Terrestrial Map 5.1B— Ranges of Two Locally Endemic Subspecies on the Edges of Their Parent Species' Ranges

Map 5. la— Ranges of two locally endemic subspecies on the edges of their parent species' ranges. Potholes meadow vole (Microtus pennsylvanicus kincaidi) and its parent species, meadow vole (M. pennsylvanicus).

Terrestrial Map 5.1b— Ranges of two locally endemic subspecies on the edges of their parent species' ranges. White Salmon pocket goph< (Thomomys talpoides limosus) and its parent species, northern pocket gopher (Thomomys talpoiaes).

Terrestrial \ Map 5.2a—Examples of vertebrate species with disjunct ranges within and adjacent to the Basin: Woodhouse's toad, green frog and Northern bobwhite.

Terrestrial Map 5.2b--Examples of vertebrate species with disjunct ranges within and adjacent to the Basin: California mountain kingsnake, ground snake, longnose snake, and wild turkey.

Terrestrial Plethodon idahoensis), no reptiles, 12 birds (includ- Species Associated with Snags, ing 5 raptors, 4 primary cavity excavators, and Including Cavity-users others), and 3 mammals (American marten, heather vole Phenacomys intermedius, and wood- Another forest component is the presence of dead land caribou) show particular correlations to standing trees or snags. Primary cavity-excavator canopy condition. birds include some 17 species (3 chickadees, 3 nuthatches, and the rest woodpeckers). Beyond Other species show correlations with other forest these 17 species are an additional 29 species of stand characteristics, including tree bark structure, vertebrates that closely associate with snag sub- tree diameter (dbh), stem density, and flora. Cer- strates. These include 3 amphibians, 3 raptors, 7 tainly, many species can be included in these lists, bats, and 3 carnivores, which use a variety of but the species listed above provide a beginning specific snag characteristics for a wide variety of point for identifying good indicators of changes in purposes. At least another 14 species are obligate canopy conditions. secondary cavity-users (see discussion below under the section Interspecies Relations). Species Associated with Mistletoe Brooms A set of species show correlations with mistletoe Species Associated with Down Wood brooms in tree crowns, including at least one Among the listed, candidate, or potential candi- invertebrate, the gossamer-winged butterfly date species of plants, 46 fungi species, the de- Mitowra johnsoni (Lycaenidae). Among verte- cayed wood bryophyte group, and five lichen brates, the marbled murrelet, spruce grouse groups were identified in the database as associated (Dendragapus canadensis), Cooper's hawk (Accipiter with down wood. At least 11 invertebrate species cooperit), long-eared owl (Asia otus), and American were identified as examples of the several thousand fisher (Martes pennant!) all use mistletoe brooms species associated with down wood. Two examples for a variety of purposes. are the checkered beetle, Enoclerus sphegeus (Cleridae), which uses both fresh down wood and Species Associated with also intact but older down wood; and the Dead Parts of Live Trees leafcutting bee Osmia bruneri (Megachilidae), which uses snags and stumps, but also uses down Among species associated with dead tree branches wood stems with abandoned beetle burrows. are the bark beetle Scolytus ventralis (Scolytidae), 6 Additionally, 82 species of amphibians, reptiles, species of woodpeckers, and 2 owls [great gray owl birds, and mammals show correlations with down (Strix nebulosa) and, though it occurs erratically wood. within the Basin, northern hawk owl (Surnia ulula)]. Interestingly, our investigations uncovered a small set of species that seem, in part, to spe- Species Associated with cialize on bark piles at the base of snags. These Trees With Exfoliating Bark include at least the invertebrate Pseudogarypus hesperus (Psendogarypidae: Chernilldal), and Some six vertebrates specialize on trees with exfoli- two amphibians, the northwestern salamander ating (peeling) bark: the white-headed woodpecker Ambystoma gracile (which also uses down logs), (Picoides albolarvatus), the brown creeper (Certhia and the Larch Mountain salamander Plethodon americana), and four bats [silver-haired bat larselli (which uses bark piles especially in the (Lasionycteris noctivagans), California myotis driest season in sites that also have moist talus (Myotis californicus), little brown myotis (Myotis and low soil content). lucifugus), and Yuma myotis (Myotisyumanensis)], although some of these species also use other substrates as roost or nest sites.

Terrestrial Species Associated with Litter And Duff the warbler more in brushland, riparian, and fire- generated shrubby montane sites with deciduous Some example species listed in the SER database trees in the understory. Lewis' (Melanerpes lewis), that are closely associated with litter and duff black-backed (Picoides arcticus), and three-toed conditions include six species of fungi, the humus/ (Picoides tridactylus) woodpeckers, along with duffbryophyte group, one lichen species and two mountain and western bluebirds (Sialia currucoides lichen groups (nitrogen-fixing soil lichens, and and Sialia mexicana), associate with sites where rotten log and tree base lichens). The carib beetle fires have killed the forest overstory. Pterostichus protractus (Carabidae) associates with fallen woody material and litter, as well as with Black-backed and three-toed woodpeckers associ- bases of herbaceous plants, rocks, and snags. ate with mid- to high-elevation forests following Among vertebrates, at least four shrews, the shrew- irruptions of insect populations such as bark mole, and three songbirds show close associations beetles. In the eastern portion of the Basin, the with litter and duff, uncommon Tennessee warbler (Vermivora peregrina) increases after outbreaks of spruce Species Associated with Fire Processes budworm. and Insect Outbreaks Species associated with Recreation Plant species show various responses to fire sup- Activities, Roads, and Trails pression, frequency, intensity, and seasonality (table 5.15). Response to fire by fungus, lichen, As with all other management activities, ground and bryophyte species generally is not known, and and soil disturbances associated with recreational response by vertebrate species is not easily de- areas, roads, and trails, can have positive, neutral, picted. and negative effects on distribution or abundance of species. Among vertebrates, black-tailed deer Among the many invertebrates that respond in may be negatively affected by the presence of various ways to fire, three examples are the lace cross-country skiers, destination resorts, logging bug Acalypta cooleyi (Tingidae), which is adversely activities, off-road vehicle activity, roads, and affected by prescribed (and unprescribed) range subdivisions. Other species also adversely affected fires that burn mosses; bark beetles including by such disturbance activities include tailed frog Dendroctonuspseudotsugae (Scolytidae), which are (Ascaphus truef), western toad (Bufo boreas), positively correlated with forest fire intensity and Woodhouse's toad (Bufo woodhousii), Great Basin that associate with forest stands with severe spadefoot, Coeur d'Alene salamander, sage grouse, drought (but not with forest stands with few to no and mountain goat. fire-injured Douglas-fir trees greater than 30 cm dbh within a 10-km radius); and the thatch ant Vertebrates benefiting from disturbance activities Formica obscuripes (Formicidae), which is posi- are species associated with building structures, tively correlated with fire frequency. including swallows and some of the bats (such as pallid bat Antrozouspallidus); gravel road and Among the vertebrates, western tanager (Piranga shoreline substrates, benefiting species such as ludoviciana) and purple martin (Progne subis) killdeer; agricultural areas, benefiting species such respond positively to recent burns (tanagers more as horned lark (Eremophila alpestris) and rosy in cottonwood riparian forest, and martins in finches (Leucosticte spp.); and human presence and locations near water). Red-headed woodpecker habitations, benefiting species such as house spar- Melanerpes erythrocephalus (rare in the Basin) and row (Passer domesticus), opossum, house mouse orange-crowned warbler (Vermivora celata) re- (Mus musculus), and black rat (Rattus rattus). spond more to older burns, the woodpecker in cottonwood/oak forests and open woodlands, and

Terrestrial Table 5.15— Plant species associated with various aspects of fire dynamics (excluding fungi). BG = bryophyte group, L = lichen, LG = lichen group, P = vascular plant, PG = vascular plant group (source: SER database).

Class Name Class Name Fire Suppression Activities Positive Effect LG Pioneer Soil Stabilizer Lichens LG Tree Crusts P Collomia renacta Negative Effect BG Aquatic Submerged Bryophyte BG Decayed Wood Bryophyte BG Epiphytic Bryophyte BG Humus Duff Bryophyte BG Rock Wet Bryophyte BG Soil Dry Bryophyte L Oceanic Fruticose L Texosporium sancti-jacobi LG Charred Snag Lichens LG Forage Lichens LG Fruticose Tree Lichens LG Leaf Lichens LG Moss and Detritus Binders Lichens LG N-fixing epiphytes lichens LG N-fixing riparian lichens LG N-fixing soil lichens LG Oceanic forage lichens LG Oceanic leaf lichens LG Oceanic tree crust lichens LG Pin Lichens LG Riparian Lichens LG Rotten Log and Tree Base Lichens LG Soil lichens P Allium dictuon P Amsinckia carinata P Arabis falcifructa P Arabis fecunda P Arabis suffrutescens var. horizontalis P Aster jessicae P Astragalus anserinus P Astragalus atratus var. inseptus P Astragalus mulfordiae P Astragalus oniciformis P Astragalus paysonii P Astragalus peckii P Astragalus solitarius P Astragalus sterilis P Astragalus tegetarioides P Astragalus tyghensis P Astragalus yoder-williamsii P Botrychium ascendens P Botrychium crenulatum P Botrychium paradoxum P Botrychium pedunculosum P Botrychium pumicola P Calochortus longebarbatus var. longebarbatus P Calochortus longebarbatus var. peckii P Calochortus nitidus P Castilleja chorotica P Castilleja christii P Chaenactis cusickii P Collomia mazama P Cypripedium fasciculatum P Douglasia idahoensis P Erigeron latus P Eriogonum novonudum P Grindelia howellii P Hackelia cronquistii P Haplopappus liatriformis P Haplopappus radiatus P Ivesia rhypara var. rhypara P Lepidium davisii P Lepidium papilliferum P Lesquerella carinata var. languida P Lesquerella paysonii P Lesquerella sp. nov. ("pulchella") P Lomatium erythrocarpum P Mentzelia mollis

