955) and Mil-stead (1960) as P fuscomaculatus (Steindachner)should be in fact called P biligonigerus (Cope) basedon the precedence of this name He suggested theaplication of the name ldquobiligonigerusrdquo to the smallerspecimens from the south whereas ldquofuscomaculatusrdquoshould be applied to the more widespread northernform

Barrio (1965) agreed with Milstead (1963) that theforms previously identified as Physalaemus fuscomac-ulatus (Steindachner) in Argentina corresponded toP biligonigerus (Cope) but disagreed with the dis-tinction between P biligonigerus (Cope) related tothe southern smaller specimens andP fuscomaculatus (Steindachner) associated to thenorthern larger specimens

Lynch (1970) indicated the absence of vomerineteeth (referred as prevomerine teeth) for all species ofpaludicoline leptodactylids except for Physalaemusbiligonigerus (Cope) which sometimes has them

Frost (1985) mentioned the occurrence of Phys-alaemus fuscomaculatus (Steindachner) in Argenti-na Cei (1987) considered it ldquoquite improbablerdquo andpresented a summarized account on the differencesbetween P biligonigerus (Cope) andP fuscomaculatus (Steindachner) He also indicatedthe absence of vomerine teeth on both forms

Cei (1990) defined and illustrated the taxon Phys-alaemus fuscomaculatus considering that ldquoCaiccedilaraacuterdquohas been misspelled and should be written ldquoCaissaraacuterdquoHe stated that P fuscomaculatus (Steindachner) andP biligonigerus (Cope) were considered as a largecomplex of species of Physalaemus biligonigerus(Cope) since the arrangement of Boulenger (1882)

He pointed out that the morphology of the former spe-cies is inaccurately determined because Stein-dachnerrsquos description of the type clearly indicated thepresence of maxillary and vomerine teeth Physalae-mus fuscomaculatus (Steindachner) was character-ized morphologically based on the holotype and on spec-imens collected at Rio Apa a tributary of Rio Para-guay not far from the type-locality adding informationto Steindachnerrsquos (1863) diagnosis The presence ofvomerine teeth is emphasized as well as the rarity ofthis species in herpetological collections

Nascimento et al (2005) diagnosed the genusPhysalaemus by the presence of quadratojugal bonesand absence of vomerine teeth among other charac-teristics These characters distinguish Physalaemusfrom Pleurodema They also associatedP fuscomaculatus (Steindachner) to the P albifronsspecies group

MATERIAL AND METHODS

Acronyms used in the text are as follow MCNAM(Museu de Ciecircncias Naturais Pontifiacutecia UniversidadeCatoacutelica de Minas Gerais Belo Horizonte Brazil)MNRJ (Museu Nacional Rio de Janeiro Brazil)MZUSP (Museu de Zoologia Universidade de SatildeoPaulo Satildeo Paulo Brazil) NHMW (NaturhistorischesMuseum of Wien Vienna Austria) UFRGS (Univer-sidade Federal do Rio Grande do Sul Porto AlegreBrazil) R and ZMUC (Zoological Museum Universi-ty of Copenhagen Denmark)

Morphometric data follow Cei (1980) and the abre-viations used for are SVL (snout-vent length) HL (headlength) HW (head width) ED (eye diameter) IOD(interorbital distance) UEW (upper eyelid width) END(eye-nostril distance) NSD (nostril-snout distance)IND (internarial distance) UL (upper arm length) AL(arm length) HAL (hand length) TL (thigh length) SL(shank length) FL (foot length from the inner meta-tarsal tubercle to the distal point of fourth toe)

RESULTS AND DISCUSSION

We examined two of the three syntypes of Gom-phobates marmoratus Reinhardt and Luumltken R 1193(formerly ZMUC 22) is in very good conditions all thedistinctive characters of the species are easily ob-served The coloration is quite faded due to the time ithas been preserved Skin on flanks is extensively fold-

Nascimento LB et al 169

ed a feature not observed on living and recently pre-served specimens An incision has been made in thepectoral girdle region but it did not cause too muchdamage Fingers and toes are slightly curved A smalllabel written ldquoType 22rdquo is tied on the right leg R 1194(formerly ZMUC 23) is not well preserved as R 1193It is flaccid and because of that it is not possible toobserve the glands and the coloration is more fadedThe third syntype R 11125 (formerly ZMUC 26) is acleaned skeleton and was not analyzed

We found verified that vomerine teeth and darkocellus on lumbar glands are not present on the typesof Gomphobates marmoratus Reinhardt and LuumltkenSteindachner (1864) diagnosed and Cei (1990) veri-fied the presence of these characters in the holotypeof Eupemphix fuscomaculatus and in additional spec-imens from Rio Apa Paraguay Additionally the pres-ence of vomerine teeth distinguishes Pleurodema fromPhysalaemus (Nascimento et al 2005) We there-fore conclude that Gomphobates marmoratus Rein-hardt and Luumltken should be removed from the synony-my with Physalaemus fuscomaculatus (Steindachner)and recognized as a full species under a new combi-nation Moreover due to the presence of vomerineteeth Eupemphix fuscomaculatus Steindachnershould be allocated to the genus Pleurodema

Species Accounts

Physalaemus marmoratus (Reinhardt and Luumltken1862 ldquo1861rdquo) n comb

Gomphobates marmoratus Reinhardt and Luumltken1862 ldquo1861rdquo

Eupemphix nattereri Steindachner 1863 (part)Paludicola albifrons ndash (not of Spix 1824) Peters

1872 (part)Paludicola biligonigera ndash Boulenger 1882 (part)Paludicola fuscomaculata ndash Meheacutely 1904Physalaemus fuscomaculatus ndash Parker 1927 (part)

Lectotype ndash R 1193 (formerly ZMUC 22) male col-lected in Lagoa Santa State of Minas Gerais Brazil inNovember 1854 by J B Reinhardt

Paralectotype ndash R 1194 (formerly ZMUC 23) malecollected in Lagoa Santa State of Minas Gerais Bra-zil in 04 August 1841 by PW Lund R 11125 (former-ly ZMUC 26) cleaned skeleton collected in Lagoa

Santa State of Minas Gerais Brazil no date by PWLund

Original description of Gomphobates marmoratus ndashldquoThis species seems not only to be much larger thanthe preceding (G notatus) but is plumper about likethe plumper of the Cystignathi The body is thickerand wider the head rounded Although the skin is looseand has many irregular folds the preceding lateral folddoes not seem to be missing besides there are the foldaround the belly the crossfold between the arms andthe inflated throat skin as in the preceding the threeavailable specimens also have openings next to thetongue which lead down to the vocal sacs What con-tributes to giving the species a different make-up isthat there are not so few large warts on the back (oneof them shows that these can disappear almost com-pletely on soft specimens) The differences are reducedto the following the small points on the outer posteriorborder of the tarsus are missing the blade of the footis wider and there is a trace of a fringe around thelarger toes The markings are really to a great extentthe same as on G notatus but much stronger and moredistinct the warts on the back are bordered by darkrings The dark lines characteristic of G notatus andthe 2 round spots on the hind part of the back are miss-ing here A white line which occupies the middle of thehind part of the back is more distinct than the preced-ing species In all other important respects they corre-spond despite the difference in their make-uprdquo (Trans-lation from old Danish by Astrid Schmidt-Nielsen)

Diagnosis ndash A species belonging to the P albifronsspecies group diagnosed by the following combinationof characters (1) size large for the group (SVL formales 333-421 mm females 330-475 mm) (2) bodyrobust (3) head slightly wider than long (4) snoutrounded in dorsal and lateral views (5) canthus ros-tralis rounded (6) vocal sac subgular well developedextending to the border of chest with belly (7) fingersand toes robust fringed with horned tips (8) tarsaltubercle present (9) metatarsal tubercles compressedprotruding with horned distal margins (10) toes webbedon base (11) large inguinal glands with the same colorpattern of dorsum (12) dorsal color pattern presentingan ldquoomegardquo-shaped mark and stripes extending toflanks and inguinal region and (13) presence of a darkbrown interorbital bar Physalaemus marmoratus canbe distinguished from all the other members of the

170 Taxonomic status of Gomphobates marmoratus

P albifrons group due to its larger size (SVL com-bined for males of the other species 282-326 mmfemales 284-341 mm) robust fingers (slender longerfingers in the other species) and dorsal color patternPhysalaemus marmoratus differs from P albifronsand P biligonigerus by the fringed fingers (fringesabsent or poorly marked in P albifrons andP biligonigerus) from P albifrons andP santafecinus by the presence of long rounded orirregular glandular ridges on dorsum (dorsal skin smoothin P albifrons and granulated in P santafecinus)from P albifrons by the presence of one tarsal tuber-cle (two in P albifrons) and large inguinal glands (in-distinct in P albifrons) from P biligonigerus by in-guinal glands with same color pattern of dorsum (in-guinal glands with color pattern darker than dorsum inP biligonigerus) from P santafecinus by the ab-sence of granules on forearms (presence of a line ofgranules on the outer margins of forearms inP santafecinus) and absence of a white longitudinalline on sacral region (evident vertebral line inP santafecinus)

Description of lectotype ndash Body robust head slightlywider than long snout rounded in dorsal (Fig 1) andlateral views (Fig 2) nostrils elliptical not protuber-ant located near the tip of snout oriented dorsolateral-ly canthus rostralis rounded loreal region slightly con-cave eyes protuberant eye diameter larger than in-terorbital distance tympanum indistinct supratympanicfold short slightly marked dorsolateral fold absentvocal sac subgular well developed extending to theborder of chest with belly choanae small oval tonguesmall narrow rounded on posterior border which isnot indented maxillary and premaxillary teeth presentvomerine teeth absent Arms short robust upper armsshorter than forearms outer margin of forearms with-out ridges or granules fingers robust fringed fingertips not expanded horned finger lengths IltIVltIIltIII(Fig 3) nuptial asperities on thumbs and on the medialborder of the inner carpal tubercle but not continuouscarpal tubercles large elliptical with the same sizesupernumerary tubercles protruding conical subartic-ular tubercles single large protruding conical Legsmoderately robust tibia slightly longer than thigh tar-sal fold absent toes slender long slightly fringed (fringeabsent on toe I) toes webbed on base tips not ex-panded horned toe lengths IltIIltVltIIIltIV (Fig 4)tarsal tubercle present metatarsal tubercles large shov-

el-shaped protruding with distal margins horned dis-tance between inner and outer metatarsal tuberclesshorter than that between the former and tarsal tuber-cles supernumerary tubercles absent subarticular tu-bercles single large protruding conical outer marginof subarticular tubercle of toe I weakly horned dorsalpattern presenting an ldquoomegardquo-shaped mark and stripes

Figure 2 Lateral view of the lectotype of Gomphobates marmora-tus Reinhardt and Luumltken 1862 ldquo1861rdquo

Figure 1 Dorsal view of the lectotype of Gomphobates marmora-tus Reinhardt and Luumltken 1862 ldquo1861rdquo

Figure 3 Ventral view of the hand of the lectotype of Gompho-bates marmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Nascimento LB et al 171

extending to flanks and inguinal region dorsum andflanks with long rounded or irregular glandular ridgesupper eyelids finely rugose ventral surfaces smoothexcept on cloacal region and thighs which are cov-ered by large granules ventral disc distinct inguinalgland large ovoid General color pattern in preserva-tive cream light brown blotches and sinuous stripesirregular generally wide bordered by brown lines lightbrown stripes continuous or interrupted tending to forman ldquoomegardquo-shaped mark under scapular region in-terorbital bar dark brown white longitudinal line onsacral region vertical bars dark brown and cream al-ternated from the snout to posterior corner of mouthtransversal bars dark brown and cream alternated ondorsum of arms and legs gular region grayish chestbelly ventral surfaces of arms and legs cream palmarand plantar surfaces grayish over cream backgroundnuptial asperities light brown horned margins of fin-gers toes and metatarsal tubercles dark brown in-guinal gland with the same color pattern of dorsum

Measurements ndash SVL 406 HW 124 HL 112 THL162 TL 173 FL 190 ED 51 ID 36 END 24UEW 39 IND 26

Variation ndash Paralectotype R 1194 shows finger tipsless horned inner border of toe I with a small fringeand subarticular tubercles on toes II III IV and Vhorned Granular ridges granules and inguinal glandsare less evident probably due to preservation Thegeneral color pattern follows that of the lectotype butcolors are more faded Measurements of paralecto-type SVL 385 HW 98 HL 106 THL 150 TL 157FL 170 ED 44 ID 43 END 29 UEW 37 IND

26 Some of the additional specimens examinedshowed a narrower head and more rugose dorsumPalmar and plantar tubercles may be less or morehorned This variation is also observed among tips offingers and toes on a single specimen General colorpattern on dorsum and nuptial asperities of recentlypreserved specimens vary from gray to dark brownthe white longitudinal line on sacral region may be fad-ed Gular region of females follows the color patternof chest and belly Morphometric variation is present-ed in Table 1

Tadpole ndash External morphology and internal oral mor-phology were described by Nomura et al (2003) asPhysalaemus fuscomaculatus

Distribution and ecology ndash Physalaemus marmora-tus breeds in temporary shallow ponds of open habi-tats in the States of Bahia Espiacuterito Santo Rio de Jan-eiro Satildeo Paulo Minas Gerais Goiaacutes Mato Grossoand Mato Grosso do Sul (Brazil) and also in Para-guay Males call floating on the water with the largevocal sac inflated During the axillary amplexus a foamnest is constructed on the water surface where theunpigmented eggs are deposited

