BIND V

F ES 

NORDISK MYKOLOGISK TIDSSKRIFT 

BIND V HEFTE 3-5 

KØBENHAVN 1956 

FESTSKRIFT TIL PROFESSOR, DR. PH IL. 0. WINGE

INDHOLD 

Side 

Catherine Roberts: The inheritance of enzymatic characters in 

yeasts .................................................................. 161 

Olof Andersson: Three rare or little known bolets in Sweden. 

Boletus pulverulentus OPAT., B. radicans PERS. ex FR. 

and Phylloporus rhodoxanthus (SCHW.) BRES. ............ 180 

Erik Bille-Hansen: The nitrogen requirements of Coprinus heptemer-us-.M. 

LANGE & SMITH .......................................... 190 

N. Fabritius Buchwald: On the dimorphism of the ascospores and 

their arrangement in the ascus of Monilinia oxycocci 

(WOR.) HONEY (syn. Sclerotinia oxycocci WOR.) ......... 196 

Kjeld Biilow: Ornamentation of spores of Russula lauroceraci 

MELZER and Russula foetens (PERS.) FR...................... 204 

M. P. Christiansen: Two new species of Corticiaceae from Denmark. 

Peniophora danica sp. n. and Corticium salicicola 

sp. n. .................................................................. 207 

M. Skytte Christiansen: A new species of the form-genus Lichenoconium 

PETR. & SYD. (Fungi imperfecti) J L. xanthoriae 

sp. n. .................................................................. 212 

E. J. H. COl'ner: A new European Clavaria: Clavulinopsis septentrionalis 

sp. nov. ................................................ 218 

Anders Danielsen: Coprinus comatus (SeRUM.) FR. sprenger asfalt 

i Bergen, Norge. (Summary: Coprinus comatus 

breaks through asphalt in Bergen, Norway) .......... ..... 221 

Finn-Egil Eckblad: Some operculate Discomycetes new to Norway 

..................................................................... 223 

K. J. Frandsen: Variations in resistance of Trifolium pratense 

to attacks of Kabatiella caulivora (KrRCHN.) KARAK.... 231 

P. Sonne Frederiksen: A new Rhodotorula species, Rhodotorula 

macerans sp. n., isolated from field-retted flax straw ... 234 

Paul Gelting: Parmelia subaurifera NYL. and P. fraudans (NYL.) 

NYL. in Greenland . ............... .................. .............. 240 

Asbjørn Hagen: Rustsopper og rustverter nye for Færøerne. 

(Summary: Two rusts and several host plants of Puccinia 

hieracii new to the Færoes) ................................. 247 

Lise Hansen: Two polyporaceous fungi with merulioid hymenophore. 

Poria taxicola (PERS.) BRES. and Polyporus 

dichrous FR. ex FR. ................................................ 251 

Louis Hal'ffisen: On Merulius silvester FALCK and M. himantioides 

FR. Preliminary remarks .......................................... 257 

Ernst Hayren: Die in FinnIand bisher gefundenen Wasserpilze... 264 

K. Hauerslev: Om Fund af Judasøre (Hirneola auricula Judae 

(L.) BERK.) i Korsør og nærmeste Omegn. (Summary: 

A mass occurrence of Hirneola auricula Judae (L.) BERK. 

in Korsør, Sealand, in Denmark) ................. ............. 266 

J. P. Jensen: Recherches sur le cycle de Coprinus disseminatus 

(PERS. ex FR.) S. F. GRAY.. ..................................... 271 

Henrik Alb. JOl'gensen: M onascus ruber van TIEGH. demonstrated 

in Denmar k ...................................................... 274 

Ivar Jørstad: On the Sonchus rust Peristemma pseudosphaeria 

(MONT.) n. comb. (syn. Puccinia sonchi ROB.) .. .......... 278 

Axel B. Klinge: Beitrage zur Pilzflora Jtitlands .............. ....... 284 

Morten Lange: Pyrenomycetes parasitic on hypogeous fungi ... 289 

Fortsættes paa Omslagets Side 3 

PDF scanning and OCR by the Danish Mycological Society 2010 - www.svampe.com

Friesia . Bind V, Hefte 3- 5, Side 161- 432, 1956. 

TIL 

PROFESSOR, DR. PHIL. ØJVIND WINGE 

PAA 70 AARSDAGEN 

DEN 19. MAJ 1956

F'RIESIA V , 1956 PLATE I

Fotograferet 1950 paa Carlsberg Laboratorium

THE INHERITANCE OF 

ENZYMATIC CHARACTERS IN YEASTS 

By CATHERINE ROBERTS 

Leeture presented in CDpenhagen at the 1951 

Microbial Genetics Symposium 

Introduetion 

In the following I would like to discuss the problem of the inheritance 

of enzymatic characters in the yeast fungi and, in doing so, to 

attempt to review critically the available evidence in support of men delian 

and non-mendelian heredity in these organisms. Such an attempt 

will necessarily involve a discussion of enzymatic adaptation in its 

bearing upon yeast genetics, especially since current views offered in 

explanation of this phenomenon are as widely divergent as those 

which attempt to explain the mechanism of gene transmission. Since 

the contributions to yeast genebcs have been exceedingly numerous, 

I propose to tre at the subject chronologically in order to trace the 

developments within the field in as clear and unambiguous a manne r 

as possible. 

