Fungi

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(EA-PS. Rolf Singer)

       

FUNGI

       

CONTENTS

	Page
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

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EA-Plant Sciences

(Rolf Singer)

       

Fungi

        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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---

BIBLIOGRAPHY

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3 vol. (Particularly the papers on fungi.)

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of Kamchatka.) Komarov, et al. Expedition a Kamtchatka ,

vol.2, pp.535-76, f.1, 1914 (German resume, p.595).

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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