Research Article |
Corresponding author: Jaime Troncoso-Palacios ( jtroncosopalacios@gmail.com ) Academic editor: Aaron Bauer
© 2015 Jaime Troncoso-Palacios, Hugo A. Díaz, Damien Esquerré, Felix A. Urra.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Troncoso-Palacios J, Díaz HA, Esquerré D, Urra FA (2015) Two new species of the Liolaemus elongatus-kriegi complex (Iguania, Liolaemidae) from Andean highlands of southern Chile. ZooKeys 500: 83-109. https://doi.org/10.3897/zookeys.500.8725
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The elongatus-kriegi complex is one of the most diverse clades of the Liolaemus (sensu stricto) subgenus of lizards. There are currently 29 species recognized in this group distributed between Chile and Argentina. Based on molecular evidence, there seem to be five main clades nested within this complex: the elongatus, leopardinus, kriegi, petrophilus and punmahuida clades. Liolaemus buergeri and L. kriegi, both of the kriegi clade, were believed to inhabit the surroundings of the Laja Lagoon, in the Biobío Region of Chile. Moreover, this Chilean population of L. kriegi was recently recognized as an undescribed taxon called “Liolaemus sp. A” based on molecular phylogenetics. In this work, we studied these two populations of the Laja Lagoon and provided the morphological diagnosis to describe them as two new species: L. scorialis sp. n. and L. zabalai sp. n., previously considered L. buergeri and “L. kriegi/Liolaemus sp. A” respectively. Additionally, we identified another population of L. scorialis in the vicinity of La Mula Lagoon in the Araucanía Region of Chile. Liolaemus scorialis differs from almost all of the species of the elongatus-kriegi complex by its considerably smaller size. Nevertheless, without molecular data we cannot assign it to any particular subclade. Liolaemus zabalai belongs to the kriegi clade based on published molecular phylogenies. Finally, we provide some natural history data on both species and we document for the first time the presence of L. neuquensis in Chile from a museum specimen from La Mula Lagoon.
Liolaemus buergeri, Liolaemus kriegi, new species, lizard, Laja Lagoon, Biobío
Liolaemus is a diverse genus of South American lizards, with currently 245 species (
The elongatus-kriegi complex
Species of the elongatus-kriegi complex grouped by clades, based on mitochondrial molecular phylogenies. (1) Species included by
elongatus clade | kriegi clade | leopardinus clade | petrophilus clade | punmahuida clade |
---|---|---|---|---|
L. antumalguen (3) | L. buergeri (1) | L. frassinettii (5) | L. austromendocinus (1) | L. flavipiceus (3) |
L. burmeisteri (4) | L. kriegi (1) | L. leopardinus (5) | L. capillitas (1) | L. punmahuida (3) |
L. elongatus (1) | L. ramonensis (5) | L. dicktracyi (2) | ||
L. smaug (4) | L. ubaghsi (5) | L. gununakuna (2) | ||
L. thermarum (3) | L. valdesianus (5) | L. parvus (3) | ||
L. petrophilus (1) | ||||
L. talampaya (2) | ||||
L. tulkas (3) | ||||
L. umbrifer (2) |
Species of the elongatus-kriegi complex by groups, based on morphological, skeletal and lifestyle traits phylogeny according to (1)
capillitas subgroup | elongatus group | kriegi group | leopardinus group |
---|---|---|---|
L. capillitas (1) | L. austromendocinus (2) | L. buergeri (1) | L. frassinettii (2) |
L. dicktracyi (1) | L. carlosgarini (3) | L. cristiani (1) | L. leopardinus (1) |
L. heliodermis (1) | L. elongatus (1) | L. kriegi (1) | L. ramonensis (1) |
L. talampaya (2) | L. flavipiceus (2) | L. valdesianus (1) | |
L. tulkas (2) | L. gununakuna (2) | ||
L. umbrifer (1) | L. parvus (2) | ||
L. petrophilus (2) | |||
L. punmahuida (2) | |||
L. thermarum (2) | |||
L. tregenzai (2) |
Currently, the elongatus-kriegi complex (
Liolaemus buergeri, of the kriegi clade, was described from El Planchón Volcano, Maule Region, Chile (
Liolaemus kriegi, also of the kriegi clade, was described from Estancia El Cóndor, Río Negro Province, Argentina (
Here, we studied the taxonomic status of the southernmost currently-recognized Chilean population of “Liolaemus buergeri”, from the vicinity of the Laja Lagoon, Biobío Region; and of “L. kriegi/Liolaemus sp. A” from the same locality. This population of “L. buergeri” is described as a new species which differs greatly from L. buergeri and almost all species of the elongatus-kriegi complex by its small snout-vent length (less than 70.0 mm). Additionally, specimens of this new species are recorded from La Mula Lagoon, Araucanía Region, Chile. For “L. kriegi/Liolaemus sp. A”, we provide a full description and diagnosis of this new species belonging to the kriegi clade.
