ABSTRACT: Liolaemus azarai is described as a new species of liolaemid lizard from northeastern. Argentina and Yacyretá Island, Paraguay. It is a member of ...
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Herpetologica, 59(2), 2003, 283–292 q 2003 by The Herpetologists’ League, Inc.
A NEW SPECIES OF LIOLAEMUS (SQUAMATA: LIOLAEMIDAE) FROM NORTHEASTERN ARGENTINA AND SOUTHERN PARAGUAY LUCIANO J. AVILA1,2 CONICET-CRILAR. Mendoza y Entre Rı´os s/n. 5301. Anillaco, La Rioja, Argentina ABSTRACT: Liolaemus azarai is described as a new species of liolaemid lizard from northeastern Argentina and Yacyreta´ Island, Paraguay. It is a member of the wiegmannii species group and may be distinguished from other members of this group by its relatively small body size, fewer scales around the midbody, and dorsal color pattern. At the type locality, on Yacyreta´ Island, L. azarai is restricted to isolated sand dune formations and is known only from two others localities in the Province of Corrientes, northeastern Argentina. The new species is the smallest member of the wiegmannii group and, like other members of the group, except L. wiegmannii, L. azarai appears to be a sand dune obligate, insectivorous, and oviparous. Key words: Argentina; Iguania; Liolaemidae; Liolaemus; Liolaemus azarai new species; Paraguay; Yacyreta´ Island
LIZARDS of the genus Liolaemus are the most species rich group of vertebrates in austral South America, and the number of described taxa is quickly approaching 180. The Liolaemus species are relatively small to moderately sized diurnal lizards, the majority of which are terrestrial, even though a few are semi-arboreal. Diets may 1 PRESENT ADDRESS: Department of Integrative Biology, Brigham Young University, 401 Widtsoe Building, Provo, UT 84602, USA. 2 CORRESPONDENCE: e-mail, lucianopjavier@hotmail. com
be insectivorous, herbivorous, or omnivorous; both oviparous and viviparous taxa are represented (Cei, 1986, 1993; DonosoBarros, 1966). Liolaemus ranges from the high Andean mountains of Peru and Bolivia in the north to northern Tierra del Fuego Island in the south and from the Pacific beaches to the sandy Atlantic beaches of Argentina, Uruguay, and Brazil (Cei, 1986; Donoso-Barros, 1966). Several different taxonomic groupings within Liolaemus have been proposed (e.g., Cei, 1979, 1986, 1993; Etheridge,
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1995; Laurent 1983, 1984; Schulte et al., 2000). The wiegmannii group is one of the smallest species groups for which there is strong evidence for monophyly based on morphological and behavioral characters (Etheridge, 2000). A cladistic analysis, based on molecular data, that included all species of the wiegmannii group except L. riojanus and L. rabinoi (Schulte et al., 2000) also found strong evidence for its monophyly. This group includes L. lutzae, L. multimaculatus, L. occipitalis, L. rabinoi (probably extinct), L. riojanus, L. salinicola, L. scapularis, and L. wiegmannii (Etheridge, 1995, 2000). The first three species are restricted to sand environments, such as the beaches and coastal dunes along the Atlantic shores of Brazil, Uruguay, and Argentina, while the latter species (except L. wiegmanii) live inland on isolated sand flats and dune systems of western Argentina (Etheridge, 2000). Liolaemus wiegmannii is considered to be the least specialized species and is not a sand obligate (Etheridge, 2000; Halloy et al., 1998; Laurent, 1983, 1984). Throughout much of central and northern Argentina, L. wiegmannii is an abundant and conspicuous lizard in a wide variety of habitats. After studies of the L. darwinii complex by Etheridge (1992, 1993), L. wiegmannii was recognized as one of the most widely distributed species of Liolaemus in South America with populations in Argentina, Uruguay, and Brazil (Achaval and Olmos, 1997; Cei, 1986, 1993). Some degree of morphological differentiation occurs between the northern populations from the Salta, Jujuy, and Catamarca provinces and the southern populations of central Argentina (L. J. Avila, unpublished data); further studies will resolve the taxonomic uncertainty of these northern populations. In the course of a revisionary study of L. wiegmannii populations of Argentina, I found that some populations previously assigned to L. wiegmannii (Alvarez and Lions, 1996; Tedesco et al., 1992) are members of a new undescribed species that appears to be restricted to sand dunes in northeastern Argentina and Yacyreta´ Island, Paraguay.
