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Overview

Distribution

California legless lizards are found in California and Mexico. They are found from western central California (San Joaquin and the coastal regions), through northwestern Baja California, and as far south as Colonia Guerrero, Mexico (Miller 1944; Stebbins 2003).

Biogeographic Regions: nearctic (Native )

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Range Description

This lizard has a spotty distribution in California (United States) and northwestern Baja California (Mexico), extending from near Antioch, California, south in the Coast Ranges, Transverse Mountains, and Peninsular Ranges, and along the coast of southern California, to Arroyo Pabellon, northwestern Baja California (Stebbins 2003), and inland in Baja California to at least La Rumarosa north of the Sierra Juarez (Hunt 1983, Grismer 2002). Scattered occurrences exist elsewhere in California, including the following: San Joaquin Valley; southern Sierra Nevada; Walker Basin; Ninemile Canyon; Paiute, Scodie, and Tehachapi mountains; desert-edge localities at the eastern end of Walker Pass in Kern County, Morongo Valley in San Bernardino County, Little San Bernardino Mountains at Whitewater in Riverside County, and the eastern slope of the Peninsular Ranges; and the Antelope Valley in the extreme western Mojave Desert (Stebbins 2003). The species also occurs on Los Coronados (Norte and Sur) and Todos Santos islands in Baja California. There are old records from Marin County and Palo Alto in the San Francisco Bay Area, California (Stebbins 2003). Its elevational range extends from sea level to about 5,100 feet (1,550 m) (Stebbins 1985). See spot map in Hunt (1983).
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National Distribution

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Global Range: (20,000-200,000 square km (about 8000-80,000 square miles)) This lizard has a spotty distribution in California and northwestern Baja California, extending from near Antioch, California, south in the Coast Ranges, Transverse Mountains, and Peninsular Ranges, and along the coast of southern California, to Arroyo Pabellon, northwestern Baja California (Stebbins 2003), and inland in Baja California to at least La Rumarosa north of the Sierra Juarez (Hunt 1983, Grismer 2002). Scattered occurrences exist elsewhere in California, including the following: San Joaquin Valley; southern Sierra Nevada; Walker Basin; Ninemile Canyon; Paiute, Scodie, and Tehachapi mountains; desert-edge localities at the eastern end of Walker Pass in Kern County, Morongo Valley in San Bernardino County, Little San Bernardino Mountains at Whitewater in Riverside County, and eastern slope of the Peninsular Ranges; and the Antelope Valley in the extreme western Mojave Desert (Stebbins 2003). The speices also occurs on Los Coronados and Todos Santos islands in Baja California. There are old records from Marin County and Palo Alto in the San Francisco Bay Area, California (Stebbins 2003). Elevational range extends from sea level to about 5,100 feet (1,550 meters) (Stebbins 1985). See spot map in Hunt (1983).

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Continent: Middle-America North-America
Distribution: USA (CW/SW California),  Mexico (NW Baja California Norte)  pulchra: Pacific slopes of C/S California, N Baja California.
Type locality: California. Restricted to San Diego, by SMITH & TAYLOR 1950.  nigra:
Type locality: Near San Diego, California, U.S.A., leg. J. BEHRENS: 2 syntypes (missing from ZMH fide HALLERMANN 1998).
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Physical Description

Morphology

California legless lizards are often mistaken for snakes because of their complete lack of limbs and elongated bodies. However, California legless lizards lack external ear openings and have unreduced eyes with moveable lids (Cogger and Zweifel, 1998). California legless lizards can also use tail autotomy to avoid predation, unlike snakes. Snout to vent length is 90 to 170 mm and total length is around 200 mm. Generally females are slightly larger, males are smaller by up to 10% of female size. Sexes are nearly impossible to distinguish without dissection.

Color varies with subspecies. In the more common Anniella pulchra pulchra, newborn lizards start out as a silvery drab color on their dorsal side and various shades of yellow on the ventral side. Anniella pulchra nigra begin life as silvery on their dorsal side and yellowy on their underside, but change with age. Adults vary in color from brown, to dark brown, to completely black.

Markings include one generally well defined line along the top of the backside and several other lateral sides. Depending of the overall darkness of the specimen, the markings vary in prominence and definition. In the blackest examples, the lines are not noticeable (Miller 1943).

