A Guide to the Amphibians
and Reptiles of California

Arboreal Salamander - Aneides lugubris

(Hallowell, 1849)

Click on a picture for a larger view

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

Red: Range in California

Click on the map for a topographical view

Map with California County Names

Adult with small spots, Contra Costa County

Adult with small spots,
Alameda County

Adult with small spots,
Santa Cruz County

Adult with small spots, Contra Costa County (missing the tip of its tail.)

Adult with small spots, Contra Costa County

Adult, found on a building in downtown San Francisco, San Francisco County
© Jackson Shedd

Adult with large spots, Monterey County

Adult male with large spots, Monterey County coastal sand dunes

Adult active at night in situ,
Marin County

Adult in situ emerging at night from a rock crevice, Marin County

Adult male with large head scar, Marin County

Adult, Santa Clara County
© aquariumkids.com
This large adult salamander crawled under the crack of the front door of a house on a rainy night in January.

Adult with few small spots,
from Mendocino County

Adult, Anchor Bay, Mendocino County
© Richard L. P. Custer

Adult, Stanislaus County © Jared Heald

This adult was rescued from the bottom of the deep end of a swimming pool in San Luis Obispo County. © Laurie Dey

Adult, Alameda County

Adult, San Mateo County © Jared Heald

Adult with small spots, found under a garbage can after the rains started in October in Santa Clara County.
© John Roberts

Adult, San Luis Obispo county,
© Andrew Harmer

Adult, San Luis Obispo County
© Henk Wallays

Adult with small spots, San Diego County © Paul Maier

Adult, Orange County © Jason Jones

Two Adults found in the Santa Ana Mountains, Orange County © Cooper Bailey

Adult, Los Angeles County
© Gregory Litiatco

Adult with small spots, San Diego County © Stuart Young

Adults with small spots, San Diego County © Jay Keller

Adult with small spots, Orange County
© Tadd Kraft

Adult, Coastal Southern California © Ivan Vershynin

Adult with a nice city view, San Luis Obispo County © Spencer Riffle

Adult hunting in situ, San Luis Obispo County © Spencer Riffle

This adult was found on a road on a rainy night in a surprisingly dry, treeless area in the foothills of far eastern Alameda County © Noah Morales

Adult, western Stanislaus County
© Noah Morales

Adult male, Santa Monica Mountains, Los Angeles County © Hugo Rojas

Adult, Sierra Nevada Mountains, Tuolumne County © Rick Staub

Subadult from the Santa Monica Mountains, Los Angeles County
© Max Roberts

Adult with heavy spotting,
Contra Costa County © Max Roberts

Adult, Los Angeles County © Max Roberts

Adult with minimal spotting found at the entrance of a crevice
in a rock face (right) in Los Angeles County © Max Roberts

Adult climbing a Coast Live Oak (Quercus agrifolia) in Los Angeles County © Max Roberts

Adult in rock crevice in Los Angeles County © Max Roberts

Arboreal Salamanders have toes
with expanded tips that help them climb vertical surfaces like tree trunks and cliffs.

Arboreal Salamanders from Southeast Farallon Island

Adult with large spots Southeast Farallon Island © Oscar Johnson

Adult with very large spots, Southeast Farallon Island © Morgan Stickrod

Juveniles

Juvenile, Santa Cruz County

Hatchling, Marin County

Hatchling, San Luis Obispo County
© Spencer Riffle

Juvenile, Marin County

Juvenile, Santa Cruz County
© Mark Gary

Hatchling, San Luis Obispo County
© Spencer Riffle

Juvenile, Santa Cruz County
© Brad Alexander

Hatchling, San Luis Obispo County
© Spencer Riffle

Juvenile, Calaveras County
© Noah Morales

Bluish juvenile, found on a rocky slope at about 3000 ft. elevation in Mariposa County © Adam Gitmed

Yearling, Los Angeles County © Max Roberts

Male Combat and Defense

Two adult males locked in bloody combat in Santa Cruz County © Sasha Robinson. The picture on the far right was taken about an hour after the first two pictures and they were still fighting.

Be careful when handling this salamander! It has lots of sharp teeth and isn't afraid to use them.

Mike Spencer found it, Val Johnson took the pictures, and Shannon Hoss lost some blood when she picked up this toothy beast in Mendocino County. © Val Johnson

Adult, Santa Barbara County © Ryan Sikola
This tough survivor is missing an eye and part of its face, but still appears healthy.
Meles compete in violent combat, which might have produced this injury, but it could also be the result of an encounter with a predator.

