Bol Zool ULISSES.pmd

Transcrição

Bol Zool ULISSES.pmd
BOLETIM DO MUSEU NACIONAL
NOVA SÉRIE
RIO DE JANEIRO - BRASIL
ISSN 0080-312X
ZOOLOGIA
N 527
o
12 DE AGOSTO DE 2010
NOTES ON THE TAXONOMIC STATUS OF
ELACHISTOCLEIS 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 the
taxon is considered a nomen dubium, referred to a species inquirenda. Five new species of the genus
Elachistocleis are described. Elachistocleis helianneae sp.nov., from Humaitá (07o35’S, 62o40’W; 90m
altitude), State of Amazonas, Brazil, is characterized by small size (SVL 22.6-28.7mm in males, 29.336.4mm in females), dorsum grayish brown with minute scattered light gray spots and a distinctive
middle longitudinal light cream stripe from the tip of snout to vent, and venter immaculate clear
cream, 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), dorsum
dark gray with small irregular clear gray spots scattered without forming defined pattern, mid dorsal
longitudinal 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 large
anastomosed whitish spots, producing a coarse marbled pattern, mainly on the chest. Elachistocleis
bumbameuboi sp.nov., from UHE Ponta da Madeira, Municipality of São Luís (02o32’S, 44o18’W; 24m
altitude), 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, extending
to the ventrolateral region. Elachistocleis matogrosso sp.nov., from Cuiabá (15o36’S, 56o06’W; 177m
altitude), 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 cream
stripe from the post-cephalic dermal fold to vent, and venter immaculate clear cream, with a poorly
defined color limit between the dorsal and ventral regions. The geographic distributions of the new
species and of Elachistocleis bicolor and E. cesarii are realized and mapped.
1
2
Received on 17 de junho de 2010. Accepted on 23 de junho de 2010.
Museu Nacional/UFRJ, Departamento de Vertebrados. Quinta da Boa Vista, São Cristóvão, 20940040 Rio de Janeiro, RJ, Brasil. E-mail: [email protected].
Fellow of Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq).
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U.CARAMASCHI
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 descrição
de cinco espécies novas de Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae).
A posição taxonômica de Elachistocleis ovalis (Schneider, 1799) é discutida e o táxon considerado
um nomen dubium, relacionado a uma species inquirenda. Cinco novas espécies do gênero
Elachistocleis são descritas. Elachistocleis helianneae sp.nov., de Humaitá (07o35’S, 62o40’W;
90m de altitude), Estado do Amazonas, Brasil, é diagnosticada pelo tamanho pequeno (CRA
22,6-28,7mm em machos, 29,3-36,4mm em fêmeas), dorso marrom acinzentado com pequenas
manchas cinza claro espalhadas e uma característica linha longitudinal mediana creme claro
desde a ponta do focinho ao ânus e ventre creme claro imaculado, com nítido limite de colorido
entre 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 tamanho
pequeno (CRA 19,6-27,0mm em machos, 23,1-26,9mm em fêmeas), dorso cinza escuro com
pequenas manchas cinza claro espalhadas sem formar padrão definido, linha longitudinal dorsal
clara 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 do
Tocantins, Brasil, é separada pelo tamanho médio (CRA 24,9-31,9mm em machos, 32,0-37,1mm
em fêmeas), dorso uniformemente cinza escuro sem desenhos ou padrão, e ventre e flancos cinza
com grandes manchas esbranquiçadas anastomosadas, produzindo um padrão marmoreado
grosseiro, principalmente na região peitoral. Elachistocleis bumbameuboi sp.nov., da UHE Ponta
da 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 em
fêmeas), dorso uniformemente cinza escuro, sem manchas nem linha clara mediana, ventre cinza,
com pequenas manchas esbranquiçadas anastomosadas, produzindo um padrão de sal-e-pimenta
que 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 marrom
acinzentado, com uma linha creme claro longitudinal mediana desde a prega dérmica pós-cefálica
até o ânus, e ventre creme claro imaculado, com limite de colorido pouco definido entre as regiões
dorsal e ventral. As distribuições geográficas das novas espécies e de Elachistocleis bicolor e
Elachistocleis 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 other
with some kind of blotches or vermiculations on belly, including E. cesarii (MirandaRibeiro, 1920), E. erythrogaster Kwet & Di Bernardo, 1998, E. magnus Toledo, 2010, E.
piauiensis Caramaschi & Jim, 1983, E. skotogaster Lavilla, Vaira & Ferrari, 2003, and
E. 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 and
have 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 in
males, 21.1-28.3mm in females) and venter mottled in cream and gray, a thin and
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NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
3
interrupted clear line on the posterior surface of thighs, and a large gland behind the
posterior corner of mouth (CARAMASCHI & JIM, 1983); E. erythrogaster, described from
São Francisco de Paula, State of Rio Grande do Sul, Brazil, presents size large (SVL
29.1-32.3mm in males, 34.0-37.6mm in females) and deep black throats in males
and females, black and blue marbled lateral surfaces, flashy red-orange venter, and
no 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 mottled
with 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, type
locality Piquete, State of São Paulo, Brazil, with medium size (SVL 22.6-26.7mm in
males, 28.6-36.0mm in females) and dorsum and limbs brownish gray with small
white dots, throat darker than venter, chest yellow with gray marks, belly white or
yellow with gray marks and reticulations reaching the flanks, orange femoral stripe,
and small postcommisural gland present (TOLEDO et al., 2010), and the recently
described 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, 39.8-43.8mm
in females) and dorsum and limbs uniform dark grayish with scarce minute brighter
dots on the outer boundaries of the dorsum, a thin mid-dorsal white stripe from the
vent to the anterior third of the dorsum, throat brownish dark, darker than chest
and belly, venter gray with minute scattered white spots mainly on the belly and
ventral surface of legs, large irregular white spots on the groin and axillary region, a
broad, 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 have
name-bearing types nor defined type localities and LAVILLA et al. (2003) showed that
two of them, E. ovalis and E. bicolor, are involved in a enormous confusion since
1841 and, in fact, later contributions not only did not solve the problem but contributed
to increase the controversy. In its turn, E. surinamensis has been treated as a junior
synonym of E. ovalis, as a senior synonym of Hypopachus pearsei Ruthven, 1914
(currently Relictivomer pearsei), or as valid species. Attempting to throw some light
on the problem, LAVILLA et al. (2003) proposed that it would be advisable to consider
the latter three species to fit with the characters that describe the genus Elachistocleis
and that E. surinamensis would have a spotted ventral color, an evident light vertebral
stripe, and would inhabit the northern portion of the generic range; E. bicolor would
have an immaculate venter and would occupy the southern portion of the generic
range (Argentina, Bolivia, Paraguay, Uruguay, and southern Brazil), restricting the
type locality of the species to Buenos Aires, Argentina; and E. ovalis would have an
immaculate, yellow ventral color and would occur in the northern portion of the
generic range. Nevertheless, in this concept E. ovalis is not associated with any existing
natural population. Additionally, LAVILLA et al. (2003) stated that these three taxa
constitute, without doubt, complexes of species and the decisions presented respecting
the definitions would be only operative frameworks for a necessary revision.
