Quinteros Et Al 2014

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A New Species ofLiolaemus(Iguania: Liolaemidae) of thealticolor-bibroniiGroup from Northern Chile

Andrs Sebastin Quinteros1,*, Pablo Valladares2, Romina Semham3, Jos Luis Acosta4,

Sebastin Barrionuevo5

, Cristian Simn Abdala3

1 Ctedra de Sistemtica Filogentica. Ctedra de Zoologa General. Facultad de Ciencias Naturales. Universidad Nacional de Salta. IBIGEO - CONICET.

Av. Bolivia 5150. Salta. Argentina.2 Departamento de Biologa, Facultad de Ciencias, Universidad de Tarapac, General Velsquez 1775, Arica, Chile.3 Instituto de Herpetologa, Fundacin Miguel Lillo, CONICET. Universidad Nacional de Tucumn. Miguel Lillo 251. 4000 - Tucumn, Argentina.4 CONICET - Universidad Nacional del Noreste, Corrientes, Argentina.5 Divisin de Herpetologa. Museo Argentino de Ciencias Naturales. CONICET.

* Corresponding Author. Email: [email protected]

Abstract.We describe a new slender species of Liolaemusof the L. alticolor-bibroniigroup of the subgenus Liolaemus.The new species is phe-netically and biogeographically close to L. alticolor, L. paulinae,and L. punabut presents a combination of character states that differentiatesit from all other species of Liolaemus.The new taxon is the first species of subgenus Liolaemus sensu strictorecorded as having supernumerarypores. The new species inhabits places where Parastrephia lucidais the predominant flora and is distributed in areas close to Putre, in theArica y Parinacota region, northern Chile. Data on its biology, complete distribution, and conservation status are lacking.

Keywords.Chile; Lizard; New species; South America; Squamata; Taxonomy.

Resumen.Describimos una nueva especie de Liolaemus,perteneciente al grupo de L. alticolor-bibroniidel subgnero Liolaemus sensu stricto.La nueva especie es fentica y biogeogrficamente cercana a L. alticolor, L. paulinaey L. puna.Pero muestra una combinacin de estados decarcter que permiten su diferenciacin con estas y con todos los dems Liolaemus.El nuevo taxon es la nica especie del subgnero Liolaemussensu strictoque exhibe poros supernumerarios. La nueva especie habita en regiones donde Parastrephia lucidaes la planta predomiante yest distribuida en reas cercanas a la localidad de Putre, en la Regin de Arica y Parinacota, en el norte de Chile. No existen datos sobre subiologa, distribucin completa ni de su estado de conservacin.

INTRODUCTION

Liolaemus comprises 255 species distributed insouthern South America, from Tierra del Fuego (south-ernmost distribution) in Argentina to La Libertad (north-ernmost distribution) in northern Peru. The L. alticolor-bibroniigroup is a member of the Liolaemus sensu strictosubgenus (Ortiz, 1981; Lobo, 2001, 2005; Espinoza et al.,2004; Lobo et al.,2010; Martinez et al.,2011; Quinteros,2012, 2013). Study of the taxonomy of the L. alticolor-bibronii group began with Ortiz (1981) and Cei (1986),and the group has grown to include 26 described species,including the new species described herein (Quinteros,2013).

In the last two years, four new species of the L. alti-color-bibroniigroup were described: L. cyaneinotatusMar-tinez et al.,2011, L. aparicioiOcampo et al.,2012, L. abda-lai,and L. pyriphlogosQuinteros, 2012. All these recentlydescribed species are distributed in Argentina and Bolivia.In the last eight years, eight species of Liolaemus weredescribed from Chile (L. carlosgarini, L. chacabucoense,L. confusus, L. filiorum, L. frassinettii, L. hermannunezi,

L. riodamas,and L. scolaroi), none of which belong to theL. alticolor-bibroniigroup. The only species described fromChile of the L. alticolor-bibronii group in recent years is

L. barbaraePincheira-Donoso and Nez, 2005, but this

taxon is a junior synonym of L. puna(see Quinteros andLobo, 2009). In the last austral summer, we conducted afield trip to northern Chile where we found a new spe-cies of Liolaemusthat we hypothesize to be a member ofthe L. alticolor-bibroniigroup. This new taxon is describedherein and compared with, among others, the phenetical-ly most similar and geographically closest taxa: L. alticolor,L. puna,and L. paulinae.

MATERIALS AND METHODS

In order to describe the new taxon, we studied themorphological characters traditionally used in Liolae-mus taxonomy, including those of Laurent (1985), Cei(1986, 1993), Etheridge (1993, 1995, 2000), Lobo (2001,2005), Abdala (2002, 2003, 2007), and Quinteros (2012,2013). Terminology for squamation follows Smith (1946)and for neck-folds follows Frost (1992). Descriptions ofbody color patterns follow Lobo and Espinoza (1999) andQuinteros (2012, 2013). Descriptions of color in life forthe new species were based on observations of freshly col-lected specimens and photographs taken at the time ofcapture. Measurements and scale counts were recorded

South American Journal of Herpetology,9(1), 2014, 2029 2014 Brazilian Society of Herpetology

