Two new species of lizards from the Liolaemus lineomaculatus section (Squamata: Iguania: Liolaemidae) from southern Patagonia

Abstract

Two new species of the lineomaculatus clade of the Liolaemus lineomaculatus section are described from southern Pa-tagonia in Argentina. Liolaemus morandae sp. nov. is found in S Chubut province and Liolaemus avilae sp. nov. inhabits NW Santa Cruz province. Several tests were performed to diagnose these new species as distinct lineages. Univariate anal-ysis of variance (ANOVA), principal component analysis (PCA), discriminant function analysis (DFA), non-parametric multivariate analysis of variance (NPMANOVA), as well as a genetic characterization through molecular analysis of vari-ance (AMOVA) were performed; genetic distances between described and these new species are reported. The new Lio-laemus species differ from other members of the lineomaculatus group in morphometric, meristic, qualitative and genetic characters; moreover they inhabit different phytogeographical provinces and districts. With these descriptions, the number of species now recognized in the lineomaculatus section is twenty one (including one more description that is in press).

Full text

Accepted by S. Carranza: 18 Oct. 2011; published: 6 Dec. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 3120: 1–28 (2011)
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1
Two new species of lizards from the Liolaemus lineomaculatus section
(Squamata: Iguania: Liolaemidae) from southern Patagonia
MARIA FLORENCIA BREITMAN1,4, MICAELA PARRA2, CRISTIAN HERNÁN FULVIO PÉREZ1
& JACK WALTER SITES, JR 3
1Centro Nacional Patagónico – Consejo Nacional de Investigaciones Científicas y Técnicas, Boulevard Almirante Brown 2915, ZC:
U9120ACD, Puerto Madryn, Chubut, Argentina. Tel: 0054 – 02965 – 451024. E-mail: chfperez@cenpat.edu.ar
2Universidad Nacional de la Patagonia San Juan Bosco, Boulevard Almirante Brown 3150, ZC: U9120ACX, Puerto Madryn, Chubut,
Argentina. Tel: 0054 – 02965 – 472885. E-mail: mica_fop@hotmail.com
3Department of Biology and Monte L. Bean Life Science Museum, Brigham Young University, 401 WIDB, Provo, UT 84602, USA.
E-mail: jack_sites@byu.edu
4Corresponding author. E-mail: breitman@cenpat.edu.ar
Abstract
Two new species of the lineomaculatus clade of the Liolaemus lineomaculatus section are described from southern Pa-
tagonia in Argentina. Liolaemus morandae sp. nov. is found in S Chubut province and Liolaemus avilae sp. nov. inhabits
NW Santa Cruz province. Several tests were performed to diagnose these new species as distinct lineages. Univariate anal-
ysis of variance (ANOVA), principal component analysis (PCA), discriminant function analysis (DFA), non-parametric
multivariate analysis of variance (NPMANOVA), as well as a genetic characterization through molecular analysis of vari-
ance (AMOVA) were performed; genetic distances between described and these new species are reported. The new Lio-
laemus species differ from other members of the lineomaculatus group in morphometric, meristic, qualitative and genetic
characters; moreover they inhabit different phytogeographical provinces and districts. With these descriptions, the number
of species now recognized in the lineomaculatus section is twenty one (including one more description that is in press).
Key words: Iguania, Liolaemidae, Liolaemus lineomaculatus section, Liolaemus morandae sp. nov., Liolaemus avilae
sp. nov., Patagonia, new species
Resumen
Se describen dos nuevas especies pertenecientes al clado lineomaculatus de la sección Liolaemus lineomaculatus de la
región sur de la Patagonia Argentina. Liolaemus morandae sp. nov. se distribuye en el sur de la provincia del Chubut,
mientras que Liolaemus avilae sp. nov. se encuentra circunscrita al Noroeste de la provincia de Santa Cruz. En este trabajo
se utilizan diferentes análisis estadísticos y moleculares para caracterizar y diferenciar las nuevas especies de las actual-
mente descritas en el grupo lineomaculatus: análisis univariados de la varianza (ANOVA), análisis de componentes prin-
cipales (PCA), análisis de funciones discriminantes (DFA), análisis multivariados no paramétricos de la varianza
(NPMANOVA); también se realiza una caracterización genética a través de un análisis molecular de la varianza (AMO-
VA) y se reportan las distancias genéticas entre especies. Las nuevas especies de Liolaemus se diferencian de los otros
miembros del grupo lineomaculatus en base a caracteres morfológicos, cualitativos y genéticos; además de habitar difer-
entes provincias fitogeográficas y distritos. El número actual de especies de la sección lineomaculatus asciende a veinti-
uno, incluyendo la presente descripción y otra que se encuentra en prensa.
Palabras clave: Iguania, Liolaemidae, sección Liolaemus lineomaculatus, Liolaemus morandae sp. nov., Liolaemus avi-
lae sp. nov., Patagonia, nueva especie

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Introduction
One of the most ecologically diverse and species-rich genera of iguanian lizards on earth is Liolaemus, with more
than 230 recognized species distributed from Perú to Tierra del Fuego (Núñez & Scolaro 2009; Abdala et al. 2010;
Avila et al. 2010a, 2010b; Lobo et al. 2010a, 2010b; Martinez et al. 2011; Quinteros & Abdala 2011; Breitman et
al. in press). These lizards extend across a wide range of latitudinal (14° ± 30’ – 52° ± 30’ S), altitudinal (0 – 4,500
m), and climatic regimes (Hellmich 1951; Donoso-Barros 1966; Etheridge & de Queiroz 1988; Frost & Etheridge
1989; Cei 1986, 1993; Etheridge & Espinoza 2000). Two main species groups have been recognized within Liolae-
mus since the first comprehensive study of the genus (Laurent 1983): Liolaemus sensu stricto (the Chilean group)
and the Eulaemus (the Argentinean group). The Liolaemus lineomaculatus section is part of the subgenus Eulae-
mus and includes 19 formally described species (Breitman et al. 2011; Breitman et al. in press). The distribution of
the L. lineomaculatus section extends from central Neuquén province in northwest Argentinean Patagonia, and
south to Tierra del Fuego Island (Donoso-Barros & Codoceo 1962; Donoso-Barros 1966; Bottari 1975; Cei 1986).
Extremely heterogeneous landscapes are occupied by lizards from the L. lineomaculatus section; thus, its phyloge-
netic/phylogeographic history has also likely been complex (Breitman et al. 2011).
Species of the L. lineomaculatus section were historically classified into three main morphological groups: lin-
eomaculatus (Etheridge 1995), kingii (Cei 1986) and archeforus (Cei 1986); plus the species L. magellanicus that
has been recognized as part of the section, but was not clearly assigned to any of these three main groups. Breitman
et al. (2011), based on nine molecular markers (mitochondrial and nuclear) and two phylogenetic approaches (con-
catenation and species tree), identified four main lineages within the L. lineomaculatus section: (1) the lineomacu-
latus group including: L. hatcheri, L. kolengh, L. silvanae and L. lineomaculatus; (2) the magellanicus group
including L. magellanicus and one more species under description (Breitman et al. in press); (3) the somuncurae
group including L. somuncurae and L. uptoni; and (4) the kingii+archeforus group including: L. baguali, L. escar-
chadosi, L. tari, L. sarmientoi, L. scolaroi, L. zullyae, L. tristis, L. archeforus, L. chacabucoense, L. kingii and L.
gallardoi (Figure 1). The four clades recovered from the molecular approach support the “traditional morphologi-
cal” lineomaculatus group, but do not support the kingii and archeforus groups; moreover they included eight
potentially new undescribed species spread through the four main clades.
Liolaemus lineomaculatus was described by Boulenger in 1885, with “Patagonia” as the type locality; later
Donoso-Barros (1966) restricted the type locality to “Puerto Deseado, Patagonia, Argentina”. Since then, speci-
mens possessing morphological features of L. lineomaculatus and collected between central Neuquén and southern
Santa Cruz were assigned to this species (e.g. Cei 1986; Williams 1997; Ibargüengoytía et al. 2001; Christie 2002).
However, recent evidence suggests that this species is a species complex comprised of at least five lineages (Breit-
man et al. 2011; Breitman unpublished data).
Here we used an integrative approach to distinguish and describe two new species from the lineomaculatus
group of the L. lineomaculatus section. Morphometric, meristic, qualitative and genetic characters, were used to
analyze this group of species through a battery of statistical tests. The two new species described here belong to the
lineomaculatus group identified as L. sp. 1 and L. sp. 2 by Breitman et al. (2011).
Material and methods
We examined series of specimens from the L. lineomaculatus clade, particularly those of the lineomaculatus group
(specimens that were used in Breitman et al. 2011). Liolaemus kolengh (n = 42), L. hatcheri (n = 20), L. silvanae (n
= 21), and L. lineomaculatus (n =17), including specimens collected from their type localities, were compared with
nine specimens of the first new species, and ten specimens of the second new species (Appendix I). Specimens
were collected by hand and sacrificed by a pericardial injection of sodium pentothal Abbot®, slightly dissected to
extract a sample of liver for molecular study, fixed in 20% formalin and later transferred to 70% ethanol. Voucher
specimens were placed in the La Plata Museum (MLP.S) and in the Herpetological Collection LJAMM-CNP of
Centro Nacional Patagónico in Puerto Madryn, Argentina.
The lineomaculatus and magellanicus groups are morphologically different (scale counts and shape, coloration
patterns, presence/absence of precloacal pores, among others; Scolaro 1992; Laurent 1995; Etheridge & Espinoza
2000; Pincheira-Donoso & Núñez 2005), and divergence of these groups is inferred to be Late Miocene (~8.46 mil-