1935-1939 

The year 1935 ean be taken as a starting point, for it was in this 

year that a paper by Professor WINGE entitled "On Haplophase and 

Diplophase in some Saccharomycetes" appeared, in which the complete 

life cycle of Saccharomyces) including sexuality, sporulation, and 

alternation of generations, was incontestably established. As LINDE 

GREN has aptly remarked in his recent book, "The Yeast Cell", it was 

WINGE "who brought yeast genetics into being", for with thi s 1935 

paper as a basis, there followed in the years 1937-1939 a series of 

now classical papers by Professor WINGE and his collaborator LAUST- 

FRIESIA V -161 - 11

- 162- 

SEN, in which the yeast fungi were, for the first time, employed in thc 

field of genetics. With the aid of the micromanipulator, they were 

able to isolate each of the 4 spores from an ascus, and in addition to 

finding that genetic segregation occurred within the ascus, the possibility 

presented itself of utilizing the micromanipulator technique 

in spore crossings and tetrad analysis. By pairing two spores and by 

observing under the microscope their subsequent copulation, the first 

artificial yeast hybrids were thus produced. These hybrids (14 interspecific 

and 1 intergeneric) were purposely produced from parent 

types which differed in their ability to ferment various sugars in 

order that the inheritance of the enzymes responsible for fermentation 

could be investigated; but although the hybrids showed heterosis and 

sporulated abundantly, their spores unfortunately germinated very 

poorly, making impossible a detailed genetic analysis of the enzymatic 

characters involved in this material. An important fact was adduced, 

however, from these experiments and that was that a hybrid 

between a fermenter of any particular sug ar and a non-fermenter 

would always be capable of fermenting the sugar, and, in addition, 

that there was always an excess of fermenters among the few viable 

spores from such hybrids. Thus the conc1usion to be drawn from 

these results was that in all instances studied, "the ability to synthesize 

a specific enzym e is dominant in the Fl." Nothing more definite 

could be stated, and such was the state of our knowledge until 

1943, when LINDE GREN began his researches in yeast genetics. 

1943-1944 

LINDEGREN'S first contribution to the field was an important 

one - namely, the discovery in 1943 of heterothallism in Saccharomyces. 

By tetrad analysis he was able to show that from each 4spored 

ascus 2 spores belong to one mating type and two to another 

and that therefore mating type specificity is controlled by a single 

pair of allelomorphs. He also described a phenomenon which he called 

"illegitimate copulation", in which he maintained that self-diploidization 

of a single-spore culture invariably lea ds to poor sporulation 

and low viability of the ascospores. Although heterothallism in 

Saccharomyces has been confirmed, it is not true that self-diploidized 

single spore cultures usually sporulate poorly and produce spores with 

poor germination; we have in our work generally found the opposite 

to be the case.

- 163- 

Figs. 1-4. WINGE and LAUSTSEN'S method for isolating the four spores 

within one ascus with the aid of a micromanipulator equipped with two 

glass needles. - (Af ter WINGE & LAUSTSEN, 1937). 

Figs. 5-6. Artificial species hybridization in yeasts by the pairing of two 

ascospores in a nutrient droplet with the aid of a micromanipulator. The 

smalle r spore is from Saccharomyces validus and the larger from S. cere 

visiae. Copulation of the two spores results in a hybrid zygote, the first bud 

of which is seen in Fig 6. - (Af ter WINGE & LAUSTSEN, 1938). 

2. 

11*

- 164 - 

These investigations of LINDEGREN led, in 1944, to a new method 

for hybridizing yeasts. Instead of employing the spore crossing tech 

nique of WINGE and LAUSTSEN, he simply mixed two haplophase cul 

tures of opposite mating type together in a small amount of nutrient 

medium, and after a period of incubation to allow for copulation, he 

transferred the mixture to a sporulating medium. Spores to be used 

in genetic analyses were then isolated from the asci which arose in this 

mixture. The advantage of this method over the spore crossing method 

is, as LINDEGREN justly claimed, the fact that the parents are not 

"used up" in the crossing and can therefore be employed in further 

study. On the other hand, the hybrids themselves are not available 

for analysis unless individual zygotes or their derivatives are isolated 

from the mixture. A more serious objection to this technique is that 

self-diploidization of haploid yeasts occurs frequently, so that the 

so-called hybrids obtained may actually represent the diplophase of 

one or the other parent. 