1602 Terrestrial Table 5.15 (continued).

Class Name Class Name P Mimulus clivicola P Mirabilis biglovii var. retrosa P Mirabilis macfarlanei P Orysopis herdersonii P Penstemon glaucinus P Penstemon lemhiensis P Penstemon peckii P Phacelia inconspicua P Phacelia lutea var. calva P Phacelia lutea var. mackenzeorum P Phacelia minutissima p Phlox idahonis P Rubus bartonianus P Silene spaldingii P Stanleya confertiflora p Thelypodium eucosmum P Trifolium thompsonii PG Botrychium Marsh Spp Group PG Botrychium Meadow Spp Group PG Carex Calcareous Peatlands Group PG Carex Dry Forest Group PG Carex Dry Meadow Low Moderate Elev Group PG Carex Ephemeral Meadow Subalpine Group PG Carex Mesic Forest Group PG Carex Mesic Meadow Low Moderate PG Carex Mesic Meadow Elev Group Subalpine Group PG Carex Noncalcareous Peatlands Group PG Carex Sagebrush Steppe Dry Group PG Carex Sagebrush Steppe Wetland PG Carex Wet Meadow Low Moderate Riparian Group Elevation Group PG Carex Wet Meadow Subalpine Group Fire Frequency Positive Effect BG Rock Wet Bryophyte BG Soil dry Bryophyte LG Charred Snag Lichens LG Pioneer Soil Stabilizers Lichens P Astragalus paysonii P Botrychuim ascendens P Botrychium crenulatum P Botrychium lineare P Botrychium paradoxum P Botrychium pedunculosm P Botrychium pumicola P Cypripedium fasciculatum P Grindelia howellii P Lesquerella sp. Nov ("pulchella") P Mimulus clivicola P Penstemon glaucinus P Penstemon lemhiensis P Silene spaldingii P Trifolium thompsonii PG Botrychium Marsh spp group PG Botrychium Meadow spp group PG Carex Dry Meadow Low Moderate Elev Group PG Carex Ephemeral Meadow Low PG Carex Ephemeral Meadow Moderate Elev Group Subalpine Alpine Group PG Carex Mesic Forest Group PG Carex Mesic Meadow Low Moderate Elev Group PG Carex Mesic Meadow Subalpine Alpine Group PG Carex Sagebrush Steppe Dry Group PG Carex Wet Meadow Subalpine Alpine Group Negative Effect BG Aquatic Submerged Bryophyte BG Decayed Wood Bryophyte BG Epiphytic Bryophyte BG Humus Duff Bryophyte

Terrestrial Table 5.15 (continued).

Class Name Class Name BG Rock Other Bryophyte L Oceanic Fruticose L Texosporium sancti-jacobi LG Fencepost Lichens LG Forage Lichens LG Fruticose Tree Lichens LG Leaf Lichens LG Moss and Ditritus Binders Lichens LG N-Fixing Riparian Lichens LG N-Fixing Soil Lichens LG Oceanic Forage Lichens LG Oceanic Tree Crust Lichens LG Pin Lichens LG Soil Lichens LG Tree Crusts P Aster jessicae P Astragalus anserinus P Astragalus atratus var. inseptus P Astragalus mulfordlae P Astragalus pulsiferae var. suksdorfii P Astragalus sinuatus P Astragalus solitarius P Astragalus sterilis P Botrychium cenulatum P Calochortus nitedus P Chaenactis cusickii P Collomia renacta P Cypripedium fasciculatum P Erigeron latus P Eriogonum novonudum P Hackelia cronquistii P Haplopappus radiatus P Howellia aquatilis P Lepidium daws/7 P Lepidium papilliferum P Lupinus cusickii P Mirabilis biglovii var. retrosa P Phancelia lutea var. mackenzeorum P Stanleya confertiflora P Thelypodium eucosmum P Thelypodium howellii ssp. spectabilis Fire Intensity —Overstor y Lethal Positive Effect

LG Aspen Specialist Lichens LG Charred Snag Lichens LG Fencepost Lichens LG Moss and Ditritus Binders Lichens LG N-Gixing Soil Lichens LG Pioneer Soil Stabilizers Lichens LG Soil Lichens P Astragalus peckii P Astragalus tyghensis P Botrychium pumicola P Caolchortus longebarbatus P Haplopappus radiatus var. longebarbatus P Lesquerella sp. nov. ("pulchella") P Penstemon glaucinus P Penstemon peckii P Trifolium thompsonii PG Carex Calcareous Peatlands Group PG Carex Ephemeral Meadow Low Moderate Elev Group PG Carex Ephemeral Meadow Subalpine PG Carex Mesic Meadow Low Alpine Group moderate Elev Group PG Carex Mesic Meadow Subalpine Alpine Group PG Carex Noncalcareous Peatlands Group PG Carex Wet Meadow Subalpine Alpine Group PG Carex Wet Meadow Low Moderate Elev Group PG Carex Sagebrush Steppe Dry Group Negative Effect

BG Aquatic Submerged Bryophyte BG Decayed Wood Bryophyte

1604 Terrestrial Table 5.15 (continued).

Class Name Class Name BG Humus Duff Bryophyte BG Rock Wet Bryophyte BG Soil Dry Bryophyte L Oceanic Fruticose L Texosporium sancti-jacobi LG Forage Lichens LG Fruticose Tree Lichens LG Leaf Lichens LG N-Fixing Epiphytes Lichens LG N-Fixing Riparian Lichens LG Oceanic Forage Lichens LG Oceanic Leaf Lichens LG Pin Lichens LG Rotten Log and Tree Base Lichens LG Tree Crusts P Astragalus pulsiferae var. suksdorfii P Astragalus sinuatus P Astragalus solitarius P Botrychium ascendens P Botrychium crenulatum P Cypripedium fasciculatum P Howellia aquatilis P Lepidium papilliferum P Mimulus clivicola P Thelypodium eucosmum P Thelypodium howellii ssp. spectabilis PG Carex Forest Riparian Group PG Carex Wet Forest Group Prescribed Fire Seasonality Spring Prescribed Fire Positive Effect LG Charred Snag Lichens LG Forage Lichens LG Oceanic Log Lichens LG Pioneer Soil Stabilizers Lichens P Grindelia howellii P Silene spaldingii PG Carex Sagebrush Steppe Dry Group Spring Prescribed Fire Negative Effect BG Decayed Wood Bryophyte BG Epiphytic Bryophyte BG Soil Dry Bryophyte L Texosporium sancti-jacobi LG Pin Lichens LG Riparian Lichens P Arabis fecunda P Astragalus peckii P Astragalus pulsiferae var. suksdorfii P Astragalus sinuatus P Astragalus solitarius P Astragalus tyghensis P Botrychium ascendens P Botrychium crenulatum P Botrychium paradoxum P Botrychium pumicola P Calochortus longebarbatus var. longebarbatus P Calochortus longebarbatus var. peckii P Cypripedium fasciculatum P Lesquerella carinata var. languida P Lesquerella paysonii P Lomatium erythrocarpum P Lupinus cusickii P Mimulus clivicola P Mimulus pulsiferae P Mirabilis biglovii var. retrosa P Mirabilis macfarlanei P Oryzopsis hendersonii P Penstemon glaucinus P Penstemon peckii P Phacelia lutea var. mackenseorum P Rubus bartonianus P Stanleya confertiflora P Thelypodium eucosmum P Thelypodium howellii ssp. spectabilis P Trifolium thompsonii

Terrestrial 1605 Table 5.15 (continued).

Class Name Class Name Late Summer or Fall Prescribed Fire Positive Effect LG Charred Snag Lichens LG Pioneer Soil Stabilizers Lichens P Astragalus applegatei P Astragalus peckii P Astragalus tyghensis P Botrychium pumicola P Calochortus longebarbatus var. longebarbatus P Cypripedium fasciculatum P Mimulus clivicola P Penstemon lemhiensis P Penstemon peckii P Silene spaldingii PG Carex Dry Meadow Low Moderate Elev Group PG Carex Ephemeral Meadow Low Moderate Elev Group PG Carex Ephemeral Meadow Sublapine PG Carex Mesic Meadow Low Moderate Alpine Group Elev Group PG Carex Mesic Meadow Subalpine Alpine Group PG Carex Sagebrush Steppe Dry Group Late Summer or Fall Prescribed Fire Negative Effect BG Aquatic Submerged Bryophyte BG Decayed Wood Bryophyte BG Epiphytic Bryophyte BG Rock Wet Bryophyte BG Soil Dry Bryophyte L Texosporium sancti-jacobi LG Oceanic Log Lichens LG Pin Lichens LG Riparian Lichens LG Rotten Log and Tree Base Lichens P Arbis fecunda P Astragalus pulsiferae var. suksdorfii P Astragalus solitarius P Botrychium ascendens P Botrychium paradoxum P Cypripedium fasciculatum P Grindelia howellii P Haploppus radiatus P Mimulus pulsiferae P Silene spaldingii P Thelypodium howellii ssp. spectabilis Early Historic Fire Suppression Activities Long-term Effects Positive Effects

BG Aquatic Submerged Bryphyte BG Decayed Wood Bryophyte BG Epiphytic Bryophyte BG Humus Duff Bryophyte BG Rock Wet Bryophyte BG Soil Bry Bryophyte L Texosporium sancti-jacobi LG N-Fixing Epiphytes Lichens LG N-Fixing Riparian Lichens LG N-Fixing Soil Lichens LG Oceanic Forage Lichens LG Pin Lichens LG Riparian Lichens LG Soil Lichens LG Tree Crusts P Rubus bartonianus Negative Effect

BG Rock Other Bryophyte L Oceanic Fruticose LG Aspen Specialist Lichens LG Charred Snag Lichens LG Excess Nitrogen Indicator Lichens LG Forage Lichens LG Fruticose Tree Lichens LG Moss and Ditritus Binders Lichens LG Oceanic Leaf Lichens LG Oceanic Log Lichens

Terrestrial Table 5.15 (continued).

