Langstroth 2011 - Liolaemus Stolzmanni L. Reichei and L. Jamesi Pachecoi-libre

8/12/2019 Langstroth 2011 - Liolaemus Stolzmanni L. Reichei and L. Jamesi Pachecoi-libre

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

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright 2011 Magnolia Press

Zootaxa 2809: 2032 (2011)www.mapress.com / zootaxa / Article

On the species identities of a complex Liolaemus fauna from the Altiplano andAtacama 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: [email protected]

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 de-scribed more than 150 years ago. Within this lineage, the Andean Liolaemus faunas have proven to be a major challengefor herpetologists. Therefore, intense research is needed in this area to clarify long-standing problems. After more than acentury of taxonomic confusion, the identity of Liolaemus stolzmanni (Steindachner, 1891) is here restored as the namethat must be applied to the lizards widely known asPhrynosaura (= Liolaemus ) reichei Werner, 1907 from the low to mid-elevation 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 typelocality for L. stolzmanni (= L. reichei ) is emended to Deserts of Iquique, Tarapac Region, Chile. Furthermore, the rec-ognition of L. pachecoi as a species distinct from L. jamesi is supported by mtDNA sequence divergence data despite theinconclusive 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 Peruand that (ii) the Chilean Altiplano populations currently recognized as L. stolzmanni are L. pachecoi, a species hithertoknown only from Bolivia. Also, I report the first confirmed specimens of L. poconchilensis from Peru, a species previouslyknown 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 amniotegenus, exceeded only by the iguanian genus Anolis from tropical America and the Caribbean. During their evolu-

tionary history, Liolaemus have extensively colonized an extraordinary diversity of environments (Espinozaet al .,2004; Pincheira-Donosoet al ., 2008a) that have forced the evolution of a remarkable diversity of morphologies andbehaviors (Schulteet al ., 2004; Pincheira-Donosoet al ., 2009). This extraordinary diversification within the genushas resulted in a substantial challenge for Liolaemus specialists aiming to identify and name these species andmajor-level groups (Etheridge & Espinoza, 2000). Part of the long-standing nomenclatural controversies within thegenus 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 inthe Altiplano (High Andes Plateau) of Argentina, Bolivia, Chile and Peru, and the adjacent lower areas of the Ata-cama desert in northern Chile and southern Peru. In this area, several nomenclatural problems have been discussedin recent syntheses (e.g. Etheridge & Espinoza, 2000; Valladareset al. , 2002; Pincheira-Donosoet al ., 2008b), and

several species have only recently been recognized as a result of historical difficulties for field explorations, isola-tion of areas, and climatic constraints. Therefore, much more systematic and geographical research is needed in this


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Zootaxa 2809 2011 Magnolia Press 21ON THE IDENTITY OF LIOLAEMUS STOLZMANNI

region to strengthen our understanding of Liolaemus assemblages. In this study, I concentrate on three taxa cur-rently recognized (e.g., Etheridge & Frost, 2010) as L. stolzmanni, L. reichei , and L. jamesi pachecoi . My aim is toprovide 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 par-ticular specimens collected long ago and housed in collections distant to the researcher, such as European or NorthAmerican collections that are difficult to access by Liolaemus specialists residing in South America. However, theadvent of the digital era has created new means of access via digital photography and Internet. At present, mostresearchers worldwide have access to collections by means of e-mail or directly through electronic platformsimplemented by institutions, where specimen-by-specimen catalogues are provided. For this study, I have utilizednewly 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, fromseveral different angles on each specimen, and with high resolution to allow detailed analyses of scale and colorvariation. 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 thespecimens, and the ease of sharing the images with colleagues. Another novel use of the Internet is to perform vir-tual lizard surveys by searching the millions of online images taken across the globe by researchers, nature enthusi-asts, tourists, and others. By this method, I obtained photographs of live lizards from locations near the typelocalities of the species of interest, which led to important questions and findings, including some of the onlyknown life habits for some taxa.

