elachistocleis bol+mus+nac+zool chi 2010

BOLETIM DO MUSEU NACIONALNOVA SÉRIE

RIO DE JANEIRO - BRASIL

ISSN 0080-312X

ZOOLOGIA No 527 12 DE AGOSTO DE 2010

1 Received on 17 de junho de 2010. Accepted on 23 de junho de 2010.2 Museu Nacional/UFRJ, Departamento de Vertebrados. Quinta da Boa Vista, São Cristóvão, 20940-

040 Rio de Janeiro, RJ, Brasil. E-mail: [email protected] of Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq).

NOTES ON THE TAXONOMIC STATUS OFELACHISTOCLEIS OVALIS (SCHNEIDER, 1799) AND DESCRIPTION

OF FIVE NEW SPECIES OF ELACHISTOCLEIS PARKER, 1927(AMPHIBIA, ANURA, MICROHYLIDAE)1

(With 6 figures)

ULISSES CARAMASCHI2

ABSTRACT: The taxonomic position of Elachistocleis ovalis (Schneider, 1799) is discussed and thetaxon is considered a nomen dubium, referred to a species inquirenda. Five new species of the genusElachistocleis are described. Elachistocleis helianneae sp.nov., from Humaitá (07o35’S, 62o40’W; 90maltitude), State of Amazonas, Brazil, is characterized by small size (SVL 22.6-28.7mm in males, 29.3-36.4mm in females), dorsum grayish brown with minute scattered light gray spots and a distinctivemiddle longitudinal light cream stripe from the tip of snout to vent, and venter immaculate clearcream, with a sharp color limit between the dorsal and ventral regions. Elachistocleis surumu sp.nov.,from Vila Surumu (04o12’N, 60o48’W; 80m altitude), Municipality of Pacaraima, State of Roraima,Brazil, is diagnosed by the small size (SVL 19.6-27.0mm in males, 23.2-26.9mm in females), dorsumdark gray with small irregular clear gray spots scattered without forming defined pattern, mid dorsallongitudinal light stripe absent, venter gray with many irregular cream spots regularly distributed,including the throat area, and undefined color transition between the dorsal and ventral regions.Elachistocleis carvalhoi sp.nov., from Aragominas (07o10’S, 48o32’W; 345m altitude), State of Tocantins,Brazil, is separated by the medium size (SVL 24.0-31.9mm in males, 32.0-37.1mm in females),dorsum uniformly dark gray without marks or pattern, and venter and flanks grayish with largeanastomosed whitish spots, producing a coarse marbled pattern, mainly on the chest. Elachistocleisbumbameuboi sp.nov., from UHE Ponta da Madeira, Municipality of São Luís (02o32’S, 44o18’W; 24maltitude), State of Maranhão, Brazil, is diagnosed by the medium size (SVL 26.9-28.8mm in males,32.8-43.4mm in females), dorsum uniformly dark gray, without spots nor light mid-dorsal stripe,venter gray, with minute anastomosed whitish blotches, producing a salt-and-pepper pattern, extendingto the ventrolateral region. Elachistocleis matogrosso sp.nov., from Cuiabá (15o36’S, 56o06’W; 177maltitude), State of Mato Grosso, Brazil, is characterized by small size (SVL 21.5-24.6mm in males,29.0-33.2mm in females), dorsum uniformly grayish brown, a mid dorsal longitudinal light creamstripe from the post-cephalic dermal fold to vent, and venter immaculate clear cream, with a poorlydefined color limit between the dorsal and ventral regions. The geographic distributions of the newspecies and of Elachistocleis bicolor and E. cesarii are realized and mapped.

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Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010

Key words: Gastrophryninae. Elachistocleis helianneae sp.nov. Elachistocleis surumu sp.nov.Elachistocleis carvalhoi sp.nov. Elachistocleis bumbameuboi sp. nov. Elachistocleis matogrosso sp.nov.

RESUMO: Notas sobre a posição taxonômica de Elachistocleis ovalis (Schneider, 1799) e descriçãode cinco espécies novas de Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae).A posição taxonômica de Elachistocleis ovalis (Schneider, 1799) é discutida e o táxon consideradoum nomen dubium, relacionado a uma species inquirenda. Cinco novas espécies do gêneroElachistocleis são descritas. Elachistocleis helianneae sp.nov., de Humaitá (07o35’S, 62o40’W;90m de altitude), Estado do Amazonas, Brasil, é diagnosticada pelo tamanho pequeno (CRA22,6-28,7mm em machos, 29,3-36,4mm em fêmeas), dorso marrom acinzentado com pequenasmanchas cinza claro espalhadas e uma característica linha longitudinal mediana creme clarodesde a ponta do focinho ao ânus e ventre creme claro imaculado, com nítido limite de coloridoentre as regiões dorsal e ventral. Elachistocleis surumu sp.nov., da Vila Surumu (04o12’N, 60o48’W;80m de altitude), Município de Pacaraima, Estado de Roraima, Brasil, é caracterizada pelo tamanhopequeno (CRA 19,6-27,0mm em machos, 23,1-26,9mm em fêmeas), dorso cinza escuro compequenas manchas cinza claro espalhadas sem formar padrão definido, linha longitudinal dorsalclara ausente, ventre cinza com muitas manchas irregulares creme regularmente distribuídas,incluindo a região gular, e transição de colorido indefinida entre as regiões dorsal e ventral.Elachistocleis carvalhoi sp.nov., de Aragominas (07o10’S, 48o32’W; 345m de altitude), Estado doTocantins, Brasil, é separada pelo tamanho médio (CRA 24,9-31,9mm em machos, 32,0-37,1mmem fêmeas), dorso uniformemente cinza escuro sem desenhos ou padrão, e ventre e flancos cinzacom grandes manchas esbranquiçadas anastomosadas, produzindo um padrão marmoreadogrosseiro, principalmente na região peitoral. Elachistocleis bumbameuboi sp.nov., da UHE Pontada Madeira, Município de São Luís (02o32’S, 44o18’W; 24m de altitude), Estado do Maranhão,Brasil, é diagnosticada pelo tamanho médio (CRA 26,9-28,8mm em machos, 32,8-43,4mm emfêmeas), dorso uniformemente cinza escuro, sem manchas nem linha clara mediana, ventre cinza,com pequenas manchas esbranquiçadas anastomosadas, produzindo um padrão de sal-e-pimentaque se estende até a região ventrolateral. Elachistocleis matogrosso sp.nov., de Cuiabá (15o36’S,56o06’W; 177m de altitude), Estado do Mato Grosso, Brasil, é caracterizada pelo tamanho pequeno(CRA 21,5-24,6mm em machos, 29,0-33,2mm em fêmeas), dorso uniformemente marromacinzentado, com uma linha creme claro longitudinal mediana desde a prega dérmica pós-cefálicaaté o ânus, e ventre creme claro imaculado, com limite de colorido pouco definido entre as regiõesdorsal e ventral. As distribuições geográficas das novas espécies e de Elachistocleis bicolor eElachistocleis cesarii são atualizadas e mapeadas.Palavras-chave: Gastrophryninae. Elachistocleis helianneae sp.nov. Elachistocleis surumu sp.nov.Elachistocleis carvalhoi sp.nov. Elachistocleis bumbameuboi sp.nov. Elachistocleis matogrosso sp.nov.

INTRODUCTION

Currently, the genus Elachistocleis Parker, 1927 is composed by eight species,distributed in two ventral color pattern groups. One has immaculate chest and belly,involving E. bicolor (Guérin-Méneville, 1838) and E. ovalis (Schneider, 1799), and otherwith some kind of blotches or vermiculations on belly, including E. cesarii (Miranda-Ribeiro, 1920), E. erythrogaster Kwet & Di Bernardo, 1998, E. magnus Toledo, 2010, E.piauiensis Caramaschi & Jim, 1983, E. skotogaster Lavilla, Vaira & Ferrari, 2003, andE. surinamensis (Daudin, 1802) (FROST, 2010; TOLEDO, 2010; TOLEDO et al., 2010).LAVILLA et al. (2003) considered that three species in the genus are well defined andhave associated name-bearing types and type localities. Elachistocleis piauiensis,described from Picos, State of Piauí, Brazil, has a small size (SVL 19.8-23.7mm inmales, 21.1-28.3mm in females) and venter mottled in cream and gray, a thin and

NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS... 3


interrupted clear line on the posterior surface of thighs, and a large gland behind theposterior corner of mouth (CARAMASCHI & JIM, 1983); E. erythrogaster, described fromSão Francisco de Paula, State of Rio Grande do Sul, Brazil, presents size large (SVL29.1-32.3mm in males, 34.0-37.6mm in females) and deep black throats in malesand females, black and blue marbled lateral surfaces, flashy red-orange venter, andno femoral posterior stripe nor postcommisural gland (KWET & DI BERNARDO, 1998);and E. skotogaster, described from Los Toldos, Departamento Santa Victoria, Salta,Argentina, has a large size (SVL 27.5-28.5mm in males, 30.3-34.4mm in females),belly and legs densely spotted in brown, dorsal region uniformly dark brown mottledwith black, without a light vertebral stripe, and postcommisural gland absent (LAVILLA

et al., 2003). To this group must be added the recently revalidated E. cesarii, typelocality Piquete, State of São Paulo, Brazil, with medium size (SVL 22.6-26.7mm inmales, 28.6-36.0mm in females) and dorsum and limbs brownish gray with smallwhite dots, throat darker than venter, chest yellow with gray marks, belly white oryellow with gray marks and reticulations reaching the flanks, orange femoral stripe,and small postcommisural gland present (TOLEDO et al., 2010), and the recentlydescribed E. magnus, type locality Fazenda Jaburi, Municipality of Espigão do Oeste,State of Rondônia, Brazil, with large size (SVL 31.8-36.6mm in males, 39.8-43.8mmin females) and dorsum and limbs uniform dark grayish with scarce minute brighterdots on the outer boundaries of the dorsum, a thin mid-dorsal white stripe from thevent to the anterior third of the dorsum, throat brownish dark, darker than chestand belly, venter gray with minute scattered white spots mainly on the belly andventral surface of legs, large irregular white spots on the groin and axillary region, abroad, not well defined femoral light stripe, and large postcommisural gland present(TOLEDO, 2010). On the other hand, the three remaining and older species do not havename-bearing types nor defined type localities and LAVILLA et al. (2003) showed thattwo of them, E. ovalis and E. bicolor, are involved in a enormous confusion since1841 and, in fact, later contributions not only did not solve the problem but contributedto increase the controversy. In its turn, E. surinamensis has been treated as a juniorsynonym of E. ovalis, as a senior synonym of Hypopachus pearsei Ruthven, 1914(currently Relictivomer pearsei), or as valid species. Attempting to throw some lighton the problem, LAVILLA et al. (2003) proposed that it would be advisable to considerthe latter three species to fit with the characters that describe the genus Elachistocleisand that E. surinamensis would have a spotted ventral color, an evident light vertebralstripe, and would inhabit the northern portion of the generic range; E. bicolor wouldhave an immaculate venter and would occupy the southern portion of the genericrange (Argentina, Bolivia, Paraguay, Uruguay, and southern Brazil), restricting thetype locality of the species to Buenos Aires, Argentina; and E. ovalis would have animmaculate, yellow ventral color and would occur in the northern portion of thegeneric range. Nevertheless, in this concept E. ovalis is not associated with any existingnatural population. Additionally, LAVILLA et al. (2003) stated that these three taxaconstitute, without doubt, complexes of species and the decisions presented respectingthe definitions would be only operative frameworks for a necessary revision.

Pending that extensive revision and in order to improve the taxonomy of the genusElachistocleis, in this paper the taxonomic status of Elachistocleis ovalis (Schneider,1799) is discussed and five new species of Elachistocleis are described from Brazil.

