Accepted by S. Carranza: 1 Jul. 2011; published: 31 Aug. 2011
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 3010: 31–46 (2011)
A new species of Liolaemus (Squamata, Iguania, Liolaemini) endemic to the Auca
Mahuida volcano, northwestern Patagonia, Argentina
LORENA ELIZABETH MARTINEZ1, LUCIANO JAVIER AVILA1, CRISTIAN HERNAN FULVIO PEREZ1,
DANIEL ROBERTO PEREZ2, JACK W. SITES, JR. 3 & MARIANA MORANDO1
1CENPAT-CONICET, Boulevard Almirante Brown 2915, U9120ACD, Puerto Madryn, Chubut, Argentina.
2Universidad Nacional del Comahue, Facultad de Ciencias del Ambiente y de la Salud, Buenos Aires 1400, 8300, Neuquén, Argentina
3Department of Biology and Monte L. Bean Life Science Museum, Brigham Young University, 401 WIDB, Provo, UT 84602, USA
We describe a new species of the Liolaemus bibronii complex. The new species differs from other members of the bi-
bronii-alticolor group in color pattern, coloration and squamation. Liolaemus cyaneinotatus is endemic to the Auca Ma-
huida volcano in northwestern Patagonia.
Key words: Iguania, Liolaemus, Auca Mahuida, Argentina, Liolaemus cyaneinotatus sp. nov., Patagonian lizards
Se describe una nueva especie del complejo Liolaemus bibronii. La nueva especie difiere de otros miembros del grupo
bibronii-alticolor por el patrón de coloración y por la escamación. Liolaemus cyaneinotatus es una población endémica
del volcán Auca Mahuida en el noroeste de Patagonia.
Palabras claves: Iguania, Liolaemus, Auca Mahuida, Argentina, Liolaemus cyaneinotatus sp. nov., lagartijas patagónicas
Liolaemus, the largest lizard genus in South America, ranges from central Peru to the northern tip of Tierra del
Fuego, and is divided in two main clades (Laurent 1983; Etheridge 1995; Schulte et al. 2000; Espinoza et al. 2004):
Liolaemus sensu stricto and Eulaemus. Within Liolaemus sensu stricto, which includes almost half of the described
species, several species groups can be recognized (Lobo et al. 2010). One group of small-sized species with slender
bodies, small limbs, and long tails is recognized as the alticolor-bibronii clade (Lobo et al. 2010). These authors
recognized in this group the following species: Liolaemus alticolor, L. araucaniensis, L. bibronii, L. bitaeniatus, L.
chaltin, L. curicensis, L. exploratorum, L. fuscus, L. gracilis, L. incaicus, L. lativitattus, L. lemniscatus, L. pagabu-
roi, L. paulinae, L. puna, L. ramirezae, L. saxatilis, L. tacnae, L. tandiliensis, L. variegatus, L. walkeri, and L.
yanalcu, but excluding L. robertmertensi and L. sanjuanensis previously related to L. bibronii (Schulte et al. 2000;
Lobo 2001; 2005; Cei 1986; 1993). In southern Argentina, a representative of this clade is the widely distributed
Liolaemus bibronii Bell 1843, a nominal species that includes several candidate species and is considered a species
complex (Morando et al. 2007). In northwestern Patagonia, the northern Neuquén and southern Mendoza provinces
are geographically very complex, with high mountains, deep valleys, and isolated plateaus and volcanic peaks,
coupled with a complex history of glaciations and pronounced climatic changes (e.g. Rabassa & Clapperton 1990;
Ramos & Kay 2006; Ramos & Folguera 2011). These characteristics most probably fostered multiple population
divergence processes across different geographic and temporal scales, and suggest that the region may be a
32 · Zootaxa 3010 © 2011 Magnolia Press
“hotspot” for Liolaemus diversification. Recent field surveys have revealed new species of Liolaemus lizards that
were previously either unknown or confused with other species (e.g.; Liolaemus thermarum, Videla & Cei 1996; L.
punmahuida, Avila et al. 2003; L. gununakuna, Avila et al. 2004; L. flavipiceus, Cei & Videla 2003; L. josei,
Abdala 2005; L. tregenzai, Pincheira- Donoso & Scolaro 2007; L. hermannunezi, Pincheira et al 2007; L. puelche,
Avila et al. 2007; L. cuyumhue, Avila et al. 2008; L. antumalguen, Avila et al. 2010). New surveys in poorly known
areas, coupled with the application of molecular techniques, have revealed additional “candidate” species of Liola-
emus that require detailed morphological study (Avila et al. 2006; Morando et al. 2003; 2004; 2007; unpublished
data). A combined morphological and molecular comparison allows us to consider an isolated population initially
identified as Liolaemus bibronii and inhabiting only the summit of the Auca Mahuida Volcanic Field, as a new spe-
cies which is described here.
