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99
Taxonomy of Hypsiboas crepitans
Available at http://www.salamandra-journal.com
© 2017 Deutsche Gesellscha für Herpetologie und Terrarienkunde e.V. (DGHT), Mannheim, Germany
SALAMANDRA 53(1) 99–113 15 February 2017 ISSN 0036–3375
Integrative taxonomy supports the existence of two distinct species
within Hypsiboas crepitans (Anura: Hylidae)
V G. D. O, I N,, C M, A F, A W. L,
M R-C, M L. L, D L, B V. S. P,
U C, M T. R C F. B. H
1) Universidade Estadual de Santa Cruz, Departamento de Ciências Biológicas, Rodovia Jorge Amado, Km 16.
CEP 45662-900 Ilhéus, BA, Brazil
2) Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Vertebrados. Quinta da Boa Vista, São Cristóvão.
CEP 20940-040, Rio de Janeiro, RJ, Brazil
3) CNRS-Guyane, USR 3456 LEEISA – Laboratoire Ecologie, Evolution Interactions des Systèmes Amazoniens,
Centre de recherche de Montabo 275 route de Montabo, BP 70620. 97334, Cayenne cedex, French Guiana
4) Pontifícia Universidade Católica do Rio Grande do Sul, Faculdade de Biociências, Laboratório de Sistemática de Vertebrados.
Av. Ipiranga 6681. CEP 90619-900, Porto Alegre, RS, Brazil
5) Universidade Estadual Paulista (UNESP), Instituto de Biociências, Departamento de Zoologia and Centro de Aquicultura da
UNESP – CAUNESP, campus Rio Claro, Av. 24A, N 1515. CEP 13506-900, Rio Claro, SP, Brazil
6) Universidade Federal do Rio Grande, Instituto de Ciências Biológicas, Laboratório de Vertebrados, Av. Itália Km 8, Carreiros.
CEP 96.203-900, Rio Grande, RS, Brazil
7) Bicho do Mato Meio Ambiente Ltda. / Bicho do Mato Instituto de Pesquisa. Rua Perdigão Malheiros 222. CEP 30380-234,
Belo Horizonte, MG, Brazil
8) Universidade de São Paulo, Instituto de Biociências, Departamento de Zoologia, Caixa Postal 11.461.
CEP 05508-090, São Paulo, SP, Brazil
9) Present address: Universidade Estadual Paulista, Campus do Litoral Paulista, Instituto de Biociências,
Laboratório de Herpetologia. Praça Infante Dom Henrique, s/n, Parque Bitarú. CEP 11.330-900, São Vicente, SP, Brazil
Corresponding author: A F, e-mail: fouquet.antoine@gmail.com
Manuscript received: 10 February 2016
Accepted on 13 September 2016 by J K
Abstract. e Neotropical treefrog Hypsiboas crepitans (W, ) has an intriguing disjunct geographic distribution
encompassing two large patches: the Atlantic Forest in southeastern South America and from the Guiana Shield to Cen-
tral America in the north, that are separated by more than km. is distribution pattern led us to review the avail-
able material and re-examine, under an integrative approach, the taxonomic status of these populations. We assessed data
using three lines of evidence: morphology, morphometry, and mitochondrial DNA. All of them suggest that the popula-
tions from the two geographical ranges are not conspecic. Given that the type material of H. crepitans is from the State of
Bahia, Brazil, and that specimens from this area cluster with the southeastern group, we resurrect Hypsiboas xerophyllus
(D B, ) for the populations of the northwestern group. Hyla levaillantii D B, , Hyla
doumercii D B, , Hyla fuentei G G, , and Hypsiboas indris C, are synonymized
with H.xerophyllus.
Key words. Amphibia, Amazonia, Atlantic Forest, Hypsiboas faber species group, revalidation, South America, synonymy.
Introduction
e Hypsiboas faber species group was proposed by F-
et al. () to accommodate a cluster of eight spe-
cies: H. albomarginatus (S, ), H. crepitans (W,
), H. exastis (C R, ), H.
faber (W, ), H. lundii (B, ), H.par-
dalis (S, ), H. pugnax (S, ) and H. rosen-
bergi (B, ). With the exception of H. albo-
marginatus (green, middle sized), all species are large,
territorial tree frogs with a lichenous colour pattern, and
rugose dorsal skin texture (F et al. , K
, L S-M ).
One of the species of this group, Hypsiboas crepitans,
exhibits an extensive and intriguing disjunct geographical
distribution (Fig. ). A southeastern group (hereaer SG)
100
V G. D. O et al.
of populations occurs along the Atlantic Forest and ad-
jacent areas, including the State of Bahia in Brazil, from
where the species was originally described (W ).
e northwestern group (hereaer NG) of populations is
distributed over the Guiana Shield, Caribbean (Tobago),
Llanos, Andes, and Middle America (Panama), from sea
level up to m a.s.l. (D , F ,
K , L , L S-M-
). ere is, thus, a > , km gap between the SG
and the NG. is unusual disjunct distribution has raised
doubts about the conspecicity of these groups of popu-
lations (C J , D , K
, L , L S-M ).
F et al. () found that the pairwise distance
among S sequences of specimens from Alagoas, Brazil,
and French Guiana was high enough to suggest the pres-
ence of two distinct species. L () retrieved the
same result comparing S sequences of specimens from
Tobago and Brazil and suggested that the population stud-
ied by C J () from the State of Bahia “may
represent a dierent, but currently undescribed, species”,
not realizing that it could be the other way around given
that this particular population is close to the type locality
of H.crepitans (see below).
However, previous works on tadpoles and calls in both
the NG and the SG provide only weak evidence support-
ing the distinction. Tadpoles of the NG present a spiracle
opening directed dorso-posteriorly, and antero-dorsal nos-
trils (see L , gure ), while tadpoles from the
SG present a spiracle directed backwards, and dorsal nos-
trils (C J , gure ). However, given the
high plasticity of anuran larvae (e.g., W ) we
remain skeptical that these slight dierences will be main-
tained in larger samples. e basic structure of the calls
of both groups is a periodic pulse train divided in one or
some notes. e number of notes from the SG (– notes:
C J , M et al. ) overlaps with
the range of number of notes of NG from Panama (–
notes; F , D , K et al.
) and from Colombia (– notes; B et al. )
(Table ). e range of pulses per second from the SG (–
; C J , M et al. ) is slightly
dierent from the value of NG recordings from Panamá
(≈; D ). e dominant frequency range of
SG recordings (.–. kHz; C J , M-
et al. ) do not dier from the values recovered
for the NG, although being individually lower than values
found for NG specimens from Panama (.–. kHz;
Figure 1. Map showing sampling localities of Hypsiboas crepitans (circles) and H. xerophyllus (triangles) specimens used in this study.
101
Taxonomy of Hypsiboas crepitans
F , D , K et al. ), and
higher than those found for Colombia (. kHz; B
et al. ). erefore, dierences between the advertise-
ment calls of specimens from the distinct groups are meag-
er. However, C J () used advertisement
call data (dominant frequency mainly) to corroborate the
hypothesis that more than one species is hidden under the
name Hypsiboas crepitans. erefore, although there is a
suspicion that the two groups may represent distinct spe-
cies, published data on advertisement calls and tadpoles
are inconclusive in this respect.
e two groups of populations are separated by the
Caatinga, Cerrado and Chaco biomes which are part of
the “Dry Diagonal” of the Neotropics (S I
, W ), a group of “dry environments” pres-
ently separating Atlantic and Amazonian Forests and to
which some cladogenic events have been attributed (C-
). Successive uctuations over time of these con-
trasting habitats help to explain relationships and faunal
exchanges between Amazonian and Atlantic Forests (see
discussions in F et al. , ). Most of the doc-
umented dispersals of terrestrial anurans between these
two biomes are rather ancient (F et al. ) and
only a few examples of Atlantic Forest species like Rhinel-
la hoogmoedi and Hypsiboas semilineatus are nested with
relatively low genetic distances (– on S) within oth-
erwise Amazonian species ( S et al. , F-
et al. ). However, no species of anuran restricted
to forest habitats is known to be distributed in both biomes
while it is the case in only a few open habitat species like
Adenomera hylae dactyla (F et al. ).
Nevertheless, although Hypsiboas crepitans is predomi-
nantly associated with forest and mesic habitats, many pop-
ulations are found in dry or xeric environments. L
M () and O J () reported
that populations from French Guiana and Suriname, re-
spectively, live in open environments such as in savannas
and inselbergs surrounded by forest. In fact, the type local-
ity of H. crepitans (Tamboril, Municipality of Condeúbas,
State of Bahia, Brazil, see B ) is situated in
a transitional area between the rocky meadows of Serra do
Espinhaço and the Caatinga where both open and forest
vegetation are present. erefore, the inuence of the dry
diagonal as a barrier for this species is not straightforward.
Given these rather wide habitat requirements, a dispersal
between Amazonia and the Atlantic Forest, recent enough
to support a conspecic status of the two groups, cannot
be excluded.
e goals of the present contribution are to review spec-
imens assigned to Hypsiboas crepitans from both distribu-
tion areas and perform analyses using an integrative ap-
proach combining molecular (mtDNA), morphology and
morphometry in order to evaluate the specic status of
these populations.
If the existence of more than one species is supported, a
number of synonyms are available for the NG and should
therefore be examined. Additionally, O J
() also reported H. fuentei (G G ), a spe-
cies known only from three localities in northeast-central
Suriname (F ), that are nested within the range of
the NG. ey also stated that they would “not be surprised,
if H. fuentei proves to be a junior synonym of H. crepitans”.
However, H () suggested that because of the
green colours of live specimens, H.fuentei seemed some-
how related to H. punctatus or H.cinera scens – both today
assigned to the H. punctatus species group. Neither H-
() nor O J () compared
their specimens with the holotype of H.fuentei, solely with
the description of G G (). Although F-
et al. () did not assign H.fuentei to any of their
species groups, they state “the angulate dentigerous proc-
ess of the vomer suggests that this species could be associ-
ated with certain Gladiator Frogs... [that] have this charac-
ter state. A study of the holotype … should clarify the mat-
ter”. Given the confusion surrounding H. fuentei and the
possible relation with H.crepitans, we include herein our
own observations of the H. fuentei holotype and evaluate
its taxonomic status.