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TWO NEW SPECIES FROM L. LINEOMACULATUS SECTION
lion years ago, Breitman et al. 2011). Because they are considered as two different morphological and molecular
clades, for the present descriptions we only performed comparisons among the species of the lineomaculatus
group. We used 18 morphometric characters and 12 meristic characters (scale counts) commonly used (defined
elsewhere, e.g. Vega et al. 2008; Avila et al. 2010a, 2010b; Martinez et al. 2011; see Appendix II for names and
definitions of characters), as well as 94 qualitative characters, 84 on squamation and 10 on body patterns. Measure-
ments were taken only on adults, with a Schwyz® electronic digital caliper to the nearest 0.1 mm; scale counts
were made on juveniles and adults with a stereoscopic microscope from fixed specimens, and qualitative characters
were observed and registered only on adults. Scale terminology, measurements, and chromatic states follow Smith
(1946), nomenclature for neck folds follows Frost (1992). Coloration in life was recorded from pictures taken at the
time of capture. Sex was determined by the thickness of the base of the tail. Where numbers of paired scales are
provided they are given as left-right.
Morphological statistical tests. We implemented statistical tests for both morphometric and meristic charac-
ters, and compared frequencies of qualitative characters. ANOVA was implemented to evaluate the significance of
differences of variable means between new and described species, DGC comparisons (Di Rienzo, Guzmán and
Casanoves test; Di Rienzo et al. 2002) were performed, and assumptions of variance equality and normality were
checked with Levene and Shapiro-Wilks tests, respectively (Montgomery 1991). When ANOVA p values were sig-
nificant but assumptions of the statistical tests were not met (p values of Levene and Shapiro-Wilks tests < 0.05),
we performed nonparametric Kruskal-Wallis tests (Kruskal & Wallis 1952) using INFOSTAT® 2009 program (Di
Rienzo et al. 2010). Sexual dimorphism was tested for all variables with either Student’s T or Kruskal-Wallis tests
(when same assumptions previously described were not met).
We performed principal component analyses (PCA) separately on the morphometric and meristic data sets; the
first six principal components (PC) were saved to perform a discriminant-function analysis (DFA) using INFOS-
TAT® 2009. The objective of this analysis was to present a visualization of differences and similarities among spe-
cies. The discriminant functions computed by this method are linear combinations of the original variables that
maximize differences between given groups (Crochet et al. 2003). Given all of the analyses performed in the pres-
ent manuscript (see below, cross-validation with morphology, genetics, and distribution), an exhaustive analysis of
PCA was considered redundant (Crochet et al. 2003).
The overall differences between the new and the described species were statistically compared using the soft-
ware PAST v2.02 (Hammer et al. 2001) for only the meristic dataset, because the scales counts are independent of
body size. One-way non-parametric MANOVA with 100,000 permutations (NPMANOVA, also known as PER-
MANOVA) was implemented, this test provides a non-parametric approximation to test for significant differences
between two or more groups based on (in our case) a Gower distance measure (Anderson 2001).
Genetic characterization. Phylogenetics relationships among all the species plus eight candidate species of
the L. lineomaculatus section were recovered by Breitman et al. (2011), and our methods are briefly described here
as follows. Sequence data for seven nuclear genes (Cmos, ACM4tg, PRLR, LDA8F, LDA1D, LDA9C and
LDA9E) and two mitochondrial genes (12S and cyt-b) were amplified (Saint et al. 1998; Wiens et al. 1999; Gam-
ble et al. 2008; Townsend et al. 2008) edited and aligned using the program Sequencher v4.8. (™Gene Codes Cor-
poration Inc. 2007) and Clustal X (Higgins & Sharp 1988, 1989; Thompson et al. 1997). The combined nuclear
alignment was 4180 bp in length, while the nuclear plus the mitochondrial alignment was 5865 bp. All sequences
are deposited in GenBank (Accession Nos. JF272765 - JF273049). The best-fitting evolutionary model for each
gene was selected using JModelTest v0.1.1 (Guindon & Gascuel 2003; Posada 2008); recombination was tested
and excluded using RDP v3.44 (recombination detection program; Martin & Rybicki 2000; Heat et al. 2006).
Bayesian analyses were conducted for the concatenated matrix (nuclear and nuclear + mitochondrial) using
MrBayes v3.1.2 (Ronquist & Huelsenbeck 2003) with four chains, run for 50 million generations and sampled at
intervals of 1,000 generations. The equilibrium samples were used to generate a 50% majority-rule consensus tree
(after a 25% burn-in), and posterior probabilities (Pp) were considered significant when 0.95 (Huelsenbeck &
Ronquist 2001). We assume convergence because all parameters had effective sample sizes greater than 200 (using
Tracer v1.5.0; Rambaut & Drummond 2009).
Genetic structure within and among the species of the lineomaculatus group was characterized using a sub-set
of our recently published molecular data set (Breitman et al. 2011); specifically the mitochondrial cytochrome b
fragment from individuals collected in the type localities of L. lineomaculatus, L. hatcheri, L. kolengh, L. silvanae,
and the new species identified as L. sp. 1 and L. sp. 2 in Breitman et al. (2011). Genetic distances between species

BREITMAN ET AL.
4 · Zootaxa 3120 © 2011 Magnolia Press
(corrected pairwise differences = intergroup distance – intragroup distance) and analyses of molecular variance
(AMOVA), between described species and L. sp. 1 and L. sp. 2, were calculated using Arlequin v3.11 (Excoffier et
al. 2005). AMOVA analysis estimates genetic structure indices using the information on the allelic content of hap-
lotypes as well as their frequencies (Excoffier et al. 1992). The information on the differences in allelic content
between haplotypes is entered as a matrix of Euclidean squared distances. The significance of the covariance com-
ponents associated with the different possible levels of genetic structure (within species, within groups of species,
among species) is tested using non-parametric permutation procedures (Excoffier et al. 1992).
Results
Morphological statistical tests
Morphological tests showed significant differences between the new species and the other taxa of the lineomacula-
tus group. Univariate tests showed that several variables differed significantly between the new species and the
named taxa, which are described in the Diagnosis section of each new species. The means, standard deviations, and
ranks for meristic and qualitative characters are summarized in Tables 1 and 2, and Levene and Shapiro-Wilks or
Kruskal-Wallis p values are summarized in Tables 3 and 4. Sexual dimorphism was not detected in most of the
twenty seven studied variables. In the first new species, similarities between males and females were rejected in
only two variables (rostral height: females 0.84–1.06, X = 0.97 vs. males 1.18–1.18, X = 1.18; p = 0.037; dorsal
scales: females 47–67, X = 52.5 vs. males 50–50, X = 50; p = 0.033), while for the second new species similarities
between males and females were rejected in only three variables (distance between fore and hind limbs: females
28.4–29.5, X = 28.4 vs. males 19.4–26.9, X = 23.44; p = 0.043; foot length: females 13.82–14.57, X = 14.08 vs.
males 15.19–5.98, X = 15.54; p = 0.0036; auditory meatus length: females 1.11–1.33, X = 1.22 vs. males 1.44–2.1,
X = 1.68; p = 0.03).
Principal component analyses (PCA, see Appendixes III to VI) recovered the first six axes with almost 80% of
the accumulated explained variance. The first axis was mainly explained by differences in size (snout-vent, foot,
tibia-fibula, radius-ulna, hand, head, eye and auditory meatus lengths; head width, eye height, rostral nasal dis-
tance, rostral height and distance from rostral to the eye), variables that explained axis number two were differ-
ences in scale numbers (scales in contact with the interparietal, scales around midbody, dorsal scales, ventral scales
and infradigital lamellae of third finger and fourth toe); the third axis was mostly explained by the number of lori-
labial, supralabial and infralabial scales. Axes 4, 5, and 6 were explained by: rostral height and number of suprala-
bial scales; rostral-nasal distance, auditory meatus length and lorilabial scales; and scales in contact with
interparietal, respectively. Discriminant-function analysis was performed with the first six principal components
recovered with the PCA (Figure 2), and revealed that size is the principal difference that separates L. sp. 1 and L.
sp. 2 from L. silvanae. Number of scales (variables that describe differences in axis number two for the PCA,
explained above) is the main variable that differentiated L. sp. 1 and L. sp. 2 from L. hatcheri and L. kolengh. We
found that the CP4 (mainly described by supralabial scales) differentiated L. sp. 1 from L. sp. 2. and L. lineomacu-
latus. Liolaemus sp. 1 differed from L. lineomaculatus based on the variables that were described by axes one and
two (number of scales and size), although some overlap was evident.
The multivariate analyses (MPMANOVA) conducted on the meristic data set revealed statistically significant
differences among all the species of the lineomaculatus group (MPMANOVA, F = 13.51, p < 0.00001; Bonferroni
corrected pairwise comparisons between species were significant at = 0.05, with two exceptions at = 0.1: L. sp.
1 vs. L. lineomaculatus, p = 0.083 and L. sp. 1 vs. L. silvanae, p = 0.051).
Genetic characterization. The AMOVA revealed that 89.48% of the variation was explained by among-spe-
cies differences, whereas intrapopulation differences explained 10.52% of the variation, p < 0.0001. The Fst value
for each species was greater than 0.893, and p values were significant (p < 0.0001) in all cases, showing that differ-
ences between species were high and the intra-species variance (between population samples within a species) was
low. The uncorrected genetic distance between the new species and the others of the lineomaculatus group, calcu-
lated using cytochrome b, was higher than 4.6% (Table 5).
Figure 1 summarizes phylogenetic relationships inferred, using the complete and nuclear-only data sets, among
species and distinct lineages within the L. lineomaculatus section, and shows the position of the new species.

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TWO NEW SPECIES FROM L. LINEOMACULATUS SECTION
FIGURE 1. Relationships between Liolaemus morandae sp. nov. and L. avilae sp. nov. within the L. lineomaculatus section,
and selected species of the subgenus Eulaemus and Liolaemus sensu stricto. On the left, phylogenetic relationships recovered
with Bayesian inference using seven nuclear genes (Cmos, ACM4tg, PRLR, LDA8F, LDA1D, LDA9C and LDA9E, 4180 bp
total alignment length), bold branches represent well supported nodes (Pp > 0.95). On the right, the Bayesian tree (modified
from Breitman et al. 2011) represents a concatenated analyses including the seven nuclear genes plus two mitochondrial genes
(12S and cyt-b; 5865 bp) and summarizes information from MP and ML methods. Nodes with high support from three methods
(MP, jackknife and ML bootstrap > 0.70; Bayesian Pp > 0.95) are identified by bold branches; solid circles show nodes with
weak MP support and open circles nodes with weak MP and ML support.
FIGURE 2. Discriminant-function analysis summarizing principal components 1 – 6, for all six species of the L. lineomacula-
tus group. Black circles: L. morandae sp. nov.; white circles: L. avilae sp. nov.; black squares: L. lineomaculatus; white
squares: L. kolengh; gray triangles: L. hatcheri; black triangles: L. silvanae.