Submitted: 12 December 2013Accepted: 21 March 2014Published: 30 April 2014

Handling Editor: Tiana Kohlsdorfdoi: 10.2994/SAJH-D-13-00040.1ZooBank: urn:lsid:zoobank.org:pub:78CEBA23-269C-4340-93A6-056408731C9A


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from specimens fixed in 10% formalin and preserved in70% ethanol. Body and scale measurements were takenwith digital calipers to the nearest 0.05 mm. When nec-essary, a binocular dissecting microscope (1040) wasused to count and characterize scales. Where bilateral,scale counts and mensural data were taken from the right

side of the lizards. Statistical summaries are reported asthe mean SD.For the purposes of diagnosing the new spe-

cies described herein, we examined the type series ortopotypes of other species of the subgenus Liolaemuswhen available, including all currently recognized spe-cies of the L. alticolor-bibronii group (preserved and inlife), with the exception of L. cyaneinotatus.For speciesthat were not collected, color pattern data were takenfrom the literature: Liolaemus fuscus, L. pseudolemnisca-tus, L. tacnae,and L. walkeri (Shreve, 1938, 1941; Cei,1986; Pincheira-Donoso and Nez, 2005; Scolaro,2005, 2006). Since L. incaicus is only known from its

type series (collection specimens), the color patterndata were those taken from preserved specimens. In to-tal, we examined 638 specimens from 17 institutions(Appendix). Institutional acronyms follow Leviton et al.(1985), with the exception of Coleccin Boliviana deFauna (CBF), La Paz, Bolivia; Coleccin Herpetolgicade la Facultad de Ciencias Exactas y Naturales (UNSJ),San Juan; Coleccin Herpetolgica del Centro de Bio-diversidad y Gentica (CBGR), Cochabamba; Museo deCiencias Naturales de la Universidad Nacional de Salta(MCN), Salta, Argentina; Coleccin Zoolgica de Zo-nas ridas y Andinas, Universidad de Tarapac (CZZA),

Tarapac, Chile.

RESULTS

Liolaemus chungarasp. nov.

Holotype

FML 26505: Male. Entrada a Putre. 181053.4S;693158.6W; 3874 m. Arica y Parinacota Regin. Ab-dala, C.S.; Acosta J.L.; Semhan R. and Valladares, P. Cols.

Paratypes

FML 26505: Female. Same data as holotype.FML 26504-508; MCN 4734-35: five males and two fe-males. Margen Sur del Lago Chungar. 181617.0S;690937.7W; 4583 m. Arica y Parinacota Regin. Ab-dala, C.S., Acosta J.L., Semhan R. and Valladares, P. Cols.CUT 1000-02, FML 26509-513: four males and four fe-males. Cerca de Putre desvo a Colpita. Arica y ParinacotaRegin. Abdala, C.S., Acosta J.L., Semhan R. and Val-ladares, P. Cols.

Diagnosis

A small (snoutvent length [SVL]


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as in L. alticolorand L. puna) and absent in L. bitaeniatus,L. incaicus, L. paulinae, L. pagaburoi, L. ramirezae, L. tacnae,

L. variegatus, and L. walkeri. Paravertebral spots are ab-sent in L. chungarabut are evident in L. aparicioi, L. bitae-niatus, L. incaicus, L. paulinae, L. pyriphlogos, L. pagaburoi,and L. variegatus.A vertebral line is distinct in specimens

of L. chungara,whereas most of the specimens of L. apari-cioilack a vertebral line (although a few specimens had atrace or fragmented line), and a vertebral line is absent inL. bitaeniatus, L. tacnae, in females of L. incaicus, and inmales of L. puna. Females of L. chungara lack precloacalpores, whereas females of L. aparicioi, L. bitaeniatus, L. in-caicus, L. ramirezae, L. variegatus, and L. yanalcu exhibitprecloacal pores. In L. chungara the throat ground coloris gray, whereas in L. bitaeniatus, L. chaltin, and L. paga-buroi it is cream-white, and in L. tacnae it is melanistic.Moreover, the throat in L. chungarapresents black spots(or lines), which differs markedly from the immaculatethroat of L. chaltin, L. incaicus,and L. ramirezae.

The new taxon differs from L. abdalai, L. bibronii,L. exploratorum, L. fuscus, L. lemniscatus, L. saxatilis, andL. tandiliensis in lacking paravertebral spots (present inthose taxa). Temporal scales (upper) are slightly keeledin L. chungara, whereas they are smooth in L. bibroniiand markedly keeled in L. lemniscatus, L. saxatilis, andL. robertmertensi.Also, most of necks scales of L. chungaraare granular and a few laminar (but this last one smooth),differing from L. abdalai, L. fuscus, L. lemniscatus, andL. saxatilis (laminar and keeled scales), L. cyaneinotatus(keeled scales) and L. bibronii, L. exploratorum,and L. san-juanensis (laminar and smooth, but without granular

scales). Liolaemus chungarahas lower scales around mid-body (3949), than L. exploratorum(5052).

