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The genus Chiasmocleis Mghely, 1904 (Anura, Microhylidae) in the Atlantic

Rain Forest of Brazil, with description of three new species.

Article · January 1997

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 INTERNATIONAL
JOURNAL OF BATRACHOLOGY
October 1997 Volume 15, No 2

The genus Chiasmocleis Mghely, 1904

(Anura, Microhylidae)
in the Atlantic Rain Forest of Brazil,
with description of three new species
Carlos Alberto Gonqalves DA CRUZ*,
Ulisses CARAMASCHI ** & Euggnio IZECKSOHN
*
* Departamento de Biologia Animal, Universidade Federal Rural do Rio de Janeiro,
2385 1-970 Seropedica, RJ, Brazil
** Departamento de Vertebradoq Museu Nacional do Rio de Janeiro, Quinta da Boa Vista,
20940-040 Rio de Janeiro, RJ, Brazil

A taxonomic study of the species of the genus Chiasmocleis occurring

in the Atlantic Rain Forest of eastern Brazil is presented. Chiasmocleis bice-
goi Miranda-Ribeiro, 1 9 20 is synonymized with C. a lbopunctata (Boettger,
1885), and rendered extralimital for the present work. The synonymization
of C. urbanae Bokermann, 1 9 5 2 with C. leucosticta (Boulenger, 1888) is
reaffirmed. Chiasmocleis leucosticta and C. schubarti Bokermann, 1 9 5 2
are redescribed, and three new species are described.

The @RSS CS4iasrwo&ls, proposed b y , M I ? ~ L (1904X

Y currenty contains 12 species,
distGbuted TromPanama to soubernSouth hme'rica, nohh anh eas'l. &%it P;nbes\Fxsm,
1985). The validity of the species included in this genus has not been assessed by a modern
review. Materials from the Atlantic Forests of Brazil indicate such an assessment is needed.
Herein, we present a review of the species of Chiasmocleis inhabiting the Atlantic Rain Forest
of eastern Brazil. This systematic review includes: (1) re-examination of previously described
taxa; (2) redescription of the valid species; and (3) description of three new species.

MATERIALS
AND METHODS

Abbreviations of the collections housing specimens are: BMNH (Natural History

Museum, London, United Kingdom); CFBH (Celio F. B. HADDADCollection, Universidade
 ALYTES 15 (2)

Estadual Paulista, Rio Claro, SP, Brazil); EI (Euggnio IZECKSOHN

Collection, Universidade
Federal Rural do Rio de Janeiro, RJ, Brazil); MNRJ (Museu Nacional do Rio de Janeiro, RJ,
Brazil); MZUSP (Museu de Zoologia, Universidade de SBo Paulo, SP, Brazil); SPCS (Sergio
POTSCHDE CARVALHO E SILVACollection, Universidade Federal do Rio de Janeiro, RJ,
Brazil); UFV (Museu de Historia Natural, Universidade Federal de Vigosa, MG, Brazil).
Specimens examined are listed in app. 1.
Measurements, in millimeters, 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); THL (thigh length); TL (tibia
length); FL (foot length).

Each of the five names proposed for a member of the genus Chiasmocleis occurring in the
Atlantic Rain Forest is discussed in chronological order.

This species was described on the basis of a female specimen (type not stated; BMNH
88.4.23.3 registered as holotype, according to FROST,1985) collected at "Sierra do Catharina,
Province Santa Catharina, Brazil" (BOULENGER, 1888). Currently this locality is in the
Catarina Mountain Range, in the state of Santa Catarina, southern ~ r a z i l PARKER
. (1934)
transferred the species to the genus Chiasmocleis, under the combination Chiasmocleis
~eucosticta.

The species was described based on a single specimen from "0s Perus, S. Paulo"
(MIRANDA-RIBEIRO, 1920). Currently, this locality is a suburb within the western side of the
city of SBo Paulo, state of SBo Paulo, Brazil. The holotype (MZUSP 924, formerly MZUSP
593, a juvenile (15.0 mm SVL) of undetermined sex, is in very poor condition, totally
discoloured, the ventral and pectoral regions dilacerated, and arms and legs almost totally
destroyed. Direct comparison of the holotype of C. bicegoi with a recently collected young
specimen of Chiasmocleis albopunctata Boettger, 1885, obtained at Iperb, state of Siio Paulo
(MNRJ 17324, 13.9 mm SVL), indicates that the former is a junior synonym of the latter.
Chiasmocleis albopunctata is an open area inhabitant, occurring in the "cerrados" of Brazil in
the states of Siio Paulo, Minas Gerais, Goias, Mato Grosso, Mato Grosso do Sul, and Distrito
Federal, and in Paraguay and Bolivia. Ipero is about 90 km from the type locality of C.
bicegoi, and it is perfectly acceptable that the species occurred near the city of Siio Paulo at the
end of the last century, when the holotype was collected by BICEGO.The shape of the snout,
although with some deformation in the holotype of C. bicegoi, is coincident in both speci-
mens; vestiges of the white blotches on the snout and dorsolateral region of the body
characteristic of C. albopunctata are still observed in the holotype of C. bicegoi. Thus,
synonymization of C. bicegoi with C. albopunctata renders it extralimital for the present work.
The citation of C. bicegoi for Itaguai, state of Rio de Janeiro, by BOGART& NELSON(1976)
applies to a new species described in the account below.

SPINULOSUS MIRANDA-RIBEIRO,
NECTODACTYLUS 1924

The genus Nectodactylus was proposed by MIRANDA-RIBEIRO (1924) to accomodate a

new species, named iV spinulosus (syntypes MNRJ 525 and 5582), from "Humboldt - Sta.
Catharina - Brasilia" (currently Corupa, state of Santa Catarina, Brazil). The genus was
synonymized with Chiasmocleisby PARKER (1934), resulting in the combination Chiasmocleis
spinulosa. Later, CARVALHO (1954) synonymized Nectodactylus spinulosus with Engystoma
leucostictum, thus recognizing Chiasmocleis leucosticta as the valid name for the species.

