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| Habitat Overview |
Vast swamps, lakes, and ponds dominated by Cypresses (Taxodium species) are hallmark habitats of the Southeastern Coastal Plain and Mississippi Embayment, supporting a diverse assemblage of plants and animals (see Blevins and Schafale, 2011). This habitat unit described here focuses just on the Cypresses and animal species that are particularly associated with Taxodium; species that are associated more generally with coastal swamps are treated in other habitat types.
For the two plant members of this habitat- Bald Cypress (Taxodium distichum) and Pond Cypress (T. ascendens) -- hydrology is a key factor: habitats occupied by these species are inundated frequently and/or for long periods. These include both riverine and non-riverine swamp forests, lake and pond shorelines, and cypress savannas associated with Carolina Bays. Water chemistry is less important, with Taxodium distichum, at least, occurring in both nutrient-rich brownwater river systems and nutrient-poor, highly acidic blackwater systems; Taxodium ascendens is found primarily in blackwater systems.
While the typically hydric swamps occupied by Bald Cypress rarely experience any fires, the shallower ponds or savannas occupied by Pond Cypresses frequently dry out, making them more likely to burn, especially given their usual occurrence within fire-maintained pine savanna and sandhill habitats. Fire, in fact, plays an important role in maintaining the more open canopy structure found in these habitats and consequently the much richer herbaceous ground cover than is found in riverine cypress swamps (Ewal, 1995).
For the animal members of this habitat, the cypresses themselves are the main habitat requirement. Anhingas are the only animal member of this habitat that actually requires standing water for foraging. Insect members of this habitat are adapted to spending their entire lives up in the trees, well above the water line. For the most part, no major differences have been observed among the animal members of this habitat between swamp forests composed primarily of T. distichum and cypress savannas dominated by T. ascendens. Neither do important differences appear to exist between cypress stands associated with peatland vs. alluvial soils, brownwater rivers vs. blackwater rivers, riverine vs. non-riverine or tidal vs. non-tidal swamps. Only Tolype minta appears to be associated primarily or possibly exclusively with cypress savannas and ponds where Pond Cypress is dominant.
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| Related NHP Natural Communities |
The Natural Heritage Program recognizes eighteen different natural communities or subtypes that contain cypress species as either dominants or important constituents: Brownwater Bottomland Hardwoods (Swamp Transition Subtype), Blackwater Bottomland Hardwoods (Swamp Transition Subtype), Cypress–gum Swamp (Brownwater Subtype), Cypress-gum Swamp (Intermediate Subtype), Cypress--gum Swamp (Blackwater Subtype), Cypress–gum Swamp (Blackwater Cove Subtype), Coastal Plain Small Stream Swamp, Coastal Plain Semipermanent Impoundment (Cypress-gum Subtype), Nonriverine Swamp Forest (Cypress-Gum Subtype), Nonriverine Swamp Forest (Mixed Subtype), Coastal Plain Depression Swamp (Mixed Subtype), Coastal Plain Depression Swamp (Cypress Dome Subtype), Cypress Savanna (Typic Subtype), Cypress Savanna (Acid Subtype), Natural Lake Shoreline Swamp (Cypress Subtype), Natural Lake Shoreline Swamp (Lake Waccamaw Subtype), Maritime Swamp Forest (Cypress Subtype), Tidal Swamp (Cypress–gum Subtype) |
| Determining Species |
Phagic and Competitory Symbioses:(Taxodium species/Acronicta perblanda-Coleotechnites variiella-Cutina albopunctella-Cutina aluticolor-Cutina arcuata-Cutina distincta-Dioryctria pygmaeella-Iridopsis cypressaria-Iridopsis pergracilis-Isoparce cupressi-Lithophane abita-Macaria aequiferaria-Nemoria elfa-Tolype minta-Inscudderia walkeri-Stictolobus minutus-Tortistilus lateralis)
In addition to the two species of Cypress themselves, which are present in 100% of the occurrences of this habitat type, all of the insect members of this habitat show >90% fidelity to cypress-containing habitats. Most appear to feed on both species of cypress, although Tolype minta may be restricted to T. ascendens (the host plants of this species still need confirmation) and Cutina arcuata and Isoparce cupressi may feed only on T. distichum. Though highly co-evolved with Cypresses, these species are more frequently mentioned in accounts as "pests" on these trees (e.g., USDA, 1971; UFL IFAS Extension 2015) than as part of the natural history of cypress swamps. We, however, prefer to recognize them as important native herbivores, i.e., as species that play an important role in the food web specifically anchored in cypress swamps.
