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Greater Short-horned Lizard - Phrynosoma hernandesi

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Species of Concern
Native Species

Global Rank: G5
State Rank: S3
(see State Rank Reason below)


Agency Status
USFWS:
USFS:
BLM: SENSITIVE
FWP SWAP: SGCN3, SGIN

External Links

State Rank Reason (see State Rank above)
This species if found across much of eastern and central Montana. It is uncommon in grasslands, shrublands, and badlands. There is anecdotal evidence of significant declines, likely from habitat loss through lands use changes. It faces threats from continued habitat loss and invasion of non-native plat species.
Greater Short-horned Lizard (Phrynosoma hernandesi) Conservation Status Summary
State Rank: S3
Review Date = 10/23/2024
How we calculate Conservation Status

See the complete Conservation Status Report
 
General Description
EGGS:
The Greater Short-horned Lizard is viviparous, eggs develop internally, and females give birth to live young.

HATCHLINGS:
Broods can include up to 30 neonates or more, averaging about 12-18 (Smith 1941, Goldberg 1971, Ashton and Ashton 1998, Hammerson 1999), although broods are smaller in the north of their range (Powell and Russell 1991b, Hammerson 1999). Clutches range from 5 to 36 young (3 to 15 in the Pacific Northwest) during July to September (Nussbaum et al. 1983, Stebbins 2003). Newborn young are generally adult-like in appearance, with a broad and flattened body shape. They are about 2.0-2.5 cm (0.8-1.0 in) snout-vent length (SVL) and weigh about 0.7 – 0.8 g (0.02-0.03 oz), growing up to 3.8 cm (1.5 in) by the time of first hibernation (Goldberg 1971, Powell and Russell 1991b, Ashton and Ashton 1998).

JUVENILES AND ADULTS:
The body is broad and flattened. The back is spiny, especially noticeable in a single row of scales fringing each side of the body. Spines at the back of the head are stubby and about as long as they are wide at their base (Hammerson 1999, Stebbins 2003). A noticeably wide and deep notch separates the right and left horns at the back of the head (St. John 2002). The coloration of the back can be shades of gray, brown, or pink, usually blending cryptically with the soil, and can vary somewhat from region to region, as well as at single localities and even single broods (Ashton and Ashton 1998). There are typically two paired rows of dark brown blotches on the back that are often edged in white. Maximum total length (TL) is approximately 15 cm (5.9 in) (Hammerson 1999, St. John 2002, Stebbins 2003). In males there is a swelling at the base of the tail, and the tail is proportionally longer than in females. Females at 18 g (0.63 oz) outweigh males at 10 g (0.35 oz) and mature females are larger on average than males (7 cm (2.8 in) SVL versus 5 cm (2.0 in) SVL) (Powell and Russell 1985b, James et al. 1997).

Diagnostic Characteristics
The broad, flattened body separates this lizard from the other three lizard species regularly documented in Montana. The range overlaps only with the Common Sagebrush Lizard (Sceloporus graciosus). The Pigmy Short-horned Lizard (Phrynosoma douglasi) is present in adjacent southeastern Idaho and has been reported once from the Centennial Valley, Beaverhead County, in extreme southwestern Montana (Maxell et al. 2003). Adults of the Pigmy Short-horned Lizard are much smaller than the Greater Short-horned Lizard (Phrynosoma hernandesi), they lack the wide notch between the horns on the back of the head that gives the head of the Greater Short-horned Lizard a “heart-shape” appearance when viewed from above, and the small horns on the back of the head project almost vertically, rather than horizontally as in the Greater Short-horned Lizard (St. John 2002, Werner et al. 2004).

Species Range
Montana Range Range Descriptions

Native

Western Hemisphere Range

 


Range Comments
The Short-horned Lizard was recently split into the Pigmy Short-horned Lizard (Phrynosoma douglasii), the Northwestern Short-horned Lizard subspecies prior to the split, and the Greater Short-horned Lizard (P. hernandesi) (Zamudio et al. 1997, Hammerson 1999, Stebbins 2003), encompassing the other five subspecies prior to the reclassification (Reeve 1952). Analyses of genetic and morphological traits (Zamudio et al. 1997) also failed to support the validity of any subspecific differentiation across the range of the Greater Short-horned Lizard.