The species was described based on a young specimen (holotype MZUSP 2309) from
"Corrego Juncado, Linhares", state of Espirito Santo, Brazil (BOKERMANN,
1952). This is a
perfectly valid and recognizable species.

The species was described on the basis of a purported male specimen (holotype MZUSP
9033) from "Ilha de SZo Sebastizo", on the coast of the state of SZo Paulo, Brazil (BOKER-
MANN, 1952). Chiasmocleis urbanae was synonymized with C. leucosticta by BOKERMANN
(1966); this action was not recognized in FROST(1985), where C. urbanae was treated as a valid
species. Examination of the holotype of C. urbanae showed it to be a female specimen and
identical to specimens of C. leucosticta from the state of Santa Catarina and the southern
portion of the state of Siio Paulo. Consequently, we agree with BOKERMANN (1966) and
consider C. urbanae to be a synonym of C. leucosticta.

Chiasmocleis leucosticta (Boulenger, 1888)

Engystoma leucostictum Boulenger, 1888.
Chiasmocleis leucosticta: PARKER,1934.
Nectodactylus spinulosus Miranda-Ribeiro, 1924.
Chiasmocleis spinulosa : PARKER,1934.
Chiasmocleis urbanae Bokermann, 1952.

Diagnosis. - A medium sized species of Chiasmocleis diagnosed by the following combination

of characters: (1) SVL 19.0-22.8 mm in males, 2 1.8-25.5mm in females; (2) body trunk ovoid;
 ALYTES 15 (2)

Fig. 1 . - Dorsal and ventral views of Clrrasmoclets leucos~!cra(Boulenger, 1888) (MNRJ 17401).

(3) snout short, tip slightly truncate in dorsal view, and slightly protruding in lateral profile;
(4) hand and foot of male webbed, of female half webbed; (5) fingers and toes lacking disks,
fringed on free parts in females;(6)fingers and toes with few, small lateral dermal spines only
in males; (7) dorsal surfaces with small dermal spines; (8) in preservative, color on dorsum
dark brown with small white dots irregularly distributed, and a light IongitudinaI mid-dorsal
line; (9) posterior sides of legs with a light longitudinal line; (10) venter roughly marbled in
dark brown and pale cream.
Description. - Size medium for the genus; body trunk avoid (fig. 1); bead short, as broad as
long; nostrils at the tip of snout, not protubrant, and directed laterally; snout short, tip
slightly truncate in dorsal view (fig. 2 A), and slightly protruding in Iateral profiie (fig. 2 B);
internaria1 distance smaller than eye to nostril distance, and only slightly smaller than eye
diameter; eye diameter less than eye to nostril distance; canthus rostraIis only slightly defined;
loreal region oblique, ff at; lips not flared; eyes small, only slightly protruding; upper eyelid
width one third of the interorbita1space; interorbital area flat; cranial crests and occipital fold
absent; postorbital fold present; tympanum absent; upper jaw projecting beyond lower;
mandible with truncate, trilobed anterior margin;tongue large, ovoid; choanae small, round-
ed, widely separated; a small, subgular vocal sac present.
Arms slender, lacking tubercles and crests on forearm. Hand of male (fig. 2 CJ webbed,
of female (fig. 3 A) half webbed; fingers lacking diskg fringsd on free parts in females, with
lateral dermal spines only in males; finger length I < I1 < TV < 111; subarticular tubercles well
developed, rounded; supernumerary tubercles absent; paImar tubercle large, divided in two
parts; thenar tubercle large, rounded, at the base of finger 1.
Legs short, robust; knee and heel lacking tubercles; tibia! and tarsal ridges absent. Foot
of male (fig. 2 D) webbed, of female (fig. 3 B) half webbed; toes lacking disks and fringed on
free parts; tm length I < I1 < V 4 111< IV; toes with few, small lateral dermai spines in males,
Fig. 2. - CIziasmocleis Ieucosticta (Boulenger, 1888), male (MNRJ 17901). Dorsal (A) and ventral (B)
views of head. Hand (C). Foot (D).
 ALYTES 15 (2)

Fig. 3. - Chiasmocleis leucosticta (Boulenger, 1888), female (MNRJ 17902). Hand (A). Foot (B).
 C. I o u c o r t i c t r
C- rchubrrti

** C - carvalhoi
C. schubarti
C. rtlantioa
+
+
C-capixaba
C . carvalhoi

-
0 100 200 300 KM
L

Fig. 4. - Geographical distribution of the species of Chiasmocleis in the Atlantic Rain Forest of Brazil
(states of southeastern and southern regions of Brazil: MG, Minas Gerais; ES, Espirito Santo; RJ,
Rio de Janeiro; SP, S5o Paulo; PR; Parana; SC, Santa Catarina; RS, Rio Grande do Sul).

lacking in females; subarticular tubercles well developed, rounded; supernumerary tubercles

absent; an oval inner, but no outer, metatarsal tubercle. Thigh length slightly less than tibia
length; knee and elbow widely separated with limbs adpressed to sides of body; combined
thigh and tibia length approximately 83 % of snout-vent length in males, 85 % in females;
heels slightly superposed when flexed legs held at right angles to body; foot length approxi-
mately 66 % of snout-vent length in males, 69 % in females.
 ALYTES 15 (2)

Skin smooth above and beneath; dorsal and ventral surfaces of body and limbs with
small, uniformly distributed dermal spines, more numerous in males. Anal opening not
modified, lacking para-anal tubercles and glands around anus.
In preservative (70 % ETOH), color on dorsum dark brown with small white dots
irregularly distributed; a light longitudinal mid-dorsal line meeting, above the anus, another
light longitudinal line present on the posterior sides of legs. Venter roughly marbled in dark
brown and pale cream; male throat infuscated.
Variation. - Variation in measurements is presented in tab. 1. Females are larger than males,
with interdigital webbing less developed. In some females, a light line is present on ventral
surface of arms and pectoral region, where it assumes a V-shape, crossed by a perpendicular
light longitudinal mid-ventral line.
Geographical distribution. - Known from the states of Siio Paulo and Santa Catarina, in
southeastern and southern Brazil (fig. 4). The occurrence of the species in the state of Parana
is predictable.
Remarks. - Specimens of C. leucosticta were collected in temporary ponds inside the forest,
from about 30 to 800 m above sea level. The reproduction is explosive, when numerous male
and female specimens congregate at the reproductive sites. KASAHARA & HADDAD(1997)
reported a population of C. leucosticta from Ribeiriio Branco, state of Siio Paulo, to be
tetraploid (4 n = 48).