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MATERIAL AND METHODS

Specimens examined, referred in the text and in the Appendix, are housed in thefollowing collections: MNRJ (Museu Nacional, Rio de Janeiro, RJ, Brazil); MPEG (MuseuParaense Emílio Goeldi, Belém, PA, Brazil); MZUSP (Museu de Zoologia, Universidadede São Paulo, SP, Brazil); MHNCI (Museu de História Natural do Capão da Imbuia,Curitiba, PR, Brazil); UFBA (Museu de Zoologia, Universidade Federal da Bahia,Salvador, BA, Brazil); MZUFV (Museu de Zoologia João Moojen de Oliveira, UniversidadeFederal de Viçosa, MG, Brazil); MCNAM (Museu de Ciências Naturais, PontifíciaUniversidade Católica de Minas Gerais, Belo Horizonte, MG, Brazil); USNM (NationalMuseum of Natural History, Smithsonian Institution, Washington, D.C., USA); CHUNB(Coleção Herpetológica da Universidade de Brasília, DF); CFBH (Célio F.B. HaddadCollection, Universidade Estadual Paulista, Campus de Rio Claro, SP, Brazil); EI(Eugenio Izecksohn Collection, Universidade Federal Rural do Rio de Janeiro,Seropédica, RJ, Brazil); AL-MN (Adolpho Lutz Collection, deposited in MNRJ); and JJ(Jorge Jim Collection, currently in the MNRJ).Measurements were taken on preserved specimens using digital callipers under adissecting microscope to the nearest 0.1mm. Abbreviations of the measurements are:SVL (snout-vent length); HL (head length); HW (head width); IND (internarial distance);END (eye to nostril distance); ED (eye diameter); UEW (upper eyelid width); IOD(interorbital distance); HAL (hand length); THL (thigh length); TL (tibia length); FL (footlength). Snout profile terminology follows HEYER et al. (1990).

RESULTS

THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS (SCHNEIDER, 1799)

Elachistocleis ovalis was proposed by SCHNEIDER (1799) as Rana ovalis, a species withsmall head, long snout, globose body, and small eyes, with gray dorsum and yellowishventer. The syntypes were originally one specimen in the “Musei Ducalis Brunovicensis”,a second specimen in the “Museo Barbyensi”, and one specimen in the “GronoviusMusei II no. 67”; the latter would have been that described by GRONOVIUS (1763,“Zoophylacii no. 65”) and illustrated by SEBA (1735, “pictura II tab. 37 f. 3”) (SCHNEIDER,1799). All syntypes are currently not known and certainly are lost and, actually, theonly figured specimen in SEBA (1735) is a Breviceps gibbosus (Linnaeus, 1758)(Brevicipitidae), a species known from Republic of South Africa (FROST, 2010). The typelocality was not stated and it is impossible to infer where the syntypes were collectedon the basis of cited collections and figures. Besides, no one of the earlier andcontemporary followers of Schneider (e.g., SHAW, 1802; DAUDIN, 1802; MERREM, 1820)stated the locality for the species; moreover, FITZINGER (1826) considered that itsdistribution would be “...Ex Asia, India...” [“E. ovalis. m. Eiförmige E. (Rana ovalis.Schneider) Ex Asia, India.”]. CUVIER (1829) associated Engystoma ovalis with the genusDactylethra ¯ “l’Engystoma ovalis Fitz., est un dactylètre” ¯, a genus created by him forspecies from Southern Africa (“le midi de l’Afrique ...”) and currently synonymizedunder Xenopus Wagler, 1827 (Pipidae) (FROST, 2010). The affirmation of CUVIER (1829)was disputed by DUMÉRIL & BIBRON (1841), who said that it was a mistake, since Fitzinger



gave, as a synonym of his Engystoma ovale, Rana ovalis Schneider; however, theyconceded that Fitzinger also included in his genus Engystoma the “Pipe lisse” of Daudin,which would be really a “Dactylèthre.” In the account on the “I. L’Engystome ovale.Engystoma ovale. Fitzinger”, DUMÉRIL & BIBRON (1841) included a list of synonyms withall names purportedly associated with that species: Rana ovalis Schneider, Rana ovalisShaw, Bufo surinamensis Daudin, Bufo ovalis Daudin, Rana bufonia Merrem, Engystomaovalis Fitzinger, Oxyrhincus bicolor Valenciennes, Oxyrhincus bicolor Guérin, Micropsunicolor Wagler, and Stenocephalus microps Tschudi. In doing that, they included underthe same concept the two known color morphs, with spotted and immaculte venters.Additionally, overlooking the statement of FITZINGER (1841) on the distribution of thespecies, they considered that Engystoma ovale occurred in South America (“l’Ameriqueméridionale”), having examined putative specimens from Surinam and from BuenosAires [Argentina]. This distributional concept was followed, without discussion, by allsubsequent authors (LAVILLA et al., 2003). Attempting to present an operative frameworkfor a necessary revision of what they considered a complex of species, LAVILLA et al.(2003) stated that E. ovalis will “fit with the characters that describe the genusElachistocleis, have an immaculate, yellow ventral coloration, and inhabit the northernportion of the generic range.” Concluding, they conceded that regarding this distributionand if the statement of FITZINGER (1826) is in error and Elachistocleis is really a neotropicaltaxon, the decision presented was based on the exclusion of those specimens consideredunder the name Elachistocleis bicolor (Guérin-Méneville, 1838).The proposition of LAVILLA et al. (2003), however, presents a central problem given thatthe name E. ovalis was not associated to any known frog population in South Americaand, if FITZINGER (1826) is correct and DUMÉRIL & BIBRON (1841) are wrong, this name isnot applicable to a neotropical taxon. Moreover, although biologically indefinite,taxonomically the name E. ovalis threatens the later congener E. bicolor, a currentlywell established species. In fact, based on the immaculate, yellow ventral color describedfor both taxa, CARCERELLI (1992), in a meeting abstract, proposed the synonymization ofE. bicolor with E. ovalis. This action was followed by KLAPPENBACH & LANGONE (1992),LANGONE (1995), and KWET & DI BERNARDO (1998), but not recognized by other authors,including FROST (2010 and earlier versions).In view of this, considering the brief and poorly informative original description, theinexistence of a name bearing type or types, the absence of a type locality, theimpossibility to associate the name with an actual population, and to prevent thethreat to a well known species, Rana ovalis Schneider, 1799 and its current combinationElachistocleis ovalis is here considered a nomen dubium, that is, a name of unknown ordoubtful application (ICZN, 1999), associated to a species inquirenda, that is, a speciesof doubtful identity needing further investigation (ICZN, 1999).

SPECIES ACCOUNTS

Elachistocleis helianneae sp.nov.(Fig.1)

Holotype – BRAZIL: AMAZONAS: Humaitá (07o35’S, 62o40’W; 90m altitude), MNRJ6989, adult , collected by Ulisses Caramaschi, 13/XII/1979.

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Paratypes – All from the type locality: MNRJ 4818-4820, collected by U.Caramaschi,26/II/1976; MNRJ 4822, MNRJ 6990-6993, collected with the holotype; MNRJ 66870-66871, collected by U.Wrani, on 17/IX/1974; MNRJ 66872-66879, collected by A.Silva,01/VI/1974; MNRJ 66880-66882, collected by A.Silva, 10/IX/1974; MNRJ 66883-66885,collected by A.Silva and C.M.Carvalho, 04/I/1979; MNRJ 66886-66887, collected byC.M.Carvalho, A.Silva and L.M.Silva, 13/I/1979; MNRJ 66888-66889, collected byL.M.Silva, A.Silva and C.M.Carvalho, 24/I/1979; MNRJ 66890-66892, collected byM.Menezes, I/1975; MNRJ 66893, collected by U.Caramaschi and C.M.Carvalho,06-07/III/1975; MNRJ 66894, collected by U.Caramaschi and C.M.Carvalho, 19/III/1975; MNRJ 66895, collected by D.Z.Araujo and V.P.Silva, 26/III/1975; MNRJ 68296-68297, collected by L.M.Silva, A.Silva and C.M.Carvalho, 13/I/1979.Diagnosis – A small sized species (SVL 22.6-28.7mm in males, 29.3-36.4mm in females),characterized by head length slightly smaller than the head width, HL about 94.6% ofHW (x̄ = 94.6; SD = 8.03; n = 33); postcommisural gland poorly developed; dorsumsmooth; in preservative grayish brown with minute scattered light gray spots; adistinctive middle longitudinal light cream stripe, from the tip of snout to vent; venterimmaculate cream; a sharp color limit between the dorsal and ventral regions; nospots on axillae or groin; a broad irregular cream line on the posterior surface of thethighs; a large light cream spot on the proximal internal surface of tibiae; a narrowlight cream stripe surrounding the knees and reaching the middle of the tarsus.

Comparisons with other species – The three species with immaculate venters are E. bicolor,E. helianneae sp.nov., and E. matogrosso sp.nov. The new species is readily distinguishedfrom E. bicolor by the longer head (HL greater than 94% of HW in H. helianneae sp.nov.;HL less than 90% of HW in E. bicolor), by the presence of the mid-dorsal stripe (absent inE. bicolor), by the presence of minute light spots on dorsum and dorsal surfaces of members(absent in E. bicolor), and by the stripe on the posterior surface of thighs broad, irregular(thin, well defined in E. bicolor). From E. matogrosso sp.nov., the new species is separatedby the mid-dorsal stripe extending from the tip of snout to vent (extending from the post-cephalic dermal fold to vent in E. matogrosso sp.nov.), by the presence of minute lightspots on dorsum and dorsal surfaces of members (absent in E. matogrosso sp.nov.), by theloreal region conspicuously white in E. helianneae (dorsal gray color of dorsum of thesnout invading the loreal region almost to the upper lip border in E. matogrosso), and bythe sharp color limit between the dorsal and ventral regions (color limit between dorsaland ventral regions poorly defined in E. matogrosso sp.nov.)

All other species in the genus Elachistocleis present some type of ventral color pattern(venter immaculate in E. helianneae sp.nov.)

Description of holotype – Body ovoid, head small, triangular, slightly broader than long;head length 97% of head width and 24.3% of SVL; head width 25.1% of SVL. Snout sub-elliptical in dorsal view, protruding in profile. Nostrils small, not protuberant, directedanterolaterally, closer to tip of snout than to eye; internarial distance smaller than the eye tonostril and interorbital distances, equalling the eye diameter, and larger than the uppereyelid width. Canthus rostralis rounded; loreal region flat, sloping abruptly to the upper lip;lips not flared. Postcommisural gland poorly developed. Eyes small, dorsolateral, only slightlyprotruding. Interorbital space slightly convex, almost two times the upper eyelid width. Nocranial crests. A poorly defined transverse skinfold across back of the head, bending



downwards slightly behind the eyes to the shoulder; a poorly defined skinfold throughdorsolateral region from the axilla to the groin. Tympanum concealed; supratympanic foldabsent. Lower jaw with truncate, trilobed anterior margin. Tongue large, oval, without anotch on posterior border. Choanae large, subcircular, widely separated. Vocal slits present.Premaxillary, maxillary, and vomerine odontophores absent. Vocal sac subgular, not expandedexternally. A weak skinfold crossing the chest between axillae.Arms moderately robust, no tubercles or crests on forearm; palmar tubercle large,divided longitudinally, twice the size of the thenar tubercle; fingers slender, free, withsubarticular tubercles developed, rounded; supranumerary tubercles absent; tip offingers not flattened or expanded; terminal grooves absent. Relative lengths of fingers,1<2<4<3. Prepollex not evident; nuptial pads or asperities absent.

Fig.1- Elachistocleis helianneae sp.nov., holotype (MNRJ 6989; SVL 26.5mm), dorsal and ventralviews of body, and head profile.

