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Accepted by S. Carranza: 30 May 2011; published: 20 Jun. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2924: 1–21 (2011)
www.mapress.com/zootaxa/Article
1
Two new mountain lizard species of the Phymaturus genus (Squamata: Iguania)
from northwestern Patagonia, Argentina
LUCIANO JAVIER AVILA1, CRISTIAN HERNAN FULVIO PEREZ1,
DANIEL ROBERTO PEREZ2 & MARIANA MORANDO1
1Centro Nacional Patagónico – Consejo Nacional de Investigaciones Científicas y Técnicas (CENPAT-CONICET), Boulevard Alm-
irante Brown 2915, U9120ACD, Puerto Madryn, Chubut, Argentina. E-mail: avila@cenpat.edu.ar; liolaemus@gmail.com;
morando@cenpat.edu.ar
2Universidad Nacional del Comahue, Buenos Aires 1400, 8300, Neuquén, Argentina. E-mail: ddneuquen@hotmail.com
Abstract
Two new species of lizards of the saxicolous and viviparous genus Phymaturus from Patagonia are described. The new
species are members of the Phymaturus patagonicus species group distributed only in volcanic plateaus and mountains of
southern Argentina. Phymaturus sitesi sp. nov. differs from all other members of the patagonicus group in its unique dor-
sal pattern of small white spots on a light-gray to brown-gray background and allopatric geographical distribution. Ab-
sence of sexual dichromatism differentiated Phymaturus sitesi sp. nov. from P. payuniae, P. zapalensis, and P. delheyi n.
sp. , and the midbody scales count differentiated from the P. delheyi sp. nov. and P. nevadoi (with some overlap). Also
Phymaturus sitesi sp. nov. presents lower count in the ventral scales than in P. delheyi sp. nov. Phymaturus delheyi sp.
nov. can be distinguished from other species of the patagonicus group by unique dorsal pattern of medium size white spots
on a dark-brown background and allopatric distribution. The presence of sexual dichromatism differentiates Phymaturus
delheyi sp. nov. from P. sitesi sp. nov. and P. nevadoi. Scales around midbody differentiated Phymaturus delheyi sp. nov.
from P. payuniae and P. sitesi sp. nov. Also ventral scale count differentiated Phymaturus delheyi sp. nov. from P. sitesi
sp. nov. and P. nevadoi (with some overlap).
Key words: Argentina, Iguanidae sensu lato, Liolaemini, patagonicus group, new species, Neuquén
Resumen
Se describen dos especies nuevas de lagartijas saxícolas y vivíparas del género Phymaturus de Patagonia. Las nuevas es-
pecies son miembros del grupo de especies Phymaturus patagonicus distribuido solamente en mesetas y montañas vol-
cánicas del sur de Argentina. Phymaturus sitesi sp. nov. difiere de todos los otros miembros del grupo patagonicus por su
patrón dorsal único de pequeñas manchas blancas sobre un fondo marrón claro y su distribución geográfica alopátrica. La
ausencia de dicromatismo sexual diferencia Phymaturus sitesi sp. nov. de P. payuniae, P. zapalensis, y P. delheyi sp. nov.
y el numero de escamas alrededor del cuerpo lo diferencia de P. delheyi sp. nov. y P. nevadoi (con alguna superposición).
También Phymaturus sitesi sp. nov. tiene menos escamas ventrales que P. delheyi sp. nov. Phymaturus delheyi sp. nov.
puede ser distinguido de otras especies del grupo patagonicus por su patrón dorsal único de manchas de mediano tamaño
sobre un fondo oscuro y distribución alopátrica. La presencia de dicromatismo sexual diferencia Phymaturus delheyi sp.
nov. de P. sitesi sp. nov. y P. nevadoi. El numero de escamas alrededor del cuerpo diferencian Phymaturus delheyi sp. nov.
de P. payuniae y P. sitesi sp. nov. También el numero de escamas ventrales diferencia Phymaturus delheyi sp. nov. de P.
sitesi sp. nov. y P. nevadoi (con alguna superposición).
Palabras claves: Argentina, Iguanidae sensu lato, Liolaemini, grupo patagonicus, nuevas especies, Neuquén
Introduction
Phymaturus is a group of saxicolous lizards found in Andean, southern Puna, Famatina, Payunia, and Patagonian
rocky environments of Argentina and central Andean mountains of Chile from 25° to 45°30’ south latitude and
AVILA ET AL.
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between 400 to 4000 m in elevation (Lobo et al. 2010, Nuñez et al. 2010). All known species are viviparous (with
a report of facultative parthenogenesis in captivity, Chiszar et al. 1999), herbivorous (but some species occasion-
ally eat insects, Scolaro et al. 2008, Perez pers. observation), and saxicolous (Cei 1986, 1993, Lobo et al. 2010,
Nuñez et al. 2010). Some characters, including wide and flattened head and body, tail with regular whorls of
spinose scales, lateral nuchal skin folds with fat-filled pouches, and a short interclaviculae, are exclusive of Phyma-
turus, providing strong evidence of monophyly (Etheridge 1995). Within Phymaturus Etheridge (1995) recognized
two groups, the palluma group, with larger species, found mainly along the eastern and western Andes slopes,
including some southern Puna regions and Famatina mountain, and the patagonicus group, smaller in size, found
mainly in extra-andean mountain chains and volcanic plateaus of Patagonian environments. This same arrangement
was recovered by Espinoza et al. (2004) in a phylogenetic study combining molecular and morphological data, but
Lobo and Quinteros (2005a) found mixed results in a morphological study, with the patagonicus group recovered
as paraphyletic in some analyses. An ongoing study (Morando et al. in review) finds strong support for Etheridge’s
proposal and additional hidden diversity within this genus. At the present time, the patagonicus group contains 19
described species (Table 1): P. patagonicus Koslowsky 1898, P. spurcus Barbour 1921, P. indistinctus Cei & Castro
1973, P. payuniae Cei & Castro 1973, P. somuncurensis Cei & Castro 1973, P. zapalensis Cei & Castro 1973, P.
nevadoi Cei & Roig 1975, P. calcogaster Scolaro & Cei 2003, P. excelsus Lobo & Quinteros 2005, P. spectabilis
Lobo & Quinteros 2005, P. tenebrosus Lobo & Quinteros 2005, P. cei i Scolaro & Ibarguengoytia 2007, P. manu e-
lae Scolaro & Ibarguengoytia 2008, P. desuetus Scolaro & Tappari 2009, P. vid elai & P. castillensis Scolaro &
Pincheira Donoso 2010, P. etheridgei Lobo et al. 2010, and P. f elixi Lobo et al. 2010. Recently P. agilis Scolaro et
al. 2008 was suggested as a synonym of P. spectabilis and P. desuetus was described based on only one specimen,
that could be a color variation from a very polymorphic population of other Phymaturus species (Lobo et al. 2010);
but until a more detailed study is available, we still accept its validity. Almost all this species diversity is recog-
nized based on external morphology and coloration coupled with the geographical isolation of populations. Within
the patagonicus group, the northernmost distributed species group is the well supported payuniae group that
includes P. nevadoi, P. payuniae, and P. zapalensis (Morando et al. in review). This clade is distributed along table-
lands, isolated volcanoes or volcanic hills, of Neuquén and Mendoza provinces, south of Atuel River and north to
Limay River. The rest of the species of the patagonicus group is distributed in volcanic tablelands or mountains in
the Patagonian Steppe region, in Rio Negro and Chubut provinces, south to the Limay River basin (P. agilis, P. cal -
cogaster, P. castillensis, P. ceii, P. desuetus, P. etheridgei, P. excelsus, P. fel ixi, P. indistinctus, P. manue lae, P.
patagonicus, P. somuncurensis, P. spurcus, P. spectabilis, P. v idelai, P. tenebrosus). Lizard surveys carried out in
recent years in northern Neuquén and southern Mendoza provinces have revealed several new species of Liolaem-
ini lizards (e.g. Liolaemus antumalguen Avila et al. 2010, L. cuyumhue Avila et al. 2009, L. puelche Avila et al.
