two new species of liolaemus lizards from northwestern argentina

89

Herpetologica, 59(1), 2003, 89–105q 2003 by The Herpetologists’ League, Inc.

TWO NEW SPECIES OF LIOLAEMUS LIZARDS FROMNORTHWESTERN ARGENTINA: SPECIATION WITHIN THE

NORTHERN SUBCLADE OF THE ELONGATUS GROUP(IGUANIA: LIOLAEMIDAE)

ROBERT E. ESPINOZA1,3,4 AND FERNANDO LOBO2

1Ecology, Evolution, and Conservation Biology and Biological Resources Research Center,University of Nevada, Reno, Reno, NV 89557, USA; and Section of Amphibians and Reptiles,Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, PA 15213-4080, USA

2Facultad de Ciencias Naturales, Universidad Nacional de Salta, Avenida Bolivia 5150,Salta 4400, Argentina

ABSTRACT: The Liolaemus elongatus group currently includes eight species (including the spe-cies described herein) of mid-sized, long-tailed, primarily saxicolous lizards. Members of this lineageare distributed in northwestern Argentina along the Andean Cordillera south to the PatagonianSteppe (27–448 S) and over a wide range of elevations (350–3900 m). Here we describe two newspecies belonging to the northern radiation of the elongatus group. The four currently recognizedmembers of this northern subclade (including the two species described herein) have fewer midbodyscales than other members of the elongatus group, brown to black heads, and lack the distinct dorsalbody or tail patterns found in adults of other species belonging to the elongatus group. Additionally,pregnant and recently spent females exhibit crimson red color in the cloacal region—a trait knownonly for members of this northern lineage and L. buergeri. The first new species, L. dicktracyi fromthe Sierra de Famatina of La Rioja Province, differs from all other members of the elongatus groupin several meristic characters and in its unique dorsal coloration: black heads, shoulders, and tails,with blue torsos and hind limbs. The second new species, L. umbrifer from the Quebrada deRandolfo in northeastern Catamarca Province, is morphologically similar to L. capillitas, but differsfrom this and other species in the elongatus group in several meristic characters, the presence ofblack shoulders and upper arms, and crimson red coloration of the cloacal region in both malesand females. The natural history of both of the new species is similar in many respects to L. capillitasand L. heliodermis, which further supports the phylogenetic affinities of members of this subcladewithin the elongatus group. Finally, we provide comparisons of recent phylogenetic hypotheses anddiscuss the content of the elongatus group.

Key words: Argentina; Liolaemidae; Liolaemus dicktracyi sp. nov.; Liolaemus elongatus group;Liolaemus umbrifer sp. nov.; Lizard; New species; Taxonomy

THROUGHOUT the arid and semiarid re-gions of South America, Liolaemus lizardsreign as the dominant components of mostreptile communities. Members of thisclade are widely distributed from the highAndes of central Peru to the shores of Ti-erra del Fuego and over a broad range ofclimates and habitats (Cei, 1986, 1993;Donoso-Barros, 1966; Etheridge and Es-pinoza, 2000). Several species of Liolae-mus even hold records among squamatesfor their latitudinal (548 S; L. magellani-cus) or elevational distributions ($5000 m;L. irregularis, L. nigriceps, and L. signifer)

3 PRESENT ADDRESS: Department of Biology, Cal-ifornia State University, Northridge, Northridge, CA91330-8303, USA.

4 CORRESPONDENCE: e-mail, [email protected]

(R. E. Espinoza, unpublished data). Thisremarkably large genus of lizards—secondonly to Anolis in number of species—in-cludes .160 recognized taxa and has con-tinued to grow at a rate of nearly four newspecies per year over the last decade (Eth-eridge and Espinoza, 2000).

Recent phylogenetic analyses of mito-chondrial DNA sequences for 60 speciesof Liolaemus (Schulte et al., 2000) indicatea well supported basal split dividing thelineage into two, similarly sized clades thatlargely correspond to groups defined pre-viously from morphological evidence (Eth-eridge, 1995). One of these lineages, thechiliensis group (sensu Etheridge, 1995;Lobo, 2001; 5 subgenus Liolaemus sensuLaurent, 1985; see also Schulte et al.,2000), includes approximately 75 species

90 [Vol. 59, No. 1HERPETOLOGICA

TABLE 1.—Maximum body sizes (SVL) and geographic distributions of the eight Liolaemus species currentlyrecognized as belonging to the elongatus group (including the two new species described herein). Data forL. thermarum are from Videla and Cei (1996) and Espinoza et al. (2000) with the inclusion of a northernrange extension for L. elongatus (Avila and Lobo, 1999) and new elevation records for L. petrophilus. The

four species belonging to the northern subclade of the elongatus groups are indicated with an asterisk.

SpeciesMaximumSVL (mm)

Latitude range(degrees S)

Elevation range(m)

Liolaemus austromendocinusL. capillitas∗L. dicktracyi sp. nov.∗L. elongatus1

L. heliodermis∗L. petrophilusL. thermarumL. umbrifer sp. nov.∗

10393918981

1008589

34–372729

28–4527

41–443527

900–23102500–39002600–2800

700–30002820

350–14002400–25003190–3490

1 See Discussion for comments on the distribution of L. elongatus.

that share two derived morphological char-acteristics: a fused Meckel’s groove andfour or fewer precloacal pores. Within thechiliensis group, several informal groupnames have been applied to morphologi-cally similar species (Cei, 1986, 1993; re-viewed in Lobo, 2001). One of these, theelongatus group as originally defined byCei (1974; see also Cei, 1975, 1986; Hulse,1979), was recently diagnosed and rede-fined to include six species: L. austromen-docinus, L. capillitas, L. elongatus, L. he-liodermis, L. petrophilus, and L. therma-rum (Espinoza et al., 2000).

The six currently recognized species ofthe elongatus group are distinguished fromother species in the chiliensis group bytheir moderate to large body size (approx-imately 80–100 mm snout–vent length:SVL); moderately slender to robust, elon-gate bodies; small, nonoverlapping to sub-imbricate body scales (.55 around mid-body) that do not terminate in a spine; andtails that are longer than 1.5 times the SVLand round in cross section (Espinoza et al.,2000). Members of this lineage are distrib-uted from near the intersection of theCumbres Calchaquıes and the Sierra delAconquija of western Tucuman Province(278 S), southward to the PatagonianSteppe of Chubut Province (448 S), andover a wide range of elevations (350–3900m; Table 1). Species in the elongatusgroup usually inhabit arid environments,including the eco-regions known as Pre-puna, Patagonian Forests, and PatagonianSteppe (Burkart et al., 1999; Cabrera and

Willink, 1980). Most are moderately to ex-clusively saxicolous, insectivorous to om-nivorous, and all species for which repro-ductive mode has been determined are vi-viparous (Cei, 1974, 1986; Espinoza et al.,2000; Hulse, 1979; Videla, 1983).

