Fungi: Encyclopedia Arctica 5: Plant Sciences (General)

Author Stefansson, Vilhjalmur, 1879-1962



(EA-PS. Rolf Singer)


Scroll Table to show more columns

Representation and Importance 2
Geographical Distribution 9
Economic Importance 13
Forestation 13
Horticulture 13
Mushrooms for Food 14
Mushrooms for Reindeer Fodder 14
Fungi Causing Deterioration 15
Bibliography 16

EA-Plant Sciences (Rolf Singer)

Fungi in the widest sense are nonflowering which, due to their dependency upon organic matter for their nutrition, do not need and are con– sequently devoid of chlorophyll; instead they live saprophytically or para– sitically, or symbiotically on or with other organisms, or on the products of decomposition or excrements of these organisms. Taxonomists now agree that the term Fungi as a systematic unit should be used only for those spore– bearing organisms which: ( 1 ) do not reproduce by fission (this eliminates the Bacteria and related groups of microorganisms); ( 2 ) have no plasmodial generation consisting of motile, multinucleate protoplasm (this removes the Myxothallophyta or slime molds); ( 3 ) are not lichenized, i.e., do not live in close association with Algae (this excludes the lichens); ( 4 ) are not so closely related to any of the autotrophic Algae that the separation from the latter would appear to be artificial from a phylogenetic point of view (this keeps the heterotrophic Algae in the appropriate group of Thallo– phytra, namely, the Algae. The Fungi proper are divided into four main groups” Phycomycetes (myceli x ^ u ^ m, where present, continuous); Ascomycetes (perfect stage an ascus, i.e., with reduction division taking place in a mother cell inside of which the spores are born); Basidiomycetes (perfect stage with spore formation on the outside of a mother cell, called a basi– dium); and the Fungi Imperfecti (perfect stage unknown — a temporary sub [: ] division).

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Representation and importance
All four classes of Fungi occur in the Arctic and Subarctic, probably in approximately the same proportion as they occur in warmer climates, but the number of species tends to be smaller than in more temperate regions. Some of the species are specifically boreal or arctic in their distribution and adaptation. Nevertheless and fungus flora of the Arctic is comparatively more important than in the warmer zones; indeed in number and diversity, Fungi in the North may even outweigh the flowering plants, though as com– ponents of vegetation they are relatively minor. In order to illustrate this numerical preponderance we may consider the flora of Greenland where E. V. Wulff indicates 416 species of flowering plants (phanerogams), whereas the known number of Fungi is about 850, i.e., there are more than twice as many Fungi — a figure that will tend to increase as mycological investiga– tions proceed. To date, most of the material studied has consisted of dead and diseased portions of flowering plants collected for general herbaria by phanerogamists, and more specific collecting of Fungi on other substrata has been neglected. Aside from that, we must keep in mind that lichens are merely a symbiotic form of life realized by close association of a fungus with an alga, and that the number of lichen species in Greenland is several hundred — a fact that should not be surprising, as many earth and rock– inhabiting lichens are particularly well adapted to the severe climatic con– ditions of the Arctic. By way of contrast, we know that the number of Fungi observed in Morocco, recently enumerated by R. Maire and R. G. Werner, is 1,200, whereas the phanerogams of the same region are estimat ^ e ^ d to number as high as 2,400.

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It is impossible to describe in detail the arctic fungus flora inde– pendently from the fungi of other regions, as it is a remarkable fact that not a single fungus genus has been discovered which is exclusively arctic in distribution or character. However, certain groups (orders, families, and genera) within the two largest groupings of Ascomycetes and Basidio– mycetes are distinctly more prominent and numerically more strongly repre– sented than others. For example, in the Ascomycetes, we find a remarkably strong representation among the Myrangiales (asci born internally, singly, in loculi), the Sphaeriales (asci born in cavities with osticles: perithecia), and the Helotiales and Pezizales (cup fungi, [: ] ^ w ^ ith asci born externally on palisades). Among the Basidiomycetes, we find the Ustilaginales (smuts), the Uredinales (rusts), and the Agaricales (mus [: ] hrooms) to be dominating. The majority of the Myrangiales and Sphaeriales grow on weakened or dead parts of flowering plants (stems and leaves)’ Ustilaginales and Uredinales are true para ti ^ sit ^ es; Agaricales are, mostly at least, seemingly soil-inhabiting, but some live in symbiosis and others on decaying cellulose or on dung, etc.
As for the Basidiomycetes (viz., Agaricales and Gasteromycetes), some unpublished work by the present author as well as some papers published recently by M. Lange (4) and some which are still unpublished in 1950 (with a summary of the results made available to the author by the cou [: ] ^ r ^ tesy of Mr. Lange) enable us now to recognize the characteristic features of the “agaric,” “bolete,” and “puffball” flora of Greenland and the alpine-arctic zones of Lapland. It appears that the most important agaricaceous genera (they are there commonly linked by symbiosis with low birches and willows) are Russula (especially Russula emetica subsp. alpestris ), Lactarius ,

