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Description and phylogenetic relationships of a new trans-Andean species of

Elachistocleis Parker 1927 (Amphibia, Anura, Microhylidae)
Article in Zootaxa · May 2020

DOI: 10.11646/zootaxa.4779.3.2
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Zootaxa 4779 (3): 323–340 ISSN 1175-5326 (print edition)
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Copyright © 2020 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4779.3.2
http://zoobank.org/urn:lsid:zoobank.org:pub:AD3DDAD2-D595-40FF-85A7-4C47783A617F
Description and phylogenetic relationships of a new trans-Andean species of

Elachistocleis Parker 1927 (Amphibia, Anura, Microhylidae)
JUAN C. SÁNCHEZ-NIVICELA1,2,3, PEDRO L. V. PELOSO4, VERÓNICA L. URGILES2,5,
MARIO H. YÁNEZ-MUÑOZ2,6, YERKA SAGREDO6,7, NADIA PÁEZ6,8 & SANTIAGO RON9*
1
Universidad Nacional de Colombia, Facultad de Ciencias, Bogotá D.C., Colombia; Grupo de Investigación Evolución y Ecología de
Fauna Neotropical. � juan.sanchezn13@gmail.com; https://orcid.org/0000-0001-7971-1216
2
Instituto Nacional de Biodiversidad, calle Rumipamba 341 y Av. de los Shyris, Casilla 17-07-8976, Quito, Ecuador
3
Laboratorio de Herpetología, Museo de Zoología, Escuela de Biología, Ecología y Gestión, Universidad del Azuay, Avenida 24 de
mayo 7-77 y Hernán Malo, Apartado 01-01-981, Cuenca, Ecuador
4
Universidade Federal do Pará, Instituto de Ciências Biológicas, Rua Augusto Correia, 01, Guamá, 66075-110, Belém, PA, Brazil
� pedropeloso@gmail.com; https://orcid.org/0000-0003-0127-8293
5
Department of Biology, University of Central Florida, Orlando, Florida
� vurgiles@knights.ucf.edu; https://orcid.org/0000-0003-4129-7122
6
Museo de Zoología, Departamento de Ciencias Biológicas, Pontificia Universidad Católica del Ecuador, Avenida 12 de octubre 1076
y Roca, Apartado 17-01-2184, Quito, Ecuador. � mayamu@hotmail.com; https://orcid.org/0000-0003-3224-1987
7�
yevasanu@gmail.com; https://orcid.org/0000-0003-3421-5870
8�
nadianuanda@gmail.com; https://orcid.org/0000-0003-0854-3993
9
Museo de Zoología, Departamento de Ciencias Biológicas, Pontificia Universidad Católica del Ecuador, Avenida 12 de octubre 1076
y Roca, Apartado 17-01-2184, Quito, Ecuador. � santiago.r.ron@gmail.com; https://orcid.org/0000-0003-3224-1987
*
Corresponding author
Abstract
Elachistocleis is a Neotropical genus of microhylid frogs with 18 species, most of which occur east of the Andes in South
America. Here, we present a new phylogeny of Gastrophryninae and describe and name a new species of Elachistocleis
from southern Ecuador—the first to be found west of the Andes and also the first from Ecuador. Our phylogeny is based
on DNA sequences of the mitochondrial genes 12S rRNA, 16S rRNA, COI, and the nuclear genes BDNF, cmyc2, H3A,
28S, SIA1, and Tyr. Elachistocleis araios sp. n., is the sister species of all other Elachistocleis. The finding of this taxon
highlights the probability of the existence of more Elachistocleis species west of the Andes.
Key words: Narrow-mouthed frog, Pacific slopes, phylogenetics, systematics, Elachistocleis araios sp. n., Ecuador.
Introduction
Microhylidae is one of the most diverse families of extant amphibians, with almost 690 named species, and dis-
tributed across most of the tropics (Frost 2019). Among New World microhylids, there are three recognized sub-
families: Adelastinae, Gastrophryninae and Otophryninae (de Sá et al. 2012; Peloso et al. 2015). One of the largest
genera of them is Elachistocleis Parker, 1927, which is distributed from Central America to Argentina. Its greatest
diversity occurs mostly in lowland ecosystems (below the 900 m), east of the Andes, with a few species occurring
in northwestern South America (Frost 2019).
Elachistocleis is characterized by the presence of a short clavicle not reaching the coracoids, a notoriously
divided procoracoid, and in having the last three vertebrae longer than wide (de Sá et al. 2012). Elachistocleis is
composed of 18 valid nominal species, E. bicolor (Guérin-Méneville, 1838), E. bumbameuboi Caramaschi, 2010,
E. carvalhoi Caramaschi, 2010, E. cesarii (Miranda-Ribeiro, 1920), E. corumbaensis Piva, Caramaschi & Albu-
querque, 2017, E. erythrogaster Kwet & Di-Bernardo, 1998, E. haroi Pereyra, Akmentins, Laufer & Vaira, 2013, E.
helianneae Caramaschi, 2010, E. magna Toledo, 2010, E. matogrosso Caramaschi, 2010, E. muiraquitan Nunes de
Almeida & Toledo, 2012, E. ovalis (Schneider, 1799), E. panamensis (Dunn, Trapido & Evans, 1948), E. pearsei
Accepted by M. Vences: 22 Apr. 2020; published: 20 May 2020 323