Apart from their exclusive (or nearly so) diet on Taxodiums, the caterpillars of the moths in this group, along with both the adults and nymphs of the Cypress Katydid, are strongly streaked with black, green, brown and white, blending in with the cypress needles on which they feed. The adult moths, in contrast, are mainly gray, tan, or reddish brown with darker longitudinal streaks, allowing them to blend in with the bark of the cypresses.
The sole vertebrate member of this habitat, the Anhinga, makes use of a wider range of swampy habitats and has a range that extends far beyond that of our Taxodium species. However, in North Carolina, its breeding range strongly coincides with that of the Cypresses and it forages primarily in deeply flooded freshwater swamps and preferentially nests in Cypresses (see Birds of North Carolina Website, 2018). Consequently, we estimate that it shows at least an 80% fidelity to cypress-containing habitats.
Except for Anhingas, which range into Central and South America, all members of this habitat are endemic to the Coastal Plains and Mississippi Embayment of the Southeast. The genus Cutina, composed of four species (Pogue and Ferguson, 1998) is entirely associated with Taxodium. The same may be true for Isoparce, which in addition to I. cupressi, also includes the recently described I. broui, which is presumed to be associated with one of the Mexican species of Taxodium. On the other hand, the other genera all have species that feed on other plant genera, including conifers but also species of hardwoods.
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| Candidates for Inclusion |
Other insects that are specialist feeders on Taxodium can be added, as can any plant or other species that are highly associated with cypress-containing habitats (i.e., >=80% fidelity). Highly specialized parasitoids associated with the insect members of this habitat -- critical for maintaining ecosystem stability -- will also be included once they are identified. |
| Habitat Sub-sets |
Only a few differences in habitat members appear to exist between the deeply flooded swamps dominated by Bald Cypress and the shallower ponds and savannas dominated by Pond Cypress — not enough to distinguish two separate habitats. |
| Distribution Map | |
| Distribution |
This map shows the counties for which the NCBP websites currently contain records for members of this habitat. The shading indicates concentrations of habitat members, weighted by their conservation significance: the darker the color, the greater the number and/or significance of the species the county contains. Although somewhat biased due to differences in survey coverage and intensity (see below), the general pattern appears accurate that there is greater concentration in the Outer Coastal Plain, where there are generally larger and more numerous stands of cypress, particularly Bald Cypress, which appears to have more associated habitat members. A secondary area of concentration is located in the Fall-line Sandhills and the Carolina-Bay-rich areas of Scotland, Robeson, Cumberland, and particularly Bladen and Columbus Counties, all where Pond Cypress Ponds and Savannas are common. Piedmont stands of Bald Cypress appear to be naturally more depauperate but the large area in the central part of the Inner Coastal Plain that shows a lower concentration of habitat members — primarily records for just the two cypress species — is likely to represent a lack of surveys in that particular region for the animal members of this habitat . |
| Survey Coverage Map |
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| Survey Coverage |
The two species of Taxodium have been well surveyed in North Carolina. Herbarium records for one or both species come from all but one of the shaded counties on the map above (see UNC Herbarium, http://www.herbarium.unc.edu/; and Vascular Plants of North Carolina, https://auth1.dpr.ncparks.gov/flora/index.php) (Warren County — which lacks herbarium records for cypress — is represented solely by one of the moth species, Macaria aequiferaria). Natural area inventories conducted by NHP have also documented the occurrences of cypress-containing natural communities throughout the Coastal Plain (indicated by the blue dots on the map above).