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Cortinarius , Hebeloma and Inocybe . The important nonmycorrhizal genera [: ] are Laccaria (especially L. laccata ), Omphalina , Melanoleuca , Marasmiellus (especially M. fibula ), Collybia , Amanita (significantly — considering the evidently faulty hypothesis of a protective value of the veils — only species with rudimentary inner or outer veil), Agaricus , Galerina , and Cystoderma . Among the Boletaceae, we encounter only Leccinum. Laccaria and Habeloma are the genera most prevalent in the extreme northern stations (around lat. 82° N. in Greenland). Occasionally, in regions with a more “Atlantic” climate, in Greenland as well as in Lapland, species of the genera Camarophyllus , Hygrocybe , and Mycena are met with, and exceptionally even Rhodophyllus (an undetermined species), Rozites caperata , and Clitocybe sp. Among the Aphyllo phorales , Leptotus lobatus seems to be widely distributed, and among the Gasteromycetes (puffballs) Lange indicates Geaster minimus , Calvatia cretacea , C. tatrensis , C. arctica , Lycoperdon umbrinum , L. nigrescens , L. spadiceum , L. pusillum , Bovista echinella , B. tomentosa , B. nigrescena , Crucibulum vulgare , and Sphaerobolus stellatus for Greenland, and Bovistella paludosa for Lapland.
Owing presumably to the absence of a large reservoir of decaying woody mat– erial, such genera as Cyttaria , Xylaria , Hypoxylon , Nectria , and allie d ^ s ^ among the Ascomycetes, and Polyporus , Poria , Fomes , Coriolus and [: ] ^ all ^ ies, as well as Stereum , Corticium , Peniophora , and Auricularia among the Basidiomycetes, and also all the imperfect forms related to these groups, are either not represented at all or are very rare in the Arctic. The same is true for Fungi normally growing in close association with the roots of living forest trees (conifers, Salicales, Fagales, Tilia , Fraxinus , etc.), such as Elaphomyces and the true truffles (Tuberales) among the Ascomycetes, and most hypogeous Gasteromycetes, as well as many boletes and agarics. With them, naturally, the fungus

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parasites of those fungi are likewise absent from the arctic regions (for example, Hypomyces lactifluorum ).
It appears thus far that the Fungi inhabiting dead stems and leaves of flowering plants are most favored under arctic conditions. This may be because of the reduced competition on the part of soil-inhabiting Fungi Imperfecti and Bacteria, both of which groups are considerably smaller in number of representa– tives, and less active than in temperate and tropical regions, owing presumably to the short northern summer. For this reason vegetative remains fail to decay rapidly but become, after overwintering, a suitable substratum for certain Sphaeriales and other Ascomycetes. This explains not only the preponderance of some of the groups enumerated above, but also the reduced geographic areas of certain fungi as compared with those of their host plants. Lind (6) in– dicates several interesting instances of definitely arctic, arctic-subarctic, or else arctic-alpine areas of parasitic or saprophytic fungi on hosts whose area extends considerably into the Temperate Zone. For instance, Rhytisma empetri and Sphaeropezia empetri were observed only in the arctic and alpine regions of the area of Empetrum nigrum , whereas other fungi ( Physalospora empetri and Chrysomyxa empetri ) follow Empetrum through its entire area. Similar in– stances have been further discussed by Lind, Komarov, Murashkinski and Ziling (9), Tranzschel (13, and Yachevski (14). Naturally, the opposite situation, where the area of the fungus is larger than that of its arctic host, also occurs. All this proves that the areas of host and parasite do not necessarily coincide — as has also been substantiated in extra-arctic regions.
The large number of fungus species as compared with the phanerogams may appear surprising at first glance even though the special condition of the over– wintering dead leaves and stems is taken into consideration; for this substratum