Pending that extensive revision and in order to improve the taxonomy of the genus
Elachistocleis, in this paper the taxonomic status of Elachistocleis ovalis (Schneider,
1799) is discussed and five new species of Elachistocleis are described from Brazil.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
4
U.CARAMASCHI
MATERIAL AND METHODS
Specimens examined, referred in the text and in the Appendix, are housed in the
following collections: MNRJ (Museu Nacional, Rio de Janeiro, RJ, Brazil); MPEG (Museu
Paraense Emílio Goeldi, Belém, PA, Brazil); MZUSP (Museu de Zoologia, Universidade
de 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, Universidade
Federal de Viçosa, MG, Brazil); MCNAM (Museu de Ciências Naturais, Pontifícia
Universidade Católica de Minas Gerais, Belo Horizonte, MG, Brazil); USNM (National
Museum of Natural History, Smithsonian Institution, Washington, D.C., USA); CHUNB
(Coleção Herpetológica da Universidade de Brasília, DF); CFBH (Célio F.B. Haddad
Collection, 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 a
dissecting 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 (foot
length). 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 with
small head, long snout, globose body, and small eyes, with gray dorsum and yellowish
venter. The syntypes were originally one specimen in the “Musei Ducalis Brunovicensis”,
a second specimen in the “Museo Barbyensi”, and one specimen in the “Gronovius
Musei 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, the
only figured specimen in S EBA (1735) is a Breviceps gibbosus (Linnaeus, 1758)
(Brevicipitidae), a species known from Republic of South Africa (FROST, 2010). The type
locality was not stated and it is impossible to infer where the syntypes were collected
on the basis of cited collections and figures. Besides, no one of the earlier and
contemporary followers of Schneider (e.g., SHAW, 1802; DAUDIN, 1802; MERREM, 1820)
stated the locality for the species; moreover, FITZINGER (1826) considered that its
distribution would be “...Ex Asia, India...” [“E. ovalis. m. Eiförmige E. (Rana ovalis.
Schneider) Ex Asia, India.”]. CUVIER (1829) associated Engystoma ovalis with the genus
Dactylethra ¯ “l’Engystoma ovalis Fitz., est un dactylètre” ¯, a genus created by him for
species from Southern Africa (“le midi de l’Afrique ...”) and currently synonymized
under 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
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NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
5
gave, as a synonym of his Engystoma ovale, Rana ovalis Schneider; however, they
conceded 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 with
all names purportedly associated with that species: Rana ovalis Schneider, Rana ovalis
Shaw, Bufo surinamensis Daudin, Bufo ovalis Daudin, Rana bufonia Merrem, Engystoma
ovalis Fitzinger, Oxyrhincus bicolor Valenciennes, Oxyrhincus bicolor Guérin, Microps
unicolor Wagler, and Stenocephalus microps Tschudi. In doing that, they included under
the same concept the two known color morphs, with spotted and immaculte venters.
Additionally, overlooking the statement of FITZINGER (1841) on the distribution of the
species, they considered that Engystoma ovale occurred in South America (“l’Amerique
méridionale”), having examined putative specimens from Surinam and from Buenos
Aires [Argentina]. This distributional concept was followed, without discussion, by all
subsequent authors (LAVILLA et al., 2003). Attempting to present an operative framework
for 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 genus
Elachistocleis, have an immaculate, yellow ventral coloration, and inhabit the northern
portion of the generic range.” Concluding, they conceded that regarding this distribution
and if the statement of FITZINGER (1826) is in error and Elachistocleis is really a neotropical
taxon, the decision presented was based on the exclusion of those specimens considered
under the name Elachistocleis bicolor (Guérin-Méneville, 1838).
The proposition of LAVILLA et al. (2003), however, presents a central problem given that
the name E. ovalis was not associated to any known frog population in South America
and, if FITZINGER (1826) is correct and DUMÉRIL & BIBRON (1841) are wrong, this name is
not applicable to a neotropical taxon. Moreover, although biologically indefinite,
taxonomically the name E. ovalis threatens the later congener E. bicolor, a currently
well established species. In fact, based on the immaculate, yellow ventral color described
for both taxa, CARCERELLI (1992), in a meeting abstract, proposed the synonymization of
E. 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, the
inexistence of a name bearing type or types, the absence of a type locality, the
impossibility to associate the name with an actual population, and to prevent the
threat to a well known species, Rana ovalis Schneider, 1799 and its current combination
Elachistocleis ovalis is here considered a nomen dubium, that is, a name of unknown or
doubtful application (ICZN, 1999), associated to a species inquirenda, that is, a species
of 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), MNRJ
6989, adult , collected by Ulisses Caramaschi, 13/XII/1979.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
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U.CARAMASCHI
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 6687066871, 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 by
C.M.Carvalho, A.Silva and L.M.Silva, 13/I/1979; MNRJ 66888-66889, collected by
L.M.Silva, A.Silva and C.M.Carvalho, 24/I/1979; MNRJ 66890-66892, collected by
M.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 6829668297, 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% of
HW (x¯ = 94.6; SD = 8.03; n = 33); postcommisural gland poorly developed; dorsum
smooth; in preservative grayish brown with minute scattered light gray spots; a
distinctive middle longitudinal light cream stripe, from the tip of snout to vent; venter
immaculate cream; a sharp color limit between the dorsal and ventral regions; no
spots on axillae or groin; a broad irregular cream line on the posterior surface of the
thighs; a large light cream spot on the proximal internal surface of tibiae; a narrow
light 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 distinguished
from 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 in
E. 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 separated
by the mid-dorsal stripe extending from the tip of snout to vent (extending from the postcephalic dermal fold to vent in E. matogrosso sp.nov.), by the presence of minute light
spots on dorsum and dorsal surfaces of members (absent in E. matogrosso sp.nov.), by the
loreal region conspicuously white in E. helianneae (dorsal gray color of dorsum of the
snout invading the loreal region almost to the upper lip border in E. matogrosso), and by
the sharp color limit between the dorsal and ventral regions (color limit between dorsal
and 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 subelliptical in dorsal view, protruding in profile. Nostrils small, not protuberant, directed
anterolaterally, closer to tip of snout than to eye; internarial distance smaller than the eye to
nostril and interorbital distances, equalling the eye diameter, and larger than the upper
eyelid width. Canthus rostralis rounded; loreal region flat, sloping abruptly to the upper lip;
lips not flared. Postcommisural gland poorly developed. Eyes small, dorsolateral, only slightly
protruding. Interorbital space slightly convex, almost two times the upper eyelid width. No
cranial crests. A poorly defined transverse skinfold across back of the head, bending
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
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downwards slightly behind the eyes to the shoulder; a poorly defined skinfold through
dorsolateral region from the axilla to the groin. Tympanum concealed; supratympanic fold
absent. Lower jaw with truncate, trilobed anterior margin. Tongue large, oval, without a
notch on posterior border. Choanae large, subcircular, widely separated. Vocal slits present.
