Zootaxa 2809: 2032 (2011)

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Copyright 2011 Magnolia Press

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Article

ZOOTAXA
ISSN 1175-5334 (online edition)

On the species identities of a complex Liolaemus fauna from the Altiplano and
Atacama Desert: insights on Liolaemus stolzmanni, L. reichei, L. jamesi pachecoi,
and L. poconchilensis (Squamata: Liolaemidae)
ROBERTO LANGSTROTH P.
Environ International, 1401 New York Ave NW, Suite 1225, Washington, DC 20005, USA. E-mail: rlangstroth@environcorp.com

Abstract
The South American lizard genus Liolaemus has undergone a complex adaptive radiation that has resulted in the evolution
of more than 200 species widely spread in an extraordinary diversity of environments, and forming a complex array of
assemblages. This evolutionary complexity has puzzled systematists and taxonomists since the first species were described more than 150 years ago. Within this lineage, the Andean Liolaemus faunas have proven to be a major challenge
for herpetologists. Therefore, intense research is needed in this area to clarify long-standing problems. After more than a
century of taxonomic confusion, the identity of Liolaemus stolzmanni (Steindachner, 1891) is here restored as the name
that must be applied to the lizards widely known as Phrynosaura (= Liolaemus) reichei Werner, 1907 from the low to midelevation deserts of Tarapac, Chile. Since 1966, the name L. stolzmanni has been erroneously assigned to populations of
Liolaemus from the high Andes of the Chile-Bolivia borderlands which, according to observations presented in this study,
correspond to Liolaemus pachecoi Laurent, 1995. A lectotype and allotype for L. stolzmanni are designated and the type
locality for L. stolzmanni (= L. reichei) is emended to Deserts of Iquique, Tarapac Region, Chile. Furthermore, the recognition of L. pachecoi as a species distinct from L. jamesi is supported by mtDNA sequence divergence data despite the
inconclusive meristic and morphometric data. In summary, I conclude that (i) the Chilean L. reichei is a synonym of L.
stolzmanni, and hence, that L. stolzmanni is a species endemic to Chile, not an element of the fauna of present-day Peru
and that (ii) the Chilean Altiplano populations currently recognized as L. stolzmanni are L. pachecoi, a species hitherto
known only from Bolivia. Also, I report the first confirmed specimens of L. poconchilensis from Peru, a species previously
known only from Chile and confused with L. reichei.
Key words: Lizards, Chile, Peru, Bolivia, Andes, taxonomy, Phrynosaura, Ctenoblepharys

Introduction
With over 220 described species, the South American lizard genus Liolaemus is the second most speciose amniote
genus, exceeded only by the iguanian genus Anolis from tropical America and the Caribbean. During their evolutionary history, Liolaemus have extensively colonized an extraordinary diversity of environments (Espinoza et al.,
2004; Pincheira-Donoso et al., 2008a) that have forced the evolution of a remarkable diversity of morphologies and
behaviors (Schulte et al., 2004; Pincheira-Donoso et al., 2009). This extraordinary diversification within the genus
has resulted in a substantial challenge for Liolaemus specialists aiming to identify and name these species and
major-level groups (Etheridge & Espinoza, 2000). Part of the long-standing nomenclatural controversies within the
genus Liolaemus has resulted from difficulties faced by modern herpetologists to access the original type series of
specimens that served as the basis for species descriptions during the XIX century.
One case of long-standing confusion involves the identities of a number of Liolaemus species widespread in
the Altiplano (High Andes Plateau) of Argentina, Bolivia, Chile and Peru, and the adjacent lower areas of the Atacama desert in northern Chile and southern Peru. In this area, several nomenclatural problems have been discussed
in recent syntheses (e.g. Etheridge & Espinoza, 2000; Valladares et al., 2002; Pincheira-Donoso et al., 2008b), and
several species have only recently been recognized as a result of historical difficulties for field explorations, isolation of areas, and climatic constraints. Therefore, much more systematic and geographical research is needed in this

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Accepted by S. Carranza: 21 Feb. 2011; published: 5 Apr. 2011

region to strengthen our understanding of Liolaemus assemblages. In this study, I concentrate on three taxa currently recognized (e.g., Etheridge & Frost, 2010) as L. stolzmanni, L. reichei, and L. jamesi pachecoi. My aim is to
provide new evidence derived from analyses of type specimens and to then suggest important rearrangements of
specific names for these species.