Analyses of high-resolution photographs were complemented with careful analyses of the original descriptionsof the species in question provided by the original authors, and based on additional later descriptions of the sametype material published in subsequent studies. Finally, as an additional source of information, and once again tak-ing advantage of modern ways of communication, I consistently discussed my observations and conclusions with anumber of herpetologists who have experience in Liolaemus research. These discussions (based on appropriate pre-sentation 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 lizardCtenoblepharis Stolzmanni from three syntypes currently housed in theNaturhistorisches Museum Wien, from the vague locality of Hoch-Peru (High Peru). According to museumrecords, theC. stolzmanni syntypes are from Coll. Stolzmann 1883 and there is no other information on the local-ity. Stolzmann (also Sztolcman) is well known as one of the most diligent ornithological collectors to haveworked in Peru, where he collected intensively in the northern part of the country, primarily in Cajamarca, Amazo-nas, and Tumbes (Stephens & Traylor, 1983), areas where there are no known liolaemid lizards. There are no local-ities from southern Peru associated with Stolzmanns ornithological field collections (Stevens & Traylor, 1983). Itis, however, very possible that Stolzmann may have received the specimens as gifts or even purchased them fromanother collector who may have collected them in the Tarapac Region, even around Iquique, which was Peruvianterritory until the aftermath of the War of the Pacific (18791884). Steindachner expressly compared his new spe-cies against a specimen cataloged asCtenoblepharis adspersus at his museum , and clearly considered hisC. stolz-manni to be congeneric with this species based on its well developed palpebral fringes.

Also, Boulenger (1891) describedCtenoblepharis 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 andC. stolz-manni ). Besides stating, shape of head and general proportions same asC. adsperus, Boulenger provided no dis-cussion as to why he considered this species comparable toC. 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), notingthat it in fact corresponds to an undescribed species ofPhrynosaura .


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LANGSTROTH22 Zootaxa 2809 2011Magnolia Press

Werner (in Brger, 1907) subsequently described the genusPhrynosaura and the speciesP. reichei from a sin-gle specimen from "Iquique" (Chile). He found the new genus to be similar to the agamidPhrynocephalus and theiguanidCtenoblepharis , but did not provide a specific diagnosis.

Besides Mllers (1928) comments onP. reichei in his description ofPhrynosaura werneri, very little was pub-lished regarding eitherC. stolzmanni or P. reichei until the printing of Donoso-Barros' (1966) Reptiles de Chile , inwhich he provided redescriptions of both species.

Although Donoso-Barros (1966:341) considered Steindachner's description ofC. stolzmanni to be "primitive",comparative, extraordinarily limited, and barely permitting one to form an opinion of the species whose typespecimens apparently were unknown to him, he did indeed form his own opinion of what specimens might beassignable to Steindachners description, which he ultimately found to fit a lizard collected from Quebrada del Incain the high Andes of the Antofagasta region. However, there are characteristics described by Steindachner whichDonoso-Barros apparently did not consider, such asC. stolzmanni having head shields larger than the largest dorsalscales and having fringed eyelids, characteristics not applicable to the lizard he collected. He also ignored the spot-ted and barred patterns described by Steindachner, which contrast with the lizard described by Donoso-Barroswhich has a brown dorsum with only small dark flecks caused by blackish pigmentation in the posterior of eachdorsal scale.

On the other hand, Donoso-Barros had obtained the holotype ofP. reichei as part of his private collection andhe 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 Brgersidiosyncratic 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 thatit had 44 scales around midbody.

During a tour of European collections, Etheridge examined theCtenoblepharis stolzmanni syntypes in Viennain 1967. In his unpublished notes he wrote, "The syntypes ofCtenoblepharis stolzmanni have the characteristics of what I have been callingPhrynosaura : a short baby bird type head with strongly projecting superciliaries, andrather large, smooth subimbricate dorsals". He also noted that the dorsal scales are equal to or slightly smaller thanthe ventral scales, there are 50 scales around midbody, and 4 precloacal pores (R.E. Etheridge,in litt. ). This visionof C. stolzmanni contrasts with that of Donoso-Barros, who evidently did not see similarities withPhrynosaura inSteindachners description.

