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Description

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Ambystoma annulatum has a slender girth, well-rounded body and, among salamanders, a generally small head and long tail. Adults grow to a range of 140-180 mm snout-vent length, with a record length of 255 mm. This species usually has 15 costal grooves (Behler 1996). It has a depressed snout that is evenly and bluntly rounded. Vomerine teeth are arrayed in two short series located entirely behind the choanae. Each series consists of about 7-11 small, blunt teeth. There are five toes on each hind foot (Bishop 1962).In life, the dorsal color of adults ranges from dark gray to blackish brown, with contrasting white to yellowish bands. Ventrally, the color varies from light gray to yellowish, peppered with light-colored spots. There is typically a short, light colored bar between the eyes that may continue below the eyes to point diagonally posterior. Recently metamorphosed juveniles have a drab green to dark gray dorsal surface, and a row of dorsolateral yellowish spots extending from the front limbs to the tip of the tail. Laterally, a broad band runs from the gills two-thirds of the way down the tail, which lacks pigmentation. Bellies are grayish-yellow. Juveniles develop the blotches or rings that characterize this species approximately two months after metamorphosis (Hutcherson et al. 1998). The sexes are monomorphic and it is unknown whether there is any geographic or seasonal variation (Bishop 1962; Johnson 1977; Petranka 1998).The specific epithet annulatum means "ringed" (Beltz 2002).

References

  • Behler, J.L. and King, F.W. (1996). National Audubon Society Field Guide to North American Reptiles and Amphibians. Knopf, New York, NY.
  • Beltz, E. (2002). ''Names of the reptiles and amphibians of North America.'' Original Description Citations for the Reptiles and Amphibians of North America.
  • Bishop, S.C. (1962). Handbook of Salamanders. Hafner, New York, NY.
  • Hutcherson, J. E., Peterson, C. L. and Wilkinson, R. F. (1989). ''Reproductive and larval biology of Ambystoma annulatum.'' Journal of Herpetology, 23, 181-183.
  • Johnson, T.R. (1977). The Amphibians of Missouri. University of Kansas Publications, Lawrence, KS.
  • Mathis, A., Murray, K. L., and Hickman, C. R. (2003). ''Do experience and body size play a role in responses of larval ringed salamanders, Ambystoma annulatum, to predator kairomones? Laboratory and field assays .'' Ethology, 109, 159-170.
  • McAllister, C. T., Trauth, S. E., and Cochran, B. G. (1995). ''Endoparasites of the ringed salamander, Ambystoma annulatum (Caudata: Ambystomatidae), from Arkansas.'' The Southwestern Naturalist, 40, 327-330.
  • Noble, G. K., and Marshall, B. C. (1929). ''The breeding habits of two salamanders.'' American Museum Novitates, 347, 1-12.
  • Nyman, S., Wilkinson, R. F., and Hutcherson, J. E. (1993). ''Cannibalism and size relations in a cohort of larval ringed salamanders (Ambystoma annulatum).'' Journal of Herpetology, 27, 78-84.
  • Petranka, J. W. (1998). Salamanders of the United States and Canada. Smithsonian Institution Press, Washington D.C. and London.
  • Spotila, J. R., and Beumer, R. J. (1970). ''The breeding habits of the ringed salamander, Ambystoma annulatum (Cope), in northwestern Arkansas.'' American Midlands Naturalist, 84, 77-89.
  • Trapp, M. M. (1956). ''Range and natural history of the ringed salamander Ambystoma annulatum, Cope (Ambystomatidae), .'' Southwestern Naturalist, 1, 78-82.
  • Trapp, M. M. (1959). Studies on the Life History of Ambystoma annulatum Cope. Master's Thesis. University of Arkansas, Fayetteville.
  • Wilson, T. J. (1993). ''.'' Predation of Ringed Salamander Larvae, Ambystoma annulatum. Southwest Missouri State University, M.S. Thesis, Springfield.

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Distribution and Habitat

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Ambystoma annulatum is native to hardwood and mixed hardwood-pine forested areas in and around the Ozark Plateau and Ouachita Mountains of Arkansas, Oklahoma, and Missouri (Petranka 1998). Most specimens are found in the vicinity of Hot Springs, Arkansas and the Missouri portion of the Ozark Plateau (Bishop 1962; Johnson 1977). Small populations have also been found in western Illinois and eastern Oklahoma (Petranka 1998). Ambystoma annulatum is found in damp forested areas, usually under leaves, rotting logs, or in abandoned ground holes of other organisms, near shallow ponds. This species is highly fossorial (adapted to digging), and adults are often found in subterranean refuges (Petranka 1998).
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Life History, Abundance, Activity, and Special Behaviors

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Although Petranka (1998) thought that this salamander was becoming increasingly rare and perhaps endangered, a more recent assessment by Hammerson (2004) concluded that the population was stable and probably exceeded 10,000 in number, putting the species at Least Concern. Threats include loss of forest habitat surrounding breeding ponds; draining or filling of breeding ponds; and the introduction of predatory fish into breeding ponds.
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Life History, Abundance, Activity, and Special Behaviors