Remarks ndash Reinhardt and Luumltken (1862 ldquo1861rdquo) des-ignated three syntypes in the original description ofG marmoratus one was collected by Reinhardt theother two were sent by Dr PW Lund one to theUniversity Museum of Copenhagen and the other tothe ldquoRoyal Museum of Natural Historyrdquo DenmarkLater on all natural history collections in Denmark wereunited into one large collection now ldquoZoological Mu-

Figure 4 Ventral view of the foot of the lectotype of Gomphobatesmarmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Table 1 Mean (x) standard deviation (SD) and range of measure-ments (in mm) of males and females of Physalaemus marmoratus(Reinhardt and Luumltken 1862 ldquo1861rdquo)

Males (N=19) Females (N=13)x SD Range x SD Range

SVL 382 21 333-421 418 42 330-475HL 102 06 92-112 106 07 90-117HW 113 08 10-128 125 10 109-138ED 45 03 40-51 46 05 37-53END 26 02 23-28 27 02 24-31IND 26 01 23-28 27 02 24-31IOD 33 04 27-43 34 04 28-41UEW 39 03 33-46 42 04 37-50THL 157 11 138-176 162 15 139-190TL 155 10 138-173 159 12 136-178FL 174 11 159-190 177 12 156-193

172 Taxonomic status of Gomphobates marmoratus

seum of the University of Copenhagenrdquo (ZMUC MAndersen pers comm) When we consulted theZMUC Curator about the loan of the syntypes he lo-cated two specimens R 1193 (here designated as thelectotype) and R 1194 (here designated as one of theparalectotypes) When asked about the third syntypethe Assistant-Curator could not trace it with certaintybecause none of the three other specimens ofG marmoratus deposited in the ZMUC collection car-ried a label indicating which was the third type How-ever he provided all the information available in thejars labels One specimen (R 1170 formerly ZMUC24) was collected by E Warming which did not col-lect any of the types Specimen R 1195 (formerlyZMUC 25) is just labeled ldquoMinas Geraisrdquo with nocollector and date The last specimen (R 11125 for-merly ZMUC 26 a cleaned skeleton loose bones andskin in alcohol) was collected by PW Lund but thelabel does not indicate the locality and date only aLundrsquos handwriting note saying ldquoNew genus of frogs(1)rdquo We considered this latter as the third syntypebecause of the evidences shown in the label

Pleurodema fuscomaculata (Steindachner 1864)n comb

Eupemphix fuscomaculatus Steindachner 1864Hiobates fuscomaculatus Steindachner 1864Gomphobates fuscomaculatus ndash Steindachner 1867Iliobates fuscomaculatus ndash Steindachner 1867Lystris fuscomaculatus ndash Cope 1869 ldquo1868rdquoPaludicola fuscomaculata ndash Boulenger 1882Physalaemus fuscomaculatus ndash Parker 1927 (part)

Holotype ndash NHMW 4316 female collected by JNatterer in Caiccedilara Municipality of Caacuteceres State ofMato Grosso Brazil (Fig 5 and Fig 6)

A very good description of the holotype of Eupem-phix fuscomaculatus as well as comparisons withother coespecific specimens and notes on distributionis provided by Cei (1990)

Remarks ndash The presence or absence of vomerine teethseems to be variable among genera of the currentlyLeiuperidae (sensu Frost et al 2006) Consequentlyit is not yet possible to determine with certainty forwhat taxon or group of taxa the presence or absenceof vomerine teeth represent a synapomorphic stateThis would require a complete sampling of all species

and all genera involved which is beyond the scope ofthis paper Until this can be accomplished we tenta-tively use the new combination Pleurodema fusco-maculata

RESUMO

A anaacutelise de exemplares depositados em coleccedilotildeesherpetoloacutegicas e identificados como Physalaemus fus-comaculatus demonstrou que estes diferem do tipodeste taacutexon Eupemphix fuscomaculatus Consequumlen-temente o status taxonocircmico de Gomphobates mar-moratus e Eupemphix fuscomaculatus foi revisadocom base nos tipos e na literatura Gomphobatesmarmoratus eacute associado ao gecircnero Physalaemus

Figure 6 Ventral view of the type of Eupemphix fuscomaculatusSteindachner 1864

Figure 5 Dorsal view of the type of Eupemphix fuscomaculatusSteindachner 1864

Nascimento LB et al 173

como Physalaemus marmoratus e se refere aosexemplares previamente identificados como Physala-emus fuscomaculatus Adicionalmente Eupemphixfuscomaculatus eacute associado ao gecircnero Pleurodemacom a combinaccedilatildeo de Pleurodema fuscomaculata

ACKNOWLEDGMENTS

We thank Giovanni Vinciprova (UFRGS) Jens Boslashdtker Ras-mussen (in memorian) and Mogens Andersen (Curator and Assis-tant-Curator of the Zoological Museum University of Copen-hagen Denmark respectively) for providing specimens under theircare Jose P Langone (Museo de Historia Natural de MontevideoUruguay) for providing photos of the type of Eupemphix fusco-maculatus Miguel A Andrade for providing photos of the lecto-type of Gomphobates marmoratus Ceacutelio FB Haddad for criti-cally reading the manuscript Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for the fellowships grant-ed to BVSP CAGC and UC and financial support

LITERATURE CITED

BARRIO A 1965 El genero Physalaemus (Anura Leptodactylidae)en la Argentina Physis 25(70)421-448

BOULENGER GA 1882 Catalogue of Batrachia Salientia siveEcaudata and Batrachia Apoda in the collection of the BritishMuseum 2nd Ed London British Museum xxx + 503p

BOULENGER GA 1886 A synopsis of the reptiles and batrachiansof the Province Rio Grande do Sul Brazil Annals and Magazineof Natural History 18(5)423-445

BOULENGER GA 1887 Description of a new or little-known SouthAmerican frogs of the genera Paludicola and Hyla Annals andMagazine of Natural History 2(5)295-300

CEI JM 1980 Amphibians of Argentina Monitore ZoologicoItaliano (NS) Firenze Monografia 2609p

CEI JM 1987 Additional notes to ldquoAmphibians of Argentinardquo anupdate 1980-1986 Monitore Zoologico Italiano 21209-272

CEI JM 1990 On a paraguayan sample of a long time confusedspecies Physalaemus fuscomaculatus (Steindachner 1864)(Anura Leptodactylidae) Bolletin di Museo Regionale diScienze Naturali di Torino 8(1)215-231

COCHRAN DM 1955 Frogs of Southeastern Brazil United StatesNatural Museum Bulletin 206 xvi + 423p

COPE ED 1869 ldquo1868rdquo Sixth contribution to the herpetology ofTropical America Proceedings of the Academy Natural Sciencesof Philadelphia 20305-313

FROST DR (ED) 1985 Amphibian species of the world Ataxonomic and geographical reference Lawrence Allen PressInc and The Association of Systematics Collections v + 732p

FROST DR 2006 Amphibian Species of the World an OnlineReference Version 40 (17 August 2006) Electronic Databaseaccessible at httpresearchamnhorgherpetologyamphibia

indexhtml American Museum of Natural History New YorkUSA [captured on 20 August 2006]

LYNCH JD 1970 Systematic status of the American leptodactylidfrog genera Engystomops Eupemphix and PhysalaemusCopeia 1970(3)488-496

MEacuteHELY L VON 1904 Investigations on paraguayan batrachiansAnnales Historico-Naturales Musei Nationalis Hungarici2207-232

MILSTEAD WW 1960 Frogs of the genus Physalaemus in SouthernBrazil with description of a new species Copeia1960(2)83-89

MILSTEAD WW 1963 Notes on brazilian frogs of the generaPseudopaludicola and Physalaemus Copeia 1963 (3)565-566

MIRANDA-RIBEIRO A 1926 Notas para servirem ao estudo dosgymnobatrachios (Anura) brasileiros Arquivos do MuseuNacional Rio de Janeiro 271-227

NASCIMENTO LB U CARAMASCHI AND CAG CRUZ 2005Taxonomic review of the species groups of the genusPhysalaemus Fitzinger 1826 with the revalidation of the generaEngystomops Jimeacutenez-de-la-Espada 1872 and EupemphixSteindachner 1863 (Amphibia Anura Leptodactylidae)Arquivos do Museu Nacional Rio de Janeiro 63(2)297-320

NOMURA F DC ROSSA-FERES AND VHM PRADO 2003 Thetadpole of Physalaemus fuscomaculatus (AnuraLeptodactylidae) with a description of internal oralmorphology Zootaxa 3701-8

PARKER HW 1927 A revision of the frogs of the generaPseudopaludicola Physalaemus and Pleurodema Annals andMagazine of Natural History ser 9(20)450-478

PETERS W 1872 Uumlber eine Sammlung von Batrachiern aus Neu-Freiburg in Brasilien Monatsberichte der Akademie derWissenschaften zu Berlin 1872680-684

REINHARDT J AND CF LUumlTKEN 1862 ldquo1861rdquo Bidgrad til Kundskabom Brasiliens Padder og Krybdyr Videnskabelige Meddelelserfra den Naturhistoriske Forening i Kjoslashbenhavn3(10-15)143-242

SPIX JB 1824 Animalia nova sive species novae Testudinum etRanarum quas it itinere per Brazilian annisMDCCCXVIII-MDCCXX jussu et auspiciis MaximillianiJosephi I Bavarrie Regis Monachii xxii + 53p

STEINDACHNER F 1863 Uumlber einige neue Batrachier aus denSammlungen des Wiener Museums Sitzungsberichte derAkademie der Wissenschaften 48186-192

STEINDACHNER F 1864 Batrachologische MitteilungenVerhandelugen Zoologisch-Botanischen Gesellschaft in Wien14239-288

STEINDACHNER F 1867 Amphibien In Reise der OesterreichischenFregatte Novara um die Erde in den Jahren 1857 1858 1859unter den Befehlen des Commodore B von Wuumlllerstorf-UrbairZoologischer Theil 70p

Received 28 April 2006Accepted 16 November 2006

174 Taxonomic status of Gomphobates marmoratus

APPENDIX I

Specimens Examined

Physalaemus albifrons BRAZIL MARANHAtildeO MNRJ 24216-24226 Barreirinha CEARAacute MNRJ 14940-14942 MNRJ 24059 MNRJ 24062-24072 Brejo Santo MNRJ 6636-6674 Mucuripe Fortaleza MNRJ 1125MNRJ 6636-6676 Fortaleza SERGIPE MNRJ 17976-17981 Santa Luzia do Itanhy BAHIA MNRJ 1094-1096Barreiras MNRJ 114-116 MNRJ 1094-1096 MNRJ 1113 MNRJ 1102-1104 MNRJ 1089-1091 Bom Jesus daLapa MNRJ 1105 Juazeiro MZUSP 76521 MZUSP 82303 MINAS GERAIS MNRJ 27180-27189 LageadoMNRJ 27179 MNRJ 21743-21745 Manga MCNAM 206-209 MCNAM 213-216 Porteirinha

Physalaemus biligonigerus BOLIacuteVIA SANTA CRUZ AMNH 144362-144370 AMNH 144371-144385 AMNH144432-144433 BRAZIL SANTA CATARINA MNRJ 31128-31153 RIO GRANDE DO SUL UFGRS 1924-1929 General Cacircmara UFGRS 1776 Uruguaiana UFGRS 1619 Tramandaiacute UFGRS 1319-20 UFRGS 1342Parque Estadual de Itapoatilde Viamatildeo PARAGUAY AMNH 23807 Paraguayan Chaco AMNH 50669-50671Colocircnia Nueva Itaacutelia Villeta URUGUAY DEPARTAMENTO RIO NEGRO MNRJ 28554

Physalaemus marmoratus BRAZIL Bahia MNRJ 28443 MNRJ 28489 Caravelas DISTRITO FEDERALMNRJ 3415 Ribeiratildeo do Torto GoiaacutesMinas Gerais MCNAM 3915-3918 MCNAM 3964-3965 MCNAM 4796region of Queimados Hydroelectric Dam (AHE Queimados) Minas Gerais MCNAM 1259 Bocaiuacuteva MCNAM3155-3157 Conselheiro Mata MNRJ 1020-1021 Lagoa Santa MNRJ 40876-40877 Pirapora MCNAM 2135-2138 MCNAM 2204-2209 MCNAM 2238 MCNAM 2604 MCNAM 2738 MCNAM 2740-2745 MCNAM2748-2751 MCNAM 3147-3148 Santana do Riacho ESPIacuteRITO SANTO MNRJ 40878-40879 Piuacutema RIO DEJANEIRO MNRJ 1015-1019 Campos dos Goytacazes MNRJ 3366 MNRJ 13872-13914 MNRJ 40587 MNRJ40880 MNRJ 40881 Satildeo Joatildeo da Barra MATO GROSSO DO SUL MNRJ 24875 Miranda MNRJ 3070 MNRJ13263 Salobra SAtildeO PAULO MNRJ 40882-40883 Botucatu MNRJ 34666-34671 Campinas MNRJ 37043-37045 Piraju PARAGUAY MNRJ 12710-12730 Asuncioacuten MNRJ 12696-12709 Brejo de Ipuatilde

Physalaemus santafecinus ARGENTINA CORRIENTES UFGRS 1943-1947 MZUSP 83258-83259 Ituza-ingoacute Estacircncia Santa Tecla

Nascimento LB et al 169

ed a feature not observed on living and recently pre-served specimens An incision has been made in thepectoral girdle region but it did not cause too muchdamage Fingers and toes are slightly curved A smalllabel written ldquoType 22rdquo is tied on the right leg R 1194(formerly ZMUC 23) is not well preserved as R 1193It is flaccid and because of that it is not possible toobserve the glands and the coloration is more fadedThe third syntype R 11125 (formerly ZMUC 26) is acleaned skeleton and was not analyzed

