2003 new species_liolaemus

March 2003] 105HERPETOLOGICA

Herpetologica, 59(1), 2003, 105–118q 2003 by The Herpetologists’ League, Inc.

A NEW SPECIES OF LIOLAEMUS FROM SOUTHERN BRAZIL(IGUANIA: TROPIDURIDAE)

LAURA VERRASTRO1,2,4, LAUREN VERONESE1, CLOVIS BUJES1, ANDMANOEL MARTINS DIAS FILHO2,3

1Setor de Herpetologia, Departamento de Zoologia, Universidade Federal do Rio Grande do Sul. Rua PauloGama, 40–Predio 12105, 90040–900, Porto Alegre–RS, Brazil

2Programa de Pos-Graduacao em Ecologia a Recursos Naturais, Universidade Federal de Sao Carlos,Rod. Washington Luiz, Km 235–13565-905–Sao Carlos–SP, Brazil

3Departamento de Ecologia e Biologia Evolutiva, Universidade Federal de Sao Carlos, Rod. WashingtonLuiz, Km 235–13565-905–Sao Carlos–SP, Brazil

ABSTRACT: We describe a new species of the genus Liolaemus from Rio Grande do Sul, Brazil.The morphology of the new species, L. arambarensis sp. nov, is compared to other species ofLiolaemus of the ‘‘wiegmannii’’ group. The new species is distinguished from others members ofthis group by a dorsal color pattern with a medium dorsal white stripe and two dorsolateral stripes,two rows of juxtaposed brown marks resembling triangles, delimited by a white bar; nasal scalesoriented dorsally; a complete row of dorsally oriented lorilabial scales between the subocular andsupralabial scales; a moderate number of scales around the mid-body; flat temporal scales; flat headscales; and a transversely divided frontal. Liolaemus arambarensis sp. nov. is omnivorous, oviparous,and exhibits sexual dimorphism in color and size.

Key words: Brazil; Karyotype; Liolaemus; Lizard; New species; Restinga habitat; Tropiduridae

LIZARDS of the genus Liolaemus (Tro-piduridae) include 150 recognized spe-cies of small to moderate size (Etheridge,1995, 2000). They live in a variety of hab-

4 CORRESPONDENCE: e-mail, [email protected]

itats from sea level to over 5000 m (Eth-eridge, 2000; Etheridge and de Queiroz,1988); in extensive areas of aeolean sand,including the sandy beaches of Chile, Ar-gentina, Uruguay, and southeastern Bra-zil; and the sand flats and dune systemsscattered throughout much of the inte-

106 [Vol. 59, No. 1HERPETOLOGICA

rior of Argentina and Chile (Etheridge,1993).

The wiegmannii group is characterizedby the presence of lorilabial scales, whichare smaller than the supralabial scales, andusually by the presence of two rows of su-pralabial scales between the subocular andthe supralabial scales. The supralabialscales are narrow, but the posterior onesare elongated. The sublabial scales are incontact with the mental scale, which iswider posteriorly. The infralabial scalesvary from flat to concave (Etheridge, 1995,2000).

According to Etheridge (1995, 2000),this group includes nine predominantlypsammophilous species: L. occipitalis,which occurs in the southern region ofBrazil and southern Santa Catarina state(de Lema, 1994; Peters and Donoso-Bar-ros, 1970); L. lutzae from the coast of Riode Janeiro, Brazil (Rocha, 1985); and L.wiegmannii, which is widely distributed inArgentina (Cei, 1986) and Uruguay (Gu-dynas, 1981a,b,c). Other species assignedto this group also occur in Argentina: L.scapularis in the arid plains of the prov-inces of Catamarca and Tucuman; L. mul-timaculatus from the coastal region ofBuenos Aires and Rıo Negro; L. rabinoifrom San Rafael Department, Mendoza;L. riojanus from La Rioja and San Juan;and L. salinicola from Catamarca (Cei,1986). Lastly, L. cranwelli occurs in SantaCruz de la Sierra, Bolivia (Donoso-Barros,1973). Etheridge (2000), in his recent pa-per on the wiegmannii group, considers L.cranwelli a synonym to L. wiegmannii.

Peters and Donoso-Barros (1970), Mull-er (1979), and de Lema (1994) recordedL. occipitalis and L. wiegmannii from RioGrande do Sul (southernmost state of Bra-zil). During field work conducted at themargins of the Laguna dos Patos in RioGrande do Sul, Brazil, lizards formerlyconsidered L. wiegmannii by Peters andDonoso-Barros (1970) and de Lema(1994) were captured. After detailed ex-amination of their dorsal color pattern andlepidosis, these specimens were judged torepresent a new species of the wiegmanniigroup (sensu Etheridge 1995, 2000).

Chile was incorrectly registered as the

type locality of L. wiegmannii by Dumeriland Bibron (1837). Bell (1843) lists thepresence of this species in Bahıa Blanca,Rıo Negro, Argentina, and Maldonado,Uruguay, as part of the sample collectedin that region by Charles Darwin. Theselocalities were confirmed by Gallardo(1966). The distribution of L. wiegmannii,according to Cei (1979a, 1986), Laurentand Teran (1981), Cabrera and Bee deSperoni (1986), and Tedesco et al. (1992),is broad and includes Argentina (EntreRıos, Buenos Aires, Bahıa Blanca, LaPampa, Corrientes, San Luis, Mendoza,Tucuman, and Jujuy), Uruguay, and south-ern Brazil. Liolaemus wiegmannii is anomnivorous species with oviparous repro-duction (Cei, 1986; Donoso-Barros, 1973).

We were unable to assign the speciesdescribed here to L. wiegmannii, or anyother species of Liolaemus. Thus, we heredescribe this new species based on mate-rial collected in southern Brazil and ana-lyzed at the Herpetology Laboratory of theFederal University of Rio Grande do Sul.

MATERIALS AND METHODS

Meristic and Morphometric Analysis

The samples examined in this study in-cluded 91 specimens of Liolaemus sp.nov., 59 of L. occipitalis, 32 of L. wieg-mannii, and 9 of L. lutzae. The mass (g)of each individual of L. arambarensis sp.nov. was recorded, and the lizards wereimmediately sacrificed with 3% Citanestanesthetic. Specimens were then fixed in10% formalin, transferred to 70% ethanolafter 72 h, and deposited at the Labora-torio de Herpetologia, Universidade Fed-eral do Rio Grande do Sul (UFRGS).

