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32 Accepted by S. Carranza: 18 Oct. 2011; published: 8 Dec. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 3123: 32–48 (2011)
www.mapress.com/zootaxa/Article
New species of lizard from the magellanicus clade of the Liolaemus lineomaculatus
section (Squamata: Iguania: Liolaemidae) from southern Patagonia
MARIA FLORENCIA BREITMAN1, CRISTIAN HERNÁN FULVIO PÉREZ1, MICAELA PARRA2,
MARIANA MORANDO1, JACK WALTER SITES, JR.3 & LUCIANO JAVIER AVILA1
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: liolaemu@criba.edu.ar; morando@cenpat.edu.ar; avila@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
A new species of the Liolaemus lineomaculatus section is described from southwestern Santa Cruz Province, Argentina.
The new species is a member of the monotypic magellanicus clade; morphological, molecular and geographical data are
sufficient to diagnose this new species as distinct form from L. magellanicus. The new species differs from L. magellani-
cus in having higher number of midbody, dorsal and ventral scales, and higher number of infradigital third finger and
fourth toe lamellae. The new species also differs in having smaller dorsal blotches on the hindlimbs and a more clearly
defined vertebral line, fewer precloacal pores and reduced dorsal scale mucronation, compared to L. magellanicus. Lio-
laemus caparensis sp. nov. is the second species described for the magellanicus group, and is geographically isolated from
L. magellanicus on the Campo Las Piedras Plateau, where it is sympatric with other endemic species of the L. lineomac-
ulatus section.
Key words: Iguania, Liolaemidae, Liolaemus lineomaculatus section, L. caparensis sp. nov., Patagonia, new species
Resumen
Se describe una nueva especie de la sección Liolaemus lineomaculatus del suroeste de la provincia de Santa Cruz. La nue-
va especie pertenece al clado monotípico magellanicus. Datos morfológicos, moleculares y geográficos ofrecen evidencia
para considerar esta nueva especie diferente a L. magellanicus. La nueva especie se diferencia de L. magellanicus por tener
un mayor número de escamas: alrededor del cuerpo, dorsales y ventrales, así también por poseer un mayor número de
lamelas subdigitales en el tercer dedo de la pata delantera y en el cuarto dedo de la pata trasera. La nueva especie también
se diferencia de L. magellanicus por tener manchas más pequeñas en las patas delanteras y una línea vertebral mejor
definida. La nueva especie posee un menor número de poros precloacales respecto de L. magellanicus. Las escamas dor-
sales de la nueva especie son menos mucronadas que las de L. magellanicus. Liolaemus caparensis sp. nov., es la segunda
especie descrita del grupo magellanicus, viviendo aislada de L. magellanicus en la Meseta Campo Las Piedras, lugar hab-
itado por otra especie endémica de la sección L. lineomaculatus.
Key words: Iguania, Liolaemidae, Sección Liolaemus lineomaculatus, L. caparensis sp. nov., Patagonia, nueva especie
Introduction
Liolaemus is one of the most ecologically diverse and species-rich genera of lizards on earth, with more than 230
recognized species (Núñez & Scolaro 2009; Abdala et al. 2010; Avila et al. 2010a, 2010b; Lobo et al. 2010a,
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A NEW SPECIES FROM L. LINEOMACULATUS SECTION
2010b; Martinez et al. 2011; Quinteros & Abdala 2011; Breitman et al. in press). Liolaemus extends from Perú to
Tierra del Fuego (14° ± 30’ – 52° ± 30’S) and ranges in altitude from sea level to almost 5,000 m, thereby inhabit-
ing many climatic regimes (Hellmich 1951; Donoso-Barros 1966; Cei 1986, 1993; Etheridge & de Queiroz 1988;
Frost & Etheridge 1989; Etheridge & Espinoza 2000).
The Liolaemus lineomaculatus section is part of the subgenus Eulaemus and includes 20 formally described
species (Breitman et al. 2011; Breitman et al. in press). This group of species has the most southerly distribution of
the genus, which extends from the Andean mountains of central Neuquén to coastal areas of Tierra del Fuego (Bot-
tari 1975; Cei 1986; Christie 2002). Historically the L. lineomaculatus section was divided morphologically into
three main groups: lineomaculatus (Etheridge 1995), kingii (Cei 1986) and archeforus (Cei 1986); plus the species
L. magellanicus that was always recognized as part of the section, but not clearly assigned to any of these three
main groups.
Liolaemus magellanicus was described by Hombron and Jacquinot in 1847 as Proctotretus magellanicus (type
locality: Havre Pecquet, in the Strait of Magellan, Chile), and is the most southerly distributed species of lizard of
the world. In 1858 Girard placed P. magellanicus in the new genus Rhytidodeira (Girard 1858) and since then, mul-
tiple taxonomic changes have been made. In 1979, Cei proposed the “L. magellanicus-lineomaculatus complex”,
and later morphological characters supported placement of the species in an ancestral line of Liolaemus called
“fueguiano (magellanicus-lineomaculatus)” (Laurent 1983). Two years later Laurent (1985) formally proposed the
magellanicus group, but with species relationships that today are not considered valid (Etheridge 1995, Schulte et
al. 2000). Cei (1986) proposed the morphological “kingii”, “archeforus” and “magellanicus” groups; the magel-
lanicus group included the species L. magellanicus, L. lineomaculatus and the genus Vilcunia (Donoso-Barros &
Cei 1971, which included L. silvanae and L. hatcheri). Cei (1986) did suggest that L. lineomaculatus probably
comprised a different group and later Laurent (1995) formally removed L. lineomaculatus from the genus Vilcunia
on the basis of its geographical distribution and several morphological characters. Laurent (1995) erected the
monotypic subgenus Austrolaemus for L. magellanicus, but later studies considered Austrolaemus as a synonym of
Liolaemus (e.g., Etheridge & Espinoza 2000; Pincheira-Donoso & Nuñez 2005; Pincheira-Donoso et al. 2008a)
and returned to the idea of the “magellanicus group” as a separate lineage from the “lineomaculatus group”. How-
ever, all these authors recognized the need for further study of relationships among these groups.