Table 1. Advertisement call data for Hypsiboas crepitans distributional patches. See text for abbreviations.
OTU Call
duration
(s)
Intercall
interval
(s)
Number
of notes
per call
Number
of pulses/
second
Note
interval
(s)
Number of
pulses/note
Pulse
duration
(s)
Dominant
frequency
(kHz)
Reference
NG – – 2–5 – – – – 2.55 F (1966)
NG 2.5–5 – 3.3–4.4 110 – 3–5 0.009 0.965–1.288 D (1970)
NG – – 2–5 – – – – 2.15 K et al. (2000)
NG 0.21–0.31 – 1–2 – – – – 0.35 B et al. (2004)
NG
Summary 0.21–5 ? 1–5 110 ? 3–5 0.009 0.965–2.55 –
SG 0.51 0.87 1–5 72–96 0.04 3–33 0.010–0.014 0.8 C J (2008)
SG 0.46 0.52 2 68–77 0.049 4–27 0.014–0.013 0.53–1.30 M et al. (2009)
SG
Summary 0.46–0.51 0.52–0.87 1–5 68–96 0.04–0.049 3–27 0.01–0.014 0.53–1.30 –
102
V G. D. O et al.
Materials and methods
General procedures
We examined specimens in zoological collections trying to
assemble material from as many populations reported as
H. crepitans as possible. Institutional abbreviations can be
found in S P (), with the addition of AL-MN
(Adolpho Lutz Collection, housed at Museu Nacional, Rio
de Janeiro, Brazil) and MTR (M T. R eld
numbers). Institutional abbreviations apply to both mor-
phological and molecular (tissue) material. A number of
specimens, published sequences, and tissues available were
selected and gathered (see below). is material was either
considered as NG or SG, allowing us to test the above-cit-
ed hypothesis that the two distributional groups of popu-
lations would represent two taxa with allopatric distribu-
tion (C J , D , K ,
L S-M ). Specimens from NG
are from Roraima state Brazil, Colombia, French Guiana
and Venezuela, while specimens from SG are from sev-
eral Brazilian localities (Supplementary document). Ad-
ditional species for comparisons were chosen based on
their availability according to the present taxonomy i.e.,
all the other species of the H.faber group, as dened by
F et al. (). To discuss the taxonomic status of
H.fuentei, we examined photos of the holotype and types
of relevant material (H. crepitans lectotype and photos of
the holo types of taxa currently considered synonyms).
Qualitative phenotypic data
We conducted the comparisons of adult specimens based
on observations of collection material (Supplementary
document ) and on literature information (see below).
Terminology of external morphology follows D
(). Standards for dorsal outline and prole of the snout
follow H et al. (). Fingers nomenclature follows
F A (). Webbing formula notation
follows S H (), as modied by M
D (). Types of vocal sac follow L (). Sex
was determined by the presence of vocal sac, vocal slits, and
developed projecting prepollex in adult males. Coloration
always refers to preserved specimens, except when stated.
Quantitative phenotypic data
Sixteen morphometric variables were used and are given in
millimeters throughout the text. Nine measurements fol-
low D (): SVL (snout–vent length), HL (head
length), HW (head width), ED (eye diameter), UEW (up-
per eyelid width), IOD (interorbital distance), IND (inter-
narial distance), TD (tympanum diameter), and TL (tib-
ia length). One measurement follows H et al. ():
THL (thigh length). Five measurements follow N
(): END (eye–nostril distance), NSD (nostril-tip of
snout distance), FL (foot length, including tarsus), FD
(fourth nger disk diameter; using nomenclature of F-
A ()), and TD (fourth toe disk di-
ameter). We included an additional measurement: FHL
(forearm+hand length: straight line distance from elbow to
the tip of the third nger). SVL, HL, HW, FHL, THL, TL,
and FL were measured with a digital caliper to the nearest
. mm. All other measurements were taken with an ocu-
lar micrometer on a Zeiss stereomicroscope.
According to B C (), specimens
preserved at dierent times can produce an articial seg-
regation among each time class when measured for mor-
phometric analyses. e reason is the gradual modication
of specimens along the years in preservative (see a brief
historic account in D et al. ). Given that
the measured specimens were collected in several dier-
ent locations and times (see Supplementary document ),
the large amount of analyzed specimens and, the normal
distribution of each measurement, we expect to minimize
the possible problems regarding artefacts of preservation.
A total of adult specimens were measured (NG: n=
males and females; SG: n = males and females).
Prior to analysis, all morphometric measures were log-
transformed to conform to requirements of normality and
homocedasticity (Z ).
Principal Component Analysis (PCA) was used to ex-
plore morphometric dierences between groups. Eigen-
vectors and associated eigenvalues were obtained from a
variance-covariance matrix. Scores of individuals were
then projected in the reduced space of the main compo-
nents of larger contributions (H et al. ). e
rst component (hereaer PC) captures the largest pos-
sible variation of the original data; the second component
(hereaer PC) is orthogonal to the rst (independent) and
provides the remaining of maximum variation (P-N-
B ).
Molecular data procedures
Our survey of GenBank sequences (performed on the th
of July ) showed that, with small additions from our
own dataset, it was possible to assemble a nearly complete
matrix using sequences of the S mitochondrial gene
(hereaer S; bp) and of the mitochondrial Cyto-
chrome c oxidase subunit (hereaer COI; bp) for all
species in the Hypsiboas faber group and for some close-
ly related species according to F et al. () to
serve as outgroups. is includes both geographical groups
of H. crepitans (see Supplementary document ), however
we were not able to gather sequences of both targeted gene
fragments for all terminals. In order to reduce missing en-
tries for outgroups, we used ve chimerical sequences (see
results). Chimerical sequences are solely of distinct indi-
viduals assigned to a same species, and no chimerical se-
quence was produced for ingroup terminals (Supplemen-
tary document ).
To amplify the S mtDNA, we used primers MVZ
(’–ATAGCACTGAAAAYGCTDAGATG–’; G
103
Taxonomy of Hypsiboas crepitans
) and S F-H (’–CTTGGCTCGTAGTTCCCT-
GGCG–’; G et al. ) following the procedures of
F et al. (). COI was amplied using primers
dgHCO- (’-TAAACTTCAGGGTGACCAAARAAY-
CA-’) and dgLCO- (’-GGTCAACAAATCATAAA-
GAYATYGG-’) described in M () following his
procedures. Fragments were sequenced in both directions
and sequencing was performed by Macrogen Inc. (Seoul,
South Korea). Data from complementary strands were
compared to generate a consensus sequence for each DNA
fragment using Sequencher . (Gene Code Corp, Ann Ar-
bor, USA).
Alignments were conducted in the online version of
MAFFT v (K et al. ), aligning each fragment
separately; both genes were aligned using MAFFT align-
ment strategy L-INS-i. e nal matrix comprises ter-
minals. e missing data are two S sequences of H. fab-
er and COI sequences of various representatives (two
H.crepitans) of the H. faber species group. We collated
newly generated sequences of S and of COI with for
S and eight for COI from GenBank. e nal matrix can
be found at BOLDSYSTEMS.
Genetic (uncorrected pairwise) distances were calculat-
ed with MEGA . (T et al. ) considering transi-
tions and transversions. Pairwise distances were computed
for a total of bp for S and bp for COI. As a rst
approximation, we considered genetic distances high (i.e.,
possibly not conspecic) when ≥ for the S and ≥
for COI. A threshold of for S is common between dis-
tinct amphibian species for which genetic distances were
studied (e.g., C-F et al. , K et
al. , R et al. ). On the other hand, divergence
values for COI have been studied for diverse animal groups
and although thresholds vary among clades (e.g., H
et al. ), a minimum value of is commonly found
between amphibian species (e.g., V et al. ).
We used JModelTest .. (G G ,
P ) to select the best nucleotide substitution
models according to the Akaike Information Criterion
(AIC) for each of the four partitions we considered (S
and each codon position for COI). We used default settings
( substitution schemes, ML optimization, NNI search).
Selected models were GTR+I+G for S and TrNef+I, F,
and TrN+G for the rst, second and third positions of the
COI respectively.
e resulting models were employed in a Bayesian anal-
ysis (BA) with MrBayes . (R et al. ). Instead
of TrNef+I and TrN+G we used the closest models (GTR+I
and GTR+G respectively) for BA. Gaps were treated as nu-
cleotides of unknown origin in Bayesian inference analyses
due to program constrains. e BA consisted of a ×
generations run and four Markov chains (one cold) sam-
pled every , generations. A conservative burn-in ()
was determined by examining stationarity of the likelihood
scores (all parameters ESS were > ,) and convergence
between the two runs using the deviation of split frequen-
cies and mixing between chains. We considered relation-
ships to be strongly supported when posterior probabilities
were equal to or higher than .. Outgroup (root) was set
as H. cinerascens + H. punctatus according to F
et al. ().
Results
Morphology
Specimens from the two groups can be distinguished based
on adults colour patterns and general morphology. Howev-
er, the two groups share the following characteristics: head
in lateral view truncated, rounded, or acuminate; head in
dorsal view rounded, truncated, or sub-elliptical; feet web-
bing formulae I[] – []II – III – IV – V. e usu-
al coloration of the body encompasses several patterns of
marbled background with or without small spots (melano-
phores) mostly resembling a bark-like pattern. e pattern
on the posterior surface of thighs is dierent between both
groups; specimens without blotches are found only in the
NG, although some specimens may present some dark lines
(Fig. C; D ). Furthermore, the presence of a
middorsal longitudinal stripe and the colour of the gular
region dier between groups (see the comparisons section).
Morphometric analyses
PCA results are congruent with the hypothesis that the
NG and the SG represent dierent species. Regarding the
males (n = ), PCA shows two major axes (PC . of
the total variation and PC .). e standardized co-
ecients and factor loadings of the Principal Component
axes are presented in Supplementary document . e two
axes show that the NG is completely separated from the
SG, and the separation of these two groups is mostly due to
both PC and PC (Fig. ). When we look at the axes indi-
vidually, PC is mostly inuenced by FL, HW, THL, SVL,
and IOD, in this order, and PC is mostly inuenced by
NSD, TD, END, FD, and TD, in this order (Supplemen-
tary document ).