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does not explain the large number of Pezizales and Agaricales, and those microscopic fungi that occur on substrata other than dead stems and leaves. However, the Arctic actually abounds in an inexhaustible reservoir of suit– able substrata. The rich lichen flora provides a great number and diversity of host plants, as is shown by the fact that a lichen collection, taken almost at random in Spitsbergen in 1921 and sent to a specialist (K. Keissler), yielded 12 species of licen parasites. Mosses, especially the Polytrichaceae, and hepatics, which are also rather numerous in arctic regions, provide the habitat for many Far Northern agarics ( Omphalina , Cortinarius , Galerina , Hy grocybe , Marasmiellus , Inocybe , etc.), and such genera as Cyphella and Lep totus . Agarics (especially the dark-spored groups) as well as some Pezizales and Helotiales, other Ascomycetes, and Phycomycetes, find a good substratum in bird droppings and other kinds of dung.
The families Gentianaceae, Ericaceae, and Primulaceae among other are known to depend largely for nomal development on endotrophic mycorrhiza. These fungi live in the roots of the plants and enter the cells, deriving for themselves as well as providing for the host some nutritive advantages by alternately absorbing cell sap by haustoria, and being digested by the host. Furthermore, even though forest trees, which form ectotrophic or peritrophic mycorrhiza (fungus hyphae surrounding or enveloping the rootlets but not entering the cells), are absent in truly arctic territories, some species belonging to those genera form extensive stands over wide regions of the Arctic and Subarctic, namely the shrubby or almost herbaceous birches and willows. There are many species of boletes and agarics connected with one species of northern birch alone, and this explains the presence, in the arctic tundra, of such large mushroom carpophores as those of Leccinum scabrum subsp.

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rotundifoliae and L. testaceoscabrum var. arcticum . Other habitats of fungi in the Arctic are diseased and dead insects, driftwood on the beaches, dead and living fishes, and dead and living algae. None of these habitats has as yet been studied extensively.
The true parasites seem to be somewhat rarer in the Arctic Zone than farther south, which may be explained by the fact that the sparse, scat– tered growth of many host plants tends to decrease direct competition between the host plants in dense populations as a preparatory factor for fungus in– fection, and also minimizes the cha ^ n ^ ce of infection of the right host by wind-blown spores. (Most true parasites such as rusts are hi hg ^ gh ^ ly specialized as to the taxonomic position of the host). This supposition seems to be sup– ported by the fact that in the Arctic the percentage of fungi with a wide host range is much larger than in the temperate zones, the number of strictly specialized forms growing exclusively on one host species being far smaller than in warmer regions. Among the rust fungi, where most species in the Tem– perate Zone ordinarily pass through several generations of spore formation, there is a tendency to abbreviate this complicated life cycle by omitting some of the spore forms. Such rusts with drastically abbreviated life cycle are called microcyclic Uredinales (1). Roughly one-third of the arctic rusts are genuine microforms, which is a much higher percentage than in the tem– perate zones, and the vast majority of the remaining species are adapted to arctic conditions either by omission of one spore form or the other, or else by the formation of a perennial mycelium. However, many of the “arctic” forms occur in the temperate zones also, and consequently cannot be inter– preted as having a secondary adaptation to arctic conditions, but rather an initial advantage enabling these forms rather than others to penetrate into arctic habitats.

