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BLM LIBRARY 88072144 QK 629 . C787 N68 2008 c. 3 Phaeocollybia of Pacific Northwest North America Lorelei L. Norvell & Ronald L. Exeter United States Department of Interior Bureau of Land Management Salem District 1717 Fabry Road SE Salem, Oregon 97306 November 2008 ISBN-13: 978-0-9791310-1-1 ISBN-10: 0-9791310-1-4 Authors: Lorelei L. Norvell is a professional mycologist, Editor-in-Chief of Mycotaxon, the International Journal of Fungal Taxonomy & Nomenclature, and president of the Pacific Northwest Mycology Service. 6720 NW Skyline Boulevard Portland, Oregon 97229 Ronald L. Exeter is a botanist for the Mary’s Peak Resource Area, Salem District Bureau of Land Management. 1717 Fabry Road SE Salem, Oregon 97306 Design and Layout: Lorelei Norvell Front Cover: Phaeocollybia kaujfmanii in Jackson State Forest, California Photograph courtesy of Scott Redhead, Agriculture & Agri-Food Canada BLM/OR/WA/GI-08/100-1792 |D Abstract Norvell, Lorelei L., Exeter, Ronald L. 2008. Phaeocollybia of Pacific Northwest North America. USDI BLM/OR/WA/GI-08/100-1792, Salem, Oregon 228p. As a result of the 1994 presidentially mandated Record of Decision, United States Bureau of Land Management (BLM) and Forest Service planning documents specified guidelines for surveying for ‘rare and ‘uncommon fungal species in United States forests within the range of the northern spotted owl. This document listed all but one of the described Phaeocollybia species then known to occur in the Pacific Northwest, believed to be the most highly diverse region for the genus in the world. Later taxonomic revision of the genus by the senior author — combined with data obtained from BLM-Pacific Northwest Mycology Service cooperative research and examination of hundreds of specimens collected by Northwest Forest Plan forest surveyors — led to the naming of ten new Phaeocollybia species. This publication offers a key to all described 25 Phaeocollybia species from Pacific Northwest United States (California, Idaho, Washington, Oregon) and Canada (British Columbia). Detailed summary descriptions are accompanied by color photos depicting each species in the field, in the lab, and under the microscope. Discussions of global distribution, ecology, development, biology, taxonomy, and suspected phylogenetic relationships offer essential background information to those working within and outside the Pacfic Northwest region. A glossary and complete bibliography to the phaeocollybia literature are also provided. Key words: Phaeocollybia, Cortinariaceae, Agaricales, fungi, Northwest Forest Plan BLM Library Denver Federal Center Bldg. 50, OC-521 P.O. Box 25047 Denver, CO 80225 ITEM HAS BEEN DIGITIZED € 4 i Color Bar Key A) Distribution & Ecology B) Development & Biology C) Taxonomy & Phylogeny D) Diagnostics E) Conspectus & Keys to species F) Species descriptions G) Bibliography H) Glossary < Phaeocollybia of Pacific Northwest North America Contents Distribution and ecology.2 Development and biology . 14 Taxonomy and phylogeny . 22 Diagnostics: key characters and identification procedures . 27 Conspectus of Pacific Northwest species of Phaeocollybia. 37 Key to Pacific Northwest Phaeocollybia species . 38 Microkey to western North American Phaeocollybia species. 41 Species descriptions . 45-204 Phaeocollybia ammiratii .45 Phaeocollybia attenuata.51 Phaeocollybia benzokauffmanii .57 Phaeocollybia californica.63 Phaeocollybia dissiliens .71 Phaeocollybia fallax.77 Phaeocollybia gregaria.85 Phaeocollybia kauffmanii .91 Phaeocollybia lilacifolia .99 Phaeocollybia luteosquamulosa .107 Phaeocollybia ochraceocana .113 Phaeocollybia olivacea.119 Phaeocollybia oregonensis .125 Phaeocollybia phaeogaleroides.131 Phaeocollybia piceae .137 Phaeocollybia pleurocystidiata 143 Phaeocollybia pseudofestiva 149 Phaeocollybia radicata .155 Phaeocollybia redheadii .161 Phaeocollybia rifflipes .169 Phaeocollybia rufotubulina .175 Phaeocollybia scatesiae .181 Phaeocollybia sipei .187 Phaeocollybia spadicea . 193 Phaeocollybia tibiikauffmanii . 199 Acknowledgments .205 Bibliography .206 Glossary .213 Tables Table i. World species of Phaeocollybia .3-4 Table 2. Phaeocollybia species according to region .6 Table 3. Global and Oregon state rarity rankings.9 Table 4. Phaeocollybia Pacific Northwest collection sites.10 Table 5. Pseudorhizal morphology in PNW phaeocollybias. 