Black Salamander, Speckled Black Salamander
|Taxonomic Notes: Rissler and Apodaca (2007, Syst Biol 56: 924-942) present arguments that the species as recognized by AmphibiaWeb should be broken into four species, with A. iecanus and A. niger recognized as full species, and a third that lacks a name in addition to A. flavipunctatus. The complex has been studied in depth by Reilly and colleagues (most recently Reilly and Wake, 2015, J Biogeog 42: 280-291), who find considerably more complexity than previously noted. No formal taxonomic changes have been made by primary researchers, but nonetheless taxonomic changes have been made by various compilers and book writers. The complex remains under study and taxonomic conclusions will be forthcoming.|
© 2012 Rob Schell (1 of 86)
Aneides flavipunctatus (Strauch, 1870)
Nancy L. Staub1
1. Historical versus Current Distribution. Black salamanders (Aneides flavipunctatus) occur in western lowland forests and meadows in northern California and extreme southern Oregon where annual precipitation is > 75 cm (Lynch, 1974). Populations generally are found at elevations below 600 m, but occur as high as 1,700 m (Lynch, 1974, 1981; Nussbaum et al., 1983). The distribution of black salamanders is disjunct; the southernmost populations (Santa Cruz Mountains) are separated from more northern populations by a gap that includes the northern part of the San Francisco Peninsula, the Marin Peninsula, and the nearly treeless area in southern Sonoma County, California. Populations south of Mt. Shasta and east of the Trinity Mountains appear to be separated from populations to the west (Larson, 1980; Lynch, 1981), although this may represent a collecting artifact. Analysis of protein variation among populations indicates a high level of genetic subdivision (Larson, 1980). Populations have been isolated from one another since the late Pliocene. Northern populations are paedomorphic in color pattern—adults retain the typical juvenile green-gray color pattern. Interestingly, in this part of the range, the salamander’s coloration matches the greenish-gray talus substrate. In other parts of the range, animals are found on dark soil (Larson, 1980; Lynch, 1981).
2. Historical versus Current Abundance. Black salamanders were once considered common in many areas of their range but have become rare in recent years (D.B. Wake, in Petranka, 1998). The proliferation of vineyards in northern California has destroyed much of the black salamander’s prime habitat (N.L.S. personal observation).
3. Life History Features.
A. Breeding. Reproduction is terrestrial.
i. Breeding migrations. Breeding migrations do not occur.
ii. Breeding habitat. There are no published data available on the courtship and breeding behavior of black salamanders (Lynch, 1974).
i. Egg deposition sites. Females probably lay eggs in July or early August in cavities below ground. Clutches have been found underground at depths of 23 and 38 cm (Van Denburgh, 1895; Storer, 1925). Eggs are attached by peduncles to moist earth. In the lab, eggs have been attached to the underside of cover objects (broken clay flowerpot pieces; N.L.S., personal observation).
ii. Clutch size. Van Denburgh (1895) described a partial clutch of 15 eggs (about twice as many eggs composed the original clutch in the field) found next to a barn in soil with numerous spaces and pieces of rotten wood. Ovarian complements range from 8–25 (mean about 12; Van Denburgh 1895; Stebbins, 1951).
C. Direct Development.
i. Brood sites. The same as egg deposition sites (underground and, in the lab, to the underside of cover objects).
ii. Parental care. Typically an egg clutch is found with a female in attendance. In the laboratory, females stayed with their clutches until eggs hatched (N.L.S., personal observation).
D. Juvenile Habitat. Juveniles are found in the same microhabitats as adults (e.g., under rocks and logs; Myers, 1930b).
E. Adult Habitat. Black salamanders occur in areas that receive > 75 cm annual precipitation (Lynch, 1974). Specific habitats include lowland forests, under rocks and logs or in wet soil along streams, under logs and rocks in grassy meadows, pastures, and burned areas, and in talus slopes (Wood, 1936; Myers and Maslin, 1948; Stebbins, 1951; Lynch, 1974, 1981; Staub, 1993). The populations in the Santa Cruz Mountains appear to prefer moister microhabitats than more northern populations. Unlike their more arboreal congeners, black salamanders are primarily ground dwellers (Myers and Maslin, 1948). Despite their ground-dwelling habits, black salamanders have a prehensile tail (Van Denburgh, 1895).
F. Home Range Size. Unknown.
G. Territories. In captivity, adults often bite one another (e.g., Myers, 1930b), and adult males and females show agonistic behavior toward conspecific intruders (Staub, 1993). Animals captured in the field are frequently scarred; males show a higher frequency of scarring than do females (Staub, 1993). This species may be territorial in the field.
H. Aestivation/Avoiding Dessication. In southern populations that are associated with streamside habitats, black salamanders are active year-round. In habitats that are not associated with permanent water, salamanders move underground during the dry season (mid April to mid October; Lynch, 1974).
I. Seasonal Migrations. Not known to occur.
J. Torpor (Hibernation). Not known to occur.
K. Interspecific Associations/Exclusions. Black salamanders occur syntopically with clouded salamanders (Aneides ferreus), wandering salamanders (Aneides vagrans), arboreal salamanders (Aneides lugubris), ensatinas (Ensatina eschscholtzii), and California slender salamanders (Batrachoseps attenuatus; Wood, 1936; Myers and Maslin, 1948; Lynch, 1974, 1985). Black salamanders also occur extensively with Plethodon elongatus in the Klamath River Valley and Trinity River drainage.
L. Age/Size at Reproductive Maturity. Reproductively mature black salamanders range in size from 60–75 mm (Lynch, 1974).
M. Longevity. In the lab, black salamanders have lived 15 yr (N.L.S., personal observation).
N. Feeding Behavior. Juveniles and adults feed on a wide variety of prey. The diet of adult salamanders consists primarily of diplopods (millipedes), coleopterans, formicans (primarily ants), and isopterans (primarily termites; Lynch, 1985). The diet of juveniles includes these prey as well as dipterans and collembolans (Lynch, 1985). Larger individuals consume larger prey items; mean and maximum prey size is correlated with body size. This correlation suggests that larger animals are selecting larger prey items and are ignoring smaller prey items. The number of prey items decreases as body size increases (Lynch, 1985).
O. Predators. Predators include western terrestrial garter snakes (Thamnophis elegans; Lynch, 1981).
P. Anti-Predator Mechanisms. When startled, juveniles generally remain immobile and adults flee (Van Denburgh, 1895; Jones, 1984). Other escape or defense behaviors include jumping (Van Denburgh, 1895), the production of sticky skin secretions (Lynch, 1981), an agonistic posture, and agonistic behaviors including biting (Lynch, 1981; Staub, 1993). The agonistic posture of black salamanders is distinctive. The animal raises its body off the substrate with the legs fully extended, the back is arched, the head elevated with the snout pointed slightly downward, and the tail undulates (Jones, 1984; Staub, 1993; Stebbins, 1954a). In the laboratory, black salamanders will bite western garter snakes, which can result in serious injuries to the snakes (Lynch, 1981).
R. Parasites. Nematodes have been found in the black salamander (Lehmann, 1954; Schad, 1960).
4. Conservation. Black salamanders have become rare in recent years due in large part to the proliferation of vineyards in northern California that has destroyed much of their prime habitat. According to Levell (1997), black salamanders are listed as Protected in Oregon.
1 Nancy L. Staub
2 David B. Wake
Literature references for Amphibian Declines: The Conservation Status of United States Species, edited by Michael Lannoo, are here.
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