Meristic data were taken under stereo-microscope, and nomenclature followsSmith (1946). The following variables weretaken for morphometric analysis: SVL(snout–vent length), HL (head length),HW (head width), AX–GR (axilla groindistance), FL (foreleg length), HLL (hindleg length), and TL (tail length). All mea-surements were taken with a 0.1-mm pre-cision caliper. The presence (and number)or absence of precloacal pores was also re-corded.


FIG. 1.—Adult specimens of Liolaemus arambar-ensis from Arambare, Rio Grande do Sul, Brazil (leftfemale, UFRGS 2951, SVL 5 55.85 mm; right male,UFRGS 2902, SVL 5 59.84 mm 5 holotype).

Data for the remaining species of thewiegmannii group (L. cranwelli, L. sca-pularis, L. multimaculatus, L. rabinoi, L.riojanus, and L. salinicola) were obtainedfrom the literature (Boulenger, 1885; Cei,1979b, 1986; Donoso-Barros, 1973; Du-meril and Bibron, 1837; Laurent, 1982,1986; Etheridge, 2000).

Cytogenetic AnalysisMaterial for cytological preparations was

obtained from the bone marrow, spleen(males and females), and testes of 10 spec-imens. The analyses were performed afterGiemsa staining. The C-banding patternswere obtained according to Sumner(1972), and the silver staining of nucleolusorganizer regions (Ag–NORs) followedHowell and Black (1980).

Natural HistoryWe examined the gonads and stomach

contents from all 91 specimens of L. ar-ambarensis. The gonads were measured,and their anatomical stage was verified bydetermining the presence of eggs in theoviducts. For observation of the reproduc-tive stages, histological sections were pre-pared, using the hematoxilin and osintechnique; histological preparations wereobserved under a microscope (25–103and 64–103). In males, the presence ofspermatozoa in the seminiferous tubulesand in the epididymis was observed inspecimens collected throughout the year,as was presence of mature and vitelogenicfollicles in females.

The digestive tract was examined understereomicroscope for the presence of preyitems. Arthropod prey items were identi-fied to Order. Frequency of occurrenceand volume of each food item was calcu-lated according to Rocha (1989). Plant ma-terial was identified as vegetative parts,fruits, and flowers.

RESULTS AND DISCUSSION

Liolaemus arambarensis sp. nov.Holotype.—UFRGS 2902, an adult

male collected by L. Veronese 14 Decem-ber 1996 in a restinga at the municipalityof Arambare, State of Rio Grande do Sul,Brazil (308 559 S, 518 309 W).

Paratypes.—All specimens from thetype locality: males: UFRGS 2903–07; fe-males: UFRGS 2908–13, collected by L.Veronese on 14 December 1996; males:UFRGS 2946, 2950, 3044, 3112–13, 3115,3118; females: UFRGS 2945, 2947, 2951,3114, 3116, 3120, collected by L. Verrastroon 25 January 1997; UFRGS 3044 femalescollected by C. Bujes on 26 April, 1997;UFRGS 3112–20 females collected by C.Bujes on 30 July 1997.

Diagnosis.—Liolaemus arambarensishas the following diagnostic characters: na-sals oriented dorsally; a complete row oflorilabial scales between subocular and su-pralabial scales; temporal and head scalessmooth; frontal usually divided transverse-ly; dorsal color pattern consisting of a mid-dorsal white stripe and two dorsolateralstripes, with two series of paravertebralbrown marks resembling triangles bor-dered by white bar (Figs. 1, 2).

Liolaemus arambarensis is referrable to


FIG. 2.—(A) Dorsal and (B) lateral view of thehead of adult male Liolaemus arambarensis (UFRGS2902; holotype). Scale 5 5 mm.

the ‘‘wiegmannii’’ group based on thepresence of lorilabial scales smaller thansupralabial scales and narrow (longer thanwide) supralabial scales, the posterior onesbeing wider; the mental scale is in contactwith the sublabial scales and is posteriorlywide; infralabial scales are concave.

Liolaemus arambarensis is distinguishedfrom all other species of the wiegmanniigroup by their distinct pattern of dorsaland ventral color and, except for L. lutzae,by having a single row of lorilabial scalesbetween the subocular and the supralabialscales. The new species is distinguishedfrom L. wiegmannii, which has laterallyoriented nostrils, by having dorsally ori-ented nostrils; the new species has headand lateral body scales that are smooth, incontrast with the keeled or rugose scalesof L. wiegmannii. The frontal scale is pre-sent and transversally divided, while it isnot divided in L. wiegmannii and is absentin L. occipitalis and L. lutzae. The dorsal

scales are markedly keeled and imbricatein the new species, similar to L. cranwelli,L. lutzae, L. occipitalis, L. riojanus, L. sca-pularis, and L. wiegmannii, but differentfrom other species (i.e., L. multimaculatus,L. rabinoi, and L. salinicola), which havesmooth or slightly crenate dorsal texture.Lateral scales in the new species are sim-ilar to those of L. lutzae, L. occipitalis, andL. wiegmannii, having a common patternwith distinct dorsal and ventral regions,while in other species of the group thispattern is not observed (Donosos-Barros,1973; Boulenger, 1885; Cei, 1979a, 1986;Laurent, 1982, 1986).

The comparison of scale counts (Table1) and biometric variables (Table 2) amongthe species of the wiegmannii group indi-cates that L. arambarensis is a small spe-cies like L. scapularis and L. wiegmannii.The statistical comparison between thenew species and the three most geograph-ically proximate species (L. lutzae, L. oc-cipitalis, and L. wiegmannii) revealed sig-nificant differences between the means ofall variables. These species are easily dis-tinguished by their the dorsal color pattern(Boulenger, 1885; Mertens, 1938; Cei,1986; Fig. 1).

