A new Andean anole species of the Dactyloa clade (Squamata: Iguanidae) from western Ecuador

Abstract

We describe a new species of Anolis from the western slopes of the Andes of Ecuador, province of Bolívar. It is referred to (1) the aequatorialis series based on its moderate size and narrow toe lamellae, and (2) the eulaemus sub-group based on having a typical Anolis digit, in which the distal lamellae of phalanx III distinctly overlap the proximal subdigital scales of phalanx II. The new species is most similar morphologically to A. otongae and A. gemmosus, both from similar elevations on the western Andean slopes of Ecuador, but differs from these species in morphology and color patterns. We present a phylogeny based on DNA sequence data as additional evidence supporting delimitation of the new species. The new species and A. gemmosus are sister taxa within the “western Dactyloa clade.”

Full text

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Amphib. Reptile Conserv. | amphibian-reptile-conservation.org May 2014 | Volume 8 | Number 1 | e76
Amphibian & Reptile Conservation [Special Section] 8(1) : 8–24.
A new Andean anole species of the Dactyloa clade
(Squamata: Iguanidae) from western Ecuador
1Fernando P. Ayala-Varela, 2Diana Troya-Rodríguez, 3Xiomara Talero-Rodríguez
and 4Omar Torres-Carvajal
1,2,3,4Escuela de Ciencias Biológicas, Ponticia Universidad Católica del ECUADOR, Avenida 12 de Octubre 1076 y Roca, Apartado 17-01-2184,
Quito, ECUADOR
Abstract.—We describe a new species of Anolis from the western slopes of the Andes of Ecuador,
province of Bolívar. It is referred to (1) the aequatorialis series based on its moderate size and
narrow toe lamellae, and (2) the eulaemus sub-group based on having a typical Anolis digit, in which
the distal lamellae of phalanx III distinctly overlap the proximal subdigital scales of phalanx II. The
new species is most similar morphologically to A. otongae and A. gemmosus, both from similar
elevations on the western Andean slopes of Ecuador, but differs from these species in morphology
and color patterns. We present a phylogeny based on DNA sequence data as additional evidence
supporting delimitation of the new species. The new species and A. gemmosus are sister taxa
within the “western Dactyloa clade.”
Key words. Clade Dactyloa, DNA, lizard, phylogeny, South America, systematics
Citation: Ayala-Varela FP, Troya-Rodríguez D, Talero-Rodríguez X, Torres-Carvajal O. 2014. A new Andean anole species of the Dactyloa clade (Squa-
mata: Iguanidae) from western Ecuador. Amphibian & Reptile Conservation 8(1) [Special Section]: 8–24 (e76).
Correspondence. 1fpayala2000@yahoo.com (Corresponding author); 2dianatr17@gmail.com; 3xiomy.talero@gmail.com;
4omartorcar@gmail.com
Copyright: © 2014 Ayala-Varela et al. This is an open-access article dis-
tributed under the terms of the Creative Commons Attribution–NonCom-
mercial–NoDerivs 3.0 Unported License, which permits unrestricted use
for non-commercial and education purposes only provided the original
author and source are credited. The ofcial publication credit source: Am-
phibian & Reptile Conservation at: amphibian-reptile-conservation.org
Introduction
With nearly 490 described species, anole lizards (Anolis)
have proliferated impressively in the Americas (Nich-
olson 2002; Poe 2004), possibly prompted by ecologi-
cal opportunity (Losos 2009). Although the diversity of
these lizards has been extensively studied in the West
Indies (Losos 2009), the same is not true for the main-
land radiation, which is probably greater than previously
thought. For example, all but two—Anolis ruibali Navar-
ro & Garrido 2004 and A. sierramaestrae Holáňová et al.
2012—of the 31 new species of Anolis described during
the last decade (2003–2013) occur in mainland Central
and South America (Uetz and Hošek 2014). Improving
knowledge concerning the diversity of mainland anoles
is crucial to understanding the nature of this radiation.
Anole lizards represent the most species-rich clade
traditionally recognized as a genus in Ecuador, with 37
species reported to date (Torres-Carvajal et al. 2014).
The diversity of anole lizards in Ecuador is remarkably
greater west of the Andes, with more than twice the num-
ber of species that occur east of the Andes (25 and 12
species, respectively). Of these, ve species have been
described during the last six years from both sides of the
Andes as a result of both careful examination of exist-
ing collections and recent collecting in poorly explored
areas. Here we contribute to that growing body of taxo-
nomic knowledge with the description of a new species
of Anolis endemic to the western slopes of the Andes in
Ecuador. We present molecular evidence supporting rec-
ognition of the new species by performing phylogenetic
analyses of mitochondrial DNA sequence data.
Materials and Methods
Morphological data
All known specimens of the new species described in
this paper are included in the type series, and were de-
posited in the Museo de Zoología, Ponticia Universi-
dad Católica del Ecuador, Quito (QCAZ). Specimens of
other species of Anolis examined in this study are listed
in Appendix 1. We follow previously proposed terminol-
ogy (Williams et al. 1995) for measurements and squa-
mation. Nine morphological measurements were taken
with digital calipers and recorded to the nearest 0.1 mm:
head length, head width, head height, forelimb length,
hindlimb length, snout-vent length, jaw length, axilla-
groin length, and snout length. In addition, tail length
measurements were taken with a ruler and recorded to
the nearest millimeter; regenerated or broken tails were
not measured. Sex was determined by noting the pres-
ence of hemipenes, which were everted in all male speci-
mens during preparation.
Statistical analyses
Given that the new species is very similar in morphol-
ogy to Anolis gemmosus and A. otongae we performed

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Ayala-Varela et al.
Fig. 1. Head of the holotype (QCAZ 3449) of Anolis poei sp. nov. in dorsal (top), ventral (middle), and lateral (bottom) views
[Scale bar = 10 mm]. Photographs by F. Ayala-Varela.

