RE-: SQUAMATA: SAURIA: IGUANIDAE DIPSOSAURUS, D. DORSALIS Catalogue of American Amphibians and Reptiles. small and keeled, with a medial row of slightly enlarged, raised scales forming a weak to moderate keel extending from the nuchal region Hulse, A.C. 1992. Dipswums, D. dorsalis. to the base of the tail. Ventral scales are smooth and imbricate, twice as large as the dorsals. Caudal scales are larger than the body scales, Dipsosaurus Hallowell keeled, equal in size, and arranged in rings. The supratemporals are Desert Iguana at the posterior surface of the supratemporal process of the parietals and the parietal foramen is in the frontal bones. The outline of the Dips-saumH allowell, 1854:92. Type-species, Cmtaphytusdotsalis osseous labyrinth is moderately distinct. The coronoid has a large Baird and Girard, 1852, by monotypy. lateral blade. The splenial is reduced and the borders of Meckel's Dipsosaums: Hallowell, 1859:7. Emendation. groove are fused to its anterior border. Meckel's groove is closed for more than halfthe distance between the splenial and the mandibular Content. One species, D. dossalis, is recognized. symphysis. Palatine teeth are lost, as may be the pterygoid teeth. The crowns of the posterior teeth are expanded and have secondary Definition. Dipsosaurusis a large lizard (maximum SVL 135 cusps. Ribs have been lost from the third and fourth cervical mm) with a short, blunt head. The tail is twice as long as the body. vertebrae. A scapular fenestra is present, but a posterior coracoid Forelegs are short, half the length of the hindlegs. Dorsal scales are fenestra is lacking. Map. The large open circles indicate type-localities or restricted type-localities, solid circles other records. The fossil locality is indicated by the star. An uncertain fossil locality from "near Agate, Nebraskan is not mapped (see text for details). Dorsal background coloration is grayish to grayish brown with inner ear. Norell (1989) provided black and white photographs of a patternof pale spotssurrounded by dark rings. Laterally this pattern Pliocene fossil and comparative Recent material. dissolves into a series of dark broken lines. The ventral surface is immaculate cream to beige. Distribution. Dipsosaums dorsalis ranges from southern Nevada, extreme southwestern Utah along the Virgin River in the Diagnosis. Dipsosaums can be distinguished from other vicinity of Beaver Dam Wash, and California east of the Sierra Nevada iguanines by the following combination of characters (de Queiroz, andcoast ranges, southwardthrough central California to theeastern 1987): one subocular scale much longer than the others; superciliary coast of Baja California del Norte and from there to the tip of Baja scales greatly elongate and strongly overlapping; pterygoid teeth California del Sur in the vicinity of Cabo San Lucas. The species is usually absent or, when present, lying along the medial edge of the restricted to the eastern side of the central mountains (Sierra San pterygoid; crowns of posterior marginal teeth with four cusps; large, Pedro MPrtir, Sierra Columbia, and Sierra de la Giganta) throughout paired openings present at or near frontonasal suture; parietal fora- most of its range in Baja California. It is present on many of the land men located entirely within frontal bone; lateral process of palatine bridges and deep water islands in the Sea of Cortez, but conspicuous behind infraorbitalf oramen small or absent; lateral pointed processes by its absence from IslaTibur6n (Murphy and Ottley, 1984). The only present on the cristae interfenestralis; posterior ends of lateral and Pacific islands on which the species has been found are Magdalena medial crests of retroarticular process divergent ontogenetically to andSta. Margarita. These lizardsare known from western andcentral form quadrangular outline. Arizona southward through Sonora, west of the Sierra Madre Occi- dental, to extreme northern Sinaloa, Mexico in the vicinity of Descriptions, Illustrtions, Distribution, Fossil Record, Tupolobampo. The northern limit of the species' range appears to and Pertinent Literature. See species account. coincide with that of Lama divaricata (Norris, 1953). Muth (1980) presented evidence that D. dorsalis may be limited by edaphic Etymology. The name Dipsosaums is derived from the conditions and associated hatching success. Greek words dipsa (- dryorthirsty) and saums(- lizard), apparently in reference to the arid habitats in which members ofthe genus occur. FossilRecord Norell (1989) reported Pliocene fossils from San Diego County, California. An early