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Article A new species of the Liolaemus darwinii group (Iguania: Liolaemidae) from Salta Province, Argentina

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  • National University of Tucuman- Unidad Ejecutora Lillo CONICET

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We describe a new species of the iguanian genus Liolaemus of Northwestern Argentina in Salta Province. This new lizard is a member of the L. boulengeri group, and within this group it is a member of the L. darwinii subgroup. With the addition of the species described here, the L. darwinii group now contains 19 species. Like most of the members of this group, the new taxon exhibits sexual dichromatism, showing a unique color pattern in males and females. The color pattern is characterized by the presence of a pre-scapular spot and lateral black stripes, unique within the L. darwinii group. Liolaemus diaguita sp nov inhabits a region where bushes are the predominant vegetation, in Quebrada de Las Conchas, in Guachipas Department, Salta Province, which has an elevation between 1200–2500 m
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26 Accepted by S. Carranza: 30 May 2011; published: 18 Jul. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2968: 2638 (2011)
www.mapress.com/zootaxa/Article
A new species of the Liolaemus darwinii group (Iguania: Liolaemidae)
from Salta Province, Argentina
CRISTIAN SIMÓN ABDALA1, ANDRÉS SEBASTIÁN QUINTEROS2,3, FEDERICO ARIAS2,
SABRINA PORTELLI2 & ANTONIO PALAVECINO2
1Instituto de Herpetología, Fundación Miguel Lillo, Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) y Cáte-
dra de la Reserva Experimental Horco Molle (REHM), Facultad de Ciencias Naturales e Instituto Miguel Lillo (IML), Universidad
Nacional de Tucumán. Miguel Lillo 251. 4000- Tucumán. Argentina. E-mail: samiryjazmin@gmail.com
2Cátedra de Anatomía Comparada and IBIGEO-CONICET, Facultad de Ciencias Naturales, Universidad Nacional de Salta, Avenida
Bolivia 5150, Salta 4400, Argentina. E-mail: squint@unsa.edu.ar; fjarias@unsa.edu.ar; sabrina.portelli@gmail.com
3Corresponding author. E-mail: squint@unsa.edu.ar
Abstract
We describe a new species of the iguanian genus Liolaemus of Northwestern Argentina in Salta Province. This new lizard
is a member of the L. boulengeri group, and within this group it is a member of the L. darwinii subgroup. With the addition
of the species described here, the L. darwinii group now contains 19 species. Like most of the members of this group, the
new taxon exhibits sexual dichromatism, showing a unique color pattern in males and females. The color pattern is char-
acterized by the presence of a pre-scapular spot and lateral black stripes, unique within the L. darwinii group. Liolaemus
diaguita sp nov inhabits a region where bushes are the predominant vegetation, in Quebrada de Las Conchas, in Guachipas
Department, Salta Province, which has an elevation between 1200–2500 m.
Key words: Argentina, Salta, Liolaemus, L. darwinii group, new species
Resumen
Describimos una nueva especie del género Liolaemus para el noroeste de Argentina en la provincia de Salta. Este nuevo
lagarto pertenece al grupo de L. boulengeri, y dentro de éste, al grupo de L. darwinii, con lo cual este grupo quedaría con-
formado hasta el momento por 19 especies. Al igual que la mayoría de los integrantes de este grupo, este nuevo taxón
presenta dicromatismo sexual evidente y el patrón de coloración tanto de machos como hembras es particular, caracter-
izado por la presencia de una mancha pre-escapular y bandas laterales negras, únicas entre las especies del grupo de L.
darwinii. Liolaemus diaguita sp nov habita en una región predominada por arbustos en la Quebrada de las Conchas, en el
departamento de Guachipas de la Provincia de Salta, en una altura entre los 1200 y 2500 m.s.n.m.
Introduction
As a consequence of numerous geological events, e. g. the formation of the Cordillera de Los Andes and the Pre-
Andean Mountains, the genus Liolaemus has undergone much fragmentation and isolation of its populations. These
events resulted in many depressions and elevations which facilitated speciation events in the genus. With the addi-
tion of this new species Liolaemus now contains by 223 species (Lobo et al., 2010), being the second largest genus
within Iguania, after Anolis.
Many studies of the taxonomy and phylogenetic relationships within Liolaemus have resulted in the proposals
of species groups and subgroups. The two species groups have been recognized as subgenera, Liolaemus (sensu
stricto) and Eulaemus (Laurent, 1983; 1985; 1995; Schulte et al., 2000; Espinoza et al., 2004). Within Eulaemus
several subgroups have been recognized: the L. lineomaculatus group, the L. boulengeri group, and the L. monta-
nus group (See Lobo et al., 2010 for a summary).
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A NEW SPECIES OF THE LIOLAEMUS DARWINII GROUP
Among the subgroups of Liolaemus is the L. darwinii group (Abdala, 2007), originally proposed by Etheridge
(1993) as the darwinii complex. This group is a member of the more inclusive L. boulengeri group (Abdala, 2007)
characterized by a patch of enlarged scales on the posterior surface of the thigh. The L. boulengeri group represents
about 30% of the known species of Liolaemus. Within the L. darwinii group there are 19 species, including the new
species here described. The darwinii complex proposed by Etheridge (1993) was supported by the presence of
straight-edged crowns on the posterior teeth and marked sexual dichromatism. Posteriorly, Abdala (2007) recover a
clade named L. darwinii group which resemble de darwinii complex of Etheridge (including 7 more new species).
