A Guide to the Amphibians
and Reptiles of California

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Long-toed Salamander - Ambystoma macrodactylum

Santa Cruz Long-toed Salamander -
Ambystoma macrodactylum croceum

Russell and Anderson, 1956

Description • Taxonomy • Species Description • Scientific Name • Alt. Names • Similar Herps • References • Conservation Status

Bright Green: Range of this subspecies in California
Ambystoma macrodactylum croceum -
Santa Cruz Long-toed Salamander

Range of other subspecies in California:

Red: Ambystoma macrodactylum sigillatum -
Southern Long-toed Salamander

Click on the top map for a topographical view.
Click on the bottom map for a larger view.

Map with California County Names

Dark Blue: Range of this subspecies

Long-toed Salamander
Eggs and Larvae

Adult, Santa Cruz County

Adult, Santa Cruz County
© Spencer Riffle

Adult, Santa Cruz County © Zach Lim

Adult, Santa Cruz County
© Brad Alexander

Adult, Santa Cruz County

Adult, Monterey County © Zach Lim

Adult, Santa Cruz County © Ryan Sikola

Adult, Santa Cruz County
© Spencer Riffle

Adult, Monterey County © Dave Feliz

Adult, Santa Cruz County
© Spencer Riffle

This dark and nearly unmarked adult was found in Santa Cruz County.
© James Maughn

This Santa Cruz County salamander was found climbing some grass at night, apparently hunting for food, probably small slugs. © Jared Heald

This Santa Cruz County salamander is displaying a defensive posture after being disturbed by bright light at night.
© Jared Heald

Adults found on a rainy night in Santa Cruz County © Adam Gitmed

This Santa Cruz County salamander was discovered eating an earthworm.
© Jared Heald

Adult, Santa Cruz County
© Ryan Sikola

Adult male, Santa Cruz County,
handled with proper federal and state permits. © Leyna Stemle

Adult female, Santa Cruz County,
handled with proper federal and state permits. © Leyna Stemle

Adult, Santa Cruz County
© Zeev Nitzan Ginsburg

Adult, dorsal pattern close-up, Santa Cruz County © Ryan Sikola

Looking at the salamander on the left and the willow leaf on the right you can see how the pattern on the salamander helps it blend into its environment to remain undetected. © Ryan Sikola

Recently-metamorphosed juvenile, Santa Cruz County, with native willow leaves they blend in with that are common in the breeding area, handled with proper federal and state permits.
© Leyna Stemle

An elongated toe (number 4) on each hind foot is the "long toe" that gives this species its common name.

Comparison of two sympatric Ambystomatid salamanders:
Left: Santa Cruz Long-toed Salamander
Right: California Tiger Salamander,
handled with proper federal and state permits. © Leyna Stemle

Transformed Juveniles

Two recently-metamorphosed juveniles, Monterey County © Dave Feliz

Recently-metamorphosed juvenile, Monterey County © Dave Feliz

Metamorph, Santa Cruz County
© Leyna Stemle

Juvenile © Jon Hirt

Juvenile, Santa Cruz County
© Jon Hirt

Larvae

Larva in water, Monterey County © Dave Feliz

Larva in water, Monterey County
© Dave Feliz

Larva, Santa Cruz County
© Aidan O'Brien

Aquatic Larvae, Santa Cruz County © Leyna Stemle

Aquatic Larvae, Santa Cruz County
The second larva from the left has almost completed metamorphosis, having reabsorbed most of its gills and excess dorsal tail. The others show no signs of metamorphosis yet.
© Leyna Stemle

Aquatic Larva, USFWS

Sympatric Salamander Larvae

Taricha granulosa

There are large dark areas on either side of the light coloring around the pupil

Ambystoma macrodactylum croceum

There are no large dark areas in the light coloring around the pupil

Ambystoma californiense

Snout is broader, more shovel-shaped

Color is mostly gray, or with a little green or blue

Toe is more wedge-shaped

Ambystoma macrodactylum croceum

Snout is more rounded

Color ranges from black to mottled green and tan to gray

Toe is a little more skinny and long

More Pictures of Eggs, Larvae, and Young of Other Subspecies of Long-toed Salamanders

Habitat

Dry breeding pond in November,
Santa Cruz County

Breeding pond, winter,
Santa Cruz County

Breeding pond, late winter
Santa Cruz County

Wildlife refuge habitat (name removed)
Santa Cruz County

Description

Size

Adults are 1 3/5 - 3 1/2 inches long (4.1 - 8.9 cm) from snout to vent, 4 - 6 2/3 inches (10 - 17 cm) in total length.

Appearance

A medium-sized salamander.
The body is stout with 12 - 13 costal grooves and a broad rounded head, a blunt snout, small protuberant eyes, and no nasolabial grooves.
The tail is flattened from side to side to facilitate swimming.

Color and Pattern

Black above with an orange dorsal stripe, broken into spots and bars.
The sides are sprinkled with whitish specks.
The venter is grey or black.

Young

Larvae have broad heads, three pairs of bushy gills and broad caudal fins that extend well onto the back.

