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The user has requested enhancement of the downloaded file. HERPETOFAUNA OF THE 100-MILE CIRCLE Desert Iguana, Dipsosaurus dorsalis (Baird and Girard, 1852) Robert L. Bezy, Natural History Museum of Los Angeles County, Los Angeles, CA 90007, USA; E-mail: [email protected]

The June sun rises like a ball of fire scorching the parched creosote flats of the Sonoran Desert. Soon cica- das commence their song of the dry, and Common Side- blotched The June sun rises (Uta stansburiana) like a ball of fire seek refuge in cool, moist underground scorching the retreats. By mid- parched creosote morning Tiger Whiptails (Aspidos- flats of the celis tigris) confine Sonoran Desert. their jerky foraging to shade patches, Soon cicadas and Zebra-tailed Figure 1. Desert Iguana (Dipsosaurus dorsalis), Yuma Co., Arizona. Photo by Erik F. Enderson. commence Lizards (Callisaurus draconoides) raise their toes off the searing desert pave- this thermophilic herbivorous (Norris 1953). their song of ment. But the large, pale “thirsty lizard” (Dipsosaurus He traveled throughout the ’ range from the the dry, and dorsalis, Desert Iguana, Figure 1) is just issuing forth Mohave Desert to Sinaloa and the cape of Baja Cali- from its burrow to bask in the hot sun. When its fornia Sur, but his work was centered in the Sonoran Common Side- body temperature reaches 40°C (104°F) the herbivore Desert of southern California where he firmly estab- blotched Lizards climbs into a Creosotebush to browse on the few lished an understanding of the basic features of the remaining yellow blossoms. Later it switches tactics, life of the Desert Iguana. Knowledge of its ecology seek refuge facing into the sun to minimize the surface area of its has been filled out by a large number of subsequent in cool, moist body that is exposed to the rays. The lizard remains workers in southern California, particularly Mayhew active into the heat of the early afternoon, and its (1965, 1971), McGinnis and Dickson (1967), Pianka underground body temperature may reach 46.4°C (116°F), the high- (1971), Berke and Heath (1975), Krekorian (1976, retreats. By est recorded for a . 1977, 1983, 1984), Muth (1977, 1980), Mautz and Nagy The importance of temperature tolerance in the (1987), and Howland (1988). As is the case for most mid-morning ecology and evolution of desert was articulat- lizards, little published research exists for Arizona Tiger Whiptails ed by Ray Cowles (1940a, 1940b). In 1944, the theme populations, and the effect of the monsoons and of behavioral thermal regulation was developed summer annuals on its ecology can be estimated only confine their jerky in grand style by Cowles and his former graduate by a few observations in Baja California Sur (Asplund foraging to shade student Chuck Bogert (see Pough 1974, Myers and 1967, Grismer 2002). Parker (1972) presented infor- Zweifel 1993, Mole and Rubio 2006), much to the mation, primarily on hatchlings, gathered incidental patches... dismay of Chuck Lowe, who also was a student of to his study of the Western Banded Gecko (Coleonyx Cowles. After Bogert (1949) introduced the Schultheis variegatus) in South Mountain Park, Maricopa Co., rapid-reading cloacal thermometer, “noose-em and Arizona, and a useful species summary and bibliogra- goose-em” became a herpetological cottage industry, phy were provided by Hulse (1988, 1992). and Lowe brought the zeal for thermal ecology to the Norris (1953) pointed out the wide spectrum of “Creosotebush League” when he joined the faculty of habitats occupied by the Desert Iguana, but empha- the University of Arizona in 1950 (see Rosen 2004). sized that the species is closely associated with the But the stellar role of the Desert Iguana in reptile Creosotebush (Larrea tridentata) in the Southwestern thermal biology remained little appreciated until Ken deserts where the flowers comprise a major part of Norris, also a former student of Cowles and later fac- the diet of this herbivorous reptile. The long tail fa- ulty member at UCLA, had the fortitude to conduct cilitates climbing in the bushes to forage or to escape and publish the first thorough study of the ecology of the extreme heat of the desert floor. The mottled