Within the elongatus group are twoclosely related, geographically isolated spe-cies, L. capillitas (Hulse, 1979) and L. he-liodermis (Espinoza et al., 2000), whichare distributed far north of most other spe-cies in the elongatus group, with the ex-ception of a few disjunct northern popu-lations of L. elongatus (Avila and Lobo,1999; but see Discussion). We refer to thesubclade including L. capillitas and L. he-liodermis as the northern radiation of theelongatus group. Members of this subcla-de, which currently includes the two spe-cies listed above plus the two new speciesdescribed below, differ from other speciesin the elongatus group in the followingways: usually fewer midbody scales 58–76(versus 71–90 in other species of thegroup), a uniformly brown to black head(dorsal head is irregularly marked in otherspecies of the group), a torso and tail thatlack distinct patterns (no stripes, bars, orrings) in adults, a darkly pigmented (usu-ally black) area surrounding the insertionof the forelimbs that is speckled with lightcolored spots, and crimson red colorationin the cloacal region of pregnant and re-cently spent females (Table 2). Here wedescribe two new species, one from LaRioja Province and another from Cata-

March 2003] 91HERPETOLOGICA

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FIG. 1.—Holotype of Liolaemus dicktracyi (FML9928; SVL 5 88.4 mm).

marca Province, that belong to this north-ern radiation of the elongatus group.

MATERIALS AND METHODS

For purposes of diagnosing the newspecies and defining the northern clade ofthe elongatus group, we examined a seriesof each of the six species currently consid-ered to be members of the elongatusgroup (sensu Espinoza et al., 2000; Table1; Appendix I). Specimens examined, in-cluding the new species described herein,are deposited in the herpetological collec-tions of the Museo Ciencias Naturales(MCN); Universidad Nacional de Salta,Argentina; the Carnegie Museum of Nat-ural History (CM), Pittsburgh, Pennsylva-nia, USA; the Fundacion Miguel Lillo(FML), Tucuman, Argentina; and the Mu-seum of Vertebrate Zoology (MVZ), Uni-versity of California, Berkeley, California,USA. All specimens examined had beencollected either at their type localities orwithin 100 km thereof. In our taxonomicanalysis, we considered external morphol-ogy, including squamation, coloration, andcolor patterns. We examined live speci-mens of all species belonging to the elon-gatus group to record color in life (exceptfor L. thermarum for which we used in-formation in Videla and Cei, 1996). AScouty global-positioning device (TrimbleNavigation Systems, Sunnyvale, California,USA) was used to record the latitude, lon-gitude, and elevation of lizards collected inthe field. Additional specimens were ex-amined after fixation in 10% formalin andpreservation in 70% ethanol. Some char-acter states were determined with the aidof a binocular dissecting microscope (10–403). Measurements were taken with dig-ital calipers to the nearest 0.01 mm. Termsfor the description of scales are those ofSmith (1946); the distribution of body pat-terns is after Lobo and Espinoza (1999);and for neck-fold terminology we followFrost (1992).

SPECIES DESCRIPTION

Liolaemus dicktracyi sp. nov.Holotype.—FML 9928 (field tag: REE

184), an adult male from Portezuelo Blan-co, 26.5 km west of Famatina on road to

Cueva de Perez, Sierra de Famatina, De-partamento Famatina, Provincia de LaRioja, Argentina (288 549 300 S, 678 419 300W; 2800 m); collected by F. B. Cruz andR. E. Espinoza, 16–17 February 1998(Fig. 1).

Paratypes.—CM 147716–17 (field tags:PT 3291; REE 201), FML 9929–33 (fieldtags: PT 3286–89, 3292), and MCN 461–62 (field tags: PT 3285, 3290) same dataas holotype.

Diagnosis.—A moderate to large bod-ied, robust lizard belonging to the Liolae-mus elongatus group (sensu Espinoza etal., 2000) that can be distinguished fromall other members of that group (and allother Liolaemus) in possessing the follow-ing characteristics: black heads (reviewedby Cei, 1998) and shoulders, indigo tolight blue torsos, and charcoal gray toblack ventral coloration (Table 2; Fig. 2).Liolaemus dicktracyi also differs from oth-er members of the elongatus group, exceptthose in the northern radiation (L. capil-litas, L. heliodermis, and L. umbrifer sp.nov., see description below), in possessing


FIG. 2.—Upper: adult male Liolaemus dicktracyiin life; lower: adult male Liolaemus umbrifer in life.

larger and, therefore, usually fewer mid-body scales (Table 2): L. dicktracyi: 60–73;versus 76–82 in L. austromendocinus; 72–90 in L. elongatus; 74–89 in L. petrophilus(Donoso-Barros and Cei, 1971); 84–89 inL. thermarum (Videla and Cei, 1996), andpresence of red coloration in the cloacalregion of pregnant and recently spent fe-males. Liolaemus capillitas, L. elongatus,and L. petrophilus have dorsal scales thatare more distinctly imbricate than those ofL. dicktracyi. Liolaemus elongatus havedark brown, irregular, longitudinal stripes(6–12 scales wide) that extend along thevertebral and lateral fields, whereas L.dicktracyi lack stripes or similar body pat-terns (Table 2). Liolaemus petrophilushave a distinct dorsal pattern of transversedark bars that form a ‘‘tigroid’’ pattern(Cei, 1975; Donoso-Barros and Cei, 1971)that is lacking in the new species. UnlikeL. dicktracyi, L. elongatus (southern pop-ulations; see Discussion), L. petrophilus,and most L. austromendocinus have dis-

tinctly ringed tails (Table 2). The tempo-rals of L. umbrifer sp. nov. (described be-low) are only slightly keeled; the head isdark brown, not black; and both males andfemales of this species have red colorationin the cloacal region. In L. dicktracyi, thetemporals are usually distinctly keeled, thehead is always black, and only females pos-sess red coloration in the cloacal region.In contrast to L. dicktracyi, which lackdorsal body patterns and possess 3–4 pre-cloacal pores, L. thermarum have dark,wide stripes on the lower flanks and lackprecloacal pores (Videla and Cei, 1996,1998).

Description of the holotype.—Adultmale (Fig. 1), 88.4 mm SVL; tail length152.1 mm, complete, not regenerated. Ax-illa–groin distance 39.6 mm. Head 19.6mm long (from anterior border of auditorymeatus to tip of snout), 15.4 mm wide (atanterior border of auditory meatus), 9.0mm high. Snout length 6.8 mm (posteriormargin of canthal to tip of snout). Inter-orbital distance (between arcs of circu-morbitals) 1.4 mm. Eye–nostril distance4.3 mm. Tibial length 19.4 mm. Footlength 25.2 mm (ankle to tip of claw onfourth toe).