Premaxillary, maxillary, and vomerine odontophores absent. Vocal sac subgular, not expanded
externally. 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, with
subarticular tubercles developed, rounded; supranumerary tubercles absent; tip of
fingers 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 ventral
views of body, and head profile.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
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U.CARAMASCHI
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% of
SVL; tibia length 38.2% of SVL. Heel of adpressed legs not reaching axilla; knee and
elbow widely separated when limbs laid along the sides; heels touching when flexed
legs held at right angle to body; knee and heel with a transverse skinfold; no tibial or
tarsal 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 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 and dorsal surfaces of limbs grayish brown, with minute
white dots scattered without forming a defined pattern; a distinctive mid-dorsal
light cream stripe, from the tip of snout to the urostile; a sharp color limit between
the dorsal and ventral regions; venter immaculate, dull white; throat grayish;
no spots on axillae and groin; an irregular broad light cream stripe on the
posterior surface of thighs; a large irregular, light cream spot on the proximal
internal surface of tibiae; a light cream stripe surrounding the knees and reaching the
middle of the tarsus.
Measurements of holotype in mm – SVL 26.5, HL 6.4, HW 6.6, IND 1.7, END 2.3, ED
1.7, 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 personal
life, besides the companionship in the home, in the field, and in the laboratory
work.
Variation – Except for minor details in corporal proportions, the descripton of the holotype
stands for males of the species. Size ranges from 23.0 to 29.5mm SVL and the intensity
of the color of the throat can vary. The females are bigger than males, size ranging from
26.3 to 37.0mm SVL. Range, mean, and standard deviation of the measurements of the
type specimens of E. helianneae are presented in table 1. In few specimens the middorsal stripe can be interrupted, but it is always present.
Geographical distribution – Northern Brazil, in the States of Amazonas, Pará, and
Rondonia, 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 dorsal
surfaces of limbs gray to dark gray, with small, scattered, irregular black spots;
longitudinal mid-dorsal stripe yellow; venter uniformly yellow to greenish yellow; a
mid-ventral white line from the chest to vent, crossed by a transversal white line
between the axillae; throat gray; stripe on posterior surface of thighs and spots on
internal surfaces of tibiae orange red; line surrounding knees and tarsus yellow; iris
brown with intense black vermiculations.
The specimen MNRJ 6949 fits the diagnosis for E. helianneae sp.nov., but has a
manuscript label containing “Engystoma ovale bicolor (Val.) / Assumpção - R. do
Paraguay / Coll. B. Schouten 1929. / Miranda-Ribeiro [signed].” The collection data
are here considered in error, due to an early labeling mistake or possible specimen
exchange in some time of these almost 90 years in the collection.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
9
TABLE 1. Range (mm), mean (x¯ ), and standard deviation (SD) of the measurements of the type
specimens of Elachistocleis helianneae sp.nov. (n = number of specimens).
C HARACTERS
R ANGE
SVL
HL
HW
IND
END
ED
UEW
IOD
HAL
THL
TL
FL
23.0-29.5
5.65-6.8
5.7-7.5
1.2-2.15
2.1-2.65
1.35-2.0
1.0-1.3
2.1-3.5
4.6-6.5
8.0-10.7
8.4-11.8
10.5-13.9
(n = 15)
x̄
SD
R ANGE
26.4
6.1
6.7
1.7
2.3
1.6
1.2
2.85
5.4
9.8
10.2
12.0
1.93
0.37
0.37
0.24
0.16
0.20
0.09
0.39
0.47
0.80
0.82
0.83
26.3-37.0
6.1-7.9
5.55-8.4
1.7-2.2
2.0-3.0
1.6-2.1
1.2-1.6
2.6-3.6
5.4-7.2
9.2-12.7
10.2-12.9
11.8-15.8
(n = 18)
x̄
31.7
7.0
7.3
2.0
2.5
1.7
1.3
3.1
6.3
11.2
11.8
13.9
SD
2.82
0.61
0.70
0.18
0.29
0.16
0.12
0.27
0.52
1.09
0.92
1.21
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, 2530725326, 25338-25344.
Diagnosis – A small sized species (SVL 19.6-27.0mm in males, 23.2-26.9mm in
females), diagnosed by the head length shorter than the head width, HL about
85.6% of HW (x¯ = 85.6; SD = 4.27; n = 24); postcommisural gland present,
small; dorsum smooth; in preservative dark gray with small irregular clear gray
spots scattered without forming defined pattern, and mid longitudinal light stripe
absent; venter gray with many irregular cream spots regularly distributed,
including the throat area; an undefined color transition between the dorsal and
ventral 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 knee
and tarsus.
Comparisons with other species – The species with some kind of ventral color pattern
are 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 distinguished
from 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 marbled
with black, blue, and some orange in males, dorsum orange marbled with black and
blue in females of E. erythrogaster; dorsum uniform dark grayish with scarce minute
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
10
U.CARAMASCHI
brighter dots in the outer boundaries in E. magnus; dorsum uniformly gray with scarce
minute irregular bright dots in E. piauiensis; dorsum dark brown mottled with black
and white spots in E. skotogaster; dorsum dark gray or black with an evident light
vertebral stripe in E. surinamensis; dorsum uniformly brown or dark gray without
marks in E. carvalhoi sp.nov. and E. bumbameuboi sp.nov.; 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 and
throat 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 reticulations
in E. cesarii; venter red orange, with hindlimbs largely black mottled in bright blue in
E. erythrogaster; venter gray with minute scattered white spots, mainly on the 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 in
E. skotogaster; venter dark gray or black with large yellow and small white spots in
E. 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-pepper
pattern, in E. bumbameuboi sp.nov.).
