Black-bellied Salamander, Common Blackbelly Salamander
© 2009 Henk Wallays (1 of 58)
Desmognathus quadramaculatus Holbrook, 1840
Mark B. Watson1
1. Historical versus Current Distribution. Bishop (1943) shows that black-bellied salamanders (Desmognathus quadramaculatus) range from northern Georgia in the southeastern United States northward through the mountains of western North Carolina, eastern Tennessee, and southeastern Virginia to southern West Virginia. Research since Bishop’s work has extended the range north into Allegheny and Franklin counties, Virginia, and upstream in the New River in West Virginia to its confluence with the Gauley River in Fayette County (Valentine, 1974). Recent surveys in West Virginia have shown that the range of black-bellied salamanders extends farther north than previously known. Turner (1997) located black-bellied salamanders in tributaries upstream in the Gauley River (Nicholas County) and T.K.P. and W.J. Humphries (unpublished data) found them in tributaries 4.8 km upstream from the mouth of the Greenbrier River. They also have been located approximately 4.8 km upstream from the mouth of the Bluestone River (Waldron et al., 2000). Black-bellied salamanders have probably been introduced into drainages where they previously were not present due to fisherman releasing individuals used as bait. Isolated records from the Piedmont of Georgia and South Carolina have been attributed to such bait–release introductions (Martof, 1953, 1955). Recent collections on the Georgia Piedmont suggest that at least one disjunct record represents a breeding, persistent population (J.B. Jensen, unpublished data).
2. Historical versus Current Abundance. Black-bellied salamanders are used widely as fish bait over the extent of their range (Camp and Lovell, 1989; Jensen and Waters, 1999). Petranka (1998) suggests that over-harvesting of black-bellied salamanders for bait may be detrimental to some local populations. Statements recorded by a local fisherman in West Virginia suggest that salamander populations have declined in that area in past years (Green and Pauley, 1987). Turner and Pauley (1992) found that black-bellied salamanders were absent at the mouth of a tributary where fisherman could easily search for salamanders as bait, but present farther upstream where fisherman had not searched.
Mitchell et al. (1999) concluded that stream pollution caused by acid mine drainage and sewage and over-collection for fish bait are threats to black-bellied salamanders. Stream acidification and contamination by metal pollutants has resulted in the elimination of black-bellied salamanders from one stream in the Great Smoky Mountains (Kucken et al., 1994). Petranka et al. (1993) demonstrated that clearcut harvesting of timber negatively affects populations of black-bellied salamanders. They estimated that timber-harvesting rates during the 1980s and early 1990s resulted in the loss of at least 14 million salamanders of all species annually in western North Carolina. Long-term (20 yr) monitoring has shown no significant changes in the abundance of this species in undisturbed locations in the Appalachian Mountains of western North Carolina (Hairston, 1996).
3. Life History Features.
A. Breeding. Reproduction is aquatic.
i. Breeding migrations. Black-bellied salamanders do not migrate.
ii. Breeding habitat. First-, second-, and third-order streams (Pauley, 1993b).
i. Egg deposition sites. Eggs of black-bellied salamanders are deposited under rocks in moving water. Austin and Camp (1992) found egg clutches in a low elevation Georgia site (Habersham County) in May and hatchlings in July. Eggs with attending females have been found in July at a high elevation site in Union County (C.D.C., personal observation). In Virginia, egg deposition occurs from June–September (Organ, 1961a). Pauley (1993b) reported a nest in West Virginia in July.
ii. Clutch size. Averages 45 eggs (range = 38–55) in black-bellied salamanders from North Carolina (Bruce, 1996). Bruce (1996) found eggs attached to large rocks in the middle of Wolf Creek, North Carolina, in June. Mills (1996) similarly found eggs in this time period from the northern portion of the range. Eggs range from an average of 3.9 mm in diameter and 188–203 mg dry weight (Tilley and Bernardo, 1993) to > 5 mm in diameter (C.D.C., unpublished data). Oocytes require 2 yr to develop, and females probably oviposit biennially (Austin, 1993).