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This species is secretive and fossorial and is generally not seen except on rainy autumn nights during breeding season. Breeding has been observed by a number of authors (Noble and Marshall 1929; Trapp 1956, 1959; Spotila and Beumer 1970; Hutcherson et al. 1989). Ambystoma annulatum breeds in shallow, fishless water (permanent or temporary ponds) and exhibits a preference for muddy or murky bodies of water. Adults migrate to breeding areas after the first heavy rains of the autumn breeding season (mid-September through November, most commonly in October, though the exact times vary depending on the local habitat). Cool temperatures and heavy rainfall stimulate breeding. This species is an explosive breeder and breeding males may try to reproduce 2-4 times in a season (Petranka 1998). Breeding occurs at night in breeding ponds, where hundreds of individuals congregate. The males usually arrive first, and can be distinguished from the females by their swollen cloacae (Bishop 1962). Males usually approach the females in shallow water and engage in courtship by nudging the female’s cloaca (Johnson 1977). They then deposit a spermatophore a few centimeters away; a male may deposit nine spermatophores in two minutes. The male may repeat this process several times; however, the female will pick up the spermatophore only after being actively courted. The more males in the pond, the less specifically they court, and they will nudge both males and females before depositing a spermatophore (Petranka 1998). Males deposit spermatophores on rocks, other spermatophores, and even each other. After breeding, the salamanders begin to move away from the pond (Bishop 1962). Fertilization is internal (Petranka 1998). A day or two after mating (24-48 hours), the female deposits between 5 and 40 eggs (mean of 10), in strings or small masses (Petranka 1998). Generally she lays the egg mass (usually about 150 eggs total) directly on the pond bottom, but she may attach them to submerged vegetation (Petranka 1998). Females may distribute the clutch in several egg masses, resorb some ova, or lay only a portion of the clutch on any given night, since Hutcherson et al. (1989) found that females in captivity had considerably more ovarian eggs than were counted in oviposited masses examined by Trapp (1959). In Missouri, permanent ponds used for breeding had a bottom substrate consisting of a thick layer of leaf litter and manure (Hutcherson et al. 1989). Eggs measure approximately 2 mm in diameter and hatch after 9-16 days (a relatively short embryonic period), again depending on the exact location (Petranka 1998). Eggs can survive temporary dry periods out of the water, with Hutcherson et al. (1989) reporting that eggs were deposited in one pond even though there was no standing water; those deposited within mud cracks or under partial shade from vegetation survived for 14-19 days until the pond refilled and hatching could occur. To get an idea of the maximum survival period out of water, Hutcherson et al. (1989) collected 879 embryos from the pond basin and incubated them at 17°C on moist soil under moist paper towels for 52 days. Twenty embryos survived and all hatched within five minutes of being submerged in water after 52 days out of water.Mature aquatic larvae are on average 48 mm in length, and have well-developed legs, toes, and a dorsal fin that extends to the head (Petranka 1998). They typically begin metamorphosis the following February through May (Petranka 1998), although Hutcherson et al. (1989) observed larvae leaving one of their Missouri study ponds during June and early July. Newly metamorphosed juveniles measure approximately 34-40 mm in length and soon begin to crawl onto land in search of underground abodes, particularly on rainy days (Hutcherson et al. 1989). Sexual maturity is attained in the second or third year of life (Petranka 1998).As an adult, this carnivore preys mostly on earthworms, insects, and land snails (Johnson 1977). Hutcherson et al. (1989) found that only 3 of 16 females examined had food in the stomach, and were unable to identify the contents. Trapp (1959) found that only 3 of 23 salamanders had prey items in the stomach (one with unidentifiable stomach contents, two containing earthworms). Newly transformed salamanders mostly had empty stomachs (24 of 30) when examined by Hutcherson et al. (1989), but some (6 of 30) had fly larvae in the stomach. Larvae eat a variety of prey, with Hutcherson et al. (1989) finding cladocerans and copepods the main components of the larval diet in autumn, dipteran larvae during winter and spring, and other occasional larval prey items including ostracods, dragonfly and damselfly nymphs, hemipterans and snails. In contrast, Trapp (1959) found that cladocerans and copepods comprised the bulk of the diet in spring (March and April), with minor components including molluscs, eggs, and Chironomus; Trapp (1959) also concluded that cladocerans and copepods were consumed preferentially, based on an analysis of food item abundance in water samples. Larval cannibalism has also been observed, both in the wild and the lab (Hutcherson et al. 1989; Nyman et al. 1993). Natural predators of adults include owls, snakes, shrews, skunks, raccoons, opossums, and other mammals. When attacked or feeling threatened, A. annulatum coils its body while tucking the head underneath the base of the tail for protection (Petranka 1998). Young larvae are preyed on by newts (Notophthalmus viridescens louisianensis, among other species) but larger ringed salamander larvae became too big for the Notophthalmus to consume at about a month prior to larval metamorphosis (Wilson 1993). In laboratory experiments comparing larval ringed salamander responses to predator (newt) vs non-predator (tadpole) chemical stimuli, larval Ambystoma annulatum were able to distinguish chemical signals of predatory newts and smaller larvae subsequently decreased activity. Other predators of ringed salamander larvae include aquatic insects, other aquatic salamanders, wading birds, and snakes (Mathis et al. 2003).Ringed salamanders can harbor a variety of endoparasites. McAllister et al. (1995) found that 83% of the salamanders in their sample (n=41) carried at least one parasite species. Endoparasites included ascarid (Cosmocercoides variabilis, in the rectum and feces), spirurid (species unknown, encysted in the stomach wall) and rhabditid nematodes (Rhabdias ranae, in the lungs and body cavity), and a myxosporean protozoan, Myxidium serotinum (in the gall bladder).
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Life Expectancy

