Mountain Lady's-slipper (Cypripedium montanum Douglas ex Lindley)
J. Seevers and F. Lang
TABLE OF CONTENTS
Species: Cypripedium montanum Douglas ex Lindley (Mountain lady's-slipper)
Taxonomic group: Vascular Plants
ROD Components: 1, 2 (West Cascades)
Other Management Status: Cypripedium montanum is a California Native Plant Society and Oregon Natural Heritage Program category 4 (watch list) species and is considered a Monitor species by the Washington Natural Heritage Program. Cypripedium montanum is a Survey and Manage species for the United States Forest Service (USFS) and the Bureau of Land Management (BLM) in Oregon.
Range: The geographic range for C. montanum, within the range of the northern spotted owl, is the Cascade Range of Washington and Oregon, the western interior valleys of Oregon, and the Klamath Mountains of southern Oregon and northern California. These Management Recommendation apply only to C. montanum in the Western Cascades as defined in the Northwest Forest Plan (USDA and USDI 1994b). The only known populations that are covered here are in Douglas, Jackson, Lane, and Marion Counties, Oregon.
Specific Habitat: Site report data indicate that C. montanum grows at elevations from 495-2146 m (1500-6500 ft.). However, most sites in the Western Cascades occur between 825-1321 m (2500-4000 ft.). Aspect is mainly northerly, occurring on slopes of 25 to 50 percent. Cypripedium montanum grows on a wide variety of substrates in wooded communities with 60-80 percent canopy closure in mixed conifer and mixed evergreen/oak woodland plant communities. Western Cascade communities most often consist of Douglas-fir (Pseudotsuga menziesii) in combination with one or more of the following: madrone (Arbutus menziesii), sugar pine (Pinus lambertiana), or ponderosa pine (Pinus ponderosa). Other associates include grand fir (Abies grandis), Pacific yew (Taxus brevifolia), incense cedar (Calocedrus decurrens), and California black oak (Quercus kelloggii).
Life History: Cypripedium montanum is a long-lived, rhizomatous, autotrophic orchid that has a symbiotic relationship with a soil fungus that begins with germination and may last until the plant dies. Asexual reproduction occurs by division of the rhizome. The species is primarily an outcrosser pollinated by small bees (Luer 1975). Dissemination is by many thousands of nearly microscopic seeds carried by wind or water, or perhaps after consumption by large herbivores (deer, elk, cattle). Fruit production does not appear to be high, 61 percent in one study (Coleman 1995). Plants may remain vegetative for more than 12 years before flowering.
Threats: Major threats to the species are loss of small, isolated populations due to activities such as timber harvest, road and trail construction, creation of recreation sites, harvesting forest products that disturbs litter and soil (herbal medicine, mushroom collecting), or fire. Threats include activities that alter soil moisture or temperature regimes, or actions that disturb the soil and litter layer, or decrease overstory canopy cover to <60 percent. Small, declining populations may indicate that specific management requirements are not being met (USDA and USDI 1994a).
- Mitigate disturbance to known sites to maintain current microclimate conditions of the habitat area until basic habitat and life history requirements are understood.
- Avoid direct mechanical damage to plants, or changes in soil moisture and temperature, or the nature of the duff layer.
- Maintain overstory canopy coverage at 60 percent or more to prevent increased sunlight to the site.
- Specific ecological requirements of C. montanum for germination, establishment, and growth.
- Role fire plays in the ecology of this species.
- Identification of pollinators and their habitat requirements.
- The identification of fungal associates, their habitat requirements, and the role they play in the life history of the species.
- Determination of the health, viability, and stability of known populations. Revisit Cypripedium sites reported more than 5 years ago to determine if the habitat is intact and if plants are present.
- Develop research and monitoring protocols in Adaptive Management Areas (USDA and USDI 1994a).
I. NATURAL HISTORY
A. Taxonomic/Nomenclatural History
Scientific name: Cypripedium montanum Douglas ex Lindley
Cypripedium montanum was described by Lindley based on a specimen collected by David Douglas in the Blue Mountains of Washington State (Cribb 1997) "on high grounds, among low brushwood," June 28 or 29, 1826 (Douglas 1959). This was the only Cypripedium collected by Douglas in the range of C. montanum.
Common name: Mountain Lady's-slipper, Moccasin slipper
Genus: Cypripedium contains about 45 species in the Northern Hemisphere. Eleven are native to North America (Cribb 1997).
