Management Recommendations
for
Clustered Lady Slipper Orchid
(Cypripedium fasciculatum Kellogg ex S. Watson)

v. 2.0

by

J. Seevers and F. Lang

December 1998

TABLE OF CONTENTS

EXECUTIVE SUMMARY

Species: Cypripedium fasciculatum Kellogg ex S. Watson (Clustered Lady Slipper/Brownie Slipper Orchid)

Taxonomic group: Vascular Plants

ROD Components: 1, 2 (Klamath Province)

Other Management Status: The Bureau of Land Management (BLM) lists Cypripedium fasciculatum as a Bureau Sensitive species in Oregon, Washington, and California. It is listed as a Sensitive species by the U.S. Forest Service, Region Six, as a threatened species in Washington State, and as threatened throughout its range by the Oregon Natural Heritage Program. Cypripedium fasciculatum is a California Native Plant Society category 4 (watch list) species.

Range: Management recommendations, as directed by the Record of Decision (ROD) for the Northwest Forest Plan only pertain to sites in the Klamath Province. The total geographic range for C. fasciculatum, within the region covered by the Northwest Forest Plan, is the Cascade Range of Washington, Oregon, and California, the western interior valleys of Oregon, and the Klamath Mountains of southern Oregon and northern California. Populations in Idaho, Montana, Wyoming, Utah, and Colorado are not covered by this document.

Specific Habitat: Cypripedium fasciculatum occurs in a variety of habitats throughout the Klamath Mountains. Within these habitats, there is great diversity in soils, elevation, aspect, and plant communities. There is no restriction to parent material. Sites vary from dry to damp, rocky to loamy, and elevations range from 330-1745 m (1000-5300 ft). Populations are found in areas with 60 to 100 percent shade provided by various plant communities including mixed evergreen, mixed conifer, Douglas-fir, pine, and black oak forest. The aspect is mainly northerly. Sites occur in several different plant associations within the Douglas-fir series.

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 1978). While it is known that a broad range of temperature and moisture regimes occurs throughout its range, extremes have not been recorded.

Life History: Cypripedium fasciculatum is a relatively long-lived perennial, living at least 30 years (Harrod 1994b), and perhaps as long as 95 years (Niehaus 1974). The orchid requires a fungal partner for seed germination and development and probably long-term maintenance (Arditti 1967; Doherty 1997; Wells 1981). Seed set, dispersal, and establishment of new populations are limited and not well understood. The pollinator is unknown.

Threats: Major threats to this species are loss of populations due to activities such as timber harvest, road and trail construction, creation of recreation sites, harvesting of forest products that disturbs litter and soil (herbal medicine, mushroom collecting), or fire. Threats include activities that alter the moisture or temperature regime, actions that disturb the soil and litter layer, or decrease vegetation cover to < 60 percent. Specific concerns associated with these activities are discussed in this text.

Management Recommendations:

1. Maintain or restore habitat conditions in areas with populations of C. fasciculatum.
2. Maintain canopy closure at 60 percent or greater (USDA and USDI 1994a).
3. 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.
4. Avoid activities that alter soil, duff, down wood, and the mycorrhizal community in the habitat area.
5. Maintain/secure known sites from prescribed burns except in Adaptive Management Areas (AMA) where research should be conducted to determine the role of fire in C. fasciculatum habitat (USDA and USDI 1994a).
6. 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).
7. 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.

Information Needs:

• Specific ecological requirements of C. fasciculatum for germination, establishment, and growth.
• The role of fire in the ecology of this species.
• Identification of pollinators and their habitat requirements.
• Identification of fungal associates, their habitat requirements, and the role they play in the life history of C. fasciculatum.
• Determination of the total number of extant populations within the west Cascades and the Klamath Mountains.
• Determination of the health, viability, and stability of known populations. Revisit C. fasciculatum sites reported more than 5 years ago to determine if the habitat is intact and if C. fasciculatum is present.

I. NATURAL HISTORY

A. Taxonomic/Nomenclatural History

Scientific name: Cypripedium fasciculatum Kellogg ex S. Watson

Cypripedium fasciculatum was originally described in the literature by Sereno Watson (1882) from a collection made by Wilhelm Suksdorf in May 1880, "on the White Salmon River, Washington Territory, above the falls." Other collections mentioned in the description were made in California in Plumas County and probably in Del Norte County. The first specimen in Oregon was collected by Thomas Howell in 1884 in Josephine County, near Grave Creek (Siddall and Chambers 1978).

Common name: Clustered Lady's Slipper, Brownie's Lady Slipper

Family: Orchidaceae

Subfamily: Cypripedioideae

Genus: Cypripedium contains about 45 species in the Northern Hemisphere. Eleven are native to North America (Cribb 1997).

Citations: Cypripedium fasciculatum Kellogg ex S. Watson, Proceedings of the American Academy of Arts and Sciences 17:380. 1882. LECTOTYPE: White Salmon River, above the falls, Washington Territory, May 1880, W. N. Suksdorf.

Synonyms: Cypripedium pusillum Rolfe, Kew Bulletin 1892:211. 1892 Cypripedium fasciculatum Rolfe var. pusillum Hooker f., Botanical Magazine plate 7275. 1893.

Cypripedium knightae A. Nelson, Botanical Gazette 42:48. 1906.

