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CORBIDI 08828 (Fig. 7), an adult male from Chambirillo close to the Checkpoint 16 of the CAZNP (07°04'8.9"S, 76°00'51.2"W, 1122 m), Provincia de Picota, Región San Martín, Perú, collected on 30 October 2010 by P. J. Venegas.
CORBIDI 08827, an adult female collected on 2 November 2010 by P. J. Venegas; CORBIDI 08786, 08787, 08788, 08789, adult females collected on 21 January 2011 by P. J. Venegas and V. Duran; CORBIDI 09215, 09216, a juvenile male and adult female, respectively, collected on 6 May 2011 by P. J. Venegas and V. Duran. All paratypes are from the same locality as the holotype.
Enyalioides binzayedi can be distinguished from other species of Enyalioides by the combination of the following characters: (1) scales posterior to the superciliaries forming a longitudinal row of strongly projecting scales across the lateral edge of the skull roof in adults of both sexes; (2) 31 or fewer longitudinal rows of strongly keeled dorsals in a transverse line between the dorsolateral crests at midbody; (3) ventral scales strongly keeled; (4) caudals increase in size posteriorly within each autotomic segment; (5) projecting scales on body or limbs absent; (6) vertebrals on neck more than five times the size of vertebrals between hind limbs in adult males.
A longitudinal row of strongly projecting scales along the lateral edge of the skull posterior and continuous with the superciliaries is also present in Enyalioides oshaughnessyi, which occurs west of the Andes in Ecuador and Colombia and differs from Enyalioides binzayedi in having smooth or slightly keeled dorsals. Species of Enyalioides occurring east of the Andes that share strongly keeled ventrals with Enyalioides binzayedi are Enyalioides azulae, Enyalioides cofanorum, Enyalioides microlepis, Enyalioides palpebralis,and Enyalioides rudolfarndti. All of these species either lack strongly projecting scales along the lateral edge of the skull roof (although they are slightly projecting in Enyalioides rudolfarndti) or have such scales but with a gap separating them from the superciliaries (Enyalioides palpebralis). Enyalioides azulae, Enyalioides cofanorum and Enyalioides microlepis differ further from Enyalioides binzayedi (character states in parentheses) in having more than 33 dorsal scales in a transverse line between the dorsolateral crests at midbody (31 or fewer), a low vertebral crest (high, with vertebrals on neck more than four times the size of vertebrals between hind limbs in both sexes), and a black gular patch (absent). The new species can be also distinguished from Enyalioides palpebralis by lacking both a superciliary triangular flap that projects posterolaterally over each eye and a small gap in the vertebral crest in the neck region, and by having femoral pores. From Enyalioides rudolfarndti (character states in parentheses), Enyalioides binzayedi also differs in having a prominent medial keel on each dorsal scale (medial keel weak or absent), dorsals nearly homogeneous in size (dorsals heterogeneous in size), and in lacking a round orange blotch in the antehumeral region (orange blotch present in adult males).
Description of holotype
Male (Fig. 7); SVL = 118 mm; TL = 180 mm; maximum head width = 25.14 mm; head length = 30.46 mm; head height = 23.70 mm; dorsal head scales uni- or multicarinate, those in parietal region strongly projected dorsally; parietal eye present; scales immediately posterior to superciliares conical and dorsolaterally projected, forming longitudinal row of seven scales that extends posteriorly over supratemporal region, with fifth anteriormost scale more than twice the size of other scales in row; temporal scales small, multicarinate, juxtaposed; two large, projected conical temporal scales dorsal to tympanum, the dorsal one in contact with the supratemporal crest, and the ventral one in contact with an enlarged pretympanic scale; 14 superciliares; four canthals; three postrostrals; 12 (left or right) supralabials counted to a point right below middle of eye; rostral (2.27 × 1.27 mm) slightly wider than adjacent supralabials; single longitudinal row of lorilabials between suboculars and supralabials at level of middle of eye, two longitudinal rows of lorilabials immediately anterior to this point; loreal region broken into small, multicarinate, and juxtaposed scales; nasal at level of supralabials III–IV; 11 (left) or 10 (right) infralabials counted to a point right below middle of eye, respectively; mental (2.47 × 1.89 mm) twice as wide and high as adjacent infralabials; postmentals three; gulars ventrally projected, those immediately anterior to gular fold keeled, mucronate, and imbricate; gular fold complete midventrally, extending dorsally and posteriorly to form antehumeral fold; neck with several longitudinal and oblique folds, and a dorsolateral row of enlarged scales.
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 between fore and hind limbs with dorsolateral crests and without folds; dorsal scales heterogeneous in size, prominently keeled, and subimbricate; scales on flanks more homogeneous in size than dorsals and less than half their size; ventral scales imbricate, keeled, subrectangular, and mucronate; ventrals as large as largest dorsals.