Description of holotype.—Adult male.SVL 59.8 mm; HL 13.7 mm; HW 10.8mm; TL 64.0 mm. Dorsal head scalessmooth. Width of rostral scale approxi-mately twice its length. Rostral length de-limited posteriorly by four postrostrals,half the lateral pair overlapping the su-pralabials. Nasals oriented dorsally; nos-trils in posterior half of nasal scale, in dis-tinct dorsal position separated by two pairsof internasals; 10 frontonasal scales rela-tively small, situated between the posteriorcanthals; supraorbital semicircles distinct,formed by irregular scales with interme-diate contact, anterior region in contactwith transversally divided frontal scale. In-terparietal polygonal, with distinct opales-cent, parietal eye in contact anteriorly withsupraorbital semicircle; parietal eye sepa-rated from suborbital semicircle by singlescale, which is limited posteriorly by threeirregular, smaller parietal scales. Mid-su-praocular large, longer than wide, mediallyseparated from the supraorbital semicircle


TA

BL

E1.

—Sc

ale

coun

tsin

spec

ies

ofth

eL

iola

emus

‘‘wie

gman

nii’’

grou

p.

Dor

sal

scal

es

Scal

esar

ound

the

body

Lon

gitu

dina

lro

ws

ofdo

rsal

scal

esV

entr

alsc

ales

Pre

-clo

acal

pore

s

fm

Infr

adig

ital

scal

esin

the

four

than

teri

oran

dpo

ster

ior

finge

r

ant

post

Ori

gin

L.

aram

bare

nsis

57–6

460

–66

15–1

851

–60

3–4

4–7

14–1

920

–25

Ara

mba

re/R

S.82

spec

imen

sex

amin

edin

this

stud

yL

.cr

anw

elli

57—

——

——

23—

Don

oso-

Bar

ros,

1973

L.

lutz

ae66

–72

58–6

822

–30

61–6

80

618

–21

24–2

9R

iode

Jane

iro/

RJ.

9sp

ecim

ens

exam

ined

inth

isst

udy

L.

mul

tim

acul

atus

—72

–74

—26

–28

08

0—

Bou

leng

er,

1885

;C

ei,

1979

aL

.oc

cipi

talis

75–8

667

–79

21–2

661

–78

07–

1018

–22

21–2

5Ja

rdim

Ede

m/R

S.59

spec

imen

sex

am-

ined

inth

isst

udy

L.

rabi

noi

—74

–78

——

06–

724

—C

ei,

1974

L.

rioj

anus

—75

——

010

22–2

3—

Cei

,19

79a

L.

salin

icol

a75

65—

890

1018

23L

aure

nt,

1986

L.

scap

ular

is26

–34

54–6

0—

23–3

02–

66–

1018

–21

22–2

6L

aure

nt,

1982

L.

wie

gman

nii

(Arg

enti

na)

50–5

252

–63

15–1

852

–60

3–5

5–6

14–1

918

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Cor

doba

/Arg

enti

na.

16sp

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ex-

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.wie

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(Uru

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713

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620

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onia

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TABLE 2.—Morphometric measurements for species of the Liolaemus ‘‘wiegmannii’’ group. SVL 5 snout–vent length; TL 5 tail length; HL 5 head length; HW 5 head width; FL 5 foreleg length; HiL 5 hind leg

length; A–G 5 axilla–groin distance.

SVL TL HL HW FL HiL A–G Origin

L. arambarensis 50.21 54.60 10.92 8.21 27.12 18.83 22.95 Arambare/RS. 69 specimensL. cranwelli 57.5 13.0 15.0 27.2 Donoso-Barros, 1973L. lutzae 63.44 79.27 14.89 10.77 37.71 24.64 29.32 Rio de Janeiro/RJ. 9 specimensL. multimaculatus 52.0 68.0 16.0 13.0 25.0 39.0 Boulenger, 1885; Cei 1979aL. occipitalis 56.67 63.35 12.59 9.80 21.53 32.24 26.76 Jardim do Edem/RS. 59 speci-

mensL. rabinoi Cei, 1974L. riojanusL. salinicola

61.573.0

72.098.0

14.514.5

12.5 22.0 36.028.0

28.033.0

Cei, 1979aLaurent, 1986

L. scapularis Laurent, 1982L. wiegmannii

(Argentina)48.93 57.73 11.78 8.81 27.12 18.2 24.32 Cordoba/Argentina. 16 speci-

mensL. wiegmannii

(Uruguay)49.03 61.2 12.13 8.81 29.02 18.99 23.4 Colonia-Montevideo/Uruguay. 16

specimens

by complete arching row of circumorbitals,laterally separated from supraciliaries bythree rows of small supraoculars. Two can-thals, posterior canthal larger and longerthen anterior canthal. Orbit bordered su-periorly by four elongate, obliquely over-lapping superciliary scales, followed by twoshort overlapping scales in the opposite di-rection. One preocular single, equal inheight and length, preceded by a curvedpreocular. Subocular elongated, borderedsuperiorly by short postocular, both scaleswith sharp keel along superior margins.Small eyelids situated immediately belowzone of small ciliaries under the eyelid,particularly in the medial portion. Internalciliaries granular; external ciliaries slightlyprojected: 3, 4 loreals. Seven lorilabials incomplete rows separate anteriorly supra-labials from loreals and posteriorly sepa-rating the supralabials from suboculars.Lorilabials separated from subocular bytwo small scales. Anterior lorilabials high,shorter than underlying supralabials,which are narrower posteriorly. All foursupralabials longer than high, with straightedges not covering anterior infralabials;posterior supralabial more elongated thanothers, preceded by two postlabials, theanterior one longer, forming curved suturewith posterior supralabial. Temporalssmooth, anteriorly juxtaposed, and sub-im-bricate posteriorly. The opening of the ex-ternal auditory meatus higher than wide,bordered anteriorly by single series of

small, projecting, convex auricular scales.Mental wider than rostral (2.4 mm wide 31.2 mm long), delimited posteriorly by twopostmentals, and laterally by the infrala-bials. Five infralabials, the second, third,and fourth with distinctively concave ex-ternal surfaces, and with keel on edge ofthe suture bordering the anterior sublabi-als, in inferior contact with sublabials 2, 3;anterior sublabials and post-mentals not incontact. Three pairs of chin shields sepa-rated anteriorly by three small scales di-verging posteriorly. Nine overlapping pos-terior gular scales between antehumeralfolds, flat, smooth; 55 ventral scales, eachlonger than wide, and larger than smooth,overlapping dorsal scales. Dorsal scales in18 longitudinal rows. Anterodorsal nuchalssmooth; dorsal scales of the body distinctlykeeled; lateral scales longer than wideoverlapping. Dorsal nuchals in 19 longi-tudinal rows. Nine posterior gulars amongthe antehumerals folds. Sixty-three scalesaround the body. Lateral superior scaleskeeled; inferior lateral scales smooth, sit-uated posteriorly in V-shape. Dorsal scalesof the limbs keeled similarly to bodyscales, becoming granular at base of limband frontal portion, larger, smooth, over-lapping ventrally; infratarsal scales withblunt keels, sub-imbricate, and obovate.Eighteen infradigital scales on the fourthright finger and 22 in the fourth right toe;subdigital lamellae on feet, with pointedkeels and spines (mucrons) strongly pro-