Dorsal head scales smooth, 17 betweenrostral and anterior border of auditory me-atus. Ten keeled temporals (left side), allbut those closest to auditory meatus with1–2 scale organs along their posterior mar-gins. Interparietal subpentagonal, smallerthan parietals in size, surrounded by sevenscales. Frontal scale azygous. Six scales be-tween frontal and rostral. Two postrostralswith 8–9 scale organs each. Circumorbitalscomplete. Five enlarged supraoculars(both sides). Five scales between frontaland supercilliaries. Six/seven (left/right)flat, elongate, imbricate supercilliaries.Canthal separated from nasal by two scales(both sides). Loreal region flat. Sevenscales surrounding nasals (both sides). Na-sals not in contact with rostral. Seven/eightlorilabials, fourth through seventh/sixththrough eighth in contact with subocular.Six/five enlarged supralabials. Fourth su-pralabial (both sides) curved upward pos-teriorly but not in contact with subocular.Five infralabials (both sides), slightly taller


than supralabials. Four internasals. Orbitwith 15 upper and 13 lower ciliaries (leftside). Orbit diameter 3.6 mm (measuredbetween upper and lower ciliaries on leftside). Subocular scale elongate. Preocularunfragmented. Longitudinal ridge alongupper margin of the three ocular scales.Rostral scale twice as wide (3.9 mm) ashigh (1.9 mm). Mental twice as wide (3.8mm) as high (1.8 mm), followed posteri-orly by two rows of 5/4 chinshields. Firstchinshield in contact with first infralabial.Scales of throat between chinshields jux-taposed in first two rows, imbricate pos-teriorly. Fifty-seven gulars between audi-tory meatus. Two outward projecting lam-inar scales along anterior border of audi-tory meatus. Auditory meatus higher (4.3mm) than wide (2.9 mm). Lateral scales ofneck granular. Subdermal fat bodies giveslightly inflated appearance to neck region.Antehumeral fold distinct. Rictal, postau-ricular, and longitudinal folds present butless conspicuous than antehumeral. Thirty-six scales between auditory meatus and an-tehumeral fold (counted along postauric-ular and longitudinal folds).

Fifty-eight dorsal scales between occi-put and anterior surface of thighs. Dorsalbody scales with round posterior margin,slightly imbricate, weakly keeled. Twenty-five longitudinal keeled scale rows overdorsum of trunk. Scales of neck regionsmaller than dorsals. Sixty-five scalesaround midbody. Ventral scales of similarsize to dorsals, 110 between mental andprecloacal pores. Three precloacal pores,each with a waxy exudate. Ventral surfaceof thighs with enlarged, laminar, imbricatescales anteriorly, abruptly changing tosmaller granular scales on posterior thirdof ventral thigh.

Fourth finger with 23 keeled, tridentate,subdigital lamellae. Claw of fourth finger1.8 mm, curved, sharp tipped, opaquebrown. Fourth toe with 28 keeled subdi-gital lamellae. Claw of fourth toe 2.2 mm,similar to that of fourth finger.

Variation.—Based on an additional sev-en adult males and two adult females. SVL72.0–91.2 mm. Head length 16.0–20.1mm, width 12.4–16.9 mm. Axilla–groindistance 31.0–45.1 mm. Tail length 138.3–

152.1 mm. Midbody scales 60–73. Dorsalscales round to rhomboidal, occasionallypointed, weakly to distinctly keeled, sub-imbricate, 56–66 between occiput and an-terior surface of thighs. Some individualswith interstitial granules along posteriorflanks. Dorsal head scales 12–17, variablein size but usually small, some irregular inshape, most convex. Ventrals 104–113.Scales around interparietal 5–8. Three tosix enlarged supraoculars. Preocular notdivided, not fused to subocular. Temporals8–10, weakly to distinctly keeled. Scalesbetween auditory meatus and antehumeralfold 30–36. Gulars 48–57, smooth, imbri-cate, circular to elliptical. Supralabials 5–7. Infralabials 4–6. Posterior tip of fourthor fifth through seventh supralabial up-turned. Scales around nasals 6–8. Nasals inslight (60%) or lacking contact (40%) withrostral. Four internasals. Rostral barely vis-ible from above. Scales between rostraland frontal 5–7. Postrostrals 2 (90%) to 3(10%), with 5–13 scale organs each. Six tonine lorilabials, fourth, fifth, or sixththrough ninth in contact with subocular.Supercilliaries 6–8. Subdigital lamellae onfourth finger 20–24; on fourth toe 26–30.Precloacal pores 3–4 in males, not presentin females.

Color in life.—Head black, abruptly fad-ing posteriorly to indigo, light blue in nu-chal region at about the level of the pos-terior border of the auditory meatus (Fig.2). Trunk (at shoulder level) to first thirdof tail indigo to light blue, most uniformin coloration in the mid-vertebral field.Light blue and black scales occasionallyspeckle the dorsal trunk, but these mark-ings are most abundant in the dorsolateralfields. Flanks slightly darker than dorsalfield. Ventrolateral field speckled with ir-regular light blue spots in most specimens.Insertion of forelimbs and antehumeraland posthumeral regions speckled withsmall blue markings (6–8 scales) on a darkindigo to black background. Tail fadesfrom indigo to light blue in anterior thirdto dark brown indigo posteriorly. Regen-erated portions of tails are black. Ventrallycharcoal gray to black. Ventral side of thetail darkens from about mid-tail to the tip.Precloacal pores yellow orange. Adult fe-


FIG. 3.—Type localities (stars) of species belongingto the northern radiation of the Liolaemus elongatusgroup in northwestern Argentina. Liolaemus heliod-ermis in Tucuman Province, L. capillitas and L. um-brifer sp. nov. in Catamarca Province, and L. dick-tracyi sp. nov. in La Rioja Province. Insert 5 Argen-tina. Some mountain ranges have been excluded forclarity.

FIG. 4.—Eroded white cliffs of Portezuelo Blanco,Sierra de Famatina, Departamento Famatina, LaRioja Province (288 549 300 S, 678 419 300 W; 2800m), the type locality of Liolaemus dicktracyi.

males had crimson red coloration in thecloacal region and at the base of the thighsand tail when collected in mid-February,otherwise sexual dichromatism is absent.

Color in preservative.—Bright blue col-or of torso in life fades and darkens tobrownish blue in alcohol and overall colordarkens slightly on trunk and ventrally.Red coloration in cloacal region of femalesfades almost completely after approxi-mately 6 mo. Precloacal pores fade fromyellow orange to cream yellow.

Etymology.—The specific name is anoun in the genitive case formed in honorof Dr. C. Richard (‘‘Dick’’) Tracy, currentlyProfessor of Biology and Director of theBiological Resources Research Center atthe University of Nevada, Reno, for hisnumerous contributions to the under-standing of the biology of lizards generallyand for inspiring, supporting, and collab-orating with us on our research with Lio-laemus specifically. With the addition of L.dicktracyi, this brings the number of Lio-laemus named after various investigators ofLiolaemus and their supporters to 60 (ap-

proximately 30% of the currently recog-nized taxa; Etheridge and Espinoza, 2000).

