Liolaemus antonietae, Troncoso-Palacios & Esquerré & Urra & Díaz & Pastene & Ruiz, 2018

Troncoso-Palacios, Jaime, Esquerré, Damien, Urra, Félix A., Díaz, Hugo A., Pastene, Carlos Castro- & Ruiz, María Soledad, 2018, Microphysogobio luhensis Huang, Chen, Zhao & Shao, 2018, n. sp., Zoological Studies 57 (22), pp. 1-19 : 11-15

Liolaemus antonietae sp. n.

( Figs. 2 , 8 )

urn:lsid:zoobank.org:pub:

2009 Liolaemus monticola chillanensis, Torres-Pérez, Méndez, Benavides, Moreno, Lamborot, Palma & Ortiz. Biol. J. Linn. Soc. , 96: 635.

2015 Liolaemus chillanensis, Escobar-Huerta, Santibáñez & Ortiz. Gayana , 79: 95.

2016 Liolaemus sp. Chillán Troncoso-Palacios, Díaz, Puas, Riveros-Riffo & Elorza. Zookeys, 632:127

2017 Liolaemus chillanensis Medina, Avila, Sites & Morando. J. Zool. Syst. Evol. Res. , 55: 238.

Material examined: Holotype: SSUC Re 697, male ( Fig. 8A - B ). Termas de Chillán, Biobío Region, Chile (3 6° 5 4'S,7 1° 2 4'W, 1 7 6 6 m). Collected by J. Troncoso-Palacios, H. Díaz and F.A. Urra. January 9, 2014.

Paratypes: SSUC Re 695-96, 698-99 ( Fig. 8C - G ), male, female, male, female, respectively. Same data as the holotype. SSUC Re 700, male. Laguna del Huemul, Shangrila, Biobío Region, Chile ( 36°52'S, 71°28'W, 1955 m). Collected by F.A. Urra. December 2014.

Specimens used in morphological and color variation: MZUC 38086 and 38090, males. Termas de Chillán.

Specimens used only in color variation: MZUC 28251, 28254 and 28569, three males; and MZUC 28249 one female. Termas de Chillán.

Diagnosis: Here we provide a diagnosis in regards of all species in the L. elongatus clade plus Liolaemus chillanensis - previously confused with L. antonietae - and L. monticola - which has a similar color pattern. Liolaemus antonietae is a medium size Liolaemus (max. SVL = 77.6 mm), with many midbody scales (86-98; 91.5 ± 3.8), lateral dark band, absence of dorsal melanism or black ventral color. The males have precloacal pores and it is the only member of the L. elongatus clade that can exhibit arboreal behavior.

Liolaemus antonietae differs from L. chillanensis because this species has olive-brown or bluish-brown dorsal color, olive or bluish ventral color and lacks precloacal pores, whereas L. antonietae never has olive or bluish coloration and the males have precloacal pores. Liolaemus antonietae resembles L. monticola in that both species have a brown dorsal color and dark lateral band. However, phylogenetic evidence shows that L. monticola does not belong in the L. elongatus-kriegi complex ( Fig. 4 ). Moreover, L. monticola is smaller (max. SVL = 65.6 mm) and has fewer midbody scales (54-66) than L. antonietae (max. SVL = 77.6 mm; midbody scales = 86-98).

Liolaemus antonietae is closely related to L. antumalguen , but is smaller (max. SVL = 77.6 mm vs. max. SVL = 107.8 mm) and has more midbody scales (86-98 vs. 72-82, Table 3). Moreover, L. antumalguen has black ventral coloration, and several specimens have black transversal dorsal spots or a very melanic dorsal coloration, whereas L. antonietae has whitish or gray ventral coloration and totally lacks black transversal dorsal spots.

Liolaemus antonietae has more midbody scales (86-98 vs 76-90), and is larger (max. SVL = 77.6 mm vs max. SVL = 69.9 mm) than L. scorialis . Dorsal coloration in L. scorialis is noticeably darker than in L. antonietae , with a marked dark occipital band, which is absent in L. antonietae .

Liolaemus antonietae differs from L. carlosgarini because this latter species features a yellowish brown dorsal coloration and a conspicuous dark occipital band, traits absent in L. antonietae . All males of L. antonietae have precloacal pores, whereas only 50% of L. carlosgarini males have precloacal pores.