Two species of Cypripedium, C. pusillum Rolfe (1892) and C. knightae A. Nelson (1906), were later described as being notably different from C. fasciculatum. However, Hitchcock et al. (1969) suggest that differences between C. knightae and C. fasciculatum do not merit specific or infraspecific designation. The name C. pusillum Rolfe was based on a cultivated plant of uncertain origin and was considered a synonym of C. fasciculatum by Hitchcock et al. (1969). Characters formerly proposed for separating eastern and western races of C. fasciculatum are of little use, and formal recognition of infra-specific taxa is not warranted on the basis of existing information (Brownell and Catling 1987).

B. Species Description: (Watson 1882; Abrams 1940; Correll 1950; Munz 1959; Peck 1961; Hitchcock et al. 1969; Luer 1975; Hickman 1993)

1. Morphology

The following description is from Hitchcock et al. (1969):

Stem 0.5-2 dm. tall, lanate-pilose, usually with a single sheathing bract near ground level, a pair of opposite leaves at to well above midlength, and often 1 or 2 lanceolate bracts near the inflorescence; leaves sessile, broadly elliptic to oblong-elliptic or elliptic-oval, mostly 4-8 cm. broad, rounded-obtuse to slightly acute; flowers (1) 2-4 in a rather tight cluster, subtended by conspicuous greenish bracts usually as long as the densely pilose ovary; sepals lanceolate-acuminate, 12-25 mm. long, greenish-brown or greenish-purple and usually purple-lined or -mottled, the lower pair fused completely or free at the tips only; petals similar to the sepals but usually somewhat broader; lip depressed-ovoid, shorter than the sepals, greenish-yellow with brownish-purple margins and often with a purplish tinge; staminodium 2.5-3 mm. long, about equaling the longest lobe of the stigma.

This species cannot be mistaken for any other Cypripedium growing in the same range, because of its small size, 2 sub-opposite leaves on a hairy stem, and the tight cluster of greenish-brown flowers with large pouches (Knight 1994). The 2 cm long, oblong capsules contain thousands of small dust-like seeds. The stem is often weak and the disproportionately large leaves may arch down to touch the ground (Luer 1975).

Cribb (1997) and Doherty (1997) have summarized what is known about the genus Cypripedium. Their reviews cover morphology, life history, cytology, phylogenetic relationships, biogeography, ecology including mycorrhizal associations, uses, culture and propagation, artificial hybridization, and taxonomy. Managers should consult these publications for details.

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 two or more subsequent vegetative seasons before blooming (Whitlow 1983; Case 1987).

Knecht (pers comm) investigated the rhizome of C. fasciculatum by excavating duff and soil away from aerial stems. Rhizomes on the east side of the Cascades were located approximately 3-7 cm (1.2-2.75 in.) below the soil's surface. On the west side they were typically deeper, but never greater than about 12 cm (4.7 in.) below the surface. Rhizomes produced buds during the growing season that remained dormant below the soil's surface and grew into shoots the following early spring. Each stem and shoot pair was attached to the distal end of a woody rhizome branch with numerous stem scars (Harrod 1994b). Some rhizomes measured over two-thirds of a meter (2 feet) long. Each shoot had corresponding adventitious roots growing down from the ventral side of the rhizome branch. Individual aerial stem scars along the branch corresponded to individual roots, many of which were senescent. According to Harrod (1994b) this species produces a dormant bud during the current growing season. This bud remains inactive through the winter, then bolts in the spring to produce an aerial stem. At the same time, a new bud is initiated and developed during the photosynthetic period. Harrod dates rhizomes by counting stem scars (assuming one stem scar per year). He estimated the rhizome he excavated to be between 25 and 30 years old. If the rhizome had broken or rotted off, only the minimum age could be determined. Stoutamire (1991) used a similar dating technique to determine the age of C. candidum. Niehaus (1974) reported studying a C. fasciculatum plant that was at least 95 years old.

2. Reproductive Biology

Several orchids, including C. calceolus, take 13 to 16 years to begin to flower (Harper and White 1974; Wells 1981). Harrod (1993b) found that small, nonflowering C. fasciculatum plants can be at least 12 years old.

As a rhizomatous perennial, C. fasciculatum may reproduce asexually by rhizome fragmentation. According to Summerhayes (1951) Cypripediums and other rhizomatous orchids reproduce when older parts of a branching rhizome dieback, leaving younger, still living branches as separate plants. This may be the case for C. fasciculatum although this has not been demonstrated in C. fasciculatum. However, Aagaard et al.s' genetic analysis (unpubl) of small C. fasciculatum populations found a level genetic variability that indicates that plants may be produced from seeds rather than by asexual reproduction.

Several stems may belong to one genet (a plant derived from a single seed) that means sites with more than one stem may actually be only one plant (Harrod et al. 1995). Several stems could belong to a ramet (individual plants derived from a single, asexually-reproducing plant) by dieback or fragmentation of a branching rhizome. Counting each stem as a plant (as was done on siting reports) could produce an inflated number of plants. However, stem counting would be a good estimate of the health (stored photosynthates) and sexual reproductive potential (flowering stems) of the population.

Cypripedium fasciculatum is self-compatible but not autogamous or apomictic and requires a biotic vector for successful pollination (Knecht 1996). The shape of C. fasciculatum flowers and position of reproductive structures suggest they have coevolved with a particular pollinator to achieve cross-fertilization (Luer 1969; Barker 1984; Harrod 1993b). Although no pollinator has been determined for C. fasciculatum, Knecht (1996) suspects the pollinator may be some type of fly (Diptera) because of the flower's musky, fetid odor, brownish color, and morphology. Other arthropods attracted to carrion, such as wasps, hornets, and beetles should not be discounted. In Europe, wasps are known to visit Epipactis helleborine and E. purpurata (Proctor et al. 1996) species with flowers similar in color to C. fascicuatum.

