Ambystoma annulatum Cope, 1886
Stanley E. Trauth
1. Historical versus Current Distribution. Ringed salamanders (Ambystoma annulatum) are endemic to the Interior Highlands (Ozark and Ouachita mountains) in Arkansas, Missouri, and Oklahoma (Anderson, 1950; Smith, 1950; Dowling, 1956, 1957; Anderson, 1965; McDaniel, 1975; Funk, 1979; Minter, 1979; Schuette, 1980; Trauth, 1980; Johnson, 1987; Turnipseed and Altig, 1991). No specimens have been reported from the Ozark Plateau of Kansas (Collins, 1993). There are no data to suggest that the current distribution differs from the pre-settlement distribution. Phillips et al. (2000) note that populations in the northeastern portion of the range (central Missouri) have less variable mitochondrial DNA components than populations to the southwest (southern Missouri, northwestern Arkansas, and eastern Oklahoma). This suggests that northern populations have been established more recently, perhaps with reforestation following the warm and dry Hypsithermal Interval (8,000–4,000 yr before present; Phillips et al., 2000).
2. Historical versus Current Abundance. Unknown. Published reports range from few to large numbers of animals. For example, while several early published records (prior to 1924) indicated few (< 10) specimens (Black and Dellinger, 1938, in Arkansas), large populations (unspecified numbers) have been mentioned (Noble and Marshall, 1929, in Missouri; Trapp, 1956; Brussock and Brown, 1982, in Arkansas). The first record reported in Oklahoma was a single specimen (Firschein and Miller, 1951). Spotila and Beumer (1970) observed 46 salamanders crossing a highway on 18 October 1966 near Fayetteville (Washington County), Arkansas, and observed 155 adults over a 3-yr period (1965–'67). Several hundred ringed salamanders were counted by McDaniel and Saugey (1977) crossing a highway on 18 October 1966 near Fayetteville (Washington County), Arkansas, and 155 adults were observed over a 3-yr period (1965–'67). Several hundred ringed salamanders were counted by McDaniel and Saugey (1977) crossing a highway on 22 October 1976 near Blanchard Springs Caverns (Stone County, Arkansas). During a salamander sting operation conducted by the Arkansas Game and Fish Commission (Arkansas Democrat, 22 October 1987), a poacher was arrested while attempting to sell about 1,060 specimens for fish bait in Hot Springs, Arkansas. Peterson et al. (1991) estimated the number of females ovipositing in each of two ponds (Stone County, Missouri) to be between 150–230 individuals, whereas Peterson et al. (1992) captured 230 males and 237 females in one of these ponds using pit fall traps. In a recent study, Briggler et al. (1999) found 1,096 specimens migrating across a highway to a single pond in northwestern Arkansas from 22 September–14 November 1998. Trauth (2000) found 17 specimens in a woodland pond in the Ozark National Forest (Stone County, Arkansas) on 26 February 2000.
3. Life History Features.
A. Breeding. Reproduction is aquatic.
i. Breeding migrations. Ringed salamanders typically breed during autumn, primarily between September and early November (Noble and Marshall, 1929; Trapp, 1956; Anderson, 1965). Trauth et al. (1989c) observed the first incidence of winter breeding; apparently, winter breeding may be a common activity in ringed salamanders in the Ozark National Forest of north-central Arkansas (Trauth, 2000). Adults are stimulated to migrate by medium to heavy rains, cool temperatures, and nighttime conditions. They travel to fishless, woodland ponds to breed; however, farm ponds (heavily used by livestock) in open pastures may also be utilized (Brussock and Brown, 1982). Hundreds of adults may be present during a single breeding episode, which normally lasts several days. Males apparently arrive before females (Spotila and Beumer, 1970). Each gravid female may be courted by 2–25 males. Males may also deposit spermatophores with or without attendant females (Spotila, 1976). Egg laying begins shortly after courtship and can last for 2 d (Johnson, 1987; Conant and Collins, 1998).
ii. Breeding habitat. Fishless woodland ponds as well as livestock ponds are suitable breeding sites for ringed salamanders (Brussock and Brown, 1982; Johnson, 1987).