Citation: Cypripedium montanum Douglas ex Lindley, Genera and Species of Orchidaceous Plants: 528. 1840.
Synonyms: Cypripedium occidentale Watson, Proceedings American Academy of Arts and Sciences 11:147. 1876.
B. Species Description (Abrams 1940; Correll 1950; Peck 1961; Munz 1968; Hitchcock et al. 1969; Luer 1975; Hickman 1993; Sheviak forthcoming).
The following description is from Hitchcock et al. (1969):
Stem (1) 2-6 dm. tall, glandular-pubescent, leafy throughout; leaves broadly elliptic to ovate-elliptic, 5-15 cm. long, up to 7 cm. broad, somewhat glandular-pubescent, sessile and usually sheathing; flowers usually 2, but occasionally 1 or 3, each subtended and usually exceeded by an erect leaf-like bract; sepals and petals light to rather deep brownish-purple, narrowly to broadly lanceolate, usually more or less twisted and wavy, the upper sepal mostly (3) 4-5 cm. long, the lower pair not quite so long, fused except for the slender terminal tooth like lobes; petals slightly longer than the sepals (up to 6.5 cm. long); lip obovoid and strongly pouched, 2-3 cm. long, dull white to purplish-tinged, usually purplish-veined; staminodium up to 10 (12) mm. long, yellowish-white and usually purplish-spotted, ovate, only rarely auriculate at the base.
Cribb (1997) and Doherty (1997) have summarized what is known about the genus Cypripedium. Their reviews cover morphology, life history, cytology, phylogenetic relationships, biogeography, and ecology including mycorrhizal associations, uses, culture and propagation, artificial hybridization, and taxonomy. Managers should consult these publications for details.
According to Sheviak (forthcoming; 1992), C. montanum is one of 4 North American members of the circumboreal C. calceolus complex. Other North American species include C. candidum, C. kentuckiense, and C. parviflorum (formerly considered C. calceolus) with 3 varieties, parviflorum, pubescens, and makasin. Cribb (1997) does not recognize Sheviak's var makasin. Sheviak restricts C. calceolus to Eurasian members of the complex. North America plants, formerly included in C. calceolus by earlier workers, are now considered to be varieties of C. parviflorum. We will follow Sheviak's treatment here.
Sheviak (pers comm) feels the taxonomy of the complex in the western United States, especially C. parviflorum (or C. calceolus in older treatments), has been totally incorrect. Hitchcock et al. (1969) note (using the old nomenclature) that there is little morphological dissimilarity between C. calceolus var. parviflorum (NOT var. pubescens as stated by Hitchcock) and C. montanum, except for the yellow instead of white, pouch-like lower lip of similar size. In spite of taxonomic confusion, some of what is known of the biology of other members of the C. calceolus complex might be cautiously applied to C. montanum.
2. Reproductive Biology
The life history of C. montanum has not been well studied. Studies of population genetics, growth patterns, and reproductive biology done for C. fasciculatum have not been done for C. montanum. However, studies of other Cypripedium species and other genera can provide some insight into the reproductive biology of this species (Luer 1975; Case 1987; Ballard 1990; Hadley 1990; Sheviak 1992; Harrod and Knecht 1994; Knecht 1996; Proctor et al. 1996).
Several orchids, including C. calceolus, take 13 to 16 years to begin to flower (Harper and White 1974; Wells 1981). Harrod (1993) found that small, nonflowering C. fasciculatum plants can be at least 12 years old.
Proctor et al. (1996) describe pollination in C. calceolus in Europe. The bright yellow bowl-like lower lip and fruity scent (without reward) attract a variety of bees. Sheviak (1992) describes the fragrance of C. parviflorum var. pubescens as ranging from a clear rose scent to rather musty, not particularly "sweet" and that of C. parviflorum var. parviflorum as "intensely sweet." Urban (1997) reports that C. montanum has a "heavenly fragrance . . . (a light vanilla scent)"; Coleman (1995) reports ". . . a pleasant fragrance, slightly reminiscent of anise."
Proctor et al. (1992) describe pollination in the European C. calceolus. There is a large opening into the bowl of the pouch-like lower lip. The column (fused stamens, stigma, and style) extends into the opening. In the bowl below, hairs beneath the column lead to a pair of smaller openings on either side of the column.
