ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by M. Heinicke: 1 Mar. 2022; published: 28 Mar. 2022 345
Zootaxa 5120 (3): 345–372
Copyright © 2022 Magnolia Press Article
Two new species of geckos of the genus Phyllopezus Peters, 1878
(Squamata: Gekkota: Phyllodactylidae) from northeastern Brazil
MARCOS J. M. DUBEUX1,2,3,*, UBIRATAN GONÇALVES2,3, CRISTIANE N. S. PALMEIRA2,4,
PEDRO M. S. NUNES1, JOSÉ CASSIMIRO5, TONY GAMBLE6,7,8, FERNANDA P. WERNECK9,
MIGUEL T. RODRIGUES5 & TAMÍ MOTT2,3
1Programa de Pós-Graduação em Biologia Animal, Departamento de Zoologia, Centro de Biociências, Universidade Federal de
Pernambuco, Avenida Professor Moraes Rego, 1235, Cidade Universitária, 50670-901, Recife, Pernambuco, Brasil.
2Setor de Herpetologia, Museu de História Natural, Universidade Federal de Alagoas, Avenida Amazonas, S/N, Prado, 57010-060,
Maceió, Alagoas, Brasil.
3Setor de Biodiversidade, Instituto de Ciências Biológicas e da Saúde, Universidade Federal de Alagoas, Av. Lourival Melo Mota, S/N,
Tabuleiro do Martins, 57072-900, Maceió, Alagoas, Brasil.
4Programa de Pós-Graduação em Desenvolvimento e Meio Ambiente, Departamento de Botânica e Zoologia, Centro de Biociências,
Universidade Federal do Rio Grande do Norte, Campus Universitário, Lagoa Nova, 59078-900, Natal, Rio Grande do Norte, Brasil.
5Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, Travessa 14, 05508-090, São Paulo,
São Paulo, Brasil.
6Department of Biological Sciences, Marquette University, Milwaukee, WI 53233, USA.
7Milwaukee Public Museum, Milwaukee, WI 53233, USA.
8Bell Museum of Natural History, University of Minnesota, St Paul, MN 55113, USA.
9Programa de Coleções Científicas Biológicas, Coordenação de Biodiversidade, Instituto Nacional de Pesquisas da Amazônia, Av.
André Araújo, 2936, Petrópolis, 69067-375, Manaus, Amazonas, Brasil.
We describe two new species of Brazilian geckos of the genus Phyllopezus based on morphological and molecular data.
The first species is currently known from a relictual Cerrado enclave—“campos rupestres”, in the mountains of the Serra
do Espinhaço in the state of Bahia. The second species is known from northeastern Atlantic Forest and transitional areas
with Caatinga biome in the state of Alagoas. The two new species are sister taxa and together are the sister clade to the
remaining species in the Phyllopezus pollicaris species complex. These new species can be morphologically distinguished
from their congeners by meristic and morphometric characters, in addition to color pattern and genetic differentiation.
Key words: Cryptic diversity; integrative taxonomy; lizard; morphological data; molecular phylogeny
The genus Phyllopezus Peters, 1878 includes saxicolous and arboreal lizards distributed predominantly in the dry-
open biomes of South America (Caatinga, Cerrado, and Chaco; Werneck et al. 2012; Cacciali et al. 2018). The genus
is morphologically diagnosed by skin with small granular scales with equidistant larger tubercles, tail with small
rhomboid scales, a single series of undivided lamellae under the base of the digits and, by the two distal phalanges
in all digits, narrowing towards the claw (Vanzolini 1953; Vanzolini et al. 1980; Rodrigues 1986). The monophyly
of the genus is supported by molecular data, including mitochondrial (16S rRNA, Cytb and ND2) and nuclear (e.g.,
RAG1, RAG2, C-MOS and ACM4) genes (Gamble et al. 2012; Werneck et al. 2012; Cacciali et al. 2018).
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Currently Phyllopezus is composed of six formally described species: P. heuteri Cacciali, Lotzkat, Gamble &
Köhler, 2018 (restricted to the Chaco biome in the Cordillera de Los Altos mountain range, Paraguay; Cacciali et al.
2018), P. lutzae (Loveridge, 1941) (restricted to the Atlantic Forest biome of northeastern Brazil, with distribution
extending from the state of Paraíba to Bahia; Albuquerque et al. 2019), P. maranjonensis Koch, Venegas & Böhme,
2006 (distributed in the dry forests of the upper Marañon basin, Peru; Koch et al. 2006), P. periosus Rodrigues,
1986 (distributed in the northern Caatinga biome of states of Ceará to Pernambuco), P. przewalskii Koslowsky, 1895
(distributed in the Chaco biome of Paraguay, Bolivia and north of Argentina and in the Cerrado biome, below the
Central Brazilian Shield, in the states of Mato Grosso and Mato Grosso do Sul (Condez et al. 2021), and P. pollicaris
(Spix, 1825) (widely distributed in the Caatinga and Cerrado biomes and entering the Atlantic Forest in the eastern
coast of Brazil; Condez et al. 2021), which includes an extremely high hidden diversity (e.g. Pellegrino et al. 1997;
Gamble et al. 2012) resulting in multiple evolutionary lineages (Gamble et al. 2012; Werneck et al. 2012) that still
await formal descriptions.
Werneck et al. (2012), using molecular data, recovered eight lineages associated to P. pollicaris sensu lato in
the Caatinga, Cerrado, and Chaco biomes (Clades I–VIII), and treated them as candidate species. In that work, the
authors resurrected the nominal taxon P. przewalskii to accommodate populations belonging to Clade V, rendering
P. pollicaris paraphyletic. Later, Cacciali et al. (2018) described a new lineage (P. heuteri) related to P. przewalskii,
which had not been included in previous studies. Combined with high genetic diversity (Gamble et al. 2012; Werneck
et al. 2012) and the presence of multiple reciprocally monophyletic groups there is a clear need of taxonomic studies
to delimit and describe the cryptic diversity within the genus.
Herein we use an integrative approach, including molecular data and external morphology to describe two new
species of Phyllopezus (currently assigned to the P. pollicaris complex) from northeastern Brazil. We also define the
P. pollicaris complex as the clade comprised of P. pollicaris sensu lato, P. przewalskii, and P. heuteri.
Material and methods
Analyzed specimens. We examined 146 specimens of Phyllopezus including representatives of all nominal taxa
occurring in the Northeast region of Brazil (P. lutzae, P. periosus, and P. pollicaris; Fig. 1; Appendix I). All P.
pollicaris specimens analyzed for external morphology come from localities represented in the molecular datasets
used in recent phylogenies (i.e., Gamble et al. 2012; Werneck et al. 2012). Specimens are deposited in the following
herpetological collections: Coleção Herpetológica do Museu de História Natural da Universidade Federal de Alagoas
(MHN-UFAL), Coleção Herpetológica da Universidade Federal da Paraíba (CHUFPB), Coleção Herpetológica da
Universidade Federal de Pernambuco (CHUFPE), Coleção Herpetológica da Universidade Federal do Rio Grande
do Norte (UFRN) and Coleção Herpetológica do Museu de Zoologia da Universidade de São Paulo (MZUSP).
Paralectotypes of Phyllopezus pollicaris (ZSM 165/0/1–2) were examined through photographs (available in
Cacciali et al. ).
Phylogenetic inference and genetic distance analyses. Phyllopezus pollicaris Clade I [sensu Werneck et al.
(2012); specimens from municipality of Mucugê, state of Bahia, hereafter treated as Phyllopezus sp.1] have been
previously considered as a candidate species (Gamble et al. 2012; Werneck et al. 2012; Cacciali et al. 2018). When
analyzing the morphology of this candidate species, we noticed its similarity with some specimens occurring in the
state of Alagoas, which although morphologically different from P. pollicaris sensu stricto, had never been included
in a molecular study. We complemented the existing dataset of partial 16S rRNA sequences with representatives of
three different locations for this population of Alagoas state [hereafter treated as Phyllopezus sp.2; see Appendix II
for GenBank accession numbers, vouchers, and localities; note that representatives of P. pollicaris Clades II and III
of Werneck et al. (2012) were not included due to the lack of 16S rRNA sequences].
DNA was extracted using phenol/chloroform protocol (Sambrook & Russel 2001). Subsequently, a fragment
of the 16S rRNA mitochondrial gene was amplified through polymerase chain reaction (PCR), using the primers
16Sar-L and 16Sbr-H (Palumbi et al. 2002). Reactions consisted of 25 µl solution containing 12.5 µl of Master Mix
PCR Buffer Promega® with 0.4 mM of each dNTP and 3 mM of MgCl2, 8.4 µl of nuclease-free water, 0.5 µl of Taq
DNA polymerase Invitrogen® (5U/µl), 0.8 µl of each primer (10pmol) plus 2 µl of DNA template (20–100 ng/µl).
Samples were amplified with: (1) initial denaturation at 94 °C for 90 sec followed by 35 cycles of denaturation at 94
°C for 45 sec, (2) annealing at 48 °C for 60 sec, and (3) extending at 72 °C for 60 sec. Samples were purified using
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isopropanol and sent to be unidirectional sequenced using Sanger method at the Laboratório Central da Universidade
Federal de Pernambuco (LABCEN), municipality of Recife, Pernambuco state, Brazil.
Sequences were aligned with other 45 sequences representative of all Phyllopezus clades (including two
sequences of Phyllopezus sp.1; Appendix II) available on GenBank using MAFFT software v.7.310 with default
parameters (Katoh & Standley 2013). Genetic distances within and between species and among clades recognized
by Werneck et al. (2012) were then estimated using Kimura 2-parameters evolutionary models (K2P; Kimura 1980)
and p-distance (p-D) with complete deletion of gaps, implemented in software MEGA X (Kumar et al. 2018).
