Resurrection and redescription of Liolaemus reichei, proposal of a neotype to stabilize its taxonomy

Resurrección y redescripción de Liolaemus reichei, propuesta de un neotipo para estabilizar su taxonomía

Table 2

Morphological measurement of Liolaemus reichei and 5 others “phrynosaurian” species. The order of the numbers corresponds to minimum, average (parenthesis) and maximum values found in the body measurements. Measures are in mm.

	L. reichei (n = 3)	L. manueli (n = 20)	L. poconchilensis (n = 4)	L. torresi (n = 8)	L. audituvelatus (n = 5)
Body length	41.5 (47.7) 50.8	42.7 (50.3) 60.7	47.17 (51.7) 53.83	53.8 (58.1) 64	50.7 (54.9) 58.3
Tail length	35.7 (39.5) 43.1	24 (46.2) 54	42.66 (47.2) 53.65	58.8 (57.4) 74	47.7 (48.9) 50.5
Head length	10 (10.9) 11.5	12.5	11.34 (13.1) 14.16	13 (13.6) 14.5	12.3 (12.9) 13.4
Head width	8.3 (8.5) 9.7	11.9	9.51 (10.9) 11.53	10.3 (11.02) 11.7	9.1 (9.8) 10.7
Forelimb length	21.4 (22.6) 24.4	22.5 (26.4) 28.8	22.2 (24.8) 27.5	26.3 (28.8) 30.5	26.7 (27.7) 30
Hindlimb length	30.1 (31.7) 33.4	31.5 (36.2) 49	31.15 (34.1) 32.62	38.6 (40.1) 42.2	34.5 (36.5) 38.1
Supralabials	9	10	12	10	12
Infralabials	6 (7) 8	7	8	7	7
Lorilabials	9	8	14	9	7 (8) 9
Scales around midbody	43 (45) 47	82	62	71	74
Scales of the body length	50 (52) 54	–	64	86	78 (83) 87
Precloacal pores	4	4	4	3	8

Diagnosis

Liolemus reichei belongs to the L. montanus group (sensu Etheridge, 1995; Schulte et al., 2000). This species differs from all other species of the L. montanus group placed in the L. boulengeri subgroup by the absence of a patch of enlarged scales on the posterior thigh of the hind limb (Etheridge, 1995; Abdala, 2007). In relation of the remaining species in the L. montanus group, L. reichei differs from L. annectens, L. cazianae, L. chlorostictus, L. disjunctus, L. dorbignyi, L. duellmani, L. eleodori, L. erguetae, L. erroneus, L. etheridgei, L. fabiani, L. famatinae, L. filiorum, L. fittkaui, L. forsteri, L. foxi, L. gracielae, L. griseus, L. hajeki, L. halonastes, L. huacahuasicus, L. huayra, L. insolitus, L. inti, L. islugensis, L. jamesi, L. juanortizi, L. lopezi, L. melanogaster, L.montanus, L. multicolor, L. orientalis, L. orko, L. ortizi, L. pachecoi, L. pantherinus, L. pleopholis, L. poecilochromus, L. polystictus, L. pulcherrimus, L. puritamensis, L. robertoi, L. robustus, L. ruibali, L. scrocchii, L. signifer, L. thomasi, L. vallecurensis, L. vulcanus, and L. williamsi by a combination of characters: prominent ciliary scales; diameter of eye greater than length between anterior border of eye and rostral; external auditory meatus without projected scales on the anterior edge; jaws isognathus; smooth dorsal scales and juxtaposed; tail shorter than snout-vent length, which are absent in the species mentioned above. Of the species that have flat, smoothly overlapping scales, L. reichei differs from L. andinus, L. audituvelatus, L. manueli, L. molinai, L. nigriceps, L. patriciaiturrae, L. poconchilensis, L. rosenmanni, L. schmidti, and L. torresi because the latter have a higher number of scales around the body (over 70 scales). From L. stolzmanni it differs by having 9 supralabials scales (6 in L. stolzmanni), the interparietal and both parietal head scales large and well formed (relatively small, fragmented and irregular in L. stolzmanni), 8 lorilabials scales (5 in L. stolzmanni), and ventral side strongly variegated (white, immaculate or lightly variegated in L. stolzmanni) (Fig. 2), and by the presence of lateral nuchal skin folds well-developed and complex (lateral fold weak in L. stolzmanni).

