Aneides flavipunctatus (Strauch 1870)

Projecting upper-jaw teeth and triangular head distinguish the Black Salamander as a climbing salamander of the genus Aneides (Dunn 1926). Aneides flavipunctatus has a characteristic black or slaty ventral coloration. Dorsal coloration varies greatly depending upon locality. The dorsum may be uniformly black, or black with very small white flecks in the extreme southern part of its range; black with large white spots in the interior Coast Range (from Alder Springs, Glenn Co., and Lucerne, Lake Co., CA south); black with pale yellow or whitish spots in the outer Coast Range (from Sonoma Co. to middle Mendocino Co.); black frosted with gray, olive, or green, with few or no light spots, in the Redwood country of Mendocino Co. and Humboldt Co.; and black with many small white spots from the Klamath Mts. to Mt. Shasta (Stebbins 1985).See other subspecies accounts at www.californiaherps.com: A. f. flavipunctatus (http://www.californiaherps.com/salamanders/pages/a.f.flavipunctatus.html) and A. f. niger (http://www.californiaherps.com/salamanders/pages/a.f.niger.html).

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 are generally found at elevations below 600 m, but occur as high as 1700 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 (Larson 1980) and mitochondrial sequence variation (Rissler and Apodaca 2007) among populations indicates a high level of genetic subdivision, with climatic factors apparently affecting the distributions of cryptic sister lineages (Rissler and Apodaca 2007). 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-grey talus substrate. In other parts of the range, animals are found on dark soil (Larson 1980; Lynch 1981). Habitat Requirements. 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 meadow 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).

This is a fully terrestrial salamander. Reproductively mature black salamanders range in size from 60 - 75 mm (Lynch 1974). 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. Staub, personal observation). 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). Typically an egg clutch is found with a female in attendance. In the laboratory, females stayed with their clutches until eggs hatched (N. L. Staub, personal observation). 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 (personal observation). Behavior. In captivity, adults often bite one another (e.g., Myers 1930) and adult males and females show agonistic behavior toward 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. Aestivation. In southern populations that are associated with streamside habitats, black salamanders are active year round. In habitats which are not associated with permanent water, salamanders move underground during the dry season (mid-April – mid-October) (Lynch 1974). Interspecific Associations/Exclusions. Black salamanders occur syntopically with clouded salamanders (Aneides ferreus), wandering salamanders (Aneides vagrans), arboreal salamanders (Aneides lugubris), ensatinas (Ensatina eschscholtzi) and California slender salamanders (Batrachoseps attenuatus) (Lynch 1974, 1985; Myers and Maslin 1948; Wood 1936). Black salamanders also occur extensively with Plethodon elongatus in the Klamath River Valley and Trinity River drainage. Feeding Behavior. Juveniles and adults feed on a wide variety of prey. The diet of adult salamanders consists primarily of diplopoda (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). Predators. Predators include western terrestrial garter snakes (Thamnophis elegans; Lynch 1981). 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 the 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 1954). In the laboratory, A. flavipunctatus will bite western garter snakes which can result in serious injuries (Lynch 1981). Parasites. Nematodes have been found in the black salamander (Lehmann 1954; Schad 1960). Life Expectancy: Staub (2016) reviews evidence of life expectancy for salamanders of the family Plethodontidae, and concludes that these generally small animals can attain relatively great age. The most direct evidence is from captive animals, and members of Plethodon, Aneides and Phaeognathus, among others, have attained ages in excess of 30 years (the individual in the photograph was collected as an adult in April 1985, and is still apparently in robust health). In a separate field study (Staub 2016), Aneides flavipunctatus longevity was estimated to be on the order of 18 or 25 years, using different methods. Other field estimates also show long lives, as great as 32 years. Methods tend to be conservative, and some individuals might be substantially older.

Little information exists concerning population trends, but those who are familiar with this salamander suggest that they have become increasingly harder to find (D. Wake, pers. comm).