(Ruthven, 1914), E. piauiensis Caramaschi & Jim, 1983, E. skotogaster Lavilla, Vaira, & Ferrari, 2003, E. surina-
mensis (Daudin, 1802), and E. surumu Caramaschi, 2010.
Until now, Elachistocleis has been unknown from the western versant of the Andes. The closest distributional
records are those of E. panamensis which occurs in central Panama, the Magdalena basin in the central Andes of
Colombia, and, marginally, in the Pacific basin in the Chocó region northwest of the Atrato river Panama (Nelson
1972). During fieldwork between 2009 and 2019 in the lowlands of southern Ecuador, west of the Andes, we found
a new species of the genus Elachistocleis. This represents the first record of this genus for Ecuador and also the first
confirmed record for the western slopes of the Andes. Here, we describe the new species and present an expanded
phylogeny of Microhylidae, highlighting the position of the new taxon.
Methods
Morphology
The description of the new species is based on two adult gravid females. Coloration patterns in life were taken
from field notes (by JCS and YS), and photographs (by JCS and SR). Specimens were euthanized with benzocaine
(2%), fixed with formaldehyde (10%) and preserved with ethanol (75%). Morphometric measurements are based
on Duellman (2001) and Peloso et al. (2014). All morphometric variables were measured three times using a digi-
tal caliper to the nearest 0.1 mm. The final value was the average among the three replicates. Abbreviations are as
follows: snout-vent length (SVL); head length (HL); head width (HW); eye diameter (ED); inter-orbital distance
(IOD), between the two medial canthi of each eye; inter-narial distance (IND); eye-nostril distance (END); thigh
length (THL); tibia length (TBL); foot length (FL); hand length from the base of the thenar tubercle to the tip of
finger III (HDL3); hand length from the base of the thenar tubercle to the tip of finger IV (HDL4); forearm length
(FAL); disc diameter of finger III (FD3); disc diameter of toe IV (FT4). Abbreviations for fingers and toes are as
follows: fingers I–IV=FI–IV; toes I–V=TI–V. Specimens examined for comparisons are deposited in the following
collections; Museo de Zoología de la Pontificia Universidad Católica del Ecuador, Quito-Ecuador (QCAZ); Labo-
ratorio de Anfibios del Instituto de Ciencias Naturales de la Universidad Nacional de Colombia (ICN)—a complete
list of examined specimens is listed in APPENDIX I.
Phylogenetic analyses
DNA extraction, amplification, and sequencing
To assess the evolutionary affinities of the new species, we sequenced DNA fragments of three mitochondrial
and one nuclear gene. DNA was extracted from muscle or liver tissue preserved in 95% ethanol using a standard
phenol–chloroform extraction protocol (Sambrook et al. 1989). We used a polymerase chain reaction (PCR) to
amplify DNA fragments for mitochondrial genes 12S rRNA (12S), 16S rRNA (16S), Cytochrome Oxidase I (CO1),
and the nuclear gene Cellular Myelocytomatosis Oncogene - exon 2 (CMYC). PCR amplifications were performed
under standard protocols and sequenced by the Macrogen Sequencing Team (Macrogen Inc., Seoul, South Korea).
Primers were 16Sc-16L, 16Sbr-H, 16L19 and 16H36E for 16S, cmyc1U and cmyc-ex2 R for C-MYC, 12Sh and
12SKH for 12S, and dgLCO1490 and dgHCO2198 for CO1 (Crawford 2003; Folmer et al. 1994; Goebel et al.
1999; Heinicke et al. 2007; Palumbi et al. 1991; Pauly et al. 2004; Wiens et al. 2005).
Phylogenetic inference
Our phylogeny is based on sequences of Microhylidae obtained from GenBank and published by Frost et al.
(2006), Wollenberg et al. (2008), Funk and Cannatella (2009), de Sá et al. (2012), Blackburn et al. (2013), and
Peloso et al. (2014, 2015) for a total of 302 terminals. In addition to the loci sequenced by us, we included Gen-
Bank sequences of the nuclear genes Brain-Derived Neurotrophic Factor (BDNF), Histone H3, 28S rRNA, Seven
in Absentia homolog (SIA1), and Tyrosinase for a total of nine loci and up to 6171 bp. As reported by Peloso et al.
(2014), the extremes of multiple 16S sequences from de Sá et al. (2012) were difficult to align and different from
sequences of all other microhylids (including congeneric species) suggesting sequencing errors. We eliminated the
5’ misaligned extremes for KC180058, KC180060, KC180062, KC180063, KC180065, KC190069, KC180071 and
KC180074. We also eliminated the 3’ extreme for KC180078.
As outgroup, we included GenBank sequences of species of the genera Afrixalus, Amietia, Arthroleptis Brevi-
324 · Zootaxa 4779 (3) © 2020 Magnolia Press SÁNCHEZ-NIVICELA ET AL.