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MATERIALS AND METHODS Measurements were taken with a dial caliper to the nearest 0.1 mm, and scale counts were made with the aid of a stereoscopic microscope from specimens fixed in 10% formalin and preserved in 70% ethanol. Definitions for scale counts and body patterns follow Lobo and Espinoza (1999), and nomenclature for neck folds follows Frost (1992). Coloration in life was observed from the holotype and photographs of live paratypes. Sex was determined by dissection or by the thickness of the base of the tail and presence and number of precloacal pores. Voucher specimens of the new species are deposited in the herpetological collection of the Universidad Nacional del Nordeste, Corrientes (UNNEC); Fundacio´n Miguel Lillo, Tucuma´n (FML); and Universidad Nacional de Rı´o Cuarto, Rı´o Cuarto (ZV-UNRC). The following acronyms are used throughout: MACN (Museo Argentino de Ciencias Naturales Bernardino Rivadavia), CENAI (Centro Nacional de Investigaciones Iolo´gicas, now housed in MACN), IMCNUNSJ (Instituto y Museo de Ciencias Naturales, Universidad Nacional de San Juan), MLP-S (Museo de La Plata, Universidad Nacional de La Plata), and LJAMM (Centro Regional de Investigaciones Cientificas y Transferencia Tecnologica La Rioja). SPECIES DESCRIPTION Liolaemus azarai sp. nov. Liolaemus wiegmanni—Tedesco et al. (1992) Liolaemus wiegmanni—Alvarez and Lions (1996) Liolaemus wiegmannii—Etheridge (2000) Holotype.—FML 8454 (Figs. 1, 2), an adult male from sand dunes from the northwestern remnant of Yacyreta´ Island (Parana´ River), Itapua´ Department, San Cosme and Damian district, Paraguay (278 349 S, 568 419 W, 243 m); 24 February 1998; collected by B. B. Alvarez, M. L. Lions, and R. Aguirre. Paratypes.—UNNEC 565, 584, FML 10372 (males), UNNEC 563, 564, 586 (females) from Curuzu´ Laurel (278 509 S, 578
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FIG. 1.—Liolaemus azarai in life, holotype (FML 8454), adult male 47.8 mm in SVL.
309 W), San Miguel Department, Corrientes Province; 14 March 1992; collected by R. Aguirre, B. B. Alvarez, A. Avalos, L. Rey, and T. Tedesco. UNNEC 588, 589, 593 (males), 590, 591, 4336 (females) from Curuzu´ Laurel (278 509 S, 578 309 W), San Miguel Department, Corrientes Province; 26 October 1992; collected by R. Aguirre. UNNEC 917 (male), 916 (female) from Curuzu´ Laurel (278 509 S, 578 309 W), San Miguel Department, Corrientes Province; 26 November 1994; collected by B. B. Alvarez, R. Aguirre, and T. Tedesco. Etymology.—Named to honor Felix de Azara, Spanish explorer and naturalist, in recognition of his contributions to the knowledge of Paraguayan natural resources in the late eighteenth century. Diagnosis.—Liolaemus azarai is the smallest species in the wiegmannii group, with a maximum adult size of 54.3 and 48.7 mm SVL in males and females, respectively; L. lutzae (80.0 and 70.2 mm; n 5 10), L. multimaculatus (72.3 and 63.5 mm; n 5 35), L. occipitalis (71.5 and 60.4 mm; n 5 9), L. rabinoi (61.0 and 68.0 mm,
n 5 9; Etheridge, 2000), L. riojanus (62.3 and 56.0 mm; n 5 20), L. salinicola (77.0 and 67.4 mm; n 5 70), L. scapularis (75.0 and 65.0 mm; n 5 40), and L. wiegmannii (59.1 and 60.0 mm; n 5 250). Midbody scale number of L. azarai does not overlap with other members of the wiegmannii group, except L. wiegmannii. Liolaemus azarai has 32–44 midbody scales while other species have the following: L. lutzae (56–75; n 5 10), L. multimaculatus (69– 93; n 5 35), L. occipitalis (66–85; n 5 9), L. rabinoi (73–81, n 5 9; Etheridge, 2000), L. riojanus (61–79; n 5 20), L. salinicola (58–78; n 5 70), L. scapularis (49– 68; n 5 40), and L. wiegmanii (38–60; n 5 198). Liolaemus azarai has a more slender body than L. multimaculatus and L. riojanus, and the dorsal color pattern of these species has several transverse rows of small rhomboidal spots that are different from the dorsal pattern of L. azarai described below. Males of L. multimaculatus and L. riojanus have a single suprascapular spot, whereas males of L. azarai have two such
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FIG. 2.—Dorsal (top) and lateral (bottom) views of the head of the holotype of Liolaemus azarai (FML 8454). Scale bar 5 2 mm.
