ELEMENT STEWARDSHIP ABSTRACT FOR TOFIELDIA RACEMOSA July 1, 1994 Stewardship Abstract No.: 017 By: Elizabeth M. Obee State of New Jersey Department of Environmental Protection Division of Parks and Forestry Office of Natural Lands Management CN 404 Trenton, New Jersey 08625 Element Stewardship Abstract Element Stewardship Abstracts (ESA's) are prepared to provide land managers and other conservation workers with current biological and management related information on those species and natural ecosystems that are most important to protect or for which control is most needed. The abstracts organize and summarize data from numerous sources, including the literature and from researchers and managers actively working with the species or ecosystem. The ESA format was originally developed by The Nature Conservancy as a starting point for the stewardship of the many species and ecosystems, or elements, protected by the Conservancy. The New Jersey Office of Natural Lands Management is developing ESA's for those elements that are of particular importance as components of the biota of the state. This includes globally rare plant species that are also listed on New Jersey's official Endangered Plant Species List. The ESA serves several important functions. It helps to identify information gaps and target future research efforts. It provides a standard format for highlighting specific information about a species or community including its management needs. It also allows information to be readily communicated among various preserves, state offices, regional centers, natural heritage programs and private organizations. The ESA is a dynamic document that is continuously updated as new information becomes available. Users are encouraged to contribute their information to the abstract. This sharing of information will benefit all land managers by ensuring the availability of up-to-date information on management techniques and knowledgeable contacts. Please contact the Office of Natural Lands Management for an ESA publication list. It will contain the date of the latest revision to each ESA. Please refer to the abstract number when ordering ESA's. The abstract is a compilation of available information and is not an endorsement of particular practices or products. Element Name: TOFIELDIA RACEMOSA (Walt.) B.S.P. (Liliaceae) Element Code: PMLIL1Y050 Preparer: Elizabeth M. Obee Common Name: False Asphodel Description: Habit: Perennial rhizomatous herb with linear basal leaves and an erect stem terminating in white flowers. For a complete description see Gleason (1952) or Fernald (1950). Stem: Sticky, pubescent, leafy only at the base. Leaves: Parallel veined, erect, grass- or iris-like, in 2 vertical rows from near base. Variable in length, to 4 dm long and 3-5 mm wide. Usually one bracteal leaf inserted somewhat below the middle of the scape. Inflorescence: A raceme or thyrse up to 15 cm long, with terminal flowers opening first. Often 3-7 flowers per node in the lower and middle of the inflorescence, and one per node near the top. Flowers: Showy, with 3 petals, 3 sepals, and 6 stamens. Small, star-like, white flowers with superior ovaries clustered at the tip of a stalk. Concave 3-nerved sepals without claws. Fruits: A septicidal capsule, many seeded, subtended by persistent perianth segments, tipped by enlarged divergent styles. Seeds are oblong and horizontal. Distinctions from Related Species: The genus is composed of about 20 species of the north temperate zone and mountains of South America, with 2 or 3 other species in the western states or artic America. Three species are generally recognized in the eastern U.S.; Tofieldia glutinosa, T. racemosa, and T. glabra. Some authors treat T. glutinosa as a variety of T. racemosa, but it is usually recognized as a distinct species (Radford, 1968). The range of T. glutinosa is more northern, occurring into Canada, Minnesota, and Ohio, but also extending south in the mountains to Georgia. The preferred habitat is also different, with T. glutinosa occurring at calcareous rock ledges, marshes, and shores, and T. racemosa in wet sand, clay, or peat (Fernald, 1950). These two species are distinguished by T. racemosa having a capsule equal or shorter than the persistent perianth and seeds with tails 0.5 or less as long as the body. T. glutinosa has a capsule longer than the perianth and seeds with long slender contorted tail or tails. T. glabra is the third eastern U.S. species in this genus, found from North Carolina to Georgia in pine savannas, flatwoods, and shrub bogs, and can be distinguished by having a glabrous scape, inflorescence axis and flower stalk. Another species T. tenuifolia, was recently transferred into the genus from Pleea tenuifolia. It occurs primarily in the coastal plain of North Carolina, but not at the same locations as T. racemosa or T. glabra (Utech, 