Description of the neotype (adult female). Small lizard, slender body, limbs long and slender, triangular head and short, distinct from neck, widest across temporal region, 0.85 times wider (as measured across widest part of temporal region) than long (as measured from inferior apex of external auditory meatus to anterior surface of rostral). Snout short (as measured from tip of snout to anterior corner or orbit), 0.33 times head length, orbit (as measured along its greatest horizontal length) short, 0.31 times head length. Nasal region swollen, convex in profile, with a transverse indentation where the postrostral scale meets the nasal and internasal scales. Frontonasal region convex in profile. Rostral narrow, 2.7 times wider than high, bordered by 4 postrostrals. Nasals scales larger, separated from rostral and anterior supralabials by anterior lorilabials. Nostril oriented posterolaterally. Dorsal head scales larger, differentiated, convex; 2 irregular prefrontals, 6 larger frontonasal scales arranged in a flower-shaped, a small medial scale surrounded by 5 large scales. All frontonasal scales irregular and convex. Two small postrostral scales. Two hexagonal internasals. Two medium scales between nasal and internasals, 1 small scale between postrostral and internasal, 2 small and irregular scales between nasal and frontonasal scales. Supraoculars large, 9 on each side, 4 irregular scale rows, mostly hexagonal, 4-5 in a horizontal line across widest part of supraocular region between superciliaries and frontals. Eleven and 12 scales form a well-defined circum orbital, somewhat irregular in size. Interparietal slightly smaller than adjacent parietal, hexagonal, bordered by 6 scales, with a distinct “eye” that correspond to pineal organ. Two elongated parietals, well-defined in their forms, posterior to interparietal. Six superciliaries larger, not keeled. Palpebrals small, convex and juxtaposed, 13 inner rectangular ciliaries, outer ciliaries of lower lid 11, triangular, sharply pointed distally; outer ciliaries of upper lid 12, third ciliary and 3 posterior ciliaries triangular, but not as strongly projecting as those of lower lid, those in middle of lid more nearly rectangular, not projecting. Preocular wider than subocular, pentagonal, preceded anteriorly by a large canthal, subocular elongated, about 8 times longer than high and postocular elongted but shorter than preocular and subocular. A single row of lorilabials, interrupted by subocular. Eight lorilabials.

