Cow Knob salamander

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Cow Knob salamander
Plethodon punctatus 02 white background 2.jpg
Status TNC G3.svg
Vulnerable  (NatureServe) [2]
Scientific classification OOjs UI icon edit-ltr.svg
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).

Contents

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.

Taxonomy

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]

Landscape at Reddish Knob, a peak on Shenandoah Mountain which delineates the two populations of Cow Knob Salamander Reddish Knob.jpg
Landscape at Reddish Knob, a peak on Shenandoah Mountain which delineates the two populations of Cow Knob Salamander

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]

Description

Plethodon punctatus as seen from below, showing the light throat coloration and grey belly Plethodon punctatus 01 underside.jpg
Plethodon punctatus as seen from below, showing the light throat coloration and grey belly

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]

Habitat

Fall colors in mixed forest on Shenandoah Mountain 2019-10-26 12 51 47 View southwest across Devrick Hollow on the west side of Shenandoah Mountain from the Highland Turnpike (U.S. Route 250) in Highland County, Virginia.jpg
Fall colors in mixed forest on Shenandoah Mountain

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]

Ecology and behavior

Seen in the wild in Pendleton County, West Virginia Plethodon punctatus 101120592.jpg
Seen in the wild in Pendleton County, West Virginia

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]

Conservation

Threats

Cow Knob salamanders are threatened by the loss of hemlock forest and other suitable habitats via logging or invasive pests (such as woolly adelgid, pictured) Adelges tsugae 1276001.jpg
Cow Knob salamanders are threatened by the loss of hemlock forest and other suitable habitats via logging or invasive pests (such as woolly adelgid, pictured)

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]

Assessments and protections

Map of national forest land in Virginia Va nationalforests.png
Map of national forest land in Virginia

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]

Related Research Articles

<i>Plethodon</i> Genus of amphibians

Plethodon is a genus of salamanders in the family Plethodontidae. They are also known as woodland salamanders or, more rarely, slimy salamanders. All members of the genus are endemic to North America. They have no aquatic larval stage. In some species, such as Plethodon cinereus, the red-backed salamander, eggs are laid underneath a stone or log. 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.

<span class="mw-page-title-main">Red-backed salamander</span> Species of amphibian

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.

<span class="mw-page-title-main">Wehrle's salamander</span> Species of amphibian

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.

<span class="mw-page-title-main">Northern slimy salamander</span> Species of amphibian

The northern slimy salamander is a species of terrestrial plethodontid salamander found throughout much of the eastern two-thirds of the United States.

<span class="mw-page-title-main">Cheat Mountain salamander</span> Species of amphibian

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 northern ravine salamander is a species of salamander in the family Plethodontidae. The species is endemic to the United States.

<span class="mw-page-title-main">Peaks of Otter salamander</span> Species of amphibian

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.

<span class="mw-page-title-main">Pigeon Mountain salamander</span> Species of amphibian

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.

<span class="mw-page-title-main">Shenandoah salamander</span> Species of amphibian

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.

<span class="mw-page-title-main">Big Levels salamander</span> Species of amphibian

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.

<span class="mw-page-title-main">Weller's salamander</span> Species of amphibian

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.

<span class="mw-page-title-main">Amphibians and reptiles of Mount Rainier National Park</span>

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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 the Appalachian Mountains in southwestern Virginia and northwestern North Carolina. It is named after the town of Blacksburg, Virginia, as its first recorded sighting was within its vicinity. Its natural habitat is temperate forest.