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Table 1: Values (in millimeters) of morphometric characters from species of the lineomaculatus group, samples sizes are shown in parenthesis, as well as mean ±
SD (min – max); abbreviations of characters are defined in Appendix II.
Var L. morandae sp. nov. (n=6) L. avilae sp. nov. (n=8) L. lineomaculatus (n=17) L. hatcheri (n=18) L. kolengh (n=25) L. silvanae (n=17)
SVL 55.67 ± 4.32 (50 - 61) 55.38 ± 3.66 (48 - 59) 54.94 ± 3.82 (46 - 60) 61.89 ± 4.27 (55 - 69) 57.48 ± 2.86 (52 - 62) 73 ± 3.64 (65 - 78)
TL 70.33 ± 9.02 (61 - 79) 68.88 ± 4.79 (64 - 77) 70.46 ± 6.69 (60 - 82) 63.3 ± 9.63 (49 - 78) 60.52 ± 5.97 (47 - 70) 76.13 ± 3.72 (72 - 84)
DFH 25.9 ± 4.33 (19.7 - 30.8) 25.3 ± 3.56 (19.4 - 29.5) 25.07 ± 2.73 (20.9 - 29.8) 28.95 ± 3.92 (19.5 - 35.3) 25.65 ± 2.49 (20.7 - 31.1) 32.38 ± 3.94 (25.1 - 39.7)
FOL 14.9 ± 1.16 (13.42 - 16.8) 14.91 ± 0.85 (13.82 - 15.98) 14.09 ± 0.87 (12.58 - 16.1) 15.02 ± 0.89 (13.5 - 16.3) 15.05 ± 0.86 (13.55 - 16.3) 18.46 ± 0.93 (16.5 - 20.1)
TFL 9.1 ± 1.26 (7.57 - 10.56) 10.16 ± 0.45 (9.72 - 11.08) 9.2 ± 1.47 (6.78 - 11.24) 8.68 ± 1.4 (5.06 - 10.47) 9.85 ± 0.71 (8.67 - 10.92) 13.37 ± 0.76 (11.77 - 14.48)
RUL 5.32 ± 0.36 (4.69 - 5.56) 5.76 ± 0.66 (4.89 - 6.81) 5.87 ± 0.54 (4.83 - 6.7) 6.44 ± 0.5 (5.67 - 7.33) 6.16 ± 0.58 (4.6 - 7.03) 7.71 ± 0.6 (6.83 - 8.82)
HAL 7.83 ± 0.79 (6.69 - 9.14) 8.02 ± 0.7 (7.27 - 9.52) 8.01 ± 0.58 (6.9 - 8.89) 9.15 ± 0.56 (8.5 - 10.57) 9.65 ± 0.63 (8.31 - 11.12) 12.23 ± 0.92 (10.65 - 13.76)
HH 7.23 ± 0.67 (6.52 - 8.21) 7.07 ± 0.51 (6.43 - 7.67) 6.99 ± 0.62 (5.77 - 8.09) 7.45 ± 0.61 (6.31 - 8.42) 6.82 ± 0.47 (6.04 - 7.5) 8.18 ± 0.58 (6.84 - 9.06)
HW 9.84 ± 0.61 (9.08 - 10.58) 9.59 ± 0.45 (9.02 - 10.19) 9.56 ± 0.71 (8.03 - 10.55) 11.22 ± 0.94 (9.7 - 13.05) 10.73 ± 0.85 (8.95 - 11.71) 13.14 ± 1.05 (11.31 - 14.71)
HL 11.96 ± 0.72 (10.96 - 13.06) 11.78 ± 0.83 (10.94 - 12.9) 11.76 ± 0.63 (10.43 - 12.71) 12.97 ± 0.92 (11.43 - 14.65) 12.04 ± 0.77 (10.58 - 13.17) 14.42 ± 1.04 (12.47 - 15.9)
EH 1.99 ± 0.23 (1.7 - 2.27) 2.2 ± 0.26 (2 - 2.62) 1.84 ± 0.22 (1.5 - 2.21) 2.05 ± 0.19 (1.72 - 2.55) 2.07 ± 0.17 (1.77 - 2.37) 2.46 ± 0.24 (2.09 - 2.9)
EL 3.15 ± 0.19 (2.91 - 3.42) 3.07 ± 0.21 (2.63 - 3.32) 2.83 ± 0.26 (2.35 - 3.3) 3.29 ± 0.3 (2.84 - 4.01) 3.37 ± 0.23 (2.85 - 3.81) 3.86 ± 0.19 (3.59 - 4.18)
RND 2.15 ± 0.14 (2.02 - 2.42) 1.95 ± 0.2 (1.69 - 2.25) 2.05 ± 0.2 (1.72 - 2.5) 2.27 ± 0.13 (1.97 - 2.48) 2.15 ± 0.16 (1.83 - 2.41) 2.38 ± 0.19 (2.08 - 2.73)
RH 1.04 ± 0.13 (0.84 - 1.18) 0.85 ± 0.2 (0.55 - 1.09) 0.93 ± 0.11 (0.76 - 1.11) 0.99 ± 0.11 (0.79 - 1.19) 1.03 ± 0.09 (0.87 - 1.2) 1.19 ± 0.1 (1.01 - 1.35)
RL 2.64 ± 0.15 (2.4 - 2.78) 2.51 ± 0.13 (2.31 - 2.69) 2.37 ± 0.18 (2.08 - 2.79) 2.7 ± 0.25 (2.14 - 3.02) 2.77 ± 0.18 (2.46 - 3.12) 3.13 ± 0.21 (2.61 - 3.47)
DRE 5.08 ± 0.26 (4.78 - 5.45) 4.9 ± 0.26 (4.53 - 5.31) 4.95 ± 0.31 (4.43 - 5.52) 5.59 ± 0.4 (4.78 - 6.41) 5.36 ± 0.35 (4.56 - 5.89) 6.18 ± 0.4 (5.47 - 6.88)
AH 1.95 ± 0.32 (1.6 - 2.44) 1.95 ± 0.3 (1.6 - 2.56) 1.83 ± 0.21 (1.5 - 2.23) 2.15 ± 0.22 (1.85 - 2.61) 2.22 ± 0.3 (1.63 - 2.95) 2.54 ± 0.26 (1.95 - 2.95)
AL 1.27 ± 0.4 (0.86 - 1.81) 1.51 ± 0.31 (1.11 - 2.1) 1.15 ± 0.14 (0.87 - 1.39) 1.77 ± 0.19 (1.38 - 2.16) 1.67 ± 0.24 (1.22 - 2.24) 2.04 ± 0.31 (1.49 - 2.57)

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All of these analyses reveal significant meristic, morphometric, nuclear and mitochondrial DNA differences
among all six species in this group, the two new species are formally named and described below.
TABLE 2. Values of meristic variables from species of the lineomaculatus group, sample sizes are shown in parentheses, as
well as mean ± SD (min – max); abbreviations for characters are defined in Appendix II.
continued.
Liolaemus morandae sp. nov.
(Figure 3–6)
2001, Liolaemus lineomaculatus, Ibargüengoytía, N., Casalins, L., Schulte II, J.A., Amico, G.A. and Sympson, L., Herpetolog-
ical Review, 32, 120.
Holotype. MLP.S 2626 (Figure 3), an adult male from Provincial Road 37, 22.8 km SW junction National Road 3,
Escalante department, Chubut province, Argentina (45°41’10,6” S; 67°53’49,9” W, 693 m) (Figures 4, 5); L.J.
Avila, M. Kozykariski, M.F. Breitman and R. Martinez collectors, 12th March 2010.
Paratypes. LJAMM-CNP 13020, adult female and LJAMM-CNP 13021, juvenile; same locality as holotype.
LJAM-CNP 9677-9679, adult females and LJAMM-CNP 9680, juvenile; from Holdich station, Escalante depart-
ment, Chubut province, Argentina (45º58’00.1” S; 68º11’58.8” W, 761 m); L.J. Avila, C.H.F. Pérez, M.F. Breitman
and N. Feltrin collectors, 2nd February 2008. LJAMM-CNP 10201, adult male and LJAM-CNP 10202, juvenile;
from Provincial Road 37, 2.5 km W junction National Road 3, Escalante department, Chubut province, Argentina
(45º37’43.4” S; 67º41’03.6” W, 637 m); L.J. Avila, C.H.F. Pérez, M.F. Breitman and N. Feltrin collectors, 2 nd Feb-
ruary 2008.
Diagnosis. Liolaemus morandae sp. nov. is a member of the L. lineomaculatus section, included in the lineo-
maculatus group, and is characterized by the absence of precloacal pores in both sexes, and presence of dorsal trifid
scales (Etheridge 1995). Molecular evidence includes this species in the lineomaculatus group, closely related to
the clade (L. sp. 2 + L. lineomaculatus) Figure 1. All the following differences are summarized in Tables 1 to 4 and
in Figure 6.
Var L. morandae sp. nov. (n=6) L. avilae sp. nov. (n=8) L. lineomaculatus (n=17)
SCI 6.33 ± 0.52 (6–7) 8.13 ± 1.25 (7–10) 7.18 ± 1.07 (6–10)
LS 4 ± 0.89 (3–5) 4.38 ± 0.52 (4–5) 4.41 ± 0.71 (4–6)
SS 7.17 ± 1.94 (5–10) 5.25 ± 0.46 (5–6) 6.71 ± 0.92 (5–8)
IS 5.33 ± 0.52 (5–6) 4.63 ± 0.74 (4–6) 4.76 ± 0.56 (4–6)
MS 59.83 ± 2.71 (55–63) 55.5 ± 2.67 (53–59) 60 ± 3.39 (54–65)
DS 51.67 ± 3.5 (47–57) 53.75 ± 2.12 (51–58) 58.41 ± 3.84 (52–68)
VS 81.33 ± 2.94 (79–85) 83 ± 6.32 (72–91) 85.29 ± 4.48 (80–95)
IL3 15.33 ± 1.21 (13–16) 16.25 ± 0.89 (15–18) 16.18 ± 1.29 (14–19)
IL4 21.17 ± 1.72 (18–23) 21.29 ± 0.95 (20–23) 21.41 ± 1.58 (19–25)
Var L. hatcheri (n=20) L. kolengh (n=31) L. silvanae (n=17)
SCI 6.85 ± 1.39 (5–10) 6.26 ± 0.89 (4–8) 6.94 ± 1.03 (5–8)
LS 4.5 ± 0.69 (4–6) 4.29 ± 0.59 (3–5) 4.18 ± 0.64 (3–5)
SS 6.85 ± 0.75 (6–8) 6.65 ± 0.8 (5–8) 6.18 ± 0.73 (5–7)
IS 5.25 ± 0.79 (4–7) 5.29 ± 0.53 (5–7) 4.88 ± 0.7 (4–6)
MS 48.65 ± 3.33 (43–57) 50.55 ± 2.55 (44–55) 59.65 ± 2.89 (55–65)
DS 48.75 ± 3.49 (43–55) 49.35 ± 2.78 (44–54) 56 ± 3.12 (52–64)
VS 67.8 ± 3.37 (62–74) 68.58 ± 3.89 (61–76) 74.59 ± 2.74 (70–80)
IL3 14.85 ± 1.39 (13–18) 14.6 ± 0.93 (13–16) 16.24 ± 1.15 (14–18)
IL4 19.45 ± 1.28 (18–22) 18.74 ± 1.46 (16–22) 20.24 ± 1.39 (18–23)

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FIGURE 3. Upper, dorsal and ventral views of male holotype in life of Liolaemus morandae sp. nov. MLP.S 2626; lower, dor-
sal and ventral views of female (LJAMM-CNP 13020).
Relative to L. lineomaculatus, L. morandae sp. nov. has fewer dorsal scales (47–57, X = 51.67 vs. 52–68, X =
58.41; p < 0.0001), fewer third finger lamellae (13–16, X = 15.33 vs. 14–19, X = 16,18; p < 0.0001), shorter fourth
toe length (13.4–16.8, X = 14.9 vs. 12.6–16.1, X = 14.09; p < 0.0001) and a larger eye (eye height: 1.7–2.27, X =
1.99 vs. 1.5–2.21, X = 1.84, eye length: 2.91–3.42, X = 3.15 vs. 2.35–3.3, X = 2.83; p < 0.0001 in both cases). Lio-
laemus morandae sp. nov. has more dorsal blotches than L. lineomaculatus, and the blotches in L. morandae sp.
nov. have (in 50% of the cases) a yellow-orange coloration while L. lineomaculatus blotches have dark gray color.