Marks (2000) describes the embryonic development of black-bellied salamanders. Turner (1997) observed hatchlings from April–September in tributaries of the Gauley River in West Virginia.
i. Length of larval stage. Bruce (1988a) found 3–4 age classes of larvae in southwestern North Carolina, depending on the population. Castanet et al. (1996) used skeletochronological aging techniques to show that larval development in one North Carolina population took 3 yr. Bruce et al. (2002) found that larvae in another North Carolina population metamorphosed in 4 yr. Austin and Camp (1992) found differences in the age and size of newly metamorphosed animals from two populations in northern Georgia. Low elevation populations had an average of 40–43 mm SVL at 3 yr, whereas hatchlings from a high elevation population averaged 54 mm SVL when 4 yr old. Mills (1996) interpreted five size classes from collections of larvae in West Virginia streams and suggested that there may be a 54–60 mo larval period. Populations in northeastern Tennessee (Csanady, 1978) and northwestern North Carolina (Beachy and Bruce, 1998) may have larval periods as short as 1 yr. Larval period in black-bellied salamanders is strongly related to rainfall patterns and is the most important determinant of adult body size (Camp et al., 2000). Bruce (1985a) found a higher frequency of young larvae in small tributary streams and fewer young larvae in larger streams, suggesting that reproduction was concentrated in headwater streams and that older larvae move downstream. Orr and Maple (1978) found that black-bellied salamander larvae hatched earlier and had more rapid yolk absorption than seal salamanders (D. monticola). They suggested this may have an adaptive advantage.
a. Food. Beachy (1997a) showed that black-bellied salamander larvae fed on Blue Ridge two-lined salamander (Eurycea wilderae) larvae in controlled experiments. Davic (1991) found that aquatic insects made up the bulk of the natural diet of larval black-bellied salamanders. In West Virginia, in the northern extreme of the range of black-bellied salamanders, larvae prey on larval dipterans and trichopterans, and plecopteran and ephemeropteran nymphs, as well as several other insect taxa (Mills, 1996). Growth of black-bellied salamander larvae is not regulated by their densities, but survival of first-year larvae is reduced by the presence of larger black-bellied salamanders under artificial, experimental conditions (Beachy, 1993b).
b. Cover. Larvae often take cover in the interstices of shallow riffle areas in streams (Austin and Camp, 1992; C.D.C., personal observation).
D. Juvenile Habitat. Juvenile habitats are similar to the adults, although juveniles tend to be farther into the middle of streams than adults (Camp and Lee, 1996). Juveniles occupy smaller substrates than adults, avoid refugia containing large salamanders (Southerland, 1986c), and flee from large conspecifics (Camp and Lee, 1996). Juveniles spend more time actively foraging outside of refugia than do adults (Camp and Lee, 1996).
E. Adult Habitat. With the exception of shovel-nosed salamanders (D. marmoratus), black-bellied salamanders are the most aquatic of the Desmognathus species. They are most often found in cool, small-volume, high-gradient perennial streams. Whitford and Hutchison (1965) suggested that black-bellied salamanders may be limited to cool streams by their limited surface area for respiration. In the Great Smoky Mountains of Tennessee, they have been found from 375–1,725 m (1150–5200 ft; Huheey, 1966). They are associated with clean water and large cobbles and flat rocks (Pope, 1924; Davic and Orr, 1987; Mills, 1996; Turner, 1997). Davic and Orr (1987) suggested there may be a positive correlation between the density of rocks and pebbles and the density of black-bellied salamanders. Black-bellied salamanders may also use burrows in stream banks as refugia or for ambushing prey (Huheey, 1964; Brandon and Huheey, 1971; Camp and Lee, 1996). Southerland (1986c) found that increasing streamside cover might influence species composition due to competition for refugia in densely populated streams. Pollio (1993) observed that adult black-bellied salamander surface density decreased as water temperature increased in a southern West Virginia stream. Mills (1996) observed a seasonal shift in the foraging location of black-bellied salamanders in West Virginia. Most (97%) black-bellied salamanders forage above the water’s edge; as summer progresses, more salamanders forage in the stream away from the bank. Post-metamorphic black-bellied salamanders forage in the water from October–April. Adults spend more time ambushing prey from refugia than do juveniles, which tend to forage more actively (Camp and Lee, 1996).