The first phylogenetic study of the Liolaemus lineomaculatus section, employing morphological and genetic
markers, showed that L. magellanicus was the sister species of the kingii and archeforus groups and a different lin-
eage from the lineomaculatus group; although, they did not show nodal-support values (Espinoza et al. 2004). New
evidence based on nine molecular markers (mitochondrial: cyt-b and 12S; and nuclear: Cmos, ACM4tg, PRLR,
LDA8F, LDA1D, LDA9C and LDA9D; 5865 bp of total alignment length) and phylogenetic inference from con-
catenation and species-tree methods (Breitman et al. 2011) identified four main clades within the L. lineomaculatus
section: (1) the lineomaculatus group (L. hatcheri, L. kolengh, L. silvanae, L. lineomaculatus); (2) the monotypic
magellanicus group; (3) the somuncurae group (L. somuncurae and L. uptoni); and (4) the kingii+archeforus group
(L. baguali, L. escarchadosi, L. tari, L. sarmientoi, L. scolaroi, L. zullyae, L. tristis, L. archeforus, L. chacabu-
coense, L. kingii and L. gallardoi); (Figure 1). The four clades recovered from the molecular work support the “tra-
ditional morphological” lineomaculatus group, but do not support the kingii and archeforus groups as separate
clades; moreover the authors proposed two different clades: the L. somuncurae and the L. magellanicus groups.
Breitman et al. (2011) also identified eight candidate species spread through the four main clades including the
magellanicus group (L. magellanicus and another terminal identified as Liolaemus sp. 3) and particularly from the
lineomaculatus group the species referred as L. sp. 1 and L. sp. 2 are under description (Breitman et al. in press).
The magellanicus group presents the southernmost distribution of Liolaemus, extending across the Strait of
Magellan to Tierra del Fuego island. This group has attracted scientific attention due to its ability to live in
extremely harsh environments (Jacksic & Schwenk 1983; Pincheira-Donoso et al. 2008b; Ibargüengoytía et al.
2010; Fernández et al. 2011). Here, we use an integrative approach based on color pattern, morphometric, meristic,
molecular and qualitative characters to describe a new species belonging to the magellanicus group (the Liolaemus
sp. 3 terminal in Breitman et al. 2011).
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Material and methods
We examined series of specimens from the type localities of the following species; lineomaculatus group: L.
kolengh (n = 42), L. hatcheri (n = 20), L. silvanae (n = 21) and L. lineomaculatus (n = 17); magellanicus group
(n = 29): eleven specimens of the new species and eighteen specimens from L. magellanicus (Appendix I); all of
these specimens were previously used in Breitman et al. (2011). Specimens were collected by hand and sacrificed
by pericardial injection of sodium pentothal Abbot®. After a liver sample was extracted for molecular analyses,
specimens were fixed in 20% formalin and later transferred to 70% ethanol. Voucher specimens were placed in La
Plata Museum, Argentina (MLP.S) and in the Herpetological collection LJAMM-CNP of Centro Nacional
Patagónico in Puerto Madryn, Argentina, http://www.cenpat.edu.ar/nuevo/colecciones03.html. Other material
examined (but not included in the analysis) is placed in the following herpetological collections: Jose Miguel Cei -
Diagnostic Collection (JMC-DC), University of San Luis, Argentina (JMCDC); Field Museum of Natural History,
Chicago, USA (FMNH); University of La Plata, Argentina (MLP.S/R); Museum of Vertebrate Zoology, University
of California, Berkeley, USA (MVZ); Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos
Aires, Argentina (MACN) (Appendix I). The general geographic distribution of L. magellanicus was obtained from
the collections listed above and bibliographic references (Cei 1971; Bottari 1975; Jaksic & Schwenk 1983; Scolaro
1992; Schulte et al. 2000).
Because there is strong morphological and molecular evidence (see introduction) to consider the magellanicus
group as a separate clade, for this description we only performed detailed comparisons among the new species and
Liolaemus magellanicus. We used 18 morphometric characters and 12 meristic characters (commonly used and
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. Mea-
surements were taken only on adults (seventeen of L. magellanicus and eight of the new species) 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), and coloration in life was observed from pictures taken at the time of capture. Sex was determined by the
thickness of the base of the tail and presence of precloacal pores. Where numbers of paired scales are provided,
they are given as left-right.
Univariate tests were performed in INFOSTAT® 2009 (Di Rienzo et al. 2010). We implemented statistical tests
for both morphometric and meristic characters, and compared frequencies of qualitative characters. A Student’s T
test was used to evaluate the significance of differences of variable means between the new species and Liolaemus
magellanicus, and assumptions (normality and variance homogeneity) were checked with the Levene and Shapiro-
Wilks tests (Montgomery 1991). When the assumptions of Student’s T test were not met, we performed a nonpara-
metric Kruskal-Wallis test (Kruskal & Wallis 1952). Sexual dimorphism was tested in for all variables with either
Student’s T or Kruskal-Wallis test (when same assumptions previously described were not met).
Phylogenetics relationships between all the species plus eight candidate species of the L. lineomaculatus sec-
tion 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; Gamble et al. 2008; Townsend et al.
2008), and then edited and aligned using the programs Sequencher v4.8 (™Gene Codes Corporation Inc. 2007) and
Clustal X (Higgins & Sharp 1988, 1989; Thompson et al. 1997). The total lengh of the nuclear alignment was 4180
bp, while the total length of the nuclear plus the mitochondrial alignment was 5865 bp; 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: Recombination Detection Program v3.44 (Martin & Rybicki 2000; Heat et al. 2006). Bayesian analyses were
conducted for the concatenated matrix using MrBayes v3.1.2 (Ronquist & Huelsenbeck 2003) with four chains and
run for 50 million generations 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 assumed convergence because all parameters had
effective sample sizes greater than 200, as determined from Tracer v1.5.0 (Rambaut & Drummond 2009).
We used the cyt-b fragment from individuals of the new species and L. magellanicus collected near the type
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A NEW SPECIES FROM L. LINEOMACULATUS SECTION
locality, to calculate genetic distance between species. Genetic distances between species (corrected pairwise dif-
ferences = intergroup distance – intragroup distance) was calculated using Arlequin v3.11 (Excoffier et al. 2005).
Results
Morphological tests showed significant differences between the new species and Liolaemus magellanicus, which
are described in the Diagnosis section of the new species. The means, standard deviations and ranks for meristic
and qualitative characters are summarized in Tables 1 and 2, and the p values of Levene, Shapiro-Wilks and
Kruskal-Wallis tests are summarized in Tables 3 and 4. Sexual dimorphism was not found in any of the meristic
variables, although some statistical differences in the morphometric variables were found. Variables that present
sexual dimorphism are mainly related to head size, for which females were smaller than males (head height: 5.65-
6.07, X = 6.16 vs. 6.72-7.77, X = 7.05; p = 0.0390, head width: 8.4-9.33, X = 8.8 vs. 9.37-11.07, X = 10.41; p =
0.013, head length: 10.43-11.4, X = 10.95 vs. 11.69-13.41, X=12.38; p = 0.018, rostral length: 2.26-2.66, X = 2.41
vs. 2.71-2.93, X = 2.83; p = 0.088, distance from rostral to eye: 4.38-4.99, X = 4.65 vs. 5.09-5.49, X = 5.23; p =
0.016, auditory meatus high: 1.65-2.09, X = 1.82 vs. 1.94-2.24, X = 2.13; p = 0.047) and the variable foot length
(11.37-13.19, X = 12.52 vs. 14.68-15.3, X = 14.99; p = 0.0068) ) (Table 5). Since these variables did not show sta-
tistical differences for discriminating the new species from L. magellanicus, and the meristic data set showed
strong evidence for discriminating species, we did not perform comparisons between species by separating sexes
for these analyses.