The Greater Short-horned Lizard is found from southeastern Alberta and southwestern Saskatchewan south through eastern Montana, the western Dakotas, Wyoming, western Nebraska, Colorado, Utah, eastern Nevada, New Mexico, Arizona, and western Texas to southern Durango (Stebbins 2003). They are found at elevations of 3,355 m (11,007 ft) or more (Montanucci 1981, Hammerson 1999). The range limit in the vicinity of southeastern Idaho, western Wyoming, northern Utah, and northern Nevada has not been precisely determined (Baxter and Stone 1985, Hammerson 1999, St. John 2002, Stebbins 2003). At the time the comprehensive summaries of amphibians and reptiles in Montana (Maxell et al. 2003, Werner et al. 2004) were published, there were 147 records from 27 counties east of the Continental Divide There was an obvious absence of records between the Missouri and Musselshell rivers in central Montana, reflecting limited survey effort in that region. Recent survey efforts have increased the number of observations across the state to 1,867 records from 38 counties east of the Continental Divide (MTNHP POD 2023).

Maximum elevation: 2,021 m (6,632 ft) in Carbon County (MTNHP POD 2023).

Observations in Montana Natural Heritage Program Database
Number of Observations: 2032

(Click on the following maps and charts to see full sized version) Map Help and Descriptions
Relative Density

Recency

 

(Observations spanning multiple months or years are excluded from time charts)



Migration
No information currently exists regarding the migration patterns of Greater Short-horned Lizards in Montana.

Habitat
This species occurs in sagebrush and grassland habitats. Occasionally in the presence of sedimentary rock outcrops (limestone, sandstone) and glacial drift. As well in areas with open stands of Limber Pine (Pinus flexilis) and Utah Juniper (Juniperus osteosperma) or Ponderosa Pine (Pinus ponderosa) (Mosimann and Rabb 1952, Werner 1974, Dood 1980, Hendricks 1999a, Vitt et al. 2005).

Favored areas in Montana tend to have a relatively high percentage of open bare ground and loose, sun-baked soils. On the southern exposures of the Pryor Mountains, Carbon County, individuals occur among limestone outcrops in canyon bottoms of sandy soil with an open canopy of Limber Pine-Utah Juniper. Individuals were also present on flats of relatively pebbly or stony soil with sparse grass and sagebrush cover (Paul Hendricks, personal observation).

Elsewhere, the Greater Short-horned Lizard is an animal of short-grass and mixed-grass (Stipa and Bouteloua) prairies, sagebrush (Artemisia sp.) and other shrubland types, and open coniferous forest in mountains. They often occur in open, shrubby or wooded areas (e.g., Piñon-Juniper, Pine-Oak Woodlands) with sparse vegetation at ground level and easy access to sunlight. Soil substrate may vary from rocky to sandy, but loose soils are usually present (Hammerson 1999, Stebbins 2003). In Alberta and Saskatchewan, Greater Short-horned Lizard is often found in coulees and small canyons associated with streams and rivers in terrain of exposed “badland” shale, with a ground cover that includes Creeping Juniper (Juniperus horizontalis) (Powell and Russell 1991a, 1998, Powell et al. 1998). In Colorado, this species was encountered frequently in both Black-tailed (Cynomys ludovicianus) and White-tailed (C. leucurus) Prairie Dog colonies (Clark et al. 1982).

Ecological Systems Associated with this Species
  • Details on Creation and Suggested Uses and Limitations
    How Associations Were Made
    We associated the use and habitat quality (common or occasional) of each of the 82 ecological systems mapped in Montana for vertebrate animal species that regularly breed, overwinter, or migrate through the state by:
    1. Using personal observations and reviewing literature that summarize the breeding, overwintering, or migratory habitat requirements of each species (Dobkin 1992, Hart et al. 1998, Hutto and Young 1999, Maxell 2000, Foresman 2012, Adams 2003, and Werner et al. 2004);
    2. Evaluating structural characteristics and distribution of each ecological system relative to the species' range and habitat requirements;
    3. Examining the observation records for each species in the state-wide point observation database associated with each ecological system;
    4. Calculating the percentage of observations associated with each ecological system relative to the percent of Montana covered by each ecological system to get a measure of "observations versus availability of habitat".
    Species that breed in Montana were only evaluated for breeding habitat use, species that only overwinter in Montana were only evaluated for overwintering habitat use, and species that only migrate through Montana were only evaluated for migratory habitat use.  In general, species were listed as associated with an ecological system if structural characteristics of used habitat documented in the literature were present in the ecological system or large numbers of point observations were associated with the ecological system.  However, species were not listed as associated with an ecological system if there was no support in the literature for use of structural characteristics in an ecological system, even if point observations were associated with that system.  Common versus occasional association with an ecological system was assigned based on the degree to which the structural characteristics of an ecological system matched the preferred structural habitat characteristics for each species as represented in scientific literature.  The percentage of observations associated with each ecological system relative to the percent of Montana covered by each ecological system was also used to guide assignment of common versus occasional association.  If you have any questions or comments on species associations with ecological systems, please contact the Montana Natural Heritage Program's Senior Zoologist.

