A new species of Liolaemus (Iguania: Liolaemidae) from the hot deserts of northern Patagonia, Argentina Author Abdala, Cristian S. CCA9FC01-29B4-4B0E-B47F-306F992B516D Consejo Nacional de Investigaciones Científicas y Técnicas, Unidad Ejecutora Lillo (CONICET-UEL), Instituto de Herpetología, Fundación Miguel Lillo, Facultad de Ciencias Naturales e IML, Universidad Nacional de Tucumán (UNT), Tucumán, Argentina samiryjazmin@gmail.com Author Chafrat, Pablo Anselmo 57AD5156-8BD0-4E39-916B-3426C621C052 Museo Patagónico de Ciencias Naturales “ Juan Carlos Salgado ”, Fundación Patagónica de Ciencias Naturales, Departamento de Biodiversidad, General Roca, Río Negro, Argentina chafrat@gmail.com Author Chaparro, Juan C. 27BEB2DB-8514-42E6-B135-04B00C8213A6 Museo de Biodiversidad del Perú, Urbanización Mariscal Gamarra A- 61, Zona 2, Cusco, Peru. & Museo de Historia Natural de la Universidad Nacional de San Antonio Abad del Cusco, Paraninfo Universitario (Plaza de Armas s / n), Cusco, Peru. jchaparroauza@yahoo.com Author Procheret, Iván Ezequiel 6E49DB1B-33D7-46ED-971B-F1CC06F3B598 Geólogos Asociados S. A., Área de Sostenibilidad Ambiental y Biodiversidad, Mendoza, Argentina. iprocheret@geologos.com.ar Author Valdes, Julián C67EA2B4-DBA4-448E-A499-395C7A4B4421 Instituto de Botánica del Nordeste (UNNE-CONICET), Corrientes Capital, Corrientes, Argentina. Julianvaldes@hotmail.com.ar Author Lannutti, Vanina 0024688B-9759-45FA-853A-A6AD181A57C1 Geólogos Asociados S. A., Área de Sostenibilidad Ambiental y Biodiversidad, Mendoza, Argentina. vlannutti@geologos.com.ar Author Perez, Laura F4F871EC-0904-488D-B875-E5DD2E9D93BA Cátedra de Ecología, Facultad de Ambiente y Arquitectura, Universidad de Congreso, Mendoza, Argentina. p_mlaura@hotmail.com Author Quinteros, Sebastián 47CB1454-8F6B-45FF-AFDC-B904C1872AE8 Instituto de Bio y Geociencias del NOA, IBIGEO-UNSa-CONICET, Cátedra de Sistemática Filogenética, Facultad de Ciencias Naturales, Universidad Nacional de Salta, Salta, Argentina. Sebasquint@gmail.com text European Journal of Taxonomy 2023 2023-08-29 890 136 164 http://dx.doi.org/10.5852/ejt.2023.890.2257 journal article 264973 10.5852/ejt.2023.890.2257 e74353f2-17c0-476c-bdda-09b64819395d 2118-9773 8305643 9604CC50-C12E-4F79-B553-E3A3560A1FE5 Liolaemus kulinko sp. nov. urn:lsid:zoobank.org:act: E97AAE2D-03AC-4A21-A4A0-A18A3C2F282B Figs 1–2 , 5–7 , Table 1–2 Diagnosis Liolaemus kulinko sp. nov. belongs to the section of Liolaemus montanus Koslowsky, 1898 of the Liolaemus boulengeri group ( Abdala 2007 ; Abdala et al. 2021c ) because it has a patch of enlarged scales on the posterior aspect of the thigh ( Etheridge 1995 ; Abdala 2007 ) ( Fig. 1D ). Within the group of L. boulengeri it belongs to the clade of L. melanops , subclade of L. goetschi , and L. cuyanus complex because it has light blue scales on the flanks of the body and tail, a black margin on the posterior border of the paravertebral spots, four to six scales in contact with mental scale ( Fig. 1E ), presence of a melanic gular ring, evident scapular spots, and the same body shape and similar lepidosis ( Abdala 2007 ; Abdala et al. 2012b , 2021c ). It differs from the species of the clades of L. anomalus and L. darwinii by having posterior teeth with crowns of expanded edges and four to six scales in contact with mental scale. It also differs from the species of the clade L. anomalus ( Abdala & Juárez Heredia, 2013 ) ( Liolaemus acostai Abdala & Juárez-Heredia, 2013 , L. anomalus , L. ditatadi Cei, 1983 , L. lentus Gallardo, 1966 , L. millcayac Abdala & Juárez-Heredia, 2013 , L. pipanaco Abdala & Juárez-Heredia, 2013 and L. pseudoanomalus Cei, 1981 ) by having less developed palpebral strap or ‘comb’, males with a greater number of precloacal pores, a greater relationship between snout–vent length (SVL) and