Miocene fossil from "near Agate, Nebraska," originally described as Tetralopbosaums minutus Dipsosaurus abrsalis (Baird and Girard) (Olson, 1937) may be referable to this species (de Queiroz, 1987; Desert Iguana Norell, 1989). Norell and de Queiroz (1991) described a fossil from the early Miocene of New Mexico which appears to represent the CmtapbytusdorsalisBairda nd Girard, 1852:126. Vpe-locality, "Des- sister group of Dipsosaums within the Iguaninae. ert of Colorado, California," restricted to Winterhaven (-Fort Yuma), Imperial County, California by Smith and Taylor (1950). Pertinent Literature. Physiology of this species has been Holotype, National MuseumofNatural History(USNM) 2699, an studied extensivelya s follows: thermoregulationb y Mosauer (19361, adult of indeterminate sex (see Comment), collected by John L. Cowles and Bogert (1944), Dawson and Bartholomew (19581, Licht Leconte, date of collection unknown (examined by author). (1%4,1965,1972), Brattstrom (1969, McGinnis and Dickson (1967), Dipso-sauna dorsalis: Hallowell, 1854:92. Porter (1967), DeWin (1967, 19711, Kemp (1969, 1972), Weathers Dipsosaurus dorsalis: Hallowell, 1859:7. (1970,1971), McGinnis and Palkenstein (19711, Kluger et al. (19731, Diplosaurus dorsalist Wong, Vorhaben, and Campbell, 1975:395. Berk and Heath (1975a,b), Callard et al. (19751, Barber and Crawford Lapsus. (1979). and Dupre and Crawford (1985); water balance and osmo- q regulation by Templeton (1960, 1966), Mimich (1970a,b), Murrish Content. Five subspecies are recognized: dorsalis, and Schmidt-Nielsen (1970). Templeton et al. (1972a). Shoemaker et carmenem& catalinensis, lucasensis, and sonon'emis. al. (1972), Munsey(1972), E Ian~d~ oertemil~(e1r9 74); and Maderson et al. (1978); metabolismby Bennett and Dawson (19721, Bennett and Definition and Diagnosis. See generic account. Licht (1972), John-Alder (1984), Gleeson (1985), and Gleeson and Dalessio (1989); endocrinology by Lisk (1967), McWhiinie and Description. The original descriptions (Baird and Girard, Cortelyou (1968), Chan et al. (19701, Bradshaw (19721, Callard and 1852; Hallowell, 1854) were brief. Detailed descriptions of external Chan (1972), Gerstle and Callard (19721, Templeton et al. (1968, characteristics are in Cope (lNO), Van Denburgh (1922), Woodbury 1972b), Walker (1973), Yousef et al. (1973), and John-Alder (1984); (1931), and Smith (1946). An extensive description of selected blood vascular properties by Dawson and Poulson (19621, Pough external and osteological characters within a phylogenetic frame- (1%9b, 1979), Dessauer (1970), Withers (1978), and Bickler (1986); work was provided by de Queiroz (1987). Stebbins (1954, 19851, immune response and fever by Evans and Cowles (19591, Evans CochranandGoin (1970), Leviton (19721, and Behler and King (1979) (l%3), Wright and Schapiro (1973), Vaughn et al. (1974), and gave general descriptions. Cope (1892) and Camp (1923) com- Bernheim and Kluger (1976); muscle physiology by Adams (1987, mented on osteology. Etheridge (1967) described caudal vertebrae. 1989), Gleeson and Johnston (1987), Gleeson and Harrison (19881, Miller (1966) described cochlear ducts and Iverson (1980) discussed and Marsh (1988); and sleep by Huntley (1987). Anatomical studies colic modifications. Peterson (1984) discussed scale microstructure. have included: osteology by Etheridge (19671, Hoffstetter and Gasc (l%9), and AveryandTamer (1971); musculature by Pough (1969a) Illustrations. Cochran and Goin (1970), Behler and King and Avery and Tanner (1971); digestive system by Iverson (1980); (1979), Smith and Brodie (1982), Burghardt and Rand (1982), Switak dentition by Taylor (1940), Hotton (1955), and Edmund (1969); (1984), and Obst et al. (1988) provided color photographs and/or epidermis by Maderson et al. (19701, Stewart and Daniel (19751, and illustrations. Black and white photographs andlor illustrations are in Peterson (1984); nasal structure by Stebbins (1948), Savage (19581, Van Denburgh (1922), Woodbury (19311, Ditmars (1936), Smith Parsons (1970), and Ulinksi and Peterson (1981); cardio-vascular (1946), Pickwell (1947), Norris (1953), Stebbins (1954,19851, Leviton system by Zug (1971); urogenital system by Bolton and Beuchat (1972), Howland (1988), and Obst et al. (1988). Black and white (191); ear by Schmidt (1964) and Wever (1978); and coloration by photographs of typical habitat are provided by Gloyd (1937) and Norris (1967). Banta (1962). Hallowell (1859), Cope (19001, and Smith (1946) Cowles and Bogert (1936), Cowles (1941), Norris (1953), Car- presented line drawings of scutellation. Osteology