Limb scales keeled and imbricate dorsally and ventrally; most scales on dorsal and posterior aspects of thighs homogeneous in size, less than half the size of scales on anterior and ventral aspects; 19 subdigitals on manual digit IV; 24 subdigitals on pedal digit IV; femoral pores on each side two; tail laterally compressed and gradually decreasing in relative height towards tip; caudal scales strongly keeled and imbricate, slightly increasing in size posteriorly on lateral and dorsal aspects of each vertebral segment; ventral caudals larger than dorsal caudals, with individual autotomic segments three scales long ventrally and four scales long dorsally.
Color in life of holotype (Fig. 7). Dorsal and lateral surface of head dark brown or black, with scattered light green scales (especially on the dorsal surface) and a dark longitudinal supratemporal stripe; supralabials greenish white intercalated with dark brown, infralabials greenish white; rostral and mental light green; gulars white, with greenish-white margins; skin between gulars dark gray; dorsal background of body, limbs, and tail light green, with a dark brown reticulation; a white blotch posterior to tympanum followed by five diffuse pale brown dorsolateral blotches extending from the neck to the base of the tail; ventral surface of body, limbs, and tail white, with a longitudinal row of 4–5 dark gray squarish marks between flanks and venter; iris coppery with a fine brown reticulation; pupil round with pale green margin.
Meristic and morphometric characters of Enyalioides binzayedi are summarized in Table 1. The holotype is the only adult male specimen available; it differs from female and subadult male paratypes in having projecting scales on each side of the vertebral crest on the neck. Additionally, female paratypes CORBIDI 08789 and 09216 are unique in having a double vertebral crest from midbody to pelvic region.
A subadult male specimen (CORBIDI 09215; Fig. 8) differs from the holotype in having scattered black spots on the ventral surface of body. All females differ from the holotype in having dorsal, broad transverse bars arranged longitudinally along the vertebral line, larger dark marks on the ventrolateral surface of body, and well defined postocular and supratemporal stripes. Dorsal background of body, limbs, and tail can be dark greenish brown (CORBIDI 08827 and 08787), as in the holotype, dark green (CORBIDI 08789), or dark brown (CORBIDI 08788) speckled with light green flecks. Females CORBIDI 08787 and 08827 have light dorsolateral blotches intercalating with dark transverse bars, which are well defined dorsolaterally and diffuse laterally (Fig. 9). Female paratypes CORBIDI 08789 and 09216 have a pale blotch behind the tympanum similar to the holotype, whereas CORBIDI 08787, 08827, and 08788 have a larger pale blotch connected to first pale dorsolateral blotch forming a continuous postympanic stripe extending from the tympanum to the scapular region. Ventrally females are white (CORBIDI 08788; Fig. 9D) or tan (CORBIDI 08787, 08789; Fig. 9B) with scattered dark brown spots or flecks. The throat in females is brown or light brown with dark flecks or diffuse reticulations, except one female (CORBIDI 08789), which has an immaculate tan throat.
Distribution and natural history
Enyalioides binzayedi is known only from its type locality in the montane rainforest of the Río Huallaga basin (Fig. 6) in northeastern Peru at an elevation of 1080 m. This locality lies within the CAZNP, in a mountain ridge between the Región San Martín and Región Loreto (Fig. 5). All individuals reported here were collected at night sleeping on vertical stems of bushes 30–230 cm above the ground. One female (CORBIDI 08788) collected on 21 January 2011 had two maturing eggs in each oviduct. Enyalioides binzayedi occurs in sympatry and possibly syntopy with Enyalioides azulae sp. n. (see above) and Enyalioides laticeps. Other species of squamate reptiles collected in the same locality include Alopoglossus angulatus, Anolis fuscoauratus, Anolis transversalis, Cercosaura manicata, Potamites ecpleopus, Potamites strangulatus, Potamites sp., Chironius fuscus, Dipsas indica, Imantodes cenchoa, Imantodes lentiferus, Micrurus obscurus, Oxyrhopus petola, and Xenopholis scalaris.