2007, L. gununakuna Avil a et al. 2004, L. punmahuida Avila et al. 2003; Phymaturus querque Lobo et al. 2010,
among other new species still in description) showing that this area is one of the most important “lizard hostspots”
of Argentina. As part of these studies we obtained several samples of Phymaturus from two mountainous areas, the
isolated Mio-Pliocene Sierra of Auca Mahuida and the Plio-Pleistocene Tromen Massif. Based on comparisons
with other species of the patagonicus species group inhabiting related mountains and plateaus (Phymaturus
payuniae, P. nevadoi and P. zapalensis), and taking into account coloration differences, geographic isolation, and
genetic differentiation (Morando et al., in review), we describe these two populations as new species of the north-
ern clade (payuniae) of the Phymaturus patagonicus species group.
Material and methods
We examined 175 specimens belonging to 14 species of the patagonicus group, including 28 specimens of the type
series of the new species described here. A more detailed comparison was made with samples of four populations
of three other species (Phymaturus nevadoi, Phymaturus payuniae, and Phymaturus zapalensis) of the payuniae
clade (Table 2, see Appendix 1). We chose these populations because they are morphologically the most similar
and could lead to misidentifications; also, based on molecular markers, they are phylogenetically most closely
related to the species described here (the payuniae group, Morando et al. in review), and they are distributed in the
same geographic area. Additional data were obtained from the original descriptions of species of the patagonicus
clade. Specimens were collected by hand, and killed in our lab by a pericardiac injection of sodium tiopenthotal
Zootaxa 2924 © 2011 Magnolia Press · 3
TWO NEW SPECIES OF PHYMATURUS FROM PATAGONIA
AVILA ET AL.
4 · Zootaxa 2924 © 2011 Magnolia Press
Pentovet®, fixed in 10–20% formalin and later transferred to 70% ethanol. Measurements were taken with a digital
caliper to the nearest 0.1 mm. The following measurements were taken with a digital caliper to the (nearest 0.1
mm): Snouth-Vent Length (SVL; from tip of snout to vent), Axilla-Groin Distance (AGD; from axilla to groin),
Foot Length (FL; from tip of claws to heel), Tibial Length (TL; greatest length of tibia, from knee to heel), Arm
Length (AL; from tip of claws to elbow), Head Length (HL; distance between anterior edge of auditory meatus and
snout tip), Head Width (HW; taken from the temporal regions), and Head High (HH; maximum height of head,
from occiput to throat). Some character states were observed with the aid of a binocular stereomicroscope. Scale
terminology follows Smith (1946), and recent treatments of related species by Cei (1986, 1993), Etheridge (1995),
Lobo and Quinteros (2005), and Lobo et al. (2010). Where numbers of paired scales are provided they are given as
left-right. Terminology of lateral neck folds follows Frost (1992). Descriptions of color in life are based on field
observations of live specimens and color photographs of recently captured animals. We examined samples of
related species of the patagonicus group (including some type series of the Museo de La Plata collection) from the
herpetological collections of Monte L. Bean Life Science Museum, Brigham Young University (BYU); Museo de
La Plata, Universidad Nacional de La Plata (MLP.S/R); Museum of Vertebrate Zoology, University of California –
Berkeley (MVZ); Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos Aires (MACN), and the
herpetological collection LJAMM-CNP of the Centro Nacional Patagónico, Puerto Madryn, Argentina (CENPAT–
CONICET). Additionally, morphological data to compare the new species were taken from original descriptions of
species of the group (Koslowsky 1898; Barbour 1921; Cei & Castro 1973; Cei & Roig 1975; Lobo & Quinteros
2005a, b; Scolaro & Cei 2003; Scolaro et al. 2005; Scolaro & Ibarguengoytia 2007; Scolaro & Ibargüengoytia
2008; Scolaro & Tappari 2009; Scolaro & Pincheira Donoso 2010; Lobo et al. 2010). Dorsal coloration of males
and females of the majority of the species of the Phymaturus patagonicus group are in Scolaro and Pincheira
Donoso (2010).
TABLE 2. Morphometric, meristic, and chromatic characteristics in species of the northern subclade of the Phymaturus patag-
onicus group.
Results
Phymaturus sitesi sp. nov.
(Figure 1, 2)
Type material. Holotype: MLP.S 2605, adult male collected on rocky cliffs on the northeastern slope of Sierra de
Auca Mahuida mountain (37º 43’ S, 68º 55’ W, 1983 m, datum = WGS 84), near Cerro de las Antenas, Auca
Mahuida Natural Protected Area, Pehuenches Department, Neuquén province, Argentina, C.H.F. Perez collector.
Characters delheyi sp. nov.
(n = 8) sitesi sp. nov.
(n = 20) nevadoi
(n = 9) payuniae
(n = 6) zapalensis
(n = 8)
SVL 78.0–93.7 81.54–92.58 82.26–92.67 84.33–86.34 88.04–94.28
Midbody scales 198–227 210–238 155–217 205–234 204–243
Dorsal scales in a head length 39–48 38–50 39–49 43–50 40–50
Ventral scales 174–202 160–190 162–179 170–200 166–200
Precloacal pores 7–8 7–12 8–11 1–7 7–9
Supralabials/infralabials 7–10/6–9 8–12/5–8 9–10/6–8 11–13/6–8 8–10/5–8
Finger/Toe lamellae 21–25/29–34 20–23/27–32 20–25/27–30 21–24/28–32 19–25/25–30
Dorsal background dark-brown gray-brown dark-brown brown-black black
Dorsal pattern white spots (1–
10 scales) uni-
formly distrib-
uted
white spots (1–2
scales) uni-
formly distrib-
uted
white spots (4–9
scales) and grow
to be larger on
flanks
irregular white
spots (4–40
scales), some-
times fused
irregular white spots
(5–14 scales),
sometimes fused
Sexual dichromatism Yes No No Yes Yes
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TWO NEW SPECIES OF PHYMATURUS FROM PATAGONIA
Paratypes: MLP.S 2606/8 males, MLP.S 2607, BYU 12590/1, LJAMM-CNP 10367/8, 10466/9, females. Same
data as holotype. LJAMM-CNP 12297-98, males, Riscos Altos, 38.6 km S junction Provincial Road 6, Auca
Mahuida Natural Protected Area, (37º 43’ S, 68º 55’ W, 1851 m, datum = WGS 84), Pehuenches Department, Neu-
quén Province, Argentina. L.J. Avila, M.L. Kozykariski, and M.F. Breitman, collectors. LJAMM-CNP 12157,
12189, males, LJAMM-CNP 12214, female, Park Ranger Post, 38.6 km S junction Provincial Road 6, Auca Mahu-
ida Natural Protected Area (37º 42’ S, 68º 51’ W, 1560 m, datum = WGS 84), Pehuenches Department, Neuquén
Province, Argentina. L.J. Avila, M.L. Kozykariski, and M.F. Breitman, collectors. LJAMM-CNP 13402, male,
road SW Park Ranger Post, Auca Mahuida Natural Protected Area (37º 42’ S, 68º 53’ W, 1659 m, datum = WGS
84), Pehuenches Department, Neuquén Province, Argentina. L.J. Avila and I. Minoli, collectors. LJAMM-CNP
12311-12312, Communication Station Site, South Slope, Auca Mahuida Natural Protected Area, (37º 46’ S, 68º
53’ W, 1935 m, datum = WGS 84), Añelo Department, Neuquén Province, Argentina. L.J. Avila, M.L. Kozyka-
riski, and M.F. Breitman, collectors.