The three species with immaculate venters and a defined color limit between the dorsal
and ventral regions are E. bicolor, E. helianneae, and E. matogrosso sp.nov. The new
species is readily distinguished from these species by having venter gray with many
irregular cream spots regularly distributed and an undefined color transition between
the dorsal and ventral regions.
Description of holotype – Body ovoid, head small, triangular, broader than long; head
length 89.7% of head width and 25.2% of SVL; head width 28.1% 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 than the eye to nostril
and 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 not
flared. Postcommisural gland present, small. Eyes small, dorsolateral, only slightly
protruding. Interorbital space slightly convex, almost two times the upper eyelid width.
No cranial crests. A transverse skinfold across back of the head, bending downwards
slightly behind the eyes to the shoulder; a poorly defined skinfold through dorsolateral
region from the axilla to the groin. Tympanum concealed; supratympanic fold absent.
Lower jaw with truncate, trilobed anterior margin. Tongue large, oval, without a notch
on posterior border. Choanae large, subcircular, widely separated. Vocal slits present.
Premaxillary, maxillary, and vomerine odontophores absent. Vocal sac subgular, not
expanded 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, with
subarticular tubercles developed, rounded; supranumerary tubercles absent; tip of
fingers 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; tibia
length 93.3% of thigh length. Sum of thigh and tibia lengths 80.8% of SVL; thigh
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
11
Fig.2- Elachistocleis surumu sp.nov., holotype (MNRJ 25210; SVL 25.4mm), dorsal and ventral
views of body, and head profile.
length 41.8% of SVL; tibia length 39.0% of SVL. Heel of adpressed legs failing to reach
axilla; knee and elbow widely separated when limbs laid along the sides; heels touching
when flexed legs held at right angle to body; knee and heel with a weak transversal
skinfold; no tibial or tarsal ridges; a small oval inner but no outer metatarsal tubercle;
plantar tubercles absent. Toes slender, free, not 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 smooth
above and beneath; anal opening not modified, no para-anal tubercles.
In preservative, dorsum dark gray with small irregular gray spots scattered without
forming defined pattern, and mid-longitudinal light stripe absent; venter gray with
many irregular cream spots regularly distributed; an undefined color transition between
the dorsal and ventral regions; small, defined white spots on axilla and groin; a short
irregular, poorly defined cream line on the posterior surface of the thighs; internal
sufaces of tibiae with white spots; no light stripe on knee and tarsus.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
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, ED
1.6, 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 type
locality, a small village currently included in the Reserva Indígena Raposa Serra do
Sol, in the State of Roraima, northern Brazil.
Variation – Except for minor details in corporal proportions, the descripton of the holotype
stands for males of the species. Size ranges from 19.6 to 27.0mm SVL and the intensity
of the color of the throat can vary. The females are slightly bigger than males, size
ranging from 23.2 to 26.9mm SVL. Range, mean, and standard deviation of the
measurements 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 specimen
not specified, presents dorsum and flanks dark gray with small irregular light gray
spots; venter, throat, and undersurfaces of limbs black with many irregular yellow
spots; spots on axilla, groin, posterior surface of tibia, and thigh stripe, orange red;
iris brown with intense black vermiculations.
TABLE 2. Range (mm), mean (x¯), and standard deviation (SD) of the measurements of the type
specimens of Elachistocleis surumu sp.nov. (n = number of specimens).
CHARACTERS
SVL
HL
HW
IND
END
ED
UEW
IOD
HAL
THL
TL
FL
(n = 20)
x̄
RANGE
19.6-27.0
4.7-5.9
5.1-7.2
1.4-2.2
1.7-2.2
1.3-1.7
0.9-1.4
2.1-2.8
4.6-5.7
7.3-10.6
7.2-9.9
8.3-11.4
23.1
5.2
6.15
1.7
2.0
1.5
1.2
2.5
5.2
8.65
8.3
9.6
SD
RAN GE
1.94
0.44
0.51
0.17
0.14
0.13
0.11
0.17
0.40
0.88
0.69
1.06
23.2-26.9
5.3-7.2
5.9-7.0
1.5-1.7
1.9-2.4
1.4-1.6
1.0-1.3
2.4-2.9
5.05-5.6
9.1-11.1
8.1-9.35
9.5-13.6
(n = 4)
x̄
25.6
5.9
6.65
1.55
2.2
1.5
1.3
2.7
5.3
9.4
8.9
11.2
SD
1.61
0.39
0.49
0.09
0.23
0.12
0.11
0.21
0.27
1.19
0.52
1.76
Elachistocleis carvalhoi sp.nov.
(Fig.3)
Holotype – BRAZIL: TOCANTINS: Aragominas (07o10’S, 48o32’W; 345m altitude), MNRJ
51384 , adult , collected by P.Fatorelli and L.Machado, 25/XI/2007.
Paratypes – BRAZIL: TOCANTINS: Collected with the holotype, MNRJ 51385; Nova
Olinda (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, collected
by E.G.Pereira and P.C.F.Carneiro, VI/2007. PARÁ: Canaã dos Carajás (06o30’S,
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
13
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 by
P.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 in
females), characterized by the head length shorter than the head width, HL about 86.6%
of HW (x¯ = 86.6; SD = 6.38; n = 9); postcommisural gland developed; dorsum smooth; in
preservative uniformly dark gray without marks or pattern; venter and flanks grayish
with large anastomosed whitish spots, producing a coarse marbled pattern, mainly on
the chest; large cream spots on axillae, groin, and posterior surfaces of tibiae; a broad
irregular, poorly defined line on the posterior surface of the thighs; no light stripe on
knee and tarsus.
