© 2014 Todd Pierson (1 of 12)
Country distribution from AmphibiaWeb's database: United States
Eurycea multiplicata (Cope, 1869)
Stanley E. Trauth1
1. Historical versus Current Distribution. Many-ribbed salamanders (Eurycea multiplicata) occur in the Ozark Plateaus and the Boston and Ouachita mountains and associated lowland rocky formations in southwestern Missouri, eastern Oklahoma, and northwestern Arkansas, at elevations of 107–763 m (Dundee, 1965a). Two subspecies are recognized: many-ribbed salamanders (E. m. multiplicata) occur in southeastern Oklahoma and west central Arkansas, while gray-bellied (or graybelly) salamanders (E. m. griseogaster) occur in northeastern Oklahoma, southwestern Missouri, and northwestern Arkansas. The type locality of E. m. multiplicata is in question (Dundee, 1950), but the type locality of E. m. griseogaster is clearly stated (Moore and Hughes, 1941). Animals are found from 107–763 m (350–2,500 ft) in elevation (Dundee, 1965a). Cope (1889) reported many-ribbed salamanders from Kansas, but this record apparently is not backed by a voucher specimen (Dundee, 1965). If Cope was correct, a range contraction for this species has occurred. Besides this mention, no evidence supports declines or shifts in distributions.
2. Historical versus Current Abundance. Generally unknown, but Dundee (1947) reports finding gray-bellied salamanders "in abundance" and "in large numbers" at certain sites and certain times of the year in Oklahoma, particularly in winter and early spring.
3. Life History Features.
A. Breeding. Reproduction is aquatic.
i. Breeding migrations. Long-distance migrations are unlikely.
ii. Breeding habitat. The mating season varies among populations of gray-bellied salamanders. Populations that inhabit thermally stable springs have a prolonged mating season compared with populations inhabiting surface streams with more variable temperatures. Based on the times when females contained spermatozoa in their reproductive tracts, mating activity could be from July–May (Ireland, 1976). Ireland (personal communication) agrees that he meant spermatheca for the term "reproductive tracts."
i. Egg deposition sites. Females lay their eggs in springs, spring-fed pools, and spring-fed ponds from autumn to early spring (Spotila and Ireland, 1970; Ireland, 1976). Eggs usually are laid on the undersides of submerged stones or beneath several layers of smaller stones (Trauth et al., 1990).
ii. Clutch size. Egg counts range from 2–21, but average 10–13 (Ireland, 1976; Trauth et al., 1990). Eggs range from 2.0–2.6 mm in diameter. Females do not brood. Hatchlings average about 10 mm SVL (Petranka, 1998), but Petranka does not document this size.
i. Length of larval stage. Larvae grow quickly, range in size from 23–85 mm TL, and metamorphose between 33–48 mm SVL, 5–8 mo after hatching.
ii. Larval requirements.
a. Food. Larvae feed most actively at night. Although larvae are described as benthic feeders and will eat isopods, ostracods, and zooplankton, aquatic insects usually are consumed (Rudolph, 1978; see also Petranka, 1998).
b. Cover. During daylight, many-ribbed salamanders are usually found under stones in slow-moving streams. Larvae will inhabit more ephemeral portions of streams than will adults (Loomis and Webb, 1951). Bragg (1955b) observed that small (< 18 mm) gray-bellied salamanders can be found in shallow water, in the open, both day and night. Larger larvae spend more time burrowed in gravel.
c. Temperature. Gray-bellied salamander larvae have been found in water as warm as 22 ˚C in summer (August) and as cold as 9.5 ˚C in winter (Dundee, 1958).
iii. Larval polymorphisms. Undescribed and unlikely.
iv. Features of metamorphosis. At metamorphosis, animals are approaching sexual maturity. Ireland (1976) found that 16% of males in the process of metamorphosing contained spermatozoa.
v. Post-metamorphic migrations. Bragg (1955b) observed that when gray-bellied salamanders begin to metamorphose, they leave the stream and complete their development along the streambank in moist leaves or under stones.
vi. Neoteny. At metamorphosis, animals are approaching sexual maturity. Gilled adults are common in gray-bellied salamander populations that inhabit caves or streams draining caves, especially on the Salem Plateau (Dundee, 1965; see also Dundee, 1947, for many-ribbed salamanders, and Petranka, 1998). Neotenic animals that inhabit caves often are pale and lack components of the normal pigment pattern, whereas neotenic animals in the vicinity of caves retain normal larval pigment patterns.