In contrast, the map also shows that surveys for the Lepidopteran members of this habitat(green dots) have been concentrated in just a few areas, particularly in the Outer Coastal Plain, Fall-line Sandhills, and three of the four brownwater river systems in North Carolina. NHP inventories documenting occurrences of these species include Fussell et al. (1995); Hall (1999a); Hall (1999b); Hall (2009); Hall and Schweitzer (1993); Hall et al. (1999); Hall et al. (2013); LeBlond et al. (1997); and LeGrand et al. (2013). Additional records come from moth sampling conducted by J.B. Sullivan at Lake Waccamaw and Lumber River State Parks and the Croatan National Forest; and from NRID records from Merchants Millpond State Park and Dismal Swamp State Natural Area. |
| Survey Priorities |
Based on the presence of high-quality natural community occurrences, the following areas should be priorities for surveys of the animal members of this habitat: Northwest River Marsh Game Land, North River Game Land, Great Dismal Swamp NWR, Meherrin River Swamp, Chowan Swamp, Cashie River Swamp, East Dismal Swamp, Pantego Swamp, Pungo Lake, Lake Mattamuskeet, Gull Rock Game Land, Scuppernong River, River Park North, Robertson’s Millpond, Howell Woods, Black Creek Swamp, Cliffs-of-the-Neuse State Park, Neuse River Game Land, Great Coharrie Creek, Carvers Creek, Antioch Bay, Goose Pond Bay, Hamby’s Bay, Upper Lumber River Swamp, Black River, South River, Waccamaw River Swamp, Lockwoods Folly River, Cape Fear River Wetlands Game Land |
| Average Imperilment of Habitat Members |
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| Habitat Conservation Status |
The average of the Imperilment Scores is 1.66. That is equivalent in value to single species elements that are secure at the Global Level but are considered to be of possible concern -- S3 -- at the state level. None of the members of this habitat are considered imperiled globally and only three – Acronicta perblanda, Iridopsis cypressaria, and Tolype minta – are considered of significant conservation at the state level. |
| High Quality Habitat Occurrence Table |
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| High Quality Habitat Occurrences |
The table above lists major areas of this habitat (aggregated across county and natural area boundaries) ranked by their combined imperilment scores, expressed as a percentage of the total possible score.
No site has a 100% score, mainly due to the extreme rarity of Acronicta perblanda in North Carolina (known from only a single site in the Core Creek area). Nonetheless, the seven areas that score above 50% meet the general criteria for high quality occurrences of this habitat: all have extensive stands of cypress and have records for both a large number of the habitat members, including some of the rarer species, such as Iridopsis cypressaria or Tolype minta. All have also been surveyed fairly intensively, which is not the case for several other large tracts of cypress such as occur along the South and Black Rivers, and Chowan Swamp.
All of these highest quality areas are probably separately viable in the short run: given the limited geographic scope and severity of past natural disturbances, populations within each of these areas should be able to resist and/or recover from most local extirpation events. The same is true for at least some forms of artificial disturbances, such as localized timber harvest or limited spraying to control Gypsy Moths. |
| Protected Habitat Occurrences |
Federal Preserves: Dismal Swamp NWR, Currituck NWR, Roanoke River NWR, Alligator River NWR, Pocosin Lakes NWR, Dare County Bombing Range, Croatan National Forest, Camp Lejeune, Eagle Island, Fort Bragg, Camp Mackall Military Reservation, Military Ocean Terminal at Sunny Point
State Parks: Merchants Millpond State Park, Dismal Swamp State Natural Area, Jockey’s Ridge State Park, Goose Creek State Park, Sandy Run Savannas State Natural Area, Singletary Lake, Jones Lake, Bay Tree Lake, Bushy Lake, Carolina Beach State Park, Lake Waccamaw, Lumber River
Other State Preserves: Roanoke River Wetlands Game Land, Holly Shelter Game Land, Suggs Millpond Game Land, Bladen Lakes State Forest, Bald Head Island Coastal Reserve, Boiling Spring Lakes PCP Preserve, Hog Branch Ponds PCP Preserve
Private Nature Preserves: Roanoke River TNC Preserve, Camassia Slopes TNC Preserve, Devil’s Gut TNC Preserve, Nag’s Head Woods TNC Preserve, Angola Creek TNC Preserve, Black River TNC Preserve, Green Swamp TNC Preserve, Myrtle Head Savanna TNC Preserve
Most of these preserves are managed at least partially to protect natural ecosystems, although several also manage some of their forest stands for timber production. Management plans, where they exist, mainly deal with the stewardship of vascular plants and vertebrates. In some cases, where targeted insect surveys have been conducted, some recognition exists that decisions concerning the use of insecticides or prescribed burning should consider impacts to non-target species, particularly those that are of conservation concern. |
| Threats and Trends |
Habitat Conversion and Exploitation
Cypress wood is valued for its rot resistance and other properties and has been heavily logged in the past. Almost all stands are second growth, with only a very small number of sites now supporting old growth trees, for example along the Black River. With the exception of Anhingas, which require fairly mature trees for nesting, members of this habitat probably all make use of even young regenerating stands. They may therefore be able to use stands that are now being exploited for wood pellet and mulch production, which are harvested at a far younger age than the mature trees that were traditionally harvested for lumber and shingles. That is likely to be true only where pesticides are not used to combat “pest” species, a category that could be applied to all of the insect members of this habitat, i.e., herbivores on cypress. Defoliating events have, in fact, been reported in Florida populations of Iridopsis cypressaria (USDA, Plant Protection Division, 1971). Unlike exotic pest species, however, native herbivores normally are well controlled by numerous parasitoid wasps and flies. The Florida outbreak, in fact, was reported as very heavy in 1968 but virtually non-existent the following year.