Regarding the females (n = ), PCA also display two
major axes (PC accounted for . of the total variation
and PC for .). e standardized coecients and fac-
tor loadings of the Principal Component axes are present-
ed in Supplementary document . e two axes show that
the NG is completely separated from the SG, and the sepa-
ration of these two is mostly due to PC (Fig. ). When we
look to the axes individually, PC is mostly inuenced by
SVL, HW, HL, FD, and THL, in this order, and PC is
mostly inuenced by IOD, NSD, FL, FD, and UEW, in this
order (Supplementary document ).
Molecular analyses
e two groups of H. crepitans are reciprocally mono-
phyletic and retrieved as strongly supported sister clades
(Fig. ). Genetic distances between them are high: in
104
V G. D. O et al.
S and in COI (Table ). A phylogeographical struc-
ture is also apparent within each of these two clades. e
NG clade comprises two main lineages segregating east-
ern populations in French Guiana and Guyana (east of Es-
sequibo river) from western populations in Guyana (west
of Essequibo river) and Roraima State (Fig. ). However,
genetic distance between them is low as well as support
for the western lineage (< .). e SG also comprises two
main lineages segregating the northern part of the Atlantic
Forest (AL, PE, BA) from the central part of the Atlantic
Forest (BA, MG). Similarly, genetic distance between them
is low (Table ) as well as support for the central Atlantic
Forest lineage (< .).
Most of the nodes of the tree are strongly supported.
However, the Hypsiboas faber species group was not re-
trieved as monophyletic with the H. albopunctatus species
group (Hypsiboas lanciformis and H. multifasciatus) being
nested within it with uncertain relationship (Fig. ). e
basal divergence within the clade H. fa ber group + H.albo-
punctatus group separates H. albomarginatus from the
rest. However, this relationship is also poorly supported.
In fact, four major groups can be recognized in this clade,
the H.albopunctatus group (widespread in South Ameri-
ca) and three strongly supported clades formed by the spe-
cies of the Hypsiboas faber group. e rst holds the two
samples of H. albomarginatus (Atlantic Forest), the second
samples of H. exastis, H. faber, H. lundii, and H.pardalis
(hereaer the H. faber clade – Atlantic Forest, Cerrado),
and the third H. crepitans, H. pugnax, and H. rosenbergi
(hereaer the H. crepitans clade – from the Guiana Shield
to Central America and the Atlantic Forest).
Observations on the holotype of Hyla fuentei
e holotype of Hyla fuentei (CM ; Fig. D) is a fe-
male in good condition of preservation. e abdomen has
a sagittal opening, possibly made for sex determination.
e specimen presents two dorsal cuts in the skin: one over
the frontoparietal fontanel and one from the right shoulder
blade to the le hip. Colours have faded and no clear pat-
tern can be seen.
Some specimens of the NG agree with the holotype of
H. fuentei. Many specimens of the NG are dark green in
Figure 2. (A) Living specimen of Hypsiboas crepitans in diurnal coloration (photo by M. S) from the UESC campus in Ilhéus, Bahia,
Brazil (specimen not collected). (B, C) igh patterns of two specimens of Hypsiboas crepitans (B = MNRJ32440 and C = MNRJ64374).
(D) Living specimen of Hypsiboas xerophyllus in diurnal coloration (photo by A. FOUQUET) from Pacaraima, Roraima, Brazil. (E,
F) igh patterns of two specimens of H. xerophyllus (E = MZUSP65854 and F = MZUSP68669). Scale bar = 1 cm.
105
Taxonomy of Hypsiboas crepitans
small melanophores on the dorsal surface, like the holo-
type of H. fuentei. e nostril shape, the reduced webbing,
and the absence of dermal ridges of the holotype conform
to what is observed in specimens of the NG.
Discussion
Taxonomic conclusions
Although previous authors have used larval morphology
and advertisement call data (C J ) to sug-
gest that populations of Hypsiboas crepitans from the two
geographical groups could represent two distinct species,
published data on this subject remained ambiguous (see
Introduction). Numerical parameters of advertisement call
overlap (or nearly so, see Table ), and call descriptions,
especially for the populations of the NG, are too succinct
and do not address intraspecic variation and other fac-
tors known to inuence amphibian calls (e.g., the social
context; see W G ). Larval morphology of
Hypsiboas species presents high levels of intraspecic vari-
ation and many closely related species have nearly indis-
tinguishable larvae (see K et al. , O et al.
). However, since important character states are some-
times overlooked in tadpole descriptions (see K et
al. ), we cannot exclude that the larvae of the two dis-
tributional groups are actually distinct. Additional studies
are needed for a comprehensive comparison of larval mor-
phology and bioacoustics of the two groups.
However, the molecular and morphological data pre-
sented herein are concordant with the recognition of the
populations from the two distributional groups as dis-
tinct species. e SG is restricted to eastern Brazil, from
the State of Paraíba to the northern State of Rio de Janeiro,
including the State of Minas Gerais. e type locality of
H.crepitans was originally designated as “Tamburil, Jiboya,
Arrayal da Conquista”, State of Bahia, Brazil (F ).
B () restricted it to “Tamburil, [Municipal-
ity of] Condeúbas, Bahia, Brazil”. Based on the aforemen-
tioned characteristics, type series data, and type locality,
the SG is considered to represent H. crepitans (Fig. ).
e NG occurrence area comprises a region where ve
taxon names are available. ree names were proposed
Figure 3. Plots of individual scores resulted from Principal Com-
ponent Analysis (PCA) of morphometric data from two groups
of adult males (A) and adult females (B) of Hypsiboas crepitans
(SG) and H. xerophyllus (NG) in the space of the rst with the
second canonical axes. Condence ellipses (95%) for the scores
of each group are shown.
Table 2. Mean intra (rst column) and inter-specic uncorrected pairwise distances among species of the Hypsiboas faber group. Above
the diagonal 12S (907 bp); below, COI (671 bp).
[COI/12S] 1 2 3 4 5 6 7 8 9
1H. crepitans [0.01/0] – 0.04 0.07 0.06 0.07 0.09 0.07 0.08 0.09
2H. xerophyllus [0.01/0] 0.13 – 0.06 0.05 0.07 0.08 0.07 0.07 0.08
3H. rosenbergi [–/0.03] – – – 0.07 0.08 0.09 0.09 0.09 0.1
4H. pugnax [–/0] – – – – 0.07 0.07 0.07 0.07 0.07
5H. faber [0.06/0.02] 0.18 0.17 – – – 0.06 0.05 0.05 0.08
6H. exastis [–/–] 0.17 0.2 – – 0.16 – 0.03 0.05 0.08
7H. pardalis [0/0] 0.19 0.2 – – 0.15 0.11 – 0.04 0.08
8H. lundii [0.01/0] 0.18 0.18 – – 0.16 0.15 0.15 – 0.08
9H. albomarginatus [0.11/0.02] 0.17 0.17 – – 0.18 0.17 0.18 0.18 –
life also, but specimens from a given population can ex-
hibit the dark green coloration while other present the pale
coloration. Moreover, many specimens of the NG present
106
V G. D. O et al.
by D B (): Hyla doumercii D
B, , from Suriname (holotype MNHN
[Fig.A], SVL . mm, adult male, vocal slits present, sup-
plementary bone on Finger II =prepollical spine; A. O-
, pers. comm.); Hyla levaillantii D B,
, from Suriname (holotype MNHN [Fig. B], SVL
. mm, adult male, vocal slits present, prepollical spine
on Finger II); Hyla xerophylla D B, ,
from “Cayenne”, French Guiana (holotype MNHN
[Fig. C], SVL . mm, probably an adult female, no male
sexual secondary character, at tubercle instead of prepol-
lical spine). Article of the International Code of Zoo-
logical Nomenclature (ICZN ) argues about the “Prec-
edence between simultaneously published names, spellings
or acts”. Following the rules of the Code, we cannot apply
the Section . about the “Determination by First Revis-
er” because this was already done (see D ,
K ).
Although we cannot establish precedence, Article of
the Code still allows us to choose which of the names of
D B () we will revalidate if applicable
to a natural population. us, any of the three names de-
scribed by D B () are available and can
be applied to the NG. Herein, we decided to consider the
Figure 4. Phylogram (50% majority rule consensus with frequencies of all observed bipartitions) hypothesized from Bayesian analysis
using 1578 bp of 12S and COI. Numbers above nodes are posterior probabilities (* indicates pp ≥ 0.99; pp < 0.5 are not indicated). Sam-
ples are labeled according to collection numbers followed by origin (country in full or Brazilian state ocial acronym if from Brazil).
107
Taxonomy of Hypsiboas crepitans
Figure 5. Dorsal and ventral views of the holotypes of (A) Hyla
doumercii [SVL = 47.2 mm, male], (B) H. levaillantii [SVL =
48.1 mm, male], (C) H. xerophyllus [SVL = 44.9 mm, likely a
female], and (D)H. fuentei [SVL = 57.0 mm, female]. Figures
are not to scale.
NG as Hypsiboas xerophyllus, resurrecting the name Hyla
xerophylla D B, from the synonymy
of Hypsiboas crepitans (W, ) and transferring the
species to the genus Hypsiboas based on phylogenetic re-
lationships.
e two remaining names, Hypsiboas indris C,
and Hyla fuentei G G, , are more recent and
based on material from Suriname. Hyla fuentei holotype
from “Suriname, Suriname District Powakka” (CM
[Fig. D], SVL . mm, adult female; G G )
presents a colour pattern in accordance with what was stat-
ed by G G (). e sagittal opening at the ab-
domen seems to be made for information on ovaries and
oocytes (see G G ). In fact, all our obser-
vations about the holotype agree well with G G
(). In addition, e holotype agreed with the NG pop-
ulations and they exhibit a dark green coloration, while
other present the pale coloration, a phenomenon proba-
bly not uncommon in Hypsiboas, as also revealed in An-
dean species of the H. pulchellus species group (K
et al. ). Aer the examination of the holotype of Hyla
fuentei we considered this species as belonging to the NG
and, thus, to be a junior synonym of the name applicable
to this group (H. xerophyllus). We have not examined the
holotype of Hypsiboas indris C, . However, the dis-
tribution of this taxa completely overlaps with the more in-
clusive distribution of the NG. In the light of the present
results – where the genetic diversity within each group is
low (Table ) – it seems very unlikely that it represents a
dierent species from H. crepitans. erefore, we provi-
sionally transfer it from the synonymy of Hypsiboas crepi-
tans to the synonymy of Hypsiboas xerophyllus.