Chiasmocleis schubarti Bokermann, 1952

Chiasmocleis schubarti Bokermann, 1952.

Diagnosis. - A medium sized species of Chiasmocleis diagnosed by the following combination

of characters: (1) SVL 19.2-26.3mrn in males, 20.4-34.5 mm in females; (2) body trunk ovoid;
(3) snout short, tip rounded in dorsal and lateral profile; (4) hand not webbed, foot only
slightly webbed at the base of toes in both sexes; ( 5 ) fingers and toes lacking disks, slightly
fringed; (6) fingers and toes of males with small, lateral dermal spines, absent in females; (7)
dorsal and ventral surfaces of body and limbs with small, uniformly distributed dermal spines
in males, absent in females except in anal region; (8) in preservative, color on dorsum
uniformly dark brown; (9) posterior sides of thighs with a light longitudinal line; (10) venter
roughly marbled in dark brown and pale cream.
Description. - Size medium for the genus; body trunk ovoid (fig. 5); head short, broader than
long; nostrils near the tip of snout, not protuberant, and directed laterally; snout short, tip
rounded in dorsal and lateral profile (fig. 6 A-B); internarial distance smaller than eye to
nostril distance, and slightly larger than eye diameter; eye diameter less than eye to nostril
distance; canthus rostralis only slightly defined; loreal region oblique, flat; lips not flared; eyes
small, only slightly protruding; upper eyelid width one third of the interorbital space;
interorbital area flat; cranial crests and occipital fold absent; postorbital fold present; tympa-
num absent; upper jaw projecting beyond lower; mandible with truncate, trilobed anterior
margin; tongue large, ovoid; choanae small, rounded, widely separated; a small, subgular
vocal sac present.
 Fig. 5. - Dorsal and ventral views of Ckiasmocleis schvharri Bokermann, 1952 (MNRJ 17538).

Arms slender, lacking tubercles and crests on forearm. Hand (fig. 6 C) not webbed in
both sexes; fingers lacking dish, slightly fringed; fingers with small, lateral dexmaI spines in
males, absent in females; finger length I < 11 < IV c 111; subarticular tubercles we11 developed,
rounded; supernumerary tubercles absent; palmar tubercle large, divided in two parts; thenar
tubercle small, rounded, at the base of finger I.
Legs short, robust; knee and heel lacking tubercles; tibia1 and tarsal ridges absent. Foot
(fig.6 D) only slightly webbed at the base of toes in both sexes; toes lacking disks, slightly
fringed; toe length I c I1 < V < TIT < IV; toes with small, lateral dermal spines jr!males, absent
in females; subarticular tubercles well developed, rounded; supernumerary tubercles absent; a
small, rounded inner, but no outer, metatarsal tubercle. Thigh length slightly less than tibia
length; knee and elbow widely separated with limbs adpresd to sides of body; combined
thigh and tibia length approximately 85 % of snout-vent length in males, 82 % in females;
heels slightly superposed when flexed legs held at right angles to body; foot length approxi-
mately 70 % of snout-vent length in males, 64 % in females
Skin smooth above and beneath; dorsal and ventral surfaces of body and limbs with
small, uniformiy distributed dermal spines in males, absent in females except in anal region.
Anal opening not mokfied, lacking para-anal tubercles and glands around anus.
In preservative (78% ETQH), color on dorsum dark brown; irregular white blotches on
outer surfaces of arms and forearms; dorsal surfaces of hands and feet with small white dots
 ALYTES 15 (2)

Fig. 6. - Chiasmocleis schubarti Bokermann, 1952 (MNRJ 17538). Dorsal (A) and ventral (B) views of
head. Hand (C). Foot (D).
Fig. 7. - Dorsal and ventral views of Chiasmocleisarlunricn sp nov. (MNRJ 17550, horotype).

irregularly distributed; a light longitudinal line on the posterior sides of thighs, Venter
roughly marbled in dark brown and pale cream; male throat infuscated.
Variation. - Variation in measurements is presented in tab. 1. In some specimens, a light
longitudinal mid-dorsal line is present.
Geographical distribution. - Known from the states of Espirito Santo and Minas Gerais in
southeastern Brazil (fig. 4).
Remarks. - Scattered specimens of C.schubarti were collected in pitfall traps install4 in low
and highland forests, 40 and 800 m above sea level. Explosive reproduction, when numerous
male and female specimens congregate, was observed in temporary ponds inside the forest.
KA~AHARA & HADDAD(1997) reported a diploid number of 24 chromosomes for this species.

Chiamodeis atlantica sp. nov.

Holoiype. - MNRJ 17550, adult male, collected at Tingui (22"36'S, 43"26W; ca. 40 m
altitude), municipality of Nova lguam, state of Rio de Jmeiro, Brazil, on 27 December 1972,
by C. A. G. DA CRUZ, S. D. L. RAIMUNW,J. G. SILVAand E. IZECKSOHN.
Pararopoiypes.- MNRJ 1755 1-1 7554 and EI 8940-8954,collected with the holotype; MNRJ
17549, on 30 January 197 1, by J. JIM, V,C. J m and E. IZECKSDHN.
Diagnosis. - A medium sized species of Chiaswtoeleishagnosed by the following combination
of characters:(1) SVL 22.0-25.0mm in males, 30.0-3 1.8 rnm in females; (2) body trunk ovoid;
(3) snout short, tip truncate in dorsal view, and slightly protruding in lateral profile; (4) hand
and foot not webbed; ( 5 ) fingersand toes lacking disks, slightly fringed; (6) fingers and toes of
60 ALYTES 15 (2)

