ZooKeys 428: 109–132, doi: 10.3897/zookeys.428.7352
A new species of Chiasmocleis (Microhylidae, Gastrophryninae) from the Atlantic Forest of Espírito Santo State, Brazil
João F. R. Tonini 1,2, Maurício C. Forlani 1, Rafael O. de Sá 1
1 Department of Biology, University of Richmond, Richmond, VA 23173, USA
2 Current Address: Department of Biological Sciences, The George Washington University, Washington, DC 20052, USA

Corresponding author: João F. R. Tonini (jfrtonini@gmail.com)

Academic editor: F. Andreone

received 20 February 2014 | accepted 14 July 2014 | Published 24 July 2014
(C) 2014 João F. R. Tonini. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
For reference, use of the paginated PDF or printed version of this article is recommended.

Citation: Tonini JFR, Forlani MC, de Sá RO (2014) A new species of Chiasmocleis (Microhylidae, Gastrophryninae) from the Atlantic Forest of Espírito Santo State, Brazil. ZooKeys 428: 109–132. doi: 10.3897/zookeys.428.7352


Among Neotropical microhylids, the genus Chiasmocleis is exceptionally diverse. Most species of Chiasmocleis were described in recent years based on external morphology, but recent studies using molecular data did not support the monophyly of the species groups clustered based on feet webbing. Furthermore, a phylogeographic study of C. lacrimae estimated high genetic divergence and low gene flow among populations across small geographic ranges. Increasing the molecular and geographic sampling, and incorporating morphological data, we identified new cryptic species. Herein, we used novel genetic and morphological data to describe a new species of Chiasmocleis.


Amphibians, Chiasmocleis quilombola sp. n. , cryptic species, phylogenetics, systematics


The diversity of evolutionary lineages with little phenotypic differences (i.e., cryptic species) might be better understood in the light of genetic delimitation of evolutionary units (Thomé et al. 2012, Hambäck et al. 2013). Recent molecular phylogenies of anuran, including work on species from the Brazilian Atlantic Forest, did not recovered as monophyletic the species groups clustered based mostly on morphology (Amaro et al. 2009, Canedo and Haddad 2012, Fouquet et al. 2012, Thomé et al. 2012).

Species are segments of population level evolutionary lineages and do not necessarily need to be phenetically distinguishable, diagnosable, monophyletic, intrinsically reproductively isolated, ecologically divergent, or anything else to be considered species, but they only have to be evolving separately from other lineages (de Queiroz 1998, de Queiroz 2007).

A recent molecular phylogeny (de Sá et al. 2012) recovered a polyphyletic Chiasmocleis and, to render the genus monophyletic, transferred one species to Elachistocleis and three species to Syncope. Recently, Peloso et al. (2014) placed Syncope in the synonymy of Chiasmocleis. Chiasmocleis is the most diverse genus of Neotropical microhylids, with 29 species distributed throughout Amazonia, Atlantic Forest, and open areas in South America, such as the Brazilian Cerrado and the Chaco of Bolivia and Paraguay (Cruz et al. 1997, de Sá et al. 2012, Peloso et al. 2014).

Tonini et al. (2013) in a phylogeographic analysis estimated high genetic divergence and low gene flow among populations of Chiasmocleis lacrimae (described as Chiasmocleis carvalhoi Cruz et al. 1997) in the Brazilian Atlantic Forest. Samples of two potential new species and of “Chiasmocleis capixaba” with less feet webbing were included as populations of Chiasmocleis lacrimae. Moreover, the study suggested that populations isolated-by-distance could represent recently diversified species, estimated to Miocene and Pliocene. Increasing the sampling along the distribution of Chiasmocleis lacrimae and Chiasmocleis capixaba and using additional molecular and morphological data, we were able to differentiate the phylogenetic structure and morphological differences associate to intraspecific and interspecific variation. We found that Chiasmocleis lacrimae and Chiasmocleis capixaba were not recovered as monophyletic, in fact populations corresponding to undescribed distinct evolutionary lineages. Although these undescribed lineages have similar body size and shape, and low nuclear divergence, they are exceptionally divergence in mitochondrial markers and are geographically structured.

Herein, we describe a new species of Chiasmocleis from the Atlantic Forest of southeastern Brazil and present a phylogenetic hypothesis for the species group.

Material and methods

Specimens and tissues used herein and comparative material are deposited in the following collections: 1) CFBH: Coleção de Anfíbios Célio Fernando Baptista Haddad, Departamento de Zoologia, Universidade Estadual Paulista Rio Claro, Rio Claro, São Paulo State, Brazil; 2) MNRJ: Museu Nacional do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro State, Brazil; 3) Museu de Zoologia, Universidade de São Paulo, São Paulo, São Paulo State, Brazil; 4) MBML: Museu de Biologia Mello Leitão, Santa Teresa, Espírito Santo State, Brazil; 5) CTA: Coleção de Tecidos e DNA da Universidade Federal do Espírito Santo (UFES) and LGA: Laboratório de Genética Animal, Vitória, Espírito Santo State, Brazil; 6) RN and CTRN: Universidade Federal Rural do Rio de Janeiro, Seropédica, Rio de Janeiro State, Brazil. Field numbers correspond to M. T. Rodrigues (MTR), Universidade de São Paulo, São Paulo, São Paulo State, Brazil; P. Rocha (PEU), Universidade Federal da Bahia, Salvador, Bahia State, Brazil; and J. F. R. Tonini (JFRT), vouchers are at UFES. Specimens examined and tissues samples are listed in Appendix 1 and 2, respectively, and sample localities are shown in Figure 1.

Figure 1.

Sample localities of A tissues and B specimens included in the present study. Sites with more than one color indicates syntopy. List of localities: 1 Porto Seguro, 2 Trancoso, 3 ReBio Córrego Veado, 4 FloNa do Rio Preto (type locality of Chiasmocleis quilombola sp. n.), 5 Parque Estadual de Itaúnas, 6 ReBio Sooretama, 7 Reserva Natural Vale, 8 Povoação, 9 FloNa dos Goytacazes, 10 Costa Bela, 11 ReBio Duas Bocas, 12 Guarapari, 13 Mata da Usina Paineiras, 14 Mimoso do Sul, 15 ReBio União, 16 Cachoeiras de Macacu, 17 Duque de Caxias, 18 Angra dos Reis, 19 Picinguaba, 20 Ilha de São Sebastião, 21 Bertioga, 22 Aracruz (type locality of Chiasmocleis capixaba), 23 Horto Florestal (type locality of Chiasmocleis lacrimae). Blue lines represent major coastal rivers, from North to South: Jequitinhonha, Mucuri, Doce, and Paraíba do Sul. BA = Bahia State, ES = Espírito Santo State, RJ = Rio de Janeiro State, SP = São Paulo State, and MG = Minas Gerais State.

The following measurements were adapted from Duellman (2001) and Peloso and Sturaro (2008); measurements were taken for 56 individuals with a digital caliper under a stereomicroscope to the nearest 0.1 mm: SVL (snout-vent length); HDL (hand length; from the base of the thenar tubercle to the tip of the third finger); HDL4 (hand length from the base of the thenar tubercle to the tip of the fourth finger); HL (head length; from snout to angle of the jaw); HW (head width; between the angle of jaws); ED (eye diameter; between anterior and posterior corner of the eye); IOD (inter-orbital distance; distance between anterior corner of the eyes); IND (inter-nostril distance); END (eye-nostril distance; from the anterior corner of the eye to the posterior margin of nostril); THL (thigh length; from the center of the cloaca opening to the outer edge of the flexed knee); TBL (tibia length; from the outer edge of the flexed knee to the heel); FAL (forearm length); FL (foot length; from tibia-tarsal articulation to tip of fourth toe); 3FD (diameter of third finger disk); 4TD (diameter of fourth toe disk). Fingers and toes are numbered and abbreviated as follows: Fingers I–IV = FI–IV, Toes I–V = TI–V.