Employing thi s new hybridization technique, LINDEGREN and 

SPIEGELMAN in 1944 were successful in producing hybrids between 

S. carlsbergensis and S. cerevisiae which were completely fertile, i. e., 

a high percentage of the spores were viable, thus making possibIe for 

the first time the tetrad analyses of artificially produced yeast hy 

brids. One of the parents, S. carlsbergensis) is able to ferment raffinose 

completely, while the other parent, S. cerevisiae) ferments only 1/3 of 

it - in other words, S. carlsbergensis is capable of fermenting meli 

biose through the activity of a specific enzyme, while S. cerevisiae is 

not. As was to be expected from the earlier researches of WINGE and 

LAUSTSEN, the hybrid itself fermented melibiose. Tetrad analyses of 

the spores from this hybrid yielded segregation ratios which excluded 

the possibility that only one gene in S. carlsbergensis was involved in 

the production of the enzyme, since in addition to ratios of 2 fermen 

ters: 2 non-fermenters, ratios of 4:0 and 3:1 were also obtained. The 

conclusion which LINDEGREN and SPIEGELMAN drew from this investi 

gation was that S. carlsbergensis contains 2 dominant genes, each of 

which is able to initiate the synthesis of the enzyme when melibiose 

is present, and that the other parent, S. cerevisiae) contains the two 

recessive alleles. In other words, this was interpreted as an example 

of true mendelian inheritance. 

SPIEGELMAN and LINDEGREN then turned to a more detailed investi 

gation of the mechanism of enzymatic adaptation in yeasts by com 

paring this process in haploid and diploid strains of the same species.

- 165 - 

A stable, homogeneous diploid was found to adapt to galactose fermentation 

in the absence of cell division, while genetically unstable 

haploids were found to adapt only during growth. This was taken to 

mean that both cytoplasmic interaction as well as mutation and 

natural selection may be operative in any particular adaptive enzyme 

system according to the strain employed, and in order to study the 

biochemistry of enzymatic adaptation, the importance of eliminating 

natural selection by employing a diploid strain of high genetic stability 

was stressed. 

Thus in the 9 years which had elapsed since the publication of 

Professor WINGE'S first paper on the life cycle of Saccharomyces 

sufficient evidence had accumulated to indicate that inheritance in the 

yeasts, just as in other organisms, followed classical mendelian laws, 

and, in addition, that the yeasts were exceedingly promising organisms 

for future investigations in the field of biochemical genetics. 

But it was during the years following 1944 that startling and revolutionary 

hypotheses based upon the yeast genetic work performed in 

the United States were proposed, which, if they are to be accepted, 

would mean profound changes in our concept of the gene. It is thi s 

work during the period 1945 to 1947 that I would like to discuss now. 

1945-1947 

,sPIEGELMAN and LINDEGREN repeated by means of the mass mating 

technique the crossing between S. carZsbergensis) a melibiose-fermenter, 

and S. cerevisiae) a non-fermenter of melibiose. The strain of 

S. carZsbergensis used in this crossing was now assumed to have only 

one gene for melibiase production, since 10 asci from the hybrid 

segregated out regularly as 2 fermenters: 2 non-fermenters. However, 

when the mating, sporulation, and planting were undertaken in the 

presence of melibiose, 6 asci gave segregation ratios of 4 fermenters: 

O non-fermenters. When melibiose was removed from the substratum, 

the 4:0 ratios reverted to the normal 2:2 ratios. However, before the 

single spore cultures carrying the recessive allele had completely lost 

their enzyme content in the melibiose-deficient medium, they were 

re-exposed to melibiose, and it was found that all cultures showed a 

marked increase in enzymatic activity. The conclusions to be drawn 

from thi s experiment were significant. First, it was assumed that the 

4:0 ratios resulted from the fact that the 2 normally negative cultures 

from each ascus were able to ferment due to the transfer of enzyme-

- 166- 

forming factors from the cytoplasm of the hybrid to that of the spores. 

Secondly, it was assumed that here was an example of the maintenance 

and synthesis of an enzyme in the complete absence of the 

gene. These conclusions, of course, were based on the implicit assumption 

that melibiase production is due to the presence of a dominant 

gene, the recessive allelomorph of which is unable to initiate the 

formation of the enzyme. As will be seen later from a discussion of 

our work on galactose fermentation, another explanation is possible. 

