A 200-year history of arctic and alpine fungi in North America: Early sailing expeditions to the molecular era

Figures & data

Figure 1. Fleshy fruiting bodies of Rocky Mountain alpine fungi. (a) Arrhenia lobata, decomposer on Salix. (b) Rhizomarasmius epidryas, decomposer on Dryas octopetala. (c) Galerina, decomposer on moss. (d) Lichenomphalia basidiolichen. (e) Melampsora epitea, rust pathogen on Salix reticulata. (f) Lactarius lanceolatus, ectomycorrhizal with S. reticulata. (g) Russula nana, ectomycorrhizal with dwarf Salix. (h) Cortinarius absarokensis, a North American ectomycorrhizal endemic

Figure 2. (a) Typical North American alpine habitat, Beartooth Plateau, Rocky Mountains. (b) Alpine bog. (c) Betula nana, ectomycorrhizal. (d) Salix arctica, ectomycorrhizal. (e) Salix reticulata, ectomycorrhizal

Table 1. Mushrooms and fleshy fungi collected during early North American Arctic expeditions, 1819–1931. Names and authorities as reported

Expedition	Species reported	Location	Citation
Captain William Edward Parry’s expedition (1819–1820)	Basidiomycota: Cantharellus lobatus Peck and Lycoperdon pratense Pers.	Melville Island, Canadian Arctic Archipelago	Brown (1823)
Captain Sir John Franklin’s second overland expedition (1825–1827)	Basidiomycota: Cantharellus canadensis Kl. MSS., Favolus (Pleuropus) humboldtii Berk., Favolus (Pleuropus) hepaticus Kl., and Favolus (Pleuropus) canadensis Kl. l. c.	Northwest Territories and arctic regions in Canada	Berkeley (1839)
Arctic expedition led by Admiral Belcher in (1852–1854)	Basidiomycota: Cantharellus lobatus Peck	Wellington Channel, Nunavut, Canada	Polunin (1940)
Sir Francis Leopold M’Clintock’s expedition in the yacht Fox (1857–1859)	Basidiomycota: Agaricus allosporus Berk., Agaricus cyathiformis Bull., Agaricus furfuraceus P., Agaricus vaginatus Bull., and Agaricus umbelliferus L.	Boothia Peninsula, Nunavut, Canada	Hooker (1860)
Arctic expedition from 1875 to 1876	Ascomycota: Peziza stercorea P. and Urnula hartii B. Basidiomycota: Agaricus (Clitopilus) undatus Fr., Agaricus (Naucoria) bellotianus B., Agaricus (Omphalia) umbelliferus L., Agaricus (Omphalia) umbilicatus Schaeff., Agaricus (Stropharia) feildeni B., Agaricus (Tubaria) furfuraceus P., Agaricus (Tubaria) pellucidus Bull., Cantharellus muscigenus Fr., Hygrophorus miniatus Fr., Hygrophorus virgineus Fr., Lycoperdon atropurpureum Vitt., Lycoperdon cretaceum B., Merulius aurantiacus Fr., and Russula integra Fr.	Northern Nunavut, Canada	Berkeley (1878)
Second voyage of the Fram Expedition (1898–1902)	Ascomycota: Mollisia graminis (Desm.), Niptera melatephra (Lasch.), and Sclerotinia vahliana Rostr. Basidiomycota: Cantharellus lobatus (Pers.), Collybia dryophila (Bull), Galera hypnorum (Batsch), Hebeloma fastibile (Fr.), Lycoperdon gemmatum (Batsch), Mycena pumila (Bull), Naucoria festiva (Fr.), Naucoria melinoides (Fr.), Naucoria nimbosa (Fr.), Omphalia umbellifera (L.), Psalliota campestris (L.), Psalliota rodmani (Peck), Psathyrella polaris n. sp., Russulina lutea (Huds.), and Tricholoma caelatum (Fr.)	Ellesmere Island, Nunavut, Canada	Rostrup and Simmons (1906)
Canadian Arctic Expedition (1913–1918)	Ascomycota: Peziza micropus Pers. var. flavida Phil., Scleroderris fuliginosa (Fr.) Karst., and Scutellina scutellata (L.) Basidiomycota: Boletus scaber Fr., Cantharellus muscigenus Fr., Clavatia creatacea (Berk.) Lloyd, Galera hypnorum Batsch., Hebeloma fastibile Fr., Hygrophorus cantharellus Fr., Hygrophorus sp., Inocybe flocculosa Berk., Lycoperdon umbrinum Pers., Naucoria sp., Omphalla umbellifera Fr., and Russula sp.	Northern regions of Nunavut, Northwest Territories, and the Yukon, Canada	Dearness (1923)
Oxford Exploration Club’s 1931 expedition to Akpatok Island	Ascomycota: Helvella lacunosa Afz., Peziza acetabulum L., and Sphaerospora asperior (Nyl.) Sacc. Basidiomycota: Boletus reticulatus (Schaeff.) Boud., Boletus sp., Cantharellus cibarius Fr., Cortinarius spp., Dictyolus muscigena (Bull.) Quél., Hygrophorus sp., Hypholoma sp., Inocybe violacea Pat., Inocybe spp., Laccaria laccata (Scop.) B. & Br., Lactarius vellereus Fr., Lycoperdon perlatum Pers., Lycoperdon sp., Mycena polygramma Quél., Mycena spp., Omphalia sp., Psilocybe semilanceata Fr., Russula emetica Fr., Russula ochroleuca Fr., and Russula sp.	Akpatok Island, Nunavut, Canada	Polunin (1934)

