Management Recommendations•Group 12

Rare Gilled Mushroom: Chroogomphus loculatus Trappe & Miller
Rare False Truffle: Gautieria magnicellaris (Pilat) Stewart & Trappe, Gautieria otthii Trog, Leucogaster citrinus (Harkness) Zeller & Dodge, Leucogaster microsporus Fogel nom. prov.
False Truffle: Nivatogastrium nubigenum (Harkness) Singer & Smith
Undescribed Taxa: Octavianina cyanescens Trappe & Castellano nom. prov. (=Octavianina sp. nov. #Trappe 7502)
Rare False Truffle: Octavianina macrospora Singer & Smith, Octavianina papyracea Singer & Smith, Sedecula pulvinata Zeller

TABLE OF CONTENTS

EXECUTIVE SUMMARY

Species: Chroogomphus loculatus Trappe & Miller, Gautieria magnicellaris (Pilat) Stewart & Trappe, Gautieria otthii Trog, Leucogaster citrinus (Harkness) Zeller & Dodge, Leucogaster microsporus Fogel in ed., Nivatogastrium nubigenum (Harkness) Singer & Smith, Octavianina cyanescens Trappe & Castellano nom. prov. (=Octavianina sp. nov. #Trappe 7502), Octavianina macrospora Singer & Smith, Octavianina papyracea Singer & Smith, Sedecula pulvinata Zeller

Taxonomic Group: Fungi

ROD Component(s): 1 & 3

Other Management Status: none

Range: Chroogomphus loculatus is known from only one site which occurs in the western Cascade Mountains in Oregon. Gautieria magnicellaris is known from 2 sites from the western Cascade Mountains in Oregon and in the Klamath Mountains in northern California. Gautieria otthii is known from 2 sites from Siskiyou Mountains in southern Oregon and Klamath Mountains in northern California. Leucogaster citrinus is known from the Klamath Mountains in northern California north to the Puget Sound Trough in western Washington. Leucogaster microsporus is known from the western Cascade Mountains in southern Washington south along the western flank of the Cascade Mountains in Oregon and also in southwest Oregon in the Coast Ranges. Nivatogastrium nubigenum is known from high elevation in Northern California north along the Cascade Mountains crest to McKenzie Pass in Oregon. Octavianina cyanescens is known from a single site located in the western Cascade Mountains. Octavianina macrospora is known from a single located near Mt. Hood in Oregon. Octavianina papyracea is known from along the coast in northwest California. Sedecula pulvinata is known from disjunct populations in Colorado, Idaho, Lassen Volcanic National Park in California, and Mt. Shasta in California.

Specific Habitat: All these taxa except Nivatogastrium nubigenum form sequestrate sporocarps in soil that develop and mature beneath the surface of the ground and form ectomycorrhizae with members of the Pinaceae. Nivatogastrium nubigenum forms sequestrate sporocarps that develop and mature on the surface of rotten Abies logs. Chroogomphus loculatus is found in association with the roots of assorted Pinaceae, particularly Tsuga mertensiana. Gautieria magnicellaris is found in association with the roots of Pinus sp. in Mexico and Abies concolor in western North America at high elevation. Gautieria otthii is found in association with the roots of Pinus ponderosa and other Pinaceae at mid to high elevation. Leucogaster citrinus is found in association with the roots of Abies lasiocarpa, Pseudotsuga menziesii, Pinus contorta, and Tsuga heterophylla at low to high elevation. Leucogaster microsporus is found in association with the roots of Pseudotsuga menziesii and Tsuga heterophylla at mid elevation. Nivatogastrium nubigenum is found on Abies logs at high elevation (above 4,000 ft.). Octavianina cyanescens is found in association with the roots of Tsuga mertensiana at high elevation (5,700 ft.). Octavianina macrospora is found in association with the roots of Tsuga heterophylla. Octavianina papyracea is found in association with the roots of Pinaceae in forests dominated by Sequoia sempervirens at low elevation (below 2,000 ft.). Sedecula pulvinata is found in association with the roots of Abies concolor, A. lasiocarpa, A. magnifica, Picea engelmannii, and Pinus contorta at high elevation (above 6,000 ft.).

Threats: Logging is the most serious threat to Chroogomphus loculatus, Gautieria magnicellaris, Gautieria otthii, Leucogaster citrinus, Leucogaster microsporus, Nivatogastrium nubigenum, Octavianina cyanescens, Octavianina macrospora, Octavianina papyracea, and Sedecula pulvinata. In addition, actions that disturb soil, coarse woody debris or remove overstory hosts could eliminate populations of some or all the above taxa.

Management Recommendations: Maintain habitat for Chroogomphus loculatus, Gautieriamagnicellaris, Gautieria otthii, Leucogaster citrinus, Leucogaster microsporus, Nivatogastrium nubigenum, Octavianina cyanescens, and Octavianina macrospora at known Federal sites by retaining forest structure and soil conditions. Avoid disturbance at known Federal sites, including modification of canopy until additional data is collected on taxon viability. There are no known sites of Octavianina papyracea and Sedecula pulvinata on Federal land so a goal for management of these taxa could be to identify likely habitat on Federal land that may support populations and protect it until sufficient surveys occur to reveal populations.

Information Needs: Revisit known sites of all taxa and collect ecological data to more completely characterize habitat. Conduct inventories, particularly in late-successional reserves, Research Natural Areas and when appropriate where management treatments or projects are scheduled or proposed.

I. NATURAL HISTORY

A. Taxonomic/Nomenclatural History

Chroogomphus loculatus was originally described by Trappe and Miller (1970) from Oregon, Lane Co., Williamette National Forest, Lamb Butte Scenic Area, on the saddle at the head of Potholes Creek. There are no known synonyms. It is a member of the family Gomphidiaceae in the order Boletales.

Gautieria magnicellaris was originally described by Pilat (1953) as Gautieria morchelliformis var. magnicellaris from Czechoslovakia. Stewart and Trappe (Cazares et. al 1992) elevated it to species rank. The only know synonym is Gautieria morchelliformis var. magnicellaris Pilat. It is a member of the Hysterangiaceae in the order Phallales.

Gautieria otthii was originally described by Trog (1857) from Hardlisberg, Switzerland. Zeller and Dodge (Dodge & Zeller 1934) reduced it to a variety as Gautieria graveolens var. otthii. We prefer to accept this taxon at the species level as Gautieria otthii. It is a member of the Hysterangiaceae in the order Phallales.

Leucogaster citrinus was originally described as Leucophleps citrina by Harkness (1899) from Mt. Tamalpais, Marin Co., California. Zeller and Dodge (1924) transferred it to Leucogaster. There are no known synonyms. It is a member of the Leucogastraceae in the order Leucogastrales.