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Yachevski (14) thinks that in the alpine Ascomycetes, the perithecia are thick-walled and of scelrotial type, and harder than those developed in the lowlands; moreover, the development of the asci and spores is allegedly slowed, the asci can be found in the most different stages of development in a single fruiting body, and the spores tend to be large and bright golden– yellow. The present author has noted that in alpine stations the spores are more often ornamented than smooth, and Heim and Singer have found that the spores tend to be larger in mountain forms and races. No corresponding facts have been established statistically for the Arctic, but one may be inclined to think that the similarity of conditions in the high mountains and the Arctic, the identity of many species in both floras, and the analo– gous case of the microcyclic rusts which are more numerous both in the Arctic and in mountains than elsewhere, would tend to make it probable that most of the peculiarities of the alpine mycoflora will also be proved to exist in the Arctic.
Geographical Distribution
There must have been an exchange of species between the large mountain massifs of North America, Europe, and Asia, and the arctic regions of those continents during the various glaciations and particularly in the interglacial periods. This does not mean that any one of these floras has necessarily been derived from any other, but merely that there must have been contact or migra– tion of floral elements, at least as far as the Fungi are concerned. Some of the species occurring both in the arctic regions and in the mountain ranges farther south are: Massaria eucarpa (Spitsbergen and Caucasus); Phyllosticta calthae (Kamchatka and Caucasus); Botrychonema juncisedum (Greenland, Iceland, Adirondack Mountains, and Alps); Phoma alpina (Ellesmere Island, Umanak, and

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Alps); Clathrospora pentamera (circumpolar: arctic and subarctic; also in the Rocky Mountains, the Pamir, and the Italian mountains); Russula emetica subsp. alpestris (Greenland eastward to Novaya Zemlya; also in the Alps, Carpathian Mountains, and Caucasus); Melanoleuca evenosa (Iceland, Alps, and Caucasus); Amanita vaginata (Greenland, Lapland, Alps); Omphalina abiegna (Lapland, Alps); certain forms of Galerina hypnorum (Lapland, Greenland, White Mountains in New Hampshire, Altai Mountains, Alps); Lactarius repraesentaneus and L. torminosus (Lapland, ?Gr f ^ e ^ enland, Altai Mountains); Leccinum scabrum subsp. rotundifoliae (Greenland westward to Novaya Zemlya; Altai Mountains); Calvatia tatrensis (Greenland, Tatra).
The majority of the species that make up the flora of the Arctic have a wide distribution; some have been demonstrated to be continuously distributed around the North Pole (for example, Mycosphaerella tassiana ^ Mycosphaerella tassiana ^ on no less than 110 different host plants ranging from Lycopodium to Hieracium ). Only a few species are so adaptable as to be almost cosmopolitan (for example, the smut Cinctractia caricis in the widest sense, and the puffball Crucibulum vulgare ), and although several species have so far been reported only from one single region (e.g., Micromyces wheldenii from the Canadian Eastern Arctic and Puc cinia rhytisomoides from the northernmost parts of the European Continent), it may be expected that most of them will be found to occur also elsewhere and thus cease to be “endemics.” The only region which may be expected to have a comparatively large degree of endemism is the northeastern Siberian Arctic.
The generally wide distribution of the arctic fungi may [: ] at first glance appear to be a puzzling fact, as land tabai table for plant life is dispersed and disrupted by wide stretches of ocean. The theory of spore-and seedcarry– ing birds as a means of plant migration over large distances has been gradually

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abandoned by most plant geographers. The hypothetical prehistoric land bridges are a matter of controversy. According to Lind (7), none of these explanations is needed as far as the fungi are concerned. During the arctic winter, fragments of plants with resistant spores or perennial mycelium (almost all arctic fungi have one or ther other, or both) intheir [: ] tissue or attached to them, may easily be carried across the ice by the wind, and will then be scattered to– gether with the snow upon the frozen ground as much as 400 miles away from their original habitat. After reaching some new arctic land, the fragments have a fair chance of falling upon or near a suitable substratum, or when the summer thaw comes of finding their way into some rivulet which will carry them to such a substratum.
However, this explanation does not take into account the fact that the almost unknown Antarctic fungus flora is so surprisingly similar to the arctic mycoflora — indeed to a degree that makes it impossible to interpret the analogy as coincidental. The same genera and sometimes the same species pre– vail in the mountain zone of Tierra del Fuego and in Greenland. For instance, the species Leptotus lobatus is equally distributed in both regions, and the Laccarias are as dominant in the Antarctic as in the Arctic. So as the Melano– leucas (having in some cases even identical or nearly identical species in both hemispheres), the Cystodermas and Omphalinas (in many cases identical species), Camarophyllus aff. lacmus , Marasmiellus fibula , Agaricus campestris , and certain species of Galerina and Naematoloma . While Russula is rare and Lactarius , Hebeloma , and the Boletaceae are unknown in the subantarctic and antarctic regions, it is remarkable that Cortinarius is very rich in species in both of these regions, and so is Inocybe — the last two general forming mycorrhize with shrubby and subherbaceous willows and birches in the North

