Phymaturus loboi, Troncoso-Palacios & Ferri-Yáñez & Laspiur & Aguilar, 2018

Troncoso-Palacios, Jaime, Ferri-Yáñez, Francisco, Laspiur, Alejandro & Aguilar, César, 2018, An updated phylogeny and morphological study of the Phymaturus vociferator clade (Iguania: Liolaemidae), Zootaxa 4441 (3), pp. 447-466 : 456-460

Phymaturus loboi sp. nov.

( Fig. 5 )

1966 Phymaturus palluma palluma (in part), Donoso-Barros, Rept. Chile: 349.

2005 Phymaturus flagellifer (in part), Pincheira-Donoso & Núñez, Pub. Ocas. Mus. Nac. Hist. Nat. 59: 24.

2005 Phymaturus cf. palluma (in part), Lobo & Quinteros, Pap. Avul. Zool. 45: 155.

2007 Phymaturus cf. palluma (in part), Lobo & Abdala, Cuad. Herpetol. 21: 113.

2012 Phymaturus maulense (in part), Troncoso-Palacios & Lobo, Cuad. Herpetol. 26: 73.

2013 Phymaturus maulense, Esquerré, Núñez & Scolaro, Zootaxa 3619: 439.

2013 Phymaturus maulense (in part), Troncoso-Palacios, Lobo, Etheridge, Acosta & Laspiur, Zootaxa 3652: 561.

2013 Phymaturus cf. palluma, Lobo & Etheridge , South Am. J. Herpetol. 8: 100.

2013 Phymaturus maulense, Garin, Troncoso-Palacios, Ferri-Yáñez & Lobos. Cuad. Herpetol. 27: 67.

2013 Phymaturus sp. 2 Morando, Avila, Pérez, Hawkins, Sites, Mol. Phyl. Evol. 66: 701.

2013 Phymaturus sp. 6 Morando, Avila, Pérez, Hawkins, Sites, Mol. Phyl. Evol. 66: 701.

2016 Phymaturus maulense Lobo, Barrasso, Hibbard & Basso. Zool. J. Linn. Soc. 176: 649.

2016 Phymaturus sp. 6 Lobo, Barrasso, Hibbard & Basso. Zool. J. Linn. Soc. 176: 653.

Holotype. SSUC Re 691. Adult male. Collected at National Route CH 115, 9 km NW of Maule Lagoon (35˚57' S, 70˚34' W), 1900 m asl , Maule Region, Chile by J. Troncoso-Palacios, A. Laspiur and R. Gómez. January 25, 2013 ( Fig. 5a,b ).

Paratypes. SSUC Re 692, male and 693, 694 females ( Fig. 5 ). Same data as the holotype . SSUC Re 410–11 (male and juvenile male). Collected road to Maule Lagoon , Maule Region, Chile, by F. Ferri-Yáñez. November 11, 2011

Other specimens referred. MNHNCL 2353, 2460–61, males. Baños del Campanario, 1500 m asl, San Clemente, Maule Region, Chile. J.C. Torres-Mura coll. February 2-3, 1992.

Etymology. The species is named after our colleague, the herpetologist Fernando Lobo, a prominent specialist in the Phymaturus genus, who has participated in the description of 16 species in this genus ( Lobo and Quinteros 2005, Lobo and Abdala 2007, Lobo and Nenda 2015, Lobo et al. 2010, 2012b, 2012c, 2013, Troncoso-Palacios and Lobo 2012) and the identification of the true type locality and redescription of P. palluma (Lobo and Etheridge 2013) . The suggested English common name for this species is "Lobo’s Rocky Lizard" and in Spanish, "Matuasto de Lobo”.

Diagnosis. Phymaturus loboi belongs to the palluma group ( Etheridge 1995) because it has short, nonimbricate superciliary scales; fragmented subocular scale; and strongly spiny tail scales. Within the palluma group, P. loboi belongs to the P. vociferator clade ( Figs. 1 and 2 ) and differs from the species within this clade in the following characteristics.