Range mass: 0.9 to 4.7 g.

Range length: 111 to 228 mm.

Range basal metabolic rate: .028 to .116 cm3.O2/g/hr.

Other Physical Features: heterothermic

Sexual Dimorphism: sexes alike; female larger

  • Cogger, H., R. Zweifel. 1998. Encyclopedia of Reptiles and Pmphibians. Sydney, Australia: Weldon Owen Pty Limited.
  • Fusari, M. 1984. Temperature responses of standard, aerobic metabolism by the California Legless Lizard Anniella pulchra. Comp. Biochem. Physiol., Vol 77A, No. 1: 97-101.
  • Fusari, M. 1985. Drinking of Soil Water by the Califonia Legless Lizard. Copeia, 4: 981-986.
  • Miller, C. 1943. An Intergradient Population Connecting Anniella pulchra pulchra and Anniella pulchra nigra. Copeia, No. 1: 2-7.
  • Wiens, J., J. Slingluff. 2001. How lizards turn into snakes: a phylogenic analysis of body-form evolution in Anguid lizards. Evolution, 55(11): 2303-2318. Accessed April 15, 2008 at http://www.blackwell-synergy.com/doi/pdf/10.1111/j.0014-3820.2001.tb00744.x.
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Size

Length: 24 cm

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Diagnostic Description

In the U.S., no other sympatric limbless reptile has movable eyelids. Differs from A. GERONIMENSIS of Baja California in having a snout that is more rounded in profile and more pointed in dorsal view, fewer longitudinal dark lines on the sides, and an orange-yellow venter. (Stebbins 1985).

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Ecology

Habitat

California Coastal Sage and Chaparral Habitat

This taxon is found in the California coastal sage and chaparral ecoregion, located along the southern coast of California and Pacific coast of Baja California, has extremely high levels of species diversity and endemism. The eight Channel Islands are also part of this ecoregion, as are Isla Guadalupe and Isla Cedros. The climate is Mediterranean, with cold wet winters and dry hot summers. Precipitation levels range between 150 to 500 millimeters per annum. Vegetation typically grows on soils made of volcanic rocks on the base of the San Pedro Martir Mountains and on soils of sedimentary origin closer to the coastal zone.

The California coastal sage and chaparral supports a diversity of habitats including montane conifer forests, Torrey pine woodland, cypress woodlands, southern walnut woodlands, oak woodlands, riparian woodlands, chamise chaparral, inland and coastal sage scrub, grasslands, vernal pools, and freshwater and salt marshes. Coastal sage scrub, chamise chaparral, and oak woodlands dominate much of the landscape. Coastal sage scrub is a diverse and globally rare habitat type occurring in coastal terraces and foothills at elevations below 1000 meters (m), interspersed with chamise chaparral, oak woodland, grasslands, and salt marsh. This habitat type is characterized by low, aromatic and drought-deciduous shrublands of Black Sage (Salvia mellifera), White Sage (Salvia apiana), Munz’s Sage (Salvia munzii), California Sage (Artemisia californica), California Buckwheat (Eriogonum fasciculatum), California Brittlebush (Encelia californica), Toyon (Heteromeles arbutifolia), Lemonade Berry (Rhus integrifolia), and a diverse assemblage of other shrubs, herbaceous plants, cacti and succulents. Opuntia, Yucca, and Dudleya are some of the most common succulent genera, with the latter represented by several species endemic to the ecoregion.

The Island Fox (Urocyon littoralis), and Santa Catalina Shrew (Sorex willetti) are endemic mammals found in the ecoregion. Some of the specialist mammalian species found in the California sage and chaparral are: San Diego Pocket Mouse (Chaetodipus fallax), Merriam's Kangaroo Rat (Dipodomys merriami), and Stephens's Kangaroo Rat (Dipodomys stephensi).

The Rosy Boa (Charina trivirgata), California Legless Lizard (Anniella pulchra), and several relict salamanders are examples of the unusual and distinctive herpetofauna. Some endemic reptile species found in the ecoregion are: San Clemente Night Lizard (Xantusia riversiana), found only on the Channel Islands; Red-diamond Rattlesnake (Crotalus ruber), San Diego Banded Gecko (Coleonyx variegatus abbotti), and Coast Horned Lizard (Phrynosoma blainvillii).

