ArticlePDF Available

Taxonomic Review Of Physalaemus Olfersii (Lichtenstein & Martens, 1856) With Revalidation Of Physalaemus Lateristriga (Steindachner, 1864) And Description Of Two New Related Species (Anura: Leiuperidae)

Authors:

Abstract

Article Taxonomic review of Physalaemus olfersii (Lichtenstein & Martens, 1856) with revalidation of Physalaemus lateristriga (Steindachner, 1864) and description of two new related species (Anura: Leiuperidae) Abstract Physalaemus olfersii was described by M.H.K. Lichtenstein and E.K. Martens in 1856 based on two syntypes collected by I.M. Olfers between the years 1818 and 1819. Due to the brevity of it's original description and high interpopulational variation, a taxonomic review of material assigned to the species is proposed. Morphological comparisons allowed us to associate the syntype of P. olfersii to populations distributed between the state of Espírito Santo and of São Paulo to which the southern limit is the to the municipality of São Paulo. Historical documents allowed us to define an area within Serra do Mar mountain range as the type locality of the species. Morphological and acoutics comparisons of P. olfersii from different localities in southern and southeastern Brazil revealed considerable variation among populations. Differences in call parameters and morphology, led us to conclude that three more species where under the name P. olfersii: Physalaemus lateristriga (Steindachner, 1864) comb. nov. and two new species, Physalaemus feioi sp. nov. and P. orophilus sp. nov.
Accepted by M. Vences: 20 Apr. 2010; published: 2 Jun. 2010 1
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2010 · Magnolia Press
Zootaxa 2491: 133 (2010)
www.mapress.com/zootaxa/Article
Taxonomic review of Physalaemus olfersii (Lichtenstein & Martens, 1856) with
revalidation of Physalaemus lateristriga (Steindachner, 1864) and description of
two new related species (Anura: Leiuperidae)
CARLA SANTANA CASSINI1,2, CARLOS ALBERTO GONÇALVES CRUZ1,3 & ULISSES
CARAMASCHI1,4
1 Universidade Federal do Rio de Janeiro, Museu Nacional, Quinta da Boa Vista, São Cristóvão, 20940–040 Rio de Janeiro, RJ,
Brasil.
2Corresponding author. E–mail: carlacassini@gmail.com
3 E–mail: cagcruz@uol.com.br
4E–mail: ulisses@acd.ufrj.br
Abstract
Physalaemus olfersii was described by M.H.K. Lichtenstein and E.K. Martens in 1856 based on two syntypes collected
by I.M. Olfers between the years 1818 and 1819. Due to the brevity of it’s original description and high interpopulational
variation, a taxonomic review of material assigned to the species is proposed. Morphological comparisons allowed us to
associate the syntype of P. olfersii to populations distributed between the state of Espírito Santo and of São Paulo to
which the southern limit is the to the municipality of São Paulo. Historical documents allowed us to define an area within
Serra do Mar mountain range as the type locality of the species. Morphological and acoutics comparisons of P. olfersii
from different localities in southern and southeastern Brazil revealed considerable variation among populations.
Differences in call parameters and morphology, led us to conclude that three more species where under the name P.
olfersii: Physalaemus lateristriga (Steindachner, 1864) comb. nov. and two new species, Physalaemus feioi sp. nov. and
P. orophilus sp. nov.
Key words: Amphibia; Physalaemus lateristriga comb. nov.; Physalaemus feioi sp. nov.; Physalaemus orophilus sp.
nov.; Atlantic Rain Forest Domain; Geographic distribution; Advertisement call
Introduction
The genus Physalaemus is currently composed by 43 species (Cruz et al. 2008; Frost 2010) and arranged in
seven fenetic groups: P. cuvieri group, P. signifer group, P. albifrons group, P. deimaticus group, P. gracilis
group, P. h en selii group, and P. o lfersii group (Nascimento et al. 2005). The P. olfersii group, as currently
recognized, is composed by five species: P. aguirrei Bokermann, 1966, P. insperatus Cruz, Cassini &
Caramaschi, 2008, P. soaresi Izecksohn, 1965, P. maximus Feio, Pombal & Caramaschi, 1999, and P. olfe rsii
(Lichtenstein & Martens, 1856).
Physalaemus olfersii occurs in open areas associated to forests (Dixo & Verdade 2006) and have been
considered a species broadly distributed in the states of Espírito Santo, Minas Gerais, Rio de Janeiro, São
Paulo, Paraná, and Santa Catarina (Nascimento et al. 2005; Frost 2010). It was described by Lichtenstein &
Martens (1856) as Phryniscus olfersii, based on two syntypes. The authors briefly characterized the species
and determined its type locality as “Brazil” (Lichtenstein & Martens 1856).
Morphological and natural history variations between populations of P. olfers ii had been observed by
some authors (Feio et al. 1999; Canelas & Bertolucci 2007; Cruz et al. 2008, Giaretta et al. 2009), which
suggest a possible complex of species.
CASSINI ET AL.2 · Zootaxa 2491 © 2010 Magnolia Press
Herein, we studied the variation of morphological and acoustic parameters from distinct populations of
Physalaemus olfersii. Based on these analyses, we present a redescription of this species, the revalidation of
Nattereria lateristriga Steindachner, 1864 under a new combination, and descriptions of two new species.
Materials and methods
Specimens examined are deposited in: AL–MN (Adolpho Lutz collection, housed in the Museu Nacional, Rio
de Janeiro, Brazil), CFBH (Célio F.B. Haddad collection, Departamento de Zoologia, Universidade Estadual
Paulista, Rio Claro, São Paulo, Brazil), CHUFSC (Universidade Federal de Santa Catarina, Florianópolis,
Santa Catarina, Brazil), DZSJRP (Departamento de Zoologia, Universidade Estadual Paulista, São José do
Rio Preto, São Paulo, Brazil), EI (Eugenio Izecksohn collection, Universidade Federal Rural do Rio de
Janeiro, Seropédica, Rio de Janeiro, Brazil), MCNAM (Museu de Ciências Naturais, Pontifícia Universidade
Católica de Minas Gerais, Belo Horizonte, Minas Gerais, Brazil), MCT–PUCRS (Museu de Ciência e
Tecnologia, Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil),
MNRJ (Museu Nacional, Rio de Janeiro, Brazil), MHNCI (Museu de História Natural Capão da Imbuia,
Curitiba, Paraná, Brazil), MZUFV (Museu de Zoologia João Moojen de Oliveira, Universidade Federal de
Viçosa, Minas Gerais, Brazil), MZUSP (Museu de Zoologia, Universidade de São Paulo, Brazil), and ZUEC
(Museu de História Natural Prof. Adão José Cardoso, Universidade Estadual de Campinas, São Paulo, Brazil).
Specimens examined are referred in the Appendix.
Analysis of one syntype of Phryniscus olfersii from the Berlin Museum of Natural History (Museum für
Naturkunde of the Humbold–University in Berlin; ZMB 3375) was done by photographs conceded by Rainer
Günther. The type specimens of Nattereria lateristriga and Phyllobates glandulosus, from the Vienna Natural
History Museum (NMW 4329 and NMW 16520, respectively) were analyzed by photographs conceded by
Franz Tiedemann and Heinz Grillitsch.
Morphological and morphometric data were obtained from 591 preserved specimens of the Physalaemus
olfersii species group. Abbreviations used for the measurements of adult specimens are: SVL (snout-vent
length), HL (head length), HW (head width), ED (eye diameter), IOD (interorbital distance), UEW (upper
eyelid width), END (eye-nostril distance), NSD (nostril-snout distance), IND (internarial distance), HAL
(hand length), TL (thigh length), SL (shank length), FL (foot length, including tarsus), IML (inner metatarsal
tubercle length), OML (outer metatarsal tubercle length), IOMD (inner-outer metatarsal tubercle distance),
and TIMD (tarsal-inner metatarsal tubercle distance). The measurements SVL, HL, HW, THL, TBL, and FL
were taken with calipers (0.01 mm, Mytutoyo), whereas other measurements were made with an ocular grid.
Morphological terminology follows Heyer et al. (1990), with exception of tibial length (instead of shank
length). Discriminant analysis (see coefficients on Table 1) was employed to study the variation between
defined groups of populations (morphological units).
Calls were recorded using a 4.5 mm/s rotation cassette recorder Panasonic model RQ L31 and cassette
recorder Marantz with a Sennheiser ME66–K6 microphone set. Recordings were digitized and analyzed using
the software Cool Edit 2000 1.1 (sample frequency: 32000 Hz; 16 bits resolution, stereo). Sonograms and
oscilograms were obtained using Sound Ruler V4. The acoustics characteristics examined were: call duration
(measured from the beginning of the first pulse to the end of the last one), call structure (harmonic, non-
harmonic) and frequency modulation (ascendant, descedent or absent), number of notes, pulses, pulse rate,
and dominant frequency. Terminology adopted for the description of acoustic properties of calls follows
McLister et al. (1995) and Gerhardt and Huber (2002).
Zootaxa 2491 © 2010 Magnolia Press · 3
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
TABLE 1—Coefficients of the first and second canonical variables (CV1 and CV2) of each morphometric variable.
Results
Morphological units
After morphological analysis of P. olfersii populations, four distinct morphological units were diagnosed: 1
“Serra do Espinhaço”, from Serra do Espinhaço mountain range; 2“Serra da Mantiqueira”, from Serra da
Mantiqueira mountain range; 3 “Serra do Mar”, from the municipality of Santa Teresa, state of Espírito
Santo, state of Rio de Janeiro to the municipality of São Paulo and central region of the state of São Paulo, in
the municipality of Botucatu; and 4 “Southern”, from southern of the state of São Paulo and states of Paraná
and Santa Catarina.
Four morphological characters separate the units: tarsal tubercle (present/weakly developed/absent), tarsal
fold (present/weakly developed/absent), lateral blak stripe on flanks (well defined/ weakly defined), white line
on dorsolateral fold (well defined/ weakly defined/absent), and annulus of tympanum (weakly marked/
indistinct).
The “Serra do Espinhaço” morphological unit is characterized by the presence of tarsal fold; absence of
tarsal tubercle; lateral black stripe weakly defined on flanks; white line on dorsolateral fold absent; and
annulus of tympanum weakly marked. The “Serra da Mantiqueira” morphological unit, by the presence of
tarsal fold; presence of tarsal tubercle; lateral black stripe weakly defined on flanks; white line on dorsolateral
fold weakly defined; and annulus of tympanum weakly marked. The “Serra do Mar” morphological unit is
characterized by the presence of tarsal tubercle weakly developed; tarsal fold weakly developed or absent;
lateral black stripe well defined on flanks; white line on dorsolateral fold well defined; and annulus of
tympanum indistinct. The “Southern” morphological unit is similar to the “Serra do Mar” morphological unit,
but tarsal tubercle is absent; white line on dorsolateral fold is weakly marked or absent; and the annulus of
tympanum is weakly marked. Morphological data and acoustic parameters between the species are shown in
Table 2 and 3 respectively (see also taxonomic accounts).
Morphometric variables Males Females
CV1 CV2 CV1 CV2
SVL 0,593585 0,200425 0,605786 0,17702
HL 0,350542 0,111240 -0,460956 0,51879
HW -0,248742 -0,150437 0,564294 0,71027
HAL 0,225402 0,172993 -0,790621 -1,17108
TL 0,350173 -0,208584 -0,476849 -0,08087
SL -0,245323 0,322937 -0,090795 0,16800
FL -0,038383 -0,135332 0,028254 -0,03400
ED 0,151044 -0,427499 0,084406 0,79402
END -0,160708 -0,337474 0,198821 0,58569
NSD 0,184751 -0,035780 -0,513378 0,06622
UEW -0,066337 0,260594 -0,282233 -0,47501
IOD -0,088025 -0,066127 -0,175539 -0,37325
IND -0,095896 0,354467 0,043834 -0,93383
IML 0,090182 0,469512 0,370677 0,02484
OML 0,110429 -0,221272 -0,394493 0,05780
IOMD 0,147205 0,192876 0,312828 0,50875
TIMD -0,317748 0,838561 0,572174 0,09189
CASSINI ET AL.4 · Zootaxa 2491 © 2010 Magnolia Press
TABLE 2—Morphological comparison between species of the Physalaemus olfersii group.
Discriminant analysis from 17 morphometric variables of males and females of the proposed
morphological units were employed (Fig. 1). The canonical plot graphs of male and female data show almost
the same discrimination among the units, except for females from “Serra do Espinhaço” which show a large
ellipse encircling due to the low sample (n= 4). In both male and female graphs, the morphological units
“Serra do Mar” and “Southern” are morphometrically more similar to each other than they are to “Serra da
Mantiqueira”. The combination of the first two functions corresponded to 93.19% and 94.99% of the entire
variation, males and females respectively. First and second canonical variables of each morphometric
parameter are shown on Table 1.
Physalaemus olfersii type locality
One of the Physalaemus olfersii syntypes is located at the Zoological Museum of Berlin (ZMB 3375, defined
here as the lectotype). There are no register on documents of that museum on whereabouts of the second
specimen (Rainer Günther, pers. comm.). The syntypes were collected by Ignaz Maria Franz von Olfers, date
of collection not available on data of that Museum (Rainer Günther, pers. comm.)
Ignaz Maria Franz Olfers (1793–1871), German naturalist, traveled to Brazil in 1816 as a diplomatic; in
the year of 1818 he traveled with Frederich Sellow (1789–1831?) during one year along his expedition
through Brazil (Papavero 1971; Straube & Scherer–Netto 2001). During this time, they collected in the states
of Rio de Janeiro, Minas Gerais and São Paulo (Fig. 2).
