Cow Knob salamander | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Amphibia |
Order: | Urodela |
Family: | Plethodontidae |
Subfamily: | Plethodontinae |
Genus: | Plethodon |
Species: | P. punctatus |
Binomial name | |
Plethodon punctatus Highton, 1971 [1972] | |
Plethodon punctatus, commonly known as the Cow Knob salamander [3] [4] [5] [6] or white-spotted salamander [7] [8] [9] (not to be confused with P. cylindraceus, the white-spotted slimy salamander), is a species of salamander in the family Plethodontidae. It is endemic to high mountain forests on the border of Virginia and West Virginia in the United States. Nearly all occurrences are on Shenandoah Mountain, Nathaniel Mountain and Great North Mountain in George Washington National Forest. Cow Knob salamanders are a member of the P. wehrlei species complex, which includes many other Appalachian salamanders historically referred to Plethodon wehrlei (Wehrle's salamander).
Major threats to Plethodon punctatus include habitat loss and climate change, [10] [11] and it is rated as Near Threatened by the IUCN. Although not protected by the Endangered Species Act, Cow Knob salamanders benefit from a 1994 conservation agreement between the U.S. Forest Service and the U.S. Fish and Wildlife Service. The species' threatened condition has helped to secure ecosystem conservation goals on Shenandoah Mountain and nearby ridges, such as redirecting the planned route of the Atlantic Coast Pipeline. Despite their restricted range, Cow Knob salamanders are abundant in localities with ideal environmental conditions, such as rocky old-growth hemlock forests.
Plethodon punctatus was named as a new species by Richard Highton in 1972. The holotype was recovered from the slope of Cow Knob, a peak on Shenandoah Mountain. Cow Knob lends its name to the common name of the species. With the use of genetic data, P. punctatus was immediately recognized as a genetically detached lineage descended from ancestral populations of the Wehrle’s salamander (P. wehrlei). [9]
Plethodon punctatus itself is divided into two populations, one occupying sites south of Reddish Knob, and the other north of Reddish Knob. [12] The species as a whole diverged from P. wehrlei between the Late Pliocene (2.84 Ma) and mid-Pleistocene (1.04 Ma), while the two populations diverged from each other in the mid- to late-Pleistocene (1.30 to 0.40 Ma). The northern population has much lower genetic diversity compared to the southern population, despite inhabiting a larger area in the present day. It is probable that the ancestral (southern) population once occupied the entire range, only for the species's suitable climate to abruptly shift south with the arrival of glacial intervals in the Pleistocene. Most of the ancestral population dispersed south and continued to occupy a broad area south of their modern-day range. The few surviving relicts which refused to move would have experienced a population bottleneck and the founder effect, diverging into a new population. With warm conditions returning in the Holocene, the southern population repopulated part of their ancestral range while the northern population quickly expanded out of its isolated refuge. It is unclear whether the two populations have regained contact at Reddish Knob. [12]
Long considered one of two valid species in the P. wehrlei group, P. punctatus has been a major player into the debate over species differentiation in that species complex. [13] [14] [15] A 2012 genetic analysis confirmed that some West Virginian populations referred to P. wehrlei were actually more closely related to P. punctatus than to other populations from southern Virginia. [13] This would render P. wehrlei a paraphyletic species, a problem which can be resolved either by lumping P. punctatus into P. wehrlei or by splitting the broader species into multiple smaller species. [14] Some of these new species have already been named ( P. dixi , [14] P. jacksoni , [14] P. pauleyi [15] ) while others are still tentatively referred to P. wehrlei. The “Northern wehrlei” clade (ranging from central West Virginia up to New York) is the closest to P. punctatus, both genetically and geographically, among all the clades historically lumped into P. wehrlei. P. pauleyi (of the Cumberland Plateau, previously the "yellow-spotted wehrlei" clade) is the next most closely related. [14] [15]
The Cow Knob salamander is a moderately large plethodontid. The largest recorded length is 17.1 cm (6.7 in), [8] [4] though most adults have a total length smaller than 15.7 cm (6.2 in). Around half of the total length is tail. [4] The base coloration is dark brown or greyish-black. The sides and legs are colored with large white or cream spots, which may extend onto a portion of the belly. The back is typically speckled with smaller and more scattered spots. [7] The species name ‘punctatus’ (“spotted”) is in reference to these dorsal spots. [7] [8] The throat has a light pink or pale coloration which transitions to the grey belly. [4]