Material and methods
We examined fifteen specimens of the new species and sample series of several populations of Liolaemus bibronii
from the type locality or surrounding areas (Appendix I), from LJAMM-CNP herpetological collection, Centro
Nacional Patagónico, Puerto Madryn, Argentina (CENPAT–CONICET), Museo de La Plata, La Plata, Argentina
(MLP.S), and Monte L. Bean Museum, Brigham Young University (BYU), Provo, UT, USA. Specimens were col-
lected by hand, sacrificed by pericardial injection of sodium pentothal Abbot®, fixed in 10–20% formalin and later
transferred to 70% ethanol. We compare the new species with the most closely distributed geographically species
of the group (Liolaemus araucaniensis, L. curicencis, L. fuscus, L. gracilis, L. lemniscatus). With the exception of
L. bibronii and L. gracilis, all other species of the group are geographically distributed far away from the Auca
Mahuida Volcanic Field, in isolated mountain systems (L. saxatilis and L. tandiliensis), more that 1000 km away in
Puna, or related environments (all the alticolor related species), and some species as L. araucaniensis, L. curicen-
sis, L. fuscus, and L. lemniscatus inhabit a very different biogeographic realm (Andean-Patagonian forest), mainly
in the western slope of Andes more than 300 km west.
Morphological data. Scale terminology and morphometric and color patterns follow Smith (1946), Etheridge
(1995), Lobo and Espinoza (1999), and Vega et al. (2008). Scale counts (meristic data) were made with a stereo-
scopic microscope, and measurements (morphometric data) were taken with a digital caliper to the nearest 0.1 mm.
Descriptions of color in life are based on notes taken in the field and color photographs of recently captured ani-
mals. The following morphometric traits were taken, as defined by Vega et al. (2008): snout–vent length (SVL),
head length (HL, from posterior edge of auricular opening to rostral scale), head width (HW, between corners of
the mouth), distance between fore and hind limbs (DFH, from axilla–groin), humerus length (HUL, from elbow to
axilla), radius-ulna length (RUL, from elbow to the internal angle between hand and forearm), hand length (HAL,
including fourth toe claw), femur length (FL, from knee to groin), tibio-fibula length (TFL, from knee to the inter-
nal angle with the foot), foot length (FOL, including fourth toe claw), fourth-toe length (4TL, including claw),
maximum body height (HMAX, at midbody), maximum body width (at midbody, WMAX), and tail length (TL).
Statistical analyses. Normal distributions and homoscedasticity of meristic and morphometric data were
examined with Shapiro-Wilk Test (W values), and Levene Test (F values), with their associated probabilities (P1
and P2). Meristic data were analyzed using a t-test for those variables which approximated normal distributions and
homocedasticity (Shapiro-Wilk P = 0.05 and Levene F = 0.05), and a Mann-Whitney U-test for variables that
departed from normality. As L. bibronii ( = 51.48 mm SVL, n = 58) and the new species ( = 49.08 mm SVL, n =
17) were not significatively different in size (t = 0.36, P = 0.7229), morphometric data were analyzed using a t-test
for all variables (Shapiro-Wilk P = 0.05; Zar 1984). Data for L. chaltin and L. puna were taken from Lobo and
Espinoza (2004) and data for L. tandiliensis from Vega et al. (2008).
Molecular procedures. Cytochrome b region (662 bp) was used for gene tree analysis, including 15 speci-
mens previously published in Morando et al. (2007), and three specimens of the new species were added for this
work (see Appendix II for details). Protocols for DNA extraction, mtDNA primer descriptions, PCR cycles, and
sequencing procedures followed Morando et al. (2003; 2004). Sequences were edited and aligned using the pro-
gram Sequencher 4.8 (™Gene Codes Corporation Inc. 1991–2007). Missing data were present in a few sequences,
and these were coded as “?”.
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A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA
Gene tree analyses. Bayesian analyses (based on TPM2uf+I+G model of evolution) determined using JMod-
eltest v0.1.1 (Guindon & Gascuel 2003; Posada 2008), were performed using MrBayes 3.1.2 (Huelsenbeck & Ron-
quist 2001). From a random starting tree, we performed four independent runs of 5 x 106 generations, and sampled
the Markov chains at intervals of 1,000 generations. We determined when stationarity was reached by plotting the
log-likelihood scores of sample points against generation time; when the values reached a stable equilibrium, sta-
tionarity was assumed. The trees were retained after a burn-in of 10,000 generations and were used to generate a
50% majority rule consensus tree. We consider P 0.95 as evidence of significant support for a clade (Huelsenbeck
& Ronquist 2001).
Liolaemus cyaneinotatus sp. nov.
Type material. Holotype. —MLP.S 2618 (Fig. 1), an adult male from Auca Mahuida (37° 41’ S, 68° 48’ W, 1332
m), Pehuenches Department, Neuquén Province, Argentina, C. Perez collector, 28 March 2008.
Paratypes.— MLP.S 2619 to 2622, LJAMM-CNP 10381, 10388 (females), LJAMM-CNP 10382-85, 10387
(male) (37° 41’ S, 68° 48’ W, 1332 m), C. Perez collector, 28 March 2008; LJAMM-CNP 10389 (male) (37° 43’ S,
68° 53’ W, 1757 m), C. Perez collector, 28 March 2008; LJAMM-CNP 10551 (male) and LJAMM-CNP 10552
(female), from Auca Mahuida (37° 43’ S, 68º 55’ W, 1983 m), C. Perez collector, 28 March 2008. All localities are
on Auca Mahuida Volcano, Pehuenches Department, Neuquén Province, Argentina.