males with lateral dermal spines, absent in females; (7) dorsal and ventral surfaces of body
and limbs with uniformly distributed dermal spines in males, absent in females except in anal
region; (8) in preservative, color on dorsum uniformly brown; (9) posterior sides of thighs
with a light longitudinal line; (10) venter roughly marbled in brown and pale cream.
Description. - Size medium for the genus; body trunk ovoid (fig. 7); head short, wider than
long; nostrils at the tip of snout, not protuberant, and directed laterally; snout short, tip
truncate in dorsal view (fig. 8 A), and slightly protruding in lateral profile (fig. 8 B); internarial
distance smaller than eye to nostril distance, and larger than eye diameter; eye diameter less
than eye to nostril distance; canthus rostralis slightly defined; loreal region almost vertical,
slightly concave; lips not flared; eyes small, only slightly protruding; upper eyelid width one
third of the interorbital space; interorbital area flat; cranial crests and occipital fold absent;
postorbital fold well developed; tympanum absent; upper jaw slightly projecting beyond
lower; mandible with truncate, trilobed anterior margin; tongue large, ovoid; choanae small,
rounded, widely separated; a small, subgular vocal sac present.
Arms slender, lacking tubercles and crests on forearm. Hand (fig. 8 C) not webbed in
both sexes; fingers lacking disks, slightly fringed; fingers with lateral dermal spines in males,
absent in females; finger length I < I1 < IV < 111; subarticular tubercles well developed,
rounded; supernumerary tubercles absent; palmar tubercle large, divided in two parts; thenar
tubercle small, rounded, at the base of finger I.
Legs short, robust; knee and heel lacking tubercles; tibia1 and tarsal ridges absent. Foot
(fig. 8 D) only slightly webbed at the base of toes in both sexes; toes lacking disks, slightly
fringed; toe length I < I1 < V < I11 < IV; toes with lateral dermal spines in males, absent in
females; subarticular tubercles well developed, rounded; supernumerary tubercles absent; a
small, oval inner, but no outer, metatarsal tubercle. Thigh length slightly less than tibia length;
knee and elbow widely separated with limbs adpressed to sides of body; combined thigh and
tibia length approximately 87 % of snout-vent length in males, 78 % in females; heels slightly
superposed when flexed legs held at right angles to body; foot length approximately 70 % of
snout-vent length in males, 67 % in females.
Skin poorly rugose above and smooth beneath; dorsal and ventral surfaces of body
and limbs with uniformly distributed dermal spines in males, absent in females except in
anal region. Anal opening not modified, lacking para-anal tubercles and glands around
anus.
In preservative (70 % ETOH), color on dorsum uniformly brown; dorsal surfaces of
hands and feet with small white dots irregularly distributed; a light longitudinal line on the
posterior sides of thighs. Venter roughly marbled in brown and pale cream; male throat
infuscated.
Measurements of holotype. - SVL 22.8; HL 6.9; HW 8.0; IND 2.0; END 2.0; ED 2.0; UEW
1.0; IOD 2.7; THL 10.4; TL 10.7; FL 16.7.
Variation. - Variation in measurements is presented in tab. 1. In some specimens, a light
longitudinal mid-dorsal line is present.
Etymology. -The specific name, a Latin adjective, refers to the Atlantic Rain Forest of eastern
Brazil,
Fig. 8. - Chiasmocleis atlantica sp. nov. ( M N R J 17550, holotype). Dorsal (A) and ventral (B) views of
head. Hand (C).Foot (D).
 ALYTES 15 (2)

Geographical distribution. - Known from the states of Rio de Janeiro and Siio Paulo in
southeastern Brazil (fig. 4).
Remarks. - Specimens of C. atlantica were collected in lowland forests, at 40 m above sea level.
Explosive reproduction, when numerous male and female specimens congregate, was obser-
ved in temporary ponds inside the forest.

Chiasmocleis capixaba sp. nov.

Holotype. - MNRJ 17514, adult male, collected at the municipality of Aracruz (19"59'S,
40°12'W; ca. 60 m altitude), state of Espirito Santo, Brazil, on 29 November 1995, by J. L.
GASPARINI.
Paratopotypes. - MNRJ 17515-17529, collected with the holotype; MNRJ 17532-17534, on
2-5 November 1994, by R. P. BASTOSand J. L. GASPARINI; MNRJ 17891-17895, on 29
November - 1 December 1995, by J. L. GASPARINI; MNRJ 17535-17537, on 15-17 January
1996, by J. P. POMBAL Jr., C. F. B. HADDADand J. L. GASPARINI; EI 8955-8956, on 20-25
November 1994, by R. P. BASTOS and J. L. GASPARINI;
CFBH 2668-2669, on 20-25 November
1994, by R. P. BASTOS and J. L. GASPARINI; CFBH 2685, on 23-27 January 1995, by J. P.
POMBAL Jr., J. L. GASPARINI and C. F. B. HADDAD;CFBH 2693-2695, on 27 November 1995,
by J. L. GASPARINI; CFBH 270 1-2702,on 30 November 1995, by J. L. GASPARINI;CFBH 27 14,
on 20 December 1995, by J. L. GASPARINI.
Diagnosis. - A small sized species of Chiasmocleis diagnosed by the following combination of
characters: (I) SVL 14.7-16.5 mm in males, 17.9-21.7 mm in females; (2) body trunk elongate
ovoid; (3) snout short, tip rounded in dorsal and lateral views; (4) hand not webbed, and foot
webbed in both sexes; ( 5 ) fingers and toes lacking disks, extensively fringed on free parts; (6)
fingers and toes of males with lateral dermal spines, absent in females; (7) dorsal and ventral
surfaces of body and limbs with small, uniformly distributed dermal spines in males, absent in
females except in anal region; (8) in preservative, color on dorsum grayish brown with gray
blotches irregularly distributed; (9) posterior sides of legs with or without a light longitudinal
line; (10) venter finely marbled in brown and pale cream.
Description. - Size small for the genus; body trunk elongate ovoid (fig. 9); head short,
approximately as broad as long; nostrils at the tip of snout, not protuberant, and directed
anterolaterally; snout short, tip rounded in dorsal and lateral views (fig. 10 A-B); internarial
distance equal eye diameter, smaller than eye to nostril distance; canthus rostralis rounded;
loreal region oblique, slightly concave; lips not flared; eyes small, only slightly protruding;
upper eyelid width smaller than half of the interorbital space; interorbital area flat; cranial
crests and occipital fold absent; post orbital fold present; tympanum absent; upper jaw slightly
projecting beyond lower; mandible with truncate, trilobed anterior margin; tongue large,
ovoid; choanae small, rounded, widely separated; a small, subgular vocal sac present.
Arms slender, lacking tubercles and crests on forearm. Hand (fig. 10 C) not webbed in
both sexes; fingers lacking disks, extensively fringed; fingers of males with lateral dermal
spines, absent in females; finger length I < I1 < IV < 111; subarticular tubercles well developed,
rounded; supernumerary tubercles absent; palmar tubercle large, divided in two parts; thenar
tubercle large, rounded, at the base of finger I.
 sp nov. (MNRJ 17514, hototype).
hg.9. -Dorsal and ventral views of Cki~smocIeiscapi.uaha