Correll (1950) and Barker (1984) indicate that C. fasciculatum has low fruit production. Barker (1984) feels pollination is an infrequent event. Harrod (1993b) found that only 31 percent of the observed flowers produced fruit capsules. The large number of seeds produced per capsule, an average 3874 per fruit (Harrod and Knecht 1994), may partially compensate for the low level of fruit production. Further investigation is needed to determine the role of pollinators in the reproductive success of C. fasciculatum.

Electrophoretic isozyme analysis of C. fasciculatum by Aagaard et al. (unpubl) indicated little genetic variation among 3 populations they studied on the Leavenworth Ranger District, Wenatchee NF. Based on their analysis, C. fasciculatum may be an outcrosser (which is consistent with Knecht et al., in prep) with significant gene flow among sampled populations. According to Aagaard et al. (unpubl) levels of genetic variability were similar to other North American species of Cypripedium reported by Case (1994). In addition, there was basically no genetic variation within "clusters" of aerial stems, suggesting that clusters of stems belong to the same individual. This is consistent with Harrod's findings (in prep) that groups of aerial stems arise from a common rhizome and are, therefore, a single genetic individual (USDA and USDI 1994a). Liston (pers comm) reports that preliminary DNA studies found plants a few centimeters apart that appeared in most cases to be raments. Plants farther apart appeared different enough to be derived from seeds. Aagaard et al. (unpubl) studied small populations and found low levels of genetic variability. This suggests that future habitat disturbances need to be reduced to prevent further loss of genetic variability.

3. Ecology

Relationships among C. fasciculatum and its fungus, fire, and ecosystem dynamics are not well understood. Careful C. fasciculatum habitat studies have begun in the last few years. Results of these studies are not known; however, some information is available from a few completed studies and observations, and studies of related Cypripedium species.

The role of fire as a component of C. fasciculatum's habitat is not clearly understood, but the shallow rhizome system may exacerbate the deleterious effects of fire. Harrod et al. (1996) studied fire effects on C. fasciculatum on the Wenatchee NF. They concluded that C. fasciculatum is a fire-intolerant species, and management of this species probably should not include prescribed fire. Their work suggests that the species cannot tolerate low-intensity fire that eliminates the duff layer, as indicated by a lack of rhizomes found in excavations after fire. Some plants survived where the duff layer was not eliminated. While hot fires have destroyed sites on the Klamath NF (Barker 1984), Knorr (pers comm) noted that sites that had burned at a low-level of intensity in 1987 showed increased numbers of individuals (stems) and expanded population areas. The plants also showed apparent good vigor, with many small individuals that were perhaps seedlings. Greenlee (1997) reported the effects of a fire on the Clearwater NF that led him to conclude that C. fasciculatum can survive low to moderate intensity fire, but not higher intensity fires. Because rhizomes are 3-12 cm (1-5 in) below the soil's surface, intense fire can damage or kill the plants at any level in the profile.

It is suggested in Appendix J2 (USDA and USDI 1994a) that the reintroduction of prescribed fire might reduce the risk of extirpation of C. fasciculatum. Because observations on the effect of fire seem to be in conflict, a carefully planned experimental prescribed burn should be conducted and analyzed before fire is used as a management tool.

Cypripedium fasciculatum's small, dust-like seeds have great potential for long-distance wind dispersal. Doherty (1997) reports that the drooping inflorescences of C. fasciculatum substantially straightened and elongated after pollination, enhancing seed dispersal. However, because air circulation is inhibited in the understory of forested habitats, seeds only disperse up to 2 m from the parent plant (Harrod 1993b). Harrod and Everett (1993) and Harrod (1994a) developed a model to predict seed dispersal. Their model suggests that C. fasciculatum seed dispersal is relatively limited (i.e., a 43-in dispersal distance in a 10 mph wind), which is consistent with preliminary results presented by Harrod (1993b). This may indicate that poor seed dispersal is one factor in the limited distribution of C. fasciculatum.

Harrod (1993b) hypothesizes that big game animals may be important for long distance dispersal and seed germination by seeds being carried in digestive tracts of animals. Deer and elk trails often occur by populations of C. fasciculatum. Elk and deer feces may contain the fungus required for germination because ungulates are known mycophagists. Observations in the Wenatchee NF (Harrod 1993b), Siskiyou NF (Kagan 1990), and Medford District BLM (Knight 1994) indicate a noticeable degree of browsing on mature fruit capsules by unknown herbivores. This hypothesis has not been tested.

Cypripedium fasciculatum has an intricate life cycle that is not fully understood. All orchids appear to require the presence of a fungus, usually a Rhizoctonia, before the seed will germinate in the wild (Arditti 1967; Doherty 1997; Wells 1981). Doherty (1997) reports that there is no question developing orchids depend on its fungal symbiont for survival. Once an orchid reaches maturity and becomes autotrophic, the degree of dependence may change.

Establishment of new populations requires suitable conditions for the fungus. What these 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) postulates that once Cypripedium plants germinate and grow to the photosynthetic state, the role of the fungus literally ceases.

Stoutamire (1991) investigated the mycorrhizal relationship and annual growth cycle of the root system of another Cypripedium species (C. candidum). He found that mycorrhizae are inhibited near the rhizome and appear irregularly in older roots. The fungal symbionts (5 unidentified fungal isolates cultured from cortical cells) were short lived and left amorphous masses in the cortical cells until senescence.