This clade is supported by a black line passing vertically through the eye (except in L. chacoensis), dark pre-scapu-
lar spot (except in species members of the L. ornatus group), the number of gular scales, the development of the
longitudinal fold, the number of scales projecting in the anterior margin of the auricular meatus, and 2 morphomet-
rics characters.
The new species belongs to the Liolaemus darwinii group, because it has a markedly sexual dichromatism, the
males exhibit a more conspicuous coloration than females, and it exhibits character states which allow us to distin-
guish it from all other members of the group. In the L. darwinii group males of different species are easily to distin-
guish, while females, mainly, show a similar color pattern. However, females of the new species are easy to
distinguish from females of other species because they exhibit a black longitudinal stripe in the upper region of lat-
eral field. Males of the new species show an evident but small pre-scapular spot, small and square-shaped paraver-
tebral spots, and the presence of a dark brown or black line in the upper region of lateral field.
In this paper we describe a new species from a sandy region, where bushes are the predominant vegetation,
located in Guachipas Department, in Central-South of Salta Province, in Quebrada de Las Conchas. Also, we pro-
vide an identification key for adult males of the species in the Liolaemus darwinii group (Sensu Abdala, 2007).
Material and methods
We studied the morphological characters traditionally used in Liolaemus taxonomy including those of Laurent
(1985), Cei (1986, 1993), Etheridge (1993, 1995, 2000), Lobo (2001) and Abdala (2002, 2003, 2007). We follow
Smith’s (1946) terminology for squamation descriptions and Frost’s (1992) for neck-fold terminology. Descriptions
of body-color patterns follow Lobo and Espinoza (1999). Descriptions of color in life were based on observations
made in the field or on photographs of specimens taken soon after capture. Determination of sexual maturity was
assessed by a combination of body size and the presence of secondary sexual characteristics (see Valdecantos et al.,
2007; Valdecantos and Lobo, 2008). Measurements and scale counts were recorded from specimens that were fixed
in 10% formalin and preserved in 70% ethanol. Body and scale measurements were taken with digital calipers to
the nearest 0.02 mm. All measurements are in mm. When necessary, a binocular dissecting microscope (10–40´)
was used to count and characterize scales. Where bilateral, scale counts and measurement data were taken from the
right side of the lizards.
For purposes of diagnosing the new species described herein, we examined type series or topotypes of other
members of the Liolaemus darwinii group whenever possible. We examined 445 specimens which correspond to
17 species. Specimens studied are listed in Appendix I. Data from L. cinereus and L. montanezi were taken from
bibliography.
Results
Liolaemus diaguita sp. nov.
(Fig. 1–5)
1993. Liolaemus quilmes—Etheridge, Boll. Mus. Reg. Sci. Nat. Torino, 11: 137–199.
2008. L. cf. quilmes 1—Morando et al., Zoologica Scripta, 37: 603–618.
Holotype. MCN 3247. 61 Km N of Cafayate, on National Road 34, 400 m S of Río Las Conchas, Quebrada de las
Conchas; Guachipas Department, Salta Province. 25°49'07.1''S; 65°41'26.1''W. 1328 m. Arias, F., Palavecino, A.,
Portelli, S. & Quinteros, S. cols. 27/01/2010.
ABDALA ET AL.
28 · Zootaxa 2968 © 2011 Magnolia Press
Paratypes. FML 23821-825. MCN 3145-50; 3243; 3248-50. Same data as Holotype.
Diagnosis. Liolaemus diaguita sp nov., belongs to the L. boulengeri group because it has a patch of enlarged
scales on posterior region of thigh (Etheridge, 1995; Abdala et al. 2006). Within this group, it belongs to the L. lau-
renti group (Abdala, 2007) because its posterior teeth have straight edged crowns and sexual dichromatism is evi-
dent. Within the L. laurenti group, Liolaemus diaguita sp nov. is a member of the L. darwinii group (Abdala, 2007)
or the darwinii complex (Etheridge, 1993) because it has a black line which crosses the eye vertically (except L.
chacoensis) and pre-scapular spots in both males and females (Abdala, 2007). Tooth shape and the line crossing the
eye distinguish the new taxon from all members of the L. melanops group [Abdala, 2007 (L. boulengeri, L can-
queli, L. cuyanus, L. donosobarrosi, L. fitzingerii, L. inacayali, L. josei, L. mapuche, L. martorii, L. melanops, L.
morenoi, L. loboi, L. sagei, L. rothi, L. senguer, L. tehuelche, L. telsen, and L. xanthoviridis)]; also the presence of
sexual dichromatism distinguish L. diaguita from L canqueli, L. cuyanus, L. chehuachekenk, L. goestchi, L. fitzing-
erii, L. martorii, L. morenoi, L. sagei, and L. xanthoviridis. It differs from L. canqueli and L. melanops, because L.
diaguita lacks cephalic melanism, and from L. canqueli, L. cuyanus, L. mapuche, L. morenoi, L. fitzingerii and L.