the tail length (TL). Liolaemus kulinko sp. nov. also differs from the species of the clade of L. darwinii ( Abdala, 2007 ) ( L. abaucan Etheridge, 1993 , L. calchaqui Lobo & Kretzschmar, 1996 , L. chacoensis Shreve, 1948 , L. cinereus Monguillot, Cabrera, Acosta & Villavicencio, 2006 , L. crepuscularis Abdala & Diaz Gómez, 2006 , L. darwinii , L. diaguita Abdala, Quinteros, Arias, Portelli & Palavecino, 2011 , L. espinozai Abdala, 2005 , L. grosseorum Etheridge, 2001 , L. koslowskyi Etheridge, 1993 , L. laurenti Etheridge, 1992 , L. lavillai Abdala & Lobo, 2006 , L. messi Ruiz, Quipildor, Ruiz-Monachesi, Escalante, Valdecantos & Lobo, 2021 , L. montanezi Cabrera & Monguillot, 2006 , L. olongasta Etheridge, 1993 , L. ornatus Koslowsky, 1898 , L. pacha Juárez Heredia, Robles & Halloy, 2013 , L. quilmes Etheridge, 1993 , and L. uspallatensi s Macola & Castro, 1982) in having less evident sexual dichromatism, fewer precloacal pores in males, and a clearly different dorsal coloration pattern. It differs from the species of the clade L. wiegmannii ( Etheridge 2000 ) ( L. arambarensis Verrastro, Veronese, Bujes & Martins Dias Filho, 2003 , L. azarai Ávila, 2003 , L. cranwelli (Donoso-Barros, 1973) , L. cuyumhue , L. lutzae Mertens, 1938 , L. multimaculatus (Duméril & Bibron, 1837) , L. occipitalis Boulenger, 1885 , L. rabinoi (Cei, 1974) , L. riojanus (Cei, 1979) , L. salinicola Laurent, 1986 , L. scapularis Laurent, 1982 , and L. wiegmannii ) in having one row of loreolabial scales (never two or three). Within the L. melanops group, L. kulinko sp. nov. differs from L. dumerili Abdala, Semhan, Moreno Azocar, Bonino, Paz & Cruz, 2012 , L. josei Abdala, 2005 , L. martorii Abdala, 2003 and L. loboi Abdala, 2003 by having a longer SVL, the presence of an evident antehumeral arch, and greater number of dorsal scales. It differs from L. tromen Abdala, Semhan, Moreno Azocar, Bonino, Paz & Cruz, 2012 by having a greater number of scales around the body, greater number of dorsal scales, and less ventral melanism. It differs from the Liolaemus telsen subclade ( Abdala 2007 ; Abdala et al. 2021c ) ( L. boulengeri , L. inacayali Abdala, 2003 , L. purul Abdala, Semhan, Moreno Azócar, Bonino, Paz & Cruz, 2012 , L. senguer Abdala, 2005 , L. tehuelche Abdala, 2003 , and L. telsen Cei & Scolaro, 1999 ) in having a greater SVL, four to six scales in contact with mental scales and a clearly different dorsal coloration pattern. It differs from the L. rothi subclade ( Abdala 2007 ; Abdala et al. 2021c ) ( L. hermannunezi Pincheira-Donoso, Scolaro & Schulte, 2007 , L. sagei Etheridge & Christie, 2003 , L. sitesi Ávila, Olave, Perez, Perez & Morando, 2013 , and L. rothi Koslowsky, 1898 ) by having light blue scales on the flanks of the body and tail, four to six scales in contact with the mental scale, and a clearly different dorsal and ventral coloration pattern. It differs from the L. fitzingerii complex ( Abdala 2007 ; Abdala et al. 2021c ) ( L. camarones Abdala, Díaz-Gómez & Juarez-Heredia, 2012 , L. canqueli Cei, 1975 , L. casamiquelai Ávila, Perez, Morando & Sites, 2010 , L. chehuachekenk Ávila, Morando & Sites, 2008 , L. fitzingerii (Duméril & Bibron, 1873) , L. melanops , L. morenoi Etheridge & Christie, 2003 , L. shehuen Abdala, Díaz-Gómez & Juarez-Heredia, 2012 , and L. xanthoviridis Cei & Scolaro, 1980 ) in having a shorter SVL, less ventral melanism, four to six scales in contact with mental scales, and a clearly different dorsal coloration pattern. Within the L. cuyanus complex, it differs from L. calliston , L. donosobarrosi , L. hugoi , and L. tirantii by having a greater SVL, four to six scales in contact with the mental scales, and a clearly different dorsal coloration pattern. It differs from L. cuyanus , and L. puelche by having a smaller SVL, four to six