Methods and material

As noted above, difficult access to type specimens has long been a limiting factor in herpetological research, in particular specimens collected long ago and housed in collections distant to the researcher, such as European or North
American collections that are difficult to access by Liolaemus specialists residing in South America. However, the
advent of the digital era has created new means of access via digital photography and Internet. At present, most
researchers worldwide have access to collections by means of e-mail or directly through electronic platforms
implemented by institutions, where specimen-by-specimen catalogues are provided. For this study, I have utilized
newly obtained digital photographs of type specimens and other material housed in European, South American,
and North American institutions (See Appendix I). All studied images were taken in ideal light conditions, from
several different angles on each specimen, and with high resolution to allow detailed analyses of scale and color
variation. There are several advantages to use of high quality photographs, such as low risk of damage to or loss of
delicate, irreplaceable specimens, the ability to mark up the images and count scales without having to touch the
specimens, and the ease of sharing the images with colleagues. Another novel use of the Internet is to perform virtual lizard surveys by searching the millions of online images taken across the globe by researchers, nature enthusiasts, tourists, and others. By this method, I obtained photographs of live lizards from locations near the type
localities of the species of interest, which led to important questions and findings, including some of the only
known life habits for some taxa.
Analyses of high-resolution photographs were complemented with careful analyses of the original descriptions
of the species in question provided by the original authors, and based on additional later descriptions of the same
type material published in subsequent studies. Finally, as an additional source of information, and once again taking advantage of modern ways of communication, I consistently discussed my observations and conclusions with a
number of herpetologists who have experience in Liolaemus research. These discussions (based on appropriate presentation of the available evidence) resulted in strong consistency among researchers.

