Lachnocaulon digynum - Koern.
Pineland Bogbutton
Other Common Names: pineland bogbutton
Taxonomic Status: Accepted
Related ITIS Name(s): Lachnocaulon digynum Koern. (TSN 39207)
Unique Identifier: ELEMENT_GLOBAL.2.139023
Element Code: PMERI02030
Informal Taxonomy: Plants, Vascular - Flowering Plants - Pipewort Family
 
Kingdom Phylum Class Order Family Genus
Plantae Anthophyta Monocotyledoneae Eriocaulales Eriocaulaceae Lachnocaulon
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Concept Reference
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Concept Reference: Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.
Concept Reference Code: B94KAR01HQUS
Name Used in Concept Reference: Lachnocaulon digynum
Conservation Status
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NatureServe Status

Global Status: G3G4
Global Status Last Reviewed: 22Mar2018
Global Status Last Changed: 22Mar2018
Ranking Methodology Used: Ranked by calculator
Rounded Global Status: G3 - Vulnerable
Reasons: Lachnocaulon digynum occurs on the Florida panhandle west to southern Mississippi and eastern Texas including Louisiana. There are over 130 occurrences but over half are on privately own lands where they are threatened by land conversion to pine plantations and urban development. Occurrences on private and public lands are threatened by fire suppression and pollution of waterways.
Nation: United States
National Status: N3N4

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Alabama (S2), Florida (S3), Louisiana (S3), Mississippi (S2S3), Texas (S1)

Other Statuses

NatureServe Global Conservation Status Factors

Range Extent Comments: This species occurs on the Florida panhandle to southern Mississippi and west to eastern Texas including Louisiana.

Area of Occupancy: 126-500 4-km2 grid cells
Area of Occupancy Comments:  

Number of Occurrences: 81 - 300
Number of Occurrences Comments: There are about 177 extant occurrences. There are ten in Alabama, sixty-nine in Florida, fifty-four in Louisiana, thirty-nine in Mississippi, and four in Texas.

Population Size Comments: Locally abundant at some sites.

Number of Occurrences with Good Viability/Integrity: Some (13-40)

Overall Threat Impact: High - medium
Overall Threat Impact Comments: Potential threats include destruction or alteration of habitats by mechanized site preparation and harvest associated with pine plantation agriculture, either directly within the site, or in the watershed of the site. The destruction of microtopographic features and increased drainage associated with mechanical site preparation would eventually eliminate Lachnocaulon digynum and all relatively habitat-specific members of the wet pine savanna and mesic pine flatwoods community. The entire range of the species is in a relatively heavily populated and expanding urban/suburban area. Therefore, habitat destruction for development is a possibility. Grazing is a threat primarily through the associated trampling which would occur. Runoff of toxic chemicals is a big threat, as the species often occurs in areas where water pools for long periods and concentrates such chemicals.The exclusion of periodic fire from its habitat is also a major threat to the species. The suppression of natural fires and the creation of firebreaks such as roads and ditches have resulted in much less frequent fire over much of the range. This results in increasing tall grasses and rapidly increasing woody growth, eventually shading out the species. Drainage efforts of all kinds are detrimental to the species, particularly since it tends to occupy the wetter areas of many of its habitats, therefore is more likely to be eliminated by a lowered water table.

Short-term Trend: Decline of 10-30%
Short-term Trend Comments: There are three occurrences where plants could not be found during the last survey. In addition, nine occurrences are historic.

Long-term Trend: Unknown

Other NatureServe Conservation Status Information

Distribution
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Global Range: This species occurs on the Florida panhandle to southern Mississippi and west to eastern Texas including Louisiana.

