Biodiversity Data Journal : Taxonomic Paper
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Taxonomic Paper
Guide to the littoral zone vascular flora of Carolina bay lakes (U.S.A.)
expand article infoNathan Howell, Alexander Krings, Richard R Braham§
‡ Department of Plant and Microbial Biology, North Carolina State University, Raleigh, United States of America
§ Department of Forestry and Environmental Resources, North Carolina State University, Raleigh, United States of America
Open Access

Abstract

Background

Carolina bays are elliptic, directionally aligned basins of disputed origin that occur on the Atlantic Coastal Plain from the Delmarva Peninsula to southern Georgia. In southeastern North Carolina, several large, natural, lacustrine systems (i.e., Carolina bay lakes) exist within the geomorphological features known as Carolina bays. Within the current distribution of Carolina bays, Bladen and Columbus counties (North Carolina) contain the only known examples of Carolina bay lakes. The Carolina bay lakes can be split into two major divisions, the “Bladen Lakes Group” which is characterized as being relatively unproductive (dystrophic – oligotrophic), and Lake Waccamaw, which stands alone in Columbus County and is known for its high productivity and species richness. Although there have been several studies conducted on these unique lentic systems, none have documented the flora comprehensively.

New information

Over the 2013−2014 growing seasons, the littoral zone flora of Carolina bay lakes was surveyed and vouchered. Literature reviews and herbarium crawls complemented this fieldwork to produce an inventory of the vascular plant species. This survey detected 205 taxa (species/subspecies and varieties) in 136 genera and 80 vascular plant families. Thirty-one species (15.2%) are of conservation concern. Lake Waccamaw exhibited the highest species richness with 145 catalogued taxa and 26 species of conservation concern. Across all sites, the Cyperaceae (25 spp.), Poaceae (21 spp.), Asteraceae (13 spp.), Ericaceae (8 spp.), Juncaceae (8 spp.), and Lentibulariaceae (6 spp.) were the six most species-rich vascular plant families encountered. A guide to the littoral zone flora of Carolina bay lakes is presented herein, including dichotomous keys, species accounts (including abundance, habitat, phenology, and exsiccatae), as well as images of living species and vouchered specimens.

Keywords

North American southeastern Coastal Plain lakes, floristics, aquatic, emersed vegetation

Introduction

Carolina bays are shallow elliptical depressions of disputed origin aligned in a northwest-southeast direction on the Atlantic Coastal Plain of the eastern United States from the Delmarva Peninsula to southern Georgia (Tuomey 1848, Glenn 1895, Melton and Schriever 1933, Prouty 1952, LeBlond 1995, Sharitz 2003). In southeastern North Carolina, several large, natural, lacustrine systems exist within the geomorphological features known as Carolina bays. Within the current distribution of Carolina bays, Bladen and Columbus counties in North Carolina contain the only known examples of Carolina bay lakes. Carolina bay lakes can be split into two major divisions, the “Bladen Lake Group”, which are dystrophic to oligotrophic and relatively unproductive, and Lake Waccamaw, which stands alone in Columbus County and is known for its high productivity, species richness, and rates of endemism (Weiss and Kuenzler 1976, Casterlin et al. 1984, LeBlond 1995, North Carolina Division of Water Quality, Environmental Sciences Section, Intensive Survey Unit 2009, North Carolina Division of Water Quality, Environmental Sciences Section, Intensive Survey Unit 2012, Schafale 2012).

Although there have been several studies conducted on these unique lentic (freshwater) systems (Prouty 1935, Eyles 1941, Hubbs and Raney 1946, Frey 1949, Frey 1951a, Frey 1951b, Frey 1954, Louder 1962, Casterlin et al. 1984, Newman and Schalles 1990; see also Suppl. material 1), none have focused comprehensively on their vegetation. Several manuals, guides, and broader floristic works are available on wetlands of North Carolina and the eastern United States (Suppl. material 2), but few floras have followed the guidelines and recommendations of Palmer et al. (1995) and Denslow et al. (2010) and documented the site-specific aquatic flora of wetlands, streams, rivers, ponds, or lakes in North Carolina (Sieren and Warr 1992, Warren et al. 2004). Nifong (1998) estimated the occurrence of 620 functionally intact, unaltered, Carolina bays remaining in the Coastal Plain of the Carolinas, and an annual rate of loss of about 36 functionally intact North Carolina bays to development, agriculture, silviculture, and other means. It is imperative that the few remaining unaltered bays be studied, especially Carolina bay lakes, considering that with increasing demotechnic growth (Wetzel 2001) and insecure protection status of isolated wetlands (Sharitz 2003), many freshwater systems, including Carolina bays and bay lakes, face an uncertain future.

A narrow time frame exists to study the few remaining natural freshwater systems not affected by severe degradation. Denslow et al. (2010) found only one aquatic flora (Sieren and Warr 1992) within the state of North Carolina between the years 1834−2009, showing a neglect of aquatic systems in floristic studies within the state. To help fill this gap in knowledge, the objectives of this study were to (1) inventory the littoral zone vascular flora of Carolina bay lakes through the collection of voucher specimens, (2) provide a comprehensive checklist of the littoral zone vascular flora based on integration of new and historic collections and reports, and (3) create an illustrated guide based on the checklist.

Background

Lake Ecosystems and Abiotic Factors

Catchment Area

Lakes (also referred to as lentic systems along with ponds) exhibit physical and chemical characteristics unique to the soils, vegetation, and land use activities present on immediately surrounding lands; thus, no two lakes are exactly the same (Moss et al. 1996, Brönmark and Hansson 2005). All lakes occur within catchment areas. A catchment area can also be referred to as a watershed or drainage basin, which is simply the zone of land surrounding a lake that drains precipitation into the lake basin (Brönmark and Hansson 2005). The area, geology, edaphic (soil) properties, land use, and vegetation of catchment areas affect the acidity, water color, nutrient input, and chemical composition of lakes (Wetzel 2001, Brönmark and Hansson 2005). Large catchment areas have a more pronounced impact on the chemical properties of lakes because they drain more precipitation, and thus the potential for more nutrients, into the lake basin. Consequently, land use activities that release excessive nutrient inputs into large catchment areas (e.g., intensive agriculture) are likely to cause eutrophication (Casterlin et al. 1984, Brönmark and Hansson 2005).

Water Color

The observed color of natural lake waters is caused by the selective absorption of wavelengths as light penetrates through the water column (Wetzel 2001). Organic matter (i.e., dead and decomposing plant and animal parts) is the principal determinant of water color in lakes (Juday and Birge 1933, Rasmussen et al. 1989, Brönmark and Hansson 2005). Due to differences in wavelength absorption, waters with little dissolved organic matter, such as hardwater lakes or glacial streams, appear blue/green, and, in contrast, lakes containing much dissolved organic matter in the form of humic substances (e.g., Carolina bay lakes and bogs) appear yellow/red or “tea-stained” in color. Humic substances are large molecules formed as a result of decomposing organic matter; they are difficult for the microbial community to degrade and are long-lived within the lake system (Brönmark and Hansson 2005).

Trophic status

Trophic status refers to the rate at which organic matter is supplied by or transported into a lake. Humic substances are the most common component in allochthonous organic matter; consequently, wetlands that receive the bulk of their organic matter from allochthonous sources (e.g., Carolina bay lakes, bogs, pocosins) are heavily “tea-stained” and are commonly referred to in the southeastern United States as “black water” lakes, streams, rivers, ponds. Lakes receiving the majority of their organic matter from allochthonous sources have been given the term dystrophic. Dystrophic lakes have low productivity and are often acidic due to large quantities of allochthonous humic input.

Phosphorous is limiting in freshwater systems and is therefore a useful determinant for production. Phosphorous concentrations are easier to quantify than carbon content and production, and, as a result, trophic status is often classified based on phosphorous content (Brönmark and Hansson 2005). Oligotrophic lakes experience low productivity associated with autochthonous carbon production and low levels of phosphorous and nitrogen. Eutrophic lakes experience high productivity associated with autochthonous production and high levels of phosphorous and nitrogen.

pH

The unit commonly used to measure acidity is pH. It is technically defined as the reciprocal of the activity of free hydrogen ions (H+; Covington et al. 1985). Because pH is measured on a logarithmic scale, a change of one unit in pH corresponds to a ten-fold increase in hydrogen ions (Brönmark and Hansson 2005). pH is measured on a scale of 1–14; most lakes possess a pH between 6 and 9, but extreme cases of acidity (1–5) and alkalinity (10–14) also exist depending upon various abiotic and biotic conditions within a lake’s catchment area (see above; Brönmark and Hansson 2005). Geological and hydrological conditions within catchment areas primarily control the pH of lakes; however, acid rain can also affect the pH of lakes. In North America, coal-fired power plants and other industries emit sulfur dioxide (SO2) into the atmosphere. As weather systems make their way across North America from west to east, they pick up this sulfur dioxide (SO2) and deposit it across the landscape in the form of precipitation (i.e., acid rain). The cumulative effects of acid rain deposition on both terrestrial and aquatic systems is known to be most severe in the eastern United States; this is due to the region's geographic location in relation to broad-scale weather paterns and industries emitting sulfur dioxides (Schindler 1988).

Photosynthesis and respiration are also known to affect the pH of waters by influencing the amount of carbon dioxide (CO2) in the water column. When CO2 is taken up and stored by aquatic macrophytes, phytoplankton, and algae during photosynthesis, free hydrogen ions (H+) are neutralized or taken up by carbonates, bicarbonates, and hydroxides, causing a reduction in H+ and thus a higher pH. Respiration adds CO2 into the system, thus releasing free H+ into the water column and lowering the pH (Brönmark and Hansson 2005). Because photosynthesis and respiration can cause fluctuating differences in pH within a 24-hour cycle, alkalinity is typically considered to be a better measurement of a lake's acidification status (Brönmark and Hansson 2005).

Alkalinity

Alkalinity refers to a lake's ability to neutralize strong inorganic acids (i.e., it is a measure of how sensitive a lake is to acidification). It is now used synonymously with acid neutralizing capacity (ANC; Wetzel 2001). Today, alkalinity is generally expressed in milliequivalents per liter (meq/L), but has commonly been recorded in the past in milligrams per liter (mg/L; Brönmark and Hansson 2005). Lakes with an alkalinity above 0.5 meq/L have good buffering capacities, whereas lakes with alkalinities below 0.01 meq/L have little or no buffering capacities (Wetzel 2001Brönmark and Hansson 2005). Lakes with low alkalinities are susceptible to drops in pH with only small additions of acid (H+), whereas lakes with high alkalinities can withstand the addition of acid (H+) into their systems without proportional drops in pH (Brönmark and Hansson 2005).

Wetzel (2001) noted that the property of alkalinity in most fresh waters is imparted by the presence of carbonates (i.e., carbonate, bicarbonate, calcium carbonate). Carbonates and hydroxides remove hydrogen ions (H+) from lakes, thus neutralizing their acidity (i.e., raising the pH to a more basic status). Lake Waccamaw, the largest Carolina bay lake, has a high alkalinity (7.0−12 mg/L or 0.14−0.24 meq/L; Weiss and Kuenzler 1976) due to the presence of both subsurface and surficial limestone deposits within and around the lake. As a result, it possesses a neutral to basic pH (6.8−8.5 s.u.) and has the ability to handle larger additions of acid.

Carolina Bays, Bay Lakes, and Pocosins

Carolina Bays

The core concentration of Carolina bays occurs in southeastern North Carolina and northeastern South Carolina (Ross 2003; Fig. 1). Although these depressions share the same elliptical shape, they vary dramatically in length along their long axis from 50 m to 8 km (with some as large as 3,600 ha; Prouty 1935, Thom 1970, Savage 1982, Sharitz and Gibbons 1982). Nifong (1982) suggested that there are fewer than 13,000 bays (unaltered and altered) left in the Coastal Plain of the Carolinas, as opposed to the 400,000 proposed by Prouty (1935). It was not until the early 20th century that researchers fully recognized the magnitude and extent of Carolina bay distribution by the use of airplanes and soon-to-be aerial imagery.

Figure 1.

Core distribution of Carolina bays. Carolina bays are known to occur from the Delmarva Peninsula south to southern Georgia. Although many historical texts frequently cite the distribution range of Carolina bays as occurring from New Jersey south to Florida, the more narrow range from the Delmarva Peninsula to southern Georgia is more accurate. Conversations with state agencies and personnel from all states included in the broader range of Carolina bays confirm their “apparent absence” in southern New Jersey and northern Florida. The core distribution of Carolina bays is located in northeast South Carolina and southeast North Carolina (darker gray). The bays in this region would be considered “classic” Carolina bays (i.e., matching all of the well-known and consistent geomorphological criteria in the literature), whose geomorphology is described well by Prouty (1952) and Ross (2003). Toward the peripheries of the known Carolina bay distribution range, the term Carolina bay tends to be used loosely and is not used in its strictest sense (i.e., depression wetlands; Chick Gaddy, pers. comm.). Figure taken from Ross (2003).

Savage (1982) declared that: “When seen from the air, Carolina bays are an astounding, unforgettable revelation. But though hundreds of thousands lie clearly visible, scattered across the Atlantic Coastal Plain from Maryland to northern Florida, they are often all but unrecognizable to the uninitiated eyes of groundlings”. The first aerial images produced of the Atlantic Coastal Plain exposed Carolina bays to both citizens and scientists on a broad scale; moreover, they initiated a flurry of scientific research on Carolina bay distribution, numbers, origin, vegetation, and soils.

The term bay is used to describe these landscape features not because they commonly contain hydric soils or are inundated with water, but because of the presence of three species of bay tree typically found within and around their elliptical boundaries (i.e., Magnolia virginiana L. [sweetbay; Magnoliaceae], Persea palustris (Raf.) Sarg. [swamp bay; Lauraceae], and Gordonia lasianthus (L.) J. Ellis [loblollybay; Theaceae]. Traditionally, the term “bay” tree has been used when speaking of the laurel trees within the Lauraceae family. While Persea palustris may be properly referred to as a “bay” tree, Gordonia lasianthus and Magnolia virginiana may not (sensu stricto), hence their common names being one word (i.e., loblollybay and sweetbay). Gordonia lasianthus and Magnolia virginiana bear a noticeable morphological resemblence to the laurels of the Lauraceae; thus, they are generally referred to as “bay” trees (sensu lato). North of Virginia, these mysterious landscape features are referred to as Delmarva potholes, bays, or basins (Tiner and Burke 1995, Lide 1997, Sharitz 2003, Tiner 2003). The inability to agree upon a clear-cut definition and universal name for these unique geological features has caused some discrepancy among estimates of bay numbers (Lide 1997).

Collectively, Carolina bays and pocosins represent the largest total acreage of palustrine wetlands in the Carolinas (Wilson 1962, Richardson 1983, Richardson and Gibbons 1993, Nifong 1998). Pocosins occur on the Atlantic Coastal Plain from southern Virginia to northern Florida (essentially the same range as Carolina bays). Unlike Carolina bays, pocosins have been poorly mapped throughout the whole of their range. Wilson (1962) and Richardson (1981) comprehensively mapped the pocosins of North Carolina. It is estimated that ca. 70% of the nation's pocosin habitat occurs in North Carolina and that over 50% of the state's palustrine wetlands are comprised of pocosins (Richardson and Gibbons 2003). Richardson (1981) suggested that ca. 8,300 km2 (3,200 mi2) of unaltered pocosins were drained for other land uses between 1962 and 1979; and ca. 3,700 km2 (1,450 mi2) of unaltered pocosins remained in North Carolina in 1980. Based on the presence of wetland soils (i.e., “soils formed under conditions of saturation, flooding, or ponding long enough during the growing season to develop anaerobic conditions in the upper part” [Vepraskas and Richardson 2001]), North Carolina is estimated to have contained nearly 7.5 million acres (3.03 million hectares) of wetlands prior to European settlement of the state; 95% of these wetlands were located in the Coastal Plain (North Carolina Division of Environmental Management 1994).

Geographic location, soil depth, soil type, surrounding land use, varying hydrology, and fire regimes interact to create vastly different vegetative and wetland assemblages within Carolina bays. Nifong (1998) summarized this diversity, noting that bays included “in some form, virtually every non-marine wetland system found on the southeastern Coastal Plain, including brackish marsh, freshwater pond, freshwater marsh, freshwater prairie, pocosin, bay forest, bog, swamp forest, depression meadow, cypress savanna, and longleaf pine savanna communities, among others”. Other communities found within Carolina bays include Pinus taeda L. (loblolly pine) plantations, cropland, and open lakes (Carolina bay lakes).

Carolina bays can be divided into two classes based on soil substrate: clay-based bays and peat-based bays. The vast majority of Carolina bay literature has referenced peat-based bays, frequently using terms such as “pocosin” or “evergreen shrub bog” to describe the vegetation growing over deep organic soils. However, there are about 27 bays (as of 1982) located in the Carolinas that contain clay subsoil not overlain with sand or peat (Kelley and Batson 1955, Nifong 1982). These clay-based bays are restricted to Cumberland, Scotland, Hoke, and Robeson Counties in North Carolina. The vegetative physiognomy of clay-based bays differs from peat-based bays in that the structure is more open in the former (i.e., they have a sparse overstory of Taxodium and an herbaceous understory composed mostly of herbaceous taxa). However, clay-based bays do share some of the classical Carolina bay morphology features (e.g., elliptical boundaries, varying size, sand rims) with peat-based Carolina bays.

Clay-based bays are species-rich communities, often supporting rare taxa within their boundaries (Nifong 1982). Clay-based bays in high quality condition typically have an open canopy with a species-rich herbaceous understory. Fire and water level fluctuations are two disturbance regimes that account for the diversity found in these bays (Sutter and Kral 1994, Nifong 1998). Peat based bays are more prevalent throughout the Coastal Plain of the Carolinas. Peat-based bays are not as restricted to the inner Coastal Plain and are not as floristically rich as high quality clay-based bays.

Bladen County, North Carolina, is well-known for its many Carolina bays. Nifong (1998) found 617 Carolina bays within Bladen County; of these, 325 were classified as fully vegetated and 292 were classified as cleared (i.e., > 50% of their natural vegetation removed). Bladen County hosts the densest cluster of unaltered bays in the state (the county is fourth densest for bays in any condition). The majority of the bays in Bladen County are found in the Cape Fear River Valley, between the Cape Fear River and the South and Black Rivers. All of these bays are considered peat-based bays. Among extent Carolina bay lakes, all but one occur in Bladen County.

Carolina bays should not be confused with pocosins; they are two distinct physiographic features that just so happen to coexist with one another on the Atlantic Coastal Plain. These two landscape features differ from one another and using the terms synonymously is a common mistake among both laymen and professionals (Ross 2003). The term pocosin originated as an eastern Algonquian term meaning “swamp-on-a-hill” (Richardson 1983). It is defined by Ross (2003) as “a Coastal Plain wetland area of variable shape and size in an area of poor surface drainage whose vegetation is mostly broad-leafed evergreen shrubs and Pinus serotina Michx. growing on organic peaty soils” and by Brinson (1991) as “ecosystems dominated by woody, predominantly evergreen species and that normally occur on histosols (organic peat or muck soils ≥ 40 cm deep) or on soils with a histic epipedon (uppermost soil horizon used to classify a soil)”. Pocosins typically are located on broad, flat, interstream areas or near estuaries where rising sea levels affect their hydrology and hinder their drainage. Although there may be “pocosin-like vegetation” within a Carolina bay, the features are structurally of different origins. Unlike Carolina bays, the origin of pocosins is generally more understood (Whitehead 1972, Whitehead 1981, Brinson 1991, Richardson and Gibbons 1993).

Brinson (1991) attributed pocosin formation and subsequent persistence to two factors: climate and topography. Climate, he attested, “determines the exchange of matter and thermal energy between pocosins and the atmosphere”. The bulk of this exhange is in the form of precipitation, much of which is lost to evapotranspiration following its input. Brinson (1991) added “while the muted topographic relief of the Atlantic Coastal Plain is probably the main contributor to pocosin formation, the feedback between climate and topography is likely essential”. In summary, pocosins have formed in landscape positions with low topgraphic relief where the regional climate and lack of surficial hydrologic connections with adjacent wetland systems interact to form ombrotrophic conditions. Here, organic matter in the form of dead terrestrial vegetation is deposited onto wetland soils and accumlates at a slow, consistent rate through geologic time, resulting in the formation of pocosins.

Historically, the Atlantic and Gulf Coastal Plains supported a heterogeneous landscape of longleaf pine savannas, xeric sandhills, upland mixed-pine hardwoods, pocosins, Carolina bays, bottomland hardwood forests, natural lakes, and black and brown-water river systems (Garren 1943, Christensen 1999). However, it is now a highly fragmented and fire-suppressed region dominated by agriculture, residential developments, and large cities with few large intact parcels of natural ecosystems remaining. Demotechnic growth (Wetzel 2001, Dudgeon et al. 2006), global warming (Smith and Tirpak 1989), increasing agricultural production (Tilman et al. 2002), fire supression (Nowacki and Abrams 2008, Palmquist et al. 2014), urbanization (Terando et al. 2014), shoreline development (Radomski and Goeman 2001, Ford and Flaspohler 2010, Frost and Hicks 2012), and introduction of invasive species (Pimentel et al. 2005) continue to threaten and encroach upon the few “natural”, intact, terrestrial and freshwater ecosystems remaining in the Southeast, including Carolina bays and bay lakes.

Carolina bays are valuable components of our national and state natural heritage (Nifong 1998). Their variable hydrology and size, presence of rare and endemic taxa, and isolated landscape position, make them valuable habitats for southeastern flora and fauna and provide important ecosystem services (Suppl. material 4). Unfortunately, Carolina bays and other palustrine wetland systems have suffered from extensive habitat loss and degradation during the past three centuries (Bennett and Nelson 1991, Mitsch and Gosselink 1993, North Carolina Division of Environmental Management 1996, Kirkman et al. 1996, Nifong 1998). Using 1988 aerial imagery, Nifong (1998) found 8,057 Carolina bays in the state of North Carolina. Of these 8,057 total bays, 6,331 (79%) had more than half of their natural vegetation removed.

Sharitz and Gibbons (1982) and Nifong (1998) suggested several ways to better preserve and manage Carolina bays in the future. For an excellent review on the copious amount of Carolina bay literature available, see Ross (2000), Ross (2003); and for detail specifically about bays in the Carolinas, see Nifong (1998).

Carolina Bay Lakes

Several Carolina bays in southeastern North Carolina contain large (i.e., > 50 hectares) natural lakes within their elliptic boundaries (Frey 1949), thereby giving them the name Carolina bay lakes. Each lake is located in the southernmost portion of the elliptical feature known as a Carolina bay (Fig. 2). The northern portions of the bays (i.e., the portions not inundated by lake waters) contain organic, peaty soils and a unique vegetative assemblage comprised of bay trees (Gordonia lasianthus, Magnolia virginiana, Persea palustris), ericaceous shrubs (e.g., Chamaedaphne calyculata (L.) Moench, Eubotrys racemosa (L.) Nutt., Kalmia L., Lyonia Nutt., Rhododendron L., Vaccinium L., Zenobia pulverulenta (W. Bartram ex Willd.) Pollard), and several other species well-associated with nutrient-poor soils (e.g., Chamaecyparis thyoides (L.) Britton, Sterns & Poggenb., Nyssa biflora Walter, Pinus serotina, and Smilax laurifolia L.).