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In general, dorsal scales of L. morandae sp. nov. are more mucronated than those of L. lineomaculatus, and dorsal
leg scales of L. morandae sp. nov. are less carinated with respect to L. lineomaculatus.
Compared to L. hatcheri, L. morandae sp. nov. has higher number of midbody scales (55–63, X = 59.83 vs.
43–57, X = 48.65 p < 0.0001), larger number of scales from occiput to rump (47–57, X = 51.67 vs. 43–55, X =
48.75; p < 0.0001), more ventral scales (79–85, X = 81.33 vs. 62–74, X = 67.8; p < 0.0001), more fourth toe lamel-
lae (18–23, X = 21.17 vs. 18–22, X = 19.45; p < 0.0001), smaller body size (snout-vent length: 50–61, X = 55.67
vs. 55–69, X = 61.89, axilla-groin distance: 19.7–30.8, X = 25.9 vs. 19.5–35.5, X = 28.95; p < 0.0001 in both
cases), shorter forelimbs (elbow-wrist length: 4.69–5.56, X = 5.32 vs. 5.67–7.33, X = 6.44, third finger length:
6.69–9.14, X = 7.83 vs. 8.5–10.57, X = 9.15; p < 0.0001 in both cases), smaller head (head width: 9.08–10.58, X =
9.84 vs. 9.7–13.05, X = 11.22, head length: 10.96–13.06, X = 11.96 vs. 11.43–14.65, X = 12.97; p < 0.0001 in both
cases), and smaller auditory meatus (auditory meatus height: 1.6–2.44, X = 1.95 vs. 1.85–2.61, X = 2.15; auditory
meatus length: 0.86–1.81, X = 1.27 vs. 1.38–2.16, X = 1.77; p < 0.0001 in both cases). Liolaemus morandae sp.
nov. has more dorsal blotches and better defined vertebral and paravertebral lines than L. hatcheri; ventral mela-
nism is more pronounced in L. hatcheri, while L. morandae sp. nov. has fewer melanophores. Dorsal scales of L.
morandae sp. nov. are less “bristly” than those of L. hatcheri. Dorsal and ventral limb scales are non-mucronate
and carinate in L. morandae sp. nov. while in L. hatcheri they are mucronate and non-carinate. Belly scales are
round-shaped in L. morandae sp. nov. and rhomboidal in L. hatcheri.
Liolaemus morandae sp. nov. differs from L. kolengh in the following traits: larger number of midbody scales
(55–63, X = 59.83 vs. 44–55, X = 50.55; p < 0.0001), larger number of scales from occiput to rump (47–57, X =
51.67 vs. 44–54, X = 49.35; p < 0.0001), more ventral scales (79–85, X = 81.33 vs. 61–76, X = 68.58; p < 0.0001),
larger number of fourth toe lamellae (18–23, X = 21.17 vs. 16–22, X = 18.74; p < 0.0001), shorter forelimbs
(elbow-wrist length: 4.69–5.56, X = 5.32 vs. 4.6–7.03, X = 6.16, third finger length: 6.69–9.14, X = 7.83 vs. 8.31–
11.12, X = 9.65; p < 0.0001 in both cases), smaller auditory meatus (auditory meatus height: 1.6–2.44, X = 1.95 vs.
1.63–2.95, X = 2.22, auditory meatus length: 0.86–1.81, X = 1.27 vs. 1.22–2.24, X = 1.67; p < 0.0001 in both
cases) and longer tail (61–79, X = 70.33 vs. 47–70, X = 60.52; p = 0.0019). Liolaemus morandae sp. nov. has more
dorsal blotches and better defined vertebral and paravertebral lines than L. kolengh; ventral melanism is stronger in
L. kolengh than in L. morandae sp. nov.; the general background coloration of L. kolengh is dark gray or black,
though in almost 50% of our samples the dorsal pattern cannot be recognized. In general, the dorsal scales of L.
morandae sp. nov. are less “bristly” than those of L. kolengh. Dorsal and ventral limbs scales are non-mucronate
and non-bristly in L. morandae sp. nov., but mucronate and “bristly” in L. kolengh.
Liolaemus morandae sp. nov. differs from L. silvanae in several characters, including: a smaller number of
scales from occiput to rump (47–57, X = 51.67 vs. 52–64, X = 56; p < 0.0001), higher number of ventral scales
(79–85, X = 81.33 vs. 70–85, X = 74,59; p < 0.0001), fewer third finger lamellae (13–16, X = 15.33 vs. 14–18, X =
16.24; p < 0.0001), smaller body size (snout-vent length: 50–61, X = 55.67 vs. 65–78, X = 73, axilla-groin dis-
tance: 19.7–30.8, X = 25.9 vs. 25.1–39.7, X = 32.38; p< 0.0001 in both cases), shorter forelimbs (elbow-wrist
length: 4.69–5.56, X = 5.32 vs. 6.83–8.82, X = 7.71, third finger length: 6.69–9.14, X = 7.83 vs. 10.65–13.76 X =
12.23; p < 0.0001 in both cases), smaller head (head height: 6.52–8.21, X = 7.23 vs. 6.84–9.06, X = 8.18, head
width: 9.08–10.58, X = 9.84 vs. 11.31–14.71, X = 13.14, head length: 10.96–13.06, X = 11.96 vs. 12.47–15.9, X =
14.42; p < 0.0001 in all cases), smaller eye (eye height: 1.7–2.27, X = 1.99 vs. 2.09–2.9, X = 2.46, eye length:
2.91–3.42, X = 3.15 vs. 3.59–4.18, X = 3.86; p < 0.0001 in both cases), and smaller auditory meatus (auditory
meatus height: 1.6–2.44, X = 1.95 vs. 1.95–2.95, X = 2.54; auditory meatus length: 0.86–1.81, X = 1.27 vs. 1.49–
2.57, X = 2.04; p < 0.0001 in both cases). The general background coloration of L. silvanae is black; there is no
recognizable dorsal pattern, and the venter is also strongly melanistic, whereas the venter is weakly melanistic in L.
morandae sp. nov. Liolaemus morandae sp. nov. possesses well-defined vertebral and paravertebral lines that are
not present in L. silvanae. In general, L. silvanae possesses a “bristly” appearance that is not present in L. moran-
dae sp. nov. Limb scales are less mucronated in L. morandae sp. nov. than in L. silvanae.
Description of holotype. Adult male. Snout-vent length 56.0 mm. Tail length (complete, not regenerated) 79.0
mm. Axilla-groin distance 22.8 mm. Auditory meatus-eye distance 4.8 mm. External auditory meatus conspicuous,
higher (2.1 mm) than wide (1.2 mm). Head length 13.0 mm (from anterior border of tympanum to tip of snout),
10.5 mm wide (at anterior border of tympanum), 7.8 mm high (at anterior border of tympanum). Snout length 4.0
mm (orbit-tip of snout distance). Interorbital distance 4.2 mm. Eye-nostril distance 3.1 mm. Forelimb length 16.7
mm. Tibial length 10.3 mm. Foot length 16.8 mm (ankle to tip of claw on fourth toe).

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FIGURE 4. Type locality of Liolaemus morandae sp. nov., Patagonic phytogeographic province, del Golfo de San Jorge Dis-
trict (45°41’10,6” S; 67°53’49,9” W, 693 m).

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TWO NEW SPECIES FROM L. LINEOMACULATUS SECTION
FIGURE 5. Distribution map for species of the lineomaculatus group, with geographical localities sampled in this study super-
imposed on regional elevation (shading). Black circles: L. morandae sp. nov.; white circle: L. avilae sp. nov.; black squares: L.
lineomaculatus; white square: L. kolengh; white triangles: L. hatcheri; black triangle: L. silvanae. Arrows indicate type locali-
ties. CH: Chubut province; SC: Santa Cruz province.
Dorsal head scales bulged, smooth, 15 between occiput, at the level of anterior border of tympanum, to rostral,
pitted with numerous scale organs in the anterior region, and reducing to a single organ, or lack, in the posterior
half of the head. Rostral scale wider (2.4 mm) than high (0.9 mm). Two postrostrals, together with anterior lorila-
bial, separate nasal scales from rostral, surrounded by six scales. Nasal scales longer than wide, irregularly hexago-
nal; nostril one-half length of nasal, posterior in position. Scales surrounding nasals 7 on the left side and 8 on the
right side. Four internasals. Frontonasals six, irregular in size and position. Prefrontals 6, a small rhomboidal scale
in the center (0.9 mm). Three dorso-lateral larger scales, one for the fragmentation of the right scale, and a pair lat-
eral medium-sized scales (1.3 mm), approximately quadrangular. Two frontal scales. Frontoparietals in two rows,
one anterior and one posterior scale, unfragmented. Interparietal pentagonal, surrounded by six scales; four smaller
and irregular in front and sides, two larger in back. Parietal eye evident. Parietals slightly bulged, irregularly
shaped, equal in size to interparietal (1.5 vs.1.5 mm). Circumorbitals:13–10. Transversally expanded supraoculars
4–3. Smaller lateral supraoculars: 16–17. One canthal higher than wide, separated from nasal by one postnasal.
Loreal scales bulged, two on the left side (by fusion of the posterior loreal scale with preocular scale) and three on
the rigth side. Lorilabials longer than wide (8–6), approximately equal to labials. Superciliaries 7–7, flattened and
elongated, anterior five broadly overlapping dorsally. Orbit with 15–17 upper and 12–13 lower ciliaries on each
side. Orbit diameter 4.5 x 1.9 mm. Preocular small, unfragmented, longer than wide. Subocular scale elongated,
approximately nine times longer than wide (3.7 x 0.6 mm). A well marked longitudinal ridge along upper margin of
preocular and subocular scales. Postocular small, slightly bulged, quarter superimposed to subocular, with a
marked longitudinal ridge. Palpebral scales small granular and bulged. Supralabials 9–10, convex. Temporals
smooth, convex, juxtaposed with one scale organ in the tip. Anterior auriculars smaller than adjacent posterior tem-
porals, slightly projecting outward (3–2). Posterior auriculars small and granular. External auditory meatus con-
spicuous, higher (2.1 mm) than wide (1.2 mm). Lateral scales of neck granular with inflated skin. Mental scale
wider (2.9 mm) than high (1.4 mm), in contact with four scales. Mental posterior followed by two postmentals, and

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two rows of three chinshields on each side. Six infralabials on each side, first on each side quadrangular two times
wider than supralabials, all others elongated, slightly smaller than supralabials. Gular scales smooth, flat, imbri-
cate, with rounded posterior margins, with melanophores. Scales of throat between chinshields slightly juxtaposed,
becoming slightly imbricate toward auditory meatus. Thirty gulars between tympanum openings. Infralabials sepa-
rated from chinshields by one to two rows of scales.
Antehumeral, longitudinal and postauricular neck folds well developed; gular, rictal, dorsolateral and oblique
not conspicuous.
Scales of dorsal neck region rhomboidal, imbricate and strongly keeled. Fifty dorsal scales between occiput
and anterior surface of thighs. Dorsal body scales rhomboidal, imbricate, strongly keeled, mucronate, very few tri-
fid scales. Dorsal scales grade laterally into slightly smaller, slightly keeled scales at midbody. Scales immediately
anterior and posterior to forelimb and hindlimb insertion small, smooth, granular, and non-overlapping. Body lat-
eral scales grading smaller to larger at midbody. Ventral body scales rhomboidal, smooth, flat, imbricate, larger
than dorsal scales. Sixty-one midbody scales; scales between mental and precloacal pores 85. Scales of cloacal
region about equal in size to ventral body scales; without precloacal pores.
Anterior suprabrachials rhomboidal, imbricate, smooth, slightly larger in size to dorsal body scales. Postabra-
chials smaller, smooth, becoming granular near axilla. Supra-antebrachials similar to suprabrachial. Infra-antebra-
chials rhomboidals, imbricate, smooth, toward the hand slightly mucronate. Supracarpals imbricated, rhomboidal,
smooth. Infracarpals strongly imbricate, rhomboidal, slightly keeled, 3-mucronate. Subdigital lamellae with 2–5
keels, each terminating in a short mucron, 2–5 mucronate, numbering: I: 8, II: 12, III: 16, IV: 17, V: 11. Claws
robust, curved and sharp, opaque brown.
Suprafemorals as large as dorsal body scales, rhomboidal, imbricated, smooth near the body, toward to the
knee slightly keeled. Postfemorals small, granular shape. Supratibials rhomboidal, imbricated, keeled, some mucr-
onate, smaller and smooth toward to the foot. Infrafemoral scales small, granular and smooth. Supratarsals rhom-
boidal, imbricated and smooth. Infratarsals small, rhomboidal, imbricate, smooth, mucronate, slightly keeled near
the digit. Subdigital scales 1–3 keeled, 1–5 mucronate, numbering: I: 9, II: 14, III: 20, IV: 21, V: 14. Claws robust,
curved and sharp, opaque brown. Tail complete, non-regenerated. Dorsal and lateral caudal scales, rhomboidal in
the first half of the tail, becoming quadrangular toward the tip, strongly keeled. Ventrals subtriangular and smooth,
toward posterior half moderate keeled.
Color of holotype in life. Dark grey dorsal background (Figure 3) that becomes lighter on the lateral region,
between axilla and groin. Dorsal pattern presents twelve paravertebral, quadrangular, black blotches, extending
from the nuchal to the postcloacal region, those series fuse into a dark line that is present to the tip of the tail. Dor-
solateral region, in between the occipital and the tip of the tail, formed by black and white-bordered blotches. Lat-
eral region, between axilla and groin, is characterized by a black and white reticulate pattern. In between the series
of blotches, one white vertebral line and two light orange paravertebral longitudinal lines appear, one and one and a
half scale wide, respectively. Vertebral line extends from the nuchal region, while the paravertebral lines extend
from the temporal region; all of them continue to the tip of the tail. Dorsal region of the limbs are dark gray with a
black reticulation.
Dark gray head background. Lateral white line along the longitudinal folds is present from the top of the audi-
tory meatus to the antehumeral fold. White subocular scale with black longitudinal ridge. Dorsal surface with black
blotches, one longitudinal blotch is present in the anterior part of the head, crossing through internasals, postnasals
and prefrontals; two transverse blotches are crossing through postfrontals and first circumorbitals, and extend back
to the last circumorbitals, forming a cross with the longitudinal blotch. The interparietal scale is surrounded by
three small dark-brown blotches and the occipitals show a white longitudinal band of the same color.
White ventral color with a black reticulate pattern over the entire body, most strongly marked in the pectoral
region and throat, and extending to the adjacent malar and maxilla region. Ventral scales with melanophores. Ven-
tral area of chest, belly, cloacal region and limbs tinged with a light yellowish color. Cloacal and postcloacal region
with few black blotches. Gray tail.
Color of holotype in preservative. After one year in preservative, the dorsal coloration of the head, dorsum,
body flanks and tail becomes darker while maintaining the contrast, but the two yellowish paravertebral lines
turned gray. Ventral scales of throat, neck, chest, belly and forelimbs retain the same coloration as in life, and the
distinctive light ventral yellowish tinge of chest, belly, cloacal region and limbs turns gray (Figure 6).