The uncorrected genetic distance between the new species and L. magellanicus was 2.99%, while the corrected
distance was > 2.6%. Figure 1 summarizes phylogenetic relationships among species and distinct lineages within
the L. lineomaculatus section, using all of the data and the nuclear-only data set, and shows the position of the new
species.
TABLE 1. Values (in millimeters) of morphometric characters from species of the magellanicus group; sample sizes are shown
in parentheses, as well as mean ± SD (min–max); abbreviations for characters are defined in Appendix II.
Var L. caparensis sp. nov. (n=8) L. magellanicus (n=17)
SVL 51.75±3.92 (47–59) 55.41±5 (49–68)
TL 67.71±8.01 (60–84) 63.58±7.28 (51–77)
DFH 24.23±1.43 (21.2–25.9) 25.96±3.52 (22.1–36.2)
FOL 13.23±1.35 (11.37–15.3) 13.91±0.72 (12.85–15.1)
TFL 7.24±0.77 (6.19–8.62) 9.09±0.91 (6.91–10.76)
EWL 5.24±0.4 (4.58–5.66) 5.98±0.52 (5.22–6.88)
HAL 7.92±0.72 (7.08–8.91) 8.24±0.57 (7.25–9.19)
HH 6.5±0.63 (5.65–7.7) 6.92±0.65 (5.53–7.89)
HW 9.41±1.02 (8.4–11.07) 9.76±0.75 (8.50–11.22)
HL 11.49±0.93 (10.43–13.41) 11.78±0.89 (10.33–13.47)
EH 1.86±0.21 (1.58–2.29) 1.98±0.31 (1.47–2.87)
EL 3.06±0.17 (2.85–3.38) 2.95±0.21 (2.6–3.28)
RND 1.95±0.25 (1.58–2.26) 2.03±0.17 (1.77–2.29)
RH 0.98±0.07 (0.9–1.12) 0.92±0.10 (0.74–1.10)
RL 2.57±0.26 (2.26–2.93) 2.75±0.26 (2.37–3.29)
DRE 4.87±0.38 (4.38–5.49) 4.72±0.32 (4.16–5.3)
AH 1.93±0.23 (1.65–2.24) 1.92±0.18 (1.53–2.3)
AL 1.6±0.19 (1.33–1.88) 1.57±0.20 (1.15–1.91)
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TABLE 2. Values of meristic variables from species of the magellanicus group; sample sizes are shown in parentheses, as well
as mean ± SD (min–max); abbreviations for characters are defined in Appendix II.
TABLE 3. Student’s T tests and related tests for means of characters showing significant differences between the two species
compared in Table 1; the Shapiro-Wilks and Levene p-values are shown when Student’s T tests were significant, as described in
the text. To assume normality and variance homogeneity Shapiro-Wilks and Levene test should not be rejected (p > 0.05).
TABLE 4. Student’s T tests and related tests for means of characters showing significant differences between the two species
compared in Table 2; the Shapiro-Wilks and Levene p-values are shown when Student’s T tests were significant, as described in
the text. To assume normality and variance homogeneity Shapiro-Wilks and Levene test should not be rejected (p > 0.05).
Kruskal Wallis p is shown for DS (dorsal scales) variable (assumptions for Student’s T test were not met).
Liolaemus caparensis sp. nov.
(Figure 2)
Holotype. MLP.S 2628 (Figure 2), an adult male from Campo Las Piedras plateau, 7 km N Punta del Lago ranch,
Lago Argentino department, Santa Cruz province, Argentina (49º 34’ 11.0” S, 72º 02’ 51.9” W, 868 m; Figures 3,
4), L.J. Avila, C.H.F. Pérez, M.F. Breitman and N. Feltrin collectors, 16th January 2008.
Paratypes. LJAMM–CNP 9379–9380 adult males, LJAMM–CNP 9383–9387 females and LJAMM–CNP
9382, 9388, 9389 juveniles; from the same locality of holotype, L.J. Avila, C.H.F. Pérez, M.F. Breitman and N. Fel-
trin collectors, 16 th January 2008.
Diagnosis. Liolaemus caparensis sp. nov. is a member of the magellanicus group, in the L. lineomaculatus sec-
tion, based on molecular evidence (Figure 1). Liolaemus caparensis sp. nov. differs from L. magellanicus in the
following traits: shorter tibia-fibula length (6.19-8.62, X = 7.24 vs. 6.91-10.76, X = 9.09; p < 0.0001), shorter
Var L. caparensis sp. nov. (n=11) L. magellanicus (n=18)
SCI 6.55±1.04 (5–8) 6.89±0.90 (6–9)
LS 4.82±0.75 (3–6) 4.28±0.67 (3–5)
SS 6.73±0.79 (6–8) 6.56±0.51 (6–7)
IS 4.73±0.65 (3–5) 4.61±0.61 (4–6)
MS 43.82±4.33 (39–55) 40.44±3.45 (34–48)
DS 41.45±2.5 (39–47) 37.50±2.62 (34–45)
VS 68.36±2.98 (64–73) 59.17±5.91 (49–77)
IL3 15±1.18 (14–18) 13.33±1.08 (11–15)
IL4 20.3±1.49 (18–23) 18.24±1.35 (15–20)
Pores 3.67±0.58 (3–4) 3.91±0.94 (3–5)
Var Student’s T test pShapiro-Wilks Levene
SVL 1.79 0.0866* 0.1869 0.5091
TL 37.37 < 0.0001*** 0.4642 0.8976
TFL 4.98 < 0.0001*** 0.9724 0.6559
RUL 3.55 0.0017** 0.2495 0.4699
Var Student’s T test pShapiro-Wilks Levene
LS -2.02 0.0539* 0.1394 0.8026
MS -2.32 0.0282** 0.0536 0.4175
DS Kruskal Wallis 0.0003 *** non-parametric
VS -5.55 < 0.0001*** 0.5038 0.4339
IL3 -3.88 0.0006*** 0.2754 0.7425
IL4 -3.69 0.0011** 0.5513 0.088
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A NEW SPECIES FROM L. LINEOMACULATUS SECTION
radius-ulna length (5.24-5.66, X = 5.24 vs. 5.22-6.88, X = 5.98; p = 0.0017), larger number of midbody scales (39-
55, X = 43.82 vs. 34-48, X = 40.44; p = 0.0282), larger number of dorsal scales (39-47, X = 41.45 vs. 34-45, X =
37.5; p = 0.0003), more ventral scales (64-73, X = 68.36 vs. 49-77, X = 59.17; p < 0.0001), more infradigital lamel-
lae of the third finger (14-18, X = 15 vs. 11-15, X = 13.33; p = 0.0006), and more infradigital lamellae of the fourth
toe (18-23, X = 20.3 vs. 15-20, X = 18.24; p = 0.0011) (for details see Tables 1 to 4). Liolaemus caparensis sp. nov.