We examined specimens of almost all Chilean species currently considered as belonging to the Liolaemus elongatus-kriegi complex. The morphological characters were examined according to
Liolaemus buergeri (in part?), Pincheira-Donoso, 2001. Not. Mens. Mus. Nac. Hist. Nat., Chile, 346: 8.
Liolaemus buergeri (in part?), Pincheira-Donoso & Núñez, 2005. Pub. Oc. Mus. Nac. Hist. Nat., Chile, 59: 285.
SSUC Re 617 (Fig.
SSUC Re 615–16 two males and 612–614 three females (Figs
The species name refers to the habitat, which is composed of accumulations of igneous rocks from the Antuco Volcano, called “scoria” from the Greek “skoria”. We propose the common name “Slag Lizard” in English and “Lagarto del escorial” in Spanish.
Liolaemus scorialis belongs to the elongatus-kriegi complex, but its specific assignation to a particular subclade is currently unknown since we have no molecular data for this new species, and molecular and morphological phylogenies for the elongatus-kriegi complex disagree in the arrangement of this complex subgroups (see discussion).
Below a wide diagnosis is provided on aspect of all species of the complex. Liolaemus scorialis differs from almost all species of the elongatus-kriegi complex by its size (maximum SVL = 69.9 mm), smaller than L. antumalguen (Table
Chilean species of the elongatus-kriegi complex that live near the distribution of Liolaemus scorialis sp. n. and L. zabalai sp. n. A L. buergeri from El Planchón (type locality, photos by J. Troncoso-Palacios) B L. buergeri from Altos de Lircay (photos by R. Díaz) C L. carlosgarini from the road to the Maule Lagoon (type locality, photos by J. Troncoso-Palacios) D L. flavipiceus from the Maule Lagoon (photos by C. Garín).
Scalation and morphological characteristics for the species of the Liolaemus elongatus-kriegi complex occurring near L. scorialis sp. n. and L. zabalai sp. n. distribution. Juvenile specimens examined are excluded. Source of data for not examined species are: L. antumalguen (
L. antumalguen | L. buergeri (M = 5, F = 9) | L. burmeisteri | L. carlosgarini (M = 6, F = 11) | L. choique | L. flavipiceus (M = 5, F = 10) | L. punmahuida | L. scorialis sp. n. (M = 7, F = 3) | L. tregenzai | L. zabalai sp. n. (M = 3, F = 5) | |
---|---|---|---|---|---|---|---|---|---|---|
Maximum SVL (mm) | 107.8 | 96.2 | 85.2 | 68.8 | 90.7 | 95.8 | 96.0 | 69.9 | 90.2 | 92.0 |
Midbody scales | 72–82 | 80–100 | 70–81 | 80–95 | 74–88 | 68–77 | 67–81 | 76–90 | 71–85 | 90–104 |
Dorsal scales | 70–78 | 78–91 | 76–85 | 68–82 | 65–81 | 60–71 | 70–78 | 74–81 | - | 86–96 |
Ventral scales | 105–118 | 111–125 | 99–110 | 112–124 | 118–135 | 93–105 | - | 115–131 | - | 116–122 |
Sexual dichromatism | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Slight | Present | Slight |
Cloacal region color (males) | Yellowish in some specimens but usually black | Yellowish | Yellowish | Yellowish | Yellowish | Reddish or yellowish in some specimens but usually black | Reddish or yellowish | Yellowish | - | Yellowish |
Tail pattern | Absent | Vertebral line with diffuse rings in the tail base | Weak rings | Rigns (marked or weak) | Absent | Absent or weak rings | Absent | Rings | Rings | Rings |
Precloacal pores on males | 3–4 | 3–4 | 0–5 | 0–3 | 3–4 | 0 | 0 | 3–4 | 0 | 3–4 (3–5*) |
Liolaemus scorialis has probably been previously confused with L. buergeri (see discussion), but in addition to the size difference, L. scorialis differs from L. buergeri because the latter has a vertebral stripe on the tail, whereas the tail is ringed in L. scorialis. Moreover, L. buergeri has more midbody scales (x = 89.4 ± 5.5, n = 14) than L. scorialis (x = 82.0 ± 4.7, n = 10) (Mann–Whitney U = 20.5, P < 0.01, DF = 21) and more dorsal scales (x = 84.1 ± 4.4) than L. scorialis (x = 76.5 ± 4.3) (Mann–Whitney U = 15.0, P < 0.01, DF = 21); but L. buergeri has fewer ventral scales (x = 118.7 ± 4.7) than L. scorialis (x = 124.0 ± 6.0) (Mann–Whitney U = 36.0, P = 0.05, DF = 21).