The specific name is a noun in the genitive case and is a patronym honoring Sheikh Mohamed bin Zayed Al Nahyan, Crown Prince of Abu Dhabi and Deputy Supreme Commander of the UAE, who created the Mohamed bin Zayed Species Conservation Fund (MBZSCF) to support species conservation projects around the globe. Field surveys leading to the discovery of the two species reported on in this paper were supported by a grant from the MBZSCF. Phylogenetic relationships Using a phylogenetic definition (de Queiroz and Gauthier 1990, 1992), Torres-Carvajal and de Queiroz (2009) applied the name Enyalioides to the crown clade originating in the most recent common ancestor of Enyalioides cofanorum Duellman 1973, Enyalioides heterolepis (Bocourt 1874), Enyalioides laticeps (Guichenot 1855), Enyalioides microlepis (O'Shaughnessy 1881), Enyalioides oshaughnessyi (Boulenger 1881), Enyalioides palpebralis (Boulenger 1883), Enyalioides praestabilis (O'Shaughnessy 1881), and Enyalioides touzeti Torres-Carvajal et al. 2008. The phylogenetic tree inferred in this study (Fig. 10) is consistent with Torres-Carvajal and de Queiroz's (2009) phylogenetic hypothesis in that species of Enyalioides are split into two primary subclades. One contains Enyalioides heterolepis and Enyalioides laticeps as sister taxa, and the other includes all remaining species of Enyalioides, as well as possibly Morunasaurus. Enyalioides azulae sp. n. is sister to the clade (Enyalioides palpebralis, (Enyalioides binzayedi sp. n., Enyalioides rudolfarndti)) with strong support (PP = 1.00), whereas Enyalioides binzayedi sp. n. is sister to Enyalioides rudolfarndti with strong support (PP = 1.00). Both species reported on in this paper, as well as Enyalioides rudolfarndti, are strongly supported (PP = 1.00) as monophyletic groups(Fig. 10). Thus, the phylogenetic tree presented here strongly supports both referral of the new species to Enyalioides and their status as different species from those recognized previously, except that the divergence between Enyalioides binzayedi and Enyalioides rudolfarndti is less than that observed within some currently recognized species (Enyalioides heterolepis and Enyalioides laticeps), which is at least partly attributable to the geographic separation of the samples. Differences in morphology and color patterns presented above provide additional evidence for recognizing Enyalioides binzayedi sp. n. and Enyalioides rudolfarndti as separate species.
|Character||Enyalioides azulae; n = 8||! Enyalioides binzayedi; n = 7|
|Vertebrals from occiput to base of tail||62–69 65.88 ± 2.70||40–55 48.00 ± 5.51|
|Dorsals in transverse row between dorsolateral crests at midbody||37–47 41.63 ± 3.20||22–31 27.57 ± 3.64|
|Ventrals in transverse row at midbody||27–33 28.75 ± 1.91||26–32 28.14 ± 2.12|
|Transverse rows of ventrals between fore and hind limb||36–44 40.38 ± 2.45||30–39 35.29 ± 2.81|
|Gulars||45–57 51.13 ± 4.05||27–31 29.14 ± 1.77|
|Infralabials||10–13 11.38 ± 1.30||10–14 11.29 ± 1.50|
|Supralabials||10–14 11.75 ± 1.28||11–15 12.00 ± 1.41|
|Canthals||4–6 4.63 ± 0.74||4–6 4.43 ± 0.79|
|Superciliaries||12–18 15.38 ± 2.07||13–14 13.57 ± 0.53|
|Subdigitals Manual Digit IV||15–22 19.25 ± 1.98||17–22 19.86 ± 1.68|
|Subdigitals Pedal Digit IV||25–28 26.50 ± 1.07||24–30 27.14 ± 2.48|
|Femoral pores in males||1 (n = 4) —||1–2 (n = 2) —|
|Femoral pores in females||1–2 (n = 4) 1.13 ± 0.35||1–3 (n = 5) 2.20 ± 0.79|
|Head length/head width||1.23–1.32 (n = 4) 1.26 ± 0.04||1.21–1.41 1.26 ± 0.07|
|Head width/head height||1.15–1.27 (n = 4) 1.20 ± 0.05||1.04–1.16 1.10 ± 0.05|
|Rostral width/rostral height||1.55–2.22 (n = 4) 1.79 ± 0.30||1.51–2.56 1.79 ± 0.36|
|Mental width/mental height||1.18–1.64 (n = 4) 1.41 ± 0.21||1.20–1.63 1.40 ± 0.17|
|Fore limb length/SVL||0.49–0.53 (n = 4) 0.51 ± 0.02||0.47–0.53 0.52 ± 0.02|
|Hind limb length/SVL||0.75–0.84 (n = 4) 0.80 ± 0.04||0.69–0.80 0.75 ± 0.04|
|Tail length/total length||0.57–0.59 (n = 5) 0.58 ± 0.01||0.56–0.60 0.58 ± 0.02|
|Maximum SVL (mm) males||96 (n = 4)||118 (n = 2)|
|Maximum SVL (mm) females||96 (n = 4)||122 (n = 5)|
Key to the species of Hoplocercinae The following key is artificial in the sense that its structure does not necessarily reflect the order of branching in the phylogeny.
- Venegas, P; Torres-Carvajal, O; Duran, V; Queiroz, K; 2013: Two sympatric new species of woodlizards (Hoplocercinae, Enyalioides ) from Cordillera Azul National Park in northeastern Peru ZooKeys, 277: 69-90. doi: 10.3897/zookeys.277.3594
- Torres-Carvajal O, de Queiroz K (2009) Phylogeny of hoplocercine lizards (Squamata: Iguania) with estimates of relative divergence times. Molecular Phylogenetics and Evolution. Molecular Phylogenetics and Evolution 50: 31-43. doi: 10.1016/j.ympev.2008.10.002
- Torres-Carvajal O, Almendáriz A, Valencia J, Yánez-Muñoz M, Reyes J (2008) A new species of Enyalioides (Iguanidae: Hoplocercinae) from southwestern Ecuador. Papéis Avulsos de Zoologia 48: 227-235. doi: 10.1590/S0031-10492008002000001