i. Egg deposition sites. Eggs are laid on submerged branches, aquatic plant stems, or on the pond bottom in loose masses (Anderson, 1965; Spotila and Beumer, 1970; Johnson, 1987). An observation of terrestrial egg laying in March as reported by Strecker (1908a) has been discounted as being erroneous (see Trauth et al., 1989c; Peterson et al., 1992). Each female either lays one or two large clusters of eggs with 75–150 eggs/cluster (Johnson, 1987) or lays several smaller clusters of from 4–31 or 2–17 eggs (Trapp, 1956).
ii. Clutch size. Average clutch size is estimated to range from 205–390 eggs/female (Trauth, 2000; Peterson et al., 1992, respectively). Eggs hatch in 2–3 wk, depending on water temperature (Johnson, 1987).
C. Larvae/Metamorphosis. Larvae hatch at 12–15 mm SVL. Coloration in newly metamorphosed individuals is olive to black dorsally and grayish white ventrally (Hutcherson et al., 1989) with a fairly broad pigment-free band on the sides of the trunk (Bishop, 1943).
i. Length of larval stage. Larvae remain in the ponds throughout the winter and metamorphose the following summer. The length of the larval period varies from ~ 6–8.5 mo (Hutcherson et al., 1989).
ii. Larval requirements.
a. Food. Small larvae feed on cladocerans, copepods, and dipteran larvae, whereas older larvae primarily consume dipteran larvae; additional prey items included ostracods, hemipterans, snails, and dragonfly and damselfly naiads (Hutcherson et al., 1989). Trapp (1959) mentions cladocerans and Chironomus sp. (Diptera) as the major components of their diet. Cannibalistic larvae are facultative and will consume the same array of invertebrate food items as non-cannibals.
b. Cover. Little information is available on the terrestrial ecology of juveniles and adults (Petranka, 1998).
iii. Larval polymorphisms. Cannibalistic larvae were reported in a Missouri pond (Nyman et al., 1993). Other than having a relatively larger body size and somewhat broader heads, most cannibals were not different in appearance from non-cannibals. Cannibals exhibited mean body lengths twice that of their conspecific prey. Nyman et al. (1993) further pointed out that cannibalism in ringed salamander larvae appears to be an opportunistic behavior, an activity related to larger body size in older cohorts that tend to metamorphose at a larger size and at an earlier date.
iv. Features of metamorphosis. Larvae average 16 mm SVL 1 mo after hatching; average length at metamorphosis varies from 34–40 mm SVL and normally occurs from late April to early June in Missouri (Hutcherson et al., 1989; Nyman et al., 1993). Large ringed salamander larvae have been observed in September in the Ozark National Forest in Arkansas (unpublished data).
v. Post‑metamorphic migrations. None observed.
vi. Neoteny. Has not been observed.
D. Juvenile Habitat. Juvenile habitat characteristics are unknown, although one juvenile reported by Trauth (1980; unpublished data) was collected on a forested hillside beneath a small flat rock.
E. Adult Habitat. Ringed salamanders are found in forested areas. Noble and Marshall (1929) found adults beneath piles of vines in a field as well as under leaves near a pond. They probably live hiding under logs and rocks or burrowing in the soil and are seldom found out or in the open (Johnson, 1987).
F. Home Range Size. Unknown.
G. Territories. Unknown.
H. Aestivation/Avoiding Dessication. Aestivation is unknown; adults likely seek shelter under cover objects or burrow when faced with dessicating conditions.
I. Seasonal Migrations. Medium to heavy autumn rains with cooler temperatures initiate breeding movements (Spotila and Beumer, 1970). The triggering mechanism for winter breeding remains unknown (Trauth et al., 1989c), although Trauth (2000) mentioned the lack of adequate rainfall during the normal breeding cycle (due to a summer/fall drought) as a possible stimulus.
J. Torpor (Hibernation). No reports on hibernating animals are available.
K. Interspecific Associations/Exclusions. Ecological associates observed by Trauth et al. (1989c) and Trauth (2000) at a winter breeding pond include spotted salamanders (Ambystoma maculatum), central newts (Notophthalmus viridescens louisianensis), wood frogs (Rana sylvatica), and spring peepers (Pseudacris crucifer).
Literature references for Amphibian Declines: The Conservation Status of United States Species, edited by Michael Lannoo, are here.
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