Zootaxa 3753 (5): 453–468 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press
Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3753.5.4 http://zoobank.org/urn:lsid:zoobank.org:pub:BBAABC94-A07E-4914-9D07-86EE07BEF11C
A new species of pine anole from the Sierra Madre del Sur in Oaxaca, Mexico (Reptilia, Squamata, Dactyloidae: Anolis) GUNTHER KÖHLER1,5, RAÚL GÓMEZ TREJO PÉREZ2, CLAUS BO P. PETERSEN1,3 & FAUSTO R. MENDEZ DE LA CRUZ4 1
Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, 60325 Frankfurt a.M., Germany Facultad de Estudios Superiores Iztacala, Universidad Nacional Autónoma de México (UNAM), Avenida de los Barrios 1, Los Reyes Iztacala, C.P. 54090, Estado de México, México 3 Zoological Museum, Natural History Museum of Denmark, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen, Denmark 4 Instituto de Biología, Universidad Nacional Autónoma de México (UNAM), A.P. 70-153, C.P. 04510, México D.F. México 5 Correspondence. E-mail:
[email protected] 2
Abstract We describe the new species Anolis peucephilus sp. nov. from the Pacific versant of southern Mexico. Anolis peucephilus differs from all congeners by having a combination of (1) smooth ventral scales; (2) usually a patch of three greatly enlarged supraocular scales; (3) extremely short hind legs, longest toe of adpressed hind leg reaching to a point between levels of axilla and ear opening, ratio shank length/snout-vent length 0.18–0.21; (4) circumnasal usually in contact with first supralabial; and (5) a large yellowish orange dewlap in males and a very small to small white dewlap in females. In external morphology, A. peucephilus is most similar to A. omiltemanus from which it differs by having even shorter hind legs with the longest toe of adpressed hind leg reaching to a point between levels of axilla and ear opening (versus usually to ear opening, occasionally to slightly beyond ear opening or to a point between shoulder and ear opening in A. omiltemanus), a slightly larger dewlap in females, to 64 mm2 (versus to 41 mm2 in A. omiltemanus), the circumnasal usually in contact with the first supralabial (versus those scales separated by the presence of a subnasal in A. omiltemanus), and 4–6 internasal scales in the new species (versus usually 6–7 in A. omiltemanus). Furthermore, A. peucephilus differs from A. omiltemanus in hemipenial morphology (no finger-like processus on asulcate side in A. peucephilus versus such a processus present in A. omiltemanus). Also, in a preliminary molecular genetic analysis of the mitochondrial CO1 gene fragment, A. peucephilus has a genetic distance of 11.5% from A. omiltemanus. Anolis peucephilus was collected at night while the lizards were sleeping in pine trees, 2–10 m above the ground. Key words: Anolis peucephilus sp. nov.; Dactyloidae; Mexico; new species; Oaxaca; Reptilia; Squamata
Resumen Describimos la nueva especie Anolis peucephilus de la vertiente Pacífico del sureste de México. Anolis peucephilus difiere de todos sus congéneres al tener una combinación de (1) escamas ventrales lisas; (2) usualmente un parche de tres escamas supraoculares fuertemente agrandadas; (3) patas traseras extremadamente cortas, el dedo más largo de la pata trasera presionado paralelo al cuerpo alcanza un punto entre el nivel de la axila y la apertura timpánica, proporción de la longitud de la pierna/longitud hocico-cloaca 0.18–0.21; (4) circumnasal usualmente en contacto con la primera supralabial; y (5) presencia de un gran abanico gular amarillo naranja en machos y un pequeño abanico gular blanco en hembras. En morfología externa, A. peucephilus es más similar a A. omiltemanus con el cual la diferencia es la presencia de patas traseras aún más cortas, con el dedo más largo de la pata trasera alcanzando un punto entre el nivel de axila y la apertura timpánica (versus usualmente alcanzando la apertura timpánica y ocasionalmente ligeramente más allá de ella o entre ella y el hombro en A. omiltemanus), un abanico gular mas grande en hembras, hasta 64 mm2 (versus hasta 41 mm2 in A. omiltemanus), la circumnasal usualmente en contacto con la primer supralabial (versus aquellas escamas separadas por la presencia de una subnasal en A. omiltemanus ), y 4–6 escamas internasales (versus 6–7 en A. omiltemanus). Anolis peucephilus también se diferencia de A.omiltemanus en la morfología de los hemipenes (sin proceso en forma de dedo en lado asulcado en A. peucephilus versus tal proceso presente en A. omiltemanus). Además, en un análisis preliminar de genética molecular de un fragmento del gen mitocondrial CO1, A. peucephilus presenta una distancia genética de 11.5% de A. omiltemanus. Los individuos de A. peucephilus fueron colectados de noche mientras dormían en pinos, entre 2–10 metros sobre el suelo.