Although it is generally assumed that fungi play an active nutritional role throughout the life of C. fasciculatum (Sheivak, pers comm; Trappe, pers comm), further research is needed to firmly establish the nature and extent of the relationship. Caution should be exercised in applying the ectomycorrhizal model of forest trees to orchids.

Kagan (1990) noted a correlation between C. fasciculatum and Douglas-fir snags or western dogwood. He speculates that in southern Oregon mycorrhizal associations may be interconnected with old Douglas-firs (Pseudotsuga menziesii), Pacific madrone (Arbutus menziesii), or Pacific dogwood (Cornus nuttallii), which are commonly found in reproducing C. fasciculatum population areas.

C. Range, Known Sites

Cypripedium fasciculatum occurs in widely disjunct locations from Washington as far south as the Santa Cruz Mountains in California and in the mountains of Idaho, Montana, Colorado, Wyoming, and Utah. Hitchcock et al. (1969) and Luer (1975) report C. fasciculatum in British Columbia, although Catling (1983) cited by Brownell and Catling (1987) discount this occurrence.

According to Brownell and Catling (1987), C. fasciculatum inhabits 6 major areas: 1) Sierra Nevada of California; 2) Coast Range and Cascade Range along the Oregon-California border (Siskiyou Mountains); 3) Santa Cruz Mountains on the central coast of California; 4) Cascade Range in Washington; 5) Rocky Mountains of northern Idaho (Darlene, Selkirk, and Clearwater Mountains), northeastern Oregon (Blue Mountains), and western Montana (Mission and Swan Ranges); and 6) Rocky Mountains of Utah (Wasatch and Uinta Mountains), Colorado (Park and Front Ranges), and Wyoming (Medicine Bow and Park Range Mountains).

Although C. fasciculatum appears to have a broad range, its distribution within the 8 western states is relatively scattered and disjunct. The northern range limit for C. fasciculatum is the Wenatchee NF in the northern Cascades of Washington. The southern range limit is the Santa Cruz Mountains of the central California coast. It occurs on 9 National Forests and 3 BLM Districts. These management recommendations pertain only to the Klamath Province as identified in the ROD. Information on habitat and observations from other areas will help to understand C. fasciculatum in the Klamath Mountains.

The following table represents the number of sites reported at the time of the viability analysis for the Northwest Forest Plan.

Table 1. Known sites by administrative units in Washington, Oregon, and California
within the range of the northern spotted owl.

* Administrative units where the Management Plan applies.

D. Habitat Characteristics and Species Abundance

Cypripedium fasciculatum occurs in a variety of habitats throughout its range that vary greatly in soils, elevation, aspect, and plant communities. Cypripedium fasciculatum is not restricted to any particular parent material and sites vary from dry to damp, rocky to loamy. Elevations range from 330-1745 m (1000-5300 ft.). Populations are found in areas with 60 to 100 percent shade provided by plant communities, including mixed evergreen, mixed conifer, Douglas-fir, pine, and black oak forest. The aspect is mainly northerly. Sites occur in several plant associations within the Douglas-fir series.

Barker (1984) found that the habitat of C. fasciculatum cannot be closely defined in the Klamath NF. Sites have been described as dry or damp, rocky to loamy, and elevations vary from 429-1750 m (1300-5300 ft.) (Barker 1984). Populations are found in areas with 60 to 100 percent shade provided by trees and shrubs in numerous plant communities, including mixed evergreen, mixed conifer, Douglas-fir (Pseudotsuga menziesii) forests, and pine/black oak (Pinus sp./Quercus kelloggii) forests (Barker 1984).

Cypripedium fasciculatum is not restricted to a particular parent material. Populations have been found on ultrabasics, granitics, schist, limestone, and quartz-diorite. Fowlie (1988) states that C. fasciculatum occurs on serpentine landslides in northwestern California. However, Fowlie also notes that the plants were "composted [growing in organic matter] in a serpentine landslide of vast extent, matted with fallen California Douglas-fir needles," and that the plants sometimes grew between the roots of the fir. This might indicate that soil organic matter is more important than parent material.

Kagan (1990) notes that in southwestern Oregon (Rogue River, Siskiyou, Umpqua and Klamath N F; and the Medford District BLM) the species occurs primarily in older Douglas-fir forests. Douglas-fir is the dominant species in the canopy (highest in frequency and cover) in the majority of plant communities in which C. fasciculatum has been reported on the Medford District BLM. Within these habitats, there is a variety of plant associations in which C. fasciculatum is found. These plant associations most often include variations of the Douglas-fir/dwarf Oregon-grape (P. menziesii/Berberis nervosa), Douglas-fir/poison oak (P. menziesii/Rhus diversiloba), and tan oak/dwarf Oregon-grape (Lithocarpus densiflorus/B. nervosa) associations. Other associations include western hemlock (Tsuga heterophylla), white fir (Abies concolor), white oak (Q. garryana), bigleaf maple (Acer macrophyllum), and tan oak (L. densiflorus). Understory associates include sword fern (Polystichum munitum), Oregon-grape (B. nervosa), dogwood (C. nuttallii), and hazel (Corylus cornuta). Elevations vary from 330-1155 m (1000-3500 ft.). The largest populations in southwestern Oregon tend to occur on or near old stream terraces (Kagan 1990).

