Habitats of North Carolina
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Upland Hardwood Forests
General Montane Mesic Forests
General Description The cove forests of the Southern Appalachians are some of the most biodiverse habitats in the Temperate Zone. Their cool, moist climates favor the growth of complex, multi-story plant communities; thick, rich litter layers; deeply-shaded rock faces and crevices; and multitudes of perennial springs and cool, clear-flowing streams. These habitats, moreover, have persisted in place for hundreds of millions of years. They survived unglaciated through the Ice Age and their steep slopes have largely protected them from permanent conversion to human uses. They provide refuges for many relicts of otherwise long-vanished floras and faunas. They are also centers of endemism in their own right, with new species being discovered on a regular basis.

Although there are few areas that still support virgin forests, these habitats continue to serve these functions during the Age of Man largely due to their steep slopes, which have resisted cultivation and large-scale development. In North Carolina, in particular, the lack of coal and other extensive mineral deposits have spared these communities from the massive strip mining and mountain-top removals that have taken place in other parts of the Appalachians. Today, these habitats are the most extensive of our remaining natural areas and retain the greatest amount of their pre-Colonial species diversity.

The habitat described here includes species that are associated with the most generalized features of the cove forests, particularly their deeply-shaded, cool, moist microclimates. It is similar, in this regard, to the General Mesic Hardwood Forests habitat, but comprises species that are restricted to the Mountains in North Carolina. Within the Mountains, however, these species occupy a wide altitudinal range, extending from the foothills of the Blue Ridge up into the areas covered with stands of Northern Hardwoods. Although the majority of these habitats occur within the steep, montane "coves" proper, higher ridgetops are also included where they are regularly swept by clouds and rain and support populations of the same mesophilic species that are more confined to ravines at lower elevations. The Determining Species of this habitat occur in both rich and acidic coves; those that are more tightly associated with either of these types are treated in other, more specifically-defined habitats.

The plant species that are restricted to this habitat are relatively few in number, with more species found only in either the rich or acidic varieties of cove forests or more widely in mesic habitats outside the mountains. They include some of the most characteristic tree species of cove forests per se, Yellow Birch and Fraser Magnolia, as well as Red Ramps, the herbaceous species made famous in the Southern Appalachians by Ramp festivals. American Ginseng -- another species famously exploited in the cove forests -- is, on the other hand, more typical of the species that may show a concentration in the mountains but that also occur more generally in mesic habitats all the way across to the eastern edge of the Coastal Plain.

Specialized herbivores are likewise few in number, consisting primarily of species associated with Sugar Maples or montane birches. Much more numerous are more generalized herbivores that feed on a variety of plant species -- most not confined to cove forests -- but which themselves have requirements for cool, moist, forests that restrict them to these habitats. Virtually all of these generalized herbivores have their center of geographic distribution located in the northern states or in Canada. The same is true for some of the predatory species of animals belonging to this habitat, particularly the birds and mammals, all of which are much more common farther north.

For the least vagile species associated with the forest floor, on the other hand, the cove forests of the Southern Appalachians are major centers of their diversity. These species are so highly dependent on specific temperature, moisture, and forest floor conditions that many of them are not only confined to the Southern Appalachians but are endemic to particular mountain massifs within this region. Although their ancestors were able to occupy much wider areas under colder, wetter climate conditions -- as prevailed in the Pleistocene -- their descendants became isolated in much smaller areas when warmer, drier habitats developed in the lowlands between these massifs (in some cases, the rivers in these valleys also act as effective barriers).

The best known of these species are the Plethodontid Salamanders, of which only the most terrestrial species -- members of the genus Plethodon -- are included in this habitat. Currently (and likely to increase), there are sixteen Determining Species for this habitat, out of the twenty that have been recorded in North Carolina. Members of this group in particular have evolved in close association with the cool, wet forest floor conditions that have allowed them to free themselves from the aquatic habitats needed by other amphibians for breeding and larval development: all members of this genus lay their eggs in underground burrows where the high soil moisture keeps them from drying out. This allows them to range freely across the forest floor, independent of surface waters, unlike most of the other members of this family (e.g., Eurycea and Desmognathus species) that are treated in habitats that contain surface waters as important features.

As described by Thomas and Hedin (2008), a number of invertebrate groups associated with the forest floors of mountain coves show similar centers of diversity in Southern Appalachian coves. These include the Xystodesmid Millipedes (Shelley and Whitehead, 1986; Marek and Bond, 2006), Phalangodid Opilionids (Thomas, 2007), Hypochilus and Nesticus spiders (Catley 1994; Hedin, 1997). Certain groups of land snails and ground beetles could also be added, along with probably many other groups of understudied invertebrates. Species that are associated with the forest floor itself are considered Determining Species of this habitat but those that are tightly associated with mesic rock faces are treated separately.