Arboreal Salamanders with an Aberrant or Unusual Appearance

Top: Typical Arboreal Salamander.
Bottom: Albino or pigmentless adult Arboreal Salamander.
Both were found in the same general location in Contra Costa County.
© 2006 John Schilling

Unusually pale adult, Marin County © Andre Giraldi

This salamander is much lighter in color than a typical Arboreal Salamander. Other typically colored salamanders were found near it. It could be missing some of its melanin or dark pigment, or its appearance could be a result of the salamander changing its color, which was described by Robert Stebbins. See description below.

These unusualy-pigmented adults were found together in San Luis Obispo County -
Unspotted (left)
Leucistic (right)
© Ryan Sikola

Leucistic adult, San Luis Obispo County © Ryan Sikola

Adult with no light spotting,
San Luis Obispo County
© Ryan Sikola

This adult from Santa Clara County has a mottled head.
© Katherine Hughes

Aestivation and Egg Brooding

During the dry period, usually from May or June to October or November, Arboreal Salamanders hide out in cool moist areas, usually underground, sometimes in tree hollows, sometimes in basements and underground garages. Females lay their eggs in late Spring and early Summer and they are sometimes found guarding their eggs along with other aestivating salamanders.

The adult Arboreal Salamander shown on the left (most likely a female) was found in an irrigation box in Sonoma County guarding a cluster of eggs attached to the box above her. The eggs were laid sometime at the beginning of July and they hatched in early October. The salamander stayed with them the entire time and continued there throughout the winter after they had hatched. The picture on the right shows the adult below a cluster of approximately ten hatchlings a day or two after hatching. They stayed in this cluster for several days. Afterwards the number of adult and juvenile salamanders observed in the box varies as they move in and out, with as many as ten observed at one time.© Sohni Yaylian

Old egg cluster that was found attached to the underside of a log on the ground in Contra Costa County. A hatchling was seen just below the egg cluster.
© Max Roberts

These eggs were found underneath some old redwood planks in a shaded area next to a house in Sonoma County. Though they were found near several aestivating Arboreal Salamanders in July when they lay their eggs, they are most likely the eggs of a slug or snail because of their opacity and because they don't appear to be attached to the wood. Gastropod eggs are sometimes confused for salamander eggs because of their similar size and because they are both laid in similar dark moist places.
© Anonymous

Eric Boyer discovered this large group of in situ estivating adult and juvenile Arboreal salamanders underneath some wood while remodeling a backyard in Santa Barbara County in June.
© Eric Boyer

This in situ Arboreal Salamander was found underneath the circular base of a large statue in a Santa Clara County backyard in August.
© Laurie Weber

Feeding

Juvenile eating an insect larva or worm in San Luis Obispo County © Ryan Sikola

Predation

An adult San Bernardino Ring-necked snake eating an adult Arboreal Salamander in Los Angeles County © Jonathan Benson

Habitat

Typical oak woodland habitat, Contra Costa County

Habitat, Contra Costa County

Arboreal salamanders inhabit this urban backyard in Oakland,
Alameda County

Coastal sand dunes habitat,
Monterey County

Rocky habitat, Marin County

Habitat at coastal marsh,
San Mateo County

Arboreal salamanders inhabit nearly-treeless, rocky, Southeast Farallon Island.

Arboreal salamanders inhabit
rocky Año Nuevo Island.

A careful look underneath the fallen branches and bark of the dead tree shown above on a wet winter afternoon turned up 16 salamanders of 4 species - one Arboreal Salamander, two Coast Range Newts, one Yellow-eyed Ensatina, and 12 California Slender Salamanders, proving that wood debris on a forest floor is an important microhabitat for salamanders. Along with fallen debris, tree bark, tree cavities, root holes, and splits in trees are also useful habitat for many kinds of wildlife, including birds, mammals, amphibians and reptiles, but dead trees and their debris are often removed indiscriminately without consideration for wildlife.

The Cavity Conservation Initiative is a group whose goal is to educate land managers and the public about the value of dead trees.

Short Video and Audio

An Arboreal Salamander
resting and running.

Several Arboreal Salamanders are seen hiding in cracks in a rock wall at night. A large one out on the rocks crawls back into a hiding spot.