Nutall’s Woodpecker (Picoides nuttallii) is endemic to the California sage and chaparral ecoregion, as are several endemic subspecies, which occur in the Channel Islands. Virtually all of the ecoregion is included in the California Endemic Bird Area. The California Gnatcatcher (Polioptila californica) is a further relict species found in the ecoregion.  The coastal populations of the Cactus Wren (Campylorhynchus brunneicapillus) are a notable occurrence of this bird, which is usually found in more arid regions.

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Unlike most lizards, California legless lizards prefer lower temperatures. In one study it was found that the average temperature of these lizards was between 21 and 28 degrees Celsius. They suggest that being adapted to this lower and wider range of temperatures allows them to remain active in the cooler temperatures of their subterranean environments. They can also remain active in cooler temperatures above ground. have “increased activity in its subterranean environment,” which is in line with observations that they are active on cooler days.

California legless lizards require loose sand for burrowing (sand, loam, or humus), moisture, warmth, and plant cover (Stebbins 2003). As a result, they are most commonly found within 100 km of the coast in dunes which harbor bush lupine (Lupinus arboreus), mock heather (Eriogonum parvilfolium), mock aster (Ericameria ericoides), and other native coastal shrubs (Jennings and Hayes 1994). These shrubs are ideal because they provide plenty of leaf litter, which helps keep temperatures in the sand relatively low and moisture content relatively high on hot days, and have extensive root systems, which attract plenty of insects for prey.

Additionally, moisture is a key aspect of their environment. Without adequate moisture they cannot shed their skin, which inhibits vision and feeding, causing them to become inactive and even starve to death (Miller 1944).

Range elevation: 0 to 1800 m.

Habitat Regions: temperate ; terrestrial

Terrestrial Biomes: desert or dune ; chaparral ; scrub forest

  • Bury, R., T. Balgooyen. 1976. Temperature Selectivity in the Legless Lizard, Anniella pulchra. Copeia, No. 1: 152-155.
  • Jennings, M., M. Hayes. 1994. Amphibian and Reptile Species of Special Concern in California. California Department of Fish and Game, Special Publication: 1-20. Accessed April 15, 2008 at http://www.dfg.ca.gov/habcon/info/herp_ssc.pdf.
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Habitat and Ecology

Habitat and Ecology
This legless lizard burrows in loose soil, especially in semi-stabilized sand dunes and in other areas with sandy soil, including habitats vegetated with oak or pine-oak woodland, or chaparral; it also occurs along wooded stream edges, and occasionally in desert-scrub (Hunt 1983, Grismer 2002, Stebbins 2003). Bush lupine and mock heather often are present in suitable dune habitats (Stebbins 2003). The species is often found in leaf litter or under rocks, logs, or driftwood.

Systems
  • Terrestrial
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Comments: This legless lizard burrows in loose soil, especially in semi-stabilized sand dunes and in other areas with sandy soil, including habitats vegetated with oak or pine-oak woodland, or chaparral; it also occurs along wooded stream edges, and occasionally in desert-scrub (Hunt 1983, Grismer 2002, Stebbins 2003). Bush lupine and mock heather often are present in suitable dune habitats (Stebbins 2003). The species is often found in leaf litter or under rocks, logs, or driftwood.

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Migration

Non-Migrant: Yes. At least some populations of this species do not make significant seasonal migrations. Juvenile dispersal is not considered a migration.

Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

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Trophic Strategy

California legless lizards forage in leaf litter for prey. Their primary diet consists of insect larvae, termites, beetles, spiders, and other invertebrates (Dudek and Associates, Inc. 2000; Stebbins 2003). Often they will not eat prey on the surface, but will burrow underground before consuming the prey (Miller 1944).

Animal Foods: insects; terrestrial non-insect arthropods

Primary Diet: carnivore (Insectivore , Eats non-insect arthropods)

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Comments: Eats insects, insect larvae, and spiders.

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Associations

Other than their immediate effect on prey and predators, not much is known about the role of Anniella pulchra in the ecosystems they inhabit. It is reported that they are parasitized by a nematode and a cestode (Miller 1944).