Leucogaster microsporus has not been formally published yet but is described in the Ph.D. thesis of R. Fogel (1975). It is described from near Sheep Creek, Linn Co., Oregon. There are no known synonyms. It is a member of the Leucogastraceae in the order Leucogastrales.

Nivatogastrium nubigenum was originally described as Secotium nubigenum by Harkness(1899) from California. Singer and Smith (1959) transferred it to the genus Nivatogastrium where it is more appropriately placed. There are no known synonyms. It is a member of the family Strophariaceae in the order Agaricales.

Octavianina cyanescens is currently in the process of being published. It is listed in the FEMAT report, ROD, and the known site database as Octavianina sp. nov. #Trappe 7502. There are no known synonyms. It is a member of the family Octavianinaceae in the order Octavianinales.

Octavianina macrospora was originally described by Singer and Smith (1960) from Twin Bridges forest camp, Clackamas Co., Oregon. There are no known synonyms. It is a member of the family Octavianinaceae in the order Octavianinales.

Octavianina papyracea was originally described by Singer and Smith (1960) from Spruce Cove, near Trinidad, Humboldt Co., California. There are no known synonyms. It is a member of thefamily Octavianinaceae in the order Octavianinales.

Sedecula pulvinata was originally described by Zeller (1941) from Mt. Shasta, California. There are no known synonyms. It is a member of the family Sedeculaceae in the order Seducuales.

B. Species Description

This is an artificial assemblage of fungal taxa grouped together that form similarly structured sporocarps.

1. Morphology

Chroogomphus loculatus is characterized by the smoky black spores, its non-gelatinous cuticle and pigmented pileus tissue which is amyloid in Melzer's solution, and the loculate to strongly intervenose nature of the hymenophore.

Pileus 2.5-7.0 broad, convex, margin expanding fully or sometimes remaining almost unexpanded, dry, with appressed or repent fibrillose scales which are dark olive over a pale orange to pale yellow ground color, pale brown. Context fleshy, pale orange. Hymenophore very irregular forming enclosed chambers due to numerous intervenose connections, clearly loculate, decurrent, pale pink at first darkening to brown orange in age. Stipe 2.5-8 cm long, 1.5-3.5 cm wide, somewhat ventricose tapering to a dull pointed base, often fused to 3/4 or more of total length, dry, pale orange or vinaceous tinged above annular zone, below pallid to yellow, streaked with olivaceous fibrils, olive tomentose over the base. Context fleshy, pale orange but often olive stained near the base. Partial veil fibrillose remaining in some as an obscure fibrillose zone. Spores (15-) 19-30 x 6-9 µm, subfusiform in profile, ovate to elongate-ovate in face view, slightly thickened walls (1 µm), deep red, ochraceous to yellow contents in Melzer's solution; brown black spore print. Basidia 48-72 x 11-15 µm, clavate, thin-walled, 4-spored, hyaline in Melzer's solution and KOH. Pleurocystidia and cheilocystidia 108-200 (-260) x (13-) 17-29 µm, numerous, nearly cylindrical, long fusiform to elongate clavate, walls thin to thick (up to 1.5-3 µm), hyaline to pale yellow contents in Melzer's solution, with pale yellow walls, protruding 1/2 to 2/3 above the hymenium, sometimes partially covered with dingy yellow brown encrusted material. Cuticle of nonviscid, innate, loosely tangled to erect, thin-walled hyphae (7.0-) 13-22 µm diam, walls somewhat roughened, either amyloid or nonamyloid to yellow brown, contents of all cells appear hyaline. Tram of pileus of more tightly interwoven hyphae 13-30 µm diam, thin-walled, amyloid. Trama of locule walls of interwoven hyphae 5-20 µm diam, thin-walled, darkly amyloid as in pileus trama. Mycelium surrounding the base of the fruiting bodies pink, hyphae 6.5-8.5 µm diam, thin- or thick-walled, amyloid. Clamp connections absent on sporocarp hyphae but present on hyphae of basal mycelium.

Gautieria magnicellaris is characterized by the ovate to elongate-ovate, large spores, and exposed gleba.

Basidiocarps as dried 11-24 x 9-27 mm, globose, subglobose, irregularly lobed, flattened or depressed at the point of rhizomorph attachment. Glebal chambers large and fully exposed in early development. Rhizomorph basal, single, ±1.5 mm diam. Peridium drying brown from the exposed spore mass. Exposed tramal tissue mostly drying thin and obscured by spores, where visible pale yellow to red brown. Chemical reactions: KOH, spore mass moderate red brown, quickly fading, trama no reaction; FeSO4, spore mass green gray, gray yellow green; Melzer's reagent and ETOH, no reaction. Glebal spore mass drying brown to red brown, trama tissue drying pale yellow and red brown. Chemical reactions same as peridium. Chambers 0.5-10 x 0.5-6 mm, labyrinthiform to rounded, some continuous through the entire sporocarp, empty. Columella in mature sporocarps not prominent, white, drying light orange yellow. Spores 17-24 x 12-15 (-18) µm including sterigmal appendage and epispore, 17-24 x 8-12 µm excluding epispore, ellipsoid to broadly ellipsoid. Spore wall ±1.5 µm thick. Sterigmal appendage 2-3 µm at the base of epispore, 1-3 µm long. Episporeprominent, of 9-13 forking and anastomosing, longitudinal to occasionally spiraling ridges, (1-) 2-3.5 (-4) µm tall, 3-5 µm broad, varying some in width on a single spore; margins smooth to cleft or humped with occasional projecting cogs; grooves smooth to slightly serrate. Ridges extending 1-3.5 µm beyond the spore apex. Neither converging at the apex nor covering the sterigmal appendage. Color as mounted in KOH, pale green yellow singly, yellow in mass; FeSO4, dark yellow singly, moderate olive in mass; Melzer's, orange singly, red orange in mass; ETOH, no reaction; cotton blue, slowly cyanophilous. Basidia 40-60 x 11-16 µm, extending up to 50 µm beyond hymenial layer, thin-walled, broadly clavate, those with immature spores attached filled with refractive inclusions that disappear after spore maturity with four conical sterigmata 2-2.5 x 2-2.5 µm. Brachybasidioles 15-33 x 7-9 µm, cystidioid elements infrequent. Subhymenium a palisade of subpolygonal cells 9-32 x 7-13 µm, thin-walled, merging into intertwined hyphae of the trama. Trama of hyaline, thin-walled, septate hyphae, 2-7 µm broad, weakly gelatinized with age; oleiferous hyphae rare, detectable in Melzer's reagent. Trama plus hymenium 200-360 µm broad. Color as mounted in KOH, hyaline; FeSO4, gray yellow green; Melzer's reagent and ETOH, no reaction; cotton blue, cyanophilous. Columella of hyaline hyphae inflated up to 10 µm adjacent to septa with spines ±1 µm projecting from the inner wall towards the interior. Peridium (margins of exposed glebal chambers) of tiers of napiform cells 7-30 x 10-25 µm. Clamp connections lacking.