SONORAN HERPETOLOGIST 23 (10) 2010 136 Unlike most other lizards, the home range of females (1,558 m2 [0.38 acre]) is nearly identical in size to that of males (1,462 m2 [0.36 acre]), and this probably reflects the near lack of size sexual dimor- phism in the species (Kreko- rian 1976). Male home ranges overlap extensively, whereas those of females do not. Al- berts (1989) has demonstrated that territories are marked by males with waxy secretions from the femoral pores that absorb UV light that is vis- Figure 2. Desert Iguana (Dipsosaurus dorsalis) in dappled sunlight beneath ible to Desert Iguanas. Gier a Creosotebush (Larrea tridentata), Yuma Co., Arizona. Photo by Erik F. Enderson. found that males who establish The daily activity pattern of the dorsal surface is well camouflaged in territories around large shade plants have access to a pattern of Desert the dappled sunlight beneath the creosote (Figure 2), greater number of females (Pianka and Vitt 2003). where the lizards’ burrows are usually located. The Estimates of population densities vary consider- Iguanas differs burrows play an important role in the ecology of the ably among localities and years. Krekorian (1983, from that of Desert Iguana. Some may have been dug originally 1984) found 300-700 individuals per hectare in the by Round-tailed Ground Squirrels (Xerospermophilus Coachella Valley, Riverside Co., California, whereas most other tereticaudus) or Desert Kangaroo Rats (Dipodomys 145 km (90 mi) to the east in the more arid Chuck- Sonoran Desert deserti), but Norris found that most were quite shallow walla Valley, Howland (1988) found only 60 lizards and appeared to have been excavated by the lizard, per hectare, presumably due to lower food availability. lizards in that particularly in sandy soils. They usually contained a The daily activity pattern of Desert Iguanas dif- it is unimodal, distinct chamber just a few inches below the surface, fers from that of most other Sonoran Desert lizards and the entrances were seen to have been plugged in that it is unimodal, centered in the mid to late centered in by sand. Smaller Desert Iguanas occupied smaller morning. In April most activity occurs from 0930 to the mid to late burrows, additional evidence that these lizards dig 1400 hrs, whereas in July it centers on 0900 to 1300 their own burrows. Some burrows are inhabited by hrs (Howland 1988). Not all lizards are active each morning. In April more than one lizard (Krekorian 1976), and individu- day, and some individuals were not observed out for most activity als have been observed to use more than one burrow several days (Krekorian 1976). (Howland 1988). Average body temperatures of foraging lizards occurs from 0930 Jeff Howland (pers. comm.) observed an adult dig range from 41.0 to 43.8°C (105.8 to 110.8°F; Nor- to 1400 hrs, its burrow. It started by scraping out a round hollow ris 1953, McGinnis and Dickson 1967, Krekorian about 30 cm (1 ft) in diameter and 1-2 cm (0.4-0.8 in) 1976, Howland 1988). The highest body temperature whereas in July it deep at the center. Then it began digging a burrow recorded was 46.4°C (115.5°F), taken with a cloa- centers on 0900 in the middle. Within about an hour, it had a burrow cal thermometer at 1030 on 10 July 1948 by Nor- deep enough to easily allow the lizard to get under ris (1953), and this appears to be a global record of to 1300 hrs. ground, and the hollow had been filled to ground voluntary thermal maximum for reptiles (summary level by soil excavated from the burrow. By this time, in Brattstrom 1965). The figure is consistent with the it was early afternoon and the lizard then went into field data of Howland (1988) in which the maximum the burrow and back-filled the entrance, as they often cloacal temperature was 46.0°C (114.8°F) and of do at the end of the daily activity period, leaving the McGinnis and Dickson (1967) who found deep body burrow and excavation area barely