Liolaemus carlosgarini, Esquerré, Damien, Núñez, Herman & Scolaro, José Alejandro, 2013
Esquerré, Damien, Núñez, Herman & Scolaro, José Alejandro, 2013, Liolaemus carlosgarini and Liolaemus riodamas (Squamata: Liolaemidae), two new species of lizards lacking precloacal pores, from Andean areas of central Chile, Zootaxa 3619 (4), pp. 428-452: 431-439
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Liolaemus carlosgarini sp. nov.
Figures 1 to 4
Holotype. MNHNCL- 4563. Male, collected by Carlos Garín at International Road 115, below the Maule Lagoon. 19 H 358560 E – 6018117 S. 1915 masl. WGS 84. On February 22, 2008.
Paratypes. MNHNCL- 4562 and one used for diaphanisation, males; MNHNCL-4561, 4564, 4565, 4566, 4567, 4568 and one used for diaphanisation, females. Same collection data as the holotype. Measurements in Table 1.
Diagnosis. Small to medium sized lizard, with a mean SVL (Snout-Vent Length) of 60.7 mm and a maximum of 68.8 mm. Slender body, moderately robust limbs, head longer than wide, tail over 1.5 times body length, and 86- 101 scales around midbody. Background dorsal coloration is ochre, with a dark occipital stripe, parietal stripes as the background color, and lateral stripes the same coloration as the occipital stripe. Black spots distributed variably between the dorsal region of the head and limbs. Individual white scales scattered on the dorsal region of the body, and a clear and mottled ventral region. It can be distinguished from almost all of the species of the elongatus-kriegi complex by its extreme reduction of precloacal pores. None of the male samples, despite being only three, had these pores, and it seems that a very low frequency of males has them (C. Garín in. litt. 2011). Because the absence of precloacal pores is not absolute in this species, minimum diagnostic traits to distinguish it from all the species of the elongatus - kriegi complex are given.
L. carlosgarini is distinguished from L. austromendocinus , L. elongatus , L. gununakuna (Avila et al. 2004), L. petrophilus , L. choique , L. shitan , L. capillitas , L. dicktracyi , L. heliodermis , L. talampaya , L. antumalguen (Avila et al. 2010), L. umbrifer and L. burmeisteri (Avila et al. 2012) because all of these species have a significantly larger SVL (with a maximal SVL ranging from 82 mm in L. heliodermis to 107.8 mm in L. antumalguen ). In addition, all of them exhibit a lower number of scales around midbody. The only species that overlaps that number with L. carlosgarini (86–101), is L. gununakuna (84–97), but the latter has iridescent yellow body coloration, with dark transversal bars, very distinguishable from the ochre coloration and longitudinal stripes in L. carlosgarini . Finally, none of the species mentioned above exhibit the design pattern of L. carlosgarini , with longitudinal stripes. It is worth noting the striking exomorphological similarity L. carlosgarini has with L. smaug , which in addition is found relatively close to the type locality of L. carlosgarini (between Las Loicas and Volcán Peteroa provincial road, Malargüe, Mendoza, Argentina: 35 ° 39 ’51,3’’S; 70 ° 12 ’00,9’’W, 1688 m), but it differs from this species because L. smaug has a lower number of scales around midbody (73–80), in addition to a constant presence of precloacal pores in males (3–4) (Abdala 2010; Abdala in. litt. 2011). Sexual dichromatism has not been observed in L. carlosgarini as in L. smaug , nor have golden yellow specimens of L. carlosgarini been observed, but this trait deserves further analysis of live specimens.
From L. cristiani it can be differentiated because L. cristiani has a very dark and pronounced melanistic stripe on the flanks (quite different from the paler stripe in L. carlosgarini ), and in the absence of an occipital stripe, present in L. carlosgarini . L. cristiani is larger (mean SVL of 70.67 vs. 60.7 mm), and has fewer number of scales around midbody (83–89 vs 86–101), although these values overlap.