Figure 2.

Position of Carolina bay lakes within Carolina bays. Carolina bay lakes are located in the southeasternmost portions of Carolina bays. The northern portions of the bays (i.e., the portion not inundated by lake waters) support shrub-bog plants over organic soils. Here, Salters (top left) and Jones (middle right) Lakes exemplify the typical bay lake position within Carolina bays. Aerial imagery, transportation, and hydrography layers obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

Nine Carolina bay lakes (i.e., Bakers Lake, Bay Tree Lake, Horseshoe Lake, Jones Lake, Lake Waccamaw, Little Singletary Lake, Salters Lake, Singletary Lake and White Lake) are known to exist within the known distribution of Carolina bays. All nine lakes occur in Bladen and Columbus counties, North Carolina (Frey 1949, LeBlond 1995, LeBlond and Grant 2005; Fig. 3). Carolina bay lakes, with the exception of Lake Waccamaw and White Lake, are nutrient poor because they receive the bulk of their hydrologic inputs in the form of precipitation. These lakes are also characteristically dystrophic due to the dominance of organic soils within their catchment area. Organic soils do not allow for the rapid decomposition of plant and animal matter, resulting in the high amount of humic substances found in the water column.

Figure 3.

Geographic location of all nine Carolina bay lakes (green text boxes). Bladen County (light yellow) supports eight of the nine Carolina bay lakes known to exist; all eight lakes occur within the Cape Fear River Valley between the Cape Fear River and South River. Bay Tree Lake is the largest Carolina bay lake in Bladen County; the smallest is Bakers Lake. Lake Waccamaw is the largest Carolina bay and bay lake in North Carolina and is the only bay lake known to exist in Columbus County (tan). Baseline vector data obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map Produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

Although some Carolina bays may contain shallow marshes or ponds (Bennett and Nelson 1991, Nifong 1998), these are not considered lakes. There is no universally accepted technical definition that distinguishes a lake from a pond (Heinonen et al. 2008); however, it seems reasonable to accept as distinguishing that lakes have a clearly defined littoral and profundal zone, a larger overall size (>8 hectares), a shoreline exposed to wave dynamics, greater water depth, a mixing of the water column by wind induced turbulence, and the ability to retain the bulk of their water volume even in years of drought (Cowardin et al. 1979, Moss et al. 1996, Williams et al. 2004, Biggs et al. 2005, Brönmark and Hansson 2005).

Carolina bays are considered to be geographically isolated wetlands with their primary water source coming directly from precipitation (Sharitz 2003, Tiner 2003). Although the vast majority of Carolina bays lack surface water connections to outside aquatic systems, Carolina bay lakes are an exception. Carolina bay lakes all contain drainage outlets--usually along their southern shorelines, but in the northwest for White Lake (Frey 1949)--that release excess water into the Cape Fear and Waccamaw River drainages during periods of high precipitation. However, during years of scarce rainfall, these lakes are more or less isolated from surrounding lotic systems and are confined to their basins (N. Howell, pers. obs.).

Lacustrine Zonation (derived from Wetzel 2001)

Lakes, including Carolina bay lakes, can be divided into distinct transitional zones, moving from the shoreline to the center of the lake (Fig. 4).

Figure 4.

Lacustrine zonation. EPI = epilittoral zone, EU = eulittoral zone. Aerial imagery obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map Produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014). Illustration (top right) by Nathan Howell.

(1) Epilittoral zone: The zone that lies entirely above the lake surface and is not influenced by the spray of surf. This zone can be thought of as the terrestrial or upland zone; the highest water levels never reach it and it is not affected by lakeshore dynamics or hydrology.

(2) Supralittoral zone: The zone that lies entirely above the lake surface and is influenced by the spray of the surf.

(3) Eulittoral zone: The zone encompassing the entire region of the shoreline from the highest and lowest seasonal water levels. This zone experiences natural disturbances such as water level fluctuations and wave dynamics.

(4) Infralittoral zone: This zone is subdivided into three zones in relation to the occurrence and distribution of the major classes of aquatic macrophytes: upper infralittoral zone where emergent rooted macrophytes persist; middle infralittoral zone where floating-leaved rooted macrophytes occur; and lower infralittoral zone where submersed-rooted, adnate, or free-floating macrophytes occur. The eulittoral and infralittoral zones collectively constitute the littoral zone.

(5) Littoriprofundal zone: The zone occupied by photosynthetic algae and bacteria, often associated with the metalimnion (i.e., the stratum between the epilimnion and hypolimnion representing a marked thermal change; also synonymous with thermocline) of stratified lakes.

(6) Profundal zone: The zone that consists of the remainder of the vegetation free sediments.

The Littoral Zone

The littoral zone of lakes (i.e., the eulittoral and infralittoral zones) is an important transition zone between adjacent uplands and the deeper pelagic area of the lake. This zone contains vascular macrophytes (i.e., aquatic vascular plants large enough to see with the naked eye) that have evolved from their terrestrial ancestors to cope with the physical and physiological demands of persisting in an aquatic environment (Sculthorpe 1967, Wetzel 2001, Brönmark and Hansson 2005, Keddy 2010). The vascular macrophytes and coarse woody debris that exist in this zone provide critical habitat for zooplankton, photosynthetic and heterotrophic microflora, macroinvertebrates, herpetofauna, avifauna, fish, and mammals (Brusnyk and Gilbert 1983, Pieczynska 1990, North Carolina Division of Environmental Management 1996, Wetzel 2001, Keddy 2010, Ewert et al. 2011). The littoral zone is characterized by having high productivity, including some of the highest rates of organic matter synthesis in the biosphere (Wetzel 2001).

Aquatic Macrophytes (derived from Wetzel 2001)

Aquatic macrophytes may be divided into four classes. Moving from the shoreline out to deeper water, these classes are as follows [taxa vouchered or reported from Carolina bay lakes are indicated by c]:

(1) Emergent macrophytes: Species rooted in saturated and inundated soils with a water depth up to 1.5 meters; root systems remain in anoxic soil conditions while leaves and reproductive organs stay above the water surface. These plants are often rhizomatous, stoloniferous, or cormous with the potential to reproduce asexually. Heterophyllous (i.e., when a plant exhibits vegetative polymorphism, having morphologically different submersed and aerial organs) species may also be emergent. Examples of genera that may be grouped in this category include Carex L.c, Cephalanthus L.c, Cladium P. Brownec, Juncus L.c, Panicum L.c, Pontederia L.c, Rhynchospora Vahlc, Scirpus L.c, and Typha L.

(2) Floating-leaved macrophytes: Species rooted in the substratum with floating leaves attached to long flexible petioles or on short petioles attached to an ascending stem.

Submersed leaves precede the floating leaves in heterophyllous species. Reproductive organs remain atop or above the water surface. Examples of genera grouped into this category include Brasenia Schreb.c, Nelumbo Adans.c, Nuphar Sm.c, Nymphaea L.c, Nymphoides Ség.c, and Potamogeton Lc.

(3) Submersed macrophytes: Species that remain completely submersed in the water column, but are rooted to the substratum. Leaf morphology is highly variable in this group, from finely dissected to very broad, and reproductive organs may be emersed, floating, or submersed. Examples of genera included in this group are Ceratophyllum L., Isoetes L., and Myriophyllum Lc.

(4) Freely floating macrophytes: Species that remain unattached to the substratum and are completely dependent upon the nutrients in the water column for survival. Reproductive organs may be floating or aerial. Examples of genera include Azolla Lam., Eichhornia Kunth, Hydrocharis L., Limnobium Rich., Trapa L., and Utricularia Lc.

Factors affecting Aquatic Macrophyte Richness in Lakes

Lacoul and Freedman (2006) provided a thorough review on how various environmental influences affect aquatic plants in freshwater systems. A few of these environmental factors are reviewed below.

Latitude

It is well known that generally the number of species occuring at the equator greatly exceeds that of the temperate and northern latitudes (Edmonds 1997). Although this general rule applies across most groups of taxa, it does not seem to apply to aquatic plants. Crow (1993) found that aquatic plants are more diverse in temperate rather than tropical latitudes. When comparing temperate wetland floras to those of tropical climes, this pattern is reinforced (Stuckey 1975, Henry and Scott 1984, Peet and Allard 1993, Ruch et al. 2009). Because Carolina bay lakes differ little in latitude, this factor does not significantly affect species richness in these systems.

pH and Alkalinity

Peat-based Carolina bays are known to have acidic (< 7 pH), nutrient poor, organic soils (Daniels et al. 1984, Leab 1990, Newman and Schalles 1990). In many respects, these isolated wetlands of the Southeast are quite similar to the peatlands of the northern United States and Canada. Floristic diversity in peatlands has been shown to increase with increased levels of calcium and alkalinity in the groundwater (Glaser et al. 1990, Vitt and Chee 1990). Similarly, aquatic macrophyte richness of lakes tends to be lower in unproductive lakes with low pH (e.g., Carolina bay lakes) and higher in more productive lakes with higher alkalinities (Roelofs 1983, Roberts et al. 1985, Rørslett 1991, Dodson et al. 2000, Vestergaard and Sand-Jensen 2000, Søndergaard et al. 2005).

Water Color

Waters with increased levels of humic substances are typically, dystrophic, acidic, and tea-stained. Tea-stained waters are not as transparent as lakes with low humic substances, thus humic lakes have a shallow euphotic zone and a narrow littoral zone, reducing the abundance and depth at which aquatic macrophytes may grow (Spence 1982). Vestergaard and Sand-Jensen (2000) also saw decreased richness in aquatic macrophytes when water transparency was low. An excellent example of how increased humic substances affect water transparency and macrophyte richness and composition can be seen when comparing White Lake to the other Carolina bay lakes. White Lake is an oligotrophic lake with transparent water due to the presence of natural springs on the lake floor. Secchi depths commonly reach to the bottom of the lake (3m/10 ft) and submerged aquatic macrophytes are able to colonize the deepest portions of the lake with ease (i.e., the euphotic zone is deep compared to the other bay lakes).

Hydrography

Frey (1949) documented the morphometry and hydrography of the Carolina bay lakes and determined that the southern portions of the lakes possessed a gentle, tapering hydrography while the northern portions possessed a steep hydrography. Floristic inventories by the first author confirm that aquatic macrophyte richness is higher along southern shorelines; so much so, that the surveying of northern shorelines was abandoned early in the life of the project. A broad sandy terrace occurring along the southern shore of Lake Waccamaw (Fig. 5) creates a wide littoral zone compared to other Carolina bay lakes. This stretch of shoreline, with its gentle hydrography, is known to support over 140 species of wetland plants, while the Bladen lakes, with their comparatively steeper hydrography, are known to support < 55 wetland plant taxa (see floristic summary).

Figure 5.

Broad, shallow, sandy terrace along Lake Waccamaw’s southern shoreline. The gentle relief of this terrace creates a wide littoral zone. Wide littoral zones are more floristically diverse and contain more available area for the establishment of aquatic macrophytes. Alternatively, narrow littoral zones do not have much area for the establishment of aquatic macrophytes and are species-poor. Aerial imagery obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

Lake Size

As a general rule, species richness usually increases with increasing area (Arrhenius 1921, Williams 1964, Connor and McCoy 1979, Rosenzweig and M.L. 1995, Søndergaard et al. 2005). Findlay and Houlahan (1997) found that species richness increased with area sampled for birds, mammals, hepertofauna, and plants in southeastern Ontario wetlands. Results from this work also support these findings with Bakers Lake (i.e., the smallest bay lake) supporting the least diverse littoral zone flora and Lake Waccamaw (i.e., the largest bay lake) supporting the most species-rich littoral zone flora. Other large natural lakes of North Carolina Coastal Plain (e.g., Lake Phelps, Lake Mattamuskeet, Lake Waccamaw) are known to support diverse shoreline floras, more so than the smaller lakes of the region (Lynch and Peacock 1982, Schafale 2012; N. Howell, pers. obs.).

Water Level Variation, Disturbance, and Soil Fertility

Keddy and Fraser (2000) summarized factors that govern littoral zone diversity irrespective of geographic location or size. Three environmental factors (i.e., water levels, soil fertility, and disturbance) govern the composition and floral diversity of littoral zones. Shorelines exposed to intermediate levels of natural disturbances will support a richer flora than those experiencing little to no disturbances and those experiencing extremely harsh disturbances. Natural disturbances may include wave action, ice scour, water level fluctuations, fire, or grazing. If water level fluctuations were absent from a lake or similar waterbody (e.g., in a permanently impounded pond), a two-staged littoral zone would result, with aquatic macrophytes in the aquatic zone and shrubs and trees in the terrestrial zone. Under long-term water level fluctuations, a multi-staged littoral zone would result, leading to increased heterogeneity and a richer flora. Keddy and Fraser (2000) attested that “simply changing water levels from one year to the next doubles the number of vegetation types”. Rørslett (1991) observed that northern European lakes experiencing water level fluctuations of 1–2 meters per year showed greater macrophyte richness than sites experiencing little or intense disturbances. Carolina bay lakes historically would have experienced long-term water level fluctuations, but the installation of water control structures (i.e., dams) in some of the lakes outlet channels has resulted in more stabilized systems (N. Howell, pers. obs.).

Shorelines exposed to frequent disturbances typically have silt and clay stripped from them; and consequently, contain few nutrients. Sheltered shorelines receive clay and silt deposits and therefore contain a higher nutrient content. Foreshores will have a distinct vegetative community characterized as having low biomass and rare species, while backshores (bays or backwater areas sheltered from disturbance) will support a higher biomass community composed of a few clonal dominants (Keddy 2010). Macrophyte richness is always higher in areas of intermediate disturbance. Eutrophification of littoral zones causes increased soil fertility, which increases biomass and negatively impacts macrophyte richness and rare plant taxa.

Study Sites

Bakers Lake

Bakers Lake (30.35 hectares; 75 acres) is a small, privately owned, Carolina bay lake, located in northwestern Bladen County between Little Singletary Lake and the Cape Fear River north of Thoroughfare Bay, ca. 1.5−2 miles east of the intersection of SR 1318 (Old River Road) and SR 1320 (Middle Road; LeBlond and Grant 2005; Fig. 6). This site is located along the northwest boundary of the Bladen Lakes Macrosite, a large tract of undeveloped and relatively unfragmented land between the Cape Fear, South, and Black River systems (LeBlond and Grant 2005; Figs 7, 8). The macrosite extends from southern Cumberland County, through Bladen County, and into southwestern Pender County. This large area is given the name “macrosite” because it contains numerous “standard sites” (i.e., smaller tracts of land with high ecological integrity) that are strongly geographically associated with one another. The majority of the macrosite is located in Bladen County and contains the largest concentration of unaltered, intact, Carolina bays.

Figure 6.

Bakers Lake and surrounding lands. Bakers Lake is located in northern Bladen County and is surrounded by a mix of agriculture and forestland. Aerial imagery, transportation, and hydrography layers obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

Figure 7.

Bladen Lakes Macrosite (vector). The Bladen Lakes Macrosite (hatched pattern) is a large area encompassing parts of southern Cumberland County, eastern Bladen County, and northwest Pender County. Historically, macrosites were established by the North Carolina Natural Heritage Program (NCNHP) in efforts to identify large, intact, natural areas that withheld numerous other smaller natural areas within their boundaries. The NCNHP no longer uses macrosites as viable natural area boundaries, but it is useful to show the extent of the Bladen Lakes Macrosite boundary. When moving from north to south, the lands are as follows: Bushy Lake State Natural Area (teal green), Suggs Mill Pond Gameland (light mint green), Bladen Lakes State Forest (forest green), Jones Lake State Park (pink), Bay Tree Lake State Park (orange), and Singletary Lake State Park (yellow). Lake Waccamaw State Park (neon green) can be seen farther south along with Friar and Brown Marsh Swamps in Columbus County. Baseline vector data obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

Figure 8.

Bladen Lakes Macrosite (ortho). The North Carolina Natural Heritage Program no longer uses macrosites as viable natural area boundaries, but here it is useful to show the extent of the Bladen Lakes Macrosite boundary. Note the large areas of fragmented land surrounding the macrosite and the relatively unfragmented land within the boundaries of the macrosite. This large tract of land contains one of the largest remaining portions of intact unaltered Carolina bay complexes known to exist. Aerial imagery, transportation, and hydrography layers obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

Dr. Clemuel Johnson and wife Nancy Johnson, of Elizabethtown, have owned Bakers Lake and surrounding lands (451.40 hectares; 1,155.45 acres) since 1980. Prior to the Johnson’s ownership, Agnes Holden Williams owned the lake and surrounding lands. Ms. Williams’ father acquired the land from an unknown seller during the early 20th century. This seller was able to successfully purchase the lake before 1929, when North Carolina legislation mandated that all lakes greater than 50 acres in size be made property of the state.

Bakers Lake forms the headwaters of Phillips Creek, which drains southward into the Cape Fear River. Bakers Lake Natural Area (i.e., Bakers Lake bay and immediate surrounding lands) is known to support five natural community types (i.e., Pond Pine Woodland – Typic Subtype (S3,G3), Peatland Atlantic White Cedar Forest (S1,G2), Low Pocosin – Gallberry/Fetterbush Subtype (S2,G2), Sand Barren – Typic Subtype (S2,G2), and Natural Lake Shoreline – Cypress Subtype (S2,G3; LeBlond and Grant 2005). Bakers Lake has been known to support heron rookeries and small populations of the state rare Anhinga (Anhinga anhinga [W2; S3B, G5]; LeGrand et al. 2014) during the spring and summer months (S. Clark, pers. comm.; N. Howell, pers. obs.). In addition, the site provides important stopover habitat for large flocks of migrating waterfowl (e.g., Aix sponsa [Wood Duck], Anas americana [American Widgeon], Anas clypeata [Northern Shoveler], Anas crecca [Green-winged Teal], Anas discors [Blue-winged Teal], Anas platyrhynchos [Mallard], Anas strepera [Gadwall], Aythya collaris [Ring-necked Duck], Aythya valisineria [Canvasback], Branta canadensis [Canada Goose], Bucephela albeola [Bufflehead], Lyphodytes cucullatus [Hooded Merganser], Oxyura jamaicensis [Ruddy Duck; G. German and S. Clark, pers. comm; N. Howell pers. obs.).

Anthropogenic disturbances (i.e., silvicultural practices, dam installation in the outflow channel, agricultural fields, confined animal feeding operations (CAFOs), fire supression, and rural residential development) have either been documented on site or on adjacent properties (LeBlond and Grant 2005; S. Clark, pers. comm.). These disturbances have lowered the integrity of several of the aforementioned natural community types within and adjacent to Bakers Lake Natural Area (N. Howell, pers. obs.), but restoration potential is still relatively high. The installation of a flashboard riser system in the outflow channel has altered the natural hydrology of the lake and caused natural water level fluctuations to essentially cease. Following the installation of the dam, the lake consistently stays at a high level, thus narrowing the littoral zone and forcing aquatic macrophytes to occur at or just below the maximum annual high water mark (N. Howell, pers. obs.).

The water quality of Baker’s Lake has not been formally tested by state agencies, but appears high in humic substances (N. Howell, pers. obs.) and the chemistry is likely similar to that of the other Bladen lakes. The lake is here considered dystrophic and relatively unproductive.

Bay Tree Lake

Bay Tree Lake (formerly Black Lake; 588.81 hectares; 1,455 acres) is a large, state-owned Carolina bay lake, located in east-central Bladen County along NC Hwy 41 east of White Lake and west of NC Hwy 210. Bay Tree Lake is part of Bay Tree Lake State Park, a 1,006.85 hectare (2,488 acre) park that includes Bay Tree Lake bay and large parcels of land lying to the north and west of Bay Tree Lake (Fig. 9).

Figure 9.

Bay Tree Lake State Park (highlighted in green) and surrounding lands. Lands surrounding Bay Tree Lake State Park to the south are privately owned and have been partially converted to agriculture. Black Creek Bay and several others in the vicinity have been cleared of their original vegetation and converted to agriculture (primarily blueberry farms in this area). Historically, Horsepen Bay was a peat-filled Carolina bay. During the development of the residential community seen along the northeast shoreline (Bay Tree Resorts), it was turned into a body of open water. Aerial imagery, transportation, and hydrography layers obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

The North Carolina General Assembly passed legislation in 1911 confirming the status of Bay Tree Lake as a state-owned public trust resource (North Carolina Division of Parks and Recreation, Planning and Development Section 2006b). Historically, Bay Tree Lake was not included within the original natural area site boundary determined by the North Carolina Natural Heritage Program (NCNHP) due to high levels of shoreline disturbance. Today, the lake is considered part of the natural area due to the presence of three rare dragonflies (Gomphus australis [Clearlake Clubtail], Gomphus cavillaris brimleyi [Brimley’s “Sandhill” Clubtail], and Progomphus bellei [Belle’s Sanddragon]) that utilize the lake throughout their life cycle.

In January 1965, a private land development group had the option to purchase 5,665.59 hectares (14,000 acres) of land surrounding Bay Tree Lake with the intent of creating an inland resort community (North Carolina Division of Parks and Recreation, Planning and Development Section 1996a). Later that year, a proposal was constructed and sent to the North Carolina Department of Conservation and Development concerning the drainage of Bay Tree Lake. The purpose for draining the lake was to improve the quality of the water and lake bottom for recreational purposes (e.g., swimming and boating). Permission to lower lake levels 4 feet was granted in 1965 and in January of 1966, the development group made a request to completely drain the lake where peat deposits and debris could be taken from the lake bottom (North Carolina Division of Parks and Recreation, Planning and Development Section 1996a).

The purpose of the drainage project was to release tannic, tea-colored, waters from the lake and divert all incoming tannic waters from a northerly adjacent swamp to below the outflow channel. Drainage of the lake was completed in the winter of 1966. The lake remained dry for 5 years while developers removed debris and peat deposits and imported large quantities of white sand, which would later be distributed around the entirety of the lakeshore. In 1970, the lakes outflow channel was plugged and the lake began to refill (North Carolina Division of Parks and Recreation, Planning and Development Section 1996a). After two years, the lake had nearly reached its original water levels. Shortly after residential lots went for sale, a breach of the lake rim occurred and tea-stained waters were allowed to re-enter the lake. The breach was plugged within 24 hours, but the lake had already returned to its original dystrophic condition (North Carolina Division of Parks and Recreation, Planning and Development Section 1996a). The lake has not been significantly altered since and remains in a dystrophic condition to this day.