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and with shrubby Nothofagus ^ Nothofagus ^ in the South. Consequently, Lind’s wind-over– ice theory, even though it correctly describes a specifically arctic way of fungus dissemination, cannot explain the wide distribution of many arctic genera and species.
It it were assumed — as indeed it is by numerous plant geographers — that migration has taken place long the mountain ranges of the Americas extending from north to south, there is no explanation for the specific identity of some floral elements in the Arctic and Antarctic, nor is there a way of overcoming the difficulty of exaplining the manner by which plants adapted to the frigid zones could have passed over the lowlands and low hills of southern Central America. It seems more logical to assume an originally homogeneous flora of which some components were able to adapt themselves to the extreme conditions of the frigid zones and to survive there, while grad– ually exposed to extinction in the moderate and warm climates because of com– petition with species which were themselves adapted more specifically to conditions prevalent in the extra-arctic regions.
This hypothesis does not by necessity deny the role of migration in specification, but it denies the specific migration from the Canadian Arctic along the Rockies and the Andes to the Antarctic in comparatively recent times, i.e., of the existing species common to both the arctic and the antarc– tic mycoflora. It implies that in the more numerous cases where only the genera l are common to both regions, the species have been born by mutations caused, for example, by climatic changes, and that specification has been analogous but slightly different in the Arctic and in the Antarctic. The fact that a large percentage of the arctic and antarctic fungi are not exclusively

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adapted to frigid conditions and extend their areas far into the warmer regions, and the fact that mycologists will readily agree that most species common to the arctic and antarctic floras may be characterized as compara– tively primitive, will tend to corroborate the author’s views.
Economic Importance
The role of fungi in nature consists of the decomposition of dead vege– table masses and their transformation into humus (saprophytes), and the transformation of nutritive elements into such a form as can be assimilated by higher plants (symbionts). Aside from these functions there are many practical uses for fungi under arctic and boreal conditions, and also much economic damage caused by them.
Forestation . Attempts at forestation in Iceland show that those trees trees which at last became established are accompanied by their mycorrhizal fungi, for example, Pinus mugho by Suillus luteus . All such experimental plantations should in the future be planned with a view to the role of the fungus partner of each tree species, its resistance to the climatic conditions being fully as important as the ecology of the tree itself. This is equally valid for the growing of berries of the [: ] Vaccinioideae group, such as cran– berries.
Horticulture . As many crops in the Arctic are greenhouse, plants, the control of their diseases should be similar to that indicated under the same conditions in other regions. One important outdoor crop of the Soviet tundra, red radishes, does not seem to be affected seriously by fungus disease. Hardy crops that might in the future be adapted to subarctic agriculture may profit by the existence of a certain margin between the [: ] ^ minimum ^ growth temperature

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of the host and that of the parasite, as has indeed been pointed out for wheat rust by Chester.
Mushrooms for Food . A successful method of growing commercial white mush– rooms in the Arctic has not been worked out because of the difficulty of ob– taining suitable manure locally, and the high cost of heating the mushroom houses to the optimum growth temperature of Agricus bisporus . Future experi– ments will undoubtedly endeavor to adapt one of the local wild species such as Melanoleuca evenosa , with an already low optimum growth temperature and more easily available substratum. If such a method could be worked out, these mushrooms would be an ideal year-round crop in the Arctic, as mushrooms do not require light for normal development.
Mushrooms collected in summer and eaten fresh, or pickled or dried for winter use, are one of the most valued vegetable crops of the arctic and especially the subarctic regions of Europe and Asia. Potentially, however, they would be equally valuable in Alaska and other parts of the American Arctic. The arctic fungus flora is particularly low in poisonous species. Aside from some very small agarics ( Clitocybe and Inocybe ), there is only the well-known, easily recognizable Amanita muscaria (fly mushroom), which is widely used by the natives of northern Siberia and Kamchatka as an intoxicating drug. All other species so far known to occur in the Arctic or Subarctic appear to be harmless.
Mushrooms as Reindeer Fodder . An important aspect of the fungi is their use as fodder either by direct pasturing of animals or by preparation of mushroom-containing fodder. The reindeer feeds on mushrooms as well as on lichens, and even in winter digs the frozen mushrooms out of the snow. The local and seasona [: ] l abundance of mushrooms is a factor in the choice of