was illustrated by penter(l961) Krekorianet al. (19681, Minnichand Shoemaker(1970), Etheridge (1967) and Avery and Tanner (1971); musculature by the Moehn (1974), Gelderloos (1976), and Alberts (1989, 1992) studied latter, Pough (1969a), and Russell (1988). Norris (1953) and Carpen- behavior. Carothers (1984) employed data on Dipsosaumsin a study ter (1961)presented behavioral postures. SchwenkandThrockmorton of sexual selection in herbivorous lizards. Ecological studies have (1989) provided an extensive series of black and white photographs included: general ecology by Norris (19531, Pianka (1971, 19861, and line drawings illustrating feeding kinematics. Iverson (1980) Parker (1972), Luckenbach and Bury (19831, and Krekorian (1984); included a photograph of colonic valves. Irish et al. (1988) provided home range and homing by Krekorian (1976,1977); chemoreception n a series of scanning electronmicrographs illustrating the skin shed- and femoralg landsecretions by Krekorian (1989),Alberts (1991), and ding cycle. Other electronmicrographso f scales are in Stewart and DussaultandKrekorian (191); reproductionby Shaw(1952), Mayhew Daniel (1975) and Peterson (1984), and of egg shells in Packard et al. (1964, 1971), Pitch (1970), Minnich (1971), Licht (19731, and Muth (1982). Savage (1958) and Parsons (1970) illustrated nasal structures. (1977, 1980, 1981); feeding and diet by Norris (1953), Minnich and Mayhew (1971) illustrated spermiogenesis and Schmidt (1964) the Shoemaker (1970), Pianka (1971), Karasov et al. (19861, Mautz and - Nagy (1987), and Cooper and Alberts (1990); physiological ecology having fewer scales around the body (X- 88.7f 2.8, N 411, from D. by Minnich (1970a), Muth (1980,1981), Bradshaw (19861, and Mautz d. lucarensis in having a single row of scales separating the rostral and Nagy (1987); and density by Krekorian (1983). Baker (1987) from the nasal ring and by having the postmentals usually separated - provided a synopsis of nematodes parasitizing the species. Lefcort by small granular scales, and from D. d. carmenensis in having a and Blaustein (1991) employed data on this species in a study of the lower number of femoral pores (X- 38.0f 1.9, N 41) and by having relationship between parasite load and sexual selection. Janzen and the postmentals usually separated by small granular scales. Paukstis (1991) included this species in a study of environmentals ex determination in reptiles. Phylogenetic relationships have recently Remark. Distribution is restricted to the type-locality. been discussed by de Queiroz (1987), Etheridge and de Queiroz (1988), Frost and Etheridge (19891, and Lang (1989). Hulse (1988) 4. Dipsosaurus dorsalis lucasensis provided a bibliography and Malkin (1956) discussed ethnozoology. Van Denburgh Etyxnology. The name dorsalis is derived from dorsum (L., Dipsosaumsdorsalis lucasenis Van Denburgh, 1920:33. Type-local- 'the back) and refers to the enlarged row of vertebral scales. The ity, "San Jose del Cabo, Lower California, Mexico." Holotype, names lucasensis, catalinensis, carmenensis,a nd sonoriensisare all California Academy of Sciences (CAS) 46090, an adult male col- geographical names referring respectively to the region around Cabo lected by Joseph R Slevin on 31J uly 1919 (not examined by au- San Lucas, Baja California del Sur; Isla Catalina; Isla Carmen; and to thor). the Mexican state of Sonora. Diagnosis. This subspeciesdiffers from D. d. dorsalisand D. Comment. Additional studies are needed to validate the - d. sonoriensisi n having a lower number of scales around the body (X taxonomic status of the Baja California del Sur forms (D. d. lucasensis, 88.8f 4.2, N- 53), from D. d. carm-e nensisinhavingalower number D. d. catalinensis, and D. d. carmenensis). of femoral pores (X- 37.5 f 2.3, N 53), and from D. d. catalinensis The holotype of D. d. dorsalis is poorly preserved and as a in having a single row of scales separating the rostral from the nasal consequence its' sex can't be determined. ring and in usually having the postmentals in contact rather than separated by small scales. 