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Longevity in ringed salamanders is unknown. Some other salamander species may live up to 10 years.

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Seaholm, L. 2002. "Ambystoma annulatum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_annulatum.html
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Associations

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Predators of ringed salamanders include owls, snakes, shrews, skunks, raccoons, opossums, and other mammals (Petranka 1998). When these salamanders are being attacked or feel threatened they will coil their bodies while tucking their heads underneath the base of their tales for protection (Petranka 1998).

Known Predators:

  • owls (Strigiformes)
  • snakes (Serpentes)
  • shrews (Sorex)
  • skunks (Mephitidae)
  • raccoons (Procyon lotor)
  • Virginia opossums (Didelphis virginiana)
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Seaholm, L. 2002. "Ambystoma annulatum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_annulatum.html
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Morphology

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Adult ringed salamanders range in length from 140 to 180 mm (5 1/2 to 7 inches). The record length is 255 mm (9 1/2 inches). Ambystoma annulatum has a slender well-rounded body with a small elongated head and a long tail. They have depressed snouts that are evenly and bluntly rounded. A. annulatum has about 15 costal grooves and 5 toes on the hind feet. They also have "vomerine teeth in two short series entirely between the inner nares, each series consisting of three rows of about 7 - 11 small blunt teeth" (Bishop 1962).

This is a striking salamander that is a dark blackish brown color with light cross bands and spots that are a buff - yellow color. There is some variation in the intensity of the markings. The belly is a pale grayish white. Usually, there is a short, light colored bar between the eyes. Someimes this continues below the eyes, pointing backwards diagnally. Looking from the top, the tail and body can appear to be completely ringed, hence the name "ringed" salamander. Interestingly enough, the rings never completely go around the body. Males and females are monomorphic and no textual evidence has been found if geographic or seasonal variation within the species exists.

The larvae of ringed salamanders are an average of 48 mm in length. They have well developed legs and toes and have a dorsal fin that extends to the head. Dorsally and on the lower sides there is uniform coloration. On the sides there is a broad, definite band lacking pigmentation from the gills to two-thirds down the tail. Juveniles get their yellow coloration after metamorphosis, and form their adult patterns within two months

(Bishop 1962, Johnson 1977, Petranka 1998).

Range length: 140 to 255 mm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: sexes alike

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Seaholm, L. 2002. "Ambystoma annulatum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_annulatum.html
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Behavior

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Ringed salamanders may communicate mainly through chemical and tactile cues during the breeding season.

Communication Channels: tactile ; chemical

Perception Channels: visual ; tactile ; chemical

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Seaholm, L. 2002. "Ambystoma annulatum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_annulatum.html
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Habitat

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Ambystoma annulatum prefers damp hardwood forests that are near shallow ponds (Bishop 1962). These salamanders are usually found hidden under rocks and logs, in piles of dead leaves, or burrowing in the soil; most of the year is spent below the surface of the ground (Bishop 1962, Johnson 1977).

Habitat Regions: temperate ; terrestrial

Terrestrial Biomes: forest

Aquatic Biomes: lakes and ponds; temporary pools

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Seaholm, L. 2002. "Ambystoma annulatum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_annulatum.html
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Distribution

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Ambystoma annulatum has been mostly found in the vicinity of Hot Springs Arkansas and throughout most of the forested Ozark Plateau in Missouri (Bishop 1962, Johnson 1977). Small populations of ringed salamanders have been found in south western Illinois and eastern Oklahoma as well (Petranka 1998).

Biogeographic Regions: nearctic (Native )

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Trophic Strategy

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Ambystoma annulatum is carnivorous, eating mostly earthworms, insects, land snails, and other invertebrates (Johnson 1977). Cannibalism has been observed in this species in both the field and the laboratory (Petranka 1998). Larvae of A. annulatum eat ostracods, hemipterans, and dragonfly and damselfly nymphs (Petranka 1998).