Legs short, robust; knee and heel lacking tubercles; tibia1 and tarsal ridges absent. Foot
of male (fig. 10 D)extensivelywebbed, of female webbed only at the base of toes; toes lacking
disks and fringed on free parts; toe length 1 < 11< V < I11 c IV; toes wilth lateral dermal spines
in males, absent in females; subarticular tubercles well developed, rounded; supernumerary
tubercles absent; an oval inner, but no outer, metatarsal tubercle. Thigh length slightly longer
than tibia length in males, and slightly less in females: knee and elbow widely separated with
limbs adpressed to sides of body; combined thigh and tibia length approximately 79 % of
snout-vent length in males, 74 % in females; heels slightly superposed when flexed legs held at
right angles to body; foot length approximately 66 % of snout-vent length in males, 59 % in
females.
Skin smooth above and beneath; dorsal and ventral surfaces of body and limbs with
small, uniformly distributed dermal spines in males, absent in females except in anal region.
Anal opening not modified, lacking para-anal tubercles and glands around anus
Jn preservative (70 % ETOH), color on dorsum grayish brown with gray blotches
irregularly distributed; loreal region dark gray; a light longitudinal mid-dorsal line on body
and a similar line on the posterior sides of thighs present or absent. Ventral surfaces of body
and limbs finely marbled in brown and pale cream; male throat infuscated.
Measurements ofholotype. - SVL 15.5; HL 5.0; HW 5.2; I N D 1.0; END 1.3; ED 1.4; UEW
1.0; IOD 1.8; THL 6.4; TL 6.6;FL 10.3.
Variation.- Variation in measurements is presented in tab. 1. Females are larger than males,
with interdigital webbing less developed. The light mid-dorsal longitudinal line on body and
the similar line on posterior surfaces of thighs are observed in approximately half of the
examined specimens.These lines appear associated with a light line on ventral surfaces of
arms and pectoral region, where it assumes a V-shape.
 ALYTES 15 (2)

Fig. 10. - Chiasmocleis capixaba sp. nov. (MNRJ 17514, holotype). Dorsal (A) and ventral (B) vien
head. Hand (C). Foot (D).
Etymology. - The name of the species, a native Brazilian word here utilized as a noun in
apposition, refers to the occurrence of the species as an inhabitant of the state of Espirito
Santo.
Geographical distribution. - Known only from the state of Espirito Santo in southeastern
Brazil (fig. 4).
Remarks. - Scattered specimens of C. capixaba were collected in pitfall traps installed in
lowland forests, at 60 m above sea level. Explosive reproduction, when numerous male and
female specimens congregate, was observed in temporary ponds inside the forest.

Chiasmocleis carvalhoi sp. nov.

Holotype. - MNRJ 17505, adult male, collected at the Horto Florestal de Santa Cruz,
municipality of Seropedica (22"44'S, 43O43'W; ca. 40 m altitude), state of Rio de Janeiro,
Brazil, on 31 January 1995, by C. A. G. DA CRUZ,L. KRAUSEand G. VINCIPROVA.
Paratopotypes. - MNRJ 17506-175 13 and 17565, collected with the holotype; MNRJ 17480-
17504, on 2 December 1968, by E. IZECKSOHN, S. T. ALBUQUERQUE and J. SAMARAO; EI
2096-2235, on 21 February 1964, by J. JIMand E. IZECKSOHN; EI 2236-2307, on November
1964, by J. JIM,S. T. ALBUQUERQUE, W. F. MENDONCA and E. IZECKSOHN; EI 2308-2399, on
December 1964, by J. JIM, S. T. ALBUQUERQUE, W. F. MENDONCA and E. IZECKSOHN; EI
438 1-4389, on December 1965, by S. T. ALBUQUERQUE, W. F. MENDONCA, A. LEBEDENCO, J.
JIMand E. IZECKSOHN; EI 439 1-4393, on 19 October 1966, by S. T. ALBUQUERQUE, W. F.
MENDONCA and E. IZECKSOHN; EI 5332, on 17 September 1975, by E. IZECKSOHN and J. G.
SILVA; EI 8937-8938, on 30 November 1979, by I. FERREIRA.
Diagnosis. - A small sized species of Chiasrnocleis diagnosed by the following combination of
characters:(1) SVL 15.5- 18.3 mm in males, 16.0-22.5mm in females; (2) body trunk ovoid; (3)
snout short, tip rounded in dorsal and lateral views; (4) hand and foot not webbed in both
sexes; (5) fingers and toes lacking disks, fringed; (6) fingers and toes with lateral dermal spines
in males, absent in females; (7) dorsal and ventral surfaces of body and limbs with small,
uniformly distributed dermal spines in males, absent in females except in anal region; (8) in
preservative, color on dorsum brown with small white dots irregularly distributed; (9)
posterior sides of legs with or without a light longitudinal line; (10) venter finely marbled in
brown and pale cream.
Description. - Size small for the genus; body trunk ovoid (fig. 11); head short, slightly broader
than long; nostrils at the tip of snout, not protuberant, and directed anterolaterally; snout
short, tip rounded in dorsal and lateral views (fig. 12 A-B); internarial distance equal eye
diameter, smaller than eye to nostril distance; canthus rostralis rounded; loreal region
oblique, slightly concave; lips not flared; eyes small, slightly protruding; upper eyelid width
smaller than half of the interorbital space; interorbital area flat; cranial crests and occipital
fold absent; postorbital fold present; tympanum absent; upper jaw slightly projecting beyond
lower; mandible with truncate, trilobed anterior margin; tongue large, ovoid; choanae small,
rounded, widely separated; a small, subgular vocal sac present.
 ALYTES 15 (2)