Bay Tree Lake State Park contains five natural community types (Mesic Pine Savanna – Coastal Plain Subtype [S2,G2G3]; Sand Barren – Typic Subtype [S2,G2]; Small Depression Drawdown Meadow – Typic Subtype [S2S3,G2?]; Small Depression Pocosin – Blueberry Subtype [S2,G3?]; and Xeric Sandhill Scrub – Typic Subtype [S3S4,G3?]. A Natural Lake Shoreline community was not assigned to Bay Tree Lake by the NCNHP due to the shoreline’s disturbance history. The present authors agree with this determination and have chosen not to assign a natural lake shoreline community to this site. However, it is worth noting that the shoreline flora of Bay Tree Lake differs only slightly from the other Bladen Lakes.

Bay Tree Lake forms the headwaters of Lake Creek, a small blackwater creek that drains southeast to the South River (the boundary between Bladen and Sampson counties). Much of the land surounding Bay Tree Lake State Park has been cleared for agriculture (particularly blueberry farms) and has limited the landscape connectivity between it and other intact natural areas. Several bay complexes occur in the immediate vicinity of Bay Tree Lake including Beagle Bay, Black Creek Bay, Causeway Bay, Cooley Bay, Horsepen Bay (now an artificially created lake/pond), Floodgate Bay, Kelso Bay, and Spring Bay. A residential resort community is located along the north and east shorelines of the lake. The boundaries of this community have continued to extend around the east and southeast shorelines. Residential development, agricultural expansion, severe offroad vehicle use, and fire supression are the primary threats to biological diversity within and around Bay Tree Lake State Park (N. Howell pers. obs.). Available water quality parameters for Bay Tree Lake are provided in Table 1.

Water Quality Data for Bay Tree Lake (Bladen County, North Carolina). Frey (1949) sampled Bay Tree Lake 6 times during the Summer and Fall of 1947. Weiss and Kuenzler (1976) sampled Bay Tree Lake twice in 1974 (March 22 and June 6) and 4 times in 1975 (April 7, June 10, August 5, October 6). North Carolina Division of Water Quality, Environmental Sciences Section, Intensive Survey Unit (2009) (DWQ) sampled Bay Tree Lake 4 times in 2008 (June 24, July 29, August 18, October 2). Value ranges have been provided where applicable to show variability. Units are as follows: km2 = squared kilometers, ha = hectares, km = kilometers, m = meters, °C = degrees celsius, mg/L – milligrams per liter, meq/L = milliequivalents per liter, s.u. = standard units, μg/L = micrograms per liter.

Frey (1949)

Weiss & Kuenzler (1976)

DWQ 2009

Trophic Status

Dystrophic

Watershed Area (km2)

10.36

Surface Area (ha)

573.84

Max Width (km)

1.77

Max Length (km)

3.05

Max Depth (m)

1.83

Mean Depth (m)

0.9

Secchi Depth (m)

0.55

0.3−0.4

1.4−1.8

Min Temp. (°C)

13.3

23.2

Max Temp. (°C)

30.5

30

Dissolved Oxygen (mg/L)

6.4

7.1−10.9

6.8−8

Alkalinity (meq/L)

0.159−0.231

pH (s.u.)

4.4

6.3−7.1

4.1−4.5

Total N (mg/L)

0.48−1.568

Total P (mg/L)

0.13−0.238

Chlorophyll-A (μg/L)

2−6

Horseshoe Lake

Horseshoe Lake (also known as Suggs Mill Pond; 109 hectares; ca. 270 acres) is an irregularly shaped Carolina bay lake located in northern Bladen County south of Bushy Lake State Natural Area, east of Little Singletary Lake, north of SR 1325 (Gum Springs Rd), and west of SR 1002 (Old Fayetteville Rd). Horseshoe Lake is one of two Carolina bay lakes within Suggs Mill Pond Game Land (4469.34 hectares; 11,044 acres; Fig. 10), the other being Little Singletary Lake. Suggs Mill Pond Game Land is owned by the State of North Carolina and the North Carolina Wildlife Resources Commission (NCWRC) and is located in northern Bladen County and southern Cumberland County. This game land is located in the northwestern portion of the Bladen Lakes Macrosite and contains one of the largest remaining examples of unaltered Carolina bay complexes.

Figure 10.

Suggs Mill Pond Game Land (outlined in green) and surrounding lands. Lands north of the red dividing line occur in Cumberland County while lands south of the red line occur in Bladen County. Suggs Mill Pond Game Land contains two large bay lakes within its boundary. Little Singletary Lake is located along the western boundary of the property and Horseshoe Lake (aka Suggs Mill Pond) is located in the center of the property. Aerial imagery, transportation, and hydrography layers obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

The state first gained rights to the property in 1994 when a 62-acre (25 ha) parcel was donated to the NCWRC from Canal Woods Industries. Thereafter, much of the remaining property was purchased from Canal Woods. The fact that Horseshoe Lake and Little Singletary lake were not owned by the state of North Carolina until the mid-1990s suggests that these lakes were involved in a similar ownership situation as Bakers Lake (i.e., these lakes must have been privately owned prior to 1929 when legislation mandated that all lakes greater than 50 acres (20.2 ha) in size be released to the state of North Carolina). Suggs Mill Pond Game Land is one of four North Carolina game lands enrolled in the Cooperative Upland habitat Restoration and Enhancement program (CURE), where management for early successional habitat is the top priority (Allen et al. 2015). Traditionally, hunters and fishermen were primary users of Suggs Mill Pond Game Land, but an increasing number of non-traditional users (i.e., birders, canoers, hikers, photographers, and researchers) visit the site regularly.

The largest bay on site contains a horsehoe-shaped artificial impoundment (Horseshoe Lake). Horseshoe Lake forms the headwaters of Ellis Creek, which drains southwest to the Cape Fear River. Although an old milldam currently maintains Horseshoe Lake, it is thought that a smaller body of open water may have been present prior to the dam’s installation in the late 19th or early 20th centuries. Horseshoe Lake was formed subsequent to the dam installation, as water levels began to rise into the peat-filled Carolina bay. Today, it is best described as a semi-permanent impoundment; however, the presence of floating bogs within the lake makes it unique from other semi-permanent impoundments in North Carolina. Parts of the lake support patches of the rare floating bog community (the largest extent known from the state), which is dominated by sedges, orchids, carnivorous plants, and ericaceous shrubs. Other portions comprise the Coastal Plain Semipermanent Impoundment community, which is characterized by open water, dominated by floating-leaved macrophytes, and a sparse overstory of Taxodium ascendens Brongn.

The floating bog community type is quite unique. Manifestations of this community type occur just above the water surface and range in size from ca. 10 × 10 m to a few hectares in size (N. Howell, pers. obs.). Some bogs may contain well-developed herbaceous vegetation in addition to small (e.g., < 3 m tall) trees of Chamaecyparis thyoides, Nyssa biflora, and Taxodium ascendens, while others contain a strictly herbaceous component. Exposed portions of peat can be seen around the peripheries of some bogs; here, Drosera intermedia Hayne, Eleocharis baldwinii (Torr.) Chapm. /E. vivipara Link, Pogonia ophioglossoides (L.) Ker Gawl, Utricularia striata Leconte ex Torr., Utricularia purpurea Walter, and other small-statured herbaceous plants can be seen colonizing the apparently young peat formations. Isolated floating bogs (i.e., bogs surrounded by open water and separated from adjacent bogs and upland habitats) of varying size show a consistent zonation pattern. Small statured herbaceous taxa colonize the outer periphery and are slowly replaced by larger herbaceous taxa (Andropogon glaucopsis, Dulichium arundinaceum (L.) Britton, Hypericum virginicum L., Rhexia nashii Small, Rhynchospora alba (L.) Vahl, Rhynchospora inundata (Oakes) Fernald, Xyris fimbriata Elliott, and Xyris smalliana Nash) and woody species (Acer rubrum L., Chamaecyparis thyiodes, Decodon verticillatus (L.) Elliott, Nyssa biflora, and Taxodium ascendens) when moving toward the center. Thus, a dome-shaped appearance is typically seen.

Few examples of floating bogs or mats of vegetation are known to science. The floating peat mats of New Hampshire are most similar to those of Horseshoe Lake. These peat mats possess the same general structure and abiotic conditions as those of Horseshoe Lake and are known to contain several overlapping taxa, inculding Drosera intermedia, Dulichium arundinaceum, Eleocharis R. Br. spp., Hypericum virginicum, Nymphaea odorata W.T. Aiton, Rhynchospora alba, and Utricularia spp. (New Hampshire Division of Forests and Lands 2015).

A separate but similar case of floating vegetation mats, forming as a result of dam installation, has been observed at Goose Creek Reservoir in South Carolina (Hunt 1943). In 1933, a dam was installed on Goose Creek, ca. 12 miles north of Charleston, subsequently flooding historic rice plantations that had reverted to brackish marsh vegetation. Hunt (1943) described the zonation (looking across to the center of the mat from the outer periphery) of a typical floating mat as follows: (1) pioneer zone (i.e., the outer margins of the mats): Alternanthera philoxeroides (Mart.) Griseb., Bidens laevis (L.) Britton, Sterns & Poggenb., Boehmeria cylindrica (L.) Sw., Habenaria repens, Hydrocotyle ranunculoides L.f., Persicaria glabra (Willd.) M. Gómez, and Sacciolepis striata (L.) Nash, (2) the cat-tail/shrub zone: Kosteletzkya pentacarpos (L.) Ledeb., Typha latifolia L., and Salix nigra Marshall, and (3) the main body: Acer rubrum, Apios americana Medik., Decodon verticillatus, Mikania scandens (L.) Willd., Panicum virgatum L. var. virgatum, Persea palustris, Rubus L. spp., and Taxodium distichum (L.) Rich.

The floating “sudd” vegetation of the upper Nile River is also somewhat similar, forming large floating mats of marsh vegetation both along the margins and within the river. Denny (1984) gave a general description of the sudd vegetation as seen only from a boat. Several taxa commonly observed along the margins of the Sudd included: Ceratophyllum demersum L., Cyperus papyrus L., Eichhornia crassipes (Mart.) Solms, Phragmites karka (Retz.) Trin. ex Steud., Typha domingensis Pers., Vossia cuspidata (Roxb.) Griff. A complete checklist of the vascular plants collected from this vegetative study can be found in the attached appendix of Denny (1984).

Eleven natural community types exist within Suggs Mill Pond Game Land, but the low and high pocosin communities are dominant, comprising 48% (2,119.74 hectares; 5,238 acres) of the site (Allen et al. 2015). Lakes and impoundments make up 8.6% (381.21 hectares; 942 acres) of the total acreage of the game land. Fair to high quality landscape connections exist between Suggs Mill Pond Game Land and adjacent natural areas within the Bladen Lakes Macrosite (i.e., Bushy Lake State Natural Area, Charlie Long Mill Pond/Big Colly Bay Natural Area, Jessups Pond, Mill Pond Bay Natural Area, and White Pond Bay Natural Area; LeBlond and Grant 2005). These connections to other large natural areas provide relatively uninterrupted habitat for the movement of plants and animals. Records of Horseshoe Lake’s water quality are lacking, but the lakes water appears high in humic substances and the chemistry is more than likely similar to the other Bladen Lakes. The lake is dystrophic and probably exhibits a pH of < 5.

Jones Lake

Jones Lake (91.05 hectares; 225 acres) is one of two dystrophic Carolina bay lakes located within Jones Lake State Park (893.54 hectares; 2,208 acres; Fig. 11), the other being Salters Lake. This lake is located in central Bladen County four miles north of Elizabethtown west of NC Hwy 242 and east of NC Hwy 53. Jones Lake State Parkforms the headwaters of an unnamed tributary of Turnbull Creek, which drains into the Cape Fear River. The state park sits on a sandy terrace (of Upper Pleistocene age) of the Cape Fear River (Soller 1988). Jones Lake was originally referred to as Woodward’s Lake, after Samuel Woodward, justice of the peace for the area in 1734 (North Carolina Division of Parks and Recreation, Planning and Development Section 2006b). It is believed that the lake later received its current name from Isaac Jones, an adjacent landowner to Samuel Wooodward, on whose land Elizabethtown was later established in 1773. Jones Lake State Park was established in 1939 and became the first state park specifically devoted to African Americans (North Carolina Department of Conservation and Development 1940).

Figure 11.

Jones Lake State Park (outlined in green) and surrounding lands. Jones Lake State Park is located between state highways 53 and 242, north of the Cape Fear River. Aerial imagery, transportation, and hydrography layers obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

LeBlond and Grant (2005) described both Jones and Salters Lakes as “among the very best examples of Carolina bay lakes in nearly pristine condition”. Jones Lake State Park is connected by fair to high quality landscape connections to Bethel Flatwoods, Cotton Bay Sand Ridge, Tatum Mill Pond/Cypress Bay, and Turnbull Creek Swamp natural areas.

Eleven natural community types have been described from Jones Lake State Park (i.e., Bay Forest, Coastal Plain Small Stream Swamp, High Pocosin, Low Pocosin, Natural Lake Shoreline, Peatland Atlantic White Cedar Forest, Pine/Scrub Oak Sandhill Mixed Oak Variant, Pond Pine Woodland, Wet Pine Flatwoods Wet Spodosol Variant, Xeric Sandhill Scrub Coastal Plain Variant, Xeric Sandhill Scrub Sandbarren Variant; LeBlond and Grant 2005, Schafale 2012), several of which are of extremely high quality and globally rare, such as the Low Pocosin, Peatland Atlantic White Cedar Forest, and Xeric Sandhill Scrub (LeBlond and Grant 2005, Schafale 2012). Available water quality parameters for Jones Lake are provided in Table 2.

Water Quality Data for Jones Lake (Bladen County, North Carolina). Frey (1949) sampled Jones Lake 9 times during the Summer and Fall of 1947. Weiss and Kuenzler (1976) sampled Jones Lake twice in 1974 (March 22 and June 6) and 4 times in 1975 (April 7, June 10, August 5, October 6). North Carolina Division of Water Quality, Environmental Sciences Section, Intensive Survey Unit 2009 (DWQ) sampled Jones Lake 5 times in 2008 (May 29, June 25, July 15, September 10, September 24). Value ranges have been provided where applicable to show variability. Units are as follows: km2 = squared kilometers, ha = hectares, km = kilometers, m = meters, °C = degrees celsius, mg/L – milligrams per liter, meq/L = milliequivalents per liter, s.u. = standard units, μg/L = micrograms per liter.

Frey (1949)

Weiss & Kuenzler (1976)

DWQ 2009

Trophic Status

Dystrophic

Watershed Area (km2)

5.18

Surface Area (ha)

90.65

Max Width (km)

0.48

Max Length (km)

0.80

Max Depth (m)

2.13

Mean Depth (m)

0.9

Secchi Depth (m)

0.73

0.3−1.22

1.3−2.4

Min Temp. (°C)

14

21.9

Max Temp. (°C)

30.5

29.6

Dissolved Oxygen (mg/L)

5.7

6.7−10.6

6.2−7.5

Alkalinity (meq/L)

0−0.002

pH (s.u.)

4.34

3.1−4.8

3.6−4.2

Total N (mg/L)

0.32−0.73

Total P (mg/L)

0.013−0.025

Chlorophyll-A (μg/L)

1−11

Lake Waccamaw

Lake Waccamaw is located south of the township of Lake Waccamaw, between Friar Swamp to the northeast, and the Waccamaw River to the south. It is the only Carolina bay lake located in Columbus County and is the largest Carolina bay and bay lake (3,617.48 hectares; 8,939 acres) in North Carolina (LeBlond 1995). Lake Waccamaw is the third largest lake in North Carolina behind Lake Mattamuskeet and Lake Phelps. The lake is part of Lake Waccamaw State Park (4,327.70 hectares; 10,694 acres; Fig. 12), which also includes lands directly abutting the lake’s southern shoreline. Stager and Cahoon (1987) estimated Lake Waccamaw to be ca. 15,000 years old or less.

Figure 12.

Lake Waccamaw State Park (outlined in green) and surrounding lands. Lake Waccamaw State Park is a large state park encompassing Lake Waccamaw and adjacent swampland and uplands. Aerial imagery, transportation, and hydrography layers obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

Prior to European civilization in the Southeast, the Waccamaw-Sioux Native American peoples, one of five Native American tribes known to inhabit the Cape Fear Region, inhabited the lands surrounding the lake (North Carolina Division of Parks and Recreation, Planning and Development Section 2006a). Native American artifacts, including dugout canoes, dating back to 1015−315 B.P. have been found within and around Lake Waccamaw. In the early 18th century, an unknown young man traveled through Columbus County on his way from north Georgia and, upon seeing Lake Waccamaw, described it as “the most pleasantest place that ever I saw in my life. It is at least eighteen miles round, surrounded with exceeding good land, as oak of all sorts, hickory and fine cypress swamps” (Gentleman 1737).

This bay lake differs from the Bladen lakes in its larger size, neutral pH, mesotrophic status, and presence of alluvial hydrologic inputs (Big Creek). Tea-stained waters from Friar Swamp are delivered into northeast Lake Waccamaw via Big Creek, the largest of several creeks draining into the lake from Friar Swamp. Lake Waccamaw forms the headwaters of the Waccamaw River, a species-rich river system known to support several rare plant (e.g., Fimbristylis perpusilla R.M. Harper ex Small & Britton, Ilex amelanchier M.A. Curtis ex Chapm., Lipocarpha micrantha (Vahl) G.C. Tucker, Oldenlandia boscii (DC.) Chapm., Rhynchospora decurrens Chapm., and Sabatia kennedyana Fernald) and animal taxa (Alligator mississipiensis [American Alligator], Elliptio folliculata [Pod Lance], Etheostoma perlongum [Waccamaw Darter], Lampsilis ochracea [Tidewater Mucket], Menidia extensa [Waccamaw Silverside], Noturus spp. 2 [Broadtail Madtom], and Procambarus leptodactylus [Pee Dee Lotic Crayfish; LeBlond 1995]).

Much of the land surrounding Lake Waccamaw has been converted to agriculture (north of the lake) and loblolly pine plantations (south of the lake). A small portion of Lake Waccamaw’s bay is still present on the northern end.

The Coastal Plain Marl Outcrop occurs along a roughly 394 m (1,000 ft.) stretch of northern shoreline and is characterized by having vertical and overhanging low cliffs in the supralittoral zone of the lake. Portions of these cliffs are submerged in the upper eulittoral zone, but local residents privately own terrestrial portions. This marl community is known for supporting the only naturally occuring population of Venus hair fern (Adiantum capillus-veneris L.) in the state.

Shoreline residential development extends along the northern shores of the lake from the lake outlet (southwest corner of lake) to just south of Big Creek. These shorelines support the globally rare Natural Lake Shoreline Marsh (Lake Waccamaw Pondlily Subtype) community. Undeveloped shorelines (i.e., Natural Lake Shoreline Swamp – Lake Waccamaw Subtype) occur from just south of Big Creek to the lake’s outlet. Historically, Lake Waccamaw experienced wide-ranging water level fluctuations determined by precipitation. In 1925, a poorly constructed dam was built at the lakes outlet in an effort to stabilize lake levels for increased recreational use. Before construction began, lake levels were so low that vehicles could be driven to the construction site on the dried lake bed (North Carolina Division of Parks and Recreation, Planning and Development Section 2006a).

The physical and hydrographic nature of Lake Waccamaw’s shoreline also differs from the other bay lakes. Lake Waccamaw’s shoreline is sandy around its entire periphery (Frey 1949), whereas the Bladen lakes may be either sandy or peaty along their shorelines.

A broad, sandy, terrace (lacking in Bladen lakes) is also present along the southeast shoreline of Lake Waccamaw (Fig. 5). This shallow underwater terrace extends perpendicularly out into the lake as far as 305 m (1,000 ft.; Frey 1949). The gentle relief of the terrace gradually extends shoreward resulting in a shallow, broad, littoral zone. This littoral zone is the most floristically rich of all Carolina bay lakes and is rivaled only by Lake Phelps in Washington County, North Carolina (N. Howell, pers. obs.). Varying water depths in the littoral zone of Lake Waccamaw result in the temporary and sometimes permanent presence of offshore sandbars and islands. This hydrographical heterogeneity in the littoral zone increases the floristic richness. A more detailed review of the lakes shoreline flora is provided in the floristic summary section and in Suppl. material 6.

The buffering effect of subsurface and surficial limestone on the naturally acidic waters of Lake Waccamaw result in an unusually diverse fauna. Lake Waccamaw contains the largest number of endemic animal species (i.e., endemic to this lake and nowhere else in the world; 10 taxa) of any site in North Carolina (Hubbs and Raney 1946, LeBlond 1995). An additional species, Fundulus waccamawensis (Waccamaw Killfish), is found only in waters within and adjacent to Lake Waccamaw and Lake Phelps (Washington County, North Carolina). Six other faunal taxa known to be rare but not endemic also occur within or adjacent to the lake. Available water quality parameters for Lake Waccamaw are provided in Table 3.

Water Quality Data for Lake Waccamaw (Columbus County, North Carolina). Frey (1949) sampled Lake Waccamaw 8 times during the Summer and Fall of 1947. Weiss and Kuenzler (1976) sampled Lake Waccamaw twice in 1974 (March 22 and June 6) and 4 times in 1975 (April 7, June 10, August 5, October 6). North Carolina Division of Water Quality, Environmental Sciences Section, Intensive Survey Unit (2012) (DWQ) sampled Lake Waccamaw 5 times in 2011 (May 4, June 8, September 1, July 20, and August 17). Value ranges have been provided where applicable to show variability. Units are as follows: km2 = squared kilometers, ha = hectares, km = kilometers, m = meters, °C = degrees celsius, mg/L – milligrams per liter, meq/L = milliequivalents per liter, s.u. = standard units, μg/L = micrograms per liter.

Frey (1949)

Weiss & Kuenzler (1976)

DWQ 2012

Trophic Status

Mesotrophic

Watershed Area (km2)

181.29

Surface Area (ha)

3617.08

Max Width (km)

5.47

Max Length (km)

8.36

Max Depth (m)

3.35

Mean Depth (m)

1.5

Secchi Depth (m)

1.34

0.61−2.38

1.1−1.9

Min Temp. (°C)

14

23.5

Max Temp. (°C)

31.5

29.9

Dissolved Oxygen (mg/L)

5.2

7.8−11

6.9−8.1

Alkalinity (meq/L)

0.14−0.24

pH (s.u.)

6.95

6.8−7.5

7.0−8.5

Total N (mg/L)

0.297−1.56

Total P (mg/L)

0.017 − .055

Chlorophyll-A (μg/L)

2.8−8

Little Singletary Lake

Little Singletary Lake (626 acres; 253.33 hectares) is located in the western half of Suggs Mill Pond Game Land (Fig. 10). Unlike Horseshoe Lake, Little Singletary Lake is natural in origin and exhibits a more “typical” bay lake physiognomy. Little Singletary Lake forms the headwaters of Lake Run, a tributary of Ellis Creek, which drains into the Cape Fear River. Relatively intact landscape connections exist to the northeast (Horseshoe Lake), southeast (Marshy Bay Natural Area), and southwest (Cedar Swamp Seep Natural Area) from Little Singletary Lake.