Molecular Analyses: Total genomic DNA was extracted from ethanol-preserved liver or muscle tissues using Qiagen DNeasy kit (Valencia, California, USA). We used four molecular markers (mtDNA: 12S, 16S, and NADH dehydrogenase subunit 2 [ND2]; nucDNA: brain-derived neurotrophic factor [BDNF]), amplified using previously published primer sets and PCR profiles (de Sá et al. 2012, Tonini et al. 2013). We performed a multiple loci alignment using an iterative procedure to compute a series of alignment/tree pairs in SATé-II (Liu et al. 2012), using default settings. GenBank accession numbers are given in Appendix 2.

The following outgroup were chosen based on published phylogenies including species of Chiasmocleis (de Sá et al. 2012): Chiasmocleis leucosticta, Chiasmocleis mantiqueira, Chiasmocleis crucis, Chiasmocleis schubarti, and Chiasmocleis cordeiroi. We selected a total of 100 samples (ingroup includes 69 samples) for a data set consisting of 2, 473 base pairs. The best partition schemes and substitution models (Table 1) were chosen using PARTITION FINDER v1.1.1 (Lanfear et al. 2012) and used in phylogenetic analysis downstream.

Table 1.

Best partition scheme and substitution models selected using Partition Finder.

Subset Best Model Subset partitions Subset sites
1 HKY+I+G 12S 1–700
2 HKY+G 16S, ND2_1 701–1044, 1661–2473\3
3 K80+I BDNF 1045–1660
4 HKY+G ND2_2 1662–2473\3
5 GTR+G ND2_3 1663–2473\3

We applied two approaches of phylogenetic estimation: 1) Maximum Likelihood (ML) and 2) Bayesian inference (BI) using the dataset containing the markers 12S, 16S, ND2, and BDNF. Maximum Likelihood in RAXML v7.2.8 (Stamatakis 2006) used a rapid-bootstrap with 1000 replications. Bayesian Inference in BEAST v1.7.4 (Drummond et al. 2012) used birth-death process as tree prior, linked tree models across partition, relaxed clock model with linked mitochondrial markers, but not the nuclear gene. The BI analysis ran for 50 million generations and the parameters were sampled every 5, 000 generations producing a total of 10, 000 trees. We discarded the first 1, 000 trees as burnin in TREEANOTATOR. The output file was checked using TRACER v1.5 and values of Effective Sample Size >200 were considered suitable. Nodes having bootstrap values >70 in ML and posterior probabilities >0.95 in BI were considered as well supported. Analyses were performed through Cipres (Miller et al. 2010) and trees were visualized and edited using FIGTREE. Data available from the Dryad Digital Repository: http:// 10.5061/dryad.gm41t. Genetic distance (p-uncorrected) was calculated in MEGA5.0 (Tamura et al. 2011). A second species awaits description (Forlani et al. submitted) and it is referred throughout this manuscript as Chiasmocleis sp.


The phylogenetic hypotheses generated through ML (Figure 2) and the BI (Figure 3) resulted in similar topology. The ML tree (Figure 2) supported two new species as sister group of Chiasmocleis capixaba, Chiasmocleis lacrimae correspond to a basal node; whereas in the BI tree (Figure 3) Chiasmocleis capixaba was estimated as sister to Chiasmocleis lacrimae, but the posterior probability of this node was lower than 0.95. Both the ML and BI trees showed clades of Chiasmocleis leucosticta, Chiasmocleis mantiqueira, Chiasmocleis crucis, Chiasmocleis schubarti, and Chiasmocleis cordeiroi, but not Chiasmocleis lacrimae and Chiasmocleis capixaba (Figure 2, 3). Populations of Chiasmocleis capixaba and Chiasmocleis lacrimae from the north of the Espírito Santo State, as well as populations of Chiasmocleis lacrimae from southern areas of the Bahia State, would represent two new cryptic lineages closely related to Chiasmocleis lacrimae and Chiasmocleis capixaba. In the ML analysis populations from southern Espírito Santo formed a clade that makes Chiasmocleis lacrimae polyphyletic (Figure 2), whereas in the BI these populations formed a clade including also populations of Chiasmocleis lacrimae from the states of São Paulo and Rio de Janeiro (Figure 3). However, given the lack of support for this node in both analyses, basing taxonomic change on the presumed polyphyly is not warranted at present.

Figure 2.

Maximum likelihood tree including 12S, 16S, ND2, and BDNF. Node numbers correspond to bootstrap, values >70 indicate good support. Although Chiasmocleis lacrimae may not represent a monophyletic species, bootstrap values are low to make further assumptions. Numbers after underscore symbol correspond to localities present in Figure 1. Scale bar represents number of substitutions/site.

Figure 3.

Phylogenetic hypothesis obtained through Bayesian Inference using 12S, 16S, ND2, and BDNF. Node numbers correspond to posterior probabilities, values >0.95 indicate good support. Numbers after underscore symbol correspond to localities present in Figure 1. Scale bar represents number of substitutions/site.

Therefore, our results show that the new species clusters within the genus Chiasmocleis.

Description of a new species
Chiasmocleis quilombola sp. n.


Figure 4

MZUSP147478, adult male, collected at the Floresta Nacional do Rio Preto (Figure 4A), Municipality of Conceição da Barra, Espírito Santo State, Brazil (18°21'19"S, 39°50'39"W), collected on December 8-16, 2009, by L. P. Costa, J. F. R. Tonini, J. Dalapicolla, R. Duda, and C. M. Mattedi.

Figure 4.

Chiasmocleis quilombola sp. n. in vivo. A male (holotype: MZUSP147478) and B female (MZUSP147479, paratopotype). Not in scale.


Males: MZUSP147471–73, MZUSP147475–76, MZUSP147494; female: MZUSP147479 (Figure 4B), Municipality of Conceição da Barra, Espírito Santo State, Brazil (18°21'19"S, 39°50'39"W), collected on December 8-16, 2009, by L. P. Costa, J. F. R. Tonini, J. Dalapicolla, R. Duda, and C. M. Mattedi.


A small-sized species of Chiasmocleis (males SVL mean= 14 ± 1.4 mm; female SVL = 17.1 mm), diagnosed by the following combination of characters: (1) body slender; (2) snout rounded in lateral and dorsal views; (3) all fingers slightly fringed, not webbed, in males and female; (4) all toes fringed and slightly webbed in males and female; (5) dermal spines on fingers and toes of males can be present or absent, absent in female; (6) dermal spines on dorsal surface of males can be present or absent, absent in female; (7) dermal spines absent on ventral surface in males and female; (8) dermal spines on chin and snout of males can be present or absent, absent in female; (9) dermal spines over outer surfaces of legs and cloaca in males can be present or absent, absent in female; (10) female has para-cloacal glands; (11) incomplete occipital fold; (12) vocal slits present in males; (13) dorsal coloration brown; (14) medial ventral body surface light cream colored, whereas ventrolateral surfaces have a light brown and cream marbled pattern; (15) ventral surfaces of fore and hind limbs with a homogeneously and finely dark pattern over a cream background; (16) dorsal surface of fore and hind limbs light brown with a few cream spots or blotches, more distinct on the fore limbs; (17) male throat infuscate; (18) mid-dorsal and/or line on posterior surface of thighs may be present; and (19) tympanum indistinct.

Description of holotype.