visible. temperatures ranging to 45.7°C (114.3°F). These The burrow is defended with an assertion display voluntary maxima are just below the critical thermal given by both sexes, and a more vigorous chal- maximum (47.5°C [117.5°F]) and lethal temperature lenge display performed mostly by males. Like those (50.5°C [122.9°F]) for the species (Cowles and Bogert of most Southwestern iguanid lizards, the display 1944), comprising an often cited example of “life on involves pushups, lateral compression of the body, the edge” in the thermal ecology of desert reptiles. and extension of the gular and ventral surfaces, but The activity season for adult Desert Iguanas in the it is species specific in format. In the most aggressive Sonoran Desert extends from March to September, display, the challenger pins his opponent against his but juveniles may be active as early as February and as body and inflicts a violent, audible tail slap (Carpenter late as October (Parker 1972, Howland 1988). In the 1961). Chuckwalla Valley, females cease activity shortly

SONORAN HERPETOLOGIST 23 (10) 2010 137 The activity season for adult Desert Iguanas in the Sonoran Desert extends Figure 3. Desert Iguanas (Dipsosaurus dorsalis) mating in the Chuckwalla Valley, Riverside Co., California. from March to Photo by Jeffrey M. Howland. September, but after egg laying, suggesting the possibility that egg Howland 1988). This is a long-lived lizard with a brooding may occur during this period (Jeff How- longevity of over 7 years in the wild (Krekorian 1984) juveniles may be land, pers. comm.). A small percentage of the females and over 14 years in captivity (Bowler 1977). Survi- active as early as become active again just before the hatchlings appear vorship is also high, averaging 50% to 60% per year in August, whereas male activity gradually decreases in both adults and subadults (Parker 1972, Krekorian February and as in August though September (Howland 1988). 1984, Howland 1988). late as October. In Courtship and mating were observed in enclosures Avoidance of predation may be one benefit of by Carpenter (1961). The males approach females the late morning-early afternoon activity period of the Chuckwalla with a rapid head bob. If she does not retreat, he bites the Desert Iguana (Pianka and Vitt 2003). Howland Valley, females her shoulder and brings his body on top of hers and (1988) witnessed five predation events on Desert attempts cloacal apposition (Carpenter 1961). Jeff Iguanas. Two of these involved moderately sized cease activity Howland (pers. comm.) observed the entire sequence Coachwhips (Coluber flagellum) that separately preyed shortly after egg of courtship and mating several times in the Chuck- on an adult male and an adult female Dipsosaurus. One walla Valley (Figure 3). In addition he observed a hatchling was observed to be taken by a large Tiger laying, suggesting strange behavior involving abdominal contact similar Whiptail (Aspidoscelis tigris) and another by a Log- the possibility that described by Carpenter (1961) for lizards in an gerhead Shrike (Lanius ludovicianus). An adult female enclosure. This consisted of young, probably imma- Long-nosed Leopard Lizard (Gambelia wislizenii) was that egg brooding ture, females crawling under adult males and flipping observed to prey on an adult female Desert Iguana may occur during over on their backs in the process. The males arched (Figure 4). Eight to nine days previous the Desert their backs a little to allow passage. In each case, the Iguana was 121 mm SVL (4.8 in) and 53.3 g (1.9 oz), this period (Jeff young female passed back and forth under the male and the leopard lizard was 146 mm SVL (5.7 in) and Howland, pers. several times, and then the interaction ended. The 80.8 g (2.9 oz). The leopard lizard consumed the males remained passive during the activity. entire Desert Iguana except of a 97-mm (1.5-g) tail tip com.). One clutch containing 2-8 (usually 3 to 5) eggs is that was broken off by biting and head shaking. Im- deposited during late May or early June (Norris 1953, mediately after the predation