has smaller dorsal blotches on the hindlimbs and a more defined vertebral line in comparison with L. magellanicus
(Figure 5), as well as fewer precloacal pores than L. magellanicus (3-4 vs. 3-5). Dorsal scales of Liolaemus capar-
ensis sp. nov. have a shorter mucron than those of L. magellanicus. The dorsal surface of limb scales is more cari-
nated and less mucronated in L. caparensis sp. nov. than in L. magellanicus (Figure 5).
FIGURE 1. Relationships between Liolaemus caparensis sp. nov. and species of the L. lineomaculatus section and selected
species of the subgenus Eulaemus and Liolaemus sensu stricto. On the left, phylogenetic relationships recovered using seven
nuclear genes (Cmos, ACM4tg, PRLR, LDA8F, LDA1D, LDA9C and LDA9E, 4180 bp total alignment length) with Bayesian
species-tree inference; 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 (5865 bp) including the seven nuclear genes plus two mitochon-
drial genes (12S and cyt-b), and summarizes information from MP and ML methods. Nodes with high support from three meth-
ods (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.
Description of holotype. Adult male. Snout-vent length 59.0 mm. Tail length (complete, not regenerated) 84.0
mm. Radius-ulna length 25.9 mm. Head length 13.4 mm (from anterior border of tympanum to tip of snout), 11.0
mm wide (at anterior border of tympanum), 7.7 mm long (at anterior border of tympanum). Snout length 4.9 mm
(orbit-tip of snout distance). Auditory meatus-eye distance 4.7 mm. Interorbital distance 4.2 mm. Eye-nostril distance
3.1 mm. Forelimb length 17.5 mm. Tibial length 8.6 mm. Foot length 14.6 mm (ankle to tip of claw on fourth toe).
Dorsal head scales slightly bulged, smooth, 14 between occiput, at level of anterior border of tympanum, to
rostral, pitted with numerous scale organs in the anterior region, and reducing to no organ in the posterior half of
the head. Rostral scale wider (2.8 mm) than long (0.9 mm). Two postrostrals that together with anterior lorilabial
separate nasal scales from rostral, surrounded by six scales. Nasal scales longer than wide, irregularly hexagonal;
nostril one-half length of nasal, posterior in position. Scales surrounding nasals 7 on each side. Four internasals.
Frontonasals and prefrontals 14, irregular in shape, size and position. Two frontal scales. Frontoparietals in two
rows, two anterior and one posterior scale. Interparietal pentagonal (1.3 mm), surrounded by five scales; three
smaller and irregular in front and sides, two larger in back. Parietal eye evident. Parietals slightly bulged, irregu-
larly shaped, equal in size to interparietal. Circumorbitals: 9 on the left side, incomplete; 10 on the right side,
incomplete. Transversally expanded supraoculars 4–3. Smaller lateral supraoculars: 16–19. Three canthals on the
left side, one posterior large, higher than wide, in contact with postnasal, two inferior, small; four canthal on the
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right side, one posterior with similar characteristics that the left side, and three inferior, small. Loreal scales slightly
bulged, 4–5 on each side. Lorilabials longer than wide (7–8), slightly smaller than labials. Superciliaries 7 on each
side, flattened and elongated, anterior four broadly overlapping dorsally. Orbit with 13–12 upper and 11–10 lower
ciliaries on each side. Orbit diameter 4.7 x 2.3 mm. Preocular small, unfragmented, longer than wide. Subocular
scale elongated, approximately nine times longer than wide (4.0 x 0.5 mm). A well marked longitudinal ridge along
upper margin of preocular and subocular scales. Postocular small, slightly bulged, quarter superimposed to subocu-
lar, with a marked longitudinal ridge. Palpebral scales, small granular and bulged. Supralabials 6 on each side, con-
vex. Temporals smooth, convex, juxtaposed with one scale organ in the tip. Anterior auriculars smaller than
adjacent posterior temporals, slightly projecting outward (1 on each side). Posterior auriculars small and granular.
External auditory meatus conspicuous, rounded (1.8 x 1.5 mm). Lateral scales of neck granular with inflated skin.
Mental scale wider (3.0 mm) than high (1.3 mm), in contact with four scales. Mental followed posteriorly by two
postmentals, and two rows of 2–1 chinshields. Five 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,
imbricate, with rounded posterior margins, with melanophores. Scales of throat between chinshields slightly imbri-
cate. Twenty-six gular scales between tympanum openings. Infralabials separated from chinshields by one to two
rows of scales. Antehumeral and longitudinal neck folds well developed; gular fold incomplete; postauricular, ric-
tal, dorsolateral and oblique folds unconspicuous.
Scales of dorsal neck region rhomboidal, imbricate, strongly keeled, mucronate. Thirty-nine scales between
occiput and anterior surface of thighs. Dorsal body scales rhomboidal, imbricate, strongly keeled and mucronate.
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 lateral scales
grading smaller to larger at midbody. Ventral body scales rhomboidal, smooth, flat, imbricate, and bigger than dor-
sal scales. Forty-four midbody scales; scales between mental and precloacal pores 71. Scales of cloacal region
about equal in size to ventral body scales. Four 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 rhomboidal, imbricate, smooth, some of
the external lateral slightly keeled. Infra-antebrachials rhomboidals, imbricate, smooth, toward the hand slightly
keeled. Supracarpals imbricated, rhomboidal, smooth. Infracarpals strongly imbricate, rhomboidal, slightly keeled
and mucronate. Infradigital lamellae with 3-keels, each terminating in a short mucron, 3-mucronate, numbering: I:
8, II: 12, III: 16, IV: 19, V: 11. Claws robust, curved and sharp, opaque brown.