Cypripedium fasciculatum usually grows in filtered light to shady areas and is rarely found growing in the open. In the Klamath NF, it is most often found in areas with 60 to 100 percent shade provided by canopy cover (Barker 1983). In the Wenatchee NF, average canopy closure is 62 percent. This species generally grows in moist areas but is found in dry sites as well.

Knecht (1996) studied C. fasciculatum at 12 unspecified sites on the Wenatchee, Umpqua, Siskiyou, and Rogue River NF, and in the Wenatchee Mountains of Washington and the Cascade Range and Klamath Mountains of Oregon. Populations grew under a wide variety of site conditions. Aspect ranged from 42-(mean 241)-352 degrees; elevation ranged from 410-(mean 761)-1900 m (1345-6234 ft.) and slope from 7-(mean 43)-84 percent. Distribution of the number of plants or populations across these ranges was not provided. Average values were provided. Consistent factors found in all plots on the east and west sides of the Cascades were presence of a Douglas-fir overstory >70 years, canopy cover >60 percent, and soil pH values between 5.7 and 6.5.

Because of the association between orchids and fungi and the heterotrophic mode of fungal nutrition, the important environmental factor controlling distribution of C. fasciculatum might be the nature of the upper organic layers of the soil profile and not the nature of the mineral soil. Some soil factors that may be important include soil depth, source, rate of decomposition, moisture content, and pH. The bryophyte communities that cover the shallow rhizome systems of C. fasciculatum may also be important. Coarse woody material may provide microsite moisture and shade and protect duff and litter layers from disturbance. Woody materials may also be evidence of lack of fire.

Abundance is difficult to ascertain. Actual number of plants at a single site cannot be determined as one rhizome often produces more than one aerial stem. Populations of C. fasciculatum consist of distinct clumps of one to a few hundred stems, each clump potentially representing a rhizomatous clone of the same genetic individual (Aagaard et al., unpubl). This indicates that the actual number of plants at each site may be much smaller than numbers shown on siting reports. There is no practical way to determine if stems are shoots from a single rhizome, raments (independent members of a clone) or genets (individuals derived from seed) short of submitting individuals to genetic analysis. Therefore, throughout this document, stem numbers will be referred to instead of plant numbers.

The abundance of stems at any one site is usually 1 to 20. Current data indicate that few sites have >50 stems per site. Ninety-six percent of the siting reports examined have <50 stems per site. Medford BLM, the administrative unit with the most sites, shows 53 percent of the sites having 10 or fewer stems. The largest reported population throughout the range of C. fasciculatum is near Taylor Creek, Siskiyou NF with more than 1000 stems.

Most sites were discovered while doing clearance work for timber sales. Siting reports indicate that many of the locations occurred within timber sale units. These reports do not state if mitigation measures were part of the plan. It is not known if C. fasciculatum survived at these sites after harvest occurred. Total number of extant sites may be considerably lower than siting reports indicate.

II. CURRENT SPECIES SITUATION

A. Why Species is Listed under Survey and Manage Standards and Guidelines

These management recommendations are limited to the Klamath Province. The Cascade Province was omitted when the ROD was published (USDA and USDI 1994c) even though C. fasciculatum was found to be at higher risk for extirpation in the Cascade Province under all alternatives of the Forest Plan.

Cypripedium fasciculatum was found to be sparsely distributed throughout its range with few stems at each site. Ninety-six percent of the sites had 50 or fewer stems. Scientists who did the viability analysis for the Northwest Forest Plan felt that most populations of C. fasciculatum had lost the potential for genetic interaction with other populations. Because of their extremely slow growth rate, complex symbiotic relationships with other organisms, and possible fire requirements it was concluded that recolonization of these species throughout their former range was unlikely (USDA and USDI 1994a).

B. Major Habitat and Viability Considerations

Certain features of C. fasciculatum's biology and ecology have important habitat and viability implications. Individuals are long-lived, may take a number of years to mature to flowering, have a symbiotic relationship with a fungus that derives nutrition from organic layers of the soil for part or all of its life, and have a shallow rhizome system sensitive to mechanical and fire disturbance. Knecht's (1996) observations indicate that plants growing in areas with <60 percent canopy closure were small and appeared faded. It is not known how long plants growing under these conditions will survive.

The major viability consideration for C. fasciculatum is loss of populations due to land management activities that directly or indirectly impact the species and its associated habitat. Populations of C. fasciculatum 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 than larger populations to succumb to natural catastrophes such as wildfire, floods, landslides, drought, and loss of pollinating insects (Falk and Holsinger 1991) or to human perturbation such as collection, and habitat alteration such as timber harvest, road building, and grazing. Smaller populations are at greater risk of extirpation through management activities occurring near or within their habitat where machinery can accidently easily destroy the entire population.

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. fasciculatum is essential for species survival. Given (1994) gives 500 individuals as an acceptable goal for minimum effective population size. He states if a population drops to 50-100 individuals, there is cause for concern. The minimum effective population size for C. fasciculatum has not been determined.

Optimum habitat conditions for C. fasciculatum do not occur in early successional communities. Most populations are found in areas with relatively closed canopies that develop later in succession. This species may require a mycorrhizal symbiont, not only as adult plants, but also for seed germination as found in other Cypripedium species (Arditti 1967). It is possible that the required symbiont(s) is (are) only present in mid- to late-successional forest communities (Harrod et al. 1996). 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.

C. Threats to the Species

Physical disturbance of sites. 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 the adventitious roots of C. candidum are particularly sensitive to disturbance. He found that damaged roots are slow to repair and are replaced slowly from the most recently produced rhizome sections.