spots. The color pattern of L. azarai differs from L. salinicola because the latter has a gray dorsal coloration with several rows of longitudinal and small paravertebral spots with white centers. Liolaemus rabinoi has gray and blue dorsal coloration and round black spots on the venter of males. These characteristics are not present in L. azarai. Liolaemus scapularis has numerous black spots in the gular region, chest, and along the sides of the body that are not present in L. azarai. The general color pattern of L. lutzae is gray with a mid-dorsal orange bar outlined laterally by smaller brown and white bars, a pattern not seen in L. azarai. Liolaemus occipitalis has an occipital scale 2–2.5 times larger than L. azarai, a series of darker spots on each side of the vertebral line, and two darker bands in the lateral field extending from axilla to groin and
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separated by a white streak; these features are not present in L. azarai. Liolaemus azarai has been previously confused with L. wiegmannii because of its similarity in external characteristics. Dorsal head scales of L. wiegmannii are rugose or slightly keeled, protruded, and rounded in shape; in L. azarai, scales are quadrangular, rectangular, or polygonal shape, with a well marked keel in internasal, frontonasal, prefrontal, and frontal scales. Nasal scales position is dorsolateral in L. azarai and lateral in L. wiegmannii. Canthal ridge is distinct and sharp in L. azarai and is distinct but blunt in L. wiegmannii. Unlike L. wiegmannii, L. azarai exhibits no strong sexual dichromatism in adult coloration; apparently intrapopulation variation in dorsal coloration is low. Dorsal background coloration is lighter in L. azarai (tan–light brown) than in L. wiegmannii (brown–dark brown, with small black areas). Prescapular and postscapular spots are very conspicuous in L. azarai, but blurred with the lateral dark marks in L. wiegmannii. Paravertebral and lateral field markings are larger in L. wiegmannii than L. azarai. In L. wiegmannii the half-moon shaped marking in the paravertebral field is well marked (3–5 scales wide, 2–4 high), enclosing almost completely (⅔) a white dot, but is smaller (2– 3 scales wide, 1–2 high) in L. azarai, enclosing only partially (⅓) a white dot. Midbody paravertebral markings in L. wiegmannii are connected by a line of black scales on each side of the mark and along edges of paravertebral fields; paravertebral markings never contact each other in L. azarai. Dorsolateral stripe is creamy white, 1–1.5 scales wide in L. azarai, and white or yellow, 1.5–2 scales wide in L. wiegmannii. Dorsal dark pigmentation is more extensive in L. wiegmannii than in L. azarai; black dots are numerous, larger, and interconnected in dorsal hind and fore legs, tail, and head sides of L. wiegmannii, but are small and sparsely spotted in L. azarai. Blue scales in males of L. wiegmannii are numerous, larger, and randomly distributed along the lateral field, but forming an irregular line 1–2 scale rows in width in the upper lateral field in L. aza-
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rai. Bright yellow or orange scales observed in L. wiegmannii males are completely absent in L. azarai. Description of the holotype.—Adult male. SVL 47.8 mm, total length 78.3 mm. Head width 8.4 mm, head length 11.0 mm (1.3 times longer than wide). Eighteen dorsal head scales (from a line drawn horizontally between anterior margin of external auditory meatus to anterior margin of rostral). Nasal scales not in contact with rostral, separated from first supralabial by one scale. Nasal bordered by eight scales. Nasal dorsolateral in orientation, separated from rostral by postrostral and from supralabials by two scales. Nares oval, occupying posterior one-half of the nasal. Nasals separated medially by nine internasals in two rows of four anterior and five posterior keeled scales each. Frontonasals (4 1 1 inconspicuous scale), prefrontals 15, frontal scale keeled. Canthal separated from nasal by one scale. Loreal scales convex, some slightly keeled. Loreal–preocular region below canthal ridge slightly concave. Canthal ridge distinct and sharp. Six enlarged, convex supralabials. Infralabials flat, with ventral ‘‘keel’’ where they meet sublabials, enlarged, nearly rectangular, four right and five left. Rostral subtriangular 2.2 times wider than high, bordered behind by four postrostrals (2.2 and 1.0 mm, respectively). Two rows of lorilabials: upper row discontinuous and with small, irregular scales, 10 on right, 8 on left; 9 on lower row. A row of scale organs located along lorilabials (one or two per scale). Subocular elongate (3.0 mm), about equal to diameter of orbit (2.7 mm from anterior to posterior margin of ciliaries). Preocular scale small, in contact with canthal. Postocular small, with sharp outer border, situated at level of sixth supralabial scale. Auditory meatus slightly oval (height 1.2, width 1.0 mm) with two (right side) and one (left side) small, white, convex nonprojecting scales along anterior margins. Seven slightly keeled, imbricate temporal scales between the upper anterior corner of auditory meatus and posterior margin of orbit. Upper temporals keeled, lower temporals smooth. Nine temporals (counting vertically from the buccal commissure to