Liolaemus curis has a yellowish dorsal coloration accompanied by black transversal dorsal spots or an overall melanic dorsal coloration, and less midbody scales (68-76) than L. antonietae ( Table 3).

Liolaemus antonietae is smaller (max. SVL = 77.6 mm vs. max. SVL = 90.7 mm) than L. choique , and although we have no data of L. choique for statistical analysis, the midbody scale count ranges show almost no overlap (86-98 vs. 74-88, Table 1). Also, L. choique has yellowish or black dorsal coloration, whereas L. antonietae has light brown dorsal coloration.

Liolaemus antonietae differs from L. elongatus because this last features a marked occipital and lateral bands or dorsal melanism, whereas only some specimens of L. antonietae have an inconspicuous occipital band and no specimen has dorsal melanism.

The dorsal pattern of Liolaemus antonietae resembles the pattern of L. smaug . However, males of L. antonietae have light brown dorsum, whereas males of L. smaug have a bright golden yellow dorsal color. Females and males of L. antonietae have white dots on the dorsum, a trait only found on males of L. smaug .

Liolaemus antonietae differs from L. janequeoae , because the latter species does not feature any dorsal pattern apart of few black or white dots and is smaller than L. antonietae (max. SVL 66.9 vs 77.6 mm).

Liolaemus antonietae differs from L. leopardinus , because it lacks the leopard-like dorsal spots on dorsum and tail present in L. leopardinus .

Liolaemus antonietae is smaller (max. SVL = 77.6 mm) than L. crandalli (max. SVL = 93.4 mm), but both exhibit a similar color pattern. However, L. crandalli has blackish dorsal head coloration, which is light brown in L. antonietae . According to the phylogenetic analysis of Avila et al. (2015) this species is the sister taxon of L. smaug + L. choique , whereas in our phylogeny L. antonietae is not closely related to L. smaug or L. choique .

Although we have no scale count data of L. burmeisteri to perform a statistical analysis, the midbody (70-81) and ventral scale count ranges (99-110) show no overlap with L. antonietae (86- 98, 118-131, respectively). Additionally, almost all specimens of L. burmeisteri lack a dorsal pattern, whereas L. antonietae always features a dorsal pattern (vertebral line, paravertebral dark spots and white dots).

Description of the holotype: Male. SVL = 61.2 mm. Horizontal diameter of the eye: 2.88 mm. Subocular length: 3.8 mm. Length of the fourth supralabial: 2.5 mm. Head length (from the anterior border of the auditory meatus to the tip of the snout): 14.3 mm. Head height (at the level of ear openings): 6.1 mm. Head width (distance between the two ear openings): 11.8 mm. Neck width: 11.7 mm. Interorbital distance: 6.3 mm. Eareye distance: 4.8 mm. Internasal scales distance: 1.8 mm. Ear width: 1.4 mm. Ear height: 1.8 mm. Axilla-groin distance: 24.4 mm. Body width: 17.2 mm. Forelimb length: 23.2 mm. Hindlimb length: 38.6 mm. Tail length (not autotomized): 99.2 mm, with relation tail length/SVL = 1.6. Rostral scale, wider ( 2.8 mm) than high ( 1.1 mm).

Two postrostrals. Four internasals. Interparietal is pentagonal, with a small central spot marking the position of the parietal eye. The interparietal is smaller than the parietals, and is surrounded by six scales. Nine scales between the interparietal and rostral. Seventeen scales between the occiput and the rostral (Hellmich index; Lobo, 2005). Orbital semicircles are complete and formed by thirteen scales on both sides. Five supraoculars on the right side and four on the left. Six superciliary scales. Frontal area divided into five scales, from back to front: 2, 1 and 2. Preocular separated from the lorilabials by one loreal scale. Two scales between nasal and canthal. Nasal in contact with the rostral, surrounded by six scales (excluding the rostral). One row of lorilabials between the supralabials and the subocular. Six supralabials, the fourth is curved upward without contacting the subocular. Five infralabials scales. Pentagonal mental scale, in contact with four scales. Five pairs of postmental shields, the second is separated by two scales. Temporal slightly keeled, subimbricated. Nine temporal scales between the level of superciliary scales and the rictal level. Four scales on the anterior edge of the ear, slightly projected, which do not cover the auditory meatus. Differentiated auricular scale, which is narrow and elongated. Forty-six gular scales between the auditory meatuses. The lateral neck fold is “Y” shaped. There is a ventrolateral fold running from the axilla to the groin. Midbody scales: 90. Dorsal scales are lanceolated, subimbricated, keeled (without mucrons) and with interstitial granules. Dorsal smaller than the ventrals. Dorsal scales: 74. Ventral scales are rounded, smooth, imbricated and without interstitial granules. Ventral scales: 118. Four precloacal pores. Supra-femoral scales are lanceolated, imbricated and smooth or slightly keeled. Infra-femoral scales are lanceolated or rhomboidal, smooth and imbricated. Supraantebrachials scales are lanceolated, imbricated and keeled. Infra-antebrachials are rounded or rhomboidal, imbricated and smooth. Dorsal scales of the first third of the tail are rhomboidal, imbricated, keeled and mucronated. Ventral scales the first third of the tail are lanceolated, smooth and imbricated. Lamellae of the fingers: I: 10, II: 14, III: 20, IV: 23 and V: 12. Lamellae of the toes: I: 10, II: 18, III: 22, VI: 29 and V: 19.