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A new species of Anolis lizard from western Ecuador
a Principal Component Analysis (PCA) to determinate
whether separation in morphological space between
those species was statistically signicant. Principal com-
ponents (PCs) were extracted from a covariance matrix
of the raw and rescaled data. The new species is most
similar to A. gemmosus, for which we also used t-tests to
evaluate quantitative differences between both species.
One of the assumptions of the t-test for two samples is
that the variances of both samples are equal: therefore,
F-tests also were performed for each character to test for
equality of variances. If the variances were not the same
(i.e., P < 0.05), an unequal variance t-statistic was used.
Statistical analyses were performed in SPSS Statistics 17
(SPSS Inc. 2008).
The distribution map was prepared in ArcMap 9.3
(ESRI, Inc.); WGS84 is the datum for all coordinates
presented below.
DNA sequence data
Total genomic DNA was digested and extracted from liv-
er or muscle tissue using a guanidinium isothiocyanate
extraction protocol. Tissue samples were rst mixed with
Proteinase K and a lysis buffer and digested overnight
prior to extraction. DNA samples were quantied using
a Nanodrop® ND-1000 (NanoDrop Technologies, Inc),
re-suspended and diluted to 25 ng/ul in ddH2O prior to
amplication.
Using primers and amplication protocols from the
literature (Folmer et al. 1994; Kumazawa and Nishida
1993; Macey et al. 1997; Schulte and Cartwright 2009)
we obtained 2807 nucleotides (nt) representing the nucle-
ar gene recombination-activating gene 1 (RAG1, 811nt),
as well as the mitochondrial genes Cytochrome c oxi-
dase I (CO1, 655nt) and a continuous fragment includ-
ing the NADH dehydrogenase subunit 2 (ND2, 1038 nt),
tRNATrp, tRNAAla, tRNAAsn, tRNACys (282nt), and
the origin of the light-strand replication (Ol, 29nt). The
new sequence data were obtained for three individuals of
the new species described herein, two of A. gemmosus,
and two of A. otongae. In addition we used sequence data
generated by Castañeda and de Queiroz (2011) for 20 in-
dividuals of the clade Dactyloa, as well as one sequence
of A. occultus, which was used as the outgroup in the
phylogenetic analysis. Gene regions of taxa included in
phylogenetic analyses along with their GenBank acces-
sion numbers are shown in Table 1.
Phylogenetic analyses
Editing, assembly, and alignment of sequences were
performed with Geneious ProTM 5.3 (Biomatters Ltd.
2010). Genes were combined into a single dataset with
eleven partitions, three per protein coding gene corre-
sponding to each codon position, one with all tRNAs, and
one with the Ol. The best partition strategy along with the
corresponding models of evolution were obtained in Par-
titionFinder 1.1.1 (Lanfear et al. 2012) under the Bayes-
ian information criterion.
Phylogenetic relationships were assessed under a
Bayesian approach in MrBayes 3.2.0 (Ronquist and
Huelsenbeck 2003). Four independent analyses were
performed to reduce the chance of converging on a lo-
cal optimum. Each analysis consisted of 20 million
generations and four Markov chains with default heat-
ing values. Trees were sampled every 1,000 generations
resulting in 20,000 saved trees per analysis. Stationarity
was conrmed by plotting the –ln L per generation in the
program Tracer 1.6 (Rambaut et al. 2013). Additionally,
the standard deviation of the partition frequencies and the
potential scale reduction factor (Gelman and Rubin 1992)
were used as convergence diagnostics for the posterior
probabilities of bipartitions and branch lengths, respec-
tively. Adequacy of mixing was assessed by examining
the acceptance rates for the parameters in MrBayes and
the effective sample sizes (ESS) in Tracer. After analyz-
ing convergence and mixing, 2,000 trees were discarded
as “burn-in” from each run. We then conrmed that the
four analyses reached stationarity at a similar likelihood
score and that the topologies were similar, and used the
resultant 72,000 trees to calculate posterior probabilities
(PP) for each bipartition on a 50% majority rule consen-
sus tree.
Systematics
The taxonomic conclusions of this study are based on the
observation of morphological features and color patterns,
as well as inferred phylogenetic relationships. We con-
sider this information as species delimitation criteria fol-
lowing the general species concept of de Queiroz (1998,
2007).
Anolis poei sp. nov.
urn:lsid:zoobank.org:act:712687F6-CF33-4969-815D-E4600D01FB4C
Proposed standard English name: Telimbela anoles
Proposed standard Spanish name: Anolis de Telimbela
Holotype
QCAZ 3449 (Figs. 1, 2), adult male, Ecuador, Provincia
Bolívar, Telimbela, 01.65789ºS, 79.15334ºW, WGS84
1,354 m, 10 June 2011, collected by Fernando Ayala-Va-
rela, Jorge H. Valencia, Diana Troya-Rodríguez, Francy
Mora, and Estefanía Boada.
Paratypes (15)
ECUADOR: Provincia Bolívar: QCAZ 3444−3448,
3451−3455, 4359, same data as holotype, ex-
cept 0.1658440ºS, 79.157150ºW, 1,310 m; QCAZ
6781−6783 Telimbela, Escuela Elisa Mariño de Carva-
jal, 0.1665857ºS, 79.172096ºW, 27 July 2004, collected
by Edwin Carrillo-Ponce and Morley Read; QCAZ 9219
Guaranda, Salinas, Recinto Tres Cruces, 01.431380ºS,

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Ayala-Varela et al.
Fig. 3. Ontogenetic transformation of color and pattern in Andinophryne olallai from Río Manduriacu, Imbabura Province, Ecuador.
(A) Froglet (11 mm SVL; in situ), (B) Froglet (15.1 mm SVL; in situ), (C) Juvenile (26.3 mm SVL; in situ), (D) Juvenile (28.1 mm
SVL; in situ), (E) Adult (44.6 mm SVL; ex situ), (F) Adult (53.3 mm SVL; in situ). Note the progressive ontogenetic change in dor-
sal patterning from heavily mottled to no pattern; lack of parotoid glands and tubercles along the ank to presence of conspicuous
parotoid glands and tubercles along the ank; a darkening of color from copper, tan, and white to dark brown; and iris color change
from vibrant crimson to copper-orange.
Fig. 2. Anolis poei sp. nov. Holotype, adult male (SVL = 59.67 mm, QCAZ 3449, A), eye close-up (SVL = 60.31 mm, QCAZ 3448,
B), subadult male (SVL = 52.12 mm, QCAZ 3455, C, D), adult male (SVL = 59.02 mm, QCAZ 3451, E, F), adult male (SVL =
60.31 mm, QCAZ 3448, G, H). Photographs by L. Bustamante (A), and O. Torres-Carvajal (B, C, D, E, F, G, H).

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A new species of Anolis lizard from western Ecuador
Fig. 3. Male dewlap of Anolis poei sp. nov. (holotype, QCAZ 3449, A; paratype, QCAZ 3455, B); A. otongae (QCAZ 4661, C;
QCAZ 11791, D); and A. gemmosus (QCAZ 4385, E; QCAZ 4352, F; QCAZ 9452, G; QCAZ 11850, H). Photographs by L. Busta-
mante (A), O. Torres-Carvajal (B, C, D, E, F, H), and S. R. Ron (G).