TABLE 3.—Morphometric measurements taken fromLiolaemus arambarensis; SVL 5 snout–vent length;TL 5 tail length; HL 5 head length; HW 5 headwidth; FL 5 foreleg length; HiL 5 hind leg length;A–G 5 axilla–groin distance; min 5 minimum; x 5mean; max 5 maximum; SD 5 standard deviation; n

5 number of individuals.

Measure-ments Min x Max SD n

SVLTLHLHWFLHiLA–G

40.7033.20

6.804.409.70

15.6017.96

50.2158.2510.92

8.2118.8327.1222.95

60.0079.8013.3010.5022.3033.6028.10

4.698.371.131.142.133.022.44

41354141414141

jected, resulting in a spiny aspect. Fourpre-cloacal pores.

Color in life.—Head predominantlybrown with some marbled orange scales.Two white stripes originating dorsally onthe supraocular scales and extending later-ally, diverging and interrupted near theneck. Two brown lines (width 1.5–2 scales),originating on the posterior border of eachorbit delimiting the area of dorsal stripes.Two circular, dark brown spots, similar toocelli, situated on each side of the inter-parietal. Two distinct, dorsolateral stripes(width 3.5–4 scales) beginning on the neckand extending to tip of tail. A distinct mid-dorsal stripe (width 1.5–2 scales) extend-ing from neck to tip of tail, where it grad-ually joins the two dorsolateral stripes. Aseries of 9–10 brown lateral marks ar-ranged paravertebrally along mid-dorsalstripe. The mid-dorsal stripe is delimitedposteriorly by a band of white scales dis-tributed along the mid-dorsal stripe, re-sembling triangles, with the hypotenuse ly-ing on the mid-dorsal stripe. From thebase to the tip of the tail, seven paraver-teral marks become nearly fused, but re-main separated by the mid-dorsal stripe,which is reduced to a thin line. Dorsalbackground coloration is dark ferruginousorange. Lateral coloration marbled withbrown lines bordering cyan-blue spots onan orange background. Coloration is moreintense near the dorsolateral stripes andprogressively fades ventrally, where whitereplaces the orange coloration. The lateralportion of the body and infrafemoral areais bright orange. Precloacal pores brightorange. Immaculate white ventrally.

Color in preservative.—Specimens fixedin 10% formalin and stored in 70% ethanollost color within 30 d. Blue areas becamegray; the orange of the pores and lateraland dorsal portions of the body disap-peared; and the brown stripes and bodylost their intensity; white areas remainedunchanged.

Variation.—Maximum adult size formales and females is 60 mm and 56 mm,respectively (Table 3). Dorsal nuchalssmooth anteriorly; dorsal scales of thebody distinctly keeled; lateral scales largerin length than in width and overlapping;

60–66 scales around the body; 57–64 dor-sal scales; 51–60 ventral scales; dorsalscales of the body with 15–18 longitudinalrows; 14–19 infradigital scales in thefourth finger and 20–25 in the fourth toe.Precloacal pores orange, 4–7 in males and3–4 in females (Table 4). Only male spec-imens have orange coloration along thesides of the body and in the infrafemoralarea, and blue marks along the body sides.In both adult females and males, the colorpattern is similar to that of recentlyhatched specimens. Liolaemus arambar-ensis is sexually dimorphic, with males be-ing larger than females, possessing morecloacal pores, and exhibiting differences incolor pattern (Tables 5, 6).

Etymology.—The name arambarensisrefers to the type locality: Arambare, stateof Rio Grande do Sul, Brazil.

Distribution and ecology.—Liolaemusarambarensis sp. nov. is known only fromthe restinga habitats associated with Ho-locene sandbanks of the Coastal Plain atthe margin of the Laguna dos Patos, fromItapua (district of the city of Viamao) tothe city of Arambare (type locality), RioGrande do Sul, Brazil.

The type locality, Arambare County (308559 S, 518 309 W) in Rio Grande do Sul,Brazil, is near the mouth of Arroio Vel-haco, on the margins of the Laguna dosPatos. The climate of Rio Grande do Sulis humid subtropical (type Cfa in Koppen’sclassification). Annual rainfall ranges1100–1300 mm, and annual temperatureis 16–18 C (Moreno, 1961). The predom-


TABLE 4.—Scale count and pre-cloacal pores in paratypes of Liolaemus arambarensis.

Dorsalscales

Scales aroundthe body

Longitudinalrows of

dorsal scalesVentralscales

Pre-cloacalpores

Infradigital scales inthe fourth anterior and

posterior finger

ant post

Males2902290329052906290729462950304431123117

60636057606462605861

63646360636563606466

19191818181716191917

55535457525656515253

4456546674

18161717171818191817

22242423242225222521

Females2908290929102911291229132945294729513120

61585962616060626362

61616060606161606161

17151816161616151618

60585957545860576059

4344334444

18181616171617161714

22202024222220222322

TABLE 5.—Morphometric data for male and femaleadults of Liolaemus arambarensis (in mm). Means,standard deviation (SD), sample size (n) and the tvalues of Student’s t-test are presented for SVL(snout–vent length), HL (head length), HW (headwidth), and TBW (tail–base width). Asterisks indicate

significant sex differences (P , 0.01).

Variable

Females(n 5 30)

Mean 6 SD

Males(n 5 30)

Mean 6 SD t

SVLHLHWTBW

49.67 6 5.08410.66 6 0.877

8.06 6 0.7055.16 6 0.685

54.57 6 4.26112.07 6 0.840

9.30 6 0.6676.93 6 0.765

3.64∗5.55∗6.15∗8.44∗

TABLE 6.—Percentages of blue area in the dorso-lat-eral region, yellow area in the infrafemural region,orange area in lateral region, and number of poresfor both sexes of Liolaemus arambarensis (n 5 60).

