Lupinus Perennis [L.]) Appears to Respond Favorably to Burning, No Detailed Studies of the Effect of Fire on This Species Have Been Published
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ABSTRACT: Although wild blue lupine (Lupinus perennis [L.]) appears to respond favorably to burning, no detailed studies of the effect of fire on this species have been published. We evaluated the effect of fire on the germination, growth, flowering, and seed set of Lupine by comparing burned and unburned populations. Fire killed most lupine seeds and seedlings located on the surface of the burned sites. At one of three study sites, survival of lupine seedlings emerging after the fire was significantly higher in burned plots than in unburned plots. Adult lupines in burned plots produced more biomass, allocated more of this biomass to leaves and stems, and had significantly higher nitrogen content than plants in unburned plots. Burned portions of three lupine populations exhibited higher percent cover and number of leaves than unburned portions of the same populations. Significantly higher percentages of flowering stems set seed, and the number of pods per T h e S h o r t - T e r m square meter increased in burned plots. These results suggest that burning lupine sites increases leaf production and seed set but also increases seed and seedling mortality. Timing burns to occur before seedlings emerge, and staggering burns a minimum of 2 E f f e c t of Fire on years apart would reduce fire damage to lupine seedlings. Lupinus perennis (L.) M i c h e l l e T r u d e a u G r i g o r e INTRODUCTION Metropolitan Park District successional processes, lupine has declined of the Toledo Area Wild blue lupine (Lupinus perennis [L.]) is in abundance and is now listed as poten- 5 1 00 West Central Avenue the sole larval food source of three species of tially threatened in the state of Ohio (Ohio Toledo, Ohio 43615 USA butterfly listed as endangered in the state of Division of Natural Areas and Preserves Ohio: the frosted elfin (Incissalia it-us 1992). Elliot J. 'Framer [Godart]), Persius dusky wing (Erynais Several preserves in the Oak Openings are Department o f Biology persius [Scudder]), and the Karner blue The University of Toledo butterfly (Lycaeides melissa samuelis being managed with prescribed burning to decrease woody plant dominance and in- To l ed o , Ohio 43606 USA [Nabokov]); the latter also is listed as a federal endangered species (U.S. Fish and crease the abundance of prairie plants such Wildlife Service 1992). The decline of these as lupine. Following fire treatments, lu- species in Ohio has been attributed to loss of pine has responded vigorously by increas- • nectaring plants and lupines (Shuey et al. ing vegetative growth, flowering, and seed 1987, Magdich 1989). Understanding the set (Mary Huffman, The Nature Conser- management needs of lupine is important for vancy, Lake Wales, Florida, and Jennifer devising an effective plan to conserve lupine Windus, Ohio Division of Natural Areas and these rare butterflies. and Preserves, Columbus, pers. com.). These observations encouraged us to con- In northwestern Ohio, lupine is found in the duct a systematic study of the effect of fire oak savanna community that is scattered on lupine. Specifically, our objectives were throughout the dry portions of a sandy soil to (1) determine if lupine germination and belt known as the Oak Openings region. seed mortality are affected by pre-scribed This community type was shaped by fire burning; (2) analyze the effect of fire on prior to settlement (May-field 1976) and mature lupine plants' growth, nutrient consisted of grasses, prairie forbs, and small content, flowering, and seed set using clumps of widely spaced oak trees, which biomass and transect studies; and (3) gave the region its name. With settlement study the effect of fire on lupine seed-ling came the suppression of fire as well as mortality. drainage, pastures, and farms. Areas of the Oak Openings not put into production were LIFE HISTORY AND FIRE colonized by woody species, which shaded EFFECTS out the lupine and other prairie plants (Campbell 1931, Mayfield 1976). As a result Information on the life history of lupine is of these scattered through the literature; we sum- marize it here for the convenience of land managers. Natural Areas Journal 16:41—48 Volume 16 (1), 1996 Natural Areas Journal 41 Lupinus perennis is a member of the Fa- The literature on the effects of fire on Populations had to be dense enough to baceae (Gleason and Cronquist 1991) and prairie plants covers species other than support a 25-m transect with a maximum a native of the eastern United States from lupine. Prairie grasses tend to flower and of 1-m spacing between lupine clumps to Minnesota to Maine and south to the Gulf fruit more heavily and exhibit increased enable consistent sampling over all six Coast (Haack 1993). Lupine grows well in growth after