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posterior corner of orbit). Scales of anterior margin of auditory meatus large; scales of the posterior margin smaller and granular. Orbit–auditory meatus distance, 4.3 mm. Orbit–rostral distance, 3.6 mm. Interparietal slightly octagonal, irregular, with a conspicuous, opalescent parietal eye. Interparietal bordered by eight scales, the parietals being the largest. Frontal region with two small azygous scales. Supraorbital semicircles complete, formed by 18 scales (right) and 16 scales (left). Five enlarged supraoculars on right side, three small in the front, laterally separated from superciliaries by three rows of 15 small, irregular scales. On left side, six enlarged supraoculars, four small in the front, separated from superciliaries by 16 small, irregular scales. Dorsal head scales with lenticular scale organs most apparent in interorbital region (5–6 per scale). Scale organs scattered (1–4 per scale) in the prefrontal and internasal region. Superciliaries four, strongly imbricate. Fourteen upper ciliaries weakly projecting and rectangular; 12 lower ciliaries of similar shape to upper ciliaries. Palpebrals small and irregular, slightly convex and juxtaposed. Mental in contact with six scales. Mental scale heptagonal (1.8 mm long and 1.3 mm wide). Chinshields forming two longitudinal rows of three enlarged scales separated one from the other by small scales. Infralabials separated from chinshield by 1–2 small scales. Scales of throat subtriangular or round, flat, imbricate. Five types of folds found along the lateral neck region: postauricular, rictal, longitudinal, gular, and antegular. The first three form a ‘‘Y’’ between auditory meatus and shoulder. Postauricular, rictal, and longitudinal folds well developed. Gular fold deep, well delimited by large, flat scales along its anterior margin and very small scales in pocket. Antegular fold reduced, formed by very small scales. Scales over longitudinal, postauricular, and rictal folds small to medium, irregular, some granular or laminar, imbricate and slightly keeled. Neck scales between auditory meatus and shoulder, 25 (counted over postauricular and longitudinal folds). Scales between posterior border of auditory meatus and antegular fold,
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15. Antegular folds separated medially by 15 scales. Supernumerary antegular folds and oblique folds absent. Dorsal scales ovate, keeled, imbricate. Scales around midbody, 41. Eighteen rows of keeled scales on dorsum at midtrunk. Dorsal body scales grade laterally at midbody into smaller, weakly keeled, subimbricate scales. Ventral body scales smooth, imbricate, and rhomboidal. Ventral scales slightly smaller than dorsals. Dorsal scales between occiput and groin, 37. Ventral scales between mental and precloacal pores, 69. Six precloacal pores. Axilla– groin distance 20.0 mm (41.8% of SVL). Brachial and antebrachial scales keeled, mucronate, imbricate, slightly smaller than dorsals. Supracarpals laminar, round, smooth, some on lateral part of the limb keeled. Supracarpals larger, fewer in number than brachial scales. Subdigital lamellae on fingers with three blunt keels, numbering: I: 7; II: 15; III: 16; IV: 12; V: 6. Forelimb length 17.5 mm (36.6% of SVL). Claws moderately long and curved. Supradigital lamellae slightly convex, smooth, imbricate. Infracarpals and infratarsal small, strongly imbricate, keeled, shortly mucronate. Supratarsals strongly keeled, lanceolate, mucronate. Scales of ventral surfaces of thighs and tibia of the same shape and size as ventrals. Number of subdigital lamellae on toes: I: 9; II: 13; III: 18; IV: 22; V: 13. Hind limb length 28.7 mm (60.0% of SVL). Convex small granules on posterior side of forelimb and hind limb. Ventral patch of about 20–23 enlarged, smooth to keeled, mucronate scales. Tail length 30.5 mm (broken and regenerated: 21.0 1 9.5 mm). Tail quadrangular in cross section at base, becoming oval distally. Dorsal and lateral caudals symmetrically rhomboidal, keeled, slightly mucronate. Ventral caudals of first 12 rows distal to vent smooth, thereafter keeled, and mucronate. Color in life.—Ground color of dorsal surfaces brown-tan. Dorsal pattern with a tan vertebral field 2–4 scales wide at midbody. No vertebral line. Paravertebral field slender, 2–3 scales wide, slightly darker brown than vertebral field. Paravertebral markings dark brown, conspicuous, 10–11
