Lupinus perennis



INTRODUCTORY


 

Fred Nation, Atlas of Florida Vascular Plants
AUTHORSHIP AND CITATION:
Meyer, Rachelle. 2006. Lupinus perennis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].

FEIS ABBREVIATION:
LUPPER

SYNONYMS:
Lupinus perennis spp. perennis var. occidentalis S. Wats. [32,52,70,73]
L. p. spp. perennis var. perennis [32,70].

NRCS PLANT CODE [70]:
LUPE3

COMMON NAMES:
wild lupine
blue lupine
sundial lupine
perennial lupine

TAXONOMY:
The scientific name of wild lupine is Lupinus perennis L. (Fabaceae) [19,32,52,73,81]. Accepted subspecies are:

L. p. spp. gracilis (Nutt.) D. Dunn
L. p. ssp. perennis [32,70]

LIFE FORM:
Forb

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
Information on state-level protected status of plants in the United States is available at Plants Database.

DISTRIBUTION AND OCCURRENCE

SPECIES: Lupinus perennis

GENERAL DISTRIBUTION:
In Canada, wild lupine occurs in Ontario and Newfoundland [20,32]. In the United States it extends from Minnesota and Iowa eastward through the Lake States and New England south through Kentucky, West Virginia, and the Atlantic coast states to northern Florida. From northern Florida its range extends westward to eastern Texas [11,25,32,70].

Lupinus perennis ssp. gracilis occurs in the southern portion of the species's range from Texas east through the Gulf states and north through Atlantic states to Virginia [32]. Lupinus p. ssp. perennis occurs in Ontario, Newfoundland, and the northeastern United States from Minnesota, Iowa and Illinois east to the Atlantic coast and south as far as Georgia. Since infrataxa are not usually distinguished in the literature, they will not be discussed further in this review. Plants Database provides a distributional map of wild lupine and its subspecies.

The following lists are based on wild lupine distribution information and the habitat characteristics and plant species composition of vegetation communities wild lupine is known to occupy. There is not conclusive evidence that wild lupine occurs in all the habitat types listed, and some community types may have been omitted.

ECOSYSTEMS [18]:
FRES10 White-red-jack pine
FRES12 Longleaf-slash pine
FRES13 Loblolly-shortleaf pine
FRES14 Oak-pine
FRES15 Oak-hickory
FRES19 Aspen-birch
FRES39 Prairie

STATES/PROVINCES: (key to state/province abbreviations)
UNITED STATES

AL CT DE FL GA IL IN IA KY LA
ME MD MA MI MN MS NH NJ NY NC
OH PA RI SC TX VT VA WV WI DC

CANADA
NF ON

BLM PHYSIOGRAPHIC REGIONS [1]:
None

KUCHLER [36] PLANT ASSOCIATIONS:
K074 Bluestem prairie
K081 Oak savanna
K082 Mosaic of K074 and K100
K095 Great Lakes pine forest
K100 Oak-hickory forest
K104 Appalachian oak forest
K106 Northern hardwoods
K110 Northeastern oak-pine forest
K111 Oak-hickory-pine
K112 Southern mixed forest
K115 Sand pine scrub

SAF COVER TYPES [14]:
1 Jack pine
14 Northern pin oak
15 Red pine
16 Aspen
20 White pine-northern red oak-red maple
21 Eastern white pine
40 Post oak-blackjack oak
42 Bur oak
43 Bear oak
44 Chestnut oak
45 Pitch pine
46 Eastern redcedar
52 White oak-black oak-northern red oak
53 White oak
69 Sand pine
70 Longleaf pine
71 Longleaf pine-scrub oak
72 Southern scrub oak
75 Shortleaf pine
76 Shortleaf pine-oak
78 Virginia pine-oak
79 Virginia pine
80 Loblolly pine-shortleaf pine
81 Loblolly pine
82 Loblolly pine-hardwood
110 Black oak

SRM (RANGELAND) COVER TYPES [60]:
601 Bluestem prairie
808 Sand pine scrub
810 Longleaf pine-turkey oak hills

HABITAT TYPES AND PLANT COMMUNITIES:
In addition to the plant communities listed above, wild lupine occurs in disturbed habitats. In power line rights-of-way of eastern New York, wild lupine occurred on sites where little bluestem (Schizachyrium scoparium), Tatarian honeysuckle (Lonicera tatarica), northern dewberry (Rubus flagellaris), and bear oak (Quercus ilicifolia) had the highest coverages and little bluestem, northern dewberry, black oak (Q. velutina), whorled yellow loosestrife (Lysimachia quadrifolia), sedges (Cyperaceae), and goldenrods (Solidago spp.) occurred at the highest frequencies [16]. In east-central New York, wild lupine was studied in 3 rights-of-way vegetation types: wild lily-of-the-valley-starflower (Maianthemum canadensis-Trientalis borealis), sweetfern-whorled yellow loosestrife (Comptonia peregrina-Lysimachia quadrifolia), and blackberry-sheep sorrel (Rubus spp.-Rumex acetosella). In the wild lily-of-the-valley-starflower type, mosses (Bryophyta, 6.9%), wild lily-of-the-valley (4.4%), grasses (Poaceae, 4.4%), and starflower (2.1%) had the highest cover. Coverage in the sweetfern-whorled yellow loosestrife type was dominated by grasses (40.9%), sweetfern (12.1%), mosses (9.4%), and whorled yellow loosestrife (5.2%). In the blackberry-sheep sorrel type, the dominants included grasses (22.7%), northern dewberry (5.0%), other blackberries (Rubus spp., 4.8%), and sheep sorrel (4.3%) [62]. Wild lupine had an average 3% cover in a 70-year-old abandoned sand mine in Indiana Dunes National Lakeshore dominated by eastern cottonwood (Populus deltoides), black oak, and sassafras (Sassafras albidum) [76]. In eastern Wisconsin, wild lupine was present in a 5-year-old prairie restoration planting with quackgrass (Elymus repens), pinnate prairie coneflower (Ratibida pinnata), and black-eyed Susan (Rudbeckia hirta) [10].

In Indiana Dunes National Lakeshore, wild lupine was not present on plots dominated by black locust (Robinia pseudoacacia), and occurred significantly (p<0.05) more often in black oak-dominated sites with basal areas of black locust <1 m/ha [51].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Lupinus perennis

Drew Feldkirchner, Wisconsin Department of Natural Resources
GENERAL BOTANICAL CHARACTERISTICS:
This description provides characteristics that may be relevant to fire ecology, and is not meant for identification. Keys for identification are available (e.g. [19,52,73,81]).