Comparisons with other species – The species with some kind of ventral color pattern
are 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 these
species by the dorsum uniformly brown or dark gray without marks (dorsal surfaces
dark 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 females
of E. erythrogaster; dorsum uniform dark grayish with scarce minute brighter dots in
the outer boundaries in E. magnus; dorsum uniformly gray with scarce minute irregular
bright dots in E. piauiensis; dorsum dark brown mottled with black and white spots in
E. 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.; 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 grayish with large anastomosed yellow or whitish yellow blotches, producing
a coarse marbled pattern, mainly in the chest area (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 in bright blue in E. erythrogaster; venter gray
with minute scattered white spots, mainly on the 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 in E. skotogaster; venter dark gray or
black with large yellow and small white spots in E. surinamensis; venter and throat
gray with many irregular yellow spots regularly distributed in E. surumu; and venter
gray with minute anastomosed whitish spots, producing a salt-and-pepper pattern, in
E. bumbameuboi sp.nov.).
The three species with immaculate venter and a sharp color limit between the dorsal
and ventral regions are E. bicolor, E. helianneae, and E. matogrosso sp.nov. The new
species is readily distinguished from these species by having venter grayish with large
anastomosed yellow or whitish yellow blotches, producing a coarse marbled pattern,
mainly in the chest area, and an undefined color transition between the dorsal and
ventral regions.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
14
U.CARAMASCHI
Description of holotype – Body ovoid, head small, triangular, slightly broader than
long; 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 than
the eye to nostril and 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 not flared. Postcommisural gland developed. Eyes small,
dorsolateral, only slightly protruding. Interorbital space slightly convex, 2.3 times
the upper eyelid width. No cranial crests. A poorly defined transverse skinfold across
back of the head, bending downwards slightly behind the eyes to the shoulder; a
skinfold through dorsolateral region from the axilla to the groin. Tympanum concealed;
supratympanic fold absent. Lower jaw with truncate, trilobed anterior margin. Tongue
large, oval, without a notch on posterior border. Choanae large, subcircular, widely
separated. Vocal slits present. Premaxillary, maxillary, and vomerine odontophores
absent. Vocal sac subgular, not expanded externally. A weak skinfold crossing the
chest between axillae.
Fig.3- Elachistocleis carvalhoi sp.nov., holotype (MNRJ 51384; SVL 31.4mm), dorsal and ventral
views of body, and head profile.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
15
TABLE 3. Range (mm), mean (x¯), and standard deviation (SD) of the measurements of the type
specimens of Elachistocleis carvalhoi sp.nov. (n = number of specimens).
C HARAC TERS
SVL
HL
HW
IND
END
ED
UEW
IOD
HAL
THL
TL
FL
RANGE
24.0-31.9
5.8-7.85
6.3-8.1
1.6-2.2
2.1-2.6
1.4-1.9
1.2-1.5
2.85-3.4
5.5-6.9
9.8-11.8
10.0-12.1
11.45-15.0
(n = 6)
x̄
28.95
6.6
7.45
1.9
2.4
1.6
1.4
3.1
6.35
10.7
10.95
13.2
SD
RANGE
3.22
0.75
0.67
0.22
0.20
0.19
0.10
0.23
0.59
0.86
0.92
1.56
32.0-37.1
6.7-7.2
7.6-8.9
1.9-2.5
2.5-3.4
1.7-2.0
1.2-1.7
2.7-3.9
6.4-8.8
12.9-14.4
11.9-14.1
14.1-16.9
(n = 3)
x̄
35.2
7.0
8.4
2.2
3.0
1.8
1.5
3.5
7.8
13.8
13.2
15.9
SD
2.96
0.25
0.70
0.28
0.49
0.18
0.27
0.65
1.27
0.82
1.13
1.52
Arms moderately robust, no tubercles or crests on forearm; palmar tubercle large,
divided longitudinally, twice as large as the thenar tubercle; fingers slender, free, with
subarticular tubercles developed, rounded; supranumerary tubercles absent; tip of
fingers 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 than
foot 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 legs
failing to reach axilla; knee and elbow widely separated when limbs laid along the
sides; heels touching when flexed legs held at right angle to body; knee and heel with
a weak transversal skinfold; no tibial or tarsal ridges; an small oval inner but no
outer metatarsal tubercle; plantar tubercles absent. Toes slender, free, not 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 smooth above and beneath; anal opening not modified, no paraanal 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 marbled
pattern, mainly on the chest; ventrolateral region with large whitish blotches; throat
grayish; large light cream spots on axillae, groin, and posterior surfaces of tibiae; a
light cream, broad, irregular stripe on posterior surfaces of thighs; no light longitudinal
stripe 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, ED
1.9,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, especially
the microhylid frogs.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
16
U.CARAMASCHI
Variation – Except for minor details in corporal proportions, the description of the
holotype stands for the male specimens examined. Size ranges from 24.0-31.9mm
SVL in males. The females are bigger than males, their size ranging from 32.0-37.1mm
SVL. Few specimens have the marbled pattern on venter and the spots on concealed
surfaces slightly coarser or thinner than the holotype.
Remarks – In specimens recently preserved it is possible to observe that the color in
life is dark gray to black on dorsum; the spots on venter are pale yellow on gray
background; the blotches on axillae, groin, and posterior surfaces of tibiae, and the
stripe on posterior surfaces of thighs are red orange.
Geographical distribution – Northern Brazil, in northwestern State of Tocantins and
southeastern State of Pará (Fig.6).
Elachistocleis bumbameuboi sp.nov.
(Fig.4)
Holotype – BRAZIL: MARANHÃO: São Luís (02o32’S, 44o18’W; 24m altitude), UHE Ponta da
Madeira, 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, MNRJ
53378, 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 in
females), 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 slightly
rugose; in preservative, dorsum uniformly dark gray, without spots nor light middorsal stripe; venter gray, with minute anastomosed whitish blotches, producing a
salt-and-pepper pattern, 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, irregular stripe on posterior surfaces of thighs; no light longitudinal stripe
on superior surfaces of tibiae and posterior surfaces of tarsus.
Comparisons with other species – The species with some kind of ventral color pattern are
E. 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 the
dorsum uniformly dark gray or black without marks (dorsum uniformly brown or dark
gray 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, dorsum
orange marbled with black and blue in females of E. erythrogaster; dorsum uniform dark
grayish with scarce minute brighter dots in the outer boundaries in E. magnus; dorsum
uniformly gray with scarce minute irregular bright dots in E. piauiensis; dorsum dark
brown mottled with black and white spots in E. skotogaster; dorsum dark gray or black
with an evident light vertebral stripe in E. surinamensis; and dorsum dark gray with small
irregular 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 whitish
spots, producing a salt-and-pepper pattern (venter grayish with large anastomosed yellow
or whitish yellow blotches, producing a coarse marbled pattern, mainly in the chest area
in E. carvalhoi; chest yellow with gray marks and belly white or yellow with gray marks
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
17
and reticulations in E. cesarii; venter red orange, with hindlimbs largely black mottled in
bright blue in E. erythrogaster; venter gray with minute scattered white spots, mainly on
the 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 in
E. 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 regularly
distributed in E. surumu).
The three species with immaculate venter and a sharp color limit between the dorsal
and ventral regions are E. bicolor, E. helianneae, and E. matogrosso. The new species is
readily distinguished from these species by having venter gray with minute anastomosed
whitish spots, producing a salt-and-pepper pattern and an undefined color transition
between the dorsal and ventral regions.