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map

U.S. & Canada State/Province Distribution
United States AL, FL, LA, MS, TX

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Baldwin (01003), Covington (01039), Escambia (01053), Mobile (01097)
FL Bay (12005), Calhoun (12013), Escambia (12033), Gulf (12045), Liberty (12077), Okaloosa (12091), Santa Rosa (12113), Walton (12131)
LA Beauregard (22011), Natchitoches (22069), Sabine (22085), St. Tammany (22103), Vernon (22115)
MS George (28039), Hancock (28045), Harrison (28047), Jackson (28059), Pearl River (28109), Perry (28111), Stone (28131)
TX Jasper (48241), Newton (48351)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
03 Lower Ochlockonee (03120003)+, Apalachicola (03130011)+, Chipola (03130012)+, St. Andrew-St. Joseph Bays (03140101)+, Choctawhatchee Bay (03140102)+, Yellow (03140103)+, Blackwater (03140104)+, Pensacola Bay (03140105)+, Perdido (03140106)+, Perdido Bay (03140107)+, Escambia (03140305)+, Mobile Bay (03160205)+*, Pascagoula (03170006)+, Black (03170007)+, Escatawpa (03170008)+, Mississippi Coastal (03170009)+, Lower Pearl. Mississippi (03180004)+
08 Upper Calcasieu (08080203)+, Whisky Chitto (08080204)+, Liberty Bayou-Tchefuncta (08090201)+
11 Saline Bayou (11140208)+
12 Toledo Bend Reservoir (12010004)+, Lower Sabine (12010005)+, Lower Angelina (12020005)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: Small, perennial, clump-forming wetland herb with leaves in a basal rosette, and a grayish head of tiny flowers on a short stalk. The tightly clustered leaves are about 1 cm long, sharp-pointed, and resemble a large moss. The 5 to 10 cm tall flowering stalks have a sheath longer than the leaves, and are terminated by a 2-3 mm wide, grayish or gray-brown globose head. Flowers are very small, with complex and difficult to distinguish parts. Generally flowering and fruiting throughout late summer and fall.
Technical Description: A clump-former, dense tufts of rosettes forming from slender ascending rhizomes which develop from axillary buds. Leaf linear-acute, 0.6-1.0 (2.0) cm long, bright yellowish green, evenly narrowing from a clasping base 1.0-2.5 mm broad, almost smooth or with a scattering of multicellular trichomes, these usually most abundant toward the margins. Sheath of the scape acute, or bifid, longer than the leaves, smooth or sparingly ciliate toward the orifice. Scape 5.0-10.0 cm long, slightly twisted, 3-ridged, smooth or with a distant scattering of filiform, multicellular trichomes. Heads grayish or dull gray-brown, globose, or hemispheric, 2.0-3.5 mm broad. The longer of the involucral bracts triangular, ca. 1.0 mm long, brownish, with translucent clavate trichomes on the backs distally. Receptacular bracts spatulate, 1.0-1.3 mm long, acute or obtusely angled, a rich brown, clavate-hairy on the backs apically. Receptacular surface densely hairy, but the hairs not so long as to obscure the female sepals. Male flower: sepals linear-spatulate, ca 1.0 mm long, curvate, a rich brown, sparingly calvate-hairy on the backs apically; receptacle hairy; androphore smooth, obpyramidal, about the length of the sepals, the apices oblique, anthers 3, yellowish, oblong, slightly exserted on filaments about as long as themselves. Female flower: sepals broadly spatulate to narrowly obovate, ca 1.0 mm long, keeled, and curvate, hence connivent over the ovulary, whitish-yellow, smooth or with a few marginal hairs distally; receptacle and short gynophore copiously pilose with pale translucent, multicellular, slightly calvate, trichomes; gynoecium 2-carpellate, 2-locular, 2-ovulate, the styles 2, bifid. Seeds ovoid to ellipsoidal, about 0.5 mm long, longitudinally striate, the connecting striae finer, almost obscure. (Godfrey and Wooten, 1979; Kral, 1966)
Diagnostic Characteristics: The family Eriocaulaceae (Hat-pins, Pipeworts, Bog buttons etc.) is easily distinguished from all other plants by even the casual observer. All members of the family have narrow leaves in basal rosettes, and flowering stalks terminating in single, button-like heads. Within the family, however, field identification immediately becomes much more difficult. There are numerous technical characters of the florets which provide the definitive separation of the genera and species, but the small size and tight clustering of floral parts make these less useful in the field. One useful field character in distinguishing Eriocaulaceae is the color of the head. Many species have conspicuously pale gray or white heads, due to the presence of milky-white, opaque trichomes. Another useful character is the pubescence of the scape, or lack of it. In the genus Eriocaulon, the lacunar tissue (air spaces) near the base of the leaves is evident, and the roots are septate and relatively unbranched. Lachnocaulon species have no evident lacunar tissue, and have a non-septate, more finely branched root system. Leaf arrangement and size can also be useful; Syngonanthus has reflexed leaves, in contrast to the ascending or spreading leaves of other family members; although leaf size in the family is extremely variable, Lachnocaulon digynum has smaller mature leaves than any other species, though there is some overlap in leaf size with the most diminutive individuals of some other species. In summary, a member of the Eriocaulaceae with the following combination of characters is probably Lachnocaulon digynum:

1) Heads grayish or gray-brown, definitely not whitish 2) Scapes smooth or nearly so, at least not densely pubescent 3) Leaves very small, averaging only 1 cm, none longer than 2 cm; sheath of the scape longer than the leaves 4) No air spaces visible in leaves 5) Roots slender fibrous, branched, not septate.

Confirmation of field identification can be made using floral characteristics.

Duration: PERENNIAL
Reproduction Comments: This species is probably wind pollinated, due to the tiny size of the flowers and well-exserted stamens and style. Kral (1966) has not observed insect pollination in any of the Eriocaulaceae. Water seems to be the obvious agent of seed dispersal, along with the random adherence of seeds to the feet of birds and mammals.
Ecology Comments: Little specific information is available on the species biology of Lachnocaulon digynum. However, there are several general characteristics of the Eriocaulaceae which we can assume to apply to this species. Seed germination probably requires some type of soil surface disturbance, and perhaps concomitant seed scarification, and copious soil moisture without much inundation of the substrate. The above statement is hypothetical, but fits well with observations of the species and its habitats. These conditions can most naturally be met by heavy rains after surface fires which expose the soil surface. Once established, Lachnocaulon digynum is one of the more stable and persistent members of the family. The plant grows and the colony expands by forming clumps of additional rosettes from short rhizomes. Its short stature and protected buds probably protect the colony from destruction by most fires, thus it can regenerate quickly after fire from both fruiting and flowering condition, depending on local weather conditions. The species is probably wind pollinated, due to the tiny size of the flowers and well-exserted stamens and style. Kral (1966) has not observed insect pollination in any of the the Eriocaulaceae. Mature fruiting bodies are evident in fall and probably persist into winter, slowly shedding seed, likely by the action of rain on the heads. Water seems the only obvious agent of dispersal, along with the random adherence of seed to the feet of birds and mammals.

The species appears to be rather long-persistent under the natural conditions where it occurs. Its clump-forming habit would tend to reduce the probability of competition within the colony, by insuring a dense cover of Lachnocaulon where establishment of other species would be difficult. However, due to its short stature, Lachnocaulon digynum can easily be shaded by taller surrounding vegetation, which would result in population decline. Without periodic fire to remove competition, the species would undoubtedly not persist for very long. With periodic fire, it most likely would remain as a part of the community indefinitely.

Riverine Habitat(s): Pool
Palustrine Habitat(s): Bog/fen, FORESTED WETLAND
Terrestrial Habitat(s): Savanna
Habitat Comments: Lachnocaulon digynum is restricted to seasonally or semipermanently saturated substrates, usually with little or no shrub or tree cover, near the southern edge of the East Gulf Coastal Plain. General habitat requirements seem to include partial or full sun, an almost permanently or at least seasonally wet substrate, little shrub or tall herb competition, and a substrate of sand, mucky sand, muck, sandy peat, or Sphagnum species.

These requirements are met in a variety of natural communities, and microsites within natural communities, of the region. An analysis of label data from 20 of the 22 known collections resulted in the following list of natural communities occupied by the species, and presumed microsites within these.