    Suggested Uses and Limitations
    Species associations with ecological systems should be used to generate potential lists of species that may occupy broader landscapes for the purposes of landscape-level planning.  These potential lists of species should not be used in place of documented occurrences of species (this information can be requested at: mtnhp.org/requests) or systematic surveys for species and evaluations of habitat at a local site level by trained biologists.  Users of this information should be aware that the land cover data used to generate species associations is based on imagery from the late 1990s and early 2000s and was only intended to be used at broader landscape scales.  Land cover mapping accuracy is particularly problematic when the systems occur as small patches or where the land cover types have been altered over the past decade.  Thus, particular caution should be used when using the associations in assessments of smaller areas (e.g., evaluations of public land survey sections).  Finally, although a species may be associated with a particular ecological system within its known geographic range, portions of that ecological system may occur outside of the species' known geographic range.

    Literature Cited
    • Adams, R.A.  2003.  Bats of the Rocky Mountain West; natural history, ecology, and conservation.  Boulder, CO: University Press of Colorado.  289 p.
    • Dobkin, D. S.  1992.  Neotropical migrant land birds in the Northern Rockies and Great Plains. USDA Forest Service, Northern Region. Publication No. R1-93-34.  Missoula, MT.
    • Foresman, K.R.  2012.  Mammals of Montana.  Second edition.  Mountain Press Publishing, Missoula, Montana.  429 pp.
    • Hart, M.M., W.A. Williams, P.C. Thornton, K.P. McLaughlin, C.M. Tobalske, B.A. Maxell, D.P. Hendricks, C.R. Peterson, and R.L. Redmond. 1998.  Montana atlas of terrestrial vertebrates.  Montana Cooperative Wildlife Research Unit, University of Montana, Missoula, MT.  1302 p.
    • Hutto, R.L. and J.S. Young.  1999.  Habitat relationships of landbirds in the Northern Region, USDA Forest Service, Rocky Mountain Research Station RMRS-GTR-32.  72 p.
    • Maxell, B.A.  2000.  Management of Montana's amphibians: a review of factors that may present a risk to population viability and accounts on the identification, distribution, taxonomy, habitat use, natural history, and the status and conservation of individual species.  Report to U.S. Forest Service Region 1.  Missoula, MT: Wildlife Biology Program, University of Montana.  161 p.
    • Werner, J.K., B.A. Maxell, P. Hendricks, and D. Flath.  2004.  Amphibians and reptiles of Montana.  Missoula, MT: Mountain Press Publishing Company. 262 p.

Food Habits
Adults and juveniles are “sit-and-wait” predators that hunt mainly by sight. Ants and beetles are the most frequent foods across the range, but several other orders of arthropods are also taken (Pianka and Parker 1975, Laird and Leech 1980, Montanucci 1981, Powell and Russell 1984, Hammerson 1999). The diet in Montana is virtually undescribed; stomach contents of three individuals from coulees near the Marias River, Toole County, included mostly ants with a few beetles, grasshoppers, and spiders (Mosimann and Rabb 1952).

Ecology
Adult Greater Short-horned Lizards are diurnal and active during the warmer daylight hours. Specific information for Montana is limited, but information from other areas within their range indicates they may appear as early as late March (Hammerson 1999), with most surface activity in the northern parts of the range occurring from mid-April to mid-September. Extreme records in Alberta extend from April 1 to November 10 (Powell and Russell 1998), but most have disappeared by the mean date of the first fall frost. Young-of-the-year are generally not active during midday hours. Small lizards appear more dependent on air temperatures than on substrate temperatures, while large ones are more dependent on substrate temperature. Predators of this species are mostly unknown, but Striped Whipsnake (Masticophis taeniatus) and Burrowing Owls (Athene cunicularia) have been reported (Hammerson 1999). Additionally, birds have been identified as the primary predatory group (Russell and Bauer 1993). The annual period of activity in Montana is poorly defined, and no predators have been reported.