Lands abutting the southern shoreline are privately owned and were once subject to residential development. Remnants of bulkheads and recreational piers can still be seen today along the southeast shoreline. The North Carolina Wildlife Resources Commission gained property rights to all remaining lands surrounding Little Singletary Lake before residential development could ensue. On June 20, 2011, a lightning caused wildfire (Simmons Road Fire) started just west of Little Singletary Lake and by August 18th, had burned over 2,023 hectares (5,000 acres) of Carolina bay and pocosin habitat, much of which surrounded Little Singletary Lake. During growing seasons of extreme drought, water levels have been known to recede low enough to reveal a clean sandy lake bottom 90−275 m (100−300 yds) out into the lake (G. Lewis, pers. comm.). Native American projectile points have been found on this lake bottom during drought years (G. Lewis, pers. comm.).

The water quality of Little Singletary Lake has not been documented by state agencies. The water appears high in humic substances and is likely similar to the other Bladen lakes (i.e., dystrophic, acidic, shallow, nutrient poor).

Salters Lake

Salters Lake (127.47 hectares; 315 acres) is the larger of the two Carolina bay lakes located in Jones Lake State Park (Fig. 11). Salters Lake was named after Sallie Salter, a revolutionary war hero who spied on the Tories while encamped at Elizabethtown. Her spying played a role in the defeat over the Tories on August 28, 1771, at the battle of Elizabethtown, where 70 Whigs defeated 400 Tories (JNorth Carolina Division of Parks and Recreation, Planning and Development Section 2006b).

Salters Lake is similar to Jones Lake in many respects, but quite possibly could be the most “pristine” of all Carolina bay lakes. Salters Lake has no shoreline development, appreciable recreational activities (e.g., outboard motor use), immediate surrounding agricultural (crop or animal production) land use, water level control structures, or historical manipulation of any kind. Natural communities and landscape features for Salters Lake are the same as those for Jones Lake (above). Available water quality parameters for Salters Lake are provided in Table 4.

Water Quality Data for Salters Lake (Bladen County, North Carolina). Frey (1949) sampled Salters Lake 7 times during the Summer and Fall of 1947. Weiss and Kuenzler (1976) sampled Salters Lake twice in 1974 (March 22 and June 6). North Carolina Division of Water Quality, Environmental Sciences Section, Intensive Survey Unit 2009 (DWQ) sampled Salters Lake 4 times in 2008 (June 25, July 15, August 20, September 24). Value ranges have been provided where applicable to show variability. Units are as follows: km2 = squared kilometers, ha = hectares, km = kilometers, m = meters, °C = degrees celsius, mg/L – milligrams per liter, meq/L = milliequivalents per liter, s.u. = standard units, μg/L = micrograms per liter.

Frey (1949)

Weiss & Kuenzler (1976)

DWQ 2009

Trophic Status

Dystrophic

Watershed Area (km2)

7.77

Surface Area (ha)

127.47

Max Width (km)

0.80

Max Length (km)

1.12

Max Depth (m)

1.82

Mean Depth (m)

2.13

Secchi Depth (m)

0.55

0.6−0.91

Min Temp. (°C)

15

21.7

Max Temp. (°C)

25.4

31.2

Dissolved Oxygen (mg/L)

6

7.9−10.1

6.5 – 8.1

Alkalinity (meq/L)

0.0019

pH (s.u.)

4.49

4.1−4.8

3.6 – 4.1

Total N (mg/L)

0.293−0.374

Total P (mg/L)

0.015−0.016

Chlorophyll-A (μg/L)

4.7 – 26

Singletary Lake

Singletary Lake (233.09 hectares; 576 acres) is located within Singletary Lake State Park (494.12 hectares; 1,221 acres; Fig. 13). This lake was named after Richard Singletary, who received the grant of land in 1729 (North Carolina Division of Parks and Recreation, Planning and Development Section 1996b). Singletary Lake State Park is located just southeast of White Lake in central-southeast Bladen County between the Cape Fear River and Colly Swamp. Singletary Lake forms the headwaters of Lake Drain Creek, which drains into Big Colly Creek, which drains to the Black River, which drains into the Cape Fear River.

Figure 13.

Singletary Lake State Park (outlined in green) and surrounding lands. Singletary Lake State Park is primarily comprised of lands immediately surrounding Singletary Lake. In addition to the lands surrounding Singletary Lake, White Lake is also managed by Singletary Lake State Park. Singletary Lake State Park is located north of the Cape Fear River and State Hwy 53 and southeast of White Lake. Aerial imagery, transportation, and hydrography layers obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

Singletary Lake is similar to the other Bladen lakes in that it is dystrophic, acidic, and nutrient poor. It contains high quality examples of the Natural Lake Shoreline Swamp (Cypress Subtype) and Natural Lake Shoreline Marsh (Typic Subtype) communities. LeBlond and Grant (2005) described this lake’s shoreline community as “one of the most aesthetically pleasing natural communities in the North Carolina Coastal Plain”. A direct landscape connection exists between Singletary Lake and Colly Swamp and the Black River to the northeast. Fair quality landscape connections exist between the state park and the Cape Fear River to the southwest. Available water quality parameters for Singletary Lake are provided in Table 5.

Water Quality Data for Singletary Lake (Bladen County, North Carolina). Frey (1949) sampled Singletary Lake 10 times during the Summer and Fall of 1947. Weiss and Kuenzler (1976) sampled Singletary Lake twice in 1974 (March 22 and June 6) and four times in 1975 (April 7, June 10, August 5, October 6). North Carolina Division of Water Quality, Environmental Sciences Section, Intensive Survey Unit (2009) (DWQ) sampled Singletary Lake 5 times in 2008 (June 25, July 15, August 20, September 24). Value ranges have been provided where applicable to show variability. Units are as follows: km2 = squared kilometers, ha = hectares, km = kilometers, m = meters, °C = degrees celsius, mg/L – milligrams per liter, meq/L = milliequivalents per liter, s.u. = standard units, μg/L = micrograms per liter.

Frey (1949)

Weiss & Kuenzler (1976)

DWQ 2009

Trophic Status

Dystrophic

Watershed Area (km2)

5.18

Surface Area (ha)

231.48

Max Width (km)

0.64

Max Length (km)

2.09

Max Depth (m)

2.74

Mean Depth (m)

2.13

Secchi Depth (m)

0.76

0.48−1.21

0.6−1

Min Temp. (°C)

13.8

24.8

Max Temp. (°C)

31

30.6

Dissolved Oxygen (mg/L)

6.6

7.3−11.2

6−7.8

Alkalinity (meq/L)

0.0019

pH (s.u.)

4.5

3.2−4.6

3.9−4.2

Total N (mg/L)

0.255−0.515

Total P (mg/L)

0.018−0.075

Chlorophyll-A (μg/L)

4.8−44

White Lake

Although not included in the sampling aspect of this study, White Lake is unique and deserves a brief summary. White Lake (432.20 hectares; 1,068 acres) is a large Carolina bay lake located in east-central Bladen County about 6 miles east of Elizabethtown, just east of the intersection of NC Hwy 53 and U.S. Hwy 701 (Fig. 14). White Lake is owned by the state of North Carolina, and is managed by Singletary Lake State park. Unlike all of the remaining bay lakes, White lake’s water is clear and not tea-stained. This feature has made it an incredibly attractive location for development and vacationers. This lake is primarily used for recreation (e.g., water sports, swimming, fishing) and essentially all of its shoreline is residentially and commercially developed.

Figure 14.

White Lake and surrounding lands. Like the majority of Carolina bay lakes, White Lake is a state-owned lake. All but a very small portion of White Lake’s shoreline has been altered. Aerial imagery, transportation, and hydrography layers obtained from NRCS Geospatial Data Gateway: https://gdg.sc.egov.usda.gov. Map produced by Nathan Howell using ArcGis Desktop: Version 10.2.2. (Environmental Systems Research Institute (ESRI) 2014).

White Lake’s remarkable water clarity is attributed to the presence of artesian springs on the lake bottom (Wells and Boyce 1953). The clarity of the lake’s water yields a deep euphotic zone (i.e., sunlight can penetrate through the entirety of the water column) with submerged aquatic macrophytes (e.g., Myriophyllum humile (Raf.) Morong; N. Howell pers. obs.) present at the lakes deepest depths. White Lake receives its hydrologic inputs principally in two forms, precipitation and groundwater (through springs). Although this lake is primarily fed by springs, its overall water levels are determined by the regional water table (i.e., during drought years, White Lake’s water levels will drop just like all other bay lakes). Another unique feature of White Lake is the location of its outlet channel. White Lake’s outlet channel is located in the northwestern section of the lake as opposed to the southeastern section where it occurs in all other bay lakes. Frey (1954) reported that William Bartram, a renowned naturalist who documented the flora, fauna, and Native American culture of the southeastern United States in the 18th century, operated a sawmill on White Lake during the 20 years following 1770. A map in Bartram and Harper (1942) shows that White Lake was formerly called Lake Bartram. Available water quality parameters for White Lake are provided in Table 6.

Water Quality Data for White Lake (Bladen County, North Carolina). Frey (1949) sampled White Lake 8 times during the Summer and Fall of 1947. Weiss and Kuenzler (1976) sampled White Lake twice in 1974 (March 22 and June 6). North Carolina Division of Water Quality, Environmental Sciences Section, Intensive Survey Unit (2009) (DWQ) sampled White Lake 5 times in 2008 (May 27, June 24, July 29, August 11, and October 2). Value ranges have been provided where applicable to show variability. Units are as follows: km2 = squared kilometers, ha = hectares, km = kilometers, m = meters, °C = degrees celsius, mg/L – milligrams per liter, meq/L = milliequivalents per liter, s.u. = standard units, μg/L = micrograms per liter.

Frey (1949)

Weiss & Kuenzler (1976)

DWQ 2009

Trophic Status

Oligotrophic

Watershed Area (mi2)

Surface Area (ha)

432.2

Max Width (km)

1.61

Max Length (km)

2.57

Max Depth (m)

3.35

Mean Depth (m)

3.04

Secchi Depth (m)

3.35

3.35

3.35

Min Temp. (°C)

15.1

22.3

Max Temp. (°C)

26.1

30.1

Dissolved Oxygen (mg/L)

6.7

8.6−10.1

6.8−8.2

Alkalinity (meq/L)

0.0019−.0099

pH (s.u.)

4.92

4.6−4.8

4.6−5.2

Total N (mg/L)

0.123−0.211

Total P (mg/L)

0.010−0.017

Chlorophyll-A (μg/L)

4.8−44

Climate

Bladen Lake Group (Bladen County, NC)

Climate data from the nearest weather station to the Bladen County bay lakes, ca. 1.6 kilometers away in Elizabethtown, North Carolina (Bladen County: 34.68° N, -78.58°W; 30.5 m elev.), show that during the thirty-year period between 1971-2000, the average annual temperature was 16.44 °C (61.6 °F) and mean annual precipitation 1,254.76 mm (49.4 in). Average daily maximum and minimum temperatures were 22.83 °C (73.1 °F) and 10.11 °C (50.2 °F; State Climate Office of North Carolina 2014; Fig. 15).

Figure 15.

Walter climate diagrams for weather stations closest to the Bladen Lakes (Bladen County, NC; a) and Lake Waccamaw (Columbus County, NC; b), based on data from the State Climate Office of North Carolina (2014). At the top left of each figure, the town closest to the weather station is listed as well as the elevation of the weather station in meters and the number of years climate data were recorded (30). At the top right of each figure, the mean annual temperature and precipitation over thirty years for each site is listed. Climate data for these figures were recorded from 1971 to 2000. Solid black areas in the diagrams represent “excess rainfall”. When the precipitation curve rises above 100 mm, there is an excess amount of precipitation present that plants do not need in order to survive. Areas marked with vertical lines between the temperature curve and the 100 mm precipitation mark on the secondary y-axis represent a “wet period”. These diagrams show that plants in these two locations are not water-stressed (i.e., the precipitation curve does not drop below the temperature curve for the 30-year climatic period).

The lowest temperature recorded for Bladen County was -14.4 °C (6 °F) on January 17, 1977 (Leab 1990). The highest recorded temperature for Bladen County was 37.7 °C (100 °F) on July 20, 1977 (Leab 1990). Monthly average temperatures were highest in July and August and lowest in December and January. Monthly precipitation amounts were also highest in July and August, while the lowest monthly precipitation amounts were in April and November (State Climate Office of North Carolina 2014; Fig. 15). The annual growing season, defined as the number of days in five out of ten years during which the daily minimum air temperature exceeds -2.2 °C (28 °F), is 243 days in Bladen County (weather data recorded from 1957-1979; Leab 1990).

Lake Waccamaw (Columbus County, NC)

Climate data from the nearest weather station to Lake Waccamaw, ca. 16 km away in Whiteville, North Carolina (Columbus County: 34.27287° N, -78.71499° W; 29.8 meters above sea level), show that for the 30-year period between 1971 and 2000, the average annual temperature was 17.16 °C (62.9 °F) and mean annual precipitation 1,275.08 mm (50.2 in). The average daily maximum and minimum temperatures over the same thirty-year period were 24.3 °C (75.8 °F) and 10 °C (50 °F; State Climate Office of North Carolina 2014; Fig. 15).

The lowest temperature recorded for Columbus County was -15 °C (5 °F) on February 12, 1973 (Spruill 1990). The highest recorded temperature for Columbus County was 40.5 °C (105 °F) on June 27, 1954 (Spruill 1990). Monthly average temperatures were highest in July and August and lowest in January and February. Monthly precipitation amounts were also highest in July and August, while the lowest monthly precipitation amounts were in April and November (State Climate Office of North Carolina 2014; Fig. 15). The annual growing season, defined as the number of days in five out of ten years during which the daily minimum air temperature exceeds -2.2 °C (28 °F), is 240 days in Columbus County (weather data recorded from 1951-1981; Spruill 1990).

Plant Communities

Four plant community types and two subtypes can be distinguished within the littoral zone of Carolina bay lakes (Schafale 2012; Table 7). Of these four community types and subtypes, three are globally critically imperiled (Natural Lake Shoreline Swamp – Lake Waccamaw Subtype; Natural Lake Shoreline Marsh – Typic Subtype; Natural Lake Shoreline Marsh − Lake Waccamaw Pondlily Subtype), while the others do not have a conservation ranking (Table 7).

Plant community types occurring within the littoral zone of Carolina bay lakes. Community types follow Schafale (2012); rank designations follow Robinson and Finnegan (2014). Community types are presented in order of increasing species richness. The Natural Lake Shoreline Marsh (Lake Waccamaw Pondlily Subtype) typically supports a couple of dominant taxa (i.e., Nuphar sagittifolia and Eriocaulon aquaticum) with several other co-dominants. The Natural Lake Shoreline Swamp (Lake Waccamaw Subtype) is known to contain 140+ taxa.

Species

Richness

Plant Community Types

State

Rank

Global

Rank

Lowest

Highest

Natural Lake Shoreline Marsh (Lake Waccamaw Pondlily Subtype)

S1

G1

Coastal Plain Semipermanent Impoundment

S4

G4G5

floating Bog

S1

G1?

Natural Lake Shoreline Swamp (Cypress Subtype)

S2

G3

Natural Lake Shoreline Marsh (Typic Subtype)

S1

G1

Natural Lake Shoreline Swamp (Lake Waccamaw Subtype)

S1

G1

S1 = Critically Imperiled, 1–5 occurrences in state; S2 = Imperiled, 6–20 occurrences in state; S4 = Apparently Secure, 101–1000 occurrences in state; G1 = Critically Imperiled, 1–5 occurrences in the world; G3 = Vulnerable, 21–100 occurrences in the world; G4 = Apparently Secure, 101–1000 occurrences in the world; G5 = Secure, 1001+ occurrences in the world.

Natural Lake Shoreline Swamp (Cypress Subtype; S2G3) [Taxodium distichumT. ascendens / Panicum hemitomon Schult. Woodland (CES203.044)].

This natural community type covers Carolina bay lake shorelines with narrow littoral zones characterized by an absent to sparse herbaceous component and a nearly closed canopy of Chamaecyparis Spach, Nyssa L., or Taxodium Rich. in the upper eulittoral zone. If a cross-section of this littoral zone were to be drawn, the epilittoral vegetation would abruptly coincide with the littoral zone (i.e., a zone of emergent herbaceous vegetation is lacking where it typically would occur between the epilittoral and infralittoral zones). This “two-staged” zonation pattern typical of this community type is directly attributable to the steeper hydrography and narrow littoral zone. The Natural Lake Shoreline Swamp (Lake Waccamaw Subtype) and the Natural Lake Shoreline Marsh community types can be distinguished from the depauperate Natural Lake Shoreline Swamp (Cypress Subtype) community type by having a broader littoral zone, a well-developed zone of herbaceous emergent macrophytes, a sparse to open canopy of Nyssa, Taxodium, or other obligate wetland hardwoods, and the absence of Nuphar sagittifolia (Walter) Pursh. Examples of this community type are found at Bakers Lake, and the western, northern, and eastern shorelines of Jones, Salters, Little Singletary, and Singletary Lakes.

Natural Lake Shoreline Swamp (Lake Waccamaw Subtype; S1G1) [Taxodium distichumT. ascendens / Panicum hemitomonSclerolepis uniflora (Walter) Britton, Sterns & Poggenb. Woodland (CEGL004465)].

This natural community type covers the southern shoreline of Lake Waccamaw located between Big Creek and the lake’s outlet on the southwest shore. This stretch of natural shoreline is characterized by gentle hydrography, which results in a broad littoral zone, and a species-rich flora dominated by emergent herbaceous macrophytes, many of which are rare. Emergent macrophytes typical of this community type include Cladium mariscoides (Muhl.) Torr., Eriocaulon aqutaicum (Hill) Druce, Panicum hemitomon, Sclerolepis uniflora, and Xyris smalliana, among others. This community type can be distinguished from the species-poor Natural Lake Shoreline Swamp (Cypress Subtype) community type by its broader littoral zone and species-rich herbaceous component (95 taxa). It can be distinguished from the Natural Lake Shoreline Marsh community types by the absence or only irregular presence of Nuphar sagittifolia and the unique assemblage of diverse herbaceous taxa (e.g., Bacopa caroliniana (Walter) B.L. Rob., Boltonia asteroides (L.) L’Hér. var. glastifolia, Cladium mariscoides, Ludwigia brevipes (B.H. Long ex Britton, A. Braun & Small) Eames, L. sphaerocarpa Elliott, and Sclerolepis uniflora).

Natural Lake Shoreline Marsh (Typic Subtype; S1G1) [Panicum hemitomonJuncus spp. Coastal Plain Lakeshore Herbaceous Vegetation (CEGL004307)].

This natural community type covers the southern shorelines of the Bladen Lakes. The southern shorelines have a broader littoral zone than the remaining portions of the lakes. Consequently, they support a more diverse emergent herbaceous component. Herbs found in this community type include Eleocharis baldwinii, E. equisetoides (Elliott) Torr., E. vivipara, Juncus pelocarpus E. Mey., Panicum hemitomon, Panicum verrucosum Muhl., Rhexia nashii, Rhynchospora distans, Saccharum giganteum (Walter) Pers., Sacciolepis striata, Scirpus cyperinus (l.) Kunth, and Xyris smalliana. This community type is also characterized as having a sparse to open canopy of Nyssa and Taxodium. This community type can be distinguished from the Natural Lake Shoreline Marsh (Lake Waccamaw Pondlily Subtype) by the absence of Nuphar sagittifolia and from the Natural Lake Shoreline Swamp (Lake Waccamaw Subtype) by the occurence of < 30 herbaceous taxa, none of which include the unique and rare herbs found at Lake Waccamaw. Examples of this community type include the southern shorelines of Jones, Little Singletary, Salters, and Singletary Lakes.

Natural Lake Shoreline Marsh (Lake Waccamaw Pond-lily Subtype; S1G1) [Nuphar sagittifolaEriocaulon aquaticum Lakeshore Herbaceous Vegetation (CEGL004297)].

This natural community type covers the western, northern, and eastern shorelines of Lake Waccamaw (i.e., where residential and commercial development is present). It is the only Natural Lake Shoreline community type dominated by Nuphar sagittifolia (a distinguishing feature) and Eriocaulon aquaticum. Nuphar sagittifolia is essentially absent from the Natural Lake Shoreline Swamp (Lake Waccamaw Subtype) community type save for small stands around the mouth of Big Creek and around the dam at the lakes outlet.

floating Bog [Rhynchospora alba Saturated Herbaceous Vegetation (CEGL004463)]

This natural community type covers the rare examples of vegetation occuring on floating peat mats in deep water of natural or artificial ponds and lakes. Horseshoe Lake is the only Carolina bay lake known to support floating bogs. The floating bogs of Horseshoe Lake are the largest in the state. These floating bogs are saturated and nutrient-poor, supporting taxa that characteristically inhabit such stressful conditions (e.g., Calopogon tuberosus (L.) Britton, Sterns & Poggenb., Drosera intermedia, Dulichium arundinaceum, Hypericum virginicum, Pogonia ophioglossoides, Rhynchospora alba, R. inundata, and Xyris fimbriata). This community type’s “floating” nature and the presence of the aforementioned plant taxa sets it apart from all others.

Coastal Plain Semipermanent Impoundment (Cypress-Gum Subtype; G4G5) [Taxodium distichum / Lemna minor L. Forest (CEGL002420)]

All portions of Horseshoe Lake not considered floating Bog fall into the Coastal Plain Semipermanent Impoundment community type. This community type is characterized by a sparse to absent canopy of Taxodium ascendens with sporadically occurring beds of floating-leaved and submersed aquatics (e.g., Brasenia schreberi J.F. Gmel, Cabomba caroliniana A. Gray, Nymphaea odorata ssp. odorata, and Utricularia spp.). This community type can be distinguished from all others by the sparse presence of Taxodium throughout the lake with floating-leaved and submersed aquatics occurring underneath.

Floristic Summary

Across All Sites

The littoral zone vascular flora of Carolina bay lakes, based on vouchered collections, reports, and personal observations, consists of 205 taxa (170 species, 4 subspecies, 30 varieties, 1 hybrid) in 136 genera and 80 vascular plant families (Table 8; Suppl. material 6). Of these 205 taxa, 186 (90.7%) are vouchered and 19 (9.3%) are known only from reports (Peet et al. 2013a, Peet et al. 2013b, North Carolina Natural Heritage Program 2014; NCSU Crop Science Department [Rob Richardson and Justin Nawrocki, pers. comm., April 9, 2015]). Of the 186 vouchered taxa, 157 (84.4%) were collected by the first author; the remaining 29 (15.6%) vouchered taxa were collected from Carolina bay lake shorelines by others and were found by completing systematic searches of major herbaria (DUKE, NCSC, and NCU). Nineteen taxa (9.3%) are listed as significantly rare and twelve taxa (5.8%) are on the NCNHP Watch List (Table 9). Four taxa (1.9%) are Federal Species of Concern (Ludwigia brevipes; Nuphar sagittifolia; Rhexia aristosa Britton; Sagittaria weatherbiana). Pair-wise comparisons of species similarity for all bays are provided in Table 10.