Phymaturus loboi differs from P. dorsimaculatus , in having higher number of dorsal scales (161.0±7.5 vs. 135.0±4.0, Tables 4 and 6), and a higher number of scales around the interparietal (9.1±1.2 vs. 6.2±2.0, Tables 4 and 6). Moreover, the males of P. loboi have green spotted shoulders and the area behind the shoulders, with an inconspicuous scapular spot ( Fig 5 ), whereas the males of P. dorsimaculatus have black color on the shoulder and in the area behind the shoulder with a bold scapular spot.

Differs from P. vociferator , in having higher number of scales around the interparietal scale (9.1±1.2 vs 7.2±1.2, Tables 4 and 6). Males of P. loboi have a green spotted shoulder with an inconspicuous scapular spot ( Fig. 5 ) whereas the males of P. vociferator have black color on the shoulders and in the area behind the shoulders, and a marked scapular spot. Additionally, P. loboi does not vocalize.

Differs from P. maulense , in having higher number of ventral scales (184.0±7.6 vs 169.8.0±7.7, Tables 4 and 6). Males of P. loboi have an inconspicuous scapular spot and females have no precloacal pores, whereas the males of P. maulense have a marked scapular spot and all females have precloacal pores.

Differs from Phymaturus damasense , in having higher number of ventral scales (184.0±7.6 vs 166.8±6.5, Tables 4 and 6) and fewer subocular scales (2.0±0.9 vs 3.6±0.5, Tables 4 and 6). Additionally, females of P. loboi never have precloacal pores, while 40% of the females of P. damasense have it.

Description of the holotype. SSUC Re 691. Adult male. Measurements in mm: snout–vent length (SVL): 102.5; trunk width: 41.7; head length: 20.7; head width: 21.1; head height: 10.9; interorbital distance: 8.4; distance between the nares: 3.9; distance between the naris and the eye: 4.9; distance between rostral and interparietal scale: 12.7; axilla-groin distance: 51.7; arm length (from tip of toe of the 3rd toe to elbow): 29.7; right foot length: 24.1 (from tip of toe of the 4th toe to heel); and tail length: 105.8 (with autotomized tip). Ten scales in contact with interparietal scale; 16 scales between interparietal and rostral (excluding both). Parietal eye inconspicuous and whitish. Supraorbital semicircles incomplete in both sides. Superciliary scales, 12/12 (left/right) juxtaposed and flat. Subocular scales, two. Posterior subocular vertically separated from supralabials by 2 row of lorilabials. Preocular scale smaller than canthal. Preocular and supralabials separated by 4 loreal scales. Canthal separated from nasal by 2 scales. Nasal in contact with 8 scales and separated from rostral by 3 scales. Rostral scale undivided. Nine rhomboidal or pentagonal, slightly keeled temporal scales. Three scales slightly projected on the anterior border of auditory meatus. Eleven supralabial scales and 10 infralabial scales. Mental pentagonal, in contact with 6 scales. Fifty six gular scales, counted transversally between mouth´s commissures. Secondary chin shields separated by 2 scales. Well-developed antehumeral and neck fold. Conspicuous gular and antegular folds. No enlarged scales on posterior margin of gular fold or on chest. Dorsal scales rounded, smooth and juxtaposed. Dorsal scales (counted between the occiput and the level of the anterior border of the hindlimbs), 157. Middorsal scales larger than those on flanks. Ventral scales larger than dorsals, quadrangular, smooth, and juxtaposed. Ventral scales (counted from mental scale to the anterior margin of cloacal opening), 175. Scales around of midbody, 216. Precloacal pores, 11. Supernumerary pores, 7 (1 above and 6 below precloacal pores). No enlarged pigal scales. Scales of the dorsal surface of the femoral area smooth, rounded to pentagonal and juxtaposed or sub-imbricated. Scales of the ventral surface of the femoral area smooth, pentagonal or hexagonal, and juxtaposed. Scales of the dorsal surface of forearm rounded, smooth, and juxtaposed or sub-imbricated. Scales of the ventral surface of forearm rounded, smooth, and juxtaposed. Subdigital lamellae of fingers: I: 9, II: 14, III: 19, IV: 21 and V: 13. Subdigital lamellae of the toes: I: 9, II: 15, III: 20, VI: 24 and V: 15. Tarsal scales smooth or with 2/3 keels. Caudal scales arranged in spiny whorls, slightly keeled with projected mucrons.