Taxonomic history of the taxa recognized as Liolaemus stolzmanni, L. reichei, and L. jamesi pachecoi
Steindachner (1891) described the lizard Ctenoblepharis Stolzmanni from three syntypes currently housed in the
Naturhistorisches Museum Wien, from the vague locality of Hoch-Peru (High Peru). According to museum
records, the C. stolzmanni syntypes are from Coll. Stolzmann 1883 and there is no other information on the locality. Stolzmann (also Sztolcman) is well known as one of the most diligent ornithological collectors to have
worked in Peru, where he collected intensively in the northern part of the country, primarily in Cajamarca, Amazonas, and Tumbes (Stephens & Traylor, 1983), areas where there are no known liolaemid lizards. There are no localities from southern Peru associated with Stolzmanns ornithological field collections (Stevens & Traylor, 1983). It
is, however, very possible that Stolzmann may have received the specimens as gifts or even purchased them from
another collector who may have collected them in the Tarapac Region, even around Iquique, which was Peruvian
territory until the aftermath of the War of the Pacific (18791884). Steindachner expressly compared his new species against a specimen cataloged as Ctenoblepharis adspersus at his museum, and clearly considered his C. stolzmanni to be congeneric with this species based on its well developed palpebral fringes.
Also, Boulenger (1891) described Ctenoblepharis jamesi from the Andes of Tarapac, Chile, from a specimen,
which bears little similarity to either of the other two species of the genus at the time (i.e., C. adspersa and C. stolzmanni). Besides stating, shape of head and general proportions same as C. adsperus, Boulenger provided no discussion as to why he considered this species comparable to C. adspersa. Note that according to Etheridge (1995:3),
Boulenger's description of Ctenoblepharis adspersus is based on a specimen from Arequipa (Arequiba), Peru,
at an altitude of 7500 ft (2286 m). Nez (2004) further commented on this specimen (BMNH 65-5-3-3), noting
that it in fact corresponds to an undescribed species of Phrynosaura.
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Werner (in Brger, 1907) subsequently described the genus Phrynosaura and the species P. reichei from a single specimen from "Iquique" (Chile). He found the new genus to be similar to the agamid Phrynocephalus and the
iguanid Ctenoblepharis, but did not provide a specific diagnosis.
Besides Mllers (1928) comments on P. reichei in his description of Phrynosaura werneri, very little was published regarding either C. stolzmanni or P. reichei until the printing of Donoso-Barros' (1966) Reptiles de Chile, in
which he provided redescriptions of both species.
Although Donoso-Barros (1966:341) considered Steindachner's description of C. stolzmanni to be "primitive",
comparative, extraordinarily limited, and barely permitting one to form an opinion of the species whose type
specimens apparently were unknown to him, he did indeed form his own opinion of what specimens might be
assignable to Steindachners description, which he ultimately found to fit a lizard collected from Quebrada del Inca
in the high Andes of the Antofagasta region. However, there are characteristics described by Steindachner which
Donoso-Barros apparently did not consider, such as C. stolzmanni having head shields larger than the largest dorsal
scales and having fringed eyelids, characteristics not applicable to the lizard he collected. He also ignored the spotted and barred patterns described by Steindachner, which contrast with the lizard described by Donoso-Barros
which has a brown dorsum with only small dark flecks caused by blackish pigmentation in the posterior of each
dorsal scale.
On the other hand, Donoso-Barros had obtained the holotype of P. reichei as part of his private collection and
he had no disparaging comments regarding Werners description, which he augmented with his own observations.
However, Werners description was no more detailed than Steindachners and was a bit clumsy due to Brgers
idiosyncratic translation, and did not include some basic elements such as the number of scales around midbody.
Donoso-Barros stated that the type had only 6 supralabials instead of the 9 reported by Werner. Also, he noted that
it had 44 scales around midbody.
During a tour of European collections, Etheridge examined the Ctenoblepharis stolzmanni syntypes in Vienna
in 1967. In his unpublished notes he wrote, "The syntypes of Ctenoblepharis stolzmanni have the characteristics of
what I have been calling Phrynosaura: a short baby bird type head with strongly projecting superciliaries, and
rather large, smooth subimbricate dorsals". He also noted that the dorsal scales are equal to or slightly smaller than
the ventral scales, there are 50 scales around midbody, and 4 precloacal pores (R.E. Etheridge, in litt.). This vision
of C. stolzmanni contrasts with that of Donoso-Barros, who evidently did not see similarities with Phrynosaura in
Steindachners description.
Donoso-Barros (1972) later provided a comparison of the type specimens of the genera Ctenoblepharis and
Phrynosaura, in which he concluded that Phrynosaura is a junior synonym of Ctenoblepharis. In this publication,
he provided new descriptive information on P. reichei based on adult specimens, as well as a detailed drawing of
the P. reichei holotype, which he considered to be a juvenile. Among the details of significance, Donoso-Barros
noted that adult P. reichei possess tricuspid lateral teeth, conditions not present in the juvenile holotype. He also
noted that P. reichei has a tripartite subocular with the middle section being the longest. He also noted that the
infradigital lamellae of the adults were much less strongly keeled than those of the juvenile. These characteristics
are also described in Steindachners description of C. stolzmanni. One interesting aspect of what appears to be a
well-proportioned and accurate drawing of the P. reichei holotype is that the dorsal scales are very small, and
would number far more than the 44 around midbody reported by Donoso-Barros (1966). It is possible that DonosoBarros based his count on the female specimen illustrated in his books photographic plates or that the drawing has
inaccurately represented the body scales.
Cei (1979) concluded that if Ctenoblepharis is to be recognized as a valid genus, it should be restricted to
include only C. adspersus, C. nigriceps, C. reichei, and C. stolzmanni. In contrast to previous authors, Cei considered C. jamesi and C. schmidti to be members of Liolaemus, a decision based on the absence of the morphological
traits which bound together his concept of Ctenoblepharis.
Five years later, Laurent (1984) revalidated Phrynosaura based on a suite of distinct morphological characteristics and found Ctenoblepharis to be a monotypic genus including only C. adsperus. As did Etheridge, Laurent
also examined the C. stolzmanni syntypes and considered them to be representative of the Phrynosaura morphotype, along with P. reichei and P. audituvelata. Laurent clearly did not find P. stolzmanni to be similar to C. aymararum, C. jamesi, or C. nigriceps, which, following Cei, he removed from Ctenoblepharis and placed in Liolaemus.
Etheridge (1995) carefully reconsidered the taxonomic history and systematics of the genus Ctenoblepharys
and other genera related to Liolaemus, and reestablished Tschudis original spelling (i.e., Ctenoblepharys
adspersa). While Etheridge did not find overriding evidence at the time to support the monophyly of Phrynosaura,