scales in contact with mental scale, presence of light blue scales in males and evident sexual dichromatism. It differs from L. goetschi by having a greater SVL, more evident sexual dichromatism, presence of an antehumeral arch, a greater quantity of light blue scales, greater number of scales around the body, greater number of dorsal scales, and fewer ventral scales. It differs from L. mapuche by having a greater number of dorsal scales between the occiput and the level of the anterior aspect of the thigh (90–96 vs 70–86), a greater number of loreolabial scales (8–10 vs 6–7), lower number of gular scales (22–25 vs 25–35), greater number of dorsal scales on the head (16–18 vs 13–16), a smooth head surface in L. kulinko sp. nov. but a wrinkled head surface in L. mapuche , and the absence of precloacal pores in females and up to six in L. mapuche . The color of the head differs in males of L. kulinko sp. nov. , that varies from light brown to gray, while in L. mapuche it is generally blue or light blue (although in some individuals there is the light gray color) and the pattern of body coloration in paravertebral and posterolateral spots of males in L. mapuche does not fade as they do in L. kulinko sp. nov. ( Figs 2–4 , see Table 1 for more differences.) Etymology The specific epithet ‘kulinko’ means ‘aguada’ in the language of the Mapuche, a group of indigenous inhabitants of south-central Chile and southwestern Argentina , including parts of Patagonia, and refers to the place where the species lives, “Aguada Pichana”. Table 1. Morphological differences between L. kulinko sp. nov. and its closest relative, L. mapuche Abdala, 2002 . * =Morphological data of number of specimens examined and traits.
species/ characters L. mapuche Abdala, 2002 (n = 26) L. kulinko sp. nov. (n = 13)*
min–max SVL (mm) 61.97–82.70 72.88–83.90
scales around body 65–76 69–82
(69.7 ± 3.0) (75.77 ± 4.6)
dorsal scales 70–86 87–97
(76.27 ± 3.8) (95.07 ± 3.94)
ventral scales 94–117 100–104
(108.87 ± 6.9) (102.57 ± 1.51)
gular scales 25–35 34–38
(29.57 ± 2.6) (35.57 ± 1.76)
precloacal pores in males 6–9 6–7
(7.87 ± 1.0) (6.57 ± 0.6)
preloacal pores in females 0–6 0–0
(1.007 ± 2.2)
scales in contact with mental scales 4–6 (5.67 ± 1.1) 4–6 (5.87 ± 0.7)
sexual dichromatism evident evident
head color in males blue, light blue, brown or gray brown, gray
body color in males yellow, light blue, brown, gray, light green brown, gray
light blue scales in males numerous few or numerous
colored scales on the sides of the body and tail in males with numerous colored scales, yellow, green and / or blue absent or some greenish
posterior paravertebral spots in males evident blurred or absent
pre scapular and post scapular spots evident variable (from evident to absent)
Material examined Holotype ARGENTINA ; Neuquén , Añelo , Bajo Añelo from south of Route 7, on the way to Aguada Pichana ; 38°14′57.87″ S , 68°56′37.38″ W ; alt. 321 m ; 9 Mar. 2021 ; I. Procheret and P. Chafrat leg.; MPCN-H-469 . Paratypes ( 12 specimens ) ARGENTINA Neuquén 2 ♀♀ ; same collection data as for holotype; 38°14′57.82″ S , 68°56′37.55″ W ; alt. 321 m ; 2 Feb. 2021 ; C. Abdala , P. Chafrat and I. Procheret leg.; MPCN-H-456–457 1 ♀ ; same collection data as for holotype; 37°59′45.32″ S , 69° 7′51.53″ W ; alt. 286 m ; 2 Feb. 2021 ; C. Abdala , P. Chafrat and I. Procheret leg.; MPCN-H-458 1 ♂ ; same collection data as for holotype; 38°20′42.28″ S , 69°4′46.55″ W ; alt. 374 m ; 2 Feb. 2021 ; C. Abdala , P. Chafrat and I. Procheret leg.; MPCN-H-459 1 ♀ ; same collection data as for holotype; 8°14′57.82″ S , 68°56′37.55″ W ; alt. 321 m ; 9 Feb. 2021 ; P. Chafrat and I. Procheret leg.; MPCN-H-468 1 ♀ , 1 ♂ ; same collection data as for holotype; 38°14′57.82″ S , 68°56′37.55″ W ; alt. 319 m ; 9 Feb. 2021 ; P. Chafrat and I. Procheret