Summary of vascular plant taxa collected or reported from Carolina bay lake littoral zones

Species and Subspecies/Varieties

Group

Families

Genera

Native

Exotic

Total

Basal Angiosperms & Magnoliids

4

6

6

0

6

Pteridophytes

6

7

7

0

7

Gymnosperms

2

3

5

0

5

Monocotyledons

17

41

84

2

86

Eudicotyledons

51

79

98

3

101

Total

80

136

200

5

205

List of North Carolina Significantly Rare and Watch List taxa collected or reported from Carolina bay lake littoral zones. Status and rank designations follow Robinson and Finnegan (2014). Taxa for which voucher specimens have been collected (by the first author or others) are indicated with a check mark (✓) in the second column. The taxonomy followed in this work and that of Robinson and Finnegan (2014) differ in one instance in the following table: Luziola fluitans (Michx.) Terrell & H. Rob. var. fluitans (as Luziola fluitans (Michx.) Terrell & H. Rob. sensu Robinson and Finnegan 2014). See Martínez-y-Pérez et al. (2008) in addition to the FNA treatment for reasons of further division to an infraspecific rank.

Taxon

Vouchered ?

State Status

Fed. Status

State Rank

Global Rank

Significantly Rare:

1

Bacopa caroliniana (Walter) B.L. Rob.

T

S1

G4G5

2

Boltonia asteroides (L.) L’Hér var. glastifolia (Hill) Fernald

SR−O

S2

G5TNR

3

Cladium mariscoides (Muhl) Torr.

SR−O

S3

G5

4

Eleocharis vivipara Link

E

S1

G5

5

Epidendrum magnoliae Muhl.

T

S1S2

G4

6

Eriocaulon aquaticum (Hill) Druce

SC−V

S2

G5

7

Ludwigia brevipes (Long) Eames

SR−T

FSC

S1S2

G2G3

8

Ludwigia sphaerocarpa Elliott

E

S1

G5

9

Luziola fluitans (Michx.) Terrell & H. Rob. var. fluitans

SR−P

S2

G4,G5

10

Lycopus angustifolius Elliott

SR−P

S1

G4?Q

11

Rhexia aristosa Britton

SC−V

FSC

S3

G3,G4

12

Rhynchospora alba (L.) Vahl

SR−P

S2

G5

13

Sagittaria filiformis J.G. Sm.

SR−P

SH

G4,G5

14

Sagittaria isoetiformis J.G. Sm.

T

S2

G4?

15

Sagittaria weatherbiana Fernald

E

FSC

S2

G3G4

16

Sclerolepis uniflora (Walter) Britton, Sterns & Poggenb.

SR−T

S2

G4

17

Spiranthes laciniata (Small) Ames

SC−V

S2

G4,G5

18

Utricularia cornuta Michx.

T

S1S2

G5

19

Utricularia resupinata B.D. Greene ex Bigelow

E

S1

G4

Watch List:

1

Dichanthelium dichotomum (L.) Gould var. roanokense (Ashe) LeBlond

W1

S2

G5T4?

2

Dichanthelium erectifolium (Nash) Gould & C.A. Clark

W1

S2

G4

3

Dryopteris ludoviciana (Kunze) Small

W1

S2

G4

4

Eleocharis equisetoides (Elliott) Torr.

W1

S3

G4

5

Habaneria repens Nutt.

W1

S2

G5

6

Nelumbo lutea Willd.

W7

S2

G4

7

Nuphar sagittifolia (Walter) Pursh

W1

FSC

S2

G5T2

8

Rhexia cubensis Griseb.

W1

S3

G4G5

9

Rhynchospora inundata (Oakes) Fernald

W1

S3

G4?

10

Rhynchospora nitens (Vahl) A. Gray

W1

S3

G4?

11

Xyris iridifolia Chapm.

W7

S2

G4G5T4T

12

Xyris smalliana Nash

W1

S3

G5

STATE STATUS: E = Endangered; T = Threatened; SC-V = Special Concern-Vulnerable; SR = Significantly Rare: −T = Throughout; −P = Periphery of Range; −O = Other; W = Watchlist: W1 = rare but relatively secure; W7 = rare and poorly known. FEDERAL STATUS: FSC = Federal Species of Concern. STATE RANK: SH = historical (known only from historical populations in the state); S1 = Critically Imperiled, 1–5 populations in the state; S2 = Imperiled, 6–20 populations in the state; S3 = Vulnerable, 21–100 populations in the state. FEDERAL RANK: G2 = Imperiled, 6–20 populations in the world; G3 = Vulnerable, 21–100 populations in the world; G4 = Apparently Secure, 101–1000 populations in the world; G5 = Secure, 1001+ populations in the world; T# = Global rank of a subspecies or variety; NR = Not Ranked; Q = Questionable taxonomy; ? = Uncertain.

Sørenson’s Similarity Index for Carolina bay lakes. Values in this table are represented as percentiles (i.e., when looking in the second column from the left under Bakers Lake, Bakers Lake is considered to be 16.4% similar to Bay Tree Lake, 23.5% similar to Horseshoe Lake, and 40.8% similar to Jones Lake). Based solely on littoral zone plant taxa, Jones Lake and Singletary Lake are 83.3% alike.

Bakers

Lake

Bay Tree

Lake

Horseshoe

Lake

Jones

Lake

Lake

Waccamaw

Little

SingletaryLake

Salters

Lake

Singletary

Lake

Bakers Lake

100

16.4

23.5

40.8

12.4

39.3

41.0

41.5

Bay Tree Lake

16.4

100

37.4

38.6

33.0

46.3

32.4

41.3

Horseshoe Lake

23.5

37.4

100

38.6

26.7

42.2

24.7

48.3

Jones Lake

40.8

38.6

38.5

100

20.5

42.5

55.6

83.3

Lake Waccamaw

12.4

33.0

26.7

20.5

100

22.5

21.3

28.9

Little Singletary Lake

39.3

46.3

42.2

42.5

22.5

100

29.5

56.0

Salters Lake

41.0

32.4

24.7

55.5

21.3

29.5

100

51.7

Singletary Lake

41.5

41.3

48.3

83.3

28.9

56.0

51.7

100

Among all taxa treated in this guide, the major vascular plant groups consisted of the following total taxa: Eudicotyledons (101 taxa; 86 species, 1 subspecies, 13 varieties, 1 hybrid), monocotyledons (86 taxa; 71 species, 1 subspecies, 14 varieties), pteridophytes (7 taxa; 6 species and 1 subspecies), gymnosperms (5 species), basal angiosperms (4 taxa; 3 species and 1 subspecies), and magnoliids (2 taxa; 1 species and 1 variety; Table 8; Fig. 16). The richest families in the eudicotyledons are Asteraceae (13 taxa; 11 species, 1 variety, 1 hybrid), Ericaceae (8 taxa; 6 species, 2 varieties), Lentibulariaceae (6 taxa), Melastomataceae (5 taxa; 4 species, 1 variety), Hypericaceae (4 taxa; 3 species, 1 variety), and Rosaceae (4 taxa; Fig. 17). The richest genera in the eudicotyledons are Utricularia (6 taxa), Rhexia L. (5 taxa), and Hypericum L. (4 taxa). The richest families in the monocotyledons are Cyperaceae (25 taxa; 20 species, 5 varieties), Poaceae (21 taxa; 17 species, 4 varieties), Juncaceae (8 taxa), Orchidaceae (5 taxa; 4 species, 1 variety), Alismataceae (4 taxa), Smilacaceae (4 taxa), and Xyridaceae (4 taxa: Fig. 17). The richest genera in the monocotyledons are Rhynchospora (9 taxa; 8 species, 1 variety), Juncus (8 taxa), Dichanthelium (Hitchc. & Chase) Gould (6 taxa; 5 species, 1 variety), Carex (4 taxa; 3 species, 1 variety), Eleocharis (4 taxa; 3 species, 1 variety), Sagittaria L. (4 taxa), Smilax L. (4 taxa), and Xyris L. (4 taxa).

Figure 16.

Distribution of plant habit across all Carolina bay lakes. Lakes dominated by herbs have broader littoral zones, which encourage the establishment of herbaceous emergent macrophytes. Lakes dominated by trees and shrubs have narrow littoral zones, which discourage the establishment of herbaceous emergent macrophytes.

Figure 17.

The thirteen most species-rich vascular plant families across all Carolina bay lakes. Cyperaceae (orange), Ericaceae (yellow), Juncaceae (dull green), Poaceae (purple), Smilacaceae (neon green), and Xyridaceae (black) consistently occur across all sites.

Among all taxa treated in this guide, the most species-rich habit is herbs (140 taxa; 119 species, 2 subspecies, 18 varieties, 1 hybrid), followed by trees and shrubs (51 taxa; 42 species, 1 subspecies, 8 varieties), and vines (14 taxa, 12 species, 2 varieties; Fig. 16). Among the herbs, Cyperaceae (25 taxa), Poaceae (20 taxa), Asteraceae (11 taxa), Juncaceae (8 taxa), Lentibulariaceae (6 taxa), Melastomataceae (5 taxa), and Orchidaceae (5 taxa) are the most species-rich families. Among trees and shrubs, the Ericaceae (8 taxa) and Rosaceae (4 taxa) were the most species-rich families. Among vines, the Smilacaceae (4 taxa) and Vitaceae (2 taxa) were the most species rich families.

Among the natural community types included in this work, the Natural Lake Shoreline Swamp (Lake Waccamaw Subtype) is the most species-rich (145 taxa) and the Natural Lake Shoreline Marsh (Lake Waccamaw Pondlily Subtype) is the least species-rich (< 10 taxa; Table 7). Five exotic taxa are known to occur in the bay lakes, four (Alternanthera philoxeroides [Amaranthaceae], Colocasia esculenta (L.) Schott [Araceae], Hydrilla verticillata (L.F.) Royle [Hydrocharitaceae], Triadica sebifera (L.) Small [Euphorbiaceae]) from Lake Waccamaw and one (Hypochaeris radicata L. [Asteraceae]) from Bay Tree Lake.

Individual Lakes

Among the lakes, the largest number of littoral zone taxa (i.e., species, subspecies, and varieties) occurred in Lake Waccamaw (145 taxa), followed by Bay Tree Lake (56 taxa) and Horseshoe Lake (52 taxa; Table 11). The least number of littoral zone taxa occurred in Bakers Lake (18 taxa).

Number of taxa (species, subspecies, and varieties) by major taxonomic group across study sites. Sites are arranged from taxonomically richest to most depauperate. BALA = Bakers Lake; BATR = Bay Tree Lake; HOLA = Horseshoe Lake; JOLA = Jones Lake; LAWA = Lake Waccamaw; LISI = Little Singletary Lake; SALA = Salters Lake; SILA = Singletary Lake.

LAWA

BATR

HOLA

LISI

SILA

JOLA

SALA

BALA

Pteridophytes

7

3

1

2

1

1

1

1

Gymnosperms

2

3

3

3

5

4

2

1

Basal angiosperms

3

--

3

--

1

--

--

--

Magnoliids

2

--

--

1

2

2

2

2

Monocots

60

23

21

17

9

10

5

3

Eudicots

71

27

24

16

18

16

12

11

Total

145

56

52

39

36

33

22

18

Bakers Lake

The littoral zone vascular flora of Bakers Lake is depauperate with respect to the other bay lakes (Table 11). A total of 18 taxa (14 species, 4 varieties) in 17 genera and 14 vascular plant families were found in this lake’s littoral zone (Suppl. material 6). All but one taxon (Tillandsia usneoides) from Bakers Lake were collected by the first author (i.e., there were no reports or historical vouchers). The richest eudicotyledonous family was Ericaceae (5 taxa; Fig. 17).

The most species-rich habit class was trees and shrubs (14 taxa; 10 species, 4 varieties), followed by herbs (3 taxa), and vines (1 taxa; Fig. 16). Among the trees and shrubs, the Ericaceae (5 taxa) is the most species-rich family. No exotic taxa or taxa of conservation concern occured at this site. One species (Rhus copallinum L.) was unique to this Carolina bay lake (i.e., it was not found/reported from any other bay lake in this study; Suppl. material 5).

Bay Tree Lake

The littoral zone vascular flora of Bay Tree Lake is comprised of 56 taxa (48 species, 2 subspecies, and 6 varieties), in 47 genera and 34 vascular plant families (Table 11; Suppl. material 6). All but 2 taxa from Bay Tree Lake were vouchered; Decodon verticillatus and Pontederia cordata L. var. cordata were personal observations. No species of conservation concern were collected or reported from Bay Tree Lake’s littoral zone. One exotic taxon (Hypochaeris radicata) was collected from this site (Suppl. material 6). Twelve taxa are unique to this bay lake (i.e., they were not found/reported from any other bay lake in this study; Suppl. material 6: [Amelanchier canadensis (L.) Medik., Carex longii Mack., Cyperus odoratus L. var. odoratus, Diodia virginiana L., Fuirena pumila (Torr.) Spreng., Hypochaeris radicata, Juncus acuminatus Michx., Krigia virginica (L.) Willd., Nuttallanthus canadensis (L.) D.A. Sutton, Panicum virgatum, Rumex hastatulus Baldwin, Smilax glauca Walter, and Stipulicida setacea Michx. var. setacea]).

The richest eudicotyledon families are Asteraceae (3 taxa), followed by Ericaceae (2 taxa) and Aquifoliaceae (2 taxa;) . The richest monocotyledonous families are Poaceae (7 taxa; 6 species, 1 subspecies), Cyperaceae (5 taxa; 4 species, 1 variety), and Juncaceae (5 taxa). The richest monocotyledon genera are Juncus (5 taxa; 3 species, 1 subspecies, 1 variety) and Panicum (3 taxa).

The most species-rich habit class was herbs (35 taxa; 29 species, 2 subspecies, 4 varieties), followed by trees and shrubs (16 taxa; 15 species, 1 variety), and vines (4 species, 1 variety; Fig. 16). Among the herbs, Poaceae (7 taxa; 6 species, 1 subspecies), Cyperaceae (5 taxa; 4 species, 1 variety), Juncaceae (5 taxa), and Asteraceae (3 taxa) are the most species-rich families. Among the trees and shrubs, Cupressaceae (3 taxa), Aquifoliaceae (2 taxa), and Ericaceae (2 taxa) are the most species-rich families.

Horseshoe Lake

The littoral zone vascular flora of Horseshoe Lake is comprised of 52 taxa (45 species, 2 subspecies, and 5 varieties), in 41 genera and 29 vascular plant families (Table 11; Suppl. material 6). All but three taxa from Horseshoe Lake were vouchered; Eleocharis baldwinii/vivipara, Rhexia aristosa, and Tillandsia usneoides were the only taxa not vouchered from the site. No exotic taxa were collected from this site. Sixteen taxa are unique to this bay lake (i.e., they were not found/reported from any other bay lake in this study; Suppl. material 6). Five taxa of conservation concern were collected or reported from this site (Rhexia aristosa, Rhynchospora alba, Rhynchospora inundata, Sagittaria isoetiformis J.G. Sm., and Xyris smalliana; Table 9).

The richest eudicotyledon families are Ericaceae (4 taxa), Lentibulariaceae (3 taxa) and Melastomataceae (3 taxa). The richest eudicotyledonous genera are Rhexia (3 taxa), Utricularia (3 taxa), followed by Hypericum (2 taxa). The richest monocotyledonous families are Cyperaceae (5 taxa), Juncaceae (4 taxa), Poaceae (3 taxa), followed by Orchidaceae (2 taxa), Smilacaceae (2 taxa) and Xyridaceae (2 taxa). The richest monocotyledonous genera are Juncus (4 taxa), followed by Rhynchospora (2 taxa), Smilax (2 taxa), and Xyris (2 taxa).

The most species-rich habit class was herbs (38 taxa; 31 species, 2 subspecies, 4 varieties), followed by trees and shrubs (11 taxa; 10 species, 1 variety), and vines (3 taxa; Fig. 16). Among the herbs, Cyperaceae (6 taxa), Juncaceae (4 taxa), followed by Lentibulariaceae (3 taxa), Melastomataceae (3 taxa), Poaceae (3 taxa), Orchidaceae (2 taxa), and Xyridaceae (2 taxa) are the most species-rich families. Among the trees and shrubs, the most species-rich family is Ericaceae (4 taxa).

Jones Lake

The littoral zone vascular flora of Jones Lake is comprised of 33 taxa (29 species, 1 subspecies, and 3 varieties), in 31 genera and 23 vascular plant families (Table 11; Suppl. material 6). All taxa, save for Cyrilla racemiflora, were vouchered by the first author or others. No exotic taxa were collected from this site. Two taxa are unique to this bay lake (i.e., they were not found/reported from any other bay lake in this study; Suppl. material 6: [Cyperus polystachyos Rottb., Rhynchospora inexpansa (Michx.) Vahl]). Xyris smalliana was the only species of conservation concern collected from this site (Table 9).

The richest eudicotyledonous family is Ericaceae (5 taxa). The richest eudicotyledonous genus is Lyonia (2 taxa; 1 species, 1 variety). The richest monocotyledonous families are Cyperaceae (3 taxa) and Poaceae (3 taxa). Monocotyledons are comprised of ten different genera.

The most species-rich habit class was trees and shrubs (20 taxa; 16 species, 1 subspecies, 3 varieties), followed by herbs (11 taxa), and vines (2 taxa; Fig. 16). Among the herbs, Cyperaceae (3 taxa) and Poaceae (3 taxa) are the most species-rich families. Among the trees and shrubs, Ericaceae (5 taxa) and Cupressaceae (3 taxa) are the most species-rich families.

Lake Waccamaw

The littoral zone vascular flora of Lake Waccamaw is comprised of 145 taxa (122 species, 3 subspecies, 19 varieties, 1 hybrid), in 111 genera and 72 vascular plant families (Table 11; Suppl. material 6). Of the 145 total catalogued taxa, 127 are vouchered and 18 are known only from reports (Suppl. material 6). Twenty-six species of conservation concern were collected or reported from Lake Waccamaw’s littoral zone. Four exotic taxa (Alternanthera philoxeroides [Amaranthaceae], Colocasia esculenta [Araceae], Hydrilla verticillata [Hydrocharitaceae], Triadica sebifera [Euphorbiaceae]) are known from this site. Ninety-five taxa are unique to Lake Waccamaw (i.e., they were not found/reported from any other bay lake in this study; Suppl. material 6).

The richest eudicotyledonous families are Asteraceae (10 taxa; 8 species, 1 variety, 1 hybrid), followed by Lentibulariaceae (4 taxa), Ericaceae (3 taxa), Rosaceae (3 taxa), and Salicaceae (3 taxa). The richest eudicotyledonous genera are Utricularia (4 taxa), Eupatorium L. (2 taxa), Hypericum (2 taxa), Ludwigia L. (2 taxa), Nyssa (2 taxa), and Salix L. (2 taxa). The richest monocotyledonous families are Poaceae (17 taxa; 13 species, 1 subspecies, 3 varieties), Cyperaceae (14 taxa; 11 species, 3 varieties), Alismataceae (4 taxa), Juncaceae (3 taxa), Orchidaceae (3 taxa), and Smilacaceae (3 taxa). The richest monocotyledonous genera are Dichanthelium (Hitchc. & Chase) Gould (6 taxa; 5 species and 1 variety), Rhynchospora (6 taxa; 5 species and 1 variety), Sagittaria L. (4 taxa), Juncus (3 taxa; 3 species, 1 subspecies, 1 variety) and Smilax L. (3 taxa).

The most species-rich habit class was herbs (96 taxa; 80 species, 3 subspecies, 13 varieties, 1 hybrid), followed by trees and shrubs (36 taxa; 32 species, 4 varieties), and vines (13 taxa; 11 species, 2 varieties; Fig. 16). Among the herbs, the Poaceae (16 taxa; 13 species, 1 subspecies, 2 varieties), Cyperaceae (14 taxa; 11 species, 3 varieties), Asteraceae (8 taxa; 6 species, 1 variety, 1 hybrid), Alismataceae (4 taxa), Lentibulariaceae (4 taxa), Juncaceae (3 taxa), and Orchidaceae (3 taxa) are the most species-rich families. Among the trees and shrubs, the Ericaceae (3 taxa), Rosaceae (3 taxa), Salicaceae (3 taxa), Betulaceae (2 taxa), Cupressaceae (2 taxa), Nyssaceae (2 taxa), and Sapindaceae (2 taxa) are the most species-rich families.

Little Singletary Lake

The littoral zone flora of Littoral Singletary Lake is comprised of 39 taxa (35 species, 1 subspecies, 3 varieties), in 32 genera and 21 vascular plant families (Table 11; Suppl. material 6). All of the 39 total catalogued taxa were vouchered (i.e., no taxa were known strictly from reports or observations; Suppl. material 6). Two species of conservation concern (i.e., Eleocharis equisetoides and Eleocharis vivipara) were collected from Little Singletary Lake’s littoral zone (Table 9). No exotic taxa are known from this site. Three taxa are unique to Little Singletary Lake (i.e., they were not found/reported from any other bay lake in this study; Suppl. material 6: [Agrostis hyemalis (Walter) Britton, Sterns & Poggenb., Rhexia virginica L., and Xyris jupicai Rich.])

The richest eudicotyledonous genus is Rhexia (2 taxa). The richest monocotyledonous families are Cyperaceae (6 taxa; 5 species and 1 variety), Juncaceae (4 taxa; 2 species, 1 subspecies, 1 variety), and Poaceae (3 taxa). The richest monocotyledonous genera are Juncus (4 taxa), Eleocharis (3 taxa), and Panicum (2 taxa).

The most species-rich habit class was herbs (23 taxa; 20 species, 1 subspecies, 2 varieties), followed by trees and shrubs (15 taxa; 14 species and 1 variety), and vines (1 taxon; Fig. 16). Among the herbs, the Cyperaceae (6 taxa), Juncaceae (4 taxa), and Poaceae (3 taxa) are the most species-rich families. Among the trees and shrubs, the Ericaceae (5 taxa) is the most species-rich family.

Salters Lake

The littoral zone flora of Salters Lake is comprised of 22 taxa (16 species, 2 subspecies, 4 varieties), in 18 genera and 16 vascular plant families (Table 11; Suppl. material 6). Twenty of the twenty-three total catalogued taxa were vouchered; Decodon verticillatus, Nyssa biflora, and Xyris iridifolia, were reports or personal observations (Suppl. material 6). Two species of conservation concern (i.e., Xyris iridifolia and Xyris smalliana) were collected/reported from Salters Lake’s littoral zone (Suppl. material 6; Table 9). No exotic taxa are known from this site. One taxon is unique to Salters Lake (i.e., not found/reported from any other bay lake in this study; Suppl. material 5: [Xyris iridifolia])

The richest eudicotyledon family is Ericaceae (5 taxa). The richest eudicotyledonous genera are Lyonia (2 taxa) and Vaccinium (2 taxa). The richest monocotyledonous family is Xyridaceae (2 taxa). The richest monocotyledon genus is Xyris (2 taxa).

The most species-rich habit class was trees and shrubs (15 taxa; 11 species, 1 subspecies, 3 varieties), herbs (5 taxa; 4 species and 1 subspecies), and vines (2 taxa; Fig. 16). Among the trees and shrubs, the Ericaceae (5 taxa) and Cupressaceae (2 taxa) are the most species-rich families. Among the herbs, the Xyridaceae (2 taxa) is the most species-rich family.