Suprafemorals as large as dorsal body scales, rhomboidal, imbricated, strongly keeled and mucronate. Post-
femorals small, granular shape. Supratibial rhomboidal, imbricated, strongly keeled and mucronate. Infrafemoral
scales small, granular and smooth. Supratarsals rhomboidals, imbricated and smooth, some lateral scales slightly
keeled. Infratarsals small, rhomboidal, imbricate, smooth, mucronate, some slightly keeled and mucronate near the
digit. Subdigital scales 3-keeled, 3-mucronate, numbering: I: 9, II: 13, III: 19, 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 and mucronate. Ventrals subtriangular and
smooth, toward posterior half moderately keeled.
Color of holotype in life. Grey dorsal background (Figure 2). The posterior borders of the dorsal scales,
between neck and pelvic region, present one or two contrasting light colors (white or light blue). Dorsal pattern
with eight series of paravertebral and quadrangular black (with a white end) blotches, appearing from the nuchal
region to about the first autotomy line, those series fuse into a dark line that is present to the tip of the tail. Dorso-
lateral region, between the occipital and the pelvic regions, has eight irregular black and white-bordered blotches.
Between the series of blotches there are three longitudinal lines, a white vertebral and two light brownish paraver-
tebral lines, the last ones are wider than the vertebral (one and half scale vs. a quarter of scale). The coloration in
the vertebral and lateral area is light brown.
Anterior region of the head shows a dark gray background with a central black blotch from the internasals to
the frontal scales. Orbitals, postfrontals and parietals are light brown, the black circumorbital scales continue in a
line of the same color until the nuchal region. Gray temporal and lateral head regions. Limbs dorsal region are gray
with a scarce black reticulation with some lighter color scales.
Ventral scales with melanophores. Black color is present throughout all the ventral body until the postcloacal
region. Dark reticulate area is present in the throat and extends to the adjacent malar and maxilla region. Cloacal
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A NEW SPECIES FROM L. LINEOMACULATUS SECTION
and postcloacal regions are black. Ventral area of lower belly and femoral region present a bright yellow color. Tail
is light gray (Figure 2).
FIGURE 2. Upper, dorsal and ventral views of male holotype in life of Liolaemus caparensis sp. nov. MLP.S 2628; below,
dorsal and ventral views of female (LJAMM-CNP 9387).
Color of holotype in preservative. After three years in preservative, the dorsal coloration of the head, dorsum,
body flanks and tail becomes darker while maintaining the contrast, but the two light brownish paravertebral lines
turned gray. Ventral scales of throat, neck, chest, belly and forelimbs maintain the same dark coloration as in life,
and the distinctive yellow ventral coloration of the femoral and belly regions turns gray (Figure 5).
Variation. Morphological and meristic characters variation, between males and females of Liolaemus caparen-
sis sp. nov. are shown in Table 5. Females in life present basically the same dorsal and lateral pattern as present in
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males, but the coloration has a light brown background; vertebral and paravertebral lines are wider and much more
notorious than the ones in males, the lines are white; although, one of the lizards presents yellowish dorsolateral
lines. Only the females present a white lateral line from the axilla to the groin. Ventral coloration shows a dark
reticulate pattern in throat, melanic chest and belly, but two specimens present a yellowish coloration in the ventro-
lateral, femoral, cloacal and postcloacal regions. All distinctive dorsal lines, and ventral coloration, femoral and
lower belly areas turns in preservative from yellowish or bright yellow to gray.
TABLE 5. Variation between females and males of Liolaemus caparensis sp. nov. summarized as mean ± SD (min–max) val-
ues for all meristic and morphometric (in millimeters) variables.
Etymology. The specific epithet of this species “caparensis” refers to the name “Capar” given to the actual
Viedma lake by the Aónikenk aborigines that inhabited this area. The Campo Las Piedras plateau is located on the
northwestern edge of Viedma Lake.
Distribution. Liolaemus caparensis sp. nov. is known only from the type locality, from Campo Las Piedras
plateau, 7 km N Punta del Lago ranch, Lago Argentino department, Santa Cruz province, Argentina. 868 m (Fig-
ures 3 and 4).
Var Females (n=5) Males (n=3)
SCI 6.75±1.04 (5–8) 6±1 (5–7)
LS 5±0.53 (4–6) 4.33±1.15 (3–5)
SS 6.88±0.83 (6–8) 6.33±0.58 (6–7)
IS 4.75±0.71 (3–5) 4.67±0.58 (4–5)
MS 44.25±4.95 (39–55) 42.67±2.31 (40–44)
DS 42.13±2.59 (39–47) 39.67±1.15 (39–41)
VS 68.75±3.01 (64–73) 67.33±3.21 (65–71)
IL3 14.88±1.36 (14–18) 15.33±0.58 (15–16)
IL4 20.25±1.67 (18–23) 20.5±0.71 (20–21)
Pores 0 3.67±0.58 (3–4)
SVL 49.8±2.77 (47–54) 55±3.61 (52–59)
TL 63±2.94 (60–67) 74±8.72 (68–84)
DFH 23.84±1.62 (21.2–25.2) 24.87±0.93 (24.1–25.9)
FOL 12.52±0.71 (11.37–13.19) 14.99±0.44 (14.68–15.3)
TFL 6.86±0.55 (6.19–7.55) 7.86±0.72 (7.19–8.62)
RUL 5.08±0.44 (4.58–5.66) 5.51±0.12 (5.38–5.61)
HAL 7.55±0.53 (7.08–8.36) 8.53±0.59 (7.85–8.91)
HH 6.16±0.41 (5.65–6.7) 7.05±0.56 (6.72–7.7)
HW 8.8±0.43 (8.4–9.33) 10.41±0.91 (9.37–11.07)
HL 10.95±0.37 (10.43–11.4) 12.38±0.91 (11.69–13.41)
EH 1.79±0.14 (1.58–1.89) 1.98±0.27 (1.77–2.29)
EL 2.99±0.1 (2.85–3.07) 3.16±0.24 (2.91–3.38)
RND 1.83±0.22 (1.58–2.15) 2.16±0.13 (2.01–2.26)
RH 0.97±0.05 (0.9–1.01) 1±0.1 (0.94–1.12)
RL 2.41±0.16 (2.26–2.66) 2.83±0.11 (2.71–2.93)
DRE 4.65±0.25 (4.38–4.99) 5.23±0.22 (5.09–5.49)
AH 1.82±0.18 (1.65–2.09) 2.13±0.17 (1.94–2.24)
AL 1.58±0.25 (1.33–1.88) 1.62±0.03 (1.59–1.64)
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A NEW SPECIES FROM L. LINEOMACULATUS SECTION
FIGURE 3. Type locality of Liolaemus caparensis sp. nov., Subantartic Phytogeographic province, Estepa de Festuca palle-
scens District (49º 34’ 11.0” S, 72º 02’ 51.9” W).