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markings present from occiput to proximal caudal annuli of tail; each marking is formed anteriorly by a more-or-less crescent-shaped gray-brown area, partly enclosing an irregular creamy white spot posteriorly. Two–three pairs of paravertebral markings on neck, nine on body, and three on tail base. Each paravertebral marking longitudinally separated from each other in front and behind by 2–4 scales. Dorsolateral stripe creamy white, narrow, 1–1.5 scales wide. Lateral field with a series of irregular crescent markings. Lateral line irregular with scattered black and white scales. Ventrolateral field with an irregular composite of white, black, and brown scales. Lateral line continues anteriorly beyond shoulder, extending above longitudinal and postauricular folds of neck, across temporal region, and terminates at postocular scale. Two black, enlarged, prescapular and postscapular spots connected by a line of black scales. Along side of the body and anterior portion of tail, scattered scales of iridescent pale blue (only observed in holotype). Top of head tanbrown but darker than body dorsal surface color, with scattered white scales. Sides of the head white with scattered black or brown scales. A narrow black line present along canthal ridge between ciliary and nasal scales. Supralabials with small black, irregular spots along posterior margin, spotted in central part of each scale. Infralabials and adjacent lateral gulars spotted, gray-black. Dorsal and lateral surfaces of forelimbs and hind limbs gray with an irregular mix of black and white scales. Posterior rows of infrafemoral scales with yellow margins. Ventral surface of body, head, neck, limb, and tail immaculate white. Precloacal pores orange. Color in preservative.—General coloration in ethanol is the same as in life, but less vibrant after 4 yr; the blue lateral scale coloration has become less brilliant. Variation in paratypes.—Variation in meristic and morphometric traits is presented in Table 1. Keel on internasal, frontonasal, and prefrontals scales varies from indistinct to distinct. Some supralabials slightly keeled in some lizards (e.g., UNNEC 5796). Interparietal scale usually ir-
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TABLE 1.—Variation in squamation and morphometrical characteristics of paratypes of Liolaemus azarai. Measurements are in mm. Numbers for each variable indicate mean 6 standard deviation, with range in parentheses. Scale counts are shown as range. * 5 scales between the upper anterior corner of auditory meatus and posterior margin of orbit. ** 5 number of scales counting between auditory meatus and shoulder. Males (n 5 7)
Variable
SVL Head length Head width Foreleg Hindleg Axilla–groin distance Midbody scales Dorsal head scales Internasal scales Frontonasal scales Prefrontal scales Infralabial scales Tympanum-orbit scales (*) Neck scales (**)
44.9 10.3 4.8 26.9 17.1 19.1
6 6 6 6 6 6
2.7 (41.2–48.3) 0.5 (9.6–11.1) 0.5 (4.2–5.6) 1.7 (25.3–30.4) 0.7 (16.4–18.6) 1.3 (18.0–22.0) 37–41 19–22 8–10 5–9 9–10 4–5 9–11 23–26
regularly shaped, bordered by 6–8 scales. Scales between posterior border of auditory meatus and antegular fold range, 14– 17. Upper temporals slightly to strongly keeled, lower temporals smooth to slightly keeled. Parietals smooth, slightly keeled or slightly rugose. Supraoculars smooth to slightly rugose, and some with a small keel. Supraorbitals smooth to slightly keeled. Loreal and lorilabials convex to keeled.
FIG. 3.—Known distribution of Liolaemus azarai. Black arrow: type locality: Isla Yacyreta´, Paraguay. Black dots: localities in Corrientes Province, Argentina (1) Curuzu´ Laurel, Departamento San Miguel and (2) San Roque, Departamento San Roque.
Females (n 5 7)
48.7 10.0 4.5 17.2 26.3 21.3
6 6 6 6 6 6
2.7 (33.3–48.7) 0.5 (9.4–10.8) 0.3 (4.2–5.1) 1.2 (15.0–18.7) 1.0 (24.6–27.7) 2.1 (19.0–23.4) 32–42 18–23 8–10 5–8 9–10 4–5 9–12 23–27