Description of holotype

Adult male. SVL 49.9 mm. Axillagroin distance21.06 mm. Head 11.35 mm long (from anterior border ofauditory meatus to tip of snout), 9.35 mm wide (at an-terior border of auditory meatus), 6.6 mm high. Interor-bital distance (between postorbital semicircles) 6.3 mm.Eyenostril distance 1.6 mm. Tibia length 7.3 mm. Footlength 16.2 mm (from ankle to tip of claw on fourth toe).Dorsal head scales smooth between rostral and anteriorborder of auditory meatus. Seven smooth temporal, up-per temporal slightly keeled. Interparietal subpentagonal,smaller than parietal in size, surrounded by six scales.Frontal azygous. Six scales between frontal and rostrals,five between frontal and supercilliaries. Two postrostralswith four scale organs each. Supraorbital semicircles com-plete, formed by 10 scales. Five enlarged supraoculars. Sixflat, elongate, imbricate supercilliaries. Canthal separatedfrom nasal by one scale. Loreal region flat. Seven scales,including rostral, surrounding nasals. Nasals in broadcontact with rostral. Eight lorilabials, sixth to eighth in

contact with subocular. Seven enlarged supralabials.Fourth supralabial curved upward posteriorly, not in con-tact with subocular. Five infralabials, slightly taller thansupralabials. Orbit with 12 upper and 11 lower ciliaries.Subocular scale elongate, length 3.27 mm, whitish (lighterthan the loreal region). Preocular unfragmented, length

0.98 mm. Postocular length 2.13 mm. Longitudinal ridgealong upper margin of the three ocular scales. Rostralscale 2.7 times wider (2.43 mm) than high (0.88 mm).Mental 1.2 times wider (2.5 mm) than high (1.9 mm), fol-lowed posteriorly by two rows of four chinshields. Twoscales in contact ventrally with second infralabial. Scalesof throat between chinshields subimbricate. Thirty-sixgulars between auditory meatus. Auricular scale evident,three (one enlarged) outward projecting laminar scalesalong anterior border of auditory meatus. Auditory me-atus higher (1.98 mm) than wide (0.92 mm). Most scalesof neck granular, some smooth and laminar. Antehumeralfold distinct. Rictal, postauricular, and longitudinal folds

present but less conspicuous than antehumeral. Thirty-one scales on neck (between posterior margin of auditorymeatus and shoulder). Nineteen scales between auditorymeatus and antehumeral fold (counted along postau-ricular and longitudinal folds). Forty-three dorsal scalesbetween occiput and anterior surface of thighs. Dorsalbody scales lanceolate, imbricate, keeled, and mucronate.Forty-three scales around midbody. Seventy ventrals be-tween mental and precloacal pores row. Four precloacalpores. Fourth finger with 17 subdigital lamellae. Fourthtoe with 22 subdigital lamellae.

Color of the holotype in ethanol

Vertebral region brownish gray with conspicuousblack vertebral line. Dorsolateral stripes evident, lightbrown, bordered by black lines. These stripes begin inposterior margin of eye, end in base of tail, thinner on an-terior region of body and becoming wider on mid-trunkand tail. Paravertebral zone absent. Lateral field the samecolor as vertebral region, speckled with black lines. Ven-trolateral line white (one scale wide). Head dorsally brownwith black line surrounding interparietal scale and blackline beginning at anterior margin of interparietal scaleand crossing frontal scale to reach rostral. Black spotsevident in superorbital semicircles and supraocular scales.Head laterally gray with dark brown spots on supra- andinfralabial scales. Temporal zone gray with white line bor-dered in dark brown. This white line begins on posteriormargin of postocular scale, crosses along dorsal margin ofauricular meatus, extends to the antehumeral fold. Necksame color as temporal zone, bearing some dark brownspots. Fore- and hind limbs light brown with dark brownand some white spots. A white line bordered in dark brownsurrounds shoulder. Ventrolateral region light brownspeckled with black spots. Throat, chest, and belly gray. A

A New Species of Liolaemus(Iguania: Liolaemidae) of the alticolor-bibroniiGroup from Northern ChileAndrs Sebastin Quinteros, Pablo Valladares, Romina Semham, Jos Luis Acosta, Sebastin Barrionuevo, Cristian Simn Abdala22

South American Journal of Herpetology,9(1), 2014, 2029


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line of black spots extends between chinshields and infral-abials. Throat speckled with black spots. Chest and bellyspotless. Fore- and hind limbs immaculate gray. Tail au-totomized, dorsally light brown with black vertebral line.

At base (anterior to autotomized region region) two blacklines parallel to vertebral line are evident. Ventral and lat-eral regions of tail light brown with light gray lines.