Listen to this sub-adult Marin County Arboreal Salamander squeak two times.
(There's a faint high-pitched squeak, then some rustling background noise, then another faint squeak.) The salamander squeaked several times when the log under which it was resting was lifted.
Sound and photo © Jonathan Hakim

An Arboreal Salamander, found sitting out on a rock at night, chirps when I grasp it across the sides firmly with my thumb and one finger. The salamander was not squeezed so hard that the sound was just the sound of air being pushed out of its lungs. This species is capable of making a fairly loud sound, but this one was very quiet, so there is a lot of background noise from a flowing creek nearby.

Description

Size

Adults measure 2 1/4 - 4 inches long (5.7 - 10.1 cm) from snout to vent and up to 7 inches (18 cm) in total length. The largest species of Aneides.

Appearance

A medium sized lungless salamander.
Toe tips are expanded and squarish.
Tail is semi-prehensile, often seen coiled.
Usually 15 costal grooves.
Two nasolabial grooves.

Color and Pattern

Color is brown above with small cream to yellow spots.
The number and size of the spots varies with location - Most Sierra Nevada and North Coast populations have weak spotting, while Gabilan Mountains and Southeast Farallon Island populations have large conspicuous spots. Other areas have populations with small spots and populations with intermediate spots.
(Morafka, 1976)

The venter is creamy white.
The undersides of the tail and feet are dull yellow.

On page 129 of his 1962 book "Amphibians of North America, Robert Stebbins writes that Aneides lugubris is:
"Capable of marked color change. When kept in a dark, cold room at 12 degrees C., 6 adults were dark-colored, with the dorsal light-colored cloudings obscure. In the laboratory at 21 degrees C., where well illuminated, the light-colored markings of the animals became conspicuous, almost dominating the dark ground color. The changes evidently are achieved principally by changes in the melanic pigmentation."

Male / Female Differences

Males have broader, more triangular heads than females.

Young

Young are dark, clouded with gray or brassy color.

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 the tissues lining their mouth. This 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 a short 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

Nocturnal.
Active when soil moisture is high after the onset of fall rains, usually in November, to May, but relatively tolerant of dry conditions compared to many other salamander species.
Strong jaws and sharp teeth are capable of producing a painful bite.
Adapted for climbing with long toes and rounded semi-prehensile.
Has been found up to 59 ft. (18 m) above the ground.

Territoriality

Both males and females are agressively territorial. Individuals covered with scars (probably from fighting) are often found, and captives kept together often bite the other salamander's tail. Since the species is often found sheltering together in large numbers in dry weather, it appears that this territorial agression is not constant.

Defense

Anti-predator behaviors include biting, a raised defensive posture, making a squeaking sound, fleeing rapidly, and jumping.

Sound

This salamander may squeak when it is picked up or disturbed.
(Here's a recorded example - There's a faint high-pitched squeak, then some rustling background noise, then another faint squeak.)
According to Stebbins and Cohen, 1997, an Arboreal Salamander "...may squeak repeatedly when caught, retracting its eyes into their sockets each time a sound is produced, When the eyeballs are depressed, their undersides protrude into the mouth cavity, thereby compressing the air in the mouth and forcing it outward. It is this movement that appears somehow to cause the sounds produced."

Diet and Feeding

Eats a variety of small invertebrates including millipedes, worms, snails and slugs, ants, termites, sowbugs, moths, and centipedes.
Also known to eat slender salamanders (Batrachoseps.)
Prey is captured by the tongue and brought into the mouth where it is crushed.
A sit-and-wait-predator, adults forage for small invertebrates and sometimes slender salamanders on the ground at night during wet weather.

Reproduction

Reproduction is terrestrial.
Adults probably reach sexual maturity in their third year.
I have not yet found any documentation that describes the time of year when this species breeds, but it is most likely during late winter and early spring, since eggs are laid in late spring to early summer.
Breeding males have a heart-shaped mental gland under their chin. Males put this gland on a female's back, stroking her back quickly with it during courtship while scratching her skin with his teeth to deliver the mental gland pheromones to the female.

Eggs

In late spring and early summer, females lay from 5 - 24 eggs in moist places, most commonly in decaying cavities of live oak trees, sometimes high off the ground, and also under rocks and other surface objects and inside logs.
Females usually remain with the eggs until they hatch, often coiled around them.
Sometimes the eggs of several salamanders are found together in large masses.
Males have also been found with females at the egg deposition site and adults and juveniles may stay together after hatching, perhaps waiting for the onset of the fall rains which will allow them to disperse from their refuge.