Diagnosis. Liolaemus cyaneinotatus is a small, slender lizard, included in the alticolor-bibronii group (Lobo et
al. 2010). The new species differs from L. araucaniensis, L. bibronii, L. curicencis, L. fuscus, and L. lemniscatus in
having conspicuous cyan spots irregularly distributed in dorsal and lateral areas of the body. Liolaemus arau-
caniensis has a greenish background coloration, with well developed but irregular and sometimes fused dorsolat-
eral transversal marks, some fused with the vertebral line, a pattern never present in L. cyaneinotatus. Liolaemus
cyaneinotatus is a slightly smaller species than L. curicensis (maximum SVL 59.0 mm vs 64.2 mm) and L. arau-
caniensis (maximum SVL 60.9); both species have higher mean of snout-vent length (60.9 mm and 60.4 mm vs.
51.0 mm) and higher mean of distance from axilla–groin (27.2 mm and 28.8 mm vs. 21.7 mm). Liolaemus arau-
caniensis, L. curicensis and L. lemniscatus have higher mean tail length than L. cyaneinotatus (104.3 mm, 112.5
mm, and 105.0 mm vs. 82.3 mm). Vertebral line is absent in Liolaemus lemniscatus unlike the new species. Liolae-
mus cyaneinotatus differs from L. araucaniensis, L. curicensis, L. fuscus, L. gracilis and L. lemniscatus in having a
higher maximum number of scales around midbody (68 vs. 58, 56, 53, 44 and 58 respectively); L. curicensis has a
greater maximum number of fourth toe infradigital lamellae on the hind foot (27 vs 24) (Table 1). Liolaemus cya-
neinotatus differs from L. gracilis in having higher mean number of scales in the following characters: dorsal
scales (62.2 vs. 51.2), dorsal head scales (12 vs. 11.5), scales around interparietal (6.6 vs. 5.9), enlarged supraocu-
lars (6.1 vs. 5.9), number of temporals (7.2 vs. 6.4), number of gulars (28.9 vs. 25.3), scales around nasal (6.8 vs.
6.4) and infradigital lamellae of 4th toe of the hind foot (23.1 vs. 21.8). Liolaemus gracilis have always well
marked dorsolateral white stripes and usually lacks of paravertebral markings. Liolaemus bibronii have always
well marked black vertebral and dorsolateral lines, as well as a pattern of paravertebral markings well defined and
larger than L. cyaneinotatus; never have brilliant cyan scales dispersed in dorsal and lateral areas, and general
background coloration is always dark, almost black in some individuals, without orange or yellow-orange clear
coloration as L. cyaneinotatus. Mean of scales counts are higher in L. cyaneinotatus than L. bibronii in: dorsal head
scale (10.75 vs. 13.07), enlarged supraoculars (3.95 vs. 4.60), scales between rostral and frontal (4.84 vs. 5.40) and
infradigital lamellae of 4th toe of the foot (21.93 vs. 23.07). Liolaemus cyaneinotatus have lower mean number of
gulars scales (30.29 vs. 28.67), neck scales (16.27 vs. 14.67), and loreolabials scales (6.40 vs 5.60) than L. bibronii
(Tables 2 and 3).
Description of holotype. Adult male (Fig. 2): snout–vent length (SVL): 55.4 mm, tail length (TL): 78.29 mm
regenerated, axilla–groin distance (DFH): 25.01 mm, head length (HL): 12.29 mm, head wide (HW): 9.69 mm,
humerus length (HUL): 6.29 mm, radius-ulna length (RUL): 5.46 mm, hand length (HAL): 7.76 mm, femur length
(FL): 8.12 mm, tibio-fibula length (TFL): 10.70 mm, foot length (FOL): 15.19 mm, fourth-toe length (4TL): 13.04
mm, maximum body height (HMAX): 6.99 mm, maximum body width (WMAX): 13.83 mm.
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FIGURE 1. Upper: Liolaemus cyaneinotatus, adult male in lateral view, showing cyan spots. Below: Liolaemus bibronii, adult
male in lateral view from Puerto Deseado, Deseado Department, Santa Cruz Province (type locality).
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A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA
TABLE 1. Diagnostic characters among geographically proximal members of the bibronii-alticolor group; data for L. arau-
caniensis, L. curicencis, L. fuscus and L. lemniscatus were taken from Piñeira-Donoso and Nuñez (2005). Ranges are given in
parentheses; snout-vent, axila-groin, and tail lengths in mm; and elevation range in m.
Dorsal head scales smooth, 10 from a line drawn horizontally between anterior margin of external auditory
meatus to anterior margin of rostral. Left nasal scale in contact with rostral. Two postrostral scales and four interna-
sals. Canthals separated from nasal by one scale. Five enlarged supralabial scales, the fourth (2.54 mm) with the
posterior margin curved upward but not contacting subocular. Four infralabial scales. Auditory meatus oval
(height: 2.39 mm; width: 1.31 mm). Scales of anterior margin of auditory meatus without keels, scales of the poste-
rior margin smaller and granular. Three enlarged, flat scales projecting over meatus; no auricular scale differenti-
ated. Seven convex, imbricate and slightly keeled temporal scales between upper anterior corner of auditory
meatus and posterior margin of orbit. Six temporals (counting vertically from bucal commisure to level of supercil-
liaries). Interparietal scale subpentagonal, bordered by six scales. Parietals and interparietal of similar size. Orbit-
auditory meatus distance (4.46 mm), almost twice to orbit-rostral distance (anterior margin of rostral: 7.01 mm).
