From the far reaches of Patagonia: new phylogenetic analyses and description of two new species of the Liolaemus fitzingerii clade (Iguania: Liolaemidae) Author Abdala, Cristian Simón Author Gómez, Juan Manuel Díaz Author Heredia, Viviana Isabel Juarez text Zootaxa 2012 3301 34 60 journal article 10.5281/zenodo.246399 eea982a6-37ad-487c-9646-a80621c74856 1175-5326 246399 Liolaemus shehuen sp. nov. ( Figs. 4–7 ) 1999 Cei and Scolaro Liolaemus fitzingerii, Revue francais de Aquariologie, 26, 79-82. 2007 Liolaemus sp7. Abdala C. S., Zootaxa , 1538, 84 pp. Holotype . ― FML 22191. Adult male. 60 km west of Telsen , Telsen Department, Chubut Province, Argentina . C. S. Abdala, J. S. Abdala and M. Juarez cols. January 23, 2004 . ( Figs. 4 , 5 ). FIGURE 4. Liolaemus shehuen . Holotype. Adult male collected 60 km west of Telsen , Telsen Department, Chubut Province, Argentina. FML 22191 Paratypes . ― FML 22192-195.4 individuals. 1 male , 3 females . Same data as holotype . FML 22204-205.2 individuals. 2 males , same data as holotype . FML 22217-219.3 individuals. 1 male , 2 females . 80 km west of Telsen , Telsen Department, Chubut Province, Argentina . C. S. Abdala, J. S. Abdala and M. Juarez cols. January 23, 2004 . Diagnosis.― Liolaemus shehuen belongs to the L. boulengeri group, characterized by having a patch of enlarged scales on the posterior medial surface of the thigh ( Etheridge 1995 ; Abdala 2007 ) and hypertrophy of the flexor tibialis internus muscle ( Abdala et al. 2006 ); tail long relative to body length, head proportionally longer than wide, palpebral ‘comb’ not developed, and more precloacal pores both in males and females than in the remaining species of the clade. Liolaemus shehuen differs from the species of the L. darwinii group ( Table 2 ) in having posterior teeth with expanded crowns and different body proportions, and in lacking a transversal black line on the eye. It differs from the species of the L. wiegmannii group ( Table 2 ) in having a row of lorilabial scales (never two or three). Liolaemus shehuen differs from L. boulengeri , L. donosobarrosi , L. goetschi , L. hermannunezi (Pincheira-Donoso, Scolaro & Schulte), L. inacayali (Abdala) , L. josei (Abdala) , L. loboi (Abdala) , L. martorii (Abdala) , L. rothi (Koslowskyi 1898) L. sagei (Etheridge & Christie) , L. senguer (Abdala) , L. tehuelche (Abdala) , and L. telsen (Cei & Scolaro) in having an evident black antehumeral arch that is absent in the aforementioned species. The species differs from all the species of the L. telsen group, except L. rothi and L. sagei , in having longer snout-vent length (max SVL 90.8 mm vs 32.3–77.1 mm ). It differs from L. mapuche (Abdala) in having different dorsal body coloration, lacking light blue scales and having more marked ventral melanism. It differs from L. cuyanus in having more marked ventral melanism and different dorsal coloration. TABLE 2. Groups and species of the Liolaemus boulengeri group, sensu Abdala (2007) plus Lobo et al. (2010), and the present study. Group Species L. anomalus L. anomalus , L. ditadai , L. lentus and L. pseudoanomalus L. abaucan , L. albiceps , L. calchaquí , L. chacoensis , L. cinereus , L. crepuscularis , L. darwinii , L. L. darwinii diaguita, L. espinozai , L. grosseorum , L. irregularis , L. koslowskyi , L. montanezi , L. laurenti , L. lavillai, L. olongasta , L. ornatus , L. quilmes and L. uspallatensis L. boulengeri , L. inacayali , L. josei , L. loboi , L. martorii , L. puelche , L. rothi , L. sagei , L. senguer , L. telsen L. tehuelche , and L. telsen L. arambarensis , L. azarai , L. cuyumhue , L. lutzae , L. multimaculatus , L. occipitalis , L. rabinoi , L. L. wiegmannii riojanus, L. salinicola , L. scapularis and L. wiegmannii L. goetschi L. cuyanus , L. donosobarrosi , L. goetschi and L. mapuche L. camarones , L. canqueli , L. casamiquelai , L. chehuachekenk , L. fitzingerii , L. melanops , L. L. fitzingerii morenoi L. shehuen and L. xanthoviridis Liolaemus shehuen sp. nov. belongs to the L. fitzingerii clade ( Abdala, 2007 ), which is characterized by its tricuspidate posterior teeth with expanded crowns, neck equal to or wider than head, cylindrical body, short limbs relative to the trunk, black antehumeral arch present and ventrally expanded, and males with ventral melanism. Within the Liolaemus fitzingerii clade, L. shehuen sp.nov. has four to six scales in contact to mental (as the southern populations of L. fitzingerii ). In L. shehuen the head melanism characteristic of L. canqueli and L. melanops is absent. Liolaemus shehuen lacks the intense green dorsum with irregular, transverse black paravertebral