Species account
Hypsiboas xerophyllus (D B, ).
New combination, revalidation
Hyla levaillantii D B, – Erp. Gen. :. New
synonymy.
Hyla doumercii D B, – Erp. Gen. :. New
synonymy.
Hyla fuentei G G, – Copeia :. New synony-
my.
Hypsiboas indris C – J. Acad. Nat. Sci. Philadelphia, Ser.
, : . New synonymy.
Holotype: MNHN , SVL . mm, probably female (no
male secondary characters), “Cayenne” [= French Guiana]
(Fig. C).
Diagnosis: Hypsiboas xerophyllus is a member of the H. fa-
ber species group (sensu F et al. ), character-
ized by: SVL .–. mm in adult males, .–. mm
in adult females; in dorsal view, head nearly rounded with
rounded, truncated, or sub-elliptical snout; single, well de-
veloped projecting prepollex in adult males; tympanum
108
V G. D. O et al.
and tympanic annulus visible externally and in contact
with the posterior margin of eye; well-developed supra-
tympanic fold covering the upper edge of the tympanum;
lower edge of the tympanum close to the mouth (at jaw
articulation level); males with vocal slits under the tongue
and parallel to the lower lip, vocal sac median, subgular
(sensu L ); presence of two groups of vomerine teeth
between choanae; dermal ridges (mbriae) absent or ru-
dimentary along arms and feet; skin smooth dorsally and
granular ventrally; colour of gular region similar to belly
colour in preserved males and females (cream); cloacal re-
gion composed by a subcloacal fold, white tubercles around
the cloacal opening, and a well-developed ap (horizontal,
above vent); anks usually show parallel transverse thin
bars; well-developed tubercles on the ventral surface of
thighs; dorsal surface of thighs usually with parallel bars,
if present, spaced and wider dorsally, thinner and lighter
posteriorly; colour on dorsum ranging from pale gray to
brown in preservative, with a X-shaped mark sometimes
over the suprascapula.
Comparisons with other species: Hypsiboas xerophyllus is
easily dierentiated from H. crepitans, its sister taxon and
morphologically most similar species, by the absence of a
mid-dorsal longitudinal dark brown stripe (present, of dif-
ferent widths, sometimes incomplete – i.e., not connect-
ing snout and vent – in H. crepitans); immaculate cream
gular region in both sexes (brown in males of H. crepi-
tans); absence of, or barely visible, bifurcated vertical dark
brown bars on ventroposterior surfaces of thighs (present
in H.crepitans; Fig. ).
From other species of the H. faber group, H. xerophyllus
diers by having a smooth texture of skin on dorsal sur-
faces, and low development of ulnar and outer tarsal der-
mal ridges (skin texture lumpy, and well developed dermal
ridges in H. pardalis, H. exastis, H. rosenbergi, and H. lun-
dii). e absence of well-developed calcars distinguishes
H. xerophyllus from H. pardalis, and H. exastis. e ab-
sence of extensive hand webbing distinguishes H. xero-
phyllus from H. pardalis, H. exastis, and H. lundii. e su-
pratympanic fold of H. albomarginatus is white (or whitish
green in live specimens) while in H. xerophyllus it presents
the same lichenous colour as the dorsum. e gular region
is immaculate cream in H. xerophyllus while it has brown
ecks in H. pugnax. In life, the iris coloration is whitish
around the pupil and yellow-green on he outer edge in
H.xerophyllus while it is golden yellow in H. pugnax.
Figure 6. Lectotype of Hypsiboas crepitans, AMNH 785, adult female, SVL ca. 69 mm, from Vitória da Conquista, State of Bahia, Brazil.
Table 3. Mean pairwise distances within species of the Hypsiboas
faber group and within other groups (gr.) of Hypsiboas. 12S se-
quences have 907 bp; COI, 671 bp.
12S COI
H. albomarginatus 0.02 0.11
H. albopunctatus gr. 0.04 0.13
H. crepitans 0.00 0.01
H. exastis – –
H. faber 0.02 0.06
H. lundii 0.00 0.01
H. pardalis 0.00 0.00
H. pellucens gr. 0.01 0.11
H. pugnax 0.00 –
H. pulchellus gr. 0.08 –
H. punctatus gr. 0.09 –
H. rosenbergi 0.03 –
H. xerophyllus 0.00 0.01
109
Taxonomy of Hypsiboas crepitans
Colour in preservative: Dorsal background light brown to
cream with dark brown blotches; an interocular stripe and
a X-shaped mark on the dorsum can be present. Flanks
with well-dened dark brown transverse stripes. Chest and
gular region beige, edge of lips green. Belly and ventral sur-
faces green, sometimes orange. Fingers beige, dark green,
or orange. Iris grey with green outer border.
Variation: Hypsiboas xerophyllus presents some variation
in size, with females usually larger than males (Table ).
Adult males have hypertrophied forearms, enlarged and
projecting prepollices, and vocal slits, all characteristics
that are absent in females. In dorsal view, shape of head
ranges from rounded to oval, and in lateral view from
rounded to truncated. Specimens usually lack any colour
pattern at inguinal region and posterior surface of thighs.
In living specimens, the dorsal pattern can vary in colour
(white, green, or brown), oen depending on light inten-
sity, and dorsal blotches can be present or absent.
Distribution: Eastern Panama, through northern Co-
lombia, Venezuela, the Guiana Shield, including adjacent
northwestern Brazil, below , m a.s.l. e species seems
absent from most French Guiana, only occurring around
inselbergs at isolated localities in the south and in dis-
turbed forest along the lower course of the Maroni River
(Apatou, Saint Laurent). e species has not been recorded
in the states of Amapá (AF pers. obs., D-L )
and Pará, north of the Amazon River (Á-P et al.
). However, it occurs throughout most Guyana (C
et al. ) and northern Suriname (O J
), although apparently absent from Kaieteur National
Park (K K ).
Natural history: According to D (; as
H. crepi tans), specimens from Panama do not exhibit
the habit to build clay nests. However, L R-
() reported this behavior as usual in Colombian
populations, and L () conrms this behav-
ior for Trinidad populations. One of us (AF) observed
building of clay nests in the state of Roraima in Brazil and
in Suriname. erefore, nest building behavior is likely
facultative as already observed by C () in
other species of Hypsiboas. Males have been observed
calling at night in small streams, ponds, and even asks
during the rainy and the dry season. e species seems
to inhabit ecotonal forest and is oen found in disturbed
habitat.
Hypsiboas crepitans (D B, )
Lectotype: AMNH , SVL ca. mm, adult female,
from Tamburil, [Municipality of] Condeúba, Bahia, Bra-
zil (Fig.).
Diagnosis: Hypsiboas crepitans is a member of the H. faber
species group (sensu F et al. ), characterized
by: SVL .–. mm in males and .–. mm in fe-
males; interdigital membranae poorly developed; mbrias
absent in arms and tibia; Dorsal coloration brown, usually
with an X-shape above the scapular region or a fragmented
mid-dorsal line; dorsal skin smooth; ank and thigh with
conspicuous transversal bars; absence of calcar ap or ap-
pendage; presence of subcloacal dermal fold, surrounded
by granules; upper cloacal ap poorly developed; absence
of lichenous subcloacal plate.
Table 4. Descriptive statistics (in millimeters) for measurements of Hypsiboas crepitans and H. xerophyllus. e results are presented
as mean ± standard deviation (range).
Measures Hypsiboas crepitans H. xerophyllus
Males (n=95) Females (n=70) Males (n=98) Females (n=13)
SVL 58.1±7.6 (33.2–71.8) 55.4±4.59 (51.0–64.7) 50.6±3.86 (42.9–63.8) 57.7±7.91 (40.9–71.8)
HL 19.2±2.2 (11.4–22.8) 18.4±2.23 (14.7–21.6) 16.6±1.5 (13.7–19.9) 19.1±2.21 (13.8–22.8)
HW 20.3±2.5 (12.0–24.1) 18.9±1.32 (17.1–21.3) 17.7±1.25 (15.4–21.7) 20.0±2.51 (14.5–24.1)
ED 6.1±0.7 (3.2–7.3) 2.2±0.15 (1.9–2.5) 5.4±0.54 (4.1–6.9) 3.7±0.43 (2.9–4.7)
END 5.6±0.7 (3.5–6.8) 3.6±0.39 (2.8–4.3) 5.5±0.72 (4.3–9.8) 1.9±0.28 (1.3–2.6)
TD 4.4±0.6 (2.1–5.5) 4.6±0.47 (4.0–5.5) 4.4±0.39 (3.4–5.3) 4.4±0.69 (2.7–6.3)
UEW 4.6±0.7 (2.7–6.3) 5.6±0.47 (4.7–6.5) 4.4±0.63 (2.2–6.0) 6.0±0.67 (4.2–7.3)
IOD 10.8±1.2 (7.0–12.8) 5.7±0.50 (5.0–6.6) 5.4±0.63 (4.0–7.0) 10.7±1.21 (7.7–12.8)
IND 3.8±0.5 (1.7–4.8) 5.6±0.44 (5.0–6.1) 3.3±0.66 (1.8–4.6) 5.5±0.69 (4.2–6.8)
NSD 1.9±0.3 (1.1–2.6) 4.5±0.50 (3.7–5.3) 2.5±0.63 (1.6–3.9) 4.4±0.63 (2.1–5.5)
FAL 27.7±3.6 (15.4–34.4) 25.7±3.46 (16.8–30.3) 23.9±2.1 (12.0–29.6) 27.4±3.55 (19.2–34.4)
4FD 2.6±0.4 (1.5–3.4) 2.7±0.38 (2.1–3.2) 2.4±0.25 (1.8–3.3) 2.6±0.38 (1.8–3.4)
THL 32.4±4.2 (18.3–40.0) 30.6±1.91 (27.7–34.1) 28.3±1.92 (23.1–34.8) 31.9±4.19 (23.2–40.0)
TL 31.7±4.1 (18.0–39.3) 30.4±1.85 (27.8–33.8) 28.0±1.99 (23.0–35.0) 31.5±4.13 (23.3–39.3)
FL 40.8±5.2 (22.5–51.9) 38.6±2.40 (34.5–42.4) 35.5±2.61 (28.8–44.3) 40.4±5.14 (30.2–51.9)
4TD 2.3±0.3 (1.2–3.0) 2.4±0.32 (2.0–3.0) 2.1±0.25 (1.5–2.9) 2.2±0.33 (1.6–2.9)
110
V G. D. O et al.