Singletary Lake

The littoral zone vascular flora of Singletary Lake is comprised of 36 taxa (32 species, 1 subspecies, 3 varieties), in 30 genera and 22 vascular plant families (Table 11; Suppl. material 6). All thirty-six total catalogued taxa were vouchered (i.e., none were reports or personal observations; Suppl. material 6). One taxon from Singletary Lake’s littoral zone is of conservation concern (i.e., Xyris smalliana; Suppl. material 6; Table 9). No exotic taxa are known from this site. One taxon is unique to Salters Lake (i.e., not found/reported from any other bay lake in this study; Suppl. material 6: [Rhododendron viscosum (L.) Torr. var. serrulatum (Small) H.E. Ahles]).

The richest eudicotyledonous families are Ericaceae (7 taxa) and Rosaceae (2 taxa). The richest eudicotyledonous genus is Vaccinium (2 taxa). The richest monocotyledonous families are Juncaceae (3 taxa), Poaceae (2 taxa), and Xyridaceae (2 taxa). The richest monocotyledonous genera are Juncus (3 taxa) and Xyris (2 taxa).

The most species-rich habit class was trees and shrubs (22 taxa; 19 species and 3 varieties), herbs (11 taxa; 10 species and 1 subspecies), and vines (3 taxa; Fig. 16). Among the trees and shrubs, the Ericaceae (7 taxa), Cupressaceae (3 taxa), Pinaceae (2 taxa), and Rosaceae (2 taxa) are the most species-rich families. Among the herbs, the Juncaceae (3 taxa), Poaceae (2 taxa), and Xyridaceae (2 taxa) are the most species-rich families.

White Lake

White Lake was not included in this study due to the severity of the lake’s shoreline development. A provisional checklist of plants known to occur within the littoral zone of White Lake (from historical vouchers, personal observation, and literature review) is provided in Suppl. material 7. The intent of the provisional checklist is to provide a baseline for future research in this lake.

Materials and methods

This work is restricted to the littoral zone vascular flora of unaltered Carolina bay lake shorelines. The littoral zone was defined as the zone of vegetation occurring between the maximum annual high water mark and the point at which submerged aquatic plants cease to persist (Fig. 4). Unaltered shorelines were defined as those lacking residential or commercial development (therefore, the entirety of White Lake and the developed shorelines of Lake Waccamaw and Bay Tree Lake were not included in this inventory).

During the 2013 and 2014 growing seasons, 36 total visits were made to the eight study sites meeting the criteria articulated above (i.e., Bakers Lake, Bay Tree Lake, Horseshoe Lake, Jones Lake, Lake Waccamaw, Little Singletary Lake, Salters Lake, Singletary Lake), resulting in 121 field hours and the identification of 204 taxa (species, subspecies, and varieties). A 10-foot aluminum boat with a transom-mounted trolling motor was used to transport equipment along Carolina bay lake shorelines. Where water was too shallow for the use of the trolling motor, we walked and pulled the boat by rope. GPS locations (NAD 83) were taken at numerous intervals and associated with all specimens collected within 30 m of each point. Digital photographs of plant habit and overall morphology were taken prior to collection using a Panasonic Lumix FZ−150. Plant specimens were pressed while in the field. Tissue samples were taken in the field and dessicated with blue indicating silica gel (purchased from Delta Enterprises Inc.) in ziploc bags. Voucher specimens and tissue samples were deposited respectively at the North Carolina State University Vascular Plant Herbarium (NCSC) and its DNA bank. The entirety of Carolina bay lake shorelines was surveyed, but it was quickly observed that all shorelines, save for the southernmost, were relatively depauperate. All taxa occurring along western, northern, and eastern shorelines could be found within the littoral zone of the southern shoreline, but the inverse did not hold true. The significantly gentler hydrography (see Frey 1949 for lake longitudinal profiles), and consequently wider littoral zone of southern shorelines, produces a more species-rich macrophyte community. Consequently, survey time was much longer on the southern, more diverse shorelines of Carolina bay lakes.

The flora is organized by the following major vascular plant groups: (1) pteridophytes, (2) gymnosperms, (3) monocots, and (4) basal angiosperms, magnoliids, and eudicotyledons. Dichotomomous keys are provided to each major group, as well as to families, genera, and species within each group. Notes are provided above some keys to aid in the identification process. Within each group, taxa are arranged alphabetically, by family, then genus, then species.

The following information is provided for each taxon account: taxon concept mapping, basionym, conservation status, habit, habitat, flowering and fruiting phenology, abundance, and presence/absence data for each site (Suppl. material 3). Unless stated otherwise, accepted taxon concepts follow Weakley (2012) and are tied to those in the following major works: RAB = Radford et al. (1968); GW = Godfrey and Wooten (1979), Godfrey and Wooten (1981); FNA = Flora of North America (pteridophytes: Blechnaceae [Cranfill 1993], Dryopteridaceae [Smith 1993b, Wagner and Montgomery 1993, Smith 1993b], Lycopodiaceae [Wagner and Beitel 1993], Osmundaceae [Whetstone and Atkinson 1993, Polypodiaceae [Andrews and Windham 1993]; gymnosperms: Cupressaceae [Michener 1993, Watson 1993, Watson and Eckenwalder 1993], Pinaceae [Kral 1993]; monocots: Alismataceae [Durand 2000], Araceae [Thompson 2000], Bromeliaceae [Luther and Brown 2000], Burmanniaceae [Lewis 2002], Cyperaceae [Ball and Reznicek 2002, Ball et al. 2002b, Kral 2002a, Kral 2002b, Kral and Persoon 2002, Mastrogiuseppe 2002, Mastrogiuseppe et al. 2002, Reznicek 2002, Reznicek and Catling 2002, Smith et al. 2002, Tucker 2002, Tucker et al. 2002], Eriocaulaceae [Kral 2000a], Haemodoraceae [Robertson 2002], Hydrocharitaceae [Haynes 2000a, Haynes 2000b], Hypoxidaceae [Herndon 2002], Juncaceae [Brooks and Clemants 2000], Mayacaceae [Faden 2000], Orchidaceae [Goldman et al. 2002, Hágsater et al. 2002, Romero-Gonzáles et al. 2002, Sheviak 2002, Sheviak and Brown 2002, Sheviak and Catling 2002], Poaceae [Barkworth 2003a, Barkworth 2003b, Campbell 2003, Clark and Triplett 2007, Daniel 2007, Freckmann and Lelong 2003a, Freckmann and Lelong 2003b, Harvey 2007, Peterson 2003, Terrell 2007, Wipff 2003], Pontederiaceae [Adanson et al. 2002], Smilacaceae [Holmes 2002], Xyridaceae [Kral 2000a]; basal angiosperms, magnoliids, and eudicots: Altingiaceae [Meyer 1997a], Amaranthaceae [Clemants 2003], Asteraceae [Bogler 2006, Chambers and O'Kennon 2006, Haines 2006, Holmes 2006, Karaman-Castro and Urbatsch 2006, Lamont 2006, Nesom 2006a, Nesom 2006b, Semple and Cook 2006, Siripun and Schilling 2006, Strother and Weedon 2006, Sundberg and Bogler 2006], Betulaceae [Furlow 1997], Cabombaceae [Wiersema 1997b], Caryophyllaceae [Swanson and Rabeler 2005], Clethraceae [Tucker and Jones 2009], Cyrillaceae [Lemke 2009], Ebenaceae [Eckenwalder 2009], Ericaceae [Dorr 2009, Fabijan 2009, Judd 2009, Judd and Kron 2009, Tucker 2009b, Tucker 2009a, Vander Kloet, S.P. 2009], Fagaeae [Jensen 1997], Iteaceae [Morin 2009], Juglandaceae [Stone 1997], Lauraceae [Wofford 1993], Magnoliaceae [Meyer 1997b], Myricaceae [Bornstein 1997], Nelumbonaceae [Wiersema 1997a], Nymphaeaceae [Wiersema and Hellquist 1997], Platanaceae [Kaul 1997], Polygonaceae [Mosyaking 2005], Ranunculaceae [Pringle 1997], Salicaceae [Argus et al. 2010], Sarraceniaceae [Mellichamp and Chase 2009], Theaceae [Prince 2009], Ulmaceae [Sherman-Broyles 1997]). Three symbols are used to relate whether our taxon concepts used here are equivalent (=), narrower (<), or broader (>) than those of other works. For example, the statement “= RAB, FNA” means that the taxon concept, as well as the species name used here, is the same as that used in RAB and FNA (see Dryopteris ludoviciana (Kunze) Small). The use of a “less than” symbol (e.g., “< Onoclea sensibilis L. – RAB, FNA”), indicates that the taxon concept used here is narrower than that used by RAB and FNA (alternatively, a “greater than” symbol would mean that the concept of a particular taxon is broader than in the cited works). An equals symbol followed by a different species name than the one bolded, indicates that the taxon concept used here is the same as in the work cited, except that the taxon was treated under a different name in the work cited (see Sagittaria filiformis J.G. Sm. vs. Sagittaria stagnorum Small).

Abundance estimates following the recommendations of Palmer et al. (1995) are provided for each lake in which a taxon was collected or observed by the current author (Table 12; Suppl. material 3). Taxa designated as “exotic” are not native to North America and are indicated by an asterisk preceding the scientific name. The conservation status and rank of species of conservation concern precede the habitat description in each taxon entry (e.g., E, FSC; S1, G2. “Habitat description”). Conservation status and rank of species are designated according to NatureServe (2012), the North Carolina Plant Conservation Program (2010), and the North Carolina Natural Heritage Program List of Rare Plants (Robinson and Finnegan 2014). Unvouchered taxa (i.e., those known only from reports or personal observations) are given one of four symbols in taxon entries (• = the first author observed the species while in the field, but was not able to collect a viable voucher specimen, ♦ = the taxon was reported by the Carolina Vegetation Survey (Peet et al. 2013a, Peet et al. 2013b), = ► the taxon was reported by the North Carolina Natural Heritage Program (North Carolina Natural Heritage Program 2014), ¤ = the taxon was reported by the North Carolina State University Crop Science Program; Rob Richardson and Justin Nawrocki, pers. comm, April 9, 2015).

Descriptions for estimating the abundance of taxa (adapted from Palmer et al. 1995)

Density

Description

Abundant

Dominant or co-dominant in one or more communities.

Frequent

Easily seen or found in one or more common communities but not dominant in any common community

Occasional

Widely scattered but not difficult to find

Infrequent

Difficult to find with few individuals or colonies but found in several locations

Rare

Very difficult to find and limited to one or very few locations or uncommon communities

When available, digital photographs and line drawings were obtained from: Britton and Brown (1913), Center for Aquatic and Invasive Plants, University of Florida, IFAS (2015), Hitchcock and Chase (1951), Mickel (1979), and United States Department of Agriculture, Natural Resources Conservation Service (USDA- NRCS) (2015).

In addition, relevant historical vouchers are cited based on systematic searches of the three major herbaria−DUKE, NCSC, and NCU. Unfortunately, it is not uncommon to find historical specimens containing vague habitat or locality descriptions. For a taxon to be included in the present study, a clear label statement referencing Carolina bay lake shoreline habitat was required (e.g., “collected from peat-drained lake bed of Suggs Mill Pond”). Herbarium vouchers meeting this criterion were annotated (following taxon concepts accepted here) and their label information was subsequently entered into spreadsheets for organization. Label information for new collections resulting from this study was captured in a DarwinCore compliant spreadsheet for upload to the online portal of the Southeastern Regional Network of Expertise and Collections (www.sernecportal.org), which feeds into iDigBio and the Global Biodiversity Data Facility (GBIF).

PTERIDOPHYTES

Families represented: 6

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Blechnaceae

Anchistea virginica (L.) C. Presl

Nomenclature: 

Basionym: Blechnum virginicum L.

Taxon concept: [= Woodwardia virginica (L.) Sm. − RAB, FNA, Weakley]

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Distribution: 

Bakers Lake (Infrequent): Howell BALA−14 (NCSC!)

Bay Tree Lake (Occasional): Howell BATR−4, 24 (NCSC!)

Jones Lake (Rare): Howell JOLA−44 (NCSC!)

Lake Waccamaw (Infrequent): Howell LAWA−59 (NCSC!)

Little Singletary Lake (Occasional): Howell LISI−42 (NCSC!)

Notes: 

Perennial herbs. Upper eulittoral zone; typically found in saturated soils or rooted on logs, stumps, and other debris (NLSS–C, NLSS–LW, NLSM–T). Jun–Sep. Fig. 18

Figure 18.

Anchistea virginica (digital photographs taken by Nathan Howell)

aSpecimen: Howell BATR24 (NCSC)
bStipe; note the dark purple coloration.
cMature frond
dPinna; note the chain-like venation pattern along the sides of leaflet midveins.

Lorinseria areolata (L.) C. Presl

Nomenclature: 

Basionym: Acrostichum areolatum L.

Taxon concept: [= Woodwardia areolata (L.) T. Moore − RAB, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Occasional): Howell BATR−5, 26 (NCSC!)

Lake Waccamaw: Wilbur 84200 (DUKE!)

Little Singletary Lake (Occasional): Howell LISI–6 (NCSC!)

Singletary Lake: Hueske s.n. (NCU!)

Notes: 

Perennial herbs. Upper eulittoral zone; typically found in saturated soils or rooted on logs, stumps, and other debris (NLSS–C, NLSS–LW, NLSM–T) . May–Sep. Fig. 19

Figure 19.

Lorinseria areolata (digital photographs taken by Nathan Howell)

aSpecimen: Howell LISI6 (NCSC)
bMature frond
cFrond underside
dPinnae

Dryopteridaceae

Dryopteris ludoviciana (Kunze) Small

Nomenclature: 

Basionym: Aspidium ludovicianum Kunze

Taxon concept: [= RAB, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Bennedict 1247 & 2298 (NCU!); Blomquist & Correll 7625 (NCU!)

Notes: 

Perennial herbs. Juncture of eulittoral and supralittoral zones (NLSS–LW). Jun–Sep. This species was not encountered by the first author, but voucher specimens (see above) place it within close proximity of Lake Waccamaw’s shoreline (i.e., it has the potential to occur at the uppermost portions of the littoral zone where the swamp forest adjoins the shoreline community on the southwest side of the lake). Fig. 20

Figure 20.

Dryopteris ludoviciana (from Mickel 1979)

Lycopodiaceae

Lycopodiella appressa (Chapm.) Cranfill

Nomenclature: 

Basionym: Lycopodium inundatum L. var. appressum Chapm.

Taxon concept: [= Lycopodium appressum (Chapm.) F.E. Lloyd & Underw. − RAB; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake: Wilbur 48656 (DUKE!)

Horseshoe Lake (Infrequent): Howell HOLA−52 (NCSC!)

Lake Waccamaw (Infrequent): Howell LAWA−110 (NCSC!)

Notes: 

Perennial herbs. Upper eulittoral zone; usually in association with saturated peaty to sandy soils (NLSS–LW, CPSI–CG). Jul–Sep. Fig. 21

Figure 21.

Lycopodiella appressa (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA110 (NCSC)
bSpecimen: Howell HOLA–52 (NCSC)
cRooting stems where making contact with soil
dTerminal strobilus

Onocleaceae

Onoclea sensibilis L.

Nomenclature: 

Taxon concept: [< O. sensibilis L. – RAB, FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Wilbur 84220 (DUKE!)

Notes: 

Perennial herbs. Upper eulittoral zone (NLSS−LW). May−Jun. This species was not encountered by the first author in the field, but a single voucher (see above) places it within close proximity to Lake Waccamaw’s southwest shoreline. Fig. 22

Figure 22.

Onoclea sensibilis (digital photograph taken by Nathan Howell)

Osmundaceae

Osmunda spectabilis Willd.

Nomenclature: 

Taxon concept: [< O. regalis L. var. spectabilis (Willd.) A. Gray − RAB, FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Infrequent): Howell LAWA−58, 87, 90 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone; sometimes establishing itself on old stumps and logs (NLSS–LW). Mar–Jun. Fig. 23

Figure 23.

Osmunda spectabilis (digital photographs taken by Nathan Howell)

aSpecimen: LAWA-90 (NCSC)
bHabit
cSterile frond
dSterile and fertile fronds

Polypodiaceae

Pleopeltis polypodioides var. michauxiana (Weath.) E.G. Andrews & Windham

Nomenclature: 

Basionym: Polypodium polypodioides (L.) Watt var. michauxianum Weath.

Taxon concept: [< Polypodium polypodioides (L.) Watt – RAB; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Infrequent): Howell LAWA−47 (NCSC!)

Salters Lake (Infrequent): Howell SALA−1 (NCSC!)

Notes: 

Perennial, frequently epiphytic, herbs. Eulittoral zone; commonly on large limbs and trunks of Taxodium and Nyssa (NLSS−C, NLSS−LW). Jun−Oct. Fig. 24

Figure 24.

Pleopeltis polypodioides var. michauxiana (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-47 (NCSC)
bSpecimen: Howell SALA-1 (NCSC)
cHabit
dHabit

GYMNOSPERMS

Families represented: 2

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Cupressaceae

Chamaecyparis thyoides (L.) Britton, Sterns, & Poggenb.

Nomenclature: 

Basionym: Cupressus thyoides L.

Taxon concept: [= RAB, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Infrequent): Howell BATR−2 (NCSC!)

Horseshoe Lake (Occasional): Howell HOLA−2, 13 (NCSC!)

Jones Lake (Occasional): Brown s.n. (NCSC!); Howell JOLA−1, 23 (NCSC!); Lance s.n. (NCU!); Russell 1304 (NCSC!)

Little Singletary Lake (Occasional): Howell LISI−8, 26 (NCSC!)

Singletary Lake (Infrequent): Howell SILA−14 (NCSC!)

Notes: 

Trees. At or just below the juncture of the supralittoral and eulittoral zones; often in saturated peaty or sandy soil (NLSS–C, NLSS–LW, NLSM–T). Mar–Apr; Oct– Nov. Fig. 25

Figure 25.

Chamaecyparis thyoides (digital photographs taken by Nathan Howell)

aSpecimen: Howell HOLA-2 (NCSC)
bBark
cLeaves and developing seed cones
dLeaves and mature seed cones

Taxodium ascendens Brongn.

Nomenclature: 

Taxon concept: [= RAB; < T. distichum L. var. imbricarium (Nutt.) Croom − FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Bakers Lake (Abundant): Howell BALA−15 (NCSC!)

Bay Tree Lake (Abundant): Howell BATR−7 (NCSC!)

Horseshoe Lake (Abundant): Howell HOLA−10 (NCSC!)

Jones Lake (Abundant): Howell JOLA−3, 22 (NCSC!); Krings 508 (NCSC!); Wilbur 57584 (DUKE!)

Lake Waccamaw (Abundant): Howell LAWA−13 (NCSC!)

Little Singletary Lake (Abundant): Howell LISI−4, 20 (NCSC!)

Salters Lake (Abundnat): Howell SALA−8 (NCSC!)

Singletary Lake (Abundant): Howell SILA−13 (NCSC!); Wilbur 27966 (DUKE!)

Notes: 

Trees. Eulittoral zone (NLSS–C, NLSS–LW, NLSM–T, NLSM−LWP, CPSI–CG). Mar– Apr; Oct. Fig. 26

Figure 26.

Taxodium ascendens (digital photographs taken by Nathan Howell)

aSpecimen: Howell BALA-15 (NCSC)
bHabit
cLeaves
dPollen cones

Taxodium distichum (L.) Rich.

Nomenclature: 

Basionym: Cupressus disticha L.

Taxon concept: [= RAB; < T. distichum (L.) Rich. var. distichum – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake: Wilbur 61464 (DUKE!)

Jones Lake: Stone 3704 (DUKE!)

Lake Waccamaw: ♦

Salters Lake: Beckman & Linnenburger 38 (DUKE!)

Singletary Lake: Crosby 4032 (DUKE!)

Notes: 

Trees. Eulittoral zone (NLSS–C, NLSS–LW, NLSM−T). Infrequent. Mar–Apr; Oct. Fig. 27

Figure 27.

Taxodium distichum (from Britton and Brown 1913)

Pinaceae

Pinus serotina Michx.

Nomenclature: 

Taxon concept: [= RAB, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Jones Lake (Rare): Howell JOLA−14 (NCSC!)

Singletary Lake (Rare): Howell SILA−37 (NCSC!)

Notes: 

Trees. Juncture of supralittoral and eulittoral zones (NLSS–C). Apr–Aug (or any time of the year in response to fire). Fig. 28

Figure 28.

Pinus serotina (digital photographs taken by Nathan Howell)

aSpecimen: Howell SILA-37 (NCSC)
bSpecimen: Howell JOLA-14 (NCSC)
cHabit
dMature seed cones

Pinus taeda L.

Nomenclature: 

Taxon concept: [= RAB, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Rare): Howell LAWA−71 (NCSC!)

Little Singletary Lake (Infrequent): Howell LISI−27 (NCSC!)

Singletary Lake (Rare): Howell SILA−12 (NCSC!)

Notes: 

Trees. Juncture of supralittoral and eulittoral zones (NLSS–C, NLSS–LW). Mar–Apr; Oct– Nov. Fig. 29

Figure 29.

Pinus taeda (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-71 (NCSC)
bSpecimen: Howell SILA-12 (NCSC)
cYoung trees
dMature seed cones

MONOCOTYLEDONS

Families represented: 17

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Alismataceae

Sagittaria filiformis J.G. Sm.

Nomenclature: 

Taxon concept: [= S. stagnorum Small – GW; S. subulata L. Buchenau var. gracillima (S. Watson) J.G. Sm.; = FNA, Weakley]

Material    Download as CSV 
Conservation status: 

SR−P; SH, G4G5.

Distribution: 

Lake Waccamaw: Blomquist & Schuster 16191 (DUKE!)

Notes: 

Perennial herbs. Eulittoral and infralittoral zones (NLSS−LW, NLSM−LWP). May−Sep. The first author has not encountered this taxon in the field, but a single voucher specimen (see above) confirms its historic presence within the lake. Fig. 30

Figure 30.

Sagittaria filiformis (from Center for Aquatic and Invasive Plants, University of Florida, IFAS 2015)

Sagittaria graminea Michx.

Nomenclature: 

Taxon concept: [= S. graminea Michx. var. graminea – RAB, GW; = S. graminea Michx. ssp. graminea – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Frequent): Howell LAWA−19, 57 (NCSC!); Radford s.n. (NCU!); ♦

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW, NLSM−LWP). May−Nov. Fig. 31

Figure 31.

Sagittaria graminea (from Center for Aquatic and Invasive Plants, University of Florida, IFAS 2015)

Sagittaria isoetiformis J.G. Sm.

Nomenclature: 

Taxon concept: [< S. teres S. Watson (misapplied) – RAB; = GW, FNA, Weakley]

Material    Download as CSV 
Conservation status: 

State T; S2, G4?.

Distribution: 

Horseshoe Lake (Infrequent): Grant s.n. (NCU!); Howell HOLA−34 (NCSC!)