Body small (SVL = 15.7 mm), slender, slightly ovoid (Figure 5); head triangular in shape, broader than long; snout short, tip of snout rounded (Figure 5A–B); nostrils located closer to the tip of snout than to eye, not protuberant, directed laterally (Figure 5C); inter-nostril distance smaller than eye–nostril distance and smaller than eye diameter; canthus rostralis slightly defined; loreal region slightly convex; lips not flared; eyes small, slightly protruding; inter-orbital area flat; incomplete occipital fold; tympanum indistinct; upper jaw projecting beyond lower one; tongue large, elongate, and laterally free; premaxillae, maxillae, and vomerine teeth absent; choanae small, rounded, widely separated, positioned anterolaterally to eye; vocal slit present.

Figure 5.

Holotype of Chiasmocleis quilombola sp. n. (MZUSP 147478). A Dorsal B ventral, and C lateral views D right hand and E right foot. White bars = 1 mm.

Arms slender, lacking tubercles on forearm. Hands not webbed (Figure 5D); fingers tips rounded, not expanded, and slightly fringed; fingers lacking dermal spines; finger lengths I<II<IV<III; thumb without nuptial asperities; subarticular tubercles well developed and rounded, proximal subarticular tubercles larger than others; supernumerary tubercles absent; thenar tubercle well developed, ovoid, and at the base of finger I; two palmar tubercles, a rounded inner tubercle and an elongated outer one (Figure 5D). Legs short, moderately robust; knee and heel lacking tubercles; tibial and tarsal ridges absent. Foot slightly webbed (Figure 5A–B, E); toes slightly fringed; toe tip rounded lacking disks; subarticular tubercles well developed, ovoid; supernumerary tubercles absent; an oval inner, but no outer, metatarsal tubercle. Toe lengths I<II<V<III<IV; toes lacking dermal spines; tibia length slightly shorter than thigh length; combined thigh and tibia lengths approximately 82.8% of snout-vent length; foot length approximately 43.9% of snout-vent length.

Skin smooth, dorsal surfaces of body lacking dermal spines. Throat black and few dermal spines found on chin and snout (Figure 5B). Cloaca lacks para-cloacal tubercles or glands.

Coloration in preservative.

Dorsum dark brown with a few small cream spots and blotches; dorsal surface of limbs dark brown with cream blotches and small spots, particularly on the proximal forelimb; palm of hands marbled brown and pale cream, foot dark brown; belly surface cream, dorsolateral and ventral surfaces with a marbled pale brown and cream pattern; throat dark brown to black. Ventral surface of thighs light brown with a finely reticulated dark pattern over a cream background cream with a few cream spots more evident close to the edges; ventral surfaces of tibia and tarsus finely marbled in light brown with cream, lighter than the dorsal surface. Absence of distinct lines on the body and limbs.

Measurements of holotype

(in mm). SVL 15.7; HDL 3.4; HDL4 2.3; HL 2.7; HW 4.5; ED 1.3; IOD 2.8; IND 1.1; END 1.2; THL 6.5; TBL 6.4; FL 6.9, FAL 3.2; 3FD 0.3; 4TD 0.4.

Variation in the type series.

Measurements data of the type series are given in Table 2 and information of the comparative material are provided in Appendix 1. Overall, the type series agrees with the holotype coloration; one specimen has a mid-dorsal line and a line on the posterior surface of the thighs and also more dermal spines (MZUSP147475). The incomplete occipital fold varied from indistinct to weakly visible laterally (= incomplete). The combined mean thigh and tibia length represents approximately 81% of mean snout-vent length in males, and 77.7% in females; foot length approximately 41.6% of snout-vent length in males and 39.7% in females.

Table 2.

Morphometric measurements (mm) of the type series of Chiasmocleis quilombola sp. n.

MZUSP147471 Paratype Male 14.4 2.8 3.8 1.0 2.6 1.1 1.5 6.1 6.3 6.2 0.3 0.3 3.0 2.8 1.8
MZUSP147472 Paratype Male 15.3 2.8 4.1 1.3 2.7 1.2 1.3 7.1 6.6 6.6 0.3 0.4 3.2 3.4 2.2
MZUSP147473 Paratype Male 16.1 3.2 3.9 1.1 2.2 1.1 1.3 6.0 6.0 6.4 0.2 0.2 3.2 3.3 2.1
MZUSP147475 Paratype Male 14.5 2.8 4.6 1.3 2.6 1.3 1.3 6.6 6.4 6.7 0.4 0.4 3.3 3.5 2.2
MZUSP147476 Paratype Male 16.1 2.7 4.4 1.2 2.7 1.3 1.4 6.7 6.3 6.1 0.3 0.5 3.0 3.3 1.8
MZUSP147478 Holotype Male 15.7 2.8 4.5 1.3 2.8 1.1 1.3 6.6 6.5 6.9 0.4 0.4 3.2 3.5 2.3
MZUSP147494 Paratype Male 16.6 3.1 4.6 1.4 2.6 1.2 1.4 7.0 6.9 7.6 0.3 0.4 3.3 3.8 2.4
MZUSP147479 Paratype Female 17.2 2.9 4.6 1.3 2.9 1.3 1.4 6.7 6.6 6.8 0.3 0.4 3.3 3.5 2.3

Abbreviations: SVL = snout-vent length; HDL = hand length; HDL4 = hand length from the base of the thenar tubercle to the tip of the fourth finger; HL = head length; HW = head width; ED = eye diameter; IOD = inter-orbital distance; IND = inter-nostril distance; END = eye-nostril distance; THL = thigh length; TBL = tibia length; FAL = forearm length; FL = foot length; 3FD = diameter of third finger disk; 4TD = diameter of fourth toe disk. For tissues numbers see Appendix 2.


The specific epithet quilombola refers to people who inhabit quilombo communities. Historically, quilombos were communities constituted by and used as refuges for escaped slaves between 1530 and 1815 during colonial Portuguese rule in Brazil. Nowadays in the north of Espírito Santo Estate quilombola communities still remain and maintain alive their traditions, such as quilombola food and craftwork. This species' name is indeclinable.


Chiasmocleis quilombola sp. n. is known from localities between the Doce River and the Mucuri River, e.g., Floresta Nacional do Rio Preto and Parque Estadual de Itaúnas, Municipality of Conceição da Barra; Reserva Biológica Córrego Veado, Municipality of Pinheiros; Reserva Natural, Reserva Biológica de Sooretama, and Cocoa plantations in Povoação, Municipality of Linhares (Appendix 2). The populations assigned to Chiasmocleis lacrimae and Chiasmocleis capixaba at northernmost of Espírito Santo State are allocated to the new taxon Chiasmocleis quilombola sp. n. (Figure 1).

Natural history.

Chiasmocleis quilombola sp. n. was collected in pitfall traps after heavy rains at Floresta Nacional do Rio Preto (Figure 6). The lines of pitfalls were installed at the vicinity of a permanent lagoon and a temporary swamp. The Floresta Nacional do Rio Preto has 2, 830 ha and an elevation between five to 50 m above the sea level. The soil is typical of coastal areas, mostly sand. The area consists of secondary forested areas and plantations with few remnants of primary Atlantic Forest.

Figure 6.

Larissa Gaigher and Dr. Yuri Leite inspecting pitfall traps installed at the type locality of Chiasmocleis quilombola sp. n., Floresta Nacional do Rio Preto, Municipality of Conceição da Barra, Espírito Santo State, Brazil.