Liolaemus scorialis is syntopic with “L. kriegi/Liolaemus sp. A”, but in addition to the size difference, the latter has more midbody scales (x = 94.3 ± 4.8, n = 8) than it (Mann–Whitney U = 1.5, P < 0.01, DF = 16). Moreover, the dorsal scale count range of L. scorialis does not overlap with the range of “L. kriegi/Liolaemus sp. A” (Table
Liolaemus scorialis differs from similar size species of the elongatus-kriegi complex as follows. Liolaemus scorialis differs from L. cristiani because the males of the latter lack precloacal pores and have reddish ventral coloration, whereas males of L. scorialis have 3–4 precloacal pores and no reddish ventral coloration.
Liolaemus scorialis differs from L. heliodermis, because the males of the latter have a black head and sulfur-yellow dorsum (
Liolaemus scorialis differs from L. parvus, because the latter has 60–77 midbody scales and 96–113 ventral scales (
Liolaemus scorialis differs from L. smaug, because the latter has marked sexual dichromatism with white spots dispersed on the dorsum of males and absent in females (
Liolaemus scorialis differs from L. tulkas, because the males of the latter have 0–1 precloacal pores (
Liolaemus scorialis differs from L. carlosgarini (Fig.
Adult male. SVL 62.3 mm. Tail length 101.5 mm (not autotomized). Axilla-groin length 26.3 mm. Head length (from the posterior border of the auditory meatus to the tip of the snout) 16.4 mm. Head width (distance between the two ear openings) 11.4 mm. Head height (at the level of ear openings) 6.9 mm. Forelimb length 21.1 mm. Hindlimb length 39.7 mm. Foot length 18.9 mm. Rostral scale wider (2.5 mm) than high (1.0 mm). Two postrostrals. Four internasals. Hexagonal interparietal scale, with a central, small, and whitish spot marking the position of the parietal eye. Interparietal smaller than parietals, surrounded by six scales; nine scales between the interparietal and rostral (both excluded); 15 scales between occiput and rostral; orbital semicircle complete on the right side, formed by 13 scales, incomplete on the left side; 6-5 supraoculars (left-right); six superciliary scales. Frontal area is divided into six scales (two posterior, one in the center and three anterior); 2 scales between nasal and canthal; preocular separated from the lorilabials by one loreal scale; nasal in contact with the rostral, surrounded by seven scales. There is one row of lorilabials between the supralabials and the subocular. Seven supralabials, the fifth is curved upward without contacting the subocular. Four infralabial scales. Mental scale pentagonal, in contact with four scales; four pairs of postmental shields, the second is separated by two scales. Temporal scales are subimbricated and slightly keeled. There are ten temporal scales between the level of superciliary scales and the rictal level. Three projected scales on the anterior edge of the ear, which are small and do not cover the auditory meatus; auricular scale is wide and is restricted to the upper third of the meatus. Forty gulars between the auditory meatuses. Well developed “Y” shaped lateral neck fold and dorsolateral fold slightly developed. Antehumeral fold present. Midbody scales 88. Dorsal scales of the vertebral zone lanceolate, imbricate, keeled and without mucrons. Dorsal scales of the paravertebral fields more rounded, subimbricate, with more poorly developed keel, without mucrons and with interstitial granules between them. Dorsal scales of the vertebral zone are larger than the ventral scales. Dorsal scales of the paravertebral fields are similar in size to the ventral scales. Dorsal scales 81. Ventral scales are rhomboidal to rounded, smooth, imbricate, and without interstitial granules. Ventral scales 131. There are four precloacal pores. The suprafemoral scales are rhomboidal to rounded, imbricate, and smooth or slightly keeled. Infrafemoral scales are rounded, smooth, and imbricate. Supra-antebrachials scales are rhomboidal to rounded, imbricate, and slightly keeled or smooth. Infra-antebrachials are rounded to rhomboidal, subimbricate with few interstitial granules, and smooth. The dorsal scales of the tail are rhomboidal, imbricate, keeled and some with mucrons. The ventral scales of the tail vary from rhomboidal to triangular, and are imbricate and smooth. Lamellae of the fingers: I: 10, II: 17, III: 21, IV: 23 and V: 13. Lamellae of the toes: I: 13, II: 18, III: 22, VI: 29 and V: 20.