Age/SexBlue(%)

Yellow(%)

Orange(%)

Numberof

pores

Juvenile maleAdult maleJuvenile femaleAdult female

28.283.3

025

50100

00

50100

00

4533

inant direction of the winds is northeast,which is related to the subtropical anticy-clone of the South Atlantic. The speed ofthe winds rarely exceeds 40 km/h; the con-stant effect gives an anemomorphic aspectto the trees and shrubs. Actually, the topof the trees become unilaterally distorted(Waechter, 1985).

The relief is basically plain, made of hol-ocenic sites of the Coast Plain. The soilsare mostly sandy and well drained withgrains between 1–2 mm of diameter,sometimes reaching 5 mm. The vegetation

gradually varies in size and density accord-ing to the distance from the margins of thelagoon (where L. arambarensis occurs);there are almost no plants in the first 5 m.Between 5–10 m, about 65% of the areais a sandy surface and the other 35% iscovered by herbaceous vegetation andsparse scrubs. The most abundant plantspecies are from the family Gramineae(Panicum racemosum, Rhynchelytrum re-pens, Paspalum notatum, Cynodon dacty-lon), followed by species from other fam-ilies like Eryngium nudicaule, Euphorbiapeploides, Oenothera sp., Polygala sp., andPetunia integrifolia. The most frequent


shrubs are Dodonaea viscosa and Cordiaverbenacea. Between 10–20 m from themargin of the lagoon, almost the same her-baceous community is present, with aslight increase of the abundance of shrubs.At 20 m from the margin, the vegetationis patchy, with a combination of herba-ceous and shrubby vegetation, and severalwoody insular vegetation clusters (‘‘mator-rais’’ and Sandbank Low Forest) and sandygrasslands. The Sandbank Low Forest is1.5–6 m high, surrounded by herbaceousspecies such as Polypodium lepidopterys,Aechmea recurvata, Peperomia peresqui-folia, Rumora sp., and shrubs like Hexach-lamis edulis, Lantana camara, Cereus hil-demanianus, Opuntia monacantha, andEupatorium sp. The epiphyte Smilax cam-pestris is also common. The most commontrees are Myrciaria cuspidata, Lithreaeabrasiliensis, Gomidesia palustris, Butiacapitata, and Myrsine umbellata (Fig. 3).

Cytogenetic data.—The karyotype of L.arambarensis (Fig. 4) indicates a diploidnumber of 2N 5 34. The karyotype has 12metacentric macrochromosomes and 22microchromosomes. The first pair of mi-crochromosomes is distinctly larger thanthe others. Analysis of chromosomes inmetaphase, stained with silver, indicatedthat NORs are located in the secondaryconstrictions of the long arm of Pair 2. Theknown diploid number for Liolaemus spe-cies varies between 2N 5 30 and 2N 5 44(Gorman, 1973). Among the species of thewiegmannii group, there are chromosomaldata for L. salinicola (2N 5 32), L. sca-pularis (2N 5 34; Navarro, 1992), L. oc-cipitalis, L. lutzae, and L. wiegmannii (2N5 34; Bertolotto et al., 1996), all with 12macrochromosomes and 20 or 22 micro-chromosomes. Thus, the karyotype ob-tained for L. arambarensis seems to beconservative with respect to other speciesof Liolaemus, but an analysis of chromo-some morphologies and banding patternsis necessary to characterize species karyo-types and contribute to the understandingof the relationship among Liolaeminaespecies.

Natural HistoryIndividuals of L. arambarensis appear to

mimic the substrate with their cryptic col-

or pattern and can easily dig holes with thehead and forelimbs to burrow themselves.The pattern used for digging the substrateand hiding corresponds to that in the sec-ond digging model described by Halloy etal. (1998). We include in this group thelizards that twist their body when burrow-ing in the sand with the help of the tailmovements. The species is able to moveunder the substrate (sand and litter) andchange position without being noticed bythe observer.

Construction of holes in the sand wasnot observed for this species in contrast tothe observations made of L. occipitalis(Verrastro and Bujes, 1998), possibly be-cause the restinga habitat provides morecover for escaping from predators and ref-uge for resting.

Liolaemus arambarensis is oviparousand the reproductive season is late Au-gust–March. The presence of spermatozoawas observed in the seminiferous tubulesfrom late August–late December, afterwhich males become reproductively inac-tive. The size of testes varies during theyear, being larger in the months of sper-matozoa production. The smallest male ex-amined with spermatozoa in the seminif-erous tubules and in the epididymis duringthe reproductive season measured 45.8mm (SVL). The smallest female with eggsin the oviducts measured 45.3 mm (SVL).Average clutch size was two eggs. Fromhistological analysis of the gonads, follicleswere present in the ovaries September–March, and the presence of eggs in theoviduct was observed October–January.Non-reproductive females (without vittel-ogenic follicles or eggs in the oviducts)were observed in March and October;non-ovigerous reproductive females (withvittelogenic follicles, but without eggs inthe oviduct) were found throughout theyear, except in January, April, August, andDecember. Ovigerous reproductive fe-males (with eggs in at least one of the ovi-ducts) were found in spring and summermonths (December–February). Field ob-servations revealed the presence of re-cently hatched individuals January–lateMarch. The reproductive cycle of L. ar-ambarensis is similar to that of other spe-


FIG. 3.—Habitat at the type locality of Liolaemus arambarensis at Arambare, RS, Brazil; note thesandy soil.

cies of the wiegmannii group. Liolaemusoccipitalis (Verrastro, 1991; Verrastro andKrause, 1994), L. lutzae (Rocha, 1989),and L. wiegmannii (Pinilla, 1991) are alsooviparous and have seasonal reproductivecycles with only one reproductive seasoneach year. Liolaemus multimaculatus (Gal-lardo, 1966), L. riojanus (Cei, 1986), andL. wiegmannii (Gallardo, 1966) are alsooviparous species. The diet of L. arambar-ensis consisted predominantly of insects,but is characterized as omnivorous be-cause of the presence of plant material inthe stomachs. The main prey items andtheir respective percent frequency of oc-currence and relative volume were: Hy-menoptera, 87.0%, 16.8%; Aranae, 80.1%,13.2%; and Coleoptera, 68.5%, 9.6%.Plant material included mainly flowers andleaves, which collectively corresponded toa frequency of 31.5% and relative volumeof 16.9%. The species of the ‘‘wiegmannii’’group are usually mentioned as insectivo-rous, but there are few detailed studies inthis respect. According to Rocha (1989,1991), L. lutzae presents an ontogenetic

shift in diet, with young specimens beinginsectivorous and adults being herbivo-rous. Another species, L. occipitalis, has adiet exclusively composed of arthropods,with no plant material (Verrastro, 1991).