Loss of microclimate to the sites. Forest structure appears to provide important microclimatic conditions for C. fasciculatum sites. Modification of forest structure (for example, canopy removal) has a profound effect on interior microclimates such as temperature and moisture (Chen et al. 1993, 1995), and ground level vegetation (Chen et al. 1992; Frost 1992). Greenlee (1997) reports a drop of 58 plants to 2 plants after a blowdown on the Nez Perce NF in Idaho.

Activities that remove canopy in large areas or patches close to C. fasciculatum populations could alter the microclimate of nearby sites by creating edge effects. Depending upon distance and exposure, there could be changes in several microclimate variables such as air temperature, relative humidity, soil temperature, and moisture, which could impact C. fasciculatum (Chen 1995).

D. Distribution Relative to Land Allocations

The following tables indicate land use designations at C. fasciculatum sites throughout the range of the Northwest Forest Plan (Table 2) and the Klamath Province (Table 3) as number of total sites and percent of total sites in each category. Land use allocation categories differ between the 2 tables.

Table 2. Known (as of 1994) Cypripedium fasciculatum Sites
Within the Range of the Northwest Forest Plan by Land Allocation

Table 3. Known Cypripedium fasciculatum Sites (as of 1994) in the Klamath Province.

III. MANAGEMENT GOALS AND OBJECTIVES

A. Management Goals for the Taxon

The management goal for C. fasciculatum is to assist in maintaining species viability within the Klamath Province.

B. Specific Objectives

• Maintain or restore ecological conditions at known sites such as hydrologic, temperature, and light regimes.
• Maintain viable populations of C. fasciculatum within occupied habitat, including potential mycorrhizal associations.
• Mitigate to avoid habitat reduction and population fragmentation.
• Maintain coarse, woody material in C. fasciculatum habitat.
• Maintain undisturbed duff and soil layers to protect mycorrhizal networks and C. fasciculatum roots and rhizomes.

IV. HABITAT MANAGEMENT

A. Lessons from History

Observation of extant sites provides information for management of C. fasciculatum. It appears that the level of damage to the site determines the impact on the plant. Some of these observations are:

Observations from Montana (Greenlee 1997) indicate that C. fasciculatum individuals in a clear-cut and in a burn with a fairly open canopy tend to dry up and turn yellow earlier than plants in less open canopy conditions.

Past forest management activities, such as timber harvest, may have contributed to declines in population numbers. This has been documented in California (USDA and USDI 1994a).

Activities that expose or excessively damage the rhizome of C. fasciculatum seem to eliminate the plant. Knecht (1996) found that plants had died in an area where fire eliminated the duff. Knecht also observed a healthy plant excavated to examine rhizome structure died. This may be related to root damage (Stoutamire 1991).

Low-intensity fire that does not eliminate the duff layer or destroy the canopy appears to have no adverse impact on C. fasciculatum (Harrod and Knecht 1996). In some cases, it appears to have benefitted the species (Knorr pers comm).

High-intensity fire that eliminates the duff layer also destroys C. fasciculatum rhizomes (Harrod and Knecht 1996).

Certain types of disturbance might be beneficial. A portion of a study plot (Knecht 1996) was run over by a bulldozer in 1993. Studies begun in 1994 found no aerial plants in the bulldozer tracks. In 1995 a number of stems appeared clustered in the track, including several that flowered.

B. Identification of Habitat Areas for Management

The habitat area identified in the ROD (USDA and USDI 1994c) for management is federal lands in the Klamath Province of southern Oregon and northern California. The original rating in the FEMAT report (USDA and USDI 1994b) indicated a 55 percent likelihood of extirpation in the Cascade Province and 20 percent in the Klamath Province (USDA and USDI 1994a) for C. fasciculatum.

C. Management within Habitat Areas

1. Maintain or restore habitat conditions in areas with populations of C. fasciculatum.
2. Maintain canopy closure at 60 percent or greater (USDA and USDI 1994a).
3. 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.
4. Avoid activities that alter soil, duff, down wood, and the mycorrhizal community in the habitat area.
5. Maintain/secure known sites from prescribed burns except in Adaptive Management Areas (AMA) where research should be conducted to determine the role of fire in C. fasciculatum habitat (USDA and USDI 1994a).
6. 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).
7. 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 Klamath Province to determine their contribution as future habitat.
• The Cascade Province was not listed as an area for management in the ROD. The Cascade Province should be included in the management area. It is necessary to manage populations, habitat, and to provide connectivity throughout the species' geographic range to sustain species viability.
• 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.

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 Klamath Province. Site visits should be made to determine stability and viability of populations reported over the last 20 years.

Revisit C. fasciculatum 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. fasciculatum seed germination and establishment needs.

Determine the role of fire (wildfire and controlled burns) C. fasciculatum 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 determined 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. fasciculatum.

B. Research Questions

What level of disturbance can C. fasciculatum plants withstand?

What is the role of fire in C. fasciculatum's life history? Viability analysis panel members indicated that investigation of the role of fire and prescribed burns could be important in reducing the likelihood of extirpation of this species (USDA and USDI 1994a). Recent research indicates that the role of fire needs to be carefully analyzed (Harrod et al. 1995). This research suggests that C. fasciculatum is a fire-intolerant species (at least with high-intensity fire) and management of this species probably should not include prescribed fire. An answer to this question is needed for management of wild fire and control burns in the known habitat areas of C. fasciculatum.