Chiasmocleis quilombola sp. n. has been misidentified as Chiasmocleis lacrimae and Chiasmocleis capixaba due to an overlap in feet webbing and body size (e.g., Tonini et al. 2013, see below). Chiasmocleis quilombola sp. n. corresponds to clade N2, Chiasmocleis sp. to clade N1, Chiasmocleis capixaba to central clade, and Chiasmocleis lacrimae to southern clades of Tonini et al. (2013). Our morphological observations and comparisons with other species combined with molecular information support Chiasmocleis quilombola sp. n. and Chiasmocleis sp. as separate evolutionary lineages (see below and Figure 1, 2). Low levels of genetic divergence in the BDNF (Table 3) between Chiasmocleis quilombola sp. n. and closely related species is consistent with recent cladogenetic events and supports a previous study that estimated initial speciation within this clade (i.e., Chiasmocleis lacrimae, Chiasmocleis capixaba, Chiasmocleis quilombola sp. n., and Chiasmocleis sp.) during the Miocene/Pliocene (Tonini et al. 2013). Chiasmocleis quilombola sp. n. and Chiasmocleis sp. corresponds to an earlier lineage split dated to approximately the Pliocene/Pleistocene (Tonini et al. 2013).

Table 3.

Genetic distance (p-uncorrected) in the BDNF (upper-right) and in the ND2 (lower-left) among Chiasmocleis quilombola sp. n. and sister species. Values at the diagonal correspond to the genetic distance within species in the ND2.

Species Chiasmocleis capixaba Chiasmocleis lacrimae Chiasmocleis cordeiroi Chiasmocleis crucis Chiasmocleis quilombola sp. n. Chiasmocleis schubarti Chiasmocleis sp.
Chiasmocleis capixaba 0.015 0.001 0.008 0.008 0.001 0.007 0.003
Chiasmocleis lacrimae 0.064 0.044 0.008 0.008 0.001 0.007 0.003
Chiasmocleis cordeiroi 0.211 0.227 0.013 0.005 0.008 0.004 0.007
Chiasmocleis crucis 0.182 0.198 0.108 0.015 0.008 0.005 0.007
Chiasmocleis quilombola sp. n. 0.071 0.082 0.217 0.209 0.006 0.007 0.004
Chiasmocleis schubarti 0.206 0.204 0.107 0.103 0.22 0.017 0.007
Chiasmocleis sp. 0.098 0.104 0.253 0.237 0.083 0.236 0.023

Chiasmocleis quilombola sp. n. is distinct from Chiasmocleis schubarti (species with which occurs in sympatry) in having smaller snout-vent length, feet slightly webbed, cream ventral surface, and marbled light brown and cream dorsolateral pattern instead of larger snout-vent length, absence of feet webbing and belly pattern roughly marbled in dark brown and light cream in Chiasmocleis schubarti (Cruz et al. 1997). Chiasmocleis quilombola sp. n. is most similar to Chiasmocleis lacrimae and Chiasmocleis capixaba, species with which it has been previously confused (e.g. Tonini et al. 2013). However, the new species is distinguished from closely relatives by the following set of characters: 1) a smaller body size, shorter head length, shorter thigh and tibia compared to Chiasmocleis lacrimae, Chiasmocleis capixaba, and Chiasmocleis sp. (Table 4), 2) smaller eye diameter, inter-orbital distance, inter-nostril distance, diameter of third finger disk, and diameter of fourth toe disk than Chiasmocleis capixaba, 3) smaller eye-nostril distance, feet length, hand length, and hand length to the tip of the fourth finger than Chiasmocleis lacrimae. Moreover, males of Chiasmocleis quilombola sp. n. have less webbing on the foot (more extensive web on the foot in Chiasmocleis capixaba, absent in Chiasmocleis sp., and ranging from little to absent in Chiasmocleis lacrimae; Cruz et al. 1997, Peloso et al. 2014, Forlani et al. submitted). The new species has slender arms, legs, finger, and toes (robust arms and legs in Chiasmocleis lacrimae; thick fingers and toes in Chiasmocleis capixaba), as well as smaller and less abundant dermal spines in males (spines larger and abundant in Chiasmocleis lacrimae; abundant in Chiasmocleis capixaba). Males of Chiasmocleis quilombola sp. n. posses less amount of fringes between fingers II and III and a slender third finger than males Chiasmocleis capixaba.

Table 4.

Differences between Chiasmocleis capixaba, Chiasmocleis lacrimae, Chiasmocleis quilombola sp. n., and Chiasmocleis sp.

Species Chiasmocleis capixaba Chiasmocleis lacrimae Chiasmocleis quilombola sp. n. Chiasmocleis sp.
Chiasmocleis capixaba SVL=15.1 (SD 0.6)
HL=2.8 (SD 0.1)
THL=6.2 (SD 0.3)
TBL=6.1 (SD 0.3)
Feet webbing head length; thickness of limbs, fingers, and toes; dermal spines Feet webbing; thickness of limbs; dermal spines
Chiasmocleis lacrimae mtDNA (ND2: 6.4%, 16S: 1.3%, 12S: 1.8%) nuDNA (BNDF: haplotype sharing) SVL=16.1 (SD 0.9)
HL=3.3 (SD 0.2)
THL=6.6 (SD 0.4)
TBL=6.5 (SD 0.3)
head and limb length; feet webbing; thickness of limbs; dermal spines body size; thickness; dermal spines
Chiasmocleis quilombola sp. n. mtDNA (ND2: 7.1%, 16S: 0.8%, 12S: 1.1%) nuDNA (BNDF: haplotype sharing) mtDNA (ND2: 8.2%, 16S: 0.8%, 12S: 1.8%) nuDNA (BNDF: haplotype sharing) SVL=14 (SD 1.4)
HL=2.6 (SD 0.2)
THL=5.8 (SD 0.7)
TBL=5.6 (SD 0.6)
Feet webbing; thickness of limbs
Chiasmocleis sp. mtDNA (ND2: 9.8%, 16S: 2.2%, 12S: 1.2%) nuDNA (BNDF: no haplotype sharing) mtDNA (ND2: 10%, 16S: 2.3%, 12S: 2.1%) nuDNA (BNDF: no haplotype sharing) mtDNA (ND2: 8.3%, 16S: 1.6%, 12S: 2.1%) nuDNA (BNDF: haplotype sharing) SVL=15.3 (SD 0.7)
HL=3.4 (SD 0.1)
THL=6 (SD 0.3)
TBL=6.2 (SD 0.3)

SVL snout-vent length; HL head length; THL tight length; TBL tibia length; SD standard deviation.

Chiasmocleis quilombola sp. n. are distinguished from other Chiasmocleis species by: 1) four externally evident fingers and five toes distinguishes it from Chiasmocleis antenori, Chiasmocleis carvalhoi, and Chiasmocleis tridactyla (digit reduction; Walker 1973, Nelson 1975, Duellman and Mendelson 1995); 2) a shorter snout-vent length differentiate it from Chiasmocleis alagoanus, Chiasmocleis albopunctata, Chiasmocleis anatipes, Chiasmocleis atlantica, Chiasmocleis avilapiresae, Chiasmocleis bassleri, Chiasmocleis centralis, Chiasmocleis cordeiroi, Chiasmocleis crucis, Chiasmocleis devriesi, Chiasmocleis hudsoni, Chiasmocleis leucosticta, Chiasmocleis magnova, Chiasmocleis mehelyi, Chiasmocleis papachibe, Chiasmocleis royi, Chiasmocleis sapiranga, Chiasmocleis shudikarensis, Chiasmocleis supercilialba, and Chiasmocleis ventrimaculata (larger snout-vent length; Dunn 1949, Bokermann 1952, Walker and Duellman 1974, Caramaschi and Pimenta 2003, Cruz et al. 1997, Cruz et al. 1999, Caramaschi and Cruz 1997, Cruz et al. 2007a, Moravec and Köhler 2007, Peloso and Sturaro 2008, Funk and Cannatella 2009, Morales and McDiarmid 2009, Peloso et al. 2014); 3) small feet webbing of males and females distinguish the new species from Chiasmocleis cordeiroi, Chiasmocleis leucosticta, Chiasmocleis mantiqueira, and Chiasmocleis sapiranga (more extensive webbed feet in males and females; Cruz et al. 1997, Cruz et al. 2007a, b); 4) a light cream belly pattern without dark spots distinguished it from Chiasmocleis alagoanus, C. atlantica, Chiasmocleis haddadi, Chiasmocleis leucosticta, and Chiasmocleis mantiqueira (belly pattern roughly marbled in dark brown and pale cream, Cruz et al.1997, Cruz et al.1999, Cruz et al. 2007b, Peloso et al. 2014); and 5) snout rounded and belly light cream colored differentiate it from Chiasmocleis gnoma (snout truncate and belly boldly marbled in brown and pale cream; Canedo et al. 2004).