Class Name Class Name LG Oceanic Tree Crust Lichens LG Pioneer Soil Stabilizers Lichens LG Rotten Log and Tree Base Lichens P Allium aaseae P Allium nevii P Arabis falcifructa P Aster jessicae P Astragalus anserinus P Astragalus atratus var. inseptus P Astragalus mulfordiae P Astragalus oniciformis P Astragallus paysonii P Astragalus peckii P Astragalus pulsiferae var. suksdorfii P Astragalus sinuatus P Astrgalus tegetariodes P Astragalus yoder-williamsii P Botrychium ascendens P Botrychium crenulatum P Botrychium pedunculosm P Botrychium pumicola P Calochortus longebarbatus var. longebarbatus P Calochortus longebarbatus var. peckii P Calochortus nitidus P Castilleja chlorotica P Castilleja christii P Castilleja pilosa var. steenesis P Chaenactis cusickii P Collomia mazama P Cypripedium fasciculatum P Douglasia idahoensis P Erigeron latus P Eriogonum novonudum P Grindelia howellii P Hackelia cronquistii P Haplopappus liatriformis P Haplopappus radiatus P Lepidium davisii P Lepidium papilliferum P Lesquerella carinata var. languida P Lesquerella paysonii P Lesquerella sp. nov. ("pulchella") P Lomatium greenmanii P Mimulus clivicola P Mirabilis biglovii var retrosa P Mirabilis macfarlanei P Penstemon glaucinus P Penstemon lemhiensis P Penstemon peckii P Phacelia inconspicua P Phacelia lutea var. calva P Phacelia lutea var. mackenxeorum P Phacelia minutissima P Phlox idahonis P Silene spaldingii P Stanleya confertiflora P Thelypodium eucosmum P Thelypodium howellii ssp. spectabilis P Trifolium douglasii P Trifolium thompsonii PG Allium Riparian Spp Group P Botrychium Forest Spp Group PG Botrychium Marsh Spp Group PG Botrychium Meadow Spp Group PG Carex Dry Forest Group PG Carex Dry Meadow Low Moderate Elev Group PG Carex Ephemeral Meadow Low Moderate PG Carex Ephemeral Meadow Subalpine Elev Group Alpine Group PG Carex Mesic Meadow Low Moderate PG Carex Mesic Meadow Subalpine Elev Group Alpine Group PG Carex Sagebrush Steppe Dry Group PG Carex Wet Meadow Low Moderate Elev Group PG Carex Wet Meadow Subalpine Alpine Group PG Penstemon cinicola complex

Terrestrial 1607 Human activities that result in carrion (fresh dead Major Biomass animals) might benefit the set of carrion-feeding species including turkey vulture (Cathartes aura), Species that predominate in or affect the biomass bald eagle, gray jay (Perisoreus canadensis), com- of an ecosystem can have a major controlling mon raven (Corvus corax), American crow (Corvus function in storage and flow of nutrients and brachyrhynchos), wolverine, and coyote, depending energy through the trophic levels. Many species, on type and location of the carrion. including plants, can dominate as major biomass of various environments. Among invertebrates, soils can be dominated numerically by mites, Key Ecological Functions of Species springtails, and other species. Western spruce This discussion presents findings on species' key budworm (Choristoneura occidentalis) and ecological functions associated with the major Douglas-fir tussock moth (Orgyiapseudotsugata) terrestrial ecological subsystems of the Basin (table can greatly affect production of major biomass, as 5.16). The major ecological functions we ad- they are key to carbon and nutrient recycling, dressed were: species that predominate in the plant biomass turnover, soil productivity, and biomass of ecosystems; herbivory; nutrient cycling energy flow dynamics in some forest ecosystems. relations; interspecies relations; soil productivity; In litter and duff layers of many forests, amphib- wood decomposition; and water quality. Under- ians often comprise either a major biomass com- standing and maintaining ecological functions, ponent or at least serve to mediate energy flow including nutrient cycling and fire relations, are through trophic levels. This may be surprising to critical facets of maintaining ecosystem health for those who would guess that only the larger or inland Northwest ecosystems (DellaSala and more visible vertebrates perform this function. others 1995). Identifying such ecological func- Specifically, throughout the Basin, at least nine tional groups of species may be more useful for salamander species and three frogs act as key management of Federal lands in the Basin than energy gates or major biomass components. Col- would attempting to identify individual keystone lectively, these amphibians are most sensitive to species. changes in down wood, litter, and duff depths and This section summarizes information derived from characteristics; still and flowing water quality and the SER database under key ecological functions quantity; and precipitation quality and weather (KEFs). Information on species' KEFs is incom- patterns. Thus, manipulating these components plete, particularly for plants and invertebrates. through management activities can influence the Information on KEFs for plant species and plant distribution and abundance of associated amphib- groups was gathered only for those species cur- ians and thus their ecological functions. rently listed as Cl or C2 candidates or for species and groups which the plant expert panels and Herbivory contractors deemed deserving of additional conser- vation focus, that is, potential Cl or C2 candi- Herbivory functions of invertebrates were dis- dates. The plant information cited below is cussed earlier. In summary, at least one example typically for plants that are scarce; other plant species of tree bole feeder, five bark beetles, and 22 species with particular functions are not discussed defoliators (including cankerworms, budworms, here. The information on invertebrates is also an leaf beetles, miners, moths, shoot borers, sawflies, incomplete representation of that fauna as the and others) can serve to alter plant succession, database contains only example species of a much create new vegetative conditions for other wildlife, larger invertebrate fauna. Our intent was to oudine decrease primary production of host plants, and some of the major categories of species functions impose other changes on vegetation substrates and within the Basin rather than to list exhaustively all ecological processes of their host environments. species with each ecological function.

1608 Terrestrial Table 5.16— A simple classification of ecological subsystems found in the Basin.

1. Terrestrial Above-Ground Environments (a) Upland terrestrial conditions - forests Tree canopy, subcanopy Shrub layer Grass/forb layer (b) Upland terrestrial conditions - shrublands and grasslands Trees Shrubs Grass/forb layer Cryptogamic crusts (soil surface) 2. Terrestrial Below-Ground Environments (a) At- and below-surface zones: coarse wood, litter, duff (b) Soil (O, A Horizons) (c) Root sphere influence zones 3. Riparian influence zone Riparian shade, leaf litter, large wood 4. Aquatic systems (a) Lotic systems Stream size scale and gradient type Bank and channel stability; side slope angle and erosion potential Reach type; habitat characteristics and processes (edge effects, storage of organisms, hydrology, particle size sorting) (b) Lentic systems Water body size Water permanence lake or pond astatic lake or pond intermittent pond wet meadow, seep, bog, fen

Vertebrates with key herbivory functions include Nutrient Cycling all the ungulates discussed previously, as well as at least five species of waterfowl (four geese and Species with key roles in determining cycling rates brant), sage grouse, at least three species of vole, of nutrients (for example, nitrogen, phosphorous, and the black-tailed jackrabbit and other rabbits. and carbon) include those involved in physical All of these major herbivores are capable of transfer of substances, nitrogen cycling, and car- altering plant community structure and plant bon sequestration. First are species that particulary species composition, particularly the ungulates affect the physical transfer of substances for nutri- as studied in the Blue Mountains by Irwin and ent cycling. In turn, this key ecological function others (1995). Even the voles can modify grass- serves to support vegetation succession and to alter land structures. the composition and enhance the productivity of the environments. This function is performed by various animals including 21 example invertebrate