In the years 1945 and 1946 there followed a series of papers by 

LINDEGREN dealing with mendelian and non-mendelian inheritance 

and with the cytogene theory, proposed to account for the behaviour 

observed. It is scarcely worthwhile to discuss in detail the experiments 

upon which the cytogene theory is based, since the theory has later on 

been completely abandoned by LINDEGREN himself, but a few general 

remarks concerning it may be of interest. In addition to the experiments 

I have aIready discussed, LINDEGREN produced hybrids heterozygous 

with respect to galactozymase, but instead of the expected 2:2 

segregations from the hybrid, 4:0 ratios were obtained. This was 

interpreted as cytoplasmic transfer - i. ,e., the adaptive enzyme, or 

"cytogene" was assumed to be transmitted through the cytoplasm 

and maintained in cells lacking the dominant gene. Thus the recessive 

cells were contaminated by the cytogene and continued to ferment as 

long as the specific substrate was present. Subsequent experimental 

work by LINDE GREN yielded a large number of irregular segregations 

which he maintained could not be accounted for in terms of multiple 

genes, but which, he claimed, gave further support for the cytogene 

theory. At the Cold Spring Harbor Symposium in 1946 this theory 

was considerably revised, in that a new phenomenon was considered 

to be involved - namely, "masked recessiveness". Again, in his genetic 

analyses, LINDE GREN had obtained a large number of irregular 

segregation ratios, the majority of which showed an excess of fermenters. 

One ascus, in particular, derived from a MmGg hybrid yielded 

4 M fermenters and 4 G fermenters. This was explained by assuming 

that the recessive alleles were "masked" by the acquisition at meiosis 

of the dominant genotype - i. e., m becomes M and g becomes G. 

When such a "masked recessive" was crossed to a normal recessive, 

2:2 ratios were obtained, but when it was crossed to a normal dominant, 

not 4 :0, but 2:2 ratios were likewise obtained. A satisfactory 

and logical explanation of this behaviour appears impossible, and the 

experimental results seem only to throw doubt upon the validity of

-167 - 

the concept of masked recessiveness. In addition to these theoreticaI 

considerations, which, it must be remembered, were primarily based 

upon the behaviour of one exceptional ascus, LINDEGREN stated that 

he had repeated the melibiose experiment to which I have aiready 

alluded - namely, that maintenance and increase of an adaptive 

enzyme in the absence of the gene occurs through cytoplasmic transfer 

from the dominant type. The original results however, could not be 

duplicated, since just as many 4:0 ratios were obtained this time in the 

absence of the substrate as were obtained in its presence ; this was 

regarded by LINDEGREN as further evidence for the existence of 

maske d recessives. It is worthy of note that in the discussion following 

the presentation of this cytogene theory with all of its complexities, 

LINDE GREN stated in response to a direct question that he would not 

even consider the multiple factor hypothesis as an explanation for 

his experimental results, as he deemed it "too elaborate to be 

justified" . 

At this same Symposium SPIEGELMAN also discussed, with special 

reference to yeast investigations, the gene-enzyme problem, wherein 

he introduced the plasmagene hypothesis. It is important to note, 

however, that SPIEGELMAN considered that inheritance in yeasts was 

completely mendelian; his "plasmagenes" were concerned therefore 

with the mechanism of gene action, rather than with gene transmission. 

Apparently unaware that LINDE GREN was unable to repeat the 

now famous experiment involving the apparent synthesis and maintenance 

of an adaptive enzyme in the absence of the gene, he endeavoured 

to account for this phenomenon in terms of plasmagenes. His 

argument was as follows: if 2:2 ratios are normally obtained from a 

heterozygous hybrid, but 4:0 ratios are obtained in the presence of 

the specific substrate, then the adaptive enzyme system must be part 

of a cytoplasmic self-duplicating mechanism, and once the enzyme is 

formed, its further produetion ean proceed independently of the presence 

of the gene. In other words, the 2 spores which carry the recessive 

alle le obtain from the cytoplasm of the hybrid replicas of the 

dominant gene, which he called the plasmagenes. These plasmagenes 

are self-duplicating and combine with precursor protein and with the 

substrate to form the enzyme, thus allowing the formation of a specific 

enzyme to occur within cells which lack the gene normally necessary 

for its synthesis. The important feature of SPIEGELMAN'S theory 

is, therefore, that the genes are continually producing nucleo-protein 

replicas of themselves which pass into the cytoplasm of the cello 

In the ensuing year, 1947, a series of papers by SPIEGELMAN and

- 168 - 

his colleagues appeared which de alt with the purely biochemical 

aspects of enzymatic adaptation. It will suffice to say that they were 

able to demonstrate the occurrence of competitive interaction between 

adaptive enzyme systems, a phenomenon which SPIEGELMAN had 

earlier predicted on the basis of the plasmagene hypothesis, since the 

different plasmagenes must necessarily com pete with one another for 

protein and energy, and as he termed it, "the outcome determines the 

enzymatic constitution of the cell". 