ceps, Callulina, Hoplophryne, Hyperolius, Heterixalus, Hemisus, Limnonectes Lithobates, Polypedates Probrevi-
ceps, Spelaeophryne, Strongylopus, Tachycnemis, and Tomopterna. Outgroup choice is based on de Sá et al. (2012)
and Peloso et al. (2015). The matrix had 302 terminals. GenBank accession numbers for newly generated se-
quences are (to be added). The aligned matrix is available in Zenodo.org under HTTPS://DOI.ORG/10.5281/ze-
nodo.3722517.
Sequences were aligned using the MAFFT 7.017 and the L-INS-i algorithm (Katoh and Standley 2013) imple-
mented as a plugin in Geneious 7.1.7 software (GeneMatters. Corp.). The alignment was visually inspected in Mes-
quite (version 3.01; Maddison and Maddison 2014), and alignment errors were adjusted manually. We partitioned
the matrix to allow separate models of evolution for 12S, 16S, 28S, and each of the codon positions of the protein-
coding genes BDNF, CMYC, CO1, Histone H3, SIA1, and Tyrosinase for a total of 21 partitions. We used software
PartitionFinder v. 1.1.1 (Lanfear et al. 2012) to simultaneously estimate both the best-fit model for each partition
and the best partition strategy.
The phylogeny was estimated using maximum likelihood (ML) as optimality criteria. Tree search was carried
out with a genetic algorithm using GARLI version 2.01 (Zwickl 2006). We conducted 20 stochastic likelihood
searches starting from stepwise trees. We chose to start from stepwise trees, instead of random trees, because in 20
preliminary searches (10 starting from stepwise and 10 from random trees), all optimal trees were found with the
stepwise searches. Search parameters were set to the default values except for “limsprrange” (10, default 6) and
genthreshfortopoterm (2500000, default 20000). Both values increase search exhaustiveness. Three out of 20 step-
wise-starting searches found best trees with likelihood values within 1 log unit of the global best tree, indicating that
the search strategy repeatedly converged on the same optimal trees.
Branch support was estimated with non-parametric bootstrap from 100 pseudoreplicate searches. Search set-
tings for each replicate were the same as those of the full search (limsprrange = 10, genthreshfortopoterm = 2500000,
other parameters default) and one search per bootstrap replicate.
To corroborate the relationships of the new species with its congeners, we made two additional phylogenetic analy-
ses for Elachistocleis using Gastrophryne as outgroup. The first analysis was based on the mtDNA genes only (2765
bp, 20 terminals) and the second on nuclear genes only (3151 bp, 20 terminals). Both analyses used maximum
likelihood (ML) as optimality criterion. Matrixes were partitioned by gene and codon position (in coding genes). To
choose the best model of evolution for each partition, we used the command MFP (Chernomor et al. 2016; Kaly-
aanamoorthy et al. 2017) in software IQ-TREE 1.6.8 (Nguyen et al. 2015). To find the best phylogeny we also used
IQ-TREE 1.6.8 under default settings. Branch support was estimated with 1000 ultrafast bootstrap searches (-bb
1000 command). Uncorrected p-distances were obtained with MEGA 7.0 (Kumar et al. 2016).
Results
Phylogeny
Our phylogeny (Fig. 1) shows strong support for the New World subfamily Gastrophryninae (bootstrap = 84).
Ctenophryne is sister to a clade composed of all remaining species of Gastrophryninae. Within the later, there are
two large clades, Chiasmocleis and a clade composed by all remaining genera. Elachistocleis, including the new
species, is sister to Gastrophryne + Hypopachus (same as in de Sá et al. 2018 and Peloso et al. 2015). The new
species is sister to a clade composed of all other species of Elachistocleis included in the analysis. Among them, E.
panamensis, is the first to diverge. The separate phylogenies for mitochondrial and nuclear genes (Fig. 2) confirm
the same relationships for E. araios sp. n. relative to other Elachistocleis. Uncorrected p-genetic distances for the
gene 16S between the new species and other Elachistocleis range from 9.6 to 14.8% (TABLE 1).
The phylogenetic position of the population from western Ecuador indicates that it represents a species distinct
from other Elachistocleis. We checked available names among all species of Elachistocleis (Frost 2019) and found
five: Engystoma cesarii mottae Miranda-Ribeiro, 1920, Engystoma ovale concolor Miranda-Ribeiro, 1920, Engys-
toma ovale lineatum Miranda-Ribeiro, 1920, and Stenocephalus microps Tschudi, 1838. Because none of them are
distributed in the Pacific Basin or Central America and they are synonymized under species markedly distinct from
the Ecuadorian populations, we conclude that it is a new species that we describe below.
a new trans-Andean species of Elachistocleis Parker 1927 Zootaxa 4779 (3) © 2020 Magnolia Press · 325
FIGURE 1. Phylogenetic relationships of the subfamily Gastrophryninae. Best maximum-likelihood tree obtained from analy-
ses of DNA sequences of mitochondrial (12S, 16S and CO1) and nuclear (BDNF, cmyc2, H3A, 28S, SIA1, and Tyr) genes from
302 terminals and up to 6171 bp. Blue numbers below branches are ML bootstrap support values. Voucher number and GenBank
accession number for the 16S gene are shown for each specimen. Outgroup and other subfamilies are not shown. The complete
phylogeny is available in Zenodo.org under HTTPS://DOI.ORG/10.5281/zenodo.3723213.