Rostral scale about twice wider than high (width: 2.66 mm; height: 1.23 mm). Mental scale subpentagonal, also
almost twice wider than high (width: 2.58 mm; length: 1.50 mm) in contact with first infralabial (on each side).
Frontal scale not divided. Five scales between frontal and rostral, five scales between frontal and supercilliaries.
Four enlarged supraoculars. Five strongly imbricate supercilliaries scales. Subocular longer (3.93 mm) than orbit
diameter (2.34 mm from anterior to posterior margin of cilliaries) and separated from supralabials by a single row
of lorilabials. Six lorilabials, with fourth through sixth contacting subocular. Preocular in contact to lorilabials. Pos-
tocular elongated, level with fifth supralabial scale. Posmental row formed by three enlarged scales, second pos-
mentals separated from each other. Scales of throat flat and imbricate. Twenty-seven gulars between auditory
Character cyaneinotatus n. sp. araucaniensis curicencis
n=15 n=1 n=34
Snout–vent length 51.0 (37.9–59.0) 60.9 60.4 (54.5–64.2)
Distance from axilla –groin 21.7 (15.3–25.8) 27.2 28.8 (27.0–31.7)
Vertebral line present present irregular
Tail length 82.3 (67.7–103.7) 104.3 112.5 (110.0–115.0)
Scales around midbody 60–68 58 49–56
Infradigital lamellae of 4th toe 21–24 22 24–27
Male precloacal pores 3–5 4 2–3
Females with precloacals pores absent absent absent
Reproductive mode ? oviparous oviparous
Elevational range 1300–2000 1400 1870–1950
Character fuscus gracilis lemniscatus
n=40 n=47 n=97
Snout–vent length 49.6 (48.2–50.5) 48.0 (39.4–57.3) 51.3 (45.8–55.7)
Distance from axilla–groin 22.4 (21.8–23.6) 22.1 (17.3–28.1) 23.7 (21.0–27.0)
Vertebral line present present absent
Tail length 85.6 (79.0–90.0) 92.8 (72.2–110.2) 105
Scales around midbody 47–53 34–44 50–58
Infradigital lamellae of 4th toe 22–25 20–26 23–24
Male precloacal pores 2–3 3–4 2–3
Females with precloacals pores absent absent absent
Reproductive mode oviparous oviparous oviparous
Elevational range 1600 0–1400 1800
36 · Zootaxa 3010 © 2011 Magnolia Press
meatus. Lateral side of the neck flat, formed by small scales, imbricate and keeled. Dorsal scales lanceolate, imbri-
cate and moderately keeled. Forty-nine scales around midbody. Sixty-three dorsal scales between occiput and ante-
rior level of thigh. Ventral scales larger than dorsals (dorsal scale length: 1.22 mm; ventral scale length: 1.39 mm).
Eigthy ventral scales counted at the midline between rostral scale and vent. Three precloacal pores. Seventeen sub-
digital lamellae in the fourth toe of the hand and twenty-four on the foot. Anterior suprabrachials rhomboidal,
imbricate, smooth, slightly larger in size to dorsal body scales. Postabrachials smaller, smooth, becoming granular
near axilla. Supra-antebrachials similar to suprabrachial. Pre-antebrachials imbricated, rhomboidal, keeled. Infra-
antebrachials rhombals, imbricate, keeled. Supracarpals imbricated, rhomboidal, smooth. Infracarpals strongly
imbricate, rhomboidal, keeled, mucronate. Subdigital lamellae with 3 keels, each terminating in a short mucron,
numbering: I: 9, II: 12, III: 15, IV: 17, V: 12. Claws robust, curved and sharp, opaque brown.
FIGURE 2. Holotype of Liolaemus cyaneinotatus MLP.S 2618, adult male in dorsal and ventral view from Auca Mahuida vol-
cano, Pehuenches Department, Neuquén province, Argentina.
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A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA
Suprafemorals as large as dorsal body scales, rhomboidal, imbricated, slightly keeled near the insertion to
strongly keeled and terminating in a short mucron. Infrafemorals and prefemorals rhomboidal, imbricated, smooth.
Postfemorals small, granular shape. Supratibial rhomboidal, imbricated, strongly keeled and with a short mucron.
Infratibials rhomboidal, imbricated, smooth. Supratarsals rhomboidal, imbricated, smooth with some scales
slightly keeled in the outer side. Infratarsal small, rhomboidal, imbricate, keeled, mucronate. Subdigital lamellaes
of fingers with three blunt keels each terminating in 3 short mucrons, numbering: I: 10, II: 14, III: 22, IV: 23, V:
broken. Claws robust, curved and sharp, opaque brown. Tail quadrangular in cross section near cloacal area,
becoming oval to round towards the tip. Dorsal and upper lateral caudal scales keeled, imbricate, mucronate.
Lower lateral scales and ventral scales weakly keeled.
