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Taxonomic review of Dendrophryniscus brevipollicatus Jiménez de la Espada,

1870, with revalidation of D. imitator (Miranda-Ribeiro, 1920) and D. lauroi
Miranda-Ribeiro, 1926, and d...
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DOI: 10.11646/zootaxa.4648.1.2
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Zootaxa 4648 (1): 027–062 ISSN 1175-5326 (print edition)
https://www.mapress.com/j/zt/
Copyright © 2019 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4648.1.2
http://zoobank.org/urn:lsid:zoobank.org:pub:ECE2A8C4-9CAA-4580-B589-D693C2F3EEB6
Taxonomic review of Dendrophryniscus brevipollicatus Jiménez de la Espada,

1870, with revalidation of D. imitator (Miranda-Ribeiro, 1920)
and D. lauroi Miranda-Ribeiro, 1926, and description of four new related species
(Anura, Bufonidae)
CARLOS ALBERTO GONÇALVES CRUZ1,4, ULISSES CARAMASCHI1,
LUCIANA ARDENGHI FUSINATTO2 & CINTHIA AGUIRRE BRASILEIRO3
1
Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Vertebrados, Quinta da Boa Vista, São Cristóvão, 20940-
040 Rio de Janeiro, RJ, Brasil. E-mail: cagcruz@uol.com.br; ulisses@acd.ufrj.br
2
Universidade do Estado do Rio de Janeiro, Departamento de Ecologia, Rua São Francisco Xavier 524, CEP 20550-013, Rio de Ja-
neiro, Brasil. E-mail: lufusinatto@gmail.com
3
Universidade Federal de São Paulo, Instituto de Ciências Ambientais, Químicas e Farmacêuticas, Departamento de Ecologia e Bio-
logia Evolutiva, Rua Professor Artur Riedel, 275, Eldorado, 09972-270 Diadema, SP, Brasil. E-mail: cabrasileiro224@gmail.com
4
Corresponding author. E-mail: cagcruz@uol.com.br
Abstract
Dendrophryniscus brevipollicatus Jiménez de la Espada is a Neotropical bufonid endemic to a small range of the Brazilian
Coastal Atlantic Forest, with reduced body size for the family and bromeligenous habit. We reviewed the taxonomic status
of populations of D. brevipollicatus from states of Rio de Janeiro and São Paulo, including some continental islands,
based on external morphology. We tested our morphological species by a DNA-barcoding approach with sequences of
16S RNA ribosomal gene fragment. DNA-barcoding analysis included other recognized Dendrophryniscus species and
was concordant with morphological species diagnosed in our review. Intraspecific genetic distances ranged from 0 to
2.72 % (± 0.91 %). Interspecific distances ranged from 3.35 % (± 0.90 %) to 20.15 (± 2.23 %). Optimal threshold values
ranged from 2.8 % to 3.0 % and barcode gap analysis showed that for all individuals the furthest intraspecific distances
was always lower than the closest non-conspecific individual. Seven distinct species were recognized. A neotype for
D. brevipollicatus was designated and described; the type locality was determined for the Açude da Solidão (22º57’S,
43º17’W, Datum WGS 84; 410 m altitude), Parque Nacional da Tijuca, Municipality of Rio de Janeiro, State of Rio
de Janeiro, Brazil. Two species were revalidated: D. imitator (Miranda-Ribeiro), with designation of a lectotype and
descriptions of the lectotype and of a topotype from the restricted type locality, Alto da Serra (23o46’S, 46o19’W, Datum
WGS 84, 800 m altitude), Municipality of Paranapiacaba, State of São Paulo, Brazil; and D. lauroi (Miranda-Ribeiro),
with descriptions of the lectotype and of a topotype from the type locality, Municipality of Angra dos Reis (22o54’S,
44o20’W, Datum WGS 84; 25 m altitude), State of Rio de Janeiro, Brazil. Four new species were described: D. davori
sp. nov., from Baixo Caledônia (22º21’S, 42º35’W, Datum WGS 84; 1600 m altitude), Municipality of Nova Friburgo,
State of Rio de Janeiro, Brazil; D. haddadi sp. nov., from Parque Estadual da Serra do Mar, Núcleo Santa Virgínia
(23o21’S, 45o08’W, Datum WGS 84; 970 m altitude), Municipality of São Luís do Paraitinga, State of São Paulo, Brazil;
D. izecksohni sp. nov., from Campo de Fruticultura da Bocaina (currently Núcleo Senador Vergueiro), Municipality of
São José do Barreiro (22º38’S, 44º34’W, Datum WGS 84, 540 m altitude), State of São Paulo, Brazil; and D. jureia sp.
nov., from Estação Ecológica da Juréia-Itatins—Núcleo Rio Verde (24º22’S, 47º04’W, Datum WGS 84; 32 m altitude),
Municipality of Iguape, State of São Paulo, Brazil. Geographical distributions of all species are provided.
Key words: Amphibia, Brazil, Atlantic Rain Forest, Dendrophryniscus, DNA-barcoding, Taxonomy
Resumo
Dendrophryniscus brevipollicatus Jiménez de la Espada é um bufonídeo neotropical endêmico a uma pequena distribuição
na Floresta Atlântica, com corpo reduzido em relação a outros membros da família e hábitos bromelígenas. Revisamos a
posição taxonômica de populações de D. brevipollicatus dos Estados do Rio de Janeiro e São Paulo, incluindo algumas
ilhas continentais, com base na morfologia externa. Testamos as espécies morfológicas obtidas através de uma abordagem
Accepted by M. Vences: 5 Jun. 2019; published: 29 Jul. 2019 27

de DNA-barcoding com sequências de fragmentos gênicos de RNA ribossômico 16S. A análise de DNA-barcoding incluiu
outras espécies conhecidas de Dendrophryniscus e foi concordante com as espécies morfológicas diagnosticadas em nossa
revisão. As distâncias genéticas intraespecíficas variaram de 0 a 2,72 % (± 0.91 %). Distâncias interespecíficas variaram
de 3,35 % (± 0.90 %) a 20,15 (± 2.23 %). Valores de limites ótimos variaram de 2,8 % a 3,0 % e a análise de espaçamento
do barcode mostrou que, para todos os indivíduos, as maiores distâncias genéticas intraespecíficas foram sempre menores
que as dos indivíduos não coespecífico mais próximo. Sete espécies distintas foram reconhecidas. Um neótipo para D.
brevipollicatus foi designado e descrito; a localidade tipo foi determinada para o Açude da Solidão (22º57’S, 43º17’W,
Datum WGS 84; 410 m altitude), Parque Nacional da Tijuca, Município do Rio de Janeiro, Estado do Rio de Janeiro,
Brasil. Duas espécies foram revalidadas: D. imitator (Miranda-Ribeiro), com designação de um lectótipo e descrições
do lectótipo e de um topótipo obtido na localidade tipo restrita, Alto da Serra (23o46’S, 46o19’W, Datum WGS 84, 800
m altitude), Município de Paranapiacaba, Estado de São Paulo, Brasil; e D. lauroi (Miranda-Ribeiro), com descrições
do lectótipo e de um topótipo obtido na localidade tipo, Município de Angra dos Reis (22o54’S, 44o20’W, Datum WGS
84; 25 m altitude), Estado do Rio de Janeiro, Brasil. Quatro novas espcies foram descritas: D. davori sp. nov., de Baixo
Caledônia (22º21’S, 42º35’W, Datum WGS 84; 1600 m altitude), Município de Nova Friburgo, Estado do Rio de Janeiro,
Brasil; D. haddadi sp. nov., do Parque Estadual da Serra do Mar, Núcleo Santa Virgínia (23o21’S, 45o08’W, Datum
WGS 84; 970 m altitude), Município de São Luís do Paraitinga, Estado de São Paulo, Brasil; D. izecksohni sp. nov., do
Campo de Fruticultura da Bocaina (atualmente Núcleo Senador Vergueiro), Muncípio de São José do Barreiro (22º38’S,
44º34’W, Datum WGS 84, 540 m altitude), Estado de São Paulo, Brasil; e D. jureia sp. nov., da Estação Ecológica da
Juréia-Itatins—Núcleo Rio Verde (24º22’S, 47º04’W, Datum WGS 84; 32 m altitude), Municpio de Iguape, Estado de São
Paulo, Brasil. As distribuições geográficas de todas as espécies são fornecidas.
Introduction
The bufonid Neotropical genus Dendrophryniscus Jiménez de la Espada, 1870 is endemic to the Coastal Atlantic
Rainforest of Brazil (Frost 2019) and almost entirely formed by toads with reduced body size (<30 mm) and spe-
cialized reproductive modes, with few large eggs that are often deposited in bromeliads (Izecksohn 1968, 1994;
Cruz & Fusinatto 2008; Carvalho-e-Silva et al. 2010; Recoder et al. 2010; Caramaschi 2012a), or small, pigmented
eggs laying in ponds (Izecksohn 1968; Izecksohn & Cruz 1972). Recent phylogenetic analysis recovered the genus
as monophyletic (Graybeal & Cannatella 1995, Fouquet et al. 2012) and sister taxon of all bufonids (Van Bocx-
laer et al. 2009; Van Bocxlaer et al. 2010; Pyron & Wiens 2011; Portik & Papenfuss 2015). Currently the genus
comprises ten recognized species (Frost 2019): D. berthalutzae Izecksohn, 1994, D. brevipollicatus Jiménez de la
Espada, 1870, D. carvalhoi Izecksohn, 1994, D. krausae Cruz & Fusinatto, 2008, D. leucomystax Izecksohn, 1968,
D. oreites Recoder, Teixeira, Cassimiro, Camacho & Rodrigues, 2010, D. organensis Carvalho-e-Silva, Mongin,
Izecksohn & Carvalho-e-Silva, 2010, D. proboscideus (Boulenger, 1882), D. skuki (Caramaschi, 2012a, 2012b), and
D. stawiarskyi Izecksohn, 1994.
Carvalho (1949) presumed a wide range for Dendrophryniscus brevipollicatus, from states of Espírito Santo to
Santa Catarina, in the Atlantic Coastal Forest of Brazil. Based on this information, Braun and Braun (1979) consid-
ered the southernmost specimens from Itaimbezinho, State of Rio Grande do Sul, as D. brevipollicatus. Posteriorly,
Izecksohn (1994) described three new species, recognizing specimens previously considered D. brevipollicatus
from Santa Teresa, State of Espírito Santo, as D. carvalhoi, specimens from Bituruna, State of Paraná, as D. stawiar-
skyi, and specimens from Joinville and Rio dos Cedros, State of Santa Catarina, as D. berthalutzae. The specimens
referred by Braun and Braun (1979) as D. brevipollicatus were identified as D. krausae by Cruz and Fusinatto
(2008). The distribution of D. brevipollicatus was then restricted to the states of Rio de Janeiro and São Paulo.
Here we reviewed the taxonomic status of populations of D. brevipollicatus from Rio de Janeiro and São Paulo.
We first diagnosed species based on external morphology and then followed a DNA-barcoding approach to estimate
genetic distances within and between diagnosed species. This integrative analysis leaded us to recognize seven
distinct species. Besides D. brevipollicatus, two species were revalidated, and four new species were described. In
this paper we present these results.
Material and methods
Morphology. Specimens used in the descriptions and for comparisons, and/or tissues collections are listed in the
Appendix 1. Acronyms for herpetological collections follow Sabaj (2016).
28 · Zootaxa 4648 (1) © 2019 Magnolia Press CRUZ et al.

TABLE 1. Species, localities, voucher numbers and GenBank Accession numbers of 16S ribosomal RNA gene sequences used for barcode analysis of Dendrophryniscus. Species
of Dendrophryniscus brevipollicatus reviewed in this study and new GenBank sequences are in bold.
Species Locality Municipality State Voucher GenBank
Dendrophryniscus berthalutzae Barragem do Rio São Bento Siderópolis SC CFBH23725 JN867552 A
Dendrophryniscus berthalutzae Barragem do Rio São Bento Siderópolis SC CFBH23724 JN867550 A
Dendrophryniscus berthalutzae Praia Grande SC CHUFSC 2271 MK140791
Dendrophryniscus berthalutzae Treviso SC CFBH10322 JN867551A
Dendrophryniscus brevipollicatus PARNA Tijuca Rio de Janeiro RJ TAD_Dendro09* MK140796
Dendrophryniscus brevipollicatus PARNA Tijuca Rio de Janeiro RJ TAD_Dendro10* MK140797
Review of Dendrophryniscus brevipollicatus

Dendrophryniscus brevipollicatus Reserva Ecológica de Guapiaçu Cachoeiras de Macacu RJ MNRJ60701 MK140795
Dendrophryniscus carvalhoi Pedra Escorada, PARNA Caparaó Divino São Lourenço ES MTR15755 JN867568A
Dendrophryniscus carvalhoi Córrego do Calçado, PARNA Caparaó Ibitirama ES MTR15757 JN867569 A
Dendrophryniscus haddadi Ilhabela SP CFBHT05308 KU495199
Dendrophryniscus haddadi PESM Santa Virgínia São Luís do Paraitinga SP CFBHT02881 (paratype) KU495198 B
Dendrophryniscus haddadi Cunha SP CAB301* MK140801
Dendrophryniscus imitator Paranapiacaba Santo André SP CFBHT14581 KU495197 B
Dendrophryniscus imitator Santos SP CFBHT05292 KU495195 B
Dendrophryniscus imitator Juquitiba SP JUQ JN867556 A
Dendrophryniscus imitator Caucaia do Alto, Reserva do Morro Grande Cotia SP AF1577 JN867553 A
Dendrophryniscus imitator São Bernardo do Campo SP TH040AF1025 JN867555 A
Dendrophryniscus imitator Estação Biológica de Boracéia Salesópolis SP AF1541 JN867554 A
Dendrophryniscus jureia Estação Ecológica Juréia-Itatins Iguape SP RJS717* MK140802
Dendrophryniscus jureia Estação Ecológica Juréia-Itatins Iguape SP RJS924* MK140803
Dendrophryniscus lauroi Ilha Grande Angra dos Reis RJ MTR15547 JN867558 A
......continued on the next page
Zootaxa 4648 (1) © 2019 Magnolia Press ·