FIGURE 2. Phylogenetic relationships of Elachistocleis based on maximum likelihood analyses of DNA sequences. Above:
phylogeny based on nuclear genes (BDNF, cmyc2, H3A, 28S, SIA1, and Tyr) . Below: phylogeny based on mitochondrial genes
(12S, 16S, and CO1). Numbers on branches correspond to ultrafast bootstrap support values (≥ 95 are considered as signifi-
cant).
TABLE 1. Matrix of pairwise uncorrected p-distances between Elachistocleis based on DNA sequences of mitochondrial gene 16S. The species name is followed by the voucher
museum number or, if unavailable, the GenBank sequence accession number.
E. araios sp. nov E. bicolor E. cesarii E. cesarii E. helianneae E. ovalis
QCAZ56485 ZVCB10599 CFBH20007 ZUEC DCC3301 MPEG29419 NKAG6488
E. araios sp. nov. QCAZ 56485 0 0.112 0.14 0.14 0.149 0.105
E. bicolor ZVCB10599 0.112 0 0.061 0.067 0.025 0.023
E. cesarii CFBH20007 0.14 0.061 0 0.006 0.099 0.065
E. cesarii ZUEC DCC3301 0.14 0.067 0.006 0 0.103 0.071
E. helianneae MPEG29419 0.149 0.025 0.099 0.103 0 0.025
E. ovalis NKAG6488 0.105 0.023 0.065 0.071 0.025 0
E. piauiensis MPEG27839 0.135 0.072 0.02 0.023 0.102 0.071
E. sp. CAS HERP231748 0.147 0.025 0.1 0.102 0.025 0.032
E. sp. E47 KC179875 0.096 0.064 0.02 0.021 0.071 0.071
328 · Zootaxa 4779 (3) © 2020 Magnolia Press

E. sp. E53 KC179876 0.12 0.029 0.072 0.078 0.021 0.035
E. surinamensis AMNH A141136 0.134 0.07 0.023 0.022 0.101 0.071
E. surinamensis KC179873 0.096 0.069 0.018 0.017 0.067 0.069
TABLE 1. (Continued)
E. piauiensis E. sp. E. sp. E. sp. E. surinamensis E. surinamensis
MPEG27839 CAS HERP231748 KC179875 KC179876 AMNH A141136 KC179873
E. araios sp. nov. QCAZ 56485 0.135 0.147 0.096 0.12 0.134 0.096
E. bicolor ZVCB10599 0.072 0.025 0.064 0.029 0.07 0.069
E. cesarii CFBH20007 0.02 0.1 0.02 0.072 0.023 0.018
E. cesarii ZUEC DCC3301 0.023 0.102 0.021 0.078 0.022 0.017
E. helianneae MPEG29419 0.102 0.025 0.071 0.021 0.101 0.067
E. ovalis NKAG6488 0.071 0.032 0.071 0.035 0.071 0.069
E. piauiensis MPEG27839 0 0.105 0.025 0.084 0.026 0.024
E. sp. CAS HERP231748 0.105 0 0.069 0.008 0.105 0.07
E. sp. E47 KC179875 0.025 0.069 0 0.08 0.018 0.02
E. sp. E53 KC179876 0.084 0.008 0.08 0 0.082 0.082
E. surinamensis AMNH A141136 0.026 0.105 0.018 0.082 0 0
E. surinamensis KC179873 0.024 0.07 0.02 0.082 0 0
SÁNCHEZ-NIVICELA ET AL.
Elachistocleis araios sp. n.
Holotype. (Figs. 3–5, 7) QCAZ 56485, an adult female, from La Independencia, 3.0583° S, 79.7171° W (WGS84),
308 m a.s.l., Ponce Enriquez canton, Azuay province, Ecuador (Fig. 3). Collected by Juan C. Sánchez-Nivicela and
Daniel Navas on December 14, 2009.
Paratype. (Figs. 3, 6–7) QCAZ 74764, an adult female from Cooperativa 23 de Noviembre, Cerro de Hayas
2.7174° S, 79.6285 W° (WGS84), 57 m a.s.l., Naranjal canton, Guayas province, Ecuador (Fig. 3). Collected by
Yerka Sagredo on February 6, 2019.
FIGURE 3. Map showing records of Elachistocleis araios sp. n. Red square = holotype, Azuay province; red circle = paratype,
Guayas province. Ecuador is shown on green on the small map. Black lines represent province boundaries.
Diagnosis. A small species of Elachistocleis (SVL in females = 25.7–26.1 mm, n = 2; males unknown), charac-
terized by an ovoid body form (Fig. 4, 6), triangular head, slightly wider than long, and a barely developed occipital
fold (Fig. 5 A–B). Snout rounded in dorsal and lateral views; canthus rostralis slightly concave and loreal region is
convex; nostrils laterally oriented (Fig. 5 A–B). Postcommissural glands inconspicuous. Tympanic annulus, tym-
panic membrane, and dentigerous process of vomer absent. Upper jaw more projected than the lower jaw, tongue
is big, ovoid, and covers the entire buccal cavity. Choanae rounded and widely separated, oriented to the anterior
region of buccal cavity. Arms slender without tubercles on forearm. Hands (Fig. 5C) lack interdigital membranes
and supernumerary tubercles; first finger almost the same size as the second, both almost as long as finger IV, fin-
ger lengths are I<II<IV<III; subarticular tubercles, ovoid and as wide as fingers, prominent thenar tubercle, palmar
tubercle longitudinally divided. Foot (Fig. 5D) lacks interdigital membranes and supernumerary tubercles, toes
lack discs. Inner plantar tubercle same size as subarticular tubercles; relative toe lengths I<II<V<III<IV. TBL is
slightly longer than THL. Dorsal surface of body and fingers lack dermal spines. Cloacal region lacks glands and
tubercles.
FIGURE 4. Holotype of Elachistocleis araios sp. n., female, in life. Above left, Lateral view; below left, dorsal view; right,
frontal view.
FIGURE 5. Holotype of Elachistocleis araios sp. n., female, in preservative. (A) Lateral view of head, (B) dorsal view of head,
(C) ventral view of right hand, (D) ventral view of right foot.