leg; MPCN-H-470–471 1 ♀ ; same collection data as for holotype; 37°59′45.32″ S , 69° 7′51.53″ W ; alt. 286 m ; 9 Feb. 2021 ; P. Chafrat and I. Procheret leg.; MPCN-H-472 2 ♀♀ ; same collection data as for holotype; 38º25′43.7″ S , 69º09′31.3″ W ; alt. 433 m ; 7 February 2021 ; P. Chafrat , A. Planchart , L. Retamal and D. Ubieta leg.; MPCN-H-473–474 1 ♀ ; same collection data as for holotype; 38°24′33.41″ S , 69° 8′15.91″ W ; alt. 402 m ; 2 Feb. 2021 ; C. Abdala , P. Chafrat and I. Procheret leg.; MPCN-H-476 1 ♂ ; same collection data as for holotype; 38°24′32.79″ S , 69° 8′24.96″ W ; alt. 405 m ; 2 Feb. 2021 ; by C. Abdala , P. Chafrat and I. Procheret leg.; MPCN-H-477 . Description ( holotype ) MEASUREMENTS . Weight 15 gm. Snout vent length 79.93 mm . Head 1.31 mm times longer ( 17.53 mm ) than wide ( 13.31 mm ). Head height 9.81 mm . Neck width 14.82 mm . Interorbital distance 8.12 mm . Eye length (horizontal) 3.07 mm . Ear–eye length 7.11 mm . Auditory meatus height 3.06 mm and width 1.6 mm . Distance from eye to upper lip 2.57 mm and distance between nostrils 2.75 mm . Subocular length 4.83 mm . Torso length 39.04 mm . Tail length 112.35 mm . Body width 26.98 mm . Thigh length 14.02 mm . Tibia length 18.41 mm . Foot length 22.47 mm . Humerus length 11.69 mm . Forearm length 9.98 mm . Hand length 13.24 mm . Pygal region length 5.98 mm . Cloaca width 6.26 mm . MORPHOLOGY . Dorsal surface of head smooth, with sixteen dorsal scales. Rostral wider than height and in contact with six scales. Mental scale same size as rostral scale, trapezoidal in shape, in contact with six scales. No contact between nasal and rostral scales, separated by scale. Two internasal scales higher than wide. Nine scales surrounding nasal that is separated from canthal by two scales. Seven scales between rostral and frontal. Front divided into two scales. Interparietal slightly smaller than parietal scales, in contact with six scales. Preocular separated from lorilabials by one scale. Six superciliaries and 15 superior ciliary scales. Anterior border of ear with two differentiated ear scales; upper border without differentiated scales. Nine temporal scales keeled. Subocular in contact with three lorilabials, without contacting fourth supralabial. Six supraoculars. Ten supralabials. Nine lorilabials. With seven infralabials, second in contact with two ventral scales. Six shield scales. Second shield separated by two scales. Scales around the body 26. With 75 dorsal scales from occiput to hind limb, with rounded posterior end, imbricated and with evident keel, without mucron. Ventral scales 105 (from mental scale to cloaca, following ventral midline of body), larger than dorsals, imbricate and laminar. Gular scales 22, imbricate and keeled. Neck with 51 scales, from posterior border of auditory meatus to shoulder, along longitudinal fold. Seven precloacal pores. Vestigial pores in gulars, pectoral, pygal, and posterior abdominal areas. Antehumeral granular scales, imbricated and well differentiated from rest. Marked atrial and longitudinal fold. Longitudinal fold scales smooth and without keel. Fourth finger with 21 scales and foot with 29 scales. A femoral patch present. Dorsal tail scales without mucron and with keel, ventral smooth. Laminar imbricate and trifid infracarpal. Infratarsals laminar imbricate and trifid. Gular fold developed. No developed postscapular fold. Patch scales rounded, with mucron, without notch. Trifid center palm scales without keel. COLORATION IN LIFE ( Fig. 1 ). Dorsal and temporal region of head, brown with few whitish scales. Supralabial and infralabial light yellow. Dark spot through the subocular, loreolabial and supralabial scales. Sides of neck with thick white line reaching forelimbs. Brown body slightly darker than head. Undefined vertebral