xanthoviridis because those species show gular and antehumeral melanism, absent in L. diaguita. The new species
has four scales contacting the mental scale, differing from L. cuyanus and all the members of the L. wiegmannii
group (L. arambarensis, L. azarai, L. cranwelli, L. cuyumhue, L. lutzae, L. multimaculatus, L. occipitalis, L. rabi-
noi, L. riojanus, L. salinicola, L. scapularis and L. wiegmannii), which have six scales contacting the mental and
from L. josei and L. mapuche where the mental scale can contact 4 to 6 scales. Also, the members of the L. wieg-
mannii group have two rows of lorilabial scales, while L. diaguita has one row. The new taxon differs from species
of the L. anomalus group (L. anomalus, L. ditadai, L. lentus, and L. pseudoanomalus - sensu Abdala, 2007)
because its tail is longer than snout-vent length (same length in L. anomalus group), head larger than wide (wider
than long in L. anomalus group), enlarged superior cilliaries absent (present in L. anomalus group), larger number
of precloacal pores in males and females, and lack of pterigoid teeth. Within the L. darwinii group (Abdala, 2007),
L. diaguita differs from the members of the L. ornatus group (L. albiceps, L. calchaqui, L. crepuscularis, L. irregu-
laris, L. lavillai, and L. ornatus) because it has an oviparous reproductive mode and it lacks precloacal pores in
females. Also it differs from L. albiceps and L. irregularis because it is smaller (Max SVL 49.7mm vs 82.5 and
86.1 mm respectively). Liolaemus diaguita differs from L. abaucan and L. uspallatensis in having imbricate and
keeled dorsal scales, while in those two species the dorsal scales are smooth or weakly keeled. Liolaemus chacoen-
sis exhibits keeled temporal scales and mucronate dorsal scales, absent in L. diaguita. The presence of a small post-
scapular spot distinguishes L. diaguita from L. abaucan, L. darwinii and L. koslowskyi which have a large and
evident postscapular spots. It differs from L. cinereus in having a black stripe in the lateral field and in lacking mel-
anism on the throat. Liolaemus darwinii, L. grosseorum, L. laurenti, L. montanezi and L. olongasta have a black
antehumeral arch, absent in L. diaguita. Liolaemus espinozai shows spots located posterior to the postscapular spot
and a lateral line in males and females, while both are absent in L. diaguita specimens. Liolaemus diaguita differs
from L. quilmes because it has a small prescapular spot and a black stripe in the lateral field, both absent in L.
quilmes.
Males and females of Liolaemus diaguita have a unique dorsal color pattern formed by paravertebral and lat-
eral spots which can join together forming paravertebral and lateral stripes respectively. Additionally, the speci-
mens lack spots on throat and belly. These character states distinguish the new taxon from all members of the L.
laurenti group.
Description of the holotype. Adult male. Snout vent length (SVL) 49.68. Head 1.3 times longer (12.48 mm)
than wide (9.21mm). Head height 6.71 mm. Interorbital distance 6.75 mm. Eye-auditory meatus distance 4.71 mm.
Auditory meatus height 2.1 mm; wide 1.61 mm. Supralabials–eye 2.3 mm; distance between nares 1.53 mm. Sub-
ocular legnth 0.72 mm. Trunk length 22.31 mm; tail length 70.32 mm. Tibia length 4.84 mm. Foot length 14.98
mm. Humerous length 3.62 mm.
Dorsal surface of head smooth, with five enlarged supraoculars. Eight scales between rostral and frontal scales.
Mental scale trapezoidal, larger than rostral, in contact with four scales. Four chinshields scales. There is no contact
between nasal and rostral scales, separated by one scale. Seven scales surround the nasal scale, which is separated
from canthal scale by four scales. Two postrostral scales, with seven scale organs. Interparietal scale larger than
paritetals, in contact with six scales. Hellmich index (dorsal scales in head, from occiput to mental scale) 17.
Preocular separated from lorilabial row by one scale. Nine supercilliaries. Sixteen upper cilliaries. Anterior
edge of the auditory meatus with three scales projecting. Auricular scale (located in antero-superior edge of the
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A NEW SPECIES OF THE LIOLAEMUS DARWINII GROUP
auditory meatus) present. Eleven smooth temporals. Subocular scale in contact with four lorilabials. Five supraoc-
ular scales. Nine supralabial and six infralabial scales. One row of nine lorilabials. Scales around midbody 57.
Sixty-three round, keeled and imbricated dorsal scales (from occiput to forelimbs). Sixty-eight ventral scales from
mental scale to cloaca, larger than dorsals, laminar and inbricate. Twenty-four imbricate and smooth gulars. Anteg-
ular fold present. Forty-five granular and smooth neck scales (counted from posterior margin of the auditory
meatus to sholuder, along the longitudinal fold). Neck folds (auricular, antehumeral and longitudinal) evident. Sev-
enteen infradigital lamellae on fourth finger and 28 on fourth toe. Femoral patch evident. Seven precloacal pores.
Dorsal caudal scales mucronate and smooth; ventral smooth.
FIGURE 1. Adult male of L. diaguita sp nov., dorsal view.
FIGURE 2. Adult male of L. diaguita sp nov., ventral view.