Life History and Behavior

A member of the Mole Salamander family (Ambystomatidae) whose members are medium to large in size with heavy, stocky bodies.

Ambystomatid salamanders have two distinct life phases:
- Larvae hatch from eggs laid in water where they swim using an enlarged tail fin and breathe with filamentous external gills. - Aquatic larvae transform into four-legged salamanders that live on the ground and breathe air with lungs.

Transformed adults are terrestrial and breathe with lungs but some gilled adults remain in the water and grow to a large size before transforming. However, neotenic adults have not been reported.

Activity

Adults spend much of their lives underground, often utilizing the tunnels of burrowing mammals such as moles and ground squirrels.

Transformed adults are rarely found outside of the breeding season.
They are mostly found under wood, logs, rocks, bark and other objects near breeding sites, or when they are breeding in the water. At other times of the year they stay in rotten logs or moist places underground such as animal burrows.

Adults migrate to breeding sites, then return to terrestrial habitats.

Longevity

Adults live to about 10 years of age.

Defense and Sound

Adults produce sticky skin secretions to deter predators, and they can vocalize with squeaks and clicks, which might startle predators who capture them. (Hossack, B. R. 2002. natural history notes: Ambystoma macrodactylum krausei (northern long-toed salamander). Vocalization. Herpetological Review 33:121.)

Diet and Feeding

Carnivorous.
Transformed adults eat small invertebrates, including worms, mollusks, insects, and spiders.
Larvae start by eating small crustaceans. As they increase in size, they gradually consume larger prey items, including crustaceans, worms, mollusks, and frog tadpoles.
Larger larvae may cannibalize smaller larvae.
Young larvae feed by sitting and waiting for prey, while larger larvae also stalk and pursue prey.

Reproduction

Reproduction is aquatic. Adults become sexually mature at 1 - 3 years, and migrate overland to the breeding site during nights with heavy rain from October through February with breeding occurring in January and February. Males enter the ponds before females.
Adults remain in the ponds from several days to more than a month.

Eggs

Females lay from 90 - 400 eggs in clusters containing from 1 - 81 eggs in shallow water, attaching them singly or in loose clusters to the undersides of logs and branches, or leaving them unattached on the bottom.
Eggs hatch in 2 - 5 weeks.

Young

Drying of ponds triggers transformation.
Larvae transform in 4 - 5 months in temporary ponds.
Larvae may not transform the first season.
Young remain at the pond sites until the first rains in the fall.

Habitat

Found in dense riparian vegetation such as willows, thick coastal scrub, and oak woodland.

Geographical Range

The species Ambystoma macrodactylum - Long-toed Salamander, is widespread in the West, occurring in California, Oregon, Idaho, and Montana, western Canada, and Southeast Alaska.

This subspecies, Ambystoma macrodactylum croceum - Santa Cruz Long-toed Salamander, is endemic to California, inhabiting a very limited range with scattered populations in a reported 11 locations (U.S.F.W. 1999.) around the coast of Monterey Bay in southern Santa Cruz County and the northern edge of Monterey County. It is thought to be a relict population, now isolated from the rest of its species.

Notes on Taxonomy

Five subspecies of Ambysoma macrodactylum are traditionally recognized, two occur in California:
A. m. sigillatum
A. m. croceum.
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"Lee-Yaw and Irwin (2012, Journal of Evolutionary Biology. 25: 2276–2287) and Lee-Yaw et al. (2014, Molecular Ecology 23: 4590–4602) evaluated geographic variation of mtDNA and nuclear genes throughout the range of the species and found the distributions of five lineages did not completely agree with those of the five presently recognized subspecies but suggested no changes in the taxonomy of the species. Raffaëlli (2022, Salamanders & Newts of the World. Plumelec, Penclen) continued to recognize the five subspecies."
(Nicholson, K. E. (ed.). 2025 SSAR Scientific and Standard English Names List)
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A 2015 study by Lee-Yaw and Irwin has identified a sixth distinct group in the Central Oregon highlands, but suggested no changes in the taxonomy of the species. (You can see this new group and my estimate of the ranges of all the subspecies described in the paper HERE.)
(J. A. Lee-Yaw & D. E. Irwin. The mportance (or lack thereof) of niche divergence to the maintenance of a northern species complex: the case of the long-toed salamander (Ambystoma macrodactylum Baird) Journal of Evolutionary Biology 28 (2015) 917-930)
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Alternate and Previous Names (Synonyms)

Ambystoma macrodactylum croceum - Santa Cruz Long-toed Salamander (Stebbins 1966, 1985, 2003, 2012)
Ambystoma macrodactylum - Long-toed Salamander (Storer 1925, Bishop 1943, Stebbins 1954)
Ambystoma macrodactyla (Baird 1850)

Conservation Issues (Conservation Status)

This subspecies is listed as federally endangered and heavily protected due to its limited range combined with loss of habitat to land development.