We found verified that vomerine teeth and darkocellus on lumbar glands are not present on the typesof Gomphobates marmoratus Reinhardt and LuumltkenSteindachner (1864) diagnosed and Cei (1990) veri-fied the presence of these characters in the holotypeof Eupemphix fuscomaculatus and in additional spec-imens from Rio Apa Paraguay Additionally the pres-ence of vomerine teeth distinguishes Pleurodema fromPhysalaemus (Nascimento et al 2005) We there-fore conclude that Gomphobates marmoratus Rein-hardt and Luumltken should be removed from the synony-my with Physalaemus fuscomaculatus (Steindachner)and recognized as a full species under a new combi-nation Moreover due to the presence of vomerineteeth Eupemphix fuscomaculatus Steindachnershould be allocated to the genus Pleurodema

Species Accounts

Physalaemus marmoratus (Reinhardt and Luumltken1862 ldquo1861rdquo) n comb

Gomphobates marmoratus Reinhardt and Luumltken1862 ldquo1861rdquo

Eupemphix nattereri Steindachner 1863 (part)Paludicola albifrons ndash (not of Spix 1824) Peters

1872 (part)Paludicola biligonigera ndash Boulenger 1882 (part)Paludicola fuscomaculata ndash Meheacutely 1904Physalaemus fuscomaculatus ndash Parker 1927 (part)

Lectotype ndash R 1193 (formerly ZMUC 22) male col-lected in Lagoa Santa State of Minas Gerais Brazil inNovember 1854 by J B Reinhardt

Paralectotype ndash R 1194 (formerly ZMUC 23) malecollected in Lagoa Santa State of Minas Gerais Bra-zil in 04 August 1841 by PW Lund R 11125 (former-ly ZMUC 26) cleaned skeleton collected in Lagoa

Santa State of Minas Gerais Brazil no date by PWLund

Original description of Gomphobates marmoratus ndashldquoThis species seems not only to be much larger thanthe preceding (G notatus) but is plumper about likethe plumper of the Cystignathi The body is thickerand wider the head rounded Although the skin is looseand has many irregular folds the preceding lateral folddoes not seem to be missing besides there are the foldaround the belly the crossfold between the arms andthe inflated throat skin as in the preceding the threeavailable specimens also have openings next to thetongue which lead down to the vocal sacs What con-tributes to giving the species a different make-up isthat there are not so few large warts on the back (oneof them shows that these can disappear almost com-pletely on soft specimens) The differences are reducedto the following the small points on the outer posteriorborder of the tarsus are missing the blade of the footis wider and there is a trace of a fringe around thelarger toes The markings are really to a great extentthe same as on G notatus but much stronger and moredistinct the warts on the back are bordered by darkrings The dark lines characteristic of G notatus andthe 2 round spots on the hind part of the back are miss-ing here A white line which occupies the middle of thehind part of the back is more distinct than the preced-ing species In all other important respects they corre-spond despite the difference in their make-uprdquo (Trans-lation from old Danish by Astrid Schmidt-Nielsen)

Diagnosis ndash A species belonging to the P albifronsspecies group diagnosed by the following combinationof characters (1) size large for the group (SVL formales 333-421 mm females 330-475 mm) (2) bodyrobust (3) head slightly wider than long (4) snoutrounded in dorsal and lateral views (5) canthus ros-tralis rounded (6) vocal sac subgular well developedextending to the border of chest with belly (7) fingersand toes robust fringed with horned tips (8) tarsaltubercle present (9) metatarsal tubercles compressedprotruding with horned distal margins (10) toes webbedon base (11) large inguinal glands with the same colorpattern of dorsum (12) dorsal color pattern presentingan ldquoomegardquo-shaped mark and stripes extending toflanks and inguinal region and (13) presence of a darkbrown interorbital bar Physalaemus marmoratus canbe distinguished from all the other members of the

170 Taxonomic status of Gomphobates marmoratus

P albifrons group due to its larger size (SVL com-bined for males of the other species 282-326 mmfemales 284-341 mm) robust fingers (slender longerfingers in the other species) and dorsal color patternPhysalaemus marmoratus differs from P albifronsand P biligonigerus by the fringed fingers (fringesabsent or poorly marked in P albifrons andP biligonigerus) from P albifrons andP santafecinus by the presence of long rounded orirregular glandular ridges on dorsum (dorsal skin smoothin P albifrons and granulated in P santafecinus)from P albifrons by the presence of one tarsal tuber-cle (two in P albifrons) and large inguinal glands (in-distinct in P albifrons) from P biligonigerus by in-guinal glands with same color pattern of dorsum (in-guinal glands with color pattern darker than dorsum inP biligonigerus) from P santafecinus by the ab-sence of granules on forearms (presence of a line ofgranules on the outer margins of forearms inP santafecinus) and absence of a white longitudinalline on sacral region (evident vertebral line inP santafecinus)

Description of lectotype ndash Body robust head slightlywider than long snout rounded in dorsal (Fig 1) andlateral views (Fig 2) nostrils elliptical not protuber-ant located near the tip of snout oriented dorsolateral-ly canthus rostralis rounded loreal region slightly con-cave eyes protuberant eye diameter larger than in-terorbital distance tympanum indistinct supratympanicfold short slightly marked dorsolateral fold absentvocal sac subgular well developed extending to theborder of chest with belly choanae small oval tonguesmall narrow rounded on posterior border which isnot indented maxillary and premaxillary teeth presentvomerine teeth absent Arms short robust upper armsshorter than forearms outer margin of forearms with-out ridges or granules fingers robust fringed fingertips not expanded horned finger lengths IltIVltIIltIII(Fig 3) nuptial asperities on thumbs and on the medialborder of the inner carpal tubercle but not continuouscarpal tubercles large elliptical with the same sizesupernumerary tubercles protruding conical subartic-ular tubercles single large protruding conical Legsmoderately robust tibia slightly longer than thigh tar-sal fold absent toes slender long slightly fringed (fringeabsent on toe I) toes webbed on base tips not ex-panded horned toe lengths IltIIltVltIIIltIV (Fig 4)tarsal tubercle present metatarsal tubercles large shov-

el-shaped protruding with distal margins horned dis-tance between inner and outer metatarsal tuberclesshorter than that between the former and tarsal tuber-cles supernumerary tubercles absent subarticular tu-bercles single large protruding conical outer marginof subarticular tubercle of toe I weakly horned dorsalpattern presenting an ldquoomegardquo-shaped mark and stripes

Figure 2 Lateral view of the lectotype of Gomphobates marmora-tus Reinhardt and Luumltken 1862 ldquo1861rdquo

Figure 1 Dorsal view of the lectotype of Gomphobates marmora-tus Reinhardt and Luumltken 1862 ldquo1861rdquo

Figure 3 Ventral view of the hand of the lectotype of Gompho-bates marmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Nascimento LB et al 171

extending to flanks and inguinal region dorsum andflanks with long rounded or irregular glandular ridgesupper eyelids finely rugose ventral surfaces smoothexcept on cloacal region and thighs which are cov-ered by large granules ventral disc distinct inguinalgland large ovoid General color pattern in preserva-tive cream light brown blotches and sinuous stripesirregular generally wide bordered by brown lines lightbrown stripes continuous or interrupted tending to forman ldquoomegardquo-shaped mark under scapular region in-terorbital bar dark brown white longitudinal line onsacral region vertical bars dark brown and cream al-ternated from the snout to posterior corner of mouthtransversal bars dark brown and cream alternated ondorsum of arms and legs gular region grayish chestbelly ventral surfaces of arms and legs cream palmarand plantar surfaces grayish over cream backgroundnuptial asperities light brown horned margins of fin-gers toes and metatarsal tubercles dark brown in-guinal gland with the same color pattern of dorsum

Measurements ndash SVL 406 HW 124 HL 112 THL162 TL 173 FL 190 ED 51 ID 36 END 24UEW 39 IND 26

Variation ndash Paralectotype R 1194 shows finger tipsless horned inner border of toe I with a small fringeand subarticular tubercles on toes II III IV and Vhorned Granular ridges granules and inguinal glandsare less evident probably due to preservation Thegeneral color pattern follows that of the lectotype butcolors are more faded Measurements of paralecto-type SVL 385 HW 98 HL 106 THL 150 TL 157FL 170 ED 44 ID 43 END 29 UEW 37 IND

26 Some of the additional specimens examinedshowed a narrower head and more rugose dorsumPalmar and plantar tubercles may be less or morehorned This variation is also observed among tips offingers and toes on a single specimen General colorpattern on dorsum and nuptial asperities of recentlypreserved specimens vary from gray to dark brownthe white longitudinal line on sacral region may be fad-ed Gular region of females follows the color patternof chest and belly Morphometric variation is present-ed in Table 1

Tadpole ndash External morphology and internal oral mor-phology were described by Nomura et al (2003) asPhysalaemus fuscomaculatus

Distribution and ecology ndash Physalaemus marmora-tus breeds in temporary shallow ponds of open habi-tats in the States of Bahia Espiacuterito Santo Rio de Jan-eiro Satildeo Paulo Minas Gerais Goiaacutes Mato Grossoand Mato Grosso do Sul (Brazil) and also in Para-guay Males call floating on the water with the largevocal sac inflated During the axillary amplexus a foamnest is constructed on the water surface where theunpigmented eggs are deposited

Remarks ndash Reinhardt and Luumltken (1862 ldquo1861rdquo) des-ignated three syntypes in the original description ofG marmoratus one was collected by Reinhardt theother two were sent by Dr PW Lund one to theUniversity Museum of Copenhagen and the other tothe ldquoRoyal Museum of Natural Historyrdquo DenmarkLater on all natural history collections in Denmark wereunited into one large collection now ldquoZoological Mu-

Figure 4 Ventral view of the foot of the lectotype of Gomphobatesmarmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Table 1 Mean (x) standard deviation (SD) and range of measure-ments (in mm) of males and females of Physalaemus marmoratus(Reinhardt and Luumltken 1862 ldquo1861rdquo)

Males (N=19) Females (N=13)x SD Range x SD Range

SVL 382 21 333-421 418 42 330-475HL 102 06 92-112 106 07 90-117HW 113 08 10-128 125 10 109-138ED 45 03 40-51 46 05 37-53END 26 02 23-28 27 02 24-31IND 26 01 23-28 27 02 24-31IOD 33 04 27-43 34 04 28-41UEW 39 03 33-46 42 04 37-50THL 157 11 138-176 162 15 139-190TL 155 10 138-173 159 12 136-178FL 174 11 159-190 177 12 156-193

172 Taxonomic status of Gomphobates marmoratus

seum of the University of Copenhagenrdquo (ZMUC MAndersen pers comm) When we consulted theZMUC Curator about the loan of the syntypes he lo-cated two specimens R 1193 (here designated as thelectotype) and R 1194 (here designated as one of theparalectotypes) When asked about the third syntypethe Assistant-Curator could not trace it with certaintybecause none of the three other specimens ofG marmoratus deposited in the ZMUC collection car-ried a label indicating which was the third type How-ever he provided all the information available in thejars labels One specimen (R 1170 formerly ZMUC24) was collected by E Warming which did not col-lect any of the types Specimen R 1195 (formerlyZMUC 25) is just labeled ldquoMinas Geraisrdquo with nocollector and date The last specimen (R 11125 for-merly ZMUC 26 a cleaned skeleton loose bones andskin in alcohol) was collected by PW Lund but thelabel does not indicate the locality and date only aLundrsquos handwriting note saying ldquoNew genus of frogs(1)rdquo We considered this latter as the third syntypebecause of the evidences shown in the label

Pleurodema fuscomaculata (Steindachner 1864)n comb

Eupemphix fuscomaculatus Steindachner 1864Hiobates fuscomaculatus Steindachner 1864Gomphobates fuscomaculatus ndash Steindachner 1867Iliobates fuscomaculatus ndash Steindachner 1867Lystris fuscomaculatus ndash Cope 1869 ldquo1868rdquoPaludicola fuscomaculata ndash Boulenger 1882Physalaemus fuscomaculatus ndash Parker 1927 (part)

Holotype ndash NHMW 4316 female collected by JNatterer in Caiccedilara Municipality of Caacuteceres State ofMato Grosso Brazil (Fig 5 and Fig 6)

A very good description of the holotype of Eupem-phix fuscomaculatus as well as comparisons withother coespecific specimens and notes on distributionis provided by Cei (1990)

Remarks ndash The presence or absence of vomerine teethseems to be variable among genera of the currentlyLeiuperidae (sensu Frost et al 2006) Consequentlyit is not yet possible to determine with certainty forwhat taxon or group of taxa the presence or absenceof vomerine teeth represent a synapomorphic stateThis would require a complete sampling of all species

and all genera involved which is beyond the scope ofthis paper Until this can be accomplished we tenta-tively use the new combination Pleurodema fusco-maculata

RESUMO

A anaacutelise de exemplares depositados em coleccedilotildeesherpetoloacutegicas e identificados como Physalaemus fus-comaculatus demonstrou que estes diferem do tipodeste taacutexon Eupemphix fuscomaculatus Consequumlen-temente o status taxonocircmico de Gomphobates mar-moratus e Eupemphix fuscomaculatus foi revisadocom base nos tipos e na literatura Gomphobatesmarmoratus eacute associado ao gecircnero Physalaemus

Figure 6 Ventral view of the type of Eupemphix fuscomaculatusSteindachner 1864

Figure 5 Dorsal view of the type of Eupemphix fuscomaculatusSteindachner 1864

Nascimento LB et al 173

como Physalaemus marmoratus e se refere aosexemplares previamente identificados como Physala-emus fuscomaculatus Adicionalmente Eupemphixfuscomaculatus eacute associado ao gecircnero Pleurodemacom a combinaccedilatildeo de Pleurodema fuscomaculata