29
TABLE 1. (Continued)
Species Locality Municipality State Voucher GenBank
Dendrophryniscus lauroi Ilha Grande Angra dos Reis RJ MTR15548 JN867557 A
Dendrophryniscus lauroi Ilha Grande Angra dos Reis RJ MNRJ57603 MK140798
Dendrophryniscus oreites Serra das Lontras Arataca BA MTR16368 JN867567 A
Dendrophryniscus organensis PARNA Serra dos Órgãos Teresópolis RJ CFBHT12350 KU495196 B
Dendrophryniscus proboscideus Serra das Lontras Arataca BA MTR_ALCX168P68 KU495200B
Dendrophryniscus proboscideus Serra das Lontras Arataca BA MTR16424 JN867565A
Dendrophryniscus proboscideus REBIO Mata Escura Jequitinhonha BA MTR17173 JN867566A
30 · Zootaxa 4648 (1) © 2019 Magnolia Press

Dendrophryniscus davori Baixo Caledônia Nova Friburgo RJ MNRJ60700 (paratype) MK140790
Legend: Source of GenBank sequences—A—Fouquet et al. 2011, B—Lyra et al. 2016. * vouchers identified by collector number, CAB—Cinthia A. Brasileiro; RJS—Ricardo J.
Sawaya; and TAD—Thiago A. Dorigo.
CRUZ et al.
Measurements of the specimens were obtained by a digital caliper under a stereomicroscope and follow Napoli
(2005). Abbreviations of the measurements (in millimeters) are: SVL (snout–vent length); HL (head length); HW
(head width); IND (internarial distance); ESD (eye to snout distance); ED (eye diameter); IOD (interorbital dis-
tance); THL (thigh length); TL (tibia length); TAL (tarsal length); FL (foot length). Snout profile terminology and
webbing formula notation follow Heyer et al. (1990). Photographs were made in a stereomicroscope Leica M205C
with a Leica DFC 450 camera.
Although Fabrezi & Alberch (1996) demonstrated the loss of finger I in the anuran evolution, we still are us-
ing the traditional nomenclature where the finger number means only the sequence of the examined fingers. If we
change the fingers nomenclature in this work, we will not match with all the previous Dendrophryniscus species
descriptions.
DNA-barcoding analysis. For the DNA-barcoding analysis, we adopted the 16S RNA ribosomal gene because
it has been comprehensively used as marker for molecular diversity assessment in amphibians (Vences et al. 2005;
Lyra et al. 2017). We amplified a fragment of 16S for 14 tissue samples collected by us or loaned from tissue col-
lections and complemented our dataset with 20 sequences of 16S available in GenBank (Table 1). New sequences
were also deposited in GenBank (Accession Number MK140790—MK140803, Table 1). For best understanding
of genetic divergences for the genus, we included data of all Dendrophryniscus species available. Only one species
described in this review located in Campo de Fruticultura da Bocaina, Municipality of São Jos do Barreiro, State
of São Paulo, Brazil, was not included in DNA-barcoding analysis (Table 1) because there was not tissue sample
available.
We extracted total genomic DNA from ethanol-preserved tissues using a DNeasy Blood & Tissue Kit (Qiagen
Inc., Valencia, CA, USA). We amplified by polymerase chain reaction (PCR) and sequenced one mitochondrial
fragment, spanning the 16S gene, from a subset of our samples using published primers (16S-AL 5-CGC CTG TTT
ATC AAA AAC AT-3 and 16S-BH 5-CCG GTC TGA ACT CAG ATC ACG T-3; Palumbi et al. 1991). PCRs were
carried out in a final volume of 20 µL containing: 20 ng template DNA, 1 X Buffer, 0.2 µm of each primer, 0.4 mΜ
dNTP mix, and 0.125 units of Taq DNA polymerase (Roche Diagnostics, Indianapolis, IN, USA). Amplification was
carried out with an initial denaturation for 5 min at 94°C, followed by 35 cycles (60 s denaturation at 94°C, 60 s an-
nealing at 50–52°C, 60 s extension at 72°C), and a final extension at 72°C for 5 min. PCR products were visualized
on an agarose gel, purified using ExoSAP-IT (USB Corp., Cleveland, OH, USA), and sequenced using a BigDye
Terminator Cycle Sequencing Kit v.3.1 (Applied Biosystems, Foster City, CA, USA) on an ABI Prism 3100 Genetic
Analyzer (Applied Biosystems). DNA sequences were edited using Sequencher 4.7 (Gene Codes Corp., Ann Arbor,
MI, USA).
Complementary sequences were assembled in the program Chromas Pro v. 2.1.8. (Technelysium Pty Ltda.
2018). Sequences alignment was computed with MAFFT v.7 (Katoh et al. 2017) following the FFT-NS-i strategy
and default parameters. After alignment, we eliminated regions of missing data and kept only the fragment ampli-
fied for all samples with 300 bp (true length from 289–296 bp because indels). We adopted the p-distance instead of
Kimura-two-parameter (K2P) as measure of genetic distance (Srivathsan & Meier 2011). We computed intraspecific
and interspecific p-distance with correspondent standard errors with 10,000 bootstrap replications and pairwise de-
letions for gaps and represented genetic distances by a Neighbor-Joining tree constructed in the program MEGA7:
Molecular Evolutionary Genetics Analysis version 7.0 (Kumar et al. 2016). We optimized the thresholds for our da-
taset and calculated barcode gaps between intraspecific and interspecific genetic distances (Meyer & Paulay 2005;
Meier et al. 2008) with spider package (Brown et al. 2012), in the R program (R Core Team 2018).
Results
Accounts on species
Dendrophryniscus brevipollicatus Jiménez de la Espada, 1870

(Figs. 1–2)
Dendrophryniscus brevipollicatus Jiménez de la Espada, 1870.

Dendrophryniscus brevipollicatus brevipollicatus—Miranda-Ribeiro 1926.
Dendrophryniscus brevipollicatus lutzi Miranda-Ribeiro, 1926.
Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 31

Dendrophryniscus b. travassoi P. Miranda-Ribeiro, 1955—nomen nudum.
Dendrophryniscus brevipollicatus—Cochran 1955 (part); Izecksohn 1994 (part).
Neotype. MNRJ 91627, adult male, collected at Açude da Solidão (22º57’S, 43º17’W, Datum WGS 84; 410 m
above sea level), Parque Nacional da Tijuca, Municipality of Rio de Janeiro, State of Rio de Janeiro, Brazil, by A.
L. Peracchi and E. Izecksohn, on 11 February1961.
Jiménez de la Espada (1870) did not designate the type specimens for D. brevipollicatus. Frost (2019) stated
that the types were presumably in the Museo Nacional de Ciencias Naturales, Madrid, Spain (MNCN). However,
according to the curator of the herpetological collection at MNCN, there are not extant type specimens of D. bre-
vipollicatus. Based on the Article 75 of the International Code of Zoological Nomenclature (ICZN 1999), we here
designate a neotype for the species.
The original type locality of D. brevipollicatus was quoted as Hab. in Brasil; prope Rio de Janeiro, in monte
Corcovado (Jiménez de la Espada 1870), and given as Corcovado, Rio de Janeiro, Guanabara by Bokermann
(1966). The current neotype designation sets the type locality for D. brevipollicatus in the same general original
region, in the Parque Nacional da Tijuca, Rio de Janeiro, RJ, Brazil.
Diagnosis. The species is characterized by: (1) medium size for the genus (SVL 18.3–25.1 mm in males, 20.9–
27.7 mm in females); (2) body slender; (3) snout mucronate in dorsal view; (4) canthus rostralis slightly curved;
(5) elliptical set of unpigmented pronounced granules posterior to the corner of mouth; (6) surfaces of upper eyelid
with numerous granules and with a conspicuous margin; (7) tip of third and fourth fingers laterally expanded; (8)
skinfold well developed on the articulation of the first and second phalanges of the fingers II, III, and IV; (9) male
with moderate nuptial pad with minuscule dark horny asperities on finger I; (10) fingers slightly fringed, webbed
only at base.
Figure 1. Dendrophryniscus brevipollicatus Jiménez de la Espada, 1870, neotype, MNRJ 91627 (SVL 20.6 mm). Dorsal and
ventral views.
Comparisons with other species. Dendrophryniscus brevipollicatus is distinguished from D. proboscideus by

the smaller size (SVL 18.3–25.1 mm in males of D. brevipollicatus; SVL 39.2–46.4 mm in males of D. probos-
cideus); D. brevipollicatus is distinguished from D. krausae and D. stawiarskyi by the slender body with uniform
shape (robust and enlarged posteriorly in those species); D. brevipollicatus is distinguished from D. berthalutzae, D.
carvalhoi, D. krausae, D. lauroi, D. oreites, D. proboscideus, D. skuki, and D. stawiarskyi by the snout mucronate in
dorsal view (snout rounded in D. berthalutzae, D. carvalhoi, D. lauroi, and D. stawiarskyi; snout truncate in dorsal
view in D. krausae, D. oreites, and D. proboscideus; snout long, narrow, spatulate, with parallel lateral borders and
rounded tip in D. skuki); by the canthus rostralis slightly curved, D. brevipollicatus is distinguished from D. davori

sp. nov., D. carvalhoi, D. krausae, D. lauroi, and D. stawiarskyi (canthus rostralis straight); D. brevipollicatus
presents an elliptical set of white granules posterior to the corner of mouth (longitudinal set in D. berthalutzae, D.
krausae, D. leucomystax, and D. oreites; two pronounced granules in D. imitator; absent in D. carvalhoi, D. haddadi
sp. nov., D. organensis, D. proboscideus, D. skuki, and D. stawiarskyi); D. brevipollicatus presents the surfaces of
upper eyelid with numerous granules densely distributed and with a conspicuous margin (few granules sparse and
external margin prominent in D. haddadi sp. nov. and D. imitator; few granules sparse and external margin con-
spicuous in D. davori sp. nov.; D. brevipollicatus is distinguished from D. berthalutzae, D. carvalhoi, D. haddadi
sp. nov., D. imitator, D. krausae, D. oreites, and D. stawiarskyi by the tip of the third and fourth fingers laterally
expanded (poorly or not expanded in those species); D. brevipollicatus is distinguished from D. davori sp. nov., D.
berthalutzae, D. carvalhoi, D. haddadi sp. nov., D. imitator, D. krausae, D. lauroi, D. leucomystax, D. oreites, and
D. stawiarskyi by the presence of a skinfold well developed on the articulation of the first and second phalanges of
the fingers II, III, and IV (poorly developed in those species); D. brevipollicatus is distinguished from D. davori
sp. nov., D. carvalhoi, D. izecksohni sp. nov., D. leucomystax, and D. stawiarskyi by the presence of moderately
developed nuptial pad in males (absent in D. leucomystax; very developed in D. davori sp. nov., D. carvalhoi, D.
izecksohni sp. nov. , and D. stawiarskyi); D. brevipollicatus is distinguished from D. berthalutzae, D. carvalhoi, D.
haddadi sp. nov., D. imitator, D. lauroi, D. oreites, D. skuki, and D. stawiarskyi by the fingers slightly fringed and
webbed only at base (fingers not fringed nor webbed in those species), from D. krausae (fingers not fringed and
webbed only at base), and from D. izecksohni sp. nov. and D. jureia sp. nov. (fingers fringed and webbed about one
third).
Figure 2. Dendrophryniscus brevipollicatus Jiménez de la Espada, 1870, neotype, MNRJ 91627 (SVL 20.6 mm). Dorsal and
lateral views of head, ventral views of hand and foot.