Table 2. Research in the proceedings of the international symposia on arctic and alpine mycology focused on North American fungi

Symposium	Research on arctic and alpine mycology in North America	Acting president	Publication Editor(s)
1st ISAM, August 1980. Barrow, Alaska	Basidiomycetes in subarctic Alaska (Miller 1982a; 1982b), Cortinarius in Alaska (Ammirati and Laursen 1982), Lactarius in Alaska (Laursen and Ammirati 1982b), lower fungi in the Arctic (Kobayasi 1982), soil fungi in arctic tundra (Laursen and Chimelewski 1982), and mycorrhizal associations with Salix rotundifolia (Linkins and Antibus 1982)	Gary Laursen	Laursen and Ammirati (1982a)
2nd ISAM, August 1984. Swiss National Park, Switzerland	Basidiomycetes in the Rocky Mountain alpine (Moser and McKnight 1987), Cortinariaceae in the Yukon Territory of Alaska (Miller 1987), and Hygrophoraceae in the arctic–alpine tundra of Alaska (Laursen, Ammirati, and Farr 1987a)	Egon Horak	Laursen, Ammirati, and Redhead (1987b)
3rd ISAM, August 1988. Svalbard	The 3rd and 4th ISAM were combined into one proceeding and the following research related to arctic and alpine mycology in North America was included: Cystoderma in Alaska (Miller 1993), Lactarius in arctic Canada (Ohenoja and Ohenoja 1993), and Myxomycetes in the Alaskan tundra (Stephenson and Laursen 1993)	Sigmund Sivertsen	Petrini and Laursen (1993)
4th ISAM, August 1992. Lanslebourg, France		Denise Lamoure	Petrini and Laursen (1993)
5th ISAM, August 1996. Polar Urals, Russia	Hebeloma from Alaska (Miller 1998), Inocybe species from arctic Canada (Ohenoja, Vauras, and Ohenoja 1998)	Viktor Mukhin	Mukhin and Knudsen (1998)
6th ISAM, August 2000. Greenland	Arrhenia auriscalpium in Colorado (Cripps and Horak 2006) and Agrocybe praemagna in the Rocky Mountain alpine region (Moser and Horak 2006)	Torbjorn Borgen	Boertmann and Knudsen (2006)
7th ISAM, August 2005. Finse, Norway	Cortinarius in the Rocky Mountain alpine (Peintner 2008) and a checklist of agarics from the Rocky Mountain alpine zone (Cripps and Horak 2008)	Gro Gulden	Gulden and Høiland (2008)
8th ISAM, August 2008. Beartooth Plateau, Montana/Wyoming	Amanita found on the Beartooth Plateau (Cripps and Horak 2010), Marasmius epidryas in the Rocky Mountain alpine (Ronikier and Ronikier 2010), Hebeloma hiemale in the Rocky Mountain alpine (Becker, Eberhardt, and Vesterholt 2010), larger fungi in arctic and subarctic Canada (Ohenoja and Ohenoja 2010), Inocybe (Mallocybe) in the Rocky Mountain alpine (Cripps, Larsson, and Horak 2010), Lycoperdaceae on the Beartooth Plateau (Kasuya 2010; Jalink 2010), and Mollisioid Ascomycetes on the Beartooth Plateau (Nauta 2010)	Cathy Cripps	Cripps and Ammirati (2010)
9th ISAM, August 2012. Utsjoki, Finland	Inocybe leiocephala, a species with an intercontinental distribution (Larsson, Vauras, and Cripps 2014), and Lactarius from the Rocky Mountain alpine zone (Cripps and Barge 2014)	Esteri Ohenoja	Ohenoja, Ruotsalainen, and Vauras (2018)
10th ISAM, August 2016. Kanazawa, Japan	Inocybe arctica and other species later reported in the North American alpine (Larsson, Vauras, and Cripps 2018)	Tamotsu Hoshino	Hoshino et al. (2018)

