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A Guide to the Amphibians
and Reptiles of California


Monterey Ensatina -
Ensatina eschscholtzii eschscholtzii

Gray, 1850
Click on a picture for a larger view



Ensatina California Range Map
Range in California: Purple

Click the map for a key to
the other Ensatina subspecies



observation link



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Monterey Ensatina Monterey Ensatina Monterey Ensatina
Adult, San Luis Obispo County Adult, Los Angeles County Adult, Santa Ana Mountains,
Orange County  © Jason Jones
Monterey Ensatina Monterey Ensatina Monterey Ensatina
  Adult, San Luis Obispo County  
Monterey Ensatina Monterey Ensatina Monterey Ensatina
Juvenile from intergrade zone,
Monterey County
Adult from intergrade zone,
Monterey County
Juvenile, San Luis Obispo County
Monterey Ensatina Monterey Ensatina Monterey Ensatina
Adult, San Gabriel Mountains, Los Angeles County © Brad Alexander Adult, San Luis Obispo County Juvenile, San Luis Obispo County
Monterey Ensatina Monterey Ensatina Monterey Ensatina
Adult, San Diego County © Stuart Young Tail is constricted at the base Adult from the intergrade zone with E. e. xanthoptica - Yellow-eyed Ensatina, coastal Santa Cruz County.
© Scott Peden
Monterey Ensatina Monterey Ensatina Monterey Ensatina
Pale, probably Leucistic, Adult, San Bernardino County. © Brian Hinds Adult from Idyllwild, Riverside County.
© Stuart Young
Monterey Ensatina Monterey Ensatinas Monterey Ensatina
Adult, Mt. Palomar, San Diego County
© Stuart Young
Adults and juvenile from Idyllwild, Riverside County. © Stuart Young Juvenile, Monterey County © Dave Feliz
     
Females With Eggs
Monterey Ensatinas
On August 3rd, Joe Garcia found these intergrade Ensatinas attending their eggs under a board underneath a
house in Monterey County.  Female Ensatinas stay with their eggs to protect them until they hatch. © Joe Garcia
Monterey Ensatina Monterey Ensatina
On September 19th, Joe returned to the crawl space, looked under the board, and found that most of
the eggs of one female had just hatched, with at least 10 hatchlings still next to the eggs. © Joe Garcia
Monterey Ensatinas Monterey Ensatinas Monterey Ensatinas
Two days later, all of the eggs of both females had hatched and the juveniles were still with the females. © Joe Garcia
 
Habitat
Monterey Ensatina Habitat Monterey Ensatina Habitat Monterey Ensatina Habitat
Coastal Redwood Forest Habitat,
Monterey County
Habitat,
Monterey County
Coastal Redwood Forest Habitat,
Monterey County
Monterey Ensatina Habitat Monterey Ensatina Habitat Monterey Ensatina Habitat
Habitat, San Luis Obispo County Habitat, 3,800 ft.,
Los Angeles County
Habitat, San Luis Obispo County
     
Short Video
  Monterey Ensatina  
  A juvenile and an adult Monterey Ensatina are uncovered in the redwoods.  
   
Description
 
Size
Adult Ensatina measure from 1.5 - 3.2 inches long (3.8 - 8.1 cm) from snout to vent, and 3 - 6 inches (7.5 - 15.5 cm) in total length.

Appearance
A medium-sized salamander.
The legs are long, and the body is relatively short, with 12 - 13 costal grooves.
Nasolabial grooves are present.
The tail is rounded and constricted at the base, which will differentiate this salamander from its neighbors.
Color and Pattern
This subspecies is reddish brown to pinkish brown above, and whitish below, with orange or reddish-orange on the base of the limbs.
The eyes are very dark with no yellow markings.
Male / Female Differences
Males have longer, more slender tails than females, and a shorter snout with an enlarged upper lip, while the bodies of females are usually shorter and fatter than the bodies of males.

Life History and Behavior
A member of family Plethodontidae, the Plethodontid or Lungless Salamanders.