Larger bees enter the pouch, then, being large enough, exit via the entrance, unrewarded. In the case of C. montanum, insects like large bumble bees may be excluded from the pouch by the size of the opening (Luer 1975). In C. calceolus and probably C. montanum, small bees enter the pouch, but cannot escape because of their size and the size and shape of the pouch lip. They wing buzz about in undirected escape efforts but soon begin to "prise" about under the stigma. In the process they slightly depress the elastic lip near the column base, creating a passageway toward the openings at the base of the column. As the small bees slip by, aided by the backward pointing hairs on the floor of the pouch, they pick up sticky pollen (no pollinia in Cypripedium) on the way out as they squeeze past the anthers. Self-fertilization rarely occurs except when little bees make several escape attempts and encounter the stigma when covered with pollen or when large bees bumble about and accidentally pick up pollen and deposit it on the stigma. Cross fertilization is the rule.
According to Dressler (1993) no member of the subfamily to which Cypripedium belongs rewards the insect visitor. Proctor et al. (1992) state that C. calceolus does not provide a nectar reward for pollinators. However, Luer (1969) thinks that once in the pouch of C. montanum, "the bee probably enjoys the minute droplets of nectar secreted on the hairs within."
Luer (1975) observed several small black bees, tentatively identified from a photograph, as a banded species of Lasioglossum pollinating a C. montanum flower. The bees entered via the lip and left by crawling out an exit under an anther. A large bumblebee (Bombus sp.) attempted entry, but departed when it found it could not squeeze into the labellum because of the small size of the opening of the pouch.
Although the Cypripedium flower is designed as a "one-way street" that ensures that the insect passes the stigma on the way out, cross fertilization, or fertilization of any kind, may not always be successful.
Ballard (1990) noted <10 percent natural fruit set in C. parviflorum from New Hampshire and that hand pollinated flowers did as poorly. Capsules with seeds from the wild and from the garden had 20 to 30 percent aborted or shriveled proembryos.
Coleman (1995) followed fruit production in 3 large California populations of C. montanum over a 4-year period in an effort to establish pollinator success. He found that 792 flowers produced 483 capsules, a 61 percent fruit set. He did not attempt to evaluate seed viability.
Orchid fruits (capsules) produce nearly microscopic seeds in large numbers. Harrod and Knecht (1994) found an average of 3874 seeds per fruit in C. fasciculatum. Correll (1950) reports an estimated 10,000 seeds per capsule in the yellow lady-slipper C. parviflorum.
Orchid seeds are usually waterproof at dispersion and are carried by water or air (Case 1987; Harrod and Everett 1993).
Steele (1996) reported great difficulty getting C. montanum seeds to germinate using methods without the fungal partner (asymbiotic). He found germination to be about 0.1 percent. This problem needs to be solved if asymbiotic methods can be used as a source of plants to reestablish extirpated populations.
Cypripedium species are dependent on mycorrhizal associations for several months or years before above-ground growth begins (Harper and White 1974).
Spring growth of orchids arises from overwintering buds produced the preceding growing season. Unlike most other plants, however, if new spring growth is destroyed by late frost, foraging animals, disease, accident, or ill-advised management practices, an orchid cannot replace the lost tissues until the following year (Sheviak 1990). Although dormant buds may be present, they will not initiate growth. The root system will remain, and a new bud may form, or a dormant bud enlarge, but the plant will suffer a major setback, and it may die (Sheviak 1990). Cypripedium plants that lose their growth before midsummer will commonly appear the next year but will not bloom (Whitlow 1983). Depending on how severely depleted their energy reserves are, they may require 2 or more subsequent vegetative seasons before blooming (Whitlow 1983; Case 1987).
Information about arrangement of roots, rhizomes, pronating buds, and growth patterns available for C. fasciculatum (Knecht, pers comm; Harrod 1994b) is not available for C. montanum.
As a rhizomatous perennial, C. montanum probably reproduces asexually by rhizome fragmentation. According to Summerhayes (1951) cypripediums and other rhizomatous orchids reproduce when older parts of a branching rhizome die back leaving younger, still living branches as separate plants. Although this is probably the case for C. montanum, as it is for many other plant species with branching rhizomes, it has not been clearly demonstrated.
