Liolaemus calliston, Avila & Perez & Minoli & Medina & Sites & Morando, 2017

Avila, Luciano Javier, Perez, Cristian Hernán Fulvio, Minoli, Ignacio, Medina, Cintia Debora, Sites, Jack W. & Morando, Mariana, 2017, New species of Liolaemus (Reptilia, Squamata, Liolaemini) of the Liolaemus donosobarrosi clade from northwestern Patagonia, Neuquén province, Argentina, Zootaxa 4362 (4), pp. 535-563 : 551-555

Liolaemus calliston sp. nov.

Holotype. LJAMM-CNP 10994 ( Figs. 10 , 11 ), an adult male from 4.4 km SE Aguada Pichana Main Camp Total Oil Company, ~ 30 km S junction Provincial Road 7, Añelo Department, Neuquén province, Argentina (38º25’43.7”S, 69º09’31.3”W, 431 m), collected by C.H.F. Perez, J. Nittmann, F. Farinaccio and D.R. Perez, 28 October 2008.

Paratypes. LJAMM-CNP 10995–10996, MLP.S 2624 (females) from same locality as holotype, collected by C.H.F. Perez, J. Nittmann, F. Farinaccio and D.R. Perez, 28 October 2008 . LJAMM-CNP 12420–12422 (females) from same locality as holotype, collected by L.J. Avila and C.H.F. Perez, 4 December 2009 . LJAMM-CNP 12417– 12419 (males) from same locality as holotype, collected by L.J. Avila and C.H.F. Perez, 4 December 2009 .

Diagnosis. Liolaemus calliston sp. nov. is a robust but small sized member of the clade of Liolaemus referred to as the boulengeri group by Etheridge (1995) and is a member of the Liolaemus donosobarrosi group ( Avila et al. 2006, 2007, Olave et. al 2014, 2016), that includes L. cuyanus , L. donosobarrosi , L. mapuche , L. josei , L. puelche and several other potential species still undescribed. Liolaemus tirantii sp. nov. can be distinguished from all other species of its group of Liolaemus by a combination of genetic and morphological characteristics.

Liolaemus calliston sp. nov. can be differentiated from L. cuyanus for having smaller size (61.0 mm vs 92.5 mm maximum SVL), have higher number of midbody scales (X = 76.8 vs X = 66.7; min.–max. 70–84 vs 60–76, Blum 2012) with overlap, higher number of dorsal scales (X = 95.3 vs 63.6; min.–max. 86–100 vs 57–74, Blum 2012) with no overlap, only four scales in contact with mental vs six in L. cuyanus , lack of prescapular marks and ventral melanism, and a very different general coloration and coloration patterns.

Liolaemus calliston sp. nov. is smaller than Liolaemus mapuche (61.0 mm vs 82.7 mm maximum SVL), have higher number of dorsal scales (X = 95.3 vs 76.2; min.–max. 86–100 vs 70–86) with almost no overlap, only four scales in contact with mental vs six in L. cuyanus (more common state, Abdala 2002), and lacks of the scapular marks, ventral melanism and lacks of the very different dorsal coloration characterized by a light blue head and scattered blue scales on a light green-blue background, ( Abdala, 2002).

Liolaemus calliston sp. nov. is smaller than L. puelche (61.0 mm vs 89 mm maximum SVL), have higher number of midbody scales (X = 76.8 vs 70.7; min.–max. 70–84 vs 67–76) with slight overlap and higher number of dorsal scales (X = 95.3 vs 76.7; min.–max. 86–100 vs 74–80) with no overlap.

Liolaemus calliston sp. nov. is smaller than L. josei (61.0 mm vs 73.1 mm maximum SVL), have higher number of midbody scales (X = 76.8 vs 69.1; min.–max. 70–84 vs 62–76), have higher number of dorsal scales (X = 95.3 vs X = 73.1; min.–max. 86–100 vs 67–81) with no overlap, lacks of the well-marked gular melanism of L. josei males as well as its more marked dorsal color pattern.