Species Annulus of tympanun Loreal region Lateral black stripe
Physalaemus olfersii Indistinct oblique well defined from the anterior
region of the eye to inguinal region
P. lateristriga weakly developed vertical well defined from the anterior
region of the eye to inguinal region
P. feioi weakly developed oblique well defined from the anterior
region of the eye to shoulder
P. orophilus weakly developed oblique well defined from the anterior
region of the eye to shoulder
Species White line on canthus rostralis,
border of upper eyelid and
dorsolateral fold
Tarsal
tubercle Tarsal fold
P. olfersii weakly developed on canthus rostralis and
well marked on upper eyelid
and on the dorsolateral fold
weakly
developed weakly developed or absent
P. lateristriga absent on canthus rostralis and
the of upper eyelid, and
weakly marked on the dorsolateral fold absent weakly developed or absent
P. feioi weakly marked on the canthus rostralis,
upper eyelid and on dorsolateral fold developed developed
P. orophilus weakly marked on the canthus
rostralis and on upper eyelid,
absent on the dorsolateral fold absent developed
Zootaxa 2491 © 2010 Magnolia Press · 5
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
FIGURE 1: Canonical analyses of 17 morphometric variables of males (A) and females (B) of the Physalaemus olfersii
mophological units groups proposed in the present study, with the respective 95% confidence limits ellipse for each
group.
CASSINI ET AL.6 · Zootaxa 2491 © 2010 Magnolia Press
FIGURE 2: Localities visited by Frederich Sellow and Ignaz Maria von Olfers during their journey in 1818 [State of Rio
de Janeiro: 1, 21, Rio de Janeiro. State of Minas Gerais: 2, Barbacena; 3, Ouro Branco; 4, 13, Ouro Preto; 5, Itambé do
Mato Dentro; 6, Ribeirão do Galheiro; 7, Serra da Moeda, municipality of Moeda; 8, Itabira; 9, Serra da Cachoeira,
municipality of Ouro Preto; 10, Serra do Itacolomi, municipality of Ouro Preto; 11, Serra da Piedade, municipality of
Caeté; 12, Serra do Caraça, municipality of Catas Altas; 14, São João Del Rei. State of São Paulo: 15, Jundiaí; 16, Itu;
17, 19, São Paulo; 18, Fazenda Ipanema, Municipality of Sorocaba; 20, Santos].
Three morphological units defined by us are distributed within the area traveled through by Olfers and
Sellow (“Serra do Espinhaço”, “Serra da Mantiqueira”, and “Serra do Mar” morphological units). Since we
could verify from the photographs that the lectotype of P. olfersii presents tarsal tubercle weakly developed;
absence of tarsal fold; lateral black stripe well defined on flanks; white line on dorsolateral fold well defined;
and annulus of tympanum indistinct, we can associate it with the populations of the "Serra do Mar"
morphological unit. Thus, these correspond to P. olfersii and the species’ type locality should be the
intersection of its geographic distribution within the area traveled by Olfers and Sellow, which is the Serra do
Mar mountain range, from the states of Rio de Janeiro to São Paulo.
Physalaemus lateristriga comb. nov. and two new species
Nattereria lateristriga, described by Steindachner (1864), is presently associated to P. olfersii. The type
locality of the species is pointed as “Curitiba, in forest land” (Steindachner 1864; Bokermann 1966).
Specimens analyzed (see Appendix) of the “Southern” morphological unit agree with the description of
Nattereria lateristriga (Steindachner, 1864). We recorded calls from a population of São José dos Pinhais,
which is the nearest city from Curitiba, distant about 10 km. The original description of Nattereria lateristriga
was based on specimen NMW 4329:1 (Fig. 3) and NMW 4329:2, females, and here we designate the
specimen NMW 4329:1 as the lectotype for Physalaemus lateristriga comb. nov.
Zootaxa 2491 © 2010 Magnolia Press · 7
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
FIGURE 3: Steindachner (1864) plate of the type of Nattereria lateristriga.
Finally, we recognize the morphological units “Serra da Mantiqueira” and “Serra do Espinhaço” as new
species: Physalaeus feioi sp. nov. and P. orophilus sp. nov., respectively.
Taxonomic Accounts
Physalaemus olfersii (Lichtenstein & Martens, 1856)
(Figs. 4, 5, 6, 7)
Phryniscus olfersii Lichtenstein & Martens, 1856.
Phrynidium olfersi—Cope, 1867.
Paludicola olfersi—Peters, 1882.
Physalaemus olfersi—Parker, 1927.
Lectotype. ZMB 3375 (Fig. 4), by present designation, female (SVL 29.4; TBL 12.3; HL 8.1; HW 6.5 mm),
collected in an undetermined locality in the Serra do Mar mountain range, from the state of Rio de Janeiro to
São Paulo (Fig. 8), by Ignaz Marian Franz Olfers and Frederich Sellow, between the years of 1818 to 1819.
Diagnosis. (1) Body length small to moderate (SVL 26.236.4 mm in males, 22.441.1 mm in females);
(2) texture of skin shagreened; (3) head longer than wide; (4) snout pointed to subelliptical in dorsal view and
acute in lateral profile; (5) lateral black stripe, from the posterior corner of the eye to the inguinal region; (6)
inguinal gland poorly evident; (7) oblique white line from the posterior corner of the eye to the arm insertion;
(8) white line weakly developed on canthus rostralis and well marked on the border of the upper eyelid and
CASSINI ET AL.8 · Zootaxa 2491 © 2010 Magnolia Press
on the dorsolateral fold; (9) annulus of tympanum indistinct; (10) vocal sac developed, dark gray; (11) tarsal
tubercle weakly developed; (12) tarsal fold weakly developed or absent; (13) advertisement call composed by
one multipulsed note (pulse rate 135 to 156 pulses per second); (14) mean dominant frequency 1.92 kHz.
FIGURE 4: Physalaemus olfersii, lectotype (ZMB 3375, adult female). A—dorsal, B—ventral, C—lateral views, and
D—foot (photographs conceded by Rainer Günther).
Comparison with other species. Physalaemus olfersii is distinguished of P. aguirrei and P. insperatus by
the absence of a white line outlining the inferior margin of the mandible and by the absence of the
supratympanic fold (both present in those species). Additionaly, P. olfersii is distinguished of P. aguirrei by
the presence of an oblique white line from the posterior corner of the eye to the arm insertion (absent in P.
aguirrei); ventral region light brown with brown and white dispersed blotches (ventral region light brown,
with white blotches, except on the ventral surface of the thigh that is uniformly white in P. aguirrei); non–
harmonic physical structure of the call (harmonic in P. aguirrei; Pimenta & Cruz 2004); and mean dominant
frequency 1.92 kHz (3.1 kHz in P. aguirrei; Pimenta & Cruz 2004). From P. insperatus, by the vocal sac not
expanded laterally (expanded laterally in P. insperatus); and ventral region light brown with brown and white
dispersed blotches (ventral region light brown with white blotches, except on the ventral surface of the thigh
that is uniformly white in P. insperatus).
Physalaemus olfersii is distinguished of P. feioi sp. nov. and P. orophilus sp. nov. by the presence of a well
marked white line on the dorsolateral fold (weakly marked in P. feioi sp. nov. and absent in P. orophilus sp.
nov.); vocal sac uniformly dark gray (with concentrated gray blotches in P. feioi sp. nov. and P. orophilus sp.
nov.); and mean dominat frequency of 1.92 kHz (2.56 kHz in P. feioi sp. nov. and 3.18 kHz in P. orophilus sp.
nov.).
Zootaxa 2491 © 2010 Magnolia Press · 9
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
Physalaemus olfersii is distinguished from P. lateristriga comb. nov.; by the loreal region oblique
(vertical in P. lateristriga comb. nov.), annulus of tympanum indistinct (distinct, weakly developed in P.
lateristriga comb. nov.); and tarsal fold weakly developed (absent in P. lateristriga comb. nov.). Moreover, it
is distinguished from P. lateristriga comb. nov. by the tibia light gray or light brown weakly marbled, except
on dorsal surface (dark gray or dark brown marked marbled in P. lateristriga comb. nov.); outer metatarsal
tubercle rounded (conic in P. lateristriga comb. nov.); and advertisement call with lower pulse rate (171 to
202 pulses per second in P. lateristriga com. nov.).
Physalaemus olfersii has smaller body length than P. maximus (SVL 41.146.5 mm in males, 42.251.5
mm in females of P. maximus); and mean dominant frequency of 1.92 kHz (two values: 0.73 or 0.78 KHz in P.
maximus; Baêta et al. 2007). Finally, P. olfersii present larger body length than P. soaresi (SVL 17.922.8 mm
in males, 19.921.52 mm in females of P. soaresi); oblique white line from the posterior corner of the eye to
the arm insertion (absent in P. soaresi); tarsal tubercle weakly developed (developed in P. soaresi); tarsal fold
absent or weakly developed (developed in P. soaresi); developed, dark gray vocal sac (weakly developed and
light brown in P. soaresi); head longer than wide (wider than long in P. s oaresi); and advertisement call with
mean dominant frequency of 1.92 kHz (3.3 kHz in P. soaresi; Weber et al. 2005).
Description. Body moderately robust (Figs. 5, 6); head longer than wide; snout pointed to subelliptical in
dorsal view, acute in lateral profile; nostrils elliptical, located and expanded laterally, closer to tip of snout
than to eye; canthus rostralis distinct; loreal region oblique; eye slightly protuberant; annulus of tympanum
and supratympanic fold indistinct; parotoid gland not developed; presence of a dorsolateral fold from the
posterior corner of eye, delimiting the dorsal region from the flank, ending at the anterior margin of the
inguinal region; vocal sac subgular, well developed, extending to the chest; choanae small, rounded, well
distinguished from each other; tongue narrow, long, free posteriorly; maxillary and premaxillary teeth visible;
vomerine teeth absent. Arms short, slender; forearms slightly more robust and as short as arms; fingers thin,
long, not fringed; finger lengths I < II IV < III; large nuptial pad divided, one part covering from 1/2 to 2/3
of the inner metacarpal tubercle and other part enclosing the thumb, except the inner side; subarticular
tubercles large, elliptical; outer metacarpal tubercle medium–sized, elliptical; inner metacarpal tubercle large,
elliptical, approximately three times the size of the outer metacarpal tubercle; absence of supernumerary
tubercles; fingers tip slightly expanded. Legs moderately robust; tibia slightly longer than thigh; sum of tibia
and thigh lengths shorter than SVL; tarsal fold absent; toes thin, long, not fringed; toe lengths I < II < III = V
< IV; subarticular tubercles large, protruding; inner metatarsal tubercle large, elliptical; outer metatarsal
tubercle medium–sized, ovoid; supernumerary tubercles absent; toes tip expanded. Dorsal surface shagreened
and ventral surfaces smooth. Ventral disc absent.
Color in life. Dorsal color pattern varying from light brown (Fig. 7) to gray with small, irregular, and
disperse scattered brown blotches; dorsal median line extending on posterior 2/3 of the dorsum; interorbital
white line interrupted at midway; loreal region dark brown with some dispersed white dots; a black stripe
from the postorbital region to the anterior margin of the inguinal gland; a white line weakly developed on
canthus rostralis and well marked on the border of the upper eyelid and on the dorsolateral fold; a dark brown
line bordering the internal margin of the ventral surface of the arms; a rectangular blotch on dorsum of the
forearm; transversal bars on dorsum of thighs; an irregular blotch on internal side of tibia, near the heel; gular
region dark gray with white dots; white dots on the anterior region of the abdomen; uniformly light brown on
the posterior region of the abdomen and ventral proximal surfaces of thighs; nuptial pads brown; color of the
inguinal gland not distinguishable of the color pattern of the dorsum; posterior region of the abdomen and
tubercles of palmar and plantar surfaces light red.
Color in preservative (70% ethanol). General color pattern maintained in preservative. Dorsal surface
light brown or gray with a light gray blotch on the center; tubercles of plantar and palmar surfaces; and
abdomen, white.
Variation (n = 151). Males are slightly smaller than females. Range, mean, and standard deviation of
SVL of males and females are in table 4. Few specimens observed have a mucronate snout in dorsal view
(sensu Heyer et al. 1990). Tarsal fold is absent in most specimens analyzed, although some specimens have a
tarsal fold weakly developed.
CASSINI ET AL.10 · Zootaxa 2491 © 2010 Magnolia Press
FIGURE 5: Physalaemus olfersii (MNRJ 56120) from municipality of Teresópolis, state of Rio de Janeiro. A, dorsal,
and B, ventral views. Bar = 20 mm.
FIGURE 6: Physalaemus olfersii (MNRJ 56120) from municipality of Teresópolis, state of Rio de Janeiro. A—dorsal,
and B—lateral view of head; C—hand, and D—foot. Bars = 10 mm.
Zootaxa 2491 © 2010 Magnolia Press · 11
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
FIGURE 7: Physalaemus olfersii (DZSJRP 13958) from Parque Nacional da Serra da Bocaina, municipality of São José
do Barreiro, state of São Paulo. Photograph by Diogo B. Provete.
FIGURE 8: Geographic distribution of Physalaemus olfersii, P. lateristriga comb. nov., P. feioi sp. nov., and P.
orophilus sp. nov.
CASSINI ET AL.12 · Zootaxa 2491 © 2010 Magnolia Press
FIGURE 9: Physalaemus olfersii advertisement call: A—oscilogram, B—audiospectrogram, C—power spectrum, D—
oscillogram of a cut of the same call evidencing pulsed structure, and E—oscillogram of a cut of the same call
evidencing the subpulsed structure. Specimen MNRJ 50312, collected at the municipality of Teresópolis (22°27'S;
42°58'W), state of Rio de Janeiro, on 20/XI/2007. Air temperature: 23°C.
Advertisement call. Advertisement calls (n = 92 calls) were obtained from eight males (MNRJ 50309–
50314; MNRJ 56119–56120) from the Parque Nacional da Serra dos Órgãos, municipality of Teresópolis,
state of Rio de Janeiro, Brazil. The advertisement call (Fig. 9, Table 3) is composed by one single multipulsed
note, each pulse composed by 2 subpulses (which can be fused or not on sonogram along the call), non–
harmonic structure, without frequency modulation; mean duration of 4.04 seconds (SD = 0.4, range = 2.6 to
5.1 seconds); pulse duration of 6 to 7 miliseconds, pulse rate ranges from 135 to 156 pulses per second (mean
= 149, SD = 5); mean duration of time between calls 2.07 seconds (SD = 0.8, amplitude = 1.072.64 seconds)
and the mean value of the dominant frequency 1.92 kHz (SD = 0.33, amplitude = 1.382.59 kHz).