Plethodontid salamanders do not breathe through lungs. They conduct respiration through their skin and their mouth tissues, which requires them to live in damp environments on land and to move about on the ground only during times of high humidity. (Plethodontid salamanders native to California do not inhabit streams or bodies of water but they are capable of surviving for some time if they fall into water.)

Plethodontid salamanders are also distinguished by their naso-labial grooves, which are vertical slits between the nostrils and upper lip that are lined with glands associated with chemoreception.

All Plethodontid Salamanders native to California lay eggs in moist places on land.
The young develop in the egg and hatch directly into a tiny terrestrial salamander with the same body form as an adult.
(They do not hatch in the water and begin their lives as tiny swimming larvae breathing through gills like some other types of salamanders.)
Activity
Ensatina live in relatively cool moist places on land becoming most active on rainy or wet nights when temperatures are moderate. They stay underground during hot and dry periods where they are able to tolerate considerable dehydration. They may also continue to feed underground during the summer months.
High-altitude populations are also inactive during severe winter cold.

Adults have been observed marking and defending territories outside of the breeding season.
Territoriality
Adults have been observed marking and defending territories outside of the breeding season.
Longevity
Longevity has been estimated at up to 15 years.
Defense
When severely threatened, an Ensatina may drop its tail to distract the attention of a predator towards the writhing tail so the animal can crawl away to safety. The tail can be re-grown. The tail also contains a high density of poison glands. When disturbed, an Ensatina will stand tall in a stiff-legged defensive posture with its back swayed and the tail raised up and secrete a milky white substance from the tail, swaying the it from side to side. This noxious substance repels predators, although some experienced predators learn to eat all but the tail. If a person gets the poison on their lips, they will experience some numbness for several hours.1
Diet and Feeding
Ensatinas eat a wide variety of invertebrates, including worms, ants, beetles, spiders, scorpions, centipedes, millipedes, sow bugs, and snails.
They expell a relatively long sticky tongue from the mouth to capture the prey and pull it back into the mouth where it is crushed and killed, then swallowed.
Typically feeding is done using sit-and-wait ambush tactics, but sometimes Ensatinas will slowly stalk their prey.
Sound
Rarely, Ensatina make a hissing sound, similar to the hissing of a snake, when threatened. (Stebbins 1951; Brodie, 1978.)
Breeding
Reproduction is terrestrial.
Breeding takes place in Fall and Spring, but may also occur throughout the winter.
Stebbins describes an elaborate Ensatina courtship involving the male rubbing his body and head against the female eventually dropping a sperm capsule onto the ground which the female picks up with her cloaca. (A description and illustration of this courtship can be seen here.) The female can store the sperm until she determines the time is right to fertilize her eggs.

At the end of the rainy season, typically April or May, females retreat to their aestivation site under bark, in rotting logs, or in underground animal burrows, and lay their eggs.
Eggs
Females lay 3 - 25 eggs, with 9 - 16 being average.
Females remain with the eggs to guard them until they hatch. (You can see pictures of two Ensatinas with their eggs and hatchlings here.)
In labs, eggs have hatched in 113 - 177 days.
Young
Young develop completely in the egg and hatch fully formed and probably leave the nesting site with the first saturating Fall rains, or, at higher elevations, after the snow melts.

Geographical Range
Monterey Ensatina are found in southern California and northern Baja California, from San Luis Obispo County south along the coast to the extreme northwest coast of Baja California. They are also found in the San Bernardino and San Gabriel mountains up to 6,000 ft., (1,800 m.)

Ensatina are the most widely-distributed plethodontid salamanders in the West, ranging from an isolated location in the mountains of Baja California north along the extreme northwest coast of Baja California, through most of California excluding the deserts, the central valley, and high elevations in the mountains, continuing north into Oregon and Washington west of the Cascades Mountains, and farther north into Canada along the coast of southern British Columbia. Also found on Vancouver Island.