ACKNOWLEDGMENTS

We thank Giovanni Vinciprova (UFRGS) Jens Boslashdtker Ras-mussen (in memorian) and Mogens Andersen (Curator and Assis-tant-Curator of the Zoological Museum University of Copen-hagen Denmark respectively) for providing specimens under theircare Jose P Langone (Museo de Historia Natural de MontevideoUruguay) for providing photos of the type of Eupemphix fusco-maculatus Miguel A Andrade for providing photos of the lecto-type of Gomphobates marmoratus Ceacutelio FB Haddad for criti-cally reading the manuscript Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for the fellowships grant-ed to BVSP CAGC and UC and financial support

LITERATURE CITED

BARRIO A 1965 El genero Physalaemus (Anura Leptodactylidae)en la Argentina Physis 25(70)421-448

BOULENGER GA 1882 Catalogue of Batrachia Salientia siveEcaudata and Batrachia Apoda in the collection of the BritishMuseum 2nd Ed London British Museum xxx + 503p

BOULENGER GA 1886 A synopsis of the reptiles and batrachiansof the Province Rio Grande do Sul Brazil Annals and Magazineof Natural History 18(5)423-445

BOULENGER GA 1887 Description of a new or little-known SouthAmerican frogs of the genera Paludicola and Hyla Annals andMagazine of Natural History 2(5)295-300

CEI JM 1980 Amphibians of Argentina Monitore ZoologicoItaliano (NS) Firenze Monografia 2609p

CEI JM 1987 Additional notes to ldquoAmphibians of Argentinardquo anupdate 1980-1986 Monitore Zoologico Italiano 21209-272

CEI JM 1990 On a paraguayan sample of a long time confusedspecies Physalaemus fuscomaculatus (Steindachner 1864)(Anura Leptodactylidae) Bolletin di Museo Regionale diScienze Naturali di Torino 8(1)215-231

COCHRAN DM 1955 Frogs of Southeastern Brazil United StatesNatural Museum Bulletin 206 xvi + 423p

COPE ED 1869 ldquo1868rdquo Sixth contribution to the herpetology ofTropical America Proceedings of the Academy Natural Sciencesof Philadelphia 20305-313

FROST DR (ED) 1985 Amphibian species of the world Ataxonomic and geographical reference Lawrence Allen PressInc and The Association of Systematics Collections v + 732p

FROST DR 2006 Amphibian Species of the World an OnlineReference Version 40 (17 August 2006) Electronic Databaseaccessible at httpresearchamnhorgherpetologyamphibia

indexhtml American Museum of Natural History New YorkUSA [captured on 20 August 2006]

LYNCH JD 1970 Systematic status of the American leptodactylidfrog genera Engystomops Eupemphix and PhysalaemusCopeia 1970(3)488-496

MEacuteHELY L VON 1904 Investigations on paraguayan batrachiansAnnales Historico-Naturales Musei Nationalis Hungarici2207-232

MILSTEAD WW 1960 Frogs of the genus Physalaemus in SouthernBrazil with description of a new species Copeia1960(2)83-89

MILSTEAD WW 1963 Notes on brazilian frogs of the generaPseudopaludicola and Physalaemus Copeia 1963 (3)565-566

MIRANDA-RIBEIRO A 1926 Notas para servirem ao estudo dosgymnobatrachios (Anura) brasileiros Arquivos do MuseuNacional Rio de Janeiro 271-227

NASCIMENTO LB U CARAMASCHI AND CAG CRUZ 2005Taxonomic review of the species groups of the genusPhysalaemus Fitzinger 1826 with the revalidation of the generaEngystomops Jimeacutenez-de-la-Espada 1872 and EupemphixSteindachner 1863 (Amphibia Anura Leptodactylidae)Arquivos do Museu Nacional Rio de Janeiro 63(2)297-320

NOMURA F DC ROSSA-FERES AND VHM PRADO 2003 Thetadpole of Physalaemus fuscomaculatus (AnuraLeptodactylidae) with a description of internal oralmorphology Zootaxa 3701-8

PARKER HW 1927 A revision of the frogs of the generaPseudopaludicola Physalaemus and Pleurodema Annals andMagazine of Natural History ser 9(20)450-478

PETERS W 1872 Uumlber eine Sammlung von Batrachiern aus Neu-Freiburg in Brasilien Monatsberichte der Akademie derWissenschaften zu Berlin 1872680-684

REINHARDT J AND CF LUumlTKEN 1862 ldquo1861rdquo Bidgrad til Kundskabom Brasiliens Padder og Krybdyr Videnskabelige Meddelelserfra den Naturhistoriske Forening i Kjoslashbenhavn3(10-15)143-242

SPIX JB 1824 Animalia nova sive species novae Testudinum etRanarum quas it itinere per Brazilian annisMDCCCXVIII-MDCCXX jussu et auspiciis MaximillianiJosephi I Bavarrie Regis Monachii xxii + 53p

STEINDACHNER F 1863 Uumlber einige neue Batrachier aus denSammlungen des Wiener Museums Sitzungsberichte derAkademie der Wissenschaften 48186-192

STEINDACHNER F 1864 Batrachologische MitteilungenVerhandelugen Zoologisch-Botanischen Gesellschaft in Wien14239-288

STEINDACHNER F 1867 Amphibien In Reise der OesterreichischenFregatte Novara um die Erde in den Jahren 1857 1858 1859unter den Befehlen des Commodore B von Wuumlllerstorf-UrbairZoologischer Theil 70p

Received 28 April 2006Accepted 16 November 2006

174 Taxonomic status of Gomphobates marmoratus

APPENDIX I

Specimens Examined

Physalaemus albifrons BRAZIL MARANHAtildeO MNRJ 24216-24226 Barreirinha CEARAacute MNRJ 14940-14942 MNRJ 24059 MNRJ 24062-24072 Brejo Santo MNRJ 6636-6674 Mucuripe Fortaleza MNRJ 1125MNRJ 6636-6676 Fortaleza SERGIPE MNRJ 17976-17981 Santa Luzia do Itanhy BAHIA MNRJ 1094-1096Barreiras MNRJ 114-116 MNRJ 1094-1096 MNRJ 1113 MNRJ 1102-1104 MNRJ 1089-1091 Bom Jesus daLapa MNRJ 1105 Juazeiro MZUSP 76521 MZUSP 82303 MINAS GERAIS MNRJ 27180-27189 LageadoMNRJ 27179 MNRJ 21743-21745 Manga MCNAM 206-209 MCNAM 213-216 Porteirinha

Physalaemus biligonigerus BOLIacuteVIA SANTA CRUZ AMNH 144362-144370 AMNH 144371-144385 AMNH144432-144433 BRAZIL SANTA CATARINA MNRJ 31128-31153 RIO GRANDE DO SUL UFGRS 1924-1929 General Cacircmara UFGRS 1776 Uruguaiana UFGRS 1619 Tramandaiacute UFGRS 1319-20 UFRGS 1342Parque Estadual de Itapoatilde Viamatildeo PARAGUAY AMNH 23807 Paraguayan Chaco AMNH 50669-50671Colocircnia Nueva Itaacutelia Villeta URUGUAY DEPARTAMENTO RIO NEGRO MNRJ 28554

Physalaemus marmoratus BRAZIL Bahia MNRJ 28443 MNRJ 28489 Caravelas DISTRITO FEDERALMNRJ 3415 Ribeiratildeo do Torto GoiaacutesMinas Gerais MCNAM 3915-3918 MCNAM 3964-3965 MCNAM 4796region of Queimados Hydroelectric Dam (AHE Queimados) Minas Gerais MCNAM 1259 Bocaiuacuteva MCNAM3155-3157 Conselheiro Mata MNRJ 1020-1021 Lagoa Santa MNRJ 40876-40877 Pirapora MCNAM 2135-2138 MCNAM 2204-2209 MCNAM 2238 MCNAM 2604 MCNAM 2738 MCNAM 2740-2745 MCNAM2748-2751 MCNAM 3147-3148 Santana do Riacho ESPIacuteRITO SANTO MNRJ 40878-40879 Piuacutema RIO DEJANEIRO MNRJ 1015-1019 Campos dos Goytacazes MNRJ 3366 MNRJ 13872-13914 MNRJ 40587 MNRJ40880 MNRJ 40881 Satildeo Joatildeo da Barra MATO GROSSO DO SUL MNRJ 24875 Miranda MNRJ 3070 MNRJ13263 Salobra SAtildeO PAULO MNRJ 40882-40883 Botucatu MNRJ 34666-34671 Campinas MNRJ 37043-37045 Piraju PARAGUAY MNRJ 12710-12730 Asuncioacuten MNRJ 12696-12709 Brejo de Ipuatilde

Physalaemus santafecinus ARGENTINA CORRIENTES UFGRS 1943-1947 MZUSP 83258-83259 Ituza-ingoacute Estacircncia Santa Tecla

170 Taxonomic status of Gomphobates marmoratus

P albifrons group due to its larger size (SVL com-bined for males of the other species 282-326 mmfemales 284-341 mm) robust fingers (slender longerfingers in the other species) and dorsal color patternPhysalaemus marmoratus differs from P albifronsand P biligonigerus by the fringed fingers (fringesabsent or poorly marked in P albifrons andP biligonigerus) from P albifrons andP santafecinus by the presence of long rounded orirregular glandular ridges on dorsum (dorsal skin smoothin P albifrons and granulated in P santafecinus)from P albifrons by the presence of one tarsal tuber-cle (two in P albifrons) and large inguinal glands (in-distinct in P albifrons) from P biligonigerus by in-guinal glands with same color pattern of dorsum (in-guinal glands with color pattern darker than dorsum inP biligonigerus) from P santafecinus by the ab-sence of granules on forearms (presence of a line ofgranules on the outer margins of forearms inP santafecinus) and absence of a white longitudinalline on sacral region (evident vertebral line inP santafecinus)

Description of lectotype ndash Body robust head slightlywider than long snout rounded in dorsal (Fig 1) andlateral views (Fig 2) nostrils elliptical not protuber-ant located near the tip of snout oriented dorsolateral-ly canthus rostralis rounded loreal region slightly con-cave eyes protuberant eye diameter larger than in-terorbital distance tympanum indistinct supratympanicfold short slightly marked dorsolateral fold absentvocal sac subgular well developed extending to theborder of chest with belly choanae small oval tonguesmall narrow rounded on posterior border which isnot indented maxillary and premaxillary teeth presentvomerine teeth absent Arms short robust upper armsshorter than forearms outer margin of forearms with-out ridges or granules fingers robust fringed fingertips not expanded horned finger lengths IltIVltIIltIII(Fig 3) nuptial asperities on thumbs and on the medialborder of the inner carpal tubercle but not continuouscarpal tubercles large elliptical with the same sizesupernumerary tubercles protruding conical subartic-ular tubercles single large protruding conical Legsmoderately robust tibia slightly longer than thigh tar-sal fold absent toes slender long slightly fringed (fringeabsent on toe I) toes webbed on base tips not ex-panded horned toe lengths IltIIltVltIIIltIV (Fig 4)tarsal tubercle present metatarsal tubercles large shov-

el-shaped protruding with distal margins horned dis-tance between inner and outer metatarsal tuberclesshorter than that between the former and tarsal tuber-cles supernumerary tubercles absent subarticular tu-bercles single large protruding conical outer marginof subarticular tubercle of toe I weakly horned dorsalpattern presenting an ldquoomegardquo-shaped mark and stripes

Figure 2 Lateral view of the lectotype of Gomphobates marmora-tus Reinhardt and Luumltken 1862 ldquo1861rdquo

Figure 1 Dorsal view of the lectotype of Gomphobates marmora-tus Reinhardt and Luumltken 1862 ldquo1861rdquo

Figure 3 Ventral view of the hand of the lectotype of Gompho-bates marmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Nascimento LB et al 171

extending to flanks and inguinal region dorsum andflanks with long rounded or irregular glandular ridgesupper eyelids finely rugose ventral surfaces smoothexcept on cloacal region and thighs which are cov-ered by large granules ventral disc distinct inguinalgland large ovoid General color pattern in preserva-tive cream light brown blotches and sinuous stripesirregular generally wide bordered by brown lines lightbrown stripes continuous or interrupted tending to forman ldquoomegardquo-shaped mark under scapular region in-terorbital bar dark brown white longitudinal line onsacral region vertical bars dark brown and cream al-ternated from the snout to posterior corner of mouthtransversal bars dark brown and cream alternated ondorsum of arms and legs gular region grayish chestbelly ventral surfaces of arms and legs cream palmarand plantar surfaces grayish over cream backgroundnuptial asperities light brown horned margins of fin-gers toes and metatarsal tubercles dark brown in-guinal gland with the same color pattern of dorsum

Measurements ndash SVL 406 HW 124 HL 112 THL162 TL 173 FL 190 ED 51 ID 36 END 24UEW 39 IND 26

Variation ndash Paralectotype R 1194 shows finger tipsless horned inner border of toe I with a small fringeand subarticular tubercles on toes II III IV and Vhorned Granular ridges granules and inguinal glandsare less evident probably due to preservation Thegeneral color pattern follows that of the lectotype butcolors are more faded Measurements of paralecto-type SVL 385 HW 98 HL 106 THL 150 TL 157FL 170 ED 44 ID 43 END 29 UEW 37 IND