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79.097970ºW, 2,628 m, 28 May 2009, collected by Eli-
cio E. Tapia, Silvia Aldás-Alarcón, and Eduardo Toral-
Contreras.
Diagnosis
We assign Anolis poei both to the aequatorialis series,
based on moderate to large body size, narrow toe lamel-
lae, small head scales, smooth ventral scales, and uni-
form dorsal scalation; and to the eulaemus-subgroup,
based on a typical Anolis digit, in which the distal la-
mellae of phalanx III distinctly overlap the rst proximal
subdigital scale of phalanx II (Williams 1976; Williams
and Duellman 1984; Castañeda and de Queiroz 2013).
At present ten species are recognized within the eulae-
mus-subgroup: Anolis anoriensis Velasco et al. 2010,
A. antioquiae Williams 1985, A. eulaemus Boulenger
1908, A. tchi Williams & Duellman 1984, A. gemmo-
sus O’Shaughnessy 1875, A. maculigula Williams 1984,
A. megalopithecus Rueda-Almonacid 1989, A. otongae
Ayala-Varela & Velasco 2010, A. podocarpus Ayala-
Varela & Torres-Carvajal 2010, and A. ventrimaculatus
Boulenger 1911. Anolis poei differs from them mostly
in dewlap features. The dewlap in males of A. poei has a
yellowish-green (or both yellow and green) gorgetal re-
Ayala-Varela et al.
gion, light blue border, and white sternal and marginal
regions (Fig. 3). It has a blackish gorgetal region, and
creamy white sternal region with light brown scales in A.
anoriensis; brown gorgetal region, and pale brown mar-
ginal region in A. eulaemus; bluish-gray gorgetal region,
orange stripes, pale bluish-rose anterior third, and white
sternal region becoming pale blue toward the belly in A.
maculigula; sepia background, with red narrow and ir-
regular stripes on each side of rows in A. megalophitecus;
white, pale yellow, or greenish-yellow gorgetal region,
with white or pale-yellow marginal and sternal regions
in A. otongae (Fig. 3); dull yellowish-green or light blue
gorgetal region, shading to dull cream, greenish yellow
or orange on the marginal region, with white or bluish
green gorgetal rows with or without brown spots and
with yellowish white, yellow or orange sternal region in
A. gemmosus (Fig. 3). The dewlap in males of A. poei
has wide rows of 3−7 scales separated by naked skin; the
width of these rows is one scale in A. tchi, 2−5 granular,
minute scales in A. podocarpus, 1–2 scales in A. ventri-
maculatus, 3−6 scales in A. otongae, and 2−3 scales in A.
gemmosus. In addition, females of the new species lack
a dewlap, which is present in females of A. anoriensis,
A. antioquiae, A. eulaemus, A. tchi, and A. podocarpus.
Anolis poei is most similar morphologically to A.
otongae and A. gemmosus (Fig. 4). From the former
species (character states in parenthesis) A. poei differs
in having small dorsal chevrons in females (large dorsal
chevrons extending onto anks), pale yellowish-brown
iris (iris dark blue), interparietal scale (if present) sur-
rounded by small swollen scales (interparietal scale
surrounded by relatively enlarged at scales), enlarged
postanal scales separated by 3−5 scales (postanal scales
separated by 1−2 scales), and in lacking a dark stripe on
side of head (dark coppery-brown stripe present). Ad-
ditionally, PCA analyses suggested that specimens of A.
poei have shorter jaws, as well as lower and narrower
heads than A. otongae (Table 2, Fig. 5), with PC1 (39%
of total variation) represented mainly by head height,
head width, and jaw length.
The new species can be distinguished from A. gem-
mosus (Table 3) in having fewer scales between sec-
ond canthals (11−14, mean = 12.08 and 12−21, mean
= 15.25, respectively; t = 5.31, P<0.005); fewer scales
between supraorbital semicircles (1−3, mean = 1.62 and
1−5, mean = 3.13, respectively; t = 4.46, P<0.005); more
lamellae under phalanges III-IV of fourth toe (18−19,
mean = 18.92 and 14−18, mean = 17.33, respectively;
t = -7.86, P<0.005); a narrower head (head width =
7.84−8.84, mean = 8.29 and 6.97−17.41, mean = 10.82,
respectively; t = -7.03, P<0.005); lower head (head
height = 6.54−7.48, mean = 6.92 and 5.42−15.96, mean =
9.51, respectively; t = -6.96, P<0.005); and shorter snout
(snout length = 6.75−7.30, mean = 6.92 and 5.79−14.95,
mean = 10.58, respectively; t = -11.74, P<0.005).
Species Voucher Locality GenBank Number
A. gemmosus QCAZ 4385 Ecuador, Car-
chi, Río San
Pablo near
Chical
ND2: KJ854205
COI: KJ854219
RAG1: KJ854212
QCAZ 4406 Ecuador, Car-
chi, Maldo-
nado, Teldibi
Ecological
Trail
ND2: KJ854206
COI: KJ854220
RAG1: KJ854213
A. otongae QCAZ 11790 Ecuador,
Pichincha,
Biological Re-
serve Otonga
ND2: KJ854207
RAG1: KJ854214
COI: KJ854221
QCAZ 11791 Ecuador,
Pichincha,
Biological Re-
serve Otonga
ND2: KJ854208
COI: KJ854222
RAG1: KJ854215
A. poei QCAZ 3444 Ecuador,
Bolívar,
Telimbela
ND2: KJ854209
COI: KJ854223
RAG1: KJ854216
QCAZ 3445 Ecuador,
Bolívar,
Telimbela
ND2: KJ854210
COI: KJ854224
QCAZ 3448 Ecuador,
Bolívar,
Telimbela
ND2: KJ854211
COI: KJ854225
RAG1: KJ854217
QCAZ 4359 Ecuador,
Bolívar,
Telimbela
RAG1: KJ854218
Table 1. Species of Anolis sequenced in this study, voucher
specimen numbers, collecting localities, and GenBank acces-
sion numbers.

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A new species of Anolis lizard from western Ecuador
Fig. 4. Part 1. Five species of Anolis from western Ecuador. A. aequatorialis: male (QCAZ 11861, A) and female (QCAZ 3443,
B); A. binotatus: male (QCAZ 3434, C, D); A. fasciatus: male (QCAZ 3450, E, F); A. otongae: male (QCAZ 11790, G) and female
(QCAZ 11791, H).

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Fig. 4. Part 2. A. gemmosus: male (QCAZ 4352, I, J), male (QCAZ 4385, K, L), male (QCAZ 11849, M, N), and female (QCAZ
4393, O, P). All photographs by O. Torres-Carvajal, except A, M, N (S. R. Ron).
Ayala-Varela et al.