Animal Foods: amphibians; insects; terrestrial non-insect arthropods; mollusks; terrestrial worms; aquatic or marine worms

Primary Diet: carnivore (Insectivore , Vermivore)

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Seaholm, L. 2002. "Ambystoma annulatum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_annulatum.html
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Associations

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Ringed salamanders are important predators of small invertebrates in the ecosystems in which they live. Some animals may depend on the dense aggregations of salamander eggs during the breeding season.

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Seaholm, L. 2002. "Ambystoma annulatum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_annulatum.html
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Benefits

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Ringed salamanders are not economically important to humans, but are of interest to scientists and to nature-oriented tourists. It is a specialized species, unique to its Ozark habitat. Because of their docility and striking appearance, they might be useful in environmental education programs (Petranka 1998).

Positive Impacts: ecotourism

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Benefits

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There are no negative impacts of ringed salamanders.

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Seaholm, L. 2002. "Ambystoma annulatum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_annulatum.html
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Life Cycle

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Development - Life Cycle: metamorphosis

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Seaholm, L. 2002. "Ambystoma annulatum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_annulatum.html
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Conservation Status

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Ringed salamanders are an increasingly rare (and probably endangered) animal, most likely because of their restricted range and specific breeding requirements (Petranka 1998). The breeding habitats for these creatures should be protected whenever possible.

US Federal List: no special status

CITES: no special status

IUCN Red List of Threatened Species: least concern

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Seaholm, L. 2002. "Ambystoma annulatum" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Ambystoma_annulatum.html
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Reproduction

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Eggs are fertilized internally via spermatophores and the eggs are deposited in shallow, fishless water in the fall, mostly in October (Petranka 1998). Courtship occurs in shallow water as well (Johnson 1977).

They reproduce at night in breeding ponds where hundreds of them congregate. Usually the males are found at the breeding ponds first, and are distinguishable from the females by their swollen cloacas (Bishop 1962). Males will usually approach females. They start off by nudging the female's cloaca, and then swim off a short distance and deposit a few spermatophores (Petranka 1998). A male may deposit nine spermataphores in two minutes (Petranka 1998). The more males that come into the breeding area, the less specific males get when depositing spermatophores, nudging both males and females before deposition (Petranka 1998). Males deposit spermatophores on rocks, on other spermatophores, and even on other individuals. At this time females are generally passive and do not pick up seminal fluid while they are actively being courted by males (Petranka 1998). Breeding lasts a few days, after which the salamanders begin to move away from the ponds (Bishop 1962).

Mating System: polygynous

Ringed salamanders reproduce in shallow water. Breeding takes place in the fall between September and November (Bishop 1962). Cool temperatures and heavy rains stimulate breeding. Females begin ovipositing within 1 - 2 days after breeding. These salamanders are usually sexually mature 2 - 3 years after metamorphosis (Petranka 1998). Eggs are laid in clusters with an average of 10 eggs in a cluster (Bishop 1962). Usually, about 150 eggs are laid in total and are sometimes attached to vegetation but are usually laid directly on the bottom of ponds (Bishop 1962). The embryonic period of A. annulatum is fairly short. Premature pond freezing and drying are the biggest risks to the embryos and larvae (Petranka 1998). A. annulatum is an explosive breeder, and at times breeding males will try to reproduce 2 - 4 times during the breeding season (Petranka 1998).

Breeding interval: Breeding occurs once yearly.

Breeding season: Breeding occurs during the fall, between September and November.

Average number of offspring: 150.

Range age at sexual or reproductive maturity (female): 2 to 3 years.

Range age at sexual or reproductive maturity (male): 2 to 3 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); oviparous

Aside from provisioning eggs before fertilization there is no parental care.

Parental Investment: no parental involvement

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Ringed salamander

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The ringed salamander (Ambystoma annulatum) is a species of mole salamander native to hardwood and mixed hardwood-pine forested areas in and around the Ozark Plateau and Ouachita Mountains of Arkansas, Oklahoma, and Missouri.[2] This species of salamander has slander body, small head, and long tail. They are usually found to have various dorsal color from dark gray to dark brown. Various close relatives are found such as Ringed salamander (A. annulatum), marbled salamander (A. opacum), and spotted salamander (A. maculatum). This species of salamander has cannibal behavior especially those in large body size.

It is found in damp, forested areas, usually under leaves, rotting logs, or in abandoned ground holes of other organisms, near shallow ponds. Highly fossorial (adapted to digging), adults are often found in subterranean refuges.[2] This salamander is increasingly rare and perhaps endangered. This is likely a result of its restricted range and specific breeding habit needs.[2] The world population is thought to be around 100,000 animals. Its conservation status is assessed as Least Concern by the IUCN.[1] This salamander can perform embryotic learning and social-facilitated learning.[3][4]