Gautieria otthii is characterized by the small spores and short sterigmata.

Basidiocarps up to 5 cm diam. when fresh, globose to subglobose, flat or depressed; immature fruiting bodies nearly smooth, but glebal cavities exposed in mature fruiting bodies. Peridium white with red tones, but emergent apical surfaces becoming brown with patches of ochre to yellow; when dry yellow brown, brown where glebal chambers are exposed. Chemical reactions: KOH, immediately a faint dark red orange; FeSO4, pale green; Melzer's reagent, quickly moderate red brown, soon fading in intensity; ETOH, no reaction. Gleba pale brown to yellow brown, drying pale brown to yellow brown; trama drying hard and somewhat brittle. Chemical reactions: KOH, spores red brown, trama no reaction; FeSO4, pale green; Melzer's reagent, immediately brown, fading in intensity; ETOH, no reaction. Chambers up to 0.5 x 2 mm, labyrinthiform, generally radiating away from the stipe, empty. Columella gray white, cartilaginous, drying red brown, up to 2 mm broad, branching near the base. Spores 13-18 x 7-10 µm including sterigmal appendage and epispore, 13-18 x 5-7 µm excluding epispore, ellipsoid, often somewhat flattened at the apex. Spore wall ±1 µm thick. Sterigmal appendage truncate, prominent, 1 µm broad a sterigmal scar, broadening to 2 µm at the base of epispore. Epispore of 9-12, sometimes forking, longitudinal ridges, 0.5-1.5 (-2) µm tall, 2-3 µm broad, ridge margins humped to cleft, grooves between ridges smooth to finely crenulate, epispore extending beyond the spore apex 1-2 µm, neither converging at the apex nor covering the sterigmal appendage. Color as mounted in KOH, pale green yellow singly, yellow in mass; FeSO4, yellow gray singly, gray olive in mass; Melzer's reagent, red orange singly, dark red orange in mass; slowly cyanophilous. Basidia (12-) 18-30 (-45) x 6-9 µm, thin-walled, clavate, with four sterigmata, usually extending not more than 15 µm beyond cystidioid elements 15-30 x 3-5 µm, clavate. Subhymenium of subpolygonal cells 10-30 x 5-13 µm, progressively smaller as the subhymenium merges into the linear hyphae composing the trama. Trama narrow, of hyaline, thin walled septate hyphae 3-5 µm diam., gelatinizing in age; oleiferous hyphae present. Trama plus hymenium 120-350 µm broad. Color as mounted in KOH, hyaline; FeSO4, yellow gray; Melzer's reagent, no reaction; cyanophilous. Columella of hyaline hyphae 1-5 µm diam, inflated to 18 µm adjacent to septa; inflated areas filled with granules and spines ±1 µm long, projecting from the inner wall surface toward the interior. Peridium 65-150 µm thick, of thick walled napiform cells 17-40 µm diam.

Leucogaster citrinus is characterized by its yellow peridium, small spores with tall ornamentation and lack of inflated cells in the trama.

Basidiocarps up to 20-45 mm, reniform, pale yellow to dark yellow, with 1 to several coarse rhizomorphs emanating from base, concolorous with peridium. KOH on peridium pale brown pinkthen brown, Ethanol negative, FeSO4 faintly gray. Gleba white, trama white, locules spherical, 0.5-3 mm broad, empty, smaller toward peridium. Consistency cartilaginous after drying. Peridium 60-200 µm thick, of periclinal, compact, pallid yellow hyphae 2-4 µm diam. Red pigment when mounted in KOH. Trama 60-80 µm thick, of subparallel, orange, thin-walled, gelatinous hyphae, 2-5 µm in diam; tramal hyphae gelatinous, hyaline, septate, 3 µm in diam, loosely interwoven. Basidioles 100-125 x 5 µm, claviform, hyaline, thin-walled. Apobasidia 100 x 5-7 µm, clavate, 4-spored, thin-walled, sterigmata lacking. Spores 8-11 x (7-) 8-9 µm, including spinose-reticulate ornamentation, subglobose, hyaline, enclosed in hyaline perisporal sac; ornamentation 1-1.5 µm high, 2-2.5 µm apart, spine base 0.2 µm wide, spore wall 1 µm thick, excluding ornamentation.

Leucogaster microsporus is characterized by its small spores, inflated cells in the peridium and trama, and a rather thick peridium.

Basidiocarps up to 40 x 50 mm, reniform, base indented with peridial tissue extending to the center of the sporocarp, glabrous, white with some yellow stains, drying pale red. Gleba white, drying pale yellow pink, exuding white, sticky latex when cut, locules spherical, 0.5-1 mm broad, empty in center of sporocarp, filled with spores next to peridium. Rhizomorphs absent. Chemical reactions; peridium KOH pale yellow, FeSO4 negative, FeSO4 + ETOH dull blue-violet. Odor sweet. Peridium 100-300 µm thick, of periclinal, pale yellow, thin-walled hyphae, 3-5 µm in diam, cells becoming inflated to 15 µm, some oleiferous, yellow, thin-walled hyphae. Outer 60 µm obscured in KOH by yellow, amorphous, pigment balls in Melzer’s reagent. Trama 60-125 µm thick, of subparallel, yellow, septate, thin-walled hyphae, 2-3 µm in diam, cells becoming inflated to 10 µm. Basidioles 30-35 x 3-4 µm, claviform, thin-walled, hyaline, some encrusted with a hyaline crystalline deposit in KOH. Apobasidia 30-60 x 6-7 µm, claviform, 4-spored, thin-walled, hyaline, sterigmata lacking. Spores 6-10 x 5-6 µm, including reticulate ornamentation, subglobose, enclosed in a hyaline perisporal sac; ornamentation of reticulate spines 0.25-0.5 µm high, 2 µm apart, with low connecting lines forming 5-6 sided alveoli.

Nivatogastrium nubigenum is characterized by its agaricoid, sweet-smelling sporocarps and inamyloid, honey colored spores which possess a germ pore.