FIGURE 1. Phymaturus sitesi, dorsal and ventral view of the holotype (MLP.S 2605), from Sierra of Auca Mahuida mountain,
Pehuenches Department, Neuquén province, Argentina.
Diagnosis. Phymaturus sitesi is a robust and medium sized member of the clade referred to as the patagonicus
group by Etheridge (1995), because it has flat imbricate superciliaries, non-rugose dorsal scales on tail, and the
subocular scale is usually not fragmented. This new species is allopatric from and differs from all other members of
the clade in its unique dorsal pattern of small white dots occupying only 1–2 scales on a light-gray to brown-gray
background. Phymaturus sitesi can be distinguished from other species of the patagonicus group (P. agilis, P. cal -
cogaster, P. castillensis, P. ceii, P. desuetus, P. etheridgei, P. excelsus, P. fel ixi, P. indistinctus, P. manue lae, P.
patagonicus, P. somuncurensis, P. spurcu s, P. spectabilis, P. videlai , P. tenebrosus) by colour pattern features and
disjunct geographical distribution. Absence of sexual dichromatism differentiated Phymaturus sitesi from P.
AVILA ET AL.
6 · Zootaxa 2924 © 2011 Magnolia Press
payuniae, P. zapalensis, and Phymaturus sp. nov. (described below). Dorsal pattern of P. sitesi is composed by
small white spots uniformly distributed (occupying 1 scales, a few occupy 2 scales) scattered along head, limbs and
trunk, a pattern never observed in P. payuniae. Ventral coloration in P. sitesi is darker than in P. payuniae, and dor-
sal coloration becomes grayish along the throat, forelimbs, and sides of chest and venter. Reticulated pattern in ven-
tral areas and gular zones observed in P. payunie is not present in P. sitesi. Dorsal pattern of P. zapalensis is
composed by irregular white spots (between 5–14 scales each, sometimes fused) scattered along head and trunk but
not in limbs and tail, white spots become smaller in lateral areas, and then lateral dark areas turn into dark bands
between shoulder and rump, a pattern never observed in P. sitesi. Ventral coloration in P. zapalensis is similar to
that observed in P. sitesi, but the gular region has irregular black spots sometimes fused to form a reticulated pat-
tern. Females of P. payuniae and P. zapalensis have dorsal or lateral patterns of bands never found in P. sitesi. Phy-
maturus nevadoi has a dorsal pattern similar to P. sitesi but dorsal dots usually occupy more scales (4–9 scales vs
1–2), they become larger on flanks (not in P. sitesi), and form a reticulated pattern in the edges of the ventral region
between limbs. Phymaturus sitesi has some overlap in midbody scale count (210–238 vs 198–217 in P. nevadoi).
Phymaturus sitesi can be distinguished from the new species described below by the dorsal pattern described
above. Scales count around midbody are higher in P. sitesi than in Phymaturus sp. nov. (210–238 vs 198–227) and
lower in ventral scales showing some overlap (160–190 vs 174–202).
FIGURE 2. Dorsal and ventral view in a paratype female of Phymaturus sitesi.
Description of the holotype. Adult male 91.1 mm (SVL). Axilla-groin distance 49.3 mm. Tail length (regener-
ated at the tip) 102.1 mm. Head length 16.1 mm; head width 15.1 mm; head depth 10.6 mm; snout length 6.4 mm
(from anterior border of the orbit to tip of snout), horizontal diameter of the orbit 4.7 mm. Arm length 28.4 mm;
tibial length 17.5 mm; foot length 25.3 mm.
Upper head scales smooth, convex, bulged, pitted with scale organs in postrostral, internasals, frontonasals,
and prefrontals. Rostral flat, two times wide as long (3.3 x 1.5 mm). Two postrostrals, different in shape and size
each other, wider than long, with three and six conspicuous scale organs each; rostral and nasal uncontacted, sepa-
rated by the anterior lorilabial scale and postrostrals. Nasal scales almost rounded (1.3 x 1.5 mm). Nostril rounded,
Zootaxa 2924 © 2011 Magnolia Press · 7
TWO NEW SPECIES OF PHYMATURUS FROM PATAGONIA
occupying almost all the nasal scale. Nasal scales in contact with eight-seven scales. Internasal, frontonasals, pre-
frontals, frontals, frontoparietals, parietals, supraoculars, and circumorbitals small, irregular in shape, almost indis-
tinguishable in size and shape from each other. Interparietal pentagonal, only distinguishable by a large and
conspicuous white cream “eye” in the middle, occupying almost half part of the scale. Twenty two dorsal head
scales between rostral and nuchals. Two scales between nasal and first canthal. First canthal small, higher than
wide. Posterior canthal larger, longer than wide. Posterior canthal slightly overlap first supercilliary. Supercilliaries
6–7 (left-right), on left side first three overlapped, an injury gap, and three overlapped; on right side all moderately
overlapped. Loreal region flat, 6 irregular scales on each side. Upper ciliary scales in two rows, those of inner rows
flat and quadrangular, those of outer row granular and compressed. Lower and upper ciliaries similar in size and
shape. Palpebral scales small, irregular, slightly granular. One preocular, small, square; one elongate subocular (5.2
x 0.9 mm), unfragmented, one small postocular; a well marked longitudinal ridge along upper margin of subocular
scale, less marked in preocular, and not marked in postocular. Two rows of lorilabials becoming only one below
subocular. Lorilabials convex, 9/8–8/8, rectangular, slightly narrow than supralabials, pitted with conspicuous and
numerous scale organs. Supralabials 10–11, flat. Temporal scales conical, smooth, swollen, juxtaposed, with a
scale organ at the tip. Auditory meatus higher than wide (4.4 x 2.0 mm) with posterior edge surrounded by granular
scales. Mental pentagonal, slightly wider than long (2.1 x 2.0) narrow than the rostral scale; in contact with anterior
infralabial and postmental but not in contact with anterior sublabials. Infralabials 8–7. Chinshields 8–7, irregular,
first 4–5 slightly quadrangular, separated from infralabials by series of 1-2-3-4 irregular, first equal in size but
becoming smaller to back. Only a few scale organs present in supralabials and infralabials. Gular scales round, flat,
and juxtaposed. Seventy-one between auditory meata. Lateral postauricular folds moderately developed, with gran-
ular scales over longitudinal fold. Ante-humeral pocket well developed. Seventy-one scales between auditory
meatus and scapula. In ventral view, gular fold not well developed with its anterior margins endowed with enlarged
scales on their borders and posterior gular fold present.