Color of the holotype in life: The head has a light brown dorsal color, with few dispersed dark brown spots. The head has a similar shade as the dorsum. The subocular is whitish with a vertical black line at the posterior end. The cheeks are whitish. The temporal zone is light brown with a dark brown stripe on the middle. The dorsum is light brown. There is a fragmented vertebral line. There are 12 series of black spots on the paravertebral fields, running from the occiput to the base of the tail. The flanks are light brown with a black lateral band that goes from the upper portion of the shoulder to the groin. There are several white dots dispersed on this black lateral band. There are white dots dispersed between the black lateral band and the paravertebral fields. The area between the belly and the dark band on the flank is yellowish with dispersed white dots. The limbs are brown, with dispersed black dots on the fore limbs and dispersed black and whitish dots on the hind limbs. The tail has a light brown dorsal coloration, with dark rings. Ventrally, the throat, the belly, the limbs and the tail are light grey, with some whitish scales dispersed. Inconspicuous yellowish color on the cloaca and thighs. Precloacal pores are orange.

Variation: In six males (including holotype): SVL: 61.2-77.6 mm. Axilla-groin distance: 24.4- 29.3 mm. Head length: 14.3-18.8 mm. Head width: 11.7-15.3 mm. Head height: 6.1-9.6 mm. Leg length: 38.6-45.7 mm. Arm length: 23.2- 26.3 mm. Foot length: 18.9-22.5 mm. Tail length: 99.2-120.0 mm ( two specimens, autotomized in the others). Tail length/SVL = 1.6. In two females: SVL: 56.6-70.5 mm. Axilla-groin distance: 24.9- 31.8 mm. Head length: 13.4-15.4 mm. Head width: 10.5-12.2 mm. Head height: 6.3-7.3 mm. Leg length: 36.9-38.7 mm. Arm length: 22.0- 24.7 mm. Foot length: 18.8-20.2 mm. Tail length: 99.0-110.0 mm (no autotomized), with relation tail length/SVL = 1.6-1.8.

The scalation variation in Liolaemus antonietae is as follows. Midbody scales: 86-98 (91.5 ± 3.8). Dorsal scales: 71-78 (74.4 ± 2.3). Ventral scales 118-131 (123.6 ± 5.4). Fourth finger lamellae: 21-24 (23.0 ± 1.1). Fourth toe lamellae: 28-30 (28.8 ± 0.8). Supralabial scales: 6-7 (6.2 ± 0.4), the fourth curved upward. Infralabial scales: 4-5 (4.7 ± 0.5). Interparietal scale pentagonal or hexagonal, bordered by 5-8 scales (5.8 ± 1.2). Nasal and rostral always in contact. Precloacal pores in males: 3-5. Precloacal pores are absent in females.

Males and females have a similar color patterns to the holotype, with small variations. Some specimens have no distinguishable vertebral line, while others have an inconspicuous occipital band. Some specimens have diffuse dark rings on the dorsal surface of the tail. In specimens with regenerated tail, there is a vertebral line on the regenerated zone. Some males have no yellowish coloration on the cloaca and thighs and have whitish color on the ventral surface of the throat, belly, limbs and the tail, with inconspicuous gray spots dispersed. Females have marked dark spots on the throat and belly and the white dorsal dots are less abundant than males, even totally absent in one female.

