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Osteology and Myology of the Head and Neck Regions of Callisaurus, Cophosaurus, Holbrookia, and Uma (Reptilia: Iguanidae)

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Osteology and Myology of the Head and Neck Regions of Callisaurus, Cophosaurus, Holbrookia, and Uma (Reptilia: Iguanidae) Great Basin Naturalist Volume 37 Number 1 Article 3 3-31-1977 Osteology and myology of the head and neck regions of Callisaurus, Cophosaurus, Holbrookia, and Uma (Reptilia: Iguanidae) Douglas C. Cox Brigham Young University Wilmer W. Tanner Brigham Young University Follow this and additional works at: https://scholarsarchive.byu.edu/gbn Recommended Citation Cox, Douglas C. and Tanner, Wilmer W. (1977) "Osteology and myology of the head and neck regions of Callisaurus, Cophosaurus, Holbrookia, and Uma (Reptilia: Iguanidae)," Great Basin Naturalist: Vol. 37 : No. 1 , Article 3. Available at: https://scholarsarchive.byu.edu/gbn/vol37/iss1/3 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. OSTEOLOGY AND MYOLOGY OF THE HEAD AND NECK REGIONS OF CALLISAURUS, COPHOSAURUS, HOLBROOKIA, AND UMA (REPTILIA: IGUANIDAE) Douglas C. Cox' and Wilmer W. Tanner^ Abstract.- A detailed study of the anterior osteology and myology of Callisatmis, Copho.saurus, Holbrookia, and Vma reveals the phylogenetic relationships among the sand lizards. An SPSS discriminant analysis of os- teological characters combined with myological characters indicates that Callimiirtis is most primitive, Cop- hosaurus and Holbrookia are most closely related, and Uma is the most distinct of the sand lizard genera. Be- cause of close relationships between Cophosaurus and Holbrookia, it is postulated that earlessness evolved once, and Cophosaurus is returned to synonymy under Holbrookia. Blainville (1835) wrote the first descrip- (1932) Holbrookia propinqua stonei; Smith tion of a sand lizard and named it Calli- (1935, 1943, 1946:137, 145) Holbrookia ele- saurus draconoides. Since then various au- gans thermophila, Holbrookia elegans ele- thors have pubhshed articles concerning gans, Holbrookia bunkeri, Holbrookia macu- sand lizards. Girard (1851) described Hol- lata ruthveni, Holbrookia maculata brookia maciilata, Trochel (1852) described dickersonae, Holbrookia maculata pulchra, Cophosaurus texanus, and that same year and Holbrookia maculata thermophila; Lins- Baird and Girard synonymized Cophosaurus dale (1940) Callisaurus draconoides myurus with Holbrookia, providing the name Hol- and Callisaurus draconoides gabbii; Heifetz brookia texana, which then remained un- (1941) Uma notata notata; Bogert and Dor- changed for over 100 years. Subsequently, som (1942) Callisaurus draconoides brevipes; Baird (1858) described Uma notata. Smith and Burger (1950) Holbrookia prop- By 1858 all genera represented in the inqua propinqua and Holbrookia propinqua sand lizard group had been described. Since piperata; Peters (1951) Holbrookia texana then new species and subspecies, as well as texana and Holbrookia texana scitula; Axtell new combinations, have been added by var- (1956) Holbrookia lacerata, Holbrookia lacer- ious authors as follows: Bocourt (1874) Hol- ata subcaudalis and Holbrookia maculata brookia elegans- Cope (1880, 1883, 1894, perspicua; Smith and Cochran (1956) Calli- 1895, 1896, 1900) Holbrookia lacerata, Hol- saurus draconoides rhodostictus; and Wil- brookia maculata fkwilenta, Uma scoparia, liams, et al. (1959) Uma paraphygus. Uma inornata, Uma rufopuncatata, Calli- Some summaries, reviews, checklists, and saurus crinitus, Callisaurus rhodostictus, comparative studies have also been written. Holbrookia maculata maculata, Callisaurus Cope (1896) synonymized Uina and Calli- draconoides ventralis, and Callisaurus ven- saurus in a short paper di.scussing the genus tralis gabbii; Stejneger (1890) Holbrookia Callisaurus. He recognized Uma again in maculata approximans and Holbrookia his large work on the crocodilians, lizards, maculata lacerata; Richardson (1915) Calli- and snakes of North America (1900) and saurus ventralis myurus; Dikerson (1919) recognized one species and three subspecies Callisaurus carmenensis; Schmidt (1921, of Callisaurus. 1922) Holbrookia maculata campi, Hol- brookia pulchra, Holbrookia dickersonae, and Smith (1946:137, 145), in his "Handbook Callisaurus ventralis inusitatus; Schmidt of Lizards," recognized in Callisaurus one and Bogert (1947) Uma exsul; Barbour species and ten subspecies and stated (p. (1921) Holbrookia thermophila; Harper 145): 'The whole group of Callisaurus of 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 'Life Science Museum, Brigham Young University, Provo, Utah 84602. 