The choice of the best evolutionary model for the character matrix was performed in PartitionFinder v.2.1.1
(Lanfear et al. 2012) using the Bayesian Information Criterion (BIC) and default settings. Bayesian analysis was
performed in MrBayes software v.3.2 (Ronquist et al. 2012). The phylogenetic analysis consisted of two independent
runs of 10 million generations each, being evaluated every 1,000 generations. The first 25% of trees were discarded
as burn-in; the majority consensus tree was visualized using the program FigTree software v.1.3.1. Nodes with
posterior probability above 0.95 were considered having high support. Phyllodactylus xanti Cope, 1863 was used
for rooting the tree following Gamble et al. (2012).
FIGURE 1. Geographic distribution of genetic and morphological vouchers of Phyllopezus pollicaris complex in northeastern
Brazil. (A) Phyllopezus pollicaris clades (following Werneck et al. ) and new samples analyzed in this study; (B) expanded
view of the known distribution of Phyllopezus sp.2 in the state of Alagoas, Brazil; (C) expanded view of the known distribution
of Phyllopezus sp.1 in the state of Bahia, Brazil. Numbers near the points correspond to the municipalities of (1) Boca da Mata,
(2) Quebrangulo, (3) Igaci, (4) Limoeiro de Anadia and (5) Coruripe, state of Alagoas, and (6) Mucugê, state of Bahia, Brazil.
Stars indicate type localities (with available morphological and molecular data) of Phyllopezus sp.1 (purple) and Phyllopezus
sp.2 (green). Locations indicated with diamond denote samples of Werneck et al. (2012) not included in this study due to the
absence of 16S rRNA data. Transparent white area [gray in the inset South America map in (A)] corresponds to Brazilian dry-
open biomes. Inset map: South America.
Species delimitation analyses. Four different methods of single locus species delimitation were performed
using the aligned 16S rRNA data (excluding external groups). (1) We calculated a cutoff point for probable
intraspecific divergence, specifically for our data set, using the Local Minima function of the Species Identity
and Evolution in R package (SPIDER; Brown et al. 2012) implemented in the R Studio software v.1.2.1 (R Core
Team 2020). This analysis is based on the concept of a barcode gap, and species identification is not provided a
priori. When the analysis detects a drop in the density of genetic distances, a possible transition between intra and
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interspecific distance values is suggested. (2) We also performed an analysis of Automatic Barcode Gap Discovery
(ABGD; Puillandre et al. 2011) implemented on the ABGD online platform (https://bioinfo.mnhn.fr/abi/public/
abgd/), which is based on the general genetic divergences between sequences of the matrix, using values from a
series of previous intraspecific divergences. For this analysis, sites with gaps were excluded, resulting in a matrix
of 347 base pairs. Previous values for minimum and maximum intraspecific divergence were defined based on the
results of Local Minima, and we also tested for values of interspecific divergence found between species already
recognized in the genus (Gamble et al. 2012; Cacciali et al. 2018), and among gekkotan sister species (Rocha et al.
2009; Fujita et al. 2010). Values between 0.001 and 1.5 of relative gap were tested. (3) To delimit species based on
the topology recovered, an analysis of Bayesian implementation of the Poisson Tree Process (bPTP; Zhang et al.
2013) was performed. This model uses a rooted phylogenetic tree to model speciation or branching events in terms
of the number of substitutions. The analysis was performed on the PTP web server (https://species.h-its.org/ptp/)
using default settings and clusters with support above 0.95 were indicated in the tree. (4) Finally, the Bayesian
implementation of the Generalized Mixed Yule Coalescent model (bGMYC; Pons et al. 2006) was conducted. This
model uses different topologies to model and to identify likely transition points between coalescence events and
cladogenesis of alleles, incorporating phylogenetic uncertainties through a Bayesian extension (Reid & Carstens
2012). The topologies were obtained from a random selection of 100 trees resulting from a Bayesian inference
implemented in the BEAST software v.1.8.4 (Suchard et al. 2018) using the GTR+Г+I model (Suchard et al. 2018)
in three independent analyses of 20 million generations each (sampling every 2 x 103) and 20% of initial burn-in.
The bGMYC analysis was performed in the R Studio software for 100,000 generations with a burn-in of 90,000
and a dilution interval of 100 samples. A cut-off point for the probability of recognition of genetic clusters of 50%
was adopted. To maximize the confidence of inferences (based on a single marker), we focus on species/lineages
congruently retrieved/delimited by all methods as they can potentially be independent, but complementary.
External morphology. Measurements and scale counts were taken under a stereomicroscope. Measurements
follow Rodrigues (1986), Cassimiro & Rodrigues (2009), and Sturaro et al. (2018), and were taken using a digital
caliper to the nearest 0.1 mm (in right side of specimens whenever possible): snout-vent length (SVL, from tip of
snout to cloacal opening), distance between limbs (DBL, from axilla to groin), tail base width (TBW, taken at the
base of the tail just posterior to the cloaca), head length (HL, from tip of snout to anterior margin of ear-opening),
head width (HW, on the widest part of head), head depth (HD, on the highest part of head), snout length (SL, from
tip of snout to anterior margin of eye), nares-snout distance (NSD, from tip of snout to anterior border of nares),
nares-eye distance (NED, to posterior border of nares to anterior edge of eye), eye-snout distance (ESD, from tip
of snout to center of eye), eye diameter (ED, in widest section of the eye), interorbital distance (IOD, between the
upper margins of eyes), internarial distance (IND, between the upper margins of nares), length of humerus (LH,
from insertion of humerus to elbow), length of forearm (LF, from tip of elbow to wrist), length of thigh (LT, from
insertion of femur to knee), length of tibia (LTB, from knee to ankle), width of mental scale (WM, between lateral
corners), length of mental scale (LM, between tip of the scale and posterior border with postmentals), width of
rostral scale (WR, between lateral corners), and length of rostral scale (LR, between anterior tip of the snout and
medial contact with nasals).
Scale counts and terminology follow Rodrigues (1986), and Cassimiro & Rodrigues (2009), and were counted
as follows: number of rostrals (R), number of postrostrals (PR), number of postnasals (PN), number of supralabials
(SL), number of infralabials (IL), number of mentals (M), number of postmentals (PM), scales that surround the
postmentals (SSP), number of ventrals in a longitudinal row (VLR, along a midventral line, from anterior margin
of forelimbs to anterior margin of hind limbs), number of dorsal tubercles in a longitudinal row (DT, along a mid-
dorsal line, from anterior margin of forelimbs to tail), number of lamellae under the fourth finger (L4F), number of
lamellae under the fourth toe (L4T), number of postcloacal tubercles at the sides of the vent (TP), and number of
postcloacal pores (CP).
Sex was identified by direct inspection of gonads (when dissected), by the presence of eggs (females) or
hemipenis (males, when everted) or by a small lateral insertion at the base of the tail when not everted (presence of
the hemipenis or the hemipenial retractor muscle).
To identify sexual dimorphism in SVL, univariate analysis of variance (ANOVA) was performed, and for
variation between sexes in the other morphometric characters we used multivariate analysis of variance (MANOVA).
The interspecific variation was summarized with Principal Component Analysis (PCA) and Discriminant Function
Analysis (DFA) for meristic and morphometric characters separately. In addition, these analyses were performed
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for the complete dataset (including all analyzed species) and for a dataset containing only the representatives of P.
pollicaris complex. All analyses were performed in the R Studio software.
For color descriptions, we used the terminology proposed by Köhler (2012) with their corresponding color
codes. For P. heuteri, P. maranjonensis and P. przewalskii morphological data for comparisons were obtained from
literature and/or photographs (Koslowsky 1895; Koch et al. 2006; Cacciali et al. 2018).
Phylogenetic inference and genetic distance analyses. The dataset included three new 16S rRNA sequences
generated in this study (486 pb) and 45 obtained from GenBank. The best evolutionary model for the dataset was
GTR + I + G. The Bayesian analysis recovered monophyly for the P. pollicaris complex (posterior probability
PP = 0.99; Fig. 2). The two individuals from municipality of Mucugê, Bahia state (Phyllopezus sp.1), present
identical haplotypes and were recovered as the sister clade of individuals from Alagoas state (Phyllopezus sp.2),
with strong support (PP = 1.0). The three individuals of Phyllopezus sp.2 from three different localities in Alagoas
state (geographic distance between collecting sites from 33 to 76 km) also present identical haplotypes. The genetic
divergence between Phyllopezus sp.1 and Phyllopezus sp.2 (GD Phyllopezus sp.1 × Phyllopezus sp.2 = K2P 0.093, p-
D 0.087) was similar to that found among other recognized species of the genus (e.g., GD P. heuteri × P. przewalskii
= K2P 0.096, p-D 0.089; see Table 1).