26 Some of the additional specimens examinedshowed a narrower head and more rugose dorsumPalmar and plantar tubercles may be less or morehorned This variation is also observed among tips offingers and toes on a single specimen General colorpattern on dorsum and nuptial asperities of recentlypreserved specimens vary from gray to dark brownthe white longitudinal line on sacral region may be fad-ed Gular region of females follows the color patternof chest and belly Morphometric variation is present-ed in Table 1

Tadpole ndash External morphology and internal oral mor-phology were described by Nomura et al (2003) asPhysalaemus fuscomaculatus

Distribution and ecology ndash Physalaemus marmora-tus breeds in temporary shallow ponds of open habi-tats in the States of Bahia Espiacuterito Santo Rio de Jan-eiro Satildeo Paulo Minas Gerais Goiaacutes Mato Grossoand Mato Grosso do Sul (Brazil) and also in Para-guay Males call floating on the water with the largevocal sac inflated During the axillary amplexus a foamnest is constructed on the water surface where theunpigmented eggs are deposited

Remarks ndash Reinhardt and Luumltken (1862 ldquo1861rdquo) des-ignated three syntypes in the original description ofG marmoratus one was collected by Reinhardt theother two were sent by Dr PW Lund one to theUniversity Museum of Copenhagen and the other tothe ldquoRoyal Museum of Natural Historyrdquo DenmarkLater on all natural history collections in Denmark wereunited into one large collection now ldquoZoological Mu-

Figure 4 Ventral view of the foot of the lectotype of Gomphobatesmarmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Table 1 Mean (x) standard deviation (SD) and range of measure-ments (in mm) of males and females of Physalaemus marmoratus(Reinhardt and Luumltken 1862 ldquo1861rdquo)

Males (N=19) Females (N=13)x SD Range x SD Range

SVL 382 21 333-421 418 42 330-475HL 102 06 92-112 106 07 90-117HW 113 08 10-128 125 10 109-138ED 45 03 40-51 46 05 37-53END 26 02 23-28 27 02 24-31IND 26 01 23-28 27 02 24-31IOD 33 04 27-43 34 04 28-41UEW 39 03 33-46 42 04 37-50THL 157 11 138-176 162 15 139-190TL 155 10 138-173 159 12 136-178FL 174 11 159-190 177 12 156-193

172 Taxonomic status of Gomphobates marmoratus

seum of the University of Copenhagenrdquo (ZMUC MAndersen pers comm) When we consulted theZMUC Curator about the loan of the syntypes he lo-cated two specimens R 1193 (here designated as thelectotype) and R 1194 (here designated as one of theparalectotypes) When asked about the third syntypethe Assistant-Curator could not trace it with certaintybecause none of the three other specimens ofG marmoratus deposited in the ZMUC collection car-ried a label indicating which was the third type How-ever he provided all the information available in thejars labels One specimen (R 1170 formerly ZMUC24) was collected by E Warming which did not col-lect any of the types Specimen R 1195 (formerlyZMUC 25) is just labeled ldquoMinas Geraisrdquo with nocollector and date The last specimen (R 11125 for-merly ZMUC 26 a cleaned skeleton loose bones andskin in alcohol) was collected by PW Lund but thelabel does not indicate the locality and date only aLundrsquos handwriting note saying ldquoNew genus of frogs(1)rdquo We considered this latter as the third syntypebecause of the evidences shown in the label

Pleurodema fuscomaculata (Steindachner 1864)n comb

Eupemphix fuscomaculatus Steindachner 1864Hiobates fuscomaculatus Steindachner 1864Gomphobates fuscomaculatus ndash Steindachner 1867Iliobates fuscomaculatus ndash Steindachner 1867Lystris fuscomaculatus ndash Cope 1869 ldquo1868rdquoPaludicola fuscomaculata ndash Boulenger 1882Physalaemus fuscomaculatus ndash Parker 1927 (part)

Holotype ndash NHMW 4316 female collected by JNatterer in Caiccedilara Municipality of Caacuteceres State ofMato Grosso Brazil (Fig 5 and Fig 6)

A very good description of the holotype of Eupem-phix fuscomaculatus as well as comparisons withother coespecific specimens and notes on distributionis provided by Cei (1990)

Remarks ndash The presence or absence of vomerine teethseems to be variable among genera of the currentlyLeiuperidae (sensu Frost et al 2006) Consequentlyit is not yet possible to determine with certainty forwhat taxon or group of taxa the presence or absenceof vomerine teeth represent a synapomorphic stateThis would require a complete sampling of all species

and all genera involved which is beyond the scope ofthis paper Until this can be accomplished we tenta-tively use the new combination Pleurodema fusco-maculata

RESUMO

A anaacutelise de exemplares depositados em coleccedilotildeesherpetoloacutegicas e identificados como Physalaemus fus-comaculatus demonstrou que estes diferem do tipodeste taacutexon Eupemphix fuscomaculatus Consequumlen-temente o status taxonocircmico de Gomphobates mar-moratus e Eupemphix fuscomaculatus foi revisadocom base nos tipos e na literatura Gomphobatesmarmoratus eacute associado ao gecircnero Physalaemus

Figure 6 Ventral view of the type of Eupemphix fuscomaculatusSteindachner 1864

Figure 5 Dorsal view of the type of Eupemphix fuscomaculatusSteindachner 1864

Nascimento LB et al 173

como Physalaemus marmoratus e se refere aosexemplares previamente identificados como Physala-emus fuscomaculatus Adicionalmente Eupemphixfuscomaculatus eacute associado ao gecircnero Pleurodemacom a combinaccedilatildeo de Pleurodema fuscomaculata

ACKNOWLEDGMENTS

We thank Giovanni Vinciprova (UFRGS) Jens Boslashdtker Ras-mussen (in memorian) and Mogens Andersen (Curator and Assis-tant-Curator of the Zoological Museum University of Copen-hagen Denmark respectively) for providing specimens under theircare Jose P Langone (Museo de Historia Natural de MontevideoUruguay) for providing photos of the type of Eupemphix fusco-maculatus Miguel A Andrade for providing photos of the lecto-type of Gomphobates marmoratus Ceacutelio FB Haddad for criti-cally reading the manuscript Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for the fellowships grant-ed to BVSP CAGC and UC and financial support

LITERATURE CITED

BARRIO A 1965 El genero Physalaemus (Anura Leptodactylidae)en la Argentina Physis 25(70)421-448

BOULENGER GA 1882 Catalogue of Batrachia Salientia siveEcaudata and Batrachia Apoda in the collection of the BritishMuseum 2nd Ed London British Museum xxx + 503p

BOULENGER GA 1886 A synopsis of the reptiles and batrachiansof the Province Rio Grande do Sul Brazil Annals and Magazineof Natural History 18(5)423-445

BOULENGER GA 1887 Description of a new or little-known SouthAmerican frogs of the genera Paludicola and Hyla Annals andMagazine of Natural History 2(5)295-300

CEI JM 1980 Amphibians of Argentina Monitore ZoologicoItaliano (NS) Firenze Monografia 2609p

CEI JM 1987 Additional notes to ldquoAmphibians of Argentinardquo anupdate 1980-1986 Monitore Zoologico Italiano 21209-272

CEI JM 1990 On a paraguayan sample of a long time confusedspecies Physalaemus fuscomaculatus (Steindachner 1864)(Anura Leptodactylidae) Bolletin di Museo Regionale diScienze Naturali di Torino 8(1)215-231

COCHRAN DM 1955 Frogs of Southeastern Brazil United StatesNatural Museum Bulletin 206 xvi + 423p

COPE ED 1869 ldquo1868rdquo Sixth contribution to the herpetology ofTropical America Proceedings of the Academy Natural Sciencesof Philadelphia 20305-313

FROST DR (ED) 1985 Amphibian species of the world Ataxonomic and geographical reference Lawrence Allen PressInc and The Association of Systematics Collections v + 732p

FROST DR 2006 Amphibian Species of the World an OnlineReference Version 40 (17 August 2006) Electronic Databaseaccessible at httpresearchamnhorgherpetologyamphibia

indexhtml American Museum of Natural History New YorkUSA [captured on 20 August 2006]

LYNCH JD 1970 Systematic status of the American leptodactylidfrog genera Engystomops Eupemphix and PhysalaemusCopeia 1970(3)488-496

MEacuteHELY L VON 1904 Investigations on paraguayan batrachiansAnnales Historico-Naturales Musei Nationalis Hungarici2207-232

MILSTEAD WW 1960 Frogs of the genus Physalaemus in SouthernBrazil with description of a new species Copeia1960(2)83-89

MILSTEAD WW 1963 Notes on brazilian frogs of the generaPseudopaludicola and Physalaemus Copeia 1963 (3)565-566

MIRANDA-RIBEIRO A 1926 Notas para servirem ao estudo dosgymnobatrachios (Anura) brasileiros Arquivos do MuseuNacional Rio de Janeiro 271-227

NASCIMENTO LB U CARAMASCHI AND CAG CRUZ 2005Taxonomic review of the species groups of the genusPhysalaemus Fitzinger 1826 with the revalidation of the generaEngystomops Jimeacutenez-de-la-Espada 1872 and EupemphixSteindachner 1863 (Amphibia Anura Leptodactylidae)Arquivos do Museu Nacional Rio de Janeiro 63(2)297-320

NOMURA F DC ROSSA-FERES AND VHM PRADO 2003 Thetadpole of Physalaemus fuscomaculatus (AnuraLeptodactylidae) with a description of internal oralmorphology Zootaxa 3701-8

PARKER HW 1927 A revision of the frogs of the generaPseudopaludicola Physalaemus and Pleurodema Annals andMagazine of Natural History ser 9(20)450-478

PETERS W 1872 Uumlber eine Sammlung von Batrachiern aus Neu-Freiburg in Brasilien Monatsberichte der Akademie derWissenschaften zu Berlin 1872680-684

REINHARDT J AND CF LUumlTKEN 1862 ldquo1861rdquo Bidgrad til Kundskabom Brasiliens Padder og Krybdyr Videnskabelige Meddelelserfra den Naturhistoriske Forening i Kjoslashbenhavn3(10-15)143-242

SPIX JB 1824 Animalia nova sive species novae Testudinum etRanarum quas it itinere per Brazilian annisMDCCCXVIII-MDCCXX jussu et auspiciis MaximillianiJosephi I Bavarrie Regis Monachii xxii + 53p

STEINDACHNER F 1863 Uumlber einige neue Batrachier aus denSammlungen des Wiener Museums Sitzungsberichte derAkademie der Wissenschaften 48186-192

STEINDACHNER F 1864 Batrachologische MitteilungenVerhandelugen Zoologisch-Botanischen Gesellschaft in Wien14239-288

STEINDACHNER F 1867 Amphibien In Reise der OesterreichischenFregatte Novara um die Erde in den Jahren 1857 1858 1859unter den Befehlen des Commodore B von Wuumlllerstorf-UrbairZoologischer Theil 70p

Received 28 April 2006Accepted 16 November 2006

174 Taxonomic status of Gomphobates marmoratus

APPENDIX I

Specimens Examined

Physalaemus albifrons BRAZIL MARANHAtildeO MNRJ 24216-24226 Barreirinha CEARAacute MNRJ 14940-14942 MNRJ 24059 MNRJ 24062-24072 Brejo Santo MNRJ 6636-6674 Mucuripe Fortaleza MNRJ 1125MNRJ 6636-6676 Fortaleza SERGIPE MNRJ 17976-17981 Santa Luzia do Itanhy BAHIA MNRJ 1094-1096Barreiras MNRJ 114-116 MNRJ 1094-1096 MNRJ 1113 MNRJ 1102-1104 MNRJ 1089-1091 Bom Jesus daLapa MNRJ 1105 Juazeiro MZUSP 76521 MZUSP 82303 MINAS GERAIS MNRJ 27180-27189 LageadoMNRJ 27179 MNRJ 21743-21745 Manga MCNAM 206-209 MCNAM 213-216 Porteirinha

Physalaemus biligonigerus BOLIacuteVIA SANTA CRUZ AMNH 144362-144370 AMNH 144371-144385 AMNH144432-144433 BRAZIL SANTA CATARINA MNRJ 31128-31153 RIO GRANDE DO SUL UFGRS 1924-1929 General Cacircmara UFGRS 1776 Uruguaiana UFGRS 1619 Tramandaiacute UFGRS 1319-20 UFRGS 1342Parque Estadual de Itapoatilde Viamatildeo PARAGUAY AMNH 23807 Paraguayan Chaco AMNH 50669-50671Colocircnia Nueva Itaacutelia Villeta URUGUAY DEPARTAMENTO RIO NEGRO MNRJ 28554

Physalaemus marmoratus BRAZIL Bahia MNRJ 28443 MNRJ 28489 Caravelas DISTRITO FEDERALMNRJ 3415 Ribeiratildeo do Torto GoiaacutesMinas Gerais MCNAM 3915-3918 MCNAM 3964-3965 MCNAM 4796region of Queimados Hydroelectric Dam (AHE Queimados) Minas Gerais MCNAM 1259 Bocaiuacuteva MCNAM3155-3157 Conselheiro Mata MNRJ 1020-1021 Lagoa Santa MNRJ 40876-40877 Pirapora MCNAM 2135-2138 MCNAM 2204-2209 MCNAM 2238 MCNAM 2604 MCNAM 2738 MCNAM 2740-2745 MCNAM2748-2751 MCNAM 3147-3148 Santana do Riacho ESPIacuteRITO SANTO MNRJ 40878-40879 Piuacutema RIO DEJANEIRO MNRJ 1015-1019 Campos dos Goytacazes MNRJ 3366 MNRJ 13872-13914 MNRJ 40587 MNRJ40880 MNRJ 40881 Satildeo Joatildeo da Barra MATO GROSSO DO SUL MNRJ 24875 Miranda MNRJ 3070 MNRJ13263 Salobra SAtildeO PAULO MNRJ 40882-40883 Botucatu MNRJ 34666-34671 Campinas MNRJ 37043-37045 Piraju PARAGUAY MNRJ 12710-12730 Asuncioacuten MNRJ 12696-12709 Brejo de Ipuatilde