Variation

Variation based on all 16 paratypes (9 males and 7 fe-males, Table 1). SVL 46.355.6 mm (X= 50.44 2.8 mm)in males and 47.352.4 m (X = 49.2 = 2.02 mm)in females; axillagroin distance 20.322.04 mm(X = 22.72 1.28 mm) in males and 22.9929.76 mm(X = 25.55 2.53 mm) in females. Head length9.8511.86 mm (X = 10.85 0.69 mm), width 7.629.45 mm (X = 8.55 0.54 mm). Tail length 71.1393.1 mm (X= 76.86 8.10 mm). Midbody scales 3849(X= 43.9 2.96 mm). Dorsal scales 4047 (X= 43.5 2.32)between occiput and anterior surface of thighs. Dor-sal head scales 1115 (X = 12.1 1.10 mm). Ventrals6284 (X= 72.4 5.54). Scales around interparietal 48(X= 6.4 1.07). Four supraoculars, 34 (X= 3.1 0.32)enlarged. Preocular not divided, not fused to subocular.Smooth temporal, upper slightly keeled (from auricu-lar scale to postorbital scale) 78 (X = 7.6 0.2). Mostof necks scales granular, some smooth and laminar;from posterior edge of auricular meatus to shoulder2733 (X= 30.3 2.4). Gulars 2937 (X= 33.4 2.06).Supralabials 68 (X = 6.3 0.67). Infralabials 45

(X= 4.8 0.42). Posterior tip of fourth through seventhsupralabial upturned, in contact with subocular in 40%of specimens. Scales around nasals 67 (X= 6.6 0.52).Four internasals. Scales between rostral and frontal 57(X= 5.7 0.67). Two postrostrals with 36 scale organs inthe left one (X= 4.6 0.96) and nine to eight in the rightone (X= 4.2 1.99). Six to nine lorilabials (X= 7 0.94),

three to five of them in contact with subocular scale. Sub-digital lamellae on fourth finger 1719 (X= 17.89 0.92);on fourth toe 2025 (X= 22.4 1.43). Precloacal pores36 in males (X= 4.28 0.95); absent in females. Amongall examined males, one possessed two supernumerariespores (Fig. 1).

Figure 1.Cloacal region of MCN 4734 showing the presence of super-numerary pores (arrows). Scale = 1 mm.

Table 1.Variation in some character-states of Liolaemus chungarasp. nov. and phylogenetically and geographically close species. Data are ranges (firstrow) and means SD (second row).

CharacterLiolaemus

alticolorn= 16

Liolaemusaparicioi

n= 12

Liolaemuschungara

n= 11

Liolaemuspaulinae

n= 15

Liolaemuspunan= 55

Liolaemustacnae

n= 5

Liolaemuswalkerin= 40

Number of midbody scales 395143.9 3.4

394643.1 2.5

284943.8 2.82

425851.6 3.96

385244.7 4.05

444846 1.8

415952 3.8

Number of dorsal scales 364841.0 2.9

404642.8 1.8

404743.45 2.2

465550.5 2.6

435164.8 2.6

475349 2.3

416150.7 4.5

Number of temporals 696.7 0.9

687.5 0.7

787.5 0.5

798.1 0.72

576.04 0.66

898.6 0.5

7108.4 0.8

Number of ventrals 628472.9 6.4

748879.9 3.6

628472.18 5.3

809285.8 3.7

789182.1 4.1

728176 3.7

609078.2 6.3

Precloacal pores in males 243 0.8

465.3 0.8

364.25 0.88

464.6 0.9

353.9 0.7

3 565.25 0.25

Precloacal pores in females Absent 031.3 1.2

Absent Absent Absent Absent Absent

Supernumerary pores Absent Absent Present Absent Absent Absent Absent

Dorsal scales Mucronate Mucronate Mucronate Non-mucronate Mucronate Non-mucronate Mucronate

Temporals Weakly keeled Keeled Smooth/upperslightly keeled

Smooth Smooth/upperslightly keeled

Smooth Smooth

Neck scales Laminar,weakly to

distinct keeled

Laminar,keeled

Granular/some laminar,

smooth

Laminar,keeled

Laminar,smooth or

slightly keeled

Granular,smooth

Laminar,smooth

Paravertebral markings Absent Evident Absent Evident Evident Absent Absent

A New Species of Liolaemus(Iguania: Liolaemidae) of the alticolor-bibroniiGroup from Northern ChileAndrs Sebastin Quinteros, Pablo Valladares, Romina Semham, Jos Luis Acosta, Sebastin Barrionuevo, Cristian Simn Abdala 23



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Color in life (Figs. 2, 3)

Vertebral zone gray with vertebral line. Paraverte-bral zone dark brown or black, without spots. Dorsolateralstripes brownish yellow, thinner at beginning (posteriorupper edge of eye), wider proximally. Dorsolateral stripes

bordered by black line. Internal line fused to paravertebralzone sometimes. Lateral field dark brown speckled withblack spots. Males and females with same color pattern,but in some males the background colors are lighter. Lat-eral field and paravertebral spots olive brown in one male.

Ventrolateral line white. In females, ventrolateral zonewith same coloration as lateral field, black spots present.Coloration of ventrolateral zone bright orange speckledwith black spots in some males. Dorsally, background ofhead same as paravertebral zone. Black line surroundinginterparietal scale and follow to the rostral scale pressent.Laterally, head with white stripe bordered by black linesbetween upper margin of the eye and dorsolateral stripes.