Terrestrial 1609 species among 13 families. Two examples are large buffer system involving cycling of carbon, in aspen tortrix (Choristoneura conflictana: various forms, from atmosphere to vegetation to Tortricidae) and pandora moth (Coloradia soils and back. Among the species listed in the pandora: Saturniidae). Many more invertebrates SER database with this function are seven Carex with this function occur within the assessment (sedge) plant groups found in calcareous and area. Vertebrates with this function number at noncalcareous peat lands; dry low-elevation least 32 species among 12 families; the species meadow vegetation communities; and mesic and include 11 amphibians [two examples are Pacific wet meadow vegetation communities at low, chorus frog (Pseudacris regilla) and rough-skinned moderate, and alpine elevations. Carbon-cycling newt (Taricha granulosa)], one bird [Canada goose models (for example, Harmon and others 1990, (Branta canadensis)~\ and 21 mammals [two ex- Turner and others 1995) can help estimate the rate amples are the bushy-tailed woodrat (Neotoma of contribution of forests to carbon stores. cinerea) and the little brown myotis (Myotis lucifugus)]. Interspecies Relations Second are species specifically affecting nitrogen Every species interacts with others. However, cycling. These include nitrogen-fixers, nitrogen- several key interactions may be of interest to immobilizers, and sources for nitrogen mineraliza- ecosystem management, including insectivory, tion. Among the species listed in the SER pollination vectors, transport of plant dissem- database, 1 bryophyte group, 7 lichen groups, and inules, nest parasitism, cavity nesting, and burrow 32 vascular plant species are mostly nitrogen- excavation. These interactions affect productivity, fixers. The dry soil bryophyte group harbors nitro- diversity, and structure of ecological communities. gen-fixing bacteria and is found in all general soil categories. The seven nitrogen-fixing lichen groups Many insectivorous (that is, insect-feeding) verte- include groups labelled in the SER database as brates within the assessment area act as biocontrol excess nitrogen indicators and nitrogen-fixing agents. While these biocontrol organisms may not groups of epiphytes, riparian and seepage species, eliminate irruptive insect populations, they may rock and soil associates, and tundra sod builders. play a significant role in reducing the amplitude of Rare vascular plants affecting nitrogen cycling, as insect outbreaks. Insectivorous vertebrates with listed in the SER database, include some 22 spe- such biocontrol potential include an amphibian cies of milk-vetch, as well as lupines, (tiger salamander Ambystoma tigrinum), 12 birds, monkeyflower, crazyweeds, clovers, and others. and 9 mammals. Insectivorous birds include (Additional leguminous and other vascular plants trumpeter swan (Cygnus buccinator), American in the Basin also perform nitrogen fixation; these kestrel (Falco sparverius), flammulated owl (Otus are not included in the SER database at this time). flammeolus), common nighthawk (Chordeiles minor), black-backed woodpecker (Picoides Invertebrates and microorganisms with nitrogen arcticus), barn swallow (Hirundo rustica), northern relation functions include the competitive bacterium rough-winged swallow (Stelgidopteryx serripennis), Bacillus thermophilus, the nitrogen-immobilizing mountain chickadee (Parus gambelt), mountain bacterium Klebsiella panticola, and many rotifers. bluebird (Sialia currucoides), hermit thrush The bacterium serves to decompose labile and inter- (Catharus guttatus), red crossbill (Loxia mediate-labile carbon: nitrogen substrates and act as a curvirostra), and evening grosbeak (Coccothraustes source for nitrogen mineralization. The rotifers vespertinus). The mammals include deer mouse participate in nitrogen-retention in soil. (Peromyscus maniculatus) and many bats. A third key function is carbon sequestration. Many trees and shrubs contribute to sequestration of atmospheric carbon, acting as part of a global

Terrestrial This list of species could have important economic least chipmunk (Tamias minimus), Douglas' squir- implications because some insectivorous species rel (Tamiasciurus douglasit), and Rocky Mountain feed on insect pests. The evening grosbeak and elk. The western red-backed vole disperses nitrogen- mountain chickadees are primary feeders of spruce fixing mycorrhizae. Douglas' squirrel and southern budworms. Red crossbill, associated with mature red-backed vole, and probably others, disperse and old conifer forests, feeds on spruce budworm lichens. Most of the waterbirds, including water- larvae. The silver-haired bat is a moth and beetle fowl, shorebirds, grebes, herons, and others, pest predator in forests. Common nighthawk, big disperse aquatic plants, algae, and invertebrates. brown bat (Eptesicus fuscus), and pale western (that The largest groups of vertebrate dispersal agents is, Townsend's) big-eared bat (Plecotus townsendii are those that disperse seeds and fruits (at least 85 pallescens) feed on insects in agricultural and urban identified vertebrate species) and those that dis- environments. Black-backed woodpeckers feed on perse plants or non-seed or fruit plant parts (60 bark beetles, especially during outbreaks. Little vertebrate species). The seed- or fruit-dispersers empirical work has been done for most of these include a wide variety of aquatic and terrestrial species to quantify rates of feeding and its specific birds and mammals, covering a wide range of effects on the amplitude of insect population environments within the Basin area. A few seem to outbreaks. Koplin (1969) suggested that black- be specialists. For example, Clark's nutcracker backed woodpeckers can suppress the amplitude of (Nucifraga columbiana) is a key element in regen- bark beetle outbreaks. Takekawa (1984) evaluated eration of whitebark pine—which is declining in the silvicultural economic advantage of providing the Basin-as well as pinyon pine, by caching (that for evening grosbeaks. is, burying) seeds in the ground. White-headed Another interspecies relation of potential interest woodpecker disperses ponderosa pine seeds and to ecosystem management is pollination vectors. California ground squirrel (Spermophilus beecheyi) Pollination vectors are species which aid produc- disperses oak (further west and south from the tion of native fruit and seed-bearing plants that, in Basin, tanoak) acorns and seeds. Rodents, espe- turn, feed mammals, birds, ants, and other insects. cially red-backed voles, play key roles in re-estab- There is no master catalogue of such vectors for lishing trees by burying seeds and spreading spores the Basin, but the SER database contains a sample of beneficial symbiotic fungi, especially after of 63 invertebrate species (among 12 families) timber harvest or fires. However, some evidence with this key function. Among vertebrates, at least suggests that if the clearing is too large, they will five species of hummingbirds and one songbird aid in re-establishment of trees only along the edge [northern oriole (Icterus galbula)} act as an- of forest cover (Mills 1995).35 giosperm pollinators. The hummingbirds pollinate Bird nest parasitism is a specialized breeding tubular flowers; the rufous hummingbird function of the brown-headed cowbird (Molothrus (Selasphorus rufus) in particular is co-evolved with ater). At least 30 other bird species suffer as nest red tubular flowers. hosts to this species. Affected species include Another interspecies relation is dispersal of fungi, forest-inhabiting vireos, warblers, and sparrows; plants, plant parts, and invertebrates. At least 153 marshland blackbirds and yellowthroats; vertebrate species engage in transportation of shrubland sparrows and flycatchers. To our knowl- seeds, spores, or other plant disseminules. Some of edge, no study has determined the extent to which the vertebrate dispersers of fungi include southern and western red-backed voles (Clethrionomys 35Also, personal communication. 1995. Allen E. Thomas, gapperi and Clethrionomys californicus), pika Fish and Wildlife Biologist. U.S. Department of Interior, (Ochotonaprinceps), flying squirrel (Glaucomys Bureau of Land Management, Idaho State Office, 3380 Americana Terrace, Boise, Idaho 83706. sabrinus), Columbian mouse (Peromyscus keenit),

Terrestrial cowbird nest parasitism is a factor in declines in beetles which create distinctive galleries of burrows other bird species. Studies in the Sierra Nevada of in the tree cambium beneath the bark. Two other California have suggested that there could be primary ground-burrowing invertebrates are the severe, negative affects on host species. thatch ant Formica obscuripes (Formicidae), whose nests are home to many arthropod species and The 17 vertebrate species that function as primary serve as wintering burrows for small mammals, cavity excavators in trees include 11 woodpeckers, andArctosa littoralis (Lycosidae: Araneida), whose 3 nuthatches, and 3 chickadees [chestnut-backed 15 to 25 centimeter deep burrows in sand, dug for chickadee (Pants rufescens), included here, is an diurnal hiding, might possibly influence the struc- occasional primary cavity excavator]. Collectively, ture of sandy beach habitats. Many other burrow- these species reside in a variety of pine, fir, and ing invertebrates occur in the Basin. hardwood woodlands and forests in the assessment area. The primary burrow-excavating vertebrates consti- tute about 41 taxa (38 species and two subspecies At least another 14 species act as secondary cavity of northern pocket gopher listed individually in users, using natural tree cavities or cavities created the SER database), which include 1 amphibian by the primary excavators, for roosting or nesting. (Great Basin spadefoot), 3 birds [bank swallow These include wood duck (buffleheads, mergan- (Riparia riparia), belted kingfisher (Ceryle alcyon), sers, and a few other waterfowl may also serve this burrowing owl (Athene cunicularia)], and the rest role), 5 small owls, 3 chickadees, 2 swallows, mammals. Great basin spadefoot is a primary white-breasted nuthatch (Sitta carolinensis), ash- excavator in sandy soil and uses mammal burrows throated flycatcher (Myiarchus cinerascens), and in other substrates. The mammals include moun- common raccoon (Procyon lotor). Doubtless, other tain beaver (Aplodontia rufa), the rare pygmy rabbit species can be added to this list. (Brachylagus idahoensis), striped skunk (Mephitis Competition for cavities may be high in forest mephitis), mink (Mustela visori), American badger conditions lacking trees or snags of adequate dbh (Taxidea taxus), 2 moles, 6 gophers, 14 squirrels or with cavities. Northern saw-whet owls (Aegolius chipmunks, 7 mice or rats, and 2 foxes. The acadicus) nest early in the season and thus may mountain beaver, skunk, badger, and foxes in displace other later-nesting secondary cavity users. particular dig large burrows used by a slightly Likewise, boreal owls (Aegolius funereus) occupy different constellation of other species. larger tree cavities and nest earlier than many The secondary burrow-using species depend on other secondary cavity users, and may displace the primary burrow excavators for their tunnels. flying squirrels. On the other hand, flammulated This set includes at least 13 vertebrate species (3 owls, our only neotropical migratory owl, are late amphibians, 3 reptiles, 1 bird, and 10 mammals): nesters and may not find adequate nest sites if they 2 salamanders and spadefoot; longnose leopard are occupied by earlier nesting species. Northern lizard, striped whipsnake, and gopher snake; pygmy-owls (Glaucidium gnoma) likely compete burrowing owl; and Virginia opossum, striped for cavities with passerines (songbirds). Western skunk, 3 mustelids (ermine, long-tailed weasel, screech owls (Otus kennicottii) may actively usurp and mink), and northern grasshopper mouse. In nests from squirrels and flickers. Among song- particular, the longnose leopard lizard depends on birds, violet-green swallows (Tachycineta thalassina) rodent burrows (kangaroo rats, pocket mice) for are an active competitor for nesting cavities. cover. Striped whipsnake uses deep underground As a parallel to the cavity-using species, another crevices in cold weather and is also a secondary species set engages in primary burrow excavation user of mammal burrows for egg-laying. in a variety of habitats and conditions. Among the invertebrates, this set includes a variety of bark