Wild lupine is a long-lived, cool-season, nitrogen-fixing forb with a thick, deep taproot [8,25,64]. The fine roots only survive about 4 weeks [8]. According to reviews and a gardening guide, wild lupine is rhizomatous [11,22,64]. In a comprehensive review of the information available on wild lupine rhizomes, Girgore and others [22] note that the perennating buds of this native perennial can be found 0 to 4 inches (0-10 cm) below the soil surface. Wild lupine grows 8 to 24 inches (20-60 cm) tall and has palmately compound leaves with 7 to 11 leaflets from 0.6 to 2.4 inches (1.5-6 cm) long. The perfect flowers are bilabiate. Petals up to 0.6 inch- (1.6 cm) long occur on erect racemes. After pollination, a 1.2- to 2-inch (3-5 cm) legume pod forms [19,52]. From information in the literature, Halpern [26] reports that pod may contain up to 7 seeds. Average seed mass is typically between 27 and 28 mg [23,83]. The seed mass of 5,839 seeds from 59 plants ranged from 8 to 41 mg [26].

RAUNKIAER [54] LIFE FORM:
Geophyte

REGENERATION PROCESSES:
According to reviews, wild lupine reproduces by production of ramets from rhizomes and germination from seed [11,16,22,26,71].

Pollination: Reviews list honey bees, bumble bees, eastern carpenter bees, and butterflies including black swallowtails, clouded sulphurs, and Karner blues as pollinators of wild lupine [11,26]. Beetles, ants, and thrips may also pollinate wild lupine [26].

Breeding system: Wild lupine is monoecious [19]. Although wild lupine can self-pollinate, most breeding occurs through cross-pollination. Self-pollinated flowers yield significantly (p≤0.0002) less fruit and seed per inflorescence and have more aborted seeds per fruit [59].

Seed production: Wild lupine may begin flowering in its 2nd year, but likely starts later in mediocre conditions [40,66,82]. It may not sprout every year. Wild lupines produced as many as 52 pods on 7 flowering stalks [82]. In west-central Wisconsin, Maxwell [47] observed increased flowering in cooler years. Number of seeds per inflorescence in self pollinated wild lupine was 40% to 60% (p<0.0001) less than outcrossed seeds [59]. The 3 highest seed yields of 4-year-old wild lupine on planted sites in Wisconsin were [66]:

Clean seeds/acre (lbs) Seeds/lb (#) Soil texture on the site
898 20,600 Fertile loam
189 22,100 Moderately fertile loam
159 23,900 Moderately fertile loamy sand

Seed dispersal: According to reviews, seed dispersal only occurs by dehiscence of the seed pod [11,22,26]. Seeds can be thrown from 3 feet (1 m) [11] to 16.4 feet (5 m) from the plant [22].

Seed banking: Based on the literature, Halpern [26] reports wild lupine seeds may germinate the summer they mature or remain dormant in the seed bank for at least 3 years.

Germination: Wild lupine seeds germinate in varying conditions. In the laboratory Mackay and others [46] found similar germination after 72 hours in seeds germinated in light and dark conditions. Zaremba and Pickering [82] observed germination throughout the year in the Albany Pine Bush of New York. However, higher germination was observed in wild lupine seeds planted in spring (59%) compared to those planted in fall (39%) [82].

Some scarification methods improve wild lupine seed germination compared to unscarified seeds, while others do not [12,22,46]. In an experiment to determine the effects various scarification treatments, 72% of seeds in the untreated group germinated over an unspecified time period. Seeds scarified with hot water exhibited an insignificant (p>0.05) decline in percent germination (57%), while those tumble scarified with pea gravel for 2 to 3 hours showed an insignificant increase (89%). Mechanical scarification using a commercial scarifier that threw seed against an abrasive drum resulted in significant (p<0.05) declines in percent germination (52%) [12]. After 1 week Mackay and others [46] reported less than 15% germination of unscarified wild lupine seeds, while percent germination of seeds scarified in sulfuric acid was about 90% for seeds soaked 15 minutes and approached 100% for seeds soaked ≥30 minutes. Seeds nicked using a razor blade exhibited 100% germination. Soaking wild lupine seeds in water of varying temperatures (72-212 F (22-100 C)) did not promote germination [46]. Exposure to fire may result in large decreases in percent germination. After 90 days, Grigore and Tremer [22] found significantly (p<0.001) lower percent germination in seeds on the surface of prescribed burned sites compared to buried and unburned seeds (see Discussion and Qualification of Plant Response).

Water availability affects percent germination. For instance, seeds that received 11 inches (28 cm, ambient) or 14 inches (35 cm, wet) of water over 3 months exhibited 92% germination, while only 62% of seeds limited to 2 inches (6 cm, dry) of water germinated. In addition, the effect of seed mass on germination varied with water availability. The probability of germination increased 3-fold with a 10-mg increase in seed mass in seeds exposed to ambient or wet conditions, while seeds in dry conditions did not show increased germination with increased seed size. In addition, the decrease in time to germination due to increased seed mass was larger in the ambient and wet treatments than in the dry treatment [26].

Temperature can also influence germination. Seeds collected in southern Ontario and stored in a freezer for 6 weeks exhibited significantly (p<0.05) higher germination after 4.5 months (99%) than those stored at room temperature (62%) [3]. In addition, at least 2 studies have cold-stratified wild lupine seeds before experimentation [26,83]. However, percent germination of both cold-treated (near freezing for 71 days) and control seeds collected in southern New England was 41% [49]. Zaremba and Pickering [82] also report germination without a cold treatment. Wild lupine seeds scarified in sulfuric acid exhibited decreased percent germination at high temperatures. From 70 to 85 F (21-29C), more than 80% of scarified seeds germinated within 54 hours. Percent germination was about 60% in scarified seeds in the 90 F (32 C) treatment and less than 4% in the ≥95 F (35 C) treatments [46].

Partial predation of seeds may reduce germination. In a laboratory study where predation was simulated by removing 30% or 60% of seed reserves, control seeds exhibited 80% emergence rates, while ≤50% of treated seeds emerged [83].

Seedling establishment/growth: Seedling survival of wild lupine is typically low. In New York, only 327 seedlings out of 1,235 germinated seeds survived their 1st winter [82]. In northwestern Ohio the highest mortality on unburned oak savanna sites was 40% [22]. In southern Ontario, only 20% of seeds planted and germinated in 1990 sprouted in 1991. In addition to predation, mortality of these and other seedlings in the area was due to desiccation, frost, and possibly shading. Increased amounts of litter and species competing for light and other resources may also reduce wild lupine establishment [3].

Mortality of lupine seedlings subjected to fire is high. Within 2 months of prescription burning in oak (Quercus spp.) savanna of northwestern Ohio, 95% of lupine seedlings on 1 site and 100% on another site died. The effect of these burns on seedlings that emerged after the fire is less clear. On 1 of 3 sites, lupine seedling survival was significantly (p<0.01) lower in burned areas than in unburned areas. The site had not been burned for 10 years before the experimental fire, while the 2 sites where wild lupine seedling survival on burned and unburned areas did not differ (p>0.1) had been burned ≤2 years prior to the experimental fires [22].