The white-spotted slimy salamander (Plethodon cylindraceus) is similar in size and color, but differs in many other respects. Plethodon punctatus can differentiated by its more slender build, slightly webbed toes, protuberant eyes, a higher number of costal grooves (usually 17–18), and a lighter throat. [4] It is very similar in appearance to closely-related salamanders in the Plethodon wehrlei group, though it is slightly larger and completely lacks distinctive coloration patterns such as red spots, brassy flecks, or large yellow dorsal spots. [7] [8] [4] [14]
Cow Knob salamanders are adapted to live in high-elevation hardwood forests with dense canopy cover and talus. Old-growth forests and hemlock stands are greatly preferred, while young pine monocultures and clearings are avoided. [3] [5] They typically occur at an elevation of 900–1,200 m (3,000–3,900 ft), [4] with some reports as low as 735 m (2,411 ft). [3] They are particularly abundant on north-facing slopes with an ample supply of rocks for cover. [3] [4] [5] Rocks with large gaps underneath are the preferred form of refuge, followed by fallen logs. [5]
The salamanders are most active in the spring (April–June) and fall (September–October), sheltering during the harsher months of the summer and winter. [4] [9] In West Virginia, their peak season extends into the early summer. [5] They forage on cool wet nights, prioritizing the forest floor but also venturing onto trees or rocks. [16] A wide range of small invertebrates are taken as prey. [17] [7] [4] [5] [9] Plethodon punctatus is a host to Cepedietta michiganensis (a ciliate) and Batracholandros magnavulvaris (a nematode), both of which are fairly benign gut parasites common among Appalachian salamanders. [5]
Cow Knob salamanders are direct developers, mating via spermatophore in the late winter or early spring. A clutch of 7-16 eggs can be laid every other year, though the precise timing is unknown. Juveniles hatch in the fall and reach sexual maturity within 3 or 4 years, with females maturing later and at a larger body size. [5] This species is not migratory on an individual basis, and most adults have a range of only a few meters throughout their lifetime.
Plethodon punctatus coexists alongside the ecologically similar Shenandoah Mountain salamander (Plethodon virginia, previously referred to Plethodon hoffmani ). The two species do not show any competitive exclusion, despite P. punctatus juveniles overlapping the size range of P. virginia adults. To reduce competition, the two species may be active at different times upon the arrival of ideal environmental conditions. P. punctatus tend to find refuge in shallow rockier soils, which may be more sensitive to surface temperature and humidity levels compared to the deep moist soils preferred by P. virginia. [17]
Plethodon punctatus occurs entirely within the protected area of George Washington National Forest, though some pressures still threaten the species. As the salamander is reliant on stable mountain forest ecosystems, the largest threat is logging within the forest, especially during peak salamander activity periods. To prevent population decline, logging should be restricted to discontinuous patches, road building should be minimized or eliminated, and replanting should emphasize trees other than white pine. [3] Another concern is defoliation from invasive insects such as woolly adelgid (Adelges tsugae) and spongy moth (Lymantria dispar). [4] [18]
Plethodon punctatus has a particularly low heat tolerance among Appalachian salamanders. [19] As a montane species with limited routes of dispersal, it is particularly vulnerable to the effects of climate change. [10] [20] [11] At intermediate rates of warming (RCP 4.5), 93 [11] –100% [10] [20] of suitable habitat is expected to be lost by 2050. Some shaded slopes may retain a suitable microclimate, but these refugia would be isolated from each other, leading to an unsustainable decrease in genetic diversity. In addition, climate change may bring Plethodon punctatus into greater competition with Plethodon cylindraceus , an aggressive species which is more dominant in warm areas at lower elevation. [11]
The IUCN Red List has listed Plethodon punctatus as a Near Threatened species since 2004. [1] NatureServe lists the species as vulnerable (G3) at a national level, and imperiled (S2) in both states considered separately. [2] The Virginia Department of Wildlife Resources classifies Plethodon punctatus as a tier Ic Species of Greatest Conservation Need. This indicates extreme risk of extirpation alongside limited potential for state conservation actions or research (due to most of its range being federally managed). [21] A 2012 petition [22] to grant federal protection to the species under the Endangered Species Act was rejected by the United States Fish and Wildlife Service (USFWS) in 2016. [23]