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which he synonymized with Liolaemus, he did recognize their prominent ciliary fringes and truncated snouts as
possible synapomorphies of a group containing the species L. audituvelatus, L. reichei, and L. stolzmanni.
More recently, Nez et al. (2003) described L. manueli and L. torresi (both as Phrynosaura) from different
areas of the Atacama Desert in Chile, which they found to be most similar to L. auditulevatus and L. reichei,
respectively. Liolaemus torresi was described from a series collected in the vicinity of Calama in the Loa river
basin to the southeast of Iquique and differs from L. reichei by its relatively longer tail, a dorsal color pattern consisting of more numerous and smaller spots, smaller and more numerous body scales, and more numerous supralabials. Liolaemus manueli is unquestionably phrynosauroid in aspect, similar to a female L. reichei with a stout,
roundish body, but differing significantly by its high number of small subimbricate to juxtaposed body scales, a
higher number of supralabials, and the presence of scales on the tympanum. Its distribution in the Diego de
Almagro area of the Atacama Region is far south of the known ranges of the other phrynosauroid species. In that
paper, Nez et al. clearly were aware of Laurents decision to include C. stolzmanni in Phrynosaura, but did not
comment on the relationship of this species to other Phrynosaura or their opinions of Laurents decision.
The most recently named phrynosauroid species is L. poconchilensis, described by Valladares (2004) from the
deserts of the Arica and Parinacota Region of far northern Chile (less than 15 km from the southern limit of Peru).
Specimens from populations likely belonging to this species were formerly reported as Phrynosaura reichei
(Donoso-Barros, 1969) or as Phrynosaura sp. from Arica (Nez et al., 2003). Valladares (2004) did not include L.
stolzmanni as a phrynosauroid Liolaemus in his diagnosis of L. poconchilensis.
In their revision of the Chilean species of Liolaemus, Pincheira-Donoso & Nez (2005) followed the interpretation of Donoso-Barros (1966) regarding Liolaemus stolzmanni and provided a more detailed description of this
poorly understood species, based on additional specimens collected by Nez. They included the first published
photograph of a lizard identified as Liolaemus stolzmanni, which clearly did not suggest phrynosauroid morphology, rather a larger, robust form with a thick neck and very large dorsal scales, which they clearly considered to be
related to L. jamesi. Pincheira-Donoso & Nez recognized Phrynosaura as a valid genus and did not include its
species in their 2005 monograph.
In the meanwhile, Laurent (1995) named Liolaemus jamesi pachecoi based on two specimens from Laguna
Colorada, Bolivia, a locality located 68 km SSE of the Salar de Ascotn where Pincheira-Donoso & Nez (2005)
would later report L. stolzmanni. Laurent was apparently so certain of this forms relationships that his remarkably
brief diagnosis compares it only to L. jamesi and L. aymararum. While finding that the number of dorsal scales
between the occiput and the thigh in the new form was intermediate (39-41) relative to L. aymararum (42-53) and
L. jamesi (30-38), Laurent assigned these lizards as a subspecies of jamesi due to their agreement in terms of the
relationship between the nostril-mouth distance and the minimum space between the supraoculars. However, he
also conceded that all three forms could be simply geographic variations of a single species (L. jamesi by seniority)
with no meaningful subspecies. No other characteristics of L. jamesi pachecoi were provided by Laurent and no
other authors have commented on this taxon.
While Cei (1979), Laurent (1984), and Etheridge (1995) have all commented on the clearly phrynosauroid
character of the lizards described by Steindachner as Ctenoblepharis stolzmanni, there has been no published discussion regarding the use of this name for lizards without any phrynosauroid characteristics by authors such as
Donoso-Barros (1966) and Pincheira-Donoso & Nez (2005), and others who have included L. stolzmanni in species diagnoses and lists based on Chilean material.
Abdala et al. (2008) included both L. reichei and L. stolzmanni in their diagnoses of L. huayra and L. inti, referring to MVZ specimens of each species from potentially novel localities: L. reichei from 4 mi N of Tacna, Peru,
and L. stolzmanni from Alta Camia, 110 mi NE Iquique, Chile. Photographs of these specimens clearly show the
Alta Camia specimens to be the nominal L. jamesi and not similar to L. j. pachecoi (confirmed independently by
H. Nez and D. Pincheira-Donoso, in litt.). The lizards from 4 mi N of Tacna are here identified as L. poconchilensis and noted as the first definitive record of the species for Peru. The similarity in color patterns with the photographs of the L. poconchilensis holotype and allotype in Valladares (2004) is striking (Figure 1).
Another Peruvian specimen cataloged as L. reichei, MVZ 84630, with the locality of 18 km (by road) Llupash, 10,000 ft in the Ancash Department, also generally agrees with a female or juvenile L. poconchilensis. However, the locality is over 1,200 km NW of the Tacna-Arica area and is being further investigated. The only
Liolaemus known from the Ancash region to date appear to be related to L. walkeri (e.g., KU 133861, MVZ Herps
84632), a species belonging to a clade which has very little relation to either L. reichei or L. poconchilensis.

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FIGURE 1. Liolaemus poconchilensis from 4 mi N of Tacna, Tacna, Peru. Top: MVZ 99663, male. Bottom: MVZ 99661,
female? Photographs Michelle Koo / Museum of Vertebrate Zoology, University of California.