Young

Young develop completely in the egg and hatch fully formed in August and September. (Staub and Wake in Lannoo 2005)
They may stay with the male and female for a time after hatching.

Habitat

Lives in moist places on land, mostly in coastal oak woodlands, but also found in yellow pine and black oak forests in the Sierra Nevada and other dryer habitats, including coastal sand dunes. Also found on moist, mossy rock faces, under rocks and woody debris on land, inside stumps, and in urban yards and buildings. In Southern California, this salamander is also associated with sycamores along seasonal streams.

Geographical Range

Endemic to California and northern Baja California.
Occurs from Humboldt county, south along the coast and coast ranges into Baja California del Norte, and in the foothills of the Sierra Nevada Mountains from El Dorado County to Madera County.

Also found on several islands off the coast of California - the Farallon Islands, Año Nuevo Island, and Catalina Island, and on Los Coronados Island off the coast of Baja California del Norte.

(The Santa Catalina Island record for this species is based on only one specimen found in 1941.
A three-year US Geological Survey biodiversity survey of the island failed to find any evidence of the species, suggesting that the single specimen was brought to the island from the Southern California mainland, probably with shipments of ranching materials, and it failed to establish a population.

[Backlin A, Compton S, Kahancza Z, Fisher R. 2005, cited in Jockusch EL, Hansen RW, Fisher RN, Wake DB. 2020. Slender salamanders (genus Batrachoseps) reveal Southern California to be a center for the diversification, persistence, and introduction of salamander lineages. PeerJ 8:e9599 https://doi.org/10.7717/peerj.9599])

Elevational Range

From sea level to 5,000 ft. (1,500 m). (Stebbins, 2003)

Notes on Taxonomy

No subspecies are currently recognized, but two subspecies have been recognized in the past - Aneides lugubris lugubris - Arboreal Salamander, and Aneides lugubris farallonensis - Farallon Salamander or Farallon Yellow-spotted Salamander (found only on South Farallon Island).

DNA studies have shown that there are genetically different groups within the species. Sean B. Reilly et al ** in a study of the phylogeographic history of Aneides lugubris published in 2015 found substantial genetic diversity in the species due to historical isolation of populations. Their summary of the results and their conclusions are reproduced below (emphasis mine), and a copy of one of the maps from the paper can be seen by clicking on the range map below, all permitted under a Creative Commons license. The entire paper is available online at biomedcentral.com.

"Results: We found six major mitochondrial clades in A. lugubris. Nuclear loci supported the existence of at least three genetically distinct groups, corresponding to populations north of the San Francisco Bay and in the Sierra Nevada, in the Santa Cruz Mountains, and in the central coast and southern California. All of the genetic breaks in mitochondrial and nuclear loci corresponded to regions where historical barriers to dispersal have been observed in other species. Geologic or water barriers likely were the most important factors restricting gene flow among clades. Climatic unsuitability during glacial maximum may have contributed to the isolation of the mitochondrial clades in the central coast and southern California. A projection of our species distribution model to a future scenario with a moderate amount of climate change suggests that most of the range of A. lugubris will remain climatically suitable, but climatic conditions in the Sierra Nevada and low elevation areas in Southern California are likely to deteriorate."

Conclusions: Aneides lugubris contains substantial cryptic genetic diversity as a result of historical isolation of populations. At least two (and perhaps three) evolutionarily significant units in A. lugubris merit protection; all six mitochondrial clades should be considered as management units within the species."

Below is a map I made showing my estimates of the ranges of the 6 clades of A. lugubris, extrapolated from the map included in the publication. Areas where I am in doubt which clade is present I have marked with gray. Approximate locations of the samples used on which I based the shaded portions of the map are shown as darker dots. Click on the map below to see the original map from the publication.

Estimated ranges of the 6 clades of Aneides lugubris (Gray = clade uncertain.)

** Sean B. Reilly, Ammon Corl, and David B. Wake. An integrative approach to phylogeography: investigating the effects of ancient seaways, climate, and historical geology on multi-locus phylogeographic boundaries of the Arboreal Salamander (Aneides lugubris). BMC Evolutionary Biology (November 2015) 15:241.