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Legs short, robust. Thigh length smaller than tibia and foot lengths; thigh length 95.8%of tibia length. Sum of thigh and tibia lengths 74.8% of SVL; thigh length 36.6% ofSVL; tibia length 38.2% of SVL. Heel of adpressed legs not reaching axilla; knee andelbow widely separated when limbs laid along the sides; heels touching when flexedlegs held at right angle to body; knee and heel with a transverse skinfold; no tibial ortarsal ridges; an oval inner but no outer metatarsal tubercle; plantar tubercles absent.Toes slender, free, weakly fringed; subarticular tubercles developed, rounded;supranumerary tubercles absent; tips of toes not flattened or expanded; terminal groovesabsent. Relative lengths of toes, 1<2<5<3<4. Skin smooth above and beneath; analopening not modified, no para-anal tubercles.In preservative, dorsum and dorsal surfaces of limbs grayish brown, with minutewhite dots scattered without forming a defined pattern; a distinctive mid-dorsallight cream stripe, from the tip of snout to the urostile; a sharp color limit betweenthe dorsal and ventral regions; venter immaculate, dull white; throat grayish;no spots on axillae and groin; an irregular broad light cream stripe on theposterior surface of thighs; a large irregular, light cream spot on the proximalinternal surface of tibiae; a light cream stripe surrounding the knees and reaching themiddle of the tarsus.Measurements of holotype in mm – SVL 26.5, HL 6.4, HW 6.6, IND 1.7, END 2.3, ED1.7, UEW 1.3, IOD 2.8, HAL 5.5, THL 9.7, TL 10.1, FL 12.1.Etymology – The name of the species is given after Helianne de Niemeyer (MNRJ),for her constant strength for the ups and downs of my professional and personallife, besides the companionship in the home, in the field, and in the laboratorywork.Variation – Except for minor details in corporal proportions, the descripton of the holotypestands for males of the species. Size ranges from 23.0 to 29.5mm SVL and the intensityof the color of the throat can vary. The females are bigger than males, size ranging from26.3 to 37.0mm SVL. Range, mean, and standard deviation of the measurements of thetype specimens of E. helianneae are presented in table 1. In few specimens the mid-dorsal stripe can be interrupted, but it is always present.Geographical distribution – Northern Brazil, in the States of Amazonas, Pará, andRondonia, and in Bolivia, in the Departments Beni and Santa Cruz (Fig.6).Remarks – Good color pictures of E. helianneae sp.nov. in life are presented in DE LA

RIVA et al. (2000) and in LIMA et al. (2006), identified as E. bicolor. Dorsum and dorsalsurfaces of limbs gray to dark gray, with small, scattered, irregular black spots;longitudinal mid-dorsal stripe yellow; venter uniformly yellow to greenish yellow; amid-ventral white line from the chest to vent, crossed by a transversal white linebetween the axillae; throat gray; stripe on posterior surface of thighs and spots oninternal surfaces of tibiae orange red; line surrounding knees and tarsus yellow; irisbrown with intense black vermiculations.The specimen MNRJ 6949 fits the diagnosis for E. helianneae sp.nov., but has amanuscript label containing “Engystoma ovale bicolor (Val.) / Assumpção - R. doParaguay / Coll. B. Schouten 1929. / Miranda-Ribeiro [signed].” The collection dataare here considered in error, due to an early labeling mistake or possible specimenexchange in some time of these almost 90 years in the collection.



Elachistocleis surumu sp.nov.(Fig.2)

Holotype – BRAZIL: RORAIMA: Municipality of Pacaraima, Vila Surumu (04o12’N,60o48’W; 121m altitude), MNRJ 25210, adult , collected by Ulisses Caramaschi,Helianne de Niemeyer, and Décio F. de Moraes Jr., 20/X-10/XI/1998.Paratypes – Collected with the holotype: MNRJ 25211-25300, 25302-25305, 25307-25326, 25338-25344.Diagnosis – A small sized species (SVL 19.6-27.0mm in males, 23.2-26.9mm infemales), diagnosed by the head length shorter than the head width, HL about85.6% of HW (x¯ = 85.6; SD = 4.27; n = 24); postcommisural gland present,small; dorsum smooth; in preservative dark gray with small irregular clear grayspots scattered without forming defined pattern, and mid longitudinal light stripeabsent; venter gray with many irregular cream spots regularly distributed,including the throat area; an undefined color transition between the dorsal andventral regions; small, defined white spots on axilla and groin; a broad irregular,poorly defined line on the posterior surface of the thighs; no light stripe on kneeand tarsus.Comparisons with other species – The species with some kind of ventral color patternare E. cesarii, E. erythrogaster, E. magnus, E. piauiensis, E. skotogaster, E. surinamensis,E. carvalhoi sp.nov., and E. bumbameuboi sp.nov. The new species is distinguishedfrom these species by the dorsal color dark gray with small irregular clear gray spots(dorsal surfaces dark gray with small white dots in E. cesarii; dorsum dark gray marbledwith black, blue, and some orange in males, dorsum orange marbled with black andblue in females of E. erythrogaster; dorsum uniform dark grayish with scarce minute

TABLE 1. Range (mm), mean (x̄ ), and standard deviation (SD) of the measurements of the typespecimens of Elachistocleis helianneae sp.nov. (n = number of specimens).

(n = 15) (n = 18) CHARACTERS RANGE x̄ SD RANGE x̄ SD SVL 23.0-29.5 26.4 1.93 26.3-37.0 31.7 2.82 HL 5.65-6.8 6.1 0.37 6.1-7.9 7.0 0.61 HW 5.7-7.5 6.7 0.37 5.55-8.4 7.3 0.70 IND 1.2-2.15 1.7 0.24 1.7-2.2 2.0 0.18 END 2.1-2.65 2.3 0.16 2.0-3.0 2.5 0.29 ED 1.35-2.0 1.6 0.20 1.6-2.1 1.7 0.16 UEW 1.0-1.3 1.2 0.09 1.2-1.6 1.3 0.12 IOD 2.1-3.5 2.85 0.39 2.6-3.6 3.1 0.27 HAL 4.6-6.5 5.4 0.47 5.4-7.2 6.3 0.52 THL 8.0-10.7 9.8 0.80 9.2-12.7 11.2 1.09 TL 8.4-11.8 10.2 0.82 10.2-12.9 11.8 0.92 FL 10.5-13.9 12.0 0.83 11.8-15.8 13.9 1.21

10 U.CARAMASCHI


brighter dots in the outer boundaries in E. magnus; dorsum uniformly gray with scarceminute irregular bright dots in E. piauiensis; dorsum dark brown mottled with blackand white spots in E. skotogaster; dorsum dark gray or black with an evident lightvertebral stripe in E. surinamensis; dorsum uniformly brown or dark gray withoutmarks in E. carvalhoi sp.nov. and E. bumbameuboi sp.nov.; color in figures and/ordescribed in KENNY, 1969, CARAMASCHI & JIM, 1983, KWET & DI BERNARDO, 1998, LAVILLA etal., 2003, NUNES et al., 2010, TOLEDO, 2010, and TOLEDO et al., 2010); by the venter andthroat gray with many irregular yellow spots regularly distributed in E. surumu sp.nov.(chest yellow with gray marks and belly white or yellow with gray marks and reticulationsin E. cesarii; venter red orange, with hindlimbs largely black mottled in bright blue inE. erythrogaster; venter gray with minute scattered white spots, mainly on the bellyand ventral surfaces of legs, in E. magnus; venter dull white or light yellow, heavilygrayish spotted, marbled, in E. piauiensis; venter gray, mottled with dark brown inE. skotogaster; venter dark gray or black with large yellow and small white spots inE. surinamensis; venter grayish with large anastomosed yellow or whitish yellow blotches,producing a coarse marbled pattern, mainly in the chest area, in E. carvalhoi sp.nov.;and venter gray with minute anastomosed whitish spots, producing a salt-and-pepperpattern, in E. bumbameuboi sp.nov.).The three species with immaculate venters and a defined color limit between the dorsaland ventral regions are E. bicolor, E. helianneae, and E. matogrosso sp.nov. The newspecies is readily distinguished from these species by having venter gray with manyirregular cream spots regularly distributed and an undefined color transition betweenthe dorsal and ventral regions.

Description of holotype – Body ovoid, head small, triangular, broader than long; headlength 89.7% of head width and 25.2% of SVL; head width 28.1% of SVL. Snout roundedin dorsal view, protruding in profile. Nostrils small, not protuberant, directed laterally,closer to tip of snout than to eye; internarial distance smaller than the eye to nostriland interorbital distances, and larger than the eye diameter and upper eyelid width.Canthus rostralis rounded; loreal region flat, sloping abruptly to the upper lip; lips notflared. Postcommisural gland present, small. Eyes small, dorsolateral, only slightlyprotruding. Interorbital space slightly convex, almost two times the upper eyelid width.No cranial crests. A transverse skinfold across back of the head, bending downwardsslightly behind the eyes to the shoulder; a poorly defined skinfold through dorsolateralregion from the axilla to the groin. Tympanum concealed; supratympanic fold absent.Lower jaw with truncate, trilobed anterior margin. Tongue large, oval, without a notchon posterior border. Choanae large, subcircular, widely separated. Vocal slits present.Premaxillary, maxillary, and vomerine odontophores absent. Vocal sac subgular, notexpanded externally. A skinfold crosses the chest between axillae.

Arms moderately robust, no tubercles or crests on forearm; palmar tubercle large,divided longitudinally, twice the size of the thenar tubercle; fingers slender, free, withsubarticular tubercles developed, rounded; supranumerary tubercles absent; tip offingers not flattened or expanded; terminal grooves absent. Relative lengths of fingers,1<2<4<3. Prepollex not evident; nuptial pads or asperities absent.

Legs short, robust. Thigh length larger than tibia and smaller than foot length; tibialength 93.3% of thigh length. Sum of thigh and tibia lengths 80.8% of SVL; thigh



length 41.8% of SVL; tibia length 39.0% of SVL. Heel of adpressed legs failing to reachaxilla; knee and elbow widely separated when limbs laid along the sides; heels touchingwhen flexed legs held at right angle to body; knee and heel with a weak transversalskinfold; no tibial or tarsal ridges; a small oval inner but no outer metatarsal tubercle;plantar tubercles absent. Toes slender, free, not fringed; subarticular tuberclesdeveloped, rounded; supranumerary tubercles absent; tips of toes not flattened orexpanded; terminal grooves absent. Relative lengths of toes, 1<2<5<3<4. Skin smoothabove and beneath; anal opening not modified, no para-anal tubercles.In preservative, dorsum dark gray with small irregular gray spots scattered withoutforming defined pattern, and mid-longitudinal light stripe absent; venter gray withmany irregular cream spots regularly distributed; an undefined color transition betweenthe dorsal and ventral regions; small, defined white spots on axilla and groin; a shortirregular, poorly defined cream line on the posterior surface of the thighs; internalsufaces of tibiae with white spots; no light stripe on knee and tarsus.

Fig.2- Elachistocleis surumu sp.nov., holotype (MNRJ 25210; SVL 25.4mm), dorsal and ventralviews of body, and head profile.

POSIÇÃO TAXONÔMICA DAS “VARIEDADES” DE B. EPHIPPIUM... 12

Measurements of holotype in mm – SVL 25.4, HL 6.4, HW 7.1, IND 1.8, END 2.1, ED1.6, UEW 1.2, IOD 2.6, HAL 5.6, THL 10.6, TL 9.9, FL 11.4.Etymology – The name of the species, a noun in apposition, is an allusion to the typelocality, a small village currently included in the Reserva Indígena Raposa Serra doSol, in the State of Roraima, northern Brazil.Variation – Except for minor details in corporal proportions, the descripton of the holotypestands for males of the species. Size ranges from 19.6 to 27.0mm SVL and the intensityof the color of the throat can vary. The females are slightly bigger than males, sizeranging from 23.2 to 26.9mm SVL. Range, mean, and standard deviation of themeasurements of the type specimens of E. surumu sp.nov. are presented in table 2.Geographical distribution – Northern Brazil, in the State of Roraima (Fig.6).Remarks – The color in life, based on a photograph of a recently preserved specimennot specified, presents dorsum and flanks dark gray with small irregular light grayspots; venter, throat, and undersurfaces of limbs black with many irregular yellowspots; spots on axilla, groin, posterior surface of tibia, and thigh stripe, orange red;iris brown with intense black vermiculations.