event the leopard lizard Howland 1988), but perhaps as late as August (May- weighed 127.2 g (4.5 oz; Jeff Howland, pers. comm). hew 1971). Hatchlings emerge during a four week pe- Diet has been investigated by Norris (1953) and riod centered in August (Howland 1988, Parker 1972). Mautz and Nagy (1987) in the Coachella Valley and Growth of Desert Iguanas is remarkably slow by Howland (1988) in Chuckwalla Valley. All three and varies with food availability (Howland 1988). In studies found that the flowers of Creosotebush Arizona, Parker (1972) estimated an average growth formed the staple during the spring. In the Coachella, rate for hatchlings of ca 20 mm/year (0.8 in/year). the spring diet also included leaves of Palmer’s Cold- Age at maturity (105 mm to 120 mm SVL [4.1 to 4.7 enia (Coldenia palmeri) and flowers of Desert Palafox in]) has been calculated at 31 to 54 months (Mayhew (Palafoxia linearis) and White Bursage (Ambrosia du- 1971, Parker 1972, Muth 1977, Mautz and Nagy 1987, mosa). The autumn diet centered on leaves of Coldenia SONORAN HERPETOLOGIST 23 (10) 2010 138 Dipsosaurus dorsalis appears to represent an ancient lineage that is now restricted to the Figure 4. Long-nosed Leopard Lizard (Gambelia wislizenii) preying on a Desert Iguana (Dipsosaurus dorsalis) in the Chuckwalla Valley, Mohave and Riverside Co., California. Photo by Jeffrey M. Howland. palmeri, Desert Dicoria (Dicoria canescens), and Indigo since Pough’s (1973) paper many exceptions have Sonoran deserts Bush (Dalea schottii), and the flowers and leaves Emory been identified to the necessity of large body size to and adjacent Dalea (Dalea emoryi). In the Chuckwalla Valley, the support herbivory (summaries in Greene 1982, Pianka spring diet also included flowers of Coulter’s Lupine and Vitt 2003). These include the switch to partial thornscrub and (Lupinus sparsiflorus), Fremont’s Pincushion (Chaenac- herbivory by Coachella Fringe-toed Lizards (Uma inor- tropical deciduous tis fremonti), and Ironwood (Olnea tesota). During the nata) in times of abundant flowers (Durtsche 1992). rest of the year the leaves, seeds, and dried flowers Iverson (1980, 1982) examined the gut structure of forest. Its of ironwood were eaten. Except for Creosotebush iguanines and determined that Dipsosaurus is the least association with flowers in the spring, there is no overlap in the main specialized for digestion of plant material. The high diet items at the two localities, and the Desert Iguana preferred body temperature of the Desert Iguana may the Creosotebush appears to be capable of subsisting on a wide variety facilitate digestion by this relatively small iguanine must be of of plants depending on the vegetation present in its herbivore (Mautz and Nagy 1987). Pianka and Vitt habitat. (2003) suggested that the consumption of mammal relatively recent In southern California, a small amount of feces (Norris 1953, Mautz and Nagy 1987) may help origin as Larrea material (1-3%) was consumed, including insects, supply the micro-organisms to the digestive tract feces, and even other Desert Iguanas (Norris needed for processing plant material. did not arrive in 1953, Mautz and Nagy 1987, Howland 1988). Remark- On the basis of phylogeny, herbivory is expected North America ably, arthropods were found to comprise 41-68% of in Dipsosaurus. The is a member of a natural the stomach contents of Desert Iguanas in the tropi- group (monophyletic clade) that includes about 30 until sometime in cal deciduous forest of Cape Region of Baja California species in seven (Frost and Etheridge 1989) or eight the Pleistocene. Sur by Asplund (1967), and Galina-Tesaro found that (Etheridge 1982) genera that are large bodied, her- on the La Paz plains the species consumed 100% bivorous, and have a mid-dorsal crest (reduced in size lepidopteran larvae during the late-summer rains in Dipsosaurus and absent in Sauromalus, chuckwallas). (Grismer 2002). Thus, herbivory does not appear to The clade is known as the Iguanidae in the taxonomic be a fixed ecological attribute, and the species may be arrangement of Frost and