Color in life. Head predominantly light brown, darker than body; some scales with a brick-orange tinge. A dis-
tinctive white line between postocular scale and antehumeral fold, a scale wide, black outlined. Subocular scale
white, with black upper border. Dorsal background coloration is ferruginous orange. Dorsal pattern with a tenuous
light brown vertebral field 2–3 scales wide at midbody. Vertebral line black, well marked, continuous from occiput
to first third of the tail, becoming broken to the tip. Paravertebral field, four-five scales wide, slightly darker than
vertebral field, with dark ferruginous orange. Paravertebral markings black, irregulars, a scale size; eight series
between shoulders and pelvic girdle, not paired. Between occiput and shoulder, paravertebral markings fused,
forming two continuous black lines; after pelvic girdle becomes regularly paired and fused to the tip of the tail.
Dorsolateral stripes whitish, one scale wide, originating on the posterior border of each orbit, and disappearing in
the first third of trunk. Iridescent cyan scales scattered at midbody following the dorsolateral stripes and contact
area between paravertebral and lateral fields. Lateral field with a light orange background slightly darker than par-
avertebral field, speckled with black, white, and iridescent cyan scales (Fig. 1). Lateral field coloration progres-
sively fades ventrally, where light gray replaces the orange background coloration, except in the proximity of the
hind limbs insertions were the orange coloration becomes more intense. Throat background light gray, with a black
reticulated pattern. Chest, belly, tail, and ventral surfaces of the thighs light gray. Lower belly, cloacal apron, ven-
tral femoral area, and first third of the tail, yellow-orange. First lateral portion of the tail bright orange. Precloacal
pores bright orange.
Color in preservative. Coloration pattern is maintained but background coloration becomes darker. All cyan
dots disappear as well as all orange or yellow coloration. Ventral coloration becomes gray, more intense in the mid-
dle of each scale.
Variation. Table 4 summarizes morphometric and scale variation data. In this section we provide information
on traits not included in the table. Dorsal head scale surface nearly uniform, smooth (or slightly wrinkled in seven
specimens); temporal scales keeled in five, smooth in ten. Fourth supralabial scale is curved upward at its posterior
margin in all except five specimens, and only contacts the subocular in only five individuals. Three individuals
with nasal scale separated from rostral. Posterior half of supraorbital semicircles complete in thirteen, incomplete
in two. Canthal scale separated from nasal by two scales, except one specimen separated by one scale. Frontal scale
divided in three, undivided in twelve. Vertebral line varies from well marked to completely absent. Paravertebral
field disappear completely in some individuals. Paravertebral marks become fused in some individuals forming a
line equal to the vertebral line, or are reduced to small dots or disappear in some individuals. Some males have a
dorsal plain pattern alive, without vertebral or paravertebral lines or paravertebral marks. Dorsal background color-
ation varies from light gray to brick orange. Lateral field varies in melanism according to the density of black
scales. Cyan dots are present in adult males, but can disappear in subadults. Density and distribution of cyan dots
varies between individuals but always are in the lateral and/or dorsolateral areas of the trunk. Usually are absent in
females. Yellow coloration is usually present in ventral areas of femurs, cloacal apron, first third of the tail and
lower belly and varies in intensity from individual to individual becoming more extensive in females that in males.
In males usually predominates yellow that orange coloration. Brick orange coloration is more intense in females
and is stronger in lateral sides of the trunk, lateral areas of the first third of the tail, and prefemoral areas. In some
females orange brick coloration form a continuous line between postoculars and antehumeral fold, upper part of the
lateral field between axilla and prefemoral region, and from postfemoral area to the tip of the tail.
Etymology. Specific epithet is in reference to the cyan dots present in adult males in dorsolateral areas of the
body, (cyanei = cyan; notatus = dotted).
40 · Zootaxa 3010 © 2011 Magnolia Press
TABLE 4. Between sex variation in meristic and morphometric characteristics of Liolaemus cyaneinotatus. Measurements are
in mm, numbers for each variable indicate mean ± standard deviation, with range in parentheses. Precloacal pores, infra and
supralabials scales are shown as ranges.
Geographic distribution. Liolaemus cyaneinotatus is known only from the type locality where the holotype
and paratypes were collected in rocky patches or open substrate outcrops above 1500 m elevation on Auca Mahu-
ida volcano, Añelo Department, Neuquén Province, (Fig. 3). Auca Mahuida Volcanic Field is part of the Payenia, a
large Quaternary volcanic province of basaltic composition in the foreland region, behind the active volcanic arc,
unique in the entire Andean Chain (Ramos & Folguera 2011). The Auca Mahuida Volcanic Field is an isolated
basaltic plateau with the Auca Mahuida volcano in its center, surrounded by dunes fields or sandy flatlands with
extremely arid characteristics, and is an island of Patagonian-like environment in the middle of Monte phytogeo-
graphic region. Liolaemus cyaneinotatus is not in contact with the geographic distribution range of other members
of the L. bibronii group, with the exception of L. gracilis, found in lower altitudes of the Auca Mahuida volcano.
But no sympatry was observed between these two species.