Clear-cut cypress stands may not regenerate to cypress unless an effort is made to re-plant them. Regeneration may be especially limited where construction of reservoirs has altered the natural flood regime and sediment deposition (Townsend, 2001; Pearsall et al., 2005). Changes in stand structure and composition may be occurring due to artificial alteration of natural flows even within well-protected preserves such as those that exist in the lower Roanoke River floodplain. If the increased flooding that appears to be developing due to climate change leads to the construction of more flood control systems either within or upstream of the Coastal Plain, these impacts are likely to increase in significance.
In the case of isolated Cypress Savannas or shallow Cypress Ponds, hydrological alteration typically takes the form of ditching and draining, followed by the conversion of the cypress stands to ether pine plantations or, in some areas, truck farms. Large numbers of Carolina Bays in the Inner Coastal Plain have been permanently converted in this way. Others have been substantially altered due to fire suppression, which is needed to keep cypress from being replaced by weedier hardwoods (Ewal, 1995). In the case of permanently flooded Carolina Bays, many have had their shorelines developed, including some state lakes, such as White Lake, Bay Tree Lake, and Lake Waccamaw. One important impact on the insect members of this habitat — all of which are nocturnal — is the disruption created by the outdoor lighting associated with shoreline development.
Climate change
Climate change is likely to have several severe effects on this habitat. Sea level rise and accompanying saltwater intrusion are likely to eliminate most, if not all cypress stands associated with Maritime Swamp Forests and Tidal Swamp Forests, all of which are located in close proximity to the sounds that line our coast. Breaches in the Outer Banks may greatly accelerate these impacts. These same effects are also likely to extend well up our coastal rivers, leading to “marshification” of what are now our most extensive areas of cypress forests, including most of those that support our highest quality occurrences of this habitat. This is already affecting the estuary of the Cape Fear, where “ghost forests” of cypress (i.e., dead trees) are easily seen from the bridges that cross what are now vast marshes along the lower portion of the river. Similar effects are visible along the canals that penetrate the peatlands of the Albemarle-Pamlico Peninsula, where large tracts of non-riverine swamp forest are likely to be ultimately lost to these impacts.
Inland areas — which will become increasingly important as refuges for this habitat — are likely to be affected by both changed flood regimes as well as increased frequency of droughts and wildfires. Riverine swamps that rarely burned in the past may be more likely to burn in the future and even the regularly burned Pond Cypress Savannas are more likely to suffer catastrophic canopy or peatland fires, greatly reducing their ability to support members of this habitat.
Exotic Invasives and Use of Pesticides
There currently do not appear to be any exotic species that represent a significant threat either to the cypresses or the animal members of this habitat. Fire ants — which are major predators on caterpillars and other insects — may be having an impact on members of this habitat that inhabit cypress savannas and the Tachinid fly, Compsilura concinnata, which was introduced to control Gypsy Moths, may have an impact on the larvae of Isoparce cupressi once they spread south into the state. However, neither of these threats has yet been investigated in the field with regard to this habitat type.