Fig.4- Elachistocleis bumbameuboi sp.nov., holotype (MNRJ 53200; SVL 26.9mm), dorsal and
ventral views of body, and head profile.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
18
U.CARAMASCHI
Description of holotype – Body ovoid, head small, triangular, broader than long; head
length 80.0% of head width and 21.2% of SVL; head width 26.4% 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 than the eye to nostril and
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 not flared.
Postcommisural gland developed. Eyes small, dorsolateral, only slightly protruding.
Interorbital space slightly convex, almost two and half times the upper eyelid width. No
cranial crests. A well defined transversal skinfold across back of the head, bending
down and backwards slightly behind the eyes to the shoulder; a strong dorsolateral skin
fold from the scapular region to the groin. Tympanum concealed; supratympanic fold
absent. Lower jaw with truncate, trilobed anterior margin. Tongue large, oval, without a
notch on posterior border. Choanae large, subcircular, widely separated. Vocal slits
present. 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, with
subarticular tubercles developed, rounded; supranumerary tubercles absent; tip of
fingers 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; thigh
length 37.8% of SVL; tibia length 37.6% of SVL. Heel of adpressed legs failing to reach
axilla; knee and elbow widely separated when limbs laid along the sides; heels touching
when flexed legs are held at right angle to body; knee and heel with a weak transversal
skinfold; no tibial or tarsal ridges; an small oval inner but no outer metatarsal tubercle;
plantar tubercles absent. Toes slender, free, not 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 smooth
above and beneath; small dermal spines scattered mainly on dorsolateral region; anal
opening 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-pepper
pattern, 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, irregular
stripe on posterior surfaces of thighs; no light longitudinal stripe on superior surfaces
of 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; ED
1.4, 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 most
popular feasts occurring in June at São Luís, Maranhão, the “Bumba-meu-boi”, which
can be “played” through several ways, with distinct characteristics of rhythms, clothes,
choreographies, characters, and musical instruments. These festivities narrate the
adventures 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.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
19
Variation – Except for minor details in corporal proportions, the description of the
holotype stands for the male specimens examined. Size ranges from 26.9-27.4mm
SVL in males. The females are bigger than males, their size ranging from 32.8-43.7mm
SVL. 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, treated
as 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-andpepper pattern, extending to the ventrolateral region; spots on groin and line on posterior
surfaces of thighs orange red; iris brown with dense black vermiculations. The
advertisement call is described and, in a note added in the proof of the paper, the
authors pointed out differences between the call they described and that described by
TOLEDO et al. (2010) for E. piauiensis based on recordings made at Pacatuba, Caucaia,
and Viçosa do Ceará, State of Ceará, Brazil.
TABLE 4. Range (mm), mean ( x¯), and standard deviation (SD) of the measurements of the type
specimens of Elachistocleis bumbameuboi sp.nov. (n = number of specimens).
(n = 3)
C HARACTERS
SVL
HL
HW
IND
END
ED
UEW
IOD
HAL
THL
TL
FL
(n = 3)
RANGE
x̄
SD
RANGE
x̄
SD
26.9-28.8
5.7-6.0
7.1-8.1
1.6-1.85
2.1-2.4
1.4-1.7
1.1-1.4
2.3-3.0
5.8-6.9
10.2-11.7
10.1-11.1
11.9-12.8
27.8
5.9
7.5
1.7
2.25
1.6
1.3
2.6
6.4
11.1
10.6
12.4
0.96
0.19
0.54
0.11
0.15
0.15
0.19
0.37
0.54
0.79
0.51
0.50
32.8-43.7
6.1-8.3
8.0-9.8
2.1-2.5
2.6-3.1
1.7-2.0
1.4-1.6
2.9-3.6
7.1-9.0
12.25-14.5
11.95-14.3
13.3-16.7
37.4
7.2
8.6
2.25
2.8
1.8
1.5
3.3
7.7
13.2
12.9
14.7
5.65
1.07
1.06
0.19
0.22
0.17
0.12
0.39
1.07
1.17
1.23
1.80
Elachistocleis matogrosso sp.nov.
(Fig.5)
Holotype – BRAZIL: MATO GROSSO: Cuiabá (15o36’S, 56o06’W; 177m altitude), MNRJ
4812, 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 by
J.Langone, 04/II/1986; CHUNB 47526, collected by LPP, 04/II/1986; Cuiabá, bank
of 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,
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
20
U.CARAMASCHI
55 o58’W; 156m altitude), RPPN Sesc Pantanal, MNRJ 32880-32882, collected by
C.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
(x¯ = 92.0; SD = 5.49; n = 8); postcommisural gland poorly developed; dorsum smooth;
in preservative, uniformly grayish brown; a thin middle longitudinal light stripe, from
the post-cephalic transverse skinfold to the vent, but absent on the head; venter
immaculate; limit between the dorsal and ventral regions poorly defined; light spots
on 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 stripe
surrounding 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. bicolor
by the longer head (HL about 92% of HW in H. matogrosso sp.nov.; HL below 90% of
HW 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 dermal
fold to vent (mid-dorsal stripe absent in E. bicolor, mid-dorsal stripe from the tip of
snout to vent in E. helianneae), by the absence of minute light spots on dorsum and
dorsal surfaces of members (present in E. helianneae), by the dorsal gray color of
dorsum of the snout invading the loreal region almost to the upper lip border (loreal
region conspicuously white in E. bicolor and in E. helianneae), and by the stripe on
the posterior surface of thighs broad, irregular (thin, well defined in E. bicolor and
in 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 than
long; head length 92.8% of head width and 22.1% of SVL; head width 23.8% of
SVL. Snout sub-elliptical in dorsal view, protruding in profile. Nostrils small, not
protuberant, directed anterolaterally, closer to tip of snout than to eye; internarial
distance smaller than the eye to nostril and interorbital distances, and larger
than the eye diameter and the upper eyelid width. Canthus rostralis rounded;
loreal region flat, sloping abruptly to the upper lip; lips not flared. Postcommisural
gland poorly developed. Eyes small, dorsolateral, only slightly protruding.
Interorbital space slightly convex, more than twice the upper eyelid width. No
cranial crests. A poorly defined transversal skinfold across back of the head,
bending downwards slightly behind the eyes to the shoulder; a poorly defined
skinfold through dorsolateral region from the axilla to the groin. Tympanum
concealed; supratympanic fold absent. Lower jaw with truncate, trilobed anterior
margin. Tongue large, oval, without a notch on posterior border. Choanae large,
subcircular, widely separated. Vocal slits present. Premaxillary, maxillary, and
vomerine odontophores absent. Vocal sac subgular, not expanded externally. A
weak skinfold crossing the chest between axillae.