Basidiocarps 10-24 mm broad, 15-40 mm tall, subglobose to more or less truncate, margin abruptly decurved, rarely umbonate, sometimes longitudinally compressed. Peridium not separating from the stipe-columella, or only rarely breaking in age and then in an irregular manner; surface glabrous, smooth, somewhat viscid; color evenly ochraceous or with streaks or darker areas of tawny, fading to white in age. Gleba loculate to sublamellate, pale red brown, not pulverulent. Stipe 5-25 mm long, 5-12 mm thick, equal to somewhat bulbous, sometimes narrowed into a cylindric columella, somewhat matted fibrillose over the exposed surface, dry and unpolished, solid or rarely tubular, with white rhizomorphs at base, columella percurrent, solid, white, veil white or whitish, arachnoid and cortina-like, causing the margin of the peridium and the apex of the stipe to be somewhat silky-shining in age, eventually evanescent. Odor fragrant sweet, taste mild. Peridium 15-50 µm thick, with an epicutis of gelatinous, narrow, interwoven, hyaline hyphae, subcutis of subparallel, thin-walled, irregularly enlarged hyphae, up to 12 µ in diam. Veil consisting of slightly gelatinized hyphae. Context of stipe and columella of interwoven hyphae with cells often enlarged up to 25 µm, hyphae in the columella hyaline, those in the lower portion of the stipe colored, throughout both structures loosely arranged and with numerous air spaces, not gelatinized. All hyphae inamyloid; clamp connections present. Trama of regularly arranged, subhyaline to yellow, interwoven hyphae, some cells irregularly enlarged. Subhymenium up to 50 µm thick, of inflated to subisodiametric cells 10-25 (-33) x 8-18.5 (-25) µm, interior to this layer occurs a layer of somewhat gelatinized hyphae, up to 15 µm in diam. Basidia 17-21 x 6-8.2 µm, hyaline, clavate, 2-4-spored, sterigmata apical or nearly so, straight to very slightly curved, cylindric to narrowly conic. Basidioles common, vesiculose, about 16 x 12.5 µm. Cystidia 60-100 x 15-25 µm, pale brown yellow to brown, thin-walled, fusoid-ventricose, the neck long and in optical section with undulating walls, the apex obtuse to subacute. Spores (3-) 7.5-9 (-12) x (3-) 5.5-6.3 µm, smooth,ellipsoid, asymmetrical, germ pore very narrow and apex not truncate; wall thick, in KOH golden ochraceous to honey colored, inamyloid.

Octavianina cyanescens is characterized the bluing reaction of the peridium and its relatively darkly pigmented and tall spore ornamentation.

Basidiocarps subglobose to ± flattened, 12-25 mm in diam, felty, yellowish white to pale yellow, in places with a scant overlay of dark brown fibrils, overnight after collection developing dark blue stains where bruised; rhizomorphs lacking. Gleba firm, moist, dark grayish brown to brownish black from the color of the massed spores, with gray tramal plates separating minute, rounded, mostly spore-filled chambers. Odor not distinctive. Taste not recorded. Peridium 150-300 µm thick; epicutis a narrow layer of appressed, light brown, thin-walled hyphae 2-7 µm in diam at the septa, occasional cells inflated to 5-10 µm; subcutis with pale brown, thin-walled hyphae near the epicutis grading to hyaline near the gleba, 2-6 µm in diam at the septa but commonly inflated to 10-30 µm. Clamp connections not found. Glebal trama of hyaline, thin-walled hyphae 2-4 µm in diam at the septa, the cells often inflated to 4-8 µm. Subhymenium of hyaline, thin-walled, ± isodiametric cells 5-10 µm in diam. Basidia in a hymenium, clavate, hyaline, thin-walled, 30-35 x 11-14 µm, the sterigmata up to 5-7 x 1.5-2 (-3) µm. Clamp connections not found. Spores dark brown, globose to occasionally broadly ellipsoid, 13-18 (-20) µm in diam excluding the ornamentation of subangular cones (3-) 4-5 (-7) x 1-6 µm in KOH, 2-4 x 1-6 µm in water, with occasional interspersed spines and smaller cones, all cones composed of agglutinated spines and forming 4-7 sided polygons at the base, sterigmata often remaining attached as an inconspicuous pedicel; in Melzer's reagent spore walls deep orange red, ornamentation hyaline and only 2-4 µm tall, heated in cotton blue the spines forming the cones separating and becoming erect, 4-8 x 1.5-3 µm.

Octavianina macrospora is characterized by thick-walled, inamyloid spores and a peridial turf of cystidioid elements.

Basidiocarps up to 2 cm diam, white when fresh, glabrous; gleba of open chambers, presumably white when fresh; sterile base present and concolor with gleba as preserved. Peridium with an epicutis consisting of a collapsed turf of clavate to cystidioid cells 18-27 x 4-8 µm or 20-30 x 3.5 µm if fusoid. Subcutis of hyaline, interwoven, subgelatinous hyphae, 3-6 µm in diam, sphaerocysts or oleiferous hyphae absent. Clamp connections absent. Basidia 1-2 spored, 29-37 x 10-13 µm, clavate, thin-walled, content granular and ochraceous in KOH. Cystidia none. Subhymenium hyphal about the diameter of the base of the basidium, 1-2 cells below the basidium the cells enlarging in age to sphaerocysts; mediostratum hyphal, thin-walled, hyaline, sphaerocysts absent. Spores 17-23 x 12-16 µm, broadly ellipsoid, wall 1.5-2 µm thick; pale buff in KOH, inamyloid; sterigmal appendage sometimes conspicuous, often slightly irregular in outline; ornamentation of spines 1-1.5 (2.5) x 0.5 µm, the spines distinct but often touching at the base.

Octavianina papyracea is characterized by inamyloid spores and a sporocarp that exudes a latex when cut.

Basidiocarps 1 cm diam, brown, globose, unchanging; gleba of minute locules, near pink buff as dried, with an abundant cream colored latex when cut, taste mild; columella absent. Peridium as dried almost like a thin nut shell, the exterior surface pale red brown, epicutis of a layer of appressed, tan hyphae with some free hyphal ends at surface as a rudimentary turf; hyphal ends 3-4 µm in diam, context of hyaline, subgelatinous hyphae with slightly thickened walls, interwoven, laticiferous hyphae present; sphaerocysts absent; clamp connections absent. Trama of interwoven, hyaline, inamyloid, subgelatinous hyphae with slightly thickened walls; subhymenium of cells 10-13 µm in diam in rows of 2-3 and rows terminated by a basidium or basidiole. Basidia 20-32 x 9-13 µm, thin-walled, hyaline, 1-2 spored, sterigmata 10 µm or more long. Cystidia scattered, 46-60 x 10-13 µm, thin-walled, hyaline, subcylindric to enlarged near apex and with an apical projection. Spores 14-17 µm, hyaline, inamyloid, globose to subglobose, sterigmal appendage inconspicuous, thick-walled ±1 µm, ornamentation of dense, unfused spines 2-3 µ long and 0.5-1 µm at the base.