spots of L. melanops , and the transverse black and yellow-orange bands of L. canqueli . Liolaemus shehuen differs from L. chehuachekenk in lacking a conspicuous vertebral band, having fewer individuals with pre- and postscapular spots, fewer ventral scales (109-121, mean=115.6 vs. 99-116, mean=106), and shorter snout-vent length (SVL 90.8 mm vs. 98.3 mm ); it differs from L. fitzingerii in having shorter snout-vent length (SVL 90.8 mm vs. 106 mm ), and lacking the bluish green or light blue dorsum coloration with red paravertebral spots and red scales speckled, as L. fitzingerii has. Liolaemus shehuen differs from L. morenoi in lacking evident scapular markings and having more ventral scales (109-121, mean=115.6 vs. 98-113, mean=107.6). L. morenoi never presents yellow-red dorsal coloration either. Liolaemus shehuen differs from L. xanthoviridis in having different dorsal and ventral coloration, with paravertebral spots and more uniform vertebral region without irregular markings. Description of the holotype . ― Adult male. SVL 90.6 mm . Trunk length 45.1 mm . Head longer ( 17.4 mm ) than wide ( 14.8 mm ). Head height 11.6 mm . Eye diameter 4.3 mm . Interorbital distance 8.7 mm . Orbit–auditory meatus distance 6.6 mm . Auditory meatus 3.1 mm high, 1.6 mm wide. Orbit–commissure of mouth distance 2.5 mm . Internares distance 3.0 mm. Subocular scale length 5.4 mm . Femur length 13.5 mm , tibia 13.7 mm , and foot 25.6 mm . Humerus length 13.1 mm . Tail length 120.2 mm . Dorsal surface of the head smooth, with 15 scales. Rostral wider than long, bordered by six scales. Mental larger than rostral, trapezoidal, bordered by four scales. Nasal not in contact with rostral. Four internasals. Nasal surrounded by eight scales, separated from canthal by two scales. Five scales between frontal and supercilliaries. Seven scales between frontal and rostral. Frontal not divided. Two postrostrals. Interparietal smaller than parietals, in contact with six scales. Orbital semicircles complete. Preocular separated from lorilabial row by one scale. Three scales in anterior margin of auditory meatus. Eleven smooth temporals. Three lorilabials in contact with subocular. Eight supralabials, none in contact with subocular. Eight supraoculars. Eight lorilabials. Five infralabials, second ventrally in contact with two scales. Six chinshields, second pair separated by three scales. Seventy-six scales around midbody. Eighty-six round, slightly imbricate, slightly keeled dorsals from occiput to hind limbs. Thirty-six scale rows in dorsum. Scales of flank same size and shape as dorsals. Ventrals same size as dorsals, flat, imbricate. Thirty-five smooth, weakly imbricate gulars. Eight precloacal pores. Antehumeral scales flat, larger than or equal in size to dorsals. Postauricular, rectal, and longitudinal folds present. Scales on the longitudinal fold granular and smooth. Fourth finger with 19 subdigital lamellae; fourth toe with 26. Infracarpals flat, imbricate, trifid; infratarsals flat, imbricate, not trifid. C olor in life .― Head grey with scattered red spots and scales on the temporal and occipital region. Conspicuous antehumeral arch. Dorsum and flanks yellow-red with faint, small paravertebral spots. Some of these spots on the posterior region have a lighter spot inside. Vertebral region orange, with two light grey stripes on each side of vertebral region. Dorsal fore- and hind limbs same color as trunk. Dorsal tail greenish grey with yellow and orange scales on flanks and at tail base. Throat pale white-yellow. Gular region, chest to cloacal region and ventral hind limbs melanic. Flanks, forelimbs and ventral tail same color as throat. FIGURE 5. Liolaemus shehuen . Holotype, ventral view. Variation .― (Table 3). Based on 10 specimens . Head longer ( 14.8–17.9 mm ; mean = 16.4 mm ) than wide ( 11.9–14.8 mm ; mean = 13.4 mm ). Head height 9.8–11.6 mm (mean = 10.7 mm ). Neck wider than head. SVL 72.7–90.8 mm (mean = 79.9 mm ), averaging 2.1 times longer than trunk. Tail length 94.9–120.3 mm (mean = 109.6 mm ). Humerus length 9.7–13.1 mm (mean = 11.4 mm ). Dorsal surface of head smooth, with 14–18 scales. One row of lorilabials, same size as supralabials. Six to nine lorilabials. Seven to 10 supralabials. Frontal unique or divided horizontally, into two or four scales. Interparietal always smaller than parietals, surrounded by 6–8 scales. Nasal surrounded by 8–9 scales. Five to seven infralabials. Mental in contact with four or six scales. Ten to 11 smooth round temporals. Longitudinal, postauricular, and antehumeral folds present. Horizontal fold Y-shaped between shoulder and auditory meatus.