Remarks. Phyllobates glandulosus, described by Steindachner (1867), was synonymized with
Physalaemus olfersii by Bokermann (1966b). According to Bokermann (1966b), the type locality of
Phyllobates glandulosus is “Brazil, probably Serra da Estrela, Inhomirim, Rio de Janeiro.”
Steindachner (1867) described the amphibians collected by the “Fregatte Novara” expedition through the
years 1857 to 1859, which ported in Brazil from August 5th to August 31th 1857. Zoologists from “Fregatte
Novara” expedition (Geroge Frauenfeld and John Zelebor) collected at the municipalities of Rio de Janeiro
and Petrópolis (Scherzer 1861). “Serra da Estrela” mentioned by Bokermann (1966b) and Scherzer (1861) is a
chain of mountains within the Serra dos Órgãos mountain range where the municipality of Petrópolis is
located. In the midway of the municipalities of Rio de Janeiro and Petrópolis, at the base of Serra da Estrela
mountains, is located “Inhomirim” mentioned by Bokermann (1966b). Scherzer (1861) didn’t specified where
the zoologists of “Fregatte Novara” had collected the Phyllobates glandulosus type, and it could be anywhere
in the municipalities of Rio de Janeiro, Petrópolis, Inhomirim or in the way from one to another (Figure 10).
In this area, are known to occur nowadays Physalaemus cuvieri, P. maculiventris, P. marmoratus, P. olfersii,
and P. signifier.
Steindachner (1867) described the specimen as having a triangular head; snout very short; tongue long
and rounded; evident tympanum; general color pattern of the body light brown; two turbercles well developed
on metacarpus and two smaller on metatarsus and small numerous tubercles on tarsus and on the sole of the
foot. Additionally, we could observe on the pictures of the type specimen (NMW 16520; Fig. 11) the presence
of large inguinal glands associated with dark ocellus; ventral surface homogeneous light brown without
blotches.
Zootaxa 2491 © 2010 Magnolia Press · 13
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
TABLE 3: Acoustic parameters of species of the Physalaemus olfersii group. olf = P. olfersii; lat = P. lateristriga; feioi = P. feioi; orop = P. orophilus; agui = P.
aguirrei; soar = P. soaresi; max = P. maximus. Values of parameters: amplitude (mean±standard deviation).Mult = multipulsed note. *What Baêta et al. (2007)
call “harmonics” are actually “side bands” sensu Vielliard (1993).
Species
olf lat feioi orop agui soar max
Locality of
populations analyzed
Teresópolis
RJ
São José dos Pinhais
PR
Viçosa and Araponga
MG
Mariana
MG
Prado
BA
Seropédica
RJ
Ouro Preto
MG
Call duration
(seconds)
2.6–5.1
(4.04±0.4)
2.3–5.5
(3.6±0.7)
2.5–5.6
(4.3±0.7)
3.1–5.2
(3.9± 0.6)
0.21–0.25
(0.23±0.01)
1.34–2.40
(1.83±?)
1.09–2.31
(2.01±0.26)
Structure NH NH NH NH H NH NH*
Frequency
modulation No No
ascendent along the call;
descendent at the end of the call ascendent descedent ascendent weak descedent
Notes 1 1 1 1 1 1 1
Pulses 519–733
(603±52)
404–990
(672±235)
30-75
(45±4)
5687
(78±5) - mult mult
Pulse duration
(ms) 6–7 4–5 40–45 42–50 - - -
Sub-pulses 2 2 1419 8-11 - - -
Pulse rate
(pulses per second)
135156
(149±5)
171–202
(183±10)
8–16
(12±1.36)
16–19
(18±0.76) - - -
Dominant
frequency
1.38–2.59
(1.92±0.33)
0.75–2.08
(1.68±0.23)
2.31–2.75
(2.56±0.0075)
3.0–3.39
(3.18±0.09 ) 3.10 3.3 0.732 or 0.775
n of calls 92 70 110 124 55 6 -
n of males 8 6 11 6 1 - -
References Present work Present work Present work Present work Pimenta & Cruz,
2004
Weber et al.
2005
Baêta et al.
2007
CASSINI ET AL.14 · Zootaxa 2491 © 2010 Magnolia Press
FIGURE 10: Area travelled by zoologists of the "Fregatte Novara" expedition (Geroge Frauenfeld and John Zelebor)
from August 5th to August 31th 1857.
Zootaxa 2491 © 2010 Magnolia Press · 15
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
FIGURE 11: Phyllobates glandulosus, holotype (NMW 16520). A—dorsal, B—ventral, and C—lateral views
(photographs conceded by Franz Tiedemman and Heinz Grillitsch). Label data: "Brasilien, Novara Expedition, 1857-
1859".
The presence of inguinal glands associated with dark ocellus and evident tympanum, distinguish the type
of Phyllobates glandulosus from Physalaemus marmoratus (Nascimento et al. 2006) and P. olfersii (present
work). Bokermann (1962) distinguished P. signifer from P. maculiventris by the presence of supranumerary
tubercles on the ventral surface of the foot in P. signifer and absence on P. maculiventris. In addition, he
figured the lateral view of these species and the tympanum annulus is more evident in P. signifer. Thus, from
those species which are distributed within the area where Geroge Frauenfeld and John Zelebor possibly
collected the type of Phyllobates glandulosus the one that better fits with the Steindachner’s descrition and
photograph of the type is Physalaemus signifer. Therefore, we transfer Phyllobates glandulosus from the
synonymy of Physalaemus olfersii to the synonymy of Physalaemus signifer (Girard, 1853).
Geographical distribution. Physalemus olfersii is distributed in the Atlantic Rain Forest Domain and its
influence areas, from the municipality of Santa Teresa, state of Espírito Santo, southern region of the state of
Minas Gerais to the municipality of São Paulo and central region of the state of São Paulo, in the municipality
of Botucatu (Fig. 8).
CASSINI ET AL.16 · Zootaxa 2491 © 2010 Magnolia Press
Physalaemus lateristriga (Steindachner, 1864), new combination
(Figs. 12, 13, 14, 15)
Nattereria lateristriga Steindachner, 1864.
Physalaemus personatus—Fitzinger in Steindachner, 1864.
Physalaemus olfersii—Nascimento, Caramaschi & Cruz, 2005 (part).
Physalaemus olfersii—Cruz, Cassini & Caramaschi, 2008 (part).
Lectotype. NMW 4329:1 (Fig. 12), by present designation, female (SVL 28.7; HL 12.2; HW 9.0; ED 0.4;
UEW 0.2; IOD 0.3; IND 0.3; END 0.3; TL 15.3; SL 14.; FL 21.0), collected at Curitiba, state of Paraná,
Brazil, on 09 November 1820, collector unknown.
Paralectotype. NMW 4329:2, male, collected with the lectotype.
Diagnosis. (1) Body length small to medium (SVL 30.3–37.8 mm in males, 32.6–40.3 mm in females);
(2) skin texture shagreened; (3) head longer than wide; (4) snout pointed in dorsal view, protruding to acute in
lateral profile; (5) lateral black stripe, from the posterior corner of the eye to the inguinal region; (6) inguinal
gland developed, not associated with dark ocellus; (7) oblique white line from the posterior corner of the eye
to the arm insertion; (8) white line absent on cantus rostralis and the border of upper eyelid, and weakly
marked on the dorsolateral fold to the inguinal region; (9) annulus of tympanum weakly developed; (10) vocal
sac developed; (11) tarsal tubercle absent; (12) tarsal fold absent or weakly developed; (13) advertisement call
composed by one multipulsed note (pulse rate 171–202 pulses per second ); (14) mean dominant frequency
1.68 kHz.
Comparison with other species. Physalaemus lateristriga comb. nov. is distinguished from P. aguirrei
and P. inspera tus by the absence of a white line outlining the inferior margin of the mandible and by the
absence of the supratympanic fold (both present in those species); by the presence of an oblique white line
from the posterior corner of the eye to the arm insertion (absent in P. aguirrei and in P. insperatus); and
ventral region light brown with brown and white dispersed blotches (ventral region light brown with white
blotches, except on the ventral surface of the thigh that is uniformly white in P. aguirrei and in P. insperatus).
Additionaly, P. lateristriga comb. nov. com. nov. is distinguished from P. aguirrei by non–harmonic physical
structure of the call (harmonic in P. aguirrei; Pimenta & Cruz 2004) and mean dominant frequency 1.68 kHz
(3.1 kHz in P. aguirrei; Pimenta & Cruz 2004). From P. insperatus, it is distinguished by the vocal sac not
expanded laterally (expanded laterally in P. insperatus).
Physalaemus lateristriga comb. nov. is distinguished from P. feioi sp. nov. and P. orophilus sp. nov. by the
vocal sac uniformly dark gray (with gray blotches in P. feioi sp. nov. and P. orophilus sp. nov.). Additionaly, it
is distinguished from P. feioi sp. nov. by the ventral surface with scattered blotches (blotches concentrated on
gular region in P. feioi sp. nov.), tarsal tubercle weakly developed or absent (developed in P. feioi sp. nov.),
and advertisement call with pulse rate from 171–202 pulses per second (8–16 pulses per sencond in P. feioi sp.
nov. and 16–19 pulses per second in P. orophilus sp. nov.). Moreover, it is distinguished from P. orophilus sp.
nov. by its smaller size (SVL 21.1–24,8 mm in males, 25.5–28.1 mm in females of P. orophilus sp. nov.).
Physalaemus lateristriga comb. nov. presents smaller body length than P. maximus (SVL 41.1–46.5 mm
in males, 42.2–51.5 mm in females of P. maximus), and advertisement call with mean dominant frequency
1.68 kHz (two values: 0.73 or 0.78 kHz in P. maximus; Baêta et al. 2007).
Physalaemus lateristriga comb. nov. is distinguished from P. o lf ersii by the presence of a white line on
canthus rostralis and on the margin of the upper eyelid, which is thicker and well marked (white line throught
canthus rostralis and on the margin of upper eyelid thin and weakly marked in P. olfersii); by the annulus of
tympanum distinct, weakly developed (indistinct in P. o lf er si i); tibia gray or brown marked marbled (weakly
marbled, except on dorsal surface in P. olfersii), tarsal fold absent (weakly developed in P. olfersii); and
advertisment call with pulse duration of 4 to 5 miliseconds (6 to 7 milisencond in P. olfersii) higher pulse rate
(131 to 159 pulses per second in P. olfersii).
Physalaemus lateristriga comb. nov. presents larger body length than P. soaresi (SVL 17.9–22.8 mm in
males, 19.9–21.5 mm in females of P. soaresi); an oblique white line from the posterior corner of the eye to
the shoulder (absent in P. s oa res i); tarsal tubercle weakly developed or absent (developed in P. soaresi);
Zootaxa 2491 © 2010 Magnolia Press · 17
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
developed and dark gray vocal sac (vocal sac weakly developed and light brown in P. soaresi); head longer
than wide (wider than long in P. soaresi); and advertisement call with mean dominant frequency 1.68 kHz (3.3
kHz in P. soaresi; Weber et al. 2005).
FIGURE 12: Physalaemus lateristriga comb. nov., lectotype (NMW 4329:1, adult female). A—dorsal, and B—ventral
views (photographs conceded by Franz Tiedemman and Heinz Grillitsch).
Description. Body robust (Figs. 13, 14); head longer than wide; snout pointed in dorsal view, protruding
in lateral profile; nostrils elliptical, not protuberant, located and oriented laterally, closer to tip of snout than to
eye; canthus rostralis distinct; loreal region vertical; eye slightly protuberant; tympanum weakly distinct;
supratympanic fold absent; parotoid glands well developed; presence of a dorsolateral fold from the posterior
corner of eye, delimiting the dorsal region from the flank, and ending at the anterior margin of the inguinal
gland; vocal sac subgular, well developed, extending to the chest; choanae small, rounded, well separated
from each other; tongue narrow, long, free posteriorly; maxillary and premaxillary teeth visible; vomerine
CASSINI ET AL.18 · Zootaxa 2491 © 2010 Magnolia Press
teeth absent. Arms short, slender; forearms slightly more robust and as short as arms; fingers thin, very long,
weakly fringed; fingers length I < II = IV < III; extensive nuptial pad divided, one part covering from 1/2 to 2/
3 of the inner metacarpal tubercle and other part enclosing the thumb, except the inner side; subarticular
tubercles single, well developed, rounded; outer metacarpal tubercle medium–sized, rounded, covering 1/3 of
carpal region; inner metacarpal tubercle large, elliptical, covering 1/2 of carpal region, approximately two
times the size of the outer metacarpal tubercle; presence of supernumerary tubercles; fingers tip slightly
expanded. Legs moderately robust; tibia slightly longer than thigh; sum of tibia and thigh lengths shorter than
SVL; tarsal fold absent; toes thin, long, slightly fringed; toes length I < II < III V < IV; subarticular tubercles
protruding, conic; inner metatarsal tubercle medium–sized, elliptical; outer metatarsal tubercle small, ovoid;
supernumerary tubercles weakly developed; toes tip slightly expanded. Dorsal surface shagreened and ventral
surface smooth. Ventral disc absent.
FIGURE 13: Physalaemus lateristriga comb. nov. (CFBH 21169) from the municipality of São José dos Pinhais, state
of Paraná. A—dorsal, and B—ventral views. Bar = 20 mm.