What is known of the ecology of C. montanum is based largely on information from cursory site reports. Detailed C. montanum habitat studies have begun in the last few years. Its fire history and interrelationships with other plants and animals are now being undertaken. Results of studies across the species' range are still unknown. Habitats vary in moisture regimes, substrates, and plant associates (Kagan 1990; Seevers, unpubl repts). Sites in the region seem to have in common a canopy coverage of 60 percent or more, including an association with conifers, often Douglas-fir.
Harrod et al. (1997) state, "Historically, suitable habitat conditions for C. fasciculatum likely shifted across the landscape over time or were found in fire refugia (Camp 1995)." Greenlee (1997) feels that C. fasciculatum in Region 1, USFS, is distributed as a metapopulation linked by recurrent extinctions and recolonizations over time. In this view, Cypripedium fasciculatum populations moved across the landscape as suitable habitat appeared and disappeared as disturbances and successional changes occurred over time. Although the effects of disturbance (type, frequency, intensity, and durations) on C. montanum and associated habitat have not been well studied, this model may apply to C. montanum as well. The role of fire needs to be investigated before it is reintroduced into C. montanum's habitat.
Urban (1997) attributes the loss of upper portions, flowers and fruits, of C. montanum on the Umatilla NF in Oregon to predation by elk and deer. He points out that this is not damaging to the orchid because lower plant parts are left intact and functioning, storing photosynthates for the next flowering period. Human predation, however, is harmful. Plants are either plucked off at ground level, leaves and all, or are dug up for garden transplant.
Orchid seeds, unlike other flowering plants, lack a differentiated embryo, endosperm and protective seed coat. A lacy, net-like outer seed coat (the testa) covers an inner undifferentiated mass of cells (the proembryo). Orchid seeds require the presence of a fungus, usually a species of Rhizoctonia, before the seeds will germinate (Arditti 1967). The embryo grows and the plant becomes established (Wells 1981; Sheviak 1990). Rhizoctonia hyphae invade and digest the seed coat and outer layers of the cell mass. Digestion of these outer layers produces sugars, which the inner cells utilize to grow and develop (Case 1987). There is little question that Rhizoctonia is essential for germination and establishment of Cypripedium species in the wild, including C. montanum. Establishment of new populations requires suitable conditions for the fungus. What these habitat conditions are is not known, but can be presumed to be moist and shady with adequate organic material to support growth of the heterotrophic fungus.
Some native orchids are completely mycotrophic when immature, spending several years in a dependent, subterranean condition, relying on the fungus for water and nutrition before sufficient growth occurs and stored food accumulates for leaf production (Case 1987). Only after adequate food storage does the plant's first green shoot appear above ground.
The relationship between fungus and orchid is not well understood, but it appears that it is an interaction easily upset (Sheviak 1990). There is some evidence that once autotrophic, the mycorrhizal relationship might not be as beneficial as normally assumed. Uninfected cortical cells contain starch; infected cells do not (Stoutamire 1991). Green protocorms supplied with all nutrients grow better with a symbiotic mycorrhizal fungus than without the nutritional supplement (Hadley 1989). Whitlow (1983) feels that once Cypripedium plants germinate and grow to the photosynthetic state the role of the fungus literally ceases.
C. Range, Known Sites
Cypripedium montanum ranges from southern Alaska, British Columbia, and western Alberta south to Montana, Idaho, Wyoming, and northern California. This species is no longer extant on Vancouver Island (Clark 1976), nor is it known from the Olympic Peninsula. It possibly is not present west of the Cascade crest in Washington, except along the Columbia River Gorge (Hitchcock et al. 1969). It is known in Asotin, Columbia, Chelan, Ferry, Douglas, Garfield, Klickitat, Kittitas, Okanogan, Pend Oreille, Spokane, Stevens, Whitman, and Yakima Counties in Washington (Gamon 1997). These known sites are outside the Western Cascade area covered by this document. Piper (1906) reported C. montanum only from east of the Cascade crest and not from western Washington. In Oregon it has been documented from the following counties: Baker, Crook, Deschutes, Douglas, Grant, Hood River, Jackson, Jefferson, Josephine, Klamath, Lake, Lane, Marion, Umatilla, Union, Wallowa, Wasco, and Wheeler (USDA and USDI 1994a). In California, C. montanum is known in Del Norte, Humboldt, Mendocino, Modoc, Mariposa, Plumas, San Francisco, San Mateo, Sierra, Siskiyou, Tehema, Trinity, and Tuolumne Counties (CalFlora Database 1997). These Management Recommendations apply to the range of the species in the Western Cascades within the range of the northern spotted owl (USDA and USDI 1994b).