It is differentiated from L. ceii , L. kriegi , L. buergeri , L. ramonensis , L. valdesianus and L. leopardinus because all of them are larger, the smallest being L. buergeri from Pichuante in Teno River, Chile, with a mean SVL of 73.2 mm and a maximum of 87.2 mm. All of these species have precloacal pores and differences in pattern, coloration and squamation. Nevertheless, L. leopardinus males lacking pores can be found, although the exomorpohological differences, especially in pattern, are very notorious. From L. coeruleus and L. neuquensis it can be distinguished by the blue or greenish ventral coloration in both species, and in having fewer scales around midbody. From L. thermarum , L. punmahuida , L. flavipiceus , L. tregenzai and L. riodamas it can be differentiated because all of these have a lower number of scales around midbody, larger size, and a uniform pattern, opposed to the well-defined design in L. carlosgarini .
Description of the holotype. Male, with SVL of 65.35 mm. Head slightly longer than wide: 15.59 mm long (from the anterior edge of the ear opening to the point of the snout), 12 .0 5 mm wide (between the anterior edges of the ear openings) and 9.91 mm high (at the level of the anterior edge of the ear openings). Snout length (from the anterior margin of the eye to the rostral scale): 4 .0 2 mm. Neck slightly wider than head, effect of the transversal folds of the neck. Hind limb extended forward barely exceeds the armpit. The tail length is 1.36 times the SVL. It lacks precloacal pores.
Rostral scale rectangular, 2.3 times wider than high and in contact with eight scales, including the nasals, which touch the rostral with only the anterior corner of the scale. Nasals polygonal, the nostril located posteriorly, occupying half of the scale’s surface. Two postrostrals elongated laterolaterally, followed by four internasal scales, being the two medial ones about four times larger than the side ones. Two frontal azygos, the anterior two times longer than the posterior. Six frontonasals in contact with the frontal azygos. Two prefrontal scales, and the frontal divided transversely; the anterior larger than the posterior. Four small postfrontals. Interpatietal pentagonal, posteriorly elongated, in contact with five scales. A gray pineal eye in the middle of interparietal scale. The two parietal scales are about twice the size of the interparietal. The scales in the nuchal and supratemporal regions are small, polymorphic, juxtaposed and generally smooth, with small ridges on some scales. 7 - 6 large supraoculars, accompanied by 21 - 18 small supraoculars. 6-8 very elongated and imbricate superciliary scales, excluding the canthal. Numerous scale organs on all the dorsal region of the head, increasing in numbers towards the anterior part of the head. 7 - 6 loreal scales, excluding the canthal, and a large and elongated subocular scale that spans the entire length of the eye. 7 - 7 lorilabial scales, in one row, being the sixth one the most elongated. 7 - 6 supralabials, fifth-fourth one more elongated and curved upwards posteriorly. It has a slightly elongated snout, with the loreal distance (distance between the anterior margin of the eye and the rostral scale) greater than the ocular diameter. The scales in the anterior border of the eye are elongated and overlapping, each one with a scale organ on it. 16–19 upper and 16 – 15 lower palpebrals, all squared and each one with a scale organ on it. Temporal scales rounded, smooth and subimbricate. Auricular scale is poorly developed but distinct.
Tympanic scales small and highly convex. Scale organs present in all the lateral region of the head, being especially numerous in the lorilabial and loreal scales. Mental scale barely wider than rostral, but notably higher, and in contact with four scales. Five pairs of postmentals, the second pair separated by two scales. 5 - 5 infralabials, with scale organs scattered on them. Scales on the gular region rounded, smooth and imbricate. Lateral region of neck with two prominent transversal folds. Neck scales small, convex, granular and with very small granular scales in the interstitial space.