2" Terrestrial Soil Productivity sphere) bacteria, bacterial pathogens, and beneficial bacteria, are responsible for nitrogen retention Species with ecological functions affecting soil and mineralization in soils. Bacteria-feeding nema- productivity include a variety of plants, inverte- todes (Acrobeloides spp.) engage in mineralization brates, and vertebrates in seven function catego- of nitrogen immobilized in bacterial biomass, and ries: physical soil turnover, nitrogen retention or may be responsible for up to 40 percent of plant uptake, soil stabilization, rock weathering, available nitrogen. Rotifers (virtually unstudied in detoxificaion of xenobiotics, metal accumulation, the assessment area) and the bacterial-predator and vegetation succession. Soil productivity can be protozoa such as Amoeba amoeba also engage in related to complex processes. For example, Seasted mineralization of nitrogen immobilized in bacte- and Knapp (1993) reported that grassland ecosys- rial biomass, and may be responsible for up to 50 tem productivity can display "transient maxima," percent or more of plant-available nitrogen in which are related to how grazing and fire affect some agricultural systems. In addition, saprophytic water, nitrogen, and light availability. fungi enter the process, as most plant-available In one sense, all plants die, decompose, and thus nitrogen must cycle through soil fungal biomass in contribute to soil organic matter and composition. forest soils. In particular, one bryophyte group (rock calcare- Many plants contribute to soil stabilization, ous bryophytes), 2 plant groups (Carex ephemeral including 6 bryophyte groups, 9 lichen groupss meadow low-moderate group and Carex subalpine 22 plant groups, and at least 1 recognized lichen groups), and 13 plant species, listed in the SER species and, among the rare plants listed in the database, significantly contribute to soil formation SER database, 76 plant species or subspecies. The and soil organic material, particularly in poorly specific functions performed by this diverse group developed or nutrient-poor soils and limestone or include: trapping of sediments (for example, rock substrates. Many other species and plant aquatic submerged bryophytes, wet rock bryophytes, groups not listed in the SER database also contrib- and wet soil bryophytes); reducing wind erosion ute to this function. (for example, dry soil bryophytes); colonization of Some 51 example invertebrate species or subspe- harsh sites or erodible slopes; and stabilization of cies distributed over seven families are included in stream banks and soils during flood events (for the database as examples of invertebrates that example, Artemesia spp., the Carex rocky stream- recycle soil layers and nutrients, and influence soil bed species group, the Mimulus guttatus species structure, porosity, and aggregation by their physi- complex and Mimulus high-elevation wet-habitat cal digging actions. Among vertebrates that per- species group, and the Penstemon foothills to form this function are a number of fossorial montane meadow species group). Species that mammals listed above as primary burrow excava- stabilize soils on post-fire sites include several tors. In particular, western and Woodhouse's toads, species of milk-vetch (Astragalus spp., Chaenactis broad-footed and coast moles (Scapanus latimanus cusickii, Penstemon peckii, and others; also see table and Scapanus orarius), shrew-mole (Neurotrichus 5.15). Many other plants not listed in the SER gibbsii), 4 ground squirrels, white-tailed antelope database likely perform this function more widely squirrel (Ammospermophilus leucurus), and water throughout the Basin. The beneficial bacteria vole (Microtus richardsont) significantly contribute Microcoleus also serve to stabilize soil and prevent to soil aeration and affect soil structure. The water erosion. vole contributes to soil aeration and drainage of Lichens excel as soil stabilizers, with 11 lichen wet soils. species groups occurring in a wide array of envi- Bacteria, nematodes, rotifers, and protozoa play ronments serving to weather rock substrates. critical roles in nitrogen retention or uptake in Additionally, seven vascular plant species in the soils. Groups of competitive rhizosphere (root

Terrestrial 1S13 SER database, including penstemons, a rockmat, pecker (feeds more frequently on large down logs an ivesia, and a phacelia, also perform this service. compared to other woodpeckers) which breaks Rock weathering is often a prelude to further soil logs apart while foraging for insects and larvae; formation and provides a substrate for other colo- and the black bear and grizzly bear, which may nizers needing mineral soils. contribute to soil nutrient cycling by breaking logs apart in search of food. The ectomycorrhizal mat-forming fungi Hysterangium may detoxify some types of pesti- cides, herbicides, and pollutants. The vesicular Bioindicators of Water Quality fungus Glomus mossae and the ectomycorrhizal Algae and aquatic invertebrate species can act as mat-forming fungus Hysterangium may sequester bio indicators of water quality. Pond- and lake- heavy metals in fungal hyphae, preventing heavy dwelling high country salamanders and frogs also metals from damaging or killing plants. are sensitive indicators of acid precipitation and By definition, vegetation succession proceeds by changes in water quality. The tailed frog is sensi- new plant species invading a site. Many plant tive to, and is probably a very good indicator of, species of the Basin are colonizers. Some bacteria increases in temperature and sedimentation in groups, exemplified by the bacterium Rhizobium mountain streams. Additionally, some 19 species meUiotii and the beneficial bacteria group of shorebirds, including sandpipers, yellowlegs, Microcoleus, aid preparation of the soil for succes- plovers, and others, probably are good indicators sion by nitrogen-fixation. of pollution of inland nesting and stopover sites.

Wood Decomposition Identifying Keystone Species Wood decomposition is a highly complex process The term keystone has been used in the ecological that can involve several thousand species of bacte- literature to refer to species whose functions di- ria, microfungi, macrofungi, nonvascular and rectly or indirectly affect the presence and abun- vascular plants, invertebrates, and vertebrates. A dance of many other species. Keystones can be single down log can undergo decay succession; plants and allies, invertebrates, or vertebrates. The many species are present only at specific wood above reviews demonstrate that species assem- decay stages. One invertebrate in particular, the blages can be identified as those closely associated carpenter ant Camponotus modoc aids in decompo- with specific environmental substrates or with sition of standing and down wood. It operates at ecological functions. Collectively, all such species the hub of a complex ecological process involving are "keystone" in that they, as a group, define the insectivorous predatory birds, primary and second- structures and processes of ecosystems. ary cavity nesters, and defoliators. Partially decom- At our present knowledge level, few species can be posed standing wood provides nest, feeding, and demonstrated as being individually "keystone" in roost sites for many other species (see above, under that their removal from the ecosystem would species associated with snags, dead parts of live greatly diminish the diversity and functions of the trees, and exfoliating bark); decomposing down system. Further, Mills and others (1993) argued wood is also used by a variety of other species (see that the concept of keystone species is wrought above, under species associated with down wood with ambiguity and imprecision, and that land and litter and duff)- managers instead should explicitly consider the Among vertebrates that might aid wood decompo- complexity of interactions in natural systems. sition are the rubber boa (Charina bottae) and Following their suggestion, we have not identified sharptail snake (Contia tenuis), which excavate a small set of species as keystones or "indicators." rotten logs for cover and prey; the pileated wood-

10jt4"/ Terrestrial One problem with the use of the indicator species egories, yes. For example, black-tailed jackrabbit concept on Federal land management has been (Lepus californicus), meadow vole (Microtus described as, "the problem of objectively defining pennsylvanicus), greater white-fronted goose (Anser which species are keystones makes it likely that albifrons), and Ross' goose (Chen rossif) all perform subjectively chosen subsets of species will be so functions of herbivory (that is, all are herbivores) labeled, whereas other species of similar impor- (maps 5.3a, 5.3b). However, each species in the tance will be ignored" (Mills and others 1993). above list performs its function under different sets of environmental conditions and geographic We use the term "key environmental correlates" areas (maps 5.3a, 5.3b) using different behaviors (KEC) here to connote the major environmental and affecting their food sources in different ways. requirements of a species. A given KEC, such as Ultimately, each species is unique in its set of cliffs, can have many associated species, and a behaviors, distribution, and functions, so that no given species can have many KECs. Thus, if a two species are exactly interchangeable. Thus, we management objective is to maintain the full set of do not suggest viewing species with common, species in an area, we recommend, along with general ecological functions such as herbivory as Mills and others (1993), that the full set of KECs equivalent and mutually replaceable in manage- of all species be provided rather than arbitrarily ment objectives. focusing on only a few species and KECs. We classified KECs and KEFs into two hierarchies Endemism, Biodiversity, of categories (appendices 5F, 5G). In these hierar- and Natural Areas chies, each KEC and KEF category is given a unique code and can be related to species in sev- With major contributions from expert panels, we eral levels of specificity in the SER database. For mapped centers of concentration of (1) species example, all species with KECs including the rarity and endemism and (2) high biodiversity. We general category of non-vegetation terrestrial defined centers of concentration of species rarity substrates (KEC level 3, appendix 5F) can be and endemism as locations with unusually high identified. A subset of these species would be those numbers of species that are rare or that are locally associated much more specifically with lithic or regionally endemic. We defined centers of (rock) substrates (KEC level 3.3). And then, a concentration of high biodiversity as locations subset of that subset of species would be those with unusally high numbers of species of all abun- associated much more specifically with cliffs (KEC dance classes and all types of geographic distribu- level 3.3.3). In this manner, general as well as tions. Within this framework, the expert panels specific associations of species with environmental identified centers of concentration variously based and habitat conditions can be depicted. The hier- on high species richness, high landscape diversity, archy of key ecological functions of species (KEFs, uniqueness of species communities, or high den- appendix 5G) operates the same way. Ultimately, sity of rare plant occurrences. Many of the high then, it is the full set of all species and their envi- biodiversity centers of concentration, especially in ronmental correlates and ecological functions that the plant panels, were the same as the centers of can be depicted in the SER database and that many rare and endemic species as initially defined collectively define biotic processess and by the plant expert panels. biodiversity of ecosystems. All panels identified centers of rarity and endemism in the southwestern Oregon portion of the Use of Ecologically Functional Equivalents Basin (Southern Cascades, Upper Klamath, and Do species with the same categories of key ecologi- Northern Great Basin ERUs), along the Snake cal functions perform the same ecological service? River (Owyhee Uplands ERU), the Columbia In a broad sense, by definition of functional cat- River Gorge (including the Northern Cascades