TABLE 2. Range (mm), mean (x̄ ), and standard deviation (SD) of the measurements of the typespecimens of Elachistocleis surumu sp.nov. (n = number of specimens).


Elachistocleis carvalhoi sp.nov.(Fig.3)

Holotype – BRAZIL: TOCANTINS: Aragominas (07o10’S, 48o32’W; 345m altitude), MNRJ51384 , adult , collected by P.Fatorelli and L.Machado, 25/XI/2007.Paratypes – BRAZIL: TOCANTINS: Collected with the holotype, MNRJ 51385; NovaOlinda (07o38’S, 48o25’W; 257m), MNRJ 51386, collected by P.Fatorelli and L.Machado,28/XI/2007; Santa Fé do Araguaia (07o09’S, 48o42’W; 190m), MNRJ 48220, collectedby E.G.Pereira and P.C.F.Carneiro, VI/2007. PARÁ: Canaã dos Carajás (06o30’S,



49o53’W; 210m), Mineração Serra do Sossego, MNRJ 52474, collected by V.B.Assis,III/2002; MPEG 16265-16266, collected by V.B.Assis, III/2003; Marabá (05o22’S,49o07’W; 84m), Reserva Mãe Maria, MNRJ 52734, collected by H.Wogel and R.Bérnils,05/II/2008; Parauapebas (06o24’S, 49o54’W; 150m), Serra dos Carajás, MNRJ 58858,no collector, 12/II/2009; Piçarra (06o26’S, 48o52’W; 215m), MNRJ 51387, collected byP.Fatorelli and L.Machado, 16/XI/2007; São Geraldo do Araguaia (06o24’S, 48o33’W;145m), MNRJ 60285, collected by P.Fatorelli and D.D.Rocha, 07/VI/2009.Diagnosis – A medium sized species (SVL 24.0-31.9mm in males, 32.0-37.1mm infemales), characterized by the head length shorter than the head width, HL about 86.6%of HW (x̄ = 86.6; SD = 6.38; n = 9); postcommisural gland developed; dorsum smooth; inpreservative uniformly dark gray without marks or pattern; venter and flanks grayishwith large anastomosed whitish spots, producing a coarse marbled pattern, mainly onthe chest; large cream spots on axillae, groin, and posterior surfaces of tibiae; a broadirregular, poorly defined line on the posterior surface of the thighs; no light stripe onknee and tarsus.Comparisons with other species – The species with some kind of ventral color patternare E. cesarii, E. erythrogaster, E. magnus, E. piauiensis, E. skotogaster, E. surinamensis,E. surumu, and E. bumbameuboi sp.nov. The new species is distinguished from thesespecies by the dorsum uniformly brown or dark gray without marks (dorsal surfacesdark gray with small white dots in E. cesarii; dorsum dark gray marbled with black,blue, and some orange in males, dorsum orange marbled with black and blue in femalesof E. erythrogaster; dorsum uniform dark grayish with scarce minute brighter dots inthe outer boundaries in E. magnus; dorsum uniformly gray with scarce minute irregularbright dots in E. piauiensis; dorsum dark brown mottled with black and white spots inE. skotogaster; dorsum dark gray or black with an evident light vertebral stripe in E.surinamensis; dorsum dark gray with small irregular clear gray spots in E. surumu;dorsum uniformly dark gray or black without marks in E. bumbameuboi sp.nov.; colorin figures and/or described in KENNY, 1969, CARAMASCHI & JIM, 1983, KWET & DI BERNARDO,1998, LAVILLA et al., 2003, NUNES et al., 2010, TOLEDO, 2010, and TOLEDO et al., 2010); bythe venter grayish with large anastomosed yellow or whitish yellow blotches, producinga coarse marbled pattern, mainly in the chest area (chest yellow with gray marks andbelly white or yellow with gray marks and reticulations in E. cesarii; venter red orange,with hindlimbs largely black mottled in bright blue in E. erythrogaster; venter graywith minute scattered white spots, mainly on the belly and ventral surfaces of legs, inE. magnus; venter dull white or light yellow, heavily grayish spotted, marbled, in E.piauiensis; venter gray, mottled with dark brown in E. skotogaster; venter dark gray orblack with large yellow and small white spots in E. surinamensis; venter and throatgray with many irregular yellow spots regularly distributed in E. surumu; and ventergray with minute anastomosed whitish spots, producing a salt-and-pepper pattern, inE. bumbameuboi sp.nov.).The three species with immaculate venter and a sharp color limit between the dorsaland ventral regions are E. bicolor, E. helianneae, and E. matogrosso sp.nov. The newspecies is readily distinguished from these species by having venter grayish with largeanastomosed yellow or whitish yellow blotches, producing a coarse marbled pattern,mainly in the chest area, and an undefined color transition between the dorsal andventral regions.

14 U.CARAMASCHI


Description of holotype – Body ovoid, head small, triangular, slightly broader thanlong; head length 97.4% of head width and 25.0% of SVL; head width 25.7% of SVL.Snout rounded in dorsal view, protruding in profile. Nostrils small, not protuberant,directed laterally, closer to tip of snout than to eye; internarial distance smaller thanthe eye to nostril and interorbital distances, and larger than the eye diameter andupper eyelid width. Canthus rostralis rounded; loreal region flat, sloping abruptly tothe upper lip; lips not flared. Postcommisural gland developed. Eyes small,dorsolateral, only slightly protruding. Interorbital space slightly convex, 2.3 timesthe upper eyelid width. No cranial crests. A poorly defined transverse skinfold acrossback of the head, bending downwards slightly behind the eyes to the shoulder; askinfold through dorsolateral region from the axilla to the groin. Tympanum concealed;supratympanic fold absent. Lower jaw with truncate, trilobed anterior margin. Tonguelarge, oval, without a notch on posterior border. Choanae large, subcircular, widelyseparated. Vocal slits present. Premaxillary, maxillary, and vomerine odontophoresabsent. Vocal sac subgular, not expanded externally. A weak skinfold crossing thechest between axillae.

Fig.3- Elachistocleis carvalhoi sp.nov., holotype (MNRJ 51384; SVL 31.4mm), dorsal and ventralviews of body, and head profile.



Arms moderately robust, no tubercles or crests on forearm; palmar tubercle large,divided longitudinally, twice as large as the thenar tubercle; fingers slender, free, withsubarticular tubercles developed, rounded; supranumerary tubercles absent; tip offingers not flattened or expanded; terminal grooves absent. Relative lengths of fingers,1<2<4<3. Prepollex not evident; nuptial pads or asperities absent.Legs short, robust. Thigh length slightly shorter than tibia length and shorter thanfoot lengths; thigh length 97.9% of tibia length. Sum of thigh and tibia lengths 70.6%of SVL; thigh length 34.9% of SVL; tibia length 35.7% of SVL. Heel of adpressed legsfailing to reach axilla; knee and elbow widely separated when limbs laid along thesides; heels touching when flexed legs held at right angle to body; knee and heel witha weak transversal skinfold; no tibial or tarsal ridges; an small oval inner but noouter metatarsal tubercle; plantar tubercles absent. Toes slender, free, not fringed;subarticular tubercles developed, rounded; supranumerary tubercles absent; tips oftoes not flattened or expanded; terminal grooves absent. Relative lengths of toes,1<2<5<3<4. Skin smooth above and beneath; anal opening not modified, no para-anal tubercles.In preservative, dorsum uniformly dark gray with no light mid-dorsal stripe or pattern;venter grayish, with large anastomosed whitish blotches, producing a coarse marbledpattern, mainly on the chest; ventrolateral region with large whitish blotches; throatgrayish; large light cream spots on axillae, groin, and posterior surfaces of tibiae; alight cream, broad, irregular stripe on posterior surfaces of thighs; no light longitudinalstripe on superior surfaces of tibiae and posterior surfaces of tarsus.Measurements of holotype in mm – SVL 31.4, HL 7.85, HW 8.1, IND 2.2, END 2.6, ED1.9,UEW 1.5, IOD 3.4,HAL6.65, THL 11.0, TL 11.2, FL 15.0.Etymology – The species is named after the late Prof. Antenor Leitão de Carvalho(MNRJ), for his contribution to the knowledge of the neotropical anurans, especiallythe microhylid frogs.

TABLE 3. Range (mm), mean (x̄ ), and standard deviation (SD) of the measurements of the typespecimens of Elachistocleis carvalhoi sp.nov. (n = number of specimens).


16 U.CARAMASCHI


Variation – Except for minor details in corporal proportions, the description of theholotype stands for the male specimens examined. Size ranges from 24.0-31.9mmSVL in males. The females are bigger than males, their size ranging from 32.0-37.1mmSVL. Few specimens have the marbled pattern on venter and the spots on concealedsurfaces slightly coarser or thinner than the holotype.Remarks – In specimens recently preserved it is possible to observe that the color inlife is dark gray to black on dorsum; the spots on venter are pale yellow on graybackground; the blotches on axillae, groin, and posterior surfaces of tibiae, and thestripe on posterior surfaces of thighs are red orange.Geographical distribution – Northern Brazil, in northwestern State of Tocantins andsoutheastern State of Pará (Fig.6).

Elachistocleis bumbameuboi sp.nov.(Fig.4)

Holotype – BRAZIL: MARANHÃO: São Luís (02o32’S, 44o18’W; 24m altitude), UHE Ponta daMadeira, MNRJ 53200 (Fig.6), adult , collected by R.R.Carvalho and others, 02-08/V/2008.Paratypes – Collected with the holotype: MNRJ 53201-53205. Type locality, MNRJ53378, collected by I.Nunes and C.Canedo, IV/2008.Diagnosis – A medium sized species (SVL 26.9-27.7mm in males, 32.8-43.7mm infemales), characterized by the head length shorter than the head width, HL about 81%of HW (x¯ = 81; SD = 6.33; n = 5); postcommisural gland developed; dorsum slightlyrugose; in preservative, dorsum uniformly dark gray, without spots nor light mid-dorsal stripe; venter gray, with minute anastomosed whitish blotches, producing asalt-and-pepper pattern, extending to the ventrolateral region; throat gray; light spotson axillae, groin, posterior surfaces of tibiae, and superior surfaces of feet absent; alight, thin, irregular stripe on posterior surfaces of thighs; no light longitudinal stripeon superior surfaces of tibiae and posterior surfaces of tarsus.Comparisons with other species – The species with some kind of ventral color pattern areE. carvalhoi, E. cesarii, E. erythrogaster, E. magnus, E. piauiensis, E. skotogaster, E.surinamensis, and E. surumu. The new species is distinguished from these species by thedorsum uniformly dark gray or black without marks (dorsum uniformly brown or darkgray without marks in E. carvalhoi; dorsal surfaces dark gray with small white dots in E.cesarii; dorsum dark gray marbled with black, blue, and some orange in males, dorsumorange marbled with black and blue in females of E. erythrogaster; dorsum uniform darkgrayish with scarce minute brighter dots in the outer boundaries in E. magnus; dorsumuniformly gray with scarce minute irregular bright dots in E. piauiensis; dorsum darkbrown mottled with black and white spots in E. skotogaster; dorsum dark gray or blackwith an evident light vertebral stripe in E. surinamensis; and dorsum dark gray with smallirregular clear gray spots in E. surumu; color in figures and/or described in KENNY, 1969,CARAMASCHI & JIM, 1983, KWET & DI BERNARDO, 1998, LAVILLA et al., 2003, NUNES et al., 2010,TOLEDO, 2010, and TOLEDO et al., 2010); by the venter gray with minute anastomosed whitishspots, producing a salt-and-pepper pattern (venter grayish with large anastomosed yellowor whitish yellow blotches, producing a coarse marbled pattern, mainly in the chest areain E. carvalhoi; chest yellow with gray marks and belly white or yellow with gray marks



and reticulations in E. cesarii; venter red orange, with hindlimbs largely black mottled inbright blue in E. erythrogaster; venter gray with minute scattered white spots, mainly onthe belly and ventral surfaces of legs, in E. magnus; venter dull white or light yellow,heavily grayish spotted, marbled, in E. piauiensis; venter gray, mottled with dark brown inE. skotogaster; venter dark gray or black with large yellow and small white spots in E.surinamensis; and venter and throat gray with many irregular yellow spots regularlydistributed in E. surumu).The three species with immaculate venter and a sharp color limit between the dorsaland ventral regions are E. bicolor, E. helianneae, and E. matogrosso. The new species isreadily distinguished from these species by having venter gray with minute anastomosedwhitish spots, producing a salt-and-pepper pattern and an undefined color transitionbetween the dorsal and ventral regions.