This clade (Phyllopezus sp.1 + Phyllopezus sp.2, hereafter called as “Group 1” due to the sharing of morphological
characteristics, see section Morphological approach) was recovered as sister of the remaining lineages belonging
to the P. pollicaris complex. The Group 1 corresponds to the sister lineage (PP = 0.99) of the clade composed by
P. heuteri, P. przewalskii and P. pollicaris Clades IV, VI–VIII (hereafter treated as “Group 2”; PP = 0.98). High
intraspecific genetic divergences in two methods were observed in P. pollicaris clades IV and VII (GD = 0.047 and
Species delimitation analyses. The ABGD analysis indicated the most likely transition point between inter
and intraspecific genetic distances around 3–7% (Fig. 3B). The Local Minima function identified a first depression
in the distance density curve at 5.73% (N = 1176, Bandwidth = 0.00975). The ABGD analyses did not show great
variation between the numbers of delimited genetic clusters and varied from 15 clusters in relative widths of the
range less than 1.0 to 16 clusters in relative widths of the range between 1.0 and 1.5 (Fig. 3). The analysis of bPTP
and bGMYC recovered 20 genetic clusters, although in bPTP the subdivisions in the clades of P. przewalskii and
P. pollicaris Clade VIII showed low support (less than 95%). Although the cut-off point of 50% was used for
the probability of recognition of genetic clusters in bGMYC analysis, it was observed that even adopting a less
conservative cut-off point of 90%, only one more genetic cluster was identified. In all analyses Phyllopezus sp.1 and
Phyllopezus sp.2 were recovered as distinct genetic clusters with high statistical support (> 95%).
External morphology. Individuals of Phyllopezus sp.1 had significant differences in SVL between sexes
(ANOVA: N = 6 ♂, 4 ♀; F1.9 = 5.43, P = 0.04). Meanwhile, Phyllopezus sp.2 do not show sexual dimorphism in
relation to the SVL (ANOVA: N = 9 ♂, 12 ♀; F1.19 = 0.15, P = 0.70). However, given the small sample size these
observations are not statistically robust. There was also no variation between sexes considering all other morphometric
characteristics: Phyllopezus sp.1 (MANOVA: F1.19 = 2.381, P = 0.105) and Phyllopezus sp.2 (MANOVA: F1.8 =
0.179, P = 0.906). Morphometric values are available in Appendix III.
When all species were analyzed together (Fig. 4A–D), P. periosus and P. lutzae were recovered in morphogroups
reciprocally segregated in the PCAs and DFAs in all datasets used, with no overlap with the other morphotypes
(exceptionally in some P. periosus outliers in the PCA for meristic data; Fig. 4A). In relation to the representatives
of the P. pollicaris complex (Fig. 4C–D), PCAs recovered morphogroups with low overlap, mainly in relation to the
representatives of P. pollicaris sensu lato.
For analyses performed only with representatives of the P. pollicaris complex (Fig. 4E–H), morphogroups with
almost no overlap in the morpho-space were recovered. Only in PCA using meristic characters Phyllopezus sp.1 and
Phyllopezus sp.2 showed some degree of overlapping (Fig. 4E).
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TABLE 1. Genetic distances between and within species and clades of Phyllopezus estimated using Kimura 2-parameters evolutionary model (K2P = bottom right diagonal) and
p-distance method (p-D = top left diagonal) with complete deletion of gaps in the 16S rRNA mitochondrial gene fragment. The lowest values of genetic distances between species
are highlighted in bold. *Intraspecific genetic distances higher than the cutoff point indicated by the Local Minima analysis. Phyllopezus pollicaris Clades follow Werneck et al.
Species/Clade 1 2 3 4 5 6 7 8 9 10 11 Intraspecific
1. P. periosus - 0.156 0.169 0.182 0.183 0.181 0.185 0.203 0.187 0.173 0.168 - -
2. P. lutzae 0.177 - 0.135 0.173 0.165 0.176 0.161 0.182 0.154 0.145 0.150 0 0
3. P. maranjonensis 0.193 0.150 - 0.181 0.164 0.157 0.170 0.192 0.167 0.156 0.171 0.002 0.002
4. P. heuteri 0.211 0.199 0.208 - 0.089 0.131 0.129 0.155 0.130 0.139 0.127 0.000 0.000
5. P. przewalskii 0.213 0.189 0.186 0.096 - 0.112 0.106 0.139 0.110 0.126 0.123 0.023 0.022
6. P. pollicaris IV 0.209 0.204 0.177 0.147 0.123 - 0.128 0.155 0.123 0.142 0.148 0.061* 0.057
7. P. pollicaris VI 0.214 0.183 0.193 0.144 0.115 0.142 - 0.125 0.097 0.154 0.145 0.004 0.004
8. P. pollicaris VII 0.240 0.212 0.224 0.178 0.156 0.177 0.139 - 0.094 0.161 0.154 0.050 0.047
9. P. pollicaris VIII 0.217 0.174 0.189 0.145 0.120 0.136 0.105 0.102 - 0.145 0.143 0.021 0.021
10. Phyllopezus sp.1 0.198 0.162 0.175 0.155 0.140 0.159 0.174 0.185 0.162 - 0.087 0 0
11. Phyllopezus sp.2 0.190 0.169 0.194 0.141 0.135 0.168 0.162 0.175 0.160 0.093 - 0 0
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FIGURE 2. Phylogram obtained by the Bayesian Inference of the16S rRNA mitochondrial gene fragment (486 pb) of genus
Phyllopezus. Posterior probabilities greater than 0.95 are indicated on the nodes. Phyllopezus pollicaris Clades I and IV–VIII
follow the definitions by Werneck et al. (2012). No representatives of the Clades II and III were included due to the lack of 16S
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FIGURE 3. (A) Summary results of Phyllopezus species delimitations based on 16S rRNA mitochondrial gene fragment using
ABGD, bPTP and bGMYC models, respectively. Total genetic breaks found in each model. Abbreviations above nodes: lut. =
P. lutzae, mar. = P. maranjonensis, per. = P. periosus, sp.1 = Phyllopezus sp.1, sp.2 = Phyllopezus sp.2, pol. VI = P. pollicaris
VI, heu. = P. heuteri, prz. = P. przewalskii, pol. VI = P. pollicaris VI, pol. VII = P. pollicaris VII, and pol. VIII = P. pollicaris
VIII. (B) Pair-to-pair genetic divergence densities indicating the most likely transition gap between intraspecific (left) and
interspecific (right) genetic distances of Phyllopezus sequences using 347 bp of 16 SrRNA mitochondrial gene fragment.
Phyllopezus pollicaris Clades numbering follow Werneck et al. (2012).
Taxonomic implications. Two main clades are recovered in the Phyllopezus pollicaris complex (Group 1 and
2) showing high genetic divergence between them (GD = K2P 0.159, p-D 0.142), similar divergence found between
more external specific lineages of the genus (e.g., P. periosus, P. lutzae, P maranjonensis; GD = 0.165–0.195).
Morphologically, the lineages belonging to Group 1 can be distinguished from those belonging to Group 2 (in
parentheses) by the presence of postmental scales hexagonal-shaped, twice as long as wide (heptagonal-shaped
with similar width and length); larger dorsal tubercles, corresponding to about six granules, elongated and generally
slightly keeled (smaller dorsal tubercles, corresponding to about four granules, subcircular or elliptical); first rows
of reduced scales that surround the enlarged scales of the postmental region do not extend beyond the posterior
margin of the third infralabial (always extending beyond the posterior margin of the third infralabial); and their
larger size: SVL 88.5 ± 7.25 / 74.4–100.2 (SVL 71.7 ± 5.3 / 59.4–87.1; ANOVA: F1,117 = 1.833, P = 0.031). For more
details on the variations see the “Comparison with Congeners” section.
Based on the external morphology and genetic data, the two lineages belonging to Group 1 of the Phyllopezus
pollicaris complex are described herein as new species.
Phyllopezus diamantino sp. nov.
(Figs. 5, 6, 9G, 10A)
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FIGURE 4. Principal Component Analysis (PCA; left) and Discriminant Function Analyzes (DFA; right) based on 13 meristic
and 20 morphometric characteristics independently for (A–D) all lineages of Phyllopezus and (E–H) only the lineages of P.
pollicaris complex occurring in northeastern Brazil.
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Phyllopezus pollicaris: Cassimiro & Rodrigues (2009); Freitas et al. (2012)
Phyllopezus pollicaris Clade A: Gamble et al. (2012)
Phyllopezus pollicaris Clade I: Werneck et al. (2012); Cacciali et al. (2018)
Phyllopezus sp.1 (aff. pollicaris): Dubeux et al. (present study)
Holotype. MZUSP 106770 (adult female) from Serra do Sincorá, Chapada Diamantina [12°59’34”S, 41°20’29”W;
935 m above sea level (a.s.l.)], municipality of Mucugê (Fig. 1C), Bahia state, Brazil, collected on 15 March 2005
by J. Cassimiro and F.S.F. Leite, field number JC 1234.
Paratypes. MZUSP 106771 (adult female; 13°00’03”S, 41°21’58”W; 999 m a.s.l.), MZUSP 106772 (adult
female; 13°01’26”S, 41°21’53”W; 980 m a.s.l.), MZUSP 106773 (adult female; 13°00’19”S, 41°21’47”W; 1010
m a.s.l.), MZUSP 106778 and MZUSP 106774 (adult male and adult female, respectively; 13°00’16”S, 41°21’
47”W; 1007 m a.s.l.), MZUSP 106775 (adult female; no coordinates), MZUSP 106776 (adult male; 13°01’08”S,
41°21’56 W; 1018 m a.s.l.), MZUSP 106778 (adult male; 13°00’18”S, 41°21’47”W; 1000 m a.s.l.), MZUSP 106779
(adult male; 13°01’02”S, 41°20’39”W; 945 m a.s.l.), MZUSP 106781 and MZUSP 106780 (juvenile unsexed and
adult male, respectively; 13°00’05”S, 41°21’57”W; 983 m a.s.l.), MZUSP 106782 (juvenile unsexed; 13°00’03”S,
41°21’58”W; 999 m a.s.l.). All paratypes are topotypes and were collected between 3–17 March 2005, by J.