Comparisons. The new species can be immediately distinguished from most Elachistocleis species in hav-
ing an immaculate venter (variably pigmented or blotched in E. bumbameuboi, E. carvalhoi, E. corumbaensis, E.
erythrogaster, E. magnus, E. ovalis, E. pearsei, E. piauiensis, E. skotogaster, E. surinamensis, and E. surumu).
Elachistocleis araios is, therefore, most similar to E. bicolor, E. cesarii, E. haroi, E. helianneae, E. matogrosso, E.
muriaquitan, and E. panamensis in a having uniform, immaculate, ventral coloration. Elachistocleis araios is distin-
guished from Elachistocleis bicolor, E. cesarii, E. corumbaensis, E. helianneae, E. matogrosso, and E. muriaquitan
by its dorsal color patters, E. araios mottled grey with a paravertebral symmetric pattern (dark dorsal coloration,
which can vary from uniformly grayish to dark grey with scattered lighter spots).
The new species is further distinguished from E. bicolor, E. corumbaensis, E. matogrosso, and E. muriaquitan
in not having any light spot in the axillae and groin (light spot present). It also differs from E. bicolor, E. corum-
baensis, and E. haroi in having grayish cream venter and hidden surfaces of limbs (bright yellow coloration on
venter and hidden surfaces of limbs on the other species). Moreover, E. bicolor also lacks a mid-dorsal stripe, which
is notorious in the new species (but note that this character might be variable in several microhylid species, and that
our sample size for E. araios is fairly small).
FIGURE 6. Paratype of Elachistocleis araios sp. n. in life. (A) Profile view, (B) ventral view, (C) dorsal view.
Elachistocleis araios differs from most species of the genus in having slender fingers and toes with slightly
expanded, rounded tips (robust fingers and toes, with tips more narrow than the width of the fingers in E. corum-
baensis, E. surinamensis, E. ovalis, E. skotogaster, E. piauiensis, E. panamensis, E. muriaquitan, E. magnus, E.
erythrogaster). In the new species, FI almost the same size as FII, both almost as long as finger IV (FI much smaller
than the second, both shorter than FIV in E. haroi). E. araios differ from several species in having the postcommis-
sural gland poorly developed (massive or large postcommissural glands in E piauiensis, E. magnus, E. erythrogaster,
helianneae, E. surumu, E. carvalhoi, E. bumbameuboi, E. cesarii / absent in E. skotogaster). The new species lack
a dorso-lateral fold (present in E. cesarii, E. helianneae, E. surumu, E. carvalhoi, E. bumbameuboi, E. matogrosso,
E. bicolor, E. skotogaster, E. magnus, E. muriaquitan, E. corumbaensis, E. pearsei). Interdigital web absent in E.
araios (present, rudiment in E. pearsei).
Elachistocleis araios (Figs. 7A-B) is most similar externally to E. panamensis (Fig. 7C), from which is differs
by the following combination of characters: a short and rounded snout in E. araios (long and truncate snout in E.
panamensis); present, although weak, lateral fringes in TIV (absent); thenar tubercle small and rounded (large and
oval); inner metatarsal tubercles small and rounded (big and oval); dorsal color patterns with dark spots without
defined patterns (generally a repetitive triangular design or a large irregular spot in the middle of the back); venter
immaculate (cream with clear spots poor visible); grayish-brown, ash-looking, throat and chin—differs from ventral
pattern (brown-cream with white spots—same as venter).
FIGURE 7. Comparison of dorsal (above) and ventral (below) views of Elachistocleis araios sp. n. in preservative. (A) holo-
type (SLV 25.7 mm); (B) paratype (SVL 26.1 mm); and, E. panamensis, (C) (ICN-48708) female (SVL 23.6 mm).
Description of the holotype. Adult female (Fig. 4–5, 7A). Body ovoid, skin smooth. Head small, HL 20.2% of
SVL, 85.2% of HW. Snout prominent, projected beyond the mouth, protruding in lateral view. Tiny eyes, slightly
prominent. Nostrils small, laterally oriented. Canthus rostralis slightly concave and convex in loreal. Complete oc-
cipital fold. Postcomissural glands poorly developed. Upper jaw projected beyond lower jaw.
Arms robust, short, without tubercles, a thick ulnar fold present. Fingers slender, short. Relative lengths of
fingers: I<II<IV<III. Subarticular tubercles round. Palmar tubercle longitudinally divided, internal section rounded
and external ovoid, four times larger than thenar tubercle. Supernumerary tubercles absent. Lateral fringe on fingers
weak. Finger tips rounded, weakly expanded. Posterior extremities without tubercles, tarsal fold present. Legs short,