Donoso-Barros (1972) later provided a comparison of the type specimens of the generaCtenoblepharis andPhrynosaura , in which he concluded thatPhrynosaura is a junior synonym ofCtenoblepharis . In this publication,he provided new descriptive information onP. 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-Barrosnoted that adultP. reichei possess tricuspid lateral teeth, conditions not present in the juvenile holotype. He alsonoted thatP. reichei has a tripartite subocular with the middle section being the longest. He also noted that theinfradigital lamellae of the adults were much less strongly keeled than those of the juvenile. These characteristicsare also described in Steindachners description ofC. stolzmanni . One interesting aspect of what appears to be awell-proportioned and accurate drawing of theP. reichei holotype is that the dorsal scales are very small, andwould number far more than the 44 around midbody reported by Donoso-Barros (1966). It is possible that Donoso-Barros based his count on the female specimen illustrated in his books photographic plates or that the drawing hasinaccurately represented the body scales.

Cei (1979) concluded that ifCtenoblepharis is to be recognized as a valid genus, it should be restricted toinclude onlyC. adspersus, C. nigriceps, C. reichei, and C. stolzmanni. In contrast to previous authors, Cei consid-eredC. jamesi and C. schmidti to be members of Liolaemus, a decision based on the absence of the morphologicaltraits which bound together his concept ofCtenoblepharis .

Five years later, Laurent (1984) revalidatedPhrynosaura based on a suite of distinct morphological character-istics and foundCtenoblepharis to be a monotypic genus including onlyC. adsperus . As did Etheridge, Laurentalso examined theC. stolzmanni syntypes and considered them to be representative of thePhrynosaura morpho-type, along withP. reichei andP. audituvelata . Laurent clearly did not findP. stolzmanni to be similar toC. ayma-rarum , C. jamesi, or C. nigriceps, which, following Cei, he removed fromCtenoblepharis and placed in Liolaemus .

Etheridge (1995) carefully reconsidered the taxonomic history and systematics of the genusCtenoblepharysand other genera related to Liolaemus , and reestablished Tschudis original spelling (i.e.,Ctenoblepharysadspersa ). While Etheridge did not find overriding evidence at the time to support the monophyly ofPhrynosaura ,


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which he synonymized with Liolaemus , he did recognize their prominent ciliary fringes and truncated snouts aspossible synapomorphies of a group containing the species L. audituvelatus, L. reichei, and L. stolzmanni .

More recently, Nezet al. (2003) described L. manueli and L. torresi (both asPhrynosaura ) from differentareas 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 riverbasin to the southeast of Iquique and differs from L. reichei by its relatively longer tail, a dorsal color pattern con-sisting of more numerous and smaller spots, smaller and more numerous body scales, and more numerous suprala-bials. 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 ofsmall subimbricate to juxtaposed body scales, ahigher number of supralabials, and the presence of scales on the tympanum. Its distribution in the Diego deAlmagro area of the Atacama Region is far south of the known ranges of the other phrynosauroid species. In thatpaper, Nezet al . clearly were aware of Laurents decision to includeC. stolzmanni in Phrynosaura , but did notcomment on the relationship of this species to otherPhrynosaura or their opinions of Laurents decision.

The most recently named phrynosauroid species is L. poconchilensis , described by Valladares (2004) from thedeserts 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 asPhrynosaura reichei(Donoso-Barros, 1969) or asPhrynosaura sp. from Arica (Nezet 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 interpre-tation of Donoso-Barros (1966) regarding Liolaemus stolzmanni and provided a more detailed description of thispoorly understood species, based on additional specimens collected by Nez. They included the first publishedphotograph of a lizard identified as Liolaemus stolzmanni , which clearly did not suggest phrynosauroid morphol-ogy, rather a larger, robust form with a thick neck and very large dorsal scales, which they clearly considered to berelated to L. jamesi . Pincheira-Donoso & Nez recognizedPhrynosaura as a valid genus and did not include itsspecies in their 2005 monograph.

