Spotted salamander | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Amphibia |
Order: | Urodela |
Family: | Ambystomatidae |
Genus: | Ambystoma |
Species: | A. maculatum |
Binomial name | |
Ambystoma maculatum (Shaw, 1802) | |
Range of A. maculatum | |
Synonyms [2] | |
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The spotted salamander or yellow-spotted salamander (Ambystoma maculatum) is a mole salamander [2] common in eastern United States and Canada. [1] It is the state amphibian of Ohio and South Carolina. The species ranges from Nova Scotia, to Lake Superior, to southern Georgia and Texas. [3] Its embryos have been found to have symbiotic algae living in and around them, [4] the only known example of vertebrate cells hosting an endosymbiont microbe (unless mitochondria are considered). [5] [6]
The spotted salamander is about 15–25 cm (5.9–9.8 in) long, [7] with females generally being larger than males. [8] They are stout, like most mole salamanders, and have wide snouts. [3] The spotted salamander's main color is black, but can sometimes be a blueish-black, dark gray, dark green, or even dark brown. Two uneven rows of yellowish-orange spots run from the top of the head (near the eyes) to the tip of the tail (dorso-lateral ranging). [9] The spotted salamander's spots near the top of its head are more orange, while the spots on the rest of its body are more yellow. The underside of the spotted salamander is slate gray and pink. Sexual dimorphism, physical differences between males and females, is displayed in the form of larger-bodied females having brighter-coloured spots. [10] Males will have a larger portion of their dorsal surface covered in spots that are less bright. There is also a correlation between body condition and spots having lower chroma. [10]
The scientific name Ambystoma maculatum comes from Ambystoma– amblys (Greek) for blunt; -stoma (Greek) meaning mouth; or anabystoma (New Latin) meaning ‘to cram into the mouth’ maculatum – macula (Latin) for spot; maculosus (Latin) for spotted. [11]
They developed larger tail fins and bodies to compete with other predators. [9] They typically live around 20 years, but some have lived up to 30 years. ("Spotted Salamander")
The spotted salamander usually lives in mature forests with ponds or ephemeral vernal pools for breeding sites. [12] Vernal pools are suitable breeding sites for these amphibians as they dry often enough to exclude fish that eat the salamander eggs and larvae, while retaining water long enough to allow amphibian larvae to complete development and metamorphose into terrestrial adults. [13] A study showed larger pools (as opposed to smaller pools) had more egg masses, higher occupancy, and higher larval survival rates for spotted salamanders. [14] Outside of the breeding season, these salamanders spend their time in forests with well-drained soils that contain many burrows dug by small mammals. [8] They have also been known to be absent from landscapes and forest with canopy cover below 30%. [15]
Salamander populations from nearby pools form genetically-distinct metapopulations. Subpopulations within 4.8 kilometers share a higher proportion of genes, while populations greater than 4.8 kilometers share a smaller proportion of genes. Inter-population dispersal is likely mediated by both species-specific behaviors and natural limitations. [16]
Spotted salamanders are fossorial, meaning they spend most of their time underground. They rarely come above ground, except after a rain or for foraging and breeding. During the winter, they brumate underground, and are not seen again until breeding season in early March–May. [17]
In North Amherst, Massachusetts, United States Spotted salamanders cross the street to get from their overwintering sites in the wooded area east of the roadway to their breeding sites to the west. Local officials built amphibian and reptile tunnels called the Henry Street salamander tunnels to help the salamanders cross the road to get to vernal pools. [18] In the spring after rain and when temperatures rise above 40 °F (4 °C), salamanders emerge from underground. They cross Henry Street to get to their breeding grounds, vernal pools which form on the other side of the road, an event common among amphibians and known as the Big Night. [19]
Ambystoma maculatum has several methods of defense, including hiding in burrows or leaf litter, autotomy of the tail, and a toxic milky liquid it excretes when perturbed. This secretion comes from large poison glands around the back and neck. The spotted salamander, like other salamanders, shows great regenerative abilities: if a predator manages to dismember a part of a leg, tail, or even parts of the brain, head, or organs, the salamander can grow back a new one, although this takes a massive amount of energy. [20] As juveniles, they spend most of their time under the leaf litter near the bottom of the pools where their eggs were laid. The larvae tend to occupy refuges in vegetation, and lower their activity in the presence of predators. [21]
Ambystoma maculatum tend to follow the same path in their migration to and from their burrows and breeding pools. [22] They accomplish their journey in conditions that lack visual cues, since it is usually during periods of cloud cover. Some studies show evidence of landmark learning in spotted salamanders. Researchers found that spotted salamanders can associate visual landmarks with food. Thus spotted salamanders may learn landmarks in their habitat that are reliable indicators of resource locations or provide orientation clues for migration to and from breeding ponds. [23]
Male salamanders come out earlier than females due to different responses in temperature than females. [24] According to a study there is a correlation between the salamander's spot coloration and body condition. Salamanders' in lower body condition had less color and those with a better body score had brighter colored spots. [10]
