Corresponding author: Omar Torres-Carvajal (
Academic editor: J. Penner
The discovery of three new species of
Reportamos el descubrimiento de tres especies nuevas de
Torres-Carvajal O, Venegas PJ, Queiroz K (2015) Three new species of woodlizards (Hoplocercinae,
The iguanian lizard clade
With nearly 40% of the total number of species described in the last seven years from Ecuador and Peru (
Snout-vent length (SVL) and tail length (TL) measurements were made with a ruler and recorded to the nearest millimeter. All other measurements (i.e., head width, length and height; rostral and mental width and height) were made with digital calipers and recorded to the nearest 0.1 mm. Sex was determined by noting the presence of hemipenes or sexually dichromatic characters. The format of
Following laboratory protocols similar to those presented by
Vouchers, locality data, and GenBank accession numbers of new DNA sequences obtained for this study.
Taxon | Voucher | Locality | GenBank number (ND4) | GenSeq nomenclature |
---|---|---|---|---|
|
QCAZ 8073 (holotype) | Ecuador: Esmeraldas: Alto Tambo, 5 km on road to Placer |
|
genseq-1 |
|
QCAZ 8395 | Ecuador: Zamora-Chinchipe: Chito, sector Los Planes |
|
genseq-2 |
|
QCAZ 8428 | Ecuador: Zamora-Chinchipe: Chito |
|
genseq-2 |
|
QCAZ 8515 | Ecuador: Zamora-Chinchipe: Chito, sector Los Planes |
|
genseq-2 |
|
QCAZ 8035 | Ecuador: Orellana: 66 km on road Pompeya-Iro |
|
genseq-4 |
|
CORBIDI 647 (holotype) | Peru: San Martín: Río Lejía on the trail La Cueva-Añazco Pueblo |
|
genseq-1 |
Editing, assembly, and alignment of sequences were performed with Geneious 7.1.7 (
Phylogenetic relationships were assessed under a Bayesian inference approach using MrBayes 3.2.2 (
The taxonomic conclusions of this study are based on the observation of morphological features and color patterns, as well as the inferred phylogenetic relationships. This information is considered as species delimitation criteria following a general lineage or unified species concept (
Holotype (QCAZ 8073, adult male, SVL = 119 mm) of
Male (Fig.
Vertebral crest strongly projected and decreasing in size posteriorly, with vertebrals on neck at least four times higher than those between hind limbs; crest bifurcates at a point approximately 10 mm posterior to the cloaca, and extends onto tail about 1/3 its length; body flanks between fore and hind limbs with slight dorsolateral fold; scales on dorsolateral fold slightly larger than adjacent scales; dorsal and flank scales small, smooth, imbricate, more or less homogeneous in size; ventral scales imbricate, keeled, rectangular or rhomboid, with a posterolateral mucron; ventrals more than twice the size (area) of dorsals.
Limb scales keeled dorsally and keeled or feebly keeled ventrally; scales on dorsal and posterior aspects of thighs heterogeneous in size, with most scales less than half the size of those scales on anterior and ventral aspects, separated from each other by skin covered with tiny granular scales; subdigitals on finger IV 25; subdigitals on toe IV 29; femoral pores on each side one; tail laterally compressed and gradually tapering posteriorly; caudal scales smooth at the base of tail, becoming keeled and imbricate towards tip, gradually increasing in size posteriorly on lateral and dorsal aspects of each caudal segment; caudals larger ventrally than dorsally; individual caudal segments three scales long ventrally and seven scales long dorsally.
(Fig.