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Description of holotype (scores for para-
types in parentheses)
Male (Figs. 1, 2); SVL 59.7 mm (46.5−60.3 mm); tail
length 150.2 mm (146.2−163.4 mm); head length 15.9
mm (14.8−16.5 mm); head width 8.4 mm (7.8−8.8 mm);
head height 7.2 mm (6.5−7.5 mm); internasal distance
2.0 mm (1.2−2.1 mm); interorbital distance 2.4 mm
(2.2−2.5 mm); interparietal absent (present, interparietal
length 0.8−0.9 mm; second largest scale length near in-
terparietal 0.3−0.4 mm); ear opening maximum length
1.6 mm (1.6−2.1 mm); snout length 6.8 mm (6.8−7.3
mm); jaw length 11.7 mm (11.7−14.4 mm); axila-groin
distance 27.7 mm (27.4−30.6 mm); femur length 14.8
mm (14.4−15.6 mm); 4th toe length 12.5 mm (10.6−12.8
mm); 4th toepad width 1.2 mm (1.0−1.3 mm); forelimb
length 36.2 mm (21.8−36.2 mm); hindlimb length 42.6
mm (42.6−52.7 mm).
Head scales multicarinate (same, unicarinate, or ru-
gose) on frontal region and unicarinate (same, multicari-
nate or rugose) on supraocular disc; 11 (10−14) scales
between second canthals; 13 (11−15) scales between rst
canthals; 6 (5−7) scales bordering the rostral posteriorly;
anterior nasal in contact with rostral (same or inferior
nasal in contact with rostral); supraorbital semicircles
separated by two (0−3) scales; supraocular disk with
scales heterogeneous in size; one elongate superciliary
followed by a series of granules (same or one small scale
instead of granules); 6 (5−8) loreal rows on left side; 49
(25−53) loreal scales; interparietal absent (same or, when
present, the interparietal smaller than ear opening, with
4−7 scales between interparietal and semicircles on each
side, and 8−15 scales between interparietal and nape
scales); suboculars in contact with supralabials; 6 (5−7)
supralabials counted up to a point below center of eye;
6 (5−7) infralabials counted up to a point below center
of eye; 7 (4−7) postmentals; one enlarged sublabial on
each side.
Dorsal crest or enlarged middorsal row absent; dorsal
scales keeled, 11 (9−11) dorsal scales in 5% the length
of SVL contained in the dorsal midline at the level of
the forelimbs; ank scales more or less separated by
skin; ventrals smaller than dorsals, 13 (8−13) longitudi-
nal rows in 5% the length of SVL; ventrals smooth and
granular, arranged in diagonal rows.
Toepads overlap the rst phalanx in all toes; 19
(18−19) lamellae under phalanges III and IV of fourth
toe (character 27 in Williams et al. 1995 and character
9 in Poe 2004); supradigitals multicarinate; tail with a
double row of middorsal scales; postanals present (same
or absent), with a slightly enlarged scale laterally on each
side.
Nuchal fold present (absent in females and juveniles);
dorsal folds absent; dewlap extending posteriorly to a
point halfway between fore and hindlimbs (absent in fe-
males); dewlap with ve longitudinal rows of 3−7 swol-
len scales, similar size to ventrals, separated by naked
skin.
Sexual variation of meristic and morphometric char-
acters in A. poei is presented in Table 4.
Color in life
Holotype (QCAZ 3449; Figs. 2, 3): background of head,
body, limbs and tail green; head with light bluish green,
dark green, and light grey irregular spots dorsally; dor-
sal surface of body with six light grey, small irregular
blotches; dorsal surface of neck with two light grey,
small irregular blotches; limbs with dark green and yel-
lowish-cream spots; lateral surface of head with a white
stripe extending posteriorly from loreal region, through
subocular region, to a point anterior to the tympanum;
white blotch with yellow center above tympanum; eye-
lids yellowish green with rst row of upper and lower
Fig. 5. Distribution of Anolis gemmosus, A. otongae and A. poei
sp. nov. along the rst and second principal components axes.
Raw Rotated
1 2 3 1 2 3
Head height -0.96 0.21 -0.02 -0.97 0.16 -0.07
Head length 0.24 0.34 0.05 0.22 0.35 0.05
Head width -0.96 0.20 -0.03 -0.96 0.15 -0.07
Jaw length 0.98 0.06 -0.06 0.98 0.11 -0.03
Snout length 0.82 0.33 -0.07 0.81 0.37 -0.06
Forelimb length -0.01 0.80 0.04 -0.05 0.80 0.00
Hindlimb length -0.01 0.85 0.02 -0.05 0.85 -0.03
Axilla-groin length -0.01 -0.50 -0.02 0.01 -0.50 0.01
Snout-vent length 0.06 -0.04 0.99 0.02 0.01 1.00
Eigenvalue 3.54 1.93 1.00 3.53 1.93 1.01
% var. explained 39.31 21.42 11.16 39.23 21.45 11.21
Table 2. PCA loadings conducted on nine morphological vari-
ables of Anolis gemmosus, A. otongae and A. poei.
A new species of Anolis lizard from western Ecuador