Commensal/Parasitic Species:

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California legless lizards avoid most predation by spending most of their time underground. However, when threatened by a predator they are able to lose their tail in order to escape. The tail will regenerate over the course of about one year, but will be much shorter, darker, and more blunt than the original. More than half of all individuals have regrown tails (Miller 1944).

Predators include ringneck snakes (Diadophis punctatus), common kingsnakes (Lampropeltis getula), deer mice (Peromyscus maniculatus), long-tailed weasels (Mustela frenata), domestic cats (Felis sylvestris), California thrashers (Toxostoma redivivum), American robins (Turdus migratorius), and loggerhead shrikes (Lanius ludovicianus) (Jennings and Hayes 1994; Miller 1944).

Known Predators:

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Population Biology

Number of Occurrences

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: 21 - 80

Comments: Jennings and Hayes (1994) mapped several dozen localities in California with extant populations; probably this translates to fewer than 100 distinct occurrences, plus a lesser number in northwestern Baja California.

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Global Abundance

10,000 - 1,000,000 individuals

Comments: Total adult population size is uncertain but probably is at least 10,000.

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Life History and Behavior

Behavior

Though it is unknown how California legless lizards communicate or interact,there is a fair bit known about how they perceive their environment. They have eyes and are believed to be relatively near-sighted. Though they have no external ear openings, they have a keen sense of mechanical disturbances and can sense vibrations well; individuals almost never feed aboveground, instead preferring to dig under ground and come up directly beneath the prey. Olfactory senses also seem likely as there are recorded instances of California legless lizards sticking their heads above ground while the body is still submerged and licking at the air, as if tasting it (Miller 1944).

Perception Channels: visual ; tactile ; vibrations ; chemical

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Cyclicity

Comments: May forage in leaf litter during the day, emerging on the surface at dusk or at night (Stebbins 1985).

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Life Cycle

Aside from size and color, newly born Anniella pulchra are nearly identical to their adult forms. As they grow, their dorsal color usually darkens: slightly in Anniella pulchra pulchra, and more significantly in Anniella pulchra nigra.

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Life Expectancy

There exists almost no data on the longevity of California legless lizards in the wild, though one adult specimen is known to have lived for six years in a lab (Jennings and Hayes 1994).

Range lifespan

Status: captivity:
6 (high) years.

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Reproduction

Not much is known about mating in California legless lizards.

Little data is available on the reproductive behaviors of California legless lizards. The breeding season is thought to be between the spring months and July and young are commonly born between September and November, suggesting a gestation period of four months. California legless lizards typically mature at two to three years of age for males and females (respectively). Maturation seems to be linked to size: 90 mm snout to vent length in males and 121 mm snout to vent length in females are the sizes at which sexual maturity is reached. Females are ovoviviparous and give live birth to litters of one to four young, most often two (Jenning and Hayes 1994; Miller 1944; Dudek and Associates, Inc. 2000).

Breeding interval: California legless lizards have been observed to breed biennially, but it is not known if this is true of all populations.

Breeding season: The breeding season probably occurs between early spring and July.

Range number of offspring: 1 to 4.

Average number of offspring: 2.

Average gestation period: 4 months.

Average age at sexual or reproductive maturity (female): 2 years.

Average age at sexual or reproductive maturity (male): 3 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; sexual ; fertilization ; ovoviviparous

As with most species of lizards, there is little to no parental investment in California legless lizards. Young are born live and are immediately independent (Miller 1944).

Parental Investment: no parental involvement; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female)

  • Dudek and Associates, Inc, 2000. "California Legless Lizard" (On-line). Western Riverside County Multiple Species Habitat Conservation Plan. Accessed April 15, 2008 at http://ecoregion.ucr.edu/full.asp?sp_num=38.
  • Jennings, M., M. Hayes. 1994. Amphibian and Reptile Species of Special Concern in California. California Department of Fish and Game, Special Publication: 1-20. Accessed April 15, 2008 at http://www.dfg.ca.gov/habcon/info/herp_ssc.pdf.
  • Miller, C. 1944. Ecological Relations and Adaptations of the Limbless Lizard of the Genus Anniella. Ecological Monographs, Vol 14, No. 3: 271-289. Accessed April 15, 2008 at http://www.jstor.org/stable/1948444.
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Ovulation occurs May-July. Oviducal eggs have been found July-October. Births occur September-November (probably mainly October). Litter size is 1-4 (usually 1-2). Not all adult females produce young each year in southern California (Behler and King 1979, Goldberg and Miller 1985).