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FIGURE 4. Distribution map for species of the magellanicus group, with localities sampled superimposed on regional eleva-
tion (shading). White square: Liolaemus caparensis sp. nov.; black squares: distribution of L. magellanicus; black and gray
squares: L. magellanicus localities used in this study from LJAMM-CNP collection. Arrow indicates type locality of L. magel-
lanicus (in Chile). SC: Santa Cruz province. Notice L. caparensis sp. nov. distribution does not overlap with the distribution of
L. magellanicus (south of the Santa Cruz river).
Natural history. The new species is viviparous and probably insectivorous (inferred from the biology of its
sister species Liolaemus magellanicus; Ibargüengoytía et al. 2010). Out of five females, only two presented a ven-
tral yellowish coloration and one of those was gravid, thus we inferred that the yellowish coloration in females is
not directly related with the reproductive state. The specimens were found in the Subantarctic Phytogeographic
province, Estepa de Festuca pallescens district, in an environment mainly characterized as Festuca grasslands
(Roig 1998). Other dominant vegetation included short bushes (Anartrophyllum desideratum, Senecio spp.) and
grasses (Stipa spp.) (Figure 3). This species was found in open substrates between rocks, sharing the habitat with
Liolaemus tari.
Concluding remarks. Lizards from the magellanicus group have a general phenotype (general body form, col-
oration, and color pattern) similar to lizards of the lineomaculatus group, but they have precloacal pores, a synapo-
morphy shared with all other species of the Liolaemus lineomaculatus section (kingii+archeforus and somuncurae
groups) but not with species from the lineomaculatus group.
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
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A NEW SPECIES FROM L. LINEOMACULATUS SECTION
(based on sister species L. archeforus vs. L. chacabucoense and L. escarchadosi vs. L. tari). Even though the
genetic distance values calculated for L. caparensis sp. nov. relative to L. magellanicus are slightly higher than val-
ues of other species from the L. lineomaculatus section, we want to highlight that the discovery of the new species
started with the genetics 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 differenti-
ation and characterization of L. caparensis sp. nov. presented here.
FIGURE 5. Morphological comparison between Liolaemus magellanicus (left) and L. caparensis sp. nov. (right). Differences
in dorsal scales are shown in blue/light blue, in L. magellanicus scales are more mucronated than in L. caparensis sp. nov.; dif-
ferences in dorsal scales of hindlimbs are shown in green/light green, in L. magellanicus scales are more mucronated and less
carinated than in L. caparensis sp. nov. Notice differences in color pattern on the tails, size of blotches on the limbs, and the dif-
ferences in vertebral lines.
Even though there is not a formal description of the morphological characteristics that define the magellanicus
group, the features that defined the species Liolaemus magellanicus do differentiate it from the other groups of the
L. lineomaculatus section (scale counts and shape, coloration patterns, presence of precloacal pores and size,
among others: Scolaro 1992; Laurent 1995; Etheridge & Espinoza 2000; Pincheira-Donoso & Núñez 2005). The
divergence of the magellanicus group from the rest of the section was inferred to have occurred during the Late
Miocene (~8.46 million years ago, Breitman et al. 2011).
Morphological, molecular, and geographic distributional data provide evidence to consider Liolaemus capar-
ensis sp. nov. a distinct species most closely related to L. magellanicus, forming the magellanicus group. Because
the aim of this work was to describe the new species and not to propose new taxonomical arrangements, we avoid
hypothesizing new taxonomies or presenting morphological features that might characterize new classifications.
Including the species described here and two more species described in another manuscript (Breitman et al. in
press), the number of species included in the L. lineomaculatus section increases to 21.
Our research group is studying the Liolaemus lineomaculatus section to delimit species boundaries and recon-
struct relationships from multiple sources of evidence. Our goal is to present monographic revisions of poorly
known groups on the basis of an “integrative taxonomy” approach (Padial et al. 2010), based on independent lines
of evidence to propose taxonomic re-arrangements and support hypotheses of species limits. In parallel, we are
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using integrative phylogeographic analyses based on molecular, morphological, ecological, and geographical data,
to reconstruct a temporal sequence of demographic histories, and to eventually test for shared patterns of diver-
gence with other Patagonian clades (Sérsic et al. 2011).
Acknowledgments
Special thanks to J.C. Bagley for help in data analyses and writing and to N. Cazzaniga for help with taxonomic
issues. We also thank to N. Frutos, M. Kozykariski, M. Nicola, R. Martinez, N. Feltrin and C. Zanotti, for assis-
tance in field collections, and other members of the Grupo de Herpetologia 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-FON-
CYT (LJA), ANPCYT-FONCYT 33789 (MM), and a doctoral fellowship (MFB) from Consejo Nacional de Inves-
tigaciones Científicas y Técnicas (CONICET), and NSF-PIRE award (OISE 0530267) for support of collaborative
research on Patagonian Biodiversity granted to the following institutions (listed alphabetically): Brigham Young
University, Centro Nacional Patagónico (AR), Dalhousie University, Instituto Botánico Darwinion (AR), Universi-
dad Austral de Chile, Universidad de Concepción, Universidad Nacional del Comahue, Universidad Nacional de
Córdoba, and University of Nebraska. We also thank the BYU Kennedy Center for International Studies, Depart-
ment of Biology, and the Bean Life Science Museum for additional funds, the Chubut province authorities for addi-
tional financial and logistic support (MFB), and fauna authorities from Chubut and Santa Cruz provinces for
collection permits.
References
Abdala, C.S., Quinteros, A.S., Scrocchi, G.J. & Stazzonelli, J.C. (2010) Three new species of the Liolaemus elongatus group
(Iguania: Liolaemidae) from Argentina. Cuadernos de Herpetología, 24, 93–109.
Avila, L.J., Pérez, C.H.F., Morando, M., & Sites, J.W., Jr. (2010a) A new species of Liolaemus (Reptilia: Squamata) from
southwestern Rio Negro province, northern Patagonia, Argentina. Zootaxa, 2434, 47–59.
Avila, L.J., Morando, M., Pérez, D.R. & Sites, J.W., Jr. (2010b) A new species of the Liolaemus elongatus clade (Reptilia: Igua-
nia: Liolaemini) from Cordillera del Viento, northwestern Patagonia, Neuquén, Argentina. Zootaxa, 2667, 28–42.
