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Phrynosoma hernandesi GIRARD, 1858

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Higher TaxaPhrynosomatidae, Phrynosomatinae, Phrynosomatini; Iguania, Sauria, Squamata (lizards)
SubspeciesPhrynosoma ornatissimum brachycercum SMITH 1942
Phrynosoma hernandesi brevirostris GIRARD 1858
Phrynosoma hernandesi diminutum MONTANUCCI 2015
Phrynosoma hernandesi hernandesi GIRARD 1858
Phrynosoma ornatissimum ornatissimum GIRARD 1858
Phrynosoma hernandesi ornatum GIRARD 1858 
Common NamesE: Mountain Short-horned Lizard; ornatum: Salt Lake Valley Short-horned Lizard
G: Berg-Kurzhornkrötenechse; ornatum: Salt Lake Valley Kurzhornkrötenechse
S: Camaleón Cuernitos de Hernandez
E: San Luis Valley Short-horned Lizard [diminutum]
E: Great Plains Short-horned Lizard [brevirostris]
E: Baur’s Short-horned Lizard [bauri]
E: Desert Short-horned Lizard [ornatissimum]
G: Wüsten-Kurzhorn-Krötenechse [ornatissimum]
E: Short-tail Short-horned Lizard [brachycercum]
G: Kurzschwanz-Kurzhorn-Krötenechse [brachycercum] 
SynonymPhrynosoma (Tapaya) hernandesi GIRARD 1858: 395
Phrynosoma hernandezi — GIRARD 1858
Tapaya ornatissima GIRARD 1858: 396
Phrynosoma douglasii ornatum GIRARD 1858: 397
Phrynosoma douglassi — GENTRY 1885: 140 (part)
Phrynosoma douglassii hernandesi — COPE 1900: 413 (part)
Phrynosoma douglassii hernandesi — COPE 1900
Phrynosoma douglasii hernandesi — VAN DENBURGH 1922
Phrynosoma douglassii ornatissimum — VAN DENBURGH 1922: 377(part)
Phrynosoma douglasii hernandesi — BURT 1935
Phrynosoma orbiculare hernandesi — KLAUBER 1939: 91
Phrynosoma orbiculare hernandesi — MARR 1944
Phrynosoma douglassii brevirostre — SMITH 1946: 302 (part)
Phrynosoma douglassii ornatissimum — SMITH 1946: 305 (part)
Phrynosoma hernandesi — SMITH & TAYLOR 1950: 100
Phrynosoma douglassii ornatissimum — REEVE 1952: 927 (part)
Phrynosoma douglassii brevirostre — REEVE 1952: 913 (part)
Phrynosoma douglassi hernandesi — VAN DEVENDER & LOWE 1977
Phrynosoma douglassi hernandesi — TANNER 1987
Phrynosoma douglasii hernandesi — CONANT & COLLINS 1991: 114
Phrynosoma douglasii hernandesi — LINER 1994
Phrynosoma hernandesi — ZAMUDIO et al. 1997: 302 (part)
Phrynosoma hernandesi — SMITH et al. 1999
Phrynosoma hernandesi — COLLINS & TAGGART 2009
Phrynosoma (Tapaja) hernandesi — CROTHER et al. 2012
Phrynosoma bauri MONTANUCCI 2015
Phrynosoma (Tapaja) hernandezi bauri — DE QUEIROZ in CROTHER et al. 2017
Phrynosoma hernandesi — KÖHLER 2021
Phrynosoma hernandesi bauri — LEACHÉ et al. 2021
Phrynosoma hernandesi — HEIMES 2022

Phrynosoma hernandesi brachycercum (SMITH 1942)
Phrynosoma ornatissimum brachycercum SMITH 1942
Phrynosoma douglassii brachycercum SMITH 1942: 362
Phrynosoma douglassii brachycercum — REEVE 1952: 916
Phrynosoma douglassii hernandesi — REEVE 1952: 922 (part).
Phrynosoma douglassii brachycercum — WEBB 1984
Phrynosoma douglassi brachycercum — TANNER 1987
Phrynosoma douglassi brachycercum — SMITH 1994: 114
Phrynosoma hernandesi brachycercum LEMOS-ESPINAL et al. 2004: 8.
Phrynosoma ornatissimum brachycercum — MONTANUCCI 2015: 75