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FIGURE 6. Morphological comparison between Liolaemus morandae sp. nov. (A), L. lineomaculatus (B) and L. avilae sp.
nov. (C). Notice differences in color pattern on the tails, size of blotches on the limbs and differences in vertebral and paraver-
tebral lines. Dorsal scales (shown on the right) are less mucronated in L. lineomaculatus. Dorsal scales of hindlimbs are less
carinated in L. morandae sp. nov. than in the other species, while they are more mucronated in L. avilae sp. nov.
Variation. Morphological and meristic character variation between males and females of Liolaemus morandae
sp. nov., are shown in Table 6. Females in life present basically the same dorsal and lateral pattern as present in
males, but females generally have a lighter brown background coloration; vertebral and paravertebral lines are
strongly defined, and in both sexes paravertebral lines vary from white to yellow. Ventrally, one male shows a light
orange color on the belly, while in other males and all females the ventral region ranged from white to gray due to
different degrees of melanism. All distinctive ventral light orange coloration changed to gray in preservative.
Etymology. The specific epithet of this species “morandae” refers to our colleague and the first author’s PhD
advisor Dr. Mariana Morando, to honor her after more than ten years of research and teaching on the Patagonian
herpetofauna.
Distribution. Liolaemus morandae sp. nov. is known only from the type locality and a few localities on the
Pampa del Castillo Plateau; and from three localities from southern Chubut and northern Santa Cruz provinces. The
holotype and some paratypes are from Provincial Road 37, 22.8 km SW junction National Road 3, Escalante
department, Chubut province, Argentina (Figures 4, 5). The other paratypes are from Provincial Road 37, 2.5 km
W junction National Road 3 in Holdich Station, Escalante department, Chubut province, Argentina (Figures 4, 5).
The distance from the type locality to the first locality is more than 17.8 km airline, and more than 38.8 km airline
to the second locality.
Natural history. Liolaemus morandae sp. nov. occurs in Patagónica province, Golfo de San Jorge District, in
environments characterized as grasslands and steppe highlands (Roig 1998), with vegetation dominated by the
shrubs Stipa spp., Senecio filaginoides, Mulinum spinosum, Nassauvia spp., among others. This species was found
in open substrates, sharing the habitat with Liolaemus kingii, L. bibronii, Diplolaemus bibronii and D. darwinii
(Figure 4). Based on the natural history of its close relative L. lineomaculatus, we hypothesize that L. morandae sp.
nov. is herbivorous and likely to also be viviparous, perhaps giving birth to between three and six neonates per
clutch (Cei 1986; Espinoza et al. 2004).

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TWO NEW SPECIES FROM L. LINEOMACULATUS SECTION
TABLE 5. Average pairwise genetic distances (expressed in percentage) calculated for cytocrhome b (659 bp). Intragroup dis-
tances are presented in the first column, while uncorrected and corrected distances (in parentheses) are present in the second
and third columns. These are comparisons among L. morandae sp. nov., L. avilae sp. nov. and all other members of the lineo-
maculatus clade.
TABLE 6. Variation between females and males of L. morandae sp. nov. and L. avilae sp. nov. Summarized as mean ± SD
(min – max) values for all meristic and morphometric (in millimeters) variables.
Intragroup distance L. morandae sp. nov. L. avilae sp. nov.
L. hatcheri 1.26 12.4 (11.3) 11.5 (10.6)
L. silvanae 1.36 9.88 (8.78) 9.29 (8.3)
L. kolengh 0.12 9.61 (9.13) 8.99 (8.65)
L. lineomaculatus 1.35 7.64 (6.54) 5.86 (4.92)
L. avilae sp. nov. 0.83 4.61 (3.91)
L. morandae sp. nov. 0.55
L. morandae sp. nov. L. avilae sp. nov.
VAR Females Males Females Males
SCI 6.25 ± 0.5 (6–7) 6.5 ± 0.71 (6–7) 9 ± 1.73 (7–10) 7.6 ± 0.55 (7–8)
LS 4 ± 0.82 (3–5) 4 ± 1.41 (3–5) 4.33 ± 0.58 (4–5) 4.4 ± 0.55 (4–5)
SS 6.75 ± 1.71 (5–9) 8 ± 2.83 (6–10) 5.33 ± 0.58 (5–6) 5.2 ± 0.45 (5–6)
IS 5 ± 0 (5–5) 6 ± 0 (6–6) 4 ± 0 (4–4) 5 ± 0.71 (4–6)
MS 60.75 ± 1.71 (59–63) 58 ± 4.24 (55–61) 55.33 ± 3.21 (53–59) 55.6 ± 2.7 (53–59)
DS 52.5 ± 4.2 (47–57) 50 ± 0 (50–50) 54.33 ± 3.51 (51–58) 53.4 ± 1.14 (52–55)
VS 81 ± 2.83 (79–85) 82 ± 4.24 (79–85) 86.33 ± 4.04 (84–91) 81 ± 6.96 (72–90)
IL3 15.25 ± 1.5 (13–16) 15.5 ± 0.71 (15–16) 16 ± 0 (16–16) 16.4 ± 1.14 (15–18)
IL4 21.25 ± 2.22 (18–23) 21 ± 0 (21–21) 20.67 ± 0.58 (20–21) 21.75 ± 0.96 (21–23)
SVL 57 ± 4.24 (52–61) 53 ± 4.24 (50–56) 56.67 ± 2.08 (55–59) 54.6 ± 4.39 (48–59)
DFH 23.73 ± 3.49 (19.7–28.1) 30.25 ± 0.78 (29.7–30.8) 28.4 ± 1.05 (27.4–29.5) 23.44 ± 3.18 (19.4–26.9)
FOL 15.19 ± 1.17 (14.09–16.8) 14.31 ± 1.26 (13.42–15.2) 14.08 ± 0.43 (13.82–14.57) 15.54 ± 0.33 (15.19–15.98)
TFL 9.23 ± 1.47 (7.57–10.56) 8.84 ± 1.1 (8.06–9.62) 9.86 ± 0.18 (9.72–10.06) 10.34 ± 0.48 (9.79–11.08)
RUL 5.33 ± 0.43 (4.69–5.56) 5.29 ± 0.32 (5.06–5.51) 5.34 ± 0.37 (4.91–5.58) 6 ± 0.7 (4.89–6.81)
HAL 7.78 ± 1.02 (6.69–9.14) 7.94 ± 0.18 (7.81–8.06) 7.91 ± 0.34 (7.67–8.3) 8.09 ± 0.89 (7.27–9.52)
HH 7.09 ± 0.58 (6.52–7.89) 7.51 ± 1 (6.8–8.21) 6.66 ± 0.32 (6.43–7.03) 7.31 ± 0.45 (6.54–7.67)
HW 9.64 ± 0.62 (9.08–10.52) 10.26 ± 0.45 (9.94–10.58) 9.25 ± 0.07 (9.2–9.33) 9.79 ± 0.47 (9.02–10.19)
HL 11.88 ± 0.87 (10.96–13.06) 12.12 ± 0.52 (11.75–12.49) 11.07 ± 0.12 (10.94–11.16) 12.2 ± 0.78 (11.01–12.9)
EH 1.93 ± 0.26 (1.7–2.27) 2.1 ± 0.19 (1.96–2.23) 2.29 ± 0.28 (2.07–2.6) 2.14 ± 0.27 (2–2.62)
EL 3.14 ± 0.21 (2.91–3.42) 3.17 ± 0.2 (3.03–3.31) 3.13 ± 0.13 (3.02–3.27) 3.03 ± 0.25 (2.63–3.32)
RND 2.19 ± 0.17 (2.02–2.42) 2.09 ± 0.07 (2.04–2.14) 1.98 ± 0.26 (1.74–2.25) 1.93 ± 0.19 (1.69–2.16)
RH 0.97 ± 0.09 (0.84–1.06) 1.18 ± 0 (1.18–1.18) 0.86 ± 0.18 (0.73–1.06) 0.84 ± 0.24 (0.55–1.09)
RL 2.61 ± 0.18 (2.4–2.76) 2.69 ± 0.13 (2.6–2.78) 2.51 ± 0.18 (2.31–2.67) 2.51 ± 0.12 (2.36–2.69)
DRE 5.06 ± 0.31 (4.78–5.45) 5.13 ± 0.2 (4.99–5.27) 4.73 ± 0.18 (4.53–4.89) 5.01 ± 0.26 (4.64–5.31)
AH 2.03 ± 0.39 (1.6–2.44) 1.8 ± 0.08 (1.74–1.86) 1.8 ± 0.15 (1.68–1.96) 2.05 ± 0.34 (1.6–2.56)
AL 1.19 ± 0.43 (0.86–1.81) 1.42 ± 0.39 (1.14–1.69) 1.22 ± 0.11 (1.11–1.33) 1.68 ± 0.26 (1.44–2.1)
TL 66 ± 7.07 (61–71) 79 ± 0 (79–79) 67.33 ± 4.93 (64–73) 69.8 ± 5.02 (65–77)