Bottari, C.V. (1975) Sobre la presencia de Liolaemus magellanicus en Tierra del Fuego, Argentina (Reptilia, Iguanidae). Physis,
34, 211–213.
Breitman, M.F., Avila, L.J., Sites, J.W., Jr. & Morando, M. (2011) Lizards from the end of the world: Phylogenetic relationships
of the Liolaemus lineomaculatus section (Squamata: Iguania: Liolaemini). Molecular Phylogenetics and Evolution, 59,
364–376.
Breitman, M.F., Parra, M., Pérez, C.H.F. & Sites, J.W., Jr. (in press) Two new species of lizards from the Liolaemus lineomacu-
latus section (Squamata: Iguania: Liolaemidae) from southern Patagonia. Zootaxa.
Cei, J.M. (1971) Herpetología Patagónica. I. Liolaemus del grupo magellanicus. Características taxonómicas y genéticas. Phy-
sis, 81, 417–424.
Cei, J.M. (1979) The Patagonian herpetofauna. In: Duellman, W. E. (Ed.). The South American herpetofauna: Its origin, evolu-
tion, and dispersal. Museum of Natural History, Univ. Kansas, Lawrence. Monograph 7, 309–329 pp.
Cei, J.M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas áridas y semiáridas. Museo
Regionale di Scienze Naturali Torino, 4, 1–527.
Cei, J.M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, Puna y Pam-
pas. Museo Regionale di Scienze Naturali Torino, 14, 1–949.
Christie, M.I. (2002) Liolaemus lineomaculatus (Sauria: Liolaemidae) en el noroeste patagónico. Cuadernos de Herpetología,
16, 79–91.
Di Rienzo, J.A., Casanoves, F., Balzarini, M.G., González, L., Tablada, M. & Robledo, C.W. (2010) InfoStat. Grupo InfoStat;
FCA, Universidad Nacional de Córdoba, Argentina. Available: http://www.infostat.com.ar (accessed 08 September 2011).
Donoso-Barros, R. (1966) Reptiles de Chile. Ediciones de la Universidad de Chile, Santiago, 458 pp.
Donoso-Barros, R. & Cei, J.M. (1971) New lizards from the Patagonian volcanic plateau of Argentina. Journal of Herpetology,
5, 89–95.
Espinoza, R.E., Wiens, J.J. & Tracy, C.R. (2004) Recurrent evolution of herbivory in small, cold-climate lizards: breaking the
ecophysiological rules of reptilian herbivory. Proceedings of the National Academy of Sciences, 101, 16819–16824.
Etheridge, R. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia:
Squamata: Tropiduridae). American Museum Novitates, 3142, 1–34.
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
Zootaxa 3123 © 2011 Magnolia Press · 45
A NEW SPECIES FROM L. LINEOMACULATUS SECTION
Etheridge, R. & de Queiroz, K. (1988) A phylogeny of Iguanidae. In: Estes R., Pregill, G. (Eds.), Phylogenetic Relationships of
Lizard Families: Essays Commemorating Charles L. Camp. Stanford Univ. Press, Palo Alto, California, pp. 283–368.
Etheridge, R. & Espinoza, R.E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semi-annotated
bibliography. Smithsonian Herpetological Information Service, 126, 1–64.
Excoffier, L., Laval, G. & Schneider, S. (2005) Arlequin ver. 3.0: An integrated software package for population genetics data
analysis. Evolutionary Bioinformatics Online, 1, 47–50.
Fernández, J.B., Smith, J., Jr., Scolaro, A. & Ibarguengontía, N.R. (2011) Performance and thermal sensitivity of the southern-
most lizards in the world, Liolaemus sarmientoi and Liolaemus magellanicus. Journal of Thermal Biology, 36, 15–22.
Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. & Gemmell, N.J. (2007) Underestimation of species richness in neo-
tropical frogs revealed by mtDNA analyses. PLoS ONE, 2, e1109.
Frost, D. (1992) Phylogenetic analysis and taxonomy of the Tropidurus group of lizards (Iguania, Tropiduridae). American
Museum Novitates, 3033, 1–68.
Frost, D.R. & Etheridge, R. (1989) A phylogenetic analysis and taxonomy of Iguanian lizards (Reptilia: Squamata). University
of Kansas Museum Natural History Miscellaneous Publication, 81, 1–65.
Gamble, T., Bauer, A.M., Greenbaum, E. & Jackman, T.R. (2008) Evidence for Gondwanan vicariance in an ancient clade of
gecko lizards. Journal of Biogeography, 35, 88–104.
Girard, C. (1858) Descriptions of some new reptiles collected by the U.S. Exploring Expedition, under the command of Capt.
Charles Wilkes, U.S.N., Fourth part. - Including the species of Saurians, exotic to North America. Proceedings of the
Academy of Natural Sciences of Philadelphia, 9, 195–199.
Guindon, S. & Gascuel, O. (2003) A simple, fast and accurate method to estimate large phylogenies by maximum-likelihood.
Systematic Biology, 52, 696–704.
Heath, L., van der Walt, E., Varsani, A. & Martin, D.P. (2006= Recombination patterns in aphthoviruses mirror those found in
other picornaviruses. Journal of Virology, 80, 11827’11832.
Hellmich, W.C. (1951) On ecotypic and autotypic characters, a contribution to the knowledge of the evolution of the genus Lio-
laemus (Iguanidae). Evolution, 5, 359–369.
Higgins, D.G. & Sharp, P.M. (1988) CLUSTAL: a package for performing multiple sequence alignment on a microcomputer.
Gene, 73, 237–244.
Higgins, D.G. & Sharp, P.M. (1989) Fast and sensitive multiple sequence alignments on a microcomputer. CABIOS, 5, 151–
153.
Hombron, J. & Jacquinot, H. (1847) Reptiles. In: Dumont d’Urville, M.J. (Ed.), Voyage au Pôle Sud et dans l’Océanie sur les
corvettes l’Astrolabe et la Zélée. Paris, 1837–1840 pp.
Huelsenbeck, J.P. & Ronquist, F. (2001) MrBayes: Bayesian inference of phylogeny. Bioinformatics, 17, 754–755.
Ibargüengoytía, N.R., Marlin Medina, S., Fernández, J.B., Gutiérrez, J.A., Tappari, F. & Scolaro, A. (2010) Thermal biology of
the southernmost lizards in the world: Liolaemus sarmientoi and Liolaemus magellanicus from Patagonia, Argentina.