Paravertebral markings vary from a light to dark brown. In some lizards, lateral field markings and lateral field merge into a mix of black, brown, and yellow-white scales. Dark spot numbers in lateral head scales vary, and in some lizards are very scarce. Only the holotype was observed alive and in good condition. Small blue scales were observed in the living holotype, but not in other preserved males. Sexual dimorphism.—Females lack the prescapular and postscapular black spots and iridiscent pale blue scales on the sides of body observed (only) in the male holotype. Ventral coloration is similar between males and females. The base of the tail of males is expanded laterally, and the precloacal pores are larger and obvious relative to those seen in females. Distribution.—Liolaemus azarai is known only from the type locality in Paraguay and two localities in Argentina: Curuzu´ Laurel, San Miguel Department (278 509 S, 578 309 W) and San Roque, San Roque Department (288 349 S, 588 429 W); both are isolated sand dune formations in northeastern Corrientes Province (Fig. 3). The distribution of L. wiegmannii in Corrientes Province (see figure 3 in Etheridge, 2000) corresponds to L. azarai. Natural history.—Liolaemus azarai is known from isolated sand dunes in the phytogeographic formation known as Sector Parque Mesopota´mico, Subdistrito
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Correntino, Distrito Oriental Chaquen˜o, Provincia Chaquen˜a (Carnevali, 1994). According to Tedesco et al. (1992), the dominant plant community on the dunes where the lizards were found at Curuzu´ Laurel is a low forest surrounding the dunes and composed of Prosopis algarrobilla, Acacia caven, and Celtis spinosa. On the sand dunes, the plant community is composed principally of shrubs and small trees of Erythroxylum cuneifolium, Ocotea acutifolia, Smilax campestris, Eugenia hyemalis, Peschiera australis var. hilariana, Elionurus muticus, and Butia paraguayensis. Active lizards were observed from morning to afternoon, and individuals encountered in the open ran toward surrounding vegetation or into burrows (Tedesco et al., 1992). Liolaemus azarai is insectivorous and oviparous (Tedesco et al., 1992). The data regarding number and characteristics of their chromosomes are under investigation (A. Hernando, personal communication). Phylogenetic affinities.—Liolaemus azarai is the most northeasterly distributed species of the wiegmannii group in Argentina. Initially, this species was confused with L. wiegmannii, to which, based on morphological similarity, it may be closely related. Liolaemus azarai is apparently endemic to the sand dunes of Corrientes Province and some islands in the Parana River. These dunes are Quaternary in origin, and these isolated formations with their herbaceous and shrubby vegetation occur only in a narrow zone in central Corrientes Province (Iriondo and Garcia, 1993). These habitats are threatened by human activities and natural catastrophic events (flooding). The lack of previous records does not allow determination of trends in population size of L. azarai. It is likely that portions of this population on Yacyreta´ Island were destroyed when the Binational Yacyreta´ Dam was erected. In fact, the type locality is a northeastern remnant of Yacyreta´ Island, which now lies partially under Yacyreta Lake. Consequently, studies on the distribution of this species are urgently needed to establish its status and possibly to prevent its extinction or local
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extirpation. Other dune formations in southern Corrientes and Entre Rios provinces have low density populations of species belonging to the wiegmannii group, but their taxonomic status is unclear because only photographs and few poorly preserved specimens were available for examination. RESUMEN Liolaemus azarai es descripto como una nueva especie de lagartija liolaemida del nordeste de Argentina y la isla Yacyreta, Paraguay. Es un miembro del grupo wiegmannii, y puede ser distinguida del resto de las especies del grupo por su menor taman˜o, bajo nu´mero de escamas alrededor del cuerpo y el patro´n de coloracio´n dorsal. En la localidad tipo, en la Isla Yacyreta, L. azarai esta limitada a formaciones aisladas de dunas. Esta especie es conocida solamente en otras dos localidades de la provincia de Corrientes, nordeste de Argentina. La nueva especie es la ma´s pequen˜a del grupo wiegmannii, y al igual que otros miembros del grupo es una lagartija exclusivamente arenı´cola, insectı´vora y ovı´para. Acknowledgments.—I thank F. Lobo for critically reading the early draft of this work; J. Sites and M. Morando for useful comments that improved the manuscript in the English version; M. Archangelsky and J. Marshall for assistance with an early translation; R. Etheridge for important suggestions about the last version of the manuscript; B. Alvarez de Avanza (UNNEC), R. Martori (ZV-UNRC), S. Kretschmar and G. Scrocchi (FML), J. Williams (MLP-S), and G. Carrizo (MACN) for the loan of specimens or/and permission to examine specimens at their institutions; B. Alvarez de Avanza, M. E. Tedesco, R. Aguirre, M. Lions (UNNEC), and S. Tiranti (UNLPam) for helpful comments on ecological and biological data; and A. Hernando (UNNEC) for helpful cytogenetic information.