In the meanwhile, Laurent (1995) named Liolaemus jamesi pachecoi based on two specimens from LagunaColorada, 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 remarkablybrief diagnosis compares it only to L. jamesi and L. aymararum . While finding that the number of dorsal scalesbetween 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 therelationship between the nostril-mouth distance and the minimum space between the supraoculars. However, healso 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 noother authors have commented on this taxon.

While Cei (1979), Laurent (1984), and Etheridge (1995) have all commented on the clearly phrynosauroidcharacter of the lizards described by Steindachner asCtenoblepharis stolzmanni , there has been no published dis-cussion regarding the use of this name for lizards without any phrynosauroid characteristics by authors such asDonoso-Barros (1966) and Pincheira-Donoso & Nez (2005), and others who have included L. stolzmanni in spe-cies diagnoses and lists based on Chilean material.

Abdalaet al. (2008) included both L. reichei and L. stolzmanni in their diagnoses of L. huayra and L. inti , refer-ring 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 theAlta Camia specimens to be the nominal L. jamesi and not similar to L. j. pachecoi (confirmed independently byH. Nez and D. Pincheira-Donoso,in litt. ). The lizards from 4 mi N of Tacna are here identified as L. poconchil-ensis and noted as the first definitive record of the species for Peru. The similarity in color patterns with the photo-graphs 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) Llu-pash, 10,000 ft in the Ancash Department, also generally agrees with a female or juvenile L. poconchilensis. How-ever, 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 Herps84632), 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 assignableto L. poconchilensis as it has scale counts (supralabials, scales around midbody, and middorsals between occiputand 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, 2150masl, just west of the city of Arequipa in 1974 and these were later cataloged in KU asPhrynosaura stolzmanni(KU 16358994). These lizards are clearly not conspecific with the type series of L. stolzmanni based on theirsmaller body scales, less conspicuous palpebral fringe, and distinctive color patterns. Also, these specimens werecollected in habitat of gray sands with granitic rocks and have cryptic coloration matching the dark sand and rocksurfaces (Figure 2). According to Duellmans field notes, the adult males have brown head, mid-dorsum, tail, andlimbs; rest of dorsum mottled orange and metallic blue; flanks pinkish tan with dark brown mottling; limbs and tailbarred with dark brown; anterior chin white; throat yellow; lateral edge of belly orange; other ventral surfaceswhite. Females have brown dorsum with dark brown and tan spots; venter white. Juveniles have dorsum likefemales 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 haslarger, 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 colorpatterns) from L. stolzmanni and were expressly considered distinct from C. stolzmanni and C. riechei (sic) byCei & Pfaur. Cei & Pfaur (1982) also discussed specimens assigned to L. insolitus from 10 km SW of Uchu-mayo, 2800 masl, which are the same lizards referred to by Pfauret al. (1978) as Ctenoblepharus sp. (sic) fromthe Batolito de la Caldera, 10 km SW Uchumayo, Arequipa. These are mentioned to be part of the series at KUfrom Uchumayo, although the stated localities are somewhat different. These specimens are clearly not L. insolitusand appear to correspond to the specimen from Arequiba, 7,500 ft mistaken by Boulenger (1885) forC. aspersa .Differences mentioned by Pfauret 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 in L. insolitus ),longer digits (i.e. long toes vs. short feet, which are clearly different in the photographs examined), distinctdorsal color pattern in males (Uchumayo males have tigered pattern of black mottling on dorsum), and possiblythe 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 clearlydistinct 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 deAscotn) form a well-supported sister species pair nested within an unnamed clade corresponding to themontanusseries, which also included a clade identified as L. reichei in a four-way polytomy with L. chlorostictus and twounnamed multispecies clades. However, the specimen identified as L. reichei from which the genetic material wasobtained, 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 sub-

ject to considerable confusion.