Sedecula pulvinata is characterized by the unique spores, powdery gleba at maturity, and pulvinate sporocarp.

Basidiocarps pulvinate, up to 6 cm wide, up to 4 cm tall, surface smooth to granular with adherent soil, white to gray. Peridium tough, leathery, 2-3 mm thick above, thin and almost evanescent below, margin of thicker peridium usually somewhat rolled under so as sometimes to elevate the basal or under side having the thinner peridial cover, composed of white parenchyma of large cells with strongly gelatinized walls; tramal plates extend centipetally to unequal depths toward the base. Gleba coarsely loculate, black except for the white to gray trama becoming powdery at maturity. odor and taste mild. Trama 1-2 mm thick. Locules labyrinthine, 1-12 x 1-3 mm, lined with powdery spores. Basidia narrow-clavate, 2-spored, sterigmata about as long as the spores, slender. Spores dark brown in KOH, nonamyloid, ovoid to somewhat ellipsoid or irregular, usually very short-pedicelate, 23-26 x 13-16.2 µm.

2. Reproductive Biology

All taxa are sequestrate fungi and thus are presumed to be dependent on mycophagy for dispersal of spores. Sequestrate fungi have sporocarps that have evolved from having exposed hymenia and forcibly discharged spores to a closed or even hypogeous habit in which the spores are retained in the sporocarp until it is consumed by an animal vector or it decays. Sequestrate fungi encompasses an artificial grouping of various genera or taxa from different families that are dependent on mycophagy for spore dispersal. Mycophagy is the consumption of fungi by animals. No specific information on reproductive biology is available for any of these taxa at this time.

3. Ecology

All taxa, except Nivatogastrium nubigenum, are presumed ectomycorrhiza formers. Mycorrhiza is the symbiotic, mutually beneficial association between a fungus and plant root. This highly interdependent relationship involves the translocation of mineral nutrients and water by the fungus to the host plant while the fungus obtains photosynthetic carbon from the host plant. Some mycorrhizal associations are strongly host specific. Many plants depend upon mycorrhizal fungi for adequate uptake of nutrients and survival in nature. Likewise mycorrhizal fungi depend upon their host plant for carbohydrate. No specific ecological information is available for any of these taxa at this time except that all these taxa form ectomycorrhiza with Pinaceae.

C. Range, Known Sites

Chroogomphus loculatus is known from a single site within the range of the northern spotted owl in Oregon: Lane Co., Williamette National Forest, Lamb Butte Scenic Area, at saddle at the head of Potholes Creek at 4,300 ft. elevation.

Gautieria magnicellaris is known from 2 sites within the range of the northern spotted owl: Oregon: Lane Co., Williamette National Forest, West Lava campground at 5,000 ft. elevation and California: Siskiyou Co., Klamath National Forest, Deadfall Meadows at 6,000 ft. elevation. This taxon was originally described from Europe and is also uncommonly found in Mexico and in the United States in Michigan and New York.

Gautieria otthii is known from 2 sites within the range of the northern spotted owl: Oregon: Josephine Co., Dutcher Creek at 2,400 ft. elevation and California: Siskiyou Co., Klamath National Forest, Marble Mt. Wilderness Area, trail to Haypress Meadows at 5,000 ft. elevation. This taxon is also rare in Switzerland.

Leucogaster citrinus is known from 7 sites within the range of the northern spotted owl: California: Siskiyou Co., Klamath National Forest, Russian Wilderness Area, near Duck Lake at 6,000 ft. elevation; Oregon: Benton Co., Siuslaw National Forest, north fork of Rock Creek at 800 ft. elevation; Benton Co., just south of the Alsea summit on highway 34 at 800 ft. elevation; Curry Co., Siskiyou National Forest, Wild Rouge Wilderness Area, Upper Stair Creek at 3,500 ft. elevation; Linn Co., Willamette National Forest, Yellow Bottom campground at 1,600 ft. elevation; Washington: Thurston Co., Fort Lewis Military Reservation, Farley Block, stand 4; Thurston Co., Fort Lewis Military Reservation, Stellar Block, stand 3.

Leucogaster microsporus is known from 7 sites within the range of the northern spotted owl: Oregon: Wasco Co., Mt. Hood National Forest, near Summit Lake rd. at 3,000 ft. elevation; Marion Co., Mt. Hood National Forest, headwaters of Clackamas River, along Cub Creek; Benton Co., Siuslaw National Forest, Mary’s Peak rd. at 2,800 ft. elevation; Lane Co., Willamette National Forest, H.J. Andrews Experimental Forest, Stand 11 at 1,950 ft. elevation; Linn Co., Willamette National Forest, Sheep Creek Canyon at 2,000 ft. elevation; Curry Co., Siskiyou National Forest, Panther Lake, Long-term Ecosystem Productivity study site ESLW block at 3,000 ft. elevation; Washington: Lewis Co., Gifford Pinchot National Forest, Quartz Creek Big Trees.

Nivatogastrium nubigenum is known from 13 sites within the range of the northern spotted owl: California: Siskiyou Co., Klamath National Forest, Marble Mt. Wilderness Area, English Peak at 5,000 ft. elevation; Siskiyou Co., Mt. Shasta, Horse camp at 8,000 ft. elevation; Siskiyou Co., Shasta-Trinity National Forest, Bunny Flat; Siskiyou Co., Shasta-Trinity National Forest, Sand Flats at 6,000 ft. elevation; Siskiyou Co., Shasta-Trinity National Forest, Panther Meadow at 7,500 ft. elevation; Siskiyou Co., Shasta-Trinity National Forest, Red Butte at 7,500 ft. elevation. Oregon: Klamath Co., Crater Lake National Park, below Goodbye Creek campground at 6,000 ft. elevation; Klamath Co., Crater Lake National Park, Mazama campground; Klamath Co., Crater Lake National Park, Lost Pond at 6,150 ft. elevation; Lane Co., Willamette National Forest, near McKenzie Pass, Frog Camp at 4750 ft. elevation; Deschutes Co., Deschutes National Forest, Tumalo Mt. trail, 0.5 miles north of highway 46 at 7,000 ft. elevation; Deschutes Co., Deschutes National Forest, Three Creeks Lake at 6,800 ft. elevation; Klamath Co., Deschutes Co., Deschutes National Forest, Odell Butte. There are also 5 sites outside the assessment area but on Federal land located in California on the Lassen Volcanic National Park.

Octavianina cyanescens is known from a single site within the range of the northern spotted owl in Oregon: Lane Co., Willamette National Forest, English Mt. above The Potholes at 5,700 ft. elevation.