Distribution: Central to eastern Brazil, from the Atlan-
tic coast of the states of Rio de Janeiro to Paraíba (present
study).
Phylogenetic relationships and biogeography
e Hypsiboas faber species group, as dened by F-
et al. (), has not been recovered as monophylet-
ic in our results – the weak relationship between H. albo-
marginatus and the remaining species of the group has
been already reported by previous authors (K et
al. , W et al. ), and the nested position of
the H. albo punctatus group within the H. faber group is
strongly supported. However, we refrain to take any for-
mal action regarding this matter as our molecular anal-
yses were simply aimed at exploring genetic divergence
within H.crepitans.
F et al. () listed the H. faber group as an
example of a clade having an Atlantic Forest (or at least
an eastern Brazilian) origin with subsequent radiations
into neighbouring regions. At that time the authors had
only Atlantic Forest dwellers in their dataset (their sam-
ple of H. crepitans was from State of Alagoas, Brazil) and
probably assumed that H. albomarginatus would be sister
to the remaining species of the group based on its unique
colour. Our results suggest a more complex history of suc-
cessive dispersals between Amazonia, Cerrados and the
Atlantic Forest. However, the lack of support at the base
of the clade prevents any further biogeographic analyses,
and additional data are needed. Nevertheless, the H. faber
clade is endemic to eastern Brazil while the H. crepitans
clade is restricted to northern South America and Panama
with the exception of H. crepitans. Such a pattern suggests
that H. crepitans originates from a dispersal event from
Amazonia to the Atlantic Forest probably via a northern
route (C ). Given the genetic distances between
H. crepitans and H. xerophyllus we hypothesize that this
dispersal occurred before the Pleistocene and was probably
favored by forest connection due to more humid climate.
Such a route is supported by biogeographical analyses of
dierent groups of organisms, as well as by climatic and
oristic evidence (B-F et al. , C et
al. , C , M S et al. , W et
al., ).
Hypsiboas pardalis and H. lundii were long consid-
ered synonyms (C N ) due to
their morphological similarity. Although the relationship
of H.exastis with H. pardalis and H. lundii was not unex-
pected (see C R ), we – unex-
pectedly – found that H. lundii is instead sister to a clade
composed of H. exastis + H. pardalis. C R-
() related H. exastis to H. pardalis and H. lun-
dii due to their similar skin texture and colour pattern; the
lichenous aspect similar to tree bark were evidence that
these species were more closely related to each other than
to all other members of the Hyla boans species group, as
dened at that time.
e IUCN distribution for Hypsiboas crepitans (L
M et al. ) and H. xerophyllus needs to be re-eval-
uated. Although F () stated that the species oc-
curs in the south of the Atlantic Forest, from São Paulo
to Santa Catarina states, we were unable to nd a speci-
mens from this region (Fig. ). We propose that the south-
ern part of the IUCN distribution is based on misidenti-
ed H. lundii or H. pardalis specimens and that the dis-
tribution of H. crepitans (as re-dened here) is restricted
to an area north of o southern latitude. Despite the fact
that the respective ranges are now conned to a smaller
portion of the formerly supposed range of H. crepitans,
both species are still widespread and, given their habitat
requirements, are likely relatively tolerant to human dis-
turbance. erefore, we suggest considering them as Least
Concern (LC).
Acknowledgements
We are grateful toward S. R, P. J. R. K and an anonymous
reviewer for valuable comments on a previous version. For their
invaluable help and company in the eld and for the gi of tis-
sue samples, we thank P. G, M. B, J. C,
C. M. C, F. C, M. D, A. C, R. D-
, D. P, R. R, M. S, S. M. S, M.
T J., V. V, H. N, and D. F. M-
J. Pictures of the holotype of Hypsiboas fuentei were
kindly provided by S. R and J. M. P. VGDO thanks
Sao Paulo Research Foundation (FAPESP; grants /-
, /-, BIOTA /-) and ANPCyT PICT
/ for nancial support and C. S. C for support
and comments on a previous versions of this manuscript. AF
beneted from an “Investissement d’Avenir” grant managed
by Agence Nationale de la Recherche (CEBA, ref.ANR--LA-
BX--), France. IN thanks Fundação Carlos Chagas Filho
de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ)
and Conselho Nacional de Desenvolvimento Cientíco e Tec-
nológico (CNPq) for support and fellowship, respectively. CM
thanks Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior (CAPES) for fellowship. UC thank CNPq for nan-
cial support. MLL thanks FAPESP (/-) and CNPq
(/-; /-) for fellowship and nancial
support. CFBH and MTR thank FAPESP (Grant /-
) and CNPq for nancial support. is study was co-funded
by NSF (DEB ) and NASA. is research was supported
by the Núcleo de Computação Cientíca (NCC/GridUNESP)
from the Universidade Estadual Paulista ‘‘Júlio de Mesquita Fil-
ho’’ (UNESP). e Centro de Estudos de Insetos Sociais (CEIS),
UNESP, Rio Claro, Brazil, allowed the utilization of the facilities
for molecular analyses.
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Supplementary material
Supplementary document . Examined specimens
Supplementary document 2. Voucher information, localities,
and GenBank accession numbers for the specimens analyzed
for this study.
Supplementary document 3. Standardized coecients and factor
loadings (r) from a Principal Component Analysis.
I
Online Supplementary data – Taxonomy of Hypsiboas crepitans
Supplementary document 1. Examined specimens
Hypsiboas albomarginatus
Brazil: Estado de Alagoas: MNRJ Fazenda Santa Justina, Pas-
so de Camaragibe; MNRJ – Murici; MNRJ , ,
– São Miguel dos Campos; MNRJ – Rio
Largo.
Brazil: Estado da Bahia: MNRJ Reserva Particular do
Patrimônio Natural Cachoeira da Água Branca, Valença; MNRJ
Mata da Cara Branca, Santa Cruz Cabrália; MNRJ Fei-
ra de Santana; MNRJ Reserva Particular do Patrimônio Nat-
ural Capitão, Itacaré; MNRJ – Fazenda Tiririca, Conde;
MNRJ Entre Rios; MNRJ Fazenda São José, Mascote;
MNRJ – Fazenda Pedra Branca, Itagibá; MNRJ
Reserva Particular do Patrimônio Natural Estação de Vera Cruz,
Porto Seguro; MNRJ Serra do Timorante, Valentim, Boa
Nova; MNRJ Reserva Particular do Patrimônio Natural Ser-
ra do Teimoso, Jussari.
Brazil: Estado do Espírito Santo: MNRJ Santa Teresa;
MNRJ , – Praia das Neves, Presidente Kennedy;
MNRJ Costa Bela, Serra; MNRJ – Parque Es-
tadual Fonte Grande, Vitória; MNRJ –, , ,
Reserva Biológica de Duas Bocas, Cariacica; MNRJ –
Barra do Riacho, Aracruz; MNRJ – Pedra do
Garrafão, Mimoso do Sul; MNRJ – Guarapari; MNRJ
, , – Reserva Biológica de Santa Lúcia,
Santa Teresa; MNRJ – Mimoso do Sul; MNRJ
Setiba, Guarapari; MNRJ Pedra Azul, Domingos Martins;
MNRJ – Ponte da Chinchinha, Povoação, Linhares.
Brazil: Estado de Minas Gerais: MNRJ – Chia-
dor; MNRJ Faria Lemos; MNRJ , – Além
Paraíba; MNRJ – Pádua; MNRJ – Faria
Lemos.
Brazil: Estado do Paraná: MNRJ RPPN Salto Mor-
ato, Guaraqueçaba; MNRJ – Estação II, Instituto
Agronômico do Paraná, Morretes.
Brazil: Estado de Pernambuco: MNRJ –, –
Refúgio Ecológico Charles Darwin, Igarassu.
Brazil: Estado de Rio de Janeiro: MNRJ Jacarepaguá, Rio
de Janeiro; MNRJ , – Pedra de Guaratiba, Rio
de Janeiro; –, –, – Ilha de Ma-
rambaia, Mangaratiba; MNRJ – Vila Dois Rios, Ilha
Grande, Angra dos Reis; MNRJ , , – Horto
Florestal de Santa Cruz, Itaguaí; MNRJ , , –,
– Duque de Caxias; MNRJ Restinga Maricá,
Maricá; MNRJ Manguinhos, Rio de Janeiro; MNRJ
Araruama; MNRJ , Fazenda Cachoeira, Morro Azul,
Engenheiro Paulo de Frontin; MNRJ – Ilha Grande,
Angra dos Reis; MNRJ Vale das Pedrinhas, Magé; MNRJ
Lídice, Rio Claro; MNRJ Grussaí, Campos dos Goyt-
acazes; MNRJ Ponta Negra, Maricá; MNRJ , ,
–, – Tanguá; MNRJ –, –
Espraiado, Maricá; MNRJ Peró, Cabo Frio; MNRJ
–, Reserva Ecológica de Guapiaçu, Cachoeiras
de Macacu; MNRJ Centro Marista São José das Paineiras,
Mendes; MNRJ – Estrada Além Paraíba – Carmo,
Carmo; MNRJ Restinga Massambaba, Arraial do Cabo;
MNRJ Itaperuna; MNRJ Itapebussus, Rio das Os-
tras; MNRJ Silva Jardim; MNRJ , São Francis-
co de Itabapoana; MNRJ Núcleo Experimental de Iguaba
Grande–UFF, Iguaba Grande; MNRJ – Pontal do Ata-
laia, Arraial do Cabo; MNRJ , –, São João
da Barra; MNRJ –, – Área da PCH Santa Fé,
Três Rios; MNRJ – Ponte Preta, Magé; MNRJ
Paracambi; MNRJ , – Saquarema; MNRJ –
Reserva Biológica União, Casimiro de Abreu; MNRJ –
Tinguá, Nova Iguaçu; MNRJ Parque Natural Mu-
nicipal da Atalaia, Macaé; MNRJ – Itaperuna; MNRJ
– Parque Natural Municipal da Prainha, Grumari, Rio
de Janeiro; MNRJ Tarituba, Paraty; MNRJ Bom Jesus
de Itabapoana; MNRJ Parque Nacional Restinga de Jurubat-
iba, Macaé; MNRJ – Parque Natural Municipal Bosque
da Barra, Rio de Janeiro; MNRJ Parque Natural Municipal
da Prainha, Rio de Janeiro.