Arguably the most spectacular, if also the most obscure and solitary, is the species that Thomas and Hedin highlight in their paper, Fumontana deprehendor. This tiny, highly weird-looking Harvestman is found only in the cove forests of the Southern Appalachians, is the sole member of its genus, and appears to be most closely related to species that are now restricted to South Africa and South America. It thus appears to be a relict of a Gondwanan fauna that was once widespread in the Paleozoic, with the implication that it has been an inhabitant of the Southern Appalachian region for hundreds of millions of years. As such it would provide a striking confirmation of the stability of the ecosystems of this region, particularly its mesic forests.

Abiotic Factors Geographic Regions: Low to High Mountains. USDA Hardiness Zones: 6. Landform: slopes, ridges, ravines, and bottomlands. Slope Aspect: all aspects, particularly above 4,000'. Soil Moisture: Mesic. Soil texture: loamy. Soil pH: acidic to circumneutral. Soil Nutrient Content: nutrient poor to rich. Microclimate: Cool and humid. Hydrological Features: springs and seeps are common, particularly at the interface between slopes and bottomlands. Fire Frequency: very rare. Drought Frequency: uncommon to rare. Ice Storm Damage: low to moderate. Wind Storm Damage: moderate. Insolation: canopies are well insolated, lower strata are deeply shaded.

Biotic Structure Vegetation Type: closed-canopy forests composed of both broadleaf, deciduous trees and Eastern Hemlocks. Strata: herb layer is typically lush in rich cove forests but sparse in acidic coves; shrub thickets are usually very well developed in acidic coves. Woody debris and leaf litter: logs and fallen branches are common; leaf litter can be plentiful, as under deciduous forests in general, but its build-up may be limited due to erosion and general downhill slippage on steep slopes

Co-evolved Species Groups Phagic and Competitory Symbioses: (Montane Betula species/Ancylis discigerana-Bucculatrix canadensisella-Cameraria betulivora-Hydrelia inornata-Peridea ferruginea-Polygonia faunus) // (Betula alleghaniensis/Parornix vicinella) // (Betula lenta/Phyllonorycter martiella) // (Hydrangea arborescens/Aspilanta hydrangaeella