TABLE 3. Variation in species of the Liolaemus fitzingerii group. Character Liolaemus Liolaemus Liolaemus Liolaemus /Species camarones canqueli casamiquelai chehuachekenk Mid-body 71-79 70-79 64-79 69-75 Scales (73.3) (75.7) (71.8) (72.0)	Liolaemus fitzingerii type locality 68-76 (71.5)	Liolaemus fitzingerii SO Santa Cruz 66-74 (71.8)	Liolaemus Liolaemus Liolaemus fitzingerii melanops morenoi CS Chubut 75-83 64-74 66-85 (78.4) (69) (73.6)	Liolaemus sp. 14 60-76 (68.2)	Liolaemus Liolaemus sp. 15 shehuen 61-74 65-80 (67.2) (73)	Liolaemus xanthoviridis 64-75 (69.2)
Ventral 109-120 115-129 98-119 99-116 Scales (114.8) (118.8) (108.8) (106) Dorsal 71-78 68-78 71-84 73-81 Scales (74.0) (73.8) (78.3) (77.5) Maximum 95.6 99.6 106.2 98.3 SVL Around Mental 4 4 4 4 Scale Precloacal 7-10 6-9 7-12 7-10 Pores in Males Sexual Yes No No No Dicrhomatism	114-136 (122.9) 68-84 (76) 110 4 8-12 No	119-134 (125.2) 68-77 (72.2) 106.5 4-6 8-10 No	116-120 95-104 98-113 (117.9) (99.6) (107.6) 73-85 73-84 69-84 (77.0) (77.5) (73.1) 96.2 79.7 88.4 4 4 4 8-10 8-10 7-10 No Yes No	99-115 (107.3) 63-85 (75.5) 88.3 4 5-11 No	102-120 109-121 (109.8) (115.6) 72-83 63-88 (77.2) (77.6) 83.7 90.8 4 4-6 4-8 7-9 No No	103-119 (113.5) 73-78 (76.2) 101.2 4 6-11 No
Head Males: greenish Black Black Ochre Color grey or brown. or or Females: grey or Grey grey reddish brown Dorsal Body Males: dark green Yellow Yellow Ochre Color or greenish or or or brown. Females: reddish grey grey greenish brown Lateral Body Males: greenish Yellow Yellow Ochre Color grey. Females: or or or light brown orange grey grey Dorsal Body Black and white Black Black Wide vertebral Pattern round transversal transversal band paravertebral bands bands with dotted spots, dorsolateral transverse redish bands dorsolateral marks	Light blue grey - reddish Greenish light blue - grey Greenish light blue to grey Red dorsolateral bands, and/or red paravertebral spots	Light blue grey with black occipital band Greenish light blue - grey Greenish light blue to grey Red dorsolateral bands, and/or red black paravertebral spots	Light blue Males: black Grey grey or grey. or to red Females: grey light brown or brown Grey -reddish Males: Green. Grey-brown Females: grey or brown Greenish light Males: green. Grey blue to grey Females: grey or or brown brown Red dorsolateral Males: Black Black transversal bands, and/or Transversal bands paravertebral red or black Females: Black spots, white paravertebral and white posterior margin spots paravertebral spots.	Greenish grey Reddish, yellowish, greenish Red and green Reddish bands, transversal or longitudindal paravertebral spots	Grey Grey or light brown Yellowish red Orange yellow or reddish yellow Yellowish red Orange yellow or reddish yellow Round Dim paravertebral paravertebral spots present or spots, White absent; dark thin posterior margin. dorsolateral bands Red bands present or absent present/absent	Grey to yellow Yellow Yellow Black spots and irregular black
Pre- and Absent Absent Absent Present Postscapular Black Spots Antehumeral Normal Wide Normal Normal Arch	Absent Normal	Absent Normal	Absent Absent Present Normal Normal Normal	Absent Normal	Absent Present or Absent Wide Wide	Absent Very wide