region, with yellowish-white circular or irregular spots. Evident paravertebral spots on neck to middle of body that gradually fade until disappearing when reaching tail. Paravertebral black spots with white posterior border. Antehumeral arch defined, black, wide, short. Faint prescapular spot, and large, evident postscapular spot, of deep black color. Sides of body same color as back, with numerous white circular spots, which disappear before reaching groin. Limbs with same color as body, with irregular black and whitish spots. Tail dorsally reddish brown without spots at its proximal end and with faint black vertebral line at its distal end. Ventral is yellowish white; yellow color is more accentuated in gular region and thighs. Gular band incomplete and two black lines extend from gular band to forelimbs. Numerous black scales distributed irregularly across venter, forming melanic spot in pectoral and anterior abdominal area. MORPHOLOGICAL VARIATION . Based on nine specimens ( 4 ♂♂ and 5 ♀♀ ). Dorsal surface of head smooth with 16–18 (mean = 16.22; SD = 0.67) scales between the rostral and occiput. Nasal surrounded by 8–9 (mean = 8.22; SD = 0.6) scales. Supralabials 8–10 (mean = 9.00; SD = 0.50); 8–10 (mean = 8.89; SD = 0.78). Lorilabials arranged in single row. Supraocular 5–7 (mean = 5.75; SD = 0.4). Parietals larger than or equal in size to interparietal, surrounded by 6–8 scales (mean = 6.89; SD = 0.78). Infralabials 6–11 (mean = 7.78; SD = 1.39). Gulars 22–25 (mean = 23.78; SD = 1.39). Temporals 8–11 (mean = 10.1; SD = 0.46) keeled. Auditory meatus higher (mean = 2.92 mm ; SD = 0.33) than wide (mean = 1.63 mm ; SD = 0.20). Developed antehumeral fold. Head longer (mean = 16.34 mm ; SD =0.98) than wide (mean = 11.98 mm ; SD = 1.18) and high (mean = 8.73 mm ; SD = 1.09). Trunk length (mean = 40.19 mm ; SD = 3.18). Maximum snout–vent length 83.98 mm . Arm length (mean = 11.45 mm ; SD = 1.48). Forearm length (mean = 9.60 mm ; SD = 0.95). Hand length (mean = 11.65 mm ; SD = 0.93). Thigh length (mean = 15.02 mm ; SD = 1.71). Number of scales around body from 69–82 (mean = 75.67; SD = 4.61). Dorsal scales between occiput and thighs 90–96 (mean = 92.78; SD = 1.86). Dorsal scales with rounded edges, imbricated, with evident keel. Fourth finger of hand with 20–24 (mean = 21.44; SD = 1.42) digital lamellae. Fourth toe with 27–31 (mean = 28.78; SD = 1.48) digital lamellae. Infracarpal with laminar, imbricate, trifid scales. Infratarsal with laminar, imbricate and smooth scales. Ventral thigh with femoral patch. Ventrals larger than dorsals 98–116 (mean = 108.67; SD = 5.53). Tail longer (mean = 82.68 mm ; SD = 16.48) than snout–vent length. Males with 6–7 (mean = 6.5 mm ; SD = 0.58) precloacal pores, and absent in females. Gular fold developed. With presence of vestigial pores in the gulars, pectoral and abdominal regions. Scales of the patch with rounded edges and mucron, without notch. Fig. 1. Details of Liolaemus kulinko sp. nov. , holotype (MPCN-H-469) (SVL 79.93 mm, tail 112.35 mm). A, C . Dorsal and ventral views of the body. B, E–F . Lateral, ventral and dorsal views of the head. D . Ventral view of precloacal pores. Photographs: P. Chafrat. Fig. 2. A . Preserved male specimens of L. kulinko sp. nov. (top) and L. mapuche Abdala, 2002 (bottom) in dorsal view showing the color pattern. B . Ventral view of L. kulinko sp. nov. (top) and L. mapuche (bottom). C . Living male specimens of L. kulinko sp. nov. (left) and L. mapuche (right) in dorsal view showing the color pattern. D . Preserved male specimens of L. kulinko sp. nov. (left) and L. mapuche (right) showing the cloaca, the arrows indicate the precloacal pores. E . Live male specimens of L. kulinko sp. nov. (top line) and L. mapuche (bottom line) exhibiting the color pattern of the head and neck