Meanwhile LINDEGREN had turne d from a consideration of the 

inheritance of the enzymes responsibIe for the fermentation of carbohydrates 

to a study of the inheritance of the genes involved in vitamin 

synthesis. Working with biotin-, pantothenic acid-, para-aminobenzoic 

acid-, and thiamine-deficient strains, and crossing these with synthesizers, 

he found for the most part that 2:2 segregations were obtained 

from the hybrid progeny. In other words, not only was mendelian 

inheritance operative in the synthesis of vitamins by yeasts, but 

the ability to synthesize appeared to be controlled in each instance 

by a single gene. However, further investigations appeared to throw 

doubt on the simplicity of the phenomenon. In a paper published 

together with RAUT in 1947, a pantothenate + type was crossed with 

a pantothenate -type. The hybrid produced was, as expected, capable 

of synthesizing pantothenate. However, 4 spores from 1 ascus derived 

from this hybrid were all pantothenate +. This case was explained 

by the authors as being due to "gene transformation", 

Still another complicating factor was introduced into yeast genetics 

by LINDE GREN in 1947 - namely, "depletion mutations". Since 

time does not permit a critical discussion of all of LINDEGREN'S 

theoreticai considerations, it will suffice in this case to state that 

he conceived a depletion mutation as a phenomenon in which a modification 

is transmitted indefinitely during vegetative reproduction but 

which is unable to be transmitted through the sexual cycle. According 

to LINDEGREN, this behaviour, which phenotypically resembles gene 

mutation, is not due merely to the effect of the environment upon the 

cytoplasm, but is actually under genetic control. 

Thus, to summarize, thi s period from 1945 to 1947 was characterized 

by the production by the American investigators of a large 

number of yeast hybrids, the progeny of which yielded both regular 

segregation ratios in accordance with classical mendelism, but also 

irregular, so-called "non-mendelian" ratios. By way of explanation, 

LINDEGREN proposed his cytogene theory and developed his concepts

- 169- 

of masked recessiveness and depletion mutations. During the same 

period SPIEGELMAN'S plasmagene theo ry was proposed in order to 

account for the maintenance and increase of an adaptive enzyme in 

the absence of the gene. Now I would like to discuss some of the work 

on adaptation and the inheritance of enzymatic character s in yeasts 

which Professor WINGE and I undertook at the Carlsberg Laborato 

rium, and which first appeared in 1948. 

1948 

By the spore-crossing technique a hybrid was produced between 

S. cerevisiae and S. chevalieri with sufficient fertility to make possibIe 

genetic investigations by tetrad analyses. The two parents are distin 

guished by the fact that S. cerevisiae is able to ferment both maltose 

and galactose, while S. chevalieri is unable to do so. Here, then, we 

had the possibility of determining what type of enzymatic inheritance 

was operative in thi s material; our experimental results established 

incontestably that it was mendelian. 

First, with regard to maltose fermentation, our original hybrid 

yielded the following segregation ratios: 9-4:0 and 3-3 :1, which 

of course excluded the possibility that 1 or 2 genes were involved in 

the production of maltase. If it is assumed that 3 genes are present, 

and that 50 % crossing over occurs between all three genes and their 

centromeres, then the theoretically expected ratios would be 52.8% 

4 :0, 44.4% 3 :1, and 2.8% 2 :2. Although agreement between the ob 

served and the expected was not close, it nevertheless was indicative 

of the fact that 3 genes were involved, and in orde r to test the validity 

of thi s hypothesis, we attempted several backcrossings of the single 

spore progeny of the hybrid to the recessive S. chevaZieri. Assuming 

that 3 genes are involved, any spore from the hybrid would contain 

either O, 1, 2, or 3 genes, and by crossing the fermenters to the reces 

sive, one should expect to obtain a 1, 2, or 3 gene segregation in the 

backcross hybrid's progeny. Such indeed was the case. One of our 

backcross hybrids yielded a typical 1 gene segregation (14-2 :2) and 

two others yielded 2 gene segregations (12-3:1 and 5-2:2; and 

4-4:0 and 16-3 :1). The conclusion to be drawn from these results 

is, therefore, that S. cerevisiae apparently contains 3 non-linked 

polymeric genes, Ml) M2) and M3) for the production of maltase. 

Now, with regard to galactose fermentation, we found at the outset 

of our experiments that although S. chevalieri had been reported to

- 170- 

be incapable of fermenting galactose, it actually was capable of doing 

so, but only slowly, after a period of adaptation. The tetrad analysis 

of 12 asci of our original hybrid showed in all cases an unmistakable 

2:2 segregation, in that 2 cultures fermented galactose rapidly (1-2 

days) and 2 slowly (5-14 days). We assumed therefore that S. cere 

visiae contained only 1 gene for the production of galactozymase. 

