Description
Slim, long-legged,and agile: an excellent climber. Reaches about 65 mm snout-vent length and about 110 mm total length. Tips of toes of adpressed limbs separated by no more than 1.5 costal folds and sometimes overlapping by as much as 1.5 folds. Usually 16 costal grooves. The toes have slightly broadened and squarish tips. Brown above, clouded with ash, greenish gray, pale gold, or reddish; dusky below. In the dark phase these salamanders may be almost solid brown above. In the light phase, pale gray color may predominate and the brown color may be reduced to a network. Some adults in Oregon and extrme northwest California are almost uniformly dark brown above with a few cream-colored spots. Hatchlings have a copper or brassy dorsal stripe that soon becomes reduced to patches on the snout, shoulders, upper surface of limbs, and tail (Stebbins 1985). As juveniles become older their color pattern become darker and duller (McKenzie and Storm 1971). Male has heart-shaped mental gland, absent in female (Stebbins 1985).

Distribution and Habitat

Country distribution from AmphibiaWeb's database: United States

U.S. state distribution from AmphibiaWeb's database: California, Oregon

View distribution map in BerkeleyMapper.

Clouded salamanders (Aneides ferreus) are found in the coastal forests of Oregon and northern California. Their range extends from the Columbia River south through the Siskiyou and Coast mountains and western Cascades of Oregon (but they are not found in the extreme northwestern part of the Coast Mountains) and northwestern California. In California their range extends to near the junctions of Hurdygurdy Creek and Goose Creek with the South Fork of the Smith River near the coast, and north of the junction of the Salmon and Klamath rivers further inland (Leonard et al., 1993; Wake and Jackman, 1999(1998)). The current distribution of the clouded salamander is much more restricted than previously thought because all British Columbia specimens and most specimens from California have been assigned to the new species Aneides vagrans (wandering salamander) (Wake and Jackman, 1999(1998)).

Clouded salamanders occur in high densities in old growth forests (Corn and Bury, 1991) in recently cut or burned areas, in association with stumps, decaying logs, and coarse, woody debris (van Denburgh, 1916; Fitch, 1936; McKenzie and Storm, 1970; Whitaker et al., 1986; Bury and Corn, 1988; Welsh and Lind, 1988, 1991). Welsh and Lind (1988) found that mesic forest stands had significantly higher capture rates of A. ferreus than did drier stands. This species tends not to be found in dense forests (Wood, 1939). Individuals have been found up to 1525m elevation (Beatty, 1979).

Populations have certainly been lost due to forestry management practices and to urban sprawl.

Abundance. Clouded salamanders are typical inhabitants of old growth forests (Corn and Bury, 1991), especially in edge habitats (e.g., forest clearings) with downed or standing decaying trees and stumps. Habitat Requirements: Juvenile. Juvenile clouded salamanders prefer bark litter over rock or leaf litter (McKenzie and Storm, 1970). Subadults select bark litter at higher temperatures (20C, 25C), and show no preference between rock or bark litter at lower temperatures (10C, McKenzie and Storm, 1970).

Habitat Requirements: Adult. Adults are found associated with stumps and logs in Douglas fir forests, in crevices in rock outcrops and road cuts, and in talus (Van Denburgh, 1916; Wood, 1939; Lowe, 1950; Maser and Trappe, 1984; Nussbaum, 1983; Leonard et al., 1993; Jackman, 1999(1998)). Animals are most commonly found in decaying logs and stumps with intact bark–either under the bark, within the log, or under the log (Bury and Corn, 1988; Corn and Bury, 1991). Habitats with such logs are often associated with forest edges (Myers and Maslin, 1948; Stebbins, 1951). For example, McKenzie and Storm (1970) found clouded salamanders in stumps and fire-charred logs of Douglas fir associated with forest clearings and road banks. The preferred type of wood is Douglas fir; individuals are less commonly found in Port Orford cedar, alder, and redwood (Myers and Maslin, 1948; Wake, 1965). Individuals have been found up to 6.1m above the ground in dead stumps (Van Denburgh, 1916; Slevin, 1928). Based on recent work documenting the presence of the wandering salamander in the forest canopy, it is probable that the clouded salamander occupies similar habitat in its range (Jim C. Spickler, personal communication). In a microhabitat selection experiment, adults consistently selected against leaf litter and showed equal preference for bark litter and rock (McKenzie and Storm, 1970). Jackman, (1999(1998)) suggest that there may be a difference in habitat preference between clouded and wandering salamanders–-in northwestern California clouded salamanders are commonly found associated with both decaying logs and rocky slopes, while wandering salamanders are almost exclusively found associated with decaying logs, not rocks.

Life History, Abundance, Activity, and Special Behaviors
Age/Size at Reproductive Maturity. Data from McKenzie (1970) suggest that males mature during their second year, at lengths greater than 36 mm SVL, and that females first reproduce in their third year, when they are approximately 55 mm SVL.