Spotted salamanders feed on earthworms, slugs, snails, spiders, millipedes, centipedes, insects, algae and other invertebrates. [25] [26] The spotted salamander larvae have been known to cannibalize others when food is scarce. [27] They sometimes also feed on smaller salamanders, such as the red-backed salamander. [28] The adult spotted salamander uses its sticky tongue to catch food. [28]
During the majority of the year, spotted salamanders live in the shelter of leaves or burrows in deciduous forests. However, when the temperature rises and the moisture level is high, the salamanders make their abrupt migration towards their annual breeding ponds. Recent studies, however, indicate that temperature may be a more important factor than precipitation, as precipitation in winter months increase, yet salamanders do not migrate. [24] In just one night, hundreds to thousands of salamanders may make the trip to their ponds for mating. [17] Males will start a dance-like behavior called a liebsspiel, where afterward they lay down their spermatophore. This dance includes the salamanders circling around one another and putting their head's on one another's tail. Then, the male will swim away wiggling his tail. If the female wants to mate she will follow him to a sperm pack he made earlier. ("Spotted Salamander Ambystoma Maculatum") [29] Males will migrate at higher rates than females early in the migration season. This could be due to different responses to temperature between males and females. [24] Mates usually breed in ponds when it is raining in the spring. Females usually lay about 100 eggs in one clutch that cling to the underwater plants and form egg masses.
The egg masses are round, jelly-like clumps that are usually 6.4–10.2 cm (2.5–4 in) long. The spotted salamander produces a unique polymorphism in the outer jelly layers of its egg masses: one morph has a clear appearance and contains a water-soluble protein, whereas the other morph is white and contains a crystalline hydrophobic protein. [30] [31] This polymorphism is thought to confer advantages in vernal pools with varying dissolved nutrient levels, while also reducing mortality from feeding by wood frog larvae. [32] [33]
Adults only stay in the water for a few days, then the eggs hatch in one to two months. When the eggs hatch depends on the water temperatures. Eggs of A. maculatum can have a symbiotic relationship with the green alga Chlorococcum amblystomatis . [34] A dense gelatinous matrix surrounds the eggs and prevents the eggs from drying out, but it inhibits oxygen diffusion (required for embryo development). [35] C. amblystomatis provides increased oxygen and supplemental nutrition from fixed carbon products via photosynthesis [36] and removes the embryo's nitrogenous waste (ammonia) in the egg capsule, aiding in the salamander's embryonic development and growth. [37] The developing salamander thus metabolizes the oxygen, producing carbon dioxide (which then the alga consumes). Photosynthetic algae are present within the egg capsule of the developing salamander embryo, enhancing growth. However, the widely used herbicide, atrazine, has been found to significantly lower hatching success rate by eliminating the symbiotic algae associated with the egg masses. [38]
As larvae, they are usually light brown or greenish-yellow. They have small dark spots and are born with external gills. In two to four months, the larvae lose their gills, and become juvenile salamanders that leave the water. Spotted salamanders have been known to live up to 32 years, [39] and normally return to the same vernal pool every year. These pools are seasonal and will usually dry up during the late spring and stay dry until winter.
Spotted salamanders are often preyed on by raccoons, skunks, turtles, and snakes. For this reason they perform an important function by connecting the food chain. [8] Moreover, one of their predators is humans since they are popularly [40] sought out through the pet trade.
Amphibians are ectothermic, anamniotic, four-limbed vertebrate animals that constitute the class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all tetrapods excluding the amniotes. All extant (living) amphibians belong to the monophyletic subclass Lissamphibia, with three living orders: Anura, Urodela (salamanders), and Gymnophiona (caecilians). Evolved to be mostly semiaquatic, amphibians have adapted to inhabit a wide variety of habitats, with most species living in freshwater, wetland or terrestrial ecosystems. Their life cycle typically starts out as aquatic larvae with gills known as tadpoles, but some species have developed behavioural adaptations to bypass this.
The mole salamanders are a group of advanced salamanders endemic to North America. The group has become famous due to the study of the axolotl in research on paedomorphosis, and the tiger salamander which is often sold as a pet, and is the official amphibian of four US states.
The tiger salamander is a species of mole salamander and one of the largest terrestrial salamanders in North America.
The California tiger salamander is a vulnerable amphibian native to California. It is a mole salamander. Previously considered to be a subspecies of the tiger salamander, the California tiger salamander was recently designated a separate species again. The California tiger salamander distinct population segment (DPS) in Sonoma County and the Santa Barbara County DPS are listed as federally endangered, while the Central California DPS is listed as federally threatened. The Sonoma County, south San Joaquin, and the Santa Barbara County DPS have diverged from the rest of the California tiger salamander populations for over one million years, since the Pleistocene and they may warrant status as separate species.