Variable Males (n = 8) Females (n = 7)
Snout–vent length 52.4 ± 6.5(37.9–59.0) 49.2 ± 3.9(41.6–53.7)
Head length 11.5 ± 1.3(8.7–13.0) 10.4 ± 0.6(9.4–11.3)
Head width 9.2 ± 1.1(7.0–10.8) 8.2 ± 0.7(7.1–9.0)
Distance between fore and hind limbs 22.8 ± 3.0(16.2–25.1) 22.2 ± 2.5(17.7–25.8)
Humerus length 5.9 ± 0.6(4.7–6.9) 5.4 ± 0.3(5.1–5.9)
Radius-ulna length 6.2 ± 0.9(4.6–7.7) 5.6 ± 0.4(4.76–6.1)
Hand length 8.1 ± 0.9(6.2–9.3) 7.4 ± 0.7(6.4–8.4)
Femur length 7.9 ± 1.1(5.5–9.5) 7.2 ± 0.5(6.5–7.7)
Tibio-fibula length 10.5 ± 1.2(7.8–11.5) 9.5 ± 0.5(8.6–10.2)
Foot length 15.4 ± 1.6(11.8–16.9) 14.2 ± 0.7(13.1–15.2)
Fourth-toe length 13.4 ± 1.4(10.2–15.1) 12.3 ± 0.6(11.6–13.1)
Maximum body height 7.2 ± 1.0(5.3–8.6) 7.0 ± 0.7(5.6–7.7)
Maximum body width 11.9 ± 2.2(7.9–15.5) 12.6 ± 1.5(10.5–15.1)
Tail length 86.8 ± 13.3(71.8–103.7) 79.3 ± 7.9(67.7–89.1)
Dorsal scales between occiput and thigh 62.7 ± 2.0(60.0–65.0) 61.8 ± 2.9(60–68)
Scales around midbody 48.6 ± 2.2(46.0–53.0) 50.1 ± 1.6(48–53)
Dorsal head scales 12.1 ± 0.3(12.0–13.0) 12
Ventral scales 83.1 ± 2.3(80.0–86.0) 84.4 ± 6.1(75–91)
Precloacal pores 3–50
Scales around interparietal 6.6 ± 1.0(6–9) 6.0 ± 0.6(5–7)
Enlarged supraoculars 4.4 ± 0.7(4–6) 4.8 ± 0.9(4–6)
Number of temporals 7.4 ± 0.5(7–8) 7.1 ± 0.9(6–9)
Number of neck scales 14.9 ± 1.5(13–17) 14.4 ± 1.1(13–16)
Number of gulars 28.4 ± 1.6(27–31) 29.0 ± 2.7(24–32)
Enlarged supralabials 5–66–7
Enlarged infralabials 4–54
Scales around nasal 6.7 ± 0.4(6–7) 7
Scales between rostral and frontal 5.2 ± 0.4(5–6) 5.6 ± 0.8(5–7)
Scales between frontal and supercilliaries 4.6 ± 0.7(3–5) 4.1 ± 0.4(4–5)
Loreolabials scales 5.9 ± 0.6 (5–7) 5.3 ± 0.5(5–6)
Infradigital lamellae of 4th toe of the hand 17.5 ± 1.4(15–19) 16.6 ± 0.5(16–17)
Infradigital lamellae of 4th toe of the pes 23.5 ± 0.9(22–24) 22.6 ± 1.1(21–24)
Zootaxa 3010 © 2011 Magnolia Press · 41
A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA
FIGURE 3. Complex landscape of northwestern Neuquén region; red line marks the Liolaemus cyaneinotatus approximate
distribution. Yellow broken line mark limits of the Auca Mahuida Volcanic Field. Major mountain ranges, cities, and roads are
also marked as reference. Inset: Region in South América.
42 · Zootaxa 3010 © 2011 Magnolia Press
FIGURE 4. Type locality of Liolaemus cyaneinotatus. Upper: general view of the area. Below: close view of the open areas
where lizards were collected.
Zootaxa 3010 © 2011 Magnolia Press · 43
A NEW SPECIES OF LIOLAEMUS FROM PATAGONIA
FIGURE 5. Bayesian consensus tree of relationships of Liolaemus cyaneinotatus with other species of the bibronii group; pos-
terior probability values are shown at each node.
Natural history. The individuals were found in active basking in the edges of shrubs (Prosopis denudans,
Senna kurtzii, S. arnottiana) or on the top of rocks. Individuals were found active between 09:00 to 19:00 in sunny
days of austral summer. Activity starts with basking behavior along the edges of the bushes or rocks. Several indi-
viduals were observed sharing basking areas and usually juveniles and adults are found under small bushes, sharing
refugee or foraging areas. They usually foraged actively in the bushes but use the sand edges or tip of branches to
bask. They seem to be territorial at some point because aggressive encounters between large males or females were
observed. This species shares its habitat with other Liolaemus species (Liolaemus austromendocinus, Liolaemus
aff. elongatus, Liolaemus aff. boulengeri, Phymaturus sitesi and P. roigorum). Liolaemus cyaneinotatus usually
occupies the microhabitat around the shrubs that grow scattered separated by an open, loose sand substrate (Fig. 4).
No data about reproduction is available, but L. cyaneinotatus is probably oviparous like L. bibronii (Medina et al.