Black-bellied salamanders inhabit cool mountain streams. Bogert (1952) observed black-bellied salamanders in streams and springs with water temperatures that varied from 10–17.7 ˚C. Spight (1968) found that black-bellied salamanders lost water as a linear function of body weight (y = 2.0031 + 0.41x at 60–70% humidity, where y = log rate loss, x = log of body weight). Black-bellied salamanders lost water at an intermediate rate compared to other species tested. Whitford and Hutchison (1965) described various measurements of metabolism of black-bellied salamanders. Black-bellied salamanders had a lower metabolic rate than spotted salamanders (Ambystoma maculatum) of comparable size. Below 10 ˚C, 80–90% of the oxygen was absorbed through the skin. The optimum temperature is from 15–20 ˚C. Booth and Feder (1991) showed that low flow rates of water result in the buildup of hypoxic layers around black-bellied salamanders that depend entirely on cutaneous breathing for gas exchange. They hypothesized that this species can survive reduced aerobic conditions using anaerobic metabolic pathways. Field studies, however, have shown that most individuals occur with at least part of their bodies above the water level (Camp and Lee, 1996; Mills, 1996), perhaps to avoid hypoxic conditions in quiet water.
F. Home Range Size. Individual black-bellied salamanders repeatedly used the same refugia or moved among a series of primary and secondary refugia (Camp and Lee, 1996). Camp and Lee (1996) found home ranges to be linear with a minimum area of 1,207 cm2. They found no correlation between home range size and body size (SVL). Home ranges often overlapped with home ranges of neighboring salamanders.
G. Territories. Black-bellied salamanders aggressively defended refugia from conspecifics (Camp and Lee, 1996). Camp and Lee (1996) rarely found two conspecifics in the same refugium, and they also found intruders absent from refugia even though the resident was away. This suggested that chemical cues might be used to mark territories. These chemical cues may be detected by smaller conspecifics to avoid predation. Roudebush and Taylor (1987b) found black-bellied salamanders were attracted to extracts of like-sized salamanders. Camp (1996) examined bite scars on black-bellied salamanders and suggested that bites frequently are inflicted by conspecifics. A high portion of these bites are on the head in larger salamanders and towards the tail on smaller ones. Males and females had similar bite frequencies, and mature animals had more scars than immature ones. These data suggest that larger animals may tend to stand their ground more against aggressors while smaller individuals may flee.
H. Aestivation/Avoiding Dessication. Aestivation is unknown. Black-bellied salamanders are associated with streams, which may buffer them from dessicating conditions.
I. Seasonal Migrations. Black-bellied salamanders do not migrate.
J. Torpor (Hibernation). Unknown.
K. Interspecific Associations/Exclusions. Black-bellied salamanders can be found associated with other members of the genus Desmognathus in streams. Seal salamanders, dusky salamanders (D. fuscus complex), and mountain dusky salamanders (D. ochrophaeus complex) are all associated with streams and seeps and typically sort by size and association with water. Desmognathus species that inhabit these streams are structured into size gradients with larger-bodied, more aquatic species found closer to water, and smaller-bodied salamanders found in more terrestrial locations (Hairston, 1949, 1987; Organ, 1961a). More terrestrial species used woody debris rather than rocks as cover objects (Southerland, 1986a). Larger black-bellied salamanders have been shown to displace smaller salamanders from refugia and feeding areas. Predation by black-bellied salamanders has been suggested to alter habitat selection of seal salamanders (Kleeberger, 1984; Carr and Taylor, 1985; Southerland, 1986b) and to be an important factor in the development of the community structure of streamside desmognathines (Hairston, 1980c). Grover (2000) reported that northern dusky salamanders and other salamanders (e.g., Plethodon species) were farther from water in stream transects where black-bellied salamanders were present than in streams where black-bellied salamanders were absent. There was not a significant difference in distance from water between seal salamanders inhabiting the same respective transects, and Grover (2000) could not rule out the possibility that abiotic differences among the streams caused the observed differences in salamander dispersion. Roudebush and Taylor (1987a,b) suggested that seal salamanders use chemical clues to avoid black-bellied salamanders. Jacobs and Taylor (1992) found that seal salamanders avoided substrates that were covered with secretions from mucous glands of black-bellied salamanders. Black-bellied salamanders did not avoid secretions from seal salamanders. Hairston (1986) suggested that predation and competition were important factors in determining species composition of streams containing desmognathine assemblages. Evidence from dietary studies, however, suggests that interspecific predation among black-bellied salamanders and other salamanders is extremely rare.
Exceptions to the body-size/moisture gradient have been recently reported. Beachy and Bruce (1998) discovered a population of small-sized black-bellied salamanders in northwestern North Carolina that is syntopic with seal salamanders that are equivalent in size but occur more terrestrially. Black-bellied salamanders in Union County, Georgia, are syntopic with a cryptic, newly discovered species of dark-ventered salamander that is approximately half the SVL of black-bellied salamanders (Camp et al., 2000). The newly discovered form, though smaller, appears to be as aquatic as black-bellied salamanders (Camp et al., 2002).