1978). Habitat: T. racemosa grows on the Coastal Plain from New Jersey to north Florida and west to southeast Texas. It is common in North Carolina, South Carolina, Georgia, Mississippi, and Florida. A disjunct population exists in Tennessee which is isolated from those in the Coastal Plain. T. racemosa occurs in wet, sandy- peaty soils, pine savannas and flatwoods, shrub bogs and seepage slopes. At the northern edge of the species' range in New Jersey there are five confirmed extant occurrences (New Jersey Natural Heritage Program, 1994). All are on state owned land in open bogs or savannas edged by white cedar swamps and pitch pine lowlands, with a total number of plants estimated at less than one thousand. There are five additional historical sites, three of which may be extant. The remaining two historical sites were destroyed by agricultural activity (New Jersey Natural Heritage Program, 1994). Within New Jersey it is often found growing with a closely related and also endangered lily species which is now restricted to the New Jersey Pine Barrens, Narthecium americanum (Stone, 1911; Schuyler, 1990). Associated herbaceous species at Narthecium locations include Aster nemoralis, Calamagrostis pickeringii, Calamovilfa brevipilis, Juncus caesariensis, Lophiola americana, Muhlenbergia torreyana, Platanthera integra, Pogonia ophioglossoides, Rhynchospora oligantha, Schizaea pusilla, and Sphagnum spp. (Schuyler, 1990). In Delaware the species is considered historical at five locations and possibly extirpated from the state. The species was once present at sites described as sea-level fens, herbaceous wetlands just above high-tide and adjacent salt marshes. These wetlands contain deep peaty muck and are formed from low- nutrient, acidic fresh ground water seepage from adjacent uplands (McAvy, pers. comm.). The historical sites are now either Spartina alterniflora or Phragmites australis marshes. In Virginia the species was always rare (Harvill, 1973). There is now only one extant site, estimated at 28 individuals in 1992 (Virginia Natural Heritage Program, 1992). The plants persist in a railroad right of way near the edge of a boggy swale in a habitat unusually dry for this species. Other species at the site include Pteridum aquilinum, Sericocarpus linifolia and Gaylussacia spp. In Tennessee there is one extant site of several hundred individuals which is disjunct from the rest of the Coastal Plain populations. The site is an open, wet savanna within the oak barrens of the southeast highland rim. This area supports several other species disjunct from the Coastal Plain including Platanthera nivia and Carex barattii (Pyne, pers. comm.). T. racemosa becomes increasingly common along the Gulf Coast. It is common throughout west Florida panhandle bogs, seepage slopes and wet prairies to just east of the Appalachicola river, at probably 50 to 100 sites (Hilsenbeck, pers. comm.). Habitat types in the Florida panhandle were described by Schwartz (1994). T. racemosa is listed as one of the species co-occurring at several sites in the Florida panhandle with a newly described species, Eriocaulon nigrobracteatum (Orzell and Bridges, 1993). The habitat that these plants grow in was characterized as deep unstable sapric muck soils of lower slope seepage fed herbaceous communities. Orzell and Bridges (1993) refer to the habitat as poor fens, which are oligotrophic to weakly minerotrophic, somewhat acidic, nutrient deficient mires poor in species of Sphagnum. The occurrences of these sites is correlated with the escarpment of sandy upland terraces reflecting various relict shorelines. Poor fens apparently form along the middle and upper reaches of small stream valleys in upland terraces. In these areas topographical relief attracts groundwater from the unconsolidated surficial aquifers (Orzell and Bridges, 1993). Other west Florida wetland habitats were described in a study of pine savannas containing Cladium mariscoides, a species recently reported for the Florida flora (Bridges and Orzell, 1993). The range of T. racemosa continues along the coast, with probably several hundred occurrences within the appropriate habitat type in southern Mississippi. In southeast Louisiana to the east of the Mississippi there are 26 confirmed occurrences in St. Tammany and Washington Parishes. These eastern Louisiana savannas support more state-rare plant species than any other habitat type in the state (Louisiana Natural Heritage Program, 1990). The westernmost edge of the species' range is in southeast Texas. Biology/Ecology: In New Jersey, T. racemosa flowers from late-June to mid-July. Little is known about the pollination biology of the species. Pollination studies of other bog species revealed that Diptera and Lepidoptera were the common pollinators of T. glutinosa and other white flowered species (Douglas, 1983). Population growth