Cassimiro, F.S.F. Leite and L.E. Lopes.
Etymology. The specific epithet “diamantino” is a latinized adjective referring to its type-locality, Parque
Nacional da Chapada Diamantina, the northern segment of the Cadeia do Espinhaço in the state of Bahia, Brazil.
Diagnosis. Phyllopezus diamantino sp. nov. is characterized by the following combination of character states:
(1) Mental scales sub-triangular, with similar length and width and posterior margin not exceeding the second
infralabial; (2) postmental scales increased, hexagonal, twice as long as wide, with broad contact each other and
previously separated by about 1/3 of its length by the mental scale; (3) up to two scales in contact with the ventral
margin of first infralabial; (4) presence of enlarged scales surrounding and separating postmental scales from the
granules of the gular region; (5) six to seven infralabial scales; (6) granular scales in the distal region of mandible,
juxtaposed, occasionally presenting tubercles of different sizes; (7) dorsal tubercles enlarged, corresponding to
about six granular scales, elongated and keeled; (8) developed pollex; (9) cycloid or triangular scales around the
auditory meatus, little bristly; (10) homogeneous scales of the same size in the region of the labial commissure;
(11) many tubercles in the angular region between the upper and lower edges of the opening of the auditory meatus
and eyes; (12) postcloacal pores always present in males and females; and (13) large sized, SVL 76.41–96.25 mm
in males, and 72.38–82.36 mm in females. See “Comparison with Congeners” section for additional diagnosis with
other genus species.
Description of holotype. Adult female, SVL 96.25, fully regenerated tail, DBL 37.97, TBW 10.12, HL 27.75,
HW 19.17, HD 8.73, SL 11.01, NSD 2.42, ESD 8.82, ED 6.94, IOD 8.87, IND 3.27, LH 21.04, LF 13.13, LT 22.02,
LTB 15.41, WM 4.41, LM 4.28, WR 4.05, LR 2.16, R 1, PR 3, SN 2, SL 7, IL 7, M 1, PM 2, SSP 7, VLR 59, DT
45, L4F 10, L4T 14, TP 3, and CP 1. Head large (SVL/HL = 3.43), distinct from neck. Mental large (HW/WM =
4.3 and HL/LM = 6.4), sub-triangular, slightly wide than longer (WM/LM = 1.03), bordered by the 1st infralabial
and in broad contact with two postmentals. A pair of postmentals, large, hexagonal-shaped, juxtaposed, longer than
wide, separated for mental by one third of their length, flanked by seven large scales with differentiated sizes, which
are replaced by granules juxtaposed that extend to the level of the labial commissure and are gradually replaced by
similar small scales, smooth and imbricate, similar to ventral scales. First five infralabials rhomboid; the first largest,
in broad contact with the postmental pair and a group of seven large, smooth, variable-shaped scales that isolate the
postmentals from the granules of the gular region. These are succeeded by small scales that undergo abrupt reduction
in size until become granules in the beginning of the gular region. First infralabial smaller than 2nd, and from the 2nd
on decreasing in size towards the labial commissure; the commissure area with granules. First to 4th infralabial scales
rhomboid. From the 2nd infralabial on, there is a group of small, elongated scales that border the infralabial row to near
of the labial commissure, which also isolates them from the granules of the gular region. Ventral scales smooth and
imbricate, cycloid, arranged in longitudinal rows. Large rostral (HW/WR = 4.7 and HL/LR = 12.8), wider than long
(WR/LR = 1.8), triangular, visible in dorsal view, with a fissure extending from the region in contact with the nasal
to half of the rostral, and a perforation in the upper left side. A pair of postrostrals protruding, separated by two tiny
scales and in contact with the one of postnasals. Large supralabials, longer than wide, decreasing in size to the end
of the labial commissure. First supralabial in broad contact with the rostral and one of the postnasals, involving part
of the nostril. Posterior snout region and interorbital region concave. Dorsal and lateral surfaces of the head covered
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with granular juxtaposed scales, with scattered tubercles on the upper surface starting at the level of interorbital
region. Granules in the snout larger than those of the occipital region. Eighteen small granules between the postnasals
and anterior ocular margin. The granules surrounding the ocular region are tiny and more spaced than those of the
snout and the dorsum. Postnasals swelling, elongated and bordering 1/4 of the nostril. The border of the auditory
meatus is surrounded by small scales and granules. In the auditory meatus, the scales are small and smooth. Dorsal
region of the body covered by granular scales and larger tubercles almost equidistant, conical and anteroposteriorly
elongated, arranged in 10 to 14 irregular lines, reaching the level of the posterior region of hindlimbs, just before
tail insertion. Postcloacal tubercles present, three on each side, easily perceived. Postcloacal pores present, one on
each side. Regenerated tail, presenting smaller overlapping cycloid scales in the dorsal region and increasing in size
in the lateral region. A row of smooth, elongated medial scales in the ventral region of the tail, two or three times
wider than long, covering to half of the ventral region of tail. Dorsal surface of the forelimbs and hindlimbs different
of the dorsum of the body, with medium scales smooth and imbricate, tubercles absent. Palmar and plantar regions
with small granules, replaced in the forearms by smooth, cycloid, and imbricated scales. Infradigital lamellae on
the 4th finger of the forelimbs and hindlimbs wider than long, wider than high, almost straight; two distal lamellae
in open V-shaped. Claws bordered by smooth and imbricate scales, composed of five scales in the ventral region,
five dorsal scales. Side of claws with two rows of scales with five scales each. Presence of sheath with three scales.
Coloration in life (Fig. 6 and 10A). Based on a not collected topotype: Body with background color Olive
Horn (16). The dorsum has irregular bands on sides beginning in the postnasal region and extending towards the
base of the tail; these bands show irregular stains in the Sepia (229) surrounded by Pale Cinnamon (55) tones. Small
irregular spots with Brownish Olive (276) tones distributed along the dorsum of the body and limbs. A lateral band
in the head beginning in the labial commissure (rather than dashes), Sepia (229) and Pale Cinnamon (55) colors,
that extends until the hindlimbs. Limbs in Sepia (229) and Sulphur Yellow (91) pattern with irregular spots in Brick
Red (36) up to the claws. Head Olive Horn (16). Irregular Brownish Olive (276) spots between the eyes and the
auditory meatus. Tail with well-defined transverse bands alternating between Olive Horn (16) and Sepia (229) with
Brownish Olive (276) spots. In the beginning of the tail, there is a Brownish Olive (276) triangular-shaped spot.
The regenerated segment of the tail is an Olive Horn (16) that do not form a distinguishable pattern. Ventral region
Sulphur Yellow (91), without spot pattern. Infradigital lamellae Pale Mauve (204).
Coloration in preservative (Fig. 5). The color pattern of holotype differs significantly from the coloration
of the topotype in life described above. The dorsal pattern is almost homogeneous with little contrast between the
irregular bands and background coloring (although these are visible when looking more closely). The background
color is Dark Drab (45) and the bars are Brunt Umber (48). The tail region (regenerated) has a lighter color in Drab
(19) and the ventral region becomes Pale Horn Color (11) slightly darker.
Intraspecific variation. All diagnostic characteristics for the new species are seen in all specimens analyzed.
The MZUSP 106772 specimen does not present postcloacal tubercles and the MZUSP 106776 lacks a postrostral
scale. Morphometric variation and the scale count range among specimens are provided in Appendix III.
Distribution, habitat, and natural history. The new species is currently known only from the mountains
of Serra do Sincorá, in the Chapada Diamantina, an area situated in the northern segment of Espinhaço mountain
range. A mosaic of vegetation types, of which “campos rupestres”, or rupestrian grasslands, a dominant open-rock
pioneer vegetation with rock-dwelling plants, are most common, characterizes the area. Notwithstanding, there are
varieties of other environments in the region, like gallery forests, “Cerrado” (savanna-like), montane forests and
semi-deciduous to deciduous forests (Giulietti & Pirani 1988).
Phyllopezus diamantino sp. nov. is nocturnal and specimens were found on rocky outcrops and in tree and shrub
trunks. Active animals were found only at night on the surface of rocks or trees or in rock crevices, and during the
day only one inactive specimen was found under a rock.
Gymnodactylus vanzolinii and Hemidactylus brasilianus were observed syntopically with P. diamantino sp.
nov. on the rocks or in rock crevices, even though the new species was also found in other microhabitats as tree
and shrub trunks. Other lizards observed and recorded at Mucugê area were: Hemidactylus mabouia (Gekkonidae),
Acratosaura mentalis, Acratosaura spinosa, Heterodactylus septentrionalis, Micrablepharus maximiliani, Psilops
mucugensis (Gymnophthalmidae), Enyalius erythroceneus (Leiosauridae), Polychrus acutirostris (Polychrotidae),
Brasiliscincus heathi (Scincidae), Ameiva ameiva, Ameivula cf. ocellifera, Tupinambis merianae (Teiidae),
Eurolophosaurus sp., Tropidurus hispidus, T. mucujensis, and T. semitaeniatus (Tropiduridae) (Freitas & Silva
2007; Cassimiro & Rodrigues 2009).
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FIGURE 5. Holotype of Phyllopezus diamantino sp. nov. (MZUSP 106770, adult female). A and B = dorsal and ventral views
of body; C = palm of hand; D = sole of the foot; E–G = dorsal, lateral and ventral views of head; H = auditory meatus; I = labial
commissure; J = cloacal region; K = tubercles and postcloacal pores on the left side of the body; L = tubercles and dorsal scales;
M = ventral scales. Scale bar = A and B (10mm), C–G and J (3mm).
