NTFP Conference Proceedings Lycopodium: Growth Form, Morphology, and Sustainability of a Non-timber Forest Product Elizabeth A. Nauertz1 and John C. Zasada2 Abstract.—Several species of Lycopodium or clubmoss belong in the category of non-timber forest products and are often gathered for a variety of traditional uses. It is important to evaluate baseline infor- mation for these species, such as abundance and frequency of occur- rence, before making any management decision. In addition, under- standing the biology of the Lycopodium group as a whole, including growth form and morphology, may enable us to make better decisions about forest management practices used and harvesting quotas allowed for sustaining these species. INTRODUCTION SPECIES DESCRIPTIONS AND USES Several Lycopodium or clubmoss species are Following is a brief description of each of the extensively collected as non-timber forest six Lycopodium species discussed in this paper. products and are marketed to industries for The descriptions are compiled from personal production of seasonal and traditional decora- observations, as well as information referenced tions and for floral and horticulture uses. In from the literature (Cody and Britton 1989; addition, medicinal (Aboriginal and homeo- Flora of North America 1993; Johnson et al. pathic), native folklore, and theatrical uses 1995; Lellinger 1985; Meeker et al. 1993; exist for various Lycopodium species (Johnson Primack 1973; Turner et al. 1983; Ullman et al. 1995; Shakhashiri 1983; Ullman 1992, 1992, 1997). 1997). Lycopodium species are considered to be fern-allies. In temperate forests they are vascu- Lycopodium species vary in growth form and lar, terrestrial, evergreen, perennial, rhizoma- morphology. The mature aerial stems of a plant tous, and clonal in nature. This group of photosynthesize. They may be either non- species has been around a long time, with branching or treelike in form. When mature, ancestors that date back to the Tertiary period the aerial stems often have cones (strobili) or (Lellinger 1985). Few studies have been done to sporangia that produce the spores necessary determine how these species can be regener- for sexual reproduction. Spores mature and are ated and how fast they recover after harvesting released in the late fall, even as late as Novem- (Matula 1995, Primack 1973). Although some ber. In general, Lycopodium aerial stems reach of the Lycopodium species are common in maturity and begin to produce spores at from 4 northern hardwood forests, the impacts of to 6 years of age, depending on the species and forest management practices on populations of local growing conditions. Lycopodium species these species are not well understood. have either aboveground or belowground running rhizomes or lateral branches. The 1Ecologist, U.S. Department of Agriculture, rhizomes have the ability to produce adventi- Forest Service, North Central Research Station, tious roots and are used for photosynthate, 410 MacInnes Drive, Houghton, Michigan 49931, water, and nutrient transport. The rhizomatous USA; Phone: 906-482-6303; e-mail: nature of these species may add to the photo- [email protected]. synthesis capacity of the plant as a whole and 2Project Leader and Research Forester, U.S. may affect the plant’s ability to get around Department of Agriculture, Forest Service, North vegetatively. A typical Lycopodium patch can Central Research Station, 1831 Highway 169 E., have multiple-aged lateral branches or rhi- Grand Rapids, Minnesota 55744, USA; Phone: zomes, and each rhizome may have ramets that 218-326-7109; e-mail: [email protected]. vary in age from 1 to 6 years. In general, the 110 bigger and more branching the patch, the older the same manner as the ground cedar. Tradi- it is. tional Aboriginal and homeopathic remedies are concocted from the dried plant parts of this The aerial stems of Lycopodium dendroideum species. L. clavatum has forking, non-branch- and L. obscurum are frequently harvested and ing aerial stems that typically bear one to two used for decorative greens. Although commonly cones when mature. The aerial stems are often found in aspen-birch forests, these two species tightly packed along the aboveground running are also found in moist rich woods and along rhizomes. In Canada, Aboriginal peoples on edges of bogs. The branching aerial stems Vancouver Island, located in British Columbia, resemble small pine trees: hence the common believed the plant should be left alone. It is said name of princess pine or ground pine. Only the that clubmoss should be left because it is individual mature aerial stems should be “something that gets you confused in the harvested, and this should be done after prime woods” or “confused and uncertain about spore release in late fall. The spores of these orientation” (Turner et al. 1983). Perhaps this two species are flammable when mature and belief stemmed from the random pattern of have been used historically for theatrical and branching that is common for this species. pyrotechnical purposes. In addition, the ground pines