L. Age/Size at Reproductive Maturity. Organ (1961a) estimated that male black-bellied salamanders from Virginia are sexually mature at 3.5 yr of age and females at 4.5 yr of age. However, because his estimates were based on specimens that were pooled from a number of different populations, their accuracy has been questioned (Austin and Camp, 1992). Bruce (1993) found that black-bellied salamander males were 64–94 mm SVL as adults compared to 73–85 mm for females in North Carolina. Males from Georgia populations may reach 120 mm SVL (Camp et al., 2000). Black-bellied salamanders in the northern portion of the range appear to reach maturity at a smaller size range (57.6–77.5 mm; Mills, 1996). Bruce (1988a) estimated that first mating occurred at 5–6 yr of age for males and 6–7 yr for females. Using data from skeletochronological analysis, Castanet et al. (1996) found that male black-bellied salamanders in North Carolina reached sexual maturity in 6–7 yr, while females matured at 7–8 yr of age. Using the same technique, Bruce et al. (2002) found that males and females at Coweeta, North Carolina, reached maturity at 7–8 yr and 9–10 yr, respectively.
M. Longevity. Skeletochronological studies have shown that natural longevity is at least 15 yr (Castanet et al., 1996; Bruce et al., 2002). Bruce et al. (2002) found individuals that were 15 yr of age.
N. Feeding Behavior. Camp and Lee (1996) showed that black-bellied salamander adults are largely ambush predators using burrows and rock crevices to wait for prey. Smaller salamanders are more likely to actively forage among refugia than adults. Adult salamanders will move away from the stream at night to feed (Hairston, 1987). During warm seasons with high relative humidity, post-metamorphic black-bellied salamanders forage at the edge of the stream and ≤ 30 cm from the edge (Mills, 1996).
Davic (1991) compared the diets of larval, juvenile, and adult black-bellied salamanders and found that there was a shift in the diet from aquatic prey to aerial prey after metamorphosis. Over 82% of prey in larvae were aquatic compared to 35% of aquatic prey in juveniles. Adult black-bellied salamanders preyed upon crayfish as well as aquatic and terrestrial insects (Hairston, 1949; Martof and Scott, 1957; Kleeberger, 1982). Mills (1996) found that the major prey items of post-metamorphic black-bellied salamanders were dipteran larvae followed by adult coleopterans and a variety of other insect taxa. A number of studies have characterized black-bellied salamanders as important predators of other salamanders (e.g., Hairston, 1980c, 1986, 1987; Southerland, 1986b; Formanowicz and Brodie, 1993; Grover, 2000). However, these conclusions have been based on anecdotal reports (e.g., Hairston, 1980c, 1986) or predation that was induced under artificial, experimental conditions (e.g., Formanowicz and Brodie, 1993). In an analysis of 659 stomachs of black-bellied salamanders from Georgia, Camp (1997b) found no evidence of predation on heterospecific desmognathines, although larval or juvenile conspecifics occasionally were taken as prey. This analysis corroborated other dietary studies (e.g., Hairston, 1949; Martof and Scott, 1957; Kleeberger, 1982; Davic, 1991; Mills, 1996) that demonstrated that predation by this species on other species of salamanders is rare. The ambush–predator strategy (Brandon and Huheey, 1971; Camp and Lee, 1996) combined with the avoidance of pheromones by smaller species (Roudebush and Taylor, 1987a; Jacobs and Taylor, 1992) probably preclude predation of salamanders under natural conditions (Camp and Lee, 1996; Camp, 1997b).
O. Predators. Egg predators probably include aquatic insects, crayfish, fishes, and other salamanders.
Larger larvae of black-bellied salamanders occasionally are cannibalistic and will prey upon smaller black-bellied salamander larvae (Beachy, 1993b; Camp, 1997b). Beachy (1991b) found that microhabitat differences resulted from experimentally induced predation of species with larger larvae (e.g., black-bellied salamanders) on smaller larvae (e.g., Blue Ridge two-lined salamanders). Beachy (1994) experimentally demonstrated that spring salamander (G. porphyriticus) larvae are more efficient predators on Blue Ridge two-lined salamanders than are black-bellied salamander larvae.