in T. racemosa is predominantly due to rhizomatous spread rather than from sexual reproduction (Summerfield, 1974). No reproduction from seed was observed in transplanted populations of the European T. pusilla, which was instead successfully established through vegetative growth in a manner similar to T. racemosa (Cranston and Valentine, 1983). Flowering is probably inhibited by shading, as for many other bog species including Narthecium americanum (Schuyler, 1990). The lack of competition with woody vegetation in open bog and savanna habitats promotes growth and flowering of bog species. Maintenance of open habitat may be due to hydrology, soil conditions, disturbance such as fire, or a combination of factors. Bog areas have very hydric, acidic and nutrient poor soils, providing poor conditions for vigorous growth of tree species. Frequent fire contributes to these edaphic factors by providing disturbance that eliminates young invading tree seedlings and keeps these areas open (MacRoberts and MacRoberts, 1993). Fire is important for the stimulation of flowering of herbs and shrubs, eliminates invading hardwoods, and exposes mineral soil necessary for seedling microsites (Louisiana Natural Heritage Program, 1990). Fire also alters the time and duration of flowering in pine flatwoods (Platt et al., 1988). In the absence of fire an open longleaf pine flatwood savanna can become overgrown with shrubs in as little as five years (Louisiana Natural Heritage Program, 1990). The pre-historic natural frequency of fires in southeastern Louisiana pine savannas has been estimated to be once every 1-4 years, with lightning fires occurring in the early growing season from late March to early June along the Gulf Coastal Plain (Louisiana Natural Heritage Program, 1990). Northern savannas in the New Jersey Pine Barrens, however, are believed to be maintained primarily by hydrology, with the effects of fire less clear. T. racemosa is apparently dependent on water moving through the substrate, as may be other species occurring in this habitat such as N. americanum (Schuyler, 1990). Permanent flooding will eliminate these species, such as that caused by beaver (Hammerson, 1994), as will alterations in hydrology by cranberry culture common in the New Jersey Pine Barrens (Stone, 1911). Natural flooding patterns will likely maintain open habitat for the species. Determination of Element Occurrence (EO) Quality: High quality occurrences exist in states where the species is common, namely North Carolina, South Carolina, Georgia, Mississippi, and Florida. Efforts to protect and manage wetland habitat in these areas is crucial to the survival of this species. Threats: Open habitat where some rare plants occur can often be a result of past human alterations of ecosystems. As human land use changes, threats may arise which require active management to simulate the past disturbance history of the site. One example is at Crystal Fen in Maine, where the hydrology and fire history of a fen containing T. glutinosa and other rare species was altered over time (Jacobsen et al. 1991). The construction of a drainage ditch at the site and the elimination of railroad associated fires led to a recolonization of the fen by woody flora. This flora was originally present at the site before the construction of the railway led to increased flooding and fires at the site. T. glutinosa occurred in the open fen and in a section of the fen with a recently closed overhead canopy, but did not occur where the fen had a completely closed canopy (Jacobsen et al., 1991). Threats to T. racemosa include factors which disrupt the hydrology of the habitat, resulting in habitat loss. Habitat destruction may result from silvicultural activities such as slash pine (Pinus ellottii var. ellottii) plantations. These plantations disrupt or destroy seepage bog and wet prairie hydrology. Fire lanes may also disrupt hydrology, but fire suppression is an even greater threat. Potential disruption of habitat may be caused by flooding by beavers, turning wetland habitat into aquatic habitat (Hammerson, 1994). Damming streams for recreational lakes is also a threat, as is disturbance by recreational canoers. In the New Jersey Pine Barrens the construction of cranberry bogs could cause detrimental changes in hydrology. Succession of bog areas by woody vegetation is a threat which requires the condition of a high water table and/or frequent fire in order to maintain habitat quality. Land Protection Specifications: Protection of land for T. racemosa requires management of hydrology and control of succession by woody vegetation. Anything that disrupts present water regimes may result in changes in vegetation. Water flow in streams and seeps should be unaltered by ditches or fire lanes. Sites near agricultural and residential development should be protected from expansion of