Etheridge (1989) or as the primarily insectivorous at more tropical latitudes. Iguaninae in that of Schulte et al. (2003). Herbivory From a physiological viewpoint, the herbivory of and large body size are thus two shared and probably the Desert Iguana is of special interest. Pough (1973) derived features of iguanine lizards. As an adaptation found that a diet consisting predominately of plant to arid environments, the Desert Iguana has either material occurs mostly in lizard species that have an retained the primitive ancestral condition of small adult body weight of greater than 100 g (3.5 oz). Adult size (relative to other iguanines) or has undergone an Dipsosaurus weigh between 25 g and 75 g (0.9 to 2.6 evolutionary reversal (i.e., re-evolved small size), but oz; Minnich 1971), and even hatchlings are primar- the species has maintained herbivory perhaps via be- ily herbivorous (Mautz and Nagy 1987). However, haviors that achieve uniquely high body temperatures. SONORAN HERPETOLOGIST 23 (10) 2010 139 Pacific (Grismer 2002). Its absence on Isla Tiburón is enigmatic. The easternmost known localities for the species in Arizona are Snyder Hill, Rillito on the Santa Cruz River, Pima Co.; Florence, Pinal Co.; Blue Point on the Salt River, Fort McDowell, Cave Creek, Maricopa Co.; 16 km (10 mi) NE Aguila, Yavapai Co.; Burro Creek, Big Sandy River, and Virgin River, Mohave Co. The locality of Audley, Yavapai Co., listed for USNM 45003, collected in 1909, may represent a rail shipping station rather than the point of collection. The distribution in the Tucson area is curious. The Avra Valley is often considered to be the eastern The distribution limit of the species in southern Arizona. But previ- ously it may have extended a short distance east of the Desert around the ends of the Tucson Mountains as is indi- Iguana in the cated by UAZ 15328, found dead on the road at the junction of Ina and Silverbell Roads by Oscar Soule Tucson area is Figure 5. Geographic distri- on 26 July 1966 (32.3374198°N, 111.0877314°W) and curious. The Avra bution of the Desert Iguana by UAZ 28748 listed as having been collected at Sny- (Dipsosaurus dorsalis; green) der Hill, Ajo Hwy, 14 km (8.7 mi) W and 7.9 km (4.9 Valley is often and the Isla Santa Catalina Desert Iguana (Dipsosaurus mi) S of Tucson on 13 Apr 1968 by James DeWeese considered to be catalinensis; red). (32.1576675°N, 111.1132820°W). As pointed out by Rosen (2007) the species penetrates farther into the the eastern limit The position of the Desert Iguana within the Arizona Upland Desert than most species centered in of the species in Iguaninae is not well resolved. On the basis of mor- the creosote flats of the Colorado and Mohave des- phology Etheridge and de Queiroz (1988) found it to erts, and he records a Desert Iguana at 1128 m (3700 southern Arizona. be a relatively early branch of the subfamily and to be ft) in desert grassland in Ban Thak Pass. But previously part of a basal trichotomy composed of (1) the Desert No range-wide studies of geographic variation in Iguana (Dipsosaurus); (2) the Fiji iguanas (Brachylophus); morphology or DNA sequences have been con- it may have and (3) all other iguanines. In a combined analysis of ducted. Lamb et al. (1992) examined mitochondrial extended a short morphology and DNA sequences, Sites et al. (1996) haplotypes of the Desert Iguana in western Arizona found it to be the earliest branch within iguanines, and southeastern California and uncovered signifi- distance east with the possible exception of the Fiji iguanas. Thus, cant geographic structure, but this did not reflect a around the ends Dipsosaurus dorsalis appears to represent an ancient break at the Colorado River as found in the Desert lineage that is now restricted to the Mohave and Tortoise (Gopherus agassizii; Lamb et al. 1989). of the Tucson Sonoran deserts and adjacent thornscrub and tropical Hulse (1992) mapped five subspecies of the Desert Mountains as is deciduous forest. Its association with the Creosote- Iguana. Grismer et al. (1994) found no consistent bush must be of relatively recent