Dorsal body scales rounded, smooth, juxtaposed. Forty-nine dorsal scales along midline of the trunk in a dis-
tance equivalent to head length. Scales around midbody 221. Mid-dorsal scales not enlarged in comparison to those
on flanks, both rounded and juxtaposed. There are 45 dorsal scales between occiput an anterior margin of hind-
limb articulations. Ventral scales larger that dorsals, rhombals and juxtaposed. Ventral scales between mental and
precloacal pores 178. Scales of the cloacal apron smaller than body scales, flat, rounded, juxtaposed. Precloacal
pores 12. Supra-brachial and ante-brachial scales smooth, slightly imbricated, with rounded posterior margins,
larger than dorsal body scales. Supracarpals laminar, round, smooth. Supradigital lamellae convex, imbricate.
Infra-brachial and ante-brachial scales smooth, with rounded posterior margins, larger that dorsal body scales, Sub-
digital lamellae with 3–5 keels (more conspicuous in proximal lamellae). Subdigital lamellae numbering I: 13;
II: 18; III: 21; IV: 23; V: 17. Claws moderately long. Infracarpals with round margins and 1–2 obtuse keels, distally
mucronate. Supra-femorals smooth, slightly imbricate, rhomboidal to rounded, few with a blunt keel. Infra-femo-
rals slightly larger and imbricate, smooth, rhomboidal. Supra-tarsals laminar, smooth, with round posterior mar-
gins. Post-femoral scales smaller and granular. Supra-tibials smooth, a few with a blunt keel, imbricate, becoming
rounded distally. Infra-tibials larger than infra-femorals, smooth, imbricate. Infra-tarsals with round margins and 3
obtuse keels, distally mucronate. Sub-digital lamellae numbering I: 13; II: 19; III: 23; IV: 28; V: 21.
Caudal scales arranged in spinose annuli, scales larger than body and limbs scales, slightly keeled, imbricated,
out-projecting.
Color in life. Dorsal coloration of head, body, limbs, and tail gray-brown, speckled with small white spots uni-
formly distributed (the dots cover mostly one scale, sometimes two of them) on a brown background. The throat,
forelimbs, and margins of chest and venter gray coloured; nevertheless, central areas of chest, abdomen, cloacal
apron, ventral sufaces of femoral and tibial region of hindlimbs, and foot, mustard.
Color in preservative. Dorsal background became a faded gray-brown with white spots uniformly distributed
on dorsal areas of the head, body, limbs, and tail. General coloration of ventral surfaces (throat, limbs, and tail)
became yellowish with a slight mustard color in the posterior region of abdomen continuous over the cloaca and
thighs.
Va ri at io n . Based on five adult males (Table 3, Figure 3): SVL 81.5–90.7 mm. Axilla groin distance 41.9–49.8
mm. Foot length 22.3–24.5 mm. Tibial length 15.8–17.5 mm. Arm length 23.0–28.5 mm. Head length 13.6–15.6
mm. Head width 13.75–14.8 mm. Head height 8.4–9.6 mm. Midbody scales 210–238. Dorsal scales in a head
length 42–50. Ventral scales 160–178. Supralabials 8–11. Infralabials 5–8. Scales around nasal 6–8. Third finger
AVILA ET AL.
8 · Zootaxa 2924 © 2011 Magnolia Press
lamellae 20–22. Fourth toe lamellae 27–30. Precloacal pores 7–12. In five adult females (Table 3, Figure 3): SVL
83.0–92.5 mm. Axilla-groin distance 43.9–51.1 mm. Foot length 21.95–24.55 mm. Tibial length 15.0–17.2 mm.
Arm length 25.1–27.5 mm. Head length 13.70–14.5 mm. Head width 12.9–13.9 mm. Head height 7.9–8.8 mm.
Midbody scales 210–229. Dorsal scales in a head length 38–47. Ventral scales 166–190. Supralabials 8–12. Infrala-
bials 6–8. Third finger lamellae 21–23. Fourth toe lamellae 28–32. We found just two lizards with five scales con-
tacted to the mental shield. Head scales becoming smaller near the occiput, anterior scales are hexagonal or
irregular in shape. Most individuals have convex scales in nuchal areas but in others these are smooth.
FIGURE 3. Dorsal color variation in the type series of Phymaturus sitesi.
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TWO NEW SPECIES OF PHYMATURUS FROM PATAGONIA
TABLE 3. Morphometric and meristic variation in 10 individuals of the Phymaturus sitesi type series. Means and standard
deviations (SD) of the main morphometric and meristic characters. Measures in mm and scales in numbers. SVL=Snouth-Vent
Length.
Dorsal background varies between individuals in intensity of color with some becoming darker than others.
White spots occupy more than 1 or 2 scales in some individuals, but usually with 1-2-3 scales more brightly than
others. Lateral areas in two females became irregularly dark by fusion of black scales. Cloacal apron slightly larger
in males with yellow coloration extended to infrafemoral scales, and very visible orange precloacal pores, unlike
females. Throat varies from plain gray to a subtle reticulated pattern.
Etymology. The species is dedicated to honor our friend and colleague Jack Walter Sites, Jr. an herpetologist
and evolutionary biologist from Brigham Young University, Provo, Utah, USA, who strongly supported and collab-
orated with us on studies of Patagonian herpetofauna for the last 10 years.
Geographic distribution. Phymaturus sitesi was only collected in rocky outcrops above 1800 m in the Sierra
of Auca Mahuida, an ancient volcanic range of northern Patagonia, Neuquén Province, Argentina (Figure 4, 5).
Sierra of Auca Mahuida is the southernmost mountain of the Retro Arc belt that borders the Payunia region. This
mountain complex comprised several volcanic cones, but the Auca Mahuida mountain is the main and more con-
spicuous geographic feature; is completely isolated from other similar mountains or Patagonian Phytogeographic
Province formations with Phymaturus species, by the valley of Colorado River by the north and by lowlands and
sand dunes fields of the Añelo basin in the west, south and southeast, with a gentle slope to Monte formation on the
northeast. It is like an island of Patagonian-like environment in the middle of Monte region and the new species
seems to be restricted only to habitats with Patagonian Phytogeographic Province vegetation above 1800 m.