Lake Waccamaw: LeBlond 5792D (NCU!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW, NLSM−LWP, CPSI−CG, FB). Jun−Sep. Fig. 32

Figure 32.

Sagittaria isoetiformis (digital photographs taken by Nathan Howell)

aSpecimen: Howell HOLA-34 (NCSC)
bLeaf
cFlower and floral buds
dFlower and floral buds
eInflorescence (note bract)
fInflorescence bract detail

Sagittaria weatherbiana Fernald

Nomenclature: 

Taxon concept: [= S. graminea Michx. var. weatherbiana – RAB, GW; = S. graminea Michx. ssp. weatherbiana – FNA; = Weakley]

Material    Download as CSV 
Conservation status: 

State E, FSC; S2, G3G4.

Distribution: 

Lake Waccamaw: Adams s.n. (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW, NLSM−LWP). Apr−Jun.

Araceae

Colocasia esculenta (L.) Schott

Nomenclature: 

Basionym: Arum esculentum L.

Taxon concept: [= GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Infrequent): Howell LAWA−93 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW, NLSM−LWP). “Generally infertile in our area” (Weakley 2012). This species is exotic and has become naturalized in roadside ditches, canals, and portions of the lakes shoreline. It spreads by way of rhizome dispersal, which is almost cartainly caused by residential homeowners digging up rhizomes from their flower beds and either tossing them into the lake or into the canals that surround the lake. Fig. 33

Figure 33.

Colocasia esculenta (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-93 (NCSC)
bHabit

Genus Wolffia Horkel ex Schleid.

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Notes: 

The first author has not encountered taxa within this genus in the field; however, the Carolina Vegetation Survey reported “Wolffia spp.” from the southwest side Lake Waccamaw. Although a species-level identification has not been made, a key to the two species most likely to inhabit this location is provided in the Identification Keys section below.

Bromeliaceae

Tillandsia usneoides (L.) L.

Nomenclature: 

Basionym: Renealmia usneoides L.

Taxon concept: [= RAB, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bakers Lake (Occasional): •

Bay Tree Lake (Frequent): Howell BATR−49 (NCSC!)

Horseshoe Lake (Occasional): •

Jones Lake (Occasional): Howell JOLA–33 (NCSC!)

Lake Waccamaw (Abundant): Howell LAWA−46, 84 (NCSC!)

Little Singletary Lake (Occasional): Howell LISI−18 (NCSC!)

Salters Lake (Occasional): Howell SALA−9 (NCSC!)

Singletary Lake (Frequent): Howell SILA−6, 20 (NCSC!)

Notes: 

Perennial, epiphytic herbs. Eulittoral zone; common in low-hanging limbs of Taxodium or Nyssa (NLSS−C, NLSS−LW, NLSM−T, CPSI−CG). Apr−Jun. Fig. 34

Figure 34.

Tillandsia usneoides (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-46 (NCSC)
bHabit (draped on branches of Taxodium ascendens)
cFlower
dCapsule

Burmanniaceae

Burmannia capitata (Walter ex J.F. Gmel.) Mart.

Nomenclature: 

Basionym: Vogelia capitata Walter ex J.F. Gmel.

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: LeBlond & Franklin 6578 (NCU!)

Notes: 

Annual herbs. Eulittoral zone (NLSS−LW). Jul−Nov. Fig. 35

Figure 35.

Burmannia capitata (digital photograph taken by Alexander Krings)

Cyperaceae

Carex alata Torr.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Rare): Howell LAWA−98 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone; usually at or just below the juncture of the supralittoral and eulittoral zones (NLSS−LW). May−Jun. Fig. 36

Figure 36.

Carex alata (digital photograph taken by Nathan Howell)

aSpecimen: Howell LAWA-98 (NCSC)
bInflorescence

Carex longii Mack.

Nomenclature: 

Taxon concept: [< C. albolutescens Schwein. – RAB, GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Rare): Howell BATR−34 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone; typically at or just below the juncture of the supralittoral and eulittoral zones. May−Jun. Fig. 37

Figure 37.

Carex longii (digital photographs taken by Nathan Howell)

aSpecimen: Howell BATR-34 (NCSC)
bInflorescence

Carex lupulina Muhl. ex Willd.

Nomenclature: 

Taxon concept: [= RAB; < C. lupulina Muhl. ex Willd. – GW (see C. lupuliformis); = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Rare): Howell LAWA−136 (NCSC!)

Notes: 

Perennial herbs. Juncture of the eulittoral and supralittoral zones (NLSS−LW). Jun−Sep. A taxon of bottomland forests throughout the state, this large-fruited sedge occurs where bottomland swamp forests abut the shoreline of Lake Waccamaw. Fig. 38

Figure 38.

Carex lupulina (digital photograph taken by Nathan Howell)

aSpecimen: Howell LAWA-136 (NCSC)
bInflorescence

Carex striata var. brevis L.H. Bailey

Nomenclature: 

Taxon concept: [< C. walteriana L.H. Bailey – RAB, GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Horseshoe Lake: Buell 2279 (DUKE!, NCSC!)

Notes: 

Perennial herbs. Eulittoral zone; typically in acidic, saturated, peaty soils (CPSI−CG, FB). May−Jun. Fig. 39

Figure 39.

Carex striata var. brevis (from Britton and Brown 1913)

Cladium mariscoides (Muhl.) Torr.

Nomenclature: 

Basionym: Schoenus mariscoides Muhl.

Taxon concept: [= RAB, FNA, Weakley]

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Conservation status: 

SR–O; S3, G5.

Distribution: 

Lake Waccamaw (Abundant): Howell LAWA−16, 146 (NCSC!); LeBlond 3862 (NCU!); Wilbur 49778, 49789 (DUKE!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Jul−Sep. This taxon is the principal sedge component of the natural shoreline community of Lake Waccamaw. Fig. 40

Figure 40.

Cladium mariscoides (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-16 (NCSC)
bSpecimen: Howell LAWA-146 (NCSC)
cInflorescence
dInflorescence

Cyperus erythrorhizos Muhl.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Horseshoe Lake: Buell 2263 (DUKE!); Rothfels, Burge, Duke Natural History Society 2403 (DUKE!)

Notes: 

Annual herbs. floating bogs; saturated, acidic, peaty soil (FB). Jul−Sep. Fig. 41

Figure 41.

Cyperus erythrorhizos (from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

Cyperus odoratus var. odoratus

Nomenclature: 

Taxon concept: [= C. odoratus L. – RAB, GW; < C. odoratus L. – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Rare): Howell BATR−63 (NCSC!)

Notes: 

Annual or short-lived perennial herbs. Eulittoral zone; typically on moist sandy beaches at or just below the maximum annual high water mark. Jul−Sep. Fig. 42

Figure 42.

Cyperus odoratus var. odoratus (digital photograph taken by Nathan Howell)

aSpecimen: Howell BATR-63 (NCSC)
bInflorescence

Cyperus polystachyos Rottb.

Nomenclature: 

Taxon concept: [> C. polystachyos Rottb. var. texensis (Torr.) Fernald – RAB; < C. polystachyos – GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Jones Lake (Rare): Howell JOLA−43 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone; usually in sandy moist soil just below the maximum annual high water mark (NLSM−T). Jul−Oct. Fig. 43

Figure 43.

Cyperus polystachyos (illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

aSpecimen: Howell JOLA-43 (NCSC)
bIllustration

Dulichium arundinaceum var. arundinaceum

Nomenclature: 

Basionym: Cyperus arundinaceus L.

Taxon concept: [< D. arundinaceum (L.) Britton – RAB, GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Horseshoe Lake (Occasional): Beal 4345 (NCSC!); Buell s.n. (DUKE!); Howell HOLA−32 (NCSC!)

Lake Waccamaw (Occasional): Howell LAWA−26, 77 (NCSC!)

Little Singletary Lake (Infrequent): Howell LISI−41 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone; calm, quiet waters along shorelines or on floating bogs (NLSS−C, NLSS−LW, CPSI−CG, FB). Jul–Oct. Fig. 44

Figure 44.

Dulichium arundinaceum (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-26 (NCSC)
bSpecimen: Howell LAWA-77 (NCSC)
cHabit
dLeaves (3-ranked)

Eleocharis baldwinii Chapm.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Infrequent): Howell BATR−36, 40 (NCSC!)

Horseshoe Lake: • (The first author has observed Eleocharis baldwinii/vivipara around the peripheries of floating bogs and along saturated peaty shores, but voucher specimens were not collected. These two species are unidentifiable from a distance and the use of a hand lens is needed to distinguish one from the other.)

Little Singletary Lake (Rare): Howell LISI−43 (NCSC!)

Notes: 

Annual (?) herbs. Eulittoral zone and infralittoral zones; typically submersed in shallow water or on sarurated organic to sandy soils above current lake levels (NLSS−C, NLSM−T). Jul−Sep. Fig. 45

Figure 45.

Eleocharis baldwinii (digital photographs taken by Nathan Howell)

aSpecimen: Howell BATR-40 (NCSC)
bSpecimen: Howell LISI-43 (NCSC)
cHabit
dHabit

Eleocharis equisetoides (Elliott) Torr.

Nomenclature: 

Basionym: Scirpus equisetoides Elliott

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Conservation status: 

W1; S3, G4.

Distribution: 

Lake Waccamaw (Infrequent): Howell LAWA−67, 155 (NCSC!)

Little Singletary Lake (Rare): Howell LISI−38 (NCSC!)

Notes: 

Perennial herbs. Eulittoral and infralittoral zones; calm, quiet waters along shorelines (NLSS−C, NLSS−LW). Jun−Sep. Fig. 46

Figure 46.

Eleocharis equisetoides (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-67 (NCSC)
bSpecimen: Howell LAWA-155 (NCSC)
cHabit
dInflorescence detail

Eleocharis olivacea var. olivacea

Nomenclature: 

Taxon concept: [< E. flavescens (Poir.) Urb. – RAB; < E. olivacea Torr. – GW; < E. flavescens (Poir.) Urb. var. olivacea (Torr.) Gleason – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Rare): Howell LAWA−78 (NCSC!); LeBlond 3987 (NCU!)

Notes: 

Perennial herbs. Eulittoral and infralittoral zones; calm, quiet waters along shorelines (NLSS−LW). Jun−Sep. Fig. 47

Figure 47.

Eleocharis olivacea var. olivacea (digital photograph taken by Nathan Howell)

aSpecimen: Howell LAWA-78 (NCSC)
bHabit

Eleocharis vivipara Link

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Conservation status: 

State E; S1, G5.

Distribution: 

Horseshoe Lake: • (The first author has observed Eleocharis baldwinii/vivipara around the peripheries of floating bogs and along saturated peaty shores, but voucher specimens were not collected. These two species are unidentifiable from a distance and the use of a hand lens is needed to distinguish one from the other.)

Little Singletary Lake (Rare): Howell LISI−53 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (calm, quiet waters) or boggy, saturated, organic soils at or just below the maximum annual high water mark (NLSS−C, NLSM−T). Jul−Sep. Fig. 48

Figure 48.

Eleocharis vivipara (digital photograph taken by Nathan Howell)

aSpecimen: Howell LISI-53 (NCSC)
bHabit

Fimbristylis autumnalis (L.) Roem. & Schult.

Nomenclature: 

Basionym: Scirpus autumnalis L.

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Radford 677 (NCU!)

Notes: 

Annual herbs. Eulittoral zone; wet, sandy, disturbed areas (NLSS−LW). Jun−Oct. Fig. 49

Figure 49.

Fimbristylis autumnalis (from Britton and Brown 1913)

Fuirena pumila (Torr.) Spreng.

Nomenclature: 

Basionym: Fuirena squarrosa Michx. var. pumila Torr.

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Rare): Howell BATR−62 (NCSC!); Wilbur 57396 (DUKE!)

Notes: 

Annual herbs. Eulittoral zone; typically in moist sandy soil at high water mark. Jul−Oct. Fig. 50

Figure 50.

Fuirena pumila (digital photograph taken by Nathan Howell)

aSpecimen: Howell BATR-62 (NCSC)
bInflorescence detail

Rhynchospora alba (L.) Vahl

Nomenclature: 

Basionym: Schoenus albus L.

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Conservation status: 

SR−P; S2, G5.

Distribution: 

Horseshoe Lake (Occasional): Howell HOLA−45 (NCSC!)

Notes: 

Perennial herbs. floating bogs of Horseshoe Lake (FB). Jul−Oct. Fig. 51

Figure 51.

Rhynchospora alba (digital photographs taken by Nathan Howell; illustration from Britton and Brown 1913)

aSpecimen: Howell HOLA-45 (NCSC)
bIllustration
cInflorescences
dInflorescences

Rhynchospora corniculata (Lam.) A. Gray

Nomenclature: 

Basionym: Schoenus corniculatus Lam.

Taxon concept: [= RAB, GW, FNA; < R. corniculata (L.) A. Gray var. corniculata − Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Occasional): Howell LAWA−135, 163 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Jul−Sep. Fig. 52

Figure 52.

Rhynchospora corniculata (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-135 (NCSC)
bSpecimen: Howell LAWA-163 (NCSC)
cInflorescence
dInflorescence detail

Rhynchospora distans (Michx.) Vahl

Nomenclature: 

Basionym: Schoenus distans Michx.

Taxon concept: [< R. fascicularis (Michx.) Vahl – RAB, GW, FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Bakers Lake (Rare): Howell BALA−2 (NCSC!)

Lake Waccamaw: Wilbur 49814 (DUKE!)

Little Singletary Lake (Rare): Howell LISI−33 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone; typically at the high water mark in moist sandy soil (NLSS−C). Jun−Sep. Fig. 53

Figure 53.

Rhynchospora distans (Howell BALA-2, NCSC)

Rhynchospora elliottii A. Dietr.

Nomenclature: 

Taxon concept: [= R. schoenoides (Elliott) Wood – RAB; = GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: ♦

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Jul−Sep.

Rhynchospora inexpansa (Michx.) Vahl

Nomenclature: 

Basionym: Schoenus inexpansus Michx.

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Jones Lake: Beal 799 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS–C). Jul–Sep. Fig. 54

Figure 54.

Rhynchospora inexpansa (illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

Rhynchospora inundata (Oakes) Fernald

Nomenclature: 

Basionym: Ceratoschoenus macrostachyus (Torr. ex A. Gray) A. Gray var. inundatus Oakes

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Conservation status: 

W1; S3, G4?

Distribution: 

Horseshoe Lake (Infrequent): Howell HOLA−53 (NCSC!); Grant s.n. (NCU!); Rothfels, Burge, Duke Nat. Hist. Soc. 2401 (DUKE!)

Notes: 

Perennial herbs. Eulittoral zone of shorelines and on floating bogs (CPSI−CG, FB). Jul−Sep. Fig. 55

Figure 55.

Rhynchospora inundata (digital photograph taken by Nathan Howell)

aSpecimen: Howell HOLA-53 (NCSC)
bInflorescence

Rhynchospora latifolia (Baldwin) W.W. Thomas

Nomenclature: 

Basionym: Dichromena latifolia Baldwin

Taxon concept: [= Dichromena latifolia Baldwin ex Elliott – RAB, GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Radford 723 (NCU!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). May−Sep. Fig. 56

Figure 56.

Rhynchospora latifolia (illustration from Britton and Brown 1913)

Rhynchospora macrostachya Torr. ex A. Gray

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Infrequent): Howell LAWA−130 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Jul−Sep. Fig. 57

Figure 57.

Rhynchospora macrostachya (digital photograph taken by Nathan Howell)

aSpecimen: Howell LAWA-130 (NCSC)
bAchene detail

Rhynchospora nitens (Vahl) A. Gray

Nomenclature: 

Basionym: Scirpus nitens Vahl

Taxon concept: [= Psilocarya nitens (Vahl) Alph. Wood – RAB, GW; = FNA, Weakley]

Material    Download as CSV 
Conservation status: 

W1; S3, G4?

Distribution: 

Lake Waccamaw: Wilbur 49781 (DUKE!)

Notes: 

Annual herbs. Eulittoral zone (NLSS−LW). Jul−Aug. Fig. 58

Figure 58.

Rhynchospora nitens (illustration from Britton and Brown 1913)

Scirpus cyperinus (L.) Kunth

Nomenclature: 

Basionym: Eriophorum cyperinum L.

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Occasional): Howell BATR−58 (NCSC!)

Jones Lake (Occasional): Howell JOLA−4, 45 (NCSC!)

Lake Waccamaw (Occasional): Howell LAWA−166 (NCSC!)

Little Singletary Lake (Occasional): •

Notes: 

Perennial herbs. Eulittoral zone (NLSS−C, NLSS−LW, NLSM−T). Jul−Sep. Fig. 59

Figure 59.

Scirpus cyperinus (digital photographs taken by Nathan Howell)

aSpecimen: Howell BATR-58 (NCSC)
bSpecimen: Howell LAWA-166 (NCSC)
cInflorescence
dInflorescence detail

Eriocaulaceae

Eriocaulon aquaticum (Hill) Druce

Nomenclature: 

Basionym: Cespa aquatica Hill

Taxon concept: [> E. pellucidum Michx. – RAB; = E. septangulare – GW; = FNA, Weakley]

Material    Download as CSV 
Conservation status: 

SC−V; S2, G5.

Distribution: 

Lake Waccamaw (Abundant): Howell LAWA−5, 52 (NCSC!); Lynch 185 (NCSC!); Wilbur 49802 (DUKE!)

Notes: 

Perennial herbs. Eulittoral and infralittoral zones (NLSS−LW, NLSM−LWP). Jul−Oct. A dominant species in the littoral zone of Lake Waccamaw. Fig. 60

Figure 60.

Eriocaulon aquaticum (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-5 (NCSC)
bSpecimen: Howell LAWA-52 (NCSC)
cHabit
dLeaves
eInflorescence
fInflorescence

Haemodoraceae

Lachnanthes caroliniana (Lam.) Dandy

Nomenclature: 

Basionym: Dilatris caroliniana Lam.

Taxon concept: [= RAB, GW, FNA, Weakley]]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Infrequent): Howell BATR−50, 51 (NCSC!)

Horseshoe Lake (Infrequent): Howell HOLA−51 (NCSC!)

Lake Waccamaw (Occasional): Howell LAWA−107 (NCSC!)

Little Singletary Lake (Infrequent): Howell LISI−25, 51 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone; typically in saturated soils at or below the maximum annual high water mark (NLSS–C, NLSS–LW, NLSM–T, CPSI–CG). Jun–early Sep; Sep–Nov. Fig. 61

Figure 61.

Lachnanthes caroliniana (digital photographs taken by Nathan Howell):

aSpecimen: Howell BATR-50 (NCSC)
bSpecimen: Howell HOLA-51 (NCSC)
cRoots
dInflorescence

Hydrocharitaceae

Hydrilla verticillata (L. f.) Royle

Nomenclature: 

Basionym: Serpicula verticillata L.f.

Taxon concept: [= GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: ¤

Notes: 

Perennial herbs. Infralittoral zone (NLSS−LW, NLSM−LWP). Jun−Aug. This exotic, invasive taxon is native to warm climates of the Old World. Hydrilla verticillata was introduced to Florida in 1950 as an ornamental and has since become a terrible aquatic invasive throughout the Southeast. Where introduced, H. verticillata chokes out native submersed aquatic vegetation (e.g., Ceratophyllum, Myriophyllum, Najas, Potomogeton, Vallisneria), negatively impacts recreational activities and alters natural hydrology and water chemistry (Ramey and Peichel 2001). Fig. 62

Figure 62.

Hydrilla verticillata (from Center for Aquatic and Invasive Plants, University of Florida, IFAS 2015)

Najas guadalupensis var. guadalupensis

Nomenclature: 

Basionym: Caulinia guadalupensis Spreng.

Taxon concept: [< N. guadalupensis (Spreng.) Magnus – RAB, GW; = N. guadalupensis ssp. guadalupensis – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Blomquist & Schuster 16190 (DUKE!)

Notes: 

Annual herbs. Infralittoral zone (NLSS−LW, NLSM−LWP). Jul−Sep. Fig. 63

Figure 63.

Najas guadalupensis (from Center for Aquatic and Invasive Plants, University of Florida, IFAS 2015)

Hypoxidaceae

Hypoxis curtissii Rose

Nomenclature: 

Taxon concept: [= H. hirsuta (L.) Coville var. leptocarpa (Engelm. & A. Gray) Fernald – RAB; = H. leptocarpa (Engelm. & A. Gray) Small – GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Rare): Howell LAWA−60 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone; at the high water mark in moist to saturated soil (NLSS−LW). Mar−Jun; May−Jul. Fig. 64

Figure 64.

Hypoxis curtissii (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-60 (NCSC)
bHabit
cFlower
dFlower

Juncaceae

Juncus acuminatus Michx.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Rare): Howell BATR−15 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone. May−Aug. Fig. 65

Figure 65.

Juncus acuminatus (illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

aSpecimen: Howell BATR-15 (NCSC)
bIllustration

Juncus biflorus Elliott

Nomenclature: 

Taxon concept: [= RAB; < J. marginatus Rostk. – GW, FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Little Singletary Lake (Rare): Howell LISI−58 (NCSC!)

Singletary Lake: Beal 796 (NCSC!)

Notes: 

Perennial herbs. Juncture of the eulittoral and supralittoral zones; usually in wet soils at or just below the high water mark (NLSM−T, NLSS−C). Jun−Oct. Fig. 66

Figure 66.

Juncus biflorus (digital photograph taken by Nathan Howell)

aSpecimen: Howell LISI-58 (NCSC)
bInflorescence

Juncus canadensis J. Gay ex Laharpe

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Infrequent): Howell LAWA−32, 170 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Jul−Oct. Fig. 67

Figure 67.

Juncus canadensis (illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

aSpecimen: Howell LAWA-167 (NCSC)
bIllustration

Juncus coriaceus Mack.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Horseshoe Lake: Beal 828 (NCSC!)

Notes: 

Perennial herbs. Juncture of eulittoral and supralittoral zones (CPSI−CG). Jun−Sep. Fig. 68

Figure 68.

Juncus coriaceus (from Britton and Brown 1913)

Juncus effusus subsp. solutus (Fernald & Wiegand) Hämet-Ahti

Nomenclature: 

Taxon concept: [< J. effusus – RAB, GW, FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Occassional): Howell BATR−6 (NCSC!)

Little Singletary Lake (Occassional): Howell LISI−3 (NCSC!)

Horseshoe Lake (Occassiona): Howell HOLA−8 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−C, NLSM−T, CPSI−CG). Jun−Sep. Fig. 69

Figure 69.

Juncus effusus (digital photographs taken by Nathan Howell; illustration from Center for Aquatic and Invasive Plants, University of Florida, IFAS 2015)

aSpecimen: Howell HOLA-8 (NCSC)
bIllustration
cHabit
dInflorescence

Juncus pelocarpus E. Mey.

Nomenclature: 

Taxon concept: [> J. abortivus Chapm. – RAB, GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Frequent): Howell BATR−61 (NCSC!); Wilbur 57415 (DUKE!)