Table 3. Current list of species of alpine Agaricales, Russulales, Boletales, and Ascomycota from the Rocky Mountains of Colorado, Montana, and Wyoming collected at elevations of ca. 3,000 to 4,200 m

BASIDIOMYCOTA

AGARICALES

AGARICACEAE

Agaricus cf. aristocratus Gulden^1

Agaricus campestris (L.) Fr.^a,1

Agaricus spp.

Bovista limosa Rostr.^a,2

Bovista nigrescens Pers.^a,2

Bovista pila Berk. & M.A. Curt.^2

Bovista plumbea Pers.^2

Bovistella utriformis (Bull.) Demoulin & Rebriev^2

Crucibulum laeve (Huds.) Kambly^3

Cystoderma sp.^1

Lepiota alba (Bres.) Sacc.^1

Lepiota magnispora Murrill^1

Calvatia booniana A.H. Smith^2

Calvatia cretacea (Berk.) Lloyd^a,2

Calvatia sculpta (Hark.) Lloyd^2

Calvatia turneri (Ellis & Everh.) Demoulin & Lange^a,2

Lycoperdon frigidum Demoulin^2

Lycoperdon norvegicum Demoulin^2

AMANITACEAE

Amanita groenlandica f. montana Cripps & Horak^a,1

Amanita nivalis Grev.^a,1,4

Amanita vaginata (Bull.) Lam.^1,4

BOLBITIACEAE

Agrocybe praecox (Pers.) Fayod^1

Agrocybe cf. praemagna M.M. Moser & E. Horak^1

Conocybe sp.