Description of neotype. Body slender, elongated (Fig. 1); head triangular, longer than wide, head length 33.5%
of SVL; snout mucronate in dorsal view, acute in lateral view (Fig. 2), snout 49.7% of head length; nostrils not pro-
tuberant, small, elliptical, located laterally at the beginning of the distal third of the eye to snout distance, slightly
below the canthus rostralis; internarial distance 65% of eye to snout distance; eye diameter 78% of eye to nostril
distance, 81% of interorbital distance; eye slightly protuberant; canthus rostralis slightly curved; loreal region
vertical; vocal sac indistinct; choanae small, circular, very far from each other; tongue long, narrow; vocal slits
present. Arms robust, forearms more robust than upper arms; hand with fingers slender, slightly fringed and webbed
only at base, distal end of the third and fourth fingers laterally expanded; finger I enlarged, covered by moderate
nuptial pad with minuscule dark horny asperities; relative lengths of fingers I<II<IV<III; subarticular tubercles
single, round; outer metacarpal tubercle large, rounded; inner metacarpal tubercle rounded, of the same size of sub-
articular tubercles; supernumerary tubercles present, small; skin fold developed on the articulation of the first and
second phalanges of the fingers II, III, and IV. Thigh length slightly shorter than tibia length; sum of thigh and tibia
lengths 80% of snout–vent length. Tarsal length 25% of the snout–vent length. Foot with toes slender, not fringed;
interdigital webbing only at the base, webbing formula: I1–2-II2-–3III2–31/2IV3–2V; distal end of toes globose, not
expanded laterally, relative lengths of toes I<II<III<V<IV; subarticular tubercles single, rounded, larger than those
of the fingers; outer metatarsal tubercle large, approximately rounded; inner metatarsal tubercle large, elliptical;
supernumerary tubercles absent. Dorsal and lateral surfaces covered by spinulose granules densely distributed, with
uniform different size; ventral surfaces rugose; presence of numerous granules on upper eyelid surface, with the
external margin prominent; elliptical set of white granules posterior to the corner of mouth.
Color in preservative. Dorsum pale brown; the characteristic pattern of a transverse brown bar on interorbital
region extending on the upper eyelids, an ‘X’ shaped brown blotch on the scapular region, an inverse ‘Y’ shaped
brown blotch on the sacral region, and two small blotches on the urostyle is only discernible.
Measurements of neotype. SVL 20.6; HL 6.9; HW 5.3; IND 1.5; ESD 3.4; ED 1.8; IOD 2.2; THL 8.0; TL 8.5;
TAL 5.2; FL 7.4.
Variation. Specimens are congruent with respect to the morphological characters. Variation of measurements
and descriptive statistics of males and females are presented in Tables 2 and 3. Sexual dimorphism can be observed
on SVL (longer in females), on arms (more robust in males), finger I (short and robust in males), and nuptial pad (on
finger I of males). Nuptial pad can be pigmented or not.
Distribution. Dendrophryniscus brevipollicatus is known from the following municipalities (Fig. 17) in the
State of Rio de Janeiro, Brazil: Cachoeiras de Macacu, Reserva Ecológica de Guapiaçu (22 o23’S, 42º43’W Datum
WGS 84), Engenheiro Paulo de Frontin (22o32’S, 43o40’W, Datum WGS 84; 400 m altitude), Magé (22o37’S,
43o03’W, Datum WGS 84; 100 m altitude), Nova Iguaçu (22o50’S, 43o37’W, Datum WGS 84; 45 m altitude), Rio
de Janeiro, Parque Nacional da Tijuca (22o57’S, 43o16’W, Datum WGS 84; 500 m altitude), Silva Jardim (22o39’S,
42o24’W, Datum WGS 84, 35m altitude).
Natural history. This species is associated to forested areas below 500 m altitude, breeding in Bromeliaceae
and Asparagaceae plants, where its larvae develop in the water deposited in leaves axillae (C.A.G. Cruz, personal
observation).
Remarks. Izecksohn (1971) described the vertebral pattern of this species.
Dendrophryniscus imitator (Miranda-Ribeiro, 1920), new status, revalidated

(Figs. 3–5)
Atelopus imitator Miranda-Ribeiro, 1920.

Dendrophryniscus brevipollicatus imitator—Miranda-Ribeiro 1926.
Dendrophryniscus brevipollicatus—Cochran 1955 (part); Heyer et al. 1990; Izecksohn 1994 (part); Malagoli et al. 2017.
Lectotype. MZUSP 847, adult female (Fig. 3), collected at Alto da Serra (23o46’S, 46o19’W, Datum WGS 84, 800
m altitude), Municipality of Paranapiacaba, State of São Paulo, Brazil, by Bicego in 1899.
Miranda-Ribeiro (1920) described Atelopus imitator based on five syntypes (MZUSP 477 and 485 from Cu-
batão, Santos; MZUSP 947, Alto da Serra; MZUSP 835, Rio Grande; MZUSP 639, Campo Grande; all localities in
the State of São Paulo). However, he described and referred measurements for one specimen (Corpo 23 mm., perna
28.), which matches the specimen MZUSP 847. This specimen is here designated lectotype for Atelopus imitator
Miranda-Ribeiro, 1920 (currently Dendrophryniscus imitator).

Currently, the color of the lectotype is fade because the time of preservation, and the body surfaces are homo-
geneous pale brown, the dorsal blotches not visible.
Measurements of lectotype. SVL 21.5; HL 7.4; HW 6.5; ED 2.4; THL 8.0; TL 8.9; TAL 4.8; FL 7.4.
Diagnosis. The species is characterized by: (1) medium size for the genus (SVL 15.1–17.4 mm in males,
14.4–21.5 mm in females); (2) body slender; (3) snout mucronate in dorsal view; (4) canthus rostralis slightly
curved; (5) presence of two pronounced granules posterior to the corner of mouth; (6) surfaces of upper eyelid with
few granules sparse and external margin prominent; (7) tip of the third and fourth fingers poorly or not expanded
laterally; (8) skinfold poor developed on the articulation of the first and second phalanges of the fingers II, III, and
IV (9) male with moderate nuptial pad with minuscule horny unpigmented asperities on finger I; (10) fingers not
fringed nor webbed.
Figure 3. Dendrophryniscus imitator (Miranda-Ribeiro, 1920), lectotype, MZUSP 847 (SVL 21.5 mm). Dorsal and ventral
views.
Comparisons with other species. Dendrophryniscus imitator is distinguished from D. proboscideus by the
smaller size (SVL 15.1–17.4 mm in males of D. imitator; SVL 39.2–46.4 mm in males of D. proboscideus); D.
imitator is distinguished from D. krausae and D. stawiarskyi by the slender body with uniform shape (robust and
enlarged posteriorly in those species); D. imitator is distinguished from D. berthalutzae, D. carvalhoi, D. krausae,
D. lauroi, D. oreites, D. proboscideus, D. skuki, and D. stawiarskyi by the snout mucronate in dorsal view (snout
rounded in D. berthalutzae, D. carvalhoi, D. lauroi, and D. stawiarskyi; snout truncate in dorsal view in D. krausae,
D. oreites, and D. proboscideus; snout long, narrow, spatulate, with parallel lateral borders and rounded tip in D.
skuki); by the canthus rostralis slightly curved, D. imitator is distinguished from D. davori sp. nov., D. carvalhoi,
D. krausae, D. lauroi, and D. stawiarskyi (canthus rostralis straight); the presence of two white pronounced gran-
ules posterior to the corner of mouth distinguishes D. imitator from D. lauroi (elliptical set of unpigmented shallow
granules posterior to the corner of mouth), D. brevipollicatus (elliptical set of white pronounced granules), and D.
berthalutzae, D. krausae, D. leucomystax, and D. oreites (longitudinal set); D. carvalhoi, D. haddadi sp. nov., D.
organensis, D. proboscideus, D. skuki, and D. stawiarskyi (set absent); D. imitator presents the surfaces of upper

eyelid with few granules sparse and external margin prominent (surfaces of upper eyelid with numerous granules
densely distributed and with a conspicuous margin in D. brevipollicatus and D. izecksohni sp. nov.; surfaces of up-
per eyelid with numerous granules with the external margin slightly prominent in D. lauroi; surfaces of upper eyelid
with few granules sparse and external margin conspicuous in D. davori sp. nov.); D. imitator is distinguished from
D. brevipollicatus, D. lauroi, and D. organensis by the tip of the third and fourth fingers poorly or not expanded
laterally (laterally expanded in those species); D. imitator is distinguished from D. brevipollicatus by the presence
of a skinfold poorly developed on the articulation of the first and second phalanges of the fingers II, III, and IV (well
developed in this species); D. imitator is distinguished from D. davori sp. nov., D. carvalhoi, D. izecksohni sp. nov.,
D. leucomystax, and D. stawiarskyi by the presence of moderate nuptial pad in male (absent in D. leucomystax; very
developed in D. davori sp. nov., D. carvalhoi, D. izecksohni sp. nov., and D. stawiarskyi); by the fingers not fringed
nor webbed, D. imitator is distinguished from D. davori sp. nov. and D. brevipollicatus (fingers slightly fringed and
webbed only at base), D. izecksohni sp. nov. and D. jureia sp. nov. (fingers fringed and webbed about one third),
and D. krausae (fingers not fringed and webbed only at base).
Original description of type series (Atelopus imitator Miranda-Ribeiro 1920, freely translated from Portu-
guese)
Elongated, with the humerus shorter than the transverse diameter of the neck. Snout compressed and depressed,
with lateral nostrils near the tip of snout, canthus rostralis evident, giving to loreal region a concave shape and
continuing by the upper eyelid. Leg carried forward, the tarsal articulation almost reaching the posterior angle of
the eye. Fingers as in A. cruciger; toes same, that is, connecting through a rudimentary membrane, between the first
extension is in the order: 1, 2, 4 and 3, and between second 1, 2, 3, 5 and 4. Body more granular in larger individuals,
smoother in the shorter. Color pale cinereous or yellowish. A wide transverse stripe between the eyes; and from the
vertex of the triangle that is posterior leave two divergent arms backwards, one or two further back, a dark stripe,
on the flanks, starting from the snout and finishing in the iliac articulation, bordered of white on its sides; members
with transversal stripes. Sometimes a narrow vertebral line, white. Abdominal side dark marbled. Body 23 mm, leg
28 mm.
Description of topotype. MZUSP 103992, adult male collected in Paranapiacaba, State of São Paulo, on March
24, 1963 by W.C.A. Bokermann and A.B. Costa.
Figure 4. Dendrophryniscus imitator (Miranda-Ribeiro, 1920), topotype, MZUSP 103992 (SVL 17.0 mm). Dorsal and ven-
tral views.

Body slender, elongated (Fig. 4); head triangular, longer than large, head length 36.4% of SVL; snout mucro-
nate in dorsal view, acute in lateral view (Fig. 5); snout 46.7% of head length; presence of a small tubercle under
the tip of snout, nostrils not protuberant, small and elliptical, located laterally near the tip of snout, below the can-
thus rostralis; internarial distance 48.2% of eye to snout distance; eye diameter 58.6% of eye to snout distance and
77.2% of interorbital distance; eye protuberant; canthus rostralis slightly curved; loreal region vertical; vocal sac
indistinct; choanae small, circular, very far from each other; tongue long, narrow, and few enlarged posteriorly;
vocal slits present. Arms robust, forearms more robust than upper arms; hand with fingers slender, not fringed nor
webbed; distal end of the third and fourth fingers slightly expanded laterally; finger I enlarged, covered by moderate
nuptial pad with minuscule horny unpigmented asperities; relative lengths of fingers I<II<IV<III; subarticular tu-
bercles single, rounded; outer metacarpal tubercle large, rounded; inner metacarpal tubercle rounded, slightly small
than subarticular tubercles; supernumerary tubercles present, small; skinfold poor developed on the articulation of
the first and second phalanges of the fingers II, III, and IV. Thigh length slightly shorter than tibia length; sum of
thigh and tibia lengths about 93.5% of snout–vent length. Tarsal length about 23% of the snout–vent length. Foot
with toes slender, not fringed; webbing formulae: I1–2+II11/2–3III2–3IV31/2–21/2V; distal end of toes globose, not
expanded laterally, relative lengths of toes I<II<III<V<IV; subarticular tubercles single, rounded, the same size of
those of the fingers; outer metatarsal tubercle with the same size of subarticular tubercles, rounded; inner metatarsal
tubercle large, approximately twice the outer, elongated; supernumerary tubercles present but scarce. Dorsal and
lateral surfaces covered by spinulose granules uniformly distributed, with different sizes, ventral surfaces covered
by numerous rounded granules; surfaces of upper eyelid with few granules sparse and external margin prominent;
presence of two pronounced granules posterior to the corner of mouth.
Figure 5. Dendrophryniscus imitator (Miranda-Ribeiro, 1920), topotype, MZUSP 103992 (SVL 17.0 mm). Dorsal and lat-
eral views of head, ventral views of hand and foot.

Color in preservative. The topotypes present dorsum pale brown; a head blotch extending on the upper eyelids
merging with an ‘X’ shaped blotch on the scapular region and an inverse ‘Y’ shaped blotch on the sacral region,
extending to the inguinal region; one transversal bar on forearms and one blotch on elbow; one transversal bar on
thigh, tibia, and tarsus, and one blotch on knee and heel; a lateral stripe starting at the snout, passing on the eye
and reaching the inguinal region, being narrower in the loreal region; all of the bars, blotches, and stripes are dark
brown; ventral surface lighter than the dorsum, with disperse small blotches.
Measurements of topotype. SVL 17.0; HL 6.2; HW 4.8; IND 1.4; ESD 2.9; ED 1.7; IOD 2.2; THL 7.7; TL 8.2;
TAL 3.9; FL 6.8.
on SVL (larger in female), on arms (slightly more robust in male), finger I (robust in male), and nuptial pad (on
finger I of male). Nuptial pad can be pigmented or not.
Distribution. Dendrophryniscus imitator is known from the following municipalities (Fig. 17) in the State of
São Paulo, Brazil: Cubatão (23o51’S, 46o27’W, Datum WGS 84), Juquitiba (23o 55’S, 47oo3’W, Datum WGS 84),
Itanhaém, Parque Estadual da Serra do Mar Núcleo Curucutu (23o59’S, 46o44’W, Datum WGS 84), Natividade da
Serra (23o22’S, 45o26’W, Datum WGS 84), Salesópolis (23o38’S, 45o52’W, Datum WGS 84, 900 m altitude), San-
to André, Paranapiacaba (23o46’S, 46o17’W, Datum WGS 84), Santos (23o56’S, 46o19’W, Datum WGS 84), São
Bernardo do Campo (23º45’S, 46º33’W, Datum WGS 84), São Paulo, Represa Billings (23o48’S, 46o37’W, Datum
WGS 84), Cotia, Reserva do Morro Grande (23o43’S, 46o57’W, Datum WGS 84).
Natural history. This species is associated to forested areas, from 100 m to 1100 m above sea level, and breeds
in bromeliads, where its larvae develop (Malagoli et al. 2017, as D. brevipollicatus).
Remarks. All individuals of the type series have tip of snout kneaded. All skin is unpigmented and surface
granulation and crenulations are few evident because time of preservation. For this reason, we presented the origi-
nal description and made a new description based on a topotype from Paranapiacaba, State of São Paulo. In the
original description the author refers to five specimens composing the type series: two from Santos, Cubatão, São
Paulo (MZUSP 477–485) collected by Bicego on December 1897; one from Alto da Serra (MZUSP 847), collected
by Bicego in 1899; one from Rio Grande, State of São Paulo (MZUSP 835), collected by Wacket on July 1902;
and one from Campo Grande, State of São Paulo (MZUSP 639), collected by Wacket on May 1902. The specimen
MZUSP 835 from Rio Grande was lost and the tag number of the specimen from Campo Grande is MZUSP 634.
We concluded that body measures presented by Miranda-Ribeiro in the original description were based on specimen
MZUSP 847 from Alto da Serra (now, Paranapiacaba).
The specimens referred by Heyer et al. (1990) and Malagoli et al. (2017) as Dendrophryniscus brevipollicatus
are now D. imitator.
Dendrophryniscus lauroi Miranda-Ribeiro, 1926, new status, revalidated

(Figs. 6–8)
Dendrophryniscus brevipollicatus lauroi Miranda-Ribeiro, 1926.