Fig. II. -Dorsal and ventral viewsol Chiamocleis carvaihoisp. nov. (MNRJ 17505, holotype).

Arms slender, lacking tubercles and crests on foream. Rand (fig. 12 C) not webbed in
both sexes; fingers lacking disks, fringed; fingers of males with lateral dermal spines, absent in
females; finger length I < I1 < IV < 111; subarticular tubercles we11 developed, rounded;
supernumerary tubercles absent; palmar tubercle large, divided in two parts; thenar tubercle
small, rounded, at the base of finger I.
Legs short, robust; knee and heel lacking tubercles; tibia1 and tarsal ridges absent. Foot
(fig. 12 D) not webbed in both sexes; toes lacking disks, fringed; toe length I < I1 < V < 111 <
TV;toes with lateral derma! spines in males absent in females; subarticular tubercles well
developed, rounded; supernumerary tubercles absent; an oval inner, but no outer, metatarsal
tubercle. Thigh length less than tibia length; knee and elbow widely separated with limbs
adpressed to sides of body; combined thigh and tibia length approximately 82 % of snout-
vent length in males, 79 % in females; heels slightly superposed when flexed l e g held at
right angles to body; foot length approximately 66 %of snout-vent length in males, 62 % in
females.
Skin smooth above and beneath; dorsal and ventral surfaces of body and limbs with
small, uniformly distributed dermal spines in maleq absent in females except in anal region.
Anal opening not modified,lacking para-anal tubercles and glands around anus.
In presemative (70 % ETOH), color on dorsum brown with small white dots irregularly
distributed; a light longitudinal mid-dorsal Iine on body, and a similar line on tbe posterior
sides of thighs, present or absent. Ventral surfaces of body and limbs finely marbled in brown
and pale cream;male throat infuscated.
Measummrs of holotype. - SVL 17.3; HL 5.6; HW 5.3; IND 1.3; END 1.6; ED 1.5; UEW
1.0; IOD 2.2; THL 7.2; TS 7.4; FL 1 1.6.
Fig. 12. - Chiasmocleis carvalhoi sp. nov. ( M N R J 17505, holotype). Dorsal (A) and ventral (B) views of
head. Hand (C).Foot (D).
68 ALYTES 15 (2)

Variation. - Variation in measurements is presented in tab. 1. A light mid-dorsal longitudinal

line on body and a similar line on posterior surfaces of thighs are observed in some specimens.
These lines appear associated with a light line on ventral surfaces of arms and pectoral region,
where it assumes a V-shape.
Etymology. - The name of the species honors the late Prof. Antenor Leitiio DE CARVALHO
(19 10-1985), an outstanding teacher, naturalist, and herpetologist.
Geographical distribution. - Known from the states of Rio de Janeiro and Siio Paulo in
southeastern Brazil (fig. 4).
Remarks. - Specimens of C. carvalhoi were collected in lowland forests, at 40 m above sea
level. Explosive reproduction, when numerous male and female specimens congregate, was
observed in temporary ponds inside the forest. BOGART & NELSON (1976) referred to a diploid
number of 24 chromosomes for specimens identified as C. bicegoi, collected at Itaguai, state of
Rio de Janeiro. This reference applies to C. carvalhoi.

Species of Chiasmocleis occurring in the Atlantic Rain Forest are readily separated in two
morphologically distinct groups. The first grouping involves C. leucosticta and C. capixaba,
characterized by the presence of well developed webbing on the feet. The second grouping
includes C. schubarti, C. atlantica, and C. carvalhoi, with no or only vestigial web on the feet.
Chiasmocleis leucosticta is larger than C. capixaba (see tab. I), and males of the former
have a fully webbed hand instead of the absence of webbing observed in C. capixaba. The
latter species has a finely marbled venter with males presenting numerous, conspicuous lateral
dermal spines on digits, whereas C. leucosticta presents a roughly marbled venter and small
dermal spines on digits. The geographical distribution of these two species is largely disjunct
(see fig. 4).
The striking webbing observed in C. leucosticta and C. capixaba is found in only one
other species of the genus, C. anatipes, described by WALKER & DUELLMAN (1974) from Santa
Cecilia, Ecuador. As noted by these authors, in the event that generic distinction should be
desirable for the fully foot-webbed species, the name Nectodactylus Miranda-Ribeiro, 1924
(type-species A! spinulosus, a junior synonym of C. leucosticta) is available.
In the second grouping, C carvalhoi is smaller than C. schubarti and C. atlantica (see
tab. l), and presents a finely marbled venter instead of the roughly marbled venter found in the
other two species. Chiasmocleis schubarti presents the snout rounded in dorsal and lateral
views, and C. atlantica has the snout truncate in dorsal and slightly protruding in lateral views.
Moreover, males of C atlantica have numerous, conspicuous lateral dermal spines on digits,
whereas C schubarti presents few, small lateral spines on those digits. Chiasmocleis atlantica
and C. carvalhoi are sympatric, and both are disjunctly distributed in relation to C. schubarti
and C. leucosticta. On the other hand, C. schubarti is sympatric with C. capixaba, and largely
disjunct from C. leucosticta (see fig. 4).
Tab. 1. - Measurements of five species of the genus Chiasmocleis Mihely, 1904 from the Atlantic Rain Forest of Brazil: Chiasmocleis leucosticta
(Boulenger , 1888); Chiasmocleis schubarti Bokermann, 1952; Chiasmocleis atlantica sp. nov. ; Chiasmocleis capljcaba sp. nov. ; Chiasmocleis
carvalhoi sp. nov. n, number of specimens; x, mean; s, standard deviation.