D. Juvenile Habitat. Similar to that of adults (cool, moist habitats near water under stones, logs, and other large materials in streams and springs, both in the open and in the twilight zone of caves; see "Adult Habitat" below), or if neotenic, similar to that of larvae (see "Neoteny" above).
E. Adult Habitat. Adults are essentially aquatic but occasionally are found on land. Many-ribbed salamanders are found in cool, moist habitats near water under stones, logs, and other large materials in streams and springs, both in the open and in the twilight zone of caves (Moore and Hughes, 1941; Dundee, 1947, 1958, 1965a; Loomis and Webb, 1951; Ireland, 1976; see also Petranka, 1998). Large numbers of gray-bellied salamanders were found under cover objects when the adjacent water had a thin film of ice after overnight subfreezing temperatures (H.A.D., field notes).
In the Ozark highlands, adults are found in caves (Dundee, 1965a). At a study site in Cherokee County, Oklahoma, gray-bellied salamanders were common where water flowed over shallow soils and exposed solid limestone in cool months, but which were dry during summer and early autumn (Dundee, 1958). Bragg (1955b) noted habitat differences between many-ribbed and gray-bellied salamanders: "Both [subspecies] occur in Cherokee County [Oklahoma] but apparently they tend to occupy different habitats. Wherever very small streams traverse solid limestone rock or soil, [many-ribbed salamanders are] very likely to be present. If a similar stream cuts through chert, flint, or granite, [gray-bellied salamanders are] the form to be expected, if either form occurs. My experience is not wide enough to say that there never are exceptions, but I can say that so far I have found none in Oklahoma." One exception is known, one of Dundee's (1958) study sites for gray-bellied salamanders was over solid limestone rock with grassy hummocks on it.
F. Home Range Size. Unknown but likely to be small. In the first collections of gray-bellied salamanders (Moore and Hughes, 1941), adults and larvae were collected together.
G. Territories. Unknown but unlikely. Loomis and Webb (1951) reported that frequently two or three, and up to four, many-ribbed salamander adults were found under a single rock. Much higher numbers of gray-bellied salamanders have been found under single cover objects (H.A.D., field notes).
H. Aestivation/Avoiding Dessication. Adult gray-bellied salamanders tend to be found in association with springs and permanent streams and within the wetter portions of such streams during drier periods (Loomis and Webb, 1951). Animals may aestivate, as suggested from at least one site in Oklahoma that would dry in the summer when no salamanders could be found (Dundee, 1958). Bragg (1955b) noted terrarium observations on many-ribbed salamanders that support the possibility of aestivation.
I. Seasonal Migrations. Individuals do not disperse far from their natal streams.
J. Torpor (Hibernation). Animals apparently stay active near the surface, being found under rocks, logs, and moss in or near the edges of streams, except during periods of extreme winter weather (Dundee, 1947).
K. Interspecific Associations/Exclusions. Many-ribbed salamanders are found in association with Ouachita dusky salamanders (Desmognathus brimleyorum) in Arkansas (Strecker, 1908a). Gray-bellied salamanders are found with grotto salamanders (Typhlotriton spelaeus) in Missouri (Noble, 1927b). Loomis and Webb (1951) noted that oftentimes stones that held many-ribbed salamanders provided a substrate for small- to medium-sized tarantulas (Aphonopelma hentzii), but tarantulas preferred the drier area under the stone.
