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Liolaemus multiformis (COPE, 1875)

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Higher TaxaLiolaemidae, Iguania, Sauria, Squamata (lizards)
Subspecies 
Common Names 
SynonymProctotretus multiformis COPE 1875: 178
Proctotretus multiformis — COPE 1878: 35
Liolaemus multiformis COPE 1876 (fide LAURENT 1992)
Liolaemus multiformis — BOULENGER 1885: 153
Liolaemus tropidonotus BOULENGER 1902: 397
Liolaemus multiformis multiformis — SHREVE 1941
Liolaemus multiformis — DONOSO-BARROS 1966: 517
Liolaemus tropidonotus — LANGSTROTH 2005
Liolaemus multiformis — LANGSTROTH 2021

from L. signifer (now = multiformis fide Lanstroth 2021)
Proctotretus signifer DUMÈRIL & BIBRON 1837: 288 (nomen dubium)
Proctotretus Signifer — DUMÉRIL, BIBRON & DUMÉRIL 1854: 269
Liolaemus signifer — DONOSO-BARROS 1966: 329
Liolaemus signifer — PETERS et al. 1970: 194
Liolaemus signifer — DIRKSEN & DE LA RIVA 1999
Liolaemus (Eulaemus) signifer — RAMIREZ LEYTON & PINCHEIRA-DONOSO 2005: 142
Liolaemus signifer — AGUILAR-PUNTRIANO et al. 2019 
DistributionW Bolivia, S Peru

Type locality: apparently not given; Burt and Burt (1931: 277) consider specimens from Juliaca, Puno, to be topotypical specimens of L. m. multiformis.  
Reproductionovovivparous. 
TypesLectotype: ANSP 13064, designated by Laurent 1992; Syntypes, now paralectotypes: ANSP 13065, 11369-70, 13098, 13104
Holotype: MNHN-RA 6860 (for a history of this specimen see Langstroth 2021); Other specimens: MUSM, BYU, FMNH. Syntypes: (6) ANSP 13064-65, 11369-70, 13098, 13104 [signifer] 
DiagnosisLectotype description: a stout female measuring 74 mm SVL with rather smooth but strongly imbricate, phylloid dorsals (DSOT=74) and with a dark dorsal ground color and scattered lighter spots (one or two scales in size), without traces of dorsal blotches. Five of the six type specimens6 have rather large, imbricate, and keeled dorsal scales (DSOT 62–76), of which ANSP 11370 has exceptionally large, imbricate, and strongly keeled dorsals (DSOT= 62), not unlike the large male syntype of L. tropidonotus; ANSP 11369 represents the other extreme among the syntypes, with somewhat smaller (DSOT= 79), more triangular, smooth dorsals that are more similar to individuals from populations from the southern end of Lake Titicaca in Bolivia (e.g., specimens described as L lenzi, L. variabilis. or L. bolivianus) (Langstroth 2021: 122).

Remarks: Compared to the lizards described as Liolaemus multiformis, L. annectens, and L. pantherinus, the lizards from the southern shores of Lake Titicaca and the Altiplano of La Paz and Oruro, including the L. lenzi holotype, the L. bolivianus syntypes, and the L. variabilis syntypes, all recognized as L. multiformis or L. signifer in the literature of 1970s onwards, tend to have more numerous, smaller, and less imbricate dorsal scales. DSOT counts of L. multiformis (n=18), including the L. multiformis type series and MNHN-RA-0.6860, range from 62 to 89 (mean= 75.6, median= 76), while DSOT counts of Bolivian specimens (n= 27) assigned previously to either L. signifer or L. multiformis, including types of L. lenzi, L. variabilis, and L. bolivianus (but not including lizards corresponding to L. forsteri), range from 74 to 110 (mean= 89.7, median= 87). Langstroth 2021: 123 suggests that the lizards of the Bolivian Altiplano with generally smaller and more numerous dorsals, can be reliably distinguished morphologically from those identified by him as L. multiformis (based on Langstroth 2021).

Boettger (1891) distinguished L. lenzi from L. multiformis based on the higher number of scales around midbody in the L. lenzi types (74–86 vs. 60 –70)7. Etheridge (in litt.) counted 72 scales around midbody and 87 paravertebrals on the L. lenzi holotype and did not mention any paratypes. My examination of the L. lenzi holotype (SMF 11110; Fig. 5) finds this specimen to have at least 92 DSOT and the dorsals to be juxtaposed to subimbricate, with very few scales showing development of keels. This evidence suggests that the L. lenzi holotype belongs to the southernmost Titicaca-Bolivian Altiplano population, not to L. multiformis. While Boettger (1891) made no specific mention of paratypes, he did describe the coloration of at least one juvenile in addition to that of the adult and gave a range of scales around midbody (74–86). Thus, Etheridge’s counts of 72 scales around midbody and 87 DSOT suggest that the L. lenzi holotype was at the lower end of Boettger’s scale counts and thus DSOT counts for the paratype(s) likely exceed 87–92 given the general positive correlation between scales around midbody and DSOT in the L. montanus group, which would only further support the separation of L. lenzi from L. multiformis based on scales around midbody and DSOT (Langstroth 2021: 123). 
CommentSynonymy: Langstroth 2021 revalidated L. multiformis and applied the name to a clade identified as Liolaemus signifer in Aguilar-Puntriano et al. (2018), with Liolaemus tropidonotus as a synonym. Donoso-Barros 1966 assigned the Liolaemus from the Antofagasta Region to L. m. multiformis, but Núñez and Fox (1989) identified these specimens as L. puritamensis. Veloso et al. (1982) reported L. multiformis from Pampa Chucuyo in the Arica y Parinacota Region, specimens that Laurent (1998) would later describe as L. pleopholis.

Distribution: see map in Langstroth 2021: 124 (Fig. 6). 
References
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