"Land use changes (especially urban and agricultural development) have reduced the amount of suitable upland habitat within migration distance of SCLTS [Santa Cruz Long-toed Salamander] breeding ponds by 35.5% across the range. Habitat fragmentation due to roads has further reduced uplands both suitable and accessible to SCLTS by another 11.8%, and sea level rise projected by 2100 reduces it another 16.2%, leaving only 36.5% of potential terrestrial habitat suitable, accessible, and unflooded. This result raises substantial concerns about the long-term viability of the majority of SCLTS populations."

(Leyna R. Stemle, David Laabs, and Christopher A. Searcy. Land use change, fragmentation, and sea level rise create escalating viability concerns for an imperiled salamander. Ecosphere. 2025;16:e70284.)

A project to rehabilitate a drying breeding pond at a National Wildlife Refuge was undertaken in 2014. Larvae were captured and raised in aquariums and then successfully released back into the wild.

Taxonomy
Family	Ambystomatidae	Mole Salamanders	Gray, 1850
Genus	Ambystoma	Mole Salamanders	Tschudi, 1838
Species	macrodactylum	Long-toed Salamander	Baird, 1849
Subspecies	croceum	Santa Cruz Long-toed Salamander	Russell and Anderson, 1956
Original Description
Ambystoma macrodactylum - Baird, 1849 - Journ. Acad. Nat. Sci. Philadelphia, Ser. 2, Vol. 1, p. 292 Ambystoma macrodactylum croceum - Russell and Anderson, 1956 - Herpetologica, Vol. 12, p. 137 from Original Description Citations for the Reptiles and Amphibians of North America © Ellin Beltz
Meaning of the Scientific Name
Ambystoma - Greek - amblys = blunt + stoma = mouth macrodactylum - Greek = long toe croceum - Latin = saffron colored, referring to the dull orange dorsal stripe. from Scientific and Common Names of the Reptiles and Amphibians of North America - Explained © Ellin Beltz
Related or Similar California Salamanders
Southern Long-toed Salamander California Tiger Salamander
More Information and References
California Department of Fish and Wildlife AmphibiaWeb Leyna R. Stemle, David Laabs, and Christopher A. Searcy. Land use change, fragmentation, and sea level rise create escalating viability concerns for an imperiled salamander. Ecosphere. 2025;16:e70284. Thelander, Carl G., editor in chief. Life on the Edge - A Guide to California's Endangered Natural Resources - Wildlife. Berkeley: Bio Systems Books, 1994. Hansen, Robert W. and Shedd, Jackson D. California Amphibians and Reptiles. (Princeton Field Guides.) Princeton University Press, 2025. 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. Flaxington, William C. Amphibians and Reptiles of California: Field Observations, Distribution, and Natural History. Fieldnotes Press, Anaheim, California, 2021. Nicholson, K. E. (ed.). 2025. Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, with Comments Regarding Confidence in Our Understanding. Ninth Edition. Society for the Study of Amphibians and Reptiles. [SSAR] 87pp. Samuel M. McGinnis and Robert C. Stebbins. Peterson Field Guide to Western Reptiles & Amphibians. 4th Edition. Houghton Mifflin Harcourt Publishing Company, 2018. 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. Robert Powell, Roger Conant, and Joseph T. Collins. Peterson Field Guide to Reptiles and Amphibians of Eastern and Central North America. Fourth Edition. Houghton Mifflin Harcourt, 2016. 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. American Museum of Natural History - Amphibian Species of the World 6.2 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.

Conservation Status
The following conservation status listings for this animal are taken from the April 2024 State of California Special Animals List and the April 2024 Federally Listed Endangered and Threatened Animals of California list (unless indicated otherwise below.) Both lists are produced by multiple agencies every year, and sometimes more than once per year, so the conservation status listing information found below might not be from the most recent lists. To make sure you are seeing the most recent listings, go to this California Department of Fish and Wildlife web page where you can search for and download both lists: https://www.wildlife.ca.gov/Data/CNDDB/Plants-and-Animals. A detailed explanation of the meaning of the status listing symbols can be found at the beginning of the two lists. For quick reference, I have included them on my Special Status Information page. If no status is listed here, the animal is not included on either list. This most likely indicates that there are no serious conservation concerns for the animal. To find out more about an animal's status you can also go to the NatureServe and IUCN websites to check their rankings. Check the current California Department of Fish and Wildlife sport fishing regulations to find out if this animal can be legally pursued and handled or collected with possession of a current fishing license. You can also look at the summary of the sport fishing regulations as they apply only to reptiles and amphibians that has been made for this website.
Organization	Status Listing	Notes
NatureServe Global Ranking	G5 T1 T2	Species Secure. Subspecies Critically Imperiled - Imperiled
NatureServe State Ranking	S2	Imperiled
U.S. Endangered Species Act (ESA)	FE	Listed as Endangered 3/11/67
California Endangered Species Act (CESA)	SE	Listed as Endangered 6/27/71
California Department of Fish and Wildlife	FP	Fully Protected
Bureau of Land Management	None
USDA Forest Service	None
IUCN

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Santa Cruz Long-toed Salamander - Ambystoma macrodactylum croceum

Santa Cruz Long-toed Salamander -
Ambystoma macrodactylum croceum