with the scapular spots. F . Live male specimens of L. kulinko sp. nov. (top line) and L. mapuche (bottom line) showing the varieties in color pattern. G . L. kulinko sp. nov. (left) and L. mapuche (right) showing the front nose. H . L. kulinko sp.nov. (left) and L. mapuche (right) exhibiting details of the color pattern of the body in lateral view. Fig. 3. A–B . Adult male specimen of Liolaemus mapuche Abdala, 2002 from the type locality. C . Adult male specimen of L. cuyanus Cei & Scolaro, 1980 from Matagusano, San Juan. D . Adult male specimen of L. goetschi Müller & Hellmich, 1938 from the type locality. E . Adult male specimen of L. puelche Ávila, Morando, Perez & Sites, 2007 from the type locality. F . Adult male specimen of L. josei Abdala, 2005 from the type locality. Photographs: C.S. Abdala. COLOR VARIATION IN LIFE ( Fig. 4A, C, E, G ; based on nine specimens in life and 18 specimens photographed; with moderate sexual dichromatism). Males with highly variable coloration pattern and more showy colors than females; color of head varies from light brown to gray. Subtle dark brown line running from eye to neck, crossing temporal region. This line absent or attenuated in some large adults. Subocular generally white or gray in color, with dark upper border. Supralabials and lorilabials always lighter in color, generally light gray or white. General body color light brown or light gray, some specimens with bluish color. Vertebral region thin and not marked, except in proximal area of neck. A wide, short, antehumeral arch, black or gray in color, not expanded towards forelimbs. In larger males the gular melanism continues. With pre- and post-scapular spots of variable size and intensity, from almost absent to black or dark brown. Post-scapular generally slightly larger than pre-scapular. Several specimens with spots posterior to post-scapular, and these always smaller, black, or dark brown. Paravertebral spots arranged in non-aligned pairs along body, generally sub-quadrangular, dark brown on front and black on back, with white or light blue back edge. In some specimens spots in shape of line or stripe across body, while in others in the shape of point or small circle. In larger males, paravertebral spots in posterior region of body fainter or absent. A few specimens with almost total absence of paravertebral and lateral spots. Lateral spots diffuse or absent, but with same shape and color as paravertebral spots. Numerous scales and white spots on sides of neck and body. Some males with light blue scales forming thin line in lateral–posterior region of body. Same design as on body continues onto tail, but with line of light blue and / or green scales, iridescent on sides of tail. In several individual this line reaches sides of body. Ventrally, males white, with melanism present in gular, pectoral and abdominal region. Females with more uniform and consistent pattern of coloration than males. Head gray, bordered by gray in degrade. Line from eye to neck generally more notable than in males. Body also varies from gray to brown in color. Vertebral region more delimited in males than females, reddish-brown on back of body and tail. Antehumeral arch more diffuse than in males. Pre- and post-scapular spots present. Generally, largest post-scapular always black or dark gray in color. With sub-quadrangular paravertebral spots that are black or brown in color and white or yellow along posterior border and may be pointed in shape. On paravertebral spots, some females have reddish-brown spot or reddish-brown anterior border. Dorsolateral bands yellow, orange or white color, which can be continuous or fragmented and usually join at base of tail. In larger females dorsolateral bands not distinguishable or very faint. With and without lateral spots. When