Distribution.—Liolaemus dicktracyi isknown only from the immediate vicinity ofthe type locality (Fig. 3): Portezuelo Blan-co, Sierra de Famatina, La Rioja (288 549300 S, 678 419 300 W; 2600–2800 m), a geo-logically well studied exposed cliff facewith strata representing Permo-Triassicand Quaternary deposits (Cei, 1982; Fig.4). Two specimens were also collectedfrom 8.4 km southeast of the type localityalong an unnamed dirt road that runsalong the Rıo Amarillo and leads to Cuevade Perez in the Sierra de Famatina. Withthe addition of L. dicktracyi, three speciesof Liolaemidae [L. famatinae and Phyma-turus mallimaccii; Cei (1980)] are nowknown to be endemic to the Sierra de Fa-matina. Currently, L. dicktracyi and L.elongatus (Avila and Lobo, 1999; but seeDiscussion) are the only species in theelongatus group known from La Rioja;however, we have examined two additionalisolated, montane populations of L. cf.capillitas from that province that also ap-pear to be undescribed species.

Natural history.—Specimens of L. dick-tracyi were found primarily on the steepand eroded cliffs and hillsides of Porte-


zuelo Blanco (Fig. 4). The two additionalspecimens were found along basaltic rockwalls that project out along the Rıo Ama-rillo and along more open, gentle slopesjust southeast of the type locality. The hab-itat at the type locality (2600–2800 m) isclassified as Prepuna (Burkart et al., 1999;Cabrera and Willink, 1980). Dominantplants species include the small spinyshrub Adesmia (Fabiaceae) and a spindly,sparsely vegetated Baccharis (Asteraceae)that grows to 1 m. Large metamorphicrock outcrops are common along the roadcut that follows along the Rıo Amarillo.When we visited the type locality in latesummer (mid-February) 1998, the lizardswere active from 0930 to approximately1800 and were most often seen baskingfrom 1–10 m above the road cut on eithersedimentary deposits (at the type locality)or on large outcrops of basaltic rock (alongdirt road). Body temperatures of two adultmale (76 and 85 mm SVL) L. dicktracyicollected on different days as they werebasking in the afternoon on the steep cliffsof the type locality averaged 35.9 C (35.8–36.0 C) despite variable weather condi-tions (sunny versus overcast and windy).The predominantly saxicolous lifestyle (al-though not always on consolidated rock) ofL. dicktracyi is consistent with the micro-habitat selection of other members of theelongatus group (Cei, 1974, 1986; Espi-noza et al., 2000), and particularly that ofL. capillitas, L. heliodermis, and L. um-brifer sp. nov. (described below), whichare strictly saxicolous (Espinoza et al.,2000; Hulse, 1979; see below) and appearto be closely related to the new species(Lobo, 2001; R. E. Espinoza and F. Lobo,unpublished data). We encountered noother species of lizards in syntopy with L.dicktracyi. However, at a slightly lower el-evation (Los Corrales, 288 499 500 S, 678389 200 W; 2200 m), we encountered L.koslowskyi, and the type locality for L. fa-matinae and Phymaturus mallimaccii isCueva de Perez [.4000 m; Cei (1980)],,20 km by road south of the type localityof L. dicktracyi.

Members of the elongatus group are vi-viparous (as far as is known) and live atmoderate to high latitudes or elevations

(Cei, 1986; Espinoza et al., 2000; Hulse,1979; Schulte et al., 2000). This correla-tion is consistent with the hypothesis thatviviparity is an adaptation to reproducingin cold climates (e.g., Guillette, 1993;Shine, 1985; Tinkle and Gibbons, 1977).We were unable to determine the repro-ductive mode of L. dicktracyi because thelizards were collected in late summer, pastthe time when other members of the elon-gatus group are known to give birth(Hulse, 1979; Ibarguengoytıa and Cussac,1998). However, because L. dicktracyilives at a relatively high elevation (2600–2800 m) and because its closest relativesare viviparous, we predict that this speciesis also live bearing. Female L. dicktracyicollected in mid-February, although nolonger pregnant, exhibited pregnant-fe-male coloration as described above and forL. capillitas (Hulse, 1979; R. E. Espinozaand F. Lobo, personal observation; see alsodescription below of L. umbrifer sp. nov.).The red color faded after the lizards werein captivity for approximately 3 wk.

Other members of the elongatus groupare primarily insectivorous (Cei, 1986;Hulse, 1979; Videla, 1983) but may peri-odically include some plant matter in theirdiets (R. E. Espinoza, unpublished data).Feces produced by L. dicktracyi (n 5 10)within 3 d of capture from the field con-tained the remains of insects, with ants ap-pearing to constitute a substantial portionof the diet. Hence, the diet of L. dicktra-cyi is similar to that of L. capillitas, whichalso feeds primarily on ants (Hulse, 1979).

Liolaemus umbrifer sp. nov.Holotype.—FML 9934 (field tag:

ACUNSa 52), an adult male from Quebra-da de Randolfo, Ruta Provincial 43, 60.3km northwest of intersection of Ruta Pro-vincial 4 and Ruta Provincial 43, 166 kmsoutheast of Antofagasta de la Sierra, De-partamento Antofagasta, Provincia de Ca-tamarca, Argentina (268 51.4569 S, 66844.8049 W; 3192 m); collected by C. Ab-dala, R. E. Espinoza, F. Lobo, and M. I.Martınez Oliver, 18 January 2001 (Fig. 5).

Paratypes.—FML 9935–45 (field tags:ACUNSa 50, 53–55, 57, 82–89) and MCN463–64 (field tags: ACUNSa 51, 56) same


FIG. 5.—Holotype of Liolaemus umbrifer (FML9934; SVL 5 88.8 mm).

data as holotype. CM 147714–15 (fieldtags: ACUNSa 58–59) 2 km southeastPena Fria, Ruta Provincial 43, 32 kmnorthwest of Quebrada de Randolfo, 105km southeast of Antofagasta de la Sierra,Departamento Antofagasta, Provincia deCatamarca, Argentina (268 43.199 S, 66857.669 W; 3487 m); collected by C. Abdala,R. E. Espinoza, F. Lobo, and M. I. Mar-tınez Oliver, 18 January 2001.

Diagnosis.—A moderate to large-bod-ied, elongate lizard belonging to the L.elongatus group (sensu Espinoza et al.,2000) that can be distinguished from allother members of that group (and all otherLiolaemus) in possessing the followingtraits: black arms (distinct from light todark brown background color of dorsum)and red coloration in the posterior abdo-men, cloacal region, ventral thighs, andventral base of the tail in both sexes (Table2). The new species can be distinguishedfrom all other members of the elongatusgroup, except the northern radiation (L.capillitas, L. dicktracyi, and L. helioder-mis) by its larger and usually fewer mid-