The tetrad analyses of our backcross hybrids which were heterozygous 

for galactose fermentation confirmed this assumption. In all cases, 

a 2:2 segregation was obtained. In the light of the earlier investiga 

tions of LINDEGREN and SPIEGELMAN on the mechanism of melibiose 

adaptation, these findings were of considerable significance. They had 

implicitly assumed that the recessive allelomorph was incapable of 

producing the specific enzyme and consequently that when a single 

ascus from a heterozygous hybrid yielded 4:0 ratios in the presence 

of the substrate, enzyme synthesis in the absence of the gene must 

be involved. We, 'On the other hand, found that in the case of galac 

tozymase, the recessive allelomorph of G was not g} but 9 s which 

controlled the slow synthesis of the enzyme; in other words, a "long 

term adaptation" was involved in the fermentation of galactose by 

the recessive alleles. Further studies of thi s long-term adaptation 

convinced us that it was not result of mutation and natural selection, 

since we were able to adapt, de-adapt, and re-adapt our gs types at 

will, depending upon whether they were grown in a galactose-contain 

ing or galactose-deficient medium. We consider, therefore, that long 

term adaptation is an expression of the fact that enzyme production 

in the presence of the sugar is gradually increased until it exists in a 

sufficiently high concentration to bring about visible fermentation. 

1949 

The year 1949 brought forth criticism of our views concerning the 

nature of long-term adaptation, as well as both criticism and confir 

mation of our views concerning the mendelian nature of the inheri 

tance of enzymatic characters in yeasts. 

MUNDKUR and LINDEGREN had reinvestigated the phenomenon of 

long-term adaptation to galactose fermentation but came to the con 

clusion that it could be explained entirely on the basis of mutation 

and natural selection. This explanation was based on the assumption 

that 3 phenotypes existed: G} a rapid fermenter; g} a non-fermenter; 

and gx a slow fermenter, and that it is through chance mutation from

- 171 - 

g -+ G that the phenotypic slow fermenter, gXJ which is actually a 

mixture of G and g cells, is produced (x signifies the number of days 

elapsing before fermentation is visible). It is true that our g s types 

adapted after varying periods of time and that it would be tempting 

to conclude that these discrepancies involved chance mutations, but, 

on the other hand, we had only 2 phenotypes in our material, a rapid 

and slow fermenter, and had never isolated a spore belonging to 

MUNDKUR and LINDEGREN'S category of "non-fermenters." Another 

point to bear in mind is that we were unable to de-adapt our G type. 

If we accept LINDEGREN'S view that long-term adaptation is due to 

mutation, then the logical consequence would be that in addition to 

the stable G gene, there also exists an unstable G gene which readily 

mutates to g. We cannot accept this view of 2 different G genes in 

S. cerevisiae for the synthesis of galactozymase. 

Through an investigation of the behaviour of non-synthesizers of 

thiamine, tryptophane, adenine, and methionine in crosses with syn 

thesizers, POMPER and BURKHOLDER found a 2:2 segregation for each 

character, demonstrating that only 1 gene was involved in each bio 

synthesis. With regard to uracil synthesis, non-allelic complementary 

genes appeared to be involved. The conclusion arrived at by these 

authors was that "the characters segregated independently and 

regularly according tomendelian principles." 

GILLILAND, using the spore crossing technique, produced a hybrid 

between S. italicus and S. chevalieri which was heterozygous in 4 

fermentation characters, and he was able to demonstrate conclusively 

by tetrad analyses that S. italicus contains 1 gene for the production 

of maltase (which has subsequently been identified as Ml) and 1 gene 

for the production of galactozymase. With regard to raffinose, 

GILLILAND obtained a 3-gene segregation from 20 asci; these asci gave 

exactly the same 3 gene segregation for sucrose fermentation, the 

only difference being that the sucrose recessives, which corresponded 

to the raffinose negatives, were actually able to ferment sucrose 

slowly, after adaptation. GILLILAND concluded that S. chevalieri 

apparently contains 3 polymeric genes for the production of (3-h 

fructosidase, an enzyme capable of hydrolyzing both sucrose and 

raffinose, but that definite proof of this assertion would have to be 

obtained from backcrosses to the recessive. An additional observation 

of interest was that all the raffinose-negative, sucrose-slow cultures 

were able to ferment maltose, indicating that a-glucosidase may be 

present in the other parent, S. italicus.

- 172- 

It was also in 1949 that LINDEGREN'S book, "The Yeast Cell", 

appeared. I will make no attempt to review thi s book, which, although 

containing linkage calculations and the first chromosome maps of the 

yeasts, deals primarily with much of the material already discussed. 

I would like to emphasize, however, that in this book LINDEGREN 

rejects both classical mendelism and his own cytogene theory, and 

proposes an elaboration of the theory involving the conversion or 

contamination of alleles to account for inheritance behaviour in 

yeasts. Since LINDE GREN is apparently convinced that all enzymatic 

characters in yeasts which have been studied are controlled by single 

genes, he should obtain only 2:2 segregations in his tetrad analyses 

of heterozygous hybrids. This, of course, he does not. Therefore the 

irregular ratios, which for the most part yield an exces s of the do 

minant type, are interpreted as being due to the contamination of the 

recessive allele by the dominant allele by contact at synapsis during 

meiosis. Gene conversion is regarded as being controlled by heredity, 

since it occurs with higher frequency in some lines; thus he would 

have, for example, a gene infIuencing the stickiness of the chromo 

some. I think it is clear that the highly speculative nature of such 

considerations, interesting as they may be, does not warrant a more 

detailed discussion. It will suffice to quote LINDE GREN directly in 

regard to his concept of the gene: "The explanation ... fits all the 

cases without any fundamental difficulty except that it requires 

alteration of our apparently well-established views concerning the 

stability of the gene." 