Color in life of the holotype. Head with black background color, spotted with green in nasal and temporal areas, without spots on the cheeks. Dorsum green, with faint black reticulation, without ocelli. Neck green with black reticulation to the level of the lower meatus (transversally), but is black with very few green spots below the level of the lower meatus (transversally). Scapular spot inconspicuous. Limbs light green, with faint black reticulation. Flanks yellowish green. Base of the tail green, the rest yellowish green (lighter than the body). Throat and chest black, extending ventrally to the insertion of the forelimbs. Abdomen with a “Π” yellowish green area (limited by the chest and flanks) with scattered black spots. Center of the belly light green with no appreciable yellow shading on the cloaca. Ventral surface of the limbs light green with few black spots. Ventral surface of the tail light green. Precloacal pores intense orange.

Variation. Measurements in mm. Adult males (n = 6): SVL, 102.1–102.8 (102.5 ± 0.3); head length, 20.3–21.8 (20.9 ± 0.8); head width, 20.3–21.5 (21.0 ± 0.4); head height, 10.9–13.2 (11.9 ± 0.9); axilla–groin distance, 50.9– 51.7 (51.4 ± 0.4); arm length, 29.7–33.0 (31.2 ± 1.7); foot length, 24.1–27.5 (25.7 ± 1.8); tibia length, 14.8–17.8 (16.3 ± 1.5); distance between the interparietal and the rostral, 11.9–12.7 (12.3 ± 0.4); distance between the nares and the eye, 4.9–5.7 (5.2 ± 0.4); and distance between the nares, 3.2–3.9 (3.6 ± 0.3). Adult females (n = 3): SVL, 93.1–98.6 (96.6 ± 3.0); head length, 19.1–19.8 (19.4 ± 0.3); head width, 17.8–18.5 (18.2 ± 0.4); head height, 9.1– 11.0 (10.1 ± 0.1); axilla–groin distance, 45.4–50.7 (48.5 ± 2.8); arm length, 28.5–30.1 (29.0 ± 0.9); foot length, 23.2–24.3 (23.8 ± 0.6); tibia length, 14.3–15.1 (14.7 ± 0.4); distance between the interparietal and the rostral, 12.3– 12.4 (12.4 ± 0.1); distance between the nares and the eye, 5.0–5.2 (5.0 ± 0.1); and distance between the nares, 3.5– 3.8 (3.6 ± 0.2). Variation in the meristic variables (scale count) is provided in the Table 6. Parietal eye inconspicuous, whitish and opaque. No enlarged scales on posterior margin of gular fold or on chest. In males, 8– 11 (9.7 ± 1.5) precloacal pores, 0–6 supernumerary pores, with no enlarged postcloacal scales. Tarsal scales smooth or bearing 2–3 keels. Males exhibit the same color pattern as holotype, with variation in color shades. One male has the limbs of the same shade of the dorsum and less yellowish on the flanks than other males. Head brown in females, slightly darker than the body. Occipital area with several dark brown spot, surrounding the interparietal (one female), on the supraocular area (“O” shaped spot), the superciliaries, the snout and temporal area (except postocular area). Scattered light brown scales on the head, between the occipital area and the snout, on the postocular area, and between the subocular and the rictal. Dorsum light brown, with a thin dark brown reticulation forming three short stripes on the neck, but not forming dark crossbars on the dorsum. A “ɔ” shaped dark brown spot on the shoulder. Dorsolateral area is darker than the occipital area. The limbs, lateral neck and flanks are light brown with scattered dark brown spots. Tail light brown with diffuse dark brown rings. Throat variegated, and one specimen has dark brown color only to the level of the meatus. The belly and ventral surfaces of the limbs and the tail are whitish with some scattered dark brown spot (especially on the chest). Flanks with oxide color. The juvenile male (SVL = 80.4 mm) has a color pattern similar to the females, but the dorsal reticulation is thicker than in females, the dark brown spot on the shoulder is “O” shaped, the head is noticeable more darker than in females, the throat is black and the flanks are light brown.