An additional specimen previously identified by Etheridge (1995) as L. reichei, LACM 9312, is also assignable
to L. poconchilensis as it has scale counts (supralabials, scales around midbody, and middorsals between occiput
and anterior edge of thigh, based on photographs and counts provided by N. Camacho, LACM) corresponding with
the L. poconchilensis holotype, despite being a juvenile with a snout-vent length of only 32.4 mm.
Duellman, Simmons & Pfaur (unpublished field notes) collected a series of lizards in near Uchumayo, 2150
masl, just west of the city of Arequipa in 1974 and these were later cataloged in KU as Phrynosaura stolzmanni
(KU 16358994). These lizards are clearly not conspecific with the type series of L. stolzmanni based on their
smaller body scales, less conspicuous palpebral fringe, and distinctive color patterns. Also, these specimens were
collected in habitat of gray sands with granitic rocks and have cryptic coloration matching the dark sand and rock
surfaces (Figure 2). According to Duellmans field notes, the adult males have brown head, mid-dorsum, tail, and
limbs; rest of dorsum mottled orange and metallic blue; flanks pinkish tan with dark brown mottling; limbs and tail
barred with dark brown; anterior chin white; throat yellow; lateral edge of belly orange; other ventral surfaces
white. Females have brown dorsum with dark brown and tan spots; venter white. Juveniles have dorsum like
females but pattern more intense; venter pale bluish gray. Cei & Pfaur (1982) described L. insolitus from a series
of species from Incln Alto in the coastal lomas north of Mollendo, Arequipa. The type series of L. insolitus has
larger, less numerous body scales, 45-53 around midbody, precisely in the range of L. stolzmanni; however, these

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lizards differ significantly (e.g., corpulent body, stout limbs, short digits, precloacal pores in females, distinct color
patterns) from L. stolzmanni and were expressly considered distinct from C. stolzmanni and C. riechei (sic) by
Cei & Pfaur. Cei & Pfaur (1982) also discussed specimens assigned to L. insolitus from 10 km SW of Uchumayo, 2800 masl, which are the same lizards referred to by Pfaur et al. (1978) as Ctenoblepharus sp. (sic) from
the Batolito de la Caldera, 10 km SW Uchumayo, Arequipa. These are mentioned to be part of the series at KU
from Uchumayo, although the stated localities are somewhat different. These specimens are clearly not L. insolitus
and appear to correspond to the specimen from Arequiba, 7,500 ft mistaken by Boulenger (1885) for C. aspersa.
Differences mentioned by Pfaur et al. (1978) for the Uchumayo lizards relative to the Cei & Pfaur (1982)
description of L. insolitus include a shorter head relative to total length (<=1/10 TL vs. >1/10 TL in L. insolitus),
longer digits (i.e. long toes vs. short feet, which are clearly different in the photographs examined), distinct
dorsal color pattern in males (Uchumayo males have tigered pattern of black mottling on dorsum), and possibly
the absence of precloacal pores in females from Uchumayo (not evident in photos examined of KU specimens).
The identity of these lizards will be addressed in a subsequent paper but it suffices to say now that they are clearly
distinct from L. stolzmanni.

FIGURE 2. KU 163589 (Image KUDA 009595). PERU: Arequipa: Alturas de Uchumayo, 3 km SW Uchumayo, 2150 m. 01
November 1974. William E. Duellman / University of Kansas Museum of Natural History.

Schulte & Moreno-Roark (2009) presented a new molecular phylogeny for the Iguania, including 96 species of
Liolaemus, where L. aymararum (accession from its type locality) and L. stolzmanni (accession from Salar de
Ascotn) form a well-supported sister species pair nested within an unnamed clade corresponding to the montanus
series, which also included a clade identified as L. reichei in a four-way polytomy with L. chlorostictus and two
unnamed multispecies clades. However, the specimen identified as L. reichei from which the genetic material was
obtained, DBCUCH 2147 collected by H. Nez & J. Sufn, is actually from the population that has been named L.
poconchilensis from Valle de Lluta y Azapa (ca. Poconchile) (H. Nez, in litt.).
In summary, the identities of the taxa recognized as Liolaemus stolzmanni, L. reichei, and L. jamesi pachecoi,
until now, have not once been critically examined in the known literature relating to Liolaemus and have been subject to considerable confusion.