BREITMAN ET AL.
16 · Zootaxa 3120 © 2011 Magnolia Press
Liolaemus avilae sp. nov.
(Figures 6, 7)
1971, Liolaemus lineomaculatus, Donoso-Barros, R. and Cei, J.M., Journal of Herpetology, 5, 89–95.
1975, Liolaemus lineomaculatus, Cei, J.M., Herpetologica, 31, 109–116.
1982, Liolaemus lineomaculatus, Cei, J.M. and Scolaro, J.A., Journal of Herpetology, 16, 354–363.
1992, Liolaemus lineomaculatus, Scolaro, J.A., Acta zoologica lilloana, 41, 287–293.
Holotype. MLP.S 2627 (Figure 7), an adult male from Lago Buenos Aires plateau, 18.7 SW Puesto Lebrun, Lago
Buenos Aires department, Santa Cruz province, Argentina (47º05’29.0” S, 71º01’12.9” W, 1154 m) (Figures 5, 8),
L.J. Avila, C.H.F. Pérez, M.F. Breitman and N. Feltrin collectors, 9th January 2008.
Paratypes. LJAMM-CNP 9250, 9253, 9274, adult males, LJAMM-CNP 9276-9277, 9399, adult females and
LJAMM-CNP 9251, 9252, juveniles; from same locality as holotype, L.J. Avila, C.H.F. Pérez, M.F. Breitman and
N. Feltrin collectors, 9 January 2008. LJAMM-CNP 9243, an adult male from Puesto Lebrun, 27.3 km W Estancia
La Vizcaina, Lago Buenos Aires Plateau, Lago Buenos Aires department, Santa Cruz province, Argentina
(46º57’51.8” S, 71º06’27.2” W, 1353 m), L.J. Avila, C.H.F. Pérez, M.F. Breitman and N. Feltrin collectors, 8th Jan-
uary 2008.
Diagnosis. Liolaemus avilae sp. nov., a member of the L. lineomaculatus section and specifically the lineo-
maculatus group, has dorsal trifid scales but lacks of precloacal pores in either sex (Etheridge 1995); molecular evi-
dence places this species in the lineomaculatus group, as the sister species to L. lineomaculatus (Breitman et al.
2011; see Tables 1 to 4 and Figure 6).
Relative to L. morandae sp. nov., L. avilae sp. nov. has more scales in contact with the interparietal scale (7–
10, X = 8,13 vs. 6–7, X = 6.33; p = 0.0027), fewer supralabial scales (5–6, X = 5.25 vs. 5–10, X = 7.17; p =
0.0008), fewer midbody scales (53–59, X = 55.5 vs. 55–63, X = 59.83; p < 0.0001), more third finger lamellae (15–
18, X = 16.25 vs. 13–16, X = 15.33; p < 0.0001), a longer elbow-wrist distance (4.89–6.81, X = 5.76 vs. 4.69–5.56,
X = 5.32; p < 0.0001), and a shorter rostral-nasal distance (1.69–2.25, X = 1.95 vs. 2.02–2.42, X = 2.15; p <
0.0001). Liolaemus avilae sp. nov. has fewer dorsal blotches than L. morandae sp. nov., and the blotches in L.
morandae sp. nov. have (50% of cases) brownish coloration, while L. avilae sp. nov. blotches have dark gray color.
Ventral hand and foot surfaces of L. morandae sp. nov. present mucronated and “bristly” scales that are less obvi-
ous in L. avilae sp. nov.
Liolaemus avilae sp. nov. differs from L. lineomaculatus in the following traits: more scales in contact with the
interparietal (7–10, X = 8,13 vs. 6–10, X = 7.18; p = 0.0027), fewer supralabial scales (5–6, X = 5.25 vs. 5–8, X =
6.71; p = 0.0008), fewer midbody scales (53–59, X = 55.5 vs. 54–65, X = 60; p < 0.0001), fewer scales from
occiput to rump (51–58, X = 53.75 vs. 52–68, X = 58.41; p < 0.0001), fewer ventral scales (72–91, X = 83 vs. 80–
95, X = 85.29; p < 0.0001), larger eyes (eye height: 2–2.62, X = 2.2 vs. 1.5–2.21, X = 1.84, eye length: 2.63–3.32,
X = 3.07 vs. 2.35–3.3, X = 2.83; p < 0.0001 in both cases), and longer auditory meatus lengths (1.11–2.1, X = 1.51
vs. 0.87–1.39, X = 1.15; p < 0.0001). Liolaemus avilae sp. nov. has fewer dorsal blotches than L. lineomaculatus.
The vertebral line is less well defined in L. avilae sp. nov. than in L. lineomaculatus. In general, dorsal scales of L.
avilae sp. nov. are more mucronated than in L. lineomaculatus. Liolaemus avilae sp. nov. possesses scales in the
dorsal surface of the forelimbs that are less carinated than those of L. lineomaculatus.
Compared with L. hatcheri, L. avilae sp. nov. has more scales in contact with the interparietal (7–10, X = 8,13
vs. 5–10, X = 6.85; p = 0.0027), fewer supralabial scales (5–6, X = 5.25 vs. 6–8, X = 6.85; p = 0.0008), more mid-
body scales (53–59, X = 55.5 vs. 43–57, X = 48.65; p < 0.0001), more scales from occiput to rump (51–58, X =
53.75 vs. 43–55, X = 48.75; p < 0.0001), more ventral scales (72–91, X = 83 vs. 62–74, X = 67.8; p < 0.0001),
more third finger lamellae (15–18, X = 16.25 vs. 13–18, X = 14.85; p < 0.0001), more fourth toe lamellae (20–23,
X = 21.29 vs. 18–22, X = 19.45; p < 0.0001), smaller body size (snout-vent length: 48–59, X = 55.38 vs. 55–69, X
= 61.89, axilla-groin distance: 19.4–29.5, X = 25.5 vs. 19.5–35.5, X = 28.95; p < 0.0001 in both cases), shorter
forelimbs (elbow-wrist length: 4.89–6.81, X = 5.76 vs. 5.67–7.33, X = 6.44, third finger length: 7.27–9.52, X =
8.02 vs. 8.5–10.57, X = 9.15; p < 0.0001 in both cases), and smaller head width (9.02–10.19, X = 9.59 vs. 9.7–
13.05, X = 11.22; p < 0.0001). Liolaemus avilae sp. nov. possesses fewer dorsal blotches and a more defined verte-
bral line relative to L. hatcheri. Ventral melanism is more pronounced in L. hatcheri than in L. avilae sp. nov.
(fewer melanophores). Dorsal scales of L. avilae sp. nov. are less “bristly” than those of L. hatcheri. In the new
species, limb surface scales (fore-hind and dorsal-ventral) are non-mucronate and carinated (in L. hatcheri the

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TWO NEW SPECIES FROM L. LINEOMACULATUS SECTION
majority are mucronated and non-carinate). Ventral scales are round-shaped in L. avilae sp. nov. and rhomboidal in
L. hatcheri.
Compared with L. kolengh, L. avilae sp. nov. has fewer scales in contact with the interparietal (7–10, X = 8,13
vs. 4–8, X = 6.26; p = 0.0027), fewer supralabial scales (5–6, X = 5.25 vs. 5–8, X = 6.65; p = 0.0008), more mid-
body scales (53–59, X = 55.5 vs. 44–55, X = 50.55; p < 0.0001), more scales from occiput to rump (51–58, X =
53.75 vs. 44–54, X = 49.35; p < 0.0001), more ventral scales (72–91, X = 83 vs. 61–76, X = 68.58; p < 0.0001),
more third finger lamellae (15–18, X = 16.25 vs. 13–16, X = 14.6; p < 0.0001), more fourth toe lamellae (20–23, X
= 21.29 vs. 16–22, X = 18.74; p < 0.0001), longer tails (64–77, X = 68.88 vs. 47–70, X = 60.52, p = 0.0019),
shorter forelimbs (elbow-wrist length: 4.89–6.81, X = 5.76 vs. 4.6–7.03, X = 6.16, third finger length: 7.27–9.52, X
= 8.02 vs. 8.31–11.12, X = 9.65; p < 0.0001 in both cases), smaller head width (9.02–10.19, X = 9.59 vs. 8.95–
11.71, X = 10.73; p < 0.0001), and smaller rostral scale (rostral height: 0.55–1.09, X = 0.85 vs. 0.87–1.2, X = 1.03,
rostral length 2.31–2.69, X = 2.51 vs. 2.46–3.12, X = 2.77; p < 0.0001 in both cases). Liolaemus avilae sp. nov.
exhibits a light background coloration, while the general background coloration of L. kolengh is dark gray or black;
however, the dorsal pattern could not be recognized in around 50% of our samples. Liolaemus kolengh exhibits
more dorsal blotches than L. avilae sp. nov. Ventral melanism is stronger in L. kolengh than in L. avilae sp. nov. In
general, the dorsal scales of L. avilae sp. nov. are less “bristly” than scales of L. kolengh. Limb scales (fore-hind
and dorsal-ventral) are non-mucronate and non-“bristly” in L. avilae sp. nov. (mucronated and “bristly” in L.
kolengh).
Liolaemus avilae sp. nov. differs from L. silvanae in several characters, including: a smaller number of scales
in contact with the interparietal (7–10, X = 8,13 vs. 5–8, X = 6.94; p = 0.0027), fewer supralabial scales (5–6, X =
5.25 vs. 5–7, X = 6.18; p = 0.0008), fewer midbody scales (53–59, X = 55.5 vs. 55–65, X = 59.65; p < 0.0001),
fewer scales from occiput to rump (51–58, X = 53.75 vs. 52–64, X = 56; p < 0.0001), more ventral scales (72–91,
X = 83 vs. 70–80, X = 74.59; p < 0.0001), smaller body size (snout-vent length: 48–59, X = 55.38 vs. 65–78, X =
73, axilla-groin distance: 19.4–29.5, X = 25.3 vs. 25.1–39.7, X = 32.38; p < 0.0001 in both cases), shorter fourth
toe length (13.82–15.98, X = 14.91 vs. 16.5–20.1, X = 18.46), shorter forelimbs (elbow-wrist length: 4.89–6.81, X
= 5.76 vs. 6.83–8.82, X = 7.71, third finger length: 7.27–9.52, X = 8.02 vs. 10.65–13.76, X = 12.23; p < 0.0001 in
both cases), smaller head (head height: 6.43–7.67, X = 7.07 vs. 6.84–9.06, X = 8.18, head width: 9.02–10.19, X =
9.59 vs. 11.31–14.71, X = 13.14, head length: 10.94–12.9, X = 11.78 vs. 12.47–15.9, X = 14.42; p < 0.0001 in all
cases), smaller eyes (eye height: 2–2.62, X = 2.2 vs. 2.09–2.9, X = 2.46, eye length: 2.63–3.32, X = 3.07 vs. 3.59–
4.18, X = 3.86; p < 0.0001 in both cases), smaller rostral scale (rostral height: 0.55–1.09, X = 0.85 vs. 1.01–1.35, X
= 1.19, rostral length: 2.31–2.69, X = 2.51 vs. 2.61–3.47, X = 3.13; p < 0.0001 in both cases), and smaller auditory
meatus (auditory meatus height: 1.95–2.56, X = 1.95 vs. 1.95–2.95, X = 2.54, auditory meatus length: 1.11–2.01, X
= 1.51 vs. 1.49–2.57, X = 2.04; p < 0.0001 in both cases). Background coloration in L. silvanae is black, there is a
no recognizable dorsal pattern, and ventral area is strongly melanistic, whereas the ventral area is weakly melanis-
tic in L. avilae sp. nov. Liolaemus avilae sp. nov. has a defined vertebral and paravertebral lines that are not present
in L. silvanae. In general, L. silvanae possesses a “bristly” appearance that is absent in L. avilae sp. nov. Scales of
limbs (fore-hind and dorsal-ventral) are less mucronated in L. avilae sp. nov. than in L. silvanae.
Description of holotype. Adult male. Snout-vent length 59.0 mm. Tail length (complete, not regenerated) 77.0
mm. Axilla-groin distance 26.9 mm. Head length 12.3 mm (from anterior border of tympanum to tip of snout), 10.1
mm wide (at anterior border of tympanum), 7.6 mm high (at anterior border of tympanum). Snout length 4.2 mm
(orbit-tip of snout distance). Interorbital distance 4.5 mm. Eye-nostril distance 3.5 mm. Auditory meatus-eye dis-
tance 4.9 mm. Forelimb length 16.3 mm. Tibial length 11.0 mm. Foot length 15.9 mm (ankle to tip of claw on
fourth toe).
Dorsal head scales bulged, smooth, 15 between occiput at level of anterior border of tympanum to rostral, pit-
ted with numerous scale organs in the anterior region, and reducing to a single organ, or absence at the posterior
half of the head. Rostral scale wider (2.8 mm) than high (0.8 mm). Two postrostrals, together with anterior lorila-
bial, separate nasal scales from rostral, surrounded by six scales. Nasal scales longer than wide, irregularly hexago-
nal, nostril one-half length of nasal, in a posterior position. Scales surrounding nasals 7 on the left side and 6 on the
right side. Four internasals. Frontonasals four, irregular in size and position. Prefrontals 5, a small quadrangular
scale in the center (1.1 mm), two dorso-lateral larger scales roughly hexagonal (1.7 mm), and a pair of lateral
medium-sized scales (1.2 mm), approximately triangular. Three frontal scales. Frontoparietals in two rows, one
anterior and two posterior scales. Interparietal pentagonal (1.3 mm), surrounded by seven scales; five in front and