Description

The Ringed Salamander has a slender body shape with a small head and a long tail. This species has 15 costal (along the ribs) grooves and a depressed round snout. There are two arrays of vomerine teeth—teeth along a thin bone that form the inferior and posterior part of the nasal septum and divide the nostrils. Each series contains 7 - 11 teeth. Each of their hind feet has five toes. The dorsal color of adults can range from dark gray to blackish brown decorated with white to yellowish bands and light-colored dots. Ventral sides are generally grayish-yellow.[5] Newly metamorphosed juveniles have black backs and white bellies. A row of light spots extended from their forelimbs to the tails. Soon after metamorphosis yellow bands started to appear, and complete adult colorations form within two months after metamorphosis.[6]

Habitat and distribution

Mixed-wood pine forest

Habitat

Ringed salamander is strictly terrestrial and has been found in hardwood and mixed hardwood-pine forests. During non-breeding seasons, adults hide under leaves, rotting logs, or abandoned holes on the ground in damp forest areas. Larvae and juveniles exist in small, fishless semi-permanent ponds.[5] Most ringed salamanders are found in the vicinity of Hot Springs, Arkansas, and the Missouri portion of the Ozark Plateau.[7][8] Small populations have also been found in western Illinois and eastern Oklahoma.[2]

Geographic distribution

Ringed salamander is endemic to the Ozark plateau and the Ouachita Mountains of southern Missouri, Arkansas, and eastern Oklahoma. This is a much more restricted range of distribution compared to other species in the same genus, such as A. maculatum and A. opacum.[9][10]

Conservation

Habitat loss

While drying of ponds during late summer has significant impacts on spring breeding species such as spotted salamanders, it does not affect ringed salamanders much. However, a reduction of rain and numbers of filling ponds during September and October can cause a great decrease in ringed salamander's breeding efforts and egg depositions.[10] Although average canopy cover and leaf litter depth do not significantly relate to ringed salamander's occupancy, continuous forests are highly associated with its breeding wetlands.[11] Clear-cut timber harvest also has a negative effect on ringed salamander distribution. Forest sites after a clear-cut harvest have more open canopies, which results in higher ground temperatures. In order to survive, ringed salamanders either retreat underground into burrows or move to other places. Thus, juveniles and adults with small body sizes are more vulnerable as they have lower dispersal abilities.[12][13]

Conservation efforts

There are two major areas of focus for maximizing the proliferation of ringed salamanders, which concentrate on minimizing the local extinction of both juveniles and adults and maintaining metapopulation dynamics respectively. Protecting permanent fishless ponds is crucial for the survival of metamorphosing juveniles. Maintaining terrestrial habitat around the breeding ponds is important for reproducing adults. A study has shown that a core terrestrial habitat with a radius of 200-500m from the pond edge must be established to provide enough space for the survival of breeding adults, especially during their first reproduction. Connectivity between ponds is critical for keeping sufficient gene flows among multiple populations. A suitable habitat should have breeding ponds no more than 300m from each other, and rescue-recolonization management is recommended.[10][11]

Ringed Salamander on leaf detritus

Population structure

Close relatives

Ringed salamander (A. annulatum), marbled salamander (A. opacum), and spotted salamander (A. maculatum) are sympatric close relatives, and they are all terrestrial. They all belong to a famous genus of salamanders called mole salamanders (Ambystoma). Spotted salamanders and marbled salamanders have much larger distributions and are more well-studied than ringed salamanders. Similar to ringed salamanders, marbled salamanders also breed in fall, while the other species spotted salamander breeds in spring.[10]

Genetic population structure

Molecular evidence from nuclear microsatellites and mitochondrial DNA analysis shows that populations of ringed salamanders in the Central Interior Highlands ecoregion are separated into two.[14] One population lives on the Ozark plateau in the north, while the other population lives in the Ouachita Mountain in the south.[15] The separation first occurred during the Cretaceous to mid-Miocene and was further bisected by the development of the Arkansas River system during the Pleistocene. Genetic differentiation is also largely influenced by human-induced habitat fragmentation since European settlement.[14]

Diet

Common preys for ringed salamander larvae include microcrustaceans, dipteran larvae, larval chironimids, and many larval and adult insects such as beetles, snails, earthworms, and eggs of other salamanders and frogs.[16][6] Ringed salamanders usually serve as the top predator in their breeding ponds.[11] This is likely due to the fall breeding timing of ringed salamanders, which allows them to prey on individuals of other spring breeding species that are still in their early life stages such as spotted salamanders, marbled salamanders, red-spotted newts, and many other anuran species, thus decreases their abundances.[10][17]

Cannibalism

Although larvae of all sizes eat other prey species, those with larger body sizes can be cannibalistic. Cannibalism can be beneficial as conspecific preys are large, might contain growth-promoting hormones, and lack toxic substances in heterospecific preys. However, there is also a risk of pathogen and endoparasite transmissions. The cannibals differ from their conspecific prey by having twice longer mean body lengths and slightly broader heads. The long breeding season leads to increasing larvae size variation, which contributes to the possibility of cannibalism. Larvae hatched later in the season have smaller sizes, so early spawn breeders may gain a selective advantage by producing large offspring with higher fitness, but this advantage may be offset by more unpredictable pond conditions. The cannibalism of ringed salamanders is an opportunistic behavior that is more likely to occur in conditions of high larvae density, the coexistence of larvae of various sizes, or insufficient alternate food choice. Larvae that commit cannibalism have been shown to have wider mouths and a larger size than their noncannibal counterparts. This adds to the evidence that cannibalism is an opportunistic enterprise based no having an advantageous size.[18] Ringed salamander larvae have intermediate-level cannibalism compared to other species in the same genus. Spotted salamanders only show cannibalism in extreme conditions, while tiger salamanders (A. tigrinum) have really high tendencies of cannibalism.[16]