Determining Species
sciNamecomNameg_ranks_rankmod_s_rankprob_of_extirpation
BEETLES
Anthophylax cyaneus
Trechus rivulisGNRS1S10.33330
Trechus luculentus luculentusa ground beetleGHTH G23T23S1S2S1S20.11107
Trechus unicoiGNRS1S2S1S20.11107
Trechus schwarzi saludaea ground beetleGNRTNRSHS1SHS1
BIRDS
Coccyzus erythropthalmusBlack-billed CuckooG5S2S20.03699
Pheucticus ludovicianusRose-breasted GrosbeakG5S3S4S3S40.00132
Bonasa umbellusRuffed GrouseG5S4S40.00041
Catharus fuscescensVeeryG5S4S40.00041
Setophaga caerulescens, summerS40.00041
Setophaga fuscaBlackburnian WarblerG5S4S40.00041
Junco hyemalis, summerS50.00000
BUTTERFLIES
Polygonia faunusGreen CommaG5S1S2S1S20.11107
Erora laetaEarly HairstreakG2G3S2S3S2S30.01230
FORBS
Trillium erectumIll-scent Trillium
Trillium grandiflorumLarge-flower Trillium
Stachys clingmaniiClingman's Hedge-nettleG2S2S20.03699
Arnoglossum reniformeGreat Indian-plantainG4S3S30.00407
Chelone lyoniiPink TurtleheadG4S3S30.00407
Euonymus obovatusRunning Strawberry-bushG5S3S30.00407
Eutrochium steeleiAppalachian Joe-pye-weed, Steele's Joe-pye-weedG4S3S30.00407
Stellaria coreiCore's StarwortG4S3S30.00407
Allium tricoccumSmall White LeekG5S4S40.00041
Claytonia carolinianaCarolina Spring-beautyG5S4S40.00041
Viola blandaSweet White VioletG5S4S40.00041
Impatiens pallidaPale Jewel-weedG5S5S50.00000
GRAMINOIDS
Carex roanensisRoan SedgeG2G3S2S20.03699
Brachyelytrum aristosumNorthern ShorthuskG5S3S30.00407
HARDWOODS
Acer pensylvanicumStriped MapleG5S4S40.00041
Betula alleghaniensisYellow BirchG5S5S50.00000
Betula lentaSweet BirchG5S5S50.00000
Magnolia fraseriFraser MagnoliaG5S5S50.00000
HARVESTMEN
Hesperonemastoma kephartiGNRS1S2S1S20.11107
Fumontana deprehendorG2G3S2S3S2S30.01230
Leiobunum hoffmaniGNRS2S3S2S30.01230
Crosbycus dasycnemusGNRS3S4S3S40.00132
Leiobunum calcara harvestmenGNRS3S4S3S40.00132
Sabacon cavicolensGNRS3S4S3S40.00132
Caddo agilisGNRSHSH
HEMIPTERAN HOPPERS
Oncopsis abietisSNR
Oncopsis citraSNR
Oncopsis deludaSNR
Oncopsis infumataSNR
Oncopsis minorSNR
MAMMALS
Napaeozapus insignisWoodland Jumping MouseG5S4S40.00041
Tamiasciurus hudsonicusRed SquirrelG5S4S40.00041
Sorex fumeusSmoky ShrewG5S4S5S4S50.00010
Myodes gapperiSouthern Red-backed VoleG5S5S50.00000
Peromyscus maniculatusDeer MouseG5S5S50.00000
MILLIPEDES
Dixioria dactyliferaa millipedeGNRS1S10.33330
Sigmoria austrimontisGNRS1S2S1S20.11107
Sigmoria stenogonGNRS1S3S1S30.03699
Dixioria coronataGNRS2S3S2S30.01230
Dixioria wataugaGNRS2S3S2S30.01230
Dixioria wrightiGNRS2S3S2S30.01230
Deltotaria brimleii brimleiiGNRTNRS3S4S3S40.00132
Falloria tuberosaGNRSHSH
Sigmoria sigirioidesGNRSHS1SHS1
Deltotaria brimleii philiaGNRTNRSHS2SHS2
Dixioria pelaG2SHS2SHS2
Falloria translineataGNRSHS2SHS2
Sigmoria nigrimontis angulosaGNRTNRSHS2SHS2
Prionogonus stibarophallusGNRSHS3SHS3
Sigmoria nigrimontis nigrimontisGNRSHS3SHS3
Sigmoria simplexGNRSHS3SHS3
MOTHS
Cameraria lentella
Platypolia anceps
Phyllonorycter martiellaGNRS1S2S1S20.11107
Pseudeva purpurigeraStraight-lined LooperG5S1S2S1S20.11107
Acrobasis betulellaBirch Tubemaker MothGNRS1S3S1S30.03699
Synanthedon fulvipesGNRS1S3S1S30.03699
Platypolia mactataAdorable Brocade MothG5S2S3S2S30.01230
Stamnodes gibbicostataShiny Gray Carpet MothG4S2S3S2S30.01230
Ancylis discigeranaYellow Birch Leaffolder MothGNRS2S4S2S40.00407
Epirrhoe alternataWhite-banded Toothed Carpet MothGNRS2S4S2S40.00407
Venusia comptariaBrown-shaded CarpetG5S2S4S2S40.00407
Acronicta funeralisFunerary DaggerG5S3S4S3S40.00132
Acronicta spinigeraNondescript DaggerG4S3S4S3S40.00132
Eosphoropteryx thyatyroidesPink-patched Looper MothG4S3S4S3S40.00132
Eugonobapta nivosaria Snowy Geometer MothG5S3S4S3S40.00132
Fishia illocataWandering Brocade MothG5S3S4S3S40.00132
Hydrelia albiferaFragile White Carpet MothG5S3S4S3S40.00132
Hyppa contrastaG3G4S3S4S3S40.00132
Idia laurentiian erebid mothGNRS3S4S3S40.00132
Leuconycta lepidulaMarbled-green Leuconycta MothG5S3S4S3S40.00132
Lithophane heminaHemina PinionG5S3S4S3S40.00132
Lithophane semiustaG4S3S4S3S40.00132
Macaria notataBirch Angle MothG4G5S3S4S3S40.00132
Olethreutes ferriferanaHydrangea Leaftier MothGNRS3S4S3S40.00132
Phlogophora irisOlive Angle Shades MothG5S3S4S3S40.00132
Trichodezia albovittataWhite-striped Black MothG5S3S4S3S40.00132
Cepphis armatariaScallop MothG5S4S40.00041
Cladara atroliturataScribbler MothG5S4S40.00041
Heterophleps refusariaThree-patched BigwingG4S4S40.00041
Plagodis serinariaLemon Plagodis MothG5S4S40.00041
Symmerista leucitysOrange-humped Mapleworm MothG5S4S40.00041
Acronicta fragilisFragile DaggerG5S4S5S4S50.00010
Acronicta innotataUnmarked DaggerG5S4S5S4S50.00010
Aethalura intertextaFour-Barred Gray MothG5S4S5S4S50.00010
Euphyia intermediataSharp-angled CarpetG5S4S5S4S50.00010
Hydrelia inornataUnadorned CarpetG5S4S5S4S50.00010
Lithophane innominataNameless Pinion MothS4S50.00010
Peridea ferrugineaChocolate Prominent MothG5S4S5S4S50.00010
Aspilanta hydrangaeellaGNRSUSU0.00202
Bucculatrix canadensisellaBirch Skeletonizer MothGNRSUSU0.00202
Cameraria betulivoraBirch-leaf Blotchminer MothGNRSUSU0.00202
Hyppa xylinoidescranberry CutwormG5SUSU0.00202
Operophtera bruceataBruce Spanworm MothGNRSUSU0.00202
Oreana unicolorellaa pyralid mothGNRSUSU0.00202
Packardia elegansElegant Tailed Slug MothGNRSUSU0.00202
Parornix vicinellaGNRSUSU0.00202
ORTHOPTERANS
Scudderia septentrionalisNorthern Bush KatydidG3S2S3S2S30.01230
Amblycorypha alexanderiClicker Round-winged KatydidGNRSUSU0.00202
SALAMANDERS
Plethodon chattahoocheeChattahoochee Slimy SalamanderG3S1S10.33330
Plethodon cheoahCheoah Bald SalamanderG1S1S10.33330
Plethodon jacksoniBlacksburg SalamanderG1G2S1S10.33330
Plethodon ventralisSouthern Zigzag SalamanderG4S1S10.33330
Plethodon amplusBlue Ridge Gray-cheeked SalamanderG2S2S20.03699
Plethodon aureolusTellico SalamanderG2S2S20.03699
Plethodon meridianusSouth Mountain Gray-cheeked SalamanderG2S2S20.03699
Plethodon jordaniJordan's SalamanderG4S3S30.00407
Plethodon richmondiSouthern Ravine SalamanderG5S3S30.00407
Plethodon shermaniRed-legged SalamanderG3S3S30.00407
Plethodon serratusSouthern Red-backed SalamanderG5S4S40.00041
Plethodon teyahaleeSouthern Appalachian SalamanderG4S4S40.00041
Plethodon yonahlosseeYonahlossee SalamanderG4S4S40.00041
Plethodon metcalfiSouthern Gray-cheeked SalamanderG4S5S50.00000
Plethodon montanusNorthern Gray-cheeked SalamanderG4S5S50.00000
Plethodon glutinosusNorthern Slimy SalamanderG5SUSU0.00202
Nr = Number of Ranked Species = 119
Ner = Number of Extant, Ranked Species = 108
Nv = Number of Historic and Extirpated Species = 2
Nar = Number of Species at Risk of Extirpation (State rank > S5) = 99
Nss = Number of Secure Species (State Rank = S5) = 9
Pss = Proportion of Secure Species (Nss/Ner) = 0.08333
ENE = Expected Number of Extirpations (Sum of PE) = 3.32452
Average PE (ENE/Ner) = 0.03078
Habitat Risk Index = (Nar+Nv) x Average PE = 101 x 0.03078 = 3.10878