Dorsal scales small, the same size as the ventral scales. They vary from rounded to subtriangular, and from juxtaposed to subimbricate. Moderate keels on the mid-dorsal scales form longitudinal lines along the dorsum, reaching the base of the tail. The keels become less prominent towards the side of the body, having already disappeared on the flanks. The scales on the flanks therefore are smooth, but in form and arrangement very similar to the dorsal scales, only being slightly smaller. Tiny granular scales are scattered in interstitial space in the dorsal region and flanks, where they are more evident. In the lateral region of the body a slight and thin fold extends longitudinally from the armpit to the groin. Ventral scales are rounded, smooth and imbricate. 95 scales around midbody. Dorsal scales of the arm are subtriangular, smooth and imbricate. Dorsal scales of forearm are very similar but more rounded, and then the more triangular shape is observed in the dorsal region of the hand. On the ventral region of the arm the scales are small, granular, juxtaposed, and surrounded by even smaller granular scales in the interstitial space. Then, the scales on the ventral region of the forearm are as large as the dorsal ones, subtriangular, imbricate, with very few granular scales in the interstitial space, and they start showing keels towards the palm, where the scales are completely keeled, triangular, imbricate and with a jagged edge. The third finger of the left hand has 20 rectangular and transversally disposed lamellae, each one provided with three keels. The scales on the dorsal femoral region are subtriangular, smooth and imbricate. In the dorsaltibial region they are more rounded, subimbricate, slightly keeled and with visible tiny granular scales in the intertstitial region. In the dorsal region of the foot the scales are subtriangular, smooth and imbricate, more similar to the scales on the femoral region rather than those on the tibial region. The scales on the ventral femoral and tibial regions are rounded, smooth and imbricate. In the plantar region scales are triangular, keeled and imbricate. Some scales in the plantar region are slightly jagged on the edges. The fourth toe has 26 rectangular and transversally disposed lamellae, each one provided with three keels. The dorsal scales on the tail are quadrangular, imbricate and keeled, with a slight mucron. Ventral scales on the tail are triangular, smooth and imbricate.
Color and pattern in preservative. The background dorsal coloration is ochre, with tiny black spots scattered in the dorsal region of the head and limbs. A well-defined black occipital stripe, although with approximately only half of the scales on the stripe being melanistic. This stripe reaches the base of the tail, and from there it continues as a thin line of tiny black spots longitudinally elongated. On the sides of the occipital stripe follows a pair of parietal stripe shaving the background ochre coloration, and then, in the flanks a pair of melanistic temporal stripes, which originate at the posterior edge of the eye, and end at the groin. The back of the body exhibits white scales evenly scattered. Ventral coloration is whitish gray, with dark and inconspicuous dots in the gular region.
Variation in the paratypes. Body measurements were divided into males and females (holotype is included within males since there are only two males in the whole sample): Mean and extreme values, in mm, for the males (two specimens) are: snout-vent length: 63.3 (61.25–65.35); axilla-groin distance: 28.36 (27.59–29.12); left forelimb length: 23.41 (21.74–25.07); left hindlimb length: 38.92 (37.23–40.6); tail length: 100 (89–111); head length: 15.13 (14.66–15.59); head width: 12.18 (12.05–12.3); and head height: 9.37 (8.82–9.91). For the females (six specimens) the measurements are: snout-vent length: 59.83 (53.46–68.8); axilla-groin distance: 26.12 (22.65–29.65); left forelimb length: 23.28 (22.65–25.11); left hindlimb length: 36.19 (32.53–39.16); tail length: 100 (85–113); head length: 13.32 (12.82–14.7); head width: 10.23 (9.93–11.25); and head height: 7.29 (6.55–7.54). Therefore we assume there is no evidence of sexual dimorphism, even though we measured only two males, the ranges of males measurements are within the range of the females. Nevertheless, a slightly larger size in the head measurements in males in relation to measurements in females is seen. Qualitatively the two males have a slightly more robust appearance than the six females.