White stripe bordered in black extending from middleposterior margin of eye to upper margin of the auditorymeatus. This stripe usually extends from auditory meatusto shoulder across horizontal fold to ventrolateral line. Insome individuals (e.g., holotype), this stripe extends toantehumeral fold. Neck same color as lateral field, with

black spots. Fore- and hind limbs light brown with darkbrown spots. Dorsally, tail same background color as dor-solateral stripes, vertebral line always present. In somespecimens, the paravertebral regions forms two blacklines parallel to vertebral line. In other specimens, theparavertebral regions are fused to vertebral line. Lateral

field of tail bright orange with black spots in some males.In some specimens the external black lines of dorsolateralstripes extend to the tip of tail. Throat with black spotsover light gray background. Chest and belly spotless. Insome males, sides of belly bright orange coloration and afew black spots in the ventrolateral field. In some males,femoral and cloacal zone bright orange. In females, lightgray to pale yellow. Pygal region of males without brightorange coloration, it remains light gray. In males, tail ven-trally orange with elongate spots. In females these spotsare present but background coloration is yellowish white.

Distribution

Liolaemus chungarais known from the vicinity of thetype locality in Putre, a small locality in northern Chile inthe Arica y Parinacota region. It also occurs around LagoChungar, and in areas close to Colpita (Fig. 4).

Natural history

The biology of Liolaemus chungara is poorly known.It inhabits sandy substrate where bushes are te predomi-nant vegetation (mainly Parastrephia lucida). In Parina-cota, it occurs in sympatry with L. pleopholis,and in areas

near Lago Chungar it occurs in sympatry with L. jamesi.The bright orange coloration present on ventrolateralzone, belly, femoral region, and ventral tail in males andabsent in females might be related to reproductive biol-ogy. Bright ventral coloration has also been reported inmales of other species of the L. alticolor-bibronii group(Quinteros, 2012, 2013).

Etymology

We name this species chungarabecause its distribu-tion includes Lago Chungar. Chungar come from the

Aymara epithet chunkara (moss on the stone).

DISCUSSION

Vidal and Labra (2008) listed 88 species of Liolaemusin Chile, including L. alticolor, L. barbarae,and L. tacnaeofthe L. alticolor-bibronii group. Liolaemus barbarae is cur-rently considered a junior synonym of L. puna(Quinterosand Lobo, 2009), and Troncoso and Etheridge (2012) con-cluded that L. tacnaedoes not occur in Chile. Troncoso andEtheridge (2012) indicated that L. alticolor is distributed

Figure 2. Dorsal view of Liolaemus chungara sp. nov. (male paratypeFML 26509, SVL = 55.56 mm).

Figure 3.Ventral view of Liolaemus chungarasp. nov. (female paratypeFML 26512, SVL = 49.07 mm).




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in Parinacota, Putre, and Caquena. However, we foundL. chungara to be distributed in Putre, Colpita (which isclose to Caquena), and around Lago Chungar (which isnear Parinacota). As such, although we did not study thespecimens that allowed Troncoso and Etheridge (2012) topropose this distribution for L. alticolor,we assume that

the distribution of L. alticolorin Chile they proposed actu-ally corresponds to L. chungara.Similarly, Troncoso and Etheridge (2012) identified

a specimen from Chapiquia that had previously beenconfused with L. tacnae.They concluded that this speci-men is not L. tacnae, because the dorsal color pattern,

Figure 4.Map showing distribution of Liolaemus chungarasp. nov. andthe closely distributed species of the L. alticolor-bibronii group. Blacktriangle: L. alticolor;black square: L. aparicioi;black circle: L. chungara;white circle: L. paulinae;white triangle: L. puna;black diamond: L. tacnae;white diamond: L. walkeri.Arrow indicates type locality.

shape of dorsal scales, and number of scales around themidbody were different. Chapiquia is located 20 kmfrom Putre, so this specimen could be L. chungara. Con-sidering this and adding the new species described since2008, the number of species of Liolaemus distributed inChile (including L. chungara) is 89.

The presence of pores varies greatly among Liolae-mus species. Precloacal pores are usually restricted tomales, but in some species females present precloacalpores (e.g., L. aparicioi, L. ramirezae, L. yanalcu), and insomespecies of the subgenus Liolaemus males lack pre-cloacal pores (L. carlosgarini, L. coeruleus, L. cristiani,L. neuquensis, L. riodamas, L. thermarum,and L. tregenzai).One male L. chungaraexhibited both precloacal pores andsupernumerary pores. Supernumerary pores have beenreported in species of the L. montanus group, Eulaemussubgenus (i.e., L. huayra, L. orientalis, and L. orko;Quin-teros and Abdala, 2011). The presence of supernumerarypores in one specimen of L. chungarais the first record of

this character-state in the entire subgenus Liolaemus.

ACKNOWLEDGMENTS

We are grateful to F. Lobo for sharing invaluabledata. S. Portelli and J.M. Daz Gmez help us in lab and indiscussion of the manuscript. For granting access to col-lections under their charge, we thank to S. Kretzschmar,G. Scrocchi, and E. Lavilla (FML); J. Williams (MLP);J. Faivovich, B. Blotto, and S. Nenda (MACN); F. Lobo(MCN); J.C. Acosta, A. Laspiur, and E. Sanabria (UNSJ);

R. Aguayo and F. Valdivia (CBGR); J. Aparicio, A. Kirigin,and M. Ocampo-Vallivian (CBF); H. Voris and A. Resetar(FMNH); L. Ford and D. Frost (AMNH); J. Cadle and J.Rosado (MCZ); J. Wiens and E. Censky (CMNH); T. Reed-er and R. Etheridge (SDSU); K. de Queiroz and R. Heyer(USNM); D. Wake and B. Stein (MVZ); and R. Etheridge(SDSU). We acknowledge the Servicio Agricola Ganaderoof Chile for providing a collecting permit (306). ASQ wassupported by PICT 2400, Agencia de Promocin Cientficay Tecnolgica. CSA was supported by PICT 2263, Agenciade Promocin Cientfica y Tecnolgica. PV was supportedby PV y CA a UTA Mayor de Investigacin Cientifica y Tec-

nologica, Project N 471313.