origin as Larrea did morphological differences between the two pen- indicated by a not arrive in North America until sometime in the insular subspecies and placed D. d. lucasensis in the specimen found Pleistocene (Van Devender 1990). synonymy of D. d. dorsalis. Similarly, Grismer (1999) The Desert Iguana ranges in Baja California Sur found that D. d. carmenensis could not be differenti- dead on the road from Cabo San Lucas to Laguna San Ignacio, north ated from peninsular populations and synonymyzed at the junction of of which it becomes restricted to the gulf side of the it with D. d. dorsalis. He found D. catalinensis to have peninsula (Grismer 2002); it extends into the Sonoran a unique darkly suffused gular surface (see Figure Ina and Silverbell Desert of Baja California, southern California, west- 4.12 in Grismer 2002) and considered it to repre- Roads by Oscar ern Arizona, Sonora, and northern Sinaloa, and into sent a distinct species. Burger and Hensley (1949) the Mohave Desert of southern California, southern redefined Dipsosaurus d. sonoriensis (Allen 1933) on Soule. Nevada, northwestern Arizona, and extreme south- the basis of color pattern (rather than scalation), and western Utah (Figure 5). The northern limit of the the subspecies was recognized by Bogert and Oliver species appears to be correlated with that of the Creo- (1945), Langebartel and Smith (1954), and Hardy and sotebush (Larrea tridentata; Norris 1953). In Sinaloa McDiarmid (1969); its status has not been appraised it is known south to near Topolobompo (25°33’N; in more recent times. Thus, the genus contains three Hardy and McDiarmid 1969). It occurs on Islas taxa: Isla Santa Catalina Desert Iguana, D. catalinensis Ángel de la Guardia, Carmen, Cerralvo, Coronados, Van Denburgh (1922); Northern Desert Iguana, D. Espíritu Santo, Monserrate, Partida Sur, San José, dorsalis dorsalis (Baird and Girard 1852), and Sonoran San Luis, and San Marcos in the Gulf of California, Desert Iguana, D. dorsalis sonoriensis Allen (1933; status and on Islas Magdalena and Santa Margarita in the unknown). SONORAN HERPETOLOGIST 23 (10) 2010 140 Baird and Girard (1852) described Crotaphytus dorsalis. Journal of Thermal Biology 1:15-22. dorsalis from the “Desert of Colorado, Cal.”, and Brattstrom, B.H. 1965. Body temperatures of reptiles. Smith and Taylor (1950) restricted the type locality to American Midland Naturalist 73:376-442. “Winterhaven (=Fort Yuma), Imperial County.” Hal- Carpenter, C.C. 1961. Patterns of social behavior in the lowell (1854) established the genus Dipso-saurus, which Desert Iguana, Dipsosaurus dorsalis. Copeia 1961:396- he amended to Dipsosaurus in 1859. 405. The Comcáac (Seri) consider the Desert Iguana to Cowles, R.B. 1940a. Observations on the winter activi- be extremely dangerous (Felger 1990, Nabhan 2003), ties of desert reptiles. Ecology 22:125-140. and its Comcáac name translates as “thing, that if Cowles, R.B. 1940b. Additional implications of reptilian it bites, prevents from arriving to home camp.” A sensitivity to high temperatures. American Naturalist legend relates how a traveler who failed to arrive at 74:542-561. his home camp was found dead in an arroyo with a Cowles, R.B., and C.M. Bogert. 1944. A preliminary Desert Iguana nearby. The pale coloration of the spe- study of the thermal requirements of desert reptiles. cies may contribute to the fear in which it is held. The Bulletin of the American Museum of Natural His- Tohono O’odham are said traditionally to have eaten tory 83:261-296. Desert Iguanas. Durtsche, R.D. 1992. Feeding time strategies of the The generic name Dipsosaurus is derived from dipsa fringe-toed lizard, Uma inornata, during breeding and (thirsty) and sauros (lizard). The species name dorsalis is non-breeding seasons. Oecologia 89:85-89. derived from dorsum (back) and refers to the enlarged Etheridge, R.E. 1982. Checklist of iguanine and Mala- row of vertebral scales. gasy iguanid lizards. Pages 60-76 in: G.M. Burghartd Acknowledgements. I thank Jeff Howland for and A. S. 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