Natural history. Little information about natural history and biology of this new species is available. Phyma-
turus sitesi was found in rocky outcrops above 1800 m, usually surrounded by vegetation characteristic of the
Payunia Phytogeographic District, Patagonian Phytogeographic Province, characterized by shrubs as Senna arnot-
tiana, S. kurtzi, Mulinum spinosum, and several species of grasses (Stipa spp.). The holotype and paratypes were
found by active search, basking on rocky outcrops or hiding in crevices. Activity depends on the season and daily
climate conditions, but lizards are usually spotted between 09:00 to 19:30. In summer they adopt a bimodal activity
pattern, and in early and late activity seasons they are unimodal. Phymaturus sitesi shares its habitat with Phymatu-
rus aff. roigorum (Phymaturus sp. 8, Morando et al., in review), a larger species of the palluma group, and other
potentially new species belonging to the bibronii, elongatus, and boulengeri groups of Liolaemus, as well as two
leiosaurids, Diplolaemus leopardinus, Leiosaurus bellii and a phyllodactylid, Homonota darwinii. Two snakes Rhi-
nocerophis ammodytoides and Pseudotomodon trigonatus were spotted in the type locality. The new species usu-
Males (N= 5) Females (N= 5)
Mean SD Mean SD
SVL 84.56 3.76 87.28 3.58
Axilla-groin distance 44.39 3.13 46.64 3.05
Head length 14.43 0.79 14.13 0.32
Head width 14.47 0.42 13.56 0.40
Head high 9.05 0.51 8.44 0.40
Foot length 23.17 0.92 23.36 0.96
Tibial length 16.52 0.67 16.17 0.80
Arm length 25.52 2.09 26.34 0.90
Midbody scales 222.60 10.38 218 8.27
Dorsal scales 45.80 3.49 42.80 3.27
Ventral scales 172.20 8.01 180.20 9.70
Fourth toe lamellae 28.20 1.09 29 1.73
Supralabial scales 9.60 1.34 9.4 1.51
Infralabial scales 7.20 1.30 7.4 0.89
Cloacal pores 8.75 2.21 - -
AVILA ET AL.
10 · Zootaxa 2924 © 2011 Magnolia Press
ally occupies microhabitats on the rocky outcrops, while the other species were found in rocky patches or open
substrate. Only Liolaemus aff. elongatus and P. aff. roigorum were observed sharing the same microhabitat of P.
sitesi but P. aff. roigorum is most common in larger outcrops with larger crevices than P. sitesi. In some areas, usu-
ally temporary creeks found between lava fields or disturbed sites along road sides, all lizard species are observed
sharing same microhabitats. No data about reproduction, diet or other natural history characteristics are available,
but as in other related species of Phymaturus, P. sitesi is viviparous and feeds on plant matter, and possibly some
arthropods.
FIGURE 4. Map of northern Neuquén and southern Mendoza region showing the complex landscape of northwestern Patago-
nia. White circle mark Phymaturus sitesi type locality in Sierra de Auca Mahuida mountain and black circle mark Phymaturus
delheyi type locality in Butacó Creek (see below), Tromen massif. Distribution of closest relatives Phymaturus nevadoi, P.
payuniae and P. zapalensis are outlined with white broken lines. Northernmost distribution limits of the Phymaturus patagoni-
cus species group in Rio Negro Province is outlined with red broken lines. Main mountains range, rivers, cities, and roads are
marked for reference. Inset: Region in South America.
Phymaturus delheyi sp. nov.
(Figures 6, 7)
Type material. Holotype: MLP.S 2609, adult male collected on rocky environments of the northern Tromen Vol-
cano massif, along Butacó Creek, on Provincial Road 37 (36º 59’ S, 69º 59’ W, 1810 m, datum = WGS 84), Pehu-
enches Department, Neuquén Province, Argentina; L.J. Avila, D.R. Pérez, and C.H.F. Pérez, collectors.
Paratypes: LJAMM-CNP 7655, MLP.S 2610 males, LJAMM-CNP 7657-7659, females. Same data as holo-
type. L.J. Avila, same as above collectors. LJAMM-CNP 5220, MLP.S 2611, females. Same data as holotype; L.J.
Avila, C.H.F. Pérez, K. Dittmar, M. Morando, and J.W. Sites, Jr., collectors.
Zootaxa 2924 © 2011 Magnolia Press · 11
TWO NEW SPECIES OF PHYMATURUS FROM PATAGONIA
FIGURE 5. Type locality of Phymaturus sitesi. Upper: general view of the area. Below: close view of the common outcrops
where lizards were collected u observed basking or hiding in crevices.
Diagnosis. Phymaturus delheyi is a robust and medium sized member of the clade of Phymaturus lizards
referred to as the patagonicus group by Etheridge (1995) because it has flat imbricate superciliaries, non-rugose
dorsal scales on tail, and subocular usually not fragmented. Phymaturus delheyi can be distinguished from other
species of the patagonicus group (P. agilis, P. calcogaster, P. castillensis, P. ceii, P. des ue tu s, P. etheridgei, P. excel-
sus, P. felixi, P. indistinctus, P. m an uela e, P. patagonicus, P. somuncurensis, P. spurcus, P. spectabilis, P. vide lai , P.
tenebrosus) by colour pattern features and geographical distribution.
AVILA ET AL.
12 · Zootaxa 2924 © 2011 Magnolia Press
FIGURE 6. Phymaturus delheyi, holotype adult male in dorsal and ventral view (MLP.S 2609), from Butacó Creek, Tromen
massif, Pehuenches Department, Neuquén Province, Argentina.
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TWO NEW SPECIES OF PHYMATURUS FROM PATAGONIA
FIGURE 7. Phymaturus delheyi, adult female in dorsal and ventral view, from Butacó Creek, Pehuenches Department, Neu-
quén Province, Argentina.
AVILA ET AL.
14 · Zootaxa 2924 © 2011 Magnolia Press
FIGURE 8. Dorsal color variation in the type series of Phymaturus delheyi.
This new species is allopatric and differs from all other members of the clade in its unique dorsal pattern of
medium size white spots covering 1–10 scales on a dark-brown background. The presence of sexual dichromatism
differentiates Phymaturus delheyi from Phymaturus sitesi sp. nov. and P. nevadoi. Dorsal pattern of P. payuniae is
composed by irregular white spots (between 4–40 scales each, sometimes fused) scattered along head and trunk,
and becoming enlarged on tail to almost reticulated on limbs, a pattern never observed in P. delheyi. Ventral color-
ation in P. payuniae is clearer than in P. delheyi. Phymaturus delheyi has a pink tinge on the chest and belly, becom-
ing an orange coloration on the lower abdomen, and cloacal and femoral regions not observed in P. payuniae.
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TWO NEW SPECIES OF PHYMATURUS FROM PATAGONIA
Scales around midbody are higher in P. payuniae than in P. delheyi and show little overlap (221–234 vs 198–227).
Dorsal pattern of P. zapalensis is composed by irregular white spots (between 5–14 scales each, sometimes fused)
scattered along head and trunk but not in limbs and tail, white spots become smaller in lateral areas, and then lateral
dark areas turn into dark bands between shoulder and rump, a pattern never observed in P. delheyi. Ventral color-
ation in P. zapalensis is greyish with a dark reticulated only in the gular region, pattern never observed in P. del-
heyi. Phymaturus nevadoi has a dorsal pattern with spots usually occupying fewer scales (4–9 scales vs 1–10),
greater density, and grow to be larger on flanks (not in P. delheyi). Phymaturus delheyi shows a higher ventral scale
count (174–202 vs 162–179 in P. nevadoi). Phymaturus delheyi can be distinguished from Phymaturus sitesi new
sp. by the dorsal pattern described above. Scales around midbody are fewer in P. delheyi than in P. sitesi (198–227
vs 210–229), and higher in ventral scales with some overlap (174–202 vs 160–190).