Wild lupine seedlings germinated from large seeds may have a better chance of surviving, at least in the short term, than seedlings from small seeds. Survival of seedlings to July was doubled with a 10-mg increase in seed mass, and larger seeds were related to greater plant size in years 1 and 2 [26].

In a simulated herbivory experiment, wild lupine seedlings subjected to 60% removal of seed reserves were significantly (p<0.05) shorter, had smaller leaf areas, and had lower dry weight than control seeds or those subject to 30% removal of seed reserves [83].

Asexual regeneration: According to reviews, a wild lupine rhizome can produce several shoots that form clumps. Ramets may be over 3 feet (1 m) from the genet, which can make identification of individual plants difficult [22,26,71].

SITE CHARACTERISTICS:
Wild lupine is commonly found in dry, sandy openings such as those found in savannas, woodland clearings, or disturbed areas [19,39,52,64,68,81].

Wild lupine typically occurs in well-drained, sandy soils with slightly acidic pH. Wild lupine's occurrence in sandy soil has been widely reported [9,26,64,68]. In southern Wisconsin, sites with wild lupine typically had more than 80% sand [41]. Wild lupine has also been reported on neutral to strongly acidic soils [33,82]. In the Albany Pine Bush of New York, soils where lupine was growing had an average pH of 5.3. This was a higher pH than found at sites throughout the Albany Pine Bush (x pH=4.8) [82]. At the Allegan State Game Area in Michigan wild lupine was found in soils with pH ranging from 4.2 to 5.6 [21]. A review states that wild lupine grows in basic conditions in the Finger Lakes region of New York [11].

Wild lupine may occur at higher frequencies on sites with disturbed soil [37,39,47]. In west-central Wisconsin, a flush of seedlings was documented after vehicular traffic disturbed the soil in a closed stand. In the same study area, experimental seeded sites prepared by grubbing trees, tilling, and herbicide application had significantly (p<0.0001) higher frequency of flowering and nonflowering lupine combined (65%) than undisturbed quadrats (40%). The author calls for more research into the importance of soil disturbance to wild lupine [47]. In Minnesota, sites associated with steep sand banks that experience sloughing were the only areas of dense wild lupine [37]. According to a review, reductions in soil disturbance may be a cause of wild lupine decline [11]. However, another review lists "excessive" soil disturbance as a possible cause for wild lupine decline [55].

Little information is available regarding other characteristics of wild lupine sites, such as elevation or precipitation. Wild lupine is included in a commercially available seed mixture meant for planting above 7,000 feet (2,000 m) [57]. Forrester and others [16] investigated wild lupine in eastern New York on sites from 200 to 400 feet (60-120 m) elevation that received average annual precipitation of 37 inches (930 mm). A site with wild lupine in southern Wisconsin receives average annual precipitation of 34 inches (858 mm), of which a 3rd typically falls during the peak of wild lupine's growing season [26]. A seed catalog recommends 12 inches (300 mm) of precipitation for wild lupine [53].

SUCCESSIONAL STATUS:
According to reviews, wild lupine is an early successional species [55,71] that prefers open and partially shaded conditions. At Indiana Dunes National Lakeshore, increases in canopy cover resulted in significant (p<0.001) declines in lupine abundance [24]. Areas of the Allegan State Game Area with low stand densities were more likely to support wild lupine than areas with high stand density, and wild lupines grown in full sun in a greenhouse were significantly (p≤0.05) larger than those grown in 35% or 65% full sunlight [21]. In rights-of-way in east-central New York, light intensity in lupine populations was significantly (p≤0.0001) greater than in adjacent areas. To favor wild lupine, investigators recommended increasing photosynthetically active radiation to at least 65% of the maximum light intensity by reducing cover [62]. Sites with wild lupine in southern Wisconsin had an average photon flux density of about 70%, with values typically ranging from 50% to 95% [41]. In southwestern Michigan, wild lupine recruitment was positively affected for 3 years after reducing cover of associated vegetation by mowing and herbicide application [23]. At a military training base in west-central Wisconsin, the median strips between tracked-vehicle ruts had significantly (p<0.05) less shrub and forest canopy and significantly (p<0.05) more wild lupine than areas 16 feet (5 m) outside the ruts [63]. In the same area, wild lupine cover was highest from 22% to 32% total daily photo flux density. Wild lupine frequency was greater near tree boles, although the shaded plants were smaller and less likely to flower [47]. In southern Wisconsin, wild lupine was most abundant in semishade and on sites in full sun with little cover of other species [40]. Observations at 4 sites in southern Ontario suggest that both very high and low light intensities may be detrimental to wild lupine [3]. In the Allegan State Game Area in Michigan, wild lupine "quality" was highest in partial sun. This was probably due to open sites being drier, which senesced wild lupine foliage [21,38]. Similar observations were made in August of 1992 at Fort McCoy, Wisconsin, where wild lupine in open areas were desiccated and those in shade looked healthy [39]. Reviews note wild lupine's intolerance of complete shade [22,25,71].

For information on the effects of canopy cover on the quality of wild lupine as a Karner blue butterfly resource see Preferred Habitat in the FEIS review of Karner blue butterfly.

SEASONAL DEVELOPMENT:
According to reviews, wild lupine in the northern portion of its range begins sprouting from rhizomes in late March or April and flowers in May and June [11,22]. Wild lupine flowers from April to July along the Blue Ridge Mountains of the Southeast [78], and in April and May in the Carolinas [52]. Fruiting occurs in June and July and senescence begins by August [11,22,25,26,52]. In study in New York, germination was observed throughout the year [82]. According to a review, wild lupine plants do not sprout every year. Length of dormancy in adult plants is unknown [71].

FIRE ECOLOGY

SPECIES: Lupinus perennis
FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: Germination of seeds in the seed bank and sprouting established plants are the most likely sources of wild lupine in the initial postfire community. Wild lupine seedling emergence after prescribed burns was documented by Grigore and Tramer [22], although possible sources of seed (seed bank or off-site) are not discussed. Given the lack of impact of prescribed burns on seeds buried to 0.4 inch (1 cm), wild lupine's short dispersal distances (see Seed dispersal), and the germination, albeit limited, of seeds exposed to moderate fires [22], the seed bank is the most likely source of postfire emergent seedlings. In addition, soil probably provides enough insulation to protect wild lupine rhizomes from fires. According to a review of wild lupine's life history, rhizomes are located from 0 to 4 inches (0-10 cm) below the soil surface [22]. It is likely that plants with relatively deep rhizomes survive more severe fires than plants with shallow rhizomes [65]. However, there have been no investigations of fire-related mortality of established lupine or the importance of rhizome depth on wild lupine survival.