development, such as cranberry farming. Recovery Potential: Reproduction from seed is believed to be negligible, as was observed for T. pusilla (Cranston and Valentine, 1983). Development from seed may be slow, as in N. ossifragum where individuals raised from seed did not produce rhizomes for two years (Summerfield, 1974). The species is probably sensitive to destruction by trampling. Rhizomatous turf of N. ossifragum damaged by trampling resulted in stunted sterile shoots (Summerfield, 1974). However, fragmentation of rhizomes may stimulate lateral buds, as for N. ossifragum (Summerfield, 1974). Attempted transplantation of T. pusilla resulted in survival of some of the transplanted plants for at least five years (Cranston and Valentine, 1983). Transplantation is, however, generally considered a risky and last-measure technique for conservation (Fahselt, 1988). Recovery from disturbance by buried rhizomes is probably good, but recovery of suitable habitat from seed is likely to meet with little success. Biological Monitoring Needs: Locations of T. racemosa which may be directly impacted by human activity should be monitored to determine potential adverse impacts. Such a situation exists in Virginia, where the only current extant site should be monitored to determine whether or not the species is being adversely affected by railroad maintenance (Ludwig, pers. comm.). Any expansion activities such as road widening or ditching would clearly be detrimental. Populations subject to potential threats such as development or cranberry farming should be monitored annually. Also, populations which have low rates of flowering should be monitored to determine if they are being impacted. Selected populations in high quality habitats should be evaluated every two or three years. Other locations which are not directly impacted by development should also be monitored to determine whether threats are present from natural succession of woody vegetation or by changes in hydrology of the habitat. Biological Monitoring Procedures: Monitoring should record changes in population sizes and flowering phenology. Habitat parameters such as changes in water level and amount of invasion by woody vegetation should also be recorded. Suggestions for intensive monitoring would include determining what proportion of new individuals were produced as a result of asexual and sexual reproduction and the average longevity of individuals. Biological Monitoring Programs: No monitoring programs exist which specifically target this species, besides tracking by Natural Heritage Programs in states where it is considered rare. However, presence or absence is monitored in programs such as that in Tennessee where there is only one extant population. This population is being monitored during management to retard woody vegetation at May Prairie. Management strategies includes burning and brush-hogging the site of approximately 40 acres. This management strategy has been implemented at variable time intervals for the last 15 years, with as much as seven years between burns (Pyne, pers. comm.). The optimal burning regime for this community type is probably shorter, possibly every three to five years (Pyne, pers. comm.). This site is seasonally wet and is subject to rapid invasion by woody plants, including numerous red maple seedlings. Several fire management programs in Florida are conducted which may affect populations of Tofieldia, including programs at Eglin Air Force Base, Apalachicola National Forest, and state forests (Hilsenbeck, pers. comm.). However, specific management for this species is not being conducted at this time. Research Needs: Information is needed on the preferred hydrology for T. racemosa, as well as the ability of the species to survive alterations of the water regime. The specific response of the species to fire is also unknown. Fire is sometimes necessary to maintain open habitat for the species, but direct burning of established populations may be detrimental. Burning of Narthecium americanum resulted in poor regeneration and growth (Summerfield, 1974). Demographic information is needed, such as the commonness of reproduction from seed. Responses to disturbance and competition such as shading and trampling should be investigated. Management Needs: Future management recommendations for the Crystal Fen site where T. glutinosa occurs include cutting, prescribed burning, and manipulating drainage to maintain the floral diversity of the site (Jacobsen et al., 1991). Locations of T. racemosa may require similar intensive management if changes in hydrology and fire frequency threaten a location with becoming dominated by woody vegetation. Fires need to be allowed to burn naturally into bogs, seepage slopes, across wet prairies and savannas, especially during the growing season. Fire is considered an important factor in keeping bogs free