"There are two geographically segregated groups of chromosomally differentiated populations of arboreal salamanders (Sessions and Kezer, 1987). These two karyotypes intergrade in south and east-central Mendocino County (Sessions and Kezer, 1987). Unpublished genetic analyses (allozymes and mitochondrial DNA sequences) show that the chromosomal units do not correlate with patterns of genetic variation (Jackman, 1993). The Farallon Island population is most similar genetically to the nearest mainland population, not populations in the Gabilan mountains to the south as suggested by Morafka (1976; see also Jackman, 1993)."

Nancy L Staub and David B. Wake. Amphibian Declines: The Conservation Status of United States Species. University of California Press, June 2005.

The Arboreal Salamanders on Southeast Farallon Island were recognized previously as the subspecies A. l. farallonensis - Farallon Salamander (Morafka, 1976), because they are more heavily spotted than mainland salamanders (except for those found in the Gabilan Mountains of Monterey County.) They have since been found to be similar genetically to the nearest mainland populations.

Alternate and Previous Names (Synonyms)

Aneides lugubris - Arboreal Salamander (Stebbins 1966, 1985, 2003, 2012)
Aneides lugubris lugubris - Arboreal Salamander (Storer 1925, Bishop 1943, Stebbins 1954)
Aneides lugubris farallonensis - Farallon Salamander (Farallon Yellow-spotted Salamander) (Bishop 1943)
Aneides lugubris farallonensis - Farallon Salamander (Storer 1925, Stebbins 1954)
Aneides lugubris lugubris - Arboreal Salamander -
California Land Salamander, Sad-colored Anaides, Spotted Autoax, Mournful Salamander, Speckled Salamander (Grinnell and Camp 1917)
Aneides lubugris farallonensis - Farallon Salamander (Grinnell and Camp 1917)
Aneides lugubris (Girard 1858)
Ambystoma punctulatum (Gray 1850)
Salamandra lugubris (Hallowell 1849)
Triton tereticauda (Eschscholtz 1833)

Conservation Issues (Conservation Status)

Because of its reliance on oak trees for nesting and aestivation, the continuing decline of live oaks and the alteration of oak woodlands may have a negative effect on this species.

Taxonomy
Family	Plethodontidae	Lungless Salamanders	Gray, 1850
Genus	Aneides	Climbing Salamanders	Baird, 1851
Species	lugubris	Arboreal Salamander	(Hallowell, 1849)
Original Description
Hallowell, 1849 - Proc. Acad. Nat. Sci. Philadelphia, Vol. 4, p. 126 from Original Description Citations for the Reptiles and Amphibians of North America © Ellin Beltz
Meaning of the Scientific Name
Aneides: Greek - lacking form or shape lugubris: Latin - gloomy, dark, possibly referring to dull coloration. from Scientific and Common Names of the Reptiles and Amphibians of North America - Explained © Ellin Beltz
Related or Similar California Salamanders
Aneides ferreus - Clouded Salamander Aneides flavipunctatus - Speckled Black Salamander Aneides iecanus - Shasta Black Salamander Aneides klamathensis - Klamath Black Salamander Aneides niger - Santa Cruz Black Salamander Aneides vagrans - Wandering Salamander
More Information and References
California Department of Fish and Wildlife AmphibiaWeb 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. 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. 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. Storer, Tracy I. A Synopsis of the Amphibia of California. University of Califonia Publications in Zoology Volume 27, The University of California Press, 1925. Jones, Lawrence L. C. , William P. Leonard, Deanna H. Olson, editors. Amphibians of the Pacific Northwest. Seattle Audubon Society, 2005. Stebbins, Robert C. and Nathan W. Cohen. A Natural History of Amphibians. Princeton University Press, 1997. David J. Morafka. Biogeographical Implications of Pattern Variation in the Salamander Aneides lugubris. Copeia, Vol. 1976, No. 3 (Aug. 20, 1976), pp. 580-586. Robert Cyril Stebbins. Amphibians of Western North America, University of California Press, 1962. Joseph Grinnell and Charles Lewis Camp. A Distributional List of the Amphibians and Reptiles of California. University of California Publications in Zoology Vol. 17, No. 10, pp. 127-208. July 11, 1917.

Conservation Status
The following conservation status listings for this animal are taken from the January 2024 State of California Special Animals List and the January 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. This salamander is not included on the Special Animals List, meaning there are no significant conservation concerns for it in California according to the California Department of Fish and Game.
Organization	Status Listing	Notes
NatureServe Global Ranking
NatureServe State Ranking
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
IUCN

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