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palpebrals black, second and third rows both yellow and
green; lateral surface of neck with dark green dots; body
anks green, with rows of yellow-centered white spots
oriented ventro-posteriorly; ventral surface of head yel-
lowish green with light yellow blotches; ventral surface
of body white with bluish-green reticulations; ventral
surface of limbs white with several transparent scales
and dark brown reticulations; ventral surface of tail white
with dark green spots anteriorly, and yellowish-green
transverse bands posteriorly; dewlap skin light blue, dark
yellowish green on gorgetal region, light blue on ster-
nal region; gorgetal scales light yellowish green; mar-
ginals and sternals white; iris dark brown with a white
inner ring. When stressed, the dorsal background color
switched from green to yellowish brown.
Subadult male (QCAZ 3455, Figs. 2,3, differences
from holotype): head with dark green and white irregu-
lar small spots dorsally; dorsal surface of body and neck
with white and dark green small spots, and larger pale
yellow spots; lateral surface of head yellowish green with
a white stripe extending posteriorly from loreal region,
through subocular region, to upper border of tympanum;
lateral surface of body with rows of white and dark green
small spots, and larger pale yellow spots; ventral surface
of head with white blotches and light blue spots; ventral
surface of body with dark green reticulations; ventral sur-
face of limbs with brown or green reticulations; ventral
surface of tail with blackish green reticulations anteri-
orly; dewlap skin white, yellow on gorgetal region, white
on sternal region; throat, edge of mouth, and tongue pink-
ish white (Fig. 6). When stressed, rust-colored blotches
appeared on dorsal surface of head, body, limbs and tail.
Adult female (QCAZ 3454, Fig. 7): dorsal surface
of head, body and tail yellowish green; dorsal surface
of body with six narrow brown chevrons, each one de-
limited posteriorly by a grayish white blotch; limbs yel-
lowish green with dark green spots arranged in bands,
and pale yellowish spots; tail with two brown chevrons
anteriorly; lateral surface of head yellowish green; loreal
region yellow; lateral surface of neck and body yellowish
green with brown dots; ventral surface of head pale yel-
low with yellowish green reticulations, short white lon-
gitudinal stripe on throat; ventral surface of body and tail
white with black reticulations laterally; ventral surface
of limbs white with some transparent scales and brown
reticulations on hindlimbs; ventral surface of tail with
brownish green reticulations anteriorly; iris brown with
a pale white ring.
Subadult female (QCAZ 3446, Fig. 7, differences with
QCAZ 3454): occipital and temporal regions with brown
and white small blotches; dorsal surface of neck with a
distinct brown chevron delimited posteriorly by a grayish
white blotch; lateral surface of body yellowish green dor-
sally and light blue ventrally, with white or cream spots;
dorsal surface of tail with two brown chevrons, each one
delimited posteriorly by a grayish white blotch.
Fig. 6. Tongue of Anolis poei sp. nov., subadult male (QCAZ
3455, top); A. gemmosus, adult male (QCAZ 4347, middle); A.
otongae, adult male (QCAZ 4661, bottom). Photographs by S.
R. Ron (top), O. Torres-Carvajal (middle, bottom).
Color in preservative
Holotype (QCAZ 3449): dorsal background of head,
body, limbs and tail grayish brown; dorsal surface of
head with metallic green, dark green, blue, gray and
white cream irregular spots; dorsal surface of body with
six black small chevrons, each delimited posteriorly by a
white irregular blotch; limbs with dark brown and white
spots; lateral surface of head with a white stripe extend-
ing posteriorly from loreal region, through subocular re-
gion, to a point anterior to the tympanum; upper border
of tympanum with a white spot; eyelids purple with rst
row of upper and lower palpebrals black, second and
third rows white and purple; neck anks with black dots;
body anks grayish brown, with dark brown diagonal
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Character A. gemmosus A. poei sp. nov. F-value P t-value P
Scales between second canthals 12–21 (24) 15.25 ± 1.98 11–14 (13) 12.08 ± 1.12 2.59 0.12 5.31 <0.005
Postrostrals 5–7 (24) 5.79 ± 0.72 5–7 (13) 5.92 ± 0.64 1.37 0.25 -0.55 0.59
Row of loreals 6–10 (24) 7.25 ± 1.15 5–8 (13) 6.31 ± 1.18 0.03 0.86 2.36 0.02
Scales between supraorbital semicircles 1–5 (24) 3.13 ± 1.23 1–3 (13) 1.62 ± 0.77 5.27 0.03 4.46 <0.005
Scales between interparietal (if present) and
semicircles 3–8 (24) 5.67 ± 1.27 4–7 (6) 5.83 ± 1.17 0.18 0.67 -0.29 0.77
Supralabials 5–7 (24) 6.08 ± 0.50 5–7 (13) 6 ± 0.41 1.29 0.27 0.51 0.61
Postmentals 4–8 (24) 6.13 ± 1.03 4–7 (13) 5.77 ± 0.93 0.18 0.67 1.03 0.31
Lamellae under phalanges III-IV of fourth toe 14–18 (24) 17.33 ± 0.92 18–19 (13) 18.92 ± 0.28 8.71 0.01 -7.86 <0.005
Head length 13.23–18.12 (94) 15.46 ± 1.07 14.79–16.5 (7) 15.67 ± 0.51 4.67 0.03 0.93 0.37
Head width 6.97–17.41 (94) 10.82 ± 3.24 7.84–8.84 (7) 8.29 ± 0.36 32.16 <0.005 -7.03 <0.005
Head height 5.42–15.96 (94) 9.51 ± 3.32 6.54–7.48 (7) 6.92 ± 0.38 31.04 <0.005 -6.96 <0.005
Jaw length 7.31–17.43 (94) 12.32 ± 3.02 11.73–14.36 (7) 12.44 ± 0.91 19.25 <0.005 0.26 0.80
Snout length 5.79–14.95 (94) 10.58 ± 2.93 6.75–7.30 (7) 6.92 ± 0.19 41.30 <0.005 -11.74 <0.005
Forelimb length 23.41–34.34 (94) 29.43 ± 2.28 21.84–36.18 (7) 28.57 ± 4.25 0.02 0.89 -0.12 0.90
Hindlimb length 41.51–63.80 (94) 52.82 ± 4.13 42.56–52.68 (7) 49.01 ± 3.33 1.00 0.32 -2.38 0.02
Axilla-groin length 20.73–33.51 (94) 26.74 ± 2.07 27.35–30.61 (7) 28.54 ± 1.30 0.95 0.33 2.26 0.03
Snout-vent length 46.71–66.21 (94) 58.34 ± 3.65 46.47–60.31 (7) 56.87 ± 4.85 0.35 0.56 -1.00 0.32
Tail length 94.94–191 (94) 154.59 ± 18.66 146.21–163.37 (7) 154.74 ± 6.32 3.82 0.05 0.02 0.98
Table 3. Summary of morphological characters of Anolis poei sp. nov. and A. gemmosus from Ecuador. For each quantitative character, the F-value,
t-value, and corresponding P-values are given. Range and sample size (in parenthesis) followed by mean ± standard deviation are given.
Character Males Females
n = 4 n = 3
Scales between second canthals 11–13 11.75 ± 0.96 12–13 12.67 ± 0.58
Postrostrals 5–6 5.75 ± 0.5 6–7 6.33 ± 0.58
Row of loreals 6–8 7 ± 1.15 5–6 5.33 ± 0.58
Scales between supraorbital semicircles 1–2 1.75 ± 0.5 1–2 1.67 ± 0.577
Scales between interparietal and semicircles Interparietal absent 6–7 6.50 ± 3.78
Supralabials to below center of eye 6 6
Postmentals 4–7 5.25 ± 1.5 6–7 6.33 ± 0.58
Lamellae under phalanges II-III of fourth toe 19 19
Head length 15.8–16.5 15.95 ± 0.38 14.8–15.62 15.29 ± 0.44
Head width 7.84–8.84 8.31 ± 0.41 8.05–8.66 8.26 ± 0.34
Head height 6.67–7.48 7.02 ± 0.39 6.54–7.27 6.8 ± 0.41
Jaw length 11.73–12.65 12.25 ± 0.38 11.86–14.36 12.70 ± 1.43
Snout length 6.75–7.04 6.87 ± 0.12 7.82–7.30 7 ± 0.26
Forelimb length 27.94–36.18 30.75 ± 3.72 21.84–28.19 25.68 ± 3.37
Hindlimb length 42.56–52.68 49.35 ± 4.59 47.50–49.56 48.57 ± 1.03
Axilla-groin length 27.35–28.17 27.76 ± 0.33 27.94–30.61 29.57 ± 1.43
Snout-vent length 58.80–60.31 59.45 ± 0.68 46.47–58.48 53.43 ± 6.22
Tail length 150.20–163.37 157.89 ± 5.89 146.21–155.38 150.53 ± 4.60
Table 4. Sexual variation in lepidosis and measurements (mm) of Anolis poei sp. nov. Range followed by mean ± standard devia-
tion are given.
A new species of Anolis lizard from western Ecuador