In the two males, the mean and extreme values of the number of scales around midbody are 92.5 (90–95). None has precloacal pores. The male paratype exhibits the following variation in squamation with respect to the holotype: Three postfrontal scales instead of four, interparietal in contact with six scales instead of five, 5 - 5 large supraoculars instead of 7 -6, 18- 16 small supraoculars instead of 21 - 18. 6 - 7 lorilabials instead of 7 - 7. Slightly more rounded dorsal scales. 21 lamellae on the third finger of the left hand instead of 20. Scales in the dorsal femoral region are rounded and slightly keeled. Plantar scales do not have a jagged edge. 27 lamellae on the fourth toe of the left foot instead of 26. In the females, the mean and extreme values of the number of scales around midbody are 92.6 (86–95). None has precloacal pores. In comparison with the holotype the females exhibit the following variation: longitudinal neck fold instead of two transversal folds in one specimen. Rostral scale in contact with six scales, and without contact with the nasals in two specimens. Undivided frontal scale in one specimen. Three postfrontals in three specimens, and two in three specimens. 4 to 6 large supraocular and 13 to 20 small supraoculars. 6 to 9 lorilabials. 5 to 8 supralabials. 4 to 6 infralabials. Four pairs of postmentals in two specimens. Dorsal scales rounded, imbricate and without visible small granular scales in the interstitial region in two specimens, and triangular, imbricate and without visible small granular scales in the interstitial region in one specimen. Dorsal scales of the forearm very slightly keeled in one specimen. Ventral scales of the forearm rounded and without visible small granular scales in the interstitial region in four specimens, and completely smooth in one specimen. Scales in the palmar region without a jagged edge in two specimens. 20 to 23 lamellae on the third finger of the left hand. Dorsal femoral scales rounded in one specimen, and slightly keeled in another specimen. Dorsal tibial scales triangular in one specimen, and without visible small granular scales in the interstitial region in four specimens. Plantar scales with a strongly jagged edge in one specimen. 27 to 30 lamellae on the fourth toe of the left foot.
There is no evidence of sexual dichromatism in the preserved specimens. This could not be inferred from the pictures because the sex of the specimens photographed is unknown. In some of the specimens the melanistic occipital stripe is weaker, consisting of only a few black scales between the ochre ones. In the specimens with a more pronounced and filled stripe, little black dots are scattered in the background colored parietal stripes. One of the female paratypes exhibits the whole ventral and gular regions mottled with black dots, while in the rest of the sample, this trait is very weak, or simply not present.
Osteological description. Based on specimen MNHNCL- 4560, female, double-stained skeleton. ( Figure 5View FIGURE 5. — A)
Cranium features: cranium length 15.26 mm (from occipital condyle to premaxilla); cranium width 9.11 mm (between sutures of jugal and maxilla); cranium height 5 .0 9 mm (the highest part of the calvarium); ocular orbit 5.55 (from the lacrimal bone to the postorbital one); rostrum 5.27 mm (from the lacrimal to the premaxilla). Rounded premaxilla with lateral process, maximum width of premaxilla half of its length. Two foramina are evident, these are the passages for the medial ethmoidal nerves (Oelrich 1956). These foramina are limited to the premaxilla and the maxilla. Nasal process of the premaxilla is acutely projected backward, lying beneath the divergence of the nasal bones. Ventrally, the premaxilla bears six acute teeth, chiseled, and caniniform on their free tip. Posterior margins of premaxilla are projected forward, almost horizontally. The premaxilla is strongly sutured to the vomer bones.
Nasal capsules (=fenestra exonarina) with medial margins limited by the premaxillar spine, the floor is formed by the septomaxilla and the premaxillar process of the maxilla. The roof is formed by a cartilage. The nasal capsule is pierced and communicates with the fenestra exochoanalis. Paired nasal bones, twice as long than wide, medially sutured, anteriorly they cover the caudal tip of the premaxillar spine as described. Laterally the nasals are sutured to the ascendant process of the maxilla, and solidly to the prefrontal bones. Nasals exhibit small foramina, scattered, variable in diameter. Caudally the nasals diverge to embrace the anterior tip of the frontal bone, where they are solidly sutured. The divergence is located at the anterior orbital level. Dorsal surface of nasal bones appear wrinkled, with depressions and scars of the scales on them. Paired and triangular prefrontal bones; medially they are sutured to the nasal bones; their caudal tips are sutured to the single frontal bone. Prefrontals are sutured to the ascendant process of the maxilla and to the tiny lachrymal bone. Prefrontals are abruptly deflected downward forming the descendant process of the prefrontals. This process is strongly sutured to the palatine bones, forming the anterior wall of the ocular capsule. The lachrymal foramen is completely limited by the descendant process of the prefrontals. Its external wall is completely formed by the lachrymal bone.