Color in life. The color pattern vary from gray to light brown (Fig. 15); dorsal median line extending on
posterior third of the dorsum; loreal region dark brown with some white dots dispersed; a black stripe under
the dorsolateral fold, extending from the postorbital region to the inguinal region; a white line well marked
and thicker on canthus rostralis and border of upper eyelid, and marked on the dorsolateral fold; a rectangular
dark brown blotch on dorsum of the forearm; transversal brown bars on dorsum of thighs; an irregular brown
blotch on internal side of tibia, near to the heel; gular region dark gray with small white dots; white dots on the
anterior region of the abdomen, large white blotches on the posterior region of the abdomen; proximal region
of ventral surfaces of limbs light brown; dark brown line laterally to the ventral surface of the arms; brown
nuptial pads; inguinal gland not distinguished from the color pattern of the dorsum; posterior region of the
abdomen and tubercles of plantar and palmar surfaces red.
Color in preservative (70% ethanol). General color pattern maintained in preservative. Dorsal surface
with a light gray blotch on the center; tubercles of plantar and palmar surfaces, and abdomen white.
Zootaxa 2491 © 2010 Magnolia Press · 19
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
TABLE 4: Mean (X), standard deviation (SD), and range of snout–vent length (SVL) of males and females of the
species of the Physalaemus olfersii group. Measurements in mm.
FIGURE 14: Physalaemus lateristriga comb. nov. (CFBH 21169) from the municipality of São José dos Pinhais, state of
Paraná. A—dorsal, and B—lateral view of head, C—hand, and D—foot. Bars = 10 mm.
Species Males Females
XSD Range XSD Range
P. aguirrei (n=86) 24.0 2.26 19.8 –27.3 24.6 2.19 21.7–28.2
P. feioi (n=151) 25.5 1.43 21.2–29.2 27.9 1.84 23.5–31.2
P. insperatus (n=6) 23.9 0.17 22.2–25.3 26.7 0.05 26.6–26.7
P. lateristriga (n=128) 32.2 1.35 30.3–37.8 35.4 2.25 32.6–40.3
P. maximus (n=29) 45.8 2.53 42.2–51.5 42.4 1.80 41.1–43.7
P. olfersii (n=151) 31.1 2.32 26.2–36.4 31.9 3.66 22.4–41.1
P. orophilus (n=26) 22.9 1.11 21.1–24.8 26.6 1.10 25.5–28.1
P. soaresi (n=14) 19.5 1.61 17.9–22.8 20.7 1.15 19.9–21.5
CASSINI ET AL.20 · Zootaxa 2491 © 2010 Magnolia Press
FIGURE 15: Physalaemus lateristriga comb. nov. (CHUFSC 1689) from the municipality of Anitápolis (27°50'23"S,
49°5'2"W), state of Santa Catarina. Photograph by Luís O.M. Giasson.
FIGURE 16: Physalaemus lateristriga comb. nov. advertisement call: A—oscilogram, B -audiospectrogram, C—power
spectrum, D—oscillogram of a cut of the same call evidencing pulsed structure, and E—oscillogram of a cut of the same
call evidencing the subpulsed structure. Specimen CFBH 21169, collected at the municipality of São José dos Pinhais,
state of Paraná, on 10/XI/2008. Air temperature: 20.8°C
Variation (n = 129). Males are slightly smaller than females. Range, mean, and standard deviation of
SVL of males and females are in table 4. Specimens analyzed present tarsal tubercle weakly developed (22%)
or absent (78%); and tarsal fold weakly developed (59%) or absent (41%). Interorbital white line complete or
incomplete. White line on dorsolateral fold weakly marked in most specimens analyzed, although in some
specimens it is absent.
Zootaxa 2491 © 2010 Magnolia Press · 21
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
Advertisement call. Advertisement calls (n = 70 calls) were obtained from six males from the
municipalities of São José dos Pinhais (n = 38) and Morretes (n = 15), state of Paraná, and São Bento do Sul,
state of Santa Catarina, Brazil (n = 17). The advertisement call (Fig. 16, table 2) is composed by one single
multipulsed note, each pulse composed by two subpulses (which can be fused on sonogram along the call);
non–harmonic structure, without frequency modulation; mean duration of the call of 3.59 seconds (SD = 0.7,
amplitude = 1.67 to 5.47 seconds); each pulse with 4 to 5 miliseconds; pulse rate ranges from 171 to 202
pulses per second (mean = 183, SD = 10); mean duration of time between calls 1.80 seconds (SD = 0.8,
amplitude = 0.8 to 3.74 seconds) and mean value of the dominant frequency 1.68 kHz (SD = 0.23, amplitude
= 0.75–2.08 kHz).
Geographical distribution. Physalemus lateristriga comb. nov. is distributed in southern state of São
Paulo, and states of Paraná and Santa Catarina (Fig. 8).
Physalaemus feioi, new species
(Figs. 17, 18, 19)
Holotype. MNRJ 5892 (Figs. 17, 18), adult male, collected at Mata da Biologia (20°45’S, 42°51’W; elevation
700 m), Campus of Universidade Federal de Viçosa, municipality of Viçosa, state of Minas Gerais, Brazil, on
15 January 2005, by Carla S. Cassini and André F. Gualberto.
FIGURE 17: Physalaemus feioi sp. nov., holotype (MNRJ 49946, adult male) from municipality of Viçosa, state of
Minas Gerais. A—dorsal, and B—ventral views. Bar = 20 mm.
Paratypes. Collected at the type locality: MNRJ 3117 (female), collected by Renato N. Feio, on 26
August 1997; MNRJ 49945–49947 (males) collected by Carla S. Cassini and André F. Gualberto, on 26
October 2007; MNRJ 5273 (male) collected by Kevin Middlebrooks, on 02 October 2003; MNRJ 5305
(male), collected by Kevin Middlebrooks and Jussara S. Dayrell, on 04 November 2003; MNRJ 5816 (male),
collected by Carla S. Cassini, Emanuel T. Silva, and Jussara S. Dayrell, on 02 December 2004; MNRJ 5825–
5829 (males), collected by Eliana F. Oliveira and Jussara S. Dayrell, on 09 December 2004, MNRJ 5890
(female), MNRJ 5891 (male), collected by André F. Gualberto and Carla S. Cassini, on 15 January 2005;
MNRJ 5996 (male), MNRJ 5997 (male), collected by André F. Gualberto, Carla S. Cassini, and Jussara S.
Dayrell, on 25 January 2004; MNRJ 6226 (female), collected by Carla S. Cassini, Eliana F. Oliveira, and
Jussara S. Dayrell, on 27 April 2005; MZUFV 2539 (male), collected by Vicente W.D. Casali, on 02 January
1996; MZUFV 2542 (male), collected by Vicente W.D. Casali, on 02 January 1996; MZUFV 3189 (male),
collected by Alexandre L. Silva, on 12 November 1997; MZUFV 3614 (female), collected by Alexandre L.
CASSINI ET AL.22 · Zootaxa 2491 © 2010 Magnolia Press
FIGURE 18: Physalaemus feioi sp. nov., holotype (MNRJ 49946, adult male) from municipality of Viçosa, state of
Minas Gerais. A—dorsal, and B—lateral view of head; C—hand, and D—foot. Bars = 10 mm.
Silva, on 13 February 1998; MZUFV 5297 (female), collected by Kevin A. Middlebrooks and Paula F. Leão,
on 31 October 2003; MZUFV 5679 (male), collected by Carla S. Cassini and Jussara S. Dayrell, on 31
October 2003; MZUFV 5680 (male), collected by Carla S. Cassini and Jussara S. Dayrell, on 31 October
2003; MZUFV 5817 (male), collected by Carla S. Cassini, on 02 December 2004; MZUFV 5894 (female),
collected by Carla S. Cassini and André F. Gualberto, on 15 January 2005; MZUFV 5998 (male), collected by
Carla S. Cassini, on 25 January 2004. Mata do Paraíso (20°48’03”S, 42°52’49”W, elevation 732 m),
municipality of Viçosa, state of Minas Gerais, Brazil: MZUFV 665 (male), collected by Renato N. Feio, on 21
September 1992; MNRJ 5688 (female), collected by Carla S. Cassini and Eliana F. Oliveira, on 21 October
2004; MNRJ 6769 (female), collected by Michael E. Nagai, on 26 January 2006.
Diagnosis. (1) Body length small to moderate (SVL 21.2–29.2 mm in males, 23.5–31.2 mm in females);
(2) skin texture shagreened; (3) head longer than wide; (4) snout sub–elliptical in dorsal view, protruding in
lateral profile; (5) lateral black stripe extending from the postorbital region to the inguinal region; (6) inguinal
gland poorly evident; (7) oblique white line from the posterior corner of the eye to the arm insertion; (8) white
line weakly marked, extending from the tip of the snout, through the canthus rostralis and margin of upper
eyelid, to the inguinal region on the dorsolateral fold; (9) annulus of tympanum weakly developed; (10) vocal
sac developed, gray with white dots; (11) tarsal tubercle developed; (12) tarsal fold developed; (13)
advertisement call composed by one note with 30 to 75 pulses (pulse rate 8 to16 pulses per second ); (14)
mean value of the dominant frequency 2.56 kHz.
Comparison with other species. Physalaemus feioi sp. nov. is distinguished from P. aguirrei and P.
insperatus by the absence of a white line outlining the inferior margin of the mandible and the supratympanic
Zootaxa 2491 © 2010 Magnolia Press · 23
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
fold (present in those species) and by the presence of an oblique white line from the posterior corner of the eye
to the arm insertion (absent in those species). It is distinguished from P. aguirrei by the advertisement call
with mean dominant frequency 2.56 kHz (3.1 kHz in P. aguirrei; Pimenta & Cruz 2004). Additionaly, it is
distinguished from P. insperatus by the vocal sac not expanded laterally (expanded laterally in P. insperatus).
Physalaemus feioi sp. nov. has smaller body length than P. maximus (SVL 41.1–46.5 mm in males, 42.2–51.5
mm in females of P. maximus); tarsal tubercle present (absent in P. maximus); advertisement call composed by
30 to 75 pulsed notes (one multipulsed note in P. maximus; Baêta et al. 2007); and mean dominant frequency
2.56 kHz (mean dominant frequency 0.73 or 0.76 kHz in P. maximus; Baêta et al. 2007).
From P. olfersii and P. lateristriga, P. feioi sp. nov. is distinguished by having a concentration of blotches
on gular region of females (females with gular region and abdomen presenting scattered blotches uniformly
distributed in those species); tarsal tubercle developed (weakly developed in those species); callosities on heel
absent (present in those species); and advertisement call with mean dominant frequency 2.56 kHz (1.24 kHz
in P. olfersii and 1.68 kHz in P. lateristriga)
Physalaemus feioi sp. nov. is distinguished from P. orophilus sp. nov. by having larger body length (SVL
21.2–31.2 mm in males, 23.5–31.2 mm in females of P. orophilus sp. nov.); tarsal tubercle present (absent in
P. orophilus sp. nov.); advertisement call with pulse emission rate of 8 to 16 pulses per second (16 to 19 pulses
per second in P. orophilus sp. nov.); each pulse with 14 to 19 subpulses (8 to 11 subpulses in P. orophilus sp.
nov.); and mean dominant frequency 2.56 kHz (3.18 kHz in P. orophilus sp. nov.)
Physalaemus feioi sp. nov. has larger body length than P. soaresi (SVL 17.9–22.8 mm in males, 19.9–
21.52 mm in females of P. soa resi), head longer than wide (wider than long in P. soaresi); an oblique white
line from the posterior corner of the eye to the arm insertion (absent in P. soaresi); vocal sac developed (vocal
sac weakly developed in P. soaresi); tarsal fold absent (developed in P. soaresi); advertisement call with mean
dominant frequency 2.56 kHz (3.3 kHz in P. soaresi; Weber et al. 2005).
Description of the holotype. Body moderately robust; head longer than wide; snout sub–elliptical in
dorsal view, protruding in lateral profile; nostrils elliptical, not protuberant, located and expanded laterally,
closer to the tip of the snout than to the eye; canthus rostralis distinct; loreal region oblique, convex; eyes
protuberant; annulus of tympanum and supratympanic fold indistinct; parotoid glands developed; presence of
a dorsolateral fold starting on the posterior corner of eye, delimiting the dorsal region from the flank, and
ending at the anterior margin of the inguinal region; vocal sac subgular, well developed, extending to the
chest; choanae small, rounded, well separated from each other; tongue narrow, long, free posteriorlly;
maxillary and premaxillary teeth visible; vomerine teeth absent. Arms short, slender; forearms slightly more
robust and longer than arms; fingers thin, long, not fringed; fingers length I < II < IV < III; nuptial pad divided
in two parts, one covering 2/3 of the inner metacarpal tubercle and other enclosing all the thumb, except the
inner side; subarticular tubercles large, single, rounded; outer metacarpal tubercle large, elliptical, covering 1/
3 of carpal region; inner metacarpal tubercle large, rounded, larger than the outer metacarpal tubercle;
supranumerary tubercles present, weakly developed; fingers tip slightly expanded. Legs moderately robust;
tibia slightly longer than thigh, the sum of tibia and thigh lengths longer than SVL; tarsal fold absent; toes
thin, long, weakly fringed; toes length I < II < III = V < IV; subarticular tubercles medium–sized, single,
protruding; inner metatarsal tubercle large, elliptical; outer metatarsal tubercle small, conic, approximately
three times smaller than the inner metatarsal tubercle; tarsal tubercle present, small; supernumerary tubercles
absent; toes tip slightly expanded. Dorsal surface shagreened and ventral surface smooth.
Color in life. Color pattern gray or brown (Fig. 19); small, irregular shaped, scattered light brown
blotches on dorsum; dorsal median line extending on posterior fourth of the dorsum; loreal region dark brown
with white dispersed dots; a black stripe under the dorsolateral fold, extending from the postorbital region to
the inguinal region; a white line weakly marked on canthus rostralis, border of upper eyelid, and dorsolateral
fold; a rectangular dark brown blotch on dorsum of the forearm; transversal brown bars on dorsum of thighs;
irregular brown blotch on internal region of tibia, near to the heel; gular region light gray with scattered white
dots; few white dots and light gray blotches on the anterior region of the abdomen; posterior region of the
abdomen light brown; a dark brown line laterally to the ventral surface of the arms; brown nuptial pads;
inguinal gland not distinguished from the color pattern of the dorsum; posterior region of the abdomen and
tubercles of palmar and plantar surfaces red.