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Fig. 7. Anolis poei sp. nov. Adult female (SVL = 46.47 mm, QCAZ 3454, A, B), subadult female (SVL = 47.99 mm, QCAZ 3446,
C, D), juvenile male (SVL = 26.85 mm, QCAZ 3453, E, F). Photographs by O. Torres-Carvajal.
bands oriented ventro-posteriorly and intercalated with
white spots; ventral surface of head white with light blue
reticulations; ventral surface of body white with faint
grayish purple reticulations; ventral surface of limbs
grayish cream with dark brown reticulations; ventral sur-
face of tail white anteriorly with a metallic green tint and
grayish purple spots, and gray posteriorly; dewlap skin
with a turquoise gorgetal region and white sternal region;
gorgetal scales light brown with a gold tint internally, and
dark brown externally; dewlap marginals and sternals
white; throat, edge of mouth and tongue white.
Adult male (QCAZ 6783): dorsal surface of head and
body dark brown with gray dots; dorsal surface of limbs
dark brown, with gray dots on forelimbs; lateral surface
of head dark brown with white cream dots dorsal and an-
terior to tympanum; body anks dark brown with faint
white dots arranged on diagonal lines that reach venter;
ventral surface of head with bluish-purple infralabial
and sublabial regions, and light purple gular region with
white irregular spots; ventral surface of body white with
faint purple reticulations; limbs creamish gray with dark
brown reticulations; ventral surface of tail white with
Ayala-Varela et al.

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purple mottling anteriorly, and gray posteriorly; dewlap
skin with a light blue gorgetal region and white ster-
nal region; gorgetal scales purple; sternal and marginal
scales white.
Adult female (QCAZ 3454): dorsal surface of head
brown with metallic blue and green frontal and supraocu-
lar regions; dorsal surface of body brown with six nar-
row black chevrons, each one delimited posteriorly by
a white blotch; forelimbs bluish brown with white spots
arranged in stripes; hindlimbs brown with dark brown
bands and dots; tail with two black chevrons anteriorly;
lateral surface of head brown with purple tint; labial re-
gion light purple; lateral aspect of neck and body purple
with black dots; ventral surface of head white with purple
brown stripes; ventral surface of body white with dark
brown dots laterally; ventral surface of limbs grayish
cream with dark brown reticulations on hindlimbs; ven-
tral surface of tail white with dark brown dots.
Subadult female (QCAZ 3446, differences with
QCAZ 3454): occipital and temporal regions with dark
brown, small blotches; dorsal surface of neck with a dis-
tinct dark brown chevron; dorsal surface of body with six
distinct, dark brown chevrons; dorsal surface of tail with
two dark brown chevrons.
Phylogenetic relationships
The data matrix analyzed in this study contained 1,065
unique site patterns. Of the 2,807 nucleotide characters
included in our analysis 1,703 were constant, 224 par-
simony uninformative, and 880 were parsimony infor-
mative. The 50% majority rule consensus tree resulting
from the Bayesian analysis (Fig. 8) is generally congru-
ent with the phylogeny of the clade Dactyloa presented
by Castañeda and de Queiroz (2011). Both the new spe-
cies described here and A. otongae are members of the
aequatorialis series of Castañeda and de Queiroz (2013),
which corresponds roughly to the “western clade” of
Castañeda and de Queiroz (2011). Our phylogeny sup-
ports strongly (PP = 0.99) a sister taxon relationship be-
tween Anolis poei and A. gemmosus, as well as the ex-
clusivity (de Queiroz and Donoghue 1990; de Queiroz
1998) of both species. They form a clade sister (PP =
0.89) to A. otongae. The clade formed by the three spe-
cies is sister (PP = 1) to a clade formed by A. aequatoria-
lis and A. anoriensis.
Distribution and ecology
Anolis poei inhabits low montane evergreen forest (Sierra
1999) on the western slopes of the Andes in central Ecua-
dor, Provincia Bolívar, between 1,310–1,354 m (Fig. 9).
Fig. 8. Phylogeny of the “western Dactyloa clade” sensu
Castañeda and de Queiroz (2011), which is part of the ae-
quatorialis series of Castañeda and de Queiroz (2013), and
representatives of the heterodermus series (A. euskalerri-
ari), punctatus series (A. transversalis), roquet series (A. lu-
ciae), latifrons series (A. agassizi), and a non-Dactyloa Ano-
lis (A. occultus). The tree is a majority rule (50%) consensus
tree of 72,000 trees obtained from a Bayesian analysis of the
mitochondrial genes COI, ND2, and adjacent tRNAs, and
the nuclear gene RAG1. Asterisks correspond to posterior
probability values ≥ 0.99. Voucher information is presented
in Castañeda and de Queiroz (2011) and Table 1.
A new species of Anolis lizard from western Ecuador

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The new species occurs in sympatry with A. aequa-
torialis, A. binotatus, and A. fasciatus at its type locality
(Fig. 4). Specimens of the new species were collected
along the border of a road, close to rivers, in second-
ary forest, and on shrubs within pastures. All individu-
als were found between 20h00 and 22h00 sleeping with
their heads up, or in a horizontal position on branches or
vines, 0.5–4.5 m above ground or streams. The smallest
individual QCAZ 3453 (SVL = 26.9 m; TL = 67.6 mm)
was collected on 11 June 2011.
Etymology
The specic name is a noun in the genitive case and is
a patronym for Steve Poe, who has published important
contributions to the systematics and evolution of Ano-
lis lizards (Poe 2004, 2011). During his collecting trips
to Ecuador in 2009 and 2010, Poe trained several young
herpetologists in eld collecting techniques and inspired
them to explore the diversity of anole lizards. This paper
is one of the products resulting from that inspiration.
Acknowledgments.—We thank Jorge H. Valencia,
Francy Mora, and Estefanía Boada for assistance in the
eld; Santiago Ron and Lucas Bustamante for the pho-
tographs; Paulina Santiana and Andrea Varela for assem-
bling some of the gures; Melissa Rodriguez for helping
with the map. Special thanks to Kevin de Queiroz and
two anonymous reviewers for commenting on previous
versions of this manuscript. OTC received funds from
Secretaría de Educación Superior, Ciencia, Tecnología
e Innovación (SENESCYT). Specimens were collected
under collection permit 001-IC-FAU/FLO/DRZCHI/MA
and 008-09 IC-FAU-DNB/MA issued by Ministerio de
Ambiente del Ecuador.
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Received: 28 April 2014
Accepted: 20 May 2014
Published: XX May 2014
A new species of Anolis lizard from western Ecuador