Light brown head, with dark brown lines: a “Ω” shaped line between nasal scales and supraocular area, two short stripes on the posterior supraocular areas, an incomplete “O” shaped dark brown line surrounding the interparietal scale, six dark brown short lines on the occipital area. The temporal area is brown with two dark brown horizontal stripes; the ocular area and the cheeks are light gray. Subocular area is gray with two dark brown vertical lines on the middle and posterior edge. Background color of the dorsum is brown. A wide occipital band on the dorsum, formed by twelve transverse dark brown bars; some white scales on the posterior border of these bars. Dark brown lateral band with few yellowish scales dispersed into it, running from the shoulder to the groin; some white scales between the occipital and lateral bands; below the lateral band the flanks are yellowish. Limbs are brown with dark brown spots and some white scales dispersed. Tail is brown with some white scales dispersed and dark brown rings. Posterior third of the tail is immaculate brown. Ventrally, the throat, belly, limbs and tail are immaculate gray. Rear portion of belly and thighs are yellowish. Precloacal pores orange.
There is no sexual dimorphism in size. In seven males: SVL: 57.4–69.9 mm. Axilla-groin distance: 21.4–28.7 mm. Head length: 15.1–17.2 mm. Head width: 11.2–13.0 mm. Head height: 6.4–8.9 mm. Foot length: 19.7–21.1 mm. Leg length: 37.1–46.2 mm. Arm length: 20.3–26.0 mm. Tail length: 101.6–111.3 mm (n = 2; autotomized in the rest). In three females: SVL: 57.3–65.6 mm. Axilla-groin distance: 25.6–32.8 mm. Head length: 15.3–15.8 mm. Head width: 11.1–12.1 mm. Head height: 6.2–6.7 mm. Foot length: 18.7–20.0 mm. Leg length: 37.2–39.0 mm. Arm length: 21.8–22.3 mm. Tail length 88.8–103.1 mm (n = 2; autotomized in the rest).
The variation of the scalation in Liolaemus scorialis is as follows. Midbody scales: 76–90 (x = 82.0 ± 4.7). Dorsal scales: 74–81 (x = 76.5 ± 4.3). Ventral scales 115–131 (x = 124.0 ± 6.0). Fourth finger lamellae: 21–24 (x = 22.7 ± 1.1). Fourth toe lamellae: 28–31 (x = 29.2 ± 1.4). Supralabial scales: 6–7 (x = 6.2 ± 0.4). Infralabial scales: 4–5 (x = 4.7 ± 0.5). Precloacal pores in males: 3–4. Interparietal scale pentagonal or hexagonal, bordered by 5–9 scales (x = 6.7 ± 1.2).
There is a slight sexual dichromatism, females have no yellowish color on the rear portion of belly and thighs. Males have the same color and pattern described for the holotype with variations only in shade. Females have the same color and pattern described for the holotype, but the background color of the dorsum can be brown or gray. One female lacks a wide occipital band because the transverse dark brown bars are not fused and it has an inconspicuous vertebral stripe. Also, in this female there are no lateral bands, since it has unfused vertical bars on the flanks. The tail has dark brown rings in both sexes. Males have orange precloacal pores. The coloration and pattern of the juveniles are unknown.
The northern known distribution limit of the new species is the type locality, near the Laja Lagoon, 1450 m, Biobío Region, Chile (37°21'S – 71°23'W; Fig.
Distributional map for Liolaemus scorialis sp. n., L. zabalai sp. n. and the species of the elongatus-kriegi complex that inhabit in proximity of its. Asterisk: L. scorialis (red = near Laja Lagoon, type locality; orange = La Mula Lagoon). Star: L. zabalai sp. n. (light green = road to Los Barros, type locality; blue = distribution in Argentina). Purple circle: L. carlosgarini. Yellow triangle: L. flavipiceus. Green pentagon: L. buergeri. Gray octagon: L. choique. Brown hexagon: L. antumalguen. Black cross: L. punmahuida. Light pink diamond: L. burmeisteri. Pink square: L. tregenzai.