Color of the holotype in ethanol. Dorsal background color dark brown. Head dorsally brown with dark brown
spots. Lateral zone of head brown with black lines. Temporal region with black spots. A black line born in the pos-
terior margin of the eye, extending and reaching the lateral field. A vertical black line crosses the eye (like all other
ABDALA ET AL.
30 · Zootaxa 2968 © 2011 Magnolia Press
members of the L. darwinii group—except L. chacoensis). Dark brown vertebral zone, without vertebral line. In the
region close to the occiput there are light brown to white lines. Eight pairs of paravertebral white spots are evident.
Two dorsolateral white stripes, bordered in light brown. Lateral fields dark brown to black, with white spots speck-
led. Ventrolateral line white and evident, bordered by a brown line ventrally. Ventrolateral field brown with
rounded white spots. Fore and hind limbs brown (lighter than dorsum) speckled with dark brown and white spots.
Ventrally throat, chest, belly, limbs and tail are immaculate, with a creamy white color. Tail dorsally, brown with
black spots forming a ring pattern.
FIGURE 3. Adult female of L. diaguita sp nov., dorsal view
FIGURE 4. Adult female of L. diaguita sp nov., ventral view.
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A NEW SPECIES OF THE LIOLAEMUS DARWINII GROUP
FIGURE 5. Map showing the distribution of L. diaguita and the species of the L. darwinii group more close distributed. Black
square: L. diaguita. Black circle: L. quilmes. Black triangle: L. espinozai. Black star: L. koslowskyi. Black diamond: L. abau-
can. White square: L. albiceps. White circle: L. lavillai. White triangle: L. calchaqui. White star: L. crepuscularis. Arrow indi-
cates the type locality of L. diaguita.
Color in life of the holotype. Dorsal background. Head with same color of dorsum with dark brown to black
lines. Temporal region same color than dorsum, with a black line which begins on the posterior margin of the eye,
and reaches the lateral region of body. Pre-scapular spot present. Paravertebral field black with ten pairs (between
fore and hind limbs) of white irregular spots. Vertebral zone without vertebral line. Dorsolateral stripes light brown
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32 · Zootaxa 2968 © 2011 Magnolia Press
with white spots. Lateral field brownish yellow. In the lateral field is evident a dark brown line bordered with white
spots. Ventrolateral line white and distinct. Ventrolateral field light gray with white spots speckled. Fore and hind
limbs grayish brown with light gray and dark brown spots. Tail dorsally, same color of dorsum of body. The black
paravertebral field of the body extends to the proximal region of tail. In distal region of tail there are black and
white lines, forming a ringed pattern. Ventrally, throat, chest, belly, fore and hindlimbs, and tail, immaculate white
cream.
Va ri at io n . Based on fifteen paratypes. Head longer (10.2–12.6 mm; Mean = 11,5 mm) than wide (8.3–12.5
mm; Mean = 9.7 mm). Head height 6.0–7.4 mm (Mean = 6.5 mm). SVL 41.5–49.6 mm (Mean = 47.2 mm), averag-
ing 2 times longer than trunk. Tail length 64.3–81.7 mm (Mean = 70.7 mm). Width of tail base 4.6–6.5 mm (Mean
= 5.9 mm). Humerus length 3.1–6.2 mm (Mean = 4.4 mm). Tibia length 4.4–6.5 mm (Mean = 5.1). Foot length
11.3–17.9 mm (Mean = 14.0 mm). Fourth finger length 4.9–7.9 mm (Mean = 6.2 mm). Fourth toe length 8.9–11.6
mm (Mean = 9.9 mm). Dorsal surface of head smooth, with 14–20 scales (Hellmich index). Seven to eight supercil-
liaries. Ten to sixteen upper cilliaries. One row of lorilabials, same size as supralabials. Eight to ten lorilabials.
Seven to eleven supralabials. Frontal divided into two scales. Interparietal always larger than parietals, surrounded
by 4–8 scales. Nasal surrounded by 6–8 scales. Four to six infralabials. Mental in contact with four scales. Ten to
thirteen smooth and round temporals. Longitudinal, postauricular and antehumeral folds present. Horizontal fold Y-
shaped between the shoulder and the auditory meatus.
Scales around midbody 52–58 (Mean = 55.5). Gulars 20–29 (Mean = 22.8). Dorsal scales between occiput and
hind limbs 57–68 (Mean = 62.6). Ventrals 58–70 (Mean = 64.6). Males with 5–7 (Mean = 5.8) precloacal pores.
Females without precloacal pores. Dorsals flat, imbricate and keeled. Sixteen to twenty infradigital lamellae on
fourth finger and 24–30 on fourth toe.
Variation in color pattern. (Fig 1–4) Sexual dimorphism evident. Males exhibit the same color pattern as the
holotype. The only difference is in the intensity of the yellow color in the lateral fieldis more intense in
males.Females with the dorsal background color gray to light brown. Head of females are similar in males and
females. The vertebral region is bordered by black rounded spots. The size and number of those spots vary and they
are absent in some specimens. Pre-scapular spot present. The lateral field exhibits two stripes, a black on the top
and white cream below, without spots. Light brown fore and hind limbs with dark brown lines and spots. Ventrally
females show the same color of the holotype.