Chiasmocleis quilombola sp. n. occurs in sympatry with Chiasmocleis schubarti at the Floresta Nacional do Rio Preto, Municipality of Conceição da Barra, and at the Reserva Biológica Córrego Veado, Municipality of Pinheiros; it also occurs with Chiasmocleis capixaba and Chiasmocleis schubarti at the Reserva Natural Vale, Reserva Biológica de Sooretama, and at Cocoa plantations in Povoação, sites in the Municipality of Linhares. The new species is allopatric to Chiasmocleis sp. and Chiasmocleis lacrimae (Figure 1). We did not have access to tissues samples of Chiasmocleis capixaba from Nova Viçosa, Bahia State (Van Sluys 1998), to include in the genetic analysis, thus the phylogenetic relationship of this population remains unclear.

Cryptic species have challenged our ability to assess current levels of biodiversity. Anuran taxonomy has used various data sources to describe the species diversity, e.g., advertisement calls, external morphology, osteology, tadpoles, ecology, molecular data, karyotypes (Duellman and Trueb 1986, Haas 2003). However, Chiasmocleis systematics has been based on external morphology from adults and behavioral information (Wogel et al. 2004, Hartmann et al. 2002, Nascimento and Skuk 2006, Oliveira Filho and Giaretta 2006, Langone et al. 2007, Peloso and Sturaro 2008, Rodrigues et al. 2008, Santana et al. 2012, but see Peloso et al. 2014). Sexual dimorphism in size, amount of webbing, and color pattern have been useful characters to diagnose species (Cruz et al. 1997). Recent molecular studies 1) demonstrated the non-monophyly of traditionally recognized species groups (de Sá et al. 2012, Peloso et al. 2014) and 2) reported high genetic divergences and low gene flow along small geographical scales, suggesting that some populations could represent new species (Tonini et al. 2013). Given the current overall biodiversity crisis and specifically the worldwide threats to amphibian biodiversity, molecular studies should move beyond the identification of genetic clades and should make every effort to formally describe those evolutionary lineages. Herein, we have taken this approach and described a new species based on a combination of morphological characters in a clade of cryptic species with shown high genetic diversity and low gene flow.

The new species occupy coastal areas North of Espírito Santo State, a region that is under strong human pressure. Therefore, marine and coastal communities are susceptible to impacts of proposed modifications in the landscape for the exploitation of mineral resources. In this context, Chiasmocleis quilombola sp. n. may face imminent threat of habitat loss, as consequence of the deforestation and intensive occupation of the space by human activities.


We are thankful to L. Costa, Y. Leite, and the Laboratório de Mastozoologia e Biogeografia team, for field assistance and tissues samples. L. Chagas and staffs of the Floresta Nacional do Rio Preto for valuable help during field surveys. M. T. Rodrigues, R. C. Amaro, C. F. B. Haddad, P. Rocha, M. Napoli, H. Zaher, M. Solé, V. Fagundes, J. L. Gasparini, J. P. Pombal Jr., and P. Passos for providing tissue samples and permission to examine specimens under their care. D. Baêta and A. C. Calijorne for hosting JFRT during visit to the Museu Nacional do Rio de Janeiro. R. Ferreira, J. L. Gasparini, M. Vences, R. A. Pyron, F. Andreone, and two anonymous referees for suggestions on the manuscript. JFRT acknowledge support by the Science without Borders program (CAPES/Brazil), The George Washington University, and award NSF-DEB 1144692 to R. O. de Sá.

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Appendix 1
Table 5.

Examined material and measurements from males included in the morphological analyses.