robust. Thigh larger than shank, but shorter than foot length. Toes slender. Relative toes lengths: I<II<V<III<IV.
Subarticular tubercles round. Supernumerary plantar tubercles absent. Lateral fringe on toes weak. Inner metatarsal
tubercle oval. Outer metatarsal tubercle absent. Toes tips rounded, weakly expanded. Tips of fingers and toes with
terminal rounded discs. Interdigital web of hands and foots absent. (Fig. 5 C–D).
Measurements of the holotype in mm: SVL 25.7; HL 5.0; HW 6.1; ED 1.8; IOD 2.7; IND 1.4; END 2.1; THL
9.9; TBL 11.0; FL 12.4; HDL3 5.2; HDL4 3.6; FAL 4.3; FD3 0.5; FT4 0.7.
Variation. Measurement from the paratype in mm: SVL 26.1; HL 7.1; HW 6.6; ED 2.8; IOD 2.6; IND 1.6; END
2.3; THL 9.7; TBL 10.7; FL 11.2; HDL3 7.1; HDL4 3.4; FAL 4.4; FD3 0.4; FT4 0.6. The patterns of coloration and
design do not represent major differences except that, in the paratype the dorsal line is thinner and less visible in the
anterior direction and absent in the head (Fig. 6, 7A, 7B).
Coloration of the holotype in life. The coloration of dorsum, thighs, shanks, and arms is light brown with
small black patches and tiny yellowish-cream dots. A narrow light cream dorsal line extends from the tip of snout
to the vent. A thin dorsolateral line resulting from the combination of brown spots extends from the nostrils to the
inguinal region and, creates a clearly marked edge between dorsum and flank colorations. Two inguinal spots are
present visible from the dorsum. The posterior surface of thighs has a light cream line extending to the inguinal
region. Irregular dark brown lines are distributed in thighs, tibia, and feet and, a slightly cream line running along
the back of the thighs and across the urostyle bump. Flanks and ventral surfaces of head are light brown with small
yellowish-cream spots. Venter is uniform light yellow anteriorly and grayish-yellow posteriorly. Ventral surface of
thighs and arms is yellowish cream with light brown (Fig. 4).
Coloration of the holotype in preservative. The coloration of dorsum and forelimbs is uniformly dark brown
with few cream spots. The dorsolateral line is not distinguishable; flanks and ventral surface of the head are gray-
ish-cream. The middorsal line is cream, the two black patches on the posterior surfaces of dorsum, the cream line on
the thighs, and the irregular dark brown lines in the forelimbs are distinct. The venter is yellowish-cream anteriorly
and grayish-cream posteriorly (Fig. 7).
Etymology. The name of the new species is a noun in apposition from the Greek araiós (αραιός) which means
scarce, alluding to the fact that it is a difficult species to find in field.
Distribution, natural history and conservation status. Elachistocleis araios is only known from two locali-
ties in the lowlands of Azuay and Guayas provinces of Ecuador. The holotype was found in a western foothill forest
fragment. The habitat is characterized by the presence of dense vegetation cover with a thick layer of leaf litter and
several small streams (Fig. 8). The holotype was collected at night while walking through the leaf litter in a trail
of cacao crops near the forest. Sympatric amphibian species included: Epipedobates anthonyi, Lithobates bwana,
Leptodactylus labrosus, and Pristimantis achatinus. The paratype and holotype were adult gravid females.
At the type locality, land has been concessioned for mining. Traditional mining is the most common type of
mineral extraction in the region. In this process, toxic compounds (e.g., cyanide) and heavy metals (e.g., mercury)
are used for extraction and refinement of minerals. Consequently, water bodies, and soils are directly and irrevers-
ibly affected. The second locality is a Provincial Natural Recreation Area of 631 ha, but it is an area with strong de-
forestation pressures. Activities such as cattle ranching and agriculture are widespread in the area as well. Therefore,
important extensions of natural vegetation have been replaced with crops and grass. Despite this, we do not have
enough evidence to define the degree of threat of this species and we highlight the need for population-level studies
to know its status, which is why we recommend that E. araios be listed as Deficient Data on the IUCN Red List.
Discussion
Few microhylids species are known from the Pacific versant of the Andes in South America, i.e. Ctenophryne ater-
rima and Ctenophryne minor. Among Elachistocleis, only E. panamensis and E. araios occur on the Pacific versant
of the Andes. However, the distribution of E. panamensis is much broader, extending from the lowlands of Panama
(Central America) to the Magdalena valley (eastern Andes) with scattered records in northern Chocó of Colombia
(Nelson 1972, Lynch & Suarez-Mayorga 2004). As far as we are aware, E. araios is the only actually known spe-
cies of the genus, endemic to the Pacific slopes of the Andes and the only member the Elachistocleis known from
Ecuador.
FIGURE 8. Habitat of the holotype of Elachistocleis araios sp. n.
Recent studies (van der Meijeden 2007, de Sá et al. 2012), suggest that the radiation and diversification of the
new-world microhylids started on the Oligocene in northern South America. The origin of the Elachistocleis may
have occurred in the Colombian Caribbean to Atlantic basins (de Sá et al. 2012). This hypothesis was inferred based
on the position of E. panamensis as diverging early within the genus. The addition of E. araios opens the possibility
of an origin in the Pacific versant of South America of Ecuador with a subsequent dispersal to the northern limit of
the Andes and then to eastern South America where the majority of extant species originated.
Our species description is based solely on two female specimens and therefore our characterization of pheno-
typic variation is limited. Although describing species based on a few specimens can be problematic, the phylo-
genetic position and geographic distribution of E. araios provides strong evidence of its uniqueness among New
World microhylids. The new species differ from congeneric species in coloration, body and snout shape, and the
presence of a small post commissural gland. Pereyra et al. (2013) suggested that coloring patterns in Elachistocleis
provide an important tool for diagnosis of species. Therefore, additional efforts should be directed to collect more
specimens and adequately characterize variation within E. araios. Given its phylogenetic position, a better charac-
terization of this species will be key to understanding patterns of phenotypic evolution in Elachistocleis.
The geographical distribution of the new species is notable for its isolation from other species. The closest,
geographically, is Elachistocleis panamensis, which is found nearly 1000 km to the north. The other species are
separated by vast distances or the formidable barrier of the Andes. Moreover, most other species of Elachistocleis
occupy the lowlands. This could suggest the existence of several species of the genus not yet discovered in the west-
ern region of the Andes in Ecuador and Colombia, a poorly known and highly threatened region of the Neotropics.
The description of a new species of Elachistocleis in southwestern Ecuador contributes to a better understand-
ing of the diversity of the genus as a whole (Nunes de Almeida & Toledo 2012, Pereyra et al. 2013), and highlight
the possible existence of additional microhylid taxa in the western Andes. This finding also raises new questions
about the complex diversification patterns of the genus in South America.