Terrestrial Map 5.3a— Vertebrate species with the key ecological function of herbivory: Meadow vole and blacktailed jackrabbit.

Terrestrial Map 5.3b—Vertebrate species with the key ecological function of herbivory: Greater white-fronted goose and Ross' goose.

Terrestrial ERU) and the desert steppe in central and south- that relatively high biodiversity can be maintained ern Washington (Columbia Plateau ERU) (table with disturbances that reduce or eliminate domi- 5.17). Centers of rarity and endemism of plants nant species. Such disturbances can include biotic and animals occurred in nearly all major land- factors such as predation, herbivory, parasitism, forms. disease, and complete emigration of a dominant competitor, or abiotic factors such as floods, fires, The centers of concentration of rarity and ende- severe windstorms, and extreme weather. mism seemed to pertain to taxonomic groups with low mobility. This may be because species with The centers of concentration of high biodiversity low mobility may be more apt to develop endemic or high occurrences of plants and aquatic species forms adapted to local conditions. Centers of were more dispersed across the Basin than were the concentration of rarity and endemism were centers of concentration of species rarity and mapped based on distributions of species of vascu- endemism, but they corresponded well to large lar plants, reptiles, amphibians, and fish. Inverte- blocks of natural areas. This may be because the brates were mapped only to family level due to large natural areas and roadless areas receive less lack of research on species taxonomy and field human disturbances and are more likely to provide ecology. Few nonvascular plant species were iden- conditions for the full set of naturally-occurring tified in this mapping exercise due to lack of field species. Two notable exceptions were the large studies. Small mammals species were included in wilderness blocks in central Idaho and the east the mapping, but large mammals were not because Cascades in Oregon, which contained few centers their larger home ranges generally precluded of high biodiversity. identifying specific sites as centers of concentra- Centers of concentration were mapped separately tion. Panelists did not identify centers of concen- for taxonomic groups of plants and animals. When tration of rarity and endemism for birds and bats overlayed, locations with three or more centers because of the high mobility of these species. defined smaller "hot spots" of rarity and ende- Instead, correlations between species and habitat mism, and high biodiversity, for plants and ani- for birds and bats were identified (SER database; mals combined. We identified 12 hot spots of TS 1995). rarity and endemism (table 5.17, map 5.4) and 7 We also listed the species associated with the hot spots of high biodiversity within the Basin centers of concentration and commented on each (table 5.18, map 5.4). Doubtless, additional hot area and possible causes for local conditions. For spots may be identified at finer scales and levels of example, we identified when an unusual soil type, geographic resolution than we used in this project. geological formation, management activity or Collectively, these hot spots mainly occurred in condition appeared to contribute to the rarity or the northern glaciated and unglaciated mountains, endemism of associated species. This documenta- central and southern Cascade mountains, Wash- tion was provided in a geographically referenced ington and Oregon plateau country, Great Basin database linked to individual centers. Some of the plains, and the Snake River valley (maps 5.4, 5.5). factors contributing to rarity and endemism were: The plant and amphibian/reptile panels were thermal hot springs, alkali pools, high topographic the only panels that identified areas of high relief that served to isolate mountain tops or biodiversity or high plant occurrences. The canyons, agricultural activities, springs, seeps, biodiversity map themes and the rarity/endemism wetlands, and riparian habitats. map themes were developed using different sets of Disturbance regimes likely play a role in maintaining criteria among the panels, for reasons explained biodiversity of some centers of concentration, above. This approach provides a starting assess- although more remains to be learned of such rela- ment for further exploration of hot spots and their tions. Huston (1994), among others, hypothesized relation to species distributions.

16:H9r"' Terrestrial Table 5.17—Hot spots of species rarity and endemism1 (see map 5.4 for location of numbered areas).

Natural Road Human Number2 Description/Location Area3 Density4 Density5 Vegetation Ownership

RE1 Washington: Columbia Basin shrub - Hanford RNA Mod., Mostly low - Desert salt shrub, Almost all private, steppe, Vantage, Washington area some low very low basin sagebrush, small BLM, no FS and Saddle Mtns. High degree of mountain big disturbance around the area due to sagebrush agriculture and grazing. Remnant steppe areas rare, populations limited. RE2 Washington/Oregon: Columbia Small: TNC High to About 1/3 in Mixed, contrast: fir, Most private, River Gorge, both sides, from reserve; SIA; west; high density, hemlock, cedar to 2 National Forests; Cascade Locks to east of Biggs, Wilderness mod.-low rest in low west; ponderosa no BLM; National north to Yakama Reservation, west north, pine, mountain big Recreation Area outside project boundary. Rapid south, and sagebrush and in Gorge elevation changes, climatic east agricultural, grass- contrasts. Gorge and river are lands north, south, corridors but highway and cliffs and east. can be barriers. RE3 Oregon: Crest of Umatilla Mtns., About 1/3 Mod-high Low-very low Grand fir, interior All Umatilla elevational changes. Many wilderness; outside ponderosa pine, National Forest, westside species are isolated 1 RNA; 1 wilderness, subalpine herb, no BLM and disjunct on mountain tops. TNC reserve none within lodgepole pine, Seeps, riparian and springs spruce/fir important. Diverse, mosaic. RE4 Oregon: Wallowa Mtn. Peaks, Almost all No roads Very low Very mixed: All Wallowa Sacajewea Peak, diverse, mosaic, wilderness; density subalpine herb, Whitman National sharp elevational ranges. Rocky Mtn. 2 RNAs within spruce/fir, grand Forest, no BLM and boreal species on Mtn. top wilderness fir, lodgepole islands, isolated. Unusual geology (limestone, greenstone, argillites), cold, snowy, many wetlands and springs. RE5 Oregon: Deschutes, Three Sisters Some Mod-low. Some areas Interior ponderosa 2 National Forests, area, special habitats due to wilderness to Small area high density pine, Douglas fir, BLM, about 1/3 volcanos and other geological west, edge of of high. east, low-vei spruce/fir, Mtn. big private/ other patterns. Wild and None in low to west sagebrush Scenic River; wild. TNC reserve; ACEC; RNA RE6 Oregon: Upper Klamath Lake, many Small: FWS High, a Mod., range Interior ponderosa Mostly National spring, alkali lakes, thermal areas, refuge; ACEC; little mod. from high to pine, agriculture, Forest, some BLM unique habitats for fish, invertebrates SIA; RNA low rangelands, and private/ other and plants. Human disturbance high sagebrush, marsh in some areas. vegetation. Table 5.17 (continued).

Natural Road Human Number2 Description/Location Area3 Density4 Density5 Vegetation Ownership

