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Xenosaurus tzacualtipantecus WOOLRICH-PIÑA & SMITH, 2012

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Higher TaxaXenosauridae, Diploglossa, Anguimorpha, Sauria, Squamata (lizards) 
Subspecies 
Common NamesZacualtipá́n Knob-scaled Lizard 
SynonymXenosaurus tzacualtipantecus WOOLRICH-PIÑA & SMITH 2012 
DistributionMexico (Hidalgo: Sierra Madre Oriental, Puebla)

Type locality: La Mojonera 7.5 km S– SW Zacualtipa ́n (20°38’29’’N, 98°36’6’’W; datum 1⁄4 WGS84), 1900 m elevation.  
Reproduction 
TypesHolotype: UBIPRO (given as LEUBIPRO) 11444, a female adult (Fig. 1), collected on 26 April 2002 by Julio A. Lemos-Espinal. This location is characterized as rain forest (Fig. 2).
Paratypes: UBIPRO (= LEUBIPRO) (17 specimens) Nine adult females: LEUBIPRO 11431, 11432, 11436, 11442, 11446, 11447, 11451, 14885, and 14886; five adult males: LEUBIPRO 14887– 14891; one juvenile: LEUBIPRO 14892; and two neonates: LEUBIPRO 14893–14894, all collected from the type locality by Julio A. Lemos-Espinal and Guillermo A. Woolrich- Piña. 
DiagnosisDiagnosis.—Xenosaurus tzacualtipantecus differs from the other species of Xenosaurus by the absence of a neck band and the presence of two pale lines that angle up from the jaws and extend parasagittally along the neck. Xenosaurus tzacualtipantecus is similar to X. grandis sanmartinensis, X. phalaroan- thereon, X. rackhami, and X. rectocollaris in having the zygomatic–postocular arches in contact, and differs from X. newmanorum, X. agrenon, X. grandis arboreus, X. grandis grandis, X. platyceps, and X. penai, in which the zygomatic–postocular arches are separat- ed. The number of longitudinal ventral scale rows in X. tzacualtipantecus overlaps exten- sively with those of X. newmanorum, X. agrenon, X. g. grandis, X. grandis sammarti- nensis, X. penai, and X. rackhami; barely overlaps with those of X. grandis arboreus and X. phalaroanthereon; and does not overlap with the higher number in X. platyceps or with the lower number in X. rectocollaris (Table 1). The number of tympanic scales in X. tzacualtipantecus is within the range seen in X. newmanorum, X. phalaroantereon, X. platyceps, and X. recto- collaris, but is lower than in X. agrenon and X. grandis grandis; no data on tympanic scales are available for X. g. arboreus, X. g. sanmartinensis, X. penai, and X. rackhami. The number of subdigital lamellae on the fourth finger in X. tzacualtipantecus is higher than in X. rectocollaris and X. phalaroanther- eon; it overlaps with the ranges in X. agrenon, X. grandis arboreus, X. g. grandis, X. platyceps, X. penai, and X. rackhami; and is lower than in X. grandis sanmartinensis and X. newmanorum. The spotted venter of X. tzacualtipantecus is shared with X. agre- non, X. g. grandis, X. grandis sanmartinensis, X. penai, and X. rackhami; it contrasts with the immaculate venters of X. newmanorum, X. grandis arboreus, X. phalaroanthereon, X. platyceps, and X. rectocollaris (Table 1).
 
CommentThis new species differs from previously described species in lacking a neck band and in having two pale lines that angle up from the jaws and extend parasagittally along the neck.

Distribution: see map in WOOLRICH-PIÑA & SMITH 2012.