CASSINI ET AL.24 · Zootaxa 2491 © 2010 Magnolia Press
Color in preservative (70% ethanol). General color pattern maintained in preservative. Dorsal surface
with light gray or brown blotch on center; tubercles of plantar and palmar surfaces and abdomen white.
Measurements of the holotype (mm). SVL 25.7; HL 9.3; HW 9.2; ED 2.9; UEW 2.0; IOD 3.1; IND 2.5;
END 2.2; TL 12.6; SL 12.0; FL 20.0.
FIGURE 19: Physalaemus feioi sp. nov. from the municipalities of A—Viçosa, and B, C—Pedra Dourada, state of
Minas Gerais. Photographs by A—Renato N. Feio and B, C—Jussara S. Dayrell.
FIGURE 20: Physalaemus feioi sp. nov. advertisement call: A—oscilogram, B—audiospectrogram, C—power
spectrum, D—oscillogram of a cut of the same call evidencing pulsed structure, and E—oscillogram of a cut of the same
call evidencing the subpulsed structure. Specimen MNRJ 49946, holotype, collected at the municipality of Viçosa
(20°45'S; 42°51'W), state of Minas Gerais, on 26/X/2007. Air temperature: 21.8°C.
Variation (n = 151). General color pattern vary of brown or gray. Females present larger body size and
color pattern of gular region lighter than males. Interorbital white line complete, incomplete or absent. Tarsal
tubercle was observed in 85.8% of the analyzed specimens, and the black stripe, well defined to 2/3 of the
anterior region of the lateral of the body, from the postorbital region to groin was observed in 73.2% of the
analyzed specimens. Few specimens have snout pointed in dorsal view. Most specimens present red belly in
life, as well as the subarticular, inner and outer metatarsal and tarsal tubercles; few specimens present these
regions cream. Range, mean, and standard deviation of SVL of males and females are in table 4.
Advertisement call. Advertisement calls (n = 110 calls) were obtained from 11 males from the
municipality of Viçosa and from the Parque Estadual da Serra do Brigadeiro, municipality of Araponga, state
of Minas Gerais, Brazil. The advertisement call of P. feioi sp. nov. (Fig. 20, table 3) is composed by one note
with 30 to 75 pulses (each pulse is composed by 811 subpulses), non–harmonic structure, with variable
Zootaxa 2491 © 2010 Magnolia Press · 25
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
frequency modulation (ascendant frequency modulation along the call and descendent frequency modulation
at the end of the call); mean duration of 4.30 seconds (SD = 0.66, amplitude = 2.48 to 5.58 seconds); pulse
rate ranges from 8 to 16 pulses per second; pulse duration of 40 to 45 miliseconds; each pulse with 14-19
subpulses; mean duration between calls is 3.72 seconds (SD = 2.74, amplitude = 0.78–12.69 seconds); mean
value of the dominant frequency 2.56 kHz (SD = 0.075, amplitude = 2.31–2.75 kHz).
Etymology: The specific epithet honors the herpetologist Dr. Renato Neves Feio (Museu de Zoologia
João Moojen de Oliveira, Universidade Federal de Viçosa, Minas Gerais, Brazil), with gratitude for his
contributions and efforts to the knowledge of anurans of Minas Gerais and pleasant friendship.
Geographical distribution. Physalaemus feioi sp. nov. is distributed along the Serra da Mantiqueira
mountain range on the states of Minas Gerais and São Paulo, Brazil (Fig. 8).
Physalaemus orophilus, new species
(Figs. 21, 22, 23)
Physalaemus olfersii—Nascimento, Caramaschi & Cruz, 2005 (part).
Holotype. MNRJ 55097 (Figs. 21, 22), adult male, collected at Reserva Particular do Patrimônio Natural da
Serra do Caraça (20°05’56”S, 43°29’15”W; elevation 1300 m), municipalityof Catas Altas, state of Minas
Gerais, Brazil, on 01 November 2008, by José P. Pombal Jr., Ana Carolina C. Lourenço, Délio P. Baêta, and
others.
FIGURE 21: Physalaemus orophilus sp. nov., holotype (MNRJ 55097, adult male) from municipality of Mariana, state
of Minas Gerais. A—dorsal, and B—ventral views. Bar = 20 mm.
Paratypes. Collected at the type locality: MNRJ 55092, MNRJ 55094, MNRJ 55096, MNRJ 55099
(females), MNRJ 55093, MNRJ 55095, MNRJ 55098, MNRJ 55100 (males), collected with the holotype;
MCNAM 3874, MCNAM 3876–3878, MCNAM 3880 (males), collected by Paula C. Eterovick, Kátia Kopp,
and Milena W. Machado, in December 2003; MCNAM 4809 (female), collected by Paula C. Eterovick and
Lilian Afonso, on 27 March 2004; MCNAM 5506 (female), MCNAM 5507, MCNAM 5509 (males),
collected by Kátia Kopp and Milena. W. Machado, in December 2003.
CASSINI ET AL.26 · Zootaxa 2491 © 2010 Magnolia Press
Diagnosis. (1) Body length small (SVL 21.1–24.8 mm in males, 25.5–28.1 mm in females); (2) skin
texture shagreened; (3) head as long as wide; (4) snout subelliptical in dorsal view, protruding in lateral
profile; (5) a lateral black stripe extending from the postorbital region to the inguinal region; (6) inguinal
gland poorly developed; (7) an oblique white line from posterior corner of the eye to the arm insertion; (8) a
white line weakly marked, extending from the tip of the snout, through the canthus rostralis and margin of
upper eyelid, absent on the dorsolateral fold; (9) annulus of tympanum weakly marked; (10) vocal sac
developed, gray marbled with scattered white dots; (11) tarsal tubercle absent; (12) tarsal fold present; (13)
advertisement call composed by one note with 56 to 87 pulses (pulse rate 16 to19 pulses per second ); (14)
mean value of the dominant frequency 3.18 kHz.
Comparison with other species. Physalaemus orophilus sp. nov. is distinguished from P. aguirrei and P.
insperatus by the absence of a white line outlining the inferior margin of the mandible (present in those
species) and by the presence of an oblique white line from the posterior corner of the eye to the arm insertion
(absent in those species). Moreover, it is distinguished from P. inspera tus by the vocal sac not expanded
laterally to the supratympanic fold (expanded laterally in P. insperatus).
Physalaemus orophilus sp. nov. is distinguished from P. feioi by having smaller body length (SVL 21.2–31.2
mm in males, 23.5–31.2 mm in females of P. feioi), tarsal tubercle absent (present in P. fei oi ), advertisement
call with pulse emission rate of 16 to 19 pulses per second (8 to 16 pulses per second in P. feioi), and mean
dominant frequency 3.18 kHz (2.56 kHz in P. feioi)
FIGURE 22: Physalaemus orophilus sp. nov., holotype (MNRJ 55097, adult male) from municipality of Mariana, state
of Minas Gerais. A—dorsal, and B—lateral view of head; C—hand, and D—foot. Bars = 10 mm.
Zootaxa 2491 © 2010 Magnolia Press · 27
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
FIGURE 23: Physalaemus orophilus sp. nov. paratype (MNRJ 55100), from the Patrimônio Natural da Serra do Caraça
(20°05'56"S, 43°29'15"W), municipalityof Catas Altas, state of Minas Gerais. Photograph by José Perez Pombal Jr.
From P. olfersii and P. lateristriga, P. orophilus sp. nov. is distinguished by having smaller body length
(SVL 22.1–37.8 mm in males, 26.9–40.3 mm in females of P. lateristriga; SVL 26.2–36.4 mm in males, 21.7–
41.1 mm in females of P. olfersii), callosities on heel absent (present in those species), and advertisement call
with mean dominant frequency 3.18 kHz (1.24 kHz in P. olfersii; 1.68 kHz in P. lateristriga).
Physalemus orophilus sp. nov. has smaller body length than P. maximus (SVL 41.1–46.5 mm in males,
42.2–51.5 mm in females of P. maximus), tarsal tubercle present (absent in P. maximus), advertisement call
with dominant frequency 3.18 kHz (mean dominant frequency 0.73 or 0.76 kHz in P. maximus; Baêta et al.
2007).
Finally, P. orophilus sp. nov. is distinguished from P. soaresi by the larger body length (SVL 17.9–22.8
mm in males, 19.9–21.5mm in females of P. soaresi), head as long as wide (wider than long in P. s oaresi), an
oblique white line from the posterior corner of the eye to the arm insertion (absent in P. so aresi), vocal sac
developed (vocal sac weakly developed in P. s oares i), tarsal tubercle absent (present in P. soaresi),
advertisement call composed by 56 to 87 pulsed notes (one multipulsed note in P. soaresi; Bokermann 1966a;
Weber et al. 2005), and longer call duration, from 3.07 a 5.18 seconds (1.34 a 2.40 seconds in P. soaresi;
Bokermann 1966a; Weber et al. 2005).
CASSINI ET AL.28 · Zootaxa 2491 © 2010 Magnolia Press
FIGURE 24: Physalaemus orophilus sp. nov. advertisement call: A—oscilogram, B—audiospectrogram, C—power
spectrum, D—oscillogram of a cut of the same call evidencing pulsed structure, and E—oscillogram of a cut of the same
call evidencing the subpulsed structure. Specimen not specified, collected at the municipality of Mariana, state of Minas
Gerais, on I/2007.
Description of the holotype. Body robust; head longer than wide; snout subelliptical in dorsal view,
protruding in lateral profile; nostril elliptical, protuberant, oriented laterally, closer to the tip of the snout than
to the eye; canthus rostralis distinct; loreal region oblique; eyes protuberant; tympanum distinct;
supratympanic fold developed; parotoid glands well developed; dorsolateral fold present, from the posterior
corner of eye, delimiting the dorsal region from flanks, and ending at the anterior margin of the inguinal
gland; vocal sac subgular, well developed, extending to the chest; choanae medium–sized, rounded, well
distinguished from each other; tongue narrow, long, free posteriorly; maxillary and premaxillary teeth visible;
vomerine teeth absent. Arms short, slender; forearms slightly more robust and as long as the arms; fingers
thin, long, not fringed; fingers length I < II < IV < III; nuptial pad divided, one part covering 2/3 of the inner
metacarpal tubercle and other part enclosing all the thumb, except the inner side; subarticular tubercles large,
single, rounded; outer metacarpal tubercle medium–sized, rounded, covering 1/3 of carpal region; inner
metacarpal tubercle medium–sized, elliptical, with the same size of the outer metacarpal tubercle; few
supernumerary tubercles well developed; fingers tip slightly expanded. Legs moderately robust; tibia slightly
longer than thigh; the sum of tibia and thigh lengths slightly shorter than SVL; tarsal fold present; toes thin,
long, weakly fringed; toes length I < II < III = V < IV; subarticular tubercles small, single, protruding; inner
metatarsal tubercle large, elliptical; outer metatarsal tubercle small, ovoid, approximately half of the size of
the inner metatarsal tubercle; tarsal tubercle absent; supernumerary tubercles absent; toes tip slightly
expanded. Dorsal surface shagreened, ventral surface smooth.
Color in preservative (70% ethanol). Color pattern gray or brown; small, irregular shaped, scattered
light brown blotches on dorsum; a light gray or light brown blotch at the center of the dorsal surface; dorsal
median line extending on posterior third of the dorsum; an interorbital dark brown line; loreal region gray
anteriorly with white dots dispersed and white on the region under the eye; a white line weakly marked on
canthus rostralis and border of upper eyelid, absent on dorsolateral fold; a rectangular dark brown blotch on
dorsum of the forearm; a dark brown line on the lateral of the ventral surface of the forearms; transversal
brown bars on dorsum of thighs; gular and anterior region of the abdomen marbled with light gray blotches
and white dots; posterior region of the abdomen and ventral surfaces of thighs white; inguinal gland not
distinguished from the color pattern of dorsum; tubercles of palmar and plantar surfaces white; nuptial pads
brown or light brown.
Zootaxa 2491 © 2010 Magnolia Press · 29
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
Measurements of the holotype (mm). SVL 25.3; HL 9.4; HW 8.3; ED 2.9; UEW 2.2; IOD 2.8; IND 2.2;
END 2.1; TL 12.2; SL 12.6; FL 20.2.
Variation (n = 26). All specimens analyzed agree in morphology and color; females are slightly larger
than males (range, mean, and standard deviation of SVL of males and females are in table 4); nuptial pad vary
in size, covering half or 2/3 of the inner metacarpal tubercle; interorbital brown line complete or incomplete;
few specimens present snout pointed in dorsal view; few individuals present vocal sac brown marbled.
Advertisement call. Advertisement calls (n = 124 calls) were obtained from six males in the
municipalityof Mariana, and in the Reserva Particular do Patrimônio Natural da Serra do Caraça, state of
Minas Gerais, Brazil. The advertisement call (Fig. 24, table 3) is composed by one note with 56 to 87 pulses
(each pulse is composed by 8 to 11 subpulses), pulse duration of 42 to 50 miliseconds; non–harmonic
structure, with an ascendant frequency modulation; mean duration of 3.85 seconds (SD = 0.58, amplitude =
3.07 to 5.18 seconds); pulse rate ranges from 16 to 19 pulses per second; mean duration between calls 1.59
seconds (SD = 0.46, amplitude = 1–2.86 seconds), and mean value of the dominant frequency 3.18 kHz (SD =
0.09, amplitude = 3–3.39 kHz).
Etymology. The specific epithet, an adjective derived from the Greek words “oros” meaning mountain
and “philos” meaning friend, composing “friend of mountains”, is given in allusion to the high elevations of
the localities where the species is found.