Phrynosoma hernandesi brevirostris GIRARD 1858
Phrynosoma (Tapaya) brevirostris GIRARD 1858: 397
Phrynosoma brevirostre — COPE 1867: 302
Phrynosoma douglassi — GENTRY 1885: 140 (part)
Phrynosoma douglassii hernandesi — COPE 1900: 413 (part)
Phrynosoma douglassii ornatissimum — VAN DENBURGH 1922: 377 (part)
Phrynosoma douglassii brevirostre — SMITH 1946: 302
Phrynosoma douglassii ornatissimum — SMITH 1946: 305 (part)
Phrynosoma douglassii brevirostre — REEVE 1952: 913
Phrynosoma douglassii hernandesi — REEVE 1952: 922 (part)
Phrynosoma douglassii ornatum — REEVE 1952: 930 (part)
Phrynosoma douglassii brevirostre — MASLIN 1964
Phrynosoma douglassi brevirostre — POWELL & RUSSELL 1985
Phrynosoma douglassii brevirostre — CONANT & COLLINS 1991: 113
Phrynosoma hernandezi — ZAMUDIO et al. 1997: 302 (part)
Phrynosoma brevirostris — MONTANUCCI 2015
Phrynosoma brevirostris — POWELL & RUSSELL 2023

Phrynosoma hernandesi hernandesi GIRARD 1858
Phrynosoma (Tapaya) hernandesi GIRARD 1858: 395
Phrynosoma (Tapaya) ornatissima GIRARD 1858: 396 (part)
Phrynosoma douglassi — GENTRY 1885: 140 (part)
Phrynosoma douglassii hernandesi — COPE 1900: 413
Phrynosoma douglassii ornatissimum — COPE 1900: 415 (part)
Phrynosoma douglassii ornatissimum — VAN DENBURGH 1922: 377 (part)Phrynosoma douglassii hernandesi — VAN DENBURGH 1922: 382
Phrynosoma douglassii hernandesi — SMITH 1946: 304
Phrynosoma douglassii ornatissimum — SMITH 1946: 305 (part)
Phrynosoma douglassii hernandesi — REEVE 1952: 922
Phrynosoma douglassii ornatissimum — REEVE 1952: 927 (part)
Phrynosoma douglassi hernandesi — ETHERIDGE 1964
Phrynosoma hernandezi — ZAMUDIO et al. 1997: 302
Phrynosoma hernandesi hernandesi — MONTANUCCI 2015: 51
Phrynosoma hernandesi hernandesi — KÖHLER 2021

Phrynosoma hernandesi diminutum MONTANUCCI 2015
Phrynosoma diminutum MONTANUCCI 2015
Phrynosoma douglassi — GENTRY 1885: 140 (part)
Phrynosoma douglassii hernandesi — COPE 1900: 413 (part)
Phrynosoma douglassii ornatissimum — VAN DENBURGH 1922: 377(part)
Phrynosoma douglassii ornatissimum — SMITH 1946: 305 (part)
Phrynosoma douglassii ornatissimum — REEVE 1952: 927 (part)
Phrynosoma hernandezi — ZAMUDIO et al. 1997: 302 (part)
Phrynosoma hernandesi diminutum — LEACHÉ et al. 2021 (see comment)