HL 5.4-7.0 6.0 0.47 6.1-7.7 6.7 0.56 5.4-8.1 6.8 0.65 4.2-5.4 4.8 0.42 4.3-6.0 5.2 0.46
HW 5.3-6.5 5.8 0.36 5.8-8.1 7.2 0.61 6.8-8.4 7.4 0.41 4.6-5.3 4.9 0.21 4.4-6.1 5.3 0.40
IND 1.2-1.6 1.5 0.11 1.4-2.1 1.9 0.20 1.2-2.0 1.6 0.21 1.0-1.4 1.2 0.14 1.0-1.4 1.3 0.14
END 1.6-2.0 1.7 0.12 1.7-2.3 2.1 0.14 1.4-2.3 1.9 0.23 1.2-2.0 1.5 0.22 1.4-2.3 1.7 0.23
ED 1.3-1.6 1.5 0.11 1.3-2.0 1.6 0.24 1.0-1.5 1.3 0.22 1.0-1.4 1.2 0.16 1.0-1.5 1.3 0.16
UEW 0.6-1.0 0.9 0.12 0.9-1.2 1.0 0.07 1.0-1.2 1.0 0.06 0.6-1.0 0.9 0.18 0.8-1.0 0.9 0.10
IOD 2.0-2.8 2.4 0.22 2.3-3.3 2.9 0.27 2.6-3.3 2.9 0.18 1.8-2.3 2.0 0.19 2.0-2.3 2.1 0.11
THL 7.8-9.1 8.3 0.33 8.1-11.1 9.4 0.78 9.0-10.7 9.9 0.46 5.4-7.0 6.3 0.48 6.0-7.3 6.8 0.41
TL 7.8-9.6 8.8 0.47 8.7-11.2 10.1 0.74 9.8-11.4 10.6 0.42 5.1-7.0 6.2 0.58 6.6-7.8 7.3 0.32
FL 11.9-15.6 13.4 0.88 14.5-17.5 15.9 1.04 11.7-17.8 16.5 1.31 9.5-11.2 10.3 0.45 10.0-12.6 11.3 0.78
Females ' n = 15 n = 12 n=3 n = 16 n = 15
Characters Range x s Range x s Range x s Range x s Range x s
SVL 21.8-25.5 23.6 1.15 20.4-34.5 28.7 4.60 30.0-31.8 30.6 1.04 17.9-21.7 20.2 1.01 16.0-22.5 20.2 1.93
HL 5.7-7.5 6.8 0.52 5.6-8.2 7.4 0.87 6.6-7.8 7.3 0.61 4.4-6.7 5.7 0.64 4.2-6.6 5.5 0.66
HW 5.7-7.4 6.4 0.45 6.2-10.2 8.0 1.15 8.1-8.2 8.1 0.06 5.4-6.2 5.7 0.22 5.0-6.4 5.6 0.40
IND 1.3-2.0 1.6 0.15 1.6-2.2 1.9 0.23 1.8-2.4 2.1 0.31 1.0-1.4 1.2 0.15 1.2-1.6 1.4 0.15
END 1.7-2.2 1.9 0.14 2.0-2.6 2.2 0.19 1.9-2.4 2.1 0.25 1.4-1.8 1.6 0.10 1.4-2.0 1.8 0.20
ED 1.4-2.0 1.6 0.18 1.4-2.0 1.8 0.24 1.6-2.0 1.8 0.21 1.0-1.9 1.3 0.21 1.0-2.0 1.4 0.30
UEW 0.7-1.3 1.0 0.12 0.9-1.4 1.1 0.16 1.1-1.4 1.3 0.15 0.7-1.0 0.9 0.09 0.8-1.0 1.0 0.07
IOD 2.5-3.0 2.7 0.18 2.7-3.7 2.2 0.29 3.6-3.7 3.7 0.06 2.0-2.9 2.4 0.20 2.1-3.0 2.4 0.28
THL 9.0-10.5 10.0 0.42 8.4-13.5 11.1 1.51 11.5-12.3 11.8 0.44 6.7-8.0 7.3 0.43 6.2-9.1 7.6 0.73
TL 9.3-11.0 10.3 0.46 9.4-13.8 11.9 1.39 11.8-12.8 12.3 0.50 6.4-8.3 7.5 0.61 6.5-9.1 7.9 0.70
FL 15.1-17.4 16.3 0.77 14.1-20.7 18.1 2.19 20.0-21.2 20.5 0.61 10.6-12.8 12.0 0.77 9.9-14.2 12.3 1.21
 ALYTES 15 (2)

L'etude taxinomique des especes du genre Chiasmocleis de la Forst Atlantique brksi-

lienne est presentee. Chiasmocleis bicegoi Miranda-Ribeiro, 1920 est mise en synonymie de C.
albopunctata (Boettger, 1885), une espece distribuee hors des limites geographiques conside-
rees dans le present travail. La synonymisation de C. urbanae Bokermann, 1952 avec C.
leucosticta (Boulenger, 1888)est confirmee. Une description detaillee de C. leucosticta et de C.
schubarti Bokermann, 1952 est fournie. Trois nouvelles especes sont decrites.

0 estudo taxonbmico das especies do gcnero Chiasmocleis que ocorrem na Floresta

Atlgntica do leste do Brasil e apresentado. Chiasmocleis bicegoi Miranda-Ribeiro, 1920 e
sinonimizada com C albopunctata (Boettger, 1885) e considerada fora da delimitaggo geo-
grafica abrangida pel0 presente trabalho. A sinonimizagBo de C. urbanae Bokermann, 1952
corn C leucosticta (Boulenger, 1888) 6 reafirmada. Chiasmocleis leucosticta e C. schubarti
Bokermann, 1952 siio redescritas e tr2s especies novas siio descritas.