Brazil: Estado de Santa Catarina: MNRJ – Palhoça.
Brazil: Estado de São Paulo: MNRJ –, Praia de
Itaguaí, Ubatuba; MNRJ Picinguaba, Ubatuba; MNRJ –
Pedro de Toledo; MNRJ – Praia de Barequeçaba,
São Sebastião; MNRJ Eldorado Paulista; MNRJ Fa-
zenda Jamapa, Jacupiranga; MNRJ Guarujá; MNRJ
Parque Estadual Intervales, Ribeirão Grande; MNRJ Barra do
Quilombo, Sete Barras; MNRJ – Estação Ecológica da
Juréia, Iguape; MNRJ , – Pariquera-Açu.
Brazil: Estado de Sergipe: MNRJ , Fazenda Cruzei-
ro, Cristinápolis; MNRJ São Cristóvão; MNRJ , –
Santa Luzia do Itanhy; MNRJ –, – Fa-
zenda CICP, Itaporanga d’Ajuda.
Hypsiboas crepitans
Brazil: Estado de Alagoas: MZUSP UHE Xingó, Vila Resi-
dencial, Piranhas; MZUSP , Rio Largo, Fazenda Canoas;
MZUSP UHE Xingó; MNRJ Fazenda São Bento, Água
Branca; MNRJ –, – São Miguel dos Campos;
MNRJ – Fazenda do Prata, São José dos Campos; MNRJ
Maceió.
Brazil: Estado da Bahia: MNRJ , – km , Ro-
dovia Jequié–Salvador, Salvador; MNRJ Salvador; MNRJ
– Fazenda Tiririca, Conde; MNRJ , ,
, , , , , , , , , ,
–, –, –, UFBA Fazenda Pe-
dra Branca, Itagibá; MNRJ Parque Municipal da Lagoa Azul,
São Desidério; MNRJ , , , , , ,
, , , –, –, , ,
, – Maracás; MNRJ , , , ,
Parque Estadual Serra do Conduru - Setor Norte, Itacaré;
MNRJ , – Boa Nova; MNRJ Mata da Cara
Branca, Santa Cruz Cabrália; MNRJ Fazenda Pedra Formo-
sa, Ibirapitanga; MNRJ –, , –, ,
UFBA , , , , , , , , , ,
Caetité; MNRJ , – Trancoso, Porto
Seguro; MNRJ , , –, –, –
Jequié; MNRJ – Fazenda Orion, Arataca; MNRJ
, CEPLAC, Ilhéus; MNRJ – Fazenda Vista
Bela, Guaratinga; MNRJ , – Januária; MNRJ
Uruçuca; MNRJ Calumbi, Macaúbas; MNRJ , UFBA
, Barreiras; MNRJ Alagoinhas; MNRJ –
RPPN Serra do Teimoso, Jussari; MNRJ , , ,
Salvador; UFBA , , Pratigi; UFBA ,
Mucugê; UFBA Bom Jesus da Lapa; UFBA , En-
cruzilhada; UFBA , Serra do Ramalho; MNRJ no
locality provided.
Brazil: Estado do Espírito Santo: MBML , Sítio do Am-
arildo, Marechal Floriano; MNRJ Alto Nova Almeida, Mare-
chal Floriano; MBML , , MZUSP , , ,
, , Colatina; MBML , , Aparecid-
inha, Santa Teresa; MBML Entorno da ESFA, Santa Teresa;
MBML Bairro Vila Nova, Santa Teresa; MBML Fazenda
Recanto da Mata, Muniz Freire; MNRJ São Simão, Muniz
II
Online Supplementary data – V G. D. O et al.
Freire; MBML , , Três Barras, Fundão; MBML Goi-
apaba-Açu, Fundão; MBML Barra de São Francisco; MBML
Cabeceira do Rio Mutum, São Roque do Canaã; MBML
Anutiba, Pedra da Severina, Alegre; MZUSP Samarco, An-
chieta; MZUSP , MNRJ , Baixo Guandu; MZUSP
, , sem maiores procedências.
Brazil: Estado de Goiás: MNRJ – Mambaí; MNRJ
, Ilha, Rio Paranã; MZUSP , , , ,
, , , PCH Santa Edwiges; MZUSP ,
, Monte Alegre de Goiás, Rio Raiz; MNRJ Cat-
alão; MNRJ , Flores de Goiás.
Brazil: Estado de Minas Gerais: MNRJ –, ,
–, –, – Mocambinho, Manga;
MNRJ , Fazenda Curral Velho, Cristália; MNRJ –
, , –, , – Peixe Cru, Turma-
lina; MNRJ –, , , Berilo; MNRJ
Veredas, Botumirim; MNRJ – Córrego Santa Rita,
Paraíso, Espera Feliz; MNRJ , – Serra do Cipó,
Santana do Riacho; MNRJ – Fazenda Cachoeira Ale-
gre, Abre Campo; MNRJ Fazenda Todos os Santos, Faria
Lemos; MNRJ , Capitão Andrade; MNRJ –
RPPN Santuário do Caraça, Catas Altas; MNRJ –,
– Fazenda do Engenho, Serra do Caraça, Catas Altas;
MNRJ Paula Cândido; MNRJ – Estrada Gov-
ernador Valadares–Mantena, km ; MNRJ Minas Novas;
MNRJ Grão Mogol.
Brazil: Estado da Paraíba: MZUSP – Fazenda Salga-
do, Gurinhém; MNRJ , , , Boqueirão.
Brazil: Estado de Pernambuco: MZUSP – Bom
Conselho; MZUSP – Serra dos Cavalos, São Caetano;
MZUSP – Engenho Sacramento; MZUSP ,
–, MNRJ – Caruaru; MNRJ Rio Bran-
co; MNRJ Serra da Borborema.
Brazil: Estado de Sergipe: MNRJ Riachuelo; MNRJ –
Fazenda Sabão, Indiaroba; MNRJ Fazenda Cruzeiro,
Cristinápolis.
Hypsiboas exastis
Brazil: Estado de Alagoas: ZUFRJ , , , Que-
brangulo.
Brazil: Estado da Bahia: UFBA Amargosa; UFBA Elí-
sio Medrado; MNRJ (parátipo) Estação Ecológica Estadual
Nova Esperança, Wenceslau Guimarães; MNRJ (parátipo),
MZUSP (parátipo), (holótipo), – (paráti-
pos) Fazenda Unacau, São José; MNRJ – Fazenda
Capitão, Itacaré; MNRJ Fazenda Serra Verde, Jussari; MNRJ
Rio Vermelho, Três Braços, Teolândia; MNRJ Amar-
gosa.
Hypsiboas faber
Brazil: Estado da Bahia: MNRJ Fazenda Serrinha, Itagibá;
MNRJ , Torrão, Prado; MNRJ , Maracás; MNRJ
Trancoso, Porto Seguro; MNRJ Itabuna; MNRJ
Jussari; MNRJ – Teolândia; MNRJ Porto Seguro.
Brazil: Estado do Espírito Santo: MNRJ Sítio do Popota,
São Lourenço, Santa Teresa; MNRJ Parque Estadual do For-
no Grande, Castelo; MNRJ Pedra do Garrafão, Mimoso do
Sul; MNRJ – Floresta Nacional do Rio Preto, Con-
ceição da Barra; MNRJ – Domingos Martins.
Brazil: Estado da Paraíba: MNRJ Reserva Biológica
Guaribas, Mamanguape.
Brazil: Estado do Paraná: MNRJ APA SEMA, Guar-
aqueçaba.
Brazil: Estado do Rio de Janeiro: MNRJ Saquarema;
MNRJ Pendotiba, Niterói; MNRJ , APA Pau
Brazil, Búzios; MNRJ Fazenda Brasiléia, Natividade; MNRJ
Quitandinha, Petrópolis; MNRJ Capela, Petrópolis;
MNRJ Ponta Negra, Maricá; MNRJ Lídice, Rio Claro;
MNRJ Pedreira Quatro Irmãos, Santo Antônio de Pádua;
MNRJ Fazenda Vargem Alegre, Porciúncula; MNRJ
Trajano de Moraes; MNRJ Ilha Grande, Angra dos Reis.
Brazil: Estado do Rio Grande do Sul: MNRJ Farroupilha.
Brazil: Estado de Santa Catarina: MNRJ – Porto
Belo; MNRJ – Santo Amaro da Imperatriz.
Brazil: Estado de Sergipe: MNRJ Mata do Castro, Santa
Luzia do Itanhy.
Brazil: Estado de São Paulo: MNRJ Lageado, Botucatu.
Hypsiboas lundii
Brazil: Estado de São Paulo: MNRJ –, –,
–, –, –, –, –
, Lageado, Botucatu; MNRJ – Chácara
Furlan, Botucatu; MNRJ Rubião Júnior, Botucatu; MZUSP
Corumbataí; MZUSP Garça; MZUSP Usina
Ester, Engenho Coelho; MZUSP PCH Eleutério, Espírito
Santo do Pinhal.
Brazil: Estado de Minas Gerais: MNRJ , – Es-
tação Biológica Vereda Grande, Presidente Olegário; MNRJ –
Fazenda Cascata (sede), Patos de Minas; MNRJ –,
, , –, – Fazenda Gameleira, João
Pinheiro; MNRJ Fazenda Rio Verde, João Pinheiro; MNRJ
– Capão da Água Limpa, João Pinheiro; MNRJ –
Uberlândia; MNRJ Alegre, Grão Mogol; MNRJ
– Fazenda Cabral, Cristália; MNRJ Córrego do
Gigante, Turmalina; MNRJ Peixe-Cru, Turmalina; MNRJ
AHE Queimado; MNRJ Usina Hidrelétrica de Mi-
randa, Indianópolis; MNRJ Ribeirão das Moendas, Pains;
MNRJ – Fazenda Índia, Brumadinho; MNRJ
Ibituruna; MNRJ – Sete Lagoas; MZUSP Lagoa
Santa (topótipo); MZUSP Belo Horizonte.