Fig.4- Elachistocleis bumbameuboi sp.nov., holotype (MNRJ 53200; SVL 26.9mm), dorsal andventral views of body, and head profile.

18 U.CARAMASCHI


Description of holotype – Body ovoid, head small, triangular, broader than long; headlength 80.0% of head width and 21.2% of SVL; head width 26.4% of SVL. Snout roundedin dorsal view, protruding in profile. Nostrils small, not protuberant, directed laterally,closer to tip of snout than to eye; internarial distance smaller than the eye to nostril andinterorbital distances, and larger than the eye diameter and upper eyelid width. Canthusrostralis rounded; loreal region flat, sloping abruptly to the upper lip; lips not flared.Postcommisural gland developed. Eyes small, dorsolateral, only slightly protruding.Interorbital space slightly convex, almost two and half times the upper eyelid width. Nocranial crests. A well defined transversal skinfold across back of the head, bendingdown and backwards slightly behind the eyes to the shoulder; a strong dorsolateral skinfold from the scapular region to the groin. Tympanum concealed; supratympanic foldabsent. Lower jaw with truncate, trilobed anterior margin. Tongue large, oval, without anotch on posterior border. Choanae large, subcircular, widely separated. Vocal slitspresent. Premaxillary, maxillary, and vomerine odontophores absent. Vocal sac subgular,not expanded externally. A skinfold crossing the chest between axillae.Arms moderately robust, no tubercles or crests on forearm; palmar tubercle large,divided longitudinally, twice as large as the thenar tubercle; fingers slender, free, withsubarticular tubercles developed, rounded; supranumerary tubercles absent; tip offingers not flattened or expanded; terminal grooves absent. Relative lengths of fingers,1<2<4<3. Prepollex not evident; nuptial pads or asperities absent.Legs short, robust. Tibia length slightly longer than thigh and shorter than foot length;thigh length 99.3% of tibia length. Sum of thigh and tibia lengths 75.4% of SVL; thighlength 37.8% of SVL; tibia length 37.6% of SVL. Heel of adpressed legs failing to reachaxilla; knee and elbow widely separated when limbs laid along the sides; heels touchingwhen flexed legs are held at right angle to body; knee and heel with a weak transversalskinfold; no tibial or tarsal ridges; an small oval inner but no outer metatarsal tubercle;plantar tubercles absent. Toes slender, free, not fringed; subarticular tuberclesdeveloped, rounded; supranumerary tubercles absent; tips of toes not flattened orexpanded; terminal grooves absent. Relative lengths of toes, 1<2<5<3<4. Skin smoothabove and beneath; small dermal spines scattered mainly on dorsolateral region; analopening not modified, no para-anal tubercles.In preservative, dorsum uniformly dark gray, without spots nor light mid-dorsal stripe;venter gray, with minute anastomosed whitish blotches, producing a salt-and-pepperpattern, extending to the ventrolateral region; throat gray; light spots on axillae, groin,posterior surfaces of tibiae, and superior surfaces of feet absent; a light, thin, irregularstripe on posterior surfaces of thighs; no light longitudinal stripe on superior surfacesof tibiae and posterior surfaces of tarsus.Measurements of holotype in mm – SVL 26.9, HL 5.7, HW 7.1, IND 1.7, END 2.1; ED1.4, UEW 1.1, IOD 2.6, HAL 5.8, THL 10.2; TL 10.1, FL 11.9.Etymology – The name of the species, a noun in apposition, is allusive to the mostpopular feasts occurring in June at São Luís, Maranhão, the “Bumba-meu-boi”, whichcan be “played” through several ways, with distinct characteristics of rhythms, clothes,choreographies, characters, and musical instruments. These festivities narrate theadventures of a cowboy and the death and resurrection of an ox. A suite follows the“ox” that, through its dance, instigates the participants to take part in the play.



Variation – Except for minor details in corporal proportions, the description of theholotype stands for the male specimens examined. Size ranges from 26.9-27.4mmSVL in males. The females are bigger than males, their size ranging from 32.8-43.7mmSVL. A few specimens have the venter pattern slightly coarser than the holotype.Geographical distribution – Known from two localities, in the State of Maranhão,northeastern Brazil (Fig.6).Remarks – A good color picture of Elachistocleis bumbameuboi sp.nov. in life, treatedas E. piauiensis, is presented by NUNES et al. (2010). Dorsum uniformly dark gray,without marks; venter gray with small clear gray small spots producing a salt-and-pepper pattern, extending to the ventrolateral region; spots on groin and line on posteriorsurfaces of thighs orange red; iris brown with dense black vermiculations. Theadvertisement call is described and, in a note added in the proof of the paper, theauthors pointed out differences between the call they described and that described byTOLEDO et al. (2010) for E. piauiensis based on recordings made at Pacatuba, Caucaia,and Viçosa do Ceará, State of Ceará, Brazil.

(n = 3) (n = 3) CHARACTERS RANGE x̄ SD RANGE x̄ SD

SVL 26.9-28.8 27.8 0.96 32.8-43.7 37.4 5.65 HL 5.7-6.0 5.9 0.19 6.1-8.3 7.2 1.07 HW 7.1-8.1 7.5 0.54 8.0-9.8 8.6 1.06 IND 1.6-1.85 1.7 0.11 2.1-2.5 2.25 0.19 END 2.1-2.4 2.25 0.15 2.6-3.1 2.8 0.22 ED 1.4-1.7 1.6 0.15 1.7-2.0 1.8 0.17 UEW 1.1-1.4 1.3 0.19 1.4-1.6 1.5 0.12 IOD 2.3-3.0 2.6 0.37 2.9-3.6 3.3 0.39 HAL 5.8-6.9 6.4 0.54 7.1-9.0 7.7 1.07 THL 10.2-11.7 11.1 0.79 12.25-14.5 13.2 1.17 TL 10.1-11.1 10.6 0.51 11.95-14.3 12.9 1.23 FL 11.9-12.8 12.4 0.50 13.3-16.7 14.7 1.80

TABLE 4. Range (mm), mean ( x̄ ), and standard deviation (SD) of the measurements of the typespecimens of Elachistocleis bumbameuboi sp.nov. (n = number of specimens).

Elachistocleis matogrosso sp.nov.(Fig.5)

Holotype – BRAZIL: MATO GROSSO: Cuiabá (15o36’S, 56o06’W; 177m altitude), MNRJ4812, adult , collected by Ulisses Caramaschi, 03/X/1987.Paratypes – BRAZIL: MATO GROSSO: Collected with the holotype, MNRJ 4813; Cuiabá,Campus of the Universidade Federal do Mato Grosso, MNRJ 6994, collected byJ.Langone, 04/II/1986; CHUNB 47526, collected by LPP, 04/II/1986; Cuiabá, bankof the Cuiabá river, MNRJ 43841, no collector, no date; Alto Paraguai (14o30’S, 56o29’W;221m altitude), Primavera, MNRJ 6977, no collector, no date; Barão de Melgaço (16o12’S,

20 U.CARAMASCHI


55o58’W; 156m altitude), RPPN Sesc Pantanal, MNRJ 32880-32882, collected byC.A.Caetano and L.F.B.Oliveira, 15/X/1999; Poconé (16o15’S, 56o37’W; 142m altitude),Base de Pesquisas do Pantanal - IBAMA, MHNCI 661, no collector, no date; Poconé,Fazenda Ipiranga, CHUNB 47531, collected by B.Duar, 14/VII/2001.Diagnosis – A small sized species (SVL 21.5-24.6mm in males, 29.0-33.2mm in females),characterized by the head length shorter than the head width, HL about 92.0% of HW(x̄ = 92.0; SD = 5.49; n = 8); postcommisural gland poorly developed; dorsum smooth;in preservative, uniformly grayish brown; a thin middle longitudinal light stripe, fromthe post-cephalic transverse skinfold to the vent, but absent on the head; venterimmaculate; limit between the dorsal and ventral regions poorly defined; light spotson axillae or groin present; a broad irregular line on the posterior surface of the thighs;a large light spot on the proximal internal surface of tibiae; a narrow light stripesurrounding the knees and reaching the middle of the tarsus.Comparisons with other species – The two species with immaculate venters are E.bicolor and E. helianneae. The new species is readily distinguished from E. bicolorby the longer head (HL about 92% of HW in H. matogrosso sp.nov.; HL below 90% ofHW in E. bicolor), but smaller than in E. helianneae (HL above 94% of HW in E.helianneae), by the presence of the mid-dorsal stripe from the post-cephalic dermalfold to vent (mid-dorsal stripe absent in E. bicolor, mid-dorsal stripe from the tip ofsnout to vent in E. helianneae), by the absence of minute light spots on dorsum anddorsal surfaces of members (present in E. helianneae), by the dorsal gray color ofdorsum of the snout invading the loreal region almost to the upper lip border (lorealregion conspicuously white in E. bicolor and in E. helianneae), and by the stripe onthe posterior surface of thighs broad, irregular (thin, well defined in E. bicolor andin E. helianneae).All other species in the genus Elachistocleis present some type of ventral color pattern(venter immaculate in E. matogrosso sp.nov.)Description of holotype – Body ovoid, head small, triangular, slightly broader thanlong; head length 92.8% of head width and 22.1% of SVL; head width 23.8% ofSVL. Snout sub-elliptical in dorsal view, protruding in profile. Nostrils small, notprotuberant, directed anterolaterally, closer to tip of snout than to eye; internarialdistance smaller than the eye to nostril and interorbital distances, and largerthan the eye diameter and the upper eyelid width. Canthus rostralis rounded;loreal region flat, sloping abruptly to the upper lip; lips not flared. Postcommisuralgland poorly developed. Eyes small, dorsolateral, only slightly protruding.Interorbital space slightly convex, more than twice the upper eyelid width. Nocranial crests. A poorly defined transversal skinfold across back of the head,bending downwards slightly behind the eyes to the shoulder; a poorly definedskinfold through dorsolateral region from the axilla to the groin. Tympanumconcealed; supratympanic fold absent. Lower jaw with truncate, trilobed anteriormargin. Tongue large, oval, without a notch on posterior border. Choanae large,subcircular, widely separated. Vocal slits present. Premaxillary, maxillary, andvomerine odontophores absent. Vocal sac subgular, not expanded externally. Aweak skinfold crossing the chest between axillae.Arms moderately robust, no tubercles or crests on forearm; palmar tubercle large,



divided longitudinally, twice as large than thenar tubercle; fingers slender, free, withsubarticular tubercles developed, rounded; supranumerary tubercles absent; tip offingers not flattened or expanded; terminal grooves absent. Relative lengths of fingers,1<2<4<3. Prepollex not evident; nuptial pads or asperities absent.Legs short, robust. Thigh length shorter than tibia and foot lengths; thigh length90.9% of tibia length. Sum of thigh and tibia lengths 67.5% of SVL; thigh length32.15% of SVL; tibia length 35.4% of SVL. Heel of adpressed legs not reaching axilla;knee and elbow widely separated when limbs laid along the sides; heels touchingwhen flexed legs held at right angle to body; knee and heel with a transversal skinfold;no tibial or tarsal ridges; an oval inner but no outer metatarsal tubercle; plantartubercles absent. Toes slender, free, weakly fringed; subarticular tubercles developed,rounded; supranumerary tubercles absent; tips of toes not flattened or expanded;terminal grooves absent. Relative lengths of toes, 1<2<5<3<4. Skin on dorsum slightlyrugose, smooth beneath; anal opening not modified, no para-anal tubercles.