body scales: L. umbrifer: 58–76; versus76–82 in L. austromendocinus; 72–90 in L.elongatus; 74–89 in L. petrophilus (Don-oso-Barros and Cei, 1971); 84–89 in L.thermarum (Videla and Cei, 1996) (Table2). Liolaemus capillitas, L. elongatus, andL. petrophilus have dorsal scales that aremore distinctly imbricate than those of L.umbrifer, and the former three specieshave keels that occupy the entire scale,whereas the scales of L. umbrifer are onlyweakly keeled (Table 2). Liolaemus elon-gatus have dark brown, irregular, longitu-dinal stripes (6–12 scales wide) that extendalong the vertebral and lateral fields,whereas L. umbrifer lack stripes or similarbody patterns (Table 2). Liolaemus petro-philus have a distinct dorsal pattern oftransverse dark bars that form a ‘‘tigroid’’pattern (Cei, 1975; Donoso-Barros andCei, 1971), which are lacking in this newspecies. Unlike L. umbrifer, L. elongatus(southern populations; see Discussion), L.petrophilus, and most L. austromendoci-nus have distinctly ringed tails (Table 2).The temporals of L. umbrifer are onlyslightly keeled, the head is dark brownwith irregular black markings, and bothmales and females of this species have redcoloration in the cloacal region, whereas inL. dicktracyi the temporals are usually dis-tinctly keeled, the head is black, and onlyfemales possess red coloration in the clo-acal region. Adult L. dicktracyi and maleL. heliodermis have blue or yellow torsos,respectively, whereas L. umbrifer have alight to dark brown trunk. In contrast toL. umbrifer, which lack dorsal body pat-terns and possess 3–4 precloacal pores, L.thermarum have dark, wide stripes on thelower flanks and lack precloacal pores (Vi-dela and Cei, 1996, 1998).

Description of the holotype.—Adultmale (Fig. 5), 88.8 mm SVL; tail length102.4 mm, 71.1 mm (69.4%) regenerated.Axilla–groin distance 41.5 mm. Head 19.2mm long (from anterior border of auditorymeatus to tip of snout), 15.9 mm wide (atanterior border of auditory meatus), 10.2mm high. Snout length 6.3 mm (posteriormargin of canthal to tip of snout). Inter-orbital distance (between arcs of circu-morbitals) 1.7 mm. Eye–nostril distance


3.3 mm. Tibial length 18.4 mm. Footlength 24.9 mm (ankle to tip of claw onfourth toe).

Dorsal head scales smooth, 18 betweenrostral and anterior border of auditory me-atus. Ten keeled temporals (left side), allwith one scale organ on their posteriormargins. Interparietal subpentagonal,slightly smaller than parietal, surroundedby seven scales. Frontal divided into twopairs of scales. Six scales between frontaland rostral. Two postrostrals with 8/10(left/right) scale organs each. Supraorbitalsemicircles complete. Three enlarged su-praoculars (both sides). Five scales be-tween frontal and supercilliaries. Six/sevenflat, elongate, imbricate supercilliaries.Canthal separated from nasal by two scales(both sides). Loreal region flat. Seven/eight scales surrounding nasals. Nasals inslight contact with rostral. Eight lorilabials(both sides), fifth through eighth in con-tact with subocular. Eight/nine enlargedsupralabials. Fifth and sixth supralabial(both sides) curved upward posteriorly andin contact with subocular. Five/six infrala-bials slightly taller than supralabials. Fourinternasals. Orbit with 15 upper and 14lower cilliaries (left side). Orbit diameter4.0 mm (measured between upper andlower ciliaries on left side). Subocularscale elongate. Preocular unfragmented.Longitudinal ridge along upper margin ofthe three ocular scales. Rostral scale morethan twice as wide (3.9 mm) as high (1.5mm). Mental nearly twice as wide (3.9mm) as high (2.2 mm), followed posteri-orly by two rows of four chinshields (bothsides). First chinshield in contact with firstinfralabial. Scales of throat between chin-shields slightly imbricate posteriorly. Sixty-six imbricate gulars between auditory me-atus. Four outward-projecting (virtuallyperpendicular) laminar scales along ante-rior border of auditory meatus. Auditorymeatus oval shaped: 4.1 3 2.6 mm. Lateralscales of neck granular. Subdermal fatbodies give slightly inflated appearance toneck region. Antehumeral fold distinct.Rictal, postauricular, and longitudinal foldspresent but less conspicuous than ante-humeral. Thirty-six scales between audi-tory meatus and antehumeral fold (count-

ed along postauricular and longitudinalfolds).

Sixty-six dorsal scales between occiputand anterior surface of thighs. Dorsal bodyscales round, slightly imbricate, weaklykeeled. Thirty-one longitudinal keeledscale rows over dorsum of trunk. Scales ofneck region smaller than dorsals. Scalesbecome increasingly rounded laterally andalong flanks. Sixty-four scales around mid-body. Ventral scales of similar size to dor-sals, 115 between mental and precloacalpores. Three precloacal pores, each with awaxy exudate. Ventral surface of thighswith enlarged, laminar, imbricate scalesanteriorly, abruptly changing to smallergranular scales on posterior third of thigh.

Fourth finger with 23 tridentate, sub-digital lamellae. Claw of fourth finger 2.0mm, curved, sharp tipped, opaque darkbrown. Fourth toe with 29 subdigital la-mellae. Claw of fourth toe 2.3 mm, similarto that of fourth finger. Hemipenes evert-ed but lobes inadequately preserved fordescribing details of ornamentation.

Variation.—Based on an additional sixadult males and eight adult females. SVL69.0–89.0 mm. Head length 14.6–19.5mm, width 11.6–16.6 mm. Axilla–groindistance 28.7–45.5 mm. Tail length 110.9–142.2 mm. Midbody scales 58–76. Dorsalscales round, not pointed, weakly keeled,subimbricate, 60–74 between occiput andanterior surface of thighs. Some individu-als with interstitial granules and/or poste-riorly notched scales along posterior flanks.Dorsal head scales 13–18, smooth, variablein size but usually small, some irregular inshape, most convex. Ventrals 98–122.Scales around interparietal 6–9. Three tosix enlarged supraoculars. Preocular notdivided, not fused to subocular. Temporals8–10, smooth to weakly keeled. Scales be-tween auditory meatus and antehumeralfold 32–42. Gulars 47–66, smooth, round,imbricate. Supralabials 5–8. Infralabials 4–7. Posterior tip of fourth through sixth su-pralabial upturned. Scales around nasals7–8. Nasals always in slight contact withrostral. Internasals 4 (93%) to 5 (7%). Ros-tral barely visible from above. Scales be-tween rostral and frontal 5–7. Two pos-trostrals with 4–19 scale organs each. Six


to nine lorilabials, fourth, fifth, or sixththrough ninth in contact with subocular.Supercilliaries 6–8. Subdigital lamellae onfourth finger 21–24; on fourth toe 25–31.Precloacal pores 2–4 in males, not presentin females.