Physalaemus santafecinus ARGENTINA CORRIENTES UFGRS 1943-1947 MZUSP 83258-83259 Ituza-ingoacute Estacircncia Santa Tecla

Nascimento LB et al 171

extending to flanks and inguinal region dorsum andflanks with long rounded or irregular glandular ridgesupper eyelids finely rugose ventral surfaces smoothexcept on cloacal region and thighs which are cov-ered by large granules ventral disc distinct inguinalgland large ovoid General color pattern in preserva-tive cream light brown blotches and sinuous stripesirregular generally wide bordered by brown lines lightbrown stripes continuous or interrupted tending to forman ldquoomegardquo-shaped mark under scapular region in-terorbital bar dark brown white longitudinal line onsacral region vertical bars dark brown and cream al-ternated from the snout to posterior corner of mouthtransversal bars dark brown and cream alternated ondorsum of arms and legs gular region grayish chestbelly ventral surfaces of arms and legs cream palmarand plantar surfaces grayish over cream backgroundnuptial asperities light brown horned margins of fin-gers toes and metatarsal tubercles dark brown in-guinal gland with the same color pattern of dorsum

Measurements ndash SVL 406 HW 124 HL 112 THL162 TL 173 FL 190 ED 51 ID 36 END 24UEW 39 IND 26

Variation ndash Paralectotype R 1194 shows finger tipsless horned inner border of toe I with a small fringeand subarticular tubercles on toes II III IV and Vhorned Granular ridges granules and inguinal glandsare less evident probably due to preservation Thegeneral color pattern follows that of the lectotype butcolors are more faded Measurements of paralecto-type SVL 385 HW 98 HL 106 THL 150 TL 157FL 170 ED 44 ID 43 END 29 UEW 37 IND

26 Some of the additional specimens examinedshowed a narrower head and more rugose dorsumPalmar and plantar tubercles may be less or morehorned This variation is also observed among tips offingers and toes on a single specimen General colorpattern on dorsum and nuptial asperities of recentlypreserved specimens vary from gray to dark brownthe white longitudinal line on sacral region may be fad-ed Gular region of females follows the color patternof chest and belly Morphometric variation is present-ed in Table 1

Tadpole ndash External morphology and internal oral mor-phology were described by Nomura et al (2003) asPhysalaemus fuscomaculatus

Distribution and ecology ndash Physalaemus marmora-tus breeds in temporary shallow ponds of open habi-tats in the States of Bahia Espiacuterito Santo Rio de Jan-eiro Satildeo Paulo Minas Gerais Goiaacutes Mato Grossoand Mato Grosso do Sul (Brazil) and also in Para-guay Males call floating on the water with the largevocal sac inflated During the axillary amplexus a foamnest is constructed on the water surface where theunpigmented eggs are deposited

Remarks ndash Reinhardt and Luumltken (1862 ldquo1861rdquo) des-ignated three syntypes in the original description ofG marmoratus one was collected by Reinhardt theother two were sent by Dr PW Lund one to theUniversity Museum of Copenhagen and the other tothe ldquoRoyal Museum of Natural Historyrdquo DenmarkLater on all natural history collections in Denmark wereunited into one large collection now ldquoZoological Mu-

Figure 4 Ventral view of the foot of the lectotype of Gomphobatesmarmoratus Reinhardt and Luumltken 1862 ldquo1861rdquo

Table 1 Mean (x) standard deviation (SD) and range of measure-ments (in mm) of males and females of Physalaemus marmoratus(Reinhardt and Luumltken 1862 ldquo1861rdquo)

Males (N=19) Females (N=13)x SD Range x SD Range

SVL 382 21 333-421 418 42 330-475HL 102 06 92-112 106 07 90-117HW 113 08 10-128 125 10 109-138ED 45 03 40-51 46 05 37-53END 26 02 23-28 27 02 24-31IND 26 01 23-28 27 02 24-31IOD 33 04 27-43 34 04 28-41UEW 39 03 33-46 42 04 37-50THL 157 11 138-176 162 15 139-190TL 155 10 138-173 159 12 136-178FL 174 11 159-190 177 12 156-193

172 Taxonomic status of Gomphobates marmoratus

seum of the University of Copenhagenrdquo (ZMUC MAndersen pers comm) When we consulted theZMUC Curator about the loan of the syntypes he lo-cated two specimens R 1193 (here designated as thelectotype) and R 1194 (here designated as one of theparalectotypes) When asked about the third syntypethe Assistant-Curator could not trace it with certaintybecause none of the three other specimens ofG marmoratus deposited in the ZMUC collection car-ried a label indicating which was the third type How-ever he provided all the information available in thejars labels One specimen (R 1170 formerly ZMUC24) was collected by E Warming which did not col-lect any of the types Specimen R 1195 (formerlyZMUC 25) is just labeled ldquoMinas Geraisrdquo with nocollector and date The last specimen (R 11125 for-merly ZMUC 26 a cleaned skeleton loose bones andskin in alcohol) was collected by PW Lund but thelabel does not indicate the locality and date only aLundrsquos handwriting note saying ldquoNew genus of frogs(1)rdquo We considered this latter as the third syntypebecause of the evidences shown in the label

Pleurodema fuscomaculata (Steindachner 1864)n comb

Eupemphix fuscomaculatus Steindachner 1864Hiobates fuscomaculatus Steindachner 1864Gomphobates fuscomaculatus ndash Steindachner 1867Iliobates fuscomaculatus ndash Steindachner 1867Lystris fuscomaculatus ndash Cope 1869 ldquo1868rdquoPaludicola fuscomaculata ndash Boulenger 1882Physalaemus fuscomaculatus ndash Parker 1927 (part)

Holotype ndash NHMW 4316 female collected by JNatterer in Caiccedilara Municipality of Caacuteceres State ofMato Grosso Brazil (Fig 5 and Fig 6)

A very good description of the holotype of Eupem-phix fuscomaculatus as well as comparisons withother coespecific specimens and notes on distributionis provided by Cei (1990)

Remarks ndash The presence or absence of vomerine teethseems to be variable among genera of the currentlyLeiuperidae (sensu Frost et al 2006) Consequentlyit is not yet possible to determine with certainty forwhat taxon or group of taxa the presence or absenceof vomerine teeth represent a synapomorphic stateThis would require a complete sampling of all species

and all genera involved which is beyond the scope ofthis paper Until this can be accomplished we tenta-tively use the new combination Pleurodema fusco-maculata

RESUMO

A anaacutelise de exemplares depositados em coleccedilotildeesherpetoloacutegicas e identificados como Physalaemus fus-comaculatus demonstrou que estes diferem do tipodeste taacutexon Eupemphix fuscomaculatus Consequumlen-temente o status taxonocircmico de Gomphobates mar-moratus e Eupemphix fuscomaculatus foi revisadocom base nos tipos e na literatura Gomphobatesmarmoratus eacute associado ao gecircnero Physalaemus

Figure 6 Ventral view of the type of Eupemphix fuscomaculatusSteindachner 1864

Figure 5 Dorsal view of the type of Eupemphix fuscomaculatusSteindachner 1864

Nascimento LB et al 173

como Physalaemus marmoratus e se refere aosexemplares previamente identificados como Physala-emus fuscomaculatus Adicionalmente Eupemphixfuscomaculatus eacute associado ao gecircnero Pleurodemacom a combinaccedilatildeo de Pleurodema fuscomaculata

ACKNOWLEDGMENTS

We thank Giovanni Vinciprova (UFRGS) Jens Boslashdtker Ras-mussen (in memorian) and Mogens Andersen (Curator and Assis-tant-Curator of the Zoological Museum University of Copen-hagen Denmark respectively) for providing specimens under theircare Jose P Langone (Museo de Historia Natural de MontevideoUruguay) for providing photos of the type of Eupemphix fusco-maculatus Miguel A Andrade for providing photos of the lecto-type of Gomphobates marmoratus Ceacutelio FB Haddad for criti-cally reading the manuscript Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for the fellowships grant-ed to BVSP CAGC and UC and financial support

LITERATURE CITED

BARRIO A 1965 El genero Physalaemus (Anura Leptodactylidae)en la Argentina Physis 25(70)421-448

BOULENGER GA 1882 Catalogue of Batrachia Salientia siveEcaudata and Batrachia Apoda in the collection of the BritishMuseum 2nd Ed London British Museum xxx + 503p

BOULENGER GA 1886 A synopsis of the reptiles and batrachiansof the Province Rio Grande do Sul Brazil Annals and Magazineof Natural History 18(5)423-445

BOULENGER GA 1887 Description of a new or little-known SouthAmerican frogs of the genera Paludicola and Hyla Annals andMagazine of Natural History 2(5)295-300

CEI JM 1980 Amphibians of Argentina Monitore ZoologicoItaliano (NS) Firenze Monografia 2609p

CEI JM 1987 Additional notes to ldquoAmphibians of Argentinardquo anupdate 1980-1986 Monitore Zoologico Italiano 21209-272

CEI JM 1990 On a paraguayan sample of a long time confusedspecies Physalaemus fuscomaculatus (Steindachner 1864)(Anura Leptodactylidae) Bolletin di Museo Regionale diScienze Naturali di Torino 8(1)215-231

COCHRAN DM 1955 Frogs of Southeastern Brazil United StatesNatural Museum Bulletin 206 xvi + 423p

COPE ED 1869 ldquo1868rdquo Sixth contribution to the herpetology ofTropical America Proceedings of the Academy Natural Sciencesof Philadelphia 20305-313

FROST DR (ED) 1985 Amphibian species of the world Ataxonomic and geographical reference Lawrence Allen PressInc and The Association of Systematics Collections v + 732p

FROST DR 2006 Amphibian Species of the World an OnlineReference Version 40 (17 August 2006) Electronic Databaseaccessible at httpresearchamnhorgherpetologyamphibia

indexhtml American Museum of Natural History New YorkUSA [captured on 20 August 2006]

LYNCH JD 1970 Systematic status of the American leptodactylidfrog genera Engystomops Eupemphix and PhysalaemusCopeia 1970(3)488-496

MEacuteHELY L VON 1904 Investigations on paraguayan batrachiansAnnales Historico-Naturales Musei Nationalis Hungarici2207-232

MILSTEAD WW 1960 Frogs of the genus Physalaemus in SouthernBrazil with description of a new species Copeia1960(2)83-89

MILSTEAD WW 1963 Notes on brazilian frogs of the generaPseudopaludicola and Physalaemus Copeia 1963 (3)565-566

MIRANDA-RIBEIRO A 1926 Notas para servirem ao estudo dosgymnobatrachios (Anura) brasileiros Arquivos do MuseuNacional Rio de Janeiro 271-227

NASCIMENTO LB U CARAMASCHI AND CAG CRUZ 2005Taxonomic review of the species groups of the genusPhysalaemus Fitzinger 1826 with the revalidation of the generaEngystomops Jimeacutenez-de-la-Espada 1872 and EupemphixSteindachner 1863 (Amphibia Anura Leptodactylidae)Arquivos do Museu Nacional Rio de Janeiro 63(2)297-320

NOMURA F DC ROSSA-FERES AND VHM PRADO 2003 Thetadpole of Physalaemus fuscomaculatus (AnuraLeptodactylidae) with a description of internal oralmorphology Zootaxa 3701-8

PARKER HW 1927 A revision of the frogs of the generaPseudopaludicola Physalaemus and Pleurodema Annals andMagazine of Natural History ser 9(20)450-478

PETERS W 1872 Uumlber eine Sammlung von Batrachiern aus Neu-Freiburg in Brasilien Monatsberichte der Akademie derWissenschaften zu Berlin 1872680-684

REINHARDT J AND CF LUumlTKEN 1862 ldquo1861rdquo Bidgrad til Kundskabom Brasiliens Padder og Krybdyr Videnskabelige Meddelelserfra den Naturhistoriske Forening i Kjoslashbenhavn3(10-15)143-242

SPIX JB 1824 Animalia nova sive species novae Testudinum etRanarum quas it itinere per Brazilian annisMDCCCXVIII-MDCCXX jussu et auspiciis MaximillianiJosephi I Bavarrie Regis Monachii xxii + 53p

STEINDACHNER F 1863 Uumlber einige neue Batrachier aus denSammlungen des Wiener Museums Sitzungsberichte derAkademie der Wissenschaften 48186-192

STEINDACHNER F 1864 Batrachologische MitteilungenVerhandelugen Zoologisch-Botanischen Gesellschaft in Wien14239-288

STEINDACHNER F 1867 Amphibien In Reise der OesterreichischenFregatte Novara um die Erde in den Jahren 1857 1858 1859unter den Befehlen des Commodore B von Wuumlllerstorf-UrbairZoologischer Theil 70p

Received 28 April 2006Accepted 16 November 2006

174 Taxonomic status of Gomphobates marmoratus

APPENDIX I

Specimens Examined

Physalaemus albifrons BRAZIL MARANHAtildeO MNRJ 24216-24226 Barreirinha CEARAacute MNRJ 14940-14942 MNRJ 24059 MNRJ 24062-24072 Brejo Santo MNRJ 6636-6674 Mucuripe Fortaleza MNRJ 1125MNRJ 6636-6676 Fortaleza SERGIPE MNRJ 17976-17981 Santa Luzia do Itanhy BAHIA MNRJ 1094-1096Barreiras MNRJ 114-116 MNRJ 1094-1096 MNRJ 1113 MNRJ 1102-1104 MNRJ 1089-1091 Bom Jesus daLapa MNRJ 1105 Juazeiro MZUSP 76521 MZUSP 82303 MINAS GERAIS MNRJ 27180-27189 LageadoMNRJ 27179 MNRJ 21743-21745 Manga MCNAM 206-209 MCNAM 213-216 Porteirinha