Distribution and natural history. In Chile Phymaturus loboi is known from the type locality and adjacent areas in the Region del Maule. This is an abundant lizard that inhabits rocky environments from the road to Maule Lagoon (35˚57' S, 70˚34' W, 1900 m asl, Fig. 6 ) and the Termas del Campanario (35˚55' S, 70˚35' W, 1800 m), at 3 km NW from type locality. In the type locality Phymaturus loboi occurs syntopically with Liolaemus buergeri , L. carlosgarini and L. flavipiceus and the vegetation is dominated by Hypochaeris tenuifolia and Festuca acanthophylla . In Argentina, P. loboi is distributed near Domuyo Volcano (36˚39' S, 70˚32' W, 2200 m) at 68 km SE from type locality.

At the date of capture (January), one female had one embryo and several small oocytes. Another female had several oocytes. The intestines of one individual (SSUC Re 410) contained plant remnants and a large number of nematodes of an unidentified species, probably of the genus Spauligodon or Parapharyngodon (C. Garin pers. comm.).

Figures

FIGURE 1. Bayesian mitochondrial consensus tree (cyt-b and 12S), showing phylogenetic relationships of Phymaturus palluma group and P. payuniae clade. Left values on nodes indicate the posterior probabilities (in percentage). Bootstrap values of the congruent clades found in the ML (middle) and maximum parsimony (right) are also shown on nodes. A = P. vociferator clade, B = P. roigorum-verdugo clade and C = P. mallimaccii clade. Outgroup (Liolaemus monticola) not shown.

FIGURE 2. Bayesian inference of all genes consensus tree (cyt-b, 12S, CMOS and PNN), showing phylogenetic relationships of Phymaturus palluma group and P. payuniae clade. Left values on nodes indicate the posterior probabilities (in percentage). Bootstrap values of the congruent clades found in the ML (middle) and maximum parsimony (right) are also shown on nodes. A = P. vociferator clade, B = P. roigorum-verdugo clade and C = P. mallimaccii clade. Outgroup (Liolaemus monticola) not shown.

FIGURE 5. Phymaturus loboi sp. nov. (A) and (B) Holotype SSUC Re 691 (SVL = 102.6 mm). (C) and (D) Paratype female SSUC Re 694 (SVL = 93.1 mm). (E) Male not collected (picture by J. E. Domínguez). (F) Female not collected (picture by C. Garin).

FIGURE 6. Distribution map for Phymaturus loboi sp. nov. and all species and candidate species of the vociferator clade. Red star: P. loboi (formerly known as P. sp. 2), Maule Lagoon, type locality. Light orange star: P. loboi (formerly known as P. sp. 6), Near Domuyo Volcano. Green square: P. damasense (Termas del Flaco and Planchón). Black square: P. maulense (Lircay). White diamond: P. vociferator (Los Barros). Green diamond: P. dorsimaculatus (Copahue). Black diamond: P. sp. 3 (Near Loncopue). Blue diamond: P. sp. 4 (Near El Cholar). Black pentagon: P. sp. 5 (Near Epulafquen).