Color in life.—Background color ofbody dark brown to brown with lightbrown to beige scales often forming irreg-ular clusters of 2–4 scales on trunk (espe-cially in the dorsolateral field) giving theappearance of faint spots (Fig. 2). Small,black (#1 scale in size) marks also dis-persed over the dorsum. Rarely, blackscales congregate posteriorly forming anirregular rectangle-shaped black field mid-dorsally. Head brown to dark brown withdiffuse, irregular black marks sometimesreaching the level of the shoulders. Pari-etal and temporal regions of head darkbrown to black. Forelimbs black at thepoint of insertion with the trunk to theforearms. On forearms, brown flecks in-crease in density towards toes. Alongflanks of trunk and surrounding the inser-tion of forelimb, black areas are speckledwith cream white scales giving the appear-ance of a dark night sky with stars. Dorsalhind limbs and tail same color as trunk inmost individuals. Distal third of tail in afew individuals becomes dark brown toblack dorsally. Ventrally light to charcoalgray with irregular darker gray markingsscattered on belly and tail. Throat usuallylighter gray than ventral torso. Cloacal re-gion, lower belly, ventral thighs, and ven-tral base of tail with crimson red colorationin females (pregnant or recently spent).Males of this species also exhibit this ven-tral coloration (Table 2), although it is usu-ally restricted to the cloacal region andventral thighs, and the color is generallyless vibrant, and sometimes orange yellowrather than crimson red. In adult males,the posterior margins of the ventral thighsare sulfur yellow. Precloacal pores yellow-orange. In one male (ACUNSa 82), thetorso is entirely black dorsally from justposterior to the shoulders to the base ofthe tail. In this individual, the forelimbsare also entirely black to the digits and thehind limbs are black to half the length ofthe tibia. The melanism on the dorsum of

this individual extends to the flanks andmost of the belly, but fades slightly to char-coal gray mid-ventrally. In juveniles, thetorso is brown with irregular, dark browntransverse bands and lacks the distinctblack pigmentation seen in adults. Theshoulders of juveniles are dark brown tonearly black, but the black coloration,when present, is not as widely distributedon the arms, suggesting the black-arm col-oration may be acquired ontogenetically.Ventrally, juveniles are light gray, almostwhite, with a few small, diffuse, darkergray markings. Juveniles also lack the redcoloration seen in the cloacal region ofadults.

Color in preservative.—As in life, over-all background color darkens slightly onhead, trunk, and ventrally. Red colorationin cloacal region fades to a pale rosy pinkafter approximately 6 mo in preservative.Precloacal pores fade from yellow orangeto cream yellow.

Etymology.—The specific epithet um-brifer is derived from the Latin word um-bra (shade) and the Latin suffix -ifer (fromthe verb fero, meaning to carry or bear).The literal translation ‘‘shade bearer’’ re-fers to the distinctly black upper arms andshoulders that are distinguishing charac-teristics of this species (Fig. 2).

Distribution.—Liolaemus umbrifer isknown primarily from the type locality(Fig. 3): Quebrada de Randolfo, alongRuta Provincial 43, Departamento Anto-fagasta, Catamarca, Argentina (268 51.4569S, 668 44.8049 W; 3192 m). Two additionalspecimens were collected 2 km southeastof Pena Fria on Ruta Provincial 43, 32 kmnorthwest of the type locality (268 43.199S, 668 57.669 W; 3487 m). Aside from therecent discovery of a population of L. elon-gatus (Avila and Lobo, 1999; but see Dis-cussion), L. umbrifer and L. capillitas arethe only members of the elongatus groupknown from Catamarca Province. Quebra-da Randolfo lies along the northern limitof the drainage of the Rıo El Bolson andwest of the Sierra Chango Real, approxi-mately 70 km (straight line) from the typelocality of L. capillitas [Mina Capillitas, De-partamento Andalgala, Catamarca; Hulse(1979); Fig. 3]. Although the distance sep-


FIG. 6.—Granite hillsides of Quebrada Randolfo,Departamento Antofagasta, Catamarca Province (26851.4569 S, 668 44.8049 W; 3192 m), the type localityof Liolaemus umbrifer.

arating these two species is relatively short,there is a vast, low elevation valley (Montehabitat) in the watershed of the Rıo Belenthat lacks suitable habitat and runs in asouthwest–northeast direction betweenthe mountain ranges separating L. capilli-tas and L. umbrifer.

Natural history.—Specimens of L. um-brifer were most commonly encounteredbasking along the steep, rocky hillsides ofQuebrada Randolfo just below the roadcut of Ruta Provincial 43 (Fig. 6). Thehabitat at the type locality (approximately3500 m) is characterized as Puna (Burkartet al., 1999; Cabrera and Willink, 1980).Dominant plant species include the spinyshrub Adesmia, the bunch grass Festuca,Senecio sp. (Asteraceae), lupines, and ad-ditional unidentified spiny shrubs to 1-mhigh. Large outcrops of light gray, brown,maroon, or black granite boulders cover

the sides of the gorge. These rocky hill-sides provide ample habitat for L. umbri-fer, which are agile saxicolous lizards that,when pursued, move quickly from boulderto boulder or retreat temporarily into crev-ices or under large rocks. Body tempera-tures of surface-active adults (n 5 9) col-lected in the afternoon (1215–1445)ranged from 31.2–35.1 C (x 5 33.1 C, SD5 1.4). At the type locality, L. umbrifer issyntopic with Phymaturus antofagastensisand sympatric with an undescribed speciesbelonging to the L. andinus group thatlives among large Adesmia shrubs in thewind-blown sand dunes at the bottom ofthe gorge. Two additional specimens of L.umbrifer were found basking in the lateafternoon (approximately 1800) sun onsmall, black, basaltic rocks (approximately0.5 3 0.3 3 0.3 m) along Ruta Provincial43 in a region of rolling hills covered withsmall pebbles, Adesmia, and sparse clumpsof Festuca some 32 km (by road) north-west of the type locality.

We were unable to determine the re-productive mode of L. umbrifer becausethe lizards were collected in late summer,past the time when other members of theelongatus group are known to give birth(Hulse, 1979; Ibarguengoytıa and Cussac,1998). However, because L. umbrifer livesat relatively high elevation (approximately3200–3500 m) and because its closest rel-atives are viviparous, we predict that thisspecies is also live bearing. Most of theadult female L. umbrifer we collected inmid-January, although no longer pregnant,exhibited pregnant-female coloration asdescribed above. Interestingly, males ofthis species also exhibit this ventral color-ation (Table 2), although the color wasgenerally less vibrant and appeared to beless widely distributed ventrally.

Feces produced by L. umbrifer (n 5 11)within 3 d of capture contained the re-mains of insects and, occasionally, parts offlowers (probably Senecio sp.).

DISCUSSION

Distribution of pregnant-female colora-tion.—A number of Liolaemus exhibitgravid- or pregnant-female coloration (e.g.,Etheridge, 2000; R. E. Espinoza and F.


FIG. 7.—Phylogenetic affinities of the elongatus group following (A) the molecular phylogeny of Schulte etal. (2000) and (B) Cei (1975). There is much congruence between the hypothesized relationships despitetheir different data sources (mitochondrial DNA versus blood serum, respectively) and methods of analysis(cladistic versus phenetic, respectively). However, the tree of Schulte et al. (2000) has members of the kriegigroup [L. buergeri, L. ceii, and L. kriegi; sensu Cei (1986)] nested within the elongatus group and positsdifferent relationships among L. austromendocinus, L. elongatus, and L. petrophilus than that of Cei (1975).See text for further discussion.