Description of the holotype. Adult male 89.5 mm (SVL). Axilla-groin distance 47.7 mm. Tail length 105.1
mm. Head length 15.7 mm; head width 15.0 mm; head depth 10.5 mm; snout length 6.6 mm, horizontal diameter of
the orbit 4.1 mm. Arm length 28.6 mm; tibial length 18.8 mm; foot length 25.9 mm.
Upper head scales smooth, convex, bulged, pitted with scale organs, in postrostral, internasals, frontonasals,
and prefrontals. Rostral flat, two times wide as high (3.24 x 1.48 mm). Four postrostrals, two on each side, small
and triangular, central scales different in shape and not similar in size each other, irregular, wider than high, with 5
and 11 conspicuous scale organs each; rostral and nasal uncontacted, separated by the anterior lorilabial scale and
postrostrals. Nasal scales almost rounded (1.52 x 1.28 mm). Nostril elongated and in angle, occupying almost all
the nasal scale. Nasal scales in contact with 9–10 scales. Internasal, frontonasals, prefrontals, frontals, frontopari-
etals, parietals, supraoculars, and circumorbitals small, irregular in shape, very homogenous in size and shape from
each other. Interparietal rhombal, only distinguishable by a large and conspicuous cream-white “eye” in the middle,
located on the half anterior part of the scale. Twenty three dorsal head scales between rostral and nuchals. Two
scales between nasal and first canthal. First canthal small, wider than long. Posterior canthal larger, longer than
wide. Posterior canthal slightly overlapping first supercilliary. Supercilliaries 6-6 (left-right); both sides all moder-
ately overlapped. Loreal region flat, 5 irregular scales on each side. Upper ciliary scales in two rows, those of inner
rows flat and quadrangular, those of outer row granular and compressed. Lower and upper ciliaries similar in shape
but lower ciliares larger. Palpebral scales small, irregular, slightly granular. One preocular, small, rectangular; one
elongate subocular (5.0 x 0.68 mm), unfragmented, one small postocular; a very evident keel in subocular, well
marked in preocular, and less marked in postocular. Two rows of lorilabials becoming only one below subocular.
Lorilabials convex, 9/9-9/7, rectangular, slightly narrow than supralabials, pitted with conspicuous and numerous
scale organs. Supralabials 10-8, flat. Temporal scales conical, smooth, swollen, juxtaposed, with a scale organ in
the tip. Auditory meatus higher than wide (4.3 x 1.9 mm) posteriorly surrounded by granular scales. Mental pentag-
onal, wider than high (2.7 x 1.9 mm); in contact with anterior infra-labial and post-mental but not in contact with
anterior sub-labials. Infralabials 7-7. Chinshields 7-8, irregular, first 4–5 slightly pentagonal, separated from infra-
labials by series of 1-2-3 irregular scales, first equal in size but becoming smaller posteriorly. Only a few scale
organs present in supra-labials and infra-labials. Gular scales round, flat, and juxtaposed. Fifty-nine between audi-
tory meata. Lateral post-auricular folds moderately developed, with granular scales over longitudinal fold. Ante-
humeral pocket well developed. Seventy-five scales between auditory meatus and shoulder. In ventral view, gular
fold not well developed at their anterior margins, but with enlarged scales on their borders and posterior gular fold
present.
Dorsal body scales rounded, smooth, juxtaposed. Forty-three dorsal scales along midline of the trunk in a dis-
tance equivalent to head length. Scales around midbody 198. Mid-dorsal scales not enlarged in comparison to those
on flanks, both rounded and juxtaposed. Dorsal scales between occiput an anterior margin of hind-limb articula-
tions 45. Ventral scales larger that dorsals, rhomboidal and juxtaposed. Ventral scales between mental and precloa-
cal pores 175. Scales of the cloacal apron smaller than body scales, flat, rhomboidal, juxtaposed. Precloacal pores
7. Supra-brachial scales rhomboidal, smooth, imbricated and ante-brachial scales smooth, slightly imbricated, with
rounded posterior margins, larger than dorsal body scales. Supra-carpals laminar, round, smooth. Supra-digital
lamellae convex, imbricate. Infra-brachial and ante-brachial scales smooth, with rounded posterior margins, larger
that dorsal body scales, Sub-digital lamellae with 3–5 keels (more conspicuous in proximal lamellae). Sub-digital
lamellae numbering I: 12; II: 18; III: 22; IV: 25; V: 18. Claws moderately long. Infra-carpals with round margins
and 2–3 obtuse keels, distally with a mucron. Supra-femorals smooth, imbricate, rhomboidal to rounded, the major-
ity with a pointed keel. Infra-femorals slightly larger and imbricate, smooth, rhomboidal. Supra-tarsals rhombals,
AVILA ET AL.
16 · Zootaxa 2924 © 2011 Magnolia Press
some with a moderate keel, slightly mucronated. Post-femoral scales smaller and rhomboidal, some pointed.
Supra-tibials imbricate, almost all with a pointed small keel. Infra-tibials larger than infra-femorals, smooth, imbri-
cate. Infra-tarsals with round margins and 3 obtuse keels, distally mucronate. Sub-digital lamellae numbering I: 15;
II: 20; III: 25; IV: 32; V: 23.
Caudal scales arranged in spinose annuli, scales larger than body and limbs scales, slightly keeled, imbricated,
out-projecting.
Color in life. Dorsal coloration of head, body and limbs speckled with white spots uniformly distributed (occu-
pying 1 scale in the head, 1–10 body scales, and 2–4 on limbs) in a darkbrown background. Tail brown, speckled
with white spots uniformly distributed and occupying 1–4 scales. In ventral view, throat, forelimbs, and margins of
chest with black reticulations on a gray venter; central areas of chest, abdomen, cloacal apron, ventral surfaces of
femoral and tibial region of hindlimbs, and foot mustard, but becoming lighter on the chest.
Color in preservative. Dorsal background became a faded gray-brown with white spots uniformly distributed
on dorsal areas of the head, body, limbs, and tail. General coloration of ventral surfaces (throat, limbs, and tail)
become yellowish with a slight mustard color in the posterior region of abdomen, and continuous over the cloaca
and thighs.
Va ri at io n . Based on three adult males (Table 4, Figure 8); SVL 87.1–89.6 mm. Axilla groin distance 44.7–47.2
mm. Foot length 25.2–28.0 mm. Tibial length 17.1–18.8 mm. Arm length 28.3–29.3 mm. Head length 14.9–15.7
mm. Head width 14.8–15.0 mm. Head high 9.6–10.6 mm. Midbody scales 198–222. Dorsal scales in a head length
43–44. Ventral scales 174–184. Supralabials 9–10. Infralabials 7–9. Scales around nasal 7–8. Third finger lamellae
21–23. Fourth toe lamellae 29–32. Precloacal pores 7–8. In five adult females (Table 4, Figure 8): SVL 78.0–93.7
mm. Axilla-groin distance 45.4–52.4 mm. Foot length 22.5–26.5 mm. Tibial length 15.3–17.8 mm. Arm length
24.2–30.1 mm. Head length 12.7–15.7 mm. Head width 12.3–14.8 mm. Head high 7.7–9.6 mm. Midbody scales
202–227. Dorsal scales in a head length 39–48. Ventral scales 189–202. Supralabials 7–10. Infralabials 6–7. Scales
around nasal 8–9. Third finger lamellae 21–25. Fourth toe lamellae 30–34.