Surviving wild lupine, emerging seedlings, and wild lupine adjacent to a burn may provide sources for secondary colonization of a burned area. However, given wild lupine's short dispersal distances, colonization from nearby areas would likely take a considerable amount of time, especially in large burned areas.

Fire regimes: According to reviews, wild lupine typically occurs in habitats subject to fairly frequent fires [11,55,71]. Fire season in areas with wild lupine varies from dormant-season to early and late-growing season fires. Fire regimes where wild lupine occurs are influenced by several factors including habitat, historical period, and weather conditions [15,79]. Information regarding fire regimes in wild lupine habitats such as savannas [7,44,79] and pitch pine (Pinus rigida) barrens [15,42,45,48] is available.

The following table provides fire return intervals for plant communities and ecosystems where wild lupine may occur. For further information, see the FEIS review of the dominant species listed below.

Community or Ecosystem Dominant Species Fire Return Interval Range (years)
bluestem prairie Andropogon gerardii var. gerardii-Schizachyrium scoparium <10 [35,50]
jack pine Pinus banksiana <35 to 200 [6,13]
shortleaf pine Pinus echinata 2-15
shortleaf pine-oak Pinus echinata-Quercus spp. <10
sand pine Pinus elliottii var. elliottii 25-45
longleaf pine-scrub oak Pinus palustris-Quercus spp. 6-10 [74]
red pine (Great Lakes region) Pinus resinosa 3-18 (x=3-10) [5,17]
red-white pine* (Great Lakes region) Pinus resinosa-P. strobus 3-200 [6,28,43]
pitch pine Pinus rigida 6-25 [4,29]
eastern white pine Pinus strobus 35-200
eastern white pine-northern red oak-red maple Pinus strobus-Quercus rubra-Acer rubrum 35-200
loblolly pine Pinus taeda 3-8
loblolly-shortleaf pine Pinus taeda-P. echinata 10 to <35
Virginia pine Pinus virginiana 10 to <35
Virginia pine-oak Pinus virginiana-Quercus spp. 10 to <35
oak-hickory Quercus-Carya spp. <35
northeastern oak-pine Quercus-Pinus spp. 10 to <35
southeastern oak-pine Quercus-Pinus spp. <10
white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra <35
northern pin oak Quercus ellipsoidalis <35
bear oak Quercus ilicifolia <35
bur oak Quercus macrocarpa <10 [74]
oak savanna Quercus macrocarpa/Andropogon gerardii-Schizachyrium scoparium 2-14 [50,74]
chestnut oak Quercus prinus 3-8
post oak-blackjack oak Quercus stellata-Q. marilandica <10
black oak Quercus velutina <35 [74]
*fire return interval varies widely; trends in variation are noted in the species review

POSTFIRE REGENERATION STRATEGY [65]:
Rhizomatous herb, rhizome in soil
Ground residual colonizer (on-site, initial community)
Secondary colonizer (on-site or off-site seed sources)


FIRE EFFECTS

SPECIES: Lupinus perennis
IMMEDIATE FIRE EFFECT ON PLANT:
Although no research reports sprouting of previously established wild lupine plants in burned areas, wild lupine morphology and generally positive or neutral response to fire [22,34,47,72] suggest that it is typically top-killed.

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
Mature plants are more likely to survive fire than seeds or seedlings. On sites in northwestern Ohio, prescribed fires killed 95% to 100% of wild lupine seedlings and resulted in significantly (p<0.001) lower germination of seeds placed on the soil surface before the fires. Adult mortality was not investigated, but established wild lupines exhibited several positive responses to prescribed burns (see Plant Response to Fire just below) [22].

PLANT RESPONSE TO FIRE:
Wild lupine plant response to fire is generally positive. Wild lupine typically increases flower and/or seed production after fire [3,22,47], and most likely sprouts from rhizomes after fire. However, presence/absence data for wild lupine rhizome sprouts on burned sites were not available as of this writing (2006). Burned portions of wild lupine populations in northwestern Ohio had significantly higher seed production (p=0.01) and increased number of pods/m (p=0.003) compared to unburned portions. In addition, wild lupine on burned sites had significantly larger biomass (p=0.035), higher nitrogen content (p=0.04), lower potassium content (p=0.008), and allocated more biomass to leaves and stems (p=0.023) than wild lupines in unburned areas [22]. On 2 burned sites in southern Ontario, the site with the most frequent burning history had the highest average percentage of flowering wild lupine, while the site that had only been burned once in over 30 years had the lowest average percentage of flowering lupine [3]. In west-central Wisconsin, prescribed burns also increased flowering in wild lupine. However, this did not translate into a noticeable increase in reproduction [47].

Wild lupine abundance is typically unaffected by fire. In oak (Quercus spp.) savannas of central Wisconsin, wild lupine percent cover did not exhibit a significant (p>0.1) response to prescribed fires conducted in July or November [34]. In upland vegetation in northwestern Wisconsin that was burned under prescription, wild lupine frequency increased an average of 1.1% on burned sites, a "neutral" response [72]. On oak savanna burned under prescription in northwestern Ohio, wild lupine exhibited an insignificant (p=0.16) increase in percent cover compared to unburned areas [22]. In west-central Wisconsin, prescribed fires in the fall of 1993 and spring of 1994 had little effect on wild lupine frequency, suggesting the fires did not reduce dormancy or increase seedling recruitment. In the year following prescribed burning, however, nonflowering (p<0.01) and flowering (p<0.05) wild lupine on burned sites experienced significantly less decline than wild lupine on unburned sites. Statistical analysis suggested increases in wild lupine in later years were due to reasons other than burning. The following table shows the number of nonflowering and flowering lupine plants in burned and control plots from 1993 to1995 and 1997 [47].

  Nonflowering wild lupine Flowering wild lupine
Fire Year Burn Control Burn Control
prefire 1226 1152 1614 1756
<1 postfire year 895 723 1485 1168
<2 postfire years 1079 1122 892 667
<4 postfire years 1318 1209 1950 1653

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Burning can influence germination and seedling mortality. Seeds placed 0.4 inch (1 cm) below the surface on a prescribed burn site with light oak litter exhibited similar germination after 90 days (75%) as seeds placed on the soil surface of an unburned prairie site (69%). Despite having significantly (p<0.001) reduced germination (7%) compared to unburned and buried seeds, seeds exposed to a moderate fire (340 F (171 C)) on a site with sweetfern and northern dewberry litter germinated significantly faster (15.29 days, p<0.01) than unburned (36.44 days) or buried (34.72 days) seeds. There was no germination of wild lupine seeds exposed to a hot fire (930 F (500C)) in clumps of little bluestem [22]. Although site histories led to confounding of treatment and site effects in field experiments at 4 sites in Pinery Provincial Park in southern Ontario, germination of seeds planted in plots after burning was higher than control plots, significantly (p<0.05) so on 2 of the sites. Germination was also generally higher on sites with a history of recent burns. Mortality of the resulting seedlings was significantly lower (39.1%) than in control plots (56.9) [3]. However, there were no significant (p>0.1) differences in mortality of seedlings that emerged after fire on burned and unburned areas on 2 oak savanna sites in northwestern Ohio, and significantly (p<0.01) more postfire emergent seedlings died in the burned area (60%) than in the unburned area (15%) of a 3rd site [22].

