Liolaemus goetschi (Iguania: Liolaemidae): redescription and phylogenetic relationships within the L. boulengeri group Author Nori, Javier Author Abdala, Cristian Simón Author Scrocchi, Gustavo José text Zootaxa 2010 2440 49 59 journal article 10.5281/zenodo.194953 714c16ae-49d5-47a0-aef6-8bef5f5daf47 1175-5326 194953 Liolaemus goetschi Müller and Hellmich, 1938 ( Fig. 1–3 ) 1938. Liolaemus goetschi. Müller and Hellmich. Zoologischer Anzeiger, 123(5–6): 130–142 . 1977 . . Cei, J. M. and J. A. Scolaro. Physis (Buenos Aires) , 36 : 225–226 . 2003. Liolaemus goetschi. Cei and Scolaro. Facena, 19: 163. 2006. Liolaemus goetschi . Avila et al . Biol. J. Linnean Soc., 89: 241–275 . 2007 . . Zootaxa , 1538 : 1–84 . 2008 . . Zootaxa , 1800 : 1–85 . 2010. Liolaemus goetschi. Nori et al. Check List, 6(1): 6-4. Holotype . ZSM 4. Adult male, Laguna Playa, approximately 20 km north of General Roca, Río Negro Province, Argentina , W. Goetsch, 1938. Paratype . ZSM 5. Adult female, Laguna Playa, approximately 20 km north of General Roca, Río Negro Province, Argentina , W. Goetsch, 1938. Type locality. In the original description of Liolaemus goetschi , the authors designated Laguna Playa as the type locality of L. goetschi (approximately 20 km north of General Roca city, the site where Prof. Dr. Goetsch collected all the individuals corresponding to the type series) as the type locality of L. goetschi . FIGURE 1 . Dorsal view of adult male of Liolaemus goetschi from the type locality FIGURE 2 . Ventral view of adult male of Liolaemus goetschi from the type locality Because the toponym “Laguna Playa” is not included in maps and records of the area and is unknown to local people, we conducted extensive surveys among rural people to locate the site. Finally, an elderly inhabitant of the nearest city (General Roca) informed us about the place and following his directions, we were able to find the exact place, located at 30º 52’ 59” S and 67º 34’ 08” W . The Liolaemus goetschi individuals were collected in subsequent intensive field surveys. Diagnosis. Liolaemus goetschi belongs to the L. boulengeri group because of the presence of the femoral patch of enlarged scales in the posterior surface of the femur (Etheridge 1995). Within this group, L. goetschi is distinguished from the L. anomalus group because of the hypertrophy of the puboischiotibialis muscle (Abdala et al . 2006), higher percentage of tail autotomy, tail longer than snout-vent length (SVL), head longer than wide, outer cilliaries not projecting and higher number of precloacal pores in males. It also differs from the species of the L. wiegmannii group in having a row of lorilabials between subocular and supralabials and four scales surrounding the mental (Etheridge 2000). FIGURE 3 . Dorsal view of adult female of Liolaemus goetschi from the type locality Liolaemus goetschi differs from species of the L. laurenti group (Abdala 2007), L. abaucan , L. albiceps , L. calchaquí , L. chacoensis , L. crepuscularis , L. darwinii , L. espinozai , L. grosseorum , L. irregularis L. koslowskyi , L. laurenti , L. lavillai , L. olongasta , L. ornatus , L. quilmes and L. uspallatensis in having posterior teeth with strongly cusped crowns and expanded margins, and in the presence of barely evident sexual dichromatism (Abdala 2007). Females of L. goetschi lack precloacal pores; this character distinguishes the species from L. albiceps , L. calachaqui , L. crepuscularis , L. irregularis , L. lavillai and L. ornatus because a high percentage of females of all these species have 1–6 precloacal pores (Abdala 2007). Within the L. melanops group (Abdala 2007), L. goetschi differs from L canqueli , L. fitzingerii , L. melanops , L. rothi , L. sagei , and L. xanthoviridis in having a shorter snout-vent-length (SVL) (max SVL 74.25 mm vs. 89.00– 106.00 mm) and because of the presence of pre and postscapular spots (absent in these species). It also differs from L. canqueli and L. melanops because of the abscence of the cephalic melanism typical of these species It differs from L. rothi and L. sagei in the more conspicuous gular melanism (Abdala 2007) and from L. morenoi in its smaller size and lower number of scales around the midbody (72–85 Mean = 79.00 vs. 62–72 Mean = 66.35). Liolaemus goetschi differs from Liolaemus inacayali by the presence of pre and postscapular spots and a lower number of scales from occiput to tigh (73–84, Mean = 77.50 vs. 83–96, Mean = 88.75) (Abdala 2003). It differs from L. cheuachekenk in having a shorter maximum SVL ( 74.25 mm vs. 98.30 mm ), in the presence of four to six scales in contact with the mental scale ( L. cheuachekenk always presents four), in the absence of abdominal and pectoral melanism and in the presence of a different color pattern (Avila et al. 2008). L. goetschi differs from L. puelche in its smaller size (max SVL 74.25 mm vs. 89.00 