Melanic No Slightly Slightly No Forelimb Jaw Melanism No Yes No No Gular Collar Complete/ Complete Complete Complete Incomplete Ventral Narrow Expanded Narrow Narrow Melanism Melanic Vent No Present Present/ Present and Hind limbs absent	No No Complete/ Incomplete Narrow No	No No Complete/ Incomplete Narrow No	No No or Slightly in No males No Males:Yes/No No Females: No Complete/ Complete Complete/ Incomplete Incomplete Narrow Narrow/ Narrow Expanded No Males:Yes/No Present/ Females: No absent	No or Slightly Yes Complete/ Incomplete Expanded Present	Slightly No Yes Yes/No Complete/ Complete/ Incomplete Incomplete Expanded Narrow Present Present/ Absent	Yes Yes Complete Expanded Present

Scales around midbody 69–80 (mean = 74). Gulars 28–35 (mean = 31.5). Dorsal scales between occiput and hind limbs 63–88 (mean = 77.6). Ventrals 110–119 (mean = 117.3). Males with 8–9 (mean = 8.2) precloacal pores. Females with 0–3 (only one female) (mean = 0.5) precloacal pores. Dorsals flat, slightly imbricate, slightly keeled. Seventeen to 20 infradigital lamellae on fourth finger and 22–29 on fourth toe. Variation in color pattern. ― No sexual dichromatism evident ( Figs. 6–7 ). Head usually grey or light brown with small yellow-orange spots or scales. Black arch ventrally expanded over antehumeral fold. Dorsal trunk and limbs yellow-orange or yellow-red, more intense in some specimens. Faint and small round paravertebral spots, generally dark, but sometimes preceded by a lighter spot. Some specimens have one or two dorsolateral whiteyellow strips and/or vertebral line of same color as paravertebral spots. Some specimens have diffuse white-yellow bars transversal to the body axis. Lateral region of the body same color as dorsum. Tail uniformly bluish grey or greenish grey with scattered spots. Gular region, chest to cloacal region, anterior forelimbs and hind limbs have ventral melanism. Males more melanic than females. In juveniles melanism expands from gular region to cloacal region as the individual grows. FIGURE 6. Liolaemus shehuen , adult female from type locality, dorsal view. Natural history .― Liolaemus shehuen sp. nov. was found in the Telsen plateau and nearby regions, along National Route 4, an area with sand dunes and abundant vegetation, an ecotone between the phytogeographic regions of Monte and Patagonia ( Cabrera & Willink 1980 ). The soil is mostly sandy with basaltic rocks, surrounding a lowland with red soils called ‘Bajo de la tierra colorada’. The vegetation consists mainly of Stipa , Verbena , Mulinum , Nassauvia and Lycium ( Cei & Scolaro 1999 ) . The Patagonian plateaus ( Telsen , Somuncurá in Río Negro and Chubut provinces, and Buenos Aires Lake, in Santa Cruz Province) are areas with high herpetofauna diversity. Liolaemus shehuen sp. nov. is sympatric with Diplolaemus darwinii (Bell) , Homonota darwinii (Boulenger) , L. bibronii (Bell) , L. petrophilus (Donoso-Barros & Cei) , L. rothi , L. telsen , Leiosaurus belli (Duméril & Bibron) , Pristidactylus nigroingulus (Cei, Scolaro & Videla) , the snakes Philodryas patagoniensis (Girard) , Philodryas trilineata (Burmesteir) , and the viperid Bothrops ammodytoides (Leybold) . Liolaemus shehuen is oviparous and insectivorous. It is active between 12.00 AM and 15:00 PM, when the sand temperature is highest. There is no more information about its biology. FIGURE 7. Liolaemus shehuen , adult female from type locality, ventral view. Distribution.― Liolaemus shehuen is distributed on the Telsen plateau and east of Río Negro Province, Argentina ( Fig. 8 ). Etymology.― shehuen is a Tehuelche word that means 'Sun', and makes reference to the dorsal coloration of Liolaemus shehuen . The tehuelches, also known as Patagones, are ancient indigenous people that have inhabited Patagonia for more than 10000 years.