Horseshoe Lake: Wilbur 2264, 81465 (DUKE!)

Jones Lake (Occasional): Howell JOLA−17, 35 (NCSC!); Wilbur 57582 (DUKE!)

Lake Waccamaw (Frequent): Howell LAWA−3 (NCSC!); Wilbur s. n., 84188 (DUKE!)

Singletary Lake (Occasional): Howell SILA−31 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−C, NLSS−LW, NLSM−T, CPSI−CG). Jul−Oct. Fig. 70

Figure 70.

Juncus pelocarpus (digital photographs taken by Nathan Howell)

aSpecimen: Howell BATR-61 (NCSC)
bSpecimen: Howell LAWA-3 (NCSC)
cFlower
dInflorescence

Juncus repens Michx.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Occasional): Howell BATR−8 (NCSC!); Wilbur 57395 (DUKE!)

Horseshoe Lake (Occasional): Beal 4348 (NCSC!); Howell HOLA−14 (NCSC!); Wilbur & Menchi Ho 83792 (DUKE!)

Lake Waccamaw (Occasional): Howell LAWA−30, 31 (NCSC!)

Little Singletary Lake (Occasional): Howell LISI−19, 44 (NCSC!)

Singletary Lake (Occasional): Howell SILA−1, 32 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−C, NLSS−LW, NLSM−T, CPSI−CG). Jun−Oct. Fig. 71

Figure 71.

Juncus repens (digital photographs taken by Nathan Howell; illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

aSpecimen: Howell SILA-32 (NCSC)
bIllustration
cHabit
dHabit

Juncus scirpoides var. scirpoides

Nomenclature: 

Taxon concept: [< J. scirpoides Lam. – RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Infrequent): Howell BATR−27, 66 (NCSC!)

Little Singletary Lake (Infrequent): Howell LISI−55 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−C, NLSM−T). Jun−Sep. Fig. 72

Figure 72.

Juncus scirpoides (digital photographs taken by Nathan Howell)

aSpecimen: Howell BATR-27 (NCSC)
bSpecimen: Howell BATR-66 (NCSC)
cInflorescence
dInflorescence

Mayacaceae

Mayaca fluviatilis Aubl.

Nomenclature: 

Taxon concept: [> M. aubletii Michx. – RAB; > M. fluviatilis Aubl. – RAB; = GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: ¤

Notes: 

Perennial herbs. Eulittoral and infralittoral zones (NLSS−LW). May−Jul. Fig. 73

Figure 73.

Mayaca fluviatilis (from Center for Aquatic and Invasive Plants, University of Florida, IFAS 2015)

Orchidaceae

Calopogon tuberosus var. tuberosus

Nomenclature: 

Basionym: Limodorum tuberosum L.

Taxon concept: [< C. pulchellus R. Brown − RAB; < C. tuberosus (L.) Britton, Sterns, & Poggenb. – GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Horseshoe Lake (Occasional): Howell HOLA−24, 39 (NCSC!)

Notes: 

Perennial herbs. floating bogs (CPSI–CG, FB). Apr–Jul. Fig. 74

Figure 74.

Calopogon tuberosus (digital photograph taken by Nathan Howell)

aSpecimen: Howell HOLA-24 (NCSC)
bFlower

Epidendrum magnoliae Muhl.

Nomenclature: 

Taxon concept: [< E. conopseum R. Br. – RAB; = FNA, Weakley]

Material    Download as CSV 
Conservation status: 

T; S1S2, G4.

Distribution: 

Lake Waccamaw: Correll & Blomquist 4900 (DUKE!)

Notes: 

Perennial, epiphytic herbs. Eulittoral zone; typically on limbs and trunks of Taxodium ascendens, Taxodium distichum, Nyssa aquatica, Nyssa biflora, Liquidambar styraciflua, and possibly other bottomland tree species in the shoreline of Lake Waccamaw (NLSS– LW, NLSS–C). Jul–Oct. This species usually co-occurs with Pleopeltis polypodiodes. The first author observed a vegetative specimen on the edge of Big Creek ca. 50–70 meters from the shoreline of Lake Waccamaw. The specimen was on a large Nyssa aquatica limb, ca. 25–30 meters above the water, and was co- occuring with Pleopeltis polypodioides.

Habenaria repens Nutt.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Conservation status: 

W1; S2, G5.

Distribution: 

Lake Waccamaw: ►

Notes: 

Perennial herbs. Eulittoral zone (NLSS–LW). Apr–Nov.

Pogonia ophioglossoides (L.) Ker Gawl.

Nomenclature: 

Basionym: Arethusa ophioglossoides L.

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Horseshoe Lake (Occasional): Howell HOLA−30 (NCSC!)

Notes: 

Perennial herbs. floating bogs (CPSI–CG, FB). Mar–Jun. Fig. 75

Figure 75.

Pogonia ophioglossoides (digital photograph taken by Nathan Howell)

aSpecimen: Howell HOLA-30 (NCSC)
bFlower

Spiranthes laciniata (Small) Ames

Nomenclature: 

Basionym: Gyrostachys laciniata Small

Taxon concept: [= RAB; < S. × laciniata – GW; = FNA, Weakley]

Material    Download as CSV 
Conservation status: 

SC−V; S2, G4,G5.

Distribution: 

Lake Waccamaw (Occasional): Howell LAWA−105, 106, 116 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS–LW). May–Aug. Fig. 76

Figure 76.

Spiranthes laciniata (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-105 (NCSC)
bLeaf
cInflorescence
dInflorescence (detail)

Poaceae

Agrostis hyemalis (Walter) Britton, Sterns & Poggenb.

Nomenclature: 

Basionym: Cornucopiae hyemalis Walter

Taxon concept: [< A. hyemalis (Walter) Britton, Sterns & Poggenb. – RAB; = A. hiemalis (Walter) Britton, Sterns & Poggenb. – GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Little Singletary Lake (Rare): Howell LISI−37 (NCSC!)

Notes: 

Perennial herbs. Juncture of supralittoral and eulittoral zones; typically in moist sandy soils (NLSM−T). Mar−Jul. Fig. 77

Figure 77.

Agrostis hyemalis (digital photographs taken by Nathan Howell; illustration from Hitchcock and Chase 1951)

aSpecimen: Howell LISI-37 (NCSC)
bIllustration
cBase of inflorescence
dInflorescence, including spikelets

Andropogon glaucopsis Steud.

Nomenclature: 

Taxon concept: [< A. virginicus L. – RAB; = GW; = A. glomeratus var. glaucopsis (Elliott) C. Mohr − FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Horseshoe Lake: Buell s.n. (DUKE!, NCSC!)

Jones Lake (Infrequent): Howell JOLA–16 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (CPSI−CG, FB). Sep−Oct. Fig. 78

Figure 78.

Andropogon glaucopsis (Howell JOLA-16, NCSC)

Andropogon virginicus var. virginicus

Nomenclature: 

Taxon concept: [< A. virginicus L. – RAB; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: ♦

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Sep−Oct. Fig. 79

Figure 79.

Andropogon virginicus (illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

Arundinaria tecta (Walter) Muhl.

Nomenclature: 

Basionym: Arundo tecta Walter

Taxon concept: [< A. gigantea (Walter) Muhl. – RAB, GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Bennedict 4350 (DUKE!)

Notes: 

Arborescent herbs. Eulittoral zone; at or just below the mean annual high water mark (NLSS−LW). Apr−Jul. The first author has not encountered this taxon in the field, but a single voucher specimen (see above) places it within the immediate vicinity. Fig. 80

Figure 80.

Arundinaria tecta (from Hitchcock and Chase 1951)

Coleataenia longifolia var. longifolia

Nomenclature: 

Basionym: Panicum longifolium Torr.

Taxon concept: [= Panicum longifolium Torr. var. longifolium – RAB; < Panicum longifolium Torr. – GW; = Panicum rigidulum Bosc ex Nees ssp. pubescens (Vasey) Freckmann & Lelong − FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Infrequent): Howell BATR – 68 (NCSC!)

Lake Waccamaw (Occasional): Howell LAWA −145, 147, 164, 168 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Jul−Oct. Fig. 81

Figure 81.

Coleataenia longifolia (illustration from Britton and Brown 1913)

Coleataenia tenera (Beyr. ex Trin.) Soreng

Nomenclature: 

Basionym: Panicum tenerum Bey. ex Trin.

Taxon concept: [= Panicum tenerum Bey. ex Trin. – RAB, GW, FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: ►

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Jun−Sep.

Dichanthelium boreale (Nash) Freckmann

Nomenclature: 

Basionym: Panicum boreale Nash

Taxon concept: [> Panicum bicknellii Nash– RAB; > D. boreale (Nash) Freckmann – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Blomquist 957 (DUKE!)

Notes: 

Perennial herbs. Eulittoral zone (NLS−LW). Apr−Sep. Fig. 82

Figure 82.

Dichanthelium boreale (illustration from Britton and Brown 1913)

Dichanthelium dichotomum var. roanokense (Ashe) LeBlond

Nomenclature: 

Basionym: Panicum roanokense Nash

Taxon concept: [< D. dichotomum (L.) Gould – RAB, GW; < D. dichotomum (L.) Gould) ssp. roanokense (Ashe) Freckmann & Lelong – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Ashe s.n. (NCU!)

Notes: 

Perennial herbs. Eulittoral zone; moist to peaty lakeshores (NLSS−LW). May−Sep.

Dichanthelium erectifolium (Nash) Gould & C.A. Clark

Nomenclature: 

Basionym: Panicum erectifolium Nash

Taxon concept: [= Panicum erectifolium Nash – RAB, GW; = FNA, Weakley]

Material    Download as CSV 
Conservation status: 

W2; S2, G4.

Distribution: 

Lake Waccamaw (Infrequent): Howell LAWA−111 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone; moist sandy to peaty shores (NLSS−LW). May−Aug. Fig. 83

Figure 83.

Dichanthelium erectifolium (digital photographs taken by Nathan Howell)

aSpecimen: Howell LAWA-111 (NCSC)
bSpecimen: Howell LAWA-127 (NCSC)
cCulm and leaf
dInflorescence

Dichanthelium species 3 = lancearium

Nomenclature: 

Taxon concept: [= Panicum lancearium Trinius – RAB; < D. portoricense (Desv. ex Ham.) B.F. Hansen & Wunderlin ssp. patulum (Scribner & Merrill) Freckmann & Lelong – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Blomquist & Correll 9383 (NCU!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). May−Sep.

Dichanthelium mattamuskeetense (Ashe) Mohlenbr.

Nomenclature: 

Basionym: Panicum mattamuskeetense Ashe

Taxon concept: [< Panicum dichotomum L. – RAB, GW; < D. dichotomum (L.) Gould ssp. mattamuskeetense (Ashe) Freckmann & Lelong – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Blomquist & Correll 9385 (DUKE!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). May−Oct. Fig. 84

Figure 84.

Dichanthelium mattamuskeetense (illustration from Britton and Brown 1913)

Dichanthelium portoricense (Desv. ex Ham.) B.F. Hansen & Wunderlin

Nomenclature: 

Basionym: Panicum portoricense Desv. ex Ham.

Taxon concept: [= Panicum portoricense Desv. ex Ham. – RAB; = D. portoricense (Desv. ex Ham.) B.F. Hansen & Wunderlin ssp. portoricense – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Infrequent): Howell BATR – 52 (NCSC!)

Lake Waccamaw: Blomquist & Correll 9383 (NCU!)

Notes: 

Perennial herbs. Eulittoral zone; moist sandy to peaty shores (NLSS−LW). May−Sep. Fig. 85

Figure 85.

Dichanthelium portoricense (Howell BATR-52, NCSC)

Eragrostis elliottii S. Watson

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Rare): Howell BATR−67 (NCSC!)

Lake Waccamaw: ►

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Sep−Oct. Fig. 86

Figure 86.

Eragrostis elliottii (Howell BATR-67, NCSC)

Eragrostis refracta (Muhl. ex Elliott) Scribn.

Nomenclature: 

Basionym: Poa refracta Muhl. ex Elliott

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: ►

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Jul−Oct.

Luziola fluitans var. fluitans

Nomenclature: 

Basionym: Zizania fluitans Michx.

Taxon concept: [= Hydrochloa carolinensis P. Beauv. – RAB, GW; = FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw (Occasional): Bolser MEH107 (NCU!); Howell LAWA−51 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Aug−Oct. Fig. 87

Figure 87.

Luziola fluitans (digital photographs taken by Nathan Howell; illustration from Hitchcock and Chase 1951)

aSpecimen: Howell LAWA-51 (NCSC)
bIllustration
cHabit
dLeaves

Panicum hemitomon Schult.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Abundant): Howell BATR−18 (NCSC!)

Horseshoe Lake (Infrequent): Howell HOLA−23 (NCSC!)

Lake Waccamaw (Abundant): Blomquist 1399 (DUKE!); Blomquist & Correll 9379 (DUKE!, NCU!); Howell LAWA−79 (NCSC!)

Little Singletary Lake (Infrequent): Howell LISI−35 (NCSC!)

Salters Lake (Occasional): Howell SALA−14 (NCSC!)

Singletary Lake (Occasional): Blomquist 1400 (DUKE!); Howell SILA−17 (NCSC!); Wilbur 60947 (DUKE!)

Notes: 

Perennial herbs. Eulittoral and infralitoral zones (NLSS−C, NLSS−LW, NLSM−T, CPSI−CG). Jun−Jul. Fig. 88

Figure 88.

Panicum hemitomon (digital photographs taken by Nathan Howell; illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

aSpecimen: Howell BATR-18 (NCSC)
bIllustration
cHabit
dInflorescences

Panicum verrucosum Muhl.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Infrequent): Howell BATR−53 (NCSC!)

Jones Lake (Rare): Howell JOLA−40, 41 (NCSC!)

Little Singletary Lake (Rare): Howell LISI−50 (NCSC!)

Notes: 

Annual herbs. Eulittoral zone (NLSM−T, NLSS−C). Aug−Oct. Fig. 89

Figure 89.

Panicum verrucosum (illustration from Britton and Brown 1913)

aSpecimen: Howell BATR-53 (NCSC)
bIllustration

Panicum virgatum var. virgatum

Nomenclature: 

Taxon concept: [< P. virgatum – RAB, GW, FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake: Wilbur 57420 (DUKE!)

Notes: 

Perennial herbs. Eulittoral zone. Jun−Oct. Fig. 90

Figure 90.

Panicum virgatum (illustration from Hitchcock and Chase 1951)

Saccharum giganteum (Walter) Pers.

Nomenclature: 

Basionym: Anthoxanthum giganteum Walter

Taxon concept: [= Erianthus giganteus (Walter) P. Beauv. – RAB, GW; = FNA. Weakley]

Material    Download as CSV 
Distribution: 

Jones Lake (Rare): Howell JOLA−37 (NCSC!)

Lake Waccamaw (Occasional): Howell LAWA−7, 160 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−C, NLSS−LW). Sep−Oct. Fig. 91

Figure 91.

Saccharum giganteum (digital photographs taken by Nathan Howell; illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

aSpecimen: Howell LAWA-7 (NCSC)
blllustration
cCulm and leaf blade
dInflorescence

Sacciolepis striata (L.) Nash

Nomenclature: 

Basionym: Holcus striatus L.

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Occasional): Howell BATR−43, 54, 55 (NCSC!); Wilbur 48657, 57394 (DUKE!)

Horseshoe Lake: Rothfels, Burge, Duke Natural History Society 2398 (DUKE!)

Lake Waccamaw (Occasional): Howell LAWA−131 (NCSC!)

Singletary Lake (Occasional): Beal 3225 (NCSC!); Frey s.n. (NCU!); Howell SILA−38 (NCSC!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−C, NLSS−LW, NLSM−T, CPSI−CG). Jul−Oct. Fig. 92

Figure 92.

Sacciolepis striata (digital photographs taken by Nathan Howell; illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

aSpecimen: Howell BATR-55 (NCSC)
bIllustration
cLeaf sheath
dInflorescence

Sphenopholis obtusata (Michx.) Scribn.

Nomenclature: 

Basionym: Aria obtusata Michx.

Taxon concept: [= RAB, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: Blomquist 1492 (DUKE!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). Apr−May. Fig. 93

Figure 93.

Sphenopholis obtusata (illustration from Hitchcock and Chase 1951)

Pontederiaceae

Pontederia cordata var. cordata

Nomenclature: 

Taxon concept: [< P. cordata − RAB; = GW; < P. cordata – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Rare): •Lake Waccamaw (Occasional): Howell LAWA−15, 50, 159 (NCSC!); Matthews s.n. (NCU!); Wilbur 59382 (DUKE!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). May−Oct. Fig. 94

Figure 94.

Pontederia cordata var. cordata (digital photographs taken by Nathan Howell; illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

aSpecimen: Howell LAWA-159 (NCSC)
bIllustration
cLeaves
dInflorescence

Pontederia cordata var. lancifolia (Muhl.) Torr.

Nomenclature: 

Taxon concept: [< P. cordata – RAB; = GW; < P. cordata – FNA; = Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: ♦

Notes: 

Perennial herbs. Eulittoral zone (NLSS−LW). May−Oct.

Potamogetonaceae

Potamogeton pulcher Tuck.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: ►

Notes: 

Perennial herbs. Eulittoral and infralittoral zones (NLSS−LW, NLSM−LWP). Jun−Sep. Fig. 95

Figure 95.

Potamogeton pulcher (illustration from Britton and Brown 1913)

Potamogeton pusillus L.

Nomenclature: 

Taxon concept: [< P. berchtoldii Fieber – RAB; = GW; > P. pusillus L. ssp. pusillus – FNA; > P. pusillus L. var. pusillus − Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: ¤

Notes: 

Annual herbs. Eulittoral and infralittoral zones (NLSS−LW, NLSM−LWP). May−Sep.

Smilacaceae

Smilax glauca Walter

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bay Tree Lake (Rare): Howell BATR−29 (NCSC!)

Notes: 

Perennial vines. Juncture of eulittoral and supralittoral zones. Late Apr−Early Jun; Sep−Nov and persisting. Fig. 96

Figure 96.

Smilax glauca (digital photographs taken by Alexander Krings)

aSpecimen: Howell BATR-29 (NCSC)
bLeaf (abaxial surface)
cLeaf showing contrast between abaxial surface (left) and adaxial surface (right)
dFruits

Smilax laurifolia L.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Bakers Lake (Frequent): Howell BALA−13 (NCSC!)

Bay Tree Lake (Occasional): Howell BATR−44 (NCSC!)

Horseshoe Lake (Occasional): Howell HOLA−3 (NCSC!)

Jones Lake (Frequent): Howell JOLA−7 (NCSC!)

Lake Waccamaw (Infrequent): Howell LAWA−34 (NCSC!)

Salters Lake (Frequent): Howell SALA−4, 15 (NCSC!)

Singletary Lake (Frequent): Howell SILA−9 (NCSC!)

Notes: 

Perennial vines. Eulittoral zone; typically at the maximum annual high water mark in saturated organic to sandy soils (NLSS−C, NLSS−LW, NLSM−T, CPSI−CG). Jul−Aug; Sep−Oct and persisting. Fig. 97

Figure 97.

Smilax laurifolia (digital photographs taken by Nathan Howell; illustrations from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

aSpecimen: Howell SALA-15 (NCSC)
bIllustration
cHabit
dInfructescence

Smilax rotundifolia L.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Lake Waccamaw: ♦

Notes: 

Perennial vines. Juncture of eulittoral and supralittoral (NLSS−LW). Apr−May; Sep−Oct and persisting. Fig. 98

Figure 98.

Smilax rotundifolia (digital photographs taken by Alexander Krings; illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

aIllustration
bLeaf (adaxial surface)
cStem (with prickles), petiole, and withered stipular tendrils (at base of petiole)
dDenticulations at base of leaf blade (arrowed)

Smilax walteri Pursh

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Horseshoe Lake (Occasional): Howell HOLA−13, 21, 25 (NCSC!)

Lake Waccamaw (Occasional): Howell LAWA−29, 55, 162 (NCSC!)

Notes: 

Perennial vines. Eulittoral zone (NLSS−LW, CPSI−CG). Late Apr−May; Sep−Nov and persisting. Fig. 99

Figure 99.

Smilax walteri (digital photographs taken by Nathan Howell [leaf and flower] and Alexander Krings [fruits])

aSpecimen: Howell LAWA-55 (NCSC)
bLeaf (adaxial surface)
cFlower
dFruits

Xyridaceae

Xyris fimbriata Elliott

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Horseshoe Lake: Rothfels, Burge, Duke Natural History Society 2400, 2404 (DUKE!)

Lake Waccamaw: ►

Singletary Lake: Frey s.n. (NCU!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−C, NLSM−T, NLSS−LW, CPSI–CG). Sep−Oct. Fig. 100

Figure 100.

Xyris fimbriata (illustration from United States Department of Agriculture, Natural Resource Conservation Service PLANTS Database / United States Department of Agriculture, Natural Resource Conservation Service 2015)

Xyris iridifolia Chapm.

Nomenclature: 

Taxon concept: [= RAB, GW; < X. laxifolia Mart. var. iridifolia (Chapm.) Kral – FNA; = Weakley]

Material    Download as CSV 
Conservation status: 

W7; S2, G4G5T4T.

Distribution: 

Salters Lake: ♦

Notes: 

Perennial herbs. Eulittoral zone (NLSS–C). Jul–Sep.

Xyris jupicai Rch.

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Distribution: 

Little Singletary Lake (Infrequent): Howell LISI−46 (NCSC!)

Notes: 

Annual, rarely biennial, herbs. Eulittoral zone (NLSS−C, NLSM−T). Jul−Sep. Fig. 101

Figure 101.

Xyris jupicai (illustration from Britton and Brown 1913)

aSpecimen: Howell LISI-46 (NCSC)
bIllustration

Xyris smalliana Nash

Nomenclature: 

Taxon concept: [= RAB, GW, FNA, Weakley]

Material    Download as CSV 
Conservation status: 

W1; S3, G5.

Distribution: 

Horseshoe Lake: R.L Wilbur 81092 (DUKE!)

Jones Lake (Occasional): Howell JOLA−42 (NCSC!)

Lake Waccamaw (Abundant): Howell LAWA−114, 125, 142, 144 (NCSC!); LeBlond 3996 (NCU)

Salters Lake: Beckman & Linnenburger 24 (DUKE!); Grant s.n. (NCU)

Singletary Lake (Occasional): Howell SILA−29 (NCSC!); Rothfels & O’ Reilly, Shaw Lab s.n. (DUKE!)

Notes: 

Perennial herbs. Eulittoral zone (NLSS−C, NLSS−LW, NLSM–T, CPSI–CG). Jul−Aug. Fig. 102

Figure 102.

Xyris smalliana (digital photographs taken by Nathan Howell; illustration from Britton and Brown 1913)

aSpecimen: Howell LAWA-114 (NCSC)
bIllustration
cInflorescence
dFlowers

BASAL ANGIOSPERMS, MAGNOLIIDS, and EUDICOTYLEDONS

Families represented: 55 (BA: 2; M: 2; E: 51)