Unspecified anecdotal reports and old herbarium records indicate that there may have been more C. montanum populations in western Oregon than there are today (Siddall et al. 1979; Meinke 1982). The accuracy of these reports should be checked by confirming the health and vigor of populations known from old herbarium records. Seventy percent of populations on BLM and USFS sighting reports contained 10 or fewer individuals; 0.6 percent have more than 30 individuals. Presently, C. montanum populations in western Oregon are small and scattered; 20 are extant in the West Cascades and represent <70 plants.
It is difficult to determine if these small populations are the norm for C. montanum (supported by Greenlee (1997), or if the small size is the result of alterations in habitat conditions. Because orchid populations may be typically small, a decline in number of populations may be more significant than the small size of populations. Small populations may reflect the slow establishment and growth rate of this species. However, small, declining populations may also indicate that specific management requirements are not being met (USDA and USDI 1994a).
D. Habitat Characteristics and Species Abundance
C. montanum occurs in a broad range of habitats. Coleman (1989, 1995) reports that C. montanum grows in both moist and dry conditions in California. A typical moist habitat is along streams under or near creek dogwood (Cornus stolonifera), western azalea (Rhododendron occidentale), with queen's cup bead-lily (Clintonia uniflora) as a ground cover. A typical dry site would be a dry hillside in coniferous forests with few to no associated understory species. The Jepson Flora (Hickman 1993) describes C. montanum habitat as moist areas, dry slopes, mixed evergreen or coniferous forests. Peck (1961) cites moist open woods, and Hitchcock et al. (1969) cite dry to fairly moist, open to shrub- or forest-covered valleys or mountainsides. Sheviak (forthcoming) reports C. montanum from mesic to dry (rarely wet) coniferous, deciduous, and broadleaf evergreen forests, openings, and thickets, and around shrubs on open slopes. These reports make it clear that C. montanum grows in a variety of habitats.
Sighting reports indicate that many known locations occurred within timber sale units because these are the areas surveyed. These reports do not state if mitigation measures were part of the timber sale plan. It is not known if the plants at these sites survived the timber harvest. Coleman's example (1995) indicates that the present total number of extant sites may be much lower than site reports indicate.
Data from sighting reports (Seevers, unpubl repts) indicate that C. montanum grows at elevations ranging from 495-2145 m (1500-6500 ft.); however, the majority of the sites occur between 825-1320 m (2500-4000 ft.). Aspect is mainly northerly, occurring on slopes of 25 to 50 percent. Location data indicates an ability to grow on a wide variety of substrates, including ultramafic and limestone. Cypripedium montanum usually occurs in wooded communities with 60-80 percent canopy closure. In the southern part of the range, communities most often consist of Douglas-fir (Pseudotsuga menziesii) in combination with one or more of the following: canyon live oak (Lithocarpus densiflora), madrone (Arbutus menziesii), sugar pine (Pinus lambertiana), or ponderosa pine (Pinus ponderosa). In the northern part of the range, plant communities consist of ponderosa pine, lodgepole pine, and Douglas-fir .
Because C. montanum and C. fasciculatum often occur sympatrically, Harrod's studies might also apply to C. montanum. Harrod et al. (1996) felt that mid- to late-successional forest communities may be necessary for C. fasciculatum. Other studies (Harrod 1994) indicate that optimum habitat conditions for C. fasciculatum are not found in early successional communities, possibly because the required fungal symbiont(s) is (are) only present in mid- to late-successional forest communities.
Although little is known about the species' ecological requirements in the field, indirect evidence is available from those who grow cypripediums in culture. Whitlow (1983) reports that information available to him indicates that C. montanum and C. fasciculatum require acidic, sandy soil, and fairly dry conditions. Doherty (1997) finds C. montanum is particularly sensitive to excessive moisture especially in the winter. He also mentions the importance of companion plantings in pots to help keep C. montanum roots cool.