The factor analysis resulted in a single factor that was positively related to the number of lamellae on the fourth finger (n = 149, r2 = 0.71, P < 0.001), the number of longitudinal ventral scale rows (n = 149, r2 = 0.78, P < 0.001), the state of the zygomatic–postocular arches (n = 149, r2 = 0.94, P < 0.001), and the number of tympanic scales (n = 149, r2 = 0.43, P < 0.001). This factor had an eigenvalue of 2.38 and explained 59.6% of the variance. The coefficients for the standardized factor scores were 0.354 for the number of lamellae on the fourth finger, 0.326 for the number of longitudinal ventral scale rows, 0.395 for the state of the zygomatic–postocular arches, and 0.180 for the number of tympanic scales. There were significant differences in the factor scores among the seven species (Table 2; F6,142 = 572.9, P < 0.001). Five of the six previously described species that we examined directly had significantly different mean factor scores from X. tzacualtipantecus; only X. phalaroanthereon did not have a significantly different mean factor score (Table 2), which supported the general uniqueness of this species independent of the two new unique traits identified above (note that X. phalaroanthereon is geographically distant from X. tzacualtipantecus). 
EtymologyEtymology.—The name tzacualtipantecus is derived from two Nahuatl words: Zacualtipan or ‘‘tzacualtipan’’ in the original Nahuatl language, referring to the closest town to the type locality and meaning ‘‘hiding place,’’ which is particularly appropriate given the crevice-dwelling habits of knob-scaled lizards; and ‘‘teca,’’ which means belonging to a place (Simeón, 1885). 
References
  • Flores-Hernández, Miguel Ángel, Leonardo Fernández-Badillo and Cristian R. Olvera-Olvera. 2016. Xenosaurus tzacualtipantecus Woolrich-Piña and Smith, 2012. Mexico, Hidalgo. Mesoamerican Herpetology 3(4): 1047–1049 - get paper here
  • Lemos-Espinal, Julio A., Geoffrey R. Smith 2015. Amphibians and reptiles of the state of Hidalgo, Mexico. Check List 11 (3): 1642 - get paper here
  • Nieto-Montes de Oca A, Sánchez-Vega H, Durán-Fuentes I 2018. A new species of knob-scaled lizard (Xenosauridae, Xenosaurus) from the Sierra Madre Oriental of Puebla, Mexico. ZooKeys 737: 141-160 - get paper here
  • Nieto-Montes de Oca, Adrián; Anthony J. Barley, Rubi N. Meza-Lázaro, Uri O. García-Vázquez, Joan G. Zamora-Abrego, Robert C. Thomson, Adam D. Leaché 2016. Phylogenomics and species delimitation in the knob-scaled lizards of the genus Xenosaurus (Squamata: Xenosauridae) using ddRADseq data reveal a substantial underestimation of diversity. Molecular Phylogenetics and Evolution 106: 241-253 - get paper here
  • Nieto-Montes de Oca, Adrián; Uri O. García-Vázquez, J. Jaime Zúñiga-Vega and Walter Schmidt-Ballardo 2013. A new species of Xenosaurus (Squamata: Xenosauridae) from the Sierra Gorda Biosphere Reserve of Querétaro, Mexico. Revista Mexicana de Biodiversidad 84: 485-498 - get paper here
  • Wilson, Larry David; Vicente Mata-Silva, Jerry D. Johnson 2013. A conservation reassessment of the reptiles of Mexico based on the EVS measure. Amphibian & Reptile Conservation 7 (1): 1–47 - get paper here
  • Woolrich-Pima, Guillermo A., Alvarado-Hernández, Adán, Mora-Guzmán, Esequiel, M., Arellano-Cárcamo. Yennifer, Olvera-Arrieta, Johhatan, González-Gonzalez, G. Yazmín and Sosa-Villanueva, Fátima 2016. Geographic Distribution: Xenosaurus tzacualtipantecus (Zacualtipan Knob-scaled Lizard) Herpetological Review 47 (4): 629
  • Woolrich-Piña, G. A., E. García-Padilla, D. L. DeSantis, J. D. Johnson, V. Mata-Silva, and L. D. Wilson. 2017. The herpetofauna of Puebla, Mexico: composition, distribution, and conservation status. Mesoamerican Herpetology 4(4): 791–884
  • Woolrich-Piña, Guillermo A. and Geoffrey R. Smith 2012. A New Species of Xenosaurus from the Sierra Madre Oriental, Mexico. Herpetologica 68 (4): 551-559. - get paper here
 
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