Reproduction and life cycle

Breeding migration and oviposition

Ringed salamander is an autumn breeding species. Males and females both reach their reproductive maturity at 1 year old, but most individuals return for breeding at 2–3 years old. Migration to breeding ponds usually started at night after or during rain in mid-September, and individuals tend to stick with specific ponds every year.[10] Annual cycle of air and soil temperatures ensures that adults enter reproductive conditions in time, but autumn rainfall is the major factor that triggers breeding migration, and the precipitation threshold must be at least 1.27 cm. A huge variation of breeding population size exists across ponds and years, which is positively correlated with the amount of rainfall during migration season.[10] Males start to migrate to breeding ponds as early as August, which is earlier than females with the earliest record in September. The end date of migration is in early November, which is similar for both sexes.[9][10]

After mating, egg laying begins on the night of courtship or the next day and is completed within two days. The female extends her rear legs laterally and arches their backs to get the cloacae onto a substrate. She stays motionless after climbing onto a suitable object until muscular contractions start. The contraction spread posteriorly from a point 2.5 from the front to the rear legs, depositing eggs from the cloaca one at a time in a row. Shapes of egg masses can be strings, clumps, or large clusters, which largely depend on what are the available supporting materials, such as sticks, for oviposition. Adults leave the pond right after breeding but could remain active above-ground for several days before getting back to their resting places.[9]

Ringed salamander larva

Larval and juvenile development

Larvae start to present in ponds in September and October. Based on eggs hatched in the laboratory environment, the average total length of larvae just hatched is 11.5mm. Unlike spring breeding species, ringed salamander larvae overwinter under the ice in their birth ponds. The larval period ranges from 7 to 9 months,[10][11] which is similar to that of marbled salamanders. Larvae mortality can be caused by freezing, drying, low pH, predation including cannibalism, and disease. Premetamorphic mortality can be as high as 99.9% and peaks during the first two months post-hatching and during metamorphic periods.[19]

Juveniles generally emerge in April the following spring. The average survival rate of juveniles from eggs to metamorphosis is only 0.2%, which is really low and doesn't vary much among ponds or years. The mean snout-vent length of metamorphosing juveniles is 39.2mm, which is about 50% of that of breeding adults, but the largest juvenile can be 76% of the adult size. Juveniles that metamorphosed in early spring are usually larger than those that metamorphose later in the summer. A larger body size corresponds to a high probability of surviving, a younger age at first reproduction, and a longer lifespan. This is the opposite trend compared to spotted salamanders, of which juveniles metamorphose later are larger, and have higher fitness.[10]

Limb regeneration loss of adults

The limb regeneration time of larval salamanders shortens with development, and postmetamorphic adult larval salamanders exhibit little or no limb regeneration.[20] Through experiments looking at regeneration times for larval salamanders at different age (40 days), axolotl (30+ days), newt (44 days) and adult salamanders (155 to 370 days), it was found that the limb regeneration time of each stage has a huge difference. Based on two criteria: each stage is similar to the sequences of previous stages of newt, while these stages must be easily identifiable by examining the external morphology of living organisms, the results revealed 11 well-defined regeneration phases.[21] There is a saying that environmental conditions cause salamanders to lose the ability to express their intrinsic ability. Regeneration was observed in the following conditions for the adult salamanders: A. annulatum, 324 to 370 dpa; A. maculatum, 255 to 300 dpa; A. texanum, 215 to 250 dpa; and A. tigranum , 155 to 180 dpa.[20]

Mate behavior

Ringed salamanders are sexually dimorphic. Breeding females are larger than males, with the mean snout-vent length being 96.3mm and 84.2mm respectively. At each breeding pond, the adult sex ratio is consistently male-biased with a 2.05:1 ratio.[10](2) A mass courtship happens when a large number of males and some females arrived at a pond. Initially, one male approaches a female and nudges her cloaca and sides of the body with his snout. After that, he quickly deposits a spermatophore somewhere nearby. A male usually deposits one spermatophore at a time but could also lay up to four in succession. This behavior is repeated and reaches a peak that lasts for 10–15 minutes. As more and more males join, a crowd of male forms around a female, where each male begin to nose the female and other males indiscriminately, and lots of spermatophores are laid at the same spot. Multiple males tend to lay their spermatophores next to or on top of each other. There can be as many as 10 or more spermatophores within a 1.6 cm2 area. Researchers didn't observe any female participation in the courtship. Females lay fertilized eggs after the mass courtship, so they might pick one or more spermatophores after the courtship as do female A. macualtum, the sympatric relative of ringed salamander. Another possibility is females may follow a chosen male and pick up his spermatophores specifically.[9]