17 Table 6. Phaeocollybia sections.26 Table 7. Spore print color, fluorescence, syringaldazine reactivity and odor in PNW phaeocollybias.29 Table 8. Major key characters of PNW phaeocollybias.33 Table 9. Morphology & ecology in P. californica, P rufotubulina & P scatesiae.69 Table 10. Diagnostic characters in the Phaeocollybia festiva complex.83 Table 11. Diagnostic characters in the Phaeocollybia kauffmanii complex.96 '’'Ron Exeter^ 2 ... Introduction — Distribution and ecology of Phaeocollybia Introduction Phaeocollybia species is provided in Tables 1 and 2. The brief summary below provides an additional Phaeocollybia is a genus of brown-spored agarics overview. easily recognized in the field by their deeply rooting Great Britain and Europe — Although the genus cartilaginous stipes and normally viscid conic- was first described from Europe, Phaeocollybia is campanulate pilei. Phaeocollybia species were represented by only six currently accepted species originally included among a heterogeneous assemblage reported from 13 countries (Austria: Moser 1950; of brown-spored species of Agaricus that Fries (1838) Czech republic & Slovakia: Vesely et al. 1972, Mann grouped within the subtribe Gymnoti in the Tribe 1991; Finland: Karsten 1879, 1881; France: Bon Naucoria. Among eleven species placed together in 1979, 1992, Konrad & Maublanc 1948, Kiihner 1980, an unnamed group characterized by gills that were Kiihner & Romagnesi 1957, Labarre 1997, Robin 2007; narrowly attached (‘nearly free’) to the stipe were five Germany: Krieglsteiner 1991,Laber 1982, 1991;great that would form the core of Phaeocollybia a century Britain: Dennis & al. 1960, Watling & Gregory 1993; later: two species first described by Fries in 1821 — Italy: Bresadola 1930; Norway: Reid 1972, Gulden Agaricus lugubris and Agaricus hilaris — and three also 1983, 1992, 2008; Romania: Pazmany & Laszlo 1987; described by him in 1838 — A. christinae, A. cidaris, Spain: Bandala & Montoya 2003, Olariaga & al. 2004; and A. festivus. When Naucoria was later elevated to Sweden: Jacobsson & Stridvall 1983; Switzerland: generic level (Kummer 1871, Quelet 1872), the genus Horak 1968, Breitenbach & Kranzlin 2000; Ukraine: retained its essentially heterogeneous character. Karpenko 1998. For the continent, see also Heim Heim (1931) segregated Phaeocollybia from Naucoria 1931; Singer 1951b, 1986; Bresinsky 1960; Lange & for brown agarics with cartilaginous ‘rooting’ stipes, Hora 1963; Poelt & Jahn 1963; Horak 1977, 2005; viscid to glutinous (‘slimy’) caps, brown ornamented Moser 1983; Moser & Jiilich 1994.) Widely distributed spores, cheilocystidia (sterile cells on gill edges), and throughout Europe, phaeocollybias are encountered gelatinous tissues. Heim transferred the ‘core’ species only sporadically and are restricted to coniferous named above to his new genus and suggested that four forests, particularly those containing spruce (Picea). other species assigned to Simocybe by Karsten (1881, Asia — Four European and five endemic species have including S. jennyae) probably should be placed in been reported from eastern Asia and India (China: Phaeocollybia as well. Heim did not select a type species Bresadola 1930, Keissler & Lohwag 1937, Liu 1995, for his genus, which another author (Konrad 1934) Teng 1996, Zang 1996; India: Horak 1974, 1980, lectotypified by designating his recently neotypified Abraham 1992; Japan: Imai 1938, Imazeki & Hongo Phaeocollybia lugubris as type species. After Heim and 1988; Siberia: Nezdoimnogo 1986, 1990. See also Konrad, the genus was critically revised by Kiihner Horak 1977; Singer 1951b, 1986). Throughout Asia, & Romagnesi (1957), Smith (1957b), Horak (1977), phaeocollybias are reported only from boreal and Singer (1970,1986,1987), Redhead & Malloch (1986), temperate coniferous forests. Bon (1992), Bandala (1994), Bandala et al. (1996), and Norvell (1998a). Australasia — There are seventeen species reported from Australasia (Australia: Fuhrer & Robinson World distribution and ecology 1992, Horak 1983, May & Wood 1997, Rees & Syme 1999, Rees & Wood 1996; malaysian archipelago: Singer was the first to recognize distribution of Corner 1994; new Zealand: Horak 1973, 1977. See Phaeocollybia species outside Europe when he also Singer 1986.) All species are endemics except transferred North American and Chinese naucorias for one possible exception. Horak (1977) placed his (i.e., Smith’s N. kauffmanii in 1940 and N. radicata, N. species Phaeocollybia longipes (Horak 1973) into attenuata, and N. similis in 1951) to Heim’s new genus. synonymy with P. festiva, despite its association with Since that time, new species have been recognized Sphagnum and Nothofagus. Future type studies may from forests on every continent except Africa and show that P. longipes is also indigenous to New Zealand. Antarctica. Distribution of currently recognized