Although soil and topography have a definite influence upon terrestrial orchid species, there is little doubt that temperature and moisture are the most essential factors that control distribution and survival of all orchids (Correll 1950). The nature of the Cypripedium habitat (moisture, temperature, light, and organic matter) may be determined by the requirements of the fungus. Doherty (1997) reports that there is no question that the developing orchid depends on its fungal symbiont for survival. Once the orchid reaches maturity and becomes autotrophic, the degree of dependence may change. The orchid will not grow successfully except where the fungus will grow, at least in its early stages of development.
II. CURRENT SPECIES SITUATION
A. Why Species Is Listed Under Survey and Manage Standards and Guidelines
Cypripedium montanum, and the habitat on which this species depends, was found to be at high risk for extirpation in the Western Cascades under all alternatives of the Forest Plan. Habitat fragmentation and alteration, fire suppression, trampling by livestock, game animals and humans, and collection of plants for horticultural and medicinal purposes have contributed to population declines. Losses due to timber harvest have been documented in Oregon (Urban 1997). Coleman (1995) cites an example from Siskiyou County, California where a prelogging survey found 2 populations of 260 and 300 C. montanum plants in an area scheduled to be clear-cut. A search of the area several years after timber harvest revealed that only 5 plants survived at the edge of the clear-cut.
Evidence of the role of fire in C. montanum ecology is mixed. Urban (1997) documented strong recolonization by C. montanum of 30-year old plantations with site preparation that used fire. The first response was a rapid increase of snowbrush (Ceanothus velutinus), with an immediate abundant and vigorous reappearance of C. montanum. Based on what we know of length of time for Cypripedium seedling establishment, the immediate reappearance of plants in the burned area suggested that the original plants were not destroyed by the fire.
In California (Nelson pers comm; Pappalardo 1997) a population of 50 plants was discovered in 1981 in a stand of Douglas-fir with 10-20 inch Diameter at Breast Height (DBH). The stand burned in 1987 and was immediately replaced by an overstory of deerbrush (Ceanothus intergerrimus). Except for one clump of 3 stems of C. montanum that survived the 1987 fire, there was no sign of the original population of 50. Conflicting reports of the impact of fire on C. montanum populations indicate that generalizations on the use of fire for management should be delayed until more evidence is available.
Only through protection of all known sites and implementation of specific management guidelines, will risk to C. montanum be reduced (USDA and USDI 1994a).
B. Major Habitat and Viability Considerations
Certain features of C. montanum's biology and ecology have important habitat and viability implications. Populations of C. montanum tend to be small and scattered, which makes them vulnerable to extirpation. Small populations are much more vulnerable to extinction from human and natural causes than are larger populations. Small populations are more likely to succumb to natural catastrophes such as wildfire, floods, landslides, drought, and loss of pollinating insects or human perturbation such as collection, timber harvest, road building, and grazing. Smaller populations are at greater risk of extirpation through management activities occurring near or within their habitat. Diminishing populations are likely to suffer reduced genetic variability and be less able to adapt to changing environmental conditions (Given 1994). Maintaining a minimum effective population size for each C. montanum is essential for species survival. Given (1994) gives 500 individuals as an acceptable goal for a minimum effective population size for vascular plant species. If a population drops to 50-100 individuals there is cause for concern. Small populations, typical of many orchid species, may reflect the slow establishment and growth rate of C. montanum, or they may indicate that specific management requirements are not being met (USDA and USDI 1994b).
Historical ecosystem processes, such as fire, that are thought to have provided habitat for C. montanum have been altered. The FSEIS (USDA and USDI 1994a) suggests that fire suppression is an important factor in the overall decline of C. montanum. Greater fire frequency east of the Cascades may correlate with the greater number of extant populations in that area (USDA and USDI 1994a), because C. montanum seems to persist in areas that have been burned.
Recent research on C. fasciculatum (Harrod et al. 1997) appears to contradict the belief that Cypripedium species are fire dependent. Harrod et al. (1997) found that C. fasciculatum is a fire-intolerant species and that management of this species probably should not include prescribed fire. The effects of disturbance (type, frequency, intensity, and duration) on C. montanum and associated habitat have not been well studied.