Distribution and natural history: Known from two localities in the Biobío Region, Chile: Termas de Chillán ( type locality, 36°54'S, 71°24'W, 1,766 m, Fig. 9 ) and Laguna del Huemul, Shangrila ( 36°52'S, 71°28'W, 1,955 m), 8 km NW from the type locality. Unlike other species of the L. elongatus clade, which are mainly saxicolous ( Morando et al. 2003; Avila et al. 2015), L. antonietae is mostly arboreal, which was seen basking on fallen trunks and on trees up approximately 4 m high. Few specimens were observed basking on rocks. Furthermore, we observed a very peculiar gregarious behavior, in which 18 specimens were observed coming out of the surrounding vegetation and entering to the same hollow of a stand tree ( Nothofagus sp. ) at sunset, in a time frame no longer than an hour. The vegetation in the type locality is dominated by Chusquea sp. and Nothofagus sp. It is an abundant lizard. At the date of capture (January), one female had three embryos and the other had several small oocytes. No remains were found in the stomach, but remnants of insects and plants along with parasitic nematodes were observed inside the intestine. In both localities, L. antonietae was found in syntopy with snake Tachymenis chilensis and lizards L. chillanensis , L. septentrionalis and L. tenuis , being recently recorded also Phymaturus vociferator ( Urra et al. 2017) .

Etymology: This species is named after Antonieta Labra Lillo, a prominent Chilean herpetologist who has made significant contributions to the eco-physiology and behavior of lizards, especially to the study of the influence of chemical signals on their behavior. She has also edited the book “Herpetología de Chile ”, which had a great impact on the Chilean herpetology. We propose the common name “Antonieta’s lizard” in English and “Lagarto de Antonieta” in Spanish. JTP gratefully thanks AL for years of teaching and collaboration.

Figures

Fig. 2. Some specimens previously misidentified as L. chillanensis in Torres-Pérez et al. (2009): (A-B) MZUC 28251 and (C-D) MZUC 28249. Here identified as L. sp. Chillán.

Fig. 8. Liolaemus antonietae sp. nov., all from type locality. (A-B) Holotype, male (SSUC Re 697), (C-E) Paratypes males (SSUC Re 698 and 695) and (F-G) Paratype female (SSUC Re 699). © 2018 Academia Sinica, Taiwan

Fig. 4. Phylogenetic relationships using Bayesian inference (50% consensus tree) based on Cyt-b (HKY + G). In color: Liolaemus sp. Chillán (red, our data from Shangrila, Termas de Chillán and GenBank data from Termas de Chillán), L. chillanensis (blue, our data from Termas de Chillán and data from GenBank) and L. elongatus from Chubut, type locality (green). Red circle on the node denotes posterior probabilities ˃ 0.95. Scale shows the number of changes per site. Numbers between parentheses indicate the number of sequences of the collapsed nodes.

Fig. 9. Distributional map for Liolaemus antonietae sp. nov. along with geographically proximate species of the L. elongatus clade. Stars: Liolaemus antonietae sp. nov (red = Termas de Chillán, type locality; white = Shangrila). Green pentagon: L. smaug (1 = near Las Leñas, 2 = between Las Loicas and Peteroa Volcano, 3 = near Las Loicas). Lilac cross: L. carlosgarini (1 = Maule Lagoon, 2 = Lircay). Black diamond: L. choique (Paso el Choique). Gray square: L. antumalguen (1= Domuyo volcano, 2= Tromen Volcano). Pink haxagon: L. burmeisteri (Caepe Malal). White asterisk: L. crandalli (Auca Mahuida Volcano). Green triangle: L. scorialis (1 = Laja Lagoon, 2 = La Mula Lagoon). Orange circle: L. elongatus (formerly L. lonquimayensis, Lonquimay Volcano). Brown circle: L. elongatus used for morphology and DNA analyses (Llaima volcano). Yellow circles: L. elongatus used for DNA analyses (1 = Pampa de Lonco Luan, 2 = Primeros Pinos, 3 = Portal La Atravesada, 4 = Laguna Blanca, 5 = near Ingeniero Jacobacci, 6 = San Carlos de Bariloche, 7 = Ojo de Agua, 8 = El Maiten, 9 = Esquel, 10 = Tecka, 11 = Gobernador Costa and 12 = Los Manantiales). Blue circles: L. shitan (1 = Estancia Piedras Blancas, type locality, 2 = near San Antonio del Cuy). Pink inverted triangle: L. janequeoae (Tolhuaca). Blue ellipse: L. sp. 6 (Copahue Volcano).