BREITMAN ET AL.
18 · Zootaxa 3120 © 2011 Magnolia Press
sides, irregular in shape and size, and two larger posterior scales. Parietal eye evident. Parietals slightly bulged,
irregularly shaped, similar in size to interparietal. Circumorbitals: 12–10. Transversally expanded supraoculars 5–
4. Smaller lateral supraoculars: 16–19. One canthal higher than wide, separate from nasal by one postnasals. Loreal
scales bulged, four on each side. Lorilabials longer than wide (7–6), approximately equal to labials. Superciliaries
7–8, flattened and elongated, anterior four broadly overlapping dorsally. Orbit with 12–15 upper and 13–11 lower
ciliaries on each side. Orbit diameter 4.3 x 2.0 mm. Preocular small, unfragmented, square. Subocular scale elon-
gated, approximately nine times longer than wide (3.8 x 0.6 mm). A well marked longitudinal ridge along upper
margin of preocular and subocular scales. Postocular small, slightly bulged, ~ 25% superimposed onto subocular,
with a marked longitudinal ridge. Palpebral scales small granular and bulged. Supralabials 6–5, convex. Temporals
smooth, convex, juxtaposed with one scale organ in the tip. Anterior auriculars smaller than adjacent posterior tem-
porals, projecting slightly outward (3–2). Posterior auricular scales small and granular. External auditory meatus
conspicuous, higher (1.7 mm) than wide (1.2 mm). Lateral scales of neck granular with inflated skin. Mental scale
wider (3.0 mm) than high (1.0 mm), in contact with four scales. Mental followed posteriorly by two postmentals,
and two rows of three chinshields on each side. Infralabials 5-4 on each side, first on each side quadrangular two
times wider than supralabials, all others elongated, slightly smaller than supralabials. Gular scales smooth, flat,
imbricate, with rounded posterior margins, with melanophores. Scales of throat between chinshields slightly juxta-
posed, becoming slightly imbricate toward auditory meatus. Thirty-seven gulars between tympanum openings.
Infralabials separated from chinshields by one to two rows of scales. Antehumeral and longitudinal neck folds well
developed; postauricular, rictal, dorsolateral, oblique and gular not conspicuous.
Scales of dorsal neck region rhomboidal, imbricate, strongly keeled. Very few trifids scales. Fifty-five dorsal
scales between occiput and anterior surface of thighs. Dorsal body scales rhomboidal, imbricate, strongly keeled.
Dorsal scales grade laterally into slightly smaller and keeled scales at midbody. Scales immediately anterior and
posterior to forelimb and hindlimb insertion small, smooth, granular, and non-overlapping. Body lateral scales
grading smaller to larger at midbody. Ventral body scales rhomboidals, smooth, flat, imbricate, larger than dorsal
scales. Fifty-eight midbody scales; scales between mental and precloacal pores 90. Scales of cloacal region similar
in size to ventral body scales; without precloacal pores.
Anterior suprabrachials rhomboidal, imbricate, smooth, slightly larger in size to dorsal body scales. Postabra-
chials smaller, smooth, becoming granular near axilla. Supra-antebrachials similar to suprabrachial. Infra-antebra-
chials rhomboidal, imbricate, smooth. Supracarpals imbricated, rhomboidal, smooth. Infracarpals imbricate,
rhomboidal, very slightly keeled. Subdigital lamellae with 2–3 keels, each terminating in a short mucron, 2–3
mucronate, numbering: I: 8, II: 14, III: 17, IV: 19, V: 12. Claws robust, curved and sharp, opaque brown. Suprafem-
orals larger as dorsal body scales, rhomboidal, imbricated, smooth upper thigh, toward to the lateral thigh keeled.
Postfemorals small, granular shape. Supratibials rhomboidal, imbricated, moderately keeled, smaller and smooth
toward to the foot. Infrafemoral scale small, granular and smooth. Supratarsals rhomboidal, imbricated and smooth.
Infratarsals small, rhomboidal, imbricate, smooth, some keeled and mucronate near the digit. Subdigital scales
have 3 keels, 3–4 mucronate, numbering: I: 9, II: 14, III: 19, IV: 22, V: 16. Claws robust, curved and sharp, opaque
brown. Tail complete, non-regenerated. Dorsal and lateral caudal scales, rhombic in the first half of the tail, quad-
rangulars toward the tip, strongly keeled. Ventral subtriangular and smooth, toward posterior half moderatly
keeled.
Color of holotype in life. Dark gray dorsal background (Figure 7) that becomes lighter on the lateral region
between axilla and groin. Dorsal pattern presents nine paravertebral, quadrangulars, dark brown blotches, extend-
ing from the nuchal to the pelvic region, these series or blotches are combined at the beginning of the pelvic region
into a pattern of transversal bands, which is present to the tip of the tail. Dorsolateral region between the occipital
and the pelvic region is characterized by eight irregular, black and white-bordered blotches that are combined with
the transversal bands of the tail. Lateral region, between axilla and groin, is characterized by a black and white
reticulate pattern. In between the series of blotches a white vertebral and two yellowish paravertebral lines, one-
scale wide, are present. Vertebral line extends from the nuchal region, while the paravertebral lines extend from the
temporal region; all are present to the pelvic region.
Dark gray head background. Two lateral white lines along the longitudinal fold are present from the top of the
auditory meatus to the antehumeral fold. White subocular scale with a black longitudinal ridge. One longitudinal
band is present in the anterior part of the head, crossing though internasals, postnasals, prefrontals, frontals, post-
frontals and interparietal scales, ending in the interparietal scale that is surrounded by small brown spots; however,

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TWO NEW SPECIES FROM L. LINEOMACULATUS SECTION
a lighter central region is observable in the frontal, postfrontal and interparietal scales. Anterior circumorbitals
present a transverse dark brown band. Nasals scales dark brown. Small dark brown blotches on postocular, tempo-
ral and occipital region. Dorsal region of the limbs are dark gray with a dark brown reticulation.
FIGURE 7. Upper, dorsal and ventral views of holotype in life of L. avilae sp. nov. MLP.S 2627; below, dorsal and ventral
views of female (LJAMM-CNP 9399).

BREITMAN ET AL.
20 · Zootaxa 3120 © 2011 Magnolia Press
FIGURE 8. Type locality of Liolaemus avilae sp. nov. Altoandina phytogeographic province, Altoandino Austral District
(47º05’29.0” S, 71º01’12.9” W, 1154 m).

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TWO NEW SPECIES FROM L. LINEOMACULATUS SECTION
Gray ventral color with a dark reticulate pattern over the entire body, it is strongly marked in throat, chest, tail
and body laterals, and extend to the adjacent malar and maxilla region. Ventral scales with melanophores. Ventral
area of chest, belly, cloacal region and limbs tinged with a light orange color. Gray tail.
Color of holotype in preservative. After three years in preservative, the dorsal coloration of the head, dorsum,
body flanks and tail become darker while maintaining the contrast, but the two yellowish paravertebral lines turned
gray. Ventral scales of throat, neck, chest, belly and forelimbs retain the same coloration as in life, and the distinc-
tive light ventral orange color of the chest, belly, cloacal region and limbs turns gray (Figure 6).
Variation. Morphological and meristic character variation between male and female Liolaemus avilae sp.
nov., is shown in Table 6. Females in life present basically the same dorsal and lateral patterns as present in males,
but females generally have a lighter brown background coloration; vertebral and paravertebral lines are variable, in
some individuals they are strongly defined while in others they are less conspicuous; lines vary from white to yel-
low in females and from light orange to pink in males. Ventrally dark reticulate pattern in the entire body is present
and is strongly marked in the throat, tail and lateral body scales. Ventral, ventro-lateral, limb, cloacal and postcloa-
cal regions vary from males to females, in the former a ventral light orange coloration is present, while in one
female (out of three) a light pink ventral coloration is present. All distinctive ventral and dorsal coloration changed
from yellow, orange or pink to gray or white in preservative.
Etymology. The specific epithet of this species “avilae” refers to our colleague and the first author’s PhD advi-
sor Dr. Luciano Javier Avila, to honor him after more than fifteen years of research and teaching on the Patagonian
herpetofauna.
Distribution. Liolaemus avilae sp. nov. is known only from the type locality and surroundings, from Lago
Buenos Aires Plateau in Puesto Lebrun and 18.7 SW from there, Santa Cruz province, Lago Buenos Aires depart-
ment, Argentina (Figures 5, 8).
Natural history. Out of three females, only one presents a ventral pinkish coloration and a different one was
pregnant; thus we infer that the pinkish coloration in females is not related to pregnancy. The species occurs in the
Altoandina Phytogeographic province, Altoandino Austral district, in environments characterized as sparse grass-
lands and chamaephytes such as Empetrum rubrum, Nassauvia pygmaea and Azorrella ameghinoi, among others
(Roig 1998); some Festuca spp. and Senecio spp. were also observed. This new species was found in open volca-
nic-sand substrates between patches of basaltic rocks, sharing the habitat with Liolaemus archeforus and Liolaemus
silvanae (Figure 8). Based on the natural history of its sister species L. lineomaculatus, we hypothesize that L. avi-
lae sp. nov. is herbivorous and likely to also be viviparous, perhaps giving birth to between three and six individu-
als per clutch (Cei 1986; Espinoza et al. 2004).
Discussion
Lizards of the L. lineomaculatus section inhabit a large, heterogeneous area in Patagonia. This area possesses an
interesting geological history, including several glacial cycles and sea level shifts, making the phylogenetic and
phylogeographic history of this section complex and interesting. The L. lineomaculatus section includes 21 nomi-
nal species and several “candidate species” that remain unstudied (Breitman et al. 2011; Breitman et al. in press).
Morando et al. (2003) have shown that, by some estimates, that the diversity of Liolaemus may be as high as twice
the number of species described in the genus, and a recent study has shown that in the past decade the rate of spe-
cies description in Liolaemus has been growing exponentially (Martinez et al. in review).
Fouquet et al. (2007), proposed for Neotropical frogs a mtDNA approach to species delimitation based on iso-
lation-by-distance population structure. The method tests for correlation of geographic distance with genetic dis-
tance, which characterized most samples up to uncorrelated values of 3%. At this value the isolation-by-distance
correlation was not significant and Fouquet et al. (2007) interpreted this break as the limit to intra-specific gene
flow.
Martinez (personal communication) found that within Liolaemus the mean genetic distance value of cyto-
chrome b between sister species was ~ 4%; moreover, she found that different groups of Liolaemus had values
ranging from 1 to 6%. In the L. lineomaculatus section, average genetic distance between described species is 1.6%
(based on the sister species L. kolengh vs. L. hatcheri), and 2.25% between species of the kingii+archeforus group
(based on sister species L. archeforus vs. L. chacabucoense and L. escarchadosi vs. L. tari). In this context we want