Home ranges of radio-tagged individuals in Alberta ranged from 4.4-2400 square meters (14.4-7,874 square feet) (James et al. 1997), with movements by females exceeding 100 m (328 ft) over a one-week period prior to mating, and as much as 266 m (873 ft) during the week prior to hibernation. Home ranges may be larger, because individuals are relatively sedentary but make irregular long movements that vary with the season (Powell and Russell 1998). The Greater Short-horned Lizard does not appear to be territorial (Powell and Russell 1998), although it exhibits some degree of site fidelity. A juvenile displaced 400 m (1,312 ft) returned to its capture site in less than a year and was subsequently captured twice more at this location (Pianka and Parker 1975). Little information is available on survival rate and longevity. Recapture data suggested high juvenile mortality in Alberta, whereas adult survival was considered relatively high (Powell and Russell 1991b, 1998). This pattern appears to be normal for other horned lizard species (Pianka and Parker 1975).

Reproductive Characteristics
Based upon information gathered from other areas within the species' range, adults mate shortly after emerging from winter dormancy in late March to early June, depending on elevation and latitude. The Greater Short-horned Lizard is viviparous, giving live birth (Nussbaum et al. 1983, Stebbins 2003). Young are born about two or three months after eggs are fertilized, mostly during early July to early August in Arizona (Smith 1941, Goldberg 1971), and late July to early August in Colorado, southern Wyoming, and Alberta (Powell and Russell 1991b, Ashton and Ashton 1998, Hammerson 1999). Size of eight litters from Alberta ranged from 6-13 young (Laird and Leech 1980, Powell and Russell 1998). One litter from Wyoming included 13 young approximately 2.3 to 2.4 cm (0.9 in) SVL at birth and consisted of two color morphs and four stillborn (Ashton and Ashton 1998). Five litters in Colorado ranged from 14-18 young (Hammerson 1999), and 12 litters from Arizona included 9-30 young (Smith 1941, Goldberg 1971). Sexual maturity of females is attained by the age of two years in Colorado (their third calendar year) and around 5-6 cm (2.0-2.4 in) SVL (Nussbaum et al. 1983, Hammerson 1999). In Alberta, males apparently reach sexual maturity in the second year, following their first winter dormancy, although females appear to mature later (Powell and Russell 1985b, 1998). Sexual maturity is reached by two years of age in the southwestern states (Pianka and Parker 1975).

Little information is available from Montana on any aspect of the reproductive biology of this species. Ten captive adult females from near Warren, Carbon County, gave birth in the first week of August (J. Barron, personal communication), and young about 3.0-3.5 cm (1.2-1.4 in) SVL have been observed in this area in August and early September (Hendricks 1999a, P. Hendricks, personal observation).

Management
The following was taken from the Status and Conservation section for the Greater Short-horned Lizard account in Maxell et al. 2009.