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Appendix 1
Additional specimens examined
Anolis gemmosus – Ecuador: Carchi: Chilma Bajo, Finca de Aníbal Pozo, 0.86397°N, 78.04723°W, 2,022 m, QCAZ
8681-82; Chilma Bajo, Finca de Aníbal Pozo, 0.86495°N, 78.04979°W, 2,071 m, QCAZ 8683; La Centella, 0.89318°N,
78.13471°W, 1,800-2,400 m, QCAZ 11784; Maldonado, Sendero Ecológico Teldibi, 0.91301°N, 78.10782°W, 1,477-
1,635 m, QCAZ 12272, 12278, QCAZ 12279-80, 4360, 4406, 4408; Río San Pablo, cerca a Chical, 0.90302°N,
78.16284°W, 1,399 m, QCAZ 4377, 4382, 4385-86,4388; Río San Pablo, cerca a Chical, 0.90327°N, 78.16201°W,
1,429 m, QCAZ 4393; Río Verde and Río Pablo, Río Estrellita, Guapil, 1,428-1,466 m, QCAZ 12289, 12294, 12302;
Cotopaxi: 115 km Oeste de Pilaló, 0.928°S, 79.057°W, 1,500 m, QCAZ 4072; 18.2 km de Quillutuña, vía a Pucayacu,
0.67843°S, 79.01565°W, 1,420 m, QCAZ 8845-49; Alrededores de San Francisco de Las Pampas, 0.42371°S,
78.96765°W, 1,800 m, QCAZ 1440-47, 2123; Bosque Integral Otonga , 0.4194°S, 79.00345°W, 1,720-2,143 m, QCAZ
2758, 2809-10, 3121, 3126-27, 3131, 3133, 3174, 3180-90, 3863-3866, 3869-71, 3940, 3974-76, 4028-34, 4224-25,
4657, 4663, 4785, 5060, 5063, 5371, 5477-79, 5482-83, 6770-73, 9888, 10424, 10438-39, 10441-42, 10452, 12057,
12060-65, 12067, 12072-73, 12075, 12077-82, Bosque Integral Otonga, a lo largo del río Esmeraldas, 0.46333°S,
79.05027°W, QCAZ 7281-89; Bosque Integral Otonga, alrededores de la estación, 0.41933°S, 79.00336°W, 1,980
m, QCAZ 10697; Bosque Integral Otonga, arriba de la estación, 0.41478°S, 79.00073°W, QCAZ 3867-68; Bosque
Integral Otonga, orillas del río Esmeraldas, 0.41932°S, 78.99396°W, 1,719 m, QCAZ 10393, 10395, 10399; Bosque
Integral Otonga, sendero a la Estación, 0.41933°S, 79.00336°W, 1,646 m, QCAZ 10696; Cerca a Naranjito,
0.41944°S, 79.00333°W, QCAZ 7825; San Francisco de Las Pampas, 0.42371°S, 78.96765°W, 1,600-1,800 m,
QCAZ 63, 68-70, 72-79, 3134-53, 3155, 3175; Vía a Otonga, 0.33183°S, 78.93791°W, 1,476-1,700 m, QCAZ 8412;
Imbabura: 6 de Julio de Cuellaje, 0.4°N, 78.525°W, QCAZ 4346-47; 6 de Julio de Cuellaje, 0.40107°N, 78.5181°W,
1,886 m, QCAZ 4349; 6 de Julio de Cuellaje, 0.40102°N, 78.51779°W, 1,897 m, QCAZ 4350; 6 de Julio de Cuel-
laje, punto 8, 0.4°N, 78.525°W, QCAZ 4348; 6 de Julio de Cuellaje, San Antonio, Cordillera de Toisán, 0.45803°N,
78.54722°W, QCAZ 9450-53; Carretera nueva vía a Cuellaje, Sector de Santa Clara, Reserva Alto Choco, 0.37603°N,
78.45857°W, 2,062 m, QCAZ 4352-54; La Mina, Junín, 0.2754°N, 78.6603°W, 1,715 m, QCAZ 3071; Manduriaco,
0.277°N, 78.873°W, 1,330 m, QCAZ 5328; Manduriacu, 7.5 km NE of Bellavista, 0.31006°N, 78.85757°W, 1,177-
1,227 m, QCAZ 11606, 12305-314; 12322, 12324, 12326, 12328, 12331; Reserva Siempre Verde, NE de Cotacachi,
0.37167°N, 78.42186°W, 2,468 m, QCAZ 8837; Reserva Alto Choco, Santa Rosa, 0.36939°N, 78.44942°W, 2,109
m, QCAZ 7330-31; Pichincha: 1-2 km oeste de Tandayapa, 0.004°S, 78.663°W, 2,000 m, QCAZ 2070-71; 2.9 km de
Tandayapa, 0.00952°S, 78.65698°W, 1,820 m, QCAZ 406-10; 5 km E Tandayapa, 0.02°S, 78.651°W, 1,975 m, QCAZ
2066-69; A orillas del Río Chisinche, en la carretera a Conchacato, 0.448°S, 78.76423°W, 1,693 m, QCAZ 6884-89;
30 km E de Santo Domingo, hacia la Reserva de Bosque Integral Otonga, 0.3884°S, 78.92995°W, QCAZ 9769-70;
9775; Bosque Protector Mindo - Nambillo, refugio, 0.106°S, 78.687°W, 1,700 m, QCAZ 2910; Cooperativa El Porve-
nir, nca El Cedral, 0.114°N, 78.56993°W, 2,297 m, QCAZ 10501-502; Desviación a Mindo, 1-5 km de la intersec-
ción hacia abajo, 0.02853°S, 78.75861°W, 1,661 m, QCAZ 9724-31; Estación Cientíca Río Guajalito, 0.22676°S,
78.82171°W, 1,791-1,814 m, QCAZ 1330, 1333, 1500, 1645, 2682-84, 2786, 2813, 2815-16, 3040-45, 3056-57, 3373,
3385, 4123-25, 4210, 4214, 6413-14, 8859, 8864-65, 9974, 11404, 12088-101; Las Tolas, 0.72818°N, 78.77792°W,
1,200-1,600 m, QCAZ 11848-49; Manuel Cornejo Astorga (Tandapi), frente a la planta de agua potable "El Placer" vía
a Conchacato, 0.42471°S, 78.78905°W, 1,500 m, QCAZ 6882; Manuel Cornejo Astorga (Tandapi), vía Atenas a 5 km
de la carretera principal, 0.40625°S, 78.83621°W, 1,671 m, QCAZ 5365-70; Mindo, 1,342-1,560 m, QCAZ 12350-
53, 12356, 12358, 12365, 12370, 12375-76; Mindo Biology Station, 0.07805°S, 78.73194°W, QCAZ 7518-20, 7522;
Mindo, camino entre Mariposas de Mindo y Mindo Garden, 0.06753°S, 78.7535°W, 1,361 m, QCAZ 6851-53, 6858;
Mindo Garden, 4 km de Mindo, 0.06901°S, 78.80166°W, QCAZ 2787; Mindo, El Monte, Road to Mindo Garden,
0.07805°S, 78.7319°W, QCAZ 7521; Mindo, Sachatamia Lodge, 0.02638°S, 78.75944°W, 1,700 m, QCAZ 11857-59;
Nanegalito, Finca El Cedral, 0.1141°N, 78.57007°W, 2,272 m, QCAZ 9462-63; Pachijal, vía Nanegalito-Los Bancos,
0.13°S, 78.72644°W, 1,741 m, QCAZ 5494-500; Palmeras, 0.244°S, 78.794°W, 1,800 m, QCAZ 871, 881-83,1351-
52, 2244, 3004-06; Recinto Chiriboga, Estación La Favorita , 0.21307°S, 78.78421°W, 1,680 m, QCAZ 5383-84;
Reserva Ecológica Bosque Nublado "Santa Lucía," 0.11928°N, 78.59647°W, 1,624-1,927 m, QCAZ 10664, 11850-52,
11888-93,11897, 11899; Tandayapa, 0.00591°N, 78.67455°W, 1,670 m, QCAZ 4086. Locality in error: Pichincha, San
Antonio de Pichincha, 0.00905°S, 78.44581°W, QCAZ 724.
Anolis otongae – Ecuador: Cotopaxi: Alrededores de San Francisco de Las Pampas, 0.42371°S, 78.96765°W, 1,800
m, QCAZ 2128; Bosque Integral Otonga , 0.41944°S, 79.00333°W, 1,900-2,300 m, QCAZ 1721, 2050-52, 3129, 3706,
3796, 3872-73, 4025, 4661, 5481, 6219, 11790-91, 12035, 12056, 12058, 12070-71; Los Libres, QCAZ 2781; Peñas
Coloradas, 0.52343°S, 79.05908°W, QCAZ 1696; Pichincha: La Victoria, 0.47747°S, 79.05336°W, 2,104 m, QCAZ
6394-96.
Ayala-Varela et al.