George Washington National Forest is managed by the United States Forest Service (USFS), providing a base level of federal protection from private enterprises. The Cow Knob salamander is an example of a Management Indicator Species (MIS), a type of animal used to evaluate and implement broader ecosystem conservation policies. [24] In 1994, the USFS and USFWS entered into a conservation agreement to maintain the species’ viability and prevent the need to list it as an endangered species. [25] [24] [26] [27] [28] Approximately 58,000 acres of salamander habitat at the crest of Shenandoah Mountain are protected from timber harvesting and road construction under this agreement. [24] Much of the protected land also overlaps with wilderness area protections and the proposed Shenandoah Mountain National Scenic Area. [24]
The Atlantic Coast Pipeline, a planned natural gas pipeline primarily funded by Dominion Energy, was initially designed to pass over Shenandoah Mountain along its route. By late October 2015, USFS concerns over the preservation of salamander habitat led the developers to propose horizontal directional drilling under the mountain. [29] In January 2016, the pipeline was forced to find a new route entirely, as even the revised construction plan would have adversely impacted wildlife. The Cow Knob salamander was the main species considered in the decision, alongside Cheat Mountain salamanders (Plethodon nettingi) and West Virginia northern flying squirrels (Glacuomys sabrinus fuscus). [30] [31] [32] [33]
Plethodon is a genus of salamanders in the family Plethodontidae. They are commonly known as woodland salamanders. All members of the genus are endemic to North America. They have no aquatic larval stage. In some species, such as the red-backed salamander. Young hatch in the adult form. Members of Plethodon primarily eat small invertebrates. The earliest known fossils of this genus are from the Hemphillian of Tennessee in the United States.
The red-backed salamander is a small, hardy woodland salamander species in the family Plethodontidae. It is also known as the redback salamander, eastern red-backed salamander, or the northern red-backed salamander to distinguish it from the southern red-backed salamander. The species inhabits wooded slopes in eastern North America, west to Missouri, south to North Carolina, and north from southern Quebec and the Maritime provinces in Canada to Minnesota. It is one of 56 species in the genus Plethodon. Red-backed salamanders are notable for their color polymorphism and primarily display two color morph varieties, which differ in physiology and anti-predator behavior.
Wehrle's salamander is a species of salamander in the family Plethodontidae. It is endemic to the Eastern United States. It is named in honor of Richard White Wehrle (1852–1937), a jeweler, naturalist, and collector of the holotype.
The Cheat Mountain salamander is a species of small woodland salamander in the family Plethodontidae. The species is found only on Cheat Mountain, and a few nearby mountains, in the eastern highlands of West Virginia. It and the West Virginia spring salamander are the only vertebrate species with geographic ranges restricted to that state.
The white-spotted slimy salamander is a species of salamander in the family Plethodontidae endemic to the Eastern United States. It is one of 55 species in the genus Plethodon, and was one of the first to be described of its cogeners. The preferred habitat of this species is under logs and leaf litter in shaded hardwood forests and wooded floodplains, and often forages on the forest floor on wet nights. It was found that with increasing temperatures, the aggression in this species also increases. In the plethodon genus, species have a lungless morphology, restricting nearly all gas and water exchange transport to the body surface. This species mainly consumes insects, including ants, centipedes, springtails, crickets, millipedes, slugs, snout-beetles, and earthworms. Common predators of this species are gartersnakes, copperheads, and birds. One of their predator defense mechanisms is the release of noxious/sticky substances through the skin by the dorsal granular glands. Another predator deterrent is when touched, this species will freeze in place and become immobile. This species of Plethodon are mostly terrestrial and deposit their direct-developing eggs on land that omits the aquatic larval stage characteristic of most amphibians, therefore this species is not restricted to aquatic habitats for reproduction and dispersal. This species, along with other Plethodontid salamanders, are frequently parasitized by Trombicula mites.