Time since last fire may influence wild lupine's response. For example, the differing responses of wild lupine seedlings on the 3 sites in northwestern Ohio (see Discussion and Qualification of Plant Response) may have been related to time since last burning. The site where significant differences were observed had not been burned in 10 years, while the sites where no significant differences were observed had been burned within the previous 2 years. The effect of time since burning on fire severity on these sites and/or differences in other site characteristics that may have influenced the response were not discussed [22].

Fire and site characteristics are likely to influence wild lupine's response to fire. However, information regarding the impact of these factors is limited. Wild lupine would likely experience greater detrimental effects when fires are relatively severe and/or perennating buds are located close to the soil surface [65]. Season of burning may influence wild lupine response. In west-central Wisconsin, cover of wild lupine differed significantly (p<0.05) between spring and fall fires, with spring fires resulting in a "more favorable" response. Frequency of wild lupine did not differ significantly between sites burned in fall and those burned in spring [47]. The response of associated vegetation is also likely to influence wild lupine's response. Prescribed fires in west-central Wisconsin only reduced canopy cover in areas with less than 50% tree canopy that were comprised primarily of pines (Pinus spp.) and multi-stemmed oaks such as black oak and northern pin oak [47].

It has been suggested that wild lupine's nitrogen-fixing ability may give it an advantage over other species on burned sites due to nitrogen volatilization during fire and nitrogen loss from thatch burn-off [39,47].

FIRE MANAGEMENT CONSIDERATIONS:
Available data suggest that wild lupine abundance is unaffected and flowering increases after fire [3,22,34,47,72]. An investigation of fire's affect on wild lupine adults, seedlings, and seeds in oak savanna of northwestern Ohio led researchers to conclude that spring prescribed fires intervals of at least 2 years provides habitat conditions necessary for lupine (see Successional Status) while minimizing recruitment losses from seed and seedling mortality [22]. However, the lack of information regarding wild lupine's response to fire, including the effects of various fire and site characteristics, precludes specific recommendations. In addition, the most appropriate method for maintaining lupine habitat will vary depending management objectives and site characteristics. For instance, in areas occupied by the Karner blue butterfly, small, patchy, less-frequent fires are typically recommended due to the importance of heterogeneity to their habitat and the possibility of high fire-related Karner blue butterfly mortality. Other methods used to maintain wild lupine habitat include mowing, cutting, and/or herbicide application (see Other Management Considerations) [47,71].

MANAGEMENT CONSIDERATIONS

SPECIES: Lupinus perennis
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Wild lupine is toxic to some livestock, but is an important food source for a variety of wildlife species.

Palatability/nutritional value: Mature plants can be toxic to domestic sheep and horses [69,80].

Insects such as beetles, butterflies, and moths feed on wild lupine [77,82]. According to reviews, wild lupine is an important larval food source for several butterflies, including the frosted elfin and the federally endangered Karner blue butterfly [2,58].

Mammals have also been reported to feed on wild lupine. Reviews report deer [22,56,75,82] and several small mammals such as rabbits, woodchucks, and chipmunks [22,82] feeding on wild lupine foliage. However, meadow voles consistently rejected wild lupine seeds in a feeding trial [30].

Herbivory can have negative impacts on wild lupine. A review implicates white-tailed deer grazing as a factor in wild lupine decline in the Albany Pine Bush [75], and excessive spring grazing may cause wild lupine mortality [56]. Increased grazing pressure by white-tailed deer was considered a concern for wild lupine in 2 sites in Pinery Provincial Park, Ontario [3]. In northwestern Ohio, "extensive" insect feeding damage was observed in wild lupine seedlings that initially survived prescribed fires but died by the end of the growing season [22]. In southern Ontario, the majority of wild lupine seedling mortality on 4 sites was due to predation. Slugs were likely responsible for the significantly (p<0.05) higher seedling mortality found on 1 of these sites [3]. Transplanted and drought-stressed wild lupine may be more susceptible to damage from herbivory [82].

It has been suggested that grazing by megafauna such as bison and elk before European contact may have prevented succession of savannas to woodland habitats and thus provided open sites for wild lupine [47,67].

Cover value: No information is available on this topic.

VALUE FOR REHABILITATION OF DISTURBED SITES:
Wild lupine shows higher production and nitrogen accumulation on mine spoils than several agricultural legume species [31]. Results of various planting methods and possible limitations of wild lupine use in prairie restoration and mitigation projects are addressed by [27,33,82]. Sperka [64] provides general information regarding planting wild lupine.

OTHER USES:
No information is available on this topic.

OTHER MANAGEMENT CONSIDERATIONS:
Literature reviews discuss the threats to and management of wild lupine and its habitat [11,25,55,71,75].

In addition to prescribed burning, mowing, herbicide application, and/or cutting may be used and, in some cases, are required to maintain or create wild lupine habitat [16,23,47]. In southwestern Michigan, a combination of mowing and application of herbicide in spring reduced overall vegetative cover, contributing to an increase in wild lupine recruitment that was observed for 3 years [23]. In rights-of-way in eastern New York, wild lupine populations were larger on herbicide, mowing, and/or cutting treatment plots than in untreated areas [16].

Direct seedling has increased wild lupine cover on sites prepared by grubbing trees, tilling, and herbicide application [47].

Swengel [67] listed plowing, frequent mowing and continuous close-cropped grazing as management practices that are incompatible with maintaining wild lupine.