TABLE 4. Morphometric and meristic variation in 8 individuals of the Phymaturus delheyi type series. Means and standard
deviations (SD) of the main morphometric and meristic characters. Measures in mm and scales in numbers. SVL=Snouth-Vent
Length.
Etymology. The species is dedicated to honor our friend Kaspar Delhey from Max Planck Institute for Orni-
thology, Germany, an Argentinean ornithologist and evolutionary biologist who helped to collected this and other
species still to be described from northern Patagonia.
Males (N= 3) Females (N= 5)
Mean SD Mean SD
SVL 88.72 1.38 89.50 6.76
Axilla-groin distance 46.11 1.50 49.86 3.22
Head length 15.45 0.46 14.44 1.08
Head width 14.9 0.13 13.76 0.93
Head high 9.99 0.51 8.67 0.85
Foot length 26.38 1.44 24.62 1.62
Tibial length 18.16 0.90 16.66 0.89
Arm length 28.73 0.48 27.42 2.09
Midbody scales 209.33 12.05 210 9.92
Dorsal scales 43.33 0.57 41.60 3.78
Ventral scales 177.66 5.50 194.20 5.26
Fourth toe lamellae 29.33 0.57 31.80 1.78
Supralabial scales 9.33 0.57 8.60 1.14
Infralabial scales 7.66 1.15 6.80 0.44
Cloacal pores 7.66 0.57 - -
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TWO NEW SPECIES OF PHYMATURUS FROM PATAGONIA
FIGURE 9. Type locality of Phymaturus delheyi. Upper: general view of the area. Below: close view of the common outcrops
where lizards were collected u observed basking or hiding in crevices.
AVILA ET AL.
18 · Zootaxa 2924 © 2011 Magnolia Press
Geographic distribution. Phymaturus delheyi was collected only in the type locality, along volcanic rims sur-
rounding the Butacó Creek, in the north part of the Tromen Massif, northern Patagonia, Pehuenches Department,
Neuquén Province, Argentina (Figure 4, 9).
Natural history. Little information about natural history and biology of this new species is available. Region
were Phymaturus delheyi was found is know as the Tromen Massif, a volcanic area were the most important volca-
noes are Tromen, Cerro Negro del Tromen and Wayle (the Punmahuida volcanic group). Vegetation is characteris-
tic of the Payunia Phytogeographic District, and including shrubs such as Senna arnottiana, S. kurtzi, Mulinum
spinosum, and several species of grasses (Stipa spp). The holotype and paratypes were found by active search; usu-
ally they were spotted basking on the rocky outcrops or hiding in crevices. Phymaturus delheyi shares its habitat
with potentially new species of the Phymaturus palluma group (P. aff. roigorum -Phymaturus sp. 10, Morando et
al., in review-), and other potentially new species belonging to the Liolaemus elongatus group, Liolaemus buergeri
and Homonota darwinii. No snakes were found in the area. The new species usually occupies the crevice micro-
habitat in the rocky outcrops, whereas the other species were found in rocky patches or open substrate. Phymaturus
delheyi is viviparous as other species of Phymaturus; a female kept in captivity gave birth two offspring, and was
observed feeding on plant matter and arthropods (Perez pers. observ.).
Discussion
The patagonicus group of Phymaturus as was defined by Etheridge (1995) is well supported by mtDNA sequences
(Morando 2004), but not clearly supported in a study using morphological characters (Lobo & Quinteros 2005). A
new molecular study based on mtDNA and multiple nuclear genes (Morando et al. in review) supports the
Etheridge (1995) division of Phymaturus into two clades. Lobo and Quinteros (2005) found that relationships
between species of this group are poorly supported and suggest that lot of work is still necessary in this group.
These authors also suggest that the radiation of this group was rapid and more recent than in the other Phymaturus
clade (the palluma group), and was characterized by too little time for morphological differentiation. Thus, mor-
phology alone must be used with caution to delimit new species and to reconstruct phylogenetic relationships in
this group.
Some areas of southwestern Rio Negro province (Central Patagonia) show a very complex and still poorly
studied set of lizard populations with extensive morphological polymorphism. Some of the species described in the
last few years were based on this set of populations, usually distributed along a same rim of a volcanic plateau, and
we suggest that caution be taken to avoid over-splitting of taxa without evidence from several sources (i.e., integra-
tive taxonomy, Padial et al. 2010). As an example, Lobo et al. (2010) recently suggested that P. agilis could be a
synonyn of P. spectabilis and in the same work they claimed that P. des ue t us could represent another case of poly-
morphism, suggesting that this clade may already be over-split in species numbers. The new species described here
are from a very geographically distant area, where all plateaus and mountain peaks are not connected between
them, thus complete allopatry is clearer than in the Rio Negro area. In the past 15 years knowledge of the diversity
of this interesting genus has increased rapidly; from the 10 species recognized by Etheridge (1995) to 35 today,
with 21 described species for the patagonicus group.
Acknowledgements
We thank M.L. Kozykariski, M.F. Breitman, K. Dittmar, J.W. Sites, C. Medina, N. Frutos, A. Cosacov, M. Nicola,
K. Delhey, M. Olave, P. Petracci, M. Hawkins, I. Minoli, R. Otteson, J.R. Goldman, C. Zanotti for help and com-
panion in field trip collections of Phymaturus species. We thank Fauna and Flora and Protected Areas authorities
(S. Di Martino and M. Aubone, M. Monteverde) of Neuquén Province for collection permits and support of our
work. Park rangers S. Goitia and F. Quiles kindly support us in the field. We thank N. Cazzaniga for help with Latin
language issues. Financial support for field work was provided by NSF “Partnership for International Research and
Education” award (OISE 0530267). Additional support was provided by an ANPCYT-FONCYT 06-00506 issued
to L. Avila and grants from Brigham Young University (the Department of Biology, M. L. Bean Museum, and Ken-
nedy Center for International Studies) issued to J. W. Sites, Jr. YPF S.A. Petroleum Company supported our
research in Auca Mahuida mountain range through a grant issued to D.R. Perez.
Zootaxa 2924 © 2011 Magnolia Press · 19
TWO NEW SPECIES OF PHYMATURUS FROM PATAGONIA
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20 · Zootaxa 2924 © 2011 Magnolia Press
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APPENDIX I
Phymaturus delheyi (8): ARGENTINA: NEUQUEN: Chos Malal Department: Northern Tromen Volcano massif, Butacó
Creek, on Provincial Road 37 (36º59’S, 69º59’W, 1810 m): MLP.S 2609 (Holotype), MLP.S 2610-2611, LJAMM-CNP
7655, 7657-7659 and 5220 (Paratypes).