Acknowledgments
Laboratory work was funded by grants from Secretaría de Educación Superior, Ciencia, Tecnología e Innovación
(SENESCYT, Arca de Noé Initiative; O. Torres-Carvajal and S. R. Ron Principal Investigators) and was carried out
at the QCAZ molecular laboratory by Sebastian Escobar. This study was founded by Universidad del Azuay (Fon-
dos UDA 2015 [39, 2016]). P.L.V. Peloso was supported by grants from the Brazilian Conselho Nacional de De-
senvolvimento Científico e Tecnológico (CNPq: grant numbers 313680/2014-0, and 434362/20182). Yerka Sagredo
was supported by ECOPAR. Special thanks to Professor John D. Lynch of the National University of Colombia for
facilitating access to the material available in his collection, and Juan David Fernández for his help in obtaining
species data. JCS thanks Manuel Sánchez for his help in the manuscript.
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APPENDIX 1. Specimens examined for morphological comparisons. For each individual we show museum number, species, locality, elevation, and sex. In specimens for which
we obtained information from the literature, a reference is provided. Museum acronyms are: ICN (Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá-
Colombia), MNRJ (Museu Nacional da Universidade Federal do Rio de Janeiro, Brazil), MZUSP (Museu de Zoologia da Universidade de São Paulo), ZUFMS-AMP (Coleção Zo-
ológica de Referência da Universidade Federal de Mato Grosso do Sul, Campo Grande), MCP (Museu de Ciências e Tecnologia, Pontifícia Universidade Católica do Rio Grande
do Sul, Porto Alegre; current for fishes & herps), FML (Fundación Miguel Lillo, San Miguel de Tucumán), ZUEC (Museu de Zoologia da Universidade Estadual de Campinas
“Adão José Cardoso”, Campinas.), AMNH (American Museum of Natural History, New York), JJ (Jorge Jim Private Collection, Deparatmento de Zoologia, IBBMA–UNESP,
Botucatu, São Paulo, Brazil.). Country abbreviations are: ARG (Argentina), BRA (Brazil), COL (Colombia), PAN (Panama).
Museum number Species Locality Elevation (m) Sex Source
MNRJ 53378 Elachistocleis bumbameuboi Säo Luís, Maranhäo, BRA 24 Male Caramaschi (2010)
MNRJ 51384 Elachistocleis carvalhoi Aragominas, Tocantins, BRA 345 Male Caramaschi (2010)
MZUSP 529 Elachistocleis cesarii Piquete, Säo Paulo, BRA 675 Male Toledo et al. (2010)
ZUFMS-AMP 924 Elachistocleis corumbaensis Corumbá, Mato Grosso do Sul, BRA ___ Male Piva et al. (2017)
MCP 2450 Elachistocleis erythrogaster Centro de Pesquisas e Conservacäo da Natureza Pró- 960 Male Kwet & Di-Bernardo (1998)
Mata, Säo Francisco de Paula, Rio Grande do Sul, BRA