Distribution. The new species is known only from its type locality. (Fig. 5)
Natural history. Liolaemus diaguita was found in a sandy area where vegetation is predominated by shrubs
(Thrichocereus sp, and Prosopis sp). The area belongs to the phytoegegraphical region of Monte (Cabrera and
Willink, 1980). It can be found basking on rocks along the road.
Etymology. The epithet diaguita comes from native people denominated Diaguitas, who inhabit Northwestern
Argentina, especially in Valles Calchaquies, since the year 850. These communities developed a rich culture, and at
present there are more than 60,000 people in Northwestern Argentina who belong to this ethnic group.
TABLE 1. Character states of squamation of the Liolaemus darwinii group. SA: number of scales around midbody; D: number
of dorsal scales from occiput to hind limbs. PPM: number of precloacal pores in males. PPF: number of precloacal pores in
females. Max. SVL: Maxim Snout Vent Length. Data from L. cinereus and from L. montanezi were taken from Monguillot et
al. (2006) and Cabrera & Monguillot (2006), respectively.
SA D PPM PPF Max. SVL
L. diaguita 5258 (55.5) 5768(62.6) 57 (5.8) 0 49.6 mm
L. abaucan 5665 (60.1) 6271 (65.7) 67 (6.5) 0 58.3 mm
L. albiceps 6572 (68.9) 7483 (77.7) 911 (10.3) 611 (7.5) 82.5 mm
L. calchaqui 6172 (64.9) 5867 (63.9) 710 (8.33) 0 57.8 mm
L. chacoensis 44–54 (49.7) 49–57 (52.6) 4–6 (5.2) 0 46.5 mm
L. cinereus 58–61 (59.3) 69–71 (69.7) 8 0 63 mm
L. crepuscularis 56–64 (59.7) 59–72 (65.8) 6–9 (7.4) 5–7 (6.5) 64.1 mm
L. darwinii 53–63 (56.6) 53–67 (61.3) 6–8 (6.7) 0–3 (0.9) 62.3 mm
L. espinozai 57–69 (63.2) 63–70(66.4) 5–8 (7.4) 0–5 (2.2) 62.8 mm
continued on the next page...
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A NEW SPECIES OF THE LIOLAEMUS DARWINII GROUP
Identification key for adult males of the Liolaemus darwinii group
1. Black vertical line to eye absent; dorsal scales markedly keeled and imbricated; inferior temporals keeled. . . . . . L. chacoensis
- Black vertical line to eye present; dorsal scales smooth or keeled and imbricate or juxtaposed; inferior temporals smooth . . .2
2. Dorsal scales smooth or slightly keeled and juxtaposed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
- Dorsal scales keeled and imbricate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
3. Scapular zone of dorsum with scapular, prescapular and postscapular spots; postscapular spot is the largest; throat immaculate
or with tiny black spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. abaucan
- Scapular zone of dorsum with prescapular spot small, postscapualar spot absent or evident, when evident smaller or equal to
prescapular spot; throat yellow, black or dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. uspallatensis
4. Antehumeral fold with spots forming a vertical arch located anterior to fore limbs (Antehumeral arch) . . . . . . . . . . . . . . . . . .5
- Antehumeral fold without spots forming a vertical arch located anterior to fore limbs (Antehumeral arch). . . . . . . . . . . . . . . .9
5. Antehumerall arch split into two on fore limb, formed by two black lines with white spots between the lines . . . L. olongasta
- Antehumeral arch never split . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
6. Scapular zone with black prescapular and postscapular spots; postscapular always larger; black spot on anterior surface of the
hind limbs present or absent; black spots in belly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. darwinii
- Scapular zone with or without prescapular or postscapular spots; without black spot on anterior surface of hind limbs; without
black spots on belly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
7. Dorsal background brown or reddish brown; without dorsolateral stripes; scapular zone with small prescapular spot; without
yellow spots on lateral region of neck. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
- Dorsal background gray; scapular zone with prescapular and postscapular spots; with yellow spot on lateral region of neck. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..L. grosseorum
8. Dorsal background brown or reddish brown; with a few or without light blue scales on sides of tail and body; throat pigmented,
usually black or dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. laurenti
- Dorsal background dark brown; with many light blues scales on sides of tail and body; throat with a few spots. . .L. montanezi
9. With small prescapular spot; postscapular spot absent; throat black or dark brown. . . . . . . . . . . . . . . . . . . . . . . . . . . L. cinereus
- With a conspicuous, large prescapular spot; postscapular spot absent; throat never black neither dark brown. . . . . . . . . . . . . 10
10. With many light blue scales on sides of tail and body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11
- With few or without blue scales on sides of tail and body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
11. Postscapular spot conspicuous, always larger than prescapular spot; without spots posterior to postscapular spot L. koslowskyi
- Postscapular spot absent, or when present never larger than prescapular spot. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12
12. Postscapular spot absent or tiny; without spot posterior to postcapular spot; lateral field of body gray or light brown; dorsolat-
eral region white or light gray. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. quilmes
- Postscapular spot absent, or when present same size as prescapular spot; with spots posterior to postscapular spot, lateral
region of body orange or red; dorsolateral region yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. crepuscularis
13. With longitudinal line or lateral spots conspicuous, same size as scapular spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- Lateral spots absent or inconspicuous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16
14. With a black line or stripe in lateral field . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. diaguita
- Without a black line or stripe in lateral field. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15
15. Head and neck darker than body; irregular shaped paravertebral spots; lateral field of body yellow or reddish yellow . . . . . . . .