Its southern limit of distribution is in La Mula Lagoon (La Araucanía Region, Chile), 48 km South from the Antuco Volcano (37°53'S – 71°22'W), 1600 m. We have no data for vegetation or environment in La Mula Lagoon. In this location, according to the Herpetological Catalog of the Museo de Historia Natural of Concepción (unpublished), L. scorialis occurs in syntopy with L. pictus (Duméril & Bibron, 1837). However, this report probably actually refers to L. septentrionalis Pincheira-Donoso & Núñez, 2005 (fide
he intestinal and stomach contents were examined: plant and insect remains were found in the intestine, along with a large number of nematodes of an unidentified species. No remains were found in the stomach. At the time of capture (January) two females had three embryos each and one female had several small oocytes.
Liolaemus kriegi, Donoso-Barros, 1974. Bol. Soc. Biol. Concepción, 47: 287.
Liolaemus kriegi (in part), Cei, 1986. Mus. Reg. Scien. Nat. Torino, 4: 230.
Liolaemus sp?, Torres-Pérez, 1997. Not. Biol., 5(4): 146.
Liolaemus kriegi (in part), Pincheira-Donoso, 2001. Not. Mens. Mus. Nac. Hist. Nat., Chile, 346: 11.
Liolaemus sp. A, Morando et al., 2003. Syst. Biol., 52: 179.
Liolaemus kriegi (in part), Pincheira-Donoso & Núñez, 2005. Pub. Oc. Mus. Nac. Hist. Nat., Chile, 59: 289.
Liolaemus kriegi (in part), Mella, 2005. Guía Camp. Rep. Chil. Zon. Cent., p. 64.
Liolaemus sp. A, Medina et al., 2013. Cuad. Herp. 27(1): 27.
Liolaemus sp. A, Medina et al., 2014. Biol. J. Linnean Soc. 113: 256.
SSUC Re 602 (Fig.
SSUC Re 598. Adult male. SSUC Re 597, 599, 600–01. Four adult females. The same data as the holotype (Figs
This species is named after Patricio Zabala, collection manager of the “Colección de Flora y Fauna Patricio Sánchez Reyes, Pontificia Universidad Católica de Chile” (SSUC). We dedicate this species to him because of his support of herpetological research in Chile, allowing us to review and deposit material in SSUC, and especially for his friendship.
Liolaemus zabalai belongs to the kriegi clade of the elongatus-kriegi complex and is closely related to some undescribed species: Liolaemus sp. C and Liolaemus sp. D; being more distant from the currently described species L. buergeri, L. kriegi and L. tregenzai (Fig.
With respect to the species species of the kriegi clade, Liolaemus zabalai differs from L. tregenzai because the latter has 71–85 midbody scales and the males have no precloacal pores (
Liolaemus zabalai differs from L. kriegi in that the latter reaches 101.1 mm SVL, has reddish cloacal coloration in both sexes and has an unringed tail (
Liolaemus zabalai differs from L. buergeri in that the latter has fewer dorsal scales (78–91; x = 84.1 ± 4.4, n = 14) than L. zabalai (86–96; x = 89.4 ± 3.2, n = 8) (Mann–Whitney U = 19.5; P = 0.01, DF = 20). Liolaemus zabalai has more loreal scales between the nasal and the subocular (4–6; x = 4.3 ± 0.6, n = 8) than L. buergeri (3–4; x = 3.3 ± 0.5, n = 14) (Mann–Whitney U = 11.0; P < 0.01, DF = 20). Also, L. buergeri has a vertebral stripe on the tail, whereas L. zabalai has a ringed original tail. The limbs in L. zabalai are black with dispersed light brown spots, whereas L. buergeri has brown limbs with dispersed black spots (Fig.