present they have same shape and color as paravertebral spots. Paravertebral spots unite on tail, where they form single stripe that extends longitudinally. White underneath with some irregularly scattered gray spots and scales. Geographic distribution The new species is restricted to the type locality and surroundings, in the localities of Aguada Pichana and Bajo de Añelo, Neuquén Province , Añelo Department, Argentina ( Fig. 5 ), at elevations between 286–433 m a.s.l. Fig. 4. A, C, E, G . Coloring patterns. Adult males of Liolaemus kulinko sp. nov. in dorsal view. B, D, F, H . Adult males of L. mapuche Abdala, 2002 in dorsal view. Photographs by I. Procheret (A, C) and C.S. Abdala (B, D–H). Phylogenetic analysis ( Figs 6–7 ) Both molecular and morphological analyses indicate that Liolaemus kulinko sp. nov. belongs to the L. boulengeri group. According to molecular data, Liolaemus kulinko is recovered within the subclade of L. goetschi , complex of L. cuyanus , as a sister species of L. mapuche . This relationship is nested within a clade made up of ( L. goetschi ( L. cuyanus ( L. josei + L. puelche )) + ( L. mapuche + L. kulinko sp. nov. )) ( Fig. 4 ). This relationship is also reflected in the table of molecular distances ( Table 2 ). The results of the morphological analysis are partially consistent with the molecular ones. In all the runs carried out, the species under study was recovered as a sister species of L. mapuche ( Fig. 7 ). Nevertheless, the L. cuyanus complex was not always recovered as monophyletic, since the species related to L. donosobarrosi (( L. tiranti + L. calliston ) + ( L. donosobarrosi + L. hugoi )) are recovered basal throughout the clade of L. melanops , very distant from the rest of species of L. cuyanus complex. In the hypothesis presented in this work, Liolaemus kulinko sp. nov. shows 28 autoapomorphies, of which nine correspond to continuous characters and 21 to discrete ones (12 of scaling and nine of coloration). The relationship between L. kulinko and L. mapuche ( Fig. 7 ), was recovered based on five characters, one continuous and four discrete (one of scales and three of coloring). Natural history Liolaemus kulinko sp. nov. , is an endemic species of the Añelo Department in the center of Neuquén Province , and whose type locality is located 30 km from the eponymous city, in the area between provincial route number 7 and Neuquén River. We consider that the river is a biogeographical barrier separating it from its sister species L. mapuche ( Fig. 5 ). The Bajo de Añelo comprises an area of 280 km ² that constitutes an extensive closed basin, displaying a convergent radial network of temporary channels (Pérez et al. 2009) and whose lowest elevation is at 223 m a.s.l. Sediments from the Cretaceous Period predominate in the area and correspond to different geological formations. These are mainly from the Neuquén group and modern sedimentary deposits of various origins (fluvial, alluvial and aeolian). The internal dune system is the environment in which most of the endemic species of the area are found. The vegetation is part of the Monte Phytogeographic Region ( Cabrera 1971 ), which constitutes an arid to semi-arid biome that extends from Salta to Chubut Province ( Abraham et al. 2009 ). The appearance of the vegetation is as islands or patches of shrubs that alternate with areas of bare soil ( Bulacios Arroyo et al. 2021 ). Each patch constitutes a microenvironment that is used in different ways by the species that inhabit it ( Bertiller et al. 2009 ). This ecosystem is characterized by the presence of numerous endemic species of insects, birds and reptiles ( Roig-Juñent et al. 2001 ; Roig et al. 2009 ). The characteristic vegetation includes Larrea divaricata Cav. , L. cuneifolia Cav. , L. nitida Cav. , Cyclolepis