The frontal bone is an unpaired, flat and tubular bone. The anterior process is sutured to the nasal and prefrontal bones. The frontal is the superior margin of the ocular orbits. Backwards it becomes broadly wider and it sutures to the parietal bone. At least in this specimen, the pineal foramen is completely included in the frontal. At both sides of that foramen there are irregular windows. The external borders of the frontal are sutured to the small postfrontal bones. Ventrally, the frontal has the olfactory channel. The postfrontals are paired bones, bridging the frontal and postorbital bones.
The parietal bone is a single one, broad, and widely sutured to the frontal. By its anterior and external process, the parietal is sutured to the postfrontal and postorbital bones. Laterally, the parietal is deflected downward; the epipterygoid bones reach these deflections. The parietal is projected towards caudal and lateral. Those projections are the supratemporal processes. These processes are sutured to the squamosal bones, to the tail of the supratemporal, and to the paraoccipital and exoccipital bones. The caudal portion of the supratemporal bones is surrounded externally by the squamosal and quadrate bones and medially by the paraoccipital and exoccipital processes. The parietal has a notch beneath the supratemporal processes, in which is houses the anterior portion of the supratemporal bones. Supratemporal fenestra measures 3.83 mm in anteroposterior axis.
Squamosal bones are paired, stick shaped, curved and with an anterior process weakly joined to the postorbital bones. Posteriad they become broader like an anchor. This caudal portion has a process inserted in a notch of the quadrate. The superior process is sutured to the supratemporal process of the parietal. Quadrates are paired, auricular-shaped bone. Ventrally they have sulcated condyles, supporting the mandibular set of bones. Dorsally they have the cephalic condyles. These condyles have a notch, and in it, is inserted the caudal portion of the squamosal bones, which are joined by cartilaginous fibers. There is a posterior crest rising from the mandibular condyle upwards. On the inferior and internal area there is a union with the posterior valves of the pterygoid bones.
Postorbitals are flattened paired bones, with three processes. The anterior one is sutured to the jugal bone, the posterior process is connected to the anterior process of the squamosal bone and the medial process is projected inward, joining the parietal and postfrontal bones. Jugals are paired bones, long and curved, slanting downward and forming the inferior margin of the ocular orbit. On their lowest part they are deflected forward and then upward riding on the maxilla, to which they are sutured, reaching the lachrymal bone. On their external surface, the jugals are pierced. On their lowest part, they are strongly sutured to the ectopterygoid.
Maxillae are paired bones, with an anterior process sutured to the premaxilla with a medial process. The ascendant process sutures to the nasal, the prefrontal and the lachrymal ones. The posterior process sutures to the jugal as described. The maxillae bear 18 pleurodont teeth, housed in a basin, the crista dentalis. Teeth 1 to 13 have tricuspidated crowns; 14 to 18 are strongly acute and caniniform. Anteriad, the maxillae are sutured by a little extension to the vomerian bones. Ventrally, the maxilla forms the outer margin of the fenestra exochoanalis, a broad arch. It features a rounded palatal process, which is sutured to the palate. Posteriorly, the maxilla forms the outer margin of the infraorbital fenestra, completed externally by the ectopterigoyd; the internal margins of the infraorbital fenestra are formed by the pterygoid and palatine bones. In dorsal view, the ectopterigoid is prolonged via an acute anterior process that reaches the level of tooth 5.
Paired vomers strongly sutured to each other. Anteriorly they are sutured to the premaxilla, and then to the palatine to which they bind obliquely. The vomer bones form the medial margin of the fenestra exochoanalis. Between them there is a shallow recess. The paired palatine bones are elongated, with three processes. The anterior one is sutured to the vomers. An anterior and external process is projected to the maxillae to which it sutures. Its caudal process is sutured to the pterygoid. Sutures are so skewed, that dorsally and forth they are sutured to the descending process of the prefrontal. On the ventral surface of the palate, the roof of the mouth, are presented noticeable longitudinal striations.