Physalaemus biligonigerus BOLIacuteVIA SANTA CRUZ AMNH 144362-144370 AMNH 144371-144385 AMNH144432-144433 BRAZIL SANTA CATARINA MNRJ 31128-31153 RIO GRANDE DO SUL UFGRS 1924-1929 General Cacircmara UFGRS 1776 Uruguaiana UFGRS 1619 Tramandaiacute UFGRS 1319-20 UFRGS 1342Parque Estadual de Itapoatilde Viamatildeo PARAGUAY AMNH 23807 Paraguayan Chaco AMNH 50669-50671Colocircnia Nueva Itaacutelia Villeta URUGUAY DEPARTAMENTO RIO NEGRO MNRJ 28554

Physalaemus marmoratus BRAZIL Bahia MNRJ 28443 MNRJ 28489 Caravelas DISTRITO FEDERALMNRJ 3415 Ribeiratildeo do Torto GoiaacutesMinas Gerais MCNAM 3915-3918 MCNAM 3964-3965 MCNAM 4796region of Queimados Hydroelectric Dam (AHE Queimados) Minas Gerais MCNAM 1259 Bocaiuacuteva MCNAM3155-3157 Conselheiro Mata MNRJ 1020-1021 Lagoa Santa MNRJ 40876-40877 Pirapora MCNAM 2135-2138 MCNAM 2204-2209 MCNAM 2238 MCNAM 2604 MCNAM 2738 MCNAM 2740-2745 MCNAM2748-2751 MCNAM 3147-3148 Santana do Riacho ESPIacuteRITO SANTO MNRJ 40878-40879 Piuacutema RIO DEJANEIRO MNRJ 1015-1019 Campos dos Goytacazes MNRJ 3366 MNRJ 13872-13914 MNRJ 40587 MNRJ40880 MNRJ 40881 Satildeo Joatildeo da Barra MATO GROSSO DO SUL MNRJ 24875 Miranda MNRJ 3070 MNRJ13263 Salobra SAtildeO PAULO MNRJ 40882-40883 Botucatu MNRJ 34666-34671 Campinas MNRJ 37043-37045 Piraju PARAGUAY MNRJ 12710-12730 Asuncioacuten MNRJ 12696-12709 Brejo de Ipuatilde

Physalaemus santafecinus ARGENTINA CORRIENTES UFGRS 1943-1947 MZUSP 83258-83259 Ituza-ingoacute Estacircncia Santa Tecla

172 Taxonomic status of Gomphobates marmoratus

seum of the University of Copenhagenrdquo (ZMUC MAndersen pers comm) When we consulted theZMUC Curator about the loan of the syntypes he lo-cated two specimens R 1193 (here designated as thelectotype) and R 1194 (here designated as one of theparalectotypes) When asked about the third syntypethe Assistant-Curator could not trace it with certaintybecause none of the three other specimens ofG marmoratus deposited in the ZMUC collection car-ried a label indicating which was the third type How-ever he provided all the information available in thejars labels One specimen (R 1170 formerly ZMUC24) was collected by E Warming which did not col-lect any of the types Specimen R 1195 (formerlyZMUC 25) is just labeled ldquoMinas Geraisrdquo with nocollector and date The last specimen (R 11125 for-merly ZMUC 26 a cleaned skeleton loose bones andskin in alcohol) was collected by PW Lund but thelabel does not indicate the locality and date only aLundrsquos handwriting note saying ldquoNew genus of frogs(1)rdquo We considered this latter as the third syntypebecause of the evidences shown in the label

Pleurodema fuscomaculata (Steindachner 1864)n comb

Eupemphix fuscomaculatus Steindachner 1864Hiobates fuscomaculatus Steindachner 1864Gomphobates fuscomaculatus ndash Steindachner 1867Iliobates fuscomaculatus ndash Steindachner 1867Lystris fuscomaculatus ndash Cope 1869 ldquo1868rdquoPaludicola fuscomaculata ndash Boulenger 1882Physalaemus fuscomaculatus ndash Parker 1927 (part)

Holotype ndash NHMW 4316 female collected by JNatterer in Caiccedilara Municipality of Caacuteceres State ofMato Grosso Brazil (Fig 5 and Fig 6)

A very good description of the holotype of Eupem-phix fuscomaculatus as well as comparisons withother coespecific specimens and notes on distributionis provided by Cei (1990)

Remarks ndash The presence or absence of vomerine teethseems to be variable among genera of the currentlyLeiuperidae (sensu Frost et al 2006) Consequentlyit is not yet possible to determine with certainty forwhat taxon or group of taxa the presence or absenceof vomerine teeth represent a synapomorphic stateThis would require a complete sampling of all species

and all genera involved which is beyond the scope ofthis paper Until this can be accomplished we tenta-tively use the new combination Pleurodema fusco-maculata

RESUMO

A anaacutelise de exemplares depositados em coleccedilotildeesherpetoloacutegicas e identificados como Physalaemus fus-comaculatus demonstrou que estes diferem do tipodeste taacutexon Eupemphix fuscomaculatus Consequumlen-temente o status taxonocircmico de Gomphobates mar-moratus e Eupemphix fuscomaculatus foi revisadocom base nos tipos e na literatura Gomphobatesmarmoratus eacute associado ao gecircnero Physalaemus

Figure 6 Ventral view of the type of Eupemphix fuscomaculatusSteindachner 1864

Figure 5 Dorsal view of the type of Eupemphix fuscomaculatusSteindachner 1864

Nascimento LB et al 173

como Physalaemus marmoratus e se refere aosexemplares previamente identificados como Physala-emus fuscomaculatus Adicionalmente Eupemphixfuscomaculatus eacute associado ao gecircnero Pleurodemacom a combinaccedilatildeo de Pleurodema fuscomaculata

ACKNOWLEDGMENTS

We thank Giovanni Vinciprova (UFRGS) Jens Boslashdtker Ras-mussen (in memorian) and Mogens Andersen (Curator and Assis-tant-Curator of the Zoological Museum University of Copen-hagen Denmark respectively) for providing specimens under theircare Jose P Langone (Museo de Historia Natural de MontevideoUruguay) for providing photos of the type of Eupemphix fusco-maculatus Miguel A Andrade for providing photos of the lecto-type of Gomphobates marmoratus Ceacutelio FB Haddad for criti-cally reading the manuscript Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for the fellowships grant-ed to BVSP CAGC and UC and financial support

LITERATURE CITED

BARRIO A 1965 El genero Physalaemus (Anura Leptodactylidae)en la Argentina Physis 25(70)421-448

BOULENGER GA 1882 Catalogue of Batrachia Salientia siveEcaudata and Batrachia Apoda in the collection of the BritishMuseum 2nd Ed London British Museum xxx + 503p

BOULENGER GA 1886 A synopsis of the reptiles and batrachiansof the Province Rio Grande do Sul Brazil Annals and Magazineof Natural History 18(5)423-445

BOULENGER GA 1887 Description of a new or little-known SouthAmerican frogs of the genera Paludicola and Hyla Annals andMagazine of Natural History 2(5)295-300

CEI JM 1980 Amphibians of Argentina Monitore ZoologicoItaliano (NS) Firenze Monografia 2609p

CEI JM 1987 Additional notes to ldquoAmphibians of Argentinardquo anupdate 1980-1986 Monitore Zoologico Italiano 21209-272

CEI JM 1990 On a paraguayan sample of a long time confusedspecies Physalaemus fuscomaculatus (Steindachner 1864)(Anura Leptodactylidae) Bolletin di Museo Regionale diScienze Naturali di Torino 8(1)215-231

COCHRAN DM 1955 Frogs of Southeastern Brazil United StatesNatural Museum Bulletin 206 xvi + 423p

COPE ED 1869 ldquo1868rdquo Sixth contribution to the herpetology ofTropical America Proceedings of the Academy Natural Sciencesof Philadelphia 20305-313

FROST DR (ED) 1985 Amphibian species of the world Ataxonomic and geographical reference Lawrence Allen PressInc and The Association of Systematics Collections v + 732p

FROST DR 2006 Amphibian Species of the World an OnlineReference Version 40 (17 August 2006) Electronic Databaseaccessible at httpresearchamnhorgherpetologyamphibia

indexhtml American Museum of Natural History New YorkUSA [captured on 20 August 2006]

LYNCH JD 1970 Systematic status of the American leptodactylidfrog genera Engystomops Eupemphix and PhysalaemusCopeia 1970(3)488-496

MEacuteHELY L VON 1904 Investigations on paraguayan batrachiansAnnales Historico-Naturales Musei Nationalis Hungarici2207-232

MILSTEAD WW 1960 Frogs of the genus Physalaemus in SouthernBrazil with description of a new species Copeia1960(2)83-89

MILSTEAD WW 1963 Notes on brazilian frogs of the generaPseudopaludicola and Physalaemus Copeia 1963 (3)565-566

MIRANDA-RIBEIRO A 1926 Notas para servirem ao estudo dosgymnobatrachios (Anura) brasileiros Arquivos do MuseuNacional Rio de Janeiro 271-227

NASCIMENTO LB U CARAMASCHI AND CAG CRUZ 2005Taxonomic review of the species groups of the genusPhysalaemus Fitzinger 1826 with the revalidation of the generaEngystomops Jimeacutenez-de-la-Espada 1872 and EupemphixSteindachner 1863 (Amphibia Anura Leptodactylidae)Arquivos do Museu Nacional Rio de Janeiro 63(2)297-320

NOMURA F DC ROSSA-FERES AND VHM PRADO 2003 Thetadpole of Physalaemus fuscomaculatus (AnuraLeptodactylidae) with a description of internal oralmorphology Zootaxa 3701-8

PARKER HW 1927 A revision of the frogs of the generaPseudopaludicola Physalaemus and Pleurodema Annals andMagazine of Natural History ser 9(20)450-478

PETERS W 1872 Uumlber eine Sammlung von Batrachiern aus Neu-Freiburg in Brasilien Monatsberichte der Akademie derWissenschaften zu Berlin 1872680-684

REINHARDT J AND CF LUumlTKEN 1862 ldquo1861rdquo Bidgrad til Kundskabom Brasiliens Padder og Krybdyr Videnskabelige Meddelelserfra den Naturhistoriske Forening i Kjoslashbenhavn3(10-15)143-242

SPIX JB 1824 Animalia nova sive species novae Testudinum etRanarum quas it itinere per Brazilian annisMDCCCXVIII-MDCCXX jussu et auspiciis MaximillianiJosephi I Bavarrie Regis Monachii xxii + 53p

STEINDACHNER F 1863 Uumlber einige neue Batrachier aus denSammlungen des Wiener Museums Sitzungsberichte derAkademie der Wissenschaften 48186-192

STEINDACHNER F 1864 Batrachologische MitteilungenVerhandelugen Zoologisch-Botanischen Gesellschaft in Wien14239-288

STEINDACHNER F 1867 Amphibien In Reise der OesterreichischenFregatte Novara um die Erde in den Jahren 1857 1858 1859unter den Befehlen des Commodore B von Wuumlllerstorf-UrbairZoologischer Theil 70p

Received 28 April 2006Accepted 16 November 2006

174 Taxonomic status of Gomphobates marmoratus

APPENDIX I

Specimens Examined

Physalaemus albifrons BRAZIL MARANHAtildeO MNRJ 24216-24226 Barreirinha CEARAacute MNRJ 14940-14942 MNRJ 24059 MNRJ 24062-24072 Brejo Santo MNRJ 6636-6674 Mucuripe Fortaleza MNRJ 1125MNRJ 6636-6676 Fortaleza SERGIPE MNRJ 17976-17981 Santa Luzia do Itanhy BAHIA MNRJ 1094-1096Barreiras MNRJ 114-116 MNRJ 1094-1096 MNRJ 1113 MNRJ 1102-1104 MNRJ 1089-1091 Bom Jesus daLapa MNRJ 1105 Juazeiro MZUSP 76521 MZUSP 82303 MINAS GERAIS MNRJ 27180-27189 LageadoMNRJ 27179 MNRJ 21743-21745 Manga MCNAM 206-209 MCNAM 213-216 Porteirinha

Physalaemus biligonigerus BOLIacuteVIA SANTA CRUZ AMNH 144362-144370 AMNH 144371-144385 AMNH144432-144433 BRAZIL SANTA CATARINA MNRJ 31128-31153 RIO GRANDE DO SUL UFGRS 1924-1929 General Cacircmara UFGRS 1776 Uruguaiana UFGRS 1619 Tramandaiacute UFGRS 1319-20 UFRGS 1342Parque Estadual de Itapoatilde Viamatildeo PARAGUAY AMNH 23807 Paraguayan Chaco AMNH 50669-50671Colocircnia Nueva Itaacutelia Villeta URUGUAY DEPARTAMENTO RIO NEGRO MNRJ 28554