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FIGURE 6. Color in life of topotype (unvouchered specimen) of Phyllopezus diamantino sp. nov.
Phyllopezus selmae sp. nov.
(Figs. 7, 8, 9H, 10B)
Hemidactylus mabouia: Roberto et al. (2015: part, p. 715, fig. 7)
Phyllopezus sp.: Gonçalves & Palmeira (2016)
Phyllopezus sp.2 (aff. pollicaris): Dubeux et al. (present study)
Holotype. MHNUFAL 13481 (adult female) from Cariri da Prensa Farm (9°41’41”S, 36°12’16”W; 83 m a.s.l.),
municipality of Boca da Mata (Fig. 1B), Alagoas state, Brazil, collected on 15 January 2014 by U. Gonçalves and
C. Palmeira, field number UGS 702.
Paratypes. Adult female (MHNUFAL 13482) collected on 3 January 2017, from the same locality of holotype
(topotypes); adult females (MHNUFAL 12169, CHUFPE-R 1002, 1003) and adult males (MHNUFAL 12168,
12172, CHUFPE-R 1004) collected on 9 July 2015, adult females (MZUSP 106766, 106767) and adult males
(CHUFPE-R 1005, MZUSP 106768) collected on 20 January 2015, and adult males (MHNUFAL 12396, 12397,
12399, 12400, MZUSP 106769) collected on 16 November 2015, from municipality of Limoeiro de Anadia (Fig.
1B), Alagoas state, Brazil (9°47’05”S, 36°28’04”W; 117 m a.s.l.); adult female (MHNUFAL 12449) and adult
male (MHNUFAL 12128) collected on 7 July 2015, from municipality of Coruripe (Fig. 1B), Alagoas state, Brazil
(10°03’17”S, 36°16’32”W; 68 m a.s.l.); adult females (MHNUFAL 10200) collected on 23 January 2015, from
municipality of Igaci (Fig. 1B), Alagoas state, Brazil (9°32’00”S, 36°36’43”W; 269 m a.s.l.), all paratypes above
were collected by U. Gonçalves and C.N.S. Palmeira; adult male (MHNUFAL 12401) collected on 13 June 1999
by Selma Torquato, adult male (MHNUFAL 16198) and juvenile unsexed (MHNUFAL 16199) collected on 21
April 2019 by M.J.M. Dubeux, from municipality of Quebrangulo (Fig. 1B), Alagoas state, Brazil (9°15’22”S,
36°25’43”W; 780 m a.s.l.).
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Etymology. The name of species is in honor of Selma Torquato, curator of Coleção Herpetológica do Museu
de História Natural da Universidade Federal de Alagoas who has generously provided many opportunities for
herpetologists to study the amphibian and reptile specimens under her care.
Diagnosis. Phyllopezus selmae sp. nov. is characterized by the following combination of characters: (1) Mental
scales in bell shaped, with concave margins and a slight central constriction, similar length and width and posterior
margin not exceeding the second infralabial; (2) postmental scales enlarged, hexagonal, twice as long as wide, with
broad contact each other and previously separated by about 1/5 of its length by the mental scale; (3) up to two scales
in contact with the ventral margin of first infralabial; (4) enlarged scales surrounding and separating postmental
scales from granules of the gular region; (5) six to seven infralabial scales; (6) cycloid and imbricated scales of
similar size in distal region of mandible; (7) enlarged dorsal tubercles, corresponding to about six granular scales,
elongated and slightly keeled; (8) developed pollex; (9) cycloid or triangular scales around the auditory meatus,
appears somewhat bristly; (10) homogeneous scales of the same size in the region of the labial commissure; (11) up
to two tubercles or tubercles absents in the angular region between the upper and lower edges of the opening of the
auditory meatus and eyes; (12) postcloacal pores not always present; and (13) large sized, SVL 89.2–100.24 mm in
males, and 83.5–99.47 mm in females. See Comparison with congeners section for more additional diagnosis with
Description of holotype. Adult female, SVL 99.47 mm, fully regenerated tail, DBL 42.82 mm, TBW 12.31
mm, HL 27.02 mm, HW 19.53 mm, HD 9.2 mm, SL 11.08 mm, NSD 2.73 mm, ESD 8.29 mm, ED 6.09 mm, IOD
8.42 mm, IND 3.79 mm, LH 20.11 mm, LF 12.23 mm, LT 20.88 mm, LTB 15.3 mm, WM 4.64 mm, LM 4.98 mm,
WR 4.54 mm, LR 2.3 mm, R 1, PR 2, SN 2, SL 8, IL 7, M 1, PM 2, SSP 7, VLR 51, DT 43, L4F 14, L4T 13, TP 2,
and CP 0. Head large (SVL/HL = 3.68), distinct from neck. Mental large (HW/WM = 4.2 and HL/LM = 5.4), bell-
shaped, slightly longer than wide (WM/LM = 0.93), narrower posteriorly and with a slight strangulation in its half,
bordered by the 1st infralabial and in broad contact with two postmentals that separated it from others infralabial and
gular scales. A pair of postmentals, large, hexagonal, juxtaposed, longer than wide, and flanked by seven large scales
with differentiated sizes, which are replaced by small scales, smooth and imbricate, similar to ventral ones. First, 2nd
and 3rd infralabials rhomboid; 1st the largest, in contact with the postmental pair, and a group of five large, smooth
and variable in shape scales that isolate the pair of postmentals from scales in the gular region. The infralabials
decrease in size towards the labial commissure; commissure area with granules. From the 2nd infralabial, there is a
group of small, elongated scales that border the infralabial row near of the labial commissure, which also isolates
them from the granules of the gular region. Ventral scales smooth and imbricate, cycloid, arranged in longitudinal
rows. Large rostral (HW/WR = 4.3 and HL/LR = 11.74), wide than longer (WR/LR = 1.97), triangular-shaped,
with a median depression at the top where there is a fissure extending from the region in contact with the nasal
to half of the rostral. Large supralabials, longer than wide, decreasing in size to the end of the labial commissure.
First supralabial in broad contact with the rostral and one of the postnasals, involving part of the nostril. Dorsal and
lateral surfaces of the head covered with granular juxtaposed scales, with scattered tubercles on the upper surface
starting at the level of interorbital region. Granules in the snout four to five times larger than those of the occipital
region. Fifteen small granules between the postnasals and anterior ocular margin. The granules surrounding the
ocular region are tiny and more spaced than those of the snout or the dorsum. The supranasal region involves half of
the nasal fossa. Postnasals swollen, elongated and bordering 1/3 of anterior portion of the nostril. The border of the
auditory meatus is surrounded by small granules. In the auditory meatus, the scales are erect and acicular, triangular,
smooth and imbricate. Dorsal region of body covered by granular scales and almost equidistant larger tubercles,
conic and elongated anteroposteriorly, arranged in 11 to 14 irregular lines, reaching the level of the posterior region
of the hindlimbs (before the tail insertion). Postcloacal tubercles present, a pair on each side, very conspicuous.
Postcloacal pores absent. Regenerated tail, presenting smaller overlapping cycloid scales in the dorsal region and
increasing in size in the lateral region. A row of smooth, elongated medial scales in the ventral region of tail, three or
four times wider than long, covering almost the entire ventral region. Dorsal surface of the forelimbs and hindlimbs,
with medium sized scales, smooth and imbricate; tubercles absent. The small granules in the palmar and plantar
regions are replaced by smooth, cycloid, and imbricated scales in the forearms. Infradigital lamellae on the fourth
finger and fourth toe wider than long, wider than high, slightly arched and becoming straighter in the distal portions.
Claws bordered by smooth and imbricate scales, composed of five scales in the ventral region, five dorsal scales.
Side of the claws with two rows of scales with five scales each. Presence of sheath with three scales.
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FIGURE 7. Holotype of Phyllopezus selmae sp. nov. (MHNUFAL 13481, adult female). A and B = Dorsal and ventral views
of body; C = palm of hand; D = sole of the foot; E–G = dorsal, lateral and ventral views of head; H = auditory meatus; I = labial
commissure; J = cloacal region; K = tubercles on the left side of the body; L = tubercles and dorsal scales; M = ventral scales.
Scale bar = A and B (10mm), C–G and J (3mm).
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Coloration in life (Fig. 8). Based on holotype: Body with background color Raw Umber (22). The dorsum with
semicontinuous longitudinal bands, on sides beginning in the postnasal region and extending towards the base of the
tail; band near the dorsal midline begins in the nuchal region; these bands show irregular dashes in the Dusky Brown
(285) surrounded by Sayal Brown (41) tones. Small irregular spots in the Pale Horn Color (11) tones distributed
along the dorsum of the body and limbs. A lateral band in the head beginning in the labial commissure (rather
than dashes), Dusky Brown (285) color, that extends until the hindlimbs. Limbs in Raw Umber (22) pattern with
irregular spots in Dusky Brown (285) and Sayal Brown (41) up to the claws. Head Raw Umber (22), superimposed
by irregular spots Dusky Brown (285). Snout with a triangular-shaped Army Brown (46) spot, surrounded by Dusky
Brown (285). Irregular Dusky Brown (285) spots between the eyes and the auditory meatus. Tail with well-defined
transverse bands alternating between Raw Umber (22) and Sayal Brown (41) with Dusky Brown (285) spots.
In the beginning of the tail, there is a Dusky Brown (285) triangular-shaped spot. The regenerated segment of
the tail is a Raw Umber (22) color that is overlaid by Sayal Brown (41) spots that do not form a distinguishable
pattern. Ventral region Pale Horn Color (11), without spot pattern. Infradigital lamellae Pale Mauve (204).