The Ensatina range map shown here shows a very large range of intergradation in Northern California between 4 subspecies as shown by Stebbins (2003 and 2012) that at one time was considered part of the range E. e. oregonensis. Molecular studies have shown complexities that make the use of intergrade innacturate (Stebbins & McGinnis 2012), but I will continue to use the term until more studies determine the taxonomy of this region, and in the other intergrade zones in the state.

Habitat
Inhabits moist shaded evergreen and deciduous forests and oak woodlands, mixed grassland, and chaparral. Found under rocks, logs, other debris, especially bark that has peeled off and fallen beside logs and trees.
Most common where there is a lot of coarse woody debris on the forest foor. In dry or very cold weather, stays inside moist logs, animal burrows, under roots, woodrat nests, under rocks.

Notes on Taxonomy
Ensatina eschscholtzii is typically treated as a "ring" species, consisting of 7 subspecies:

E. e. croceater
E. e. eschscholtzii

E. e. klauberi

E. e. oregonensis

E. e. picta

E. e. platensis

E. e. xanthoptica


These subspecies ring the Central Valley but do not interbreed where the rings overlap in Southern California (and possibly in the central Sierra Nevada.) These contact zones are still under study.

Some researchers see Ensatina eschscholtzii as two or more species forming a superspecies complex, recognizing E. e. klauberi, found at the southern end of the ring, as a separate species - Ensatina klauberi.
E. e. eschscholtzii
has been found to hybridize with intergrades of E. e. croceator and E. e. klauberi.

Charles W. Brown explains the taxonomy of the Ensatina complex in detail, describing it as "a classical example of Darwinian evolution by gradualism; an accumulation of micro mutations that is now leading to the formation of a new species."


Ensatina in San Francisco


Determining the taxonomy of Ensatina in San Francisco County and the west side of the San Francisco Bay has been challenging:


The range map in Stebbins, Robert C. Amphibians and Reptiles of Western North America. McGraw-Hill, 1954, shows them to be intergrades, but does not indicate whether they itergrade with E. e. eschscholtzii or with E. e. oregonensis.

The range map in Stebbins, Robert C. A Field Guide to Western Amphibians and Reptiles. Houghton Mifflin. 1985. shows them to be intergrades, but does not indicate whether they itergrade with E. e. eschscholtzii or with E. e. oregonensis.

The large color range map in Thelander, Carl G., editor in chief. Life on the Edge - A Guide to California's Endangered Natural Resources - Wildlife. Berkeley: Bio Systems Books, 1994. based on two of Stebbins works appears to show them as intergrades with E. e. eschscholtzii but that is not clear either.

The range map in Petranka, James W. Salamanders of the United States and Canada. Smithsonian Institution, 1998. shows them to be E. e. oregonensis.

The range map in Stebbins, Robert C. A Field Guide to Western Amphibians and Reptiles Third Edition. Houghton Mifflin. 2003. is too small to show that part of the peninsula.

The range map in Joao Alexandrino, Stuart J. E. Baird, Lucinda Lawson, J. Robert Macey, Craig Moritz, and David B. Wake.  Strong Selection Against Hybrids at a Hybrid Zone in the Ensatina Ring Species Complex and Its Evolutionary Implications.  Evolution, 59(6), 2005, pp. 1334–1347. shows them to be E. e. oregonensis

David B. Wake. Problems with Species: Patterns and Processes of Species Formation in Salamanders. Annals of the Missouri Botanical Garden. 93: 8–23. Published on 31 May 2006. shows a range map (click here to see it) based on a preliminary analysis from a study in progress using mitochondrial DNA in which Ensatina on the SF peninsula are shown as
E. e. xanthoptica
but there are E. e. oregonensis clades show on the coast south of there. (A color map in the same paper shows the entire area west of the S.F. Bay to be E. e. oregonensis.)

A range map in Shawn R. Kuchta, Duncan S. Parks, David B. Wake. Pronounced phylogeographic structure on a small spatial scale: Geomorphological evolution and lineage history in the salamander ring species Ensatina eschscholtzii in central coastal California. Molecular Phylogenetics and Evolution 50 (2009) 240–255 shows them to be E. e. xanthoptica (with E. e. oregonensis to the southwest.) (Click here to see the map.)