mm in L. puelche ), lower number of scales around midbody (62–72, Mean = 66.35 vs. 67–76, Mean = 70.75), gular melanism and two series of well defined black paraventral spots, absent in L. puelche (Avila et al. 2007) . Within the Liolaemus cuyanus clade (Abdala 2007), composed of L. mapuche , L. cuyanus , and L. donosobarrosi , L. goetschi differs from the first two species in having smaller SVL (max SVL 74.25 mm vs. 79.00 and 102.00 mm, respectively) and a clearly different color pattern that never has a black antehumeral arch (Abdala 2002). It differs from L. donosobarrosi in having longer SVL (max SVL 74.25 mm vs. 60.80 mm ), a lower number of scales around the midbody (62–72, Mean = 66.30 vs. 79–95, Mean = 85.40) and a different color pattern (Abdala 2005, 2007). Liolaemus goetschi differs from L. boulengeri , L. josei , L. loboi , L. senguer , L. tehuelche and L. telsen , in having a faint prescapular spot and a larger postscapular spot; two black spots usually band -or line- shaped on each side of the gular region; belly in males and females white or light pink, never yellow, red or bright blue; anterior throat in males never melanic and sexual dichromatism absent or barely marked. It also differs from L. martorii in having a longer SVL (max SVL 74.25 mm vs. 67.10 mm ), and four to six scales in contact with mental scale ( L. martorii always has four); L. goetschi also has a higher number of light blue scales in tail and body; also scapular spots and spots on the sides of body that are larger and more marked ( Table 1 ). TABLE 1 . Main differences between Liolaemus goestchi , L.melanops and L. martorii . Character L. goetschi L. martorii L. melanops External morphology. Maximum SVL 74.25 mm ( 56.20 mm – 74.25 mm ; Mean = 65.80 mm ); tail length (93.00 mm – 105.90 mm ; Mean = 98.20 mm ); 1.4 times longer than SVL. Head 1.28 times longer ( 12.10 mm – 17.55 mm ; x = 14.54 mm ) than wide ( 9.20 mm – 12.55 mm ; Mean = 11.31 mm ). Head height ( 6.70 mm – 14.10 mm , Mean = 8.59 mm ). Humerus length ( 7.10 mm – 10.70 mm , Mean = 8.27 mm ). Radius length ( 6.60 mm – 9.20 mm ; Mean = 8.11 mm ). Hand length ( 7.60 mm – 9.90 mm , Mean = 8.26 mm ). Tibial length (10.00 mm – 14.15 mm , Mean = 11.68 mm ).

(n = 20)	(n = 13)	(n = 13)
Maximum SVL	74.25 mm	67.10 mm	79.66 mm
Cephalic melanism	Absent	Absent	Present

Light blue scales on body sides Abundant	Scarce	Variable
Size and color of scapular Large and marked spots of body	Middle-sized and light	Variable
Number of scales in contact 4 to 6 with the mental	4 (always)	6 (always)
Tail length 93.00 to 105.90 mm, (Mean = 98.20 mm)	86.10 to 92.30 mm, (Mean = 87.30 mm)	81.90 to 116.50 mm, (Mean = 106.10 mm)
Head length 13.40 to 17.50 mm, (Mean = 15.50 mm)	13.00 to 15.80 mm, (Mean = 14.60 mm)	13.30 to 16.70 mm, (Mean = 14.60 mm)
Hand length 7.60 to 9.90 mm, (Mean = 8.26mm)	6.90 to 8.20 mm, (Mean = 7.40 mm)	7.40 to 10.90 mm, (Mean = 8.48 mm)
Gular scales 30 to 36, (Mean = 32.80)	32 to 39, (Mean = 34.6)	33 to 42, (Mean = 36.5)
Dorsal body scales 68 to 78, (Mean = 72.70)	64 to 80, (Mean = 76.20)	73 to 84, (Mean = 77.50)

Dorsal head scales smooth, convex and bulged. Scale organs distributed in postrostrals, internasals, frontonasals and prefrontals. Rostral pentagonal wider than long. Two postrostrals in contact with anterior supralabials. 15–18, (Mean = 16.25) dorsal head scales, from rostral to occiput (Hellmich Index). Nasal scales subpentagonal and in dorsolateral position. Eigth to 12 scales surrounding nasal. Frontal scale irregularly shaped, slightly longer than prefrontals. Interparietal pentagonal with a conspicuous pineal eye and surrounded by seven scales. Two parietals of irregular shape and larger than interparietals. Interparietal in contact with 6–10 (Mean = 7.30) scales. 9–14 (Mean = 12.30) circumorbitals. 6–8 (Mean = 7.15) laterally expanded supraoculars. Two rows of small scales between supraoculars and supercilliaries. 6–10 (Mean = 7.80) elongated and overlapped supercilliaries. One preocular longer than wide; one elongated subocular and one small postocular. A single row of 6–7 (Mean = 6.9) convex lorilabials. Supralabials 7–10 (Mean = 8.50) subquadrangular and elongate. Temporal scales 17–22 (Mean = 19.10) smooth and juxtaposed. Auditory meatus 1.86 times higher ( 2.30 mm – 4.00; Mean = 3.08) than wide ( 1.30 mm – 2.10 mm ; Mean = 1.68 mm ) surrounded by small granular scales. Mental scale pentagonal, wider than high in contact with 4–6, (Mean = 4.70) scales. Infralabials, 6–8,(Mean = 6.80) large and subquadrangular. Gular scales 30–36 (Mean = 32.80) smooth and imbricate. Y-shaped horizontal fold. Dorsal scales at midbody subtriangular, moderately imbricate and keeled, arranged in 22 longitudinal rows. Ventral scales along body midline: 79–89 (Mean = 74.15), smooth and moderately imbricate, subtriangular, of equal size or slightly larger than dorsal scales at midbody. 