At the time the comprehensive summaries of amphibians and reptiles in Montana (Maxell et al. 2003, Werner et al. 2004) were published, there were about 130 total records for Greater Short-horned Lizard from 27 counties, with records generally widely scattered across the state east of the Continental Divide. Cope (1879) considered the Greater Short-horned Lizard about the most abundant reptile (second only to the Prairie Rattlesnake, (Crotalus viridis)) along the Missouri River, but it is no longer thought common anywhere in Montana, with the possible exception of some of the counties bordering Wyoming (Maxell et al. 2003). Populations in Carbon County appear to be robust (Vitt et al. 2005, J. Barron, personal communication), with many recent sightings (Barron reported 140 individuals near Warren in 2004) during favorable conditions. However, trend estimates remain unavailable for any locality in Montana, including areas of recent surveys. Connectivity of populations is unknown. Large gaps of a 100 km (62 mi) or more exist between some documented occurrences north of the Missouri River, and status is uncertain west of the Mussellshell River and south of the Missouri River. At the local scale, limited data suggest this species is relatively sedentary, with some degree of site fidelity, although females may move 100 m (328 ft) or more prior to mating or winter dormancy (Pianka and Parker 1975, James et al. 1997, Powell and Russell 1998). Thus, populations appear vulnerable to habitat fragmentation. Densities of this species have not been determined with any degree of accuracy but are thought to be generally low (considerably <50 individual/ha) (Pianka and Parker 1975, Powell and Russell 1985b), although perhaps locally fairly common (Hammerson 1999). Risk factors relevant to the viability of populations of this species are likely to include habitat loss/fragmentation, grazing, fire, road and trail development, on- and off-road vehicle use, use of pesticides and herbicides, oil and gas development, and surface mining. However, perhaps the greatest risk to maintaining viable populations of Greater Short-horned Lizard in Montana is the lack of baseline data on its distribution, status, habitat use, and basic biology (Maxell and Hokit 1999), which are needed to monitor trends and recognize dramatic declines when and where they occur. Few studies address or identify risk factors. In an Idaho study (Reynolds 1979), the closely related Pigmy Short-horned Lizard was four times more abundant in sagebrush habitat than in sites dominated by exotic Crested Wheatgrass (Agropyron cristatum), and more abundant in sagebrush habitat grazed by sheep than in ungrazed sagebrush and either grazed or ungrazed Crested Wheatgrass. A greater abundance of invertebrates may explain the preference for sagebrush over Crested Wheatgrass habitat (Reynolds 1979). More open ground for basking in grazed sagebrush may explain its preference over ungrazed sagebrush (Reynolds 1979, Hammerson 1999). Local extirpations of Greater Short-horned Lizard in Colorado have occurred in areas of intense cultivation and urban expansion, although this species seems to do well in areas used for livestock grazing (Hammerson 1999), perhaps because grazing reduces vegetative cover. Increased vehicular traffic in areas of oil and gas exploration and development have probably reduced populations in parts of Colorado (Hammerson 1999). Use of insecticides to control grasshopper infestations could also depress populations of this species, which feeds on these and other ground-dwelling insect species.