35 36 Great Basin Naturalist Vol. 37, No. 1 western United States is in need of revision. U. n. rufopunctata, U. n. inornata, U. sco- The subspecies are not adequately charac- paria, and U. exsul. He discussed the evolu- terized, nor are their ranges well worked tion of Uma and its relationship to other out. There is very little information on the sand lizards. life history." Although we now know much The conflict relative to the classification more about the life history of CaUisaurus, it of the notata-scoparia group was discussed is still in need of a comprehensive tax- by Mayhew (1964b). He recognized U. in- onomic review and remains a monotypic ornata, U. notata, and U. scoparia all as full genus. (A careful revision for CaUisaurus species on the basis of temperature toler- has not yet been attempted.) ance and reproductive data. The genus Holbrookia has had two revi- Peters (1951) reviewed Holbrookia texana sions. Schmidt (1922) made the first; a sec- {Cophosaurus texanus). He described two sub- ond was by Axtell (1958). Schmidt (p. 709) species, but mentions little concerning rela- stated: "The taxonomy of the North Ameri- tionships with other sand lizards. can lizards of the genus Holbrookia Girard Clarke (1965) revived Cophosaurus texan- offers one of the most interesting and diffi- us on the basis of behavioral data collected cult problems in North American herpeto- in a large comparative study of the sand logy." fizard group. He recognized several species and states Ecological and behavioral studies con- that (p. 712) ^'Holbrookia is obviously di- cerning sand lizards have also rectly related to CaUisaurus, from which it been pub- differs only in the concealed tympanum and lished by Burt (1931a, 1931b), Stebbins 1954, with which it agrees in general features of (1944, 1966), Ramsey (1948, 1949), color pattern and scutellation." Cagle (1950), Williams and Smith (1958), Smith (1946:137, 145) doubts that a prac- Axtell (1960), Lannom (1962), Carpenter tical means of characterizing the species ex- (1963, 1967), Clarke (1965), Mayhew (1964a, 1964b, Pianka ists and believes that "until such a means is 1966), and Parker found there will remain indefinitely a prob- (1972), Tanner and Krogh (1975), and Judd lem in defining the ranges of the several (1974, 1975). subspecies, or in defending their actual The anatomy of these lizards has not validity. Accordingly the genus, par- been thoroughly studied. Earle, in a series ticularly the maculata group, merits a care- of articles (1961a, 1961b, 1961c, 1962), de- ful study perhaps more than any other in scribed in detail the comparative anatomy the United States." of the middle ear of sand lizards. Stebbins Axtell (1958) recognized only three spe- (1943, 1944) described the nasal structures cies, HoUjrookia lacerata, H. propinqua, and and some aspects of the ecology of Uma, H. maculata. He considered H. texana to be then (1948) described the nasal structures of a species of the genus CaUisaurus and lizards in general, which included the sand therefore did not discuss it. He did list two lizards. Axtell (1958) described the osteology subspecies for lacerata, two for propinqua, of Holbrookia and stated that it is essentially and 11 for maculata. the same as found in all sand lizards. Eth- Uma has been reviewed by Heifetz ridge (1964) studied the skeletal morphology (1941), Norris (1958), and Mayhew (1964b). of sceloporine lizards, which includes sand Heifetz recognized U. notata notata, U. no- lizards, and compared their relationships. tata cowlesi, U. inornata, and U. scoparia. Savage (1958) studied Urosaurus and Uta He also referred to the taxonomic confusion and made remarks concerning related gen- that exists because of erroneous type local- era, which included sand lizards. A few ref- ities in this genus. In his checklist Schmidt erences to sand lizards were made by Lar- (1953) recognized only one species (notata), sen and Tanner (1974) while studying with three subspecies {notata, inornata, and Sceloporus, and Guttman (1970b) also refers scoparia). to them in his electrophoretic study of the Norris (1958) recognized Uma n. notata. hemoglobins of sand lizards. He found that March 1977 Cox: Lizard Morphology 37 all genera possessed the same major and mi- from San Bernardino Co., California: (BYU) nor protein components. 11389, (CAS) 42072. Only portions of the osteology have been Skeletal material was prepared by careful adequately treated; the myology is essen- dissection. Bones were cleaned with forceps tially untouched. The objectives of this and dissecting needles and soaked in Clorox study are to: (1) describe the skull osteology bleach for several minutes to loosen soft tis- and branchiomeric myology of sand lizards; sues, after which hirther picking and clean- (2) identify osteological and myological ing was done. Skulls were not allowed to characteristics that distinguish the sand liz- dry, but were preserved
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