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A Guide to the Amphibians
and Reptiles of California


Arboreal Salamander - Aneides lugubris

(Hallowell, 1849)
Click on a picture for a larger view



Arboreal Salamander Range Map
Range in California: Red

Dot-locality Range Map.



observation link





Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
Adult, Contra Costa County Adult, Alameda County Adult, Santa Cruz County
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
Adult, Contra Costa County (missing the tip of its tail.) Adult, Mendocino County
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
  Adult, Contra Costa County   Adult, found on a building in downtown San Francisco, San Francisco County
© Jackson Shedd
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
Adult male, Monterey County coastal sand dunes Adlt male, Monterey County coastal sand dunes
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
Adult active at night in situ, Marin County Adult active at night in situ, Marin County Adult active at night in situ, Marin County Adult active at night in situ, Marin County
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
Adult emerging at night from a rock crevice, Marin County Adult male with large head scar, Marin County Adult, Orange County
© Jason Jones
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
Adult, Alameda County Adult, San Luis Obispo county,
© Andrew Harmer
Adult, Southeast Farallon Island.
© Oscar Johnson
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
Two adults found in the Santa Ana Mountains, Orange County © Cooper Bailey Adult, San Luis Obispo County
© Henk Wallays
Adult, Sierra Nevada Mountains, Tuolumne County © Rick Staub
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
Adult, Los Angeles County
© Gregory Litiatco
Adult, San Diego County © Stuart Young This adult was rescued from the bottom of the deep end of a swimming pool in San Luis Obispo County. © Laurie Dey This adult in Santa Clara County has a mottled head. © Katherine Hughes
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
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
Albino or pigmentless adult, from Contra Costa County, found with the normal adult shown above the albino in the left picture. © 2006 John Schilling
Arboreal Salamander Arboreal Salamander foot  
Heavily-spotted 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.
Arboreal Salamanders have toes
with tips that help them climb vertical surfaces like tree trunks and cliffs.
   
       
Aestivation
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.

Arboreal Salamanders Arboreal Salamanders    
Eric Boyer discovered this large group of estivating adult and juvenile Arboreal salamanders underneath some wood while remodeling a backyard in Santa Barbara County in June.
© Eric Boyer
This salamander was found underneath the circular base of a large statue in a Santa Clara County backyard in August.
© Laurie Weber
   
       
Juveniles
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
Juvenile, Marin County Juvenile, Santa Cruz County Juvenile, Santa Cruz County
© Brad Alexander
Hatchling, Marin County
       
Predation
San Bernardino Ring-necked snake eating an adult Arboreal Salamander San Bernardino Ring-necked snake eating an adult Arboreal Salamander    
An adult San Bernardino Ring-necked snake eating an adult Arboreal Salamander
in Los Angeles County © Jonathan Benson
   
     
Habitat
Arboreal Salamander Habitat Arboreal Salamander Habitat Arboreal Salamander Habitat Arboreal Salamander Habitat
Typical oak woodland habitat, Contra Costa County Habitat, Contra Costa County Arboreal salamanders inhabit this urban backyard in Oakland, Alameda County
Arboreal Salamander Habitat Arboreal Salamander Habitat Arboreal Salamander Habitat Arboreal Salamander Habitat
Coastal sand dunes habitat,
Monterey County
Rocky habitat, Marin County Habitat at coastal marsh,
San Mateo County
Arboreal salamanders inhabit barren, rocky Southeast Farallon Island.
Arboreal Salamander Habitat Arboreal Salamander Habitat    
Arboreal salamanders inhabit barren, rocky Año Nuevo Island A careful look underneath the fallen bark of this dead tree turned up one Arboreal Salamander, two Coast Range Newts, one Yellow-eyed Ensatina, and 12 California Slender Salamanders, illustrating how dead wood and bark on a forest floor is an important microhabitat for salamanders and other wildlife.    
       
Short Video and Audio
Arboreal Salamander Arboreal Salamander Arboreal Salamander Arboreal Salamander
speaker icon speaker icon 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 are two high-pitched, bird-like squeaks with some rustling background noise.) 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 prehensile, often coiled.
Usually 15 costal grooves.
Two nasolabial grooves.
Color and Pattern
Color is brown above with small cream to yellow spots.
The number of spots varies with location - Sierra Nevada populations have weak spotting, Gabilan Mountains and Farallon Islands populations have large conspicuous spots.
The venter is creamy white.
The undersides of the tail and feet are dull yellow.
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 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
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 tail.
Has been found up to 59 ft. (18 m) above the ground.
Territoriality
Both males and females are agressively territorial. Scarred individuals are often found, and captives kept together often bite each others' tails.
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.)
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, 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. 
Breeding
Reproduction is terrestrial.
Adults probably reach sexual maturity in their third year.
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.
Multiple salamanders sometimes lay their eggs 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.

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.
Full Species Range Map
Elevational Range
From sea level to 5,000 ft. (1,500 m).

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.

Notes on Taxonomy
One species of A. lugubris is recognized, but several DNA studies have shown that there are genetically different groups within the species.


The most recent research I am aware of is that of Sean B. Reilly et al ** in a study of the phylogeographic history of Aneides lugubris published in 2015. They 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.

Arboreal Salamander Range Map
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 (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, 1849
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
Speckled Black Salamander
Santa Cruz Black Salamander
Wandering Salamander
Clouded 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.

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.

1976 Morafka, David J. and Benjamin H. Banta. Biogeographical implications of pattern variation in the Salamander Aneides lugubris. Copeia 1976(3):580-586.

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, meaning there are no significant conservation concerns for it in California according to the California Department of Fish and Game.


Organization
Status Listing
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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