References

  1. 1 2 IUCN SSC Amphibian Specialist Group (2022). "Plethodon punctatus". IUCN Red List of Threatened Species . 2022: e.T59351A118995830. doi: 10.2305/IUCN.UK.2022-2.RLTS.T59351A118995830.en . Retrieved 28 August 2023.
  2. 1 2 "Plethodon punctatus". NatureServe Explorer. 4 August 2023.
  3. 1 2 3 4 5 Buhlmann, Kurt A.; Pague, Christopher A.; Mitchell, Joseph C.; Glasgow, Robert B. (1988). "Forestry operations and terrestrial salamanders: techniques in a study of the Cow Knob salamander, Plethodon punctatus". Management of Amphibians, Reptiles, and Small Mammals in North America: Proceedings of the Symposium (USDA Forest Service General Technical Report RM-166). Flagstaff: USDA Forest Service. pp. 38–44.
  4. 1 2 3 4 5 6 7 8 9 10 11 Mitchell, Joseph C. (1991). "Amphibians and Reptiles". In Karen, Terwilliger (ed.). Virginia's Endangered Species: Proceedings of a Symposium. Blacksburg: McDonald and Woodward Publishing Company. pp. 437–439.
  5. 1 2 3 4 5 6 7 8 Tucker, Robert B. (1998). "Ecology and natural history of the Cow Knob salamander, Plethodon punctatus Highton, in West Virginia". Marshall University Graduate Thesis.
  6. ↑ Powell, Robert; Conant, Roger; Collins, Joseph T. (2016). Peterson Field Guide to Reptiles and Amphibians of Eastern and Central North America (4th ed.). Boston: Houghton Mifflin Harcourt. p. 88.
  7. 1 2 3 4 5 Green, N. Bayard; Pauley, Thomas K. (1987). Amphibians and reptiles in West Virginia. Pittsburgh: University of Pittsburgh Press. pp. 75–76.
  8. 1 2 3 4 Highton, Richard (1988). "Plethodon punctatus". Catalogue of American Amphibians and Reptiles. p. 414.
  9. 1 2 3 4 Petranka, James W. (1998). Salamanders of the United States and Canada. Washington, London: Smithsonian Institution Press. pp. 390–392. ISBN   9781560988281.
  10. 1 2 3 Milanovich, Joseph R.; Peterman, William E.; Nibbelink, Nathan P.; Maerz, John C. (2010-08-16). "Projected Loss of a Salamander Diversity Hotspot as a Consequence of Projected Global Climate Change". PLOS ONE. 5 (8): e12189. Bibcode:2010PLoSO...512189M. doi: 10.1371/journal.pone.0012189 . ISSN   1932-6203. PMC   2922335 . PMID   20808442.
  11. 1 2 3 4 Jacobsen, Carl D.; Brown, Donald J.; Flint, William D.; Pauley, Thomas K.; Buhlmann, Kurt A.; Mitchell, Joseph C. (2020-03-01). "Vulnerability of high-elevation endemic salamanders to climate change: A case study with the Cow Knob Salamander (Plethodon punctatus)". Global Ecology and Conservation. 21: e00883. doi: 10.1016/j.gecco.2019.e00883 . ISSN   2351-9894. S2CID   211145164.
  12. 1 2 Graham, Matthew R.; Flint, William D.; Powell, Alexsis M.; Fet, Victor; Pauley, Thomas K. (2023). "Phylogeography of the Cow Knob Salamander (Plethodon punctatus Highton): populations on isolated Appalachian mountaintops are disjunct but not divergent". Frontiers in Amphibian and Reptile Science. 1. doi: 10.3389/famrs.2023.1175492 . ISSN   2813-6780.
  13. 1 2 Highton, Richard; Hastings, Amy Picard; Palmer, Catherine; Watts, Richard; Hass, Carla A.; Culver, Melanie; Arnold, Stevan J. (2012). "Concurrent speciation in the eastern woodland salamanders (Genus Plethodon): DNA sequences of the complete albumin nuclear and partial mitochondrial 12s genes". Molecular Phylogenetics and Evolution. 63 (2): 278–290. doi:10.1016/j.ympev.2011.12.018. ISSN   1055-7903. PMID   22230029.
  14. 1 2 3 4 5 6 Kuchta, Shawn R.; Brown, Ashley D.; Highton, Richard (2018). "Disintegrating over space and time: Paraphyly and species delimitation in the Wehrle's Salamander complex". Zoologica Scripta. 47 (3): 285–299. doi:10.1111/zsc.12281. S2CID   90215639.