Palatine bones are not sutured together, and form the anterior piriform recess. This recess is continued, consisting of pterygoid divergent branches. The pterygoid bones are paired, anteriorly joining the palatines. In the middle of the pterygoid there is an extension that is projected obliquely outwards to join solidly to the pterygoid process of the ectopterygoid bone. Pterygoids are caudally projected in two valves to reach the base of the quadrate. The pterygoids in the middle of their length have two basins that host the basipterygoid processes of the basisphenoid bone. Immediately above them, on the dorsal side of the pterygoid there is a notch from which rises a slender rod that is the epipterygoid. In this individual at least, there are no pterygoid teeth.
The basisphenoid is a single bone, projecting forward it has basipterygoid processes that attach to the pterygoid bones. The basioccipital is fused to the basipterygoid. On the anterior part is the basisphenoid process, projected deep into the piriform recess. The basioccipital is a medial bone carrying the cephalic condyle. There is a pair of lateral eminences projecting to the ventral side, in front and sides of the occipital condyle: the spheno-occipital tubercles. The recesses for the jugular vein open like winged bodies. The otic occipital portion is projected forward as a rounded body.
The mandible: the dentary bears 23 teeth. Teeth 1 to 16 (17 missing) are tricuspid, 18 to 23 are caniniform, housed in a deep basin, typically pleurodont. The dentary at its distal end ventrally presents a discrete Meckel’s groove. Caudally and via lingual view, the dentary diverges to house the splenial bone with a strong suture. Together, both bones form the anterior inferior alveolar foramen (see Oelrich 1956) that extends at the level of teeth 3 and 4. The dentary is deflected upward by the dorsum of the splenial process of the coronoid. The dentary bone, by ventral view, is projected backward, beyond the coronoid bone, to suture with both the angular and suprangular bones.
The dentary via ventral and lingual view is projected backward until the ventral mandibular coronoid, beyond the coronoid it sutures with the angular and suprangular bones.
The coronoid bone rides on both the dentary and splenial bones. Posteriad it is sutured to the suprangular bone. The latter has a large basin, the mandibular foramen. The articular is the most proximal bone and articulates with the quadrate as described.
The sternal apparatus: paired clavicles, flat and dorsally curved at the tips. They are widened as a spatula, without processes along the bone. The single interclavicular bone is arrow-shaped. Two lateral processes at the anterior end projecting to the side and backward, and a medial process that is projected straight caudally, reaching the middle of the sternal fontanelle; the angle between the two processes is 66 º. The medial process is flat with a widening in the middle that ends in a point with the appearance of a necktie.
The epicoracoid is centered, covered by interclavicular bone. With the coracoid they form two fossae: the anterior coracoid fenestra and the smaller posterior coracoid fossa. The coracoids are paired bones that come together with the epicoracoid as described. They bear the glenoid fossa in connection with the humerus. The coracoids exhibit a foramen. The sternum is a triangular-shaped shield featuring a sternal fossa. The outer edges of the sternum have jagged ends to contact with four pairs of ribs. The last two are the xiphisternum.
The tibia of this species lacks the presence of a blade-shaped tibial process, therefore this indicates it belongs to Liolaemus (Liolaemus) sensu strictu .
Distribution. The vicinity of the Maule Lagoon, the type locality, is the only place where this species’ presence can be assured. Very similar specimens also lacking precloacal pores have been found at the base of the Copahue Volcano, Argentina, at 2000 masl, very close to the Chilean border, and also in Caviahue, Argentina (J.A. Scolaro). It is also possible to find similar specimens at the Puelche River, Chile, slightly north of the type locality, although that must be confirmed with further sampling (C. Garín in. litt. 2011).
Natural History. Probably a viviparous species, just like similar species that inhabit high altitudes. Saxicolous. Diet unknown. The sympatric herpetofauna consists of L. buergeri , L. flavipiceus , Phymaturus maulense (Núñez et al. 2010) and Pleurodema bufonina .
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