Phymaturus sitesi (20): ARGENTINA: NEUQUEN: Pehuenches Department: Auca Mahuida mountain near Cerro de las
Antenas, Auca Mahuida Natural Protected Area (37º43’S, 68º55’W, 1983 m), MLP.S 2605 (Holotype), MLP.S 2606-2608,
BYU 12590-12591, LJAMM-CNP 10367-10368, 10466-10469 (Paratypes); Riscos Altos, 38.6 km S junction Provincial
Road 6, Auca Mahuida Natural Protected Area (37º 43’ S, 68º 55’ W, 1851 m): LJAMM-CNP 12297-12298 (Paratypes).
Park Ranger Post, 38.6 km S junction Provincial Road 6, Auca Mahuida Natural Protected Area (37º 42’ S, 68º 51’ W,
1560 m), LJAMM-CNP 12157, 12189, 12214 (Paratypes); road SW Park Ranger Post, Auca Mahuida Natural Protected
Area (37º 42’ S, 68º 53’ W, 1659 m), LJAMM-CNP 13402 (Paratypes). Añelo Department: Auca Mahuida Natural Pro-
tected Area, Communication Station Site, South Slope, Auca Mahuida Natural Protected Area (37º 46’ S, 68º 53’ W, 1935
m), LJAMM-CNP 12311-12312 (Paratypes).
Phymaturus nevadoi (9): ARGENTINA: MENDOZA: Malargüe Department: Provincial Road 180, 22.9 km N junction Pro-
vincial Road 186, 3.1 km S La Ventana post (35º55’S, 68º36’W, 1888 m): LJAMM-CNP 4431-4433. Provincial Road 186,
25.5 km NE Mina Ethel, 4 km S Marfil Post (35º55’S, 68º36’W, 1873 m): LJAMM-CNP 7933-7938.
Phymaturus payuniae (6): ARGENTINA: MENDOZA: Malargüe Department: unnamed road, 2.8 km E junction Provincial
Road 183, 13.8 km E road to El Clavado Post, Payun Plateau, (36º39’S, 69º16’W, 1731 m): LJAMM-CNP 4436-4438.
Provincial Road 183, piedmont of Volcan Payun Liso, Malargüe Department, Mendoza Province, Argentina (36º29’S,
69º22’W, 2134 m): LJAMM-CNP 7975-7977.
Phymaturus zapalensis (9): ARGENTINA: NEUQUEN: Catan Lil Department: Provincial Road 46, 9.5 km SW gate Laguna
Blanca National Park (39º08’S, 70º25’W, 1401 m): LJAMM-CNP 8067-8074. Unpaved road, 2 km W National Road 40,
18,3 km SW junction Provincial Road 24, near El Salitral (39º49’S, 70º38’W, 1139 m): LJAMM-CNP 8916.
Phymaturus ceii (11): ARGENTINA: RIO NEGRO: 25 de Mayo Department: Provincial Road 8, 17 km S Antonio del Cuy
(40º17’S, 68º27’W): LJAMM-CNP 1628-1630, 1713, and MCN 908, MCN 910-913, MCN 916, MCN 918.
Phymaturus excelsus (15): ARGENTINA: RIO NEGRO: Ñorquinco Department: Provincial Road 6,1 km NW Ojo de Agua
(41º32’S, 69º51’W, 1141 m): LJAMM-CNP 2136-2137, 2266, 2355-6, 2652-2654. Provincial Road 6, 2.2 km NE Ojos de
Agua, (41º32’S, 69º51’W, 1105 m): LJAMM-CNP 3534-3535, 3622-3626.
Phymaturus spectabilis (22): ARGENTINA: RIO NEGRO: Ñorquinco Department: Provincial Road 6, 17.4 km NE Ojos de
Agua, 28 km SW Ingeniero Jacobacci, (41º26’S, 69º46’W, 1014 m): LJAMM-CNP 3600-3621.
Phymaturus spurcus (10): ARGENTINA: RIO NEGRO: Ñorquinco Department: National Road 1s40, 2.5 km N de Chen-
queniyen, volcanic rim 0.5 km N Chenqueniyen post (41º33’S, 70º40’W, 1123 m): LJAMM-CNP 3504-3508. Provincial
Road 6, 2.2 km NE Ojos de Agua (41º32’S, 69º51’W, 1105 m): LJAMM-CNP 3536, 3627. 25 de Mayo Department:
National Road 23, hill near Ranch Huanuluan, 25.1 km W Ingeniero Jacobacci (41º21’S, 69º48’W, 942 m): LJAMM-CNP
3586, 3629-3630.
Phymaturus somuncurensis (12): ARGENTINA: RIO NEGRO: Valcheta Department: Somuncurá Plateau (41º11’S, 66º53’W,
1002 m): LJAMM-CNP 4453-4456. Corona Chico Hill (41°31' S, 67°00' W, 1420 m): LJAMM-CNP 6022. ARGEN-
TINA: RIO NEGRO: 9 de Julio Department: 65.6 km police station El Rincon, near Corona Hill, between Corona Grande
Hill and Corona Chico Hill (41°23’S, 66°57’W, 1425 m): LJAMM-CNP 6826-6832.
Phymaturus tenebrosus (3): ARGENTINA: RIO NEGRO: Pilcaniyeu Department: National Road 40, 2,7 km S San Pedro
Ranch (40º52’S, 70º34’W, 1096 m): LJAMM-CNP 5426, BYU 47978. National Road 40, Cerro Alto (40º46’S, 70º34’W,
1203 m): LJAMM-CNP 8661.
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TWO NEW SPECIES OF PHYMATURUS FROM PATAGONIA
Phymaturus calcogaster (11): ARGENTINA: CHUBUT: Telsen Department: Crossroads to De la Vaca Lagoon, 16.2 km Pro-
vincial Road 4, near edge of the lagoon, from Elio Calfuquir’s post (42º30’S, 67º21’W, 660 m): LJAMM-CNP 6855-6857,
8125-8132.
Phymaturus felixi (15): ARGENTINA: CHUBUT: Paso de Indios Department: Provincial Road 24, 110 km S (44º31’S,
6911’W, 560 m): LJAMM-CNP 3717, 3823-3830, 3832-3837.
Phymaturus indistinctus (5): ARGENTINA: CHUBUT: Rio Senguer Department: Provincial Road 20, San Bernardo Moun-
tains, 19 km W Los Manantiales (45º27’S, 69º42’W, 669 m): LJAMM-CNP 2124, 2238, 2273. Pampa Lehman, on the
borders of Provincial Road 20 (45°24’S, 69°52’W, 458 m): LJAMM-CNP 8198. Provincial Road 20, 4 km N junction Pro-
vincial Road 22 (45º25’S, 69º50’W, 502 m): LJAMM-CNP 2650-2651; MCN 810.
Phymaturus patagonicus (6): ARGENTINA: CHUBUT: Gaiman Department: National Road 25, 40 km WSW Dolavon
(43º27’S, 66º07’W, 125 m): LJAMM-CNP 3205-3210.
Phymaturus videlai (3): ARGENTINA: CHUBUT: Sarmiento Department: Buen Pasto, (45º04’S, 69º27’W, 652 m): LJAMM-
CNP 9084-9086.