Lupinus perennis: REFERENCES


1. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]
2. Black, Scott Hoffman; Vaughan, D. Mace. 2005. Species Profile: Lycaeides melissa samuelis Nabokov, 1944: Karner blue (Lycaenidae: Polyommatinae: Polyommatini), [Online]. In: Shepherd, Matthew D.; Vaughan, D. Mace; Black, Scott Hoffman, eds. Red list of pollinator insects of North America. CD-ROM Version 1 (May 2005). Portland, OR: The Xerces Society for Invertebrate Conservation (Producer). 9 p. Available: http://www.xerces.org/Pollinator_Red_List/Leps/Lycaeides_melissa_samuelis.pdf [2006, June 6]. [63161]
3. Boyonoski, Anna May. 1992. Factors affecting the establishment and maintenance of Lupinus perennis (wild lupine). Guelph, ON: University of Guelph. 167 p. Thesis. [62566]
4. Buchholz, Kenneth; Good, Ralph E. 1982. Density, age structure, biomass and net annual aboveground productivity of dwarfed Pinus rigida Moll. from the New Jersey Pine Barren Plains. Bulletin of the Torrey Botanical Club. 109(1): 24-34. [8639]
5. Clark, James S. 1990. Fire and climate change during the last 750 yr in northwestern Minnesota. Ecological Monographs. 60(2): 135-159. [11650]
6. Cleland, David T.; Crow, Thomas R.; Saunders, Sari C.; Dickmann, Donald I.; Maclean, Ann L.; Jordan, James K.; Watson, Richard L.; Sloan, Alyssa M.; Brosofske, Kimberley D. 2004. Characterizing historical and modern fire regimes in Michigan (USA): a landscape ecosystem approach. Landscape Ecology. 19: 311-325. [54326]
7. Cole, Kenneth L.; Taylor, Robert S. 1995. Past and current trends of change in a dune prairie/oak savanna reconstructed through a multiple-scale history. Journal of Vegetation Science. 6: 399-410. [50258]
8. Craine, J. M.; Tilman, D.; Wedin, D.; Reich, P.; Tjoelker, M.; Knops, J. 2002. Functional traits, productivity and effects on nitrogen cycling of 33 grassland species. Functional Ecology. 16(5): 563-574. [62178]
9. Curtis, John T. 1959. Savanna. In: Curtis, John T. The vegetation of Wisconsin. Madison, WI: The University of Wisconsin Press: 325-351. [60528]
10. Diboll, Neil. 1986. Mowing as an alternative to spring burning for control of cool season exotic grasses in prairie grass plantings. In: Clambey, Gary K.; Pemble, Richard H., eds. The prairie: past, present and future: Proceedings of the 9th North American Prairie Conference; 1984 July 29 - August 1; Moorhead, MN. Fargo, ND: Tri-College University Center for Environmental Studies: 204-209. [3574]
11. Dirig, Robert. 1994. Historical notes of wild lupine and the Karner blue butterfly at the Albany Pine Bush, New York. In: Andow, David A.; Baker, Richard J.; Lane, Cynthia P., eds. Karner blue butterfly: a symbol of a vanishing landscape. Miscellaneous Publication 84-1994. St. Paul, MN: University of Minnesota, Minnesota Agricultural Experiment Station: 23-36. [63142]
12. Dreesen, David R.; Harrington, John T. 1997. Propagation of native plants for restoration projects in the southwestern U.S.--preliminary investigations. In: Landis, Thomas D.; Thompson, Jan R., tech. coords. National proceedings: forest and conservation nursery associations--1997; Regeneration, reforestation, restoration: The seedling is the key; 1997 August 11-14; Bemidji, MN; 1997 August 19-21; Boise, ID. Gen. Tech. Rep. PNW-GTR-419. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station: 77-88. [29122]
13. Duchesne, Luc C.; Hawkes, Brad C. 2000. Fire in northern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 35-51. [36982]
14. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]
15. Forman, Richard T. T.; Boerner, Ralph E. 1981. Fire frequency and the pine barrens of New Jersey. Bulletin of the Torrey Botanical Club. 108(1): 34-50. [8645]
16. Forrester, Jodi A.; Leopold, Donald J.; Hafner, Sasha D. 2005. Maintaining critical habitat in a heavily managed landscape: effects of power line corridor management on Karner blue butterfly (Lycaeides melissa samuelis) habitat. Restoration Ecology. 13(3): 488-498. [54703]
17. Frissell, Sidney S., Jr. 1968. A fire chronology for Itasca State Park, Minnesota. Minnesota Forestry Research Notes No. 196. St. Paul, MN: University of Minnesota. 2 p. [34527]
18. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]
19. Gleason, Henry A.; Cronquist, Arthur. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York: New York Botanical Garden. 910 p. [20329]
20. Goodban, Anthony G.; Bakowsky, Wasyl D.; Bricker, Bradlay D. 1996. The historical and present extent and floristic composition of prairie and savanna vegetation in the vicinity of Hamilton, Ontario. In: Warwick, Charles, ed. Proceedings of the 15th North American prairie conference; 1996 October 23-26; St. Charles, IL. Bend, OR: The Natural Areas Association: 87-103. [30255]
21. Greenfeld, Lisa Michelle. 1997. Habitat quality and utilization analysis in a spatial context: the case of Lupinus perennis L. and Lycaeides melissa samuelis Nabokov, (Lepidoptera: Lycaenidae). East Lansing, MI: Michigan State University. 67 p. Thesis. [62840]
22. Grigore, Michelle Trudeau; Tramer, Elliot J. 1996. The short-term effect of fire on Lupinus perennis (L.). Natural Areas Journal. 16(1): 41-48. [26757]
23. Gross, Katherine L.; Mittelbach, Gary G.; Reynolds, Heather L. 2005. Grassland invasibility and diversity: responses to nutrients, seed input, and disturbance. Ecology. 86(2): 476-486. [52898]
24. Grundel, Ralph; Pavlovic, Noel B.; Sulzman, Christina L. 1998. Habitat use by the endangered Karner blue butterfly in oak woodlands: the influence of canopy cover. Biological Conservation. 85(1/2): 47-53. [62202]
25. Haack, Robert A. 1993. The endangered Karner blue butterfly (Lepidoptera: Lycaenidae): biology, management considerations, and data gaps. In: Gillespie, Andrew R.; Parker, George R.; Pope, Phillip E., eds. Proceedings, 9th central hardwood forest conference; 1993 March 8-10; West Lafayette, IN. Gen. Tech. Rep. NC-161. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station: 83-100. [27003]
26. Halpern, Stacey L. 2005. Sources and consequences of seed size variation in Lupinus perennis (Fabaceae): adaptive and non-adaptive hypotheses. American Journal of Botany. 92(2): 205-213. [52937]
27. Harrington, John A. 1995. Planning and implementation of a right-of-way native planting for Wisconsin Highway 51. In: Hartnett, David C., ed. Prairie biodiversity: Proceedings, 14th North American prairie conference; 1994 July 12-16; Manhattan, KS. Manhattan, KS: Kansas State University: 175-179. [28255]