Compared to the other species of the elongatus-kriegi complex that occur near the known distribution of Liolaemus zabalai, the new species may be diagnosed as follows. Males of L. zabalai have precloacal pores, whereas males of L. flavipiceus and L. punmahuida lack them (Table
Adult male. SVL: 90.3 mm. Tail length: 92.3 mm (autotomized). Axilla-groin length 39.7 mm. Head length (from the posterior border of the auditory meatus to the tip of the snout) 22.2 mm. Head width (distance between the two ear openings) 16.5 mm. Head height (at the level of ear openings) 11.7 mm. Forelimb length 28.5 mm. Hindlimb length 47.1 mm. Foot length 23.4 mm. Rostral scale wider (4.5 mm) than high (2.2 mm). Two postrostrals. Four internasals. Heptagonal interparietal scale, with a central, small, and whitish central spot marking the position of the parietal eye. Interparietal smaller than right parietal, but bigger than left parietal, surrounded by eight scales: nine scales between the interparietal and the rostral; 14 scales between occiput and rostral; orbital semicircle complete on both sides (formed by 13 scales); 5 supraoculars on both sides; seven superciliary scales. Frontal area is divided into six scales (three posterior, one anterior-left, two anterior-rigth); 2 scales between nasal and canthal; preocular separated from the lorilabials by one loreal scale; nasal in contact with the rostral, surrounded by six scales. There is one row of lorilabials between the supralabials and the subocular. Seven supralabials, the fourth is curved upward without contacting the subocular. Five infralabial scales. The mental scale is pentagonal and is in contact with four scales. Four pairs of postmental shields, the second is separated by two scales. Temporal scales are subimbricated and smooth or slightly keeled. Nine temporal scales between the level of superciliary scales and the rictal level. Two projected scales on the anterior edge of the ear, which are small and do not cover the auditory meatus. There is no differentiated auricular scale. Forty-two gulars between auditory meatus. Well developed “Y” shaped lateral neck fold with antehumeral and posthumeral folds developed. Dorsolateral fold slightly developed. Midbody scales 90. Dorsal scales on the vertebral zone are lanceolate to rounded, subimbricate, keeled and without mucrons. Dorsal scales on the paravertebral fields are more rounded, subimbricate, smooth or with less developed keels, without mucrons and there are interstitial granules between them. Dorsal scales are smaller than the ventral scales. Dorsal scales 86. Ventral scales are rhomboidal, smooth, subimbricate, and with few interstitial granules. Ventral scales 122. There are three precloacal pores. The suprafemoral scales are rhomboidal, imbricate, and smooth or keeled. Infrafemoral scales are lanceolate to rhomboidal, smooth, and subimbricate and with few interstitial granules. Supra-antebrachials scales are rhomboidal to rounded, subimbricate, and keeled or smooth. Infra-antebrachials are rounded to rhomboidal, subimbricate, and smooth. The dorsal scales of the tail are lanceolate to rectangular, subimbricate, keeled or smooth and with few interstitial granules. The ventral scales of the tail vary from lanceolate to triangular, and are subimbricate and smooth. Lamellae of the fingers: I: 11, II: 16, III: 20, IV: 22 and V: 15. Lamellae of the toes: I: 12, II: 16, III: 21, VI: 27 and V: 18.
Black head, with some light brown spots on the supraocular and snout areas. The scales located behind the orbital semicircles are light brown; but the interparietal scale, parietal scales and the scales in contact with the parietal scales are black. Superciliary scales are light brown with black spots. Temporal scales are light brown; cheeks light gray with some black spots. Subocular is gray with a black vertical line on the middle. Background color of the dorsum is light brown. Wide occipital band on the dorsum, formed by twelve transverse black bars. Very few whitish scales dispersed on the dorsum. Black lateral band bearing a few dispersed whitish scales, running from the tip of snout to the groin. Flanks below lateral band are light brown. Limbs black with dispersed light brown spots. Tail light brown with inconspicuous vertebral stripe in the regenerated zone; occipital black band ends in the first fifth of the tail, remainder with some dispersed black spots and a black vertebral stripe. Throat, belly and ventral surfaces of limbs whitish with dispersed inconspicuous dark dots. Rear portion of the belly and the thighs are yellowish. Ventrally, tail is whitish with a dark gray ventral stripe and diffuse dark gray rings from the cloaca to the midpoint of the tail. Precloacal pores orange.