The Clarification

While working on a review of the state of the knowledge and taxonomic problems of the Liolaemus found to thenorth of the Tropic of Capricorn, I came across online photographs of unusual Liolaemus from SW Bolivia in thevicinity 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 juxta-posed 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, num-bered 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 agreedthat these appear to be what he has referred to as L. stolzmanni and has collected from Ollage and the Salar deAscotn, suggesting that the Chilean L. stolzmanni and the Bolivian L. jamesi pachecoi might be conspecific. How-ever, this question could only be answered by the examination of the syntypes ofCtenoblepharis stolzmanni .

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

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 MiguelLillo.


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Discussion: Comparison of L. stolzmanni and L. reichei

Upon receiving photographs of the exceptionally well-preserved L. stolzmanni type series from the Naturhisto-risches Museum Wien (Figures 57), it was immediately evident that Etheridge and Laurent were correct in theirplacement ofC. stolzmanni within their concepts ofPhrynosaura and that Donoso-Barros had misapplied the nameC. 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 ofP. reichei as being lanceolate anteriorly and becomingquadrangular posteriorly and Werner mentioned that they form an angular pattern, both of which are evident in the

L. stolzmanni allotype. As noted by Werner forP. reichei , the toes of the L. stolzmanni type series are distinctly yel-lowish. Steindachner had noted that there were three subocular scales and while Donoso-Barros reported only sin-gle subocular inP. 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 onthe same individual. Also, Donoso-Barros (1972) reported only conical teeth in the juvenileP. reichei holotype heredescribed 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 cityof 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-Donosoet al., 2008b). This question is irrelevant as there is noevidence 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 evi-dence of any Liolaemus , phrynosauroid or otherwise, onlyPhyllodactylus 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 cityof Iquique, the L. reichei type locality, are strikingly similar to the L. stolzmanni type series (Figures 8 and 9). Boththe live animals and the preserved type series show somewhat raised and outwardly projecting imbricate scales onthe 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, threedark lines radiating from the orbit through the labials onto the maxilla, and paired series of rounded blackish spotson the dorsum, with intervening lighter zones. The Alto Patache lizards have been reported asP. reichei (Larranet 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 Lio-laemus from the Iquique area, here referred to L. stolzmanni and hitherto referred to L. reichei, can clearly be dis-tinguished from all other phrynosauroid Liolaemus (equivalent to the reichei group of Pincheira-Donosoet al .,2008). It has fewer and larger scales around midbody (ca. 4450) than other phrynosauroid species (>60). Its dor-sal scales are strongly imbricate to subimbricate or juxtaposed in the posterior of some females versus subimbricateto 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 non-phrynosauroid Liolaemus.


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FIGURE 7. Female syntype here designated allotype (NMW 13580:3) of Liolaemus stolzmanni (Steindachner, 1891). Photo-graphs 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 /Naturhisto-risches Museum Wien.

In summary, I hereby propose thatPhrynosaura reichei Werner, 1907, be considered a junior subjective syn-onym of Liolaemus stolzmanni (Steindachner, 1891). Based upon external characteristics, the syntypes of L. stolz-manni cannot be distinguished from lizards from the Iquique area recognized historically asP. reichei or L. reichei .Although the namePhrynosaura reichei has been more widely used, Steindachners description is unquestionablyvalid, the type series remains preserved in excellent condition, and the name has been appropriately applied byauthors such as Cei (1979), Laurent (1984), and Etheridge (1995) who each recognized it as a phrynosauroid spe-cies but in no case intended to compare it againstP. 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 dayPeru 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 stolz-manni (Steindachner, 1891) be emended to Deserts of Iquique, Tarapac Region, Chile in recognition that thespecimens 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 preclo-

acal pores, as lectotype of Liolaemus stolzmanni (Steindachner, 1891), specimen NMW 13580:3, a female with adistinctly broader thorax than either male, as allotype, and NMW 13580:2, a male with four evident, but less welldeveloped precloacal pores, as paralectotype.