Lobo, personal observation; Lobo and Es-pinoza, 1999). In most species, this consistsof bright yellow to orange or salmon coloralong the sides of the head, throat, andneck, which sometimes extends along theflanks or belly. In members of the north-ernmost clade of the elongatus group, thiscolor consists of a patch of crimson red inthe cloacal region that extends to the lowerbelly and, in some species, along the thighsand onto the proximal quarter of the tail.Within this group, this coloration is knownfrom pregnant and recently spent femaleL. capillitas (Hulse, 1979; F. Lobo and R.E. Espinoza, personal observation), re-cently spent female L. dicktracyi (de-scribed above), males and recently spentfemales of L. umbrifer (described above),and suspected but not confirmed for preg-nant and recently spent female L. heliod-ermis (Table 2). Additionally, this colora-tion was recently observed in male andpregnant female L. buergeri (M. Christie,personal communication; F. Lobo and R.E. Espinoza, personal observation), a spe-cies that may be nested within the elon-gatus group (Schulte et al., 2000; and seebelow). When we collected L. dicktracyiand L. umbrifer, females were no longerpregnant but still exhibited this coloration.We hypothesized that, owing to the timingof reproduction among members of theelongatus group (Hulse, 1979; Ibarguen-goytıa and Cussac, 1998; F. Lobo and R.E. Espinoza, unpublished data), the fe-males were collected soon after parturition

and simply retained the coloration for sev-eral days thereafter as is typical of L. cap-illitas (F. Lobo and R. E. Espinoza, per-sonal observation) and other iguanians(Cooper and Greenberg, 1992; Cooperand McGuire, 1993). However, the pres-ence of this trait in male L. umbrifer poseseither an exception or an interesting chal-lenge to linking this coloration with preg-nancy. Moreover, it is not known how, oreven whether, this coloration is used forcommunication among species of Liolae-mus. In other iguanian lizards, gravid-fe-male coloration is usually distributed alongthe flanks, head, or neck and is thought toprovide a visual signal which, when cou-pled with aggressive behavior, deters malesfrom attempting to court or copulate withfemales (Cooper and Greenberg, 1992). Incontrast, the ventral distribution of thiscoloration seems less conspicuous in mem-bers of the northern radiation of the elon-gatus group, which would suggest theneed for specialized behaviors associatedwith exhibiting the red patches. Behavioralstudies are sorely needed to test these pre-dictions.

Phylogenetic relationships within theelongatus group.—A recent cladistic anal-ysis of mitochondrial DNA gene sequenc-es for 60 species of Liolaemus (Schulte etal., 2000) found strong bootstrap support(100%) for a clade including the elongatusgroup members (sensu Cei, 1974, 1986)plus L. buergeri, L. ceii, and L. leopardi-nus (Fig. 7A). In a phenetic analysis based


on serological evidence, Cei (1975) alsoconsidered the elongatus group (which, atthat time, consisted only of L. austromen-docinus, L. elongatus, and L. [elongatus]petrophilus) to be closely related to thekriegi group (see also Cei, 1986), whichincludes L. buergeri, L. ceii, and L. kriegi(Fig. 7B). It is worth noting that these twohypotheses of relationships are remarkablysimilar despite the difference in the kindsof data used and types of analyses con-ducted. The only incongruence betweenthe hypotheses that cannot be attributedto differences in taxon sampling lies in thedetermination of the sister species of L.elongatus. Cei (1975) allied this specieswith L. [elongatus] petrophilus (Fig. 7B),whereas Schulte et al. (2000) found L.elongatus to be most closely related to L.austromendocinus (bootstrap 100%; Fig.7A). However, the L. elongatus used in theSchulte et al. (2000) analysis was collectednear Uspallata, in Mendoza Province, andthe type locality of this species is ChubutProvince (although not precisely identi-fied; Koslowsky, 1896), at least 1300 kmsouth. Other researchers have noted thevariability in size, squamation, body pat-terns, and color among populations of thewidely distributed L. elongatus (L. Avila,personal communication; Cei, 1986; R. E.Espinoza and F. Lobo, unpublished data;Espinoza et al., 2000; Quatrini et al.,1998). Some of these populations are al-lopatric and readily diagnosable; in fact,our preliminary examination of specimensassigned to L. elongatus suggests thatthere are at least three cryptic specieswithin the complex (R. E. Espinoza and F.Lobo, unpublished data). Hence, the taxonin the tree of Schulte et al. (2000) mostlikely represents an undescribed speciesrather than L. elongatus sensu stricto. In-deed, recent morphological and molecularanalyses, which include specimens of L.elongatus from Chubut, support Cei’s(1975) sister grouping of L. elongatus andL. petrophilus (R. E. Espinoza and F.Lobo, unpublished data).

Schulte et al. (2000) also identified L.capillitas as the sister taxon to the remain-ing members of their more inclusive elon-gatus group (i.e., including the kriegi

group; Fig. 7A). Thus, if L. dicktracyi andL. umbrifer form a clade with L. capillitasand L. heliodermis, as is supported by ourmorphological and molecular evidence (R.E. Espinoza and F. Lobo, unpublisheddata), then this northern radiation consti-tutes the sister lineage of the more inclu-sive elongatus group. Additionally, L. leo-pardinus, a species from moderate eleva-tions (1800–2750 m) in north central Chile(33–348 S) was placed as the outgroup ofL. buergeri 1 L. ceii [the kriegi group ofCei (1986), less L. kriegi] (Schulte et al.,2000; Fig. 7A). Given that L. ramonensisand L. valdesianus are both morphologi-cally similar to, and were originally de-scribed as, subspecies of L. leopardinus(Hellmich, 1950; Muller and Hellmich,1932), future analyses of the elongatusgroup should consider these taxa as well.Finally, L. coeruleus, L. cristiani, L. isa-belae, and L. neuquensis share a numberof morphological traits in common withspecies in the elongatus group (e.g., elon-gate body and long tail, small body scales,lack of precloacal pores) and may, there-fore, represent additional members of thislineage (Lobo, 2001; F. Lobo, unpublisheddata; Videla and Cei, 1998). Determiningthe taxonomic status of the various morphsof L. ‘‘elongatus’’ and the phylogenetic re-lationships among the numerous putativemembers of the elongatus group are thesubjects of detailed molecular and mor-phological analysis currently under inves-tigation (L. Avila, personal communica-tion; R. E. Espinoza and F. Lobo, unpub-lished data).