Table 1 continued
SA D PPM PPF Max. SVL
L. grosseorum 52–59 (56) 54–59 (56.3) 6–8 (7.5) 0 55.1 mm
L. irregularis 62–71 (66.7) 67–83 (75.4) 7–11 (8.6) 4–9 (7.3) 86.1 mm
L. koslowskyi 51–61 (56.2) 50–65 (59.6) 5–7 (5.57) 0–1 (0.2) 68.5 mm
L. laurenti 52–57 (54.8) 54–60 (56.9) 6–8 (6.8) 0–3 (0.4) 58.9 mm
L. lavillai 63–76 (69.2) 67–74 (70.1) 6–9 (7.1) 3–9 (5.6) 64.5 mm
L. montanezi 61–62 (61.5) 69 6 0 63 mm
L. olongasta 52–58 (54.9) 56–71 (65.2) 6–8 (7.1) 0–2 (0.5) 65.6 mm
L. ornatus 55–62 (58.3) 62–71 (66.2) 6–9 (7.5) 0–6 (4.2) 64.6 mm
L. quilmes 56–66 (61.5) 57–69 (64.7) 5–7 (6.1) 0–1 (0.3) 61.1 mm
L. uspallatensis 67–78 (71.4) 76–84 (80.3) 5–7 (6) 0 63.8 mm
ABDALA ET AL.
34 · Zootaxa 2968 © 2011 Magnolia Press
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. ornatus
- Body darker than head and neck; square shaped paravertebral spots; dorsal background orange or yellow; lateral field of body
orange or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. espinozai
16. Dorsal color pattern with irregular spots irregularly spread on dorsum; throat melanic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Dorsal color pattern with spots regularly distributed on dorsum; throat never melanic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18
17. Head dorsally light gray, never melanic; dorsal background of body dark or light gray, never light blue . . . . . . . . . ..L. albiceps
- Head dorsally dark brown or black; dorsal background of body light blue or bluish gray. . . . . . . . . . . . . . . . . . . . L. irregularis
18. Wide dorsolateral stripes; square shaped or absent paravertebral spots; wide vertebral zone . . . . . . . . . . . . . . . . . . . . L. lavillai
- Wide or narrow dorsolateral stripes; square or line shaped (wider than long) paravertebral spots always present, when line
shaped the spots cross the dorsolateral stripes; narrow vertebral zone. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. calchaqui
Discussion
The Liolaemus darwinii complex was proposed by Etheridge (1993). In that contribution, four new species were
described (L. abaucan, L. koslowskyi, L. olongasta and L. quilmes), L. darwinii and L. ornatus were re-described,
and the L. darwinii complex characterized. Etheridge’s contribution is fundamental to the group taxonomy, and the
genus, because his proposal included the separation of species based on color pattern characters, at aspect not con-
sidered previously.
Until 1992, the Liolaemus darwinii group included only four species (L. darwinii, L. irregularis, L. ornatus and
L. uspallatensis), but with the contributions of Etheridge (1992, 1993) the number of species which belong to the
group was increased. Most of those species were previously confused with L. darwinii. Moreover, more new spe-
cies were subsequestly described: L. albiceps (Lobo and Laurent, 1995), L. calchaqui (Lobo and Kretzschmar,
1996), L. grosseorum (Etheridge, 2001), L. espinozai (Abdala, 2005), L. crepuscularis (Abdala and Diaz Gomez,
2005), L. lavillai (Abdala and Lobo, 2006), L. cinereus (Monguillot et al. 2006), and L. montanezi (Cabrera and
Monguillot, 2006) all of them were mainly based on color pattern characters. At present the L. darwinii group
includes 19 species, considering L. diaguita and L. chacoensis which were proposed inside the Liolaemus sensu
stricto subgenus by Etheridge (1995), but according to other phylogenetic analyses (Schulte et al, 2000; Morando
et al., 2004; and Abdala, 2007) it is a member of the L. darwinii group.
The Liolaemus darwinii group was studied in detail by Abdala (2007), who performed inclusive phylogenetic
and taxonomic analyses, studying the L. boulengeri group, or “patch group”, and in that context he defined the L.
darwinii group (Fig. 6). According to Abdala’s (2007) proposal, the L. darwinii group belongs to the L. boulengeri
group, and within it, it is included in the L. laurenti group. In the L. laurenti group, the L. darwinii group is sister of
the L. wiegmannii group, which is composed of sand lizards, with unique morphological and ethological characters
(Etheridge, 2000). The L. darwinii group is characterized as males and females of the species members have a dark
(black) vertical line crossing the eye (absent only in L. chacoensis) and due to the presence of the prescapular spot
-absent in species of the L. ornatus clade-(Abdala, 2007). Etheridge (1993) proposed as synapomorphies of the
group the presence of straight edge crowns in posterior teeth and a marked sexual dichromatism, where males are
more conspicuous than females. In Abdala’s (2007) phylogeny, those character-states are present in the members of
the L. darwinii group but they are not exclusive, they are (together with species of the L. wiegmannii group) in the
more inclusive L. laurenti group. Also, the species members of the L. anomalus group share this character state.