Liolaemus calliston sp. nov. can be only be differentiate from L. donosobarrosi in its particular dorsal coloration of colorful transversal bands in live animals, lack of transversal dark bands on the sides of the head, an oval-shaped tympanic opening vs a more quadrangular in L. donosobarrosi , and supralabials and lorilabials scales quadrangular shaped in L. calliston sp. nov. than in L. donosobarrosi . Dorsal coloration pattern in L. tirantii lacks of the banded pattern of L. calliston and the colorful general coloration; series of transversal half-moon dots almost disappear in live animal in L. calliston sp. nov. and are fused each other laterally along the body, given it a ¨wavy¨ appearance. L. calliston sp. nov. have a higher average number of dorsal scales than in L. tirantii sp. nov. (X = 95.3 vs 87.2). Infralabials, supralabial and lorilabials scales more quadrangular shaped in L. calliston sp. nov. than in L. tirantii sp. nov.

Description of holotype. Adult male ( Fig. 10 , 11 ) 60.4 mm snout-vent length (SVL); tail length 103.4 mm. Axilla-groin distance 27.2 mm. Head length 13.1 mm (from anterior border of tympanum to tip of snout), 11.5 mm wide (at anterior border of tympanum), 7.5 mm high (at anterior border of tympanum). Snout length 4.9 mm (orbittip of snout distance). Interorbital distance 6.0 mm. Eye-nostril distance 3.9 mm. Orbit-auditory meatus distance 4.7 mm. Forelimb length 17.4 mm. Tibial length 12.6 mm. Foot length 18.5 mm (ankle to insertion of claw on fourth toe). Dorsal head scales something bulged, smooth, 16 between occiput at level of anterior border of tympanum to rostral, pitted with numerous scale organs in the anterior region, and reducing to a single organ, or lack, in the posterior half of the head. Rostral scale wider (2.5 mm) than high (0.8 mm). Two postrostrals, together with anterior lorilabial, separate nasal scales from rostral. Nasal scales longer than wide, irregularly heptagonal; nostril one-half length of nasal, posterior in position. Scales surrounding nasals 7–7. Two internasals. Frontonasals four, irregular in size and position. A row with three scales between frontonasals and prefrontals. Prefrontals 5, a small pentagonal scales in the center (1.2 mm), and two pairs scales: a pair dorso-lateral larger scales, irregularly hexagonal (1.6 mm), and a pair lateral medium-sized scales (1.4 mm), aproximatly pentagonal. Two frontal scale. Frontoparietals in two rows, two anterior and posterior scales. Interparietal subpentagonal, surrounded by six scales; four smaller and irregular in front and sides, two larger in back. Parietal eye evident. Parietals some bulged, irregularly shaped, similar in size to interparietal (1.8 vs. 1.6 mm). Circumorbitals: 13–12. Transversally expanded supraoculars 6–6. Smaller lateral supraoculars: 14–19. Anterior canthal wider than long, separate from nasal by one postnasals. Posterior canthal longer than wide. Loreal scales, bulged (6–7). Lorilabials longer than wide (7–8), smaller than labials. Superciliaries 6–7, flattened and elongated, anterior four broadly overlapping dorsally. Orbit with 14–15 upper and thirteen lower ciliaries on each side. Orbit diameter 3.3 x 2.1 mm. Preocular small, unfragmented, longer than wide. Subocular scale elongated, nine times longer than wide (4.2 x 0.4 mm). A wellmarked longitudinal ridge along upper margin of preocular and subocular scales. Postocular small, slightly bulged, quarter superimposed to subocular, with a marked longitudinal ridge. Palpebral scales small granular and bulged. Supralabials six on each side, convex. Temporals smooth, convex, juxtaposed with one scale organ in the tip. Anterior auriculars smaller than adjacent posterior temporals, something projecting outward (3–2). Posterior auriculars small and granular. External auditory meatus conspicuous, higher (1.9 mm) than wide (1.1 mm). Lateral scales of neck granular with inflated skin. Mental scale wider (2.3 mm) than high (1.4 mm), in contact with five scales. Mental followed posteriorly by two postmentals, and two rows of four chinshields on right side (left side with scar damage). Five infralabials on each side, stretched, slightly smaller than supralabials. Gular scales smooth, flat, imbricate, with rounded posterior margins, with melanophores. Scales of throat between chinshields slightly juxtaposed, becoming slightly imbricate toward auditory meatus. Thirty-three gulars between tympanum openings. Infralabials separated from chinshields by one to three rows of scales.