Several native species undergo periodic defoliating outbreaks in cypress stands. These include two of the current members of this habitat, Iridopsis cypressaria and Iridopsis pergracilis (USDA, 1971; xx). Other species include Cypress Leaf Beetles (Systena marginalis and/or S. plicata) and the Baldcypress Leafroller (Archips goyerana) (refs), both of which could be added to this habitat if they prove to be present in North Carolina and monophagous on Taxodium species. Unlike exotic species, however, these native species have long-evolved biological controls, particularly parasitoid wasps and flies (which if host specific, would also be members of this habitat). Normally, herbivore outbreaks — in their native habitats — are only periodic, with the control species quickly reducing the populations of their host species. Artificial control measures are rarely needed, with cypresses normally quickly re-foliating. Application of Btk or other broad spectrum pesticides is more likely to have highly detrimental impacts on this habitat, far outweighing any conservation benefit that would be obtained.
Pesticide impacts on this animals in this habitat type are more likely to result as collateral damage from spraying campaigns to control Gypsy Moth populations. While that species attacks hardwoods preferentially, they often co-occur with cypress stands in floodplain forests. The largest single Gypsy Moth eradication campaign in North Carolina involved the use of Btk, which was sprayed over many sites containing stands of cypress (mostly Pond Cypress). In a study of the impacts to non-target species of Lepidoptera conducted as part of that project (Hall et al., 1999), five of the habitat members — Iridopsis pergracilis, Isoparce cupressi, Macaria aequiferaria, Nemoria elfa and Tolype minta — were estimated to be at moderate to high risk of Btk impacts (based primarily on their presence as early instar larvae at the time of spraying).
In addition to impacts to the insect members of this habitat, Anhingas – as fish-eating species – is affected by the biological accumulation and magnification of persistent insecticides, herbicides, or other pollutants entering the water (Frederick et al., 2000). These include agricultural chemicals originating from farms and tree plantations located even at some distance from a protected area of cypress (Anhingas themselves may forage in areas located well-away from a preserve where they nest). Other sources include industrial sites which release pollutants – e.g., mercury and dioxins -- into the air as well as the water.
Loss of Connectivity
With extensive swamps along numerous creeks and rivers in the Coastal Plain, tidal swamp forests and maritime swamp forests along the sounds, and numerous ponds and savannas in the interbasin areas, and this habitat has been well-connected in the past, allowing any local extirpation to be recovered through re-colonization from off-site populations. Sea level rise and saltwater intrusion, however, are likely to lead to loss of connections along the coast and ongoing habitat conversion may do the same for the areas in between the river basins. Increased isolation of populations, coming at a time of increasing environmental disturbances, may substantially reduce the viability of this habitat, even where individual sites maintain their stands of cypress. |
| Status Summary |
Cypress swamps still exist as large tracts of good quality habitat in North Carolina and the majority of the habitat members have been recorded at numerous locations, many of them well protected at least from development or exploitation. While this habitat as a whole appears to be relatively secure for the next several decades, the combined effects of climate change and exploitation and habitat conversion by human activity are likely greatly reduce the overall range of this habitat in the state and make individual occurrences far less viable. Habitat members associated with Carolina Bays, Cypress Savannas, Non-Riverine Swamp Forests, and Tidal Swamp Forests are likely to be at particular risk.
State Rank: S4S5
This habitat is currently secure but closer monitoring will likely be needed in the future, i.e., moving its rank more towards S3. |
| Stewardship Recommendations |
Restore/Maintain Natural Hydrology Within preserves, plug ditches previously used to attempt to drain swamps and remove or thin former pine plantations or hardwood stands that may have prospered due to the prior draining efforts but which may now be affecting the natural hydrology of the site. Outside of preserves, work with federal and other agencies to ensure that reservoirs operate in run-of-river mode or otherwise maintain a natural flood regime (following the model of the Roanoke River restoration — see Pearsall et al., 2005).