Geographical distribution. Physalaemus orophilus sp. nov is distributed in the municipalities of
Mariana, Caeté, Catas Altas, Barão de Cocais, Guanhães, Peçanha, and Santa Bárbara, in the Espinhaço
mountain range in the state of Minas Gerais, Brazil (Fig. 8).
Discussion
Calls of different populations of P. o lfersii have been described (Heyer et al. 1990; Weber et al. 2005; Giaretta
et al. 2009) and doubts to attribute names to populations have been reported (Feio et al. 1999; Canelas &
Bertolucci 2007; Giaretta et al. 2009). Although, nomenclature of some call parameters are not standardized
among publications. Heyer et al. (1990) defined the call of P. olfersii from Boracéia, state of São Paulo, as
‘complexly partially pulsed’. Giaretta et al. (2009) described the call from Atibaia, state of São Paulo
populations, as composed by a single ‘pseudo-pulsed note’, referring to the same call structure observed by
Heyer et al. (1990). Herein, we followed McLister et al. (1995) and Gehardt and Huber (2002), where ‘note’
is considered as the sound unit produced by a single respiratory event, ‘pulses’ a subunit of a note and
produced by contraction of the body wall during a single respiratory event (McLister et al., 1995) and
‘subpulses’ as subunits of a pulse produced by passive amplitude modulation resulted from one body wall
contraction (Gerhardt & Huber, 2002). Our field observations of the species P. olfersii, P. lateristriga, and P.
feioi seemed enough to apply these definitions because movements of contractions of the body wall and
inflation/deflation of the vocal sac during the calls of the species are evident (e.g. Orrico et al., 2009).
In this way, the main structure of the call of P. ol fer si i shown in Heyer et al. (1990), Giaretta et al. (2009) and
herein are subpulsed (two subpulses within one pulse). In the oscillogram figured in Giaretta et al. (2009), and
herein it is possible to see that each pulse is composed by two subpulses and they may be fused together along
the call of P. olfersii.
Calls of P. olfersii and P. lateristriga are distinguished by differences in pulse rate (135–156 pulses per
second in P. olfersii and 171–202 pulses per second in P. lateristriga) and pulse duration (6 to 7 ms in P.
olfersii and 4 to 5 ms in P. lateristriga). Acoustics parameters of P. feioi and P. orophilus are well
distinguished from each other by dominant frequency (2.31–2.75 kHz in P. feioi and 3.0–3.39 kHz in P.
orophilus) and pulse rate (8–16 pulses per second in P. f eioi and 16–19 pulses per second in P. orophilus) .
Calls of P. olfersii and P. lateristriga distinguish from those of P. feioi. and P. orophilus by number of
subpulses (two subpulses sused or not along the call of P. olfersii and P. lateristriga; 14–19 subpulses in P.
feioi and 8–11 subpulses in P. orophilus) and pulse rate (8–16 pulses per second in P. feioi and 16–19 pulses
per second in P. orophilus).
CASSINI ET AL.30 · Zootaxa 2491 © 2010 Magnolia Press
Morphological and acoustic parameters (dominant frequency and pulse rate) allowed us to name those
populations of Physalaemus aff. olfersii pointed by Feio et al. (1999) and Canelas and Bertolucci (2007) as P.
feioi and P. orophilus sp. nov. respectively. Morphological comparisons led us to conclude that the population
from Boracéia, state of São Paulo mentioned by Heyer et al. (1990) is more similar to Physalaemus olfersii.
Bokermann (1966a) mentioned a population of P. olfersii from the municipality of Campo Grande, state of
São Paulo, and found similar dominant frequency of the calls, from 1.0 to 2.0 kHz (we found the dominant
frequency for P. olfersii from 1.38 to 2.56 kHz). Although, he also mentioned that the mean duration of the
call of that population is 2 seconds and figured the call (we found that the mean duration of call of P. olfersii is
4.04 seconds and the shortest call is 2.6 seconds). However, it should be emphasized that it was dificult to
measure call duration from sonograms, because at least untill the year 1986 the maximum time of analysis
was 2.4 seconds (Cardoso 1986); and all sonograms presented by Bokermann (1966a) had maximum time of
analysis of 2.2 seconds.
Calls of populations of P. olfer si i from Boracéia (Heyer et al. 1990) and Atibaia (Giaretta et al. 2009) are
in congruence with some call parameters of P. olf er si i presented herein (call composed by one multipulsed
note with two subpulses without frequency modulation and dominant frequency around 1.002.70 kHz).
However, they show a significant difference on pulse rate (200225 and 220-250 pulses per second in
populations from Boracéia and Atibaia respectively) from present work (135–156 pulses per second). The
pulse duration found in the population of P. olfersii from Atibaia (4 to 4.4 ms; Giaretta et al. 2009) and the
pulse duration of P. lateristriga comb. nov. (4 to 5 ms) are different from the pulse duration of Teresópolis,
state of Rio de Janeiro, P. olfersii population (6 to 7 ms). Although the population from Atibaia, state of São
Paulo (Giaretta et al. 2009) is located southern to Serra da Mantiqueira mountain range, these calls are more
similar to P. olfersii and P. lateristriga comb. nov. than they are to P. feioi. However, calls of populations from
Boracéia (Heyer et al. 1990) and Atibaia (Giaretta et al. 2009) are more similar to each other than they are to
P. olfersii or P. lateristriga comb. nov.
The dominant frequency of the advertisement calls of P. lateristriga comb. nov., populations of P. ol fe rs ii
from Boracéia, state of São Paulo (Heryer et al. 1990) and Teresópolis, state of Rio de Janeiro (present work),
presented a high intrapopulational amplitude variation (over 1.0 kHz). Although, we found a small coefficient
of variation (CV) for this parameter within the population from Teresópolis (CV = 5,9%). However, Giaretta
et al. (2009) found a smaller amplitude of dominant frequency among the calls they analyzed (2.24–2.50 kHz)
of the population from Atibaia, state of São Paulo.
Because no morphological differences were found between populations of P. olfersii from Boracéia and
Teresópolis; similarities were found in calls parameters (dominant frequency and pulse rate) of populations of
P. olfersii from Boracéia (Heyer et al. 1990) and Atibaia (Giaretta et al. 2009); and call parameters (pulse rate)
suggest some diference between populations from Boarcéia (Heyer et al. 1990)/Atibaia (Giaretta et al. 2009)
and Teresópolis, it is possible that as more calls of more males of populations within the distribution range of
P. olfersii and P. lateristriga comb. nov. are analyzed, more cryptic species (sensu Bickford et al. 2007) can
still be found.
The P. olfersii species group is associated with the Atlantic Rain Forest Domain (sensu Ab’Sáber 1977),
from the state of Bahia to state of Santa Catarina. Physalaemus aguirrei ranges from southern state of Bahia
and northern state of Espírito Santo to northeastern state of Minas Gerais; P. orophilus sp. nov. occurs on
south of the Espinhaço mountain range, middle east state of Minas Gerais; P. feioi sp. nov. is distributed in the
Mantiqueira mountain range from southeastern and south of the state of Minas Gerais to northeastern state of
São Paulo; P. lateristriga comb. nov. is distributed in southern state of São Paulo and states of Paraná and
Santa Catarina; P. maximus occurs in southeastern and central regions of the state of Minas Gerais; P. olfer si i
is broadly distributed in Serra do Mar mountain range, from the state of Espírito Santo to the state of Santa
Catarina and the most continental register is in the municipality of Botucatu, state of São Paulo; and P. soaresi
is known from the municipalities of Seropédica (type locality), Duque de Caxias and Rio de Janeiro,
southeastern state of Rio de Janeiro (Pontes et al., 2010). Only two species, among those just listed, are known
to occur in simpatry and sintopy: P. feioi and P. maximus at the Parque Estadual da Serra do Brigadeiro,
municipality of Araponga, state of Minas Gerais (Feio et al. 1999).
Zootaxa 2491 © 2010 Magnolia Press · 31
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
The transition zone between P. ol fe rs ii and P. lateristriga found herein, located between the states of
Paraná and southern state of São Paulo, is coincident with transitions between other morphological species
within anurans—e.g., Phyllomedusa burmeisteri (Pombal and Haddad, 1992), Rhinella crucifer (Baldissera et
al. 2004) and Hypsiboas bischoffi (Marcelino et al. 2009). Additionally, a phylogeographic break between
distinct mitochondrial lineages of the snake Bothropoides jararaca was found by Grazziotin et al. (2006) in
this same area. These concordant patterns suggest a common origin for the geographic differentiation patterns
that may be associated with geographical barriers or fluctuations associated with climatic cyclical alterations
of the Tertiary and Quaternary Periods. Both phenotypic and genetic diversity studies on a variety of
complexes of species of different organisms will be needed to investigate these general hypotheses.
Acknowledgments
We are grateful to Célio F.B. Haddad (CFBH), Denise de C. Rossa–Feres (DZSJRP), Fátima M. de Souza and
Elizabeth R. Bilo (ZUEC), Hussam Zaher and Carolina C. Mello (MZUSP), Luciana B. Nascimento
(MCNAM), Renato Neves Feio (MZUFV), Oswaldo L. Peixoto (EI), Julio César de Moura–Leite and Magno
V. Segalla (MHNCI), and Gláucia Pontes (MCT–PUCRS) for permitted the examination and/or loan of
specimens; Rainer Günther (ZMB), Franz Tiedemann and Heinz Grillitsch (NMW), José Perez Pombal Jr.
(MNRJ), Luiz Menta Giasson (CFBH), and Diogo Borges Provete (DZSJRP) for the photographs conceded of
the syntype of Phryniscus olfersii, the types of Nattereria lateristriga and Phyllobates glandulosus,
Physalaemus orophilus, P. lateristriga and P. olfersii respectively; Ana Carolina C. Lourenço, André F.
Gualberto, Bianca von Berneck, Carlos Eduardo Conte, Délio Baêta, Eliana O. Faria, Emmanuel Teixeira,
Jussara S. Dayrel, Magno V. Segalla, Marcos Bilate, Mariane Targino, Monica Carolina da Silva, and Luiz N.
Weber for helping in fieldworks; Felipe Leite and Magno V. Segalla for providing call records; Victor G. D.
Orrico for reviewing the manuscript; and Conselho Nacional de Desenvolvimento Científico e Tecnológico
(CNPq) for financial supports.
References
Ab’Sáber, A.N. (1977) Os domínios morfoclimáticos na América do Sul. Primeira aproximação. Geomorfologia, 52, 1
23.
Baêta, D., Lourenço, A.C.C., Pezzuti, T.L. & Pires, M.R.S. (2007) The tadpole, advertisement call, and geographic
distribution of Physalaemus maximus Feio, Pombal & Caramaschi, 1999 (Amphibia, Anura, Leiuperidae). Arquivos
do Museu Nacional, 65, 27–32.
Baldissera, F.A., Caramaschi, U. & Haddad, C.F.B. (2004) Review of the Bufo crucifer species group, with descriptions
of two new related species (Amphibia, Anura, Bufonidae). Arquivos do Museu Nacional, 62, 255–282.
Bickford, D., Lohman, D.J., Sodhi, N.S., Ng, P.K.L., Mier, R., Winker, K., Ingram, K.K. & Das, I. (2007) Cryptic species
as a window on diversity and conservation. Trends in Ecology and Evolution, 22, 148–155.
Bokermann, W.C.A. (1962) Notas sobre três espécies de Physalaemus (Amphibia, Salientia, Leptodactylidae). Anais da
Academia Brasileira de Ciências, 4: 563–568.
Bokermann, W.C.A. (1966a) Dos nuevas especies de Physalaemus de Espírito Santo, Brasil (Amphibia,
Leptodactylidae). Physys, 26, 193–202.
Bokermann, W.C.A. (1966b) Lista Anotada das Localidades Tipo de Anfíbios Brasileiros. RUSP, São Paulo, 183p.
Canelas, M.A.S. & Bertoluci, J. (2007) Anurans of the Serra do Caraça, southeastern Brazil: species composition and
phenological patterns of calling activity. Iheringia, Série Zoologia, 97, 21–26.
Cardoso, A.J. (1986) Utilização de recursos para reprodução em comunidades de anuros do sudeste do Brasil. Doctoral
thesis. Universidade Federal de Campinas, Campinas, 216pp.
Cope, E. D. (1867) On the families of the raniform Anura. Journal of the Academy of Natural Sciences of Philadelphia,
2, 189-206.
Cruz, C.A.G., Cassini, C.S. & Caramaschi, U. (2008) A new species of Physalaemus Fitzinger, 1826 (Anura,
Leiuperidae) from southern Brazil. South American Journal of Herpetology, 3, 239–243.
Dixo, M. & Verdade, V.K. (2006) Herpetofauna de serrapilheira da Reserva Florestal de Morro Grande, Cotia (SP). Biota
Neotropica, 6, 1–10.
CASSINI ET AL.32 · Zootaxa 2491 © 2010 Magnolia Press
Feio, R.N., Pombal, Jr., J.P. & Caramaschi, U. (1999) New Physalaemus (Anura: Leptodactylidae) from the Atlantic
Forest of Minas Gerais, Brazil. Copeia, 1999, 141–145.
Frost, Darrel R. (2010) Amphibian Species of the World: an Online Reference. Version 5.4 (8 April, 2010). American
Museum of Natural History, New York, USA. Available from: http://research.amnh.org/vz/herpetology/amphibia/
(13 Abril, 2010). http://research.amnh.org/herpetology/amphibia/index.php. (01 March 2009).
Gerhardt, H.C. & Huber, F. (2002) Acoustic comunnication in insects and anurans. The University of Chicago Prees,
Chicago, 531p.
Giaretta, A.A., Martins, L.B. & Santos, M.P. 2009. Further notes on the taxonomy of four species of Physalaemus
(Anura, Leiuperidae) from the Atlantic Forest of Southeastern Brazil. Zootaxa, 2166, 51-60.