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feeding grounds in Lapland and northern U.S.S.R. According to Rautavaara (12), the weight of the reindeer is on an average 20% higher than normal in a good mushroom year, and 12% lower than normal in a year when mushrooms are scarce. Fodder containing mushrooms accounts for about 20% of the meat production, valued at about 10 million Finnish marks per year, in Finnish Lapland alone.
Fungi Causing Deterioration . Some fungi, such as the domestic fungus Serpula lacrimans (also incorrectly called “ Merulius lacrymans ”) and Lentinus lepideus , cause the deterioration of the wooden frames, wharves, railway ties, and all kinds of wooden structures, even in mines. These fungi are not so danger– ous in the Arctic as they are in warmer regions, as they do not grow fast enough outdoors; yet indoors they may be very destructive. The deteriora– tion of fabrics in the tropics, which is mostly due to Fungi Imperfecti, has lately received much attention. Data on fungus destruction of fabrics in the Arctic are not available at present, but it is expected to be less severe than in the topics or even in the temperate zones.

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1. Arthur, J.C., “Notes on arctic Uredinales,” Mycologia , vol.20, no.1, pp.41-43, 1928.

2. Berkeley, M.J. “Enumeration of the fungi collected during the Arctic Expedition 1875-76,” Linncan Soc. J . vol.17, pp.13-17, 1878.

3. Dearness, J. “The fungi of the arctic coast of American west of the looth meridian,” Canadian Arctic Expedition, 1913-18. Report. Botany . Pt.C. Fungi , vol.4, 1923.

4. Lange, M. “Macromycetes. Part I: The Gasteromycetes of Greenland in The Botanical Exploration of West Greenland 1946,” Medd . Grønland , vol.147, no.4.

5. Larsen, Poul. “Fungi of Iceland,” Botany of Iceland vol.2, pp.451-607, 1931. (With literature.)

6. Lebedeva, L.A. Griby Arkticheskogo Poberezhia Sibiri . (Mushrooms of the Arctic Littoral.) Leningrad, 1928. Akad.Nauk Komissiia po Izucheniiu Yakutskoi Avtonomnoi Sovets. Sotsial.Resp. Trudy , vol.12.

7. Lind, J. “Studies on the geographical distribution of arctic circumpolar micromycetes,” Danske Vidensk. Selsk. Biologiske Medd . vol.11, no.2, pp.1-152, 1934. (Includes fairly complete literature on the subject.)

8. Linder, D.H. “Fungi,” Polunin, Nicholas. Botany of the Canadian Eastern Arctic . Ottawa, 1947, pt.2, pp.234-83, Nat.Mus.Can. Bull . no.97. Biological Ser . no.26.

9. Murashkinski, K.E., and Ziling, M.K. “Materialy po Mikoflore Altaya i Sayana.” (Materials on the mycroflora of the Altai and Saian.) Sibirskii Inst. Selskogo Khoz. i Lesovod., Omsk. Trudy , vol.10, no.4, pp.1-31, 1928.

10. Nannfeldt, J.A. “Contributions to the mycoflora of Sweden,” Svensk . Bot.Tidskr . vol.22, no.1-2, pp.115-39, 1928.

11. Norwegian Expedition to Novaya Zemlya 1921. Report of the Scientific Results . Ed. by Olaf Holtedahl. Oslo, Brøggers, 1924-30. 3 vol. (Particularly the papers on fungi.)

12. Rautavaara, Toivo. Suomen Sienisato, Porwoo . Helsinki, 1947.

13. Tranzschel, W.H. “Griby i Myxomycety Kamtchatki.” (Mushrooms and mycomycetes of Kamchatka.) Komarov, et al. Expedition a Kamtchatka , vol.2, pp.535-76, f.1, 1914 (German resume, p.595).

EA-PS. Singer: Fungi - Bibliography

14. Yachevski, A.A. Osnovy Mikologii . (Foundations of Mycology.) Moscow, Leningrad, 1933, (Especially Chap.XI, 5, 5A, pp.809-825. With list of literature.)

Rolf Singer