  15. 1 2 3 Felix, Zachary I.; Wooten, Jessica A.; Pierson, Todd W.; Camp, Carlos D. (2019-05-24). "Re-evaluation of the Wehrle's salamander (Plethodon wehrlei Fowler and Dunn) species group (Caudata: Plethodontidae) using genomic data, with the description of a new species". Zootaxa. 4609 (3): zootaxa.4609.3.2. doi:10.11646/zootaxa.4609.3.2. ISSN   1175-5334. PMID   31717092. S2CID   181533495.
  16. ↑ Jacobsen, Carl D.; Flint, William D.; Newman, Jillian C.; Brown, Donald J. (2019). "Plethodon punctatus (Cow Knob salamander). Arboreal behavior". Herpetological Review. 50 (1): 110.
  17. 1 2 Fraser, Douglas F. (1976). "Coexistence of Salamanders in the Genus Plethodon: A Variation of the Santa Rosalia Theme". Ecology. 57 (2): 238–251. doi:10.2307/1934813. ISSN   0012-9658. JSTOR   1934813.
  18. ↑ Mitchell, Joseph; Pauley, Thomas; Withers, David; Roble, Steven; Miller, Brian; Braswell, Alvin; Cupp, Paul; Hobson, Christopher (1999-04-01). "Conservation Status of the Southern Appalachian Herpetofauna". Virginia Journal of Science. 50 (1). doi:10.25778/33tx-fx91. ISSN   0042-658X.
  19. ↑ Markle, Tricia M.; Kozak, Kenneth H. (2018). "Low acclimation capacity of narrow‐ranging thermal specialists exposes susceptibility to global climate change". Ecology and Evolution. 8 (9): 4644–4656. doi:10.1002/ece3.4006. ISSN   2045-7758. PMC   5938462 . PMID   29760904.
  20. 1 2 Sutton, William B.; Barrett, Kyle; Moody, Allison T.; Loftin, Cynthia S.; DeMaynadier, Phillip G.; Nanjappa, Priya (2015). "Predicted Changes in Climatic Niche and Climate Refugia of Conservation Priority Salamander Species in the Northeastern United States". Forests. 6 (1): 1–26. doi: 10.3390/f6010001 . ISSN   1999-4907.
  21. ↑ "2015 Virginia Wildlife Action Plan". bewildvirginia.org. Retrieved 2023-08-28.
  22. ↑ Center for Biological Diversity (11 July 2012), PETITION TO LIST 53 AMPHIBIANS AND REPTILES IN THE UNITED STATES AS THREATENED OR ENDANGERED SPECIES UNDER THE ENDANGERED SPECIES ACT (PDF)
  23. ↑ "Evaluation of a Petition To List the Cow Knob Salamander as an Endangered or Threatened Species Under the Act" (PDF). Federal Register. 81 (51): 14064. 16 March 2016.
  24. 1 2 3 4 Forest Service (2014). George Washington National Forest Revised Land and Resource Management Plan (R8-MB 143 A) (PDF). Roanoke: U.S. Department of Agriculture.
  25. ↑ U.S. Forest Service, U.S. Fish and Wildlife Service (25 January 1994), Conservation Agreement for the Cow Knob Salamander (PDF)
  26. ↑ Springston, Rex (26 January 1994). "From under a rock, a pioneer emerges". Richmond Times-Dispatch . pp. B1, B4.
  27. ↑ McCue, Cathryn (27 January 1994). "Agencies agree to protect rare species". The Roanoke Times . pp. C4.
  28. ↑ "Safety designed for salamander". Daily Press . 28 January 1994. pp. C5 S.
  29. ↑ Springston, Rex (31 October 2015). "Gas pipeline will avoid salamander". Richmond Times-Dispatch . pp. A1, A8.
  30. ↑ Atkinson, Kathleen; Tooke, Tony (19 January 2016), Forest Service Permit Letter 1-19-2016, File Code: 2700 (PDF)
  31. ↑ Baars, Samantha (21 January 2016). "Cow knob salamander reroutes Atlantic Coast Pipeline". C-VILLE Weekly.
  32. ↑ Schwaner, Jeff; Peters, Laura (22 January 2016). "Forest Service denies pipeline route". The News Leader .
  33. ↑ Martz, Michael (22 January 2016). "Forest Service rejects latest proposal for pipeline route". Richmond Times-Dispatch . pp. A1, A7.
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