Females lay eggs in late June and July and hatchlings emerge in late August or September (Petranka, 1998). Egg clutches have been found in rotting logs (Storm, 1947). Reported clutch sizes vary from 9-17 eggs (Storm, 1947). An ovarian count reported 12 eggs (Fitch, 1936). If the reproductive biology of the clouded salamander is similar to that of the wandering salamander, brood sites occur in the forest canopy as well as in decaying logs (Welsh and Wilson, 1995). McKenzie (1970) found little seasonal variation in nale reproductive characteristics. Sperm can be found in vasa deferentia in every month except September.

Clutches have been found without a parent in attendance (A. ferreus, Storm, 1947), with a female in attendance (A. vagrans, Dunn, 1942), and with both a male and a female in attendance (A. ferreus, Storm, 1947). Behavior. In contrast to the wandering salamander, the clouded salamander appears to be relatively aggressive. Thirty-two percent of specimens from an Oregon population had scars, presumably from conspecific attacks, and males had a higher percentage of scars than did females (Staub, 1993). Studies on A. vagrans indirectly suggest that this species may not be as aggressive as is A. ferreus—A. vagrans does not use the chemical signals of fecal pellets to delimit territory boundaries, as other plethodontids tend to do (Ovaska and Davis, 1992).

Interspecific Associations/Exclusions. Clouded and wandering salamanders overlap in a zone < 15 km wide in northwestern California. Jackman, (1999(1998)) report slight evidence of introgression, but clear hybrid individuals have never been identified. The range of the clouded salamander also overlaps with its congener, the black salamander (A. flavipunctatus) (Fitch, 1936; Myers and Maslin, 1948; Stebbins, 1985), especially in the Klamath River valley, but the black salamander occurs in hotter, drier regions (for example, further up the Klamath River valley). Feeding Behavior. As with most salamanders, clouded salamanders are generalist feeders. Adults primarily eat isopods (sowbugs), hymenopterans (ants), and coleopterans but their diet also includes an important assortment of other insects (e.g., dipterans, isopterans (termites)), and mites, spiders, pseudoscorpions, centipedes, and millipedes (Fitch, 1936; Storm and Aller, 1947; Bury and Martin, 1973; Whitaker et al., 1986). Whitaker et al. (1986) found no significant differences between the diets of adult males and females. Hatchlings (< 20 mm SVL) eat small prey, primarily mites, springtails, flies, and small beetles (Whitaker, et al., 1986). As juveniles get larger, they switch to eating larger prey items such as sowbugs, larger beetles, and earwigs (Whitaker et al., 1986). Adults occasionally consume shed skin (Whitaker et al., 1986).

Predators. Poorly documented, but Petranka (1998) suggests predators of Aneides sp. include mammals, woodland birds, and snakes.

Anti-Predator Mechanisms. Several anti-predatory behaviors have been observed in the clouded or wandering salamander when individuals are startled or attacked: crawling away rapidly, immobility, a defensive posture (raising the body and undulating the tail), and flipping around followed by immobility (Fitch, 1936; Brodie, 1977). Skin secretions are thought to be noxious (Brodie, 1977).

Parasites. One species of nematode has been identified in the clouded salamander (Lehman, 1954; Goldberg et al., 1998).

Trends and Threats
Populations associated with post-logged areas eventually decline, and Corn and Bury (1991) doubt that this species can survive in areas where forests are intensively managed on short rotation cycles because of severe reduction in moisture conditions and in the amount of large woody debris. The amount of coarse woody debris retained after timber harvesting under current forest management guidelines probably does not provide adequate habitat for the clouded salamander (Butts and McComb, 2000).

Possible reasons for amphibian decline

General habitat alteration and loss
Habitat modification from deforestation, or logging related activities
Secondary succession

Comments
Mitochondrial, allozyme, and karyotypic data distinguish A. ferreus from A. vagrans (Sessions and Kezer, 1987; Jackman, 1999(1998). Morphologically, these two species are similar (Beatty, 1979).

The species epithet, "ferreus," is Latin and means “iron colored,” and is presumably a reference to the species' metallic dorsal coloration (Tighe 2023).

See another account at californiaherps.com.

References

Brodie, E. D., Jr. (1977). "Salamander antipredator postures." Copeia, 1977, 523-535.

Corn, P. S. and Bury, R. B. (1991). ''Terrestrial amphibian communities in the Oregon Coast Range.'' Wildlife and Vegetation of Unmanaged Douglas-fir Forests. L. F. Ruggiero, K. B. Aubry, A. B. Carey, and M. H. Huff, eds., USDA Forest Service General Technical Report PNW-GTR-285, 304-317.

Tighe, K.A. (2023). Catalog of type specimens of recent Caudata and Gymnophiona in the National Museum of Natural History, Smithsonian Institution. Smithsonian Contributions to Zoology 654.

Species Account Citation: AmphibiaWeb 2023 Aneides ferreus: Clouded Salamander <https://amphibiaweb.org/species/3935> University of California, Berkeley, CA, USA. Accessed Nov 25, 2024.

Citation: AmphibiaWeb. 2024. <https://amphibiaweb.org> University of California, Berkeley, CA, USA. Accessed 25 Nov 2024.

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