What mycorrhizal fungi are associated with C. fasciculatum? Is this association species-specific and/or codependent?

What are the requirements for the fungal host; what soil environment is needed to support the fungal mycobiont?

What are the biotic and abiotic requirements for each stage of C. fasciculatum's life history?

What are critical microclimatic elements? How do edge effects modify the microclimate of the habitat (temperature and moisture), and how does this affect the plant?

What stand characteristics are conducive to this species' establishment?

Is there a correlation between stand age and the age of C. fasciculatum plants in the stand?

What role does coarse woody material at sites play in mycorrhizal association and moisture retention?

What specific site characteristics are necessary to maintain existing populations?

Are remnant late-successional and old-growth forests necessary for establishment of new populations? If so, is this because of mycorrhizae requirements or the orchid's requirements, or both?

Who are the pollinators, and what are their habitat needs?

What are the requirements for seed germination and seedling establishment?

What is the role of ungulates and other mammals in the ecology and distribution of this species?

What are the requirements for seed germination and seedling establishment?

Do plants increase in size with increasing age of the rhizome? What is the longevity of individual rhizomes?

How many years can C. fasciculatum remain vegetative underground?

What factor(s) trigger flowering and flowering periodicity?

What is the genetic variability of C. fasciculatum throughout its range?

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 recommended mitigating measures are being implemented and 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. fasciculatum plants will require long- term (>20 yrs) to determine vigor, reproduction, and survival of the populations.

VI. REFERENCES

Aagaard J. E., Shea K., Harrod R. J. (unpubl). Genetic variation within and among populations of the rare orchid, Cypripedium fasciculatum. 23 p. Available from: J. E. Aagaard, 16524 104th NE , Bothell, WA 98011.

Abrams L. 1940. Illustrated flora of the Pacific states, Vol. 1. Stanford(CA): Stanford Univer. Pr. 557 p.

Arditti J. 1967. Factors affecting the germination of orchid seeds. Bot Rev 33:1-97.

Barker L. M. 1984. Botanical investigation and management guidelines for Cypripedium species. [unpubl report]. Yreka (CA): USDA, Forest Service, Region 5, Klamath NF.

Brownell V. R., Catling P.M. 1987. Notes on the distribution and taxonomy of Cypripedium fasciculatum Kellogg ex Watson (Orchidaceae). Lindleyana 2(1):53-57.

Camp A. 1995. Predicting late-successional fire refugia from physiography and topography. [PhD dissertation] Seattle: University of Washington. 137 p. Available from: UMI, Ann Arbor, MI.

Case M. 1994. Extensive variation in the levels of genetic diversity and degree of relatedness among five species of Cypripedium (Orchidaceae). Am J Bot 8(2):175-184.

Catling P. 1983. Terrestrial Orchids in Canada. In: Plaxton EH, editor. North American Terrestrial Orchids. Symposium II, Proceedings and Lectures. Southfield: Michigan Orchid Soc. p. 87-132.

Chen J., Franklin J., Spies T. 1992. Vegetation responses to edge environments in old-growth Douglas-fir forests. Ecol Appl 2:387-396.

_________ 1993. Contrasting microclimates among clear-cut, edge, and interior old-growth Douglas-fir forest. Agric Forest Meterol 63:219-237.

_________ 1995. Growing-season microclimatic gradients from clear-cut edges into old-growth Douglas-fir forests. Ecol Appl 5(1):74-86.

Correll D. S. 1950. Native orchids of North America north of Mexico. Waltham (MA): Chronica Botanica. 400 p.

Cribb P. 1997. The genus Cypripedium. Portland (OR): Timber Pr. 301 p.

Doherty J. W. 1997. The genus Cypripedium: a botanical and horticultural overview. North American Native Orchid J. March. 5-116.

Falk D., Holsinger K. 1991. Genetics and conservation of rare plants. New York: Oxford Univer Pr. 283 p.

Fowlie J. A. 1988. Cypripedium fasciculatum on serpentine landslides of northwestern California. Orchid Digest (July-Aug.-Sept.): 136-139.

Frost E. J. 1992. The effects of forest clear-cut edges on the structure and composition of old-growth mixed conifer stands in the western Klamath Mountains. [MSc thesis]. Arcata (CA): Humboldt State University.

Given D. R. 1994. Principles and practice of plant conservation. Portland OR: Timber Pr. 292 p.

Greenlee J. 1997. Cypripedium fasciculatum conservation assessment. [unpubl rept]. Missoula (MT): USDA, Forest Service, Region 1, Lolo N F.

Harper J. L., White J. 1974. The demography of plants. Ann Rev Ecol Syst. 5: 419-463.

Harris L. D. 1984. The fragmented forest. Chicago: Univer Chicago Pr. 211 p.

Harrod R. J. 1993a. Habitats of Cypripedium fasciculatum on the Wenatchee National Forest. [unpubl study]. USDA, Forest Service, Region 6, Wenatchee N F, Leavenworth Ranger District.

Harrod R. J. 1993b. Studies relating to Cypripedium fasciculatum. [report]. USDA, Forest Service, Region 6, Wenatchee NF, Leavenworth Ranger District.

Harrod R. J. 1994a. Characteristics and Dispersal of Cypripedium fasciculatum seeds. [abstract]. Northw Sci Assoc Meeting, 23-26 March 1994. Northw Sci. 68(2): 129

Harrod R. J. 1994b. Rhizome morphology and sensitivity to ground disturbance of Cypripedium fasciculatum. [report]. USDA, Forest Service, Region 6, Wenatchee NF, Leavenworth Ranger District.