Grazziotin, F.G., Monzel, M., Echeverrigaray, S. & Bonatto, S.L. (2006) Phylogeography of the Bothrops jararaca
complex (Serpentes: Viperidae): past fragmentation and island colonization in the Brazilian Atlantic Forest.
Molecular Ecology, 15, 3969–3982.
Haddad, C.F.B. & Martins, M. (1994) Four species of Brazilian poison frogs related to Epipedobates pictus
(Dendrobatidae): taxonomy and natural history observations. Herpetologica, 50, 282–295.
Izecksohn, E. (1965) Uma nova espécie de Physalaemus Fitzinger, do estado do Rio de Janeiro (Amphibia, Anura).
Revista Brasileira de Biologia, 25:165–168.
Lichtenstein, M.H.K. & Martens, E.K. (1856) Namenverzeichnis in der zoologischen Sammlung der Königlichen
Universität zu Berlin. Nomenclator Reptilium et Amphibiorum. Musei Zoologici Berolinensis. 40 p.
Marcelino, V.R., Haddad, C.F.B. & Alexandrino, J. (2009) Geographical distribution and morphological variation of
striped and non-striped populations of the Brazilian atlantic Forest treefrog Hypsiboas bischoffi (Anura: Hylidae).
Journal of Herpetology, 43, 351–361.
McLister, J.D., Stevens, E.D. & Bogard, J. (1995) Comparative contractile dynamics of calling and locomotor muscles in
three hylid frogs. Journal of Experimental Biology, 198: 1527–1538.
Nascimento, L.B., Caramaschi, U. & Cruz, C.A.G. (2005) Taxonomic review of the species group of the genus
Physalaemus Fitzinger, 1826 with revalidation of the genera Engystomops Jimenéz–de–la–Espada, 1872 and
Eupemphix Steindachner,1863 (Amphibia, Anura, Leptodactylidae). Arquivos do Museu Nacional, 63, 297–320.
Nascimento, L.B., Pimenta, B.V.S., Cruz, C.A.G & Caramaschi, U. (2006) Taxonomic status of Gomphobates
marmoratus Reinhardt and Lütken, 1862 “1861” and Eupemphix fuscomaculatus Steindachner, 1864 (Amphibia,
Anura, Leptodactylidae). South American Journal of Herpetology, 1:166–174.
Orrico, V.G.D., Lingnau, R. & Giasson, L.O.M. (2009) The advertisement call of Dendropsophus nahdereri (Anura,
Hylidae, Dendropsophini). South American Journal of Herpetology, 4, 295–299.
Papavero, N. (1971) Essays on the history of neotropical dipterology with special comments on reference to collectors
(1750–1905). Museu de Zoologia da Universidade de São Paulo, São Paulo, 1, 216 p.
Parker, H.W. (1927) A revision of the frogs of the genera Pseudopaludicola, Physalaemus, and Pleurodema. Annals and
Magazine of Natural History, 9, 450–478.
Peters, W.C.H. (1882) Über Batrachier, insbesondere über die so eben veröffentlichte sweite Auflage des Catalogs der
Batrachia salientia s. Ecaudata des British Museum von Herrn G. A. Boulenger. Sitzungsberichte der Gesellschaft
Naturfforschenden Freunde zu Berlin, 1882, 60-62.
Pimenta, B.V.S. & Cruz, C.A.G. (2004) The tadpole and advertisement call of Physalaemus aguirrei Bokermann, 1966
(Amphibia, Anura, Leptodactylidae). Amphibia–Reptilia, 25, 204–210.
Pombal Jr., J.P. & Haddad, C.F.B. (1992) Espécies de Phyllomedusa do grupo burmeisteri do Brasil Oriental, com
descrição de uma espécie nova (Amphibia, Hylidae). Revista Brasileira de Biologia, 52, 217–229.
Pontes, J.A.L., Pontes, R.C., Santa-Fé, C.P., Lima, V.M., Rocha, C.F.D. (2010) Amphibia, Anura, Leiuperidae,
Physalaemus soaresi Izecksohn, 1965: New record, distribution extension and geographic distribution map. Check
List, 6, 159–161.
Scherzer, K. (1861) Narrative of the circumnavigation of the globe by the austrian frigate Novara (Commodore B. Von
Wullerstorf-Urbair) undertaken by order of the imperial government, in the years of 1857, 1858, & 1859, under the
immediate auspices of His I. and R. Highness the archduke Ferdinand Maximilian, Commander-in-chief of the
austrian navy. Saunders, Otley & Co., London, 578p.
Steindachner, F. (1864) Batrachologische Mitteilungen. Verhandlungen Zoologisch–Botanischen Gesellschaft in Wien,
14, 239–288.
Steindachner, F. (1867) Amphibien. In: Reise der Oesterreichischen Fregatte Novara um die Erde, in den Jahren 1857,
1858, 1859, unter den Befehlen des Commodore B. von Wüllerstorf–Urbair. Zoologische Theil. 70 p., 5 pls.
Straube, F.C. & Scherer–Neto, P. (2001) História da Ornitologia no Paraná. In: Straube, F.C. (Ed.) Ornitologia sem
fronteiras, incluindo os Resumos do IX Congresso Brasileiro de Ornitologia. Fundação O Boticário de Proteção à
Natureza, Curitiba, p. 43–116.
Vielliard, J. (1993) "Side-bands" artefact and digital sound processing. Bioacoustics, 5,159–162.
Weber, L.N., Carvalho–e–Silva, S.P. & Gonzaga, L.P. (2005) The tadpole of Physalaemus soaresi (Anura,
Leptodactylidae), with comments on taxonomy, reproductive behavior, and vocalizations. Zootaxa, 1072, 35–42.
Zootaxa 2491 © 2010 Magnolia Press · 33
TAXONOMIC REVIEW OF PHYSALAEMUS OLFERSII
Appendix
Additional specimens examined
Physalaemus aguirrei. BRAZIL, Bahia: Mucuri MNRJ 19275–19282; Nova Viçosa MNRJ 18917–18919; Parque
Nacional do Descobrimento, Prado MCNAM 3311–3315. Minas Gerais: Nanuque MNCAM 3316–3324. Espírito
Santo: Conceição da Barra MZUFV 3230–3233; Itaúnas MNRJ 28431, 39862–39864; Linhares MNRJ 22753–
22800, 28490–28494.
Physalaemus feioi. BRAZIL, Minas Gerais: Parque Estadual Serra do Brigadeiro, Araponga MZUFV 2211–2212,
2214, 2215, 2217, 2489, 2541, 2724, 2726–2728, 2730, 3298, 3913–3916, 5334, 6715, 2213, 2216, 3299–3300,
3900, 5333, 5335, 6717–6720; Barão de Cocais MCNAM 1471; Caeté MCNAM 865–867; Camanducaia ZUEC
11535; Pedra Dourada MZUFV 6392–6395, 7322, 7369–7372, 6714, 6716, 6732–6733; Tombos MZUFV 5503–
5505, 5510; Viçosa MNRJ 49995–49997, MZUFV 267, 268, 664, 665, 667, 2513, 2750, 3614, 4040, 5688, 5872,
5995, 6008–6009, 6160, 6672, 6735–6739, 6769, 6794, 6805–6806, 6838–6839, 6840, 6903, 6919, 7011–7012,
ZUEC 6162; Mata da Biologia, Campus da Universidade Federal de Viçosa, Viçosa MZUFV 2540, 3116, 3118–
3119, 3189–3190, 5229, 5272, 5759, 5818, 5889, 5893, 5997, 6714, 6716, 6732–6733, 6769, 6806, 7224. São
Paulo: Santo Antônio do Pinhal CFBH 14641; Socorro ZUEC 4488–4489, 4555–4557, 4559–4565.
Physalaemus insperatus. BRAZIL, Paraná: Serra Branca do Araraquara, Guaratuba MNRJ 10332 (holotype), MNRJ
1917, 10328–10331 (paratypes).
Physalaemus lateristriga. BRAZIL, São Paulo: Buri MZUSP 134904–134906; Capão Bonito ZUEC 8523, 9000,
11982; Iporanga CFBH 6314, 8281; Piedade CFBH 15987, 22264, MZUSP 123525–123527; Pilar do Sul CFBH
6106–6110; Ribeirão Branco CFBH 246, 1826, 2334, 6824–6825, 6828, 6832–6833, 16567, 16585, 26566, ZUEC
6557, 6559, 6561, 6578–6579; Fazendinha São Luiz, Ribeirão Branco CFBH 16564–16565, 16568, 16570–16575,
16578–16582, 16587; Ribeirão Grande AL–MN 2820–2823, MZUSP 299, 117436, 117486–117487; Parque
Estadual Carlos Botelho, São Miguel Arcanjo MNRJ 41741, MCNAM 4807, 7308–7313. Paraná: Cerro Azul
MCNAM 4805–4806, MZUFV 4805–4807; São José dos Pinhais DZSJRP 6237, 8851–8853, 8858, 8864–8865,
CFBH 21169–21170; Guaratuba MNRJ 35098, MZUSP 7623–7624; Morretes CFBH 12385, 12388, 12389, MNRJ
31084, 57046, 57047. Santa Catarina: Mina Rocha, Adrianópolis MNRJ 49766–49768, MHNCI 113, 1606, 2975,
4740, 3760; Anitápolis: CHUFSC 1689–1690; Blumenau MCT–PUCRS 8726; Corupá MCT–PUCRS 8736; Gruta
de Botuverá MZUSP 91849; Ilhota MCT–PUCRS 1354–1355; Joinville MCT–PUCRS 8652; Rio Vermelho
MZUSP 82371, MZUSP 82755–82756; São Bento do Sul CFBH 3019, 5268–5277, 5440, 21164–21170, MNRJ
12823, AL–MN 2824–2825, ZUEC 5264.
Physalaemus maximus. BRAZIL, Minas Gerais: Parque Estadual Serra do Brigadeiro, Araponga MNRJ 18810
(holotype), MNRJ 18811, 21736–21738 (paratypes), MZUFV 1442–1443, 2723, 3917, 3719, 3873–3875, 4121
4122, 6655, 8272–7823; Ouro Branco MZUFV 7091–7093, 7120, 7245–7246, 7248, 7552; Ouro Preto MNRJ
46692–46693, MZUFV 6597.
Physalaemus olfersii. BRAZIL, Espírito Santo: Santa Teresa MZUSP 53561–53564. Rio de Janeiro: Parati MNRJ
2428; Teresópolis MNRJ 482, 12826–12827, 2098, 33117, 44567–44568, 5520, 5525, 50309–50314, 56119–56120,
AL–MN 2098, 2836, MZUSP 53446, 53454–53455, 82376; Tinguá MNRJ 2414. São Paulo: Arapeí ZUEC 6991;
Parque das Neblinas, Bertioga MZUSP 136809–136010, 136825, 136839; Botucatu MNRJ 18812–18813; Reserva
Florestal Morro Grande, Cotia MZUSP 132268–132269, 134387–134389; Embu MZUSP 26031; Fazenda do
Veado, Serra da Bocaina, São José do Barreiro MZUSP 52966–52969, 53194–53199; Campo Grande da Serra
MZUSP 105, 134, 36897, 84441–84445; Salesópolis ZUEC 6402; Estação Ecológica de Boracéia, Salesópolis
MZUSP 18467, 25843–25844, 28199, 60238, 60567–60568, 137469–137470; Santo André ZUEC 6138;
Paranapiacaba, Santo André MZUSP 8850, 10785, 13865, 82837; São José do Barreiro MZUSP 82425–82426,
ZUEC 941, 6354–6358, 6544; São Paulo CFBH 13566, MZUSP 9590, 10018, 77032–77034, ZUEC 1359–1360;
Serra da Bocaina, São José do Barreiro MZUSP 82873–82889; Campo da Fruticultura, Serra da Bocaina, São José
do Barreiro MNRJ 55565, MZUSP 84846, 84848–84849, 84853–84854, 84858, 84865–84867, 84869–84870,
84872–84875; Fazenda do Papagaio, Serra da Bocaina, São José do Barreiro MZUSP 84454–84462; São Luiz do
Paraitinga CFBH 5815–5816, 9867, 9885–9886, 10046–10047, 10786, 16262–16271.
Physalaemus orophilus. BRAZIL, Minas Gerais: Guanhães MCNAM 971; Nova Era MCNAM 1114; Peçanha
MCNAM 1132, 1134; Taboleiro MCNAM 3224, 7477, 7762.
Physalaemus soaresi. BRAZIL, Rio de Janeiro: Horto Florestal Mário Xavier, Seropédica EI 1943–1944 (paratypes),
MNRJ 17837–17848.
... cuvieri group; Iglesias & Natale 2013), P. jordanensis (P. gracilis group; Bokermann 1967; Giaretta et al. 2009), P. feioi (Cassini et al. 2010), P. olfersii (Giaretta et al. 2009), and P. lateristriga (Cassini et al. 2010), where the last three are members of the P. olfersii group. Call duration of P. centralis is similar to that of P. albifrons (Pederassi et al. 2015), P. ephippifer (Kaefer et al. 2011), P. erikae , and P. fernandezae (Barrio 1965); pulse number is similar as in P. feioi (Cassini et al. 2010), and pulse rate is similar as in P. henselii (Barrio 1965). ...
... cuvieri group; Iglesias & Natale 2013), P. jordanensis (P. gracilis group; Bokermann 1967; Giaretta et al. 2009), P. feioi (Cassini et al. 2010), P. olfersii (Giaretta et al. 2009), and P. lateristriga (Cassini et al. 2010), where the last three are members of the P. olfersii group. Call duration of P. centralis is similar to that of P. albifrons (Pederassi et al. 2015), P. ephippifer (Kaefer et al. 2011), P. erikae , and P. fernandezae (Barrio 1965); pulse number is similar as in P. feioi (Cassini et al. 2010), and pulse rate is similar as in P. henselii (Barrio 1965). ...