Dendrophryniscus b. travassoi P. Miranda-Ribeiro, 1955—nomen nudum.
Dendrophryniscus brevipollicatus—Cochran 1955 (part); Izecksohn 1994 (part).
Lectotype. MNRJ 394, adult female (Fig. 6), collected at Municipality of Angra dos Reis (22o54’S, 44o20’W, Da-
tum WGS 84; 25 m altitude), State of Rio de Janeiro, Brazil, by Lauro Travassos.
Miranda-Ribeiro (1926), in the original description, referred to two typos (syntypes). P. Miranda-Ribeiro (1955)
found only one specimen, which was designated lectotype. Currently the lectotype (SVL 18.2 mm) is in poor condi-
tion, fade, with a strong transversal sulcus caused by an early line tied behind the head, and right leg broken.
Diagnosis. The species is characterized by: (1) medium size for the genus (SVL 14.5–19.9 mm in males, 19.0
mm in female); (2) body slender; (3) snout rounded in dorsal view; (4) canthus rostralis straight; (5) elliptical set of
unpigmented shallow granules posterior to the corner of mouth; (6) presence of numerous granules on upper eyelid
surfaces, with the external margin slightly prominent; (7) tip of the third and fourth fingers laterally expanded (8)
skinfold poor developed on the articulation of the first and second phalanges of the fingers II, III, and IV; (9) male
with moderate nuptial pad with minuscule and pigmented horny asperities on finger I; (10) fingers not fringed nor
webbed.

Figure 6. Dendrophryniscus lauroi Miranda-Ribeiro, 1926, lectotype, MNRJ 394 (SVL 18.2 mm). Dorsal view.
Comparisons with other species. Dendrophryniscus lauroi is distinguished from D. proboscideus by the smaller
size (SVL 14.5–19.9 mm in males of D. lauroi; SVL 39.2–46.4 mm in males of D. proboscideus); D. lauroi is
distinguished from D. krausae and D. stawiarskyi by the slender body with uniform shape (robust and enlarged
posteriorly in those species); D. lauroi is distinguished from D. davori sp. nov., D. brevipollicatus, D. haddadi sp.
nov., D. imitator, D. izecksohni sp. nov., D. krausae, D. oreites, D. organensis, D. proboscideus, and D. skuki by
the snout rounded in dorsal view (snout mucronate in dorsal view in D. davori sp. nov., D. brevipollicatus, D. had-
dadi sp. nov., D. imitator, D. izecksohni sp. nov., and D. organensis; snout truncate in dorsal view in D. krausae,
skuki); by the canthus rostralis straight, D. lauroi is distinguished from D. berthalutzae, D. brevipollicatus, D. had-
dadi sp. nov., D. imitator, D. izecksohni sp. nov., D. oreites, D. organensis, D. proboscideus, and D. skuki (canthus
rostralis slightly curved); D. lauroi presents an elliptical set of unpigmented shallow granules posterior to the corner
of mouth (elliptical set of white, pronounced granules in D. brevipollicatus; longitudinal set in D. berthalutzae, D.

krausae, D. leucomystax, and D. oreites; two pronounced granules in D. imitator; absent in D. carvalhoi, D. had-
dadi sp. nov., D. organensis, D. proboscideus, D. skuki, and D. stawiarskyi); D. lauroi presents the surfaces of up-
per eyelid with numerous granules and with the external margin slightly prominent (surfaces of upper eyelid with
numerous granules densely distributed and with a conspicuous margin in D. brevipollicatus and D. izecksohni sp.
nov.; surfaces of upper eyelid with few granules sparse and external margin prominent in D. haddadi sp. nov. and D.
imitator; surfaces of upper eyelid with few granules sparse and external margin conspicuous in D. davori sp. nov.);
D. lauroi is distinguished from D. davori sp. nov., D. berthalutzae, D. carvalhoi, D. haddadi sp. nov., D. imitator,
D. krausae, D. oreites, and D. stawiarskyi by the tip of the third and fourth fingers laterally expanded (poorly or not
expanded in those species); D. lauroi is distinguished from D. brevipollicatus by the presence of a skinfold poorly
developed on the articulation of the first and second phalanges of the fingers II, III, and IV (well developed in this
species); D. lauroi is distinguished from D. davori sp. nov., D. carvalhoi, D. izecksohni sp. nov., D. leucomystax,
and D. stawiarskyi by the presence of moderate nuptial pad in male (absent in D. leucomystax, very developed in
D. davori sp. nov., D. carvalhoi, D. izecksohni sp. nov., and D. stawiarskyi); D. lauroi is distinguished from D.
brevipollicatus by the fingers not fringed nor webbed (slightly fringed and webbed only at base in this species), from
D. krausae (fingers not fringed and webbed only at base), and from D. izecksohni sp. nov. and D. jureia sp. nov.
(fingers fringed and webbed about one third).
Original description (freely translated from Portuguese). “B—D. b. lauroi—Specimens from Angra dos Reis,
collected by Dr. Lauro Travassos present two types, with weakening of xyphisternum, absence of supranasal groove,
and body more slender.
Description of topotype. MNRJ 58298, adult male, collected at Ilha Grande (23o10’S, 44o09’W, Datum WGS
84, 100 m altitude), Municipality of Angra dos Reis, State of Rio de Janeiro, Brazil, on 03 January 1997.
Figure 7. Dendrophryniscus lauroi Miranda-Ribeiro, 1926, topotype, MNRJ 58298 (SVL 19.2 mm). Dorsal and ventral
views.
Body slender, elongated (Fig. 7); head triangular, longer than large, head length 30.7% of SVL; snout rounded
in dorsal view, acute in lateral view (Fig. 8); snout 49.1% of head length; nostrils not protuberant, small, ellipti-
cal, located laterally near the tip of snout, slightly below the canthus rostralis; internarial distance 55.1% of eye
to snout distance; eye diameter 75.8% of eye to snout distance and 91.6% of interorbital distance; eye slightly
protuberant; canthus rostralis straight; loreal region vertical; vocal sac indistinct; choanae small, circular, very far

from each other; tongue long, narrow. Vocal slits present. Arms robust, forearms more robust than upper arms; hand
with fingers slender, not fringed nor webbed, distal end of III and IV fingers slightly expanded laterally; finger I
enlarged, covered by moderate nuptial pad with minuscule pigmented horny asperities; relative lengths of fingers
I<II<IV<III; subarticular tubercles single, rounded; outer metacarpal tubercle large, rounded; inner metacarpal tu-
bercle rounded, slightly smaller than subarticular tubercles; supernumerary tubercles present, small and scarce,
skin fold poorly developed on the articulation of the first and second phalanges of the fingers II, III, and IV. Thigh
length slightly shorter than tibia length; sum of thigh and tibia lengths 83.5% of snout–vent length. Tarsal length
28% of the snout vent length. Foot with slender toes, not fringed; interdigital webbing only at the base, webbing
formulae: I1–2II11/2–3III2–21/2IV21/2–2V; distal end of toes globose, not expanded laterally, relative lengths of toes
I<II<III<V<IV; subarticular tubercles single, rounded, larger than those of fingers; outer metatarsal tubercle large,
approximately rounded; inner metatarsal tubercle large, elliptical; supernumerary tubercles present, small, scarce.
Dorsal and lateral surfaces covered by numerous spinulose shallow granules, uniformly distributed, ventral surfaces
rugose; presence of numerous granules on upper eyelid surfaces, with the external margin slightly prominent; pres-
ence of elliptical set of unpigmented shallow granules posterior to the corner of mouth.
Figure 8. Dendrophryniscus lauroi Miranda-Ribeiro, 1926, topotype, MNRJ 58298 (SVL 19.2 mm). Dorsal and lateral
views of head, ventral views of hand and foot.

Color in preservative. The color of the lectotype is fade because of the time of preservation, and the body
surfaces are homogeneously pale brown, being not possible to observe the body blotches. The topotype presents
dorsum pale brown background and brown blotches and bars; a head blotch extending on the upper eyelids, an ‘X’
shaped blotch on the scapular region, and an inverse ‘Y’ shaped blotch on the sacral region; on forearms, one trans-
versal bar and one blotch on elbow; one transversal bar on thigh, tibia, and tarsus, and one blotch on knees and heels;
a lateral dark brown stripe from the snout, passing on the eye and reaching the inguinal region, being narrower in
the loreal region; ventral surface pale cream, with disperse small brown blotches.
Measurements of topotype. SVL 19.2; HL 5.9; HW 5.7; IND 1.6; ESD 2.9; ED 2.2; IOD 2.4; THL 8.1; TL 8.4;
TAL 5.2; FL 6.7.
Variation. Specimens are congruent respecting morphological characters. Variation in size and shape of the
brown blotches on dorsal surfaces can be observed. Variation of measurements and descriptive statistics of males
and females are presented in Tables 2 and 3. Sexual dimorphism can be observed on SVL (longer in females), on
arms (more robust in males), finger I (robust in males), and nuptial pad (on finger I of males). Nuptial pad can be
pigmented or not.
Distribution. Dendrophryniscus lauroi is known from the following municipalities (Fig. 17) in the State of Rio
de Janeiro, Brazil: Angra dos Reis, Ilha Grande (22o10’S, 44o09’W, Datum WGS 84; 100 m altitude), Angra dos
Reis, Fazenda Floresta (22o54’S, 44o20’W, Datum WGS 84; 25 m altitude), and Mangaratiba (22o59’S, 44o06’W,
Datum WGS 84; 80 m altitude); and State of São Paulo, Municipality of Bananal (22o41’S, 44o19’W, Datum WGS
84; 500 m altitude).
Natural history. This species is associated to forested areas and restinga formations, from few meters to 500 m
above sea level, and breeds in bromeliads plants, where its larvae develop (D. Vrcibradic and C.A.G. Cruz personal
observations). Tadpoles were described in Izecksohn and Cruz (1972) as D. brevipollicatus.
Remarks. An illustration of D. lauroi was presented by Miranda-Ribeiro (1926, fig.72, pg. 138).
Dendrophryniscus davori sp. nov.

(Figs. 9–10)
Holotype. MNRJ 58300, adult male, collected at Baixo Caledônia (22º21’S, 42º35’W, Datum WGS 84; 1600 m
altitude), Municipality of Nova Friburgo, State of Rio de Janeiro, Brazil, by T.A. Dorigo, C.C. Siqueira and D.
Vrcibradic, on 17 March 2009.
Paratypes. MNRJ 58301, adult female, and MNRJ 58302, adult male, collected with the holotype; MNRJ
60700, juvenile, collected at the type locality, by D. Vrcibradic, on 01 November 2009.
Diagnosis. The species is characterized by: (1) medium size for the genus (SVL 18.9–22.1 mm in males, 24.2
mm in female); (2) body slender, elongated; (3) snout mucronate in dorsal view; (4) canthus rostralis straight with a
tubercle nearer to the eye than to the snout; (5) presence of a longitudinal set of white granules behind the corner of
the mouth; (6) surfaces of upper eyelid with few granules sparse and external margin conspicuous; (7) tip of the third
and fourth fingers poor expanded laterally; (8) skinfold poor developed on the articulation of the first and second
phalanges of the fingers II, III, and IV ; (9) male with a very developed nuptial pad with horny asperities conspicu-
ous and pigmented on finger I (10) fingers slightly fringed and webbed only at base.
Comparisons with other species. Dendrophryniscus davori sp. nov. is distinguished from D. proboscideus by
the smaller size (SVL 18.9–22.1 mm in males; SVL 39.2–46.4 mm in males of D. proboscideus); D. davori sp. nov.
is distinguished from D. krausae and D. stawiarskyi by the slender body with uniform shape (robust and enlarged
posteriorly in those species); D. davori sp. nov. is distinguished from D. berthalutzae, D. carvalhoi, D. krausae,
skuki); by the canthus rostralis straight D. davori sp. nov. is distinguished from D. berthalutzae, D. brevipollicatus,
D. haddadi, D. imitator, D. izecksohni, D. jureia, D. oreites, D. organensis, D. proboscideus, and D. skuki (canthus
rostralis slightly curved); by the canthus rostralis with a tubercle nearer to the eye than the nostril, D. davori sp.
nov. differs from all congeneric species (canthus rostralis without tubercle); the presence of longitudinal set of
pronounced granules posterior to the corner of mouth distinguishes D. davori sp. nov. from D. lauroi (elliptical