Lachnocaulon digynum is restricted to seasonally or semipermanently saturated substrates, usually with little or no shrub or tree cover, near the southern edge of the East Gulf Coastal Plain. It seems to have been found only in the Coastal Pine Meadows (=Gulf Coastal Lowlands in Florida) and Longleaf Pine Belt (=Western Highlands in Florida) sections of the East Gulf Coastal Plain (Powell 1983), and never more than 90 miles from the Gulf of Mexico, most often within 40 miles of the Gulf. 90% of the known sites are within an area only 150 miles east to west, centered on Mobile Bay, Alabama.
1) Hillside seepage herb bog - East Gulf Coastal Plain type - generally on lower slopes and bowl-shaped streamheads in the Lower Longleaf Pine Belt or on the escarpment between the Longleaf Pine Belt and Coastal Pine Meadows from the Apalachicola River westward to the Florida parishes of Louisiana. Maintained by seepage from adjacent uplands, and kept open by frequent fire. Have been studied by Eleutarius and Jones (1969). The surrounding uplands can be either pine flatwoods or longleaf pine/scrub oak sandhills, possibly resulting in differing hydrological conditions within the bog. Three Lachnocaulon digynum sites could be definitely referred to this type, two surrounded by pine flatwoods and one by sandhills.

2) Hillside seepage shrub bog - East Gulf Coastal Plain type - This is basically the previous type without frequent fire, and succeeding towards a seep forest climax. The one Lachnocaulon digynum site from this type had an overstory of Ilex sp., Titi (Cliftonia monophylla or Cyrilla racemiflora) and Bay (Magnolia virginiana or possibly Persea sp. or Gordonia lasianthus). This is presumably not a stable site for Lachnocaulon, and it may only be persisting from a time when the community had a more open character, and may be present only in openings.

3) Wet Pine Savannah (after Duever (1984) ) or Mesic to Wet Pine Savannah (after Walker and Peet (1983) ) - East Gulf Coastal Plain Type - Restricted to the Coastal Pine Meadows (Gulf Coastal Lowlands) region from the Florida panhandle to eastern Louisiana. On generally flat land, maintained by seasonal saturation or inundation due to high water table, and by frequent fire (Duever 1984, Folkerts 1982). Have been studied by Norquist (1984), and the related Atlantic Coastal Plain type has been studied by Walker and Peet (1983), among many others. Within this type, Lachnocaulon digynum has been reported once with no distinction as to microhabitat, but has more often been associated with one of two types of microsites within this type. One of these includes seepage areas, pond margins, and swales, most likely inundated for longer periods than the surrounding savannah, and exposed only in summer and fall. Lachnocaulon digynum has been reported three times from this microhabitat, where it probably has less competition than in the "drier" savannah. Another microsite (with three reports of Lachnocaulon digynum) includes "bogs" within this community type, presumably with Sphagn um and probab ly Sarrac enia sp.

4) Mesic (Longleaf) Pine Flatwoods - East Gulf Coastal Plain Type - Found in the Longleaf Pine Belt from Florida to Louisiana; on flat uplands with sand substrate, frequent fire. In this type, Lachnocaulon digynum (2 sites) is restricted to spagnous or sandy bogs or swales within the community, which would have a hydrologic regime resembling that of the Wet Pine Savannah. The farthest inland sites for the species are believed to be in these microsites within this community.

5) Spring-fed Stream - Western Highlands Type - Range not determined, apparently numerous in the western Florida panhandle (Wilhelm 1984). Characterized by clear water and usually sandy-bottomed, perennial or seasonal. Seepage stream of Brinson (1982), at least in part. The one collection of Lachnocaulon digynum from this type is described as "rooted in muck in shallow water."

An additional five sites are described as being in bogs or sphagnum bogs, with no futher specifications. These could be referred to communities 1,2, or 3. One other collection, and references in general habitat descriptions in Kral (1966), are from wet sands and sandy peats, ditches, and roadbanks. Most likely these represent less natural ecological equivalents of swales within wet pine savannahs described above, but do indicate the persistance of the species after some disturbance.

Associated species have rarely been mentioned, but are most obviously the herbaceous plants of the above communities. Specifically mentioned as associated with Lachnocaulon digynum by Ken Gordon (pers. comm.) is Xyris drummondii, which is described by Kral (1966a) as occurring in the same habitats as the former, and which has an almost identical range.

Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: Management of this species should include 1) biological monitoring and species biology research to determine relationship of the species to fire regimes and microhabitat conditions, 2) inventory of areas of potential habitat within the range for additional sites, particularly on public lands, 3) determination of the highest quality occurrences and natural communities for protection, 4) negotiation of management agreements for sites on public lands, and 5) to continue to manage preserved sites through prescribed burning and protection from drainage and other threats.
Restoration Potential: The recovery potential of Lachnocaulon digynum is probably dependent almost exclusively on the degree of habitat alteration which has occurred. I believe that establishment or reestablishment of the species is probably possible in any area of habitat which has not been excessively drained, leveled by bulldozing, or chemically poisoned. Fire would remove woody and tall herb competition and expose the soil surface. Seed could be sown in a variety of apparently suitable microsites, both before and after fires in various seasons. The opportunity for successful establishment is probably good, given the proper combination of soil exposure, stratification, and moisture has been achieved by these attempts to simulate the probable natural conditions for establishment. In excessively altered sites, it would probably be difficult to find a microsite suitable for the species, and recovery potential would be poor. Declining sites could be restored by burning, with clipping of woody vegetation if necessary, as long as the soil surface or drainage pattern had not been altered.
Management Requirements: Lachnocaulon digynum requires active management, most importantly the maintenance of its habitat through prescribed burning. In addition to prescribed burning, it is also necessary to prevent drainage of the habitat by adjacent or upslope ditching. No grazing should be allowed in the habitat, and no major disturbances of the soil surface within the population. Minor surface disturbances, however, could provide sites for establishment of new colonies.

There are numerous studies documenting the role of fire in the maintenance of mesic pine flatwoods (Lemon 1949, Garren 1943, Christensen 1981) wet pine savannahs (Folkerts 1982, Kologiski 1977, Walker and Peet 1983), and hillside seepage bogs (Eleutarius and Jones 1969). However, there is no specific information on the fire regime most conducive to growth of Lachnocaulon digynum. Probably late fall or winter fire every few years would be best for Lachnocaulon; annual fire may be appropriate for Lachnocaulon, although it could reduce overall species diversity. Summer fires may be more beneficial in some cases, particularly if the competing vegetation is becoming too dense. Komarek (1965) believes summer fires caused by lightning to have been the prevalent regime in presettlement times, and Folkert (1982) suggests that it may be more effective at opening up space for germination and establishment of species such as Lachnocaulon digynum. Perhaps fall or winter fires are effective in maintenance of existing populations, but summer fires may be better in the creation of new sites for colonization. Further observations and research will be necessary to clarify this question.

Monitoring Requirements: Lachnocaulon digynum is in definite need of biological monitoring to determine basic information about its stability at a site, responses to competition and fire regime, and appropriate management techniques to perpetuate the species.

In monitoring this species, the definition and distinction of individual plants is impossible, therefore percent cover data is more appropriate. Monitoring should take place in late summer or early fall, when the species has reached its maximum development for the year. Measurements, probably within permanently located 1/2 sq m or smaller quadrats, should consist of percent cover and height of competition by stratum (low herb, tall herb, shrubs) and list of species present, and observations on soil moisture.


Management Programs: Management at the Buttercup Flats site should have begun in 1985 by the Crosby Arboretum Foundation. Presumably, this will consist of prescribed burning, perhaps filling or blocking ditches or ruts which may be draining the area slightly, and monitoring for signs of effects of nearby development. No other sites are known to be specifically managed for Lachnocaulon, although it is likely that some are burned periodically for timber management purposes.
Monitoring Programs: Monitoring should be beginning for this species at Buttercup Flats Preserve, Mississippi, this year. No other monitoring programs are known to exist.
Population/Occurrence Delineation Not yet assessed
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Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 22Mar2018
NatureServe Conservation Status Factors Author: Morse, L. & K. Gordon; rev. VEC, rev. Treher (2018)
Management Information Edition Date: 24Oct1986
Management Information Edition Author: EDWIN BRIDGES
Element Ecology & Life History Edition Date: 07Jul1992

Botanical data developed by NatureServe and its network of natural heritage programs (see Local Programs), The North Carolina Botanical Garden, and other contributors and cooperators (see Sources).