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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage: Anniella pulchra

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 0
Specimens with Barcodes: 1
Species With Barcodes: 1
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Conservation

Conservation Status

The main threat facing California legless lizards is human habitat disturbance. Harmful activities include the destruction of natural habitat for agriculture, housing developments, sand mining, golf courses, and off-road vehicle activities (Stebbins 2003). Additionally, exotic plant species, such as ice plants (Carpobrotus edulis and Mesembryanthemum crystallinum), Marram grass (Ammophila arenaria), veldt grass (Ehrharta calycina) and eucalyptus trees (Eucalyptus spp.) support much smaller populations of insects and arthropods, which these lizards depend on for food (Jennings and Hayes 1994).

In 1998 to 1999, a moderate-high impact search was done at Moss Landing dune, California, over an area of 1.57 hectares. Previous studies based on lower impact searches would suggest a population of about 170, though researchers found nearly 3600. These results suggest that California legless lizards may be locally abundant (Kuhnz et al. 2005).

Though California legless lizards are listed as a species of special concern in California, they are not protected by federal or international regulations.

In 1998, a rule was proposed to the Department of the Interior Fish and Wildlife Service to list Anniella pulchra nigra as endangered, but it was withdrawn based on the amount of currently protected habitat and conservation efforts to restore native vegetation to dunes colonized by alien vegetation.

US Federal List: no special status

CITES: no special status

IUCN Red List of Threatened Species: least concern

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IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2007

Assessor/s
Hollingsworth, B. & Hammerson, G.A.

Reviewer/s
Cox, N., Chanson, J.S. & Stuart, S.N. (Global Reptile Assessment Coordinating Team)

Contributor/s

Justification
Listed as Least Concern in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category.
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National NatureServe Conservation Status

United States

Rounded National Status Rank: N3 - Vulnerable

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NatureServe Conservation Status

Rounded Global Status Rank: G3 - Vulnerable

Reasons: Spotty occurrence in California and Baja California; much habitat (about 20%) has been destroyed and fragmented by urbanization and agriculture; other habitat has been degraded by human activities and invasive plants; conservation status is not well known due to the difficulty in adequately censusing this fossorial species.

Intrinsic Vulnerability: Moderately vulnerable

Environmental Specificity: Very narrow to narrow.

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Population

Population
Jennings and Hayes (1994) mapped several dozen localities in California with extant populations; probably this translates to fewer than 100 distinct occurrences, plus a lesser number in northwestern Baja California. The total adult population size is uncertain but is probably at least 10,000. The species has been eliminated from probably about 20% of its historical range in California (Jennings and Hayes 1994).

Population Trend
Decreasing
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Global Short Term Trend: Relatively stable to decline of 30%

Global Long Term Trend: Decline of 30-50%

Comments: This species has been eliminated from probably about 20% of its historical range in California (Jennings and Hayes 1994). Additional habitat has been degraded, and abundance of the species undoubtedly has been reduced in those areas.

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Threats

Major Threats
The species has been exterminated from much of southern California as a result of urbanization and agricultural development (Jennings and Hayes 1994). These factors have fragmented the habitat. Excessive human recreational use and invasive alien plants (e.g., ice plant) may degrade the habitat of coastal dune populations. Other activities that have detrimentally affected populations include sand mining, golf course construction, and off-road vehicle use (Stebbins 2003). In parts of California there is now better conservation of the remaining dunes. In Baja California, the population is being negatively impacted by urbanization between Tijuana and Ensenada.
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Degree of Threat: High

Comments: The species has been exterminated from much of southern California as a result of urbanization and agricultural development (Jennings and Hayes 1994). These factors have fragmented the habitat. Excessive human recreational use and invasive exotic plants (e.g., "ice plant") may degrade the habitat of coastal dune populations. Other activites that have detrimentally affected populations include sand mining, golf course construction, and off-road vehicle use (Stebbins 2003).