set of shallow granules), D. brevipollicatus, D. izecksohni., and D. jureia (elliptical set of pronounced granules),
D. imitator (two pronounced granules), and absent in D. carvalhoi, D. haddadi, D. organensis, D. proboscideus,
D. skuki, and D. stawiarskyi; D. davori sp. nov. presents the surfaces of upper eyelid with few granules sparse and
external margin conspicuous (surfaces of upper eyelid with numerous granules and with a conspicuous margin in
D. brevipollicatus and D. izecksohni); D. davori sp. nov. is distinguished from D. brevipollicatus, D. lauroi, and D.
organensis by the tip of the third and fourth fingers poorly or not expanded laterally (laterally expanded in those
species); D. davori sp. nov. is distinguished from D. brevipollicatus by the presence of a skinfold poorly developed
on the articulation of the first and second phalanges of the fingers II, III, and IV (well developed in this species);
D. davori sp. nov. is distinguished from D. brevipollicatus, D. imitator, D. jureia, D. lauroi, and D. leucomystax
by the presence of very developed nuptial pad in male (absent in D. leucomystax, poor developed in D. jureia, and
moderate in D. brevipollicatus, D. haddadi, D. imitator, and D. lauroi); D. davori sp. nov. is distinguished from D.
berthalutzae, D. carvalhoi, D. haddadi, D. izecksohni, D. krausae, D. lauroi, D. oreites, D. skuki, and D. stawiarskyi
by the fingers slightly fringed and webbed only at base (fingers fringed and webbed about one third in D. izecksohni
and D. jureia; fingers not fringed and webbed only at base in D. krausae; not fringed nor webbed in D. berthalutzae,
D. carvalhoi, D. haddadi, D. imitator, D. lauroi, D. oreites, D. skuki, and D. stawiarskyi).
Figure 9. Dendrophryniscus davori sp. nov., holotype, MNRJ 58300 (SVL 22.1 mm). Dorsal and ventral views.
Description of holotype. Body slender, elongated (Fig. 9); head triangular, longer than large, head length 34.8%
of SVL; snout mucronate in dorsal view, acute in lateral view (Fig. 10); snout 51.9% of head length; presence of a
small tubercle under the tip of snout in ventral view; nostrils not protuberant, small and elliptical, located laterally
near the tip of snout, below the canthus rostralis; internarial distance 47.5% of eye to snout distance; eye diameter
50% of eye to snout distance and 71% of interorbital distance; eye protuberant; canthus rostralis almost straight
with a tubercle nearer to the eye than the nostril; loreal region vertical; choanae small, circular, and very far from
witch one; tongue long and few enlarged posteriorly; vocal sac indistinct; vocal slits present. Arms strongly robust;
hand with fingers robust, slightly fringed and webbed only at base, distal end of fingers globose, tip of the third and
fourth finger slightly expanded laterally; male with a very developed nuptial pad with conspicuous and pigmented
horny asperities on finger I; relative lengths of fingers I<II<IV<III; subarticular tubercles single, rounded; outer
metacarpal tubercle large, oval; inner metacarpal tubercle elliptical, almost the same size of outer metacarpal tu-
bercle; supernumerary tubercles few, small, and disperse; skinfold few developed on the articulation of the first and
second phalanges of the fingers II, III, and IV. Thigh length slightly shorter than tibia length; sum of thigh and tibia
lengths about 81% of snout–vent length. Tarsal length about 25% of the snout vent length. Foot with toes robust,
not fringed; interdigital webbing moderately developed, webbing formulae: I1–2II1–3III2–4IV3–21/2V; distal end of
toes globose, not expanded laterally, relative lengths of toes I<II<III<V<IV; subarticular tubercles single, rounded,
bigger than those of the fingers; outer metatarsal tubercle small, elliptical; inner metatarsal tubercle oval and large,

approximately twice the outer; supernumerary tubercles numerous. Dorsal and lateral surfaces covered by spinulose
granules, irregularly distributed, with different sizes; ventral surfaces densely covered by prominent white granules,
regularly distributed; surfaces of upper eyelid with few granules sparse and external margin conspicuous; presence
of a longitudinal set of white granules behind the corner of the mouth.
Color in preservative. Dorsum grayish light to dark brown , delimited by a whitish margin; a head blotch ex-
tending on the upper eyelids merging with an ‘X’ shaped blotch on the scapular region and an inverse ‘Y’ shaped
blotch on the sacral region (fade on the holotype) extending to the inguinal region; one transversal bar on forearm
and one blotch on elbow; one transversal bar on thigh, tibia, and tarsus, and one small blotch on heel; a lateral
stripe starting at the snout, passing on the eye and reaching the inguinal region; all bars, blotches, and stripes are
dark brown; a marked white blotch under the eye and white tubercles at the corner of the mouth; ventral surface
uniformly whitish.
Figure 10. Dendrophryniscus davori sp. nov., holotype, MNRJ 58300 (SVL 22.1 mm). Dorsal and lateral views of head,
ventral views of hand and foot.

Measurements of holotype. SVL 22.1; HL 7.7; HW 6.7; IND 1.9; ESD 4.0; ED 2.0; IOD 2.8; THL 8.8; TL 9.2;
TAL 5.6; FL 9.2.
Variation. Specimens are congruent with respect to the morphological characters. Measurements of male and
female are presented in Tables 2 and 3; juvenile paratype measured 15.7 mm SVL. Sexual dimorphism can be ob-
served on SVL (larger in female), on arms (slightly more robust in male), finger I (robust in male), and nuptial pad
(on finger I of male). On the holotype the dorsal blotches are less evident.
Distribution. Dendrophryniscus davori is known only from the type locality (Fig. 17), Baixo Caledônia, Mu-
nicipality of Nova Friburgo, State of Rio de Janeiro, Brazil.
Natural history. This species is associated to forested areas, at 1600 m above sea level.
Etymology. The name of the species honors our friend Dr. Davor Vrcibradic, from Universidade Federal do
Estado do Rio de Janeiro (UNIRIO), by his contribution to the knowledge of Brazilian amphibians and reptiles.
Dendrophryniscus haddadi sp. nov.

(Figs. 11–12)
Holotype. CFBH 09300, adult male, collected at Parque Estadual da Serra do Mar, Núcleo Santa Virgínia (23o21’S,
45o08’W, Datum WGS 84; 970 m altitude), Municipality of São Luís do Paraitinga, State of São Paulo, Brazil, by
Luís O.M. Giasson, on 16 June 2005.
Paratypes. All specimens collected at the type locality by Luis O.M. Giasson, on several dates: CFBH 09302,
adult female, on 16 June 2005; CFBH 09872, adult female; CFBH 09874, adult female, both on 30 October 2005;
CFBH 10780, adult female, on 23 January 2006; CFBH 12647, young female, on February 2005; CFBH 14877,
adult male; CFBH 14878, adult female; CFBH 14879, young female, all collected on 19 December 2006; CFBH
16320, adult male, on 13 March 2005; CFBH 16321, adult male; on 17 August 2005;16322, adult male, on 01 Oc-
tober 2005; CFBH 16325, adult male, on 26 November 2005. ,
(5) absence of a set of pronounced granules posterior to the corner of mouth; (6) surfaces of upper eyelid with few
granules sparse and external margin prominent; (7) tip of the third and fourth fingers poorly or not expanded later-
ally; (8) skinfold poor developed on the articulation of the first and second phalanges of the fingers II, III, and IV; (9)
male with moderate nuptial pad with minuscule light brown horny unpigmented asperities on finger I; (10) fingers
not fringed nor webbed.
Comparisons with other species. Dendrophryniscus haddadi sp. nov. is distinguished from D. proboscideus by
the smaller size (SVL 15.7–16.8mm in males of D. haddadi sp. nov.; SVL 39.2–46.4 mm in males of D. probosci-
deus); D. haddadi sp. nov. is distinguished from D. krausae and D. stawiarskyi by the slender body with uniform
shape (robust and enlarged posteriorly in those species); D. haddadi sp. nov. is distinguished from D. berthalutzae,
D. carvalhoi, D. krausae, D. lauroi, D. oreites, D. proboscideus, D. skuki, and D. stawiarskyi by the snout mucro-
nate in dorsal view (snout rounded in D. berthalutzae, D. carvalhoi, D. lauroi, and D. stawiarskyi; snout truncate
in dorsal view in D. krausae, D. oreites, and D. proboscideus; snout long, narrow, spatulate, with parallel lateral
borders and rounded tip in D. skuki); by the canthus rostralis slightly curved, D. haddadi sp. nov. is distinguished
from D. davori sp. nov., D. carvalhoi, D. krausae, D. lauroi, and D. stawiarskyi (canthus rostralis straight); the
absence of a set of white pronounced granules posterior to the corner of mouth distinguishes D. haddadi sp. nov.
from D. lauroi (elliptical set of unpigmented shallow granules posterior to the corner of mouth), D. brevipollicatus
(elliptical set of white pronounced granules), D. berthalutzae, D. krausae, D. leucomystax, and D. oreites (longitu-
dinal set), from D. imitator (two pronounced granules); D. haddadi sp. nov. presents the surfaces of upper eyelid
with few granules sparse and external margin prominent (surfaces of upper eyelid with numerous granules densely
distributed and with a conspicuous margin in D. brevipollicatus and D. izecksohni sp. nov.; surfaces of upper eyelid
with numerous granules with the external margin slightly prominent in D. lauroi; surfaces of upper eyelid with few
granules sparse and external margin conspicuous in D. davori sp. nov.); D. haddadi sp. nov. is distinguished from
D. brevipollicatus, D. lauroi, and D. organensis by the tip of the third and fourth fingers poorly or not expanded
laterally (laterally expanded in those species); D. haddadi sp. nov. is distinguished from D. brevipollicatus by the
presence of a skinfold poorly developed on the articulation of the first and second phalanges of the fingers II, III,

and IV (well developed in this species); D. haddadi sp. nov. is distinguished from D. davori sp. nov., D. carvalhoi,
D. leucomystax, and D. stawiarskyi by the presence of moderate nuptial pad in male (absent in D. leucomystax; very
developed in D. davori sp. nov., D. carvalhoi, D. izecksohni sp. nov., and D. stawiarskyi); by the fingers not fringed
nor webbed, D. haddadi sp. nov. is distinguished from D. davori sp. nov. and D. brevipollicatus (fingers slightly
fringed and webbed only at base), D. izecksohni sp. nov. and D. jureia sp. nov. (fingers fringed and webbed about
one third), D. krausae (fingers not fringed and webbed only at base); by the forearms as robust as upper arms D.
haddadi sp. nov. is distinguished from D. imitator (forearms more robust than upper arms).
Figure 11. Dendrophryniscus haddadi sp. nov., holotype, CFBH 09300 (SVL 16.8 mm). Dorsal and ventral views.
Description of the holotype. Body slender, elongated (Fig. 11); head triangular, longer than large, head length
36.3% of SVL; snout mucronate in dorsal view, acute in lateral view (Fig. 12); snout 49.2% of head length; nostrils
not protuberant, small and elliptical, located laterally near the tip of snout, below the canthus rostralis; internarial
distance 43.3% of eye to snout distance; eye diameter 56.6% of eye to snout distance and 73.9% of interorbital
distance; eye protuberant; canthus rostralis slightly curved; loreal region vertical; choanae small, circular, and
very far from which one; tongue long, narrow, and few enlarged posteriorly; vocal sac indistinct; vocal slits pres-
ent. Arms robust, forearms as robust as upper arms; hand with fingers slender, not fringed nor webbed; distal end
of the third and fourth fingers poorly expanded laterally; finger I enlarged, covered by moderate nuptial pad with
minuscule light brown horny unpigmented asperities; relative lengths of fingers, I<II<IV<III; subarticular tubercles
single, rounded; outer metacarpal tubercle large, rounded; inner metacarpal tubercle rounded, slightly smaller than
subarticular tubercles; supernumerary tubercles present, small; skinfold poorly developed on the articulation of
the first and second phalanges of the fingers III and IV. Thigh length slightly larger than tibia length; sum of thigh
and tibia lengths 85.1% of snout–vent length. Tarsal length 26.2% of the snout vent length. Foot with toes slender,
not fringed; interdigital webbing only at the base, webbing formulae: I1–2+II1–3III2-–3IV3–2V; distal end of toes
globose, not expanded laterally, relative lengths of toes, I<II<III<V<IV; subarticular tubercles single, rounded, the
same size of those of the fingers; outer metatarsal tubercle slightly larger than subarticular tubercles, rounded; inner
metatarsal tubercle large, approximately 1.5 times the outer, elliptical; supernumerary tubercles present but scarce.
Dorsal and lateral surfaces covered by numerous, prominent spinulose granules uniformly distributed, with uniform
size, ventral surfaces covered by numerous rounded granules; surfaces of upper eyelid with few granules sparse and
external margin prominent; absence of a set of pronounced granules posterior to the corner of mouth.