C. Threats to the Species
Physical disturbance of microhabitat (population sites). Loss or drying of the duff layer and of larger down woody debris may affect mycorrhizal symbionts. Although the relationship is not fully understood, symbionts may be necessary for adult plants as well as for establishment of seedlings. Site disturbance may occur with timber harvesting activities, raking for Matsutake mushrooms, road building activities, recreational activities, wildfire, prescribed burning, and trampling by livestock, game animals, and humans. Harrod (1994b) and Knecht (1996) found that activity that exposes or damages the rhizome appears to kill the plant. Physical disturbance of the site may affect the mycorrhizal fungus. Stoutamire (1991) reports that adventitious roots of C. candidum are particularly sensitive to disturbance. Damaged roots are slow to repair and are replaced slowly from the most recently produced rhizome sections.
Adverse edge-effects alter interior habitat conditions and microclimate due to fragmentation of habitat. Edge effects are of particular importance in Pacific Northwest forests because much of the landscape is fragmented by timber harvest practices. Chen et al. (1995) quantified distances of edge influence within forests for several microclimatic variables and assessed the influence on edge effects of edge exposure orientation, time of day, and management of organisms and conditions that appeared to be associated with interior conditions of old-growth forest, including decaying litter and woody debris. The depth of the edge influence into the forest of these microclimatic variables differed with edge orientation, time of day, and local weather. Edge effects were strongest on southwest-facing edges, in the early afternoon, and under partially clear, hot and windy weather conditions. Small forest patches can be essentially "all edge" (Olsen et al. 1995; Chen et al. 1995).
Structural changes in forested areas from edge effects and canopy removal and associated microclimatic alterations may make sites uninhabitable for forest adapted C. montanum populations.
Decrease in pollinator abundance. Not all C. montanum pollinators are known. Pollinators and their habitats could be impacted by unknown factors.
Collection of plants. Collection from the wild for horticultural or medicinal purposes continues to be a threat, and collection for scientific use could pose a threat. The severity of these threats depends on the size of the population and the intensity and method of collection. In 1995 the World Wildlife Fund rated lady-slipper orchids (Cypripedium spp.) among the world's 10 most wanted plants or animals, threatened by illegal and unsustainable trade.
D. Distribution Relative to Land Allocations
Table 1. Known ( as of 1994) C. montanum sites
within the range of the Northwest Forest Plan by land allocation.
Percent of Populations
|Adaptive Management Areas (AMA)
|Administratively withdrawn/Late-Successional Reserves (LSR)
III. MANAGEMENT GOALS AND OBJECTIVES
A. Management Goals for the Taxon
Management goal for C. montanum is to assist in maintaining species viability throughout its range in the Western Cascades.
B. Specific Objectives
- Maintain or restore ecological conditions at known sites such as hydrologic, temperature, and light regimes.
- Maintain viable populations of C. montanum within occupied habitat, including potential mycorrhizal associations.
- Mitigate to avoid habitat reduction and population fragmentation.
- Maintain coarse, woody material in C. montanum habitat.
- Maintain undisturbed duff and soil layers to protect mycorrhizal networks and C. montanum
IV. HABITAT MANAGEMENT
A. Lessons from History
Observation of extant sites provides us with information on management of C. montanum. Some of these observations are:
- Anecdotal reports (Siddall et al. 1979; Meinke 1982) and herbarium records suggest the species was more common in the past. Past forest management activities have contributed to declines in population numbers (Urban 1997).
- Survival after timber harvest appears to be poor (Coleman 1995).
- Conflicting evidence of the effect of fire on population survival. Urban (1997) found the immediate and vigorous reappearence of plants after fire. Nelson (pers comm) and Pappalardo (1997) found no survival after fire.
- Urban (1997) followed the health of C. montanum populations on the Umatilla NF where the overstory had been removed. He reported that the species declined over 5 years. During that time plants became less robust and turned from "bronze" to a light yellow.
B. Identification of Habitat Areas for Management
The habitat area identified in the ROD for management is federal lands in the Western Cascades (USDA and USDI 1994c).
C. Management Within Habitat Areas
- Maintain or restore habitat conditions in areas with populations of C. montanum.
- Maintain canopy closure at 60 percent or greater (USDA and USDI 1994a).
- Maintain down logs, snags, and duff layer within the habitat area for soil moisture and mycorrhizal associates. Provide for future recruitment of coarse woody debris.
- Avoid activities that alter soil, duff, down wood, and the mycorrhizal community in the habitat area.
- Maintain/secure known sites from prescribed burns.