RE7 Oregon: Summer Lake Slight: edge High, very Low Interior ponderosa National Forest, (westshore), marsh areas, fresh ofWSAand little low pine, rangelands, a little BLM, and alkali spring, important to fish, SIA; RNA wheatgrass, small private/other invertebrates, plants. sagebrush, marsh vegetation. RE8 Oregon: Warner Lakes, Crump FWS refuge Mod-low Mod-low Agriculture, salt Most FWS, BLM, Lake, Hart Mtn. Refuge. Lakes, small ACEC, desert shrub, no National Forest, springs, alkali areas, important RNAs and sagebrush, small private/other to fish, invertebrates and plants. WSA edge wheatgrass, marsh vegetation. RE9 Oregon: Alvord Lake, Steens Some WSA, 2 Mod-low, Low density Salt desert shrub, Mostly 2 BLM, Mtns., alkali lakes, springs, lake, ACEC, RNA small area wheatgrasses, no National Forest, marshes, Great Basin species, of high antelope bitterbrush, some private/other insect research, high human sagebrush steppe, disturbance. agric. and rangelands RE10 Oregon: Ontario north to SmalrRNA Mod Very low Mostly agricultural, 2 BLM, Huntington, Snake River, Weiser to high, rangelands, salt private/ other, River. Sand pockets, gravels, lake about 1/3 desert shrub, no National Forest beds, extremely low diversity in in mod-high basin sagebrush, some areas, limestones, Great Basin wheatgrass arid grass/shrub communities, Snake River Canyon. RE11 Idaho: Snake River from Oregon Small ACEC, Mod-low Very low Desert salt shrub, A lot of BLM, Idaho border to Grand View, part of TNG to high, basin and Wyoming no National Forest. Bird's of Prey Area. High contrast in recreational big sagebrush, Some private/ other river canyon to surrounding. River is use agriculture and corridor and unique habitat, geology, wheatgrass steep slopes, disjunct populations from south, limestone, sand, ash, unique soils. Grazing and mining conflicts. RE12 Idaho: Hot Springs south to Bruneau. ACEC, WSA, Mod Low-very low Wheatgrass, A lot of BLM. Some Dept. of Defense. Hard bottom WSR rangelands, No National Forest. playa, clay soils. May have been desert salt shrub, Dept. of Defense, isolated in past. Some highly adapted sagebrush, private species, hot spring important to agriculture snail, inverts. 'Hot spots are defined as intersection areas of three or more centers of concentration of species' rarity or endemism among all taxonomic groups mapped. 2See map 5.4 for specific locations. 'Natural Areas: RNA = Research Natural Area, SIA = Special Interest Area, WSA = Wilderness Study Area, TNC = The Nature Conservancy reserve, FWS = Fish and Wildlife Service, WSR = Wild and Scenic River, ACEC = Areas of Critical Environmental Concern. 4Road Density: From predicted road density classes [ICBEMP GIS data (1 km2 rater data)]: None = 0.00-0.01 km/sq km, very low = 0.01-0.06 km/sq km, low = 0.06-0.44 km/sq km, moderate = 0.44-1.06 km/sq km, high = 1.06-2.92 km/sq km, extremely high = 2.92+ km/sq km. 5Human density: Low = 0-3 people/sq km, moderate = 3-6 people/sq km, high = 6-30 people/sq km. Table 5.18—Hot spots of species biodiversity1 (see map 5.4 for locations of numbered areas).

Natural Road Human Number2 Description/Location Area3 Density4 Density5 Vegetation Ownership

B1 Washington: North Cascade, eastern 1/3 None in Low Subalpine fir/spruce, Mostly National edge, Pasayten Wilderness tied into Wilderness, north, low Douglas Fir, alpine Forest, and state Cathedral Park (BC), North Cascade WSA, TNC, in south and western larch, forest, a little BLM and Manning (BC) National Parks. 2 RNA (FS, lodgepole, subalpine and private. Glaciated landscape, rounded peaks, Washington herb, Mtn. sagebrush, elevation and climatic contrast to State) interior ponderosa surroundings, wetlands, bog, pine meadows, many rare plants. Large carnivores, amphibians, BC linkage. B2 Washington: Saddle Mtn. to Yakima, A little FWS Low in 1/4 high Sagebrush/ BLM, Dept. of part of Hanford (DOE) and Yakima refuge, a east, mod- (SW), bunchgrass, Defense, Firing Range (DOD). Remnant little WSR, high in rest low- wheatgrass, Dept. of Energy, sage/steppe, desert salt shrub, a lot Hanford and south, none mod. agricultural, private/ other of reptiles, breeding birds, Yakama in middle. rangelands, amphibians. Active sand dunes, Firing Range Some desert salt shrub, seeps, talus, riverine, slopes with have some restricted bitterbrush deep soils, playa bottoms. reserve areas

B3 Washington/Oregon: Columbia Gorge Small part of High 1/4 or Fir, hemlock, cedar, 2 National Forests; from Cascade Locks to Mosier, Mt. wilderness, more in Douglas fir, oak Columbia Gorge Adams to north. Highly diverse, high FWS refuge, mod-high, woodlands, grand fir, NRA; Yakama elevation and climate contrast. SIA. Gorge rest in ponderosa pine, Reservation; Gorge/river is corridor and barrier. National low rangelands, private/other Unique plant communities, geology, Recreation agricultural, soils. Volcanic activity, spring, falls, Area; RNA sagebrush highly disturbed. West side outside ICBEMP boundary.

B4 Oregon/Idaho: Wallowa - Whitman WSR, Mod-low Low Wyoming big Mostly National National Forest to Salmon River, wilderness, recreation sagebrush, Forest, some BLM Hell's Canyon, elevation and climate RNA pressure wheatgrass, and private/other. extremes, refuge and corridor. mod-high bunchgrasses, Basalt, limestone, granite, steep. some subalpine Dry with springs/seeps/creeks as herb micro-sites. Very diverse. Heavy grazing in some areas. Table 5.18 (continued).

Natural Road Human Number2 Description/Location Area3 Density4 Density5 Vegetation Ownership

B5 Idaho: Upper Snake River from ACEC, TNC Mod, none 1/4 mod Wyoming big BLM, Dept. of Idaho Oregon border to Mt. Home in canyon high, rest sagebrush, Idaho Defense, A.F.B. Birds of Prey area. High low fescue, desert salt private/other elev. and micro-climate contrast. shrub, agriculture, High diversity, as canyon acts as wheatgrass, corridor between basin, rangelands mountains and valleys. B6 Idaho: Panhandle, St. Joe River, WSR, RNA Mod Low, Interior ponderosa Almost all Clearwater. Some checkerboard some mod. pine, cedar, Douglas National Forest, ownership. High amphibian diversity, fir, lodgepole some BLM, canyon refuge for many species private/ other from coast. Some warm, moist, low elevation. Ferns, red alder, cedar forests. Some heavy logging. B7 Montana: Flathead, Glacier Park, Most in East: low Low Spruce/fir, interior Most National Blackfoot Reservation into Alberta natural areas, to none, except in Douglas fir, Park Service, to the north. Eastern portion out of 1/2 National far west: far west, lodgepole, serai some Blackfoot ICBEMP boundary. Large carnivores, Park; WSR; mod-high and park shrub, subalpine Reservation, intact plant communities, subalpine RNA; visitor areas herb, alpine tundra, National Forest, and alpine Rocky Mtn, relatively wilderness. some agric. in west a little private. undisturbed except in lower elevations. 'Intersection areas of 3 or more centers of concentration of biodiversity among all taxonomic groups mapped. 2See map 5.4 for specific locations. 'Natural Areas: RNA = Research Natural Area, SIA = Special Interest Area, WSA = Wilderness Study Area, TNC = The Nature Conservancy reserve, FWS = Fish and Wildlife Service, WSR = Wild and Scenic River, ACEC = Areas of Critical Environmental Concern. 4Road Density: From predicted road density classes [ICBEMP CIS data (1 km2 rater data)]: None = 0.00-0.01 km/sq km, very low = 0.01-0.06 km/sq km, low = 0.06-0.44 km/sq km, moderate = 0.44-1.06 km/sq km, high = 1.06-2.92 km/sq km, extremely high = 2.92+ km/sq km. 5Human density: Low = 0-3 people/sq km, moderate = 3-6 people/sq km, high = 6-30 people/sq km. Fewer centers of concentration and hot spots were shrub steppe communities, which have declined delineated in the eastern portion of the Basin than rapidly in this century due to agriculture, livestock in the western portion for several reasons. First, grazing, roads, noxious weeds, seeding, and hu- there were fewer available inventories and research man settlement. There also appear to be some studies of rare and endemic species in the eastern critical habitat areas in southern Oregon that may portion, especially northeast Washington and have little protection for maintaining hot spots of central Idaho, than in the western portion. Sec- rarity and endemism. ond, there appear to be more centers of concentra- The Columbia River Gorge had the greatest con- tion of species rarity and endemism in transition vergence of centers of concentration of rarity and areas between the Pacific coast and the east side of endemism, as well as many centers of high the Cascades Mountains, and between the north- biodiversity. The unique topographic, geologic, ern Great Basin and the southern Rocky Moun- and climatic extremes in this area and the dispersal tains, with fewer areas in the heart of the Rocky corridor and barrier caused by the Columbia River Mountians and central Idaho which have more and Gorge have created high species diversity and uniform environments. Third, the panel members a high concentration of rare and endemic species. may have had greater experience in Washington The area also may have received a higher degree of and Oregon environments than further east in the scientific inventory and study than other areas assessment area. Although an effort was made to because of its proximity to Portland, and its scenic represent expertise from Idaho, Montana, Utah, beauty, accessibility, and history. Tracts of private and Wyoming, many panelists were from Oregon. lands have been acquired by the Forest Service for Of the 12 hot spots of rarity and endemism, nine addition to the National Recreation Area. Inten- occurred wholly or partly in Oregon, two on the sive visitor use could threaten this sensitive area. Upper Snake River of the Owyhee Uplands and The rugged terrain provides some degree of natu- one in the Columbia steppe in Washington (map ral protection to sites within the gorge, but the 5.4). Only three of these hot spots of rarity and adjacent uplands have experienced greater human edemism (Umatilla Plateau Mountains, Wallowa disturbance. Mountains, and Snake River) had 25 percent or The Upper Snake River and Saddle Mountain areas more of their area within designated existing of the Owhyee Uplands also appear as hot spots for natural areas. The remaining nine hot spots had rarity, endemism, and biodiversity. The Upper Snake about 10 percent or less within natural areas. This River is similar to the Columbia Gorge, it has high means that most of the hot spots of rarity and topographic diversity, high contrast in climate and endemism are not necessarily being managed to elevation, and unique geology. Like the Columbia protect the species and conditions identified by River and Gorge, the Snake River and Gorge the panels. also serve as both a major dispersal corridor The Umatilla Mountains, Wallowa Mountains, and a dispersal barrier for various species. and Deschutes and Snake River areas appeared to be rarity and endemism hot spots because topographic, climatic, and geologic conditions have created unique and isolated habitats. The remaining areas also have some unique habitat features, especially thermal springs, alkali lakes, and marshes in southern Oregon, but they also contained rare remnants of native desert scrub and

Terrestrial 1S23 Map 5.4— Distribution of hot spots of species rarity and endemism and centers of biodiversity, with Ecological Reporting Units. These are locations that are particularly rich in species of plants, invertebrates, and vertebrates. See tables 5.17 and 5.18 for descriptions of each site. Map 5.5-- Distribution of centers of concentration of species rarity and endemism of plants and animals.