Octavianina macrospora is known from a single site within the range of the northern spotted owl in Oregon: Clackamas Co., Mt. Hood National Forest, Twin Bridges campground.

Octavianina papyracea is known from 2 sites within the range of the northern spotted owl in California: Humboldt Co., near Trinidad, Spruce Cove and Humboldt Co., Redwoods State Park, near Orick, Fern Canyon.

Sedecula pulvinata is known from a single site within the range of the northern spotted owl in California: Siskiyou Co., Mt. Shasta at 7,000 ft. elevation. This taxon is also known from Lassen Volcanic National Park, Lassen National Forest and Sierra National Forest in California; Payette National Forest in Idaho; and from Enchanted Meadow in Boulder Co., Colorado.

D. Habitat Characteristics and Species Abundance

All these taxa, except Nivatogastrium nubigenum, form sequestrate sporocarps in soil that develop and mature beneath the surface of the ground.

Nivatogastrium nubigenum forms sequestrate sporocarps that develop and mature on the surface ofrotten Abies logs.

Chroogomphus loculatus is found in association with the roots of assorted Pinaceae, particularly Tsuga mertensiana.

Gautieria magnicellaris is found in association with the roots of Pinus sp. in Mexico and Abies concolor in the western North America at high elevation.

Gautieria otthii is found in association with the roots of Pinus ponderosa and other Pinaceae at mid to high elevation.

Leucogaster citrinus is found in association with the roots of Abies lasiocarpa, Pseudotsuga menziesii, Pinus contorta, and Tsuga heterophylla at low to high elevation.

Leucogaster microsporus is found in association with the roots of Pseudotsuga menziesii and Tsuga heterophylla at mid elevation.

Nivatogastrium nubigenum is found in association with the roots of Abies logs at high elevation (above 4,000 ft.).

Octavianina cyanescens is found with Tsuga mertensiana at high elevation (5,700 ft.).

Octavianina macrospora is found in association with the roots of Tsuga heterophylla.

Octavianina papyracea is found in association with the roots of Pinaceae in forests dominated by Sequoia sempervirens at low elevation (below 2,000 ft.).

Sedecula pulvinata is found in association with the roots of Abies concolor, A. lasiocarpa, A. magnifica, Picea engelmannii, and Pinus contorta at high elevation (above 6,000 ft.).

II. CURRENT SPECIES SITUATION

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

Potentially all taxa are at risk from management and recreational activities that remove the mycorrhizal host or disturb the soil.

Chroogomphus loculatus is found within the range of the northern spotted owl from a single population. Under Option 9, this taxon was considered to have a 5 percent likelihood of being well-distributed throughout its range 50 percent likelihood of being locally restricted, 28 percent likelihood of restriction to refugia, and 18 percent likelihood of extirpation on Federal lands. This taxon is believed to be at high risk under the Northwest Forest Plan because of its rarity and dependent mycorrhizal association with old-growth legacy Tsuga mertensiana at high elevation.

Gautieria magnicellaris is found within the range of the northern spotted owl from 2 populations. Under Option 9, this taxon was considered to have a 0 percent likelihood of being well-distributed throughout its range, 35 percent likelihood of being locally restricted, 50 percent likelihood of restriction to refugia, and 15 percent likelihood of extirpation on Federal lands. This taxon is believed to be at high risk under the Northwest Forest Plan because of its rarity and dependent mycorrhizal association with old-growth legacy Abies concolor at high elevation.

Gautieria otthii is found within the range of the northern spotted owl from 2 populations. Under Option 9, this taxon was considered to have a 0 percent likelihood of being well-distributed throughout its range, 35 percent likelihood of being locally restricted, 50 percent likelihood of restriction to refugia, and 15 percent likelihood of extirpation on Federal lands. This taxon is believed to be at high risk under the Northwest Forest Plan because of its rarity and dependentmycorrhizal association with old growth legacy, mid to high-elevation Pinus ponderosa and other Pinaceae.

Leucogaster citrinus is found within the range of the northern spotted owl from 7 populations. Under Option 9, this taxon was considered to have a 0 percent likelihood of being well-distributed throughout its range, 35 percent likelihood of being locally restricted, 50 percent likelihood of restriction to refugia, and 15 percent likelihood of extirpation on Federal lands. This taxon is believed to be at high risk under the Northwest Forest Plan because of its rarity and dependent mycorrhizal association with old growth legacy, low to high-elevation Abies lasiocarpa, Pseudotsuga menziesii, Pinus contorta, and Tsuga heterophylla.

Leucogaster microsporus is found within the range of the northern spotted owl from 7 populations. Under Option 9, this taxon was considered to have a 0 percent likelihood of being well-distributed throughout its range, 35 percent likelihood of being locally restricted, 50 percent likelihood of restriction to refugia, and 15 percent likelihood of extirpation on Federal lands. This taxon is believed to be at high risk under the Northwest Forest Plan because of its rarity and dependent mycorrhizal association with old growth legacy, mid-elevation Pseudotsuga menziesii and Tsuga heterophylla.

Nivatogastrium nubigenum is found within the range of the northern spotted owl from 13 populations. Under Option 9, this taxon was considered to have a 55 percent likelihood of being well-distributed throughout its range, 25 percent likelihood of being locally restricted, 20 percent likelihood of restriction to refugia, and 0 percent likelihood of extirpation on Federal lands. This taxon is believed to be at high risk under the Northwest Forest Plan because of its rarity and dependent saprophytic association with old growth legacy, high elevation Abies spp.

Octavianina cyanescens is found within the range of the northern spotted owl from a single population. Under Option 9, this taxon was considered to have a 0 percent likelihood of being well-distributed throughout its range, 0 percent likelihood of being locally restricted, 60 percent likelihood of restriction to refugia, and 40 percent likelihood of extirpation on Federal lands. This taxon is believed to be at high risk under the Northwest Forest Plan because of its rarity and dependent mycorrhizal association with old growth legacy, high-elevation Tsuga mertensiana.

Octavianina macrospora is found within the range of the northern spotted owl from a single population. Under Option 9, this taxon was considered to have a 0 percent likelihood of being well-distributed throughout its range, 2 percent likelihood of being locally restricted, 83 percent likelihood of restriction to refugia, and 15 percent likelihood of extirpation on Federal lands. This taxon is believed to be at high risk under the Northwest Forest Plan because of its rarity and dependent mycorrhizal association with old growth legacy, high-elevation Tsuga heterophylla.

Octavianina papyracea is found within the range of the northern spotted owl from 2 populations. Under Option 9, this taxon was considered to have a 0 percent likelihood of being well-distributed throughout its range, 2 percent likelihood of being locally restricted, 83 percent likelihood of restriction to refugia, and 15 percent likelihood of extirpation on Federal lands. This taxon is believed to be at high risk under the Northwest Forest Plan because of its rarity and dependent mycorrhizal association with old growth legacy, low-elevation Sequoia sempervirens.