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Management

Conservation Actions

Conservation Actions
Protected habitats exist in California in Asilomar State Beach, Carrizo Plain Preserve, Morro Bay State Park, Point Dume State Park (Jennings and Hayes 1994). The species also receives some protection on Pendleton Marine Corps Base and Vandenberg Air Force Base. In general, its coastal sand dune habitat is now better conserved in the United States.
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Global Protection: Few to several (1-12) occurrences appropriately protected and managed

Comments: Protected habitats exist in California in Asilomar State Beach, Pendleton Marine Corps Base, Carrizo Plain Preserve, Morro Bay State Park, Point Dume State Park, and Vandenberg Air Force Base (Jennings and Hayes 1994).

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Relevance to Humans and Ecosystems

Benefits

There are no known adverse effects of Anniella pulchra on humans.

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California legless lizards provide no known direct benefit to humans, although all species could play important roles in ecosystem stability.

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Wikipedia

Anniella pulchra

The California Legless Lizard (Anniella pulchra) is a limbless, burrowing lizard often mistaken for a snake.

Description[edit]

These lizards are around 7 inches (18 cm) long from snout to vent (not including tail). They have small, smooth scales typically colored silvery above and yellow below, although black or dark brown forms exist in Monterey County, California which were thought to be a separate subspecies at one point.[1]

Taxonomy[edit]

There were formerly two subspecies of California Legless Lizard recognized based on individual color morphs: the Silvery Legless Lizard, A. p. pulchra, and the Black Legless Lizard, A. p. nigra. However, contemporary taxonomy considers them simply a melanistic morph.[2] More recently, A. pulchra has been split up into 5 different species: A. pulchra, A. alexanderae, A. campi, A. grinnelli, and A. stebbinsi.[3][4]

Distribution and habitat[edit]

They live in loose, sandy soils or leaf litter, typically in sand dunes along the coast. They are found from Contra Costa County in northern California, all the way south to Baja California, although occurrences are often scattered. They require moisture to aid in shedding their skin. Without it, their vision and feeding can be affected, potentially starving the animal.[5]

Diet[edit]

Their diet consists of mainly beetles, larval insects, termites, ants, and spiders.

Reproduction[edit]

Males are slightly smaller than females[citation needed], otherwise there is no discernible difference between the two sexes. Females are ovoviviparous and probably breed between early spring and July, with 1 to 4 young born September-November. Young lizards resemble their parents except look like smaller versions of them.

References[edit]

Anniella.org[6]

  1. ^ CaliforniaHerps.com on Anniella pulchra
  2. ^ CaliforniaHerps.com on the former A. p. nigra "subspecies"
  3. ^ Theodore J. Papenfuss and James F. Parham (2013) Four New Species of California Legless Lizards (Anniella). Breviora Sep 2013: 1-17
  4. ^ Anniella in the Reptile Database
  5. ^ Animal Diversity Web: Anniella pulchra
  6. ^ http://www.anniella.org
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Names and Taxonomy

Taxonomy

Comments: A proposed change in the specific name from pulchra to nigra (Hunt 1983) was rejected by most herpetologists (see Murphy and Smith 1985, 1991; Ballinger et al. 1992; Jennings et al. 1992); 1993 rulings by the ICZN conserved the name Anniella pulchra for this species and placed the name argentea (as published in the trinomen Anniella nigra argentea, on the list of rejected and invalid names (Bull. Zool. Nomen. 50(2):186-187).

Phylogenetic analysis of mtDNA data clearly separates Anniella pulchra into distinct northern and southern clades, but these clades do not correspond with the black and silver nominal subspecies nigra and pulchra (Pearse and Pogson 2000). The analyses strongly indicate that the two disjunct populations of A. p. nigra are not monophyletic; they "may have arisen independently from different ancestral populations in a parallel evolutionary repsonse to selection in cool, coastal habitats" (Pearse and Pogson 2000). Accordingly, de Queiroz and Reeder (in Crother 2008) did not recognize subspecies in A. pulchra.

Molecular data support recognition of the family Anniellidae as a monophyletic group (Macey et al. 1999).

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