2008). Data about diet came from the analysis of two guts, ant heads and beetle legs were found, with partially
digested small leaves and vegetable scraps. This suggests that L. cyaneinotatus is similar to other lizards in its use
of invertebrates and plants as food resources (Videla 1983; Acosta et al. 1996; Quatrini et al. 2001; Belver & Avila
44 · Zootaxa 3010 © 2011 Magnolia Press
Remarks. Lizards of the Liolaemus alticolor-bibronii group have the largest latitudinal distribution in a Lio-
laemus species group, with L. walkeri from Junín Province in central Peru to L. bibronii in central Santa Cruz prov-
ince in southern Argentina. Some species are found only in arid and semiarid environments, including Puna of
Peru, Argentina, Bolivia, and Chile as L. alticolor, L. bitaeniatus, L. chaltin, L. incaicus, L. lativittatus, L. pagabu-
roi, L. paulinae, L. puna, L. ramirezae, L. tacnae, L. variegatus, L. walkeri, L. yanalcu, meanwhile others are
restricted to some mountain ranges as L. saxatilis (Sierras de Cordoba and San Luis), L. tandiliensis (Tandil Moun-
tains), or particular biogeographic regions as L. araucaniensis. Liolaemus lemniscatus is found in several habitats
of the western slopes of Andean mountains and in northern Andean-Patagonian forest; L. curicensis and L. fuscus
are found only in central Chile region. Liolaemus gracilis is found in environments of central Argentina, but L.
exploratorum is an enigmatic species described based on museum specimens and never found again in its pur-
ported range of distribution. Liolaemus bibronii is the species with probably the largest geographic range in lati-
tude, but Morando et al. (2007) found that this may be a species complex with several candidate species.
A mtDNA gene tree analysis, including the new described species as well as other related species and candi-
date species of the group is depicted on Fig. 5. This tree is based on the mitochondrial gene fragment cyt-b (662
bp). Liolaemus cyaneinotatus is recovered within the bibronii north clade, that also includes L. gracilis, L. ramir-
ezae, L. saxatilis, L. robertmertensi and several candidate species (Posterior Probability PP: 1). The bibronii south
clade includes the haplotypes from the type locality of L. bibronii and also includes candidate species and is recov-
ered with strong support (PP: 1). The objective of this tree is to show the position of this new species in relation
with other terminal taxa of the L. bibronii complex. Species limits based on only one gene have limitations and
although Liolaemus cyaneinotatus haplotypes are recovered as monophyletic, which strongly supports this new
species; an integrative approach using a multi-loci species tree method and morphological analyses are needed in
order to fully understand the phylogenetic relationships of the bibronii complex. A detail study with these charac-
teristics is being carried out by our research group and will be published elsewhere.
We thank N. Feltrin, M. Breitman, M. Kozykariski, N. Frutos, C. Zanotti, R. Otteson, M. Hawkins, K. Dittmar, J.
Goldman and A. Cosacov for help in field collections of L. bibronii species group, and N. Cazzaniga for help with
Latin language issues. We thank REPSOL YPF for support the project “Biodiversidad en el Area Sensible Auca
Mahuida. Evaluación de la presencia de posibles especies endémicas y lineamientos de gestión para la prevención,
minimización y mitigación de impactos sobre la biodiversidad” (Convenio Biodiversidad VAM CM 4900047452),
issued to D.R. Perez. We acknowledge the NSF “Partnership for International Research and Education” award
(OISE 0530267) for support collaborative research on Patagonian biodiversity, granted to the following institutions
(listed alphabetically): BYU, CENPAT, Dalhousie University, Darwinion Botanical Institute, George Washington
University, Universidad Nacional de Cordoba, Universidad Austral de Chile, Universidad Nacional del Comahue,
and Universidad de Concepción. Additional support was provided by a CONICET PIP 6469 issued to L. Avila. We
thank fauna and conservation authorities of Neuquén province for collection permits and work authorizations.
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APPENDIX I. Specimens examined.
Liolaemus bibronii (n=59): ARGENTINA: SANTA CRUZ: Deseado Department: National Road 3, 6 km N Tres Cerros (48°
04’ 15.2” S, 67° 37’ 05.6” W): LJAMM-CNP 4558/59-4680-4724. National Road 3, 30 km S Caleta Olivia, (46° 40’ 56.4”
S, 67° 23’ 23.7” W): LJAMM-CNP: 4563/64. National Road 281, 45 km NW Tellier, (47° 28’ 20” S, 66° 33’ 14” W):
LJAMM-CNP 5909/10. National Road 281 junction National Road 3, 7 km NW Jaramillo, (47° 08’ 17.9” S, 67° 12’ 44.1”
W): LJAMM-CNP 5912. Provincial Road 43, 16 km E Las Heras, (46° 33’ 50.5” S, 68° 40’ 16.3” W): LJAMM-CNP
5913. 1 km W Tellier, (47° 39’ 12.5” S, 66° 03’ 05.8” W): LJAMM-CNP 5915. 5.5 km N Puerto Deseado by coastal road,
(47° 43’ 01”S -65 50’ 28” W): LJAMM-CNP 7475-7476. Road to Los Perales oil camp, 3 km S junction National Road 26
to Provincial Road 18, (46° 03’ 22” S, -69° 20’ 31” W): LJAMM-CNP 9159 to 61. 5.5 km N Puerto Deseado, (47° 42’
54”S, 65° 50’ 21” W): LJAMM-CNP 9886- 9887 to 96. Provincial Road 47, 55.4 km SW Tellier, 3 km S bridge on Rio
Deseado, juction Provincial Road 89, (47° 51’ 01” S, 66° 37’ 20” W): LJAMM-CNP 9897 to 9903. Provincial Road 75,
2.8 km W junction Provincial Road 87, (48° 08’ 34” S, 68 12’ 47” W): LJAMM-CNP 9919/20. Provincial Road 12, 9.8
km N junction Provincial Road 75, 3 km S Gobernador Moyano, (47° 47’ 37” S, 68° 35’ 32” W): LJAMM-CNP 9921/23.