1. Dipsosaurus dorsalis dorsalis (Baird and Girard) Remarks. Asplund (1967) discussed reproduction, food habits and behavior. Murray (1955) discussed coloration and Scudder Ctvtaphytusd orsalis Baird and Girard, 1852:126. See species synon- et al. (1983) compared mainland populations with those from Isla ymy. Cerralvo. Dipsosaums dorsalis dorsalis: Van Denburgh, 1920:33. First use of The distribution of the subspecies is restricted to the southern trinomial. two thirds of Baja California del Sur, several islands in the Sea of Cortez (Murphy and Onley, 1984) and the Pacific Coast islands of Diagnosis. This subspecies differs from D. d. sonoriensisin Magdalena and Sta. Margarita. usually having the rostrals and nasals separated by -tw o rows of small scales, in having 2-3 rows of scales (X- 2.8f 0.3, N 31) between the 5. Dipsosaurus dorsalis sonoriensis Allen suboculars and supralabials, a-n d in having a greater number of su-p ralabials (X- 10.1 + 0.79, N 31) and infralabials (X- 10.3f 0.83, Dipso-saumsdorsalissonoriensisA llen, 1933:4. Type-locality,' Her- N 31); and differs from all other subsp-e cies in having- a much higher mosillo, Sonora, Mexico," restricted to 15misoutheastofHermo- number of scale rows at midbody (X 107 f 4.6, N 31). sillo by Peters (1952). Holotype, University of Michigan Muse- um of Zoology (UMMZ) 72121, an adult male collected by 2. Dipsosaurus dorsalis carmenensis J. Piatt, J. Scofield, and M.J. Allen on 1J uly 1932 (examined by Van Denburgh author). Dipsosaums dorsalis sonoraensk Malkin, 19%:80. Lapsus. Dipsosaums carmenensis Van Denburgh, 1922:81. Type-locality, 'Near Puerto Bellandro, Carmen Island, Gulf of California, Mex- Diagnosis. This subspecies differs from D. d. dorsalisby the ico." Holotype, California Academy of Sciences (CAS) 50504, an following combination of characters: usually one row of scales - adult male collected by Joseph R. Slevin on 21 May 1921 (not separatingthe rostral from the nasals, 1-2 rows ofscales (X- 1.5f 0.5, examined by author). N 34) separating the suboculars fro-m the supralabials, a lower Dipsosaums dorsalis carmenensis: Savage and Cliff, 1954:74. First number- o f supralabials( X- 8.4 f 0.6, N 34) and infralabials (X = 8.3 use of trinomial. f 0.8, N 34); and differs from all other -su bspecies by h-a ving a higher number of scales around the body (X 108 f 4.9, N 34). Diagnosis. This subspeciesdiffers fromD. d. dorsalisand D. -d . sonoriensisi n -h aving a lower number of scales around the body (X Remarks. Some confusion exists concerning the type- 92.9 f 4.9, N 12); and differs from D. d. luca-s ensis in having a locality and date of collection of the type-specimen. Peters (1952) higher number of femoral pores (X- 43.1 f 2.1, N 121, and from the restricted the type-localityto 15 mi southeast of Hermosillo; however, former in having the postmentals in contact rather than being sepa- the locality most recently recognized by the University of Michigan rated by small scales. Museum of Zoology is 8.1 km SE of Hermosillo (Kluge, 1984). In addition, Morrow gave the date of collection as July 1, 1932, but the Remark. Distribution is restricted to Isla Carmen and Isla date in both the museum records and Peters (1952) is June 21-24, Coronados in the Sea of Cortez. 1932. This subspecies is restricted to the arid lowland regions of 3. Dipsosaurus dorsalis catalinensis central and southern Sonora and extreme northern Sinaloa, Mkxico, west of the Sierra Madre Occidental. Bogert and Oliver (1945), Van Denburgh Langebartel and Smith (19541, and Hardy and McDiarmid (1969) discussed the range of the subspecies; the latter provided a locality Dipsosaums catalinensis Van Denburgh, 1922:83. Type-locality, map for Sinaloa. "Santa Catalina Island, Gulf of California, Mexico." Holotype, California Academy of Sciences (CAS) 50505, an adult male col- Literature Cited lected by Joseph R. Slevin on 12 June 1921 (not examined by author). Adams, BA. 1987. Thermal dependence of passive electrical proper- Dipsaaums dorsalis catalinensis: Soule and Sloan, 1966:141. First ties of lizard muscle fibres. J. Exp. Biol. 133:169-182. use of trinomial. -1 989.T emperature effects on membrane chloride conductance and electrical excitability of lizardskeletal muscle fibers. J. Exp. Diagnosis. This subspecies differs from all others by having Biol. 144551-564. the medial region of the chin heavily suffused with dark brown or Alberts, A.C. 1989. Ultraviolet visual sensitivity in Desert Iguanas: im- chocolate pigmentation which extends into the anterior gular region; plications for pheromone detection. Anim. Behav. 38:129-137. and differs specifically from D. d. dorsalis and D. d. sonoriensis in -1 991. Phylogenetic andadaptivevariation inlizard femoral gland secretions. Copeia 191:69-79. stimuli by an herbivorous actively foraging lizard, Dipsosaums -. 1992. Pheromonal self-recognition in Desert Iguanas. Copeia dorsalis. Herpetologica 46:259-266. 1392:229-232. Cope, E.D. 1892. The osteology of the Lacertilia. Proc. Amer. Philos. Allen, M.J. 1933. 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