REFERENCES

Abdala C.S. 2002. Nuevo Liolaemus (Iguania: Liolaemidae)perteneciente al grupo boulengeri de la provincia de Neuqun,Argentina. Cuadernos de Herpetologa16:313.

Abdala C.S. 2003. Cuatro nuevas especies del gnero Liolaemus(Iguania: Liolaemidae), pertenecientes al grupo boulengeri, de laPatagonia Argentina. Cuadernos de Herpetologa17:332.

Abdala C.S. 2007. Phylogeny of the boulengeri group (Iguania:Liolaemidae, Liolaemus) based on morphological and molecularcharacters.Zootaxa1538:184.




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APPENDIX

Specimens examined

Voucher numbers for lots are followed by the number of specimens in parentheses.

Liolaemus alticolor.BOLIVIA: Departamento de La Paz:Near Tiaguanacu, MCZR 169064 (lectotype), MCZR 7287 (para-lectotype); Rio Huarocondo, MCZ 12409; Tiahuanaco, near Lake Titicaca, MCZR 128518525; Tiahuanaco, 16330S,68420W, CBF 2925, 28932896.

Liolaemus abdalai.ARGENTINA: Provincia de Neuqun:Ruta Provincial 23.8 km N de Pilolil, Orillas del Rio Alumin,392229S, 705721W, MCN 2741 (holotype), MCN 273940, 274243, FML 784344; Ruta Provincial 11, 0.2 km aloeste de Arroyo Remec, 3902S; 7121W, MCN 274450; Arroyo Quilanlahue, Parque Nacional Lanin, Lacar, FML 1776.

Liolaemus bibronii.ARGENTINA: Provincia de Santa Cruz:Gruta de Lourdes, 2 km E Ruta Provincial 281, 11.2 km NWPuerto Deseado, FML 10106107.

Liolaemus bitaeniatus.ARGENTINA: Provincia de Catamarca:Agua de las Palomas, FML 1932, 3593; Quebrada Pea LaHorqueta - Distrito Espinillo, FML 6347; Confluencia de ros Quebrada y Candado, Andalgal, FML 7137 38. Provincia

de Salta:San Fernando de Escoipe, Chicoana FML 1655 (6). Provincia de Tucumn:Sierra de Medina, FML 2237 (2),2345 (2); Dique La Angostura, El Mollar, FML 2384 (4), 2499 (2); El Mollar, FML 2462, 2475 (5); Cerro Las Botijas, Sierrade Medina, MCN 900 901.

Liolaemus chaltin.ARGENTINA: Provincia de Jujuy:Departamento Cochinoca: 3 km NO de Abrapampa, FML 1461 (5);Abrapampa FML 1871 (4), 2513 (2); Ruta Provincial 71, 4.2 km W de Abrapampa, 224224.4S, 654312.4W, FML 9874(holotype); 2 km N Abrapampa, 2319673S, 6605399W, MCN 235; 2.9 km de la interseccin entre ruta 7 y 71, caminoa Cochinoca, 6,8 km W de Abra Pampa, 224559.4S, 654454.7W, MCN 222131.

Liolaemus chungara. CHILE: Regin de Arica y Parinacota: Entrada a Putre, 181053.4S, 693158.6W, 3874 m,FML 26505 (holotype), FML 26506 (paratype); southern shore of Lago Chungar, 181617.0S; 690937.7W, 4583 m,FML 26504508, MCN 473435 (paratypes); Provincia de Parinacota, km 12.5 de la ruta andina A93 Parinacota, Visviri,

1808.115S, 6918.000W, CZZA 346350 (paratypes); cerca de Putre desvo a Colpita, FML 26509513 (paratypes).

Liolaemus exploratorum.ARGENTINA: Provincia de Santa Cruz:MLP.S 571 (holotype), 567, 570, 573 (paratypes).

Liolaemus fuscus.CHILE: MACN 1671823, 21621. Regin de Coquimbo:Coquimbo, AMNH 131833834, MCZ 165146.Regin Metropolitana:El Cerezo, MCZ 65395; La Calera, Aconcagua, MCZ 165150; road to La Disputada, 1.5 km pastturn off Forest Los Farellones, 33.33333S, 70.36667W, MVZ 187797; on road to La Disputada Mines (33.36667S,70.38333W, MVZ 187804; road to Farellones, 33.35S, 70.35W, MVZ 196546548, 196550, 196559, 196562, 196565,196574575, 196581. Regin de Valparaiso:Baha Oscuro, FML 1592(2); Parque Nacional Campana, SDSU 1866; Val-paraso, MCZ 38621626, MCZ 165147.