Color in preservative. Dorsum brown ; a head blotch extending on the upper eyelids merging with an ‘X’ shaped
blotch on the scapular region and an inverse ‘Y’ shaped blotch on the sacral region, extending to the inguinal region;
one transversal bar on forearms and one blotch on elbow; one transversal bar on thigh, tibia, and tarsus, and one
blotch on knee and heel; a lateral stripe starting at the snout, passing on the eye and reaching the inguinal region,
being narrower in the loreal region; all of the bars, blotches, and stripes are dark brown; ventral surface lighter than
the dorsum, with disperse small brown blotches.
TAL 4.4; FL 6.1.
on SVL (larger in female), on arms (slightly more robust in male), finger I (robust in male), and nuptial pad (on
finger I of male).
Distribution. Dendrophryniscus haddadi is known from the following municipalities (Fig. 17) in the states of
Rio de Janeiro (Paraty) and São Paulo (Caraguatatuba, Cunha, Picinguaba, São Luiz do Paraitinga, São Sebastião,
Ubatuba).
Natural history. This species is found in forested areas to bromeliads where they probably breed (L. Giasson
and C. Brasileiro, personal observation)
Etymology. The name of the species honors our friend Dr. Célio F.B. Haddad, from Universidade Estadual Pau-
lista (UNESP—Campus Rio Claro), by his huge contribution to the knowledge of Brazilian amphibians.
Figure 12. Dendrophryniscus haddadi sp. nov., holotype, CFBH 09300 (SVL 16.8 mm). Dorsal and lateral views of head,

Dendrophryniscus izecksohni sp. nov.
(Figs 13–14)
Holotype. MZUSP 103536, adult male, collected at Campo de Fruticultura da Bocaina (currently Núcleo Senador
Vergueiro), Municipality of São José do Barreiro (22o38’S, 44o34’W, Datum WGS 84, 540 m altitude), State of São
Paulo, Brazil, by F. M. Oliveira, O. Oliveira and W.C.A. Bokermann, in November 1968.
Paratypes. All specimens collected at type locality: MNRJ 2416, adult female, collected by J.L.A. Feio and
P. Miranda-Ribeiro, no date; MZUSP 103516, 103517, adult males, MZUSP 103518, MZUSP 103519, adult male,
collected by F.M. Oliveira, O. Oliveira, C.A. Seabra and W.C.A Bokermann, in November 1965; MZUSP 103520,
adult male, MZUSP 103537, 103538, adult females, MZUSP 103539, 103541, adult males, MZUSP 103542, ju-
venile male, MZUSP 103543, adult male, MZUSP 103544, 103545, juvenile males, collected with the holotype;
MZUSP 79104, 78508, juvenile males, collected, by F.M. Oliveira, O. Oliveira e W.C.A. Bokermann, from October
to November 1968.
24.2 mm in females); (2) body slender; (3) snout mucronate in dorsal view; (4) canthus rostralis slightly curved; (5)
elliptical set of pronounced granules posterior to the corner of mouth; (6) surfaces of upper eyelid with few granules
and with a conspicuous margin; (7) tip of the third and fourth fingers poor or not expanded laterally; (8) skinfold
poor developed on the articulation of the first phalange of the fingers II, III, and IV; (9) male with large white nuptial
pad with minuscule light brown horny asperities on finger I; (10) fingers fringed and webbed about one third.
Figure 13. Dendrophryniscus izecksohni sp. nov., holotype, MZUSP 103536 (SVL 21.3 mm). Dorsal and ventral views.
Comparisons with other species. Dendrophryniscus izecksohni sp. nov. is distinguished from D. proboscideus
by the smaller size (SVL 19.4–24.6 mm in males of D. izecksohni sp. nov.; SVL 39.2–46.4 mm in males of D. pro-
boscideus); D. izecksohni sp. nov. is distinguished from D. krausae and D. stawiarskyi by the body with uniform
shape (enlarged posteriorly in those species); D. izecksohni sp. nov. is distinguished from D. berthalutzae, D. car-
valhoi, D. krausae, D. lauroi, D. oreites, D. proboscideus, D. skuki, and D. stawiarskyi by the snout mucronate in
dorsal view (snout rounded in D. berthalutzae, D. carvalhoi, D. lauroi, and D. stawiarskyi; snout truncate in dorsal
view in D. krausae, D. oreites, and D. proboscideus; snout long, narrow, spatulate, with parallel lateral borders and
rounded tip in D. skuki); by the canthus rostralis slightly curved, D. izecksohni sp. nov. is distinguished from D.
davori sp. nov., D. carvalhoi, D. krausae, D. lauroi, and D. stawiarskyi (canthus rostralis straight); D. izecksohni
sp. nov. presents an elliptical set of pronounced granules posterior to the corner of mouth (elliptical set of unpig-
mented shallow granules posterior to the corner of mouth in D. lauroi; elliptical set of white, pronounced granules
in D. brevipollicatus and D. jureia sp. nov.; longitudinal set in D. berthalutzae, D. krausae, D. leucomystax, and D.

oreites; two pronounced granules in D. imitator; absent in D. carvalhoi, D. haddadi, D. organensis, D. probosci-
deus, D. skuki, and D. stawiarskyi); D. izecksohni sp. nov. presents the surfaces of upper eyelid with few granules
sparse and external margin conspicuous (surfaces of upper eyelid with numerous granules and with the external
margin slightly prominent in D. lauroi; surfaces of upper eyelid with numerous granules densely distributed and
with a conspicuous margin in D. brevipollicatus; D. izecksohni sp. nov. is distinguished from D. brevipollicatus
and D. lauroi by the tip of the third and fourth fingers poorly or not expanded (laterally expanded in those species);
D. izecksohni sp. nov.is distinguished from D. brevipollicatus by the presence of a skinfold poorly developed on
the articulation of the first and second phalanges of the fingers II, III, and IV (well developed in this species); D.
izecksohni sp. nov. is distinguished from all species of the genus , except D. davori sp. nov., D. carvalhoi, D. leu-
comystax, and D. stawiarskyi by the presence of large nuptial pad in male (absent in D. leucomystax, moderate in
all other species; D. izecksohni sp. nov. is distinguished from D. brevipollicatus, D. haddadi, D. krausae, and D.
lauroi by fingers fringed and webbed about one third (slightly fringed and webbed only at base in D. brevipollicatus;
fingers not fringed and webbed only at base in D. krausae; and fingers not fringed nor webbed in D. berthalutzae,
D. carvalhoi, D. haddadi, D. imitator, D. lauroi, D. oreites, D. skuki, and D. stawiarskyi.
Figure 14. Dendrophryniscus izecksohni sp. nov., holotype, MZUSP 103536 (SVL 21.3 mm). Dorsal and lateral views of
head, ventral views of hand and foot.

Description of holotype. Body slender, elongated (Fig. 13); head triangular, longer than large, head length
33.5% of SVL; snout mucronate in dorsal view, acute in lateral view (Fig. 14); snout 45.9% of head length; pres-
ence of a small tubercle under the tip of snout in ventral view; nostrils not protuberant, small and elliptical, located
laterally near the tip of snout, below and leaning the canthus rostralis; internarial distance 52.9% of eye to snout
distance; eye diameter 64.7% of eye to snout distance and 73.3% of interorbital distance; eye protuberant; canthus
rostralis slightly curved; loreal region vertical; choanae small, circular, and very far from witch one; tongue long,
narrow, and few enlarged posteriorly; vocal sac indistinct; vocal slits present. Arms robust, forearms more robust
than upper arms; hand with fingers slender, fringed and webbed about one third; tip of the third and fourth finger
poorly expanded laterally; finger I enlarged, covered by large white nuptial pad with minuscule light brown horny
asperities ; relative lengths of fingers I<II<IV<III; subarticular tubercles single, rounded, except at the third subar-
ticular tubercles of fingers II and III where they are horizontally elliptical; outer metacarpal tubercle large, rounded;
inner metacarpal tubercle rounded, approximated a quarter of the outer metacarpal tubercle; few, supernumerary
tubercles absent; skinfold poor developed on the articulation of the first phalanges of the fingers II, III, and IV. Thigh
length as large as tibia length; sum of thigh and tibia lengths about 83.5% of snout vent length. Tarsal length 22.5%
of the snout vent length. Foot with toes slender, not fringed; interdigital webbing developed, webbing formulae: I1–
1II1–2III1–3- IV3-–2- V; distal end of toes globose, not expanded laterally, relative lengths of toes I<II<III<V<IV;
subarticular tubercles single, rounded, the same size of those of the fingers; outer metatarsal tubercle rounded, twice
the size of subarticular tubercle; inner metatarsal like the outer; few, small and sparse supernumerary tubercles. Dor-
sal, ventral, and lateral surfaces covered by small and shallow spinulose granules irregularly distributed; surfaces
of upper eyelid with few granules sparse and external margin poorly marked; external margin of tibia and tarsus
smooth.
Color in preservative. Dorsal surfaces grayish brown; poorly defined brown transversal bar on thigh, tibia, and
tarsus; ventral surface whitish, with numerous irregular brown marbling.
TAL 4.8; FL 8.8.
and descriptive statistics of males and females are presented in Tables 2 and 3. Some specimens present on the dor-
sum an ‘X’ shaped brown blotch on the scapular region and an inverse ‘Y’ shaped brown blotch on the sacral region
extending to the inguinal region. Sexual dimorphism can be observed on arms (slightly more robust in males), finger
I (enlarged in males), and nuptial pad (on finger I of male).
Distribution. Dendrophryniscus izecksohni is known only from the type locality (Fig. 17), Campo de Fruticul-
tura (about 600 m above sea level), Municipality of São José do Barreiro, State of São Paulo, Brazil.
Natural history. Unknown.
Etymology. The name of the species honors the late Dr. Eugenio Izecksohn, from Universidade Federal Rural
do Rio de Janeiro (UFRRJ), our special friend and one of the most important Brazilian herpetologists.
Dendrophryniscus jureia sp. nov.

(Figs. 15–16)
Holotype. MNRJ 91510, adult male, collected at Estação Ecológica da Juréia-Itatins—Núcleo Rio Verde (24º22’S,
47º04’W, Datum WGS 84; 32 m altitude), Municipality of Iguape, State of São Paulo, Brazil, by J.P. Pombal Jr. and
M. Gordo, on 03 September 1988.
Paratypes. All specimens collected at the type locality: MNRJ 91511–91512, adult males, collected with ho-
lotype; MNRJ 91513, juvenile, collected by J.P. Pombal Jr., E.C.P. Pombal, and P.C. Bauduim, on 17 July 1990;
MNRJ 40007, collected by M. Gordo and Wolfgang, in 16–20 November 1989; MNRJ 92455 collected by C.A.
Brasileiro and M.T.C. Thom, on 01 November 2006; CFBH 373, collected by M. Gordo, on 01 October 1986;
ZUEC 9755, collected by J.P. Pombal Jr., M. Gordo & E.P. Pombal, on 07 September 1989.
(5) elliptical set of pronounced white granules posterior to the corner of mouth; (6) surfaces of upper eyelid with
few granules sparse and external margin poorly marked; (7) tip of the third and fourth fingers poor or not expanded
laterally; (8) skinfold poor developed on the articulation of the first phalange of the fingers II, III, and IV; (9) male

with poor developed nuptial pad with minuscule light brown horny asperities on finger I; (10) fingers not fringed
and webbed about one third.
Comparisons with other species. Dendrophryniscus jureia sp. nov. is distinguished from D. proboscideus by the
smaller size (SVL 16.0–18.7 mm in males of D. jureia sp. nov.; SVL 39.2–46.4 mm in males of D. proboscideus);
D. jureia sp. nov. is distinguished from D. krausae and D. stawiarskyi by the body with uniform shape (enlarged
posteriorly in those species); D. jureia sp. nov. is distinguished from D. berthalutzae, D. carvalhoi, D. krausae,
skuki); by the canthus rostralis slightly curved, D. jureia sp. nov. is distinguished from D. davori sp. nov., D. carv-
alhoi, D. krausae, D. lauroi, and D. stawiarskyi (canthus rostralis straight); D. jureia sp. nov. presents an elliptical
set of pronounced white granules posterior to the corner of mouth (elliptical set of unpigmented shallow granules
posterior to the corner of mouth in D. lauroi; longitudinal set in D. berthalutzae, D. krausae, D. leucomystax, and
D. oreites; two pronounced granules in D. imitator; absent in D. carvalhoi, D. haddadi, D. organensis, D. probosci-
deus, D. skuki, and D. stawiarskyi); D. jureia sp. nov. presents the surfaces of upper eyelid with few granules sparse
and external margin conspicuous (surfaces of upper eyelid with numerous granules and with the external margin
slightly prominent in D. lauroi; surfaces of upper eyelid with numerous granules densely distributed and with a
conspicuous margin in D. brevipollicatus; D. jureia sp. nov. is distinguished from D. brevipollicatus and D. lauroi
by the tip of the third and fourth fingers poorly or not expanded (laterally expanded in those species); D. jureia sp.
nov. is distinguished from D. brevipollicatus by the presence of a skinfold poorly developed on the articulation of
the first and second phalanges of the fingers II, III, and IV (well developed in this species); D. jureia sp. nov. is
distinguished from D. davori sp. nov., D. carvalhoi, D. izecksohni, D. leucomystax, and D. stawiarskyi by the pres-
ence of moderate nuptial pad in male (absent in D. leucomystax, very developed in D. davori sp. nov., D. carvalhoi,
D. izecksohni, and D. stawiarskyi); D. jureia sp. nov. is distinguished from D. berthalutzae, D. brevipollicatus, D.
carvalhoi, D. haddadi, D. imitator, D. krausae, D. lauroi, D. oreites, D. skuki, and D. stawiarskyi. by fingers fringed
and webbed about one third (slightly fringed and webbed only at base in D. brevipollicatus; fingers not fringed and
webbed only at base in D. krausae; and fingers not fringed nor webbed in D. berthalutzae, D. carvalhoi, D. haddadi,
D. imitator, D. lauroi, D. oreites, D. skuki, and D. stawiarskyi.
Figure 15. Dendrophryniscus jureia sp. nov., holotype, MNRJ 91510 (SVL 17.4 mm). Dorsal and ventral views.
Description of the holotype. Body slender, elongated (Fig. 15); head triangular, longer than large, head length
36.8% of SVL; snout mucronate in dorsal view, acute in lateral view (Fig. 16); snout 43.7% of head length; presence

of a small tubercle under the tip of snout, nostrils not protuberant, small and elliptical, located laterally near the tip
of snout, below the canthus rostralis; internarial distance 50.0% of eye to snout distance; eye diameter 64.3% of eye
to snout distance and 75.0% of interorbital distance; eye protuberant; canthus rostralis slightly curved; loreal region
vertical; choanae small, circular, and very far from witch one; tongue long, narrow, and few enlarged posteriorly;
vocal sac indistinct; vocal slits present. Arms slender, forearms more robust than upper arms; hand with fingers
slender, not fringed and webbed about one third; distal end of the third and fourth fingers slightly expanded laterally;
finger I enlarged, covered by moderate nuptial pad with minuscule light brown horny asperities; relative lengths of
fingers I<II<IV<III; subarticular tubercles single, rounded; outer metacarpal tubercle large, rounded; inner meta-
carpal tubercle rounded, slightly small than subarticular tubercles; supernumerary tubercles absent; skinfold poor
developed on the articulation of the first and second phalanges of the fingers II, III, and IV. Thigh length slightly
shorter than tibia length; sum of thigh and tibia lengths about 86.7% of snout–vent length. Tarsal length about 24.1%
of the snout vent length. Foot with toes slender, poor fringed; webbing formulae: I1–2II1–3III21/2–3-IV31/2–2V; dis-
tal end of toes globose, not expanded laterally, relative lengths of toes I<II<III<V<IV; subarticular tubercles single,
rounded, the same size of those of the fingers; outer metatarsal tubercle with the same size of subarticular tubercles,
rounded; inner metatarsal tubercle large, approximately twice the outer, elongated; supernumerary tubercles present
but scarce. Dorsal and lateral surfaces covered by tubercles and spinulose granules uniformly distributed, with dif-
ferent sizes, ventral surfaces rugose; surfaces of upper eyelid with few granules sparse and external margin poorly
marked; elliptical set of pronounced white granules posterior to the corner of mouth.
Figure 16. Dendrophryniscus jureia sp. nov., holotype, MNRJ 91510 (SVL 17.4 mm). Dorsal and lateral views of head,