In three males: SVL: 72.6–90.3 mm. Axilla-groin distance: 32.7–38.6 mm. Head length: 17.6–22.2 mm. Head width: 14.2–16.5 mm. Head height: 9.2–11.7 mm. Foot length: 21.5–23.0 mm. Leg length: 42.1–47.2 mm. Arm length: 24.6–28.5 mm. Tail length: 102.0 mm in one specimen (autotomized in the rest). In three females: SVL: 71.8–90.2 mm. Axilla-groin distance: 32.9–42.7 mm. Head length: 17.9–19.5 mm. Head width: 13.9–16.6 mm. Head height: 9.4–11.1 mm. Foot length: 20.6–24.2 mm. Leg length: 41.5–48.8 mm. Arm length: 24.8–29.4 mm. Tail length: 105–115 mm (in two specimens without autotomized tails).
The variation of the scalation in Liolaemus zabalai is as follows. Midbody scales: 90–104 (x = 94.3 ± 4.8). Dorsal scales: 86–96 (x = 89.4 ± 3.2). Ventral scales 116–122 (x = 119.5 ± 2.1). Fourth finger lamellae: 19–22 (x = 20.9 ± 1.0). Fourth toe lamellae: 26–27 (x = 26.8 ± 0.5). Supralabial scales: 6–7 (x = 6.6, ± 0.5). Infralabial scales: 4–5 (x = 4.6 ± 0.5). Interparietal scale pentagonal, hexagonal or heptagonal, bordered by 5–8 scales (x = 7.3 ± 1.1). Precloacal pores in males: 3–4.
There is slight sexual dichromatism; males are slightly darker than females. In general, all specimens have the pattern and color described for the holotype. One female has rusty-colored scales dispersed on the flanks, paravertebral fields and groin. In all specimens, the ventral surface of the throat, belly and limbs are whitish with dark marked or inconspicuous dots dispersed; there is a fragmented midventral stripe on the belly of two specimens. Males and females have a yellowish coloration in the posterior portion of the belly and the thighs (faint in some females). The tail has black rings, marked or diffuse, with a fragmented vertebral stripe in all specimens with complete original tails. Males have orange precloacal pores. The coloration and pattern of the juveniles are unknown.
To our knowledge, in Chile this species is only found in the surroundings of the Laja Lagoon. The type locality is near Los Barros, Laja Lagoon, Biobío Region, Chile (37°31'S – 71°15'W, 1460 m, Fig.
Liolaemus zabalai is also found in Argentina (where it has been called “Liolaemus sp. A”) at several localities in Neuquén Province (
An analysis of the intestinal contents performed on one specimen, showed that this species is omnivorous, but feeds mainly on plants. At the time of capture (January) the females had no embryos, but three had several small oocytes.
In this work, the taxonomic status of two Chilean populations of the Liolaemus elongatus-kriegi complex from the Laja Lagoon have been clarified, here newly described as L. zabalai (previously confused with L. kriegi and also designed as Liolaemus sp. A) and L. scorialis.
Assigning Liolaemus scorialis to any of the groups (
In the case of Liolaemus zabalai of the kriegi clade, the uncorrected pairwise differences between it and other species of the kriegi clade are 2.94–3.79%, almost at the limit of the value (3%) proposed for identify candidate species in Liolaemus (
In this study, Liolaemus ceii is considered a junior synonym of L. kriegi. This synonymy was recommended by
Liolaemus chillanensis was included in the elongatus clade by
In summary, this work describes two new species of the elongatus-kriegi complex lizards from the vicinity of the Laja Lagoon, in southern Chile, one probably confused with L. buergeri: L. scorialis and the other with a history of mis-identifications as L. kriegi or Liolaemus sp. A, for which we provide the formal name L. zabalai. Nonetheless, there is certainly still much to discover about the diversity of this group of Patagonian lizards.
We thank P. Zabala (Pontificia Universidad de Católica de Chile) for allowing us to review and deposit material in the collection under his care. We are grateful to the following colleagues (and museums) for allowing us to study specimens: H. Núñez (Museo Nacional de Historia Natural), M. Lamborot (Laboratorio de Citogenética, Facultad de Ciencias, Universidad de Chile), J.F. Troncoso (Museo de Historia Natural de Concepción), J. Artigas and J.C. Ortiz (Museo de Zoología de la Universidad de Concepción). F. Lobo, C. Roman, L.J. Avila, F. Tillack, C. Garín and A. Laspiur for sending literature. C.F. Garín and R. Díaz for providing photographs. J. Troncoso-Palacios thanks M. Penna for his support. HAD thanks CONICYT for the support with the master's fellowship FAU thanks the PhD fellowship from CONICYT. D. Esquerré is supported by a Becas Chile-CONICYT scholarship. Two anonymous reviewers greatly improved the work. Thanks to the Servicio Agrícola y Ganadero (SAG) for the collecting permit (N°4468/2014).