Vernal pools, also called vernal ponds or ephemeral pools, are seasonal pools of water that provide habitat for distinctive plants and animals. They are considered to be a distinctive type of wetland usually devoid of fish, and thus allow the safe development of natal amphibian and insect species unable to withstand competition or predation by fish. Certain tropical fish lineages have however adapted to this habitat specifically.
The Jefferson salamander is a mole salamander native to the northeastern United States, southern and central Ontario, and southwestern Quebec. It was named after Jefferson College in Pennsylvania.
Eastern newts are 2-4'inch long in length. These animals are common aquarium pets, being either collected from the wild or sold commercially. The striking bright orange juvenile stage, which is land-dwelling, is known as a red eft. Some sources blend the general name of the species and that of the red-spotted newt subspecies into the eastern red-spotted newt.
Lithobates sylvaticus or Rana sylvatica, commonly known as the wood frog, is a frog species that has a broad distribution over North America, extending from the boreal forest of the north to the southern Appalachians, with several notable disjunct populations including lowland eastern North Carolina. The wood frog has garnered attention from biologists because of its freeze tolerance, relatively great degree of terrestrialism, interesting habitat associations, and relatively long-range movements.
The blue-spotted salamander is a mole salamander native to the Great Lakes states and northeastern United States, and parts of Ontario and Quebec in Canada. Their range is known to extend to James Bay to the north, and southeastern Manitoba to the west.
The ringed salamander 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 marbled salamander and spotted salamander. This species of salamander has cannibal behavior especially those in large body size.
The streamside salamander is a species of mole salamander from North America, occurring in several Midwestern states of the US.
The long-toed salamander is a mole salamander in the family Ambystomatidae. This species, typically 4.1–8.9 cm (1.6–3.5 in) long when mature, is characterized by its mottled black, brown, and yellow pigmentation, and its long outer fourth toe on the hind limbs. Analysis of fossil records, genetics, and biogeography suggest A. macrodactylum and A. laterale are descended from a common ancestor that gained access to the western Cordillera with the loss of the mid-continental seaway toward the Paleocene.
The Santa Cruz long-toed salamander is an endangered subspecies of the long-toed salamander, which is found only close to a few isolated ponds in Santa Cruz and Monterey Counties in California. It has a black body, broken yellow or orange irregular striping along its spine, and a tail fin well evolved for swimming. Like other mole salamanders, it is found near pools or slow-moving streams and has a very secretive lifestyle, making it difficult to find.
The marbled salamander is a species of mole salamander found in the eastern United States.
The blunt-headed salamander is a mole salamander endemic to Mexico. It is only known from the vicinity of its type locality, near Morelia, in Michoacán state in Southwestern Mexico. It inhabits a landscape consisting of a mosaic of natural grasslands and pine-oak forests at elevations of about 2,000 m (6,600 ft) asl. Breeding takes place in ponds. An average adult has a mass of 6.18 grams while wet. Adult females of the species range from 42-93 mm and males range from 45.4-70.5 mm in standard length.
The small-mouth salamander is a species of mole salamander found in the central United States, from the Great Lakes region in Michigan to Nebraska, south to Texas, and east to Tennessee, with a population in Canada, in Pelee, Ontario. It is sometimes referred to as the Texas salamander, porphyry salamander, or the narrow-mouthed salamander. The Kelley's Island salamander was synonymized with A. texanum in 1995.
The silvery salamander is a hybrid species of mole salamander from the United States of America and Canada. It is usually between 5.5–7.75 in (14.0–19.7 cm) long and slender, with many small silvery-blue spots on its back and sides. It is brownish grey, and the area around its vent is grey. A unisexual Ambystoma hybrid species, A. platineum has been grouped with other unisexual ambystomatids that take genetic material from Jefferson salamanders, streamside salamanders, small-mouthed salamanders, tiger salamanders and the blue-spotted salamander.
Chlorococcum amblystomatis,, is a species of single-celled green algae known for its symbiotic relationship with the spotted salamander, Ambystoma maculatum. It grows symbiotically inside salamander eggs, primarily in the eggs of the spotted salamander, Ambystoma maculatum. It has also been reported in other salamander species, such as the Japanese black salamander, Hynobius nigrescens, which is endemic to Japan.
The Henry Street salamander tunnels are two amphibian tunnels in Amherst, Massachusetts, United States, built in 1987 to assist salamander migration between their overwintering grounds and their breeding grounds, an annual courtship event known as the Big Night. Previously, spotted salamanders were often crushed by vehicles as they crossed Henry Street. They winter on its east side and cross westward in the spring to breed in what is known as a salamander congress in the vernal pools that form there.
Salamander congress is a salamander mating ritual which occurs in spring on the Big Night. It is a gathering of salamanders in vernal pools for the purpose of breeding. When male and female salamanders meet in the vernal pools it is called a congress.