genistoides D.Don , Suaeda divaricata Moq. , Atriplex lampa (Moq.) Gillies ex Small , Prosopidastrum globosum Burkart , Neltuma alpataco Phil. (C.E. Hughes & G.P. Lewis 2022) , Hyalis argentea var. latisquama Cabrera , Parkinsonia praecox (Ruiz & Pav.) Hawkins , Schinus johnstonii F.A. Barkley , Erodium cicutarium (L.) L’Hér. ex Aiton, Maihuenia patagonica Phil. (Britton & Rose) , Grindelia chiloensis (Cornel.) Cabrera , Senecio filaginoides var. lobulatus (Hook. & Arn.) Cabrera , Acantholippia seriphioides (A.Gray) Moldenke , Ephedra ochreata Miers. , Monttea aphylla (Miers.) Benth. & Hook.f. ( Burkart 1964 ; Cabrera 1971 ; Roig 1987 ; Arbo 1999 ; Gandullo et al. 2004 , 2016 ; Riveros et al. 2011 ). Liolaemus kulinko sp. nov. is sympatric with L. grosseorum , L. calliston , L. cuyumhue , L. quinterosi Ruiz, Quipildor, Bulacios-Arroyo, Chafrat & Abdala, 2019 , L. darwinii , L. gracilis , Aurivela longicauda Bell, 1843 , Leiosaurus bellii Duméril & Bibron, 1837 , Homonota darwinii Boulenger, 1885 , H. horrida (Burmeister, 1861) , and H. underwoodi Kluge, 1964 . Snakes seen during several visits to the locality were Bothrops ammodytoides Leybold, 1873 , B. diporus Cope, 1862 , Erythrolamprus sagittifer (Jan, 1863) , Philodryas patagoniensis Girard, 1858 , Ph. trilineata (Burmeister, 1861) , Ph. psammophidea Günther, 1872 , Micrurus pyrrhocryptus Wagler, 1824 , and Pseudotomodon trigonatus (Leybold, 1873) , all of which represent potential predators of Liolaemus kulinko . Table 2. Pairwise uncorrected p-distance for cyt- b rRNA between L. kulinko sp. nov. and related species in the L. boulengeri group. The maximum and minimum values from the terminal taxa are included for each species. * = morphological data of number of specimens examined and traits.
species L. cuyanus L. josei L. puelche L. mapuche
Cei & Scolaro, 1980 Abdala, 2005 Avila, Morando, Fulvio-Pérez & Sites, 2007 Abdala, 2002
L. cuyanus
L. josei 9.36–10.17
L. puelche 9.64–10.36 6.24
L. mapuche 7.04–9.20 8.51–9.58 8.65–8.94
L. kulinko sp. nov. 7.48–12.02 7.56–9.60 8.23–9.93 8.51–9.79
Liolaemus kulinko sp. nov. , is a robust, psammophilic species, adult females weigh between 9.5 and 12.5 gm while adult males between 14 and 15 gm showing sexual dimorphism in other measurements. They are oviparous. Two females observed in November were pregnant, while two adult females collected in February had three and four shelled eggs, another female collected in March had one shelled egg, suggesting two clutches per year with one to four eggs per clutch. Fig. 5. Map of the distribution area of Liolaemus kulinko sp. nov. (red star), type locality of the holotype (black arrow with red star) and type locality of Liolaemus mapuche Abdala, 2002 (yellow square). Fig. 6. Molecular phylogenetic tree showing the relationships between the species of the Liolaemus boulengeri group, including L. kulinko sp. nov. Based on the analysis of stomach content and feces, its diet consists mainly of insect species, with a high content of ants of the genera Acromyrmex Mayr, 1865 ( Acromyrmex lobicornis Emery, 1888 ) and Camponotus Mayr, 1861 ( Camponotus sp. ), and, to a lesser extent, Hemiptera, Diptera , Coleoptera (Fam. Buprestidae Leach, 1815 ) , and spiders (indeterminate). Seeds and other plant remains were also found. This species is abundant and easy to observe, between 11 and 16 hrs. It takes refuge mainly under large bushes of Neltuma Raf. ( Neltuma alpataco ) and Schinus L. ( S. johnstonii F.A. Barkley ). It frequents dunes and sandy areas, although it also lives in higher stony areas, on the lower slopes of Añelo. In March 2021 , at twilight time, females were identified in patches of vegetation along with newborn juveniles that may be their progeny; this suggests possible maternal care (Procheret personal observation).