Phrynosoma hernandesi ornatum GIRARD 1858
Phrynosoma ornatum GIRARD 1858: pl. 21
Phrynosoma douglassi — GENTRY 1885:140 (part)
Phrynosoma douglassii hernandesi — COPE 1900:413 (part)
Phrynosoma douglassii hernandesi — RICHARDSON 1915:423 (part)
Phrynosoma douglassii ornatum — STEJNEGER 1919:3.
Phrynosoma douglassii ornatissimum — VAN DENBURGH 1922:377 (part)
Phrynosoma douglassii ornatum — SMITH 1946:307.
Phrynosoma douglassii ornatissimum — SMITH 1946:305 (part)
Phrynosoma douglassii douglassii — REEVE 1952:918 (part)
Phrynosoma douglassii hernandesi — REEVE 1952:922 (part)
Phrynosoma hernandezi — ZAMUDIO et al. 1997: 302 (part)
Phrynosoma hernandesi ornatum — MONTANUCCI 2015: 61
Phrynosoma hernandesi ornatum — KÖHLER 2021

Phrynosoma hernandesi ornatissimum GIRARD 1858
Phrynosoma (Tapaya) hernandesi GIRARD 1858: 395 (part)
Phrynosoma (Tapaya) ornatissima GIRARD 1858: 396
Phrynosoma ornatissima — COPE 1871: 305
Phrynosoma Douglassi subsp. ornatissima COPE 1878: 49
Phrynosoma douglassi — GENTRY 1885: 140 (part)
Phrynosoma douglassii hernandesi — COPE 1900: 413 (part)
Phrynosoma douglassii ornatissimum — COPE 1900: 415
Phrynosoma douglassii ornatissimum VAN DENBURGH 1922: 377
Phrynosoma douglassii hernandesi — VAN DENBURGH 1922: 382 (part)
Phrynosoma douglassi ornatissimum — KNOWLTON & JANES 1934
Phrynosoma douglassii hernandesi — SMITH 1946: 304 (part)
Phrynosoma douglassii ornatissimum — SMITH 1946: 305
Phrynosoma douglassii ornatissimum — REEVE 1952: 927
Phrynosoma hernandezi — ZAMUDIO et al. 1997: 302 (part)
Phrynosoma ornatissimum ornatissimum — MONTANUCCI 2015: 67
Phrynosoma hernandezi ornatissimum — DE QUEIROZ in CROTHER et al. 2017
Phrynosoma ornatissimum — HEIMES 2022 
DistributionCanada (SE Alberta, SW Saskatchewan)
USA (New Mexico, W Texas, Utah, Colorado, Arizona, South Dakota, Idaho, Nevada, Oregon)
Mexico (NE Sonora, Chihuahua)

bauri: USA (Colorado, Nebraska, SE Wyoming, NE New Mexico); Type locality: 12.8 km north of Orchard, Morgan County, Colorado

brachycercum: Mexico (Durango, Chihuahua, Zacatecas); Type locality: “Durango” (restricted to Durango City, Durango, Mexico by REEVE 1952)

brevirostris: Canada (S Alberta, S Sasketchewan),
USA (Wyoming, W North Dakota, W South Dakota, W Nebraska, Utah, Colorado); Type locality: Restricted to 9.6 km E of Agate, Sioux County, Nebraska by Montanucci 2015. Reeve (1952:913) emended the name of the type locality (Pole Creek) in accordance with its current geographic name, Lodgepole Creek, and restricted the type locality to near Dix, Kimball County, Nebraska.

diminutum: USA (Colorado, probably N New Mexico); Type locality: Medano Road, just outside Medano Ranch, Alamosa County, Colorado.

hernandesi: Canada (Alberta, Saskatchewan), USA (New Mexico, W Texas, Utah, Colorado, Arizona, South Dakota), Mexico (NE Sonora, Chihuahua; Type locality: New Mexico and Sonora. Restricted to Santa Fe, New Mexico by SMITH & TAYLOR 1950. Resetricted to Fort Huachuca, Cochise County, Arizona by MONTANUCCI 2015.