CORTINARIACEAE

Cortinarius absarokensis M.M. Moser & McKnight^1,5

Cortinarius adelbertii J. Favre^1,6

Cortinarius albonigrellus J. Favre^1,5

Cortinarius alpinus Boud. = Cortinarius favrei D.M. Henderson^a,b,c,1,5

Cortinarius anomalus (Fr.) Fr.^a,b,c,1

Cortinarius atroalbus M.M. Moser^1,6

Cortinarius caninus Fr.^b,c,1

Cortinarius chrysomallus Lamoure^1,5

Cortinarius cinnamomeoluteus P.D. Orton^a,1

Cortinarius expallens M.M. Moser^1,6

Cortinarius ferrugineifolius M.M. Moser^1,6

Cortinarius fuscoflexipes M.M. Moser & McKnight^1,5

Cortinarius galerinoides Lamoure^1,6

Cortinarius hemitrichus (Pers.) Fr.^3

Cortinarius hinnuleus Fr. s.l.^1,6

Cortinarius hinnuleus var. favreanus Bon^c,3

Cortinarius hybospermus M.M. Moser^1,6

Cortinarius inops J. Favre^1,6

Cortinarius laetus M.M. Moser^1,6

Cortinarius minutalis Lamoure^a,^1,6

Cortinarius mucronatus M.M. Moser & McKnight^1,5

Cortinarius paraphaeochrous M.M. Moser^1,6

Cortinarius parvannulatus Kühner^a,^3

Cortinarius pauperculus J. Favre^1,5

Cortinarius phaeochrous J. Favre^1,6

Cortinarius phaeopygmaeus J. Favre^1,6

Cortinarius polaris Høiland^a,b,c,1

Cortinarius pusillus M.M. Moser, McKnight & Sigl^1

Cortinarius rufoanuliferus M.M. Moser & McKnight^1,5,6

Cortinarius stenospermus Lamoure^1,5,6

Cortinarius subrigidipes M.M. Moser^1,6

Cortinarius subtorvus Lamoure^a,b,c,1,6

Cortinarius tenebricus J. Favre^1,5

Cortinarius cf. umbilicatus P. Karsten^3

Cortinarius vibratilis (Fr.) Fr.^1

Cortinarius vulpicolor M.M. Moser & McKnight^1,5,6

ENTOLOMATACEAE

Entoloma alpicola (Favre) Bon and Jamoni^a,1

Entoloma cf. atrosericeum (Kühner) Noordel.^3

Entoloma cetratum (Fr.) M.M. Moser^1

Entoloma juncinum (Kühner & Romagn.) Noordel.^1

Entoloma sericeum Quél.^a,1

Rhodocybe cf. popinalis (Fr.) Singer^a,^1

HYMENOGASTRACEAE

Alnicola sp.^1

Galerina annulata (J. Favre) Singer^1

Galerina arctica (Singer) Nezdojm.^1

Galerina atkinsoniana A.H. Smith^1

Galerina clavata (Velen.) Kühner^1

Galerina subclavata Kühner^3

Galerina cf. pseudomycenopsis Pilát^a,^1

Galerina unicolor (Vahl.) Singer = Galerina marginata (Batsch) Kühner^1

Galerina spp.