Variation in meristic and morphometric characters of
Paratype (QCAZ 6671, adult female, SVL = 132 mm) of
Summary of morphological characters and measurements (mm) of
Character | |||
---|---|---|---|
Dorsals in transverse row between dorsolateral crests at midbody | 39–40 |
28–35 |
22–26 |
Ventrals in transverse row at midbody | 31–33 |
23–29 |
23–26 |
Vertebrals from occiput to base of tail | 50–51 |
43–62 |
51–57 |
Gulars | 47 | 30–35 |
36 |
Infralabials | 11 | 9 | 9–11 |
Supralabials | 13 | 10–12 |
9–12 |
Canthals | 5 | 5–6 |
5 |
Superciliaries | 14–17 |
13–18 |
13–15 |
Transverse rows of ventrals between fore and hind limbs | 47–49 |
38–46 |
37–40 |
Subdigitals finger IV | 23–25 |
15–20 |
18–19 |
Subdigitals toe IV | 27–29 |
24–27 |
22–27 |
Femoral pores | 1–2 |
0–3 |
3–4 |
Tail length/Total length | 0.57–0.60 |
0.59–0.71 |
0.60–0.61 |
Distribution of
The specific epithet is a noun in apposition and refers to Alto Tambo, Provincia Esmeraldas, Ecuador, a village on the Ibarra-San Lorenzo road where
Although previously referred to
Holotype of
Holotype of
Paratypes of
Close-up of left dorsum of
The only other species of
Male (Fig.
Vertebral crest strongly projected and decreasing in size posteriorly, with vertebrals on neck at least three times higher than those between hind limbs; crest bifurcates at a point approximately 10 mm posterior to the cloaca, and extends onto tail about ¼ its length; flanks between fore and hind limbs with dorsolateral and ventrolateral longitudinal folds, as well as several oblique folds; axillary region with three vertical folds; scales on dorsolateral folds slightly larger than adjacent scales giving the fold the appearance of a low crest; scales between dorsolateral folds and vertebral crest heterogeneous in size, prominently keeled, and imbricate, with largest scales twice as large as smallest ones; neck and scapular region with scattered, large conical scales; flank scales ventral to dorsolateral folds similar to those dorsal to folds, with largest scales four times as large as smallest ones (Fig.
Limb scales keeled dorsally and ventrally, homogeneous in size on fore limbs; scales on dorsal and posterior aspect of thighs heterogeneous in size, with most scales less than half the size of those on anterior and ventral aspects; scales on dorsal surface of shanks heterogeneous in size, with granular scales between large keeled scales; subdigitals on finger IV 17; subdigitals on toe IV 25; three femoral pores on left leg, two on right leg; tail laterally compressed and gradually tapering posteriorly; caudal scales strongly keeled and imbricate, increasing in size posteriorly on lateral and dorsal aspects of each caudal segment; caudals larger ventrally than dorsally; individual caudal segments three scales long ventrally and six scales long dorsally.
(Fig.
Variation in meristic and morphometric characters of
This species has a marked sexual dichromatism in background colors (green in males, brown in females; see Fig.
Adult females share similar color patterns with juveniles (Fig.
The specific epithet
Holotype of
Male (Fig.
Vertebral crest strongly projected and decreasing in size posteriorly, with vertebrals on neck at least four times higher than those between hind limbs; crest bifurcates posteriorly and extends onto tail less than ¼ its length; body flanks between fore and hind limbs with slight dorsolateral and ventrolateral longitudinal folds; scales on dorsolateral folds similar in size to adjacent scales; dorsal and flank scales small, keeled, imbricate, more or less homogeneous in size, and separated from each other by skin covered with tiny granular scales; ventral scales imbricate, smooth or slightly keeled, rectangular or rhomboid, with a posterolateral mucron; ventrals more than twice the area of dorsals.
Limb scales keeled dorsally and smooth or slightly keeled ventrally; scales on dorsal and posterior surfaces of thighs heterogeneous in size, with most scales less than half the size of those on anterior and ventral surfaces, separated from each other by skin covered with tiny granular scales; subdigitals on finger IV 17; subdigitals on toe IV 25; femoral pores on each side four; tail laterally compressed and gradually tapering posteriorly; caudal scales strongly keeled and imbricate, not gradually increasing in size posteriorly on lateral and dorsal aspects of each caudal segment; caudals larger ventrally than dorsally; individual caudal segments three scales long ventrally and six scales long dorsally.
(Fig.
Variation in meristic and morphometric characters of
Individuals of
The specific epithet is a noun in the genitive case and is a patronym honoring Sophia Rothschild in recognition of her financial support for the improvement of the herpetological collection of CORBIDI through the BIOPAT Program.
The phylogenetic tree inferred in this study (Fig.
50% Majority rule consensus tree of hoplocercine lizards (
The following key is artificial in the sense that its structure does not necessarily reflect the order of branching in the phylogeny.