Coloration in preservative (Fig. 7). In general, the coloration in preservative does not differ substantially
from life coloration. The background color becomes similar to Beige (254), tending to a more grayish tone. The
longitudinal bands retain their color; however, they lose the Sayal Brown color (41) that surrounds them in life,
becoming more prominent in relation to the background. On the dorsal surface of the limbs and in the regenerated
portion of the tail, the Sayal Brown color (41) becomes Fawn Color (258) and the ventral region becomes Pale Horn
Color (11) slightly darker.
Intraspecific variation. All diagnostic characters used for describing the new taxon are present in all specimens
analyzed. However, different dorsal background coloration in life were observed, depending on the time and type
of the substrate of capture, ranging from Raw Umber (22) to Drab (19). Ontogenetic variations of color were also
observed; a juvenile (MHNUFAL 16199; Fig. 8C) presents a more demarcated dark longitudinal bands and Pale
Buff (1) lighter color background. Morphometric and meristic variation are provided in Appendix III.
FIGURE 8. Coloration in life of Phyllopezus selmae sp. nov. (A) Holotype [MHNUFAL 13481], (B and C) paratypes
[MHNUFAL 16198 and MHNUFAL 16199, respectively], (C) variation of juvenile coloration.
Distribution, habitat, and natural history. Phyllopezus selmae sp. nov. is a nocturnal species found in rocky
outcrops and trees, in heights up to 10 m. In the daytime, specimens were found sheltering either under tree bark,
clumps of epiphytes or bromeliad roots. Animals were mainly observed active in the early evening when foraging
in forested sites near rivers with rocky bed. The species was also found sharing bromeliads with P. lutzae. When
specimens were captured, they twisted their body by turning quickly to the side, and when attempting to bite would
produce an agonistic “squeaking” sound (not recorded). The distribution of the species is only known for the state
of Alagoas, with altitudes ranging from 68 m in the municipality of Coruripe to 780 m a.s.l. at the top of the rock
formation of Pedra Talhada, municipality of Quebrangulo (Fig. 1).
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Comparisons with congeners. Phyllopezus diamantino sp. nov. and Phyllopezus selmae sp. nov. are
morphologically more similar to each other than to the other representatives in the genus and together are
distinguished from the congeners mainly by characters in the gular region (Fig. 9). Both new species (Fig. 9G–H)
differ from P. periosus (Fig. 9A) by the presence of increased scales separating the postmentals granules in the
gular region (absent in P. periosus), posterior margin of the mental scale not exceeding the anterior margin of the
second infralabial scale (exceeding the anterior margin of the second infralabial in P. periosus) and postmental
scales in direct contact (separated by the mental scale in P. periosus). Both new species differ from P. lutzae (Fig.
9B) by presenting a long mental scale, with similar length and width (short mental with a length corresponding
to half the width in P. lutzae), posterior margin of the posmental scales exceeding half of the second infralabial
(not reaching the second infralabial in P. lutzae) and up to two scales in contact with the first infralabial (three to
four scales in contact with the ventral margin of the first infralabial in P. lutzae). Both new species differ from P.
maranjonensis (Fig. 9C) in having the central pair of postmentals distinctly larger than the scales that surround them
and in contact with the first infralabial (almost the same size and separated from the first infralabial by one or two
scales in P. maranjonensis). Both new species differ from P. maranjonensis and P. heuteri in having six or seven
infralabial scales (7–10 in P. maranjonensis and 8–9 in P. heuteri). Both new species differ from P. maranjonensis,
P. heuteri (Fig. 9F), P. pollicaris sensu stricto (Fig. 9D) and P. przewalskii (Fig. 9E) in having postmental scales
twice longer than wide (postmentals with similar width and length in P. maranjonensis, P. heuteri, P. pollicaris and
FIGURE 9. Scheme of the arrangement of scales in the gular region of Phyllopezus species. (A) P. periosus [based on CHUFPB
1936], (B) P. lutzae [based on CHUFPB 19518], (C) P. maranjonensis [based on ZFMK 84997], (D) P. pollicaris stricto sensu
[ZSM 165/0/1, paralectotype], (E) P. przewalskii [based on SMF 100496], (F) P. heuteri [based on SMF 100494], (G) Phyllopezus
diamantino sp. nov. [based on MZUSP 106770] and (H) Phyllopezus selmae sp. nov. [based on MHNUFAL 13481]. Figures
D–F adapted of the photographs provided by Cacciali et al. (2018). Horizontal dashed gray line marks the posterior margin of
the third infralabial scale. Scale bars = 3mm.
The two new species can also be distinguished from P. periosus in having a color pattern in longitudinal or
transverse dark irregular dorsal bars (6 to 7 well-defined light-colored transverse bands limited anteriorly and
posteriorly by bars dark in P. periosus). The two new species can be distinguished from P. lutzae in being distinctly
larger (72.38–96.25 mm in Phyllopezus diamantino sp. nov. and 83.50–100.24 mm in Phyllopezus selmae sp. nov.
versus a maximum of 62.77 mm in P. lutzae), in having a coloration pattern in irregular longitudinal or transverse
dorsal dark bars (homogeneous orange marbled pattern in P. lutzae). Both new species can be distinguished from P.
lutzae and P. maranjonensis in having larger dorsal tubercles, corresponding to about six granules (indistinct dorsal
tubercles in P. lutzae and few slightly enlarged tubercles on the back, rarely forming rows on P. maranjonensis).
Both new species can be distinguished from P. lutzae in having developed pollex (absent or poorly developed in
P. lutzae). The two new species can be distinguished from P. maranjonensis in having a pattern of coloration in
irregular longitudinal or transverse dorsal dark colored bars (four regular dark colored cross bars between the neck
and vent in P. maranjonensis), and six to eight supralabial scales (8–10 in P. maranjonensis). Both new species
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can be distinguished from P. heuteri in having cycloid or triangular scales around the auditory meatus, little bristly
(spiny and bristling scales in P. heuteri).
FIGURE 10. Coloration in life of dorsal view of the body and dorsolateral view of head of (A) Phyllopezus diamantino sp. nov.
[topotype (unvouchered specimen)] and (B) Phyllopezus selmae sp. nov. [paratype (MHNUFAL 16198)].
Phyllopezus diamantino sp. nov. can be differentiated from Phyllopezus selmae sp. nov. due to the presence
of granular and juxtaposed scales in the distal region of mandible, and may present tubercles of different sizes
(cycloid and imbricated scales, of similar size in Phyllopezus selmae sp. nov.), mental scale triangular with almost
straight lateral edges (bell-shaped with concave margins and a slight strangulation in its half in Phyllopezus selmae
sp. nov.; Fig. 9), and anterior portion of the postmental scales separated by almost 1/3 of the length by the mental
scale (versus separated by 1/5 in Phyllopezus selmae sp. nov.; Fig. 9), dorsal coloration pattern in dark transverse
bands interrupted by a light cervical band (dark bands arranged in well-defined or transverse longitudinal rows
showing interruptions, light cervical band absent or not evident in Phyllopezus selmae sp. nov.; Figs. 5–8 and 10),
four to six tubercles in the angular region between the upper and lower edges of the opening of the auditory meatus
and eyes (up to two tubercles or tubercles absents in this region in Phyllopezus selmae sp. nov.; Figs. 5F and 7F),
homogeneous scales of the same size in the region of the labial commissure (increased scales on the upper and lower
sides of the labial commissure in Phyllopezus selmae sp. nov.; Figs. 5I and 7I), and a pair of postcloacal pores is
always present (not always present in Phyllopezus selmae sp. nov., Figs. 5K and 7K). Morphometric variations and
the scale counts range among specimens analyzed are provided in Appendix III.
In addition to the cryptic diversity already recognized for the Phyllopezus pollicaris complex (Gamble et al. 2012;
Werneck et al. 2012; Cacciali et al. 2018), the loss of the lectotype (ZSM 2510/0; Michael Franzen comm. pers.
in Cacciali et al. 2018) has hampered taxonomic arrangements involving the group. Phyllopezus pollicaris was
described by Spix (1825) as Thecadactylus pollicaris almost 200 years ago, to accommodate a population in the
interior of the state of Bahia (“in sylvis interioris Bahiae campestribus”), with no further locality data (Spix 1825;
In Bahia state, a high diversity of lineages belonging to the P. pollicaris complex were found, some of them
are not closely related (P. diamantino sp. nov. [previously P. pollicaris Clade I] and P. pollicaris Clades III, VI and
VIII; Werneck et al. 2012; Cacciali et al. 2018; present study). All the diagnostic characters for both of the new
species and the comparisons with congeners were based on the photographs of the paralectotypes of P. pollicaris
[ZSM 165/0/1–2; according to Müller & Brongersma (1933) those specimens were in the original type series], in the
TWO NEW PHYLLOPEZUS FROM BRAZIL Zootaxa 5120 (3) © 2022 Magnolia Press · 363
detailed description of these provided by Müller & Brongersma (1933) and Cacciali et al. (2018), and in the original
description by Spix (1825). We observed that the P. pollicaris paralectotypes are morphologically similar to lineages
belonging to Group 2 (P. heuteri + P. przewalskii + P. pollicaris Clades IV, VI–VIII). They share characteristics
such as number, size, format and disposition of the gular scales and its small size (see “Taxonomic Implications”
and “Comparison with Congeners” sections). Although the municipality of Mucugê is located in Bahia, the same
Brazilian state where the type-locality of P. pollicaris sensu stricto is located, based on the morphological comparisons
explicated above proves that the new species is not the same phylogenetic lineage of P. pollicaris of Spix.