BREITMAN ET AL.
22 · Zootaxa 3120 © 2011 Magnolia Press
to highlight that the discovery of the new species described here started with the molecular analyses presented by
Breitman et al. (2011), in which several clades were recognized as candidate species, continued with the nuclear
data here presented, and concluded with the morphological differentiation and characterization documented here
for each new species.
Even though the first attempt of classification of the L. lineomaculatus section placed these species in the Lio-
laemus sensu stricto group (Etheridge 1995), recent studies support the placement of the L. lineomaculatus section
within the Eulaemus group, sister to the montanus section within the broader Liolaemus phylogeny (Young
Downey 1998; Schulte et al. 2000; Morando 2004; Breitman et al. 2011).
In contrast to this well-supported position of the L. lineomaculatus section in the generic phylogeny, this clade
has until recently been in a state of taxonomic flux, with several authors interpreting this section nested within the
subgenus of Liolaemus. Laurent (1985, 1995) proposed the subgenus Rhytidodeira Girard (1858) for the L. lineo-
maculatus section, whereas Pincheira-Donoso and Núñez (2005) considered the use of Rhytidodeira inappropriate.
The lineomaculatus, kingii and archeforus groups have all been subject to several taxonomic rearrangements.
Before the description of these new species (L. morandae sp. nov. and L. avilae sp. nov.) the lineomaculatus group
included: L. lineomaculatus, L. hatcheri, L. kolengh and L. silvanae, and depending on the author, also L. magel-
lanicus. Some authors have considered the species L. silvanae, L. hatcheri and L. kolengh as members of a differ-
ent genus: Vilcunia (Donoso-Barros & Cei 1971) or later as a subgenus (Pincheira-Donoso et al. 2008) including L.
lineomaculatus, although this classification has been discouraged (Lobo et al. 2010a). Etheridge (1995) recognized
the lineomaculatus group (characterized morphologically by the absence of precloacal pores in males and the pres-
ence of some dorsal trifid scales) formed by the silvanae group (L. silvanae, L. hatcheri and L. kolengh) and the
species L. lineomaculatus. The silvanae group is characterized by the presence of keeled and imbricated nuchal
scales and post-femoral sub-imbricated scales (Abdala & Lobo 2006), but the use of “sub-imbricated” as a charac-
ter for classification seems ambiguous and in practice is not useful. The species L. magellanicus has been consid-
ered by Pincheira-Donoso and Nuñez (2005) and Breitman et al. (2011) a separate evolutionary lineage, since the
latter hypothesis was proposed based on molecular data, morphological work is needed to test it.
Analyses of all species of the L. lineomaculatus section using molecular sequence data from nine genes,
defined four well-supported clades: lineomaculatus, magellanicus, somuncurae and kingii+archeforus (Breitman
et al. 2011). Based on the previous morphology-based proposals and molecular data, the lineomaculatus group is
the only group that is clearly distinct from all other members of the section. Genetically, two hypotheses have been
proposed for species relationships within the lineomaculatus group: (1) a species tree approach (((L. lineomacula-
tus + L. morandae sp. nov. + L. avilae sp. nov.) (L. silvanae + L. kolengh)) L. hatcheri); versus (2) a concatenation
approach (L. hatcheri ((L. kolengh + L. silvanae) (L. morandae sp. nov. (L. avilae sp. nov. + L. lineomaculatus))))
(Breitman et al. 2011). Neither of these topologies support the previously defined silvanae group (L. silvanae + L.
kolengh + L. hatcheri) within the lineomaculatus group (Breitman et al. 2011) because it recovers L. hatcheri as
the sister taxa of all the other species of the lineomaculatus group. Based on these conflicts between the morpho-
logically proposed group and the molecular hypotheses, the need of a taxonomic revision of the lineomaculatus
group based on additional lines of evidence is now necessary.
At a more inclusive level, the entire L. lineomaculatus section is in need of a more detailed study to evaluate
each of its groups and taxonomic affinities. Our longer term goal is to prepare monographic revisions of poorly
known groups using an “integrative taxonomy” approach (Padial et al. 2010). We are currently using molecular,
morphological, ecological and geographical data, to reconstruct a temporal sequence of demographic histories, spe-
cies limits and relationships, and to eventually test for shared patterns of divergence with other Patagonian clades
(Sérsic et al. 2011).
Acknowledgments
Thanks to J.C. Bagley for help in data analyses and writing, and to N. Cazzaniga for help with the taxonomic
issues. We thank: L.J. Avila, M. Kozykariski, R. Martinez and N. Feltrin, for assistance in field collections, and
other members of the Grupo de Herpetología Patagónica for assistance in animal curation procedures. This
research benefitted from valuable discussions and comments from J.C. Bagley and two anonymous reviewers.
Financial support was provided by the following grants: PICT 2006-506 ANPCYT-FONCYT, ANPCYT-FONCYT

Zootaxa 3120 © 2011 Magnolia Press · 23
TWO NEW SPECIES FROM L. LINEOMACULATUS SECTION
33789, and a doctoral fellowship (MFB) from Consejo Nacional de Investigaciones Científicas y Técnicas
(CONICET), and NSF-PIRE award (OISE 0530267) for support of collaborative research on Patagonian Biodiver-
sity granted to the following institutions (listed alphabetically): Brigham Young University, Centro Nacional
Patagónico (AR), Dalhousie University, Instituto Botánico Darwinion (AR), Universidad Austral de Chile, Univer-
sidad de Concepción, Universidad Nacional del Comahue, Universidad Nacional de Córdoba, and University of
Nebraska. For additional financial and logistical support we thank the BYU Kennedy Center for International Stud-
ies, Department of Biology, and the Bean Life Science Museum. We also thank the Chubut province authorities for
financial/logistical support (to MFB), and the fauna authorities of Chubut and Santa Cruz provinces for collection
permits.
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APPENDIX II. Acronyms used for characters examined in this study; for definitions see Avila et al. (2010a, 2010b), Vega et
al. (2008), Martinez et al. (2011) and Abdala and Lobo (2006)
APPENDIX III. Eigenvalues, percent of explained variance, cumulated percent of explained variance for the first six principal
components (PC) and the contribution of each variable to the PCs (see Appendix I for names definitions)
Morphometric characters Meristic characters
SVL: Snout-vent length
TL: Tail length
DFH: Distance between fore and hind limbs
FOL: Foot length
TFL: Tibia-fibula length
RUL: Radius-ulna length
HAL: Hand length
HH: Head height
HW: Head wide
HL: Head length
EH: Eye height
EL: Eye length
RND: Rostral-nasal distance
RH: Rostral height
RL: Rostral length
DRE: Distance from rostral to the eye
AH: Auditory meatus height
AL: Auditory meatus length
SCI: Scales in contact with the interparietal
LS: Lorilabial scales
SS: Supralabial scales
IS: Infralabial scales
MS: Scales around midbody
DS: Dorsal scales
VS: Ventr al sc ale s
IL3: Infradigital lamellae (3rd on left hand)
IL4: Infradigital lamellae (4rd on left foot)
PC1 PC2 PC3 PC4 PC5 PC6
Eigenvalue 11.76 3.79 1.71 1.19 1.08 0.89
Percent of explained variance 45 15 7 5 4 3
Cumulated percent of explained variance 45 60 66 71 75 78
Contributions of the variables to the factors:
SVL 0.27 0.07 -0.10 0.10 -0.13 0.06
DFH 0.15 0.10 -0.21 0.23 -0.29 0.27
FOL 0.26 0.09 -0.07 -0.07 0.13 -0.07
TFL 0.22 0.18 -0.14 -0.06 0.22 -0.10
RUL 0.26 0.03 0.04 -0.15 0.01 -0.05
HAL 0.26 0.00 -0.08 -0.01 0.19 0.03
HH 0.23 0.13 0.11 0.04 -0.22 -0.08
HW 0.28 -0.01 0.06 0.01 -0.08 -0.03
HL 0.27 0.05 0.10 0.01 -0.12 -0.04
EH 0.21 0.03 -0.09 -0.16 0.15 0.20
EL 0.25 -0.06 0.02 -0.02 0.07 0.20
RND 0.22 -0.04 0.15 0.02 -0.31 -0.25
RH 0.19 -0.03 -0.09 0.36 0.06 -0.07
RL 0.25 -0.08 -0.05 0.05 -0.08 -0.05
DRE 0.27 -0.03 0.09 0.00 -0.15 -0.07
AH 0.24 -0.08 0.04 -0.04 0.12 0.00
AL 0.22 -0.11 0.06 -0.14 0.29 0.18
SCI -0.01 0.21 0.24 -0.18 -0.25 0.74
LS 0.00 0.03 0.43 -0.36 -0.30 -0.33
SS 0.00 -0.07 0.43 0.62 -0.04 0.06
IS 0.03 -0.15 0.49 0.21 0.45 0.06
MS 0.01 0.44 -0.13 0.16 0.06 -0.21
DS -0.02 0.42 -0.08 0.21 0.07 0.00
VS -0.10 0.42 -0.03 0.11 -0.04 0.00
IL3 0.02 0.35 0.21 -0.20 0.33 -0.05
IL4 -0.04 0.39 0.32 -0.05 0.02 -0.02

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APPENDIX IV. Discriminant-function analysis. Contribution of the principal components to the Canonical Axis, standarized
by the variance.
APPENDIX V. Position of the centroids in the discriminant space. Notice that the centroid of L. avilae sp. nov. is the only one
present in the (+ ; +); while the centroid of L. morandae sp. nov. is in the same discriminant quadrant (- ; +) than L. lineomacu-
latus, but its position is considerably lower respect to the axis 2.
APPENDIX VI. Cross-validation obtained from the discriminant-function analysis.
CA1 CA2
PC 1 1.03 -0.58
PC 2 0.69 0.83
PC 3 -0.83 0.00
PC 4 -0.2 0.16
PC 5 0.33 -0.12
PC 6 0.29 -0.26
Species Axis 1 Axis 2
L. hatcheri -1.63 -1.51
L. kolengh -1.63 -1.66
L. lineomaculatus -0.93 3.28
L. silvanae 5.11 -0.61
L. morandae sp. nov. -1.5 1.63
L. avilae sp. nov. 0.91 1.68
Species L. hatcheri L. kolengh L. lineomaculatus L. silvanae L. morandae sp. nov. L. avilae sp. nov. Error
(%)
L. hatcheri 10 7 0 0 1 0 44
L. kolengh 4200 00 0 17
L. lineomaculatus 00 13 03 1 24
L. silvanae 00 0 170 0 0
L. morandae sp. nov. 00 2 04 0 33
L. avilae sp. nov 00 0 00 7 0