Phylogenetic RelationshipsA phylogeny for the L. wiegmannii

group has been recently proposed by Eth-eridge (2000) and used herein to hypoth-esize the phylogenetic relationships of L.arambarensis. Among the characters ana-lyzed, Etheridge included several meristicand morphometric multistate characters,not discussed herein.

Liolaemus arambarensis appears to be-long to the L. wiegmannii group, definedby Etheridge (2000, his figure 9), by shar-ing the following characters with all spe-cies in his Branch 19 of his cladogram(numbers correspond to character list giv-en by Etheridge, 2000): Lorilabial scalessmall, more than seven between the pos-terior canthals (Character 4: State 1); su-pralabials narrow (Character 6: State 1);no contact between postmentals and the


FIG. 4.—Karyotype and site of active NORs in Liolaemus arambarensis (UFRGS 2369, male) from Ar-ambare, Rio Grande do Sul, Brazil. 2N 5 34 (12 M 1 22 m). Scale 5 10 mm.

infralabials (Character 11: State 1); snoutprofile flat (Character 32: State 1); and di-urnal escape behavior includes rapid burialunder sand (Character 36: State 1).

Among the species in the L. wiegmannigroup, L. arambarensis appears to be re-ferable to the ‘‘sand lizards’’ group (sensuEtheridge, 2000, his figure 9, Branch 20)by sharing the following characters: dorsalposition of the nasal scale (Character 2:State 1); proximal prebranchial scales withdistal granules below the apex, which caus-es the scales to project outward from thesurface to form a tridentate margin (Char-acter 15: State 1); and gravid female col-oration absent (Character 30: State 1).

Liolaemus arambarensis differs fromthe species in Branch 20 given in Ether-idge (2000, his figure 9) by the presenceof a different state for Character 28. Al-though State 2 (adult male coloration fine-

ly checkered dark gray on throat and ab-domen) is assigned as a synapomorphy ofBranch 20 and State 4 as a synapomorphyof Branch 21, L. arambarensis has a dif-ferent pattern, herein considered as auta-pomorphic of this species (State 3: adultmale ventral surfaces immaculate, or near-ly so).

Liolaemus arambarensis appears refer-able to Branch 21 by sharing the followingcharacters with those species: upper tem-poral scales smooth (Character 1: State 0);outer surface of second, third, and fourthinfralabial scales concave with a keelwhere they suture with adjacent sublabialscales (Character 9: State 2); subdigital la-mellae of pes with blunt keels, withoutstrongly projecting mucrons (Character21: State 1).

Branch 22 of the cladogram from Eth-eridge (2000, his figure 9) is defined by


FIG. 5.—Phylogenetic hypothesis proposed by Eth-eridge (2000) for the ‘‘wiegmannii’’ group, which sug-gests Liolaemus arambarensis as sister group for Lio-laemus Branch 24; future studies will be required toconfirm this hypothesis.

four polymorphic characters, not analyzedherein. Liolaemus arambarensis, however,appears referable to Branch 23 by sharingthe following synapomorphies: frontonasalscales small (more than seven between theposterior canthals; Character 3: State 1),although these scales are not as small as inL. occipitalis, there are more than sevenpresent; infratarsal scales surface smooth(Character 17: State 2); infratarsal scalessub-imbricate (Character 19: State 2);shape infratarsal scales of obovate (Char-acter 20: State 1).

All the characters discussed above in-clude L. arambarensis in a clade formedby this species and L. occiptalis, L. rabi-noi, L. multimaculatus, and L. riojanus.Relationships of the new species to thespecies of this clade must be further in-vestigated. Liolaemus arambarensis, how-ever, does not possess any of the autapo-morphies described by Etheridge for theseknown species. Liolaemus arambarensisalso does not possess the synapomorphiesdefining Branch 25, which includes L.multimaculatus and L. riojanus. Amongthe synapomorphies described for Branch24 (Fig. 5), L. arambarensis possesses two:Character 13: State 1, almost all posteriorgulars without an apical notch and scalesensory organ; and Character 26: State 1,most blue scales in clumps on sides ofbody. It lacks, however, synapomorphies

Character 8: State 1 (inferior margin of an-terior supralabial scales distinctively con-cave, all together forming a scalloped low-er border that overlaps anterior infralabialscales) and Character 25: State 1 (irides-cent blue scales present in both fixedmales and females) that also defines thisbranch. All the information given abovesuggests that L. arambarensis is a sistergroup for Branch 24 (L. rabinoi, L. mul-timaculatus, and L. riojanus), but furtherinvestigation is needed to better supportsuch a hypothesis.

After our analysis and with consider-ation of Peters and Donoso-Barros (1970),Muller (1979), and de Lema (1994), whocite L. wiegmannii for southern RioGrande do Sul (specifically for Tapes), weconclude that all these populations corre-spond to L. arambarensis and that there isno record of L. wiegmannii in Brazil.

RESUMO

Uma nova especie do genero Liolaemusdo Rio Grande do Sul, Brazil e descritaneste trabalho. A morfologia da nova es-pecie, L. arambarensis sp. nov, e compar-ada as especies de Liolaemus do grupo‘‘wiegmannii.’’ A nova especie se distinguedos demais membros desse grupo pelopadrao de coloracao dorsal com uma faixabranca medio dorsal e duas faixas dorso-laterais, duas fileiras justapostas de marcasmarrons lembrando triangulos, limitadaspor uma barra branca; escamas nasais or-ientadas dorsalmente; um linha completade escamas lorilabiais entre as escamas su-bocular e supralabial orientada dorsal-mente; um numero moderado de escamasao redor do corpo escamas temporais plan-as; escamas da cabeca planas; e, frontaldividida transversalmente. Liolaemus ar-ambarensis sp. nov. e omnıvoro, ovıparo ecom marcante dimorfismo sexual em cor etamanho.