Brazil: Estado de Goiás: MNRJ Catalão; MNRJ –
, – EFLEX (Estação Florestal Experimental), Sil-
vânia; MNRJ – Santo Antônio do Descoberto; MNRJ
, MZUSP –, Chapada dos Veadeiros;
MZUSP , –, , – Luziânia;
MZUSP – PCH Santa Edwiges I, Mambaí; MZUSP
, , , – PCH Santa Edwiges II,
Buritinópolis; MNRJ – Unaí (MG) e Formoso (GO);
MZUSP , –, , , Serra da Mesa;
MZUSP –, –, , , , ,
, UHE Corumbá, Caldas Novas.
Brazil: Estado de Mato Grosso: MZUSP , , –
Alto Araguaia.
Brazil: Distrito Federal: MNRJ Mata do Gedige, Brasília.
Brazil: Estado de Tocantins: MZUSP Paranã.
Hypsiboas pardalis
Brazil: Estado da Bahia: MNRJ Guaratinga.
Brazil: Estado do Espírito Santo: MNRJ , , –
, MBML , , MZUSP , – Santa Ter-
esa; MBML , , , , , Nova Lombardia,
Santa Teresa; MNRJ , MBML , , , , Es-
tação Biológica de Santa Lúcia, Santa Teresa; MBML Bairro do
Eco, Santa Teresa; MBML , , Alto Rio Saltinho, Santa
Teresa; MBML Sítio Pousada Canaã, Santa Teresa; MBML
Vila Nova, Santa Teresa; MBML Associação do Banestes,
Santa Teresa; MNRJ , MBML , , , , ,
Reserva Biológica de Duas Bocas, Cariacica; MBML ,
, , , , , , Alto, Reserva Biológica
de Duas Bocas, Cariacica; MBML Cariacica; MNRJ ,
III
Online Supplementary data – Taxonomy of Hypsiboas crepitans
, , MBML Parque Estadual do Forno Grande,
Castelo; MBML , Castelo; MNRJ FLONA Goytacaz-
es, Linhares; MNRJ Fazenda Oliveira, Guaçuí; MNRJ
Fazenda dos Japoneses, Vargem Alta; MNRJ , MBML ,
Parque Municipal Goiapaba-açu, Fundão; MNRJ Alto
Nova Almeida, Marechal Floriano; MBML , , ,
Sítio Amarildo, Marechal Floriano; MNRJ Afonso Claudio;
MNRJ –, Domingos Martins; MBML Pedra
Azul, Domingos Martins; MBML , , , RPPN
Oiutrem, Matilde, Alfredo Chaves; MBML Povoação de Bau-
nilha, Córrego Santinho, Colatina; MBML Crubixá-Mirim,
Santa Leopoldina; MBML Pousada Vila Suíça, Santa Leopol-
dina; MZUSP , , Rio Doce; MZUSP Sooretama.
Brazil: Estado de Goiás: MNRJ “Goyaz”.
Brazil: Estado de Minas Gerais: MNRJ –, MZUSP
, Fazenda Montes Claros, Reserva Ecológica de Carat-
inga, Caratinga; MNRJ , Sítio , Chiador; MNRJ
Sítio , Chiador; MNRJ – Mata dos Pena, Eugenópolis;
MNRJ Paula Cândido; MNRJ Sítio , Além Paraíba;
MNRJ Sítio , Além Paraíba; MNRJ Sítio , Além
Paraíba; MNRJ – Fazenda Cachoeirão, Além Paraíba;
MNRJ Além Paraíba; MNRJ UHE Guilman-Amorim,
Antônio Dias; MZUSP PCH Cachoeira Grande, Antônio
Dias; MZUSP – PCH Cocais Grande, Antônio Dias;
MNRJ Estação Biológica Mata do Sossego, Simonésia;
MNRJ , , Fazenda Olinda, São José, Belmiro Bra-
ga; MNRJ Água Limpa, Juiz de Fora; MNRJ Torrões,
Juiz de Fora; MNRJ Lagoa do Maximus, próximo a Santa
Rita de Ouro Preto, Ouro Preto; MNRJ , , ,
Viçosa; MNRJ Muriaé; MNRJ Reserva Biológica de
Mar de Espanha, Mar de Espanha; MNRJ Fazenda da Prata,
Sapucaia; MNRJ Mata de Gruta, Santana do Deserto; MNRJ
, Eugenópolis; MNRJ – Fazenda Cachoeira
Alegre, Abre Campo; MZUSP –, PARNA do
Caparaó; MZUSP , – UHE Fumaça, Mariana.
Brazil: Estado do Rio de Janeiro: MNRJ Paracambi;
MNRJ Teresópolis; MNRJ – Reserva Ecológica
de Guapiaçu, Cachoeiras de Macacu; MNRJ , , ,
Morro Azul, Engenheiro Paulo de Frontin; MNRJ –
Tinguá, Nova Iguaçu; MNRJ , –, –
Taquara, Duque de Caxias; MNRJ , , ,
Visconde de Mauá, Resende; MNRJ – Fazenda
Marimbondo, Resende; MNRJ – Fazenda Barreto,
Nova Friburgo; MNRJ – eodoro de Oliveira, Nova
Friburgo; MNRJ – Fazenda João Fernando, RJ ,
Valença; MNRJ Capela, Petrópolis; MNRJ ,
CEDEA, Rocio, Petrópolis; MNRJ , Petrópolis; MNRJ
k UTM , Sapucaia, Três Rios ou Mar de
Espanha; MNRJ – Fazenda Santa Bárbara, P.E. Três
Picos, Cachoeiras de Macacu; MNRJ –, Duas
Barras; MNRJ , Volta Redonda; MNRJ –
Vale da Revolta, Teresópolis; MNRJ , MZUSP –
Teresópolis; MNRJ PNM Fazenda Atalaia, Macaé; MNRJ
, , Comendador Levy Gasparian; MNRJ
Rialto, Barra Mansa; MNRJ Floriano, entrada do km ,
Via Dutra, Barra Mansa; MNRJ Patronato, Três Rios; MNRJ
Acampamento Batista Carioca, Areal, Três Rios; MNRJ
Área da PCH Santa Fé, Três Rios; MNRJ – Fa-
zenda Canaã, Três Rios; MNRJ Lídice, Rio Claro; MNRJ
Itaperuna; MNRJ Trajano de Moraes; MNRJ –
Paraty; MNRJ Fazenda Barra da Água Limpa, Bom
Jesus do Itabapoana, MZUSP – Itatiaia; MZUSP
Miguel Pereira.
Brazil: Estado de São Paulo: MNRJ Aparecida; MZUSP
– Parque das Neblinas, Bertioga; MZUSP –
Praia de Boracéia, Bertioga; MZUSP , –
Juquitiba; MZUSP – Parque Estadual de Carlos Bo-
telho, São Miguel Arcanjo; MZUSP São Miguel Arcanjo;
MZUSP , , Boracéia, Salesópolis; MZUSP
Boracéia; MZUSP E.E. de Bananal; MZUSP ,
Piedade; MZUSP , , , – Campo Grande
da Serra; MZUSP Fazenda Intervales; MZUSP Serra
da Bocaina; MZUSP Ferraz de Vasconcelos; MZUSP ,
Ribeirão Grande.
Hypsiboas pugnax
Venezuela: ZUEC Mantecal; MZUSP – Guárico
El Corozo Ranch, km N de Calabozo; MZUSP –
Guárico Espino.
Colombia: MZUSP – Meta: Villavicencio; MHUA
, , , Antioquia, Gomez Plata, Hacienda Vegas de
La Clara U de A; MHUA Córdoba, Ayapel, Vinícolas ICA;
MHUA Antioquia, Maceo, Hacienda Santa Barbara.
Trinidad: MZUSP Saint Benedict NR Saint Joseph
Northern Range.
Hypsiboas rosenbergi
Ecuador: Bulin: AL –.
Ecuador: Província de Esmeraldas: MZUSP San Javier;
MZUSP Cachavi; MZUSP – Cachavi.
Ecuador: Província de Pichincha: MZUSP – Centro
Cientíco Rio Palenque.
Panama: Província do Panamá: MZUSP —Canal Zone
mi N Miraores.
Hypsiboas xerophyllus
Brazil: Roraima: MNRJ –, –, –,
–, MZUSP , , Pacaraima; MZUSP
Serra da Saracura; MZUSP , , , ,
, Ilha de Maracá; MZUSP Marco de Fronteira
BV, Uiramutã; MZUSP , , , , , ,
Mucajaí; MZUSP , , Tepequem, Amajari;
MZUSP Fazenda Salvamento; MZUSP , , ,
, , , MPEG , , Caracaraí; MZUSP
, MPEG , , , , , Boa Vista; MZUSP
Bonm; MZUSP Uranduíque; MZUSP Fa-
zenda São Marcos; MZUSP , , , ,
Serra do Tepequem; INPA , , , , , ,
, , PARNA do Viruá; MPEG , , Rio
Ajarani, BR , Iracema.
Colombia: Meta: MZUSP , , Villavicencio;
AL-MN “Palomina” (Possibly Palomino, Colombia); MZUSP
Magdalena, Cañaveral, Parque Nacional Tayrona; MHUA–
A , Santander, Carmen de Chuqurí, Vereda La Bode-
ga; MHUA-A Antioquia, Puerto Berrio, Hacienda la Suiza;
MHUA-A , , , , Antioquia, Maceo, Vere-
da Las Brisas; MHUA-A Caldas, Norcasia, Hacienda El Val-
le; MHUA-A Antioquia, San Roque, Vereda La Providencia;
MHUA-A Antioquia, Nariño, Vereda Puente Linda; MHUA-
A Antioquia, San Francisco, Vereda La Lora; MHUA-A
Tolima, Falan, Vereda La Roca; MHUA-A Caldas, Pensilva-
nia, Vereda La Esperanza; MHUA-A – Antioquia, Nariño,
Vereda Puente Linda.
Panama: AL-MN San Pablo.
Trinidad: MZUSP Saint Joseph.