TABLE 2. Range and mean ± standard deviation (in mm) of the measurements of the species of Dendrophryniscus (abbreviations of the measurements explained in the Material
and methods). Males.
SVL HL HW IND ESD ED IOD THL TL TAL FL
D. brevipollicatus 18.6‒25.1 6.1‒7.5 4.8‒6.3 1.2‒1.7 2.9‒3.9 1.8‒2.2 2.2‒3.0 6.7‒9.1 7.5‒9.4 4.2‒6.5 6.5‒8.7
N. 12 (20.7 ± 1.88) (6.7 ± 0.39) (5.5 ± 0.40) (1.5 ± 0.14) (3.3 ± 0.32) (2.0 ± 0.10) (2.6 ± 0.23) (8.0 ± 0.68) (8.3 ± 0.59) (5.1 ± 0.57) (7.3 ± 0.64)
D. imitator 15.1‒17.4 5.3‒6.2 4.3‒5.1 1.3‒1.5 2.7‒3.0 1.5‒2.0 2.2‒2.6 6.5‒7.7 7.0‒8.2 3.9‒4.6 5.3‒6.8
N. 5 (16.6 ± 0.89) (5.8 ± 0.45) (4.7 ± 0.26) (1.4 ± 0.06) (2.9 ± 0.11) (1.7 ± 0.18) (2.4 ± 0.15) (7.2 ± 0.54) (7.6 ± 0.51) (4.2 ± 0.30) (6.2 ± 0.60)
D. lauroi 14.5‒19.9 5.1‒7.0 4.1‒5.9 1.0‒1.8 2.3‒3.3 1.4‒2.1 1.9‒2.7 5.6‒7.8 6.4‒8.5 3.7‒5.1 5.2‒7.8
N. 17 (16.7 ± 2.06) (5.9 ± 0.69) (4.9 ± 0.68) (1.3 ± 0.20) (2.7 ± 0.35) (1.7 ± 0.19) (2.3 ± 0.22) (6.6 ± 0.73) (7.2 ± 0.73) (4.3 ± 0.45) (6.3 ± 0.84)
D. davori 18.9–22.0 5.8–7.7 5.7–6.7 1.8–1.9 3.0–4.0 1.9–2.0 1.9–2.8 8.4–8.8 7.9–9.2 4.7–5.6 7.6–9.2
N. 2
D. haddadi 15.7‒16.8 5.2‒ 6.1 4.7‒5.0 1.2‒1.4 2.9‒3.1 1.7‒1.9 2.2‒2.4 6.6‒7.2 7.0‒7.6 3.8‒4.9 5.8‒6.6

N. 6 (16.3 ± 0.44) (5.7 ± 0.31) (4.8 ± 0.12) (1.3 ± 0.06) (3.0 ± 0.09) (1.8 ± 0.07) (2.2 ± 0.15) (6.9 ± 0.25) (7.3 ± 0.21) (4.4 ± 0.48) (6.2 ± 0.25)
D. izecksohni 19.4‒24.6 6.7‒7.8 5.6‒7.1 1.5‒1.8 3.3‒3.8 1.8‒2.3 2.0‒3.4 8.4‒9.8 8.7‒10.1 6.0‒6.5 6.9‒9.0
N. 7 (21.5 ± 1.76) (7.3 ± 0.45) (6.3 ± 0.50) (1.7 ± 0.12) (3.5 ± 0.16) (2.0 ± 0.19) (2.7 ± 0.42) (8.9 ± 0.43) (9.0 ± 0.48) (5.5 ± 0.65) (8.3 ± 0.76)
D. jureia 16.0‒18.7 4.7‒6.6 4.4‒6.4 1.2‒1.7 2.7‒3.2 1.2‒2.0 2.1‒2.5 6.6‒8.0 7.2‒8.4 4.2‒4.8 6.1‒7.5
N. 6 (17.3 ± 0.97) (5.5 ± 0.88) (5.6 ± 0.77) (1.5 ± 0.18) (2.9 ± 0.18) (1.9 ± 0.10) (2.3 ± 0.11) (7.5 ± 0.54) (7.9 ± 0.43) (4.5 ± 0.24) (6.9 ± 0.49)

53
TABLE 3. Range and mean ± standard deviation (in mm) of the measurements of the species of Dendrophryniscus (abbreviations of the measurements explained in the Material
and methods). Females.
SVL HL HW IND ESD ED IOD THL TL TAL FL
D. brevipollicatus 20.9‒27.7 6.2‒7.8 6.0‒6.9 1.6‒1.7 3.3‒3.8 1.8‒2.3 2.8‒3.5 8.9‒11.2 9.0‒11.2 5.4‒6.9 7.3‒11.0
N. 3 (24.1 ± 3.41) (7.0 ± 0.77) (6.3 ± 0.41) (1.7 ± 0.06) (3.5 ± 0.25) (2.0 ± 0.22) (3.2 ± 0.31) (9.9 ± 0.95) (10.0 ± (6.0 ± 0.62) (8.8 ± 1.60)
0.99)
D. imitator 14.4‒20.7 5.8‒6.8 4.8‒5.4 1.4‒1.7 2.7‒3.2 1.6‒2.1 2.3‒2.6 6.7‒8.4 7.3‒9.1 4.2‒5.1 6.4‒8.9
N. 8 (18.1 ± 1.94) (6.3 ± 0.34) (5.0 ± 0.23) (1.5 ± 0.11) (3.0 ± 0.15) (1.8 ± 0.16) (2.5 ± 0.11) (7.5 ± 0.50) (8.0 ± 0.52) (4.6 ± 0.26) (7.2 ± 0.75)
D. lauroi 18.0‒20.3 6.1‒6.6 4.7‒5.7 1.3‒1.5 2.6‒3.3 1.7‒2.3 2.1‒2.5 7.2‒8.0 7.3‒8.0 4.7‒5.5 6.0‒7.0
N. 6 (19.1 ± 0.95) (6.2 ± 0.19) (5.4 ± 0.37) (1.4 ± 0.08) (3.0 ± 0.24) (1.9 ± 0.22) (2.3 ± 0.14) (7.6 ± 0.31) (7.8 ± 0.25) (5.0 ± 0.28) (6.5 ± 0.36)
D. davori 24.2 8.4 7.1 1.9 4.0 2.2 3.2 10.1 10.3 5.8 9.9
N. 1
54 · Zootaxa 4648 (1) © 2019 Magnolia Press

D. haddadi 17.5‒22.8 6.0‒7.3 4.5‒6.1 1.4‒1.8 2.9‒3.6 1.8‒2.1 2.2‒2.7 7.0‒9.0 7.2‒9.1 4.6‒6.0 6.4‒7.9
N. 6 (20.5 ± 1.97) (6.7 ± 0.44) (5.5 ± 0.56) (1.5 ± 0.15) (3.4 ± 0.26) (1.9 ± 0.09 (2.4 ± 0.19) (8.1 ± 0.68) (8.2 ± 0.63) (5.2 ± 0.50) (7.2 ± 0.50)
D. izecksohni 19.6‒24.2 6.1‒8.1 5.7‒7.0 1.3‒1.9 3.1‒3.8 2.0‒2.4 2.5‒3.4 8.7‒9.6 8.9‒9.7 5.5‒5.8 7.2‒10.6
N. 5 (21.5 ± 1.81) (7.2 ± 0.74) (6.4 ± 0.47) (1.6 ± 0.23) (3.5 ± 0.26) (2.1 ± 0.19) (3.0 ± 0.40) (9.2 ± 0.32) (9.3 ± 0.29 (5.6 ± 0.11) (9.0 ± 1.26)
D. jureia 17.6‒19.6 5.0‒5.0 6.4‒6.4 1.5‒1.6 3.1‒3.1 1.7‒2.2 2.5‒2.6 8.3‒8.4 8.6‒8.6 5.1‒5.3 7.3‒7.8
N. 2
CRUZ et al.
Color in preservative. Dorsum light brown, with fading blotches, a head blotch extending on the upper eyelids
merging with a ‘X’ shaped brown blotch on the scapular region and an inverse ‘Y’ shaped brown blotch on the
sacral region extending to the inguinal region; on forearms, one transversal fading brown bar; one transversal fad-
ing brown bar on thigh, tibia, and tarsus; whitish tubercles in the corner of the mouth; lateral sides of the body dark
brown; ventral surface whitish marbled with numerous irregular brown blotches.
TAL 4.2; FL 6.9.
on arms (slightly more robust in males), finger I (enlarged in males), and nuptial pad (on finger I of male).
TABLE 4. Intraspecific genetic distances expressed in values of p-distance and corresponding standard errors for the 16S
RNA ribosomal gene fragment of species of Dendrophryniscus. Not computed (n/c) in cases with only one sample for
species. Species of D. brevipollicatus group reviewed in this study are in bold.
Species p-distance Standard Error
D. proboscideus 0.00% 0.00%
D. oreites n/c n/c
D. carvalhoi 0.34% 0.33%
D. davori n/c n/c
D. brevipollicatus 1.60% 0.56%
D. lauroi 0.34% 0.23%
D. haddadi 1.83% 0.62%
D. jureia 2.72% 0.91%
D. organensis n/c n/c
D. imitator 2.30% 0.56%
D. berthalutzae 0.10% 0.09%
Figure 17. Map of geographic distribution of species of Dendrophryniscus brevipollicatus group reviewed in this study. RJ,
State of Rio de Janeiro; SP, State of São Paulo.

Distribution. Dendrophryniscus jureia is known only from the type locality, Estação Ecológica Juréia-Itatins
(Fig. 17) (about 32 m above sea level), Municipality of Iguape, State of São Paulo, Brazil.
Natural history. Unknown.
Etymology. The name of the species is an allusion to the type locality, in the Estação Ecológica da Juréia-Itatins,
Municipality of Iguape, State of São Paulo, Brazil.
Figure 18. Neighbor joining tree of genetic distances between species of Dendrophryniscus included in DNA-barcoding
analysis based in 16S RNA ribosomal gene fragment. Species of Dendrophryniscus brevipollicatus group reviewed in this study
are in black and lighted by colors correspondent to their distribution on the map (see Figure 17). Other Dendrophryniscus spe-
cies are in grey. Scale bar corresponding to substitutions per site (p-distance).
DNA-barcoding analysis
Our barcode analysis included 34 sequences of a 300 bp fragment of the 16S rRNA gene, comprising eleven Den-
drophryniscus species (Table 1). Intraspecific genetic distances ranged from 0 % to 2.72 (± 0.91 %) (Table 4).
Interspecific distances ranged from 3.35 % (± 0.9 %) between D. brevipollicatus and D. lauroi to 20.15 (± 2.23 %)
between D. berthalutzae and D. carvalhoi (Table 5). When considering only species of D. brevipollicatus group
reviewed in this study, the greater distance (12.25 ± 1.82 %) was found between D. brevipollicatus and D. davori
(Table 5). Optimal threshold values ranged from 2.8 % to 3.0 %, for total absence of false positives (no conspe-
cific matches within threshold distance) or false negatives (non-conspecific species within threshold). Barcode gap

TABLE 5. Genetic distance between species expressed in values of p-distance (below diagonal) and corresponding standard errors (above diagonal) for the 16S RNA ribosomal
gene fragment of species of Dendrophryniscus. Genetic distance and S.E. between species of D. brevipollicatus group reviewed in this study are in bold.
Species 1 2 3 4 5 6 7 8 9 10 11
1 D. proboscideus 2.11% 2.02% 2.04% 2.09% 2.09% 2.10% 1.97% 2.11% 2.06% 2.23%
2 D. oreites 16.72% 2.24% 2.01% 1.93% 1.95% 1.91% 1.92% 2.01% 1.86% 2.12%
3 D. carvalhoi 15.87% 19.97% 2.06% 2.11% 2.15% 2.06% 2.12% 2.25% 2.17% 2.23%
4 D. davori 15.92% 15.92% 16.09% 1.82% 1.83% 1.69% 1.50% 1.84% 1.69% 1.69%
5 D. brevipollicatus 17.67% 14.59% 17.73% 12.25% 0.90% 1.28% 1.41% 1.67% 1.69% 1.57%
6 D. lauroi 17.27% 14.61% 18.46% 11.83% 3.35% 1.40% 1.42% 1.70% 1.68% 1.60%
7 D. haddadi 18.26% 13.93% 17.07% 10.30% 6.98% 7.58% 1.32% 1.51% 1.47% 1.53%
8 D. jureia 16.38% 14.68% 18.54% 7.79% 8.04% 7.78% 7.42% 1.47% 1.49% 1.45%

9 D. organensis 16.78% 15.75% 19.22% 12.50% 10.64% 10.89% 8.59% 9.22% 1.38% 1.79%
10 D. imitator 17.92% 14.33% 19.86% 10.94% 11.96% 11.97% 8.88% 9.58% 8.23% 1.71%
11 D. berthalutzae 18.90% 17.18% 20.15% 9.74% 9.12% 8.87% 9.01% 7.61% 12.12% 11.41%