We acknowledge Celio F. B. HADDAD (CFBH), Renato N. FEIO(UFV), Ana Maria R. COSTAand

Paulo E. VANZOLINI (MZUSP), and Sergio I? DE CARVALHO E SILVA (SPCS) for the loan of specimens
under their care; Paulo Roberto NASCIMENTO, for the drawings, and Cassia Satie Y MURAMATSU, for
the measurements of specimens; JosC I? POMBAL Jr. and Hussam ZAHER,for critically reviewing the
manuscript. Aracruz Celulose kindly permitted the searching for specimens in the preserved forests under
its care. The authors were partially supported by CNPq (Conselho Nacional de Desenvolvimento
Cientifico e Tecnologico).

BOETTGER, O., 1885. - Liste von Reptilien und Batrachiern aus Paraguay. Z. Naturw., 58: 213-248.
BOGART, J. P. & NELSON, C. E., 1976.- Evolutionary implications from karyotypic analysis of frogs of the
families Microhylidae and Rhinophrynidae. Herpetologica, 32 (2): 199-208.
BOKERMANN, W. C. A., 1952. - Microhylidae da coleggo do Departamento de Zoologia (Arnphibia -
Anura). Papkis Avulsos do Departamento de Zoologia, S6o Paulo, 10 (16): 27 1-292.
----- 1966. - Lista anotada das localidades tipo de anfbios brasileiros. Serviqo de DocumentagBo, Reitoria
da Universidade de SBo Paulo: 1- 183.
BOULENGER, G. A., 1888. - A list of batrachians from the province Santa Catharina, Brazil. Ann. Mag.
nat. Hist., (6), 1 (6): 415-417.
CARVALHO, A. L., 1954. - A preliminary synopsis of the genera of American microhylid frogs. Occ. Pap.
Mus. 2001. Univ. Michigan, 555: 1-21.
FROST, D. R. (ed.), 1985. - Amphibian species of the world. A taxonomic and geographical reference.
Lawrence, Allen Press, Inc., and The Association of Systematics Collections: i-v +1-732.
KASAHARA, S. & HADDAD, C. F. B., 1997. - Karyotypes of two Brazilian microhylid frogs of the genus
Chiasmocleis, including a new case of polyploidy. 1 Herp., 31 (1): 139-142.
MEHELY, L. VON,1904. - Investigations on Paraguayan batrachians. Ann. Mus. nut. hungarici, 2: 207-232,
1 pl.
MIRANDA-RIBEIRO, A., 1920. - 0 s engystomatideos do Museu Paulista (corn urn genero e tres especies
novos). Rev. Mus. Paulista, 12 (2): 28 1-288,2 pl.
----- 1924. - De Batrachorum generibus speciebusque duobus in collectio Musei Nationalis servatis. Bol.
Mus. nac. Rio de Janeiro, 1 (4): 255-257.
PARKER, H. W., 1934. - A monograph of the frogs of the family Microhylidae. London, Trustees of the
British Museum: i-viii + 1-208.
WALKER, C. F. & DUELLMAN, W. E., 1974. - Description of a new species of microhylid frog, Chiasmo-
cleis, from Ecuador. Occ. Pap. Mus. nat. Hist. Univ. Kansas, 26: 1-6.

Chiasmocleis leucosticta. - BRAZIL: S&o Paulo: Casa Grande (MNRJ 17564); Ribeiriio Branco
(MNRJ 17900-17904;CFBH 268,2136,2229-2232,2234,2236-2237,2239-2243,2245-2247,2251-2652;
EI 8957-8958);Ilha de Siio Sebastiiio (MZUSP 9033, holotype of Chiasmocleisurbanae); Santa Catarina:
Corupa (MNRJ 525,5582, syntypes of Nectodactylus spinulosus); Santa Luzia e Araujos (MNRJ 17563).
Chiasmocleis schubarti. - BRAZIL: Espirito Santo: Linhares, Corrego Juncado (MZUSP 2309,
holotype); Linhares, Sooretama (EI 2095; MNRJ 17548); Santa Teresa (El 8939); Aracruz (MNRJ
17538, 17539, 17542-17545, 17546, 17547, 17896-17899; CFBH 2667, 2703-2704, 2710-2713; EI 8959-
8960); Minus Gerais: Marlieria, Parque Estadual do Rio Doce (MNRJ 17883-17890; UFV 2600-2601,
2603-2608,2610,2612-26 15,26 17-2618).
Chiasrnocleis atlantica. - BRAZIL: Rio de Janeiro: Nova Iguaqu, Tingua (MNRJ 17550, holotype;
MNRJ 17549, 17551-17554, paratypes; EI 8940-8954, paratypes); SGo Paulo: Ubatuba, Picinguaba
(SPCS 5605-5607).
Chiasmocleis capixaba. - BRAZIL: Espirito Santo: Aracruz (MNRJ 17514,holotype; EI 8955-8956,
paratypes; MNRJ 17515-17529, 17532-17534, 17535-17537, 17891-17895, paratypes; CFBH 2668-2669,
2685,2693-2695,2701-2702,2714, paratypes).
Chiasmocleis carvalhoi. - BRAZIL: Rio de Janeiro: Seropedica, Horto Florestal de Santa Cruz
(MNRJ 17505, holotype; MNRJ 17480-17504,17506-175 13,17565, paratypes; EI 2096-2235,2236-2307,
2308-2399, 4381-4389, 4391-4393, 5332, 8937-8938, paratypes); Arraial do Cabo, Ilha de Cabo Frio
(MNRJ 17555-17562); Duque de Caxias (MNRJ 17566-17567, 17568-17571, 17572, 17573-17575,
17576); Niteroi (MNRJ 17577, 17578); Nova Iguaqu, Tingua (EI 4378, 4379, 4380, 4390, 4394); SGo
Paulo: Ubatuba (CFBH 1322, 1571); Ubatuba, Picinguaba (SPCS 5608-5610, 5612-5613, 5617, 5620-
5621).

Corresponding editor: W. Ronald HEYER.

O ISSCA 1997

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