Arms moderately robust, no tubercles or crests on forearm; palmar tubercle large,
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
21
divided longitudinally, twice as large than thenar tubercle; fingers slender, free, with
subarticular tubercles developed, rounded; supranumerary tubercles absent; tip of
fingers 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 length
90.9% of tibia length. Sum of thigh and tibia lengths 67.5% of SVL; thigh length
32.15% 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 touching
when 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; plantar
tubercles 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 slightly
rugose, smooth beneath; anal opening not modified, no para-anal tubercles.
Fig.5- Elachistocleis matogrosso sp.nov., holotype (MNRJ 4812; SVL 33.2mm), dorsal and ventral
views of body, and head profile.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
22
U.CARAMASCHI
In preservative, dorsum and dorsal surfaces of limbs uniformly grayish brown; a middorsal longitudinal light cream stripe, from the post-cephalic dermal fold to vent; a
poorly defined color limit between the dorsal and ventral regions; venter immaculate
clear cream; throat grayish; few, small spots on axillae and groin; an irregular broad
light cream stripe on the posterior surface of thighs; a large irregular, light cream spot
on the proximal internal surface of tibiae; a light cream stripe surrounding the knees
and 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, ED
1.6, 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 Mato
Grosso and Mato Grosso do Sul, Brazil, where the species occurs.
Variation – Except for minor details in corporal proportions, the descripton of
the holotype stands for females of the species. Size ranges from 29.0 to 33.2mm
SVL and the intensity of the color of the throat can vary. The males are smaller
than females, size ranging from 21.5 to 24.6mm SVL. Range, mean, and standard
deviation of the measurements of the type specimens of E. matogrosso are
presented in Table 5. In a few specimens the mid-dorsal stripe can be interrupted
or almost absent.
Geographical distribution – Central Brazil, in southwestern State of Mato Grosso
and northwestern State of Mato Grosso do Sul (Fig.6).
TABLE 5. Range (mm), mean (x¯ ), and standard deviation (SD) of the measurements of the type
specimens of Elachistocleis matogrosso sp.nov. (n = number of specimens).
C HARACTERS
SVL
HL
HW
IND
END
ED
UEW
IOD
HAL
THL
TL
FL
RANGE
21.5-24.6
5.0-6.1
5.55-6.6
1.3-1.8
1.9-2.3
1.3-1.5
0.8-1.4
2.3-2.7
4.5-5.3
6.9-9.5
8.3-9.85
8.8-11.8
(n = 5)
x̄
23.0
5.5
6.05
1.6
2.1
1.4
1.1
2.4
4.9
8.4
8.9
10.2
SD
RANGE
1.51
0.41
0.46
0.19
0.12
0.11
0.21
0.14
0.31
1.06
0.66
1.13
29.0-33.2
6.2-7.3
6.3-7.9
1.4-2.1
2.1-2.6
1.5-1.6
1.15-1.4
2.5-3.0
5.1-6.2
9.7-10.7
9.8-11.7
11.6-13.9
(n = 3)
x̄
30.6
6.65
7.1
1.8
2.4
1.6
1.2
2.75
5.5
10.2
10.65
12.6
SD
2.25
0.60
0.78
0.36
0.27
0.06
0.13
0.23
0.60
0.48
0.98
1.18
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
Fig.6- Geographic distribution of species of Elachistocleis.
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
23
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 affect
the 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) to
Buenos Aires, Argentina. The species, in their conception, would encompasses frogs
that, filling the characters that describe the genus Elachistocleis, had an immaculate
venter 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 presented
decision 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 length
below 90% of head width), dorsum smooth, uniformly grayish brown, without light spots
nor a distinctive middle longitudinal light stripe, venter immaculate yellow or greenish
yellow, a sharp color limit between the dorsal and ventral regions, no spots on axillae or
groin, a thin, well defined line on the posterior surface of the thighs, and a narrow light
stripe surrounding the knees and reaching the middle of the tarsus. The geographical
distribution (Fig.6) comprises southwestern and southern Brazil, in the states of Mato
Grosso do Sul, southern São Paulo, Paraná, Santa Catarina, and Rio Grande do Sul,
Paraguay (for detailed distribution see BUSQUETTI & LAVILLA, 2006), Uruguay (for detailed
distribution see NÚÑEZ et al., 2004), and northern Argentina.
Elachistocleis surinamensis was originally characterized by DAUDIN (1802) as having a
brown 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 of
an evident light vertebral stripe. The type locality was originally stated as “Surinam”, a
broad 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 probably
originally deposited in the Museum Nationalle d’Histoire Naturelle de Paris, is currently
lost. As an operative framework for a necessary revision of this species complex
represented by E. surinamensis, LAVILLA et al. (2003) considered that it would be
represented 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), who
reported the geographical distribution for the species from a few localities in the states
of São Paulo, Minas Gerais, and Goiás, Brazil. Actually, the geographical distribution of
E. 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 southeastern
Brazil, in the states of Minas Gerais, Espírito Santo, Rio de Janeiro, and São Paulo. It is
possible that more than one species is involved, but at first not discriminated by the
used characters.
The remaining species, including those described herein, have modern descriptions, defined
name bearing types, and precise type localities. Elachistocleis piauiensis is distributed in
the states of Piauí, Maranhão, Ceará, and Tocantins (see map in NUNES et al., 2010, excluding
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
25
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 das
Araucárias, Serra Geral, State of Rio Grande do Sul, southern Brazil, at 900-1200m of
altitude (FROST, 2010) (Fig.6). Elachistocleis skotogaster is known from the type locality,
Los Toldos, 1100m altitude, Departamento Santa Victoria (LAVILLA et. al., 2003), and from
one locality near to Isla de Cañas, 800m altitude, Departamento Iruya, and two localities
in the Departamento Orán, 350 and 450m altitude, all in the Provincia Salta, northern
Argentina (see map in CAJADE et al., 2009, and Fig.6). Elachistocleis magnus is known from
three localities in the State of Rondônia, Brazil (Fig.6). The geographic distributions of the
new species are treated in the respective accounts.
The present observations and descriptions contribute to a better undertanding of the
composition of Elachistocleis. Notwithstanding, the actual diversity encompassed by
this genus is far from known, as can be seen, only as examples, the pictures of the
species, 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 Carolina
Mello (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 allowing
examination 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 Nacional
de Desenvolvimento Científico e Tecnológico (CNPq) provided financial support.