28. Heinselman, Miron L. 1970. The natural role of fire in northern conifer forests. In: The role of fire in the Intermountain West: Symposium proceedings; 1970 October 27-29; Missoula, MT. Missoula, MT: Intermountain Fire Research Council: 30-41. In cooperation with: University of Montana, School of Forestry. [15735]
29. Hendrickson, William H. 1972. Perspective on fire and ecosystems in the United States. In: Fire in the environment: Symposium proceedings; 1972 May 1-5; Denver, CO. FS-276. [Washington, DC]: U.S. Department of Agriculture, Forest Service: 29-33. In cooperation with: Fire Services of Canada, Mexico, and the United States; Members of the Fire Management Study Group; North American Forestry Commission; FAO. [17276]
30. Howe, H. F.; Brown, J. S. 2000. Early effects of rodent granivory on experimental forb communities. Ecological Applications. 10(3): 917-924. [62179]
31. Jefferies, R. A.; Bradshaw, A. D.; Putwain, P. D. 1981. Growth, nitrogen accumulation and nitrogen transfer by legume species established on mine spoils. Journal of Applied Ecology. 18(3): 945-956. [62181]
32. Kartesz, John T.; Meacham, Christopher A. 1999. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Available: North Carolina Botanical Garden. In cooperation with: The Nature Conservancy, Natural Resources Conservation Service, and U.S. Fish and Wildlife Service [2001, January 16]. [36715]
33. Kearns, S. Kelly. 1986. A comparison of transplanting times and methods for salvaging prairie forbs and grasses. In: Clambey, Gary K.; Pemble, Richard H., eds. The prairie: past, present and future: Proceedings of the 9th North American Prairie Conference; 1984 July 29 - August 1; Moorhead, MN. Fargo, ND: Tri-College University Center for Environmental Studies: 197-200. [3572]
34. King, Richard S. 2003. Habitat management for the Karner blue butterfly (Lycaeides melissa samuelis): Evaluating the short-term consequences. Ecological Restoration. 21(2): 101-106. [62557]
35. Kucera, Clair L. 1981. Grasslands and fire. In: Mooney, H. A.; Bonnicksen, T. M.; Christensen, N. L.; Lotan, J. E.; Reiners, W. A., tech. coords. Fire regimes and ecosystem properties: Proceedings of the conference; 1978 December 11-15; Honolulu, HI. Gen. Tech. Rep. WO-26. Washington, DC: U.S. Department of Agriculture, Forest Service: 90-111. [4389]
36. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384]
37. Lane, Cynthia P. 1999. Benefits of heterogeneous habitat: oviposition preference and immature performance of Lycaeides melissa samuelis Nabokov (Lepidoptera: Lycaenidae). St. Paul, MN: University of Minnesota. 185 p. Dissertation. [62850]
38. Lawrence, William S. 1994. Karner blue butterfly populations in the Allegan State Game Area, Michigan. In: Andow, David A.; Baker, Richard J.; Lane, Cynthia P., eds. Karner blue butterfly: a symbol of a vanishing landscape. Miscellaneous Publication 84-1994. St. Paul, MN: University of Minnesota, Minnesota Agricultural Experiment Station: 53-62. [63144]
39. Leach, Mark K. 1993. Status and distribution of the Karner blue butterfly at Fort McCoy, Wisconsin: final report on a two-year study. [Madison, WI]: The Nature Conservancy, Wisconsin Chapter. [Prepared for: U.S. Army, Fort McCoy Military Reservation, Natural Resource Management Division, Fort McCoy, WI]. Unpublished report on file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 50 p. [+ appendix]. [62631]
40. Leach, Mark K. 1994. Savanna plant species distributions along gradients of sun-shade and soils, [Online]. In: Fralish, James S.; Anderson, Roger C.; Ebinger, John E.; Szafoni, Robert, eds. Living in the edge: 1994 proceedings of the North American conference on savannas and barrens; 1994 October 15-16; Normal, IL. [Chicago, IL]: U.S. Environmental Protection Agency (Producer). 6 p. Available: http://www.epa.gov/greatlakes/ecopage/upland/oak/oak94/Proceedings/Leach.html [2006, August 22]. [63163]
41. Leach, Mark K.; Givnish, Thomas J. 1999. Gradients in the composition, structure, and diversity of remnant oak savannas in southern Wisconsin. Ecological Monographs. 69(3): 353-374. [52701]
42. Little, Silas. 1998. Fire and plant succession in the New Jersey Pine Barrens. In: Forman, Richard T. T., ed. Pine Barrens: ecosystem and landscape. New Brunswick, NJ: Rutgers University Press: 297-314. [50781]
43. Loope, Walter L. 1991. Interrelationships of fire history, land use history, and landscape pattern within Pictured Rocks National Seashore, Michigan. The Canadian Field-Naturalist. 105(1): 18-28. [5950]
44. Lorimer, Craig G. 1985. The role of fire in the perpetuation of oak forests. In:, Johnson, J. E., ed. Challenges in oak management and utilization. Madison, WI: University of Wisconsin, Cooperative Extension Service: 8-25. [19543]
45. Lorimer, Craig G.; White, Alan S. 2003. Scale and frequency of natural disturbances in the northeastern US: implications for early successional forest habitats and regional age distributions. Forest Ecology and Management. 185(1-2): 41-64. Available: http://www.sciencedirect.com/jfind?query=forest+ecology+and+management. [41377]
46. Mackay, Wayne A.; Davis, Tim D.; Sankhla, Daksha; Riemenschneider, Don E. 1996. Factors influencing seed germination of Lupinus perennis. Journal of Environmental Horticulture. 14(4): 167-169. [62217]
47. Maxwell, Judith Ann. 1998. The conservation of the Karner blue butterfly (Lycaeides melissa samuelis Nabokov): ecological studies on habitat creation and management. Madison, WI: University of Wisconsin. 193 p. Dissertation. [62841]
48. Motzkin, G.; Patterson, W. A., III; Foster, D. R. 1999. A historical perspective on pitch pine - scrub oak communities in the Connecticut Valley of Massachusetts. Ecosystems. 2(3): 255-273. [60894]
49. Nichols, G. E. 1934. The influence of exposure to winter temperatures upon seed germination in various native American plants. Ecology. 15(4): 364-373. [55167]
50. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; Gottfried, Gerald J.; Haase, Sally M.; Harrington, Michael G.; Narog, Marcia G.; Sackett, Stephen S.; Wilson, Ruth C. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-volume 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. [36978]
51. Peloquin, Robert L.; Hiebert, Ronald D. 1999. The effects of black locust (Robinia pseudoacacia L.) on species diversity and composition of black oak savanna/woodland communities. Natural Areas Journal. 19(2): 121-131. [42543]
52. Radford, Albert E.; Ahles, Harry E.; Bell, C. Ritchie. 1968. Manual of the vascular flora of the Carolinas. Chapel Hill, NC: The University of North Carolina Press. 1183 p. [7606]
53. Rainier Seeds, Inc. 2003. Catalog, [Online]. Davenport, WA: Rainer Seeds, Inc., (Producer). Available: http://www.rainerseeds.com [2003, February 14]. [27624]
54. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]
55. Savignano, Dolores A. 1994. The distribution of the Karner blue butterfly in Saratoga County, New York. In: Andow, David A.; Baker, Richard J.; Lane, Cynthia P., eds. Karner blue butterfly: a symbol of a vanishing landscape. Miscellaneous Publication 84-1994. St. Paul, MN: University of Minnesota, Minnesota Agricultural Experiment Station: 73-80. [63145]
56. Schweitzer, Dale F. 1994. Recovery goals and methods for Karner blue butterfly populations. In: Andow, David A.; Baker, Richard J.; Lane, Cynthia P., eds. Karner blue butterfly: a symbol of a vanishing landscape. Miscellaneous Publication 84-1994. St. Paul, MN: University of Minnesota, Minnesota Agricultural Experiment Station: 185-193. [63147]
57. Sharp Brothers Seed Company. 1989. Catalog of wildflowers and forbs. Amarillo, TX: Sharp Brothers Seed Company. 20 p. [18001]
58. Shepherd, Matthew D. 2005. Species Profile: Callophrys irus (Godart), 1824: Frosted elfin (Lycaenidae: Theclinae: Eumaeini), [Online]. In: Shepherd, Matthew D.; Vaughan, D. Mace; Black, Scott Hoffman, eds. Red list of pollinator insects of North America. CD-ROM Version 1 (May 2005). Portland, OR: The Xerces Society for Invertebrate Conservation (Producer). 5 p. Available: http://www.xerces.org/Pollinator_Red_List/Leps/Callophrys_irus.pdf [2006, August 17]. [63162]
59. Shi, X. J.; Michaels, H. J.; Mitchell, R. J. 2005. Effects of self-pollination and maternal resources on reproduction and offspring performance in the wild lupine, Lupinus perennis (Fabaceae). Sexual Plant Reproduction. 18: 55-64. [63128]
60. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]
61. Shuey, John A. 1997. Dancing with fire: ecosystem dynamics, management, and the Karner blue (Lycaeides melissa samuelis Nabokov) (Lycaenidae). Journal of the Lepidopterists' Society. 51(3): 263-269. [62208]
62. Smallidge, Peter J.; Leopold, Donald J.; Allen, Craig M. 1996. Community characteristics and vegetation management of Karner blue butterfly (Lycaeides melissa samuelis) habitats on rights-of-way in east-central New York, USA. Journal of Applied Ecology. 33(6): 1405-1419. [62172]
63. Smith, Mark A.; Turner, Monica G.; Rusch, Donald H. 2002. The effect of military training activity on eastern lupine and the Karner blue butterfly at Fort McCoy, Wisconsin, USA. Environmental Management. 29(1): 102-115. [62210]
64. Sperka, Marie. 1973. Growing wildflowers: A gardener's guide. New York: Harper & Row. 277 p. [10578]
65. Stickney, Peter F. 1989. FEIS postfire regeneration workshop--April 12: Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. 10 p. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. [20090]
66. Stoeckeler, J. H. 1958. Preliminary observations of perennial lupine as a cover crop in field-planting sites and forest nurseries in Wisconsin. Soil Science Society Proceedings. 22(2): 170-173. [62221]
67. Swengel, Ann B. 1993. Observations of Karner blues and the barrens butterfly community in Wisconsin 1987-1993. Report to National Biological Survey and U. S. Fish and Wildlife Service. Baraboo, WI: North American Butterfly Association. 81 p. On file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. [62613]
68. Taft, John B. 1997. Savanna and open-woodland communities. In: Schwartz, Mark W., ed. Conservation in highly fragmented landscapes. New York: Chapman & Hall: 24-54. [51500]
69. U.S. Department of Agriculture, Agricultural Research Service, Animal Disease and Parasite Research Division. 1964. 16 plants poisonous to livestock in the western states. Farmers' Bulletin No. 2106. Washington, DC: U.S. Department of Agriculture. 49 p. [46538]
70. U.S. Department of Agriculture, Natural Resources Conservation Service. 2006. PLANTS database (2006), [Online]. Available: http://plants.usda.gov/. [34262]
71. U.S. Department of the Interior, Fish and Wildlife Service. 2003. Karner blue butterfly recovery plan (Lycaeides melissa samuelis), [Online]. Fort Snelling, MN: U.S. Department of the Interior, Fish and Wildlife Service, Karner Blue Butterfly Recovery Team (Producer). 273 p. Available: http://ecos.fws.gov/docs/recovery_plans/2003/030919.pdf [2006, August 23]. [63164]
72. Vogl, Richard John. 1961. The effects of fire on some upland vegetation types. Madison, WI: University of Wisconsin. 154 p. Dissertation. [52282]
73. Voss, Edward G. 1985. Michigan flora. Part II. Dicots (Saururaceae--Cornaceae). Bull. 59. Bloomfield Hills, MI: Cranbrook Institute of Science; Ann Arbor, MI: University of Michigan Herbarium. 724 p. [11472]
74. Wade, Dale D.; Brock, Brent L.; Brose, Patrick H.; Grace, James B.; Hoch, Greg A.; Patterson, William A., III. 2000. Fire in eastern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 53-96. [36983]
75. Wagner, David L.; Nelson, Michael W.; Schweitzer, Dale F. 2003. Shrubland Lepidoptera of southern New England and southeastern New York: ecology, conservation, and management. Forest Ecology and Management. 185(1-2): 95-112. [61257]
76. Wilcox, Christina A.; Young-Moon, Chun; Choi, Young D. 2005. Redevelopment of black oak (Quercus velutina Lam.) savanna in an abandoned sand mine in Indiana Dunes National Lakeshore, USA. The American Midland Naturalist. 154(1): 11-27. [54842]
77. Williams, Andrew H.; Young, Daniel K. 1999. Attraction of Pedilus lugubris (Coleoptera: Pyrochroidae) to Epicauta murina and Epicauta fabricii (Coleoptera: Meloidae) and new food plant records for Epicauta spp. The Great Lakes Entomologist. 32(1/2): 97-99. [62219]
78. Wofford, B. Eugene. 1989. Guide to the vascular plants of the Blue Ridge. Athens, GA: The University of Georgia Press. 384 p. [12908]
79. Wolf, Joy. 2004. A 200-year fire history in a remnant oak savanna in southeastern Wisconsin. The American Midland Naturalist. 152(2): 201-213. [55434]
80. Woodcock, E. F. 1925. Observations on the poisonous plants of Michigan. American Journal of Botany. 12(2): 116-131. [62183]
81. Wunderlin, Richard P. 1998. Guide to the vascular plants of Florida. Gainesville, FL: University Press of Florida. 806 p. [28655]
82. Zaremba, Robert E.; Pickering, Michael. 1994. Lupine ecology and management in New York State. In: Andow, David A.; Baker, Richard J.; Lane, Cynthia P., eds. Karner blue butterfly: a symbol of a vanishing landscape. Miscellaneous Publication 84-1994. St. Paul, MN: University of Minnesota, Minnesota Agricultural Experiment Station: 87-93. [63146]
83. Zhang, Jianhua; Maun, M. A. 1991. Effects of partial removal of seed reserves on some aspects of seedling ecology of seven dune species. Canadian Journal of Botany. 69(7): 1457-1462. [35459]

FEIS Home Page