Estimated Risk to the Determining Species
Estimated Risk to the Co-evolved Species Groups
Estimated Security of the Habitat
Index of Habitat Imperilment
Identified Risks
Observed Trends
Distribution Map
Distribution Almost all of the records for the species associated with this habitat come from the Mountains or nearby areas in the western Piedmont. Records from further east mainly represent rare, sporadic records of Black-billed Cuckoos from the breeding season. Records for both Red Squirrels and Unmarked Dagger Moths from Wake County are highly unlikely to represent resident, reproducing populations.
Major Conservation Reserves
Priority Areas for Surveys and Conservation Protection
Stewardship and Management Recommendations
References Thomas, S.M. and Hedin, M., 2008. Multigenic phylogeographic divergence in the paleoendemic southern Appalachian opilionid Fumontana deprehendor Shear (Opiliones, Laniatores, Triaenonychidae). Molecular Phylogenetics and Evolution, 46(2):645-658. Available online at: https://www.sciencedirect.com/science/article/pii/S1055790307003521?casa_token=r1KZfWySwYQAAAAA:geMEAJBIss8ImV1KPTaPvYvqTs6Su9c-_hjDPqZ9AOIUWLDAWHkMPfsxbMwiwvAS4JmWBCL_Dw
Updated on 2021-12-31 14:04:13