Hebeloma alpinicola A.H. Sm., Evenson & Mitchel^b,7

Hebeloma alpinum (J. Favre) Bruchet^a,b,c,1,7

Hebeloma aurantioumbrinum Beker, Vesterh. & U. Eberh.^b,c,7

Hebeloma avellaneum Kauffman^b,7

Hebeloma excedens (Peck) Sacc.^b,7

Hebeloma dunense L. Corb. & R. Heim^b,c,7

Hebeloma hiemale Bres.^b,c,7

Hebeloma hygrophilum Poumarat & Corriol^b,c,7

Hebeloma marginatulum (J. Favre) Bruchet^b,c,7

Hebeloma mesophaeum (Pers.) Quél.^a,b,c,1,7

Hebeloma nigellum Bruchet^a,c,7

Hebeloma oreophilum Beker & U. Eberh.^b,c,7

Hebeloma spetsbergense Beker & U. Eberh.^b,c,7

Hebeloma subconcolor Bruchet^b,c,7

Hebeloma vaccinum Romagn.^b,c,7

Hebeloma velutipes Bruchet^b,c,7

Phaeogalera stagnina (Fr.) Kühner^3

INOCYBACEAE

Inocybe alpinomarginata C.L. Cripps, E. Larss. & Vauras^b,c,8

Inocybe arctica E. Larss., Vauras & C.L. Cripps^b,c,8

Inocybe cf. bulbosissima (Kuehner) Bon^1

Inocybe canescens J. Favre^1

Inocybe dulcamara (Pers.) P. Kumm.^a,b,c,1,9

Inocybe fraudans (Britzelm.) Sacc.^1

Inocybe giacomi J. Favre^b,c,1,7

Inocybe lacera (Fr.) P. Kumm.^a,^1

Inocybe leiocephala D.E. Stuntz^a,b,c,1,10

Inocybe leonina Esteve-Rav. & A. Caball.^b,c,7

Inocybe leucoblema Kühner^b,c,9

Inocybe cf. maculipes J. Favre^1

Inocybe murina E. Larss., Cripps, & Vauras^b,c,8

Inocybe occulta Esteve-Rav., Bandini, B. Oertel & G. Moreno^a,b,c,8

Inocybe paragiacomi E. Larss., C.L. Cripps & Vauras^b,c,8

Inocybe phaeocystidiosa Esteve-Rav., G. Moreno & Bon^b,c,8

Inocybe purpurpeobadia Esteve-Rav. & A. Caball.^b,c,8

Inocybe cf. rhacodes J. Favre^1

Inocybe subgiacomi C.L. Cripps, Vauras & E. Larss.^b,c,8

Inosperma calamistratum (Fr.) Matheny & Esteve-Rav.^a,^1

Inosperma cf. subhirsutum (Kühner) Matheny & Esteve-Rav.^1

Mallocybe arthrocystis (Kühner) Matheny & Esteve-Rav.^b,c,9

Mallocybe fulvipes (Kühner) Matheny & Esteve-Rav.^b,c,9

Mallocybe fuscomarginata (Kühner) Matheny & Esteve-Rav.^b,c,1,9

Mallocybe leucoloma (Kühner) Matheny & Esteve-Rav.^b,c,9

Mallocybe terrigena (Fr.) Matheny, Vizzini & Esteve-Rav.^a,b,c,9

Mallocybe substraminipes (Kühner) Matheny & Esteve-Rav.^b,c,9

Pseudosperma cf. rimosum (Bull.) Matheny & Esteve-Rav.^1

HYDNANGIACEAE

Laccaria nobilis A.H. Sm. In G.M. Mueller^b,1,11

Laccaria laccata var. pallidifolia (Peck) Peck = L. tetraspora Singer = L. laccata (Scop.) Cooke^a,b,1,11

Laccaria montana Singer^a,b,1,11

Laccaria pumila Fayod^a,b,1,11

Laccaria pseudomontana Osmundson, C.L. Cripps & G.M. Mueller^b,1,11

HYGROPHORACEAE

Arrhenia acerosa (Fr.) Kühner^1

Arrhenia auriscalpium (Fr.) Fr.^a,1,12

Arrhenia lobata (Pers.) Kühner & Lamoure ex Redhead^a,1

Arrhenia velutipes (P.D. Orton) Redhead, Lutzoni, Moncalvo & Vilgalys^a,1

Gliophorus psittacinus (Schaeff.) Herink = Hygrocybe psittacina (Schaeff.) P. Kumm.^1