Social behavior

Ringed salamanders can learn what are predators and foods from chemical stimuli as early as embryos. This allows larvae to avoid predation and have a higher foraging efficiency right after hatching. Lab experiments have proved that embryos exposed to odors of prey species like shrimp and mussel show attraction to those stimuli post-hatching.[3] Embryotic exposures to chemical stimuli from cannibalistic larvae or predators, such as Eastern newt (Notophthalmus viridescens) and dragonfly nymphs, resulted in increasing use of habitat refuge (vegetation cover) and decreasing activities post-hatching.[3][4] These are signs of anti-predator responses. Decreasing activity is especially effective for avoiding the detection of visual predators and reducing encountering possibilities.[4] The underlying physiological mechanism might be that the stimuli can shape the developing olfactory system in a different way to cause either olfactory imprinting or sensitization. Because of the plasticity of ringed salamander's diet, researchers think that sensitization is a more plausible explanation as it means a temporary response to a stimulus, while olfactory imprinting means showing a long-term response.[3]

Young larvae are also able to distinguish between predator and prey after observing other larvae that are known as conspecific models. Social learning is commonly thought to exist only in group-living organisms, but it can happen in non-gregarious species like ringed salamanders as well. Ringed salamander larvae can form temporary high-density groups during a short period after hatching, which provides the environment for social learning. A study on ringed salamanders has demonstrated that a larva can successfully recognize dragonfly nymph as a danger after watching the anti-predator reactions of four conspecific demonstrators.[17] This socially facilitated learning and the former embryotic learning are both beneficial in a way that it minimizes costs due to direct samplings of different foods and random encounters with predators.[3]

Enemies

Fish is a predator of ringed salamanders at all life stages in ponds.[17] Aquatic insects, salamanders, snakes, and wading birds, such as, can all be potential predators of ringed salamander larvae.[17] Studies have recorded many insect predators including Odonata, Hemiptera, and Thanmophis Proximus. Bird predators include great blue herons and American bittern, snake predators include Nerodia sipedon, and turtle predators include Chelydra serpentina.[19]

Larvae's anti-predator responses gradually decrease with increasing body sizes. Large larvae are less vulnerable to gap-limited predators such as newts. They are either too big as preys or have swifter fleeing reactions with stronger muscles. Ringed salamanders that are one month before metamorphosis are completely immune to newt predation.[4]

Ringed salamanders also suffer from lots of endoparasites. The most common parasite species is ascarid nematode (Cosmocercoides variabilis). This parasite is not only commonly found in ringed salamander, but also spreads widely in the family Ambystomatidae. Rhabditid nematodes (Rhabdias ranae) are the second most common parasite that infests A. annulatum's lungs and body cavities. Other relatively rare parasites include gall bladder myxosporean (Myxidium serotinum) and tissue-dwelling spirurids that are usually en-cysted in ringed salamanders' stomach walls.[15]