Sedecula pulvinata is found within the range of the northern spotted owl from a single population. Under Option 9, this taxon was considered to have a 0 percent likelihood of being well-distributed throughout its range, 35 percent likelihood of being locally restricted, 50 percent likelihood of restriction to refugia, and 15 percent likelihood of extirpation on Federal lands. This taxon is believed to be at high risk under the Northwest Forest Plan because of its rarity and dependent mycorrhizal association with old growth legacy, high-elevation Abies concolor, A. lasiocarpa, A. magnifica, Picea engelmannii, and Pinus contorta.

B. Major Habitat and Viability Considerations

The major viability consideration for Chroogomphus loculatus, Gautieria magnicellaris, Gautieria otthii, Leucogaster citrinus, Leucogaster microsporus, Nivatogastrium nubigenum, Octavianina cyanescens, Octavianina macrospora, Octavianina papyracea, and Sedecula pulvinata is loss of the known populations within the range of the northern spotted owl. Considerations include all management or recreational activities that disturb the soil or duff.

Relatively little is known about the autecology of Chroogomphus loculatus, Gautieria magnicellaris, Gautieria otthii, Leucogaster citrinus, Leucogaster microsporus, Nivatogastrium nubigenum, Octavianina cyanescens, Octavianina macrospora, Octavianina papyracea, and Sedecula pulvinata. They are presumed mycorrhiza formers of a restricted group of Pinaceae species. Therefore disturbance that affects the host will potentially strongly affect these taxa.

Fire is a potential threat to populations of Chroogomphus loculatus, Gautieria magnicellaris, Gautieria otthii, Leucogaster citrinus, Leucogaster microsporus, Nivatogastrium nubigenum, Octavianina cyanescens, Octavianina macrospora, Octavianina papyracea, and Sedecula pulvinata because particularly a hot ground fire could harm populations from disturbance to soil or by damaging or killing host trees.

Climate change may result in decline in vigor of these taxa and may result in the extirpation of these taxa from the range of the northern spotted owl. Climate change could potentially impact all populations of these taxa. An increase in temperature or a decrease in precipitation could affect disjunct populations.

C. Threats to the Species

Threats to Chroogomphus loculatus, Gautieria magnicellaris, Gautieria otthii, Leucogaster citrinus, Leucogaster microsporus, Nivatogastrium nubigenum, Octavianina cyanescens, Octavianina macrospora, Octavianina papyracea, and Sedecula pulvinata are those actions that disrupt stand conditions necessary for their survival, particularly damage to host trees and soil disturbance. These include logging that removes its presumed mycorrhizal or saprophytic host and other actions that cause disturbance to the soil or large woody debris, particularly road, trail, and campground construction.

These taxa are not routinely harvested for use as food.

D. Distribution Relative to Land Allocations

The only known population of Chroogomphus loculatus is in a late-successional reserve: Lamb Butte Scenic Area, Williamette National Forest.

Gautieria magnicellaris is known from one site on the Klamath National Forest that is congressionally withdrawn. The other site within the assessment area is from the Williamette National Forest and is designated as a late-successional reserve.

Gautieria otthii is known from one site on the Klamath National Forest that is congressionally withdrawn. The other known site is not on Federal land.

Leucogaster citrinus is known from 4 sites that are congressionally withdrawn: Russian Wilderness Area, Wild Rouge Wilderness Area, and both sites on the Fort Lewis Military Reservation. One site is located within a late-successional reserve on the Willamette national Forest. One site is administratively withdrawn; Siuslaw National Forest. The site near the Alsea summit is not on Federal land.

Leucogaster microsporus is known from 5 sites that are located within late-successional reserves: Siuslaw National Forest, Sheep Creek Canyon on the Willamette National Forest, Cub Creek on the Mt. Hood National Forest, Siskiyou National Forest, and Gifford Pinchot National Forest. One site is on matrix land: Summit Lake rd. on the Mt. Hood National Forest. One site is in an adaptive management area on the H.J. Andrews Experimental Forest.

Nivatogastrium nubigenum is known from 5 sites that are congressionally withdrawn: all 3 sites on Crater Lake National Park, Red Butte on the Shasta-Trinity National Forest, and English Peak within the Marble Mt. Wilderness Area on the Klamath National Forest. Three sites are within late-successional reserves; Bunny Flat and Sand Flats on the Shasta-Trinity National Forest and Three Creeks Lake on the Deschutes National Forest. Three sites are on matrix land: Panther Meadow on the Shasta-Trinity National Forest, Odell Butte and Tumalo Mt. trail on the Deschutes National Forest. One site is on administratively withdrawn land on the Willamette National Forest. One site is not on Federal land: Horse camp on Mt. Shasta.

Octavianina cyanescens is known from one site located within a late-successional reserve.

Octavianina macrospora is known from one site that is administratively withdrawn.

Octavianina papyracea is known from 2 sites, both are on non-Federal land.

Sedecula pulvinata is known from one site on Mt. Shasta but the specific locality is not mentioned thus land allocation can not be determined.

III. MANAGEMENT GOALS AND OBJECTIVES

A. Management Goals for Taxon

The goal for management of Chroogomphus loculatus, Gautieria magnicellaris, Gautieria otthii, Leucogaster citrinus, Leucogaster microsporus, Nivatogastrium nubigenum, Octavianina cyanescens, Octavianina macrospora, Octavianina papyracea, and Sedecula pulvinata is to assist in maintaining viable populations of these taxa within the assessment area. Known sites of these rare taxa should be protected until sufficient information is generated to suggest management will not result in extirpation of these taxa.

B. Specific Objectives

Maintain habitat conditions at all known sites on Federal land for Chroogomphus loculatus, Gautieria magnicellaris, Gautieria otthii, Leucogaster citrinus, Leucogaster microsporus, Nivatogastrium nubigenum, Octavianina cyanescens, Octavianina macrospora, Octavianina papyracea, and Sedecula pulvinata.

IV. HABITAT MANAGEMENT

A. Lessons from History

There has not been any management of sites for any of these taxa. Since all taxa are presumptive mycorrhiza formers, an abundance of potential hosts must be protected where fungal populations exist. When mycorrhiza host trees are damaged or removed a negative impact is usually reflected in the population of the fungal partner. Although not documented for these taxa, many fungi are harmed by air pollution, acid deposition, N deposition, and SOx (Gulden et al., 1992).

B. Identification of Habitat Areas for Management

The only known site of Chroogomphus loculatus has good potential to be managed to maintain viability of the population.