I would like at this time to mention the work of EPHRUSSI and his 

colleagues which also appeared in 1949 and which de alt for the first 

time with the inheritance of respiratory enzymes in yeasts. Mutants 

deficient in cytochrome oxidase and characterized by the production 

of small colonies were induced through the action of acrifIavine. 

These mutant s were employed in extensive crossing experiments with 

the result that only a very small percentage of them reappeared in 

the progeny of the hybrids. If the results are to be explained on the 

basis of multiple genes, EPHRUSSI has calculated that at least 16 

dominant genes must be involved. Since this seems highly improb 

able, it is assumed that the acriflavine-induced mutants have lost 

self-duplicating cytoplasmic particles which are essential for enzyme 

synthesis. Accordingly, these first investigations on the inheritance 

of respiratory enzymes point to the interesting fact that cytoplasmic, 

rather than mendelian, inheritance is involved.

- 173- 

1950-1951 

One of the most interesting of the recent contributions is a paper 

by SPIEGELMAN, SUSSMAN, & PINSKA dealing with the nature of longterm 

adaptation in yeasts. It will be recalled that we have explained 

the phenomenon by assuming that a type containing the recessive 

allele is a slow synthesizer of the enzyme and that it becomes adapted 

through a gradual accumulation of enzyme in the presence of the 

substrate; MUNDKUR and LINDEGREN, however, consider that the 

phenomenon is due entirely to the occurrence of chance mutations 

from non-fermenters to fermenters. SPIEGELMAN, on the other hand, 

considers both explanations inadequate and assumes that long-term 

adaptation is cytoplasmic in nature. His experimental results indicate 

that during the period of adaptation, the population of a gs type 

actually becomes heterogeneous through production of so-calle d 

"positive" cells by the original "negatives". These modifications are 

not gene mutations; they were found to be induced only by the 

presence of the substrate, and in its absence, the positive cells were 

stated to produce negative buds. SPIEGELMAN believes that such a 

phenomenon can be explained only on a cytoplasmic basis, and furthermore, 

that it represents additional evidence in support of the 

plasmagene hypothesis. The theory is admittedly attractive, but 

similar investigations of adaptive enzym e systems other than galacto· 

zymase are necessary to prove its universal validity. 

The final contribution I would like to discuss deals with the genes 

responsibIe for maltose fermentation. Professor WINGE and I undertook 

a further study of this problem in order to establish conclusively 

that polymeric genes are involved in the production of maltase, and 

that the inheritance of fermentative characters in yeasts is mendelian 

in nature. This I believe we have done. The work involved the isolation 

of the 3 M genes, as well as a fourth M gene, M4) which arose 

both spontaneously and as aresult of Rontgen irradiation. The identit 

Y of these 4 genes was firmly established by extensive hybridization 

work, involving the artificial production of 81 new hybrids. In 

all hybrids in which 2 different M genes were present, the progeny 

yielded 2·gene segregations, whereas the progeny of hybrids between 

types containing the same gene yielded no segregation. In addition 

to these investigations, work is now in progress on the inheritance 

of the ability to ferment raffinose and sucrose. It can be stated that 

we have completely confirmed GILLILAND'S findings with regard to

- 174- 

the existence of 3 polymeric genes for these fermentations, and at 

present we have aiready isolated 2 of the 3 genes involved - namely, 

R2 and Ra. 

Finally, to summarize the developments which have taken place in 

the field of yeast genetics, it ean be said that the work was initiated 

here in Denmark on the assumption that inheritance in yeasts was 

mendelian in nature. Subsequent investigations by LINDEGREN demonstrated 

the occurrence of apparent non-mendelian segregation ratios 

with regard to fermentation characters and the synthesis of vitamins 

and amina acids. Accordingly, his concepts of the nature of inheritance 

in yeasts passe d through three distinct phases: first, classical 

mendelism; second, cytoplasmic inheritance, embodied in thc 

cytogene theory; and finally, his current view of the instability of 

the gene. It is therefore a matter of considerable interest that the 

recent investigations of EPHRUSSI have indicated that cytoplasmic as 

well as mendelian inheritance is involved in the synthesis of respiratory 

enzym es in yeasts. I think that it is evident, however, that the 

continuation of the work at the Carlsberg Laboratorium, complicated 

as it may be by the related problem of enzymatic adaptation, has 

nevertheless conclusively established that the inheritance of fermentative 

characters in the yeasts which we have investigated is mendelian 

and in some cases involves the existence of polymeric genes. 