References

  1. ^ a b IUCN SSC Amphibian Specialist Group (2014). "Ambystoma annulatum". The IUCN Red List of Threatened Species. IUCN. 2014: e.T59052A56219662. doi:10.2305/IUCN.UK.2014-1.RLTS.T59052A56219662.en. Retrieved 4 January 2018.
  2. ^ a b c d Petranka, James W. (2010). Salamanders of the United States and Canada. Washington, D.C.: Smithsonian Institution Press. ISBN 9781588343086.
  3. ^ a b c d e Crane, Adam L., et al. "Learning to find food: evidence for embryonic sensitization and juvenile social learning in a salamander." Animal Behaviour 142 (2018): 199-206.
  4. ^ a b c d Mathis, Alicia, Kevin L. Murray, and Caleb R. Hickman. "Do experience and body size play a role in responses of larval ringed salamanders, Ambystoma annulatum, to predator kairomones? Laboratory and field assays." Ethology 109.2 (2003): 159-170.
  5. ^ a b "AmphibiaWeb - Ambystoma annulatum". amphibiaweb.org. Retrieved 2022-11-13.
  6. ^ a b Hutcherson, Jackie E.; Peterson, Chris L.; Wilkinson, Robert F. (June 1989). "Reproductive and Larval Biology of Ambystoma annulatum". Journal of Herpetology. 23 (2): 181. doi:10.2307/1564028. ISSN 0022-1511. JSTOR 1564028.
  7. ^ Bishop, Sherman C. (1962). Handbook of Salamanders. New York: Hafner Publishing Company.
  8. ^ T., T. (1977). The Amphibians of Missouri. Lawrence, KS: University of Kansas Publications.
  9. ^ a b c d Spotila, James R.; Beumer, Ronald J. (July 1970). "The Breeding Habits of the Ringed Salamander, Ambystoma annulatum (Cope), in NorthWestern Arkansas". American Midland Naturalist. 84 (1): 77. doi:10.2307/2423728. ISSN 0003-0031. JSTOR 2423728.
  10. ^ a b c d e f g h i j k Semlitsch, Raymond D.; Anderson, Thomas L.; Osbourn, Michael S.; Ousterhout, Brittany H. (2014-03-01). "Structure and Dynamics of Ringed Salamander (Ambystoma annulatum) Populations in Missouri". Herpetologica. 70 (1): 14. doi:10.1655/herpetologica-d-13-00074. ISSN 0018-0831. S2CID 85776338.
  11. ^ a b c d Crawford, John A.; Tunnage, Joshua A.; Wright, Eric M. (July 2017). "Breeding Pond Occupancy of the Ringed Salamander (Ambystoma annulatum) in East-central Missouri". The American Midland Naturalist. 178 (1): 151–157. doi:10.1674/0003-0031-178.1.151. ISSN 0003-0031. S2CID 90145558.
  12. ^ Ousterhout, Brittany Hunter (2016). Effects of phenotype- and condition-dependent factors on juvenile dispersal of the ringed salamander (Ambystoma annulatum) (Thesis). University of Missouri Libraries. doi:10.32469/10355/60415.
  13. ^ B., Semlitsch, Raymond D. Conner, Christopher A. Hocking, Daniel J. Rittenhouse, Tracy A. G. Harper, Elizabeth. Effects of timber harvesting on pond-breeding amphibian persistence : testing the evacuation hypothesis. OCLC 1231915862.
  14. ^ a b Burkhart, Jacob Jonathon. Genetic diversity and distribution of the ringed salamander (Ambystoma annulatum) across multiple spatial scales. Diss. University of Missouri--Columbia, 2018.
  15. ^ a b McAllister, Chris T., Stanley E. Trauth, and Betty G. Cochran. "Endoparasites of the ringed salamander, Ambystoma annulatum (Caudata: Ambystomatidae), from Arkansas." The Southwestern Naturalist 40.3 (1995): 327-330.
  16. ^ a b Nyman, Stephen, Robert F. Wilkinson, and Jackie E. Hutcherson. "Cannibalism and size relations in a cohort of larval ringed salamanders (Ambystoma annulatum)." Journal of Herpetology (1993): 78-84.
  17. ^ a b c d Crane, Adam, Alicia Mathis, and Carly McGrane. "Socially facilitated antipredator behavior by ringed salamanders (Ambystoma annulatum)." Behavioral Ecology and Sociobiology 66.5 (2012): 811-817.
  18. ^ Nyman, Stephen; Wilkinson, Robert F.; Hutcherson, Jackie E. (1993). "Cannibalism and Size Relations in a Cohort of Larval Ringed Salamanders (Ambystoma annulatum)". Journal of Herpetology. 27 (1): 78–84. doi:10.2307/1564909. ISSN 0022-1511. JSTOR 1564909.
  19. ^ a b Peterson, Chris L., et al. "Premetamorphic survival of Ambystoma annulatum." Herpetologica (1991): 96-100.
  20. ^ a b Young, Henry E.; Bailey, Claudia F.; Dalley, Bernell K. (July 1983). "Envirnmental conditions prerequisite for complete limb regeneration in the postmetamorphic adult land-phase salamander,Ambystoma". The Anatomical Record. 206 (3): 289–294. doi:10.1002/ar.1092060307. ISSN 0003-276X. PMID 6614511. S2CID 24854488.
  21. ^ Young, Henry E.; Bailey, Claudia F.; Dalley, Bernell K. (July 1983). "Gross morphological analysis of limb regeneration in postmetamorphic adultAmbystoma". The Anatomical Record. 206 (3): 295–306. doi:10.1002/ar.1092060308. ISSN 0003-276X. PMID 6614512. S2CID 46190433.
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Ringed salamander: Brief Summary

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The ringed salamander (Ambystoma annulatum) is a species of mole salamander native to hardwood and mixed hardwood-pine forested areas in and around the Ozark Plateau and Ouachita Mountains of Arkansas, Oklahoma, and Missouri. This species of salamander has slander body, small head, and long tail. They are usually found to have various dorsal color from dark gray to dark brown. Various close relatives are found such as Ringed salamander (A. annulatum), marbled salamander (A. opacum), and spotted salamander (A. maculatum). This species of salamander has cannibal behavior especially those in large body size.

It is found in damp, forested areas, usually under leaves, rotting logs, or in abandoned ground holes of other organisms, near shallow ponds. Highly fossorial (adapted to digging), adults are often found in subterranean refuges. This salamander is increasingly rare and perhaps endangered. This is likely a result of its restricted range and specific breeding habit needs. The world population is thought to be around 100,000 animals. Its conservation status is assessed as Least Concern by the IUCN. This salamander can perform embryotic learning and social-facilitated learning.

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