LITERATURE CITED ACHAVAL, F., AND A. OLMOS. 1997. Anfibios y Reptiles del Uruguay. Serie Fauna Nu´mero 1:1–128. Montevideo, Uruguay. ALVAREZ, B., AND M. L. LIONS. 1996. Geographic distribution: Liolaemus wiegmannii. Herpetological Review 27:32. CARNEVALI, R. 1994. Fitogeografı´a de la Provincia de Corrientes. Gobierno de la Provincia de Corrientes—Instituto Nacional de Tecnologı´a Agropecuaria. Corrientes, Argentina. CEI, J. M. 1979. A reassessment of the genus Cte-
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noblepharis (Reptilia, Sauria, Iguanidae) with description of a new subspecies of Liolaemus multimaculatus from western Argentina. Journal of Herpetology 13:297–302. . 1986. Reptiles del centro, centro-oeste y sur de la Argentina. Museo Regionale di Scienze Naturali, Torino. Monografie 4:1–527. . 1993. Reptiles del noroeste, nordeste y este de la Argentina. Museo Regionale di Scienze Naturali, Torino. Monografie 14:1–929. DONOSO-BARROS, R. 1966. Reptiles de Chile. Ediciones Universidad de Chile, Santiago, Chile. ETHERIDGE, R. 1992. A new psammophilus lizard of the genus Liolaemus (Squamata: Tropiduridae) from northwestern Argentina. Bolletino del Museo Regionale di Scienze Naturali, Torino 10:1–19. . 1993. Lizards of the Liolaemus darwinii complex (Squamata: Iguania: Tropiduridae) in northern Argentina. Bolletino del Museo Regionale di Scienze Naturali, Torino 11:137–199. . 1995. Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates 3142:1–34. . 2000. A review of lizards of the Liolaemus wiegmannii group (Squamata, Iguania, Tropiduridae), and a history of morphological change in the sand-dwelling species. Herpetological Monographs 14:293–352. FROST, D. 1992. Phylogenetic analysis and taxonomy of the Tropidurus group of lizards (Iguania, Tropiduridae). American Museum Novitates 3033:1– 68. HALLOY, M., R. ETHERIDGE, AND G. M. BURGHARDT. 1998. To bury in sand: phylogenetic relationships among lizard species of the boulengeri group, Liolaemus (Reptilia: Squamata: Tropiduridae), based on behavioral characters. Herpetological Monographs 12:1–37. IRIONDO, M. H., AND N. O. GARCIA. 1993. Climatic variations in the Argentine plains during the last 18,000 years. Palaeogeography, Palaeoclimatology, Palaeoecology 101:209–220. LAURENT, R. F. 1983. Contribucio´n al conocimiento de la estructura taxono´mica del ge´nero Liolaemus Wiegmann (Iguanidae). Boletı´n de la Asociacio´n Herpetolo´gica Argentina 1:16–18. . 1984. On some iguanid genera related to or previously confused with Liolaemus Wiegmann. Journal of Herpetology 18:357–373. LOBO, F., AND R. E. ESPINOZA. 1999. Two new cryptic species of Liolaemus (Iguania: Tropiduridae) from northwestern Argentina: resolution of the purported reproductive bimodality of Liolaemus alticolor. Copeia 1999:122–140. SCHULTE, J. A., J. R. MACEY, R. E. ESPINOZA, AND A. LARSON. 2000. Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariances and dispersal. Biological Journal of the Linnean Society 69:75–102. TEDESCO, M. E., A. HERNANDO, AND B. B. ALVAREZ. 1992. Hallazgo de Liolaemus wiegmanni (Dume´ril y Bibron 1837) (Iguania, Liolaemidae) en la pro-
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vincia de Corrientes (Repu´blica Argentina). Facena 9:117–122. Accepted: 9 October 2002 Associate Editor: Joseph Mendelson III
APPENDIX I Specimens Examined Liolaemus azarai (n 5 15): PARAGUAY: MISIONES: Yacyreta´ Island, Parana´ River: UNNEC 5796– 97, 5892–95. ARGENTINA: CORRIENTES: San Miguel Department: Curuzu´ Laurel (278 509 S, 578 309 W): UNNEC 562, 582–83, 585, 4332–35. San Roque Department: San Roque (288 349 S, 588 429 W): ZV-UNRC 2218. Liolaemus wiegmannii (n 5 260): ARGENTINA: BUENOS AIRES: Partido Bahı´a Blanca: Bahı´a Blanca (388 449 S, 628 169 W): MACN 1670–72; Ingeniero White (388 479 S, 628 169 W): MACN 24820. Partido Coronel Dorrego: Farola de Monte Hermoso, Monte Hermoso (388 599 S, 618 179 W): CENAI 850. Partido General Lavalle: Mar de Ajo (368 439 S, 568 419 W). MACN 25234; Santa Teresita (368 339 S, 568 419 W): CENAI 188; Laguna Chasico´, Campo de los Guijarros: MACN 31606–07. Partido Guaminı´: Laguna Alsina (Estacio´n Bonifacio) (368 499 S, 628 159 W): MACN 20938–40. Partido Patagones: Carmen de Patagones (408 489 S, 638 009 W): MACN 15191, 31620– 22, 31762, 31876; Estancia El Me´dano, Carmen de Patagones (408 489 S, 638 009 W): MACN 31876; Bahı´a San Blas (408 339 S, 628 169 W): MACN 6406. CATAMARCA: Andalgala Department: Agua de las Palomas (278 379 S, 668 079 W): LJAMM 4299–300. La Paz Departament: Ican˜o (288 559 S, 658 199 W), Escuela Hogar Nro. 4—Ministerio de Educacio´n: CENAI 117. CORDOBA: Punilla Departament: Los Cocos (308 569 S, 648 309 W): MACN 8919–22; Cruz Grande (308 569 S, 648 299 W): MACN 21443; La Cumbre (308 599 S, 648 309 W): MACN 22384–85, 24158–61, 31611–17; 2 km NE La Cumbre (308 599 