The only known sites of Gautieria magnicellaris within the assessment area have good potential to be managed to maintain viability of the population.

Only the site of Gautieria otthii located on the Klamath National Forest has good potential to be managed to maintain viability of the population. The other known site is on non-Federal land.

Six of the 7 sites of Leucogaster citrinus within the assessment area have good potential to be managed to maintain viability of the population. There is one site that is not on Federal land.

Six of the 7 sites of Leucogaster microsporus within the assessment area have good potential to be managed to maintain viability of the population. One site, Summit Lake rd. on the Mt. Hood National Forest, is on matrix land.

Six of 13 sites of Nivatogastrium nubigenum within the assessment area have good potential to be managed to maintain viability of the population. Three sites are on National Park land. One site is not on Federal land. The 3 sites on matrix land should also be strongly considered for management as they include Panther Meadows on Mt. Shasta that harbors a large population, and two sites on the Deschutes National Forest that are on the periphery of the taxon range.

The only known site of Octavianina cyanescens has good potential to be managed to maintain viability of the population.

The only known site of Octavianina macrospora has good potential to be managed to maintain viability of the population.

Octavianina papyracea is not known from any Federal lands within the range of the northern spotted owl..

Only one known site of Sedecula pulvinata occurs within the range of the northern spotted owl. The exact location of the this population is unknown.

The seemingly preferred habitat of these taxa are also somewhat under-collected and in critical need of survey. New populations may be found with additional surveys.

C. Management Within Habitat Areas

Status of management activities is unknown for extant sites. However, at and around known sites, it is recommended that current habitat conditions and micro-climatic conditions be maintained, impacts from soil disturbing activities minimized, and damage or removal of host trees prevented.

The few known locations of Chroogomphus loculatus, Gautieria magnicellaris, Gautieria otthii, Leucogaster citrinus, Leucogaster microsporus, Nivatogastrium nubigenum, Octavianina cyanescens, Octavianina macrospora, Octavianina papyracea, and Sedecula pulvinata should be managed to include an area that is large enough to maintain the habitat and associated micro-

climate of the populations. The Regional mycologist is available to consult with field staff and managers on the size of the appropriate area for management.

• Maintain dominance of specific host tree associates as outlined in section 1D, which are necessary for mycorrhizal association.
• Minimize loss, disruption or compaction of soil particularly from management, road construction or recreational activities.
• Manage tree diseases in the area to minimize loss of host trees.

D. Other Management Issues and Considerations

No additional management issues or considerations are identified at this time.

V. RESEARCH, INVENTORY, AND MONITORING NEEDS

A. Data Gaps and Information Needs

Revisit known sites of all taxa and collect ecological data to more completely characterize habitat. Conduct surveys to locate additional populations of Chroogomphus loculatus, Gautieria magnicellaris, Gautieria otthii, Leucogaster citrinus, Leucogaster microsporus, Nivatogastrium nubigenum, Octavianina cyanescens, Octavianina macrospora, Octavianina papyracea, and Sedecula pulvinata particularly in late-successional reserves, Research Natural Areas and when appropriate where management treatments or projects are scheduled or proposed.

Data are lacking regarding the specific response of these taxa to management practices such as logging, road, trail, and campground construction, prescribed fire and collection of secondary forest products. Also needed are information on each fungus taxon concerning the area required to support viable populations, population age structure, dispersal requirements and maximum distance over which populations can interact. Exact host tree associations for each fungus taxon need documentation.

B. Research Questions

• What is the ecological amplitude of Chroogomphus loculatus, particularly is it restricted to association with Tsuga mertensiana?
• What is the ecological amplitude of Gautieria magnicellaris, particularly is it restricted to association with Abies concolor?
• What is the ecological amplitude of Gautieria otthii, particularly is it restricted to association with Pinus ponderosa?
• What is the ecological amplitude of Leucogaster citrinus, particularly is it restricted to association with Abies lasiocarpa, Pseudotsuga menziesii, Pinus contorta, and Tsuga heterophylla?
• What is the ecological amplitude of Leucogaster microsporus, particularly is it restricted to association with Pseudotsuga menziesii and Tsuga heterophylla?
• What is the ecological amplitude of Nivatogastrium nubigenum, particularly is it restricted to association with Abies logs?
• What is the ecological amplitude of Octavianina cyanescens, particularly is it restricted to association with Tsuga mertensiana?
• What is the ecological amplitude of Octavianina macrospora, particularly is it restricted to association with Tsuga heterophylla?
• What is the ecological amplitude of Octavianina papyracea, particularly is it restricted to association with Sequoia sempervirens?
• What is the ecological amplitude of Sedecula pulvinata, particularly is it restricted to association with Abies concolor, A. lasiocarpa, A. magnifica, Picea engelmannii, and Pinus contorta?
• How do all taxa respond to manipulation or modification of the micro-climate?
• What is the density of tree hosts needed to sustain viability of the populations of all taxa?
• What is the range of suitable microclimate conditions, particularly humidity, soil moisture, and soil temperature for these taxa?
• What is the relationship of stand age to population viability for all taxa?

C. Monitoring Needs and Recommendations

Known sites on Federal land of all taxa should be revisited periodically to assess compliance with management guidelines and evaluate impacts.

VI. REFERENCES

Cázares, E., J. García, J. Castillo, and Trappe, J.M. 1992. Hypogeous fungi from northern Mexico. Mycologia 84:341-359.

Dodge, C.W., and Zeller, S.M. 1934. Hymenogaster and related genera. Ann. Mo. Bot. Gard. 21:625-708.

Gulden, G., K. Hoiland, K. Bendiksen, T.E. Brandrud, B.S. Foss, H.B. Jenssen, and D. Laber. 1992. Macromycetes and Air Pollution: Mycocoenological studies in three oligotrophic spruce forests in Europe. Bibliotheca Mycologica 144: 1-81.

Harkness, H.W. 1899. Californian hypogeous fungi. Proc. Calif. Acad. Sci. Third series 1(8) 241-292.

Miller, Jr., O.K., and Trappe, J.M. 1970. A new Chroogomphus with loculate hymenium and a revised key to section Floccigomphus. Mycologia 62:831-836.

Pilat, A. 1953. Sydowia 7:10.

Singer, R. and Smith, A.H. 1959. Studies on secotiaceous fungi - V. Nivatogastrium gen. nov. Brittonia 11: 224-228.

Trog, J.G. 1857. Naturforsch. Ges. Bern Mitt. p. 43.

Zeller, S.M., and Dodge, C.W. 1924. Leucogaster and Leucophleps in North America. Ann. Mo. Bot. Gard. 11:389-410.