Antehumeral and longitudinal neck well developed; postauricular very distinctive; gular incomplete; rictal, dorsolateral, oblique unconspicuous.

Scales of dorsal neck region rhombals, imbricate, smooth. Ninety-nine dorsal scales between occiput and anterior surface of thighs. Dorsal body scales rhomboidal, imbricate, smooth to very slightly keeled towards the posterior half of body. Dorsal scales grade laterally into slightly smaller, smooth scales at midbody. Scales immediately anterior and posterior to forelimb and hindlimb insertion small, smooth, granular, and nonoverlapping. Body lateral scales grading smaller to larger at midbody. Ventral body scales rhombals, smooth, flat, imbricate, larger than dorsal scales. Eighty-four scales around midbody; scales between mental and precloacal pores 94. Scales of cloacal region about equal in size to ventral body scales. Six conspicuous precloacal pores.

Anterior suprabrachials rhomboidal, imbricate, smooth, slightly larger in size to dorsal body scales. Postabrachials smaller, smooth, becoming granular near axilla. Supra-antebrachials similar to suprabrachial. Infraantebrachials rhombals, smooth. Supracarpals imbricated, rhomboidal, smooth. Infracarpals strongly imbricate, rhomboidal, keeled, 1–3 mucronate. Subdigital lamellae with 2–3 keels, each terminating in a short mucron, numbering: I: 9, II: 14, III: 19, IV: 20, V: 12. Claws robust, curved and sharp, opaque brown.

Suprafemorals as large as dorsal body scales, rhomboidal, imbricated, smooth, some very slightly keeled. Postfemorals small, granular shape. Supratibial rhomboidal, imbricated, smooth. Infrafemorals scale small, granular and smooth with enlarged and bulged scales patch, with thirty scales (femoral patch), mucronate. Supratarsal rhombals, imbricated and smooth. Infratarsal small, rhomboidal, imbricate, smooth. Subdigital scales keeled, each terminating in 1–3 short mucron, numbering: I: 11, II: 16, III: 20, IV: 25, V: 15. Claws robust, curved and sharp, opaque brown. Tail complete, non-regenerated. Dorsal and lateral caudals scales, quadrangular and moderatly keeled. Ventral subtriangular and smooth, toward posterior half moderatly keeled.

Color in life. Head dorsal coloration darker than body coloration, homogenously Prout´s Brown (47) with darker areas Burnt Umber (48); all this pattern, extend to the sides of the head. Dorsal background coloration between occiput to rump with five Dark Salmon Color (59) transverse bands separated by similar wide bands of Light Yellow Orcher (13). Each Dark Salmon Color (59) band posteriorly bordered by slightly evident half-moon Tawny (60) dots barely encircling Cream White (52) scales. Two more Dark Salmon Color (59) bands continuous along the first quarter of the tail, then bands became smaller and encircle the tail as incomplete and broken rings, and almost disappear in the last third to the tip. From the posterior area of the tympanum to the cloacal region of the tail, a lateral band of Dark Salmon Color (59), sometimes dotted of Cream White (52). From the lateral sides of the neck to the first third of the tail, iridescent Caribbean Blue (168) dots are homogeneously distributed, reaching some middorsal areas of the trunk and becoming faded and less evident in the last two thirds of the tail. Limbs with incomplete rings of Burnt Umber (48) on a Dark Salmon Color (59) background. Ventral areas Cream White (52) with a reticulated dark on the mandibular, gular and ventral neck regions. Some Sulphur Yellow (80) speckles on the chest and upper belly becoming less evident on the lower belly; continuous Spectrum Yellow (79) coloration along the infrafemoral region.

Color in preservative. In preservative the general brightly color pattern disappear or fade, all dorsal bands of Light Yellow Orcher (13) and Sulphur/Spectrum Yellow (80/79) coloration disappear, Sulphur Yellow (80) speckles fade but are evident.