Fig.5- Elachistocleis matogrosso sp.nov., holotype (MNRJ 4812; SVL 33.2mm), dorsal and ventralviews of body, and head profile.

22 U.CARAMASCHI


In preservative, dorsum and dorsal surfaces of limbs uniformly grayish brown; a mid-dorsal longitudinal light cream stripe, from the post-cephalic dermal fold to vent; apoorly defined color limit between the dorsal and ventral regions; venter immaculateclear cream; throat grayish; few, small spots on axillae and groin; an irregular broadlight cream stripe on the posterior surface of thighs; a large irregular, light cream spoton the proximal internal surface of tibiae; a light cream stripe surrounding the kneesand reaching the middle of the tarsus.Measurements of holotype in mm – SVL 33.2, HL 7.3, HW 7.9, IND 2.1, END 2.6, ED1.6, UEW 1.4, IOD 3.0, HAL 6.2, THL 10.7, TL 11.7, FL 13.9.Etymology – The specific name, a noun in apposition, honors both states of MatoGrosso and Mato Grosso do Sul, Brazil, where the species occurs.Variation – Except for minor details in corporal proportions, the descripton ofthe holotype stands for females of the species. Size ranges from 29.0 to 33.2mmSVL and the intensity of the color of the throat can vary. The males are smallerthan females, size ranging from 21.5 to 24.6mm SVL. Range, mean, and standarddeviation of the measurements of the type specimens of E. matogrosso arepresented in Table 5. In a few specimens the mid-dorsal stripe can be interruptedor almost absent.Geographical distribution – Central Brazil, in southwestern State of Mato Grossoand northwestern State of Mato Grosso do Sul (Fig.6).


TABLE 5. Range (mm), mean (x̄ ), and standard deviation (SD) of the measurements of the typespecimens of Elachistocleis matogrosso sp.nov. (n = number of specimens).



Fig.6- Geographic distribution of species of Elachistocleis.

24 U.CARAMASCHI


DISCUSSION

The confusion involving the taxonomy of Elachistocleis ovalis, E. bicolor, and E. surinamensis,the three oldest species currently in the genus Elachistocleis, was summarized by LAVILLA

et al. (2003). Elachistocleis ovalis, presently considered a species inquirenda, will not affectthe taxonomical composition of the genus until its actual status can be settled.Elachistocleis bicolor had its type locality convincingly restricted by LAVILLA et al. (2003) toBuenos Aires, Argentina. The species, in their conception, would encompasses frogsthat, filling the characters that describe the genus Elachistocleis, had an immaculateventer and would occupy the southern part of the generic range. The authors, however,stated that frogs with this set of characters form a complex of species and their presenteddecision constituted an operative framework for a necessary revision (LAVILLA et al., 2003).Elachistocleis bicolor is characterized by the medium size (SVL 22.9-31.5mm in males,27.2-35.5mm in females; RODRIGUES et al., 2003), head wider than long (head lengthbelow 90% of head width), dorsum smooth, uniformly grayish brown, without light spotsnor a distinctive middle longitudinal light stripe, venter immaculate yellow or greenishyellow, a sharp color limit between the dorsal and ventral regions, no spots on axillae orgroin, a thin, well defined line on the posterior surface of the thighs, and a narrow lightstripe surrounding the knees and reaching the middle of the tarsus. The geographicaldistribution (Fig.6) comprises southwestern and southern Brazil, in the states of MatoGrosso do Sul, southern São Paulo, Paraná, Santa Catarina, and Rio Grande do Sul,Paraguay (for detailed distribution see BUSQUETTI & LAVILLA, 2006), Uruguay (for detaileddistribution see NÚÑEZ et al., 2004), and northern Argentina.Elachistocleis surinamensis was originally characterized by DAUDIN (1802) as having abrown belly, mottled with gray, which was also reported by KENNY (1969), RIVERO et al.(1986), CARCERELLI (1992), and MURPHY (1997); LAVILLA et al. (2003) added the presence ofan evident light vertebral stripe. The type locality was originally stated as “Surinam”, abroad geographical concept during the XVIII-XIX centuries according to LAVILLA et al.(2003); the name bearing type, a specimen donated to Daudin by M. de Bèze and probablyoriginally deposited in the Museum Nationalle d’Histoire Naturelle de Paris, is currentlylost. As an operative framework for a necessary revision of this species complexrepresented by E. surinamensis, LAVILLA et al. (2003) considered that it would berepresented by those frogs that fit the characters that describe the genus Elachistocleis,have a spotted ventral coloration, and inhabit the northern portion of the generic range,namely Trinidad, northern Venezuela, and northern Surinam.Elachistocleis cesarii was revalidated and well characterized by TOLEDO et al. (2010), whoreported the geographical distribution for the species from a few localities in the statesof São Paulo, Minas Gerais, and Goiás, Brazil. Actually, the geographical distribution ofE. cesarii (Fig.6) involves northeastern Brazil, in the states of Ceará, Sergipe, and Bahia,central Brazil, in the states of Mato Grosso, Goiás, and Federal District, and southeasternBrazil, in the states of Minas Gerais, Espírito Santo, Rio de Janeiro, and São Paulo. It ispossible that more than one species is involved, but at first not discriminated by theused characters.The remaining species, including those described herein, have modern descriptions, definedname bearing types, and precise type localities. Elachistocleis piauiensis is distributed inthe states of Piauí, Maranhão, Ceará, and Tocantins (see map in NUNES et al., 2010, excluding



the locality record for São Luís, State of Maranhão), and Mato Grosso (present data; Fig.6).Elachistocleis erythrogaster is restricted to the southeastern border of the Planalto dasAraucárias, Serra Geral, State of Rio Grande do Sul, southern Brazil, at 900-1200m ofaltitude (FROST, 2010) (Fig.6). Elachistocleis skotogaster is known from the type locality,Los Toldos, 1100m altitude, Departamento Santa Victoria (LAVILLA et. al., 2003), and fromone locality near to Isla de Cañas, 800m altitude, Departamento Iruya, and two localitiesin the Departamento Orán, 350 and 450m altitude, all in the Provincia Salta, northernArgentina (see map in CAJADE et al., 2009, and Fig.6). Elachistocleis magnus is known fromthree localities in the State of Rondônia, Brazil (Fig.6). The geographic distributions of thenew species are treated in the respective accounts.The present observations and descriptions contribute to a better undertanding of thecomposition of Elachistocleis. Notwithstanding, the actual diversity encompassed bythis genus is far from known, as can be seen, only as examples, the pictures of thespecies, all identified as E. ovalis, presented by KENNY (1969, plate XIIIb) from Trinidad,MURPHY (1997, plate 50) from Manzanilla-Cocos Bay, Trinidad, GORZULA & SEÑARIS (1999,fig.63) from El Manteco, Venezuelan Guayana, DE LA RIVA et al. (2000) from Rio Seco,Santa Cruz, Bolivia, and LESCURE & MARTY (2001, p.272) from Guyane.

ACKNOWLEDGMENTS

I deeply acknowledge Ana Maria Costa Prudente (MPEG), Hussam Zaher and CarolinaMello (MZUSP), Júlio César Moura Leite (MHNCI), Marcelo F. Napoli (UFBA), Renato N.Feio (MZUFV), Luciana B. Nascimento (MCNAM), W. Ronald Heyer (USNM), Guarino R.Colli (CHUNB), Célio F.B. Haddad (CFBH), and Oswaldo Luiz Peixoto (EI) for allowingexamination of specimens under their care and for data access on related species.Helianne de Niemeyer photographed the holotypes. Carlos Alberto G. Cruz (MNRJ)and W. Ronald Heyer (USNM) critically read the manuscript. The Conselho Nacionalde Desenvolvimento Científico e Tecnológico (CNPq) provided financial support.

REFERENCES

BUSQUETTI, F. & LAVILLA, E.O., 2006. Lista comentada de los anfibios de Paraguay. Cuadernosde Herpetología, 20(2):3-79.

CAJADE, R.; BARRASSO, D.A. & NENDA, S.J., 2009. Amphibia, Anura, Microhylidae, Elachistocleisskotogaster: Map of geographic distribution, distribution extension, and new altitudinal records.Check List, 5(2):418-421.

CARAMASCHI, U. & JIM, J., 1983. A new microhylid frog, genus Elachistocleis (Amphibia, Anura),from Northeastern Brasil. Herpetologica, 39(4):390-394.

CARCERELLI, L.C., 1992. Revisão taxonômica do gênero Elachistocleis Parker, 1927. XII CongresoLatino-Americano de Zoologia e XIX Congresso Brasileiro de Zoologia, Resumos, Belém(PA):124.

CUVIER, B., 1829. Le Règne Animal Distribué d’Après son Organisation, pour servir de basea l’Histoire Naturelle des Animaux et Introduction a l’Anatomie Comparée. Paris: Détervilleet Crochard Libraires. xv + 406p.

DAUDIN, F.M., 1802 (An XI). Histoire Naturelle des Rainettes, des Grenouilles et des Crapauds.Paris: Bertrandet. 108p.

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DE LA RIVA, I.; KÖHLER, J.; LÖTTERS, S. & REICHLE, S., 2000. Ten years of research onBolivian amphibians: updated checklist, distribution, taxonomic problems, literature andiconography. Revista Española de Herptologia, 14(2000):19-164.

DUMÉRIL, A.M.C. & BIBRON, G., 1841. Erpétologie Générale ou Histoire Naturelle Complètedes Reptiles. Tome 8. Paris: Librairie Encyclopédique de Roret. ii + 792p.

FITZINGER, L.J., 1826. Neue Classification der Reptilien nach ihren NatürlichenVerwandtschaften. Nebst einer Verwandtschafts-Tafel und einem Verzeichnisse der Reptilien-Sammlung des K. K. Zoologischen Museum’s zu Wien. Wien: J.G. Heubner. 66p. 1pl.

FROST, D.R., 2010. Amphibian Species of the World: an Online Reference. Version 5.4 (8April 2010). Eletronic Database accessible at: http://research.amnh.org/vz/herpetology/amphibia/. American Museum of Natural History, New York, USA. Accessed: 24/IV/2010.

GORZULA, S. & SEÑARIS, J.C., 1999 [1998]. Contributions to the herpetofauna of theVenezuelan Guayana. I. A data base. Caracas: Scientia Guayanae, 8. xviii + 270p. 129pls.

GRONOVIUS, L.T., 1763. Zoophylacii Gronoviani. Fasciculus Primus exhibens AnimaliaQuadrupedia, Amphibia atque Pisces quae in Museo suo adservat, rite examinavit, systematicedisposuit, descripsit, atque iconibus ilustravit. Lugduni Batavorum, Sumptibus Auctoris.

HEYER, W.R.; RAND, A.S.; CRUZ, C.A.G.; PEIXOTO, O.L. & NELSON, C.E., 1990. Frogs ofBoracéia. Arquivos de Zoologia, 31(4):231-410.