... gracilis group; Bokermann 1967; Giaretta et al. 2009), P. feioi (Cassini et al. 2010), P. olfersii (Giaretta et al. 2009), and P. lateristriga (Cassini et al. 2010), where the last three are members of the P. olfersii group. Call duration of P. centralis is similar to that of P. albifrons (Pederassi et al. 2015), P. ephippifer (Kaefer et al. 2011), P. erikae , and P. fernandezae (Barrio 1965); pulse number is similar as in P. feioi (Cassini et al. 2010), and pulse rate is similar as in P. henselii (Barrio 1965). Dominant frequency of P. centralis is similar to P. lisei (Morais & Kwet 2012) and P. maximus (Baêta et al. 2007;Peres & Simon 2012). ...
Article
Full-text available
Acoustic signals are employed to support the correct designation of species and enables the discrimination between individuals. We analyzed both intra and interindividual variability of the advertisement call of Physalaemus centralis on the basis of specimens recorded in eight localities of the Cerrado in central Brazil. Within-individual variation, most of the call parameters were static properties. Both spectral and temporal call properties were dynamic between individuals. All call proprieties varied more between individuals than within an individual. Our study adds important data for future studies on the taxonomy and phylogeny of the P. cuvieri clade.
... mm of snout-vent length in males and 22.4-41.1 mm in females (Cassini et al. 2010), and is distributed in the Atlantic Rain Forest domain and its influence areas, from the municipality of Santa Teresa, state of Espírito Santo, southern region of the states of Minas Gerais and São Paulo (Cassini et al. 2010). Ctenid spiders of the genus Phoneutria have already been reported as anuran predators (Rego et al., 2005, Santana et al., 2009, Caldart et al., 2011, Pacheco et al., 2016, and are nocturnal with wandering habits, which actively seek their prey (Lucas, 1988). ...
... mm of snout-vent length in males and 22.4-41.1 mm in females (Cassini et al. 2010), and is distributed in the Atlantic Rain Forest domain and its influence areas, from the municipality of Santa Teresa, state of Espírito Santo, southern region of the states of Minas Gerais and São Paulo (Cassini et al. 2010). Ctenid spiders of the genus Phoneutria have already been reported as anuran predators (Rego et al., 2005, Santana et al., 2009, Caldart et al., 2011, Pacheco et al., 2016, and are nocturnal with wandering habits, which actively seek their prey (Lucas, 1988). ...
... For both these species we encourage the need to gather more information (Bland et al., 2012, Morais et al., 2013. Using only the approximation of the known geographic range size of each species (Kolenc et al., 2007;Cassini, Cruz and Caramaschi, 2010;Pereyra et al., 2012;Batista and Bastos, 2014) both species should probably be categorized as Least Concern (more than 20.000 km 2 , IUCN, 2013, Morais et al., 2013). However, it is important to note that population studies of amphibians employing accurate census techniques are still scarce in Brazil (Vasconcelos and Colli, 2009;Hiert, Hoper and Moura, 2012) therefore a more accurate analyses should be performed by IUCN specialists. ...
Article
Full-text available
Our goal was to assess the species composition, reproductive modes and conservation status of the anurans of Araucaria Forest and Atlantic Rain Forest in the Campina Grande do Sul (CGS) and Bocaiúva do Sul (BS), in the state of Paraná, Brazil. The list of species was assembled based on field and museum records. We recorded 35 species that showed 11 different reproductive modes. Thirty five species occurred in CGS and 20 in BS, one species is classified as “data deficient” under the IUCN criteria while just two species hold this classification within the state. Amphibian richness in these areas correspond to 24% of the known species diversity known for Paraná. Species richness and community composition correspond to the regional phytophysiognomy and local environmental heterogeneity.
Article
Anurans show a wide variety of anti-predator mechanisms, and the species of the Neotropical clade Leiuperinae display several of them. Most species of Edalorhina, Physalaemus and Pleurodema show eyespots, hidden bright colours, macroglands in a inguinal/lumbar position, defensive behaviours and/or chemical defence. We conducted a histological analysis of dorsal and lumbar skin and revised the colour patterns, defensive behaviours and glandular secretions to study the diversity and evolution of anti-predator mechanisms associated with macroglands. We describe 17 characters and optimize these in a phylogenetic hypothesis of Leiuperinae. In the most recent common ancestor of Edalorhina + Engystomops + Physalaemus + Pleurodema, a particular type of serous gland (the main component of macroglands) evolved in the lumbar skin, along with the absence of the Eberth–Katschenko layer. A defensive behaviour observed in leiuperines with macroglands includes four displays (‘crouching down’ behaviour, rear elevation, body inflation and eye protection), all present in the same ancestor. The two elements associated with aposematism (hidden bright colours and eyespots) evolved independently in several species. Our results provide phylogenetic evidence for the startle-first hypothesis, which suggests that behavioural displays arise as sudden movements in camouflaged individuals to avoid predatory attacks, before the origin of bright coloration.
Article
Full-text available
We report the first record of Physalaemus albifrons in the state of Alagoas, in the municipality of Boca da Mata, Brazil.
Article
Full-text available
Faunal surveys are important tools to enhance biological conservation strategies in different regions. This study presents information on amphibian species richness and species composition in Parque Nacional das Araucárias; the largest conservation unit of mixed ombrophilous forest in Santa Catarina State, Southern Brazil. Active searches were conducted in October 2008 and March 2009 in sites potentially used for reproduction of species. We recorded a total of twenty nine species, being 28 native and one exotic species, and belonging to 16 genera and eight families. The high richness recorded, as well as the occurrence of species associated with nearly pristine habitats, suggest that Parque Nacional das Araucárias is an important area for amphibian conservation in Santa Catarina State.
Article
Full-text available
This study provides the first record of Physalaemus maximus in the state of Espírito Santo, southeastern Brazil. Physalaemus maximus was recorded in the Municipality of Castelo (1100 m a.s.l.), where it was observed in explosive reproduction in an Atlantic Forest remnant on 06 January 2010. The advertisement call of P. maximus consisted of a single multipulsed note, with carrier frequency emitted in 1,250 Hz and mean duration of 2.10 s. The record from Castelo extends in approximately 140 km eastward from the previous geographic distribution admitted for the species.
Article
Full-text available
We present a list of the anuran amphibians of the state of Espírito Santo, southeastern Brazil. The list was compiled from data gathered from fieldwork over the course of the past 20 years in different localities, and from literature records and voucher specimens deposited in scientific collections. Our list comprises 133 species. Patchy sampling efforts and recent species descriptions suggest that the list may increase significantly with further sampling, considering the presence of several geographical gaps. The need for adequate sampling in these gap areas is highlighted.
Article
Full-text available
The Bufo crucifer species group is revised on the basis of external morphological and morphometrical characteristics, evidencing variation in size, shape of the parotoid gland, width of the head, cranial crests, and presence or absence of yellow spots near the cloaca and hind limbs. Five species are recognized: B. crucifer Wied-Neuwied, 1821, B. ornatus Spix, 1824 (revalidated), B. henseli A.Lutz, 1934 (revalidated), B. abei sp.nov., and B. pombali sp.nov. The geographic distribution of the species is associated with the Atlantic Rain Forest and adjacent areas: B. crucifer occurs from the State of Ceará to southern State of Espírito Santo and northeastern State of Minas Gerais; B. ornatus is distributed from southern State of Espírito Santo, through the states of Rio de Janeiro and São Paulo to northern State of Paraná, and possibly in northeastern Argentina, in the provinces Misiones and Corrientes; B. henseli is found from southern State of Santa Catarina to the coast of the State of Rio Grande do Sul; B. abei sp.nov., described from Córrego Grande, Municipality of Florianópolis, State of Santa Catarina, is distributed from the State of Paraná to southern State of Santa Catarina and areas of the northern State of Rio Grande do Sul; and B. pombali sp.nov., described from the Reserva Biológica de Peti, Municipality of São Gonçalo do Rio Abaixo, State of Minas Gerais, occurs in transitional areas between the Atlantic Rain Forest and the “cerrados” in the State of Minas Gerais. Additionally, Bufo crucifer var. pfrimeri Miranda-Ribeiro, 1926, currently in the synonymy of B. crucifer, is transfered to the synonymy of Bufo guttatus Schneider, 1799. Bufo levicristatus Boettger, 1885 is considered a species inquirenda and removed from the synonymy of any species included in the B. crucifer group. Bufo spixii Fitzinger, 1826 is transfered from the synonymy of Bufo margaritifer (Laurenti, 1768) to the synonymy of Bufo ornatus Spix, 1824.
Article
Full-text available
Herein we describe the tadpole and the advertisement call of Physalaemus maximus, based on specimens from the Municipality of Ouro Preto, State of Minas Gerais, Brazil. Our data are compared with the information available on the P. olfersii species group. Updated data on geographic distribution are also presented.
Article
Full-text available
We analyzed the advertisement calls of four species of Physalaemus (P. olfersii, P. moreirae, P. bokermanni, and P. jordanensis) from the Brazilian Atlantic Forest. The tadpole of P. olfersii is described for the first time. Calls of all species are composed by just one kind of note. Those of P. olfersii are long (4035 ms), pseudo-pulsed, with dominant frequency around 2375 Hz; 7–10 sidebands are well distinguishable. The calls of P. moreirae are pulsed, with duration about 604 ms, mean dominant frequency around 1369 Hz and 3–5 sidebands. Physalaemus bokermanni has a call structurally different from the other species by the absence of sidebands or harmonics, larger inter-pulse breaks (around 34–42 ms) and a wider frequency range (around 3700 Hz). Calls of P. jordanensis are pulsed and have 7–8 harmonics, last about 1068 ms and dominant frequency around 2742 Hz. Physalaemus olfersii tadpoles show typical features of other Physalaemus larvae, measuring about 19 mm in total length. Our data on call of P. olfersii suggest that a broader taxonomic survey would be worthwhile by revealing that different species are included under this name. Calls of P. moreirae and P. bokermanni, in general, are in agreement with those previously described from other sites, but complementary features are given. Those of Physalaemus jordanensis show some differences in relation to the calls from the type locality, but additional data are still necessary for a taxonomic conclusion.
Article
Full-text available
A new species of the frog genus Physalaemus, belonging to the P. signifer species group, is described from the Atlantic Forest of the Serra do Brigadeiro (20°42′S, 42°29′N), Municipality of Araponga, State of Minas Gerais, Brazil. The new species is readily separated from all other species included in the P. signifer group by its large size (43.3-48.5 mm SVL). /// Uma nova espécie de anuro do gênero Physalaemus, pertencente ao grupo de P. signifer, é descrita da Floresta Atlântica da Serra do Brigadeiro (20°42′S, 42°29′W), Município de Araponga, Estado de Minas Gerais, Brasil. A nova espécie é facilmente separada de todas as outras espécies do grupo de P. signifer pelo seu maior tamanho (43,3-48,5 mm SVL).
Article
Full-text available
We present an annotated list of anurans, lizards, and snakes registered at the Reserva Florestal de Morro Grande, Cotia, State of São Paulo. The list includes 27 anurans, five lizards and three snakes captured mainly in pitfall traps and encountered during field surveys. Eighteen species of anurans captured from pitfall traps were used to compare six areas sampled inside the Reserve, three in secondary forest and three in mature forest. The analysis showed significant differences among secondary and mature forest. We also compared the similarity of the anurofauna of the Reserve to those of six other Atlantic Forest localities inside the State. The anurofauna found at the Reserve is more similar to that found in other localities at the Atlantic plateau than those from coastal plains. The small number of reptiles specimens captured in pitfall traps precluded the same analyses made for anurans. The low abundance of these animals inside the forest limited our ability to compare different areas. There is an evident lack of information on the biology, distribution and conservation status on Brazilian herpetofauna that could be filled by inventory and monitoring of this fauna. Only natural history knowledge can lead to efficient conservation planning of these species.
Article
Full-text available
The species groups of the genus Physalaemusare redefined based on morphometrics, external morphology, color patterns, and osteological characters. Seven species groups of Physalaemus are recognized: P. cuvieri group (eight species), P. signifergroup (ten species), P. albifronsgroup (four species), P. deimaticus group (three species), P. gracilis group (five species), P. henselii group (three species), and P. olfersii group (four species). The genus Engystomops Jiménez-de-la-Espada, 1872 is revalidated to include the former P. pustulosus group (sensuLynch, 1970). It consists of seven species:E. petersi Jiménez-de-la-Espada, 1872 (type-species by monotypy), E. coloradorum (Cannatella & Duellman, 1984), E. guayaco (Ron, Coloma & Cannatella, 2005), E. montubio (Ron, Cannatella & Coloma, 2004), E. pustulatus (Shreve, 1941), E. pustulosus (Cope, 1864), and E. randi (Ron, Cannatella & Coloma, 2004). The genus EupemphixSteindachner, 1863 is revalidated to include E. nattereriSteindachner, 1864. Key words: Anura. Leptodactylidae. Physalaemus. Engystomops.Eupemphix,. RESUMO: Revisão taxonômica dos grupos de espécies do gênero Physalaemus Fitzinger, 1826 com revalidação dos gêneros Engystomops Jiménez-de-la-Espada, 1872 e Eupemphix Steindachner, 1863 (Amphibia, Anura, Leptodactylidae). Os grupos de espécies do gênero Physalaemus são redefinidos com base em características de morfometria, morfologia externa, padrão de desenho e osteologia. Os sete grupos propostos são: grupo P. cuvieri (oito espécies), grupo P. signifer (dez espécies), grupo P. albifrons (quatro espécies), grupo P. deimaticus (três espécies), grupo P. gracilis (cinco espécies), grupo P. henselii (três espécies) e grupo P. olfersii (quatro espécies). Ogênero Engystomops Jiménez-de-la-Espada, 1872 é revalidado para incluir as espécies previamente pertencentes ao grupo P. pustulosus (sensu Lynch, 1970), sendo composto por sete espécies: E. petersi Jiménez-de-la-Espada, 1872 (espécie-tipo por monotipia), E. coloradorum (Cannatella & Duellman, 1984), E.