a fire. Most studies (Curtis populations. Soils at all six sites are clas- full sun or partial shade, but shows limited and Partch 1950, Robocker et al. 1953, sified as Ottokee and Tedrow series, survival under totally shaded conditions Hulbert 1969, Peet et al. 1974) point to a formed in sandy material on postglacial (Bess 1989, cited in Haack 1993). Lupines reduction in litter and increased light avail- beach ridges and dunes (Stone et al. 1980). frequently are found in poor soils, benefit- ability as the reasons for this increased The three burned sites were utilized in ting from nitrogen fixation by Rhizobium vigor, although higher spring temperatures 1992 to continue the transect studies on bacteria that form nodules in their roots due to blackening of the soil and the re- burned and unburned portions of the same (Allen and Allen 1981). The rhizome of lease of minerals from ash have also been lupine population. wild blue lupine has "eyes" that send up suggested. Schivarin et al. (1987) con- new shoots to form clumps (Foster 1984); cluded that fire does not affect mycorrhizal Germination Studies these perennating buds are located 0–10 fungi and nutrient storage in fire-treated cm below the soil surface (Harshberger little bluestem (Schizachyrium scoparium Only cream-colored L. perennis seeds were 1970). The shoots that emerge are ramets Michx). Reichert and Reeder (1971) noted used for germination experiments to elim- (Harper 1977) that produce clonal individ- that the soil temperature 1.0 cm below the inate possible permeability differences uals through asexual reproduction. Zarem- surface remained unchanged by a prairie based on pigment variation in the seed ba et al. (1991) noted that 2-year-old lupine fire. These latter two studies are significant coat (Werker et al. 1979, Egley et al. 1983). plants can have up to 5 vegetative shoots, in light of the potential effect of heat on To test the effect of fire on hardened lu- and 3-year-old plants may support up to 17 the Rhizobium bacterium involved in root pine seed, and to remove any seed that had stems under ideal conditions. nodule formation and nitrogen fixation in not developed a hard seed coat, the meth- lupine. od outlined by the International Seed Test- Lupine leaves develop from the taproot in ing Association for germinating seeds of April in northwest Ohio. Flowers bloom in METHODS the genus Lupinus (Anonymous 1966) was mid-May and seed pods form by mid-June. employed on 500 cream-colored seeds. A plant will produce all cream or all Study Sites Fifty of the seeds (10%) germinated dur- mottled seeds, with two to six seeds per ing the test, which meant they had perme- pod, depending upon the growing condi- The Oak Openings geologic region ex- able seed coats; the other 450 seeds were tions. The hairy seed pods dry and dehisce tends through portions of Henry, Lucas, impermeable to water. These "hard" seeds by twisting in early July, sending seeds and Fulton counties in northwestern Ohio were used for testing the effect of fire on distances of up to 5 m from the plant and continues into southeastern Michigan germination; seeds were not cold stratified (authors, pers. obs.). (Figure 1). Several public agencies have prior to heat treatments. preserved portions of this region in west- Seeds of the genus Lupinus exhibit dor- ern Lucas County, including The Nature To simulate the various fuel types that lu- mancy characteristics that may be altered Conservancy, the Ohio Department of pine seeds would be subjected to under by fire or scarification. The seeds can lie Natural Resources–Division of Natural natural conditions, two mesh envelopes, dormant for many years because they are Areas and Preserves, and the Metropolitan each containing 50 seeds, were randomly impermeable, and they will germinate Park District of the Toledo Area. Pre- placed at the base of little bluestem (Schiza- under increasing relative humidity (Quin- serves owned by these agencies are being chyrium scoparium), two in a leaf litter mix livan 1971) or when the seed coat is cracked managed with prescribed burning to re- of sweet-fern (Comptonia peregrina L.) and by pressure or temperature fluctuations store degraded oak savanna communities dewberry (Rubus flagellaris Wind.), and (Bewley and Black 1982). Seeds of wild to presettlement conditions; these areas two in light oak leaf litter at 1 cm below the blue lupine do not require cold treatment provided the burned sites used in this study. surface to simulate lupine seed that had for germination (Nichols 1934, Zaremba et Several private landowners granted per- become stratified in the soil. Two envelopes al. 1991); however, Boyonoski (1992) mission to study lupine populations in the of 50 cream-colored lupine seeds each were found that stratified lupine seed exhibits Oak Openings that are not subjected to placed on the ground in an unburned prai- 100% germination versus 65% for unstrat- fire.