RESUMEN

El grupo de especies de Liolaemus elon-gatus actualmente incluye ocho especies(incluyendo las especies que aquı se des-criben) de lagartos de mediano tamano,con colas largas, y primariamente saxıcolas.Los miembros de este linaje se distribuyendesde el noroeste de Argentina hacia elsur, a lo largo de la Cordillera de los Andeshasta la estepa Patagonica (27–448 S) y so-bre un amplio rango de elevacion (350–3900 m). Aquı nosotros describimos dosespecies nuevas que pertenecen a la radia-cion nortena del grupo elongatus. Los cua-


tro miembros actualmente reconocidos deeste subclado norteno (incluyendo las dosespecies que aquı se describen) se diferen-cian de los otros miembros del grupo elon-gatus, por contar con un menor numerode escamas alrededor del cuerpo, cabezasmarron a negras, y por carecer de los dis-tintos patrones dorsales de cola o cuerpoencontrados en adultos de otras especiesque pertenecen al grupo elongatus. Ade-mas, las hembras que recientemente hanparido exponen color rojo vivo en la regioncloacal, una caracterıstica unica conocidapara miembros de este linaje norteno y L.buergeri. La primera especie nueva que sedescribe, L. dicktracyi, de la Sierra de Fa-matina de la Provincia de La Rioja, difierede todo los otros miembros del grupo elon-gatus en varios caracteres merısticos y ensu coloracion dorsal unica: posee cabeza,hombros y cola negros, y el torso y extre-midades posteriores azules. La segunda delas especies nuevas que se describen, L.umbrifer de la Quebrada de Randolfo enel noreste de la Provincia de Catamarca,es morfologicamente parecido a L. capilli-tas, pero difiere de esta y otras especies enel grupo elongatus en varios caracteresmerısticos, la presencia de brazos y hom-bros negros, y en la coloracion roja viva dela region cloacal en ambos, machos y hem-bras. La historia natural de ambas especiesnuevas es similar en muchos aspectos a L.capillitas y L. heliodermis debido a sus afi-nidades filogeneticas de los miembros deeste subclado dentro del grupo elongatus.Finalmente, nosotros proveemos compa-raciones de recientes hipotesis filogeneti-cas y discutimos el contenido del grupoelongatus.

Acknowledgments.—We appreciate the assistanceof C. Abdala, F. Cruz, and M. I. Martınez Oliver whohelped collect the types. The senior author thanks F.Cruz and G. Perotti for lodging and companionshipand M. Huniken for providing a place to study spec-imens at CRILaR (Anillaco, La Rioja). We thank S.Murua and E. Fra of the Provincial Flora and Faunaoffices of La Rioja and Catamarca, respectively, fortheir assistance with collecting permits (issued to F.Cruz and F. Lobo). J. Wiens and S. Rogers of theCarnegie Museum of Natural History, G. Scrocchiand S. Kretzschmar of the Fundacion Miguel Lillo,and D. Wake and B. Stein of the Museum of Verte-brate Zoology (MVZ) provided specimens for com-parisons and logistical and curatorial support. We are

grateful to A. Summers who housed us during ourvisit to MVZ. J. Rawlins provided invaluable advicein forming the epithet for L. umbrifer, and C. Hitch-cock repeatedly revised the map to suit our specifi-cations. We thank R. Etheridge and an anonymousreviewer for critically reading and improving earlyversions of the manuscript and C. R. Tracy for hisenduring support and critical feedback of our studies.R. E. Espinoza was partially supported by grants fromthe American Society of Ichthyologists and Herpe-tologists, the National Geographic Society, the Soci-ety of Comparative and Integrative Biology, and theUpstate [NY] Herpetological Association. The Biolo-gy Department, Graduate School, and Biological Re-sources Research Center at the University of Nevada,Reno, and the Carnegie Museum of Natural Historyalso provided additional logistical and financial sup-port. A fellowship from CONICET of Argentina andthe Universidad Nacional de Salta supported F. Lobo.

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Accepted: 11 June 2002Associate Editor: Stephen Tilley

APPENDIX I

Specimens Examined

Liolaemus austromendocinus (10).—ARGENTI-NA: Provincia de Mendoza: Departamento Malargue,on Ruta Nacional 40, 47 km S Malargue (358 46910.40 S, 698 389 46.80 W; 1920 m): FML 3432 (1–7);Laguna Nina Encantada (358 099 37.40 S, 698 52909.00 W; 1680 m): FML 7189–91 (erroneously iden-tified as L. thermarum in Espinoza et al. [2000]).

L. capillitas (16).—ARGENTINA: Provincia deCatamarca: Departamento Andalgala, Mina Capillitas(3000–3600 m): FML 1229 (1, 2, 4, 6, 7, 11, 12, 16,18–21); Morro El Arenal (El Ingenio) (3100 m):FML 2029 (1–4).

L. dicktracyi (10).—ARGENTINA: Provincia deLa Rioja: Departamento Famatina, 26.5 km W of Fa-matina on road to Cueva de Perez, Portezuelo Blanco(288 549 300 S, 678 419 300 W; 2800 m): FML 9928


(holotype); CM 147716–17 (paratypes); FML 9929–33 (paratypes); MCN 461–62 (paratypes).

L. elongatus (10).—ARGENTINA: Provincia deNeuquen: Departamento Alumine, along Arroyo Ru-caco, SE end of Lago Choroi, 3.5 km E and 6.5 kmN Cerro Ruca Choroi (39.128 S, 71.178 W; 1250 m):MVZ 188764. Provincia de Rıo Negro: DepartamentoBariloche, ridge above Refugio Neumeyer, 15 km SBariloche (41.268 S, 71.318 W; 1600 m): MVZ 188781.Departamento Norquinco, along Rıo Chenqueniyen,10 km E and 3 km S Cerro Pico Quemada (41.538 S,71.008 W; 1150 m): MVZ 188732–33, 188739; rim-rock, 4 km S and 1 km E Alto del Escorial (41.598 S,70.768 W; 1100 m): MVZ 188743; Laguna de Los Jun-co, Escorial de Chenqueniyen, 5 km N Alto de Es-corial (41.518 S, 70.788 W; 1150 m): MVZ 188758.Departamento Pilcaniyeu, 1.5 km N Estacion PeritoMoreno, approximately 28 km ENE Bariloche(41.058 S, 71.018 W; 900 m): MVZ 180069; CanadonBonito, 23 km NE Pilcaniyeu (40.988 S, 70.428 W):MVZ 188727–28.

L. heliodermis (3).—ARGENTINA: Provincia deTucuman: Departamento Tafı del Valle, Ruta Provin-cial 307, approximately 32 km (by road) from Tafı delValle at km marker 95 (268 40.829 S, 658 48.749 W;2820 m): FML 6006–07 (paratypes), FML 7196 (ho-lotype).

L. petrophilus (3).—ARGENTINA: Provincia deChubut: Ruta Nacional 25, 10 km S Puesto Espinal(350 m): FML 793 (1–3).

L. umbrifer (16).—ARGENTINA: Provincia deCatamarca: Departamento Antofagasta, Quebrada deRandolfo, Ruta Provincial 43, 60.3 km north of in-tersection of Ruta Provincial 4 and Ruta Provincial43, 166 km southeast of Antofagasta de la Sierra (26851.4569 S, 668 44.8049 W; 3192 m): FML 9934 (ho-lotype), FML 9935–45 (paratypes), MCN 463–64(paratypes); 2 km southeast Pena Fria, Ruta Provin-cial 43, 32 km northwest of Quebrada de Randolfo,105 km southeast of Antofagasta de la Sierra (26843.199 S, 668 57.669 W; 3487 m): CM 147714–15 (pa-ratypes).

Documents

two new species of liolaemus lizards from northwestern argentina