The Clarification
While working on a review of the state of the knowledge and taxonomic problems of the Liolaemus found to the
north of the Tropic of Capricorn, I came across online photographs of unusual Liolaemus from SW Bolivia in the
vicinity of the type locality of L. jamesi pachecoi, but also in the vicinity of the Salar de Ascotn from which L.
stolzmanni has been reported. These robust lizards have the very broad, large dorsal scales, which are often juxtaposed and separated by black skin (Figure 3).
No photographs or other illustrations of L. jamesi pachecoi were available in the literature, which led to the
obtainment of digital photographs of the L. jamesi pachecoi paratype, a female here designated as allotype, numbered FML 02788H from the Fundacin Miguel Lillo (Figure 4). The L. jamesi pachecoi allotype and the specimen

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identified as L. stolzmanni in Figure 16 of Pincheira-Donoso & Nez (2005) are strikingly similar. Herman Nez
(in litt.) reviewed the photographs of the Desierto de Siloli lizard and the L. jamesi pachecoi allotype and agreed
that these appear to be what he has referred to as L. stolzmanni and has collected from Ollage and the Salar de
Ascotn, suggesting that the Chilean L. stolzmanni and the Bolivian L. jamesi pachecoi might be conspecific. However, this question could only be answered by the examination of the syntypes of Ctenoblepharis stolzmanni.

FIGURE 3. Liolaemus lizard from Desierto de Siloli, Sud Lpez, Potos, Bolivia, January 2010. Photograph Edgardo Milla
Anacona.

FIGURE 4. Allotype (female) of Liolaemus jamesi pachecoi, FML 02788, Laguna Colorada (67 44O, 22 13S), 4270 m,
Prov Sud Lipez, Dto Potosi, Bolivia, 29oct89, cols. Salazar y Pacheco. Photographs Monique Halloy/Fundacin Miguel
Lillo.

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LANGSTROTH

Discussion: Comparison of L. stolzmanni and L. reichei

Upon receiving photographs of the exceptionally well-preserved L. stolzmanni type series from the Naturhistorisches Museum Wien (Figures 57), it was immediately evident that Etheridge and Laurent were correct in their
placement of C. stolzmanni within their concepts of Phrynosaura and that Donoso-Barros had misapplied the name
C. stolzmanni to his lizard from Quebrada del Inca.

FIGURE 5. First male syntype here designated as lectotype (NMW 13580:1) of Liolaemus stolzmanni (Steindachner, 1891).
Photographs Richard Gemel/Naturhistorisches Museum Wien.

Donoso-Barros (1966) described the ventral scales of P. reichei as being lanceolate anteriorly and becoming
quadrangular posteriorly and Werner mentioned that they form an angular pattern, both of which are evident in the
L. stolzmanni allotype. As noted by Werner for P. reichei, the toes of the L. stolzmanni type series are distinctly yellowish. Steindachner had noted that there were three subocular scales and while Donoso-Barros reported only single subocular in P. reichei in 1966, he later reported three suboculars in 1972. However, examination of the L.
stolzmanni type series shows the subocular to be variably entire or fragmented into two or three sections, even on
the same individual. Also, Donoso-Barros (1972) reported only conical teeth in the juvenile P. reichei holotype he
redescribed in 1966 but later reported tricuspid lateral teeth from adults.
There is some minor debate as to whether the type locality is in the immediate vicinity of the present-day city
of Iquique near the Pacific coast or whether it should amended to "Oasis de Pica", an inland locality at ca. 1200
masl and to the east of Iquique proper (Pincheira-Donoso et al., 2008b). This question is irrelevant as there is no
evidence to suggest that the coastal lizards are not conspecific with the inland lizards. Herman Nez (in litt.),
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however, has searched intensively for L. reichei around Oasis de Pica with his colleagues and have found no evidence of any Liolaemus, phrynosauroid or otherwise, only Phyllodactylus and Microlophus.

FIGURE 6. Second male syntype here designated paralectoype (NMW 13580:2) of Liolaemus stolzmanni (Steindachner,
1891). Photographs Richard Gemel/Naturhistorisches Museum Wien.

Lizards from the Oasis de Niebla de Alto Patache (20.824209S, 70.156204W; 780 masl), 60 km S of the city
of Iquique, the L. reichei type locality, are strikingly similar to the L. stolzmanni type series (Figures 8 and 9). Both
the live animals and the preserved type series show somewhat raised and outwardly projecting imbricate scales on
the body and limbs, similar scale sizes and counts, as well as very similar patterns of darker pigmentation, such as
a line extending from the loreal or preocular, through the orbit, and posteriorly through the temporal zone, three
dark lines radiating from the orbit through the labials onto the maxilla, and paired series of rounded blackish spots
on the dorsum, with intervening lighter zones. The Alto Patache lizards have been reported as P. reichei (Larran et
al., 2001) and no other liolaemid is in fact reported from the Iquique coastal zone or the interior upland plain (e.g.,
the Oasis de Pica area) west of the Andean Cordillera.
Based on the information provided by Nez & Yez (1983) and Valladares (2004), the phrynosauroid Liolaemus from the Iquique area, here referred to L. stolzmanni and hitherto referred to L. reichei, can clearly be distinguished from all other phrynosauroid Liolaemus (equivalent to the reichei group of Pincheira-Donoso et al.,
2008). It has fewer and larger scales around midbody (ca. 4450) than other phrynosauroid species (>60). Its dorsal scales are strongly imbricate to subimbricate or juxtaposed in the posterior of some females versus subimbricate
to juxtaposed in other species. It has fewer supralabials than the other species. Also, all currently recognized
phrynosauroid Liolaemus are allopatric in relation to each other, although some are known to co-occur with nonphrynosauroid Liolaemus.