ornatissumum: USA (New Mexico, Texas); Type locality: restricted to the Rio Grande Valley at Albuquerque, Bernalillo County, New Mexico

ornatum: Idaho, Nevada, Oregon, Utah; Type locality: valley of the Great Salt Lake, Utah  
Reproductionovovivparous (Lambert & Wiens 2013). 
TypesLectotype: USNM 197; paralectotype: USNM 198 (Stejneger 1890:113 referred to USNM 107 and 198 as “Girard’s types” of P. hernandesi but the number 107 is a typographical error fide Reeve 1952:923, Montanucci 2015).
Holotype: UCM 11356, adult male, collected from 12.8 km north of Orchard, Morgan County, Colorado, by V. Janzen on 10 May 1958 (Fig. 11). Paratopotypes. UCM 11358–59, both males, collected by V. Janzen on 10 May 1958. [bauri]
Holotype: USNM 23993 [brachycercum]
Type: USNM 208; Lectotype: USNM 4592c (designated by Montanucci 2015: 28) [brevirostris]
Holotype: UCM 61895, adult female, collected from Medano Road, just outside The Nature Conservancy’s Medano Ranch, 2,308 m., Alamosa County, Colorado, by A. Schneider and A. Mitchell on 4 August 2006 (Fig. 14). Paratypes. LSUM 13834–37, 13839–40, MEL 1013, 1015, 1016, MVZ 27042, UCM 3894, 3898, 48465– 66, 51268, 61896, UMMZ 62242, 62244, 62247–52, 62255–58, 62261–64, USNM 8558, 44888, 44890. See Appendix I for locality data. [diminutum]
Holotype: USNM 234, a female; UMMZ 3849 fide Banta 1970 but in error fide Montanucci 2010 who concluded that USNM 234 is the holotype [ornatum]
Syntypes: (3) ANSP 8700-02. Lectotype: USNM 204 (designated by Montanucci 2015: 68);Girard (1858a) described this species on the basis of specimens in the USNM, but he did not designate a type specimen. Other syntypes: USNM 576284 [Tapaya ornatissima] 
DiagnosisDiagnosis: Phrynosoma h. hernandesi can be distinguished from other members of the P. douglasii species complex by the following combination of adult characters: (1) snout protruding, 49.4% ± 1.56 (42.5–57.3%) of orbit to rostral scale distance; (2) rostrofrontal profile rather flat, gradually sloping; (3) frontal rim usually elevated above the occipital shelf; (4) enlarged frontal rim scales 2.96 ± 0.15 (2–4) / 2.93 ± 0.13 (2–4); (5) temporal shelf long, 25.7% ± 0.96 (18.9–32.3%) in males, 26.1% ± 0.79 (17.2–36.2%) in females; (6) temporal shelf flat to weakly convex; (7) cephalic horns moderately long, third temporal horn length 18.2% ± 0.46 (10.3–24.1%); (8) cephalic horns directed horizontally or only slightly elevated; (9) tympanum elliptic, moderately broad to broad; (10) tympanum exposed; (11) tail moderately long, 247% ± 8.87 (187–301%) in males, 219% ± 4.84 (175–272%) in females; (12) dorsal spots wedge-shaped or forming transverse bands; (13) light-colored borders of dorsal spots confined to posterior edges; (14) dorsolateral white spots absent; (15) gular area with melanistic vermiculations and relatively few spots, with or without gray suffusion (melanin-dispersed phase); (16) abdomen with large melanistic spots or coalescing into dark suffusion (melanin-dispersed phase); (17) interrupted melanistic subcaudal bands present; bands may be complete distally (melanin-dispersed phase) [Montanucci 2015: 53].


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CommentSubspecies: The subspecies brevirostre, hernandesi, and ornatissimum are artificial assemblages of populations (Zamudio et al. 1997). Nuclear and mitochondrial DNA yields different tree topologies for this species. Leaché et al. 2021 found that diminutum nests within hernandesi in a mitochondrial tree as well as a nuclear tree, but the latter had only one sample while in the mitochondrial tree diminutum formed a monophyletic clade, hence we consider it as a subspecies, rather than synonym of hernandesi. The former subspecies P. d. brevirostre, ornatissimum, and P. d. ornatum have been synonymized with P. hernandesi by Zamudio et al. 1997 but brevirostris and ornatissimum were revalidated by Montanucci 2015 but Leaché et al. 2021 merged bauri and brevirostris into hernandesi. Based on geography and morphological similarities (Montanucci, 2015), it is likely P. o. brachycercum will be verified as conspecific with P. ornatissimum (Leaché et al. 2021).