Venezuela: Guárico: MZUSP Calabozo; MZUSP
Espino; Sucre: MZUSP Playa Colorada; Tabay: MZUSP
Mérida; Aragua: AL-MN Maracay; ZUEC Santa
Bárbara; Táchira: ZUEC Uribante; ZUEC –, No
further locality.
IV
Online Supplementary data – V G. D. O et al.
Supplementary document 2. Voucher information, localities, and GenBank accession numbers for the specimens analyzed for this
study. [a] Voucher information for 12S ; [b] Voucher information for COI.
Sample ID Locality Voucher 12S COI
H. crepitans Grão Mogol, Minas Gerais, Brazil CFBH 10243 KX697938 KX697985
H. crepitans “Bahia”, Brazil CFBH 13275 KX697941 KX697988
H. crepitans Campo Alegre, Alagoas, Brazil CFBH 16361 KX697945 KX697992
H. crepitans Aurelino Leal, Bahia, Brazil CFBH 18728 KX697950 KX697997
H. crepitans Caetité, Bahia, Brazil CFBH 21073 KX697948 KX697995
H. crepitans Dom Basílio, Bahia, Brazil CFBH 21090 KX697949 KX697996
H. crepitans Jaboticatubas, Minas Gerais, Brazil CFBH 24368 KX697953 KX698000
H. crepitans Camamu, Bahia, Brazil CFBH 27826 KX697959 KX698006
H. crepitans Itabuna, Bahia, Brazil CFBH 2885 KX697939 KX697986
H. crepitans Piranhas, Alagoas, Brazil CFBH 2966 AY843621 –
H. crepitans Mucugê, Bahia, Brazil JC 1273 KX697967 KX698013
H. crepitans Jussari, Bahia, Brazil MRT 5785 KX697969 KX698015
H. crepitans Januária, Minas Gerais, Brazil MTJ 384 KX697971 KX698017
H. crepitans Buíque, Pernambuco, Brazil MTR 15366 KX697972 KX698018
H. crepitans Miguel Calmon, Bahia, Brazil MTR 20070 KX697976 KX698022
H. “xerophyllus” Haute Wanapi, French Guiana, France 552PG KX697932 KX697979
H. “xerophyllus” Boa Vista, Roraima, Brazil CFBH 29144 KX697961 KX698008
H. “xerophyllus” Mont Saint Marcel, French Guiana, France isolate BM095 KX697962 KX698011
H. “xerophyllus” E.E.Maracá, Roraima, Brazil MTR 20407 KX697977 KX698023
H. “xerophyllus” Pacaraima, Roraima, Brazil MTR 20730 KX697978 KX698024
H. “xerophyllus” Kurupukari, Guyana ROM 20560 JN970517 JN970772
H. “xerophyllus” Paramakatoi, Guyana ROM 28436 JN970518 JN970773
H. “xerophyllus” Parish_Hill, Guyana ROM 44089 JN970519 JN970774
H. “xerophyllus” “Igarapé Cocal”, Brazil USNM 302400 DQ380354 –
H. albomarginatus Ilhéus, Bahia, Brazil MTR 16695 KX697975 KX698021
H. albomarginatus_q [a] Caruaru, Pernambuco, Brazil;
[b] Juiz de Fora, Minas Gerais, Brazil
[a]USNM 284519; [b]CF-
BHT13378
AY549316 KX698009
H. cinerascens [a] Iwokrama, Guyana;
[b] Unknown
[a]AMNH A-164105; [b] JF
isolate 2371
AY549336 KX698010
H. ericae [a;b] Alto Paraiso de Goiás, Goiás, Brazil [a]CFBH 3599; [b]CFBH 6764 AY549332 KX698012
H. exastis Arataca, Bahia, Brazil MTR 16289 KX697973 KX698019
H. faber Uruçuca, Bahia, Brazil CFBH 13030 –KX697980
H. faber São Carlos, São Paulo, Brazil isolate 118 – JQ627303
H. faber San Vicente, Guarani, Argentina MACN 36999 AY549333 –
H. faber San Vicente, Guarani, Argentina MACN 37000 AY549334 –
H. faber Varzedo, Bahia, Brazil MTR 16691 KX697974 KX698020
H. faber Salesópolis, São Paulo, Brazil USNM 303034 DQ380356 –
H. lanciformis [a] Alpahuayo, Loreto, Peru;
[b] Shaime, Zamora-Chincipe, Ecuador
[a] MJH 564; [b] isolate 18237 AY843636 JN970768
H. lundii Assis, São Paulo, Brazil CFBH 20059 KX697944 KX697991
H. lundii Brasília, Distrito Federal, Brazil CFBH 22795 KX697956 KX698003
H. lundii Belo Horizonte, Minas Gerais, Brazil CFBH 22814 KX697957 KX698004
H. lundii Campo Limpo de Goiás, Goiás, Brazil CFBH 23593 KX697951 KX697998
H. lundii Lagoa Santa, Minas Gerais, Brazil CFBH 24355 KX697952 KX697999
H. lundii São Carlos, São Paulo, Brazil CFBH 26853 KX697954 KX698001
H. lundii Rio Claro, São Paulo, Brazil CFBH 4000 AY843639 –
H. lundii Costa Rica, Mato Grosso do Sul, Brazil CFBH 7280 KX697934 –
H. lundii São Roque de Minas, Minas Gerais, Brazil CFBH 9154* KX697936 KX697983
H. lundii Alto Paraíso, Goiás, Brazil CFBH 9324 KX697937 KX697984
V
Online Supplementary data – Taxonomy of Hypsiboas crepitans
Sample ID Locality Voucher 12S COI
H. lundii Paranã, Tocantins, Brazil MRT 4332 KX697968 KX698014
H. lundii Petrolina de Goiás, Goiás, Brazil MRT 8417 KX697970 KX698016
H. multifasciatus Demerara, Guyana AMNH A-141040 AY843648 AY843648
H. pardalis Domingos Martins, Espírito Santo, Brazil CFBH 11334 KX697940 KX697987
H. pardalis Petrópolis, Rio de Janeiro, Brazil CFBH 13962 KX697942 KX697989
H. pardalis São Luis do Paraitinga, São Paulo, Brazil CFBH 16020 KX697935 KX697982
H. pardalis Maringá, Rio de Janeiro, Brazil CFBH 17542 KX697946 KX697993
H. pardalis Santa Leopoldina, Espírito Santo, Brazil CFBH 18002 KX697947 KX697994
H. pardalis Teresópolis, Rio de Janeiro, Brazil CFBH 22027 KX697955 KX698002
H. pardalis Mimoso do Sul, Espírito Santo, Brazil CFBH 25503 KX697958 KX698005
H. pardalis Cachoeiras de Macacu, Rio de Janeiro, Brazil CFBH 27405 KX697960 KX698007
H. pardalis São Luis do Paraitinga, São Paulo, Brazil CFBH 6499 KX697933 KX697981
H. pardalis Itatiaia, Rio de Janeiro, Brazil CFBHT4595 KX697943 KX697990
H. pardalis Salesópolis, São Paulo, Brazil USNM 303046 AY843651 –
H. pellucens San Juan, Pichincha, Ecuador WED 53621 AY326058 AY326058
H. pugnax Ayapel, Córdoba, Colombia MRC 510 KX697963 –
H. pugnax Gómez Plata, Antioquia, Colombia MRC 513 KX697964 –
H. pugnax Norcasia, Caldas, Colombia MRC 514 KX697965 –
H. pulchellus Buenos Aires, Buenos Aires, Argentina MACN 37788 AY549352 –
H. punctatus Resistencia, Chaco, Argentina MACN 37792 AY549353 –
H. rosenbergi E Pedernales, Manabi, Ecuador KU 217629 AY819438 –
H. rosenbergi Maceo, Antioquia, Colombia MRC 508 KX697966 –
H. rutelus [a;b] El Cope, Cocle, Panama [a] KRL 798; [b] USNM 572699 AY843662 FJ766740
Supplementary document 3. Standardized coecients and factor loadings (r) from a Principal Component Analysis (CDA; Fig.3)
for 16 morphometric characters of two Operational Taxonomic Units (OTUs) of Hypsiboas crepitans. Cum. Prop. = cumulative
proportion of eigenvalues.
Measurements Males Females
PC1 PC2 r (PC1) r(PC2) PC1 PC2 r (PC1) r(PC2)
SVL -0.051259 -0.019337 -0.903869 -0.340971 -0.056931 0.001937 -0.975977 0.033204
HL -0.049143 -0.017838 -0.871643 -0.316396 -0.049324 0.002171 -0.946597 0.041662
HW -0.049520 -0.017450 -0.910738 -0.320932 -0.052074 -0.002213 -0.967214 -0.041107
NSD 0.033115 -0.092269 0.312513 -0.870765 -0.032079 0.042403 -0.470261 0.621610
IND -0.055426 0.017232 -0.625668 0.194523 -0.037001 0.002607 -0.738501 0.052041
UEW -0.034914 -0.040303 -0.494884 -0.571283 -0.047864 0.019396 -0.718680 0.291226
ED -0.043462 -0.020399 -0.777362 -0.364856 -0.043967 -0.003470 -0.891364 -0.070353
IOD -0.142854 0.058591 -0.896194 0.367574 -0.057721 -0.092086 -0.522806 -0.834058
END -0.032284 -0.022338 -0.586094 -0.405529 -0.046775 0.006113 -0.882090 0.115281
TD -0.027915 -0.031178 -0.513080 -0.573040 -0.054854 0.010522 -0.835374 0.160242
FHL -0.052215 -0.018928 -0.855917 -0.310264 -0.054793 -0.003064 -0.919238 -0.051396
4FD -0.041548 -0.029878 -0.693932 -0.499026 -0.053960 0.018977 -0.835099 0.293686
4TD -0.039945 -0.031080 -0.648998 -0.504972 -0.052089 0.000047 -0.946296 0.000861
THL -0.050017 -0.015955 -0.907767 -0.289567 -0.051758 0.000801 -0.943330 0.014607
CL -0.047836 -0.015795 -0.891682 -0.294418 -0.050472 -0.000944 -0.946317 -0.017696
FL -0.050864 -0.014523 -0.911669 -0.260300 -0.048540 0.026065 -0.758781 0.407444
Eigenvalue 0.050425 0.019683 0.039746 0.011885
Cum. Prop 57.2663 79.6201 63.0218 81.8662