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27
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APPENDIX
ADDITIONAL SPECIMENS EXAMINED
Elachistocleis bicolor – BRAZIL: MATO GROSSO DO SUL: Bela Vista (EI 1400-1418); Bonito (CHUNB
49299); Maracaju (EI 4010-4013); Ponta Porã, Inhuverá (MNRJ 6955); Três Lagoas (CFBH 1360813610). 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 48154817); Curitiba (MZUSP 13659-13661); Jaguariaíva (MNRJ 66159); Piraquara, Mananciais da
Serra (MHNCI 1181). SANTA CATARINA: (MHNCI 1181a,b); Faxinal dos Guedes, Represa Santa
Laura (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 (MZUSP
21727); 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, EI
756); 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-
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
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 (USNM
94101); Entre Independencia y Villarica, Arroio Ytá (MNRJ 6975-6976); Brejo de Ipuã, near
Nueva Italia (MNRJ 66903). URUGUAY: ARTIGAS: Barra del Arroio Yacuí (MNRJ 6982-6983).
ARGENTINA: CHACO: Resistencia (MZUSP 36386). CORRIENTES: Ituzaingó, Santa Tecla (MNRJ
39940-39942). 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 4754747548, 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); Flores
de 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); Rio
Verde (CHUNB 49435); Santa Tereza (MNRJ 53149-53150); São Domingos (CHUNB 32223-32224,
35568-35576, 47527-47528, 47530); São João da Aliança, Jatobazinho (MNRJ 6974); Três
Ranchos (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, Fazenda
Cuiabá (MZUFV 2973); Andrequicé, UHE Formoso (MNRJ 13891); Araponga (MZUFV 8279-8280);
Astolfo Dutra, Triunfo (MCNAM 6542); Belmiro Braga, Brumadinho (MNRJ 27503); Betim (MNRJ
66849, 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 (MNRJ
56303); Catas Altas, Serra do Caraça (MNRJ 44708, 60489-60493, 66029); Catas Altas (MCNAM
12884-12885); 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 (MCNAM
9546-9547); 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 5070550707); João Pinheiro, Fazenda Veredas (MNRJ 38813); Juiz de Fora, Água Limpa (MNRJ 69606961, 6971-6972); Manga, Mocambinho (MNRJ 40589, MZUFV 2838-2839); Mariana, Samitri
(MZUFV 578-583); Marliéria, Parque Estadual do Rio Doce (MNRJ 13980, 51092, 51093, MZUFV
4839-4840); Mocambinho, Jaíba (MZUFV 2838-2839, 3029); Nova Era (MCNAM 1112); Ouro
Branco, Serra do Ouro Branco (MZUFV 7102); Pains (MCNAM 7434-7435); Palmital, UHE
Queimados (MCNAM 2894, 4001-4004, 5851-5853, 8981); Paracatu (MCNAM 10593, CHUNB
25365-25445, 25447-25483, 25485-25500, 26915); Paracatu/Pirapora, Linha de Transmissão
(MCNAM 11012-11013); PARNA Serra do Cipó (MCNAM 2697-2698, 2732, 2762,
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
NOTES ON THE TAXONOMIC STATUS OF ELACHISTOCLEIS OVALIS...
29
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 1038510386, 10389, 10391, 10419); Rio Preto, PCH Mato Limpo (MCNAM 2304); RPPN Serra
do Caraça (MCNAM 5796); Sacramento (CHUNB 49400); Santana do Riacho, Serra do
Cipó (MCNAM 1851-1854, 2458, 6955); Santa Rita de Jacutinga (MCNAM 2060); São
Gonçalo do Rio Abaixo (MCNAM 13725-13732); São Gonçalo do Rio Preto (MCNAM
8563-8564); São João Nepomuceno (MNRJ 30488, 32185-32186, 49639-49642); São
José do Goiabal (MCNAM 12843); São José do Mantimento/Durandé, PCH Varginha e
Várzea Alegre (MCNAM 11218); Tocantins (MZUFV 7591-7592); Tombos (MZUFV 53205321); Três Pontas (MCNAM 7871); Urucânia (MCNAM 8364); Urucuia (MNRJ 734),
Viçosa, UFV-Piscicultura (MZUFV 640, 8044); Viçosa, UFV-Mata do Paraíso (MZUFV
2525, 2531, 6138); Viçosa, Sítio do Fiúza (MZUFV 5309). ESPÍRITO SANTO: Itaperuna
(MNRJ 60299). RIO DE JANEIRO: Atafona (MNRJ 7978-6981); Campo Belo (AL-MN
075, 405, 407, 814, 842); Itamonte, Planalto do Itatiaia (MNRJ 13978-13979); Itaperuna
(MNRJ 54366); Lídice (MNRJ 66675); São João da Barra (MNRJ 6968-6970). SÃO
PAULO: Angatuba (CFBH 23136); Botucatu (EI 4403); Botucatu, Fazenda Dinucci (MNRJ
61116, MNRJ 66851); Botucatu, Fazenda Edgardia (MNRJ 66860); Botucatu, Fazenda
Lageado (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 (CFBH
4997); 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 ovale
concolor); São Paulo (CHUNB 47543); São Simão (EI 1396-1397); Taubaté (EI 12341236); Ubatuba (CFBH 10907).
Elachistocleis erythrogaster – RIO GRANDE DO SUL: São Francisco de Paula, Centro de
Pesquisas e Conservação da Natureza Pró-Mata (MNRJ 39098, paratype, ex-MCP 3142).
Elachistocleis helianneae – BOLIVIA: BENI: Guayaramerim (USNM 123964). SANTA
CRUZ: 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: Santa
Maria (MNRJ 2918). PARAGUAY: Assunción (MNRJ 6949) [in error].
Elachistocleis piauiensis – BRAZIL: PIAUÍ: Brejo do Piauí (MNRJ 42073); Castelo do
Piauí (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
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
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 4953049531); Corumbá (MCNAM 7404, 7634); Coxim (MZUSP 61040-61041); Miranda (MNRJ
1858, 6964, 6965-6967, MZUSP 65107-65155); Nioaque (EI 4014); Salobra (MNRJ
6950-6954); 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 Bom
Intento (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).
Bol. Mus. Nac., N.S., Zool., Rio de Janeiro, n.527, p.1-30, ago.2010
BOLETIM DO MUSEU NACIONAL, NOVA SÉRIE, ZOOLOGIA ISSN 0080-312X
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