11–15 pigals (Mean = 12.55). 4–8 (Mean =4.70) precloacal pores in males, absent in females. 62–72 (Mean = 66.35) scales around midbody. 68–78 (Mean = 72.70) dorsal scales of midbody from occiput to hind limbs. Suprahumerals and supraradials keeled and imbricate. Supratarsals smooth and imbricate. Infratarsals strongly imbricate. Supracarpals smooth and imbricate. Infracarpals keeled and imbricate. Supradigital lamellae smooth and imbricate. Infradigital lamellae tricarinate and imbricate. Fourth finger with 17–22 (Mean = 19.30) scales. Suprafemorals imbricate and rhomboidal. Infrafemorals smooth, imbricate and rhomboidal. Supratibials keeled and imbricate. Bulged femoral patch with 20–25 scales. Infratibials and supradigitals smooth and imbricate. Infradigitales tricarenate. Coloration . Males and females with slightly different coloration pattern. In most males, numerous light blue scales can be observed on the sides of tail and fewer on the sides of body and head. In females, these colors are absent in the scales. Dorsal color of females is more yellowish and usually white immaculate ventrally. FIGURE 4 . Map showing the known distribution of Liolaemus goetschi and L. martorii . Stars: localities where L. goetschi were located. Rhombuses: localities where L. martorii were located. Dorsum of head brown, with irregularly spread black spots. Nasal scale slightly lighter. A black band extends from the posterior zone of the eye to the upper lip margin. This band is bordered anteriorly and posteriorly by white spots. Also, a dark line posterior to the eye is present. Dorsal background coloration light brown. Also, has pre and postscapular spots, the latter larger than the former. Four series of black subquadrangular spots are present, two paravertebral and two lateral; both edged posteriorly by a white band. Nine to 12 black paravertebral spots are present, which are larger and more marked than the lateral spots. Paravertebral spots extend to the dorsal tail proximally to fuse, forming irregular spots less marked than those in the body. Anteriorly to the paravertebral spots, a reddish subquadrangular spot may be present. In many specimens, the vertebral region has isolated yellowish scales. Sides of body brown, slightly lighter than dorsum. Lateral region of tail and dorsal region of limbs light brown, similar to that of the body sides and slightly lighter than dorsum. Background of throat, chest and belly white or light pink. Ventral gular region is melanic or dark, reticulate anteriorlly, variegated with gray to black lines. Natural history. This is a typical psammophylus lizard of the Monte phytogeographical region (Cabrera, 1976). Vegetation in the area is typical of the Monte, with dominance of Atriplex lampa , Cercidiurn praecox , Larrea divaricata , L. nítida and L. cuneifolia . Liolaemus goetschi seems to be most related to Larrea species. The species is difficult to observe and capture; individuals were found mainly at the time of the day when sand surface temperatures reach the highest values, between 13:00 and 15:00 hours. It is a relatively quiet species that, when chased, tends to hide and stay still or seek refuge in small caves. Most of the species studied in the group are oviparous and omnivorous (Abdala 2007); we suppose that this species shares the same characteristics. During field work, we observed some specimens feeding on ants and insect larvae. Distribution ( Fig. 4 ). Liolaemus goetschi inhabits the province of Río Negro. This species was found in its type locality (Laguna Playa), in Cerro Policia Town (Nori et al . 2010) and at 20 and 50 km south of the locality of Catriel. The specimens collected south of Catriel did not show morphological differences that enabled us to assume that they belong to a species different from the species of those specimens collected in Laguna Playa. All the indivuals are representatives of Liolaemus goetschi . Müller and Hellmich (1938) mentioned the existence of Liolaemus populations south of Catriel that might correspond to the same species. This assumption is supported by the specimens studied in the present work. Phylogenetic position. The results of the phylogenetic analysis based on the morphological data matrix provided by Abdala (2007) and the incorporation of the 128 characters of Liolaemus goetschi from the type locality are presented in Figure 5 . Here Liolaemus goetschi is included in the L. telsen group, which comprises: L. boulengeri L. goetschi , L. josei , L. loboi L. martorii , L. tehuelche , L. telsen and L. senguer . In our analysis, L.goetschi and L. martorii appear as sister species; this hypothesis suggests a close phylogenetic relationship between the two species. L. telsen group is part of the L. melanops group, which is included in the L. chacoensis group, all of them within the L. boulengeri group.