(24)Amphib. Reptile Conserv. | amphibian-reptile-conservation.org May 2014 | Volume 8 | Number 1 | e76
Fernando Ayala-Varela is the director of the herpetology collection at the Ponticia Universidad Católica
del Ecuador in Quito. He received his diploma at the Ponticia Universidad Católica del Ecuador, Quito in
2004. He has been interested in herpetology since childhood and has dedicated a lot of time studying the
lizards of Ecuador, specically the taxonomy and ecology of Anolis species. His current research interests
include reproductive biology and ecology of lizards and snakes in Ecuador.
Diana Troya-Rodríguez received a B.Sc. in Biology from Ponticia Universidad Católica del Ecuador
(PUCE) in 2013. As a student, she joined the Museo de Zoología QCAZ, Ponticia Universidad Católica
del Ecuador in Quito, where she developed a great interest in reptiles. She has been studying anole lizards
for the last four years. For her undergraduate thesis, Diana worked on the “Comparative phylogeography
of two sympatric species of Anolis (Squamata: Iguanidae) and the impact of global warming on their dis-
tribution.”
Omar Torres-Carvajal graduated in Biological Sciences from Ponticia Universidad Católica del Ecua-
dor (PUCE) in 1998, and in 2001 received a Master’s degree in Ecology and Evolutionary Biology from
the University of Kansas under the supervision of Dr. Linda Trueb. In 2005 he received a Ph.D. degree
from the same institution with the thesis entitled “Phylogenetic systematics of South American lizards
of the genus Stenocercus (Squamata: Iguania).” Between 2006–2008 he was a postdoctoral fellow at the
Smithsonian Institution, National Museum of Natural History, Washington DC, USA, working under the
supervision of Dr. Kevin de Queiroz. He is currently Curator of Reptiles at the Zoology Museum QCAZ of
PUCE and an Associate Professor at the Department of Biology in the same institution. He has published
more than 30 scientic papers on taxonomy, systematics and biogeography of South American reptiles,
with emphasis on lizards. He is mainly interested in the theory and practice of phylogenetic systematics,
particularly as they relate to the evolutionary biology of lizards.
Xiomara Talero-Rodríguez is an undergraduate biology student at Ponticia Universidad Católica del
Ecuador in Quito. She joined Museo de Zoología QCAZ last year and has been helping with several anole
lizard projects ever since. She is currently interested in studying ecology and behavior of anoles.
A new species of Anolis lizard from western Ecuador
.
In accordance with the International Code of Zoological Nomenclature new rules and regulations (ICZN 2012), we have deposited this paper in publicly accessible institutional libraries.
Digital archiving of this paper are found at the following institutions: ZenScientist (http://www.zenscientist.com/index.php/ledrawer); American Museum of Natural History, New York,
New York (USA); Ernst Mayr Library, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (USA); Smithsonian Institution Libraries, Washington, D.C.
(USA); Florida Museum of Natural History, Gainesville, Florida (USA); Instituto Nacional de Pesquisas da Amazônia, Manaus (BRAZIL); Universidade Federal do Rio de Janeiro, Rio
de Janeiro (BRAZIL).
The new species described herein has been registered in ZooBank (Polaszek 2005a, b), the ofcial online registration system for the ICZN. The ZooBank publication LSID (Life Science
Identier) for the new species described herein can be viewed through any standard web browser by appending the LSID to the prex ‘‘http://zoobank.org/’’. The LSID for this publication
is: urn:lsid:zoobank.org:pub:61380956-F1AC-46C0-84F3-C1ED545C46DC.
Separate print-only edition of paper(s) (reprint) are available upon request as a print-on-demand service. Please inquire by sending a request to: Amphibian & Reptile Conservation
(amphibian-reptile-conservation.org; arc.publisher@gmail.com).
Complete journal archiving is found at: ZenScientist (http://www.zenscientist.com/index.php/ledrawer); American Museum of Natural History, New York, New York (USA); Florida Mu-
seum of Natural History, Gainesville, Florida (USA); Instituto Nacional de Pesquisas da Amazônia, Manaus (BRAZIL); Universidade Federal do Rio de Janeiro, Rio de Janeiro (BRAZIL).
Amphibian & Reptile Conservation is a Content Partner with the Encyclopedia of Life (EOL); http:///www.eol.org/ and submits information about new species to the EOL freely.
Citations
ICZN. 2012. Amendment of Articles 8,9,10,21 and 78 of the International Code of Zoological Nomenclature to expand and rene methods of publication. Zootaxa 3450: 1–7.
Polaszek A et al. 2005a. Commentary: A universal register for animal names. N a t u r e 437: 477.
Polaszek A et al. 2005b. ZooBank: The open-access register for zoological taxonomy: Technical Discussion Paper. Bulletin of Zoological Nomenclature 62(4): 210–220.