Terrestrial 1625 The Upper Snake area, where Great Basin species Some Ecological Reporting Units (for example, converge with Columbia Basin species, has high Central Idaho Mountains and Northern Glaciated biodiversity, but it also has a high degree of species Mountains) have large natural areas, whereas rarity and endemism because of the unique habi- others (for example, Columbia Plateau, Upper tats and the effects of management activities in Snake) have only very small and isolated natural surrounding uplands. The Saddle Mountain Area areas, and throughout the Basin the locations of is a refuge for some of the last intact areas of desert natural areas do not consistently correspond to salt shrub and sagebrush steppe, which have been centers of concentration or hot spots of converted within the Basin to agriculture and biodiversity or species rarity or endemism. livestock range. The Handford Energy Reserve and Based on home range sizes of vertebrates, even the Yakima Firing Range have served to protect some smallest natural areas (less than or equal to 50 ha) areas of native sagebrush steppe from develop- could support at least one individual, and perhaps ment, but this area contains virtually no BLM- or small populations, of 70 percent of vertebrate FS-administered lands. species (fig. 5.8). Based on area alone, natural areas Hot spots for biodiversity overlapped more with larger than 650 hectares could support 90 percent existing natural areas, especially wilderness areas, of the vertebrate species from the region; approxi- than did hot spots of species rarity and endemism. mately 50 percent of the natural areas are larger The natural areas tended to be in the upper eleva- than 650 hectares. Natural areas would have to be tion, forested portions of the hot spots, leaving at at least 10,000 hectares in area before 99 percent least some of the lower elevation portions of the of the vertebrate species could be supported. Only hot spots unprotected from ground-disturbing 16 percent of the natural areas are larger than human activities. For plants and large carnivores, 10,000 hectares. there is likely a correlation between centers of biodiversity and natural areas or roadless areas.

Natural Areas Characterization Natural areas fall into 26 categories of existing land allocations on FS- and BLM-administered lands, with varying management objectives, selec- tion criteria, allowed uses, size ranges, and agency or organization administration. In the four western states most encompassed by the Basin (that is, eastern Oregon, eastern Washington, most of Idaho, and western Montana), Federal land management agencies administer a total of 41.63 million hectares. On these lands, 28 percent or 11.72 million hectares occur in natural areas (TS 1995).

Natural areas throughout the Basin were desig- Figure 5.8 — Cumulative percentages of vertebrate species nated under a single management objective or by home range area, as compared with cumulative per- strategy, in contrast to more consistent approaches centages of natural areas by size. to natural area management in British Columbia, Canada, and in New Zealand, India, or other countries (Joshi and Gadgil 1991, Pressley 1994).

1826 Terrestrial Expectations of species occurrence based on home classes. Some species are closely associated with range size does not necessarily mean that a particu- single biophysical factors, although many species lar size natural area would contain viable popula- are likely correlated with multiple factors. Most tions of all associated species. Natural areas would ERUs had at least some unique species, although have to be several times larger than the area of an many species overlapped several units. A few individual home range of most species to support additional examples here help illustrate some enough individuals for a viable population. The general principles of species biogeography within area multiplier necessary for sustaining viable the Basin. populations would depend on the population structure, life history, habitat relation, and behav- Landforms ior of the species. Habitat composition, pattern, disturbance regimes, and connectivity through- Nine general landform categories have been identi- out surrounding landscapes also are critical fied in the Basin. In some cases, significant biogeo- considerations. graphic correlates with some species distributions are readily apparent by simply matching species Of the 389,009 kilometers of streams in the Basin, distribution with these landforms. An example of 51,293 kilometers (13%) occur within existing a species that tightly associates with highland natural areas. Most streams are within wilderness landforms (glaciated and unglaciated mountains or wilderness study areas with the headwater, high and breaks) is the carpenter ant (map 5.6). Like- gradient and steep portions of streams over- wise, pronghorn shows alignment of its distribu- represented as compared to the low gradient, lower tion with lowland landforms (map 5.7) as well as elevation portions. Only 7 of the 132 4th-field with areas of low precipitation (map 5.8); and Hydrologic Units [HUC] inventoried did not have lowland landforms and low precipitation are at least 1.6 kilometers of stream within a natural correlated. The distribution of pronghorn also area. An average of 386 kilometers of stream demonstrates another simple biogeographical occurred per 4th-field HUC. Some categories of principle, that species can be shown to be distrib- natural areas, namely the research natural areas, uted according to several biophysical factors, areas of environmental concern, special interest themselves correlated. areas, and wild and scenic river areas, have been established specifically to protect only a portion of Disjunct Distributions a stream, lake or river. These areas are generally small and linear and do not include protection of Another example of biogeography of species in the uplands that may be critical to maintaining the Basin pertains to disjunct distributions. In some health of the aquatic system. There are 2,150 lakes cases, species such as mountain goat (maps 5.9a, b, c) covering 97,242 hectares within natural areas. that are associated with highland landforms appear as several disjunct populations because of the Biogeography disjunct distribution of such landforms within the Basin. The mountain goat has been mapped as Much of the above discussion of species groups has having 12 disjunct population centers in the touched upon various aspects of biogeography of Basin. The mountain goat seems closely tied to species. In particular, reasons for distribution and high elevation and high topographic relief, specifi- abundance and response of population to broad- cally with the highest elevation marked in each scale biophysical attributes has been discussed. We 6th-field HUC as mapped in the Basin. identified species with disjunct, peripheral, scattered, regionally endemic, and locally endemic distributions. We also identified some species closely associated with some of the nine landform

Terrestrial 162t The map of mountain goat distribution and high- in map 5.10a and 5.10b (appendix 5H). In these lands (map 5.9a) also illustrates a lesson in how examples, many of the species are restricted to map resolution affects interpretation of species specific ranges because of their high habitat speci- biogeography. At the broadscale used in this assess- ficity. One example is Coeur d'Alene salamander, ment (1 km2 map cells), east-central Idaho is which selects sites with talus, seeps, and spray zones shown as very high elevation and thus as poten- of waterfalls, with a slope of 10 to 60 percent, tially suitable habitat for, and within the range of, above elevations of 400 meters. Suitable sites mountain goat. At a finer resolution, however, this usually have at least 25 percent canopy cover. At area contains three valleys separated by narrow the broadscale, they are found in fractured belt mountain ranges (Big Lost River-Challis Creek rock formations in the northern Rocky Mountain valleys, Little Lost River-Pahsemeroi River valleys, extension in northern Idaho and northwestern and Birch Creek-Lemhi River valleys). It is un- Montana. At the mid-scale within these sites, they likely that any mountain goats that may occur in select for appropriate elevations, slopes, vegetation the inner ranges would travel between these three cover, and stream or waterfall proximity. ranges.36 Thus, at a finer resolution, there may Ultimately, distributions of local endemics can be appear smaller and more localized, disjunct goat a result of contracted ranges due to habitat loss or populations than were indicated by the broad-scale extirpations, overall scarcity of suitable environ- information. ments, or other factors. The extent to which gla- Species with disjunct ranges pose interesting bio- ciation and paleoclimates played a role in geographic questions. Some, such as the longnose determining current distributions of locally en- snake (see map 5.2b), are poorly known. The demic populations, particularly of invertebrates longnose snake's disjunct pattern in the southern and plants, in the Basin is not well studied. periphery of the Basin and along the Snake River Endemics tend to be less studied than other spe- is as yet unexplained. It selects for open or closed, cies, at least among vertebrates. The median confi- low and medium shrub environments, sometimes dence levels in scientific knowledge as ranked by near agricultural land and urban areas, and prob- the expert panelists were significantly lower for the ably occurs in a wider range of habitat types than 39 locally or regionally endemic vertebrate species current literature suggests. It may be that its appar- than for all other vertebrates (P<0.001, log- ent disjunct distribution is an artifact of inad- likelihood ratio test, G=859.27, DF=3). equate sampling. The same may be true, to some extent, with the California mountain kingsnake and western ground snake (map 5.2b). On the Other Patterns of Endemism other hand, the disjunct pattern of wild turkey and Distribution (map 5.2b) most likely is due to selective introduc- Plants and allies and vertebrates were specifically tions in the Basin. coded in the SER database according to the five classes of species distribution (that is, scattered, Local Endemics regionally endemic, locally endemic, peripheral, Distributions of collective "hot spots" of locally and disjunct). Species of plants and allies included endemic species were discussed above. Specific in this study are those listed as threatened, examples of locally endemic vertebrates are shown endangered, or candidate, or those suggested by the plant expert panels for such listings. 36Personal communication, 1995. Allen E. Thomas, Fish and Wildlife Biologist. U.S. Department of Interior, Bureau of Land Management, Idaho State Offic, 3380 Americana Terrace, Boise, Idaho 83706.

Terrestrial