The 16S rRNA was the most conservative mitochondrial gene used in the delimitation of genetic clusters of
Phyllopezus (Cacciali et al. 2018) and recovered reciprocally monophyletic P. diamantino sp. nov. and P. selmae
sp. nov. In addition, it was efficient in recovering the phylogenetic relationships of the Phyllopezus lineages even
among the more recent lineages, as already identified in previous studies (Gamble et al. 2012; Cacciali et al.
FIGURE 11. Typical environments of the type localities of (A and B) Phyllopezus diamantino sp. nov. in the municipality of
Mucugê, Bahia State and (C and D) P. selmae sp. nov. in the municipality of Boca da Mata, Alagoas State, Northeastern Brazil.
The high cutoff point for the transition between intra and interspecific variation identified by the analysis
of Local Minima (5.73%) and reinforced by ABGD is probably the result of the long and complex evolutionary
history of the genus (Werneck et al. 2012). According to Werneck et al. (2012) the populations of the municipality
of Mucugê (Bahia state) (P. diamantino sp. nov.), together with those of the municipalities of São Geraldo do
Araguaia (Pará state) and Barra do Garça (Mato Grosso state) (Clade II; not included in the present study), and
now also the lineage of P. selmae sp. nov. for Alagoas state (present study), correspond to the sister clade to all the
remaining representatives of the P. pollicaris complex. These lineages highly distinct from the other congeners and
that diverged earlier (~11.5 Ma). Although related, the lineages within this clade have a widely disjunct distribution
and are surrounded by areas of occurrence of other lineages belonging to the complex (Werneck et al. 2012). Such
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364 · Zootaxa 5120 (3) © 2022 Magnolia Press
a scenario may represent sampling gaps in some regions or a wide historical extinction of these lineages in the
intermediate regions, which may be key to understanding the diversification and current distribution of the oldest
lineages of P. pollicaris complex.
The type locality of P. diamantino sp. nov. is located in the northern portion of the Serra do Espinhaço mountain
range, a region of “campos rupestres” with more than 1,000 meters of elevation (Fig. 11A and B). Such mountain
regions are characterized by unique floras and a high rate of endemism. Past climates altering and isolating habitats
in those mountain ranges may have been related to events that led to the divergence of lineages within the P.
pollicaris complex. Similar biogeographic scenario has already been reported for frogs (e.g., Lugli & Haddad
2006a, b; Cassimiro et al. 2008; Faivovich et al. 2009; Pombal Jr et al. 2012; Leite et al. 2012; Carvalho et al. 2013;
Trevisan et al. 2020), lizards (e.g., Rodrigues et al. 2006; Cassimiro & Rodrigues 2009; Rodrigues et al. 2009a, b;
Rodrigues et al. 2017), and amphisbaenians (e.g., Mott et al. 2008).
Although Magalhães et al. (2015) has not registered any species of Phyllopezus during the inventory for the
Parque Nacional da Chapada Diamantina, P. diamantino sp. nov. (identified as P. pollicaris) had already been
registered for the municipality of Mucugê, state of Bahia (Cassimiro & Rodrigues 2009; Freitas et al. 2012). As it
is a generally common species where it occurs and of easy observation (MJMD pers. obs.) the absence of records
in other regions of the park, despite the considerable sampling effort realized by Magalhães et al. (2015) (30 days
of sampling effort, 26,640 hours of trap arrays, and 960 hours of active random searches), may be an indication of
a restricted distribution of this species. This fact reinforces the need to identify and attribute names to this cryptic
species, since the current taxonomic context can give the false impression of a species with wide distribution to
lineages that, in fact, are geographically restricted and that may be under different degrees of threat. This fact
sometimes hampers the understanding of the evolutionary processes responsible for the pattern of diversification
of organisms, in addition to underestimating the degree of threat of a given species, hindering the development of
consistent and reliable conservation plans (Meiri et al. 2018).
Although P. selmae sp. nov. is presently know only from state of Alagoas, it is likely that its distribution is
underestimated, since the state still has a high level of Wallacean deficit (lack of knowledge of the geographic
distribution of species; Whittaker et al. 2005). Almost all records for the species are associated with the Coruripe
River basin, and the samples were obtained during one of the few herpetofauna inventories carried out in the Agreste
region of the state, which made the transition between the Atlantic Forest and the semiarid Caatinga biomes (Fig.
11C and D; Gonçalves & Palmeira 2016).
Although we are far from resolving the taxonomy of P. pollicaris species complex, the present work is a step
forward in that direction formally describing its concealed diversity and unveiling crucial data for conservation
policy in a megadiversity but still unknown and so threatened country (Moura & Jetz 2021).
The authors thank S. Torquato (MHN-UFAL), F. Delfim and D. Mesquita (CHUFPB), and E. Freire (UFRN) for
allowing us to study specimens under their care; to the Seresta S.A, specifically G. Neto, J. Prado and M. Daher for
logistic support along the Coruripe River basin; to Centro Nacional de Pesquisa e Conservação de Répteis e Anfíbios
of Instituto Chico Mendes de Conservação da Biodiversidade (RAN/ICMBio) for collection permits; to L. Lima, L.
Lobo and F. Soares for field work help; F. Magalhães, I. Costa-Silva and A. Mello for help in the analysis. MJMD
thanks Fundação de Amparo à Ciência e Tecnologia do Estado de Pernambuco - FACEPE (IBPG-1117-2.04/19),
FPW, MTR, PMSN and TM thank Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq
(311504/2020-5 to FPW; 302864/2020-2 to MTR; 313622/2018-3 and 432506/2018-7 to PMSN; and 309904/2015-
3 and 312291/2018-3 to TM) for financial support. JC thanks F.S.F. Leite and L.E. Lopes for their valuable help
and company during the fieldworks in Mucugê. MTR thanks FAPESP (2011/50146-6 and 2003/10335-8), and all
students of his lab for support during field work.
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APPENDIX I. Additional specimens examined
Phyllopezus lutzae: BRAZIL: Paraíba: Pedras de Fogo (CHUFPB 19517, 19518, 19519, 24979); Alagoas: Maceió (UFRN 36,
236; LABI 181, 182, 183); Quebrangulo ([MHNUFAL] = LABI 653).
Phyllopezus periosus: BRAZIL: Paraíba: Cabaceiras (topotypes: CHUFPB 1930, 1931, 1934, 1936, 1939, 1940, 1947); Rio
Grande do Norte: Serra Negra do Norte (MHNUFAL 12426, 12428); Currais Novos (UFRN 5551, 5552).
Phyllopezus pollicaris: BRAZIL: Bahia: Paulo Afonso (CHUFPB 11294, 11295, 11296, 11297, 11298, 11299, 11300, 12058,
22330, 25018, 25111, 25137, 25569); Ceará: Aiuaba (CHUFPB 5145, 5147, 5153, 5159, 5160, 5161, 5162, 5163, 5165,
5324, 5325, 5326, 5327, 5328, 5329, 13767, 13768, 13769, 13778, 13780, 13788); Ubajara (CHUFPB 26129, 26894,
26916); Paraíba: Cabaceiras (CHUFPB 10224, 10225, 10228, 10229, 10230, 10231); Pernambuco: Buíque (CHUFPB
23509, 25026, 25562, 25973; [CHUFPE-R] = CAT 272, 391, 271,388, 392); Piauí: Uruçuí (CHUFPB 8697); Floriano
(CHUFPB 8699); Coronel José Dias (CHUFPB 14603, 14606, 14608, 14611, 14614, 14616, 14617, 14618); Andorinha
(CHUFPB 22324, 22349, 22364, 22450, 22473, 22603); São Raimundo Nonato (CHUFPB 25050, 25153); Sergipe:
Canindé de São Francisco (CHUFPB 18559, 18562, 18563, 18567, 18568, 18570, 18573, 18581, 18587, 18592, 18594,
18595, 18602, 18605, 18626, 18633).
APPENDIX II. 16S rRNA mitochondrial gene fragment sequences of Phyllopezus samples used in the study. Clades
previously delimited by Werneck et al. (2012) are identified by Roman numerals.