Acknowledgments.—We thank B. Koelln, E. Her-mel, F. Maraschin, G. Severino, G. Melamed, J. D.Denardin, Jr., M. Piccoli, M. Freitas, R. Bernhard,and R. Caruccio for assistance in collecting; P. Brackfor vegetation descriptions; T. O. de Freitas for cy-togenetic analyses; L. Avila and J. Langone for pro-viding material on loan; R. Etheridge for helpful sug-gestions; and L. Malabarba for valuable comments ofthe manuscript. We also thank to Instituto de Bio-


ciencias of Universidade Federal do Rio Grande doSul for travel support.

LITERATURE CITED

BELL, T. 1843. The Zoology of the Voyage of H.M.S.Beagle, Under Command of Captain Fitzroy, R.N.,During the Years 1832 to 1836. Part V. Reptiles.Smith, Elder and Company, London, U.K.

BERTOLOTTO, C., M. RODRIGUES, G. SKUK, AND Y.YONENAGA-YASSUDA. 1996. Comparative cytoge-netic analysis with differential staining in three spe-cies of Liolaemus (Squamata, Tropiduridae). Her-editas 125:257–264.

BOULENGER, G. A. 1885. Catalogue of the Lizards inthe British Museum (Natural History), SecondEdition, Vol. II. London, U.K.

CABRERA, M., AND N. T. BEE DE SPERONI. 1986.Composicion y distribucion de la lacertofauna dela provincia de Cordoba, Argentina. II. Amphis-baenidae, Anguidae e Iguanidae. Historia Natural(Corrientes, Argentina) 6:1–12.

CEI, J. M. 1974. Two new species of Ctenoblepharis(Reptilia, Iguanidae) from the arid environments ofthe central Argentina (Mendoza Province). Journalof Herpetology 8:71–75.

. 1979a. A reassessment of the genus Cten-oblepharis (Reptilia, Sauria, Iguanidae) with a de-scription of new subspecies of Liolaemus multi-maculatus from western Argentina. Journal of Her-petology 13:297–302.

. 1979b. Nota preliminar sobre la distribuciongeografica de Liolaemus wiegmanni (Dumeril etBibron) (Sauria, Iguanidae). Publicaciones Oca-sionales del Instituto de Biologıa Animal, Univer-sidad Nacional de Cuyo, Serie Cientıfica 14:1–4.

. 1986. Reptiles del centro, centro-oeste y surde la Argentina. Museo Regionale di Scienze Na-turali, Torino, Monografie 4:1–527.

DE LEMA, T. 1994. Lista comentada dos repteis ocor-rentes no Rio Grande do Sul, Brasil. Comunicacaodo Museu de Ciencias e Tecnologia da PontifıciaUniversidade Catolica do Rio Grande do Sul, SerieZoologia 7:41–150.

DONOSO-BARROS, R. 1973. Un nuevo saurio de Bo-lıvia (Lacertilia, Iguanidae). Neotropica, BuenosAires 19(60):132–134.

DUMERIL, A. M. C., AND G. BIBRON. 1837. Erpeto-logie General ou Histoire Naturelle Complete desReptiles, Vol. 4. Roret, Paris, France.

ETHERIDGE, R. 1993. Lizards of the Liolaemus dar-winii complex (Squamata: Iguania: Tropiduridae)in northern Argentina. Bollettino del Museo Re-gionale di Scienze Naturali, Torino 11:137–199.

. 1995. Redescription of Ctenoblepharys ad-spersa Tschdi 1845, and the taxonomy of Liolae-minae (Reptilia: Squamata: Tropiduridae). Ameri-can Museum of Natural History, Novitates 3142:1–34.

. 2000. A review of lizards of the Liolaemuswiegmannii group (Squamata, Iguania, Tropiduri-dae), and a history of morphological change in thesand-dwelling species. Herpetological Monographs14:293–352.

ETHERIDGE, R., AND K. DE QUEIROZ. 1988. A phy-

logeny of Iguanidae. Pp. 283–367. In R. Estes andG. Prigill (Eds.), Phylogenetic Relationships of theLizard Families—Essays Commemorating CharlesL. Camp. Stanford University Press, Stanford, Cal-ifornia, U.S.A.

GALLARDO, J. M. 1966. Liolaemus lentus nov. sp. (Ig-uanidae) de la Pampa y algunas observaciones so-bre los saurios de dicha Provincia Argentina y deloeste de Buenos Aires. Neotropica, Buenos Aires12(37):15–29.

GORMAN, G. C. 1973. The chromosomes of the Rep-tilia, a cytotaxonomic interpretation. Pp. 349–424.In A. B. Chiarelli and E. Campanna (Eds.), Cyto-taxonomy and Vertebrate Evolution. AcademicPress, London, U.K.

GUDYNAS, E. 1981a. Comentarios sobre biotopos,habitats, herpetofauna y la biogeografia del Uru-guay y areas vecinas. Jornadas de Ciencias Natur-ales, Centro Educativo Don Orione, Montevideo 2:7–8.

. 1981b. Consideraciones sobre la herpetofau-na del Uruguay y areas vecinas con una aproxima-cion biogeografica. Jornadas de Ciencias Naturales,Centro Educativo Don Orione, Montevideo 2:5–6.

. 1981c. New departmental records of Liolae-mus wiegmannii from Uruguay. Herpetolgical Re-view 12:84.

HALLOY, M., R. ETHERIDGE, AND G. M. BUR-GHARDT. 1998. To bury in the sand: phylogeneticrelationships among lizard species of the boulengerigroup, Liolaemus (Reptilia: Squamata: Tropiduri-dae), based on behavioral characters. Herpetologi-cal Monographs 12:1–37.

HOWELL, W. M., AND D. A. BLACK. 1980. Controlledsilver-staining of nucleolus organizer regions with aprotective colloidal developer: a 1 step-method.Experientia 36:1014–1015.

LAURENT, R. F. 1982. Description de trois especiesnouvellae du genere Liolaemus (Sauria, Iguanidae).Spixiana, Munchen 5:139–147.

. 1986. Descripciones de nuevos Iguanidae delgenero Liolaemus. Acta Zoologica Liolloana 38:87–105.

LAURENT, R. F., AND E. TERAN. 1981. Lista de losanfibios y reptiles de la provincia de Tucuman.Ministerio de Cultura y Educacion, Fundacion Mi-guel Lillo, Tucuman, Miscelanea 71:1–15.