Posterior to subocular, 6 scales smaller than lorilabials and supralabials, rounded and convex. Nine supralabials, equal in size to lorilabials. No supralabials in contact with subocular. One anterior and rectangular loreal, equal in size to second loreolabial, surrounded by preocular, lorilabials and 2 canthals. Temporals larger, convex, juxtaposed, 6 between postocular and anterior margin of ear. External auditory meatus large, rounded, 1.1 times higher than width, vertical diameter 0.56 times longitudinal diameter of orbit, bordered by irregular scales, 2 smaller anteriorly, 1 largest on the upper size and not differentiated from posterior temporals. Mental large 0.92 times as wide as rostral, bordered by 2 infralabials and 2 postmentals, not in contact with anterior sublabials. Infralabials 5, gulars medium size, smooth, rounded, convex, imbricated. Ventral scales triangular, similar in size to dorsal scales, imbricate and smooth. Posterior abdominal scales semirectangular. Scales of precloacal region similar to anterior ventrals, but wider than long. No precloacal pores. Dorsal scales of neck small, flat, slightly concave, triangular, nonoverlaping, no interstitial granules. Dorsal scales of body larger, rounded or semicircular. Some scales semitriangular, smooth and slightly imbricate. Lateral scales wider than long. Scales around midbody 47. Middorsal scales from occiput to point even with anterior margin of thigh 51. Lateral nuchal skin folds well-developed and complex. Two short folds, one originating at superior and other at inferior margin of auditory meatus, converging posteriorly to form a V-shaped fold, continuing posteriorly as longitudinal neck fold, intercepted by oblique neck fold and antehumeral fold. Scales of lateral neck, flat or slightly concave, nonoverlapping. Immediately below the fold, triangular scales, imbricate and larger, similar to the gular scales. Limbs slender and long. Hindlimbs 0.7 times snout-vent lenght. Hindlimbs reaches the posterior edge of the eye. Scales are smooth and imbricate. Brachial scales triangular, about equal in size to dorsal body scales. Antebrachials tend to be rectangular. Elbow scales rounded, and wider than large. Preantebrachials flat, and semitriangular scales. Prefemorals small, convex, juxtaposed. Suprafemorals and postfemorals larger and triangular, smooth and imbricated. Supratibials and pretibials longer than wide, imbricate and rugous. Infratibials smooth, larger and triangular. Supracarpals smooth, rounded, imbricate, supratarsales smooth, triangular and imbricate. Infracarpals imbricate, somewhat projecting, mucronate. Infratarsals rounded, mucronate, imbricate; supradigitals imbricate, keeled and triangular, with distal margin slightly concave. Lateral digitals triangular, smooth, imbricate. Subdigital lamellae of fourth toe 18, claws long and slender. Tail short and thin, slightly thick at the base and somewhat depressed. The rest is slender and rounded distally. Tail 0.85 percent body length. Dorsal and lateral caudal scales tend to be rectangular, smooth and imbricate and wider than long on middle third of tail. Ventral caudal scales triangular on middle third of tail, but then are rectangular and strongly imbricate, longer than wider. Autotomic region with 12 scales on dorsal and lateral tail, and eigth
ventral scales.

The specimen designated as neotype has a dorsum color as light brown or gray color, similar to the sand on which they live. On the dorsal neck are 2 small black spots which are in parallel along to the dorsal body, where they get larger. Each side of the neck has black spots of the medium size, forming 2 parallel bands along the dorsum similar to those previously described. Scapular area shows a black spot subsequently followed a yellow spot, a pattern which is along the dorso-thoracic region; subsequently the yellow spots are located to the ventral side, which is repeated to the inguinal region. The black spots on median dorsal side area accompanied by small white spots. Along the dorsum 4 bands of rounded and black spots, the 2 most lateral with larger spots can be observed. On the tail there is a pattern of black spots laterally disposed, a pattern repeated to the end of tail. Ventrally there is a heavily variegated pattern, more intense on the lateral side and throat. Fore limbs without spots, hind limbs with white scales on the anterior thigh, accompanied with some black scales, without yellow spots.

Variation in morphological measures and scaling. It is based on 2 specimens, a male and a female collected from the same site of the neotype. Variation can be seen in table 1. In relation to the head, there is a low variation in the number and form of the scales, as rostral and postrostral (all with 2 scales), 9 supralabials, circum orbital well defined, large and no fragmented subocular, parietals large, pentagonal and well defined, number of scales between posterior edge of the eyes and ear edge (all with 7 scales), and large antehumeral folds. Difference was observed mainly in the form of interparietal, in both females they are hexagonal, while in the male it is pentagonal, in the latter there are small scales between the interparietal and postfrontals. Furthermore, there is variation in the number of infralabials, scales around and along the body. Variation is also observed in the number of internasals, which are fragmented in females. The juvenile male has only 2 parallel bands of the black spots on the dorsal zone, which are subsequently accompanied by yellow spots. In the middle of the body there are small white spots, which are found together with small black spots in the lumbar region. The tail has a spotted pattern similar to females. Forelimb with small white spots, and hindlimbs with black and white spots. Ventrally there is a heavily variegated pattern, similar to the females. The color of the neotype in alcohol is dark gray in dorsum, and clear gray, with variegated form maintained.