Liolaemus gracilis.ARGENTINA: Provincia de Buenos Aires:Mar del Sur, General Alvarado, MCN 215658, UNMDP 320,326, 474. Provincia de San Luis:Estancia el Centenario, MLP.R 5306. Provincia de Chubut:40 km N de Trelew, MCN1345. Provincia de La Pampa:km 38, Ruta Nacional 28, Curac, FML 8371. Provincia de Mendoza:Tupungato, FML00963 (3); Malarge, FML 02731; San Rafael, FML 723436, 7238. Provincia de Rio Negro:Adolfo Alsina, Caleta de LosLoros, FML 2970 (2); El Cuy, FML 8399; Valcheta, MLP.R 1692.

Liolaemus incaicus.PERU: Departamento de Calca:near Calca, Hacienda Urco FMNH 266542 (holotype), FMNH 34104,FMNH 34127 (14) (paratypes); Sicuani, AMNH 38068070.

Liolaemus lemniscatus.CHILE: Regin de BioBio:Concepcin, CMNH 64727, 64730, MCZ 164037038; 164041; 164045;164047; 164049; 164056; 164059060; 164062064; FMNH 214220230; Escuadrn, Concepcin, CMNH 64728; Cura-nilahue, Arauco, CMNH 64729, USNM 58710. Regin de Coquimbo:Coquimbo, FML 1559 (2). Regin Metropolitana:Santiago, USNM 165620.




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Liolaemus pagaburoi.ARGENTINA: Provincia de Tucumn: Trancas FML 1613233; Huacahuasi, Tafi del Valle, FML16838; Tafi del Valle, FML 1829 (4); Puesto el Muoz, Taf del Valle, FML 2435(9); Hualinchai, Trancas, FML 2722 (4) 2746(11) FML 2633 (8).

Liolaemus paulinae.CHILE: Regin de Antofagasta:Orillas del Rio Loa, Calama, FML 1196 (paratype), 1341 (2); shore ofLoa River, SDSU 190911, MZUC 19360, 19362367, 193671, 19370, 19382.

Liolaemus pseudolemniscatus.CHILE: Regin de Coquimbo:MNHNC 137677, 1501, 1531.

Liolaemus puna.ARGENTINA: Provincia de Jujuy:Camino a Laguna Blanca, FML 929; Susques, FML 1265; Laguna lar-ga, Rinconada, FML 1512; Cuesta de Fundiciones, camino a Mina Pirquitas, Rinconada, FML 1517 (3); Rinconada, FML1519 (2); Pampa de los Pozuelos, Abra Pampa, Rinconada, FML 1533 (8); Abdn Castro Tolay, Cochinoca, FML 1874;

Abdn Castro Tolay, 231967.3S, 660539.9W, MCN 229232; Casa Mocha, subiendo al NW del Nevado del Chai,MCN 69899; 2.5 km al SE de Susques, Por Ruta 16 hacia Salinas Grandes, MCN 171819. Provincia de Salta:QuebradaLos Berros, 5 km E of Olacapato, Los Andes, 240835S, 664205W, FML 1364 (holotype), FML 991427 (paratypes);Cuesta del Acay, La Poma, FML 1661 (5), 1663 (9); Quebrada de Los Berros, Olacapato, FML 2779 (2); Campo Amarillo,Los Andes, FML 3647; Camino a Sey, La Poma, FML 3348 (2); Cerro Verde, Los Andes, FML 3649; Camino al Abra del Acay,MCN 94950; km 210, Ruta Nacional 51. 0.6 km de ruta 51 hacia las rocas. A 6,4 km Sur de ruta de ingreso a Olacapato,24.2410147S, 66.67711W, MCN 189092, 189497; 10 km oeste de Escuela Las Arcas, Cachi Adentro, camino al cerro de

la virgen, 250240.2S, 661642.0W, MCN 217779; Olacapato, Los Andes, 240821.3S, 66423.71W, SDSU 357982.CHILE: Regin de Tarapaca:Chiapa, SDSU 169799, MCZ 149852, 14985456, 149858; Volcn Tato, USNM 165641,MZUC 19392 (3). Regin de Atacama:San Pedro de Atacama, MNHN 583, 585, 588.

Liolaemus pyriphlogos. ARGENTINA: Provincia de Jujuy: vicinity of Laguna Leandro, Humahuaca Department,230150S; 651446,8W, FML 18199 (holotype), FML 18198, 18200201(paratypes); 10 km antes de Aparzo, desdeHumahuaca (230950,5S, 651148W), FML 18208210; afueras de Chorcn, camino a Laguna Leandro, FML 18236;camino a Mudana desde Uqua, (232030S, 651327,5W), FML 18250252; entre Aparzo y Humahuaca (231009.3S,651101.4W), FML 18258259; entre Chorcn y Laguna Leandro (230157.5S, 651414.3W), FML 18260262; Cami-no de Humahuaca a Chorcn, 2310761S, 6511709W, MCN 226, 228, 58998; Laguna Leandro, W of Chorcn, Huma-huaca, FML 1463 (32); Camino a Laguna Leandro, Humahuaca, FML 348889.