Chiasmocleis Numbers Locality State SVL HL HW ED IOD IND END THL TBL FL 3FD 4TD FAL HDL HDL4
Chiasmocleis capixaba MNRJ17514 Aracruz ES 14.77 2.91 4.04 1.18 2.57 1.16 1.21 6.19 6.34 6.43 0.35 0.44 2.98 3.36 2.08
Chiasmocleis capixaba MNRJ17515 Aracruz ES 13.98 2.84 4.44 1.28 2.71 1.21 1.39 6.17 6.13 6.58 0.29 0.46 3.07 3.28 1.94
Chiasmocleis capixaba MNRJ17516 Aracruz ES 14.47 2.93 3.98 1.18 2.64 1.19 1.32 6.08 5.96 6.64 0.30 0.42 3.10 3.36 2.16
Chiasmocleis capixaba MNRJ17517 Aracruz ES 14.70 3.00 4.10 1.36 2.61 1.29 1.26 6.23 6.10 6.74 0.33 0.51 3.15 3.55 2.32
Chiasmocleis capixaba MNRJ17518 Aracruz ES 16.02 3.13 4.39 1.32 2.64 1.21 1.45 6.61 6.80 7.45 0.38 0.48 3.62 3.86 2.33
Chiasmocleis capixaba MNRJ17519 Aracruz ES 15.40 2.99 3.95 1.36 2.63 1.23 1.56 6.05 6.31 6.28 0.33 0.45 3.38 3.17 2.09
Chiasmocleis capixaba MNRJ17520 Aracruz ES 14.94 2.76 3.82 1.26 2.66 1.20 1.36 5.71 6.06 5.90 0.41 0.47 2.96 3.15 1.80
Chiasmocleis capixaba MNRJ17535 Aracruz ES 15.23 2.96 3.83 0.97 2.39 1.18 1.07 6.18 5.76 5.97 0.38 0.42 2.76 3.05 2.13
Chiasmocleis capixaba MNRJ17536 Aracruz ES 15.24 2.50 3.92 1.23 2.60 1.16 1.41 5.83 5.79 6.63 0.33 0.45 2.93 3.52 2.19
Chiasmocleis capixaba MNRJ17895 Aracruz ES 15.46 2.89 4.32 1.12 2.56 1.22 1.21 6.62 6.30 6.46 0.33 0.47 3.25 3.34 2.15
Chiasmocleis capixaba MNRJ22962 Reserva Natural Vale ES 14.83 3.02 4.44 1.26 2.67 1.18 1.38 6.29 6.08 6.37 0.36 0.44 3.12 3.34 2.08
Chiasmocleis capixaba MNRJ22966 Reserva Natural Vale ES 15.05 2.99 4.41 1.37 2.71 1.19 1.43 6.71 6.20 6.25 0.36 0.52 3.18 3.15 1.72
Chiasmocleis capixaba MZUSP147468 ReBio Duas Bocas ES 15.88 3.05 4.40 1.27 2.87 1.25 1.41 6.84 6.53 6.92 0.43 0.43 3.06 3.44 2.14
Chiasmocleis capixaba MZUSP147469 ReBio Duas Bocas ES 16.23 3.02 4.61 1.05 2.99 1.31 1.68 6.80 6.71 7.05 0.48 0.48 3.28 3.29 2.30
Chiasmocleis lacrimae MNRJ17480 Horto Florestal RJ 16.20 3.40 4.40 1.10 2.70 0.90 1.40 6.50 6.50 7.30 0.30 0.40 3.30 3.60 2.50
Chiasmocleis lacrimae MNRJ17481 Horto Florestal RJ 15.60 3.30 4.10 1.10 2.60 1.00 1.40 6.20 6.30 7.20 0.40 0.40 3.50 3.60 2.60
Chiasmocleis lacrimae MNRJ17482 Horto Florestal RJ 16.20 3.50 2.60 1.30 2.60 1.00 1.90 6.50 6.50 7.10 0.40 0.50 3.30 3.80 2.50
Chiasmocleis lacrimae MNRJ17484 Horto Florestal RJ 19.15 3.57 4.82 1.35 2.84 1.44 1.69 7.45 7.35 8.23 0.34 0.36 3.56 4.23 2.69
Chiasmocleis lacrimae MNRJ17485 Horto Florestal RJ 16.10 3.70 4.30 1.20 2.50 0.90 1.60 6.90 6.70 7.50 0.30 0.40 3.40 3.60 2.40
Chiasmocleis lacrimae MNRJ17486 Horto Florestal RJ 15.07 3.00 4.33 1.22 2.52 1.22 1.50 6.30 6.22 6.68 0.30 0.43 3.19 3.44 2.24
Chiasmocleis lacrimae MNRJ17487 Horto Florestal RJ 16.10 3.33 4.38 1.19 2.91 1.30 1.68 6.94 6.78 7.16 0.27 0.39 3.46 3.53 2.08
Chiasmocleis lacrimae MNRJ17488 Horto Florestal RJ 15.90 3.20 3.80 1.10 2.50 0.90 1.30 5.80 6.20 6.60 0.30 0.40 3.00 3.30 2.20
Chiasmocleis lacrimae MNRJ17489 Horto Florestal RJ 15.40 3.10 3.90 1.20 2.20 0.90 1.30 6.10 6.10 6.90 0.30 0.40 3.10 3.40 2.10
Chiasmocleis lacrimae MNRJ17490 Horto Florestal RJ 16.10 3.30 4.00 1.10 2.50 0.90 1.40 6.70 7.10 7.70 0.40 0.40 3.50 4.20 2.70
Chiasmocleis lacrimae MNRJ17491 Horto Florestal RJ 15.26 2.90 4.75 1.21 2.92 1.47 1.49 6.59 6.46 7.13 0.32 0.41 3.34 3.78 2.41
Chiasmocleis lacrimae MNRJ17492 Horto Florestal RJ 15.50 3.90 4.60 1.30 2.50 1.20 1.50 6.10 6.30 6.50 0.40 0.40 3.10 3.70 2.50
Chiasmocleis lacrimae MNRJ17498 Horto Florestal RJ 17.10 3.70 4.70 1.20 2.50 1.10 1.40 6.70 6.80 6.90 0.40 0.50 3.40 3.40 2.50
Chiasmocleis lacrimae MNRJ17505 Horto Florestal RJ 16.80 3.42 4.62 1.41 2.84 1.32 1.71 7.19 6.88 7.43 0.33 0.38 3.50 3.87 2.42
Chiasmocleis lacrimae MNRJ17506 Horto Florestal RJ 16.95 3.30 4.84 1.36 3.00 1.35 1.62 7.19 6.80 7.32 0.41 0.43 3.60 4.09 2.63
Chiasmocleis lacrimae MNRJ17507 Horto Florestal RJ 15.16 3.14 4.43 1.29 2.71 1.24 1.45 6.06 6.13 6.28 0.34 0.41 3.12 3.51 2.22
Chiasmocleis lacrimae MNRJ17565 Horto Florestal RJ 16.66 2.97 4.42 1.36 2.79 1.30 1.47 7.05 6.69 7.14 0.49 0.43 3.23 3.97 2.63
Chiasmocleis lacrimae MNRJ66497 Mimoso do Sul ES 16.60 3.14 4.77 1.40 2.89 1.31 1.48 6.69 6.49 6.90 0.39 0.43 3.55 3.80 2.32
Chiasmocleis quilombola sp. n. MNRJ29057 Povoação ES 12.72 2.70 3.89 1.05 2.42 1.08 1.20 5.35 5.05 5.24 0.30 0.39 2.65 2.79 1.79
Chiasmocleis quilombola sp. n. MNRJ29058 Povoação ES 13.23 2.68 3.90 1.12 2.47 1.10 1.12 5.28 4.91 5.17 0.34 0.42 2.52 2.78 1.73
Chiasmocleis quilombola sp. n. MNRJ29059 Povoação ES 13.56 2.53 3.63 1.15 2.37 1.08 1.12 5.36 5.25 5.33 0.29 0.39 2.80 3.00 1.89
Chiasmocleis quilombola sp. n. MNRJ29060 Povoação ES 12.59 2.55 3.79 1.08 2.41 1.08 1.16 5.12 4.67 4.56 0.32 0.32 2.48 2.48 1.67
Chiasmocleis quilombola sp. n. MNRJ29073 Povoação ES 13.76 2.53 3.94 1.25 2.43 1.18 1.29 5.67 5.63 5.88 0.41 0.41 2.88 3.07 1.85
Chiasmocleis quilombola sp. n. MNRJ29074 Povoação ES 13.47 2.33 3.72 1.18 2.27 1.08 1.15 5.24 5.27 5.60 0.32 0.40 2.73 3.02 1.81
Chiasmocleis quilombola sp. n. MBML2858 Povoação ES 13.47 2.54 3.94 0.88 2.04 0.82 1.03 5.11 5.39 5.50 0.33 0.43 2.61 2.91 1.72
Chiasmocleis quilombola sp. n. MBML2866 Povoação ES 12.15 2.24 3.34 1.05 2.16 0.72 0.95 5.26 5.11 5.33 0.27 0.39 2.55 2.75 1.62
Chiasmocleis quilombola sp. n. MBML2863 Povoação ES 12.30 2.34 3.75 0.82 2.00 0.83 0.93 5.22 5.32 5.42 0.31 0.48 2.50 2.63 1.59
Chiasmocleis quilombola sp. n. MZUSP147473 Flona Rio Preto ES 16.07 3.18 3.87 1.09 2.18 1.09 1.33 6.01 6.00 6.41 0.24 0.23 3.15 3.30 2.07
Chiasmocleis quilombola sp. n. MZUSP147478 Flona Rio Preto ES 15.71 2.75 4.52 1.30 2.80 1.13 1.25 6.55 6.47 6.91 0.35 0.44 3.21 3.45 2.33
Chiasmocleis quilombola sp. n. MZUSP147471 Flona Rio Preto ES 14.43 2.75 3.83 1.03 2.57 1.11 1.51 6.07 6.27 6.23 0.25 0.26 2.99 2.84 1.83
Chiasmocleis quilombola sp. n. MZUSP147475 Flona Rio Preto ES 14.48 2.79 4.55 1.28 2.55 1.28 1.25 6.63 6.41 6.73 0.36 0.36 3.26 3.48 2.16
Chiasmocleis quilombola sp. n. MZUSP147494 Flona Rio Preto ES 16.55 3.14 4.63 1.36 2.55 1.24 1.36 7.02 6.89 7.59 0.25 0.37 3.33 3.80 2.40
Chiasmocleis quilombola sp. n. MZUSP147476 Flona Rio Preto ES 16.09 2.70 4.36 1.24 2.65 1.31 1.36 6.73 6.26 6.11 0.32 0.47 2.95 3.33 1.83
Chiasmocleis quilombola sp. n. MZUSP147472 Flona Rio Preto ES 15.28 2.80 4.08 1.33 2.67 1.24 1.33 7.08 6.55 6.60 0.34 0.41 3.20 3.40 2.22
sp. MTR13495 Trancoso BA 15.92 3.39 4.07 1.06 2.65 0.94 1.43 6.37 6.16 6.77 0.27 0.42 3.60 3.45 2.06
sp. MTR13547 Trancoso BA 13.80 3.53 4.09 0.97 2.36 1.00 1.35 5.59 5.68 5.72 0.27 0.35 2.95 3.18 2.27
sp. MTR13545 Trancoso BA 15.07 3.51 4.24 1.10 2.43 1.03 1.35 5.91 6.41 6.78 0.29 0.38 2.95 3.40 2.35
sp. MTR13590 Trancoso BA 15.48 3.48 3.99 1.17 2.51 0.80 1.32 6.01 6.22 6.69 0.30 0.45 3.11 3.73 2.41
sp. MTR13565 Trancoso BA 16.27 3.78 4.31 0.95 2.59 1.04 1.61 5.64 6.33 6.56 0.33 0.46 3.20 3.68 2.67
sp. MTR13548 Trancoso BA 15.36 3.34 4.57 1.06 2.52 1.12 1.09 6.44 6.62 6.72 0.30 0.40 3.19 3.64 2.36
sp. MTR13546 Trancoso BA 15.53 3.10 4.40 1.24 2.30 0.91 1.46 6.02 5.96 6.56 0.31 0.40 2.90 3.30 2.14
sp. MTR13489 Trancoso BA 15.34 3.53 4.05 1.16 2.35 1.00 1.36 6.67 6.54 6.73 0.27 0.44 3.40 3.46 2.40
Appendix 2
Table 6.