ICZN – International Commission on Zoological Nomenclature, 1999. International Code ofZoological Nomenclature. 4th Edition. London: International Trust for Zoological Nomenclature.xxx + 306p.

KENNY, J.S., 1969. The Amphibia of Trinidad. Studies on the Fauna of Curaçao and OtherCaribbean Islands, 29:1-78, xv pls.

KLAPPENBACH, M.A. & LANGONE, J.A., 1992. Lista sistematica y sinonimica de los anfibios delUruguay, con comentarios y notas sobre su distribución. Anales del Museo Nacional de HistoriaNatural de Montevideo, 8:163-222.

KWET, A. & DI BERNARDO, M., 1998. Elachistocleis erythrogaster, a new microhylid speciesfrom Rio Grande do Sul, Brazil. Studies on the Neotropical Fauna and Environment, 33:7-18.

LANGONE, J.A., 1995. Ranas y sapos del Uruguay (reconocimiento y aspectos biológicos). MuseoA.D. Larrañaga, Série Divulgación, 5:1-123.

LAVILLA, E.O.; VAIRA, M. & FERRARI, L., 2003. A new species of Elachistocleis (Anura:Microhylidae) from the Andean Yungas of Argentina, with comments on the Elachistocleis ovalis –E. bicolor controversy. Amphibia-Reptilia, 24(3):269-284.

LESCURE, J. & MARTY, C., 2001 [2000]. Atlas des Amphibiens de Guyane. Paris: PatrimoinesNaturels 45. 390p.

LIMA, A.P.; MAGNUSSON, W.E.; MENIN, M.; ERDTMANN, L.K.; RODRIGUES, D.J.; KELLER, C. &HÖDL, W., 2006. Guia de Sapos da Reserva Adolpho Ducke, Amazônia Central – Guide to theFrogs of Reserva Adolpho Ducke, Central Amazonia. Manaus: Áttema Design Editorial. 168p.

MERREM, B., 1820. Versuch eines System der Amphibien (Tentamen SystematisAmphibiorum). Marburg: J.C. Krieger.

MIRANDA-RIBEIRO, A., 1920. Os engystomatideos do Museu Paulista (com um gênero e trêsespécies novos). Revista do Museu Paulista, 12:281-288, 2 pls.

MURPHY, J.C., 1997. Amphibians and Reptiles of Trinidad and Tobago. Malabar: Krieger



Publishing Company. xiii + 245p.

NUNES, I.; CANEDO, C. & CARVALHO JR., R.R., 2010. Advertisement call and geographicdistribution of Elachistocleis piauiensis Caramaschi & Jim, 1983 (Amphibia, Microhylidae), withnotes on the presence of post-commisural gland in the genus. South American Journal ofHerpetology, 5(1):30-34.

NÚÑEZ, D.; MANEYRO, R.; LANGONE, J. & DE SÁ, R.O., 2004. Distribución geográfica de lafauna de anfíbios del Uruguay. Smithsonian Information Herpetological Service (134):1-34.

RIVERO, J.A.; LANGONE, J.A. & PRIGIONI, C.M., 1986. Anfibios anuros colectados por laespedición del Museo Nacional de Historia Natural de Montevideo al Rio Caura, Estado Bolivar,Venezuela; con la descripción de una nueva especie de Colostethus (Dendrobatidae).Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo, 11(157):1-15.

RODRIGUES, D.J.; LOPES, F.S & UETANABARO, M., 2003. Padrão reprodutivo de Elachistocleisbicolor (Anura, Microhylidae) na Serra da Bodoquena, Mato Grosso do Sul, Brasil. Iheringia,Série Zoologia, 93(4):365-371.

SCHNEIDER, J.G., 1799. Historiae Amphibiorum Naturalis et Literariae. Fasciculus PrimusContinens Ranas, Calamitas, Bufones, Salamandras et Hydros in Genera et Species DescriptosNotisque Suis Distinctos. Jena: Friederici Frommanni. i-xiii + 264p.

SEBA, A., 1735. Locupletissimi Rerum Naturalium Thesauri Accurata Descriptio, et IconibusArtificiosissimis Expressio, per Universsam Physices Historiam. Opus, cui, in hoc RerumGenere, Nullum par Exstitit. Ex Toto Terrarum Orbe Collegit, Digessit, Descripsit, etDepingendum Curavit. Tomus II. Amstelaedami: J. Wetstenium, & Gul. Smith, & Janssonio-Waesbergios.

SHAW, G., 1802. General Zoology or Systematic Natural History. London: T. Davidson, 3:1-615.

TOLEDO, L.F., 2010. A new species of Elachistocleis (Anura; Microhylidae) from the BrazilianAmazon. Zootaxa, 2496:63-68.

TOLEDO, L.F.; LOEBMANN, D. & HADDAD, C.F.B., 2010. Revalidation and redescription ofElachistocleis cesarii (Miranda-Ribeiro, 1920) (Anura: Microhylidae). Zootaxa, 2418:50-60.

APPENDIX

ADDITIONAL SPECIMENS EXAMINED

Elachistocleis bicolor – BRAZIL: MATO GROSSO DO SUL: Bela Vista (EI 1400-1418); Bonito (CHUNB49299); Maracaju (EI 4010-4013); Ponta Porã, Inhuverá (MNRJ 6955); Três Lagoas (CFBH 13608-13610). SÃO PAULO: Castilho (MZUSP 36500, 36501-36502); Fartura (UFBA 7956-7958); Piraju,Pedreira (MNRJ 30404,66897-66902); Tabapuã (CFBH 4225). PARANÁ: Bituruna (MNRJ 2745,3717, 3909, 6932-6947, 17135-17137); Bituruna, 80km from União da Vitória (MNRJ 4815-4817); Curitiba (MZUSP 13659-13661); Jaguariaíva (MNRJ 66159); Piraquara, Mananciais daSerra (MHNCI 1181). SANTA CATARINA: (MHNCI 1181a,b); Faxinal dos Guedes, Represa SantaLaura (MNRJ 48298); Florianópolis (MNRJ 13975-13977); Lagoa (MZUSP 12527-12528, 36384,36385); Novo Horizonte (MZUSP 35445-35449); São Bento do Sul (MZUSP 55933); São Domingos(CFBH 3841, 3859, 4010). RIO GRANDE DO SUL: (MNRJ 2087); Bom Jesus (CFBH 18199);Emboaba (EI 5614); Gramado (MZUSP 16032-16036); Itaqui (MZUSP 0511); Osório (MZUSP21727); Porto Alegre (MNRJ 3658, MZUSP 21724-21726); Porto Alegre, Vila Florida, Rio Guaíba(MZUSP 16055); Santo Augusto (MNRJ 6984); São Francisco de Paula (MNRJ 3654, 6973, EI756); Severiano de Almeida, Cerro do Meio Dia (MNRJ 4518-4520); Torres (AL-MN 549); Viamão(MZUSP 64753). PARAGUAY: AMAMBAY (USNM 253204-253205). Itapúa (USNM 253509, 253510-

28 U.CARAMASCHI


253516, 253517, 253518, 253519, 253520-253521, 253522-253523, 253524, 253525).CAAGUAZU: Ruta Ciudad del Este-Assunción, Km 217 (MNRJ 66896); Puerto Bertoni (USNM94101); Entre Independencia y Villarica, Arroio Ytá (MNRJ 6975-6976); Brejo de Ipuã, nearNueva Italia (MNRJ 66903). URUGUAY: ARTIGAS: Barra del Arroio Yacuí (MNRJ 6982-6983).ARGENTINA: CHACO: Resistencia (MZUSP 36386). CORRIENTES: Ituzaingó, Santa Tecla (MNRJ39940-39942). TUCUMÁN: Tucumán (MNRJ 3462, 12490-12491).

Elachistocleis bumbameuboi - BRAZIL: MARANHÃO: Carolina (CHUNB 51588-51590, 51649-51656).

Elachistocleis carvalhoi - BRAZIL: PARÁ: Carajás (CHUNB 47529).

Elachistocleis cesarii - BRAZIL: CEARÁ: Caucaia, Estação Ecológica do Pecém (MNRJ 55891);Fortaleza, Mucuripe (MNRJ 6956-6959). SERGIPE: Brejo Grande, Fazenda Capivara (MNRJ 49529,49532-49536). BAHIA: Barreiras (UFBA 8635-8646, 9325-9329, 9854-9859); Camaçari, Arembepe(UFBA 6233-6235); Cocos (CHUNB 50142, 50146, 50171, 51551); Correntina (CHUNB 47547-47548, 47550); Jaborandi (CHUNB 51009, 51013-51014); Juazeiro (UFBA 6776); Porto Seguro(UFBA 9199-9200); Riachão das Neves (CHUNB 47552); São Desidério (CHUNB 51010-51012).DISTRITO FEDERAL: Brasília (MNRJ 7018-7019); Planaltina, Lagoa Bonita (MNRJ 18330-18331).GOIÁS: Alvorada do Norte (CHUNB 38038-38043); Aporé, UHE Espora (MNRJ 41395-41396);Aruanã (CHUNB 42710); Catalão (CHUNB 50334); Colinas do Sul (CHUNB 48336, 50333); Floresde Goiás (CHUNB 38421-38428); Goiânia (MNRJ 6682, 66850); Minaçu, UHE Serra da Mesa(MNRJ 20203-20207, MPEG 8944-8948, CHUNB 07137-07138); Mineiros (CHUNB 28126-28127);Pires do Rio (CHUNB 38686); Pontalina, Fazenda Lagoa Grande (MNRJ 32396-32398); Pontalina(MNRJ 66853, CFBH 3768); Porangatu (MNRJ 53151-53152); Rio São Miguel (MNRJ 2772); RioVerde (CHUNB 49435); Santa Tereza (MNRJ 53149-53150); São Domingos (CHUNB 32223-32224,35568-35576, 47527-47528, 47530); São João da Aliança, Jatobazinho (MNRJ 6974); TrêsRanchos (CHUNB 44730). MATO GROSSO: Barra do Tapirapé (MNRJ 7021-7037); Diamantino,Fazenda São João, Km 200 da BR 163 (MNRJ 49659-49660); Pontes e Lacerda (MZUSP 61219).MATO GROSSO DO SUL: Xavantina, Rio Areões (MNRJ 66856). MINAS GERAIS: Abre Campo,Fazenda Cachoeira Alegre (MNRJ 43302-43306); Alto Caparaó (CHUNB 46259); Araguari, FazendaCuiabá (MZUFV 2973); Andrequicé, UHE Formoso (MNRJ 13891); Araponga (MZUFV 8279-8280);Astolfo Dutra, Triunfo (MCNAM 6542); Belmiro Braga, Brumadinho (MNRJ 27503); Betim (MNRJ66849, MCNAM 2047, 13060); Botumirim, Veredas de Botumirim (MCNAM 5671); Brumadinho(MNRJ 12495); Brumadinho, Inhotim (MCNAM 13040-13044); Brumadinho, Serra da Calçada(MCNAM 3684); Buritizeiro (CHUNB 44628); Caeté (MCNAM 11468, MZUFV 427); Capim Branco(MCNAM 6307); Carangola (MZUFV 3782, 4824); Catas Altas, Serra do Caraça, Campinho (MNRJ56303); Catas Altas, Serra do Caraça (MNRJ 44708, 60489-60493, 66029); Catas Altas (MCNAM12884-12885); Chapada Gaúcha (CHUNB 34082-34090); Conceição do Mato Dentro, Taboleiro(MCNAM 3115); Conceição do Mato Dentro, Itacolomi (MCNAM 11767, 12114-12115, 12189,13240); Conselheiro Mata (MCNAM 605, 3153); Divinópolis, Parque Ecológico Gafanhoto (MCNAM9546-9547); Esmeraldas (MCNAM 2375, MZUFV 1713); Faria Lemos, Fazenda Todos os Santos(MNRJ 41599-41601); Grão Mogol/Berilo, UHE Irapé (MCNAM 3840, 6633, 9819); Guanhães(MCNAM 1092-1095, 3186); Guaraciaba, PCH Jurumirim (MZUFV 5545); Itambé do Mato Dentro(MCNAM 5670); Itumirim (MCNAM 3559); Jaboticatubas, Serra do Cipó (MNRJ 45346); Januária,RPPN Porto Cajueiro (MCNAM 13425-13427); João Pinheiro, Fazenda Fruta Danta (MNRJ 50705-50707); João Pinheiro, Fazenda Veredas (MNRJ 38813); Juiz de Fora, Água Limpa (MNRJ 6960-6961, 6971-6972); Manga, Mocambinho (MNRJ 40589, MZUFV 2838-2839); Mariana, Samitri(MZUFV 578-583); Marliéria, Parque Estadual do Rio Doce (MNRJ 13980, 51092, 51093, MZUFV4839-4840); Mocambinho, Jaíba (MZUFV 2838-2839, 3029); Nova Era (MCNAM 1112); OuroBranco, Serra do Ouro Branco (MZUFV 7102); Pains (MCNAM 7434-7435); Palmital, UHEQueimados (MCNAM 2894, 4001-4004, 5851-5853, 8981); Paracatu (MCNAM 10593, CHUNB25365-25445, 25447-25483, 25485-25500, 26915); Paracatu/Pirapora, Linha de Transmissão(MCNAM 11012-11013); PARNA Serra do Cipó (MCNAM 2697-2698, 2732, 2762,