1 | Dorsal head scales flat, smooth, juxtaposed; vertebral crest absent or composed of a discontinuous row of enlarged scales that are longer than tall |
|
– | Dorsal head scales conical; vertebral crest present, composed of projecting scales that are taller than long |
|
2 | Tail depressed, short (tail length < snout-vent length), with enlarged spiny scales dorsally and laterally |
|
– | Tail nearly round, moderate (tail length > snout-vent length), with rings of enlarged spiny scales |
|
3 | Vertebral region of trunk without enlarged scales; tail with three scale rows separating the spiny whorls ventrally |
|
– | Some vertebral scales in trunk region enlarged forming a discontinuous longitudinal row; tail with two scale rows separating the spiny whorls ventrally |
|
4 | Usually two femoral pores on each leg; two postmentals; females without streaks on throat |
|
– | Femoral pores 3–4 on each leg; usually four postmentals; females with dark streaks on throat |
|
5 | Caudal scales homogeneous in size within each autotomic segment |
|
– | Caudal scales increase in size posteriorly within each autotomic segment |
|
6 | Gular region in males white with a black medial patch |
|
– | Gular region in males orange or dirty cream, with longitudinal brown, reddish-brown, bluish, or orange streaks, and a large brown or black medial blotch at the level of the gular fold |
|
7 | Lateral superciliary projection present; vertebral crest usually discontinuous (absent on posterior part of neck) |
|
– | Lateral superciliary projection absent; vertebral crest continuous |
|
8 | Scattered, conspicuous large scales on dorsum, flanks, and hind limbs present |
|
– | Scattered, conspicuous large scales on dorsum, flanks, and hind limbs absent |
|
9 | Scattered large scales tetrahedral in shape; vertebrals on neck maximum twice as high as those between hind limbs |
|
– | Scattered large scales strongly keeled, not tetrahedral in shape; vertebrals on neck at least three times higher than those between hind limbs |
|
10 | Ventrals smooth or slightly keeled |
|
– | Ventrals conspicuously keeled |
|
11 | Gulars in males cream or yellow without black margins; usually one femoral pore on each leg |
|
– | Gulars in males bright orange or red, with black margins; usually two femoral pores on each leg |
|
12 | Dorsals heterogeneous in size, with scattered, tetrahedral, projecting scales (sometimes absent in males or juveniles); dorsolateral crests well developed between hind limbs |
|
– | Dorsals homogeneous in size, without projecting scales; dorsolateral crests inconspicuous or absent between hind limbs |
|
13 | Dorsals smooth or slightly keeled |
|
– | Dorsals conspicuously keeled |
|
14 | Scales on flanks heterogeneous in size, with a few enlarged, circular, keeled scales; iris bright red in both sexes; black patch under gular fold extending dorsally to form a short antehumeral bar in males |
|
– | Scales on flanks almost homogenous in size; iris brown in both sexes; black medial patch on gular region not extending dorsally to form an antehumeral bar in males |
|
15 | Dorsals in transverse row between dorsolateral crests at midbody 31 or fewer |
|
– | Dorsals in transverse row between dorsolateral crests at midbody more than 31 |
|
16 | Scales along the lateral edge of the skull roof strongly projected; dorsal scales homogeneous in size, with prominent median keel; antehumeral orange blotch in adult males absent |
|
– | Scales along the lateral edge of the skull roof slightly projected; dorsal scales heterogeneous in size, without prominent median keel; distinct antehumeral orange blotch in adult males |
|
17 | White or cream spot posterior to tympanum usually present; 41–54 (mean = 45.96 ± 3.49) dorsals in transverse row between dorsolateral crests at midbody; gular background in adult males light blue |
|
– | White or cream spot posterior to tympanum absent; 37–47 (means = 41.63 ± 3.20 in |
|
18 | Vertebral scales in neck region in adult males similar in size to vertebrals in pelvic region; 45–57 (mean = 51.13 ± 4.05) gulars |
|
– | Vertebral scales in neck region in adult males more than twice as high as vertebrals in pelvic region; 42–48 (mean = 44.40 ± 2.22) gulars |
|
For the type specimens of the species described in this paper we thank all collectors for their help in the field. Venegas is indebted to R. Wagter for logistic support in the field. We thank A. Varela and P. Santiana for editing the photographs; L.A. Coloma for providing photographs of the type specimens of