Spring-fed headwaters in eastern Oklahoma often contain assemblages of 3–5 species of salamander larvae, including long-tailed salamanders (E. longicauda), cave salamanders (E. lucifuga), many-ribbed salamanders, Oklahoma salamanders (E. tynerensis), and grotto salamanders (Typhlotriton spelaeus) that segregate by distance from the spring head. This habitat segregation may be due to competitive factors (Rudolph, 1978; see also Petranka, 1998).
L. Age/Size at Reproductive Maturity. Reproductive maturity is reached at metamorphosis, or shortly thereafter (Ireland, 1976), although gilled adults are common in some populations (Dundee, 1947; see also Petranka, 1998). Ireland (1976) found that all metamorphosed males have spermatozoa in their seminiferous tubules; 16% of males in the process of metamorphosing also contained spermatozoa. Size varies across populations, with transformed animals averaging 28–45 mm SVL (Moore and Hughes, 1941; Loomis and Webb, 1951; Dundee, 1965). Males and females vary little in size, form, or color (Moore and Hughes, 1941). Little growth occurs after metamorphosis (Dundee, 1965a). Neotenic animals can exceed metamorphosed animals in size, reaching 54 mm SVL and 160 mm TL (Dundee, 1965a).
M. Longevity. Unknown.
N. Feeding Behavior. Undescribed, but adults likely feed on a variety of aquatic and semi-aquatic vertebrates associated with springs and permanent streams. The diet of neotenic adults likely resembles that of larvae (see "Larval requirements" above), although adults, being larger, may take larger prey. Terrestrial adult gray-bellied salamanders consume some aquatic arthropods but primarily eat terrestrial arthropods, snails, and oligochaetes (Dundee, 1958).
O. Predators. According to Petranka (1998), few data are available on natural predators, but they undoubtedly include crayfish and raccoons. Terrestrial stages probably are preyed upon by large beetles, other salamanders, and frogs. Fishes will feed on larvae and appear to exclude them in downstream sections of streams (Petranka, 1998).
P. Anti-Predator Mechanisms. In daylight, both larvae and adults seek cover under stones in slowly moving water. If their stone is removed and animals feel threatened, they will seek cover under a nearby stone (Moore and Hughes, 1941).
Q. Diseases. Undescribed.
R. Parasites. McAllister et al. (1995d) described the metazoan parasites of 50 larval and adult gray-bellied salamanders from seven Arkansas counties. In general, these parasites are typical of parasites reported from other plethodontid salamanders and exhibit little or no host specificity. Seven (14%) animals were infected with ≥ 1 parasites as follows: a seuratoid nematode (Desmognathinema nantahalaensis; three animals), an ancanthocephalan (Fessisentis vancleavei; two animals [this species also reported by Malewitz (1956) from Oklahoma specimens and from Madison and Benton counties, Arkansas, by Saltarelli (1977) and Buckner and Nickol (1978)]), larval intradermal mites (Hannemania sp.; two animals), and a plagiorchid nematode (Brachycoelium salamandrae; one animal). Ectoparasitic flukes (Sphyranra euryceae) occur in larval and neotenic gray-bellied salamanders (Dundee, 1958; McAllister et al., 1991) and nematodes in the larvae (Dundee, 1958). In terrestrial gray-bellied salamanders, cysts, probably of nematodes and flukes, are found in the skins of terrestrial individuals and cestodes occur in the gut (Dundee, 1958).
4. Conservation. Many-ribbed salamanders inhabit a fairly large region of rugged, hilly, and mountainous terrain, most of which is not amenable to agriculture or developing urban settings. Despite considerable logging over much of the terrain and use of flatter areas for cattle grazing, substantial populations of this salamander are found in many settings, even within properties that people have developed. The only threats to the species are an increasing number of recreational homes scattered throughout the region and some improvements to caves that they inhabit being turned into commercial caves. Some of the stream areas that the species inhabits might become polluted if improper sanitary facilities are constructed. Overall, we do not visualize much impact on the species.
1Stanley E. Trauth
2Harold A. Dundee
Literature references for Amphibian Declines: The Conservation Status of United States Species, edited by Michael Lannoo, are here.
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