Species Clade Voucher Locality GenBank
Phyllopezus diamantino sp. nov. I (MTR) JC 1185 Mucugê, Bahia, Brazil JN935553
Phyllopezus diamantino sp. nov. I (MTR) JC 1219 Mucugê, Bahia, Brazil JN935554
Phyllopezus selmae sp. nov. - MHNUFAL 10200 Igaci, Alagoas, Brazil OM530506
Phyllopezus selmae sp. nov. - MHNUFAL 13482 Boca da Mata, Alagoas, Brazil OM530507
Phyllopezus selmae sp. nov. - MZUSP 106768 Limoeiro de Anadia, Alagoas, Brazil OM530508
Phyllopezus pollicaris IV LG 1309 Serra da Mesa, Goiás, Brazil JN935569
Phyllopezus pollicaris IV LG 1792 Palmas, Tocantins, Brazil JN935572
Phyllopezus pollicaris IV LG 1845 Lajeado, Tocantins, Brazil JN935574
Phyllopezus pollicaris VI MTR 3074 Santo Inácio, Bahia, Brazil JN935584
Phyllopezus pollicaris VI MTR 3287 Santo Inácio, Bahia, Brazil JN935586
Phyllopezus pollicaris VI MTR 3263 Gentio do Ouro, Bahia, Brazil JN935585
Phyllopezus pollicaris VII LG 1310 Niquelândia Goiás, Brazil JN935568
Phyllopezus pollicaris VII CHUNB 36991 Paranã, Tocantins, Brazil JN935559
Phyllopezus pollicaris VII CHUNB 36992 Paranã, Tocantins, Brazil JN935560
Phyllopezus pollicaris VII CHUNB 37001 Paranã, Tocantins, Brazil JN935561
Phyllopezus pollicaris VII CHUNB 43850 São Domingos, Goiás, Brazil JN935563
Phyllopezus pollicaris VII CHUNB 43852 São Domingos, Goiás, Brazil JN935564
Phyllopezus pollicaris VIII MTR 887020 Cabaceiras, Paraíba, Brazil JN935558
Phyllopezus pollicaris VIII LG 1011 Porto Seguro, Bahia, Brazil JN935566
Phyllopezus pollicaris VIII LG 807 Xingó, Alagoas, Brazil JN935575
Phyllopezus pollicaris VIII LG 808 Xingó, Alagoas, Brazil JN935576
Phyllopezus pollicaris VIII LG 1342 Campo Formoso, Bahia, Brazil JN935570
Phyllopezus pollicaris VIII LG 1343 Campo Formoso, Bahia, Brazil JN935571
Phyllopezus pollicaris VIII MTR 2346 Uruçuí-Una, Piauí, Brazil JN935580
Phyllopezus pollicaris VIII MTR 2807 Uruçuí-Una, Piauí, Brazil JN935581
Phyllopezus pollicaris VIII MTR 2958 Uruçuí-Una, Piauí, Brazil JN935582
Phyllopezus pollicaris VIII MTR 2959 Uruçuí-Una, Piauí, Brazil JN935583
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TWO NEW PHYLLOPEZUS FROM BRAZIL Zootaxa 5120 (3) © 2022 Magnolia Press · 369
APPENDIX II. (Continued)
Species Clade Voucher Locality GenBank
Phyllopezus pollicaris VIII MTR 3681 Ilha do Gado Bravo, Bahia, Brazil JN935587
Phyllopezus pollicaris VIII MTR 3748 Alagoado, Bahia, Brazil JN935588
Phyllopezus pollicaris VIII MTR 4960 Serra das Confusões, Piauí, Brazil JN935589
Phyllopezus pollicaris VIII MZUSP 92491 Serra das Confusões, Piauí, Brazil JN935590
Phyllopezus przewalskii V TG 00105 unknown, Paraguay JN935565
Phyllopezus przewalskii V LG 1093 Fuerte Esperanza, Chaco, Argentina JN935567
Phyllopezus przewalskii V MTD 43490 Fortin Toledo, Boqueron, Paraguay JN935578
Phyllopezus przewalskii V MTD 43492 Fortin Toledo, Boqueron, Paraguay JN935579
Phyllopezus przewalskii V MNCN 5903 Serranía Aguarague, Tarija, Bolivia JN935577
Phyllopezus przewalskii - SMF 100495 Paraguay MF278834
Phyllopezus przewalskii - MNHNP 11957 Paraguay MH397465
Phyllopezus przewalskii - MNHNP 11958 Paraguay MH397466
Phyllopezus heuteri - MNHNP 2 39 Cordillera Department, Paraguay MH397468
Phyllopezus heuteri - MNHNP 2 40 Cordillera Department, Paraguay MH397467
Phyllopezus lutzae - CHUNB 50461 Mata de São João, Bahia, Brazil JN935548
Phyllopezus lutzae - CHUNB 50462 Mata de São João, Bahia, Brazil JN935549
Phyllopezus lutzae - CHUNB 50463 Mata de São João, Bahia, Brazil JN935550
Phyllopezus periosus - MTR 887022 Cabaceiras, Paraíba, Brazil JN935552
Phyllopezus maranjonensis - ZFMK 84995 Balsas, Peru JN935555
Phyllopezus maranjonensis - ZFMK 84997 Balsas, Peru JN935557
Phyllopezus maranjonensis - ZFMK 84996 Balsas, Peru JN935556
Phyllodactylus xanti - ROM 38490 Baja California Sur, Mexico AY763284
APPENDIX III. Morphometrics (in mm) and meristic data [mean ± standard deviation (range)] of Phyllopezus occurring
in northeastern Brazil. Details of acronyms of characteristics are mentioned in “Material and methods” section.
P. lutzae P. periosus P. pollicaris sensu lato
Male(n = 3) Female(n = 7) Male(n = 10) Female(n = 4) Male(n = 43) Female(n = 44)
SVL 61 ±1.61
DBL 26.12 ±1.1
TBW 6.95 ±0.44
HL 17.58 ±0.67
HW 11.6 ±0.45
HD 6.1 ±0.43
SL 7.33 ±0.13
NSD 1.65 ±0.22
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APPENDIX III. (Continued)
P. lutzae P. periosus P. pollicaris sensu lato
Male(n = 3) Female(n = 7) Male(n = 10) Female(n = 4) Male(n = 43) Female(n = 44)
ESD 5.43 ±0.2
ED 3.34 ±0.22
IOD 5.91 ±0.3
IND 2.48 ±0.07
LH 11.49 ±0.8
LF 7.09 ±0.44
LT 12.24 ±0.51
LTB 8.51 ±0.18
WM 2.46 ±0.02
LM 1.19 ±0.15
WR 2.68 ±0.05
LR 1.55 ±0.16
R1 ±0(1─1) 1 ±0(1─1) 1 ±0(1─1) 1 ±0(1─1) 1 ±0(1─1) 1 ±0(1─1)
PR 3 ±1(2─3) 3 ±0(2─3) 2 ±0(2─2) 2 ±0(2─2) 2 ±0(2─3) 2 ±0(2─3)
PN 2 ±0(2─2) 2 ±0(2─3) 3 ±1(2─3) 2 ±1(2─3) 2 ±0(2─2) 2 ±0(2─2)
SL 8 ±1(8─9) 8 ±1(8─9) 8 ±1(7─9) 7 ±0(7─7) 7 ±0(6─8) 7 ±0(6─8)
IL 7 ±1(7─8) 7 ±1(7─8) 7 ±1(6─8) 6 ±1(6─7) 6 ±0(5─7) 6 ±0(6─7)
VLR 59 ±4(56─63) 60 ±3(56─64) 43 ±3(38─48) 44 ±1(42─46) 47 ±4(40─60) 45 ±3(39─55)
DT 31 ±11(20─42) 33 ±8(20─43) 45 ±4(37─49) 43 ±4(40─50) 38 ±4(34─47) 37 ±4(30─49)
L4F 10 ±1(9─10) 10 ±1(9─11) 14 ±1(13─15) 13 ±1(12─14) 11 ±1(10─14) 11 ±1(10─14)
L4T 11 ±2(9─12) 11 ±1(9─12) 14 ±1(13─16) 14 ±0(13─14) 12 ±1(10─13) 11 ±1(10─13)
TP 0 ±0(0─0) 0 ±0(0─0) 2 ±1(2─3) 2 ±1(1─2) 3 ±1(0─5) 2 ±1(0─4)
CP 0 ±1(0─1) 0 ±1(0─1) 1 ±0(1─1) 1 ±0(1─1) 0 ±0(0─1) 0 ±1(0─1)
APPENDIX III. (Conitnued)
Phyllopezus diamantino sp. nov. Phyllopezus selmae sp. nov.
Male(n = 5) Female(n = 6) Male(n = 9) Female(n = 13)
SVL 84.66 ±6.7
DBL 35.55 ±2.87
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APPENDIX III. (Continued)
Phyllopezus diamantino sp. nov. Phyllopezus selmae sp. nov.
Male(n = 5) Female(n = 6) Male(n = 9) Female(n = 13)
TBW 8.46 ±1.2
HL 24.1 ±2.05
HW 16.45 ±1.84
HD 8.07 ±0.73
SL 9.98 ±0.71
NSD 2.08 ±0.19
ESD 7.76 ±0.72
ED 5.24 ±1.11
IOD 7.48 ±0.79
IND 2.98 ±0.35
LH 18.07 ±1.64
LF 11.89 ±0.72
LT 20.24 ±1.24
LTB 14.06 ±0.91
WM 3.76 ±0.21
LM 1.93 ±0.19
WR 4.38 ±0.41
LR 4.07 ±0.32
R1 ±0(1─1) 1 ±0(1─1) 1 ±0(1─1) 1 ±0(1─1)
PR 2 ±1(2─3) 2 ±0(2─2) 2 ±0(2─2) 2 ±0(2─2)
PN 2 ±0(2─2) 2 ±0(2─2) 2 ±0(2─3) 2 ±0(2─2)
SL 7 ±0(6─7) 7 ±0(7─7) 7 ±0(6─7) 7 ±0(7─8)
IL 7 ±0(6─7) 7 ±1(6─7) 6 ±1(6─7) 7 ±0(6─7)
VLR 56 ±3(51─59) 55 ±2(52─57) 46 ±1(44─48) 46 ±2(45─51)
DT 45 ±5(38─52) 47 ±3(43─50) 45 ±2(41─47) 44 ±1(42─46)
L4F 12 ±1(11─13) 12 ±1(11─13) 13 ±1(12─14) 13 ±1(11─14)
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APPENDIX III. (Continued)
Phyllopezus diamantino sp. nov. Phyllopezus selmae sp. nov.
Male(n = 5) Female(n = 6) Male(n = 9) Female(n = 13)
L4T 13 ±1(12─14) 13 ±1(12─14) 14 ±1(13─15) 13 ±1(12─14)
TP 2 ±0(2─3) 2 ±0(2─2) 2 ±0(2─3) 2 ±1(0─3)
CP 1 ±0(1─1) 1 ±0(1─1) 0 ±0(0─0) 0 ±0(0─0)