Accepted by S. Carranza: 11 Nov. 2013; published: 9 Jan. 2014
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Introduction The Pacific versant of southern Mexico (states of Oaxaca and Guerrero) is home to a distinct assemblage of beta anoles (fide Etheridge 1959). The majority of species found in this region shares the presence of an oval patch of usually three greatly enlarged supraocular scales (all Mexican Pacific species except for A. macrinii Smith 1968 and A. unilobatus Köhler & Vesely 2010). Recently, several nominal species of Mexican anoles have been placed into the synonymy of other species (Köhler 2011, 2012a; Nieto Montes de Oca et al. 2013). The currently recognized species known to occur along the Pacific versant of southern Mexico can be divided roughly into species with moderately to strongly keeled midventral scales (i.e., A. forbesi Smith & Van Gelder 1955, A. isthmicus Fitch 1978, A. megapholidotus Smith 1933, A. microlepidotus Davis 1954, A. nebuloides Bocourt 1873, A. nebulosus (Wiegmann 1834), A. quercorum Fitch 1978, A. subocularis Davis 1954, A. unilobatus) and those with usually perfectly smooth (faintly to weakly keeled in some individuals of A. macrinii) midventral scales (i.e., A. dunni Smith 1936, A. gadovii Boulenger 1905, A. liogaster Boulenger 1905, A. macrinii, A. omiltemanus Davis 1954, A. taylori Smith & Spieler 1945). Köhler et al. (2013) proposed that A. omiltemanus in Guerrero is an arboreal pine forest specialist that prefers open pine forests with little undergrowth. Recent field work in Oaxaca produced a series of anoles that also seem to be ecologically restricted to pine forests in the Sierra Madre del Sur. A comparison of external morphology and molecular data of this series with that for all known Mexican and Central American species of Anolis confirmed our initial assumption that these anoles represent an undescribed species. Therefore, we describe it as a new species below.
Materials and methods In evaluating species boundaries within the populations of anoles found in western Mexico, we follow the unified species concept (de Queiroz 2007). As lines of evidence for species delimitation, we apply a phenotypic criterion (external morphology: coloration, morphometrics, and pholidosis) and a criterion for reproductive isolation (genetic distinctness of the mitochondrial CO1 gene). Recently, Nicholson et al. (2012) proposed a new classification for the anoles, seriously questioned by several researchers (Castañeda & de Queiroz 2013; Poe 2013). We choose to be conservative until this debate has been settled and use the name Anolis in this article. Prior to preservation of collected specimens in the field, we took color photographs of each individual’s extended dewlap. For this purpose, we preferably utilized the standard forceps of genuine Swiss Army Knives since their broad, flat apex prevents even thin-skinned dewlaps from damage and functions as an approximate scale (width = 3 mm in the models of both suppliers). Immediately after euthanasia, relative hind limb length was determined by recording the point reached by the tip of the fourth toe when the extended hind limb was adpressed along the straightened specimen. Whenever possible, we everted the hemipenes of male specimens by injecting 70% ethanol into the hemipenial pockets after manually pre-everting the hemipenes. Specimens were then preserved by injecting a solution of 5–10 mL absolute (i.e., 36%) formalin in 1 L of 96% ethanol into the body cavity and thighs, preferably also sprinkling everted hemipenes and extended dewlaps with this solution, and stored in 70% ethanol. The collected specimens have been deposited in the collection of the Senckenberg Forschungsinstitut Frankfurt (SMF), and the remaining specimens, here indicated with GK field numbers, will be deposited in the Instituto de Biología (IBH), Universidad Nacional Autónoma de México (UNAM), México D.F., Mexico. Abbreviations for museum collections follow Sabaj Pérez (2010). Coordinates and elevation were recorded using Garmin GPS receivers with built-in altimeters. All coordinates are in decimal degrees, WGS 1984 datum, and rounded to the fifth decimal place. The capitalized colors and color codes (the latter in parentheses) are those of Köhler (2012b). Terminology of markings used in color descriptions follow Köhler (2012b). Head length was measured from the tip of the snout to the anterior margin of the ear opening, with the calipers held in a vertical position relative to the head. Snout length was measured from the tip of the snout to the anterior border of the orbit, with the calipers held in a horizontal position relative to the head. Head width was determined with the broad tips of the calipers aligned with the levels of posterior margin of eye and supralabial scales, respectively, with the calipers held in a vertical position relative to the head. Dorsal and ventral scales were counted at midbody along the midline. Tail height and width were measured at the point reached by the heel of the extended hind leg. Subdigital lamellae were counted on Phalanges II to IV of Toe IV of the hind limbs, and separately on distal phalanx.We considered the scale directly
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anterior to the circumnasal to be a prenasal. Abbreviations used are AGD (axilla-groin distance), dorsAG (number of medial dorsal scales between levels of axilla and groin), dorsHL (number of medial dorsal scales in one head length), HDT (horizontal diameter of tail), HL (head length), HW (head width), IFL (infralabial scales), IP (interparietal plate), SAM (scales around midbody), ShL (shank length), SL (snout length), SO (subocular scales), SPL (supralabial scales), SS (supraorbital semicircles), SVL (snout–vent length), TL (tail length), VDT (vertical diameter of tail), ventrAG (number of medial ventral scales between levels of axilla and groin), ventrHL (number of medial ventral scales in one head length). In reporting the frequencies of character states, we used the following terminology: if a character state was present in more than 65% of the examined specimens, we coded it as “usually”; 20% “commonly”; 5% “occasionally”; and 80 m SVL (versus