MERTENS, R. 1938. Bemerkungen uber die brasilian-ischen Arten der Gattung Liolaemus. ZoologischerAnzeiger, Leipzig 123(7/9):220–222.

MORENO, J. A. 1961. Clima do Rio Grande do Sul.Secretaria da Agricultura, Porto Alegre, Brazil.

MULLER, P. 1979. The evolution of the Liolaemuswiegmanni-complex and the dispersal centers inBrazil. Amazoniana VI(4):537–555.

NAVARRO, J. 1992. Cariotipos de trece species de la-gartijas del noroeste argentino de los grupos Lio-laemus, Eulaemus y Ortholaemus. Acta ZoologicaLilloana 41:225–230.

PETERS, J. A., AND R. DONOSO-BARROS. 1970. Cat-alogue of the Neotropical Squamata. Part II—Liz-ards and Amphisbaenians. Smithsonian InstitutionPress, Washington, D.C., U.S.A.

PINILLA, M. P. R. 1991. Reproductive and fat body


cycle of the lizard Liolaemus wiegmanni. Amphib-ia-Reptilia 12:195–202.

ROCHA, C. F. D. 1985. Ecologia de Liolaemus lutzae(Sauria-Iguanidae) na Restinga da Barra de Marica-RJ. B.S. Thesis, Universidade do Estado do Rio deJaneiro, Rio de Janeiro, Brazil.

. 1989. Estrategia e ciclo reprodutivo de Lio-laemus lutzae (Sauria: Iguanidae) na restinga deBarra do Marica, RJ. M.S. Thesis, Departamentode Zoologia, Instituto de Biociencias, UniversidadeEstadual de Campinas, Sao Paulo, Brazil.

. 1991. Composicao do habitat e uso do es-paco por Liolaemus lutzae (Sauria: Tropiduridae)em uma area de restinga. Revista Brasileira doBiologia 51:839–46.

SMITH, H. M. 1946. Handbook of Lizards. Comstock,Ithaca, New York, New York, U.S.A.

SUMNER, A. T. 1972. A simple technique for dem-onstrating centromeric heterochromatin. Experi-mental Cell Research 75:303–306.

TEDESCO, M. E., A. HERNANO, AND B. B. ALVAREZ.1992. Hallazgo de Liolaemus wiegmanni (Dumerilet Bibron, 1837) (Iguania, Liolaemidae) en la prov-incia de Corrientes (Republica Argentina). Facena,Corrientes, Argentina 9:117–121.

VERRASTRO, L. 1991. Aspectos ecologicos e biologi-cos de uma populacao de Liolaemus occipitalisBoul. 1885, nas dunas costeiras da praia Jardim At-lantico, Tramandaı, RS. (Reptilia-Iguanidae). M.S.Thesis, Universidade Federal do Rio Grande doSul, Porto Alegre, Brazil.

VERRASTRO, L., AND C. S. BUJES. 1998. Ritmo deatividade de Liolaemus occipitalis Boulenger, 1885(Sauria, Tropiduridae) na praia de Quintao, RS—Brasil. Revista Brasileira de Zoologia 15:907–914.

VERRASTRO, L., AND L. KRAUSE. 1994. Analysis ofgrowth in a population of Liolaemus occipitalisBoul. 1885, from the coastal sand-dunes of Tra-mandaı—RS, Brazil (Reptilia: Tropiduridae). Stud-ies on Neotropical Fauna and Environment 29:99–111.

WAECHTER, J. L. 1985. Aspectos ecologicos da ve-

getacao de restingas no Rio Grande do Sul, Brasil.Comunicacoes do Museu de Ciencias da PUCRS,Serie Botanica, Porto Alegre 33:49–68.

Accepted: 22 April 2002Associate Editor:Joseph Mendelson III

APPENDIX ISpecimens Examined

Liolaemus wiegmanni, Rio Cuarto, Argentina (ZV-UNRC) Males: 3325, 3327, 3336, 3338, 3346, 3366,3369, 3378; Females: 3324, 3328, 3330, 3348, 3367,3375, 3377, 3380. Colonia, Uruguay (UFRGS) Males:3182, 3300–01, 3322–26; Female: 3183. Montevideo,Uruguay (MNHN) Males: 154, 196; Females: 137,266, 3306, 3310, 3316, 5838.

Liolaemus arambarensis, Arambare, Rio Grandedo Sul, Brasil (UFRGS) Males: 2369, 2682, 2694–95, 2728, 2799, 2748, 2750, 2769–72, 2774, 2776,2778, 2781, 2785, 2788–89, 2799, 2817, 2819–20,2822–25, 2902–03, 2905–07, 2946–48, 2950–51,3044, 3112–13, 3117–19, 2369, 2695, 2728–29,2748–49, 3186, 3221, 3267; Females: 2750–51,2773, 2775, 2779, 2782–84, 2786–87, 2790, 2794,2796, 2800, 2802, 2818, 2821, 2826–32, 2908–13,2945, 2947, 2951, 3114, 3120, 3268. Arambare, RioGrande do Sul, Brasil (MCTPUCRS) 4475, 4525,4652–54. Barra do Ribeiro, Rio Grande do Sul,Brasil (UFRGS) Male: 3184. Females: 3185, 3222.Tapes, Fz. Formosa, Rio Grande do Sul, Brasil(MCTPUCRS) 5121, 5362–65.

Liolaemus occipitalis, Cidreira, Rio Grande do Sul,Brasil (UFRGS) Males: 2497, 2504–05, 2513, 2519,2522, 2524, 2540, 2545–46, 2556, 2559–61, 2564–65,2567, 2628, 2630, 2632–33, 2635, 2637–38, 2753,2755, 2760, 2765–67; Females: 2498–2501, 2503,2510–12, 2514, 2521, 2541–44, 2548, 2555, 2557–58,2562–63, 2566, 2619, 2623–24, 2629, 2754, 2757,2764, 2768.

Liolaemus lutzae, Restinga do Marica, Rio de Ja-neiro, Brasil (UFRGS) Males: 227–28, 825; Females:203–04. Praia de Grumari, Rio de Janeiro, Brasil(UFRGS) Male: 2482; Females: 2483, 2487–88.

2003 new species_liolaemus

Education