Restore/Maintain Natural Burn Regime Restore burn regimes within Cypress Savannas and surrounding fire-maintained habitats (e.g., sand ridges and flatwoods). Most of the members of this habitat are likely to be unaffected by light ground fires, but could be drastically affected by fires reaching the canopy. Fires hot enough to ignite underlying peat may also affect the roots of the cypresses, posing a risk to the entire habitat. Wherever possible, burn only a portion of stand in any one year, leaving at least some cypress-containing areas as refuges during the burn and as re-colonization sources following the burn.
Exclude or Minimize the Use of Insecticides Most of the members of this habitat are likely to be adversely affected by use of aerial application of broad-spectrum insecticides. The Lepidoptera, in particular, are also likely to be affected by the use of Bacillus thuringienesis kurstaki (Btk), a bacterial preparation that affects the majority of moth and butterfly species. When outbreaks of Gypsy Moths are discovered outside areas where they have become established, eradication efforts within preserves should only be conducted using Lymantria-specific control agents, including Gypchek or pheromone flakes (Disparlure). Efforts to control mosquito populations should similarly make use of specific control agents, such as Bacillus thuringiensis israelensis (Bti). Occasional defoliating outbreaks of habitat members themselves are best left untreated, but if they persist, investigations should be made to determine if their specific natural control agents (e.g., parasitic wasps and flies) have somehow been eliminated from the preserve due to environmental changes.
Preserve managers need to work with neighboring landowners to reduce the use of pesticides on adjoining lands. Environmentally persistent pesticides should be particularly discouraged. Since this habitat includes at least one likely pollinating species (Isoparce cupressi), use of pesticides or GMO-produced toxins that accumulate in nectar and pollen should also be curtailed.
Limit Intrusive Uses The insect members of this habitat are probably fairly insensitive to human presence but are likely to be adversely affected by outdoor lighting, particularly high intensity lights that give off high levels of UV. Anhingas are more sensitive to frequent human activities, particularly around their nesting sites (Frederick et al., 2000). Trails (including canoe trails) should be routed to avoid both nests and important foraging areas.
Maintain Landscape Connectivity The best preventative or restorative for local extirpation is to maintain connections to other areas occupied by the habitat members. Within preserves, support this principle by dividing cypress habitats, as much as possible, into multiple management units, not all of which will be affected by a given management action during any one year. Outside of preserves, work with adjoining landowners to make sure that timber operations or habitat conversions do not permanently eliminate connections between habitat units. Assign a high priority for conservation to preserving connections between habitat units, even if only as series of disjunct stepping stones. State agencies and large land conservation organization should give particular emphasis in maintaining connections between river basins.
Monitor Habitat Member Populations Following any particular management action or major natural disturbance, monitor the effects on the habitat members themselves — including the insects — not just indirectly on supposed surrogates, such as vegetation. Give special attention to habitat members with the highest degree of imperilment, which in this habitat type includes Acronicta perblanda and Iridopsis cypressaria. Populations should be periodically surveyed over multiple areas to determine whether the conservation status of either the habitat or its individual members is changing due to climate change or other widespread events. Further surveys are needed in any case to document the presence and conservation status of at least a few members of this habitat, including A. perblanda, I. cypressaria, Lithophane abita, and Tolype minta.
Educate the Public and Support Scientific Research The Lepidopteran members of this habitat are easily observed using blacklight/sheet set-ups. Moth nights held in areas within or adjacent to cypress stands can be used to educate about habitat specialization, co-evolution, and the relationship between habitat/ecosystem conservation and species conservation. On good nights, when hundreds of cypress moths show up at the sheets, the role cypress-feeding species play in providing food for species higher up in the food web should be readily apparent. In outbreak years, the role of native control organisms in bringing populations back to normal levels would be an important topic.
Scientific research into any of these subjects should be welcomed, as should any effort to monitor the long-term changes in this habitat as a reflection of environmental change. Each management effort should be regarded as an experiment, with the results monitored and fed-back into the decision-making process, helping bring about adaptive management. |
| References |
Ewal, K.C. 1995. In Fire in wetlands: a management perspective. Proceedings of the Tall Timbers Fire Ecology Conference (No. 19, pp. 111-116). Available online at: http://talltimbers.org/wp-content/uploads/2014/03/Ewel1995_op.pdf
Frederick, P. C. and D. Siegel-Causey (2000). Anhinga (Anhinga anhinga), version 2.0. In The Birds of North America (A. F. Poole and F. B. Gill, Editors). Cornell Lab of Ornithology, Ithaca, NY, USA
Fussell, J.O.; Webster, W.D.; Hall, S.P.; LeGrand, H.E.; Schafale, M.P.; and Russo, M.J. 1995. Ecosystem survey of Dare County Air Force Range, North Carolina. Rep. to N.C. Natural Heritage Program, Division of Parks and Recreation, Department of Environment and Natural Resources; Raleigh, NC.