A second conclusion is that the populations of lizards inhabiting the Ollage, Salar de Carcote, and Salar deAscotn areas of easternmost Antofagasta, Chile, referred to asC. stolzmanni by Donoso-Barros (1966) or as L.stolzmanni by Valladareset al. (2002), Pincheira-Donoso & Nez (2005), and other subsequent authors, are partof 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 Wernerto be a dangerous competitor and denied him access to the collections. This very fact is the simple explanation fora 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 geo-graphic or other boundaries. Both Etheridge (1995) and Valladareset 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 princi-pal 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-Donosoet al.(2008) recognized L. pachecoi as a distinct species without comment. Schulte & Moreno-Roarks (2009) phyloge-netic 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 . Mostrecently, 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 bySchulte & Moreno-Roark is approximately 4% (J. Schulte,in litt .). Considering that the mtDNA divergencereported by Valladareset al. (2002) between L. audituvelatus and L. fabiani is 3.5% and that between L. famatinaeand L. aff.andinus (reported as L. andinus AF099245) is 3.3%, the 4% divergence betweenL. 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 delInca (identified previously as L. stolzmanni , but treated under the name L. pachecoi from now on) has not beendescribed, the photograph of the yellow lizard from Desierto del Siloli (Figure 3) was independently identified asL. stolzmanni by D. Pincheira-Donoso (in litt. ) and H. Nez (in litt. ) and is rather distinct from lizards recog-nized as L. jamesi (see Pincheira-Donoso & Nez, 2005). Biogeographically, the Desierto de Siloli is the corridorwhich links the Salar de Ascotn with Laguna Colorada, the type locality of L. jamesi pachecoi located some 60km from Salar de Ascotn. These populations occupy endorrheic basins of the Altiplano and are isolated from theother populations recognized as L. jamesi on the Pacific slope by high volcanic ranges. Based on the above consid-erations, I agree with the recognition of L. pachecoi as a species distinct from L. jamesi by Pincheira-Donosoet al.(2008).

Acknowledgements

I am greatly indebted to Richard Etheridge and Herman Nez for their comments on the manuscript, their unpub-lished museum notes, their deep insights, and their impartial objectivity. I am also very grateful to Richard Gemelof 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 forproviding the photographs of the L. jamesi pachecoi paratype, as well to Edgardo Milla Anacona for his photo-graph of the live L. jamesi pachecoi and to Luis Prez Reyes for his photograph of the live L. stolzmanni from nearIquique. 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, scalecounts, and measurements. I thank Jaime E. Pfaur for providing the description of L. insolitus . I thank BillDuellman for more than 30 years of generosity and support, and for facilitating images of University of Kansasspecimens. Finally, I thank Daniel Pincheira-Donoso for his knowledge on the Chilean Liolaemus and his encour-agement to follow through on my inclinations.

References

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Cei, J.M. (1979) A reassessment of the genusCtenoblepharis (Reptilia, Sauria, Iguanidae) with a description of a new subspe-cies of Liolaemus multimaculatus from western Argentina. Journal of Herpetology , 13, 297302.

Cei, J.M. & Pfaur, J.E. (1982) Una especie nueva de Liolaemus (Iguanidae: Squamata): su sistemtica, ecologa y distribucin. In: Actas 8vo Congreso Latinoamericano de Zoologa, Mrida, Venezuela, Octubre 1980 , pp. 573586.

Donoso-Barros, R. (1966) Reptiles de Chile . Universidad de Chile, Santiago de Chile, 458+cxlvi pp.Donoso-Barros, R. (1969) Observaciones in vivo sobrePhrynosaura reichei Werner (Sauria-Iguanidae). Boletn Sociedad de

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Etheridge, R. (1995) Redescription ofCtenoblepharys adspersa Tschudi 1845, and the taxonomy of the Liolaeminae (Reptilia:Squamata: Tropiduridae). American Museum Novitates , (3142), 134.

Etheridge, R. & Espinoza, R.E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semi-annotatedbibliography.Smithsonian Herpetological Information Service , 126, 164.

Etheridge, R. & Frost, D.R. (2010) Liolaemidae Frost and Etheridge . Available from http://research.amnh.org/vz/herpetology/ f/Liolaemidae.pdf (accessed 17 September 2010)

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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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