Brodie et al. (1989) showed that garter snakes (Thamnophis sp.) prey on black-bellied salamanders. Adult black-bellied salamanders occasionally are cannibalistic on smaller conspecific juveniles and larvae (Camp, 1997b). In Georgia, black-bellied salamanders may occur in streams that are also occupied by potentially predaceous fishes (e.g., banded sculpins [Cottus carolinae], bluehead chubs [Nocomis leptocephalus], creek chubs [Semotilus atromaculatus]), and northern watersnakes [Nerodia sipedon] have been observed foraging in optimum habitat for black-bellied salamanders [C.D.C., personal observation]).
P. Anti-Predator Mechanisms. Eggs of black-bellied salamanders frequently are deposited beneath large rocks in streambeds, obscuring them from potential predators. Females of many plethodontid salamanders exhibit brooding behavior (Jaeger and Forester, 1993). Female black-bellied salamanders have been observed tending clutches of eggs (T.K.P., personal observation).
Adults do not exhibit noxious skin secretions. They remain motionless when touched or show other passive defensive postures. They will bite and use pseudoaposematic coloration for defense. Members of the genus Desmognathus seem to be palatable to many species. The aggressive nature and biting ability of black-bellied salamanders are used as anti-predator defenses. Brodie (1978) showed that biting was used as a defense against shrew attacks. Brodie et al. (1989) observed three anti-predator responses during encounters with snakes (Thamnophis sp.): flipping, biting, and tail autotomy. Biting appeared to be a successful defense against snake attacks, and all salamanders that were bitten by a snake on the tail were not eaten by the snake (Brodie et al., 1989). Pope (1928) describes a black-bellied salamander from Highlands, North Carolina, as assuming a defensive posture and opening its mouth threateningly when he attempted to capture it. Brodie (1977) lists members of genus Desmognathus as using biting and pseudoaposematic coloration as antipredator defenses.
Q. Diseases. None reported.
R. Parasites. Rankin (1937) lists the following species collected from black-bellied salamanders: Protozoa—Cryptobia borreli, Cytamoeba bacterifera, Eutrichomastix batrachorum, Hexamastix batrachorum, Hexamitus batrachorum, Hexamitus intestinalis, Karotomorpha swezi, Prowazekella longifilis, and Tritrichomonas augusta; Trematoda—Brachycoelium hospitale, Diplostomulum desmognathi, and numerous unidentified metacercariae; Cestoda—Crepidobothrium cryptobranchi and proteocephalid cysts; Nematoda—Capillaria inequalis, Omeia papillocauda, Oxyuris magnavulvaris, and spirurid cysts; Acanthocephala—cysts.
Huheey (1966) observed leeches attached to black-bellied salamanders in the Great Smoky Mountains National Park, Tennessee, and stated that leeches were a common parasite. Leeches also are common on specimens from Georgia (C.D.C., personal observation). Baker (1987) lists Batracholandros magnavulvaris, Falcaustra plethodontis, Desmognathinema nantahalaensis, and Omeia papillocauda as parasitic nematodes in black-bellied salamanders. Goater et al. (1987) lists adult nematodes (Capillaria inequalis, Thelandros magnavulvaris, Omeia papillocauda, Desmognathinema nantahalaensis, Falacustra plethodontis, and Cosmoceroides dukae), trematodes (Brachycoelium elongatum, Gorgoderina bilobata, and Phyllodistomum solidum), and cestodes (Cylindrotaenia americana) in black-bellied salamanders. They also found larval nematodes (Ascaridoidea sp.), cestodes (proteocephalan plerocercoids), and acanthocephalans (Centrorynchus conspectus) in black-bellied salamanders.
4. Conservation. Stream pollution caused by acid mine drainage and sewage, clearcut forestry practices, and overcollection for fish bait are threats to black-bellied salamanders. In contrast, black-bellied salamanders are used widely as fish bait over the extent of their range and because of this, they have been introduced into drainages where they previously were not present. While long-term (20 yr) monitoring has shown no substantive changes in the abundance of this species in undisturbed locations in the Appalachian Mountains of western North Carolina (Hairston, 1996), black-bellied salamanders are listed as a Species of Special Concern in West Virginia.
1Mark B. Watson
2Thomas K. Pauley
3Carlos D. Camp
Literature references for Amphibian Declines: The Conservation Status of United States Species, edited by Michael Lannoo, are here.
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