Samples included in the molecular analyses and genbank numbers. Numbers in bold were retrieved from previous studies.

Chiasmocleis Voucher Tissue Locality State 12S 16S ND2 BDNF
Chiasmocleis capixaba MZUSP147497 CTA1861 ReBio Duas Bocas ES KM111721 KM111817 JQ410706 KM111908
Chiasmocleis capixaba MZUSP147498 CTA1863 ReBio Duas Bocas ES KM111722 KM111818 JQ410707 KM111909
Chiasmocleis capixaba MZUSP147499 CTA1864 ReBio Duas Bocas ES KM111723 KM111819 JQ410708 KM111910
Chiasmocleis capixaba MZUSP147468 CTA1865 ReBio Duas Bocas ES KM111724 KM111820 KM111992 KM111911
Chiasmocleis capixaba MZUSP147482 CTA1869 ReBio Duas Bocas ES KM111725 KM111821 KM111993 KM111912
Chiasmocleis capixaba MZUSP147469 CTA1870 ReBio Duas Bocas ES KM111726 KM111822 KM111994 KM111913
Chiasmocleis capixaba MZUSP147500 CTA1871 ReBio Duas Bocas ES KM111727 KM111823 JQ410709 KM111914
Chiasmocleis capixaba MZUSP147510 CTA1872 Serra ES KM111728 KM111824 JQ410685 KM111915
Chiasmocleis capixaba MZUSP147512 CTA1874 Serra ES KM111729 KM111825 JQ410690 KM111916
Chiasmocleis capixaba MZUSP147513 CTA1875 Serra ES KM111730 KM111826 JQ410691 KM111917
Chiasmocleis capixaba JFT479 CTA1876 Serra ES KM111731 KM111827 JQ410692 KM111918
Chiasmocleis capixaba MZUSP147514 CTA1877 Serra ES KM111732 KM111828 JQ410693 KM111919
Chiasmocleis capixaba JFT483 CTA1879 Serra ES KM111733 KM111829 JQ410694 KM111920
Chiasmocleis capixaba JFT499 CTA1881 Serra ES KM111734 KM111830 JQ410695 KM111921
Chiasmocleis capixaba MZUSP147520 CTA1893 Serra ES KM111735 KM111831 JQ410700 KM111922
Chiasmocleis capixaba MZUSP147521 CTA1894 Serra ES KM111736 KM111832 JQ410701 KM111923
Chiasmocleis capixaba MZUSP147523 CTA1896 Serra ES KM111737 KM111833 JQ410702 KM111924
Chiasmocleis capixaba MTR12276 CTMZ6907 FloNa dos Goytacazes ES KM111738 KM111834 - KM111925
Chiasmocleis capixaba MTR12296 CTMZ6908 FloNa dos Goytacazes ES KM111739 KM111835 JQ410688 KM111926
Chiasmocleis capixaba MTR12407 CTMZ6912 Reserva Natural Vale ES KM111740 KM111836 - -
Chiasmocleis capixaba MTR12484 CTMZ6920 Reserva Natural Vale ES KM111741 KM111837 - KM111927
Chiasmocleis capixaba MTR12485 CTMZ6921 Reserva Natural Vale ES KM111742 KM111838 - KM111928
Chiasmocleis capixaba - CTMZ6939 Guarapari ES KM111743 KM111839 - KM111929
Chiasmocleis capixaba MTR12076 MTR12076 Reserva Natural Vale ES KM111744 KM111840 JQ410687 KM111930
Chiasmocleis capixaba MTR12297 MTR12297 FloNa dos Goytacazes ES KM111745 KM111841 JQ410689 KM111931
Chiasmocleis cordeiroi CFBH32057 CFBH15784 Ilhéus BA KM111759 KM111852 KM111995 KM111939
Chiasmocleis cordeiroi MZUSP147496 CTA1935 Ituberá BA KM111760 KM111853 KM111996 KM111940
Chiasmocleis cordeiroi MTR22122 MTR22122 EE Wenceslau Guimarães BA KM111761 KM111854 KM111997 KM111941
Chiasmocleis cordeiroi MTR22123 MTR22123 EE Wenceslau Guimarães BA KM111762 KM111855 KM111998 KM111942
Chiasmocleis cordeiroi PEU137 PEU137 Jaguaripe BA KM111763 KM111856 KM111999 KM111943
Chiasmocleis cordeiroi PEU146 PEU146 Jaguaripe BA KM111764 KM111857 KM112000 KM111944
Chiasmocleis crucis MTR6001 CTMZ6898 Serra do Teimoso BA KM111765 KM111858 KM112001 KM111945
Chiasmocleis crucis - CTMZ6900 Ilhéus BA KM111766 KM111859 KM112002 KM111946
Chiasmocleis crucis - CTMZ6901 Ilhéus BA KM111767 KM111860 KM112003 KM111947
Chiasmocleis crucis MTR16070 MTR16070 Serra Bonita BA KM111768 KM111861 KM112004 KM111948
Chiasmocleis lacrimae - CFBH73 Picinguaba SP KM111748 KC180040 JQ410715 KC180202
Chiasmocleis lacrimae CFBH17495 CFBH7361 Ilha de São Sebastião SP KM111749 KM111844 -
Chiasmocleis lacrimae - CFBH76 Picinguaba SP KM111750 KC180063 JQ410714 KC180163
Chiasmocleis lacrimae JFT981 CTA1934 Matada Usina Paineiras ES KM111751 KM111845 JQ410710 KM111932
Chiasmocleis lacrimae - CTMZ6946 Bertioga SP KM111752 KM111846 - KM111933
Chiasmocleis lacrimae RN7003 CTRN173 Angra dos Reis RJ KM111753 KM111847 - -
Chiasmocleis lacrimae RN7004 CTRN174 Angra dos Reis RJ KM111754 KM111848 - KM111934
Chiasmocleis lacrimae RN7005 CTRN175 Angra dos Reis RJ KM111755 KM111849 - KM111935
Chiasmocleis lacrimae MNRJ47477 MNRJ47477 ReBio União RJ KM111746 KM111842 - -
Chiasmocleis lacrimae MNRJ48415 MNRJ48415 ReBio União RJ KM111747 KM111843 - -
Chiasmocleis lacrimae MNRJ49302 MNRJ49302 Cachoeiras de Macacu ES KM111756 KM111850 JQ410712 KM111936
Chiasmocleis lacrimae MNRJ60744 MNRJ60744 Duque de Caxias RJ KM111757 - JQ410713 KM111937
Chiasmocleis lacrimae MNRJ66494 MNRJ66494 Mimoso do Sul ES KM111758 KM111851 JQ410711 KM111938
Chiasmocleis leucosticta CFBH19029 CFBH8594 PE Ilha do Cardoso SP KM111769 KM111862 - -