The Liolaemus darwinii group includes species distributed in Central-West Argentina, and it is formed by two
monophyletic groups. One of them is the L. ornatus clade, which includes species inhabiting high elevations,
where they are viviparous, mainly herbivorous and the females exhibit a large number of precloacal pores. The
other group is the L. grosseorum clade (Fig. 6), inhabiting Central-West and South Argentina. It shares morphomet-
ric character states from head, neck, auditory meatus and trunk. Results from Abdala (2007) are congruent with
previous contributions, like molecular based phylogenies of Schulte et al. (2000) and Morando et al. (2004). The
last one, particularly, although it didn’t include species of the L. ornatus group, it recovered the L. grosseorum
group and the stem species to those clades of Abdala (2007).Morando et al., (2008) perform a phylogeographic
analysis of populations of L. kowslowskyi. There, the author identified six populations close to L. quilmes, the sam-
ple named L. cf quilmes 1, correspond to L. diaguita. Also, Morando et al. (2008) founded two clades of L.
koslowskyi (named L. koslowskyi North and L. koslowskyi East) which differs from the L. koslowskyi sensu stricto.
Zootaxa 2968 © 2011 Magnolia Press · 35
A NEW SPECIES OF THE LIOLAEMUS DARWINII GROUP
FIGURE 6. Cladogram of the L. darwinii group, proposed for Abdala (2007), based on total evidence analysis of the L. bou-
lengeri group.
The new species here described—like Liolaemus cinereus and L. montanezi—were not included in any phylo-
genetic analysis, so their phylogenetic position still remains unknown. However, based on morphological charac-
ters, geographic distribution, the previous confusion with L. quilmes (Etheridge, 1993), and the assignment to a
population close to L. quilmes (Morando et al., 2008) we can conclude that it is possible that the new species is a
stem member of the L. darwinii group.
Within the Liolaemus darwinii group, L. diaguita is the only species where both males and females show a
black stripe in the lateral field of the body. And like all other members of the L. darwinii group, it exhibits a marked
sexual dichromatism. Most of the species of the group (except for L. albiceps and L. irregularis) males exhibit the
diagnostic character states of the species; usually those character states referred to color pattern. Although females
exhibit the same color pattern as males—but less conspicuous—in many cases it is very difficult to distinguish
between females of different species of the L. darwinii group. This situation is more complicated in areas where
two species of the L. darwinii group inhabit in sympatry, like L. darwinii and L. grosseorum (Nihuil, Mendoza
Province, Argentina), L. olongasta and L. laurenti (Villa Unión, La Rioja Province, Argentina) and L. koslowskyi
and L. espinozai (Campo el Arenal, Catamarca Province, Argentina). In all cases the correct identification of males
is necessary, and for this reason we provide an identification key for adult males of the known species. Remaining,
at least, 6 candidate species closely related to L. quilmes (Morando et al., 2008).
Acknowledgments
We thank F. Lobo for his critical and very helpful review of a first draft. We thank E. Lavilla and S. Kretzschmar
(FML), F. Lobo (MCN), J. Williams (MLP), J. Faivovich and B. Bloto (MACN) and R. Etheridge (SDSU) for giv-
ing us access to collections under their charge. This study received financial support from CONICET (PIP 2841 to
F. Lobo) and Universidad Nacional de Salta (ASQ grant CIUNSA 1919). Permits for field research and collection
of specimens were extended by the Secretaría de Recursos Naturales y Medio Ambiente of Salta Province, Argen-
tina.
ABDALA ET AL.
36 · Zootaxa 2968 © 2011 Magnolia Press
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APPENDIX I
List of specimens studied. Acronyms correspond to Fundación Miguel Lillo (FML) and Museo de Ciencias Naturales – Univer-
sidad Nacional de Salta (MCN). All localities are from Argentina. Numbers in parentheses following each taxon name are
the number of individuals examined. Note that some specimen numbers from FML represent lots of more than one individ-
ual. In such cases, the number of individuals in the lot is indicated in brackets following the catalog number.
Liolaemus abaucan (24). CATAMARCA. Departamento Tinogasta. FML 2638 (holotype); FML 2639 [4] (paratypes): 12
km S of Palo Blanco, Ruta Provincial 34. FML 16557-58 [2]: 24 km al W de Fiambalá por ruta provincial 45. 27°41'18,6"
S 67°47'48,6" W. 2104 m.; FML 2585 [17]: Fiambalá.
Liolaemus albiceps (28). SALTA. Departamento Los Andes. FML 3370 (holotype); FML 1824 [11] (paratypes): 10 km S of
Estación Muñano, San Antonio de los Cobres. Departamento de Rosario de Lerma. FML 1865 [12]: Estación Cachiñal.
FML 3331 [4] (paratypes): Santa Rosa de Tastil.
Liolaemus calchaqui (19). TUCUMÁN. Departamento Trancas. FML 3082 (holotype); FML 2750 [9]: Punta de Agua. FML
9745: W of Hualinchay Ruta Provincial 352, Cumbres Calchaquíes. Departamento Tafí del Valle. FML 2425 [8]: Puesto
El Muñoz.