References
Help
  • Brinson, S. 1982. Draft Florida riverine community classification. Florida Natural Areas Inventory, Tallahassee. 1 p. Unpublished report.

  • Carr, S. 1991. PETS Plant species listed by ranger district. Table sent over Data General computer mail system to V. Crouch from Forest Botanist Susan Carr, Kisatchie National Forest, Pineville, Louisiana. 3 pp.

  • Christensen, N. L. 1981. Fire regimes in southeastern ecosystems. Pages 112-136 in: H.A. Mooney, T. M. Bonnicksen, N. L. Christensen, J. E. Lotan, and W. A. Reiners, technical coordinators. Proceedings of the Conference Fire Regimes and Ecosystem Properties, December 11-15, 1978, Honolulu, HI. General Technical Report WO-GTR-26. USDA Forest Service, Washington, DC. 594 pp.

  • Clewell, A.F. 1985. Guide to vascular plants of the Florida panhandle. Florida State Univ. Press, Tallahassee, Florida. 605 pp.

  • Duever, L. 1984. Draft palustrine and terrestrial community classifications, and descriptions. Unpublished draft report by the Florida Natural Areas Inventory, Tallahassee, FL. 20 pp.

  • Eleuterius, L. N. and S. B. Jones, Jr. 1969. A florisitic and ecological study of pitcher plant bogs in south Mississippi. Rhodora 71:29-34.

  • Flora of North America Editorial Committee. 2000. Flora of North America north of Mexico. Vol. 22. Magnoliophyta: Alismatidae, Arecidae, Commelinidae (in part), and Zingiberidae. Oxford Univ. Press, New York. xxiii + 352 pp.

  • Folkerts, G.W. 1982. The Gulf Coast pitcher plant bogs. American Scientist 70: 260-267.

  • Garren, K. H. 1943. Effects of fire on vegetation of the southeastern United States. Bot. Rev. 9:617-654.

  • Godfrey, R.K., and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United States: Monocotyledons. Univ. Georgia Press, Athens. 712 pp.

  • Hatch, S.L., K.N. Gandhi, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agricultural Experiment Station, College Station. 158 pp.

  • Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.

  • Kologiski, R.L. 1977. The phytosociology of the Green Swamp, North Carolina. Tech. Bull. 250. North Carolina Agriculture Experiment Station, North Carolina State University. Raleigh, NC. 101 pp.

  • Komarek, E.V., Sr. 1965. Fire ecology: grasslands and man. Pages 169-220 in Proc. 4th Annual Tall Timbers Fire Ecology Conference.

  • Kral, R. 1966a. Xyris (Xyridaceae) of the continental U.S. and Canada. Sida 2(3): 177-260.

  • Kral, R. 1966b. Eriocaulaceae of continental North America north of Mexico. Sida 2:285-332.

  • Lemon, P. C. 1949. Successional responses of herbs in the longleaf-slash pine forest after fire. Ecology 30:135-145.

  • Norquist, H.C. 1984. A comparative study of the soils and vegetation of savannas in Mississippi. M.S. thesis, Mississippi State Univ., Mississippi State. 110 pp.

  • Poole, J. M., W. R. Carr, D. M. Price, and J. R. Singhurst. 2007. Rare plants of Texas. Texas A&M University Press, College Station. 640 pp.

  • Powell, R.L. 1983. Potential geological Natural Landmarks of the East Gulf Coastal Plain. Report to Natural Landmarks Branch, U.S. Dept. of the Interior. 434 pp.

  • Small, J.K. 1933. Manual of the southeastern flora. Two volumes. Hafner Publishing Company, New York.

  • Walker, J. and R.K. Peet. 1983. Composition and species diversity of pine-wiregrass savannas of the Green Swamp, North Carolina. Vegetatio 55:163-179.

  • Wilhelm, G.S. 1984. Vascular flora of the Pensacola region. Ph.D. dissertation, Southern Illinois Univ., Carbondale. 213 pp.

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