Harrod R. J., Everett R. 1993. Preliminary observations on seed dispersal and seed production of Cypripedium fasciculatum. [abstract]. Northw Sci Assoc Meeting, March 1993. Northw Sci 67(2): 131.

Harrod R. J., Knecht D. 1994. Preliminary observations of the reproductive ecology of Cypripedium fasciculatum. [abstract]. North Sci Assoc Meeting, 23-26 March 1994. Northw Sci. 68(2):129.

Harrod R. J., Knecht D., Kuhlmann E., Ellis M., Davenport R. 1997. Effects of the Rat and Hatchery Creek fires on four rare plant species. In: Proceedings of the Conference on Fire Effects on Rare and Endangered Species and Habitats. Nov. 13-16, 1995, Coeur d'Alene (ID): IAWF. Fairfield, WA.

Hickman J. C., editor. 1993. The Jepson manual: higher plants of California. Berkeley: Univer. California Pr. 1400 p.

Hitchcock C. L., Cronquist A., Ownbey M., Thompson J. W. 1969. Vascular plants of the Pacific Northwest, Vol. 1. Seattle: Univer Washington Pr. p 832-833.

Hooker J. D. 1893. Cypripedium fasciculatum var. pusillum, Tab. 7275. Curtis' Botanical Magazine 49 (3rd series): 119.

Johnson C. J. 1969. Migration and dispersal of insects by flight. London (UK): Methuen. 763 p.

Kagan J. 1990. Draft species management guide for Cypripedium fasciculatum for south western Oregon. Portland: Oregon Natural Heritage Data Base.

Knecht D. 1996. The Reproductive and population ecology of Cypripedium fasciculatum (Orchidaceae) throughout the Cascade Range. [MSc thesis]. Ellensberg: Central Washington University. 64 p.

Knight L. 1994. Draft habitat management guide for Cypripedium fasciculatum. [report]. Medford (OR): Bureau of Land Management, Medford District.

Luer C. A. 1969. The genus Cypripedium in North America. Am Orchid Soc Bull. October: 903-908.

_________. 1975. The native orchids of the United States and Canada excluding Florida. New York Bot Garden. 361 p.

Munz P. A. 1959. A flora of California. Berkeley: Univer California Pr. 1681 p.

Nelson A. 1906. Contribution from the Rocky Mountain herbarium, VII. Botan Gaz 42:48-54.

Niehaus T. F. 1974. Sierra wildflowers: Mt. Lassen to Kern Canyon. California Natural History Guides. Berekeley: Univer California Pr. 223 p.

Noss R., Cooperrider A. 1994. Saving nature's legacy: protecting and restoring biodiversity. Washington D. C. and Covelo (CA): Island Pr. 416 p.

Peck M. 1961. A manual of the higher plants of Oregon. 2d ed. Portland (OR): Binfords and Mort. 936 p.

Proctor M., Peyo A., Lack A. 1996. The Natural History of Pollination. Portland (OR): Timber Pr. 479 p.

Rolfe R. A. 1892. Kew Bulletin 1892:211.

Seevers J. Unpublished reports. Medford (OR): Bureau of Land Management, Medford District.

Sheviak C. J. 1990. Biological considerations in the management of temperate terrestrial orchid habitats. NY State Mus Bull 471. p 194-196.

Siddall J. L., Chambers K. L., Wagner D. 1979. Rare, threatened and endangered vascular plants in Oregon --an interim report. Salem: Oregon State Land Board, Division of State Lands. 109 p.

Stoutamire W. P. 1991. Annual growth cycles of Cypripedium fasciculatum Muhl. Roots systems in an Ohio prairie. Lindleyana 6:235-240.

Summerhayes V. S. 1951. Wild Orchids of Great Britain. London (UK): Collins. 366 p.

Tamm C. O. 1972. Survival and flowering of perennial herbs, II. The behavior of some orchids on permanent plots. Oikos 23:274-292.

USDA, Forest Service, and USDI, Department of Interior, Bureau of Land Management. 1994a. Final Supplemental Environmental Impact Statement on Management of Habitat for Late-Successional and Old-growth Forest Related Species within the Range of the Northern Spotted Owl. Appendix J2, Results of Additional Species Analysis. Portland, OR.

USDA, Forest Service, and USDI, Bureau of Land Management. 1994b. Final Supplemental Environmental Impact Statement on Management of Habitat for Late-successional and Old-Growth Forest Related Species within the Range of the Northern Spotted Owl, Appendix A, Forest Ecosystem Management: An Ecological, Economic, and Social Assessment. Portland, OR.

USDA, Forest Service, and USDI, Bureau of Land Management. 1994c. Record of Decision for Amendments to Forest Service and Bureau of Land Management Planning Documents within the Range of the Northern Spotted Owl and Standards and Guidelines for Management of Habitat for Late-successional and Old-Growth Forest Related Species within the Range of the Northern Spotted Owl. Portland, OR.

Watson S. 1882. Proc Am Acad. 17:380. 1882.

Wells T. C. E. 1981. Population ecology of terrestrial orchids. In: Synge H, editor. The Biological Aspects of Rare Plant Conservation. New York: J Wiley. p 281-295.

Whitlow C. E. 1983. Cypripedium Culture. In: Proc. N. Am. Terrestrial Orchids Symposium II and lectures. Livonia (MI): Mich. Orchid Soc.