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FIGURE 7. Female syntype here designated allotype (NMW 13580:3) of Liolaemus stolzmanni (Steindachner, 1891). Photographs Richard Gemel/Naturhistorisches Museum Wien.

FIGURE 8. Liolaemus stolzmanni, in life, from Oasis de Niebla de Alto Patache, 60 km S of Iquique, Chile. Photograph
Luis Prez Reyes.

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FIGURE 9. Lateral aspect of Liolaemus stolzmanni paralectotype (NMW 13580:2). Photograph Richard Gemel /Naturhistorisches Museum Wien.

In summary, I hereby propose that Phrynosaura reichei Werner, 1907, be considered a junior subjective synonym of Liolaemus stolzmanni (Steindachner, 1891). Based upon external characteristics, the syntypes of L. stolzmanni cannot be distinguished from lizards from the Iquique area recognized historically as P. reichei or L. reichei.
Although the name Phrynosaura reichei has been more widely used, Steindachners description is unquestionably
valid, the type series remains preserved in excellent condition, and the name has been appropriately applied by
authors such as Cei (1979), Laurent (1984), and Etheridge (1995) who each recognized it as a phrynosauroid species but in no case intended to compare it against P. reichei for purposes of taxonomic revision.
Given that there is no known specimen of a lizard assignable to L. stolzmanni from a locality in modern day
Peru or anywhere in the high Andes (>3000 masl), and that the only known lizards matching the type series of L.
stolzmanni are from coastal Chile near Iquique, I recommend that the type locality associated with Liolaemus stolzmanni (Steindachner, 1891) be emended to Deserts of Iquique, Tarapac Region, Chile in recognition that the
specimens were most likely obtained by Stolzmann from third party who collected them from a location in the
Tarapac Department or Iquique itself, which was Peruvian territory prior to the end of War of the Pacific in 1884,
which is consistent with the date of 1883 in the museum records.
In the interest of further clarity, I designate specimen NMW 13580:1, a male with four well-developed precloacal pores, as lectotype of Liolaemus stolzmanni (Steindachner, 1891), specimen NMW 13580:3, a female with a
distinctly broader thorax than either male, as allotype, and NMW 13580:2, a male with four evident, but less well
developed precloacal pores, as paralectotype.
A second conclusion is that the populations of lizards inhabiting the Ollage, Salar de Carcote, and Salar de
Ascotn areas of easternmost Antofagasta, Chile, referred to as C. stolzmanni by Donoso-Barros (1966) or as L.
stolzmanni by Valladares et al. (2002), Pincheira-Donoso & Nez (2005), and other subsequent authors, are part
of the metapopulation of lizards from adjacent areas of Bolivia recognized as L. jamesi pachecoi Laurent, 1995.
One may ask if Werner, also a Viennese zoologist and a contemporary of Steindachner, had ever examined the
L. stolzmanni specimens housed in that city. As described by Werners student Wettstein (1940), Steindachner,
Director of the Naturhistoriches Museum Wien until shortly before his death in 1919, considered the young Werner
to be a dangerous competitor and denied him access to the collections. This very fact is the simple explanation for
a great deal of taxonomic confusion.

Discussion: On the status of the taxa Liolaemus aymararum, L. jamesi and L. jamesi pachecoi
In his description of Liolaemus jamesi pachecoi, Laurent (1995) posited that L. aymararum, L. jamesi, and L. j.
pachecoi may merely represent clinal or patternless variations of a widely distributed L. jamesi with no clear geographic or other boundaries. Both Etheridge (1995) and Valladares et al. (2002) commented on the possibility that
L. aymararum was simply a synonym of L. jamesi. The latter authors also demonstrated graphically, but without
comment, that the lizards they recognized as L. stolzmanni from Ollage could not be distinguished in their principal components plot from L. jamesi based on 16 morphometric characters. Pincheira-Donoso & Nez (2005) for-