Specimens examined. Museum codes are as follow: LCUC (Laboratorio de Citogenética, Facultad de Ciencias, Universidad de Chile), MNHN-CL (Museo Nacional de Historia Natural, Chile), MRC (Museo de Historia Natural de Concepción), MZUC (Museo de Zoología de la Universidad de Concepción) and SSUC (Colección de Flora y Fauna Patricio Sánchez Reyes, Pontificia Universidad Católica de Chile).
Liolaemus buergeri. LCUC 2311. El Planchón, 2370 m. M. Lamborot & M.E. Manzur colls. 07/01/1996. SSUC Re 434–37. El Planchón, road to Teno Lagoon. J. Troncoso-Palacios, L. Negrete & R. Barros colls. January, 2012. SSUC Re 171–180. Maule Lagoon. F. Ferri coll. 20/02/2011.
Liolaemus carlosgarini. MNHN-CL 4531–67. Road to Maule Lagoon. C. Garín coll. 22/02/2008. SSUC Re 181–189, 349. Road to Maule Lagoon. F. Ferri coll. 20/02/2011.
Liolaemus cristiani. SSUC Re 537. El Peine. J. Troncoso-Palacios coll. 28/11/2011.
Liolaemus flavipiceus. MNHN-CL 2118, 2120. Maule Lagoon. C. Veloso & S. Silva colls. MNHN-CL 2167, 2170. Maule Lagoon. J.C. Torres-Mura & H. Núñez. MNHN-CL 4399–07. Laguna del Maule, aguas abajo, 2153 m. C. Garín & G. Lobos colls. 03/03/2008. SSUC Re 169–70. Maule Lagoon. F. Ferri coll. 20/02/2011.
Liolaemus frassinettii. LCUC 800–01. Cantillana. Unknown coll. 14/04/1983. SSUC Re 80. Altos de Cantillana. F. Torres coll.
Liolaemus leopardinus. MNHN-CL 3437–3439. El Colorado. H. Núñez, C. Garín, V. Meriggio, S. Fox & S. Perea colls. 06/01/2001. MNHN-CL 4025, 4027–28. Farellones. C. Veloso coll. 11/01/1988. MNHN-CL 4890–91. El Colorado. D. Esquerré, M. Palma, S. Fox & E. Santoyo colls. February, 2012. SSUC Re 364. Farellones. F. Ferri coll. 12/10/2010. SSUC Re 365. Farellones. F. Ferri coll. 13/02/2011. SSUC Re 366–67. Farellones. F. Ferri, M.L. Carrevedo & J. Troncoso-Palacios colls. 25/01/2012.
Liolaemus neuquensis. MRC 676. La Mula Lagoon, Araucanía Region, Chile. Unknown coll.
Liolaemus ramonensis. MNHN-CL 4007–08, 4012, 4015–17. Quebrada de Macul. C. Veloso & P. Espejo colls. 06/03/1987.
Liolaemus scorialis. SSUC Re SSUC Re 612-17. 7 km NW of the summit of the Antuco Volcano, near the Laja Lagoon, Biobío Region, Chile. J. Troncoso-Palacios, F. Urra & H. Díaz colls. 08/01/2014. MRC 675, 677, 680, 682. La Mula Lagoon, Ralco National Reserve. Unknown coll. 01/12/2001.
Liolaemus ubaghsi. MNHN-CL 3808–16. Chapa Verde. H. Núñez, C. Garín & D. Pincheira-Donoso colls. 22–23/05/2003. MNHN-CL 1601. Chapa Verde. M. Elgueta coll. SSUC Re 491–92. Tranque Barahona, O’Higgins Region, Chile. R. Thomsom & G. Ugalde colls. 15/04/2008.
Liolaemus valdesianus. SSUC Re 129. Cajón del Maipo. Unknown coll. SSUC Re 363. Lo Valdés. F. Ferri coll. 10/01/2011. SSUC Re 559. El Yeso. C. Garín coll. 20/02/2013.
Liolaemus zabalai. SSUC Re 597–602. Near Los Barros, Laja Lagoon, Biobío Region, Chile. Collected by J. Troncoso-Palacios, F. Urra and H. Díaz. 07/01/2014. MZUC 35607, 39567. Malleco, Volcán Antuco, Los Barros. Unknown coll.