The northern ravine salamander is a species of salamander in the family Plethodontidae. The species is endemic to the United States.
The Peaks of Otter salamander is a species of salamanders in the family Plethodontidae. It is endemic to the Peaks of Otter area in the Blue Ridge Mountains of Virginia. It is a montane salamander found at elevations above 442 m (1,450 ft), but more commonly above 760 m (2,490 ft). It can be locally common, but its distribution is small and patchy. This makes it vulnerable to local threats such as timber harvesting, recreational development, defoliation by gypsy moths, and spraying to control the latter.
The Kiamichi slimy salamander is a species of salamander in the family Plethodontidae endemic to the United States, has a natural habitat of temperate forests, and is found over a small range. This nocturnal species is mainly threatened by habitat loss and was first described by Highton in 1989. It is rated as a vulnerable species by the International Union for Conservation of Nature.
The Pigeon Mountain salamander is a species of salamander in the family Plethodontidae. It is endemic to Pigeon Mountain in the US state of Georgia.
The Shenandoah salamander is a small, terrestrial salamander found exclusively in Shenandoah National Park in Virginia. The Shenandoah salamander inhabits a very small range of land on just three mountain peaks. Due to the small habitat range, interspecies competition, and climate change, the population of the Shenandoah salamander is vulnerable to extinction. Mitigating human effects on the habitat of the species will be essential in attempting to preserve and grow the population.
The Big Levels salamander is a species of salamander in the family Plethodontidae. It is endemic to Virginia in the eastern United States. First described in 2004, it derives its specific name, sherando, from Sherando Lake in the George Washington National Forest. Its common name refers to the Big Levels area of southeastern Augusta County, Virginia, a series of flat to gently rolling mountain tops in the Blue Ridge Mountains, where it was found.
The Shenandoah Mountain salamander is a species of salamander in the family Plethodontidae native to the eastern United States. It should not be confused with the Shenandoah salamander, which inhabits Shenandoah National Park, east of Shenandoah Mountain.
Weller's salamander is a species of salamander in the family Plethodontidae. This species in endemic to the southeastern mountain range of the United States. It is mainly found in North Carolina near Grandfather Mountain. The salamanders have a unique metallic spotting which distinguishes them from other Plethodon species and other salamanders in the area. They mainly inhabit cool forests with rocky areas.
The green salamander is a species of lungless salamander in the family Plethodontidae. It and the Hickory Nut Gorge green salamander are the only currently-described members of the genus Aneides that inhabit any areas in the eastern half of United States. Rarely seen in the field, the green salamander is an extremely habitat-specific species that is seldom found away from its preferred surroundings: moist, shaded rock crevices. Green salamanders have one of the most specialized niches of any salamander.
The Dixie Caverns salamander is a species of salamander in the family Plethodontidae. It is endemic to the state of Virginia in the United States. It is most often found in Roanoke County, but several specimens have been recorded in surrounding counties. It was commonly confused with Wehrle's salamander and was merged with that species shortly after description, but a 2019 study reaffirmed it as a distinct species.
The Blacksburg salamander is a species of salamander in the family Plethodontidae. It is endemic to the Southeastern United States, where it is restricted to a portion of the Appalachian Mountains in southwestern Virginia. Its common name refers the town of Blacksburg, Virginia, as many specimens were initially found in the vicinity of the town.
The yellow-spotted woodland salamander is a species of salamander in the family Plethodontidae. It is endemic to the United States, where it is distributed throughout the Cumberland Plateau in the states of Kentucky and Tennessee. Its natural habitat is temperate forest. It was long considered to be both an isolated western population and a unique yellow-spotted color morph of the Wehrle's salamander, but a study published in 2019 found it to be a distinct species.