57
analysis showed that for all individuals the furthest intraspecific distances was always lower than the closest non-
conspecific individual. Genetic distances between all individuals analyzed are illustrated in the Neighbor Joining
tree (Figure 18).
Discussion
Morphological comparisons between populations of Dendrophryniscus brevipollicatus lead us to recognize seven

diagnosable lineages. Regarding the genetic distance, divergences between the fragments of 16S for Dendrophrynis-
cus brevipollicatus populations were coherent with interspecific values found for other frogs (Vences et al. 2005;
Lyra et al. 2017). Based on these evidences, we proposed the revalidation of two species and the description of four
new species beyond D. brevipollicatus. However, the genus Dendrophryniscus is a group of difficult taxonomy.
The taxonomic history of the species can illustrate the complexity. The original description of the genus by
Jiménez de la Espada (1870) was based on specimens collected in monte Corcovado (current Floresta da Tijuca),
Rio de Janeiro, designated as Dendrophryniscus brevipollicatus. In 1920, Miranda-Ribeiro described Atelopus imi-
tator, based on five specimens from some localities in the State of São Paulo (Santos, Alto da Serra, Rio Grande,
and Campo Grande). A few years later, Miranda-Ribeiro (1926) considered A. imitator as a synonymous form of
D. brevipollicatus and proposed three subspecies: Dendrophryniscus brevipollicatus lutzi, from Corcovado, Rio de
Janeiro; D. b. lauroi, from Angra dos Reis, State of Rio de Janeiro, and D. b. imitator, replacing A. imitator from
State of São Paulo. These subspecies were placed in synonymy of D. brevipollicatus by Cochran (1955). In 1971,
Izecksohn studied the vertebral pattern of D. brevipollicatus from four populations (Paranapiacaba and Serra da
Bocaina, São Paulo; Floresta da Tijuca, Rio de Janeiro; and Santa Teresa, Espírito Santo), but the constant pattern
found for São Paulo State populations in contrast to high polymorphism found to Floresta da Tijuca lead to incon-
clusive use of vertebral morphology in taxonomy of these populations. In 1994, Izecksohn presented a complete
diagnosis of D. brevipollicatus for populations from Rio de Janeiro to São Paulo State, and observed that the exam-
ined material seemed to gather some vicariant forms that would probably constitute different species.
Some particularities of the genus can explain their taxonomic complexity. It is almost entirely formed by bu-
fonids with reduced body size, sometimes inside the criteria proposed to Clarke (1996) and Yeh (2002) to include
miniaturized amphibians. The evolutionary tendency of reduction in body size is recurrent in amphibians (Hanken
& Wake 1993; Clarke 1996; Yeh 2002) and can have some morphological consequences like oversimplification of
external morphology and diversification of some bony structures if compared to relatives of bigger body size (Han-
ken & Wake 1993; Clarke 1996; Yeh 2002). These aspects can difficult the taxonomy, as example of Dendrophrynis-
cus brevipollicatus. For these reasons, we recognize the limitation of external morphology to define species, but
consider it a good starting point since it is supposed to be conservative, the number of specimens per locality is
often low (limiting some approaches of internal morphology that can damage collection specimens), and records
of larvae or advertisement calls are scarce. Future contributions concerning of bioacoustics, reproductive biology,
and larvae data can increase our ability to diagnose species inside D. brevipollicatus group. Primarily, the advance
of noninvasive techniques for internal morphology study, like X-ray microtomography (Micro CT-scan) (Metscher
2009), can bring new insights about internal morphology, which seems to be a valuable source of variation.
Acknowledgements
We thank Hussam Zaher and Taran Grant (MZUSP), Felipe Toledo (ZUEC), Marcelo F. Napoli (MZUFBA), Re-
nato N. Feio (MZUFV), Célio F.B. Haddad (CFBH), Glucia Pontes (MCP), Mrcio Borges Martins (MCN), Selvino
Neckel de Oliveira (UFSC), Oswaldo Luiz Peixoto (EI), and Thiago Arnt Dorigo (UERJ) for loan of specimens
and/or tissue samples. To Marta Calvo Revuelta (MNCN) for the information on the inexistence of type specimens
of Dendrophryniscus brevipollicatus. To Daniel B. Maciel for the photographs and Kelly R. Zamudio for helping
in sequences of DNA. LAF received postdoctoral fellowship from Fundação de Amparo a Pesquisa do Estado de
São Paulo (FAPESP,# 2013/21174-7) and Fundação Carlos Chagas Filho de Amparo Pesquisa do Estado do Rio
de Janeiro (FAPERJ, process number E-26/202.775/2016). CAB was granted by Fundação de Amparo Pesquisa
do Estado de São Paulo (FAPESP grants #08/50928-1 and #13/50741-7). We also received financial support from
Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and Fundação O Boticário.

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Appendix
Additional specimens examined:

Dendrophryniscus berthalutzae—Paran: MCP 7690, Guaratuba; MZUSP 137962, Guaraqueçaba. Santa Catarina: CFBH 10322-
10323, Treviso; MNRJ 31809, Itapoá; MNRJ 2131 (holotype), Joinville; MNRJ 49537, São Bento do Sul; MZUSP 138813,
Apiúna; MZUSP 35450–35458, Nova Horizonte; MZUSP 55851, Pirabeiraba; MCP 8273, Santo Amaro da Imperatriz.
Dendrophryniscus brevipollicatus—Rio de Janeiro: EI 1963–1967, 4016–4022, 4193–4194, 4203, 4948–4949, MNRJ 22065–
22067, 42861–42865, 42874–42878, MZUSP 103905–103957, ZUEC 3715, Parque Nacional da Floresta da Tijuca, Rio de
Janeiro; MNRJ 86388–86389, 58294–58295, 60701–60702, 72310, Reserva Ecológica de Guapiaçu, Cachoeiras de Macacu;
EI 4948–4949, 5187, Engenheiro Paulo de Frontin; MNRJ 22074, 59011, 60220, Mag; MNRJ 1943, 53520–53522, Serra dos
Gaviões, Silva Jardim; MNRJ 8156, Nova Iguaçu.
Dendrophryniscus carvalhoi—Espírito Santo: EI 4127 (holotype), 4124–4126, 4128 (paratypes), MNRJ 1287 (2 specimens),
MNRJ 38364–38365, 40678, MZUSP 104124–104135, Santa Teresa.
Dendrophryniscus haddadi—Rio de Janeiro: MNRJ 2430, 11292–11294, 2438, 89864–89865, Parque Nacional da Serra da Bo-
caina, Parati. São Paulo: CFBH 03952, 03963, 04323, 12648–12649, 12699, 13839–13840, 15616, 15674–15675, MNRJ 92461,
92462, 92463, 92464-92465, Ilha do Prumirim, Ubatuba; CFBH 1389, 1415, 1713, 1714, 2528, MNRJ 22077, 92123, MZUSP
69813, ZUEC 1389, 1713–1714, 1415, 2528, Picinguaba, Ubatuba; MNRJ 92458, 1762–1763, Ilha do Mar Virado, Ubatu-
ba; CFBH 13834–13838, MNRJ 92459-92460, Ilha dos Porcos, Ubatuba; CFBH 12687–12692, 12694–12696, 15216–15219,
17410–17411, MZUSP 51667, Ilha de São Sebastião, Ilhabela; MZUSP 30585–30586, 30567, 30569–30572, 30583–30584,
30587–30589, Ilha dos Búzios, Ilhabela; MZUSP 1280, Caraguatatuba; CFBH 10712, 10743, 12636–12638, 12640–12641,
Parque Estadual da Serra do Mar, Cunha; CFBH 11632, Natividade da Serra.
Dendrophryniscus imitator—São Paulo: MNRJ 42866, ZUEC 11331, São Paulo; CFBH 11624, Represa Billings, São Paulo;
CFBH 11164, 11166, 11175, 12659, 12666, 12683, 12291, MNRJ 92456–92457, Parque Estadual da Serra do Mar Núcleo Curu-
cutu; CFBH 12670–12672, CFBH 12674–12675, Biritiba Mirim; MZUSP 134375, 134407–134408, Reserva do Morro Grande,
Cotia; CFBH 24122, EI 4148–4152, MZUSP 127645–127648, 134781–134784, Juquitiba; MNRJ 22054–22060, 22061–22062,
MZUSP 103991, 103996–103999, 104007–104013, 133209, EI 4129–4137, Paranapiacaba, Santo André; MNRJ 42867–42873,
Alto da Serra de Cubatão (=Paranapiacaba, Santo André); CFBH 15955, Santos; MZUSP 37786, 61126, Estação Biológica de
Boracéia, Salesópolis.
Dendrophryniscus krausae—Rio Grande do Sul: MCN 13809 (holotype), MCN 13808, 13810, 13812, MNRJ 50290 (para-
types), Maquiné; MCN 11028–11030 (paratypes), Cambará do Sul.
Dendrophryniscus lauroi—Rio de Janeiro: MNRJ 2446, 11305–11306, 2019, 10501–10512, 42879–42882, MZUSP 9744–
9754, Angra dos Reis; MNRJ 48516–48518, 52723, 57603–57605, 57799–57800, 58296, 58297, 58299, 58303, 58304, 58304,
90575–90587, Ilha Grande, Angra dos Reis; MNRJ 679, 5757–5762, 2415, 11275–11279, 71511–71512, Mangaratiba; MNRJ
76647, Rio Claro. São Paulo: EI 1187–1190, 1429–1436, 1963, Bananal.
Dendrophryniscus leucomystax—Rio de Janeiro: EI 4045–4046, EI 4047–4067 (paratypes), EI 4068 (allotype), EI 4069 (holo-
type), EI 5425–5433, MNRJ 22063, MNRJ 60131 (paratype), Tinguá, Nova Iguaçu; MNRJ 2246, 2945, Barro Branco, Duque
de Caxias. São Paulo: EI 5420–5424, Sete Barras; MNRJ 505, Sororocaba; MZUSP 92521, Ilha do Cardoso, Cananéia; MZUSP
138277, Registro.
Dendrophryniscus oreites—Bahia: MZUSP 142493 (holotype), Parque Nacional de Serra das Lontras, Arataca.
Dendrophryniscus organensis—Rio de Janeiro: MNRJ 58291–58293 (topotypes), Teresópolis.

Dendrophryniscus proboscideus—Bahia: MNRJ 26455, Arataca; MNRJ 2733–2734, Salvador; UFBA 10086–10087, 10551,
Catu; UFBA 10108, Vera Cruz; UFBA 10470, Wenceslau Guimarães; MZUFV 6001, Macarani; MNRJ 2733–2734. Minas
Gerais: MZUFV 4134–4138, 5929–5934, Almenara.

Dendrophryniscus skuki—Bahia: MNRJ 74619 (holotype), Itacar.
Dendrophryniscus stawiarskyi—Paraná: MNRJ 2650 (holotype), Bituruna.
Dendrophryniscus spp.—Rio de Janeiro: MNRJ 3581, 22068, 22069–22073, ZUEC 11506, Parque Nacional do Itatiaia, Ita-
tiaia. São Paulo: CFBH 24123, Piquete; MNRJ 28204, ZUEC 11331, Parque Estadual Intervales, Ribeirão Grande; MZUSP
125443–125444, Tapiraí.
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Taxonomic review of Dendrophryniscus brevipollicatus Jiménez de la Espada,

Article in Zootaxa · July 2019

Ulisses Caramaschi Luciana Ardenghi Fusinatto

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Taxonomic review of Dendrophryniscus brevipollicatus Jiménez de la Espada,

Accepted by M. Vences: 5 Jun. 2019; published: 29 Jul. 2019 27

Material and methods

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Review of Dendrophryniscus brevipollicatus

Zootaxa 4648 (1) © 2019 Magnolia Press ·

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Dendrophryniscus brevipollicatus Jiménez de la Espada, 1870

Dendrophryniscus brevipollicatus Jiménez de la Espada, 1870.

Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 31

Comparisons with other species. Dendrophryniscus brevipollicatus is distinguished from D. proboscideus by

32 · Zootaxa 4648 (1) © 2019 Magnolia Press CRUZ et al.

Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 33

Dendrophryniscus imitator (Miranda-Ribeiro, 1920), new status, revalidated

Atelopus imitator Miranda-Ribeiro, 1920.

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Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 35

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Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 37

Dendrophryniscus lauroi Miranda-Ribeiro, 1926, new status, revalidated

Dendrophryniscus brevipollicatus lauroi Miranda-Ribeiro, 1926.

38 · Zootaxa 4648 (1) © 2019 Magnolia Press CRUZ et al.

Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 39

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Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 41

Dendrophryniscus davori sp. nov.

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Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 43

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Dendrophryniscus haddadi sp. nov.

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46 · Zootaxa 4648 (1) © 2019 Magnolia Press CRUZ et al.

Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 47

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Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 49

Dendrophryniscus jureia sp. nov.

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Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 51

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Review of Dendrophryniscus brevipollicatus

Zootaxa 4648 (1) © 2019 Magnolia Press ·

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Review of Dendrophryniscus brevipollicatus

Zootaxa 4648 (1) © 2019 Magnolia Press ·

Morphological comparisons between populations of Dendrophryniscus brevipollicatus lead us to recognize seven

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Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 59

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Additional specimens examined:

Dendrophryniscus organensis—Rio de Janeiro: MNRJ 58291–58293 (topotypes), Teresópolis.

Review of Dendrophryniscus brevipollicatus Zootaxa 4648 (1) © 2019 Magnolia Press · 61

Dendrophryniscus stawiarskyi—Paraná: MNRJ 2650 (holotype), Bituruna.

62 · Zootaxa 4648 (1) © 2019 Magnolia Press CRUZ et al.

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