S, 648 309 W). MACN 24634–36. Rı´o Cuarto Departament: Rı´o Cuarto (338 089 S, 648 229 W): ZV-UNRC 828, 1602; 20 km W Chaja´n (338 339 S, 658 019 W): ZV-UNRC 3638; Coronel Moldes (338 389 S, 648 339 W): ZV-UNRC 3644, 3690; Laguna Oscura (338 539 S, 648 529 W): ZV-UNRC 2475, 2478, 2504, 3074, 3082–84, 3086, 3090–91, 3093, 3097, 3123, 3125, 3129, 3131–32, 3139–40, 3154, 3159, 3165–66, 3169, 3175, 3177, 3180–81, 3186, 3207, 3209, 3214, 3217– 19, 3224, 3229, 3231, 3233, 3235–36, 3238–40, 3242– 47, 3249, 3250, 3252–58, 3262–70, 3273–78, 3280– 88, 3453–59, 3493, 3497. San Javier Departament: Tilquicho (328 119 S, 658 139 W): ZV-UNRC 299, 301. ENTRE RIOS: Islas del Ibicuy Departament: Me´danos (328 259 S, 598 049 W): CENAI 856, 858, 868. LA PAMPA: ‘‘Pampa Central’’: MACN 1474–80, 1482. Capital Departament: Colonia Ine´s y Carlota (368 189 S, 638 559 W): MACN 23257–59; airport, 3 km NE Santa Rosa (368 379 S, 648 179 W): MACN 23281, 23284; Santa Rosa (368 379 S, 648 179 W): CENAI 58(2). Catrilo´ Departament: Lonquimay (368 289 S, 638 379 W): MACN 31187–96; 3 km SW Uriburu (368 309 S, 638 519 W): MACN 31197–205. Conelo Departament: Conelo (368 009 S, 648 369 W):
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MACN 6635, 6911–12, 6698. Guatrache´ Departament: ‘‘Don Amelio’’, Peru´ (378 409 S, 648 069 W): MACN 22035–36. Hucal Departament: Abramo (378 549 S, 638 519 W): MACN 34440. Loventue´ Departament: Loventue´ (368 119 S, 658 179 W): MACN 7193–94; Las Vertientes, 35 km SW Carro Quemado (368 289 S, 658 219 W): MACN 23263–65; Victorica (368 129 S, 658 279 W): CENAI 851. Lihue´ Calel Departament: Lihue´ Calel (388 029 S, 658 339 W): CENAI 840; Lihue´ Calel, (388 029 S, 658 339 W): MACN 35354. Mara´ Co Departament: General Pico (358 409 S, 638 469 W): MACN 30960–61. MENDOZA: La Paz Department: National Highway 146, km 276, 74 km NE Monte Coman (348 179 S, 678 149 W). LJAMM 4039–44. SAN JUAN: San Martı´n Departament: San Martı´n, Calle Nacional s/n (318 319 S, 688 249 W): IMCN-UNSJ 103, 104. Angaco Departament: Angaco, Velazquez Street s/n, intersection of provincial routes 59 and 230 (318 289 S, 688 209 W): IMCN-UNSJ 105–07. SAN LUIS: Chacabuco Departament: 7 km N Naschel (328 569 S, 658 289 W) on National Highway 148: ZV-UNRC 662. Departamento Coronel Pringles: Saladillo (Rı´o Quinto) (338 129 S, 658 519 W): CENAI 839. La Toma (338 049 S, 658 389 W): ZV-UNRC 1483–84, 4064. Departamento General Pedernera: Estancia La Emboscada, El Morro (338 139 S, 658 299 W): MACN 10209(2); El Morro (338 139 S, 658 299 W): MACN 25055; 20 km W La Punilla (338 099 S, 658 059 W): ZV-UNRC 3655; Juan Jorba (338 379 S, 658 169 W): ZV-UNRC 3693; La Punilla (338 099 S, 668 059 W): ZV-UNRC 4098– 99. Gobernador Dupuy Departament: Bagual (338 099 S, 658 359 W): MACN 15265. SANTA FE: Garay Departament: Colonia Mascias (Isla Garay) (308 479 S, 608 009 W): MACN 7846–47. TUCUMA´N: Burruyacu´ Departament: Agua Negra, Sierra de Medina: FML 769(4), 780, 823(6), 2393(8), 2393(2), 2571(2),
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2577, 2704(3), 2721(7), 2801(3). BOLIVIA: SANTA CRUZ: Nueva Moka: MACN 15233. Liolaemus lutzae (n 5 9): BRASIL: ‘‘Brasil,’’ CENAI (o/n). RIO DE JANEIRO: Cabo Frı´o: FML 1287(7), MLP-S 920–21. Liolaemus multimaculatus (n 5 50): ARGENTINA: BUENOS AIRES: Partido Claromeco´: FML 1596(10). Partido Coronel Dorrego: MLP-S 137, 141–46, 155, 372. Partido General Alvarado: Miramar: MLP-S 138–39, 486–503. Partido General Lavalle: Las Toninas: MACN 33491–92. Lucila del Mar: MACN 32419. Partido San Antonio Norte: Cabo San Antonio: MACN 29705–08, 29714. Municipio Urbano Monte Hermoso: MLP-S 141–46, 149, 155, 157, 370–72. RIO NEGRO: Adolfo Alsina Department: Bahia Creek (418 049 S, 638 579 W): LJAMM 2559. Liolaemus scapularis (n 5 68): ARGENTINA: CATAMARCA: Santa Maria Department: Campo Los Pozuelos, National Road 40, between km 894 and 904: FML 2022(18), 5 km E Santa Maria: FML: 2932(10). SALTA: Cafayate Department: Cafayate: Los Me´danos: FML 1819(4), 1823(10), 2203(11), 2221(4), 2232(11). Liolaemus occipitalis (n 5 3): BRASIL: ‘‘Brasil,’’ CENAI (o/n). BRASIL: Rı´o Grande do Sul: Tramandaı´: Praia Jardim Atlantico: FML 2618, 2620. Liolaemus riojanus (n 5 32): ARGENTINA: LA RIOJA: ‘‘La Rioja’’: MLP-S 995. Independencia Department: 110.7 km S Villa Unio´n, on Ruta Provincial 26: FML 3005(18). San Martin Department: 5 km N Baldecito: 2626(4), 2378(1), 2876(8). Liolaemus salinicola (n 5 37): ARGENTINA: CATAMARCA: Tinogasta Department: road between Taton and Medanitos: FML 8441–42. Medanitos: FML 1886–87, 1907(4), 1909(10), 2079, 2101(5). 10 km N Taton (278 289 S, 678 359 W): LJAMM 2347– 54, 2373–75. 4 km N Palo Blanco (278 179 S, 678 459 W): LJAMM 2379–80.
DATE OF PUBLICATION Herpetologica, Vol. 59, No. 1, was mailed 25 February 2003.