FML 24900 Elachistocleis haroi El Algarrobal, Jujuy, ARG 1211 Male Pereyra et al. (2013)
MNRJ 6989 Elachistocleis helianneae Humaitá, Amazonas, BRA 90 Male Caramaschi (2010)
ZUEC 11384 Elachistocleis magnus Fazenda Jaburi, Espigäo do Oeste, Rondönia, BRA 280 Male Toledo (2010)
MNRJ 4812 Elachistocleis matogrosso Cuiabá, Mato Grosso, BRA 177 Female Caramaschi (2010)
ZUEC 5666 Elachistocleis muiraquitan Xapuri, Acre, BRA 172 Male Nunes de Almeida & Toledo (2012)
ICN 56702 Elachistocleis ovalis Acácias, Meta, COL 514 Male This work
ICN 56704 Elachistocleis ovalis Acácias, Meta, COL 514 Female This work
ICN 56706 Elachistocleis ovalis Acácias, Meta, COL 514 Female This work
ICN 11707 Elachistocleis panamensis 10km between Villa Vieja-Baraya, Huila, COL ___ Juv This work
ICN 11708 Elachistocleis panamensis 10km between Villa Vieja-Baraya, Huila, COL ___ Male This work
......continued on the next page
APPENDIX 1. (Continued)
ICN 11713 Elachistocleis panamensis 10km between Villa Vieja-Baraya, Huila, COL ___ Male This work
ICN 11714 Elachistocleis panamensis 10km between Villa Vieja-Baraya, Huila, COL ___ Female This work
ICN 43419 Elachistocleis panamensis Cispata, Córdova, COL ___ Juv This work
ICN 43420 Elachistocleis panamensis Cispata, Córdova, COL ___ Male This work
ICN 43421 Elachistocleis panamensis Cispata, Córdova, COL ___ Juv This work
ICN 44699 Elachistocleis panamensis Puerto Boyacá, Boyacá, COL ___ Female This work
ICN 48687 Elachistocleis panamensis Montería, Córdoba, COL 64 Male This work
ICN 48688 Elachistocleis panamensis Montería, Córdoba, COL 64 Female This work
a new trans-Andean species of Elachistocleis Parker 1927

ICN 48699 Elachistocleis panamensis Montería, Córdoba, COL ___ Female This work
ICN 48700 Elachistocleis panamensis Montería, Córdoba, COL ___ Male This work
ICN 48702 Elachistocleis panamensis Montería, Córdoba, COL 64 Juv This work
ICN 48703 Elachistocleis panamensis Santa Cruz de Lorica, Córdoba, COL 20 Female This work
ICN 48705 Elachistocleis panamensis Santa Cruz de Lorica, Córdoba, COL 20 Male This work
ICN 48706 Elachistocleis panamensis Santa Cruz de Lorica, Córdoba, COL 20 Male This work
ICN 48709 Elachistocleis panamensis Montería, Córdoba, COL ___ Female This work
ICN 48921 Elachistocleis panamensis Montería, Córdoba, COL ___ Male This work
Zootaxa 4779 (3) © 2020 Magnolia Press ·

......continued on the next page
339
APPENDIX 1. (Continued)
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ICN 49060 Elachistocleis panamensis Juan de Acosta, Atlántico, COL 15 Female This work
ICN 49061 Elachistocleis panamensis Baraona, Atlántico, COL 50 Female This work
ICN 49062 Elachistocleis panamensis Tubará, Atlántico, COL 100 Juv This work
ICN 49343 Elachistocleis panamensis San Marcos, Sucre, COL 45 Female This work
ICN 50104 Elachistocleis panamensis Zambrano, Bolívar, COL 70 Juv This work

ICN 50107 Elachistocleis panamensis Zambrano, Bolívar, COL 70 Male This work
AMNH 52741 Elachistocleis panamensis Old Panama, PAN ___ ___ Dunn et al. (1948)
ICN 14043 Elachistocleis pearsei Turbaco, Bolívar, COL 100 Male This work
ICN 52778 Elachistocleis pearsei Cármen de Apicala, Tolima, COL ___ Female This work
ICN 56413 Elachistocleis pearsei Cármen de Apicala, Tolima, COL ___ Male This work
JJ 6024 Elachistocleis piauiensis Picos, Piauí, BRA ___ Male Caramaschi & Jim (1983)
FML 07927 Elachistocleis skotogaster Los Toldos, Santa Victoria, Salta, ARG 1100 Male Lavilla et al. (2003)
MNRJ 25210 Elachistocleis surumu Pacaraima, Roraima, BRA 121 Male Caramaschi (2010)

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a new trans-Andean species of Elachistocleis Parker 1927

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340 · Zootaxa 4779 (3) © 2020 Magnolia Press

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