- Manage population sites to include an area large enough to maintain current habitat and associated microclimate, primarily temperature and moisture. The size should be determined by a field visit and should consider factors such as canopy cover, slope, aspect, topographic position, vegetation structure (growth form, stratification, and coverage), and species composition (Chen et al. 1995; Harris 1984).
- Given the long life-span of individuals, manage C. montanum and associated communities to be responsive to short-term (wildfire, soil disturbance) and long-term (ecological succession) environmental changes and maintain the species evolutionary potential.
- Manage for biological (mycorrhizae and pollinators) and ecological (soil temperature, moisture, and organic matter) requirements at each life stage. Each life stage may require specific mitigation. Ensure that indiscriminate insecticide spraying does not affect the populations of bees or other insects this species depends on for pollination.
D. Other Management Issues and Considerations
- Evaluate late successional and old-growth stands within watersheds in the West Cascades to determine their contribution as future habitat.
- Develop a conservation strategy to conserve the best populations to maintain viability of the species. As more information becomes available, management within some sites may be possible.
- Consider restoration activities to attempt reintroduction into historic locations with potential habitat.
V. RESEARCH, INVENTORY, AND MONITORING NEEDS
The objective of this section is to identify opportunities for additional information which could contribute to more effective species management. The content of this section has not been prioritized or reviewed as to how important the particular items are for species management. While the inventory, research, and monitoring identified below are not required, these recommendations should be addressed by a regional coordinating body at the Northwest Forest Plan level.
A. Data Gaps and Information Needs
- Determine the total number of extant populations within the West Cascades . Site visits should be made to determine stability and viability of populations reported over the last 20 years.
- Revisit C. montanum sites in timber sale units. These sites need to be revisited to collect data on populations trends that have occurred since the last visit. Specific information on stand age, fire history, duff layer, coarse woody material, snags, percent canopy, plant association, and abiotic factors should be collected.
- Inventory reserve areas (Research Natural Areas, Areas of Critical Environmental Concern, Late-Successional Reserves, Wilderness Areas).
- Determine ecological requirements for C. montanum seed germination and establishment needs.
- Determine the role of fire (wildfire and controlled burns) C. montanum habitat needs.
- Identify pollinators and their habitat requirements to determined if this is the limiting factor in fruit production.
- Identify fungal associates and their habitat requirements to determine if this is the limiting factor in population size and recruitment.
- Identify specific ranges of site characteristics for reproductively successful populations.
- Determine effects of kinds and intensities of actions on rhizome survival, growth and health.
- Determine the minimum effective population size for C. montanum.
B. Research Questions
Specific research questions:
- What are the edge effects on C. montanum habitat temperature, light, and moisture? How do they affect the plant?
- What role does disturbance ecology, including fire, play in germination, establishment, and long-term survival of this species (USDA and USDI 1994a)?
- What stand characteristics are necessary for this species' establishment?
- What are the biotic and abiotic requirements for each stage of C. montanum's life history?
- What is the role of coarse woody debris at sites? Is its role a relationship with the mycorrhiza at the site, or is it retention of moisture on the site (or is it both)?
- What mycorrhizal fungus is associated with C. montanum, and what are its habitat requirements? Is this association species-specific and codependent?
- What are the necessary biological and ecological requirements at each life stage of C. montanum?
- What are the C. montanum pollinators, and what are their habitat requirements?
- What is the role of ungulates and other mammals in the ecology of this species (if any)?
- What are the factor(s) that trigger flowering and flowering periodicity?
- How many years does/can C. montanum remain vegetative underground?
- What specific habitat characteristics are necessary to maintain populations? What essential habitat features must be maintained in forest corridors and matrix to ensure species survival?
- Are remnant late-successional and old-growth forest parcels necessary for establishment of new populations?
- What are the effects of disturbance, isolation, and size on population genetics?
- What are the growth characteristics of the species: rhizome depth, branching, bud production?
C. Monitoring Needs and Recommendations
Monitor demographically to determine patterns of recruitment (births and immigration), mortality (deaths), survivorship, longevity, and population trends (Wells 1981).
Monitor to determine if design features have been implemented and they are effective.
Determine and monitor microclimate conditions to see if they are being maintained at optimum levels at habitat sites.
Monitor long-term effects of particular types of management practices to determine the impact of action on survival. Longevity of C. montanum plants will require long- term (>20 yrs) to determine vigor, reproduction, and survival of the populations.
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