2763,2779,11767); Patrocínio, Mata da Roda d’Água (MCNAM 13820, 13842-13843); Peçanha,Lagoa da Serra Negra (MCNAM 1131); Periquito, Fazenda Bonaparte (MCNAM 2540);Pirapora (MNRJ 12493-12494); Pompéu/Curvelo, UHE Retiro Baixo (MCNAM 10385-10386, 10389, 10391, 10419); Rio Preto, PCH Mato Limpo (MCNAM 2304); RPPN Serrado Caraça (MCNAM 5796); Sacramento (CHUNB 49400); Santana do Riacho, Serra doCipó (MCNAM 1851-1854, 2458, 6955); Santa Rita de Jacutinga (MCNAM 2060); SãoGonçalo do Rio Abaixo (MCNAM 13725-13732); São Gonçalo do Rio Preto (MCNAM8563-8564); São João Nepomuceno (MNRJ 30488, 32185-32186, 49639-49642); SãoJosé do Goiabal (MCNAM 12843); São José do Mantimento/Durandé, PCH Varginha eVárzea Alegre (MCNAM 11218); Tocantins (MZUFV 7591-7592); Tombos (MZUFV 5320-5321); Três Pontas (MCNAM 7871); Urucânia (MCNAM 8364); Urucuia (MNRJ 734),Viçosa, UFV-Piscicultura (MZUFV 640, 8044); Viçosa, UFV-Mata do Paraíso (MZUFV2525, 2531, 6138); Viçosa, Sítio do Fiúza (MZUFV 5309). ESPÍRITO SANTO: Itaperuna(MNRJ 60299). RIO DE JANEIRO: Atafona (MNRJ 7978-6981); Campo Belo (AL-MN075, 405, 407, 814, 842); Itamonte, Planalto do Itatiaia (MNRJ 13978-13979); Itaperuna(MNRJ 54366); Lídice (MNRJ 66675); São João da Barra (MNRJ 6968-6970). SÃOPAULO: Angatuba (CFBH 23136); Botucatu (EI 4403); Botucatu, Fazenda Dinucci (MNRJ61116, MNRJ 66851); Botucatu, Fazenda Edgardia (MNRJ 66860); Botucatu, FazendaLageado (MNRJ 66858, 66866-66867, 66868-66869); Botucatu, Fazenda Monjolão(MNRJ 66869); Botucatu, Fazenda Santa Maria do Araquá (MNRJ 66862-66865);Campinas, Souzas (MNRJ 34698); Cubatão (MZUSP 37, 2023, paralectotypes); Cubatão,Raiz da Serra (MZUSP 2, two syntypes of Engystoma ovale lineata); Itirapina (CFBH4997); Paranapiacaba, Alto da Serra (MZUSP 715, paralectotype); Pedro de Toledo(MNRJ 7020); Perús (MZUSP 36, 2024-2027, paralectotypes); Piquete (MZUSP 529,lectotype; MZUSP 264); Rio Claro (CFBH 4132-4137, 4147-4148, 4209, 4230-4231,4233-4237, 4256-4257, 6575-6578); Rubião Júnior (MNRJ 49643, MNRJ 66854, 66855,66859); Santo André (EI 1398); São Manuel, Estação Experimental de São Manoel(MNRJ 66857); São Paulo, Belém (MZUSP 38, paralectotype); São Paulo, Ipiranga(MZUSP 33, 42, 264, paralectotypes; MZUSP 41, two syntypes of Engystoma ovaleconcolor); São Paulo (CHUNB 47543); São Simão (EI 1396-1397); Taubaté (EI 1234-1236); Ubatuba (CFBH 10907).

Elachistocleis erythrogaster – RIO GRANDE DO SUL: São Francisco de Paula, Centro dePesquisas e Conservação da Natureza Pró-Mata (MNRJ 39098, paratype, ex-MCP 3142).

Elachistocleis helianneae – BOLIVIA: BENI: Guayaramerim (USNM 123964). SANTACRUZ: Cercado (USNM 142132-142133). BRAZIL: AMAZONAS: Manaus (MNRJ 7796,MPEG 16103, MZUSP 58869-58870); Puruzinho (MZUSP 56900). PARÁ: Ananindeua(MPEG 3726-3728); Belém, Campus de Pesquisa do MPEG (MPEG 6353, 6875, 6929,6930-6931, 6934, 6939, 20566); Belém (MPEG 1767; MZUSP 1113, 36473). RONDÔNIA:Calama (MZUSP 56901); Foz do Jamari (MZUSP 56904); Porto Velho (MNRJ 3656,MZUSP 16622-16628, 16630-16631, 16633-16639, 16519-16562, 16564-16578,16642-16649, 16580-16619); São Carlos (MZUSP 56902-56903). MATO GROSSO: SantaMaria (MNRJ 2918). PARAGUAY: Assunción (MNRJ 6949) [in error].

Elachistocleis piauiensis – BRAZIL: PIAUÍ: Brejo do Piauí (MNRJ 42073); Castelo doPiauí (MPEG 19823); Picos, BR 316, Km 312 (MNRJ 66848, holotype ex-JJ 6024;MNRJ 14253, 60086, paratypes ex-JJ 6025-6026). MATO GROSSO: Cuiabá, Campus

30 U.CARAMASCHI


da UFMT (MHNCI 610); Cuiabá (MNRJ 4814).

Elachistocleis magnus – BRAZIL: RONDÔNIA: Costa Marques (CHUNB 29202-29203);Pimenteiras do Oeste (CHUNB 58882).

Elachistocleis matogrosso – BRAZIL: MATO GROSSO: Mato Grosso (MZUSP 52105).MATO GROSSO DO SUL: Aquidauana (MNRJ 2372); Corumbá, Fazenda Nhumirim(MNRJ 33045); Corumbá, Passo do Lontra, Base de Estudos do Pantanal (MNRJ 49530-49531); Corumbá (MCNAM 7404, 7634); Coxim (MZUSP 61040-61041); Miranda (MNRJ1858, 6964, 6965-6967, MZUSP 65107-65155); Nioaque (EI 4014); Salobra (MNRJ6950-6954); Taunay (MNRJ 6962-6963).

Elachistocleis surumu – BRAZIL: RORAIMA: (MPEG 7637, 7660, 7673, 7712, 7785,7801, 7849, 7850, 7858, 7895, 7926, 7927, 7929, 7936); Boa Vista, Fazenda BomIntento (MPEG 1265-1267, 1269-1272, 1292); Pacaraima, Vila Surumu (MNRJ 27316).

Elachistocleis sp. – BRAZIL: AMAZONAS: Humaitá (JJ 6671-6672, MNRJ 4821). GOIÁS:Alto Paraíso de Goiás, Estrada para o PARNA Chapada dos Veadeiros (MNRJ 27742);Chapada dos Veadeiros (MNRJ 722, 6924-6931); “Planalto de Goiás” (MNRJ 6948);Veadeiros (MNRJ 720). TOCANTINS: Porto Alegre do Tocantins (MNRJ 41397-41398).BOLIVIA: COCHABAMBA: Chapare (USNM 146599).

BOLETIM DO MUSEU NACIONAL, NOVA SÉRIE, ZOOLOGIA ISSN 0080-312X

Universidade Federal do Rio de JaneiroReitor – Aloísio Teixeira

Museu NacionalDiretora – Claudia Rodrigues Ferreira de Carvalho

Editores – Miguel Angel Monné Barrios, Ulisses Caramaschi

Editores de Área – Adriano Brilhante Kury, Ciro Alexandre Ávila, Claudia Petean Bove,Débora de Oliveira Pires, Guilherme Ramos da Silva Muricy, Izabel Cristina Alves Dias,João Alves de Oliveira, João Wagner de Alencar Castro, Marcela Laura Monné Freire,Marcelo de Araújo Carvalho, Marcos Raposo, Maria Dulce Barcellos Gaspar de Oliveira,Marília Lopes da Costa Facó Soares, Rita Scheel Ybert, Vânia Gonçalves Lourenço Esteves

Conselho Editorial – André Pierre Prous-Poirier (Universidade Federal de Minas Gerais),David G. Reid (The Natural History Museum - Reino Unido), David John Nicholas Hind(Royal Botanic Gardens - Reino Unido), Fábio Lang da Silveira (Universidade de SãoPaulo), François M. Catzeflis (Institut des Sciences de l’Évolution - França), GustavoGabriel Politis (Universidad Nacional del Centro - Argentina), John G. Maisey (AmericamMuseun of Natural History - EUA), Jorge Carlos Della Favera (Universidade do Estadodo Rio de Janeiro), J. Van Remsen (Louisiana State University - EUA), Maria Antonietada Conceição Rodrigues (Universidade do Estado do Rio de Janeiro), Maria CarlotaAmaral Paixão Rosa (Universidade Federal do Rio de Janeiro), Maria Helena PaivaHenriques (Universidade de Coimbra - Portugal), Maria Marta Cigliano (UniversidadNacional La Plata - Argentina), Miguel Trefaut Rodrigues (Universidade de São Paulo),Miriam Lemle (Universidade Federal do Rio de Janeiro), Paulo A. D. DeBlasis(Universidade de São Paulo), Philippe Taquet (Museum National d’Histoire Naturelle -França), Rosana Moreira da Rocha (Universidade Federal do Paraná), Suzanne K. Fish(University of Arizona - EUA), W. Ronald Heyer (Smithsonian Institution - EUA)

Normalização – Edson Vargas da Silva

Diagramação e arte-final – Lia Ribeiro

Indexação – Biological Abstracts, ISI – Thomson Scientific, Ulrich’s InternationalPeriodicals Directory, Zoological Record, NISC Colorado, Periodica, C.A.B. International

Tiragem – 600 exemplares

Disponível em: <http://www.publicacao.museunacional.ufrj.br>

Biblioteca/MN, e-mail: [email protected]

MUSEU NACIONALUniversidade Federal do Rio de Janeiro

Quinta da Boa Vista, São Cristóvão20940-040 – Rio de Janeiro, RJ, Brasil

Impressão:SERMOGRAF Artes Gráficas e Editora Ltda

Petrópolis, RJ

elachistocleis bol+mus+nac+zool chi 2010

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