Biology. The knowledge of this species is very poor. Apparently, its distribution is restricted to the Tillansials formation (Pinto, 2005), found on the coastal dunes of Iquique, Chile. Ruiz-de Gamboa and Ferrú (2013) confirmed their distribution on 9 localities, all corresponding to “absolute desert” and “desert scrub”, but they did not make an analysis of inter-population variation. Their activity is crepuscular, and this species regulate its temperature at midday under rocks or Tillansials. Reproductive phenology, as well as diet, is unknown. Liolaemus reichei is consumed sporadically by the fox Lycalopex culpaeus (Guzmán-Sandoval et al., 2007)

Discussion

In proposing the synonymy of L. reichei with L stolzmanni, Langstroth (2011) described the similarity of a single specimen of L. reichei, but failed to point out their differences. Furthermore, the site of collection of his specimen was not provided, and his photograph of the specimen does not show a number of critical characters for a complete comparison.

From the original photographs of L. stolzmanni shown in Langstroth (2011), it is possible to distinguish the difference with L. reichei in number of supralabials, numbers of lorilabials, and the presence of lateral nuchal skin folds well-developed and complex. Interparietal and both parietal head scales large and well formed (Fig. 3A), and ventral side strongly variegated. However, it is not possible to see more details of external morphology, for example the rostral and postrostral scales and how these meet with the nasals and internasals scales. The scales of the frontonasal region, orientation of the nostril, dorsal head scales and circum orbital are poorly observed; superciliaries and outer ciliaries of the upper and lower lids are not distinguishable, subocular and the canthal is difficult to see. The scales around midbody are undistinguished and it is very difficult to count scales along the body. Carpal and tarsal scales are not clearly visible.

Langstroth (2011) considered appropriate to use the description of Donoso-Barros (1966) of L. reichei to compare with L. stolzmanni, for example with ventral scales, number of subocular or conical teeth, but Donoso-Barros did not have in hand specimens of L. reichei sensu stricto and the holotype mentioned there is from Antofagasta region (Herpetological Collection of the Zoology Museum of the University of Concepción, MZUC 011736, ex 5452 private collection of Donoso-Barros, specimens collected near Antofagasta city on January, 1970) here recognized as L. cf torresi. Donoso-Barros (1972) proposed the “Oasis de Pica”, an inland locality at ca. 1,200 m and to the east of Iquique, the type locality of L. reichei.

Steindachner (1891) and Langstroth (2011) mentioned that the subocular is fragmented in 2 or 3 parts in L. stolzmanni, but our specimens of L. reichei have only 1 subocular, similar to the original “big sub-ocular” in Werner (1907) (Fig. 3B). Other similarities between L. reichei and its original description are the hindlimbs extend forward beyond the auditory meatus to the posterior edge of the eye. In view of these observations we propose a revalidation of L. reichei. With regard to L. stolzmanni it will be important to determine the true range. According to Langstroth (2011), no phrynosaurians species are known from high elevations (> 3,000 m), and it is not possible to determine the elevation for L. stolzmanni in “high Perú” or (“hoch Perú” as in the original description of Steindachner). However, it is possible to find L. audituvelatus, L. torresi and L. manueli in the prepuna ecoregion (~ 2,500 m) from Antofagasta and Atacama regions (Núñez & Veloso, 2001; Núñez & Yáñez, 1983; Valladares, 2011) and a similar elevation for L. stolzmanni in Tarapacá, Arica and Parinacota regions is possible, or even in the vicinity of Arequipa or Moquegua, southern Peru.

Acknowledgements

We thank Margarita Ruiz de Gamboa for help in collecting specimens of Liolaemus reichei, to Dr. Juan Carlos Ortiz for permission to one of us (PVF) to analyze specimens of collection of lizards from University of Concepción. We thank Dr. Paul Ramsay from Plymouth University for his revision of the manuscript. To Servicio Agrícola y Ganadero (SAG) for permission to capture No. 239, April 2010, and No. 306, January 2013. PVF and CSA were supported by University of Tarapacá / UTA Mayor de Investigación Científica y Tecnológica Project No. 4713-13 and No. 4717/17.

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