References
  • Literature Cited AboveLegend:   View Online Publication
    • Ashton, K.G. and K. L. Ashton. 1998. Phrynosoma douglasii (short-horned lizard). Reproduction. Herpetological Review 29(3):168-169.
    • Baxter, G.T. and M.D. Stone. 1985. Amphibians and reptiles of Wyoming. Second edition. Wyoming Game and Fish Department. Cheyenne, WY. 137 p.
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    • Cope, E. D. 1879. A contribution to the zoology of Montana. American Naturalist 13(7): 432-441.
    • Dood, A.R. 1980. Terry Badlands nongame survey and inventory final report. Montana Department of Fish, Wildlife, and Parks and Bureau of Land Management, Helena, MT. 70 pp.
    • Goldberg, S.R. 1971. Reproduction in the short-horned lizard Phrynosoma douglassi in Arizona. Herpetologica 27(3): 311-314.
    • Hammerson, G.A. 1999. Amphibians and reptiles in Colorado. University Press of Colorado & Colorado Division of Wildlife. Denver, CO. 484 p.
    • Hendricks, P. 1999a. Amphibian and reptile survey of the Bureau of Land Management Miles City District, Montana. Montana Natural Heritage Program, Helena, MT. 80 p.
    • James J.D., A.P. Russell, and G.L. Powell. 1997. Status of the eastern short-horned lizard (Phrynosoma douglassii brevirostre) in Alberta. Alberta Environmental Protection, Wildlife Management Divison, Wildlife Status Report No. 5, Edmonton, AB. 1-20.
    • Laird, M. and R. Leech. 1980. Observations on the short-horned lizard in southeastern Alberta. Blue Jay 38(4): 214-218.
    • Maxell, B.A. and D.G. Hokit. 1999. Amphibians and reptiles. Pages 2.1– 2.30 In G. Joslin and H. Youmans, committee chairs. Effects of recreation on Rocky Mountain wildlife: a compendium of the current state of understanding in Montana. Committee on Effects of Recreation on Wildlife, Montana Chapter of the Wildlife Society.
    • Maxell, B.A., J.K. Werner, P. Hendricks, and D.L. Flath. 2003. Herpetology in Montana: a history, status summary, checklists, dichotomous keys, accounts for native, potentially native, and exotic species, and indexed bibliography. Society for Northwestern Vertebrate Biology, Northwest Fauna Number 5. Olympia, WA. 135 p.
    • Maxell, B.A., P. Hendricks, M.T. Gates, and S. Lenard. 2009. Montana amphibian and reptile status assessment, literature review, and conservation plan, June 2009. Montana Natural Heritage Program. Helena, MT. 643 p.
    • Montanucci, R.R. 1981. Habitat separation between Phrynosoma douglassi and P. orbiculare (lacertilia: iguanidae) in Mexico. Copeia 1981(1): 147-153.
    • Mosimann, J.E. and G.B. Rabb. 1952. The herpetology of Tiber Reservoir Area, Montana. Copeia(1): 23-27.
    • Nussbaum, R.A., E.D. Brodie, Jr. and R.M. Storm. 1983. Amphibians and reptiles of the Pacific Northwest. University of Idaho Press. Moscow, ID. 332 pp.
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    • Powell, G.L. and A.P. Russell. 1984. The diet of the eastern short-horned lizard (Phrynosoma douglassi brevirostre) in Alberta and its relationship to sexual size dimorphism. Canadian Journal of Zoology 62(3): 428-440.
    • Powell, G.L. and A.P. Russell. 1985b. Growth and sexual size dimorphism in Alberta (Canada) populations of the eastern short-horned lizard, Phrynosoma douglassi brevirostre. Canadian Journal of Zoology 63(1): 139-154.
    • Powell, G.L. and A.P. Russell. 1991b. Parturition and clutch characteristics of short-horned lizards (Phrynosoma douglassii brevirostre) from Alberta. Canadian Journal of Zoology 69(11): 2759-2764.
    • Powell, G.L. and A.P. Russell. 1998. The status of short-horned lizards (Phrynosoma douglasi) and (P. hernandezi) in Canada. Canadian Field Naturalist 112(1):1-16.
    • Powell, G.L., A.P. Russell, and P.J. Fargey. 1998. The distribution of the Short-horned Lizard Phrynosoma hernandesi in Saskatchewan, Canada. Northwestern Naturalist 79: 19-26.
    • Powell, G.L., and A.P. Russell. 1991a. Distribution of the eastern short-horned lizard (Phrynosoma douglasi brevirostre) in Alberta, Canada. Northwestern Naturalist. 72(1): 21-26.
    • Reeve, W.L. 1952. Taxonomy and distribution of the horned lizard genus Phrynosoma. Kansas University Science Bulletin 34: 817-960.
    • Reynolds, T.D. 1979. Response of reptiles populations to different land management practices on the Idaho national engineering laboratory site. Great Basin Naturalist 39(3): 255-262.
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    • Vitt, L.J., J.P. Caldwell, and D.B. Shepard. 2005. Inventory of amphibians and reptiles in the Billings Field Office Region, Montana. Sam Noble Oklahoma Museum of Natural History and Department of Zoology, University of Oklahoma, Norman, OK. 33 pp.
    • Werner, J.K. 1974. Geographic distribution, Phrynosoma douglassi brevirostre. Herpetological Review 5(1):20.
    • Werner, J.K., B.A. Maxell, P. Hendricks and D.L. Flath. 2004. Amphibians and Reptiles of Montana. Mountain Press Publishing Company: Missoula, MT. 262 pp.
    • Zamudio, K.R., K.B. Jones, and R.H. Ward. 1997. Molecular systematics of short-horned lizards: biogeography and taxonomy of a widespread species complex. Systematic Biology 46:284-305.
  • Additional ReferencesLegend:   View Online Publication
    Do you know of a citation we're missing?
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    • Cope, E.D. 1875. Check-list of North American Batrachia and Reptilia; with a systematic list of the higher groups, and an essay on geographical distribution. Based on the specimens contained in the U.S. National Museum. U.S. Natioanl Museum Bulletin 1: 1-104.
    • Cope, E.D. 1900. The crocodilians, lizards, and snakes of North America. Report of the U.S. National Museum 1898: 153-1270.
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    • Crother, B.I. (ed.) 2008. Scientific and standard English names of amphibians and reptiles of North America north of Mexico. SSAR Herpetological Circular No. 37:1-84.
    • Dammann, J. 1949. Birth of eighteen young Phrynosoma douglassi Hernandes. Herpetologica 5: 144.
    • Dumas, P.C. 1964. Species-pair allopatry in the genera Rana and Phrynosoma. Ecology 45(1): 178-181.
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    • Martin, P.R. 1980a. Terrestrial wildlife habitat inventory in southeastern Montana. Montana Department of Fish, Wildlife and Parks and Bureau of Land Management, Helena MT. 114 p.
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Greater Short-horned Lizard — Phrynosoma hernandesi.  Montana Field Guide.  .  Retrieved on , from