Variation. Table 4 summarizes the variation of males and females of the type series for some scale and morphometric characters. Precloacal pores are present only in males. In males, dorsal series of roughly half-moon shaped dots became more evident in some specimens. Almost in all individuals observed, dots fused with the nearest on the dorsolateral area given and appearance of three spots, with a posterior Pale Sulphur Yellow (92) border. A male lacks of symmetrically arranged left-right half-moon dots series, with left series side missing. Patterns of transversally arranged dots are not regularly distributed. Density pattern of Caribbean Blue (168) dots varies between individuals. Ventral dark reticulation varies in density and arrangement but are limited to gular and neck region. Dark dots are dispersed on chest and belly, with color variation between Maroon (39) to Jet Black (300). Sulphur Yellow (80)/Spectrum Yellow (79) coloration is limited to gular/upper chest region and femoral region, in some individuals extended to the foot. Females ( Fig. 12 ) usually have wider bands of Light Yellow Orcher on the dorsal area; ventral surface of one female was completely and homogeneously cover with Salmon Color (83), except in mandibular and gular region. Iridescent Caribbean Blue (168) dots are absent in females.

Etymology. The specific epithet is an adjective of Greek origin ( calliston = very beautiful), in reference to the distinct and brightly colorful pattern of dorsal and lateral body of this new species.

Distribution. This species is restricted to the type locality.

Natural history. No studies on the basic biology or ecology of this new species are available; we have only anecdotal data from the original collecting trip. The type locality of Liolaemus calliston sp. nov. is characterized by sedimentary rocks separated by barren soil and sparse clumps of low bushes that have been heavily impacted by heavy road machinery. Despite our efforts, we were unable to observe any animal in less disturbed environments that surround the type locality. Lizards were collected late in the afternoon, and only a few were basking on rocks. Lizards were wary and dropped into burrows below the stones when the intruder is still at a large distance. If pressed, they ran from rock to rock across patches of bare soil or between rocks accumulated on the edges of the disturbed area, sometimes seeking cover between roots or basal root clumps. They are fast and heavy runners and tend to return to their burrow or seek shelter under rocks.

Liolaemus calliston sp. nov. is oviparous; adult females captured in December had large eggs that extended their abdominal walls, and had conspicuously slow escape runs. They appear to exist at a very low density but ongoing studies in the area will be crucial to elucidate some biological and ecological characteristics of this species. Vegetation is typical of Monte Sur Patagónico ( Roig 1998); predominant vegetation formation is a low to medium shrubs steppe characterized by bushes between 0.50 to 2.00 m. Soils are subject to wind erosion which form dwarf dunes interspersed between rock outcrops separated by bare sand dunes ( Fig. 13 ). Common plants include: Grindelia chiloensis, Senecio filaginoides , Acantholippia seriphioides , Ephedra ochreata , Chuquiraga erinacea , Larrea nitida , L. divaricata , L. cuneifolia , Atriplex lampa , Prosopis alpataco , Fabiana peckii and Adesmia guttulifera , Cercidium praecox , Sporobolus rigens , Hyalis argéntea , Monttea aphylla , Neosparton ephedroides and Senna aphylla . Lizard fauna in this general region include up to six congeners: Liolaemus darwinii , L. grosseorum , L. gracilis , L. cuyumhue , L. goetschi , L. austromendocinus , the three phyllodactylid geckos Homonota darwinii , H. horrida , H. underwoodi , two leiosaurids, Leiosaurus belli and Pristidactylus fasciatus , and the teiid Aurivela longicauda . Snakes observed at the type locality include Philodryas trilineata , Bothrops diporus , Xenodon semicinctus , Siagonodon borrichianus ; at least the first two of these are likely predators of L. calliston sp. nov.

Figures

FIGURE 10. Liolaemus calliston sp. nov., holotype adult male from 4.4 km SE Aguada Pichana Main Camp Total Oil Company, ~30 km S junction Provincial Road 7, Añelo Department,Neuquén province, Argentina.

FIGURE 11. Liolaemus calliston sp. nov., holotype adult male in dorsal and ventral view.

FIGURE 12. Liolaemus calliston sp. nov., undetermined paratype adult female from 4.4 km SE Aguada Pichana Main Camp Total Oil Company, ~30 km S junction Provincial Road 7, Añelo Department, Neuquén province, Argentina.

FIGURE 13. Type locality of Liolaemus calliston sp. nov. Upper: general view of the area. Below: close view of the microhabitat where lizards were collected.