Hygrocybe conica (Schaeff.) P. Kumm.^a,1

Hygrocybe marchii (Bres.) F.H. Møller^1

Lichenomphalia sp.^a,1

LYOPHYLLACEAE

Calocybe chrysenteron (Bull.) Singer^1

Calocybe onychina (Fr.) Donk^1

MARASMIACEAE

Hydropus sp.^1

MYCENACEAE

Hemimycena sp.^1

Mycena pura (Pers.) P. Kumm. s.l.^1

NIACEAE

Flagelloscypha sp.^1

PHYSALACRIACEAE

Flammulina sp.^1

Rhizomarasmius epidryas (Kühner ex A. Ronikier) A. Ronikier & Ronikier^a,1

STROPHARIACEAE

Deconica chionophila (Lamoure) Noordel.^1

Hypholoma elongatum (Pers.) Ricken^3

Stropharia alpina M. Lange^1

TRICHOLOMATACEAE

Clitocybe dryadicola (J. Favre) Harmaja^1

Clitocybe festivoides Lamoure^3

Collybia cookei (Bres.) J.D. Arnold^1

Dermoloma sp.^1

Gamundia leucophylla (Gillet) H.E. Bigelow = Fayodia leucophylla (Gillet) M. Lange & Sivertsen^1

Gymnopus alpicola (Bon & Ballarà) Esteve-Rav., V. González, Arenal & E. Horak^3

Loreleia marchantiae (Singer & Clémençon) Redhead, Moncalvo, Vilgalys & Lutzoni^1

Lepista cf. ricekii Bon^3

Melanoleuca cognata (Fr.) Konrad & Moutbl. s.l.^a,^b,c,1

Melanoleuca substrictipes Kühner^3

Mycenella sp.^1

Omphalina rivulicola (J. Favre) Lamoure^1

RUSSULALES

RUSSULACEAE

Lactarius glyciosmus (Fr.) Fr.^a,c,1,13,14

Lactarius lanceolatus O.K. Miller & Laursen^a,b,c,13,14

Lactarius nanus J. Favre^a,b,c,1,5,13,14

Lactarius pallidomarginatus Barge & C.L. Cripps^b,13,14

Lactarius repraesentaneus Britzelm.^b,c,1,13,14

Lactarius salicis-reticulatae Kühner^b,c,1,13,14

Russula altaica (Singer) Singer^a,b,c,15 similar to Russula gracillima Jul. Schäff.

Russula amoenipes var. kuehneriana^15

Russula cf. heterochroa Kühner^c,15

Russula intermedia P. Karst.^b,c,15

Russula laevis Kälviäinen, Ruotsalainen & Taipale^a,b,c,15

Russula laccata Huijsman^a,b,c,1,15

Russula nana Killerm.^a,b,c,1,5,15

Russula purpureofusca Kühner^b,c,15

Russula saliceticola (Singer) Kühner ex Knudsen & T. Borgen^b,c,15

Russula subrubens (J.E. Lange) Bon^b,c,1,15

BOLETALES

BOLETACEAE

Leccinum cf. rotundifoliae (Singer) A.H. Smith, Thiers & Watling^b,c,1

CANTHARELALES

CANTHARELLACEAE

Rickenella fibula (Bull.) Raithelh.^a,^1

Rickenella swartzii (Fr.) Kuyper^1

ASCOMYCOTA

Bryoglossum gracile (P. Karst.) Redhead^a,3

Ciborinia ciborium (Vahl) T. Schumach. & L.M. Kohn^3

Morchella sp.^b,3

Notes. ^aReported from Canadian Arctic, not molecularly confirmed (Ohenoja and Ohenoja 2010).

^bGenbank sequence available.

^cIntercontinental distribution molecularly confirmed.

¹Cripps and Horak (2008).

²Kasuya (2010).

³Cripps, Unpublished. Representative specimens deposited in the fungal section of the MONT Herbarium, Bozeman, Montana.

⁴Cripps and Horak (2010).

⁵Moser and McKnight (1987).

⁶Moser, McKnight, and Ammirati (1995).

⁷Cripps et al. (2019).

⁸Cripps, Larsson, and Vauras (2020).

⁹Cripps, Larsson, and Horak (2010).

¹⁰Larsson, Vauras, and Cripps (2014).

¹¹Osmundson, Cripps, and Mueller (2005).

¹²Cripps and Horak (2006).

¹³Barge, Cripps, and Osmundson (2016).

¹⁴Barge and Cripps (2016).

¹⁵Noffsinger (2020).

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