Provincial Road 12, 86.5 km N junction Provincial Road 49, 87.5 km SW Pico Truncado, (46° 58’ 03” S, 68° 24’ 58” W):
LJAMM-CNP 9924/25. Provincial Road 75, 2.8 km W junction Provincial Road 87, (48° 08’ 34” S, 68° 12’ 47” W):
LJAMM-CNP 10072 to 82. Provincial Road 12, 25.8 km N junction Provincial Road 49, 27 km N Aguada del Cuero
Ranch, (47° 26’ 05” S, 68° 34’ 59” W): LJAMM-CNP 10123/24. Liolaemus gracilis (n=45): ARGENTINA. MENDOZA:
Malargüe Department: Provincial Road 186, 70.8 km E junction National Road 40, 2 km SW El Molino post, (35° 55’
48.8” S, 69° 04’ 47.8” W): LJAMM-CNP 7932. Provincial Road 180, 15 km S La Cortadera, (36° 39’ 44” S, 68° 40’ 32”
W): LJAMM-CNP 5060.LA PAMPA: Curacó Department: Provincial Road 23, 40 Km N Casa de Piedra: LJAMM-CNP
120. Provincial Road N° 23, N Casa de Piedra, (38° 07’ S, 67° 06’ W): LJAMM-CNP 144- 147- 149 to 152. Puelén
Department, Provincial Road 23, 6.6 km S Provincial Road 26, (37° 56’ 23” S, 67° 06’ 32.1” W): LJAMM-CNP 8432.
NEUQUEN: Añelo Department, Provincial Road. 7, 28.7 km N Añelo., (38° 11’ 04.9” S, 69° 01’ 22.5” W): LJAMM-CNP
5699- 5711 to 13. Collón Curá Department, Provincial Road 47, 38,1 km N junction National Road 237, 18.1 km N Santo
Tomás, (39° 40’ 52.7” S, 70° 00’ 13.5” W): LJAMM-CNP 8907- 8908- 8909. RIO NEGRO: El Cuy Department, El Cuy,
(39° 55’ 33” S, 68° 20’ 37” W): LJAMM-CNP 1589/90. Provincial Road 71, 41.7 km SW junction Provincial Road 6, (39°
39’ 58” S, 68° 24’ 36.5” W): LJAMM-CNP 7114 to 17. San Antonio Department, Las Grutas. Piedra Colorada Beach,
(40° 50’ 26” S, 65° 07’ 04” W): LJAMM-CNP 2572. General Roca Department: 25 Km N Cervantes, near salty lagoon:
LJAMM-CNP 5170. 38 km N National Road 22 to General Roca from junction to Casa de Piedra: LJAMM-CNP 5175 to
78. Valcheta Department, 10.5 km NE Bajo Rico, (40° 07’ 9.8” S, 66° 25’ 54.2” W): LJAMM-CNP 6289. El Cuy, 13.5 km
W junction Provincial Road 6 and Provincial Road 71 to Cerro Policia: LJAMM-CNP 7113. CHUBUT: Biedma Depart-
ment: San Pablo Ranch, (42° 36’ 54” S, 64° 10’ 26” W): LJAMM-CNP 3330 to 37- 3340. Puerto Madryn, (42° 46’ S, 65°
03’ W): LJAMM-CNP 3344- 3698- 4499. Telsen Department, Provincial Road 4, 3.5 km W Telsen, (42° 26’ 27.8” S, 66°
58’ 50.6” W): LJAMM-CNP 5486. Provincial Road 61 40.3 km junction Provincial Road 11 between Ranquilhuao and
San Manuel Ranchs, (42° 44’ 48.8” S, 66° 59’ 54.8” W): LJAMM-CNP 5947. Liolaemus cyaneinotatus (n=15): ARGEN-
TINA. NEUQUEN. Pehuenche Department. Auca Mahuida volcano, (37° 41’ 27”S, 68° 48’ 11”): LJAMM-CNP 10375 to
78- 10381 to 88. (37° 43’ 23” S, 68° 53’ 18” W): LJAMM-CNP 10389. (37° 43’ 33” S, 68° 55’ 34” W): LJAMM-CNP
APPENDIX II. Specimens used in molecular analysis.
Liolaemus punmahuida LJAMM-CNP 2626; Liolaemus neuquensis LJAMM-CNP 2665; Liolaemus robertmertensi LJAMM-
CNP 1961; Liolaemus saxatilis LJAMM-CNP 8456; Liolaemus gracilis LJAMM-CNP 8432; Liolaemus ramirezae
LJAMM-CNP 4416; Liolaemus cyaneinotatus LJAMM-CNP 10380- 10381- 10382; Liolaemus bibronii: type locality:
LJAMM-CNP 9924; Lb 1 LJAMM-CNP 8124; Lb 2 LJAMM-CNP 9054; Lb 3 LJAMM-CNP 8686; Lb 4 LJAMM-CNP
5654; Lb 8 LJAMM-CNP 8011; Lb 9 LJAMM-CNP 6426; Lb 10 LJAMM-CNP 5387; Lb 11 LJAMM-CNP 5323.