Hybridization: Phrynosoma hernandesi hybridizes with P. ornatissimum in Arizona and with P. bauri in Colorado (Montanucci 2015).

Habitat: montane

Group: Belongs to the Tapaja clade fide LEACHE & MCGUIRE 2006.

Distribution: see maps 3, 5, 7, 9, 14, 18, 19 in Montanucci 2015: 154 ff and Leaché et al. 2021: 3 (fig. 1 and Supplementary Figure S5). See also locality lists in Montanucci 2015: 137 (hernandesi) and Montanucci 2015: 151 (ornatum). Not in Durango fide Lemos-Espinal (2018). See maps 1,2 in Montanucci 2015: 163 ff and Leaché et al. 2021.

Synonymy: after Montanucci 2015. Köhler synonymized brevirostris with P. hernandesi, which was confirmed by Leaché et al. 2021. Köhler 2021 also included the following species as synonym of Phrynosoma hernandesi: Phryno- soma ornatissima, Phrynosoma brevirostris, Phrynosoma douglassii brachycercum, Phrynosoma bauri, and Phrynosoma diminutum. Köhler 2021 synonymized Phrynosoma ornatissimum and brachycercum with Phrynosoma hernandesi hernandesi.

NCBI taxonID: 2816098 [ornatissima] 
EtymologyNamed after Francisco Hernandez (1514-1587), Spanish physician who travelled in Mexico to study Aztec herbal medicine and to collect biological specimens. The correct spelling of the specific epithet is with an “s” (see Smith et al. 1999).

The subspecific epithet diminutum, Latin perfect participle of dēmĭnŭo, dēmĭnuěre, meaning “diminutive,” is in reference to the small adult size of this species.

ornatum: The Latin word ornatus –a-um, (participle of the verb orno), meaning “adorned” or “decorated” or “embellished”. As no type description exists, one can only speculate as to Girard’s intended reference, but possibly to contrasting dark and light-colored markings on the dorsum.

The specific epithet brevirostris comes from the Latin adjective brevis,-e, meaning “short”, and a modified version of the second declension Latin neuter noun rostrum, meaning “snout”, in reference to the short or abbreviated snout of this species. Apparently, Girard’s (1858a) intent was to create a compound noun in apposition to the subgeneric name Tapaya (see Smith & Reeve, 1951 for origin of the name), crafting his new name in third declension feminine gender since Tapaya is apparently feminine. Although Girard’s root word rostris does not exist in Latin, Cope (1866:302) modified Girard’s name to brevirostre as though it were an adjective needing to agree with the neuter gender of Phrynosoma. However, since nouns used in apposition to other nouns retain their own gender (only adjectives must agree in gender with the name of the genus), Cope’s alteration of the name was an unnecessary and incorrect subsequent spelling. Therefore, Montanucci 2015 retained Girard’s original spelling for the available name. See ICZN articles 31.2.1. and 34.2.1.

The subspecific epithet bauri is a Latin patronym in the genitive singular, honoring the late Bertrand E. Baur, a longtime friend and avid student of Phrynosoma. Baur died in November of 2013 due to injuries sustained in a car accident the previous year.

ornatissima: The Latin word ornatus –a – um, (participle of the verb orno), with the neuter superlative suffix,-issimum, meaning “the most highly adorned” or “...decorated” or “... embellished.”

brachycercum: The Greek word roots βραχύς = brachys an adjective meaning “short” and κέρκος = kerkos or cercus, a feminine noun meaning “tail”, in reference to the very short tail in this race. Smith (1942) modified brachycercus (used as a trinomial) to brachycercum as though it were an adjective needing to agree with the neuter gender of Phrynosoma. Although nouns used in apposition to other nouns retain their own gender (only adjectives must agree in gender with the name of the genus) such that Smith’s modification of the name was unnecessary, his use of the word as an adjective must be followed. See ICZN articles 31.2.1 and 34.2.1. 
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