have belowground rhizomes Lycopodium annotinum or stiff clubmoss is not running between 2 and 6 inches below the soil traditionally gathered or harvested. Although surface (Nauertz and Zasada, personal observa- considered evergreen, this species tends to dry tions); consequently, a new aerial stem may not out quickly and does not remain green and appear above the soil surface until sometime in pliable, as do the other species harvested for the second year of growth. decorative greens. Lycopodium annotinum has forking, non-branching aerial stems that Lycopodium complanatum, also known as produce a single strobilus, or cone, when ground cedar, is most commonly found in pine mature. The stems are typically densely packed forest communities, typically grows in clumps, along aboveground running rhizomes. and can cover large areas. It has tree-like, branching aerial stems with adpressed and Shining clubmoss or Lycopodium lucidulum is tightly packed microphylls or leaves that most common to northern mesic forests domi- strongly resemble cedar boughs. When mature, nated by sugar maple, preferring the embedded the aerial stems host a “candelabra” of cones acid, wet woods or rocky areas. Native Ameri- for spore production. This species of Lycopo- cans traditionally gathered it for use as pad- dium differs from the ground pines L. ding for cradleboards. L. lucidulum does have dendroideum and L. obscurum, in that it has some features that are unique to this group of aboveground running rhizomes growing clubmoss species. It has forking, non-branch- through the litter layer on the forest floor. ing aerial stems that, when mature at around 6 Lycopodium complanatum is also harvested for years of age, produce sporangia that contain decorative greens and has some homeopathic spores in the leaf axils. In addition, gemmae or and Aboriginal medicinal uses. Harvesting for bulblets may be produced in the upper leaf greens is different for this plant in that the axils. They mature usually in late summer or entire aboveground runner, or rhizome, is early fall, and when bumped will catapult off yanked from the surface of the soil; hence, the the aerial stem and, upon landing in proper entire plant or section of a clone is harvested, growing media and conditions, have the poten- rather than just the mature aerial stems as tial for growing into a new, genetically identical done for the ground pines. Optimal lateral plant. This is a means of vegetatively increasing extention or growth for each rhizome has been plant frequency, but not genetic variation. recorded to occur in year 1 (most recent year Lycopodium lucidulum does not technically growing tip) to year 6, with maximum extension form rhizomes; instead the aerial stems tend to reaching 400 to 600 cm. Greatest aerial stem layer, and then branch and fork. Each fall the mass was recorded in years 3 to 6, with peak leaf litter from the overstory canopy falls, weights reaching 13 to 25 grams (Nauertz and anchoring the layered aerial stems to the Zasada, unpublished data). surface of the forest floor. Adventitious roots for water and nutrient uptake will form along the Running clubmoss, Wolf’s paw, or Lycopodium layered aerial stems. clavatum also has aboveground running rhi- zomes and is harvested for decorative greens in 111 NTFP Conference Proceedings LYCOPODIUM REPRODUCTIVE be very dense and vast, or it may be thin and CHARACTERISTICS sparse. Patches of Lycopodium may be located near each other and occur frequently within an Reproduction method, both sexual and area, or there may be large areas where no asexual, may help explain the frequency and Lycopodium occurs at all. cover patterns of individual Lycopodium spe- cies. All ferns and fern-allies produce spores Lycopodium, especially ground pine, is often and experience an alternation in generations, found in northern hardwood forests in which is a means of dividing, segregating, and Michigan’s Upper Peninsula. Nauertz (1999) re-combining chromosomes. This allows for conducted a study in these forests to gain genetic variability to occur and for evolution to insight on the species biology and presence in proceed (Lellinger 1985). Spores vary in where managed and unmanaged forests. In addition, they germinate within the soil surface layers as part of a larger study (USDA, FS, NCRS, NC- and were found to germinate at deeper depths 4153-94-03), basal area was measured and as well as on the soil surface (Cobb 1963, recorded in all of these stands in 1995-1996. Freeberg 1962). The spores produced by fern- Four forest types were studied: (1) Unmanaged allies such as Lycopodium may remain viable old-growth forest with trees as old as 250+ for many years and could take up to 7 or more years that have not been disturbed by logging. years to develop into a gametophyte (Cobb Basal area was 23 - 47 m2/ha. (2) Unmanaged 1963). Gametophytes may remain alive from second-growth forest that was clearcut in the only a few weeks or months (Lellinger 1985) to 1910s and now remains unmanaged with trees as long as 10 or more years (Cobb 1963).