The range map in Stebbins, Robert C., and McGinnis, Samuel M.  Field Guide to Amphibians and Reptiles of California: Revised Edition (California Natural History Guides) University of California Press, 2012. shows them to be E. e. xanthoptica.

There are certainly more studies with more maps which I have not seen, but since recent research shows them as E. e. xanthoptica (and photos corroborate this) I will show them as such until further research shows otherwise.

Conservation Issues  (Conservation Status)
None
Taxonomy
Family Plethodontidae Lungless Salamanders Gray, 1850
Genus Ensatina Ensatinas Gray, 1850
Species Eschscholtzii Ensatina Gray, 1850
Subspecies

eschscholtzii Monterey Ensatina Gray, 1850
Original Description
Ensatina eschscholtzii - Gray, 1850 - Cat. Spec. Amph. Coll. Brit. Mus., Batr. Grad., p. 48

from Original Description Citations for the Reptiles and Amphibians of North America © Ellin Beltz

Meaning of the Scientific Name
Ensatina: Latin - sword shaped/similar to, possibly referring to the teeth.
eschscholtzii: honors Johann F. Eschscholtz.
eschscholtzii: honors Johann F. Eschscholtz.

from Scientific and Common Names of the Reptiles and Amphibians of North America - Explained © Ellin Beltz

Alternate Names
None

Related California Salamanders
Large-blotched Ensatina
Oregon Ensatina
Painted Ensatina
Sierra Nevada Ensatina

Yellow-eyed Ensatina
Yellow-blotched Ensatina

More Information and References
California Department of Fish and Wildlife

AmphibiaWeb

Charles W. Brown's Ensatina Web Site

Stebbins, Robert C., and McGinnis, Samuel M.  Field Guide to Amphibians and Reptiles of California: Revised Edition (California Natural History Guides) University of California Press, 2012.

Stebbins, Robert C. California Amphibians and Reptiles. The University of California Press, 1972.

Stebbins, Robert C. A Field Guide to Western Reptiles and Amphibians. 3rd Edition. Houghton Mifflin Company, 2003.

Behler, John L., and F. Wayne King. The Audubon Society Field Guide to North American Reptiles and Amphibians. Alfred A. Knopf, 1992.

Powell, Robert., Joseph T. Collins, and Errol D. Hooper Jr. A Key to Amphibians and Reptiles of the Continental United States and Canada. The University Press of Kansas, 1998.

Bartlett, R. D. & Patricia P. Bartlett. Guide and Reference to the Amphibians of Western North America (North of Mexico) and Hawaii. University Press of Florida, 2009.

Bishop, Sherman C. Handbook of Salamanders. Cornell University Press, 1943.

Lannoo, Michael (Editor). Amphibian Declines: The Conservation Status of United States Species. University of California Press, June 2005.

Petranka, James W. Salamanders of the United States and Canada. Smithsonian Institution, 1998.

Joao Alexandrino, Stuart J. E. Baird, Lucinda Lawson, J. Robert Macey, Craig Moritz, and David B. Wake.  Strong Selection Against Hybrids at a Hybrid Zone in the Ensatina Ring Species Complex and Its Evolutionary Implications.  Evolution, 59(6), 2005, pp. 1334–1347.

Shawn R. Kuchta, Duncan S. Parks, David B. Wake. Pronounced phylogeographic structure on a small spatial scale: Geomorphological evolution and lineage history in the salamander ring species Ensatina eschscholtzii in central coastal California. Molecular Phylogenetics and Evolution 50 (2009) 240–255

Conservation Status

The following status listings come from the Special Animals List and the Endangered and Threatened Animals List which are published by the California Department of Fish and Wildlife.


This salamander is not included on the Special Animals List, which indicates that there are no significant conservation concerns for it in California.
Organization
Status Listing
U.S. Endangered Species Act (ESA) None
California Endangered Species Act (CESA) None
California Department of Fish and Wildlife None
Bureau of Land Management None
USDA Forest Service None
 

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