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mally synonymized L. aymararum with L. jamesi but did not mention L. jamesi pachecoi. Pincheira-Donoso et al.
(2008) recognized L. pachecoi as a distinct species without comment. Schulte & Moreno-Roarks (2009) phylogenetic hypothesis of the Iguania, including 96 species of Liolaemus, places L. aymararum and L. stolzmanni
(from Salar de Ascotn) in a well supported sister-species relationship, but they did not include jamesi. Most
recently, Etheridge & Frost (2010) recognized L. jamesi as a polytypic species with three named subspecies (L. j.
aymararum, L. j. jamesi, and L. j. pachecoi).
However, the mtDNA sequence divergence between the taxa identified as L. aymararum and L. stolzmanni by
Schulte & Moreno-Roark is approximately 4% (J. Schulte, in litt.). Considering that the mtDNA divergence
reported by Valladares et al. (2002) between L. audituvelatus and L. fabiani is 3.5% and that between L. famatinae
and L. aff. andinus (reported as L. andinus AF099245) is 3.3%, the 4% divergence between L. aymararum (=L.
jamesi) and L. stolzmanni (=L. pachecoi) is enough to support the recognition of two distinct species. Also,
morphometric analyses may not discriminate biologically or ecologically significant difference between species.
While the coloration in life of the Chilean lizards from the Salar de Ascotn, Salar de Carcote and Quebrada del
Inca (identified previously as L. stolzmanni, but treated under the name L. pachecoi from now on) has not been
described, the photograph of the yellow lizard from Desierto del Siloli (Figure 3) was independently identified as
L. stolzmanni by D. Pincheira-Donoso (in litt.) and H. Nez (in litt.) and is rather distinct from lizards recognized as L. jamesi (see Pincheira-Donoso & Nez, 2005). Biogeographically, the Desierto de Siloli is the corridor
which links the Salar de Ascotn with Laguna Colorada, the type locality of L. jamesi pachecoi located some 60
km from Salar de Ascotn. These populations occupy endorrheic basins of the Altiplano and are isolated from the
other populations recognized as L. jamesi on the Pacific slope by high volcanic ranges. Based on the above considerations, I agree with the recognition of L. pachecoi as a species distinct from L. jamesi by Pincheira-Donoso et al.
(2008).

Acknowledgements
I am greatly indebted to Richard Etheridge and Herman Nez for their comments on the manuscript, their unpublished museum notes, their deep insights, and their impartial objectivity. I am also very grateful to Richard Gemel
of the Naturhistorisches Museum Wien for his prompt cooperation in providing the excellent photographs of the L.
stolzmanni syntypes and data from the collections. Also, I thank Monique Halloy of the Fundacin Miguel Lillo for
providing the photographs of the L. jamesi pachecoi paratype, as well to Edgardo Milla Anacona for his photograph of the live L. jamesi pachecoi and to Luis Prez Reyes for his photograph of the live L. stolzmanni from near
Iquique. Also, I thank Michelle Koo and staff at the Museum of Vertebrate Zoology for photographs of specimens.
I also am indebted to Neftali Camacho of the Los Angeles County Museum for last minute photographs, scale
counts, and measurements. I thank Jaime E. Pfaur for providing the description of L. insolitus. I thank Bill
Duellman for more than 30 years of generosity and support, and for facilitating images of University of Kansas
specimens. Finally, I thank Daniel Pincheira-Donoso for his knowledge on the Chilean Liolaemus and his encouragement to follow through on my inclinations.

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APPENDIX I. Museum specimens examined.

Liolaemus jamesi (2). CHILE: TARAPAC: Alta Camia, 110 mi NE Iquique: MVZ Herps 6680506.
Liolaemus pachecoi (1). BOLIVIA: POTOS: Laguna Colorada (67 44O, 22 13S), 4270 m, Prov. Sud Lpez: FML
02788;
Liolaemus poconchilensis (3). CHILE: ARICA Y PARINACOTA: (Pampa) Chaca: LACM 9312; PERU: TACNA: 4 mi N
of Tacna: MVZ Herps 99661, 99663.
Liolaemus stolzmanni (3). CHILE: TARAPAC: Vicinity of Iquique: NMW 13580:13.
Liolaemus aff. walkeri (1). PERU: ANCASH: 7 km E Laguna Conococha: KUH 133861.
Liolaemus species 1 (1). PERU: ANCASH: 18 km (by road) Llupash: MVZ Herps 84630.
Liolaemus species 2 (3). PERU: AREQUIPA: 3 km SW Uchumayo, Alturas de Uchumayo: KUH 163589, 163592, 163594.
Liolaemus species 3 (1). PERU: AREQUIPA: 18 km N Matarani, Quebrada de Guerrero: KUH 163595.

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