Physalaemus marmoratus BRAZIL Bahia MNRJ 28443 MNRJ 28489 Caravelas DISTRITO FEDERALMNRJ 3415 Ribeiratildeo do Torto GoiaacutesMinas Gerais MCNAM 3915-3918 MCNAM 3964-3965 MCNAM 4796region of Queimados Hydroelectric Dam (AHE Queimados) Minas Gerais MCNAM 1259 Bocaiuacuteva MCNAM3155-3157 Conselheiro Mata MNRJ 1020-1021 Lagoa Santa MNRJ 40876-40877 Pirapora MCNAM 2135-2138 MCNAM 2204-2209 MCNAM 2238 MCNAM 2604 MCNAM 2738 MCNAM 2740-2745 MCNAM2748-2751 MCNAM 3147-3148 Santana do Riacho ESPIacuteRITO SANTO MNRJ 40878-40879 Piuacutema RIO DEJANEIRO MNRJ 1015-1019 Campos dos Goytacazes MNRJ 3366 MNRJ 13872-13914 MNRJ 40587 MNRJ40880 MNRJ 40881 Satildeo Joatildeo da Barra MATO GROSSO DO SUL MNRJ 24875 Miranda MNRJ 3070 MNRJ13263 Salobra SAtildeO PAULO MNRJ 40882-40883 Botucatu MNRJ 34666-34671 Campinas MNRJ 37043-37045 Piraju PARAGUAY MNRJ 12710-12730 Asuncioacuten MNRJ 12696-12709 Brejo de Ipuatilde

Physalaemus santafecinus ARGENTINA CORRIENTES UFGRS 1943-1947 MZUSP 83258-83259 Ituza-ingoacute Estacircncia Santa Tecla

Nascimento LB et al 173

como Physalaemus marmoratus e se refere aosexemplares previamente identificados como Physala-emus fuscomaculatus Adicionalmente Eupemphixfuscomaculatus eacute associado ao gecircnero Pleurodemacom a combinaccedilatildeo de Pleurodema fuscomaculata

ACKNOWLEDGMENTS

We thank Giovanni Vinciprova (UFRGS) Jens Boslashdtker Ras-mussen (in memorian) and Mogens Andersen (Curator and Assis-tant-Curator of the Zoological Museum University of Copen-hagen Denmark respectively) for providing specimens under theircare Jose P Langone (Museo de Historia Natural de MontevideoUruguay) for providing photos of the type of Eupemphix fusco-maculatus Miguel A Andrade for providing photos of the lecto-type of Gomphobates marmoratus Ceacutelio FB Haddad for criti-cally reading the manuscript Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for the fellowships grant-ed to BVSP CAGC and UC and financial support

LITERATURE CITED

BARRIO A 1965 El genero Physalaemus (Anura Leptodactylidae)en la Argentina Physis 25(70)421-448

BOULENGER GA 1882 Catalogue of Batrachia Salientia siveEcaudata and Batrachia Apoda in the collection of the BritishMuseum 2nd Ed London British Museum xxx + 503p

BOULENGER GA 1886 A synopsis of the reptiles and batrachiansof the Province Rio Grande do Sul Brazil Annals and Magazineof Natural History 18(5)423-445

BOULENGER GA 1887 Description of a new or little-known SouthAmerican frogs of the genera Paludicola and Hyla Annals andMagazine of Natural History 2(5)295-300

CEI JM 1980 Amphibians of Argentina Monitore ZoologicoItaliano (NS) Firenze Monografia 2609p

CEI JM 1987 Additional notes to ldquoAmphibians of Argentinardquo anupdate 1980-1986 Monitore Zoologico Italiano 21209-272

CEI JM 1990 On a paraguayan sample of a long time confusedspecies Physalaemus fuscomaculatus (Steindachner 1864)(Anura Leptodactylidae) Bolletin di Museo Regionale diScienze Naturali di Torino 8(1)215-231

COCHRAN DM 1955 Frogs of Southeastern Brazil United StatesNatural Museum Bulletin 206 xvi + 423p

COPE ED 1869 ldquo1868rdquo Sixth contribution to the herpetology ofTropical America Proceedings of the Academy Natural Sciencesof Philadelphia 20305-313

FROST DR (ED) 1985 Amphibian species of the world Ataxonomic and geographical reference Lawrence Allen PressInc and The Association of Systematics Collections v + 732p

FROST DR 2006 Amphibian Species of the World an OnlineReference Version 40 (17 August 2006) Electronic Databaseaccessible at httpresearchamnhorgherpetologyamphibia

indexhtml American Museum of Natural History New YorkUSA [captured on 20 August 2006]

LYNCH JD 1970 Systematic status of the American leptodactylidfrog genera Engystomops Eupemphix and PhysalaemusCopeia 1970(3)488-496

MEacuteHELY L VON 1904 Investigations on paraguayan batrachiansAnnales Historico-Naturales Musei Nationalis Hungarici2207-232

MILSTEAD WW 1960 Frogs of the genus Physalaemus in SouthernBrazil with description of a new species Copeia1960(2)83-89

MILSTEAD WW 1963 Notes on brazilian frogs of the generaPseudopaludicola and Physalaemus Copeia 1963 (3)565-566

MIRANDA-RIBEIRO A 1926 Notas para servirem ao estudo dosgymnobatrachios (Anura) brasileiros Arquivos do MuseuNacional Rio de Janeiro 271-227

NASCIMENTO LB U CARAMASCHI AND CAG CRUZ 2005Taxonomic review of the species groups of the genusPhysalaemus Fitzinger 1826 with the revalidation of the generaEngystomops Jimeacutenez-de-la-Espada 1872 and EupemphixSteindachner 1863 (Amphibia Anura Leptodactylidae)Arquivos do Museu Nacional Rio de Janeiro 63(2)297-320

NOMURA F DC ROSSA-FERES AND VHM PRADO 2003 Thetadpole of Physalaemus fuscomaculatus (AnuraLeptodactylidae) with a description of internal oralmorphology Zootaxa 3701-8

PARKER HW 1927 A revision of the frogs of the generaPseudopaludicola Physalaemus and Pleurodema Annals andMagazine of Natural History ser 9(20)450-478

PETERS W 1872 Uumlber eine Sammlung von Batrachiern aus Neu-Freiburg in Brasilien Monatsberichte der Akademie derWissenschaften zu Berlin 1872680-684

REINHARDT J AND CF LUumlTKEN 1862 ldquo1861rdquo Bidgrad til Kundskabom Brasiliens Padder og Krybdyr Videnskabelige Meddelelserfra den Naturhistoriske Forening i Kjoslashbenhavn3(10-15)143-242

SPIX JB 1824 Animalia nova sive species novae Testudinum etRanarum quas it itinere per Brazilian annisMDCCCXVIII-MDCCXX jussu et auspiciis MaximillianiJosephi I Bavarrie Regis Monachii xxii + 53p

STEINDACHNER F 1863 Uumlber einige neue Batrachier aus denSammlungen des Wiener Museums Sitzungsberichte derAkademie der Wissenschaften 48186-192

STEINDACHNER F 1864 Batrachologische MitteilungenVerhandelugen Zoologisch-Botanischen Gesellschaft in Wien14239-288

STEINDACHNER F 1867 Amphibien In Reise der OesterreichischenFregatte Novara um die Erde in den Jahren 1857 1858 1859unter den Befehlen des Commodore B von Wuumlllerstorf-UrbairZoologischer Theil 70p

Received 28 April 2006Accepted 16 November 2006

174 Taxonomic status of Gomphobates marmoratus

APPENDIX I

Specimens Examined

Physalaemus albifrons BRAZIL MARANHAtildeO MNRJ 24216-24226 Barreirinha CEARAacute MNRJ 14940-14942 MNRJ 24059 MNRJ 24062-24072 Brejo Santo MNRJ 6636-6674 Mucuripe Fortaleza MNRJ 1125MNRJ 6636-6676 Fortaleza SERGIPE MNRJ 17976-17981 Santa Luzia do Itanhy BAHIA MNRJ 1094-1096Barreiras MNRJ 114-116 MNRJ 1094-1096 MNRJ 1113 MNRJ 1102-1104 MNRJ 1089-1091 Bom Jesus daLapa MNRJ 1105 Juazeiro MZUSP 76521 MZUSP 82303 MINAS GERAIS MNRJ 27180-27189 LageadoMNRJ 27179 MNRJ 21743-21745 Manga MCNAM 206-209 MCNAM 213-216 Porteirinha

Physalaemus biligonigerus BOLIacuteVIA SANTA CRUZ AMNH 144362-144370 AMNH 144371-144385 AMNH144432-144433 BRAZIL SANTA CATARINA MNRJ 31128-31153 RIO GRANDE DO SUL UFGRS 1924-1929 General Cacircmara UFGRS 1776 Uruguaiana UFGRS 1619 Tramandaiacute UFGRS 1319-20 UFRGS 1342Parque Estadual de Itapoatilde Viamatildeo PARAGUAY AMNH 23807 Paraguayan Chaco AMNH 50669-50671Colocircnia Nueva Itaacutelia Villeta URUGUAY DEPARTAMENTO RIO NEGRO MNRJ 28554

Physalaemus marmoratus BRAZIL Bahia MNRJ 28443 MNRJ 28489 Caravelas DISTRITO FEDERALMNRJ 3415 Ribeiratildeo do Torto GoiaacutesMinas Gerais MCNAM 3915-3918 MCNAM 3964-3965 MCNAM 4796region of Queimados Hydroelectric Dam (AHE Queimados) Minas Gerais MCNAM 1259 Bocaiuacuteva MCNAM3155-3157 Conselheiro Mata MNRJ 1020-1021 Lagoa Santa MNRJ 40876-40877 Pirapora MCNAM 2135-2138 MCNAM 2204-2209 MCNAM 2238 MCNAM 2604 MCNAM 2738 MCNAM 2740-2745 MCNAM2748-2751 MCNAM 3147-3148 Santana do Riacho ESPIacuteRITO SANTO MNRJ 40878-40879 Piuacutema RIO DEJANEIRO MNRJ 1015-1019 Campos dos Goytacazes MNRJ 3366 MNRJ 13872-13914 MNRJ 40587 MNRJ40880 MNRJ 40881 Satildeo Joatildeo da Barra MATO GROSSO DO SUL MNRJ 24875 Miranda MNRJ 3070 MNRJ13263 Salobra SAtildeO PAULO MNRJ 40882-40883 Botucatu MNRJ 34666-34671 Campinas MNRJ 37043-37045 Piraju PARAGUAY MNRJ 12710-12730 Asuncioacuten MNRJ 12696-12709 Brejo de Ipuatilde

Physalaemus santafecinus ARGENTINA CORRIENTES UFGRS 1943-1947 MZUSP 83258-83259 Ituza-ingoacute Estacircncia Santa Tecla

174 Taxonomic status of Gomphobates marmoratus

APPENDIX I

Specimens Examined

Physalaemus albifrons BRAZIL MARANHAtildeO MNRJ 24216-24226 Barreirinha CEARAacute MNRJ 14940-14942 MNRJ 24059 MNRJ 24062-24072 Brejo Santo MNRJ 6636-6674 Mucuripe Fortaleza MNRJ 1125MNRJ 6636-6676 Fortaleza SERGIPE MNRJ 17976-17981 Santa Luzia do Itanhy BAHIA MNRJ 1094-1096Barreiras MNRJ 114-116 MNRJ 1094-1096 MNRJ 1113 MNRJ 1102-1104 MNRJ 1089-1091 Bom Jesus daLapa MNRJ 1105 Juazeiro MZUSP 76521 MZUSP 82303 MINAS GERAIS MNRJ 27180-27189 LageadoMNRJ 27179 MNRJ 21743-21745 Manga MCNAM 206-209 MCNAM 213-216 Porteirinha

Physalaemus biligonigerus BOLIacuteVIA SANTA CRUZ AMNH 144362-144370 AMNH 144371-144385 AMNH144432-144433 BRAZIL SANTA CATARINA MNRJ 31128-31153 RIO GRANDE DO SUL UFGRS 1924-1929 General Cacircmara UFGRS 1776 Uruguaiana UFGRS 1619 Tramandaiacute UFGRS 1319-20 UFRGS 1342Parque Estadual de Itapoatilde Viamatildeo PARAGUAY AMNH 23807 Paraguayan Chaco AMNH 50669-50671Colocircnia Nueva Itaacutelia Villeta URUGUAY DEPARTAMENTO RIO NEGRO MNRJ 28554

Physalaemus marmoratus BRAZIL Bahia MNRJ 28443 MNRJ 28489 Caravelas DISTRITO FEDERALMNRJ 3415 Ribeiratildeo do Torto GoiaacutesMinas Gerais MCNAM 3915-3918 MCNAM 3964-3965 MCNAM 4796region of Queimados Hydroelectric Dam (AHE Queimados) Minas Gerais MCNAM 1259 Bocaiuacuteva MCNAM3155-3157 Conselheiro Mata MNRJ 1020-1021 Lagoa Santa MNRJ 40876-40877 Pirapora MCNAM 2135-2138 MCNAM 2204-2209 MCNAM 2238 MCNAM 2604 MCNAM 2738 MCNAM 2740-2745 MCNAM2748-2751 MCNAM 3147-3148 Santana do Riacho ESPIacuteRITO SANTO MNRJ 40878-40879 Piuacutema RIO DEJANEIRO MNRJ 1015-1019 Campos dos Goytacazes MNRJ 3366 MNRJ 13872-13914 MNRJ 40587 MNRJ40880 MNRJ 40881 Satildeo Joatildeo da Barra MATO GROSSO DO SUL MNRJ 24875 Miranda MNRJ 3070 MNRJ13263 Salobra SAtildeO PAULO MNRJ 40882-40883 Botucatu MNRJ 34666-34671 Campinas MNRJ 37043-37045 Piraju PARAGUAY MNRJ 12710-12730 Asuncioacuten MNRJ 12696-12709 Brejo de Ipuatilde

Physalaemus santafecinus ARGENTINA CORRIENTES UFGRS 1943-1947 MZUSP 83258-83259 Ituza-ingoacute Estacircncia Santa Tecla