of woody vegetation (MacRoberts and MacRoberts, 1993). Southeast pine savannas should be burned every one to three years, during the early growing season (Louisiana Natural Heritage Program, 1990). Burns early in the growing season are expected to cause greater mortality of shrubs and trees and stimulate flowering of herbaceous species (Louisiana Natural Heritage Program, 1990). Summary of Stewardship Needs: Loss of habitat is one important factor threatening T. racemosa. Less than 10% of the original pine flatwood savannas of Louisiana, for example, remains in an undisturbed condition (Louisiana Natural Heritage Program, 1990). However, indirect effects of changes in human land use and resulting shifts in hydrology and community structure are also important threats to the survival of T. racemosa. Alteration of the water regime is a serious threat and, therefore, T. racemosa is vulnerable to flooding or desiccation of its habitat. Reproduction from seed is low, restricting the colonization of new habitat and increasing the importance of maintaining habitat quality. Locations of T. racemosa should be monitored for population size changes, water level changes, and the number of flowering plants present, especially in locations on the periphery of the species' range. Specific protection of habitat is needed to maintain the hydrology and community structure necessary for survival of T. racemosa. It is likely to be negatively impacted by any human development or disturbance due to the specificity of its habitat requirements. This species is currently considered globally secure and not tracked by Natural Heritage Programs over most of its relatively limited range. However, it is hopeful that its occurrence in species rich and botanically unusual habitat sites may result in the inadvertent protection of this species before it becomes seriously endangered. Bibliography for Tofieldia racemosa (Walt.) B.S.P. : Bridges, E.L. and S.L. Orzell. 1993. Cladium mariscoides (Cyperaceae) in the western Florida panhandle and its phytogeographic significance. Phytologia 74: 35-42. Cranston, D.M., and D.H. Valentine. 1983. Transplant experiments on rare plant species from Upper Teesdale. Bio. Cons. 26: 175- 191. Douglas, S. 1983. Floral color patterns and pollinator attraction in a bog habitat. Can. J. Bot. 61: 3494-3501. Fahselt, D. 1988. The danger of transplantation as a conservation technique. Natural Areas Journal 8: 238-244. Fernald, M.L. 1950. Gray's Manual of Botany. 8th ed. American Book Co., New York. Gleason, H.A. 1952. The New Britton and Brown illustrated flora of the Northeastern United States and adjacent Canada. D. Van Nostrand Co., Inc., Princeton. Hammerson, G. A. 1994. Beaver (Castor canadensis): Ecosystem alterations, management, and monitoring. Natural Areas Journal 14: 44-57. Harvill, A.M. 1973. Some new and very local populations of rare species in Virginia. Castanea 38:305-307. Jacobsen, G.L., H. Almquist-Jacobsen, and J.C. Winne. 1991. Conservation of rare plant habitat: Insights from the recent history of vegetation and fire at Crystal Fen, northern Maine, USA. Bio. Cons. 57: 287-314. Louisiana Natural Heritage Program. 1990. The longleaf pine flatwood savannahs of southeastern Louisiana. Louisiana Natural Heritage Program. Department of Wildlife and Fisheries. Baton Rouge, Louisiana. MacRoberts, M.H., and B.R. MacRoberts. 1993. Why don't west Louisiana bogs and glades grow up into forests? Phytologia 74: 26-34. New Jersey Natural Heritage Program. 1994. Element occurrence records for Tofieldia racemosa. New Jersey Natural Heritage Program, Trenton, New Jersey. Orzell, S.L. and E.L. Bridges. 1993. Eriocaulon nigrobracteatum (Eriocaulaceae), a new species from the Florida panhandle, with a characterization of its poor fen habitat. Phytologia 74: 104-124. Platt, W.J., G.W. Evans, and M.M. Davis. 1988. Effects of fire season on flowering of forbs and shrubs in longleaf forests. Oecologia 76: 353-363. Radford, A.E., Ahles, H.E. and C.R. Bell. 1964. Manual of the vascular flora of the Carolinas. University of North Carolina Press, Chapel Hill, North Carolina. Schuyler, A.E., 1990. Element stewardship abstract for Narthecium americanum. New Jersey Department of Evironmental Protection and Energy, Division of Parks and Forestry, Office of Natural Lands Management. Trenton, New Jersey. Schwartz, M.W. 1994. Natural distribution and abundance of forest species and communities in northern Florida. Ecology 75: 687-705. Stone, W.H. 1911. Abama americana (Ker.) Morong. Bartonia. 4: 1- 5. Summerfield, R.J. 1974. Narthecium ossifragum (L.) Huds. J. Ecol. 62(1): 325-339. Utech, F.H. 1978. Floral vascular anatomy of Pleea tenuifolia Michx. (Liliaceae-Tofieldieae) and its reassignment to Tofieldia. Ann. Carn. Mus. 47: 423-454. Virginia Natural Heritage Program, 1992. Element occurrence record for Tofieldia racemosa.