Journal of Thermal Biology, 35, 21–27.
Jacksic, F.M. & Schwenk, K. (1983) Natural history observations on Liolaemus magellanicus, the southernmost lizard in the
world. Herpetologica, 39, 457–461.
Kruskal, W.H. & Wallis, W.A. (1952) Use of ranks on one-criterion variance analysis. Journal of the American Statistical Asso-
ciation, 47, 583–621.
Laurent, R.F. (1983) Contribución al conocimiento de la estructura taxonómica del género Liolaemus Wiegmann (Iguanidae).
Boletín de la Asociación Herpetológica Argentina, 1, 16–18.
Laurent, R.F. (1985) Segunda contribución al conocimiento de la estructura taxonómica del género Liolaemus Wiegmann
(Iguanidae). Cuadernos de Herpetología, 1, 1–37.
Laurent, R.F. (1995) A tentative arrangement of subgenera of the genus Liolaemus Wiegmann (Reptilia: Squamata: Tropiduri-
dae). Bulletin of the Maryland Herpetological Society, 31, 10–14.
Lobo, F., Espinoza, R.E. & Quinteros, S. (2010a) A critical review and systematic discussion of recent classification proposals
for liolaemid lizards. Zootaxa, 2549, 1–30.
Lobo, F., Slodki, D. & Valdecantos, S. (2010b) Two new species of lizards of the Liolaemus montanus group (Iguania: Liolae-
midae) from the northwestern uplands of Argentina. Journal of Herpetology, 44, 279–293.
Martin, D. & Rybicki, E. (2000) RDP: detection of recombination amongst aligned sequences. Bioinformatics, 16, 562–563.
Martinez, L.E., Avila, L.J., Pérez, C.H.F., Perez, D.R., Sites, J.W., Jr. & Morando, M. (2011) A new species of Liolaemus
(Squamata, Iguania, Liolaemini) endemic to the Auca Mahuida Mountain, northwestern Patagonia, Argentina. Zootaxa,
3010, 31–46.
Montgomery, D.C. (1991) Diseño y Análisis de Experimentos. Grupo Editorial Iberoamérica. 585 pp.
Núñez, H. & Scolaro, J.A. (2009) Liolaemus (Donosolaemus) chacabucoense, nueva especie de lagartija para la región de
Aisén, Chile (Reptilia, Sauria). Boletín del Museo Nacional de Historia Natural de Chile, 58, 67–74.
Padial, J.M., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, 1–
14.
Pincheira-Donoso, D. & Núñez, H. (2005) Las especies chilenas del género Liolaemus (Iguania, Tropiduridae, Liolaeminae).
Taxonomía, sistemática y evolución. Publicación Ocasional del Museo Nacional de Historia Natural de Chile, Santiago,
486 pp.
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
BREITMAN ET AL.
46 · Zootaxa 3123 © 2011 Magnolia Press
Pincheira-Donoso, D., Scolaro, A. & Sura, P. (2008a) A monographic catalogue on the systematics and phylogeny of the South
American iguanian lizard family Liolaemidae (Squamata, Iguania). Zootaxa, 1800, 1–85.
Pincheira-Donoso, D., Hodgson, D.J., & Tregenza, T. (2008b) The evolution of body size under environmental gradients in
ectotherms: why should Bergmann’s rule apply to lizards?. BMC Evolutionary Biology, 8, 68–78.
Posada, D. (2008) jModelTest: Phylogenetic model averaging. Molecular Biology and Evolution, 25, 1253–1256.
Quinteros, A.S. & Abdala, C.S. (2011) A new species of Liolaemus of the Liolaemus montanus section (Iguania: Liolaemidae)
from Northwestern Argentina. Zootaxa, 2789, 35–48.
Roig, A. F. (1998) La vegetación de la Patagonia. In: Correa M.N. (Ed.), Flora Patagónica. Colección científica del INTA,
Tomo VIII, parte I, Buenos Aires, Argentina, pp. 48–174.
Rambaut, A. & Drummond, A. J. (2009) Tracer v1.5, Available at http://beast.bio.ed.ac.uk/Tracer (accessed 08 September
2011).
Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics,
9, 1572–1574.
Saint, K.M., Austin, C.C., Donnellan, S.C. & Hutchinson, M.N. (1998) C-mos, a nuclear marker useful for squamate phyloge-
netic analysis. Molecular Phylogenetics and Evolution, 10, 259–263.
Schulte, J.A., Macey, J.R., Espinoza, R.E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolae-
mus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariances and dispersal. Biological
Journal of the Linnean Society, 69, 75–102.
Scolaro, J.A. (1992) Morphological differences between Vilcunia and L. magellanicus and L. lineomaculatus: Discriminant
analysis. Acta Zoológica Lilloana, 41, 287–293.
Sérsic, A.N., Cosacov, A., Cocucci, A.A., Johnson, L.A., Pozner, R., Avila, L.J., Sites, J.W., Jr., & Morando, M. (2011) Emerg-
ing phylogeographic patterns in plants and terrestrial vertebrates from Patagonia. Biological Journal of the Linnean Soci-
ety, 110, 475–494.
Smith, H.M. (1946) Handbook of lizards. Comstock Publishing Company, Ithaca, New York, U.S.A, 557 pp.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The Clustal X windows interface: flexible
strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876–4882.
Townsend, T.M., Alegre, R.E., Kelley, S.T., Wiens, J.J. & Reeder, T.W. (2008) Rapid development of multiple nuclear loci for
phylogenetic analysis using genomic resources: an example from squamate reptiles. Molecular Phylogenetics and Evolu-
tion, 47, 129–142.
Vega, L.E., Bellagamba, P. & Lobo, F. (2008) A new endemic species of Liolaemus (Iguania: Liolamidae) from the mountain
range of Tandilia, Buenos Aires province, Argentina. Herpetologica, 64, 81–91.
Wiens, J.J., Reeder, T.W. & Montes de Oca, A.N. (1999) Molecular phylogenetics and evolution in sexual dichromatism among
population of the Yarrow’s spiny lizard (Sceloporus jarrovii). Evolution, 53, 1884–1897.
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A NEW SPECIES FROM L. LINEOMACULATUS SECTION
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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), and Martinez et al. (2011).
Morphometric 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
Meristic characters
SCI: Scales in contact with interparietals
LS: Lorilabial scales
SS: Supralabial scales
IS: Infralabial scales
MS: Midbody scales
DS: Dorsal scales
VS: Ventral scales
IL3: Infradigital lamellae (3rd on left hand)
IL4: Infradigital lamellae (4rd on left foot)
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