Liolaemus chacoensis (9). SANTIAGO Del ESTERO. Departamento Jiménez. FML 3063, [3], 15 km E of Comandante Pie-
drabuena. SALTA. Departamento de Anta. FML 2798 [4]; FML 6748: Finca Pozo Largo, 8 km S of Joaquín V.
González. LA RIOJA. Departamento Castro Barros. FML 7736: 33 km E of Anillaco.
Liolaemus crepuscularis (11): CATAMARCA. Departamento Andalgalá: FML 16622 (holotype) Puesto Flores, 2 km al S de
Mina Capillitas; FML 1623-29 [7] (paratypes): Puesto Flores, 2 km al S de Mina Capillitas; FML 21545-47 [3]: Cruce
entre camino a mina Santa Rita y Campo El Arenal, 3058 m.
Liolaemus darwinii (12). LA PAMPA. Departamento Curacó. FML 8373 [2]: 9 km E of Puelches. FML 8306 [2]: 40 km N of
Casa de Piedra, Ruta Provincial 23. MENDOZA. Las Heras. FML 2733 [2]: 27 km W of Mendoza, Ruta Nacional 7.
FML 263 [2]: Rodeo del Medio. NEUQUÉN. Departamento Zapala. FML 8316 [2]: between Zapala and Bosque Petri-
ficado. FML 8579: Cerro Lotena. RÍO NEGRO. Departamento General Roca. FML 8313: 20 km N of Cervantes.
Liolaemus diaguita (17). SALTA. Departamento Guachipas. MCN 3247 (holotype); MCN 3145-50, 3243, 3248-50; FML
3241-46 (paratypes): 61 km N of Cafayate, 400 m S of Río Las Conchas, Quebrada de las Conchas; 25°47’07.1’’S;
65°41’26.1’’ W. 1328 m.
Liolaemus espinozai (26): CATAMARCA. Departamento Andalgalá: FML 15527 (holotype); FML 15528-31 [4] (para-
types): El Ingenio, 67 km al S de Santa María por ruta provincial 47, 2620 m.; FML 3604 [21] (paratypes): El Ingenio, 67
km al S de Santa María por ruta provincial 47 (laderas al E del Campo de Arenal), 2620 m.
Liolaemus grosseorum (9). MENDOZA. Departamento San Rafael. FML 2972 [9]: Lago embalse El Nihuil.
Liolaemus irregularis (8). JUJUY. Departamento Tumbaya. FML 1313: Arroyo Aguas Blancas, Abra de Pives. Departa-
mento de Susques. FML 444 [2]: Sey. SALTA. Departamento Los Andes. FML 440 [5]: San Antonio de los Cobres.
Liolaemus koslowskyi (7). LA RIOJA. Departamento Famatina. FML 2659, (holotype); FML 2660 [6] (paratypes): 9,2 km E
of Central Plaza of Pituil, Ruta Provincial 11.
Liolaemus lavillai (25). SALTA. Departamento La Poma. FML 16597 (holotype); FML 16598-621 [24] (paratypes): 26,1 km
al N de La Poma, sobre ruta nacional 40.
Liolaemus laurenti (20). LA RIOJA. Departamento Famatina. FML 02518 (holotype); FML 02519 [4] (paratypes): 12 km E
of Central Plaza of Pituil, Ruta Provincial 11. FML 2524 [15]: 9,9 km al W de Antinaco.
ABDALA ET AL.
38 · Zootaxa 2968 © 2011 Magnolia Press
Liolaemus olongasta (13). LA RIOJA. Departamento General Lavalle. FML 2667 (holotype); FML 2668 [2] (paratipos):
51,3 km S of Villa Unión, Ruta Provincial 26. FML 2669 [10]: 51,3 km al S de Villa Unión, sobre ruta provincial 23.
Liolaemus ornatus (146). JUJUY. Departamento Rinconada. FML 1523 [56]: Mina Pirquitas. Departamento Cochinoca.
FML 1875 [11]; FML 2099 [16]: Abra Pampa. Departamento Humahuaca. FML 2076 [15]: Chuapi Rodeo. FML 2089
[9]: Mudana. FML 2100 [40]: Abra de Coranzuli. SALTA. Departamento Los Andes. FML 1366 [10]: Olacapato.
Liolaemus quilmes (54). CATAMARCA. Departamento Santa María. FML 1159 [6]; FML 1158 [5]: Santa María. FML
1838 [8]: El Puesto. SALTA. Departamento Cafayate. FML 2644 (holotype), FML 2645 [6] (paratypes): 3,2 km S of
Animaná, Ruta Nacional 40, Departamento Cachi. FML 1665 [17]: 3 km E of Cachi. TUCUMÁN. Departamento Tafí
del Valle. FML 2467 [6]: km 98 Ruta Provincial 307. FML 2291 [5]: Tafí del Valle.
Liolaemus uspallatensis (18). MENDOZA. Departamento Las Heras. FML 1541 [12]: 33 km N of Uspallata. FML 3466 [4]:
20 km N of Uspallata. FML 1773 [2]: Pampa de Yalguaraz.
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