Chiasmocleis leucosticta MZUSP136053 CTMZ2485 PE Carlos Botelho SP KM111770 KM111863 - -
Chiasmocleis leucosticta MZUSP136055 CTMZ2493 PE Carlos Botelho SP KM111771 - - KM111949
Chiasmocleis leucosticta MZUSP136059 CTMZ2497 PE Carlos Botelho SP - KM111864 - KM111950
Chiasmocleis leucosticta MTR7128 CTMZ6943 Fazenda Intervales SP - - KM112005 KM111951
Chiasmocleis leucosticta - CTMZ6944 Piedade SP KM111772 - KM112006 -
Chiasmocleis mantiqueira - CTMZ6891 Serra do Brigadeiro MG KM111773 KM111865 - KM111952
Chiasmocleis mantiqueira UFMG-A9643 UFMG-T1802 Ouro Branco MG KM111774 KM111866 KM112007 -
Chiasmocleis mantiqueira UFMG-A9659 UFMG-T1804 Ouro Branco MG KM111775 KM111867 - KM111953
Chiasmocleis mantiqueira UFMG-A9651 UFMG-T1810 Ouro Branco MG KM111776 KM111868 - KM111954
Chiasmocleis mantiqueira UFMG-A9656 UFMG-T1815 Ouro Branco MG KM111777 KM111869 - KM111955
Chiasmocleis quilombola sp. n. - CFBH1437 ReBio Sooretama ES KM111778 KC180044 - KC180193
Chiasmocleis quilombola sp. n. - CFBH1438 ReBio Sooretama ES KM111779 KC179977 - KC180168
Chiasmocleis quilombola sp. n. CFBH19471 CFBH9055 Povoação ES KM111780 KM111870 KM112008 KM111956
Chiasmocleis quilombola sp. n. CFBH18076 CFBH9082 Povoação ES KM111781 KM111871 KM112009 KM111957
Chiasmocleis quilombola sp. n. CFBH18077 CFBH9083 Povoação ES KM111782 KM111872 KM112010 KM111958
Chiasmocleis quilombola sp. n. JFT831 CTA1906 FloNa do Rio Preto ES KM111783 KM111873 JQ410669 KM111959
Chiasmocleis quilombola sp. n. MZUSP147471 CTA1907 FloNa do Rio Preto ES KM111784 KM111874 JQ410670 KM111960
Chiasmocleis quilombola sp. n. MZUSP147472 CTA1908 FloNa do Rio Preto ES KM111785 KM111875 JQ410671 KM111961
Chiasmocleis quilombola sp. n. MZUSP147473 CTA1909 FloNa do Rio Preto ES KM111786 KM111876 JQ410672 KM111962
Chiasmocleis quilombola sp. n. MZUSP147474 CTA1918 FloNa do Rio Preto ES KM111787 KM111877 JQ410673 KM111963
Chiasmocleis quilombola sp. n. MZUSP147475 CTA1919 FloNa do Rio Preto ES KM111788 KM111878 JQ410674 KM111964
Chiasmocleis quilombola sp. n. MZUSP147494 CTA1923 FloNa do Rio Preto ES KM111789 KM111879 JQ410677 KM111965
Chiasmocleis quilombola sp. n. MZUSP147479 CTA1929 FloNa do Rio Preto ES KM111790 KM111880 JQ410679 KM111966
Chiasmocleis quilombola sp. n. MZUSP147480 CTA1931 FloNa do Rio Preto ES KM111791 KM111881 JQ410680 KM111967
Chiasmocleis quilombola sp. n. MZUSP147493 CTA1933 FloNa do Rio Preto ES KM111792 KM111882 JQ410681 KM111968
Chiasmocleis quilombola sp. n. JFT990 CTA1938 PE de Itaúnas ES - KM111883 - KM111969
Chiasmocleis quilombola sp. n. MTR12017 CTMZ6903 Reserva Natural Vale ES KM111793 KM111884 - KM111970
Chiasmocleis quilombola sp. n. MTR12077 CTMZ6905 Reserva Natural Vale ES KM111794 KM111885 - -
Chiasmocleis quilombola sp. n. MTR12470 CTMZ6916 Reserva Natural Vale ES KM111795 KM111886 - KM111971
Chiasmocleis quilombola sp. n. MTR12471 CTMZ6917 Reserva Natural Vale ES KM111796 KM111887 - KM111972
Chiasmocleis quilombola sp. n. MTR21527 LGA3267 ReBio Córrego Veado ES KM111797 KM111888 JQ410668 KM111973
Chiasmocleis schubarti CFBH9331 CFBH2078 ReBio Sooretama ES KM111798 KC180071 KM112011 KC180122
Chiasmocleis schubarti CFBH18075 CFBH9060 Povoação ES KM111799 KM111889 KM112012 KM111974
Chiasmocleis schubarti CFBH 22501 CTA1860 ReBio Duas Bocas ES KM111800 KM111890 KM112013 KM111975
Chiasmocleis schubarti MZUSP147485 CTA1887 FloNa do Rio Preto ES KM111801 KM111891 KM112014 KM111976
Chiasmocleis schubarti MZUSP147487 CTA1924 FloNa do Rio Preto ES KM111802 KM111892 KM112015 KM111977
Chiasmocleis schubarti MTR12094 CTMZ6906 FloNa dos Goytacazes ES KM111803 KM111893 KM112016 KM111978
Chiasmocleis schubarti LGA2630 LGA2630 ReBio Córrego Veado ES KM111804 KM111894 JQ410661 KM111979
Chiasmocleis schubarti MTR12266 MTR12266 FloNa dos Goytacazes ES KM111805 KM111895 KM112017 KM111980
Chiasmocleis schubarti MTR17524 MTR17524 PE do Rio Doce MG KM111806 KM111896 KM112018 KM111981
Chiasmocleis schubarti MTR17571 MTR17571 PE do Rio Doce MG KM111807 KM111897 KM112019 KM111982
sp. - CFBH15818 Porto Seguro BA - KM111898 - KM111983
sp. MTR13466 CTMZ6923 Trancoso BA KM111808 KM111899 JQ410665 KM111984
sp. MTR13489 CTMZ6924 Trancoso BA KM111809 KM111900 JQ410666 KM111985
sp. MTR13495 CTMZ6925 Trancoso BA KM111810 KM111901 - KM111986
sp. MTR13545 CTMZ6927 Trancoso BA KM111811 KM111902 - KM111987
sp. MTR13547 CTMZ6929 Trancoso BA KM111812 KM111903 - KM111988
sp. MTR13548 CTMZ6930 Trancoso BA KM111813 KM111904 - KM111989
sp. MTR13565 CTMZ6932 Trancoso BA KM111814 KM111905 - KM111990
sp. MTR13579 CTMZ6933 Trancoso BA KM111815 KM111906 - KM111991
sp. MTR13589 CTMZ6935 Trancoso BA KM111816 KM111907 KM112020 -