Hall, S.P. 1999a. Inventory of the moths, butterflies, and grasshoppers of Pettigrew, Goose Creek, and Jockey's Ridge State Parks and Nag's Head Woods TNC Preserve. Unpubl. Rep., NC NHP; Raleigh, NC
Hall, S. P. 1999b. Inventory of the macrolepidoptera of the Devil’s Gut Preserve. North Carolina Natural Heritage Program, Raleigh, North Carolina, 110 pp
Hall, S.P. 2009a. Landscape inventory of the Fall-Line Sandhills. Unpubl. Rep. to US Fish and Wildlife Service Raleigh Field Office; Raleigh. North Carolina Natural Heritage Program, Raleigh
Hall, S.P. and Schweitzer, D.F. 1993. A survey of the moths, butterflies, and grasshoppers of four Nature Conservancy Preserves in Southeastern North Carolina. Unpubl. Rep. to TNC, NC Field Office; Durham, NC
Hall, S.P.; LeGrand, H.E.; and Sullivan, J.B. 2013. A Natural Heritage Inventory of the Tar River Floodplain, North Carolina. 2013 field data and moth collection records.
Hall, S.P.; Sullivan, J.B.; and Schweitzer, D.F. 1999. Eradication of the Asian-strain of The Gypsy Moth from the Cape Fear Region of North Carolina: Assessment of Risk to Nontarget Macro-Lepidoptera. USDA Forest Service Technical Publication Series; Morgantown, WV. 95 pp. Available online at: http://nc-biodiversity.com/sites/default/files/AGM%20Non-Target%20Impact%20Study%2C%20edited%20June%202017.pdf
Hinsley, L.E. 2002. Research at N. C. State University related to regeneration of Atlantic White Cedar (AWC) and Baldcypress. USFWS, Raleigh Ecological Services Field Office. Available online at: https://www.fws.gov/raleigh/coastal/plnwrawc/atlanticwhitecedarresearch.html
LeBlond, R.J.; Fussell, J.O.; Braswell, A.L.; Grant, G.S.; Hall, S.P.; and Sullivan, J.B. 1997. Inventory of the Rare Species, Natural Communities, and Critical Areas of Camp Lejeune Marine Corps Base, North Carolina. Phase III. NC Natural Heritage Program, Div. Parks and Recreation; Raleigh, NC; 1131 pp.
LeGrand, H.E.; Hall, S.P.; and Sullivan, J.B. 2013. A Natural Heritage Inventory of the Roanoke River Floodplain, North Carolina. NC Natural Heritage Program, Raleigh.
Pearsall, S.H., McCrodden, B.J. and Townsend, P.A., 2005. Adaptive management of flows in the lower Roanoke River, North Carolina, USA. Environmental management, 35(4), pp.353-367. Available online at: ftp://kshsrv.fgg.uni-lj.si/students/podipl/UVR/Pearsall_et_al_2005.pdf
Popenoe, J., Warwick, C.R., and Kjelgren R. 2018. Key Plant, Key Pests: Baldcypress (Taxodium distichum). UFL IFAS Extension, ENH1293.
Townsend, P.A., 2001. Relationships between vegetation patterns and hydroperiod on the Roanoke River floodplain, North Carolina. Plant Ecology, 156(1), pp.43-58.
http://edis.ifas.ufl.edu/topic_cypress_pest_insects
US Fish and Wildlife Service 2013. Habitat management plan for Roanoke River National Wildlife Refuge. Available online at: https://www.fws.gov/uploadedFiles/Roanoke%20River%20HMP%20signed-sm.pdf
USDA, Plant Protection Division. 1971. Cooperative economic insect report 21:283. Available online at: https://www.biodiversitylibrary.org/page/43510618#page/31/mode/1up
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