Liolaemus ramirezae.ARGENTINA: Provincia de Catamarca:Mina Capillitas, Andalgal, FML 3612; Morro El Arenal, elIngenio, Andalgal, FML 2561 (3). Provincia de Salta:La Poma, FML 1658, MCN 173335; 21 km N de La Poma, FML3006; Santa Rosa de Tastil, Rosario de Lerma, FML 3335. Provincia de Tucumn:km 98, ruta provincial 307, Amaichadel Valle, Taf del Valle FML 2240, 1367, 2275 (4), 2279 (2), 2330 (4), 2383 (2), 238486, 2436, 2463, 2481, 2486; Rutaprovincial 307, O de El Infiernillo, Tafi del Valle, FML 2715; km 98.5 ruta provincial 307, Tafi del Valle, FML 8182; km 95de Ruta provincial 307, Tafi del Valle, FML 6012, 17438; km 95, Ruta Provincial 307 (2640.82S, 6548.74W), MCN 466,46970.

Liolaemus robertmertensi.ARGENTINA: Provincia de Catamarca:Extremo septentrional del Bolson de Pipanaco, PuestoRo Blanco, a 3 km del Salar, 30 km S de Andalgal, Pomn, FML 1308 (3), 1482; Salar de Pipanaco, Pomn, FML 1478(2); Los Nacimientos, Beln, FML 16442; Estancia Ro Blanco, Salar de Pipanaco, Pomn, FML 6786 87, 16791; Hualfn,Belen, FML 1753; Toma del Ro Andalgal, Andalgal, FML 6782 83, FML 6785; Quebrada al N de Termas de Fiambal,Tinogasta, FML 7710; Salar de Pipanaco, MLPS 987, MCN 1713. Provincia de La Rioja:Quebrada de Santa Cruz, Cas-tro Barros, FML 944143; a 600 m de Puesto Vallecito, camino a la Mexicana, Famatina, MCN 163839; 290723.4S,673846.1W, MCN 174748; 290723.4S, 673846.1W, MCN 175455; Sierra de Velasco, MCN 195456.

Liolaemus sanjuanensis.ARGENTINA: Provincia de San Juan:Sierra de Pie de Palo, FML 1016 (paratype), UNSJ 73549,766.

Liolaemus saxatilis.ARGENTINA: Provincia de Crdoba:Achiras, Ro Cuarto, AMNH 65193199, MCN 90305; Achiras,Departamento Ro IV, SDSU 173637; Achiras (3310S; 5769W), SDSU 126616, MLP.S 1166167.

Liolaemus tacnae.PERU: Departamento Arequipa:Ro que cruza el camino a 15 mts de Arequipa (camino Arequipa-Puno), Arequipa, FML 1544. Departamento Tacna:Mina Toquepala, MCZ 45806 (holotype), 4580708 (paratypes).




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Liolaemus tandiliensis.ARGENTINA: Provincia de Buenos Aires:Sierra de los Padres, MCN 160405, 1612, 161415;Sierra de los Difuntos, MCN 160611, 161617; Sierra La Brava, MCN 1613.

Liolaemus variegatus.BOLIVIA: Departamento de Cochabamba:FML 1210 (2) (paratypes), CBGR S/N (4), CBGR 116,118, 121, 124, 130, 13239, 145, 15053, 9092, 122, MLP.S 841.

Liolaemus walkeri.PERU: AMNH 88324326. Departamento de Ayacucho:Rapi, Ayacucho, 13.0975S, 73.8136W, MCZ45850; Huancavelica, FMNH 81380389; 81395396. Departamento de Junin:6 km NE Paccha, SDSU 1937; Junn,AMNH 63389390 (paratypes); Llocllapampa, 11.82014275S, 75.62435913W, MCZ 43770775, 43777; Mina Juanchis-cochas, 40 km N de Jauja, 11.59924507S, 75.46884918W, MCZ 43779781; Maraynioc, 45 mi NE Tarma, 11.3667S,75.4W, MCZ 45887888; Casa Pato, MCZ 100111, FML 371 (2). Departamento de Apurimac:Puna cerca de Abancay,FML372; Runa ms arriba de Abancas, FML 1283. Departamento de Lima:Ticlio, 11.57843876S; 76.1938858W, MCZ45783.

Liolaemus yanalcu.ARGENTINA: Provincia de Salta:Camino al Acay desde Estacin Muano (7 km), 56 km de San Anto-nio de los Cobres, MCN 95560, 1038, 1635; Camino al Acay desde Estacin Muano, 56 km, 2418316S 6609070W,MCN 36061, 53536, 541; Camino al Acay desde Estacin Muano, 89 km, MCN 702, 70507; Camino al Acay desdeEstacin Muano, MCN 72526; km 148, Ruta Nacional 51, E de San Antonio de los Cobres, 241315.9S, 661546.4W,MCN 72829; 5 km al S de Estacin Muano, camino al Acay, MCN 1449; 6.7 km O de Estacin Muano, Camino al

Acay (242047.5S, 660933.9W, MCN 1750; 7 km desde Estacin Muano hacia el Nevado del Acay, 242051.8S,660927.2W, MCN 223639, 2501, 261316.



Quinteros Et Al 2014

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