Whatever the reasons for the disparity between LINDEGREN'S and our 

experimental results may be, and the occurrence of self-diploidization, 

overlapping of generations, and 8-nucleate asci may be mentioned 

in this connection, the faet remains that we have not obtained segregation 

ratios in disaccord with MENDEL'S laws. We have no choice, 

therefore, but to consider the concepts of cytogenes, depletion mutations, 

masked recessives, and gene conversions to be superfluous, and 

to regard the inheritance of fermentative characters in yeasts as 

completely mendelian. 

Postscript added January 1956. 

Limitation of space precludes more than a brief mention of some 

of the major contributions to the field made during the past five 

years. 

Interest in the problem of irregular segregation of yeast tetrads 

has continued unabated, and several different explanations of this 

phenomenon have been advanced. In 1953 MUNDKUR observed the

- 175- 

localization and movement of mitochondria within the living yeast 

cell, and knowledge of the localization of enzymes within mitochondria 

led him to con cl ude that the random distribution of mitochondria 

within mother cells and buds could be responsibie for vegetative 

mutants and genetic irregularities. Another explanation, based upon 

both genetic and cytological evidence, has been found in the occurrenee 

of supernumerary mitoses in the ascogenous cells of yeast 

monohybrids (WINGE & ROBERTS 1954). If the normal number of 4 

nuclei in the immature ascus is increased to 8 with consequent duplication 

of the genotypes, and if some of the nuclei (or spores) do not 

survive, then it ean be a matter of chance which genotype s the surviving 

spores contain. In addition, these extra mitoses may lead to 

the formation of binucleate heterozygous or homozygous spores, and 

examples of irregular segregation in LINDEGREN'S haploid material 

revealed that one of the 4 single-spore cultures in each of 3 irregular 

tetrads was diploid and heterozygous. Still another explanation - 

and one which enjoys many adherents - involves the occurrence of 

polyploidy in yeasts. It is obvious that polyploid inheritance in a 

yeast would result in segregations quite different from the 2:2 

expected from a monohybrid, and claims of its existence were set 

forth in 1951 by LINDEGREN & LINDEGREN and by ROMAN, HA WTHORNE, 

& DOUGLAS. In 1954 further evidence for polyploid segregation was 

presented by POMPER & McKEE and by LEUPOLD & HOTTINGUER, while 

a still more detailed study by ROMAN, PHILLIPS, & SANDS appeared 

the following year. It should be noted that these investigators all 

worked with haploid material, whose single spore progeny could, as a 

rare event, diploidize, and by mixing such a clone with another 

haploid or diploid of opposite mating type, triploids or tetraploids 

were obtained in small numbers. Diploid material, on the other hand, 

contains the D-gene which brings about diploidization whenever 

present and also appears to abolish the expression of mating type. 

The fusion of diploid cells to form tetraploids or the fusion of a 

diploid and a haploid cell has not been found in this material. 

Another important advance has been the demonstration of the 

existence of complementary genes in the inheritance of fermentative 

characters in yeasts. LEUPOLD & HOTTINGUER (1954), working on 

galactose fermentation, obtained segregation ratios which led them 

to conclude that 2 complementary genes were involved in the fermentation 

of this monosaccharide. The elegant investigations of GILLILAND 

(1954) on Saccharomyces diastaticus have elucidated the genetics of

- 177- 

super-attenuation, or dextrin fermentation, which is of practical significance 

in brewing and, at the same time, have demonstrated the 

occurrence of complementary genes in this material. He isolated the 

gene S) which produces amylase, responsibIe for the hydrolysis of 

dextrin to maltose, Ml) which is responsibIe for the rapid fermentation 

of maltose, and a new M-gene, M5) which is responsibIe for the 

slow fermentation of maltose. For super-attenuation to occur, therefore, 

the presence 'Df both S and an M-gene is required. Finally, with 

regard to complementary gene action, it has been demonstrated 

(WINGE & ROBERTS 1956) that raffinose fermentation may be brought 

about not only by the R-genes responsibIe for the synthesis of (3-hfructosidase 

which hydrolyzes the raffinose molecule into melibiose 

and fructose, but also by the combined action of the melibiase gene, 

Me) and the galactozymase gene, G. It was demonstrated that melibiase 

alone is able tD hydrolyze the raffinose molecule into sucrose 

and galactose, but if galactozymase is also present, then the galactose 

portion is fermented away, resulting in a 1/3 fermentation of the 

raffinose molecule. 

Finally, mention should be made of the significant contribution of 

WINGE (1955) in the form of a new hypothesis termed interallelic 

crossing-over, an hypothesis which was not evolved in order to. account 

for irregular segregation but to account for the occurrence in 

yeasts of extremely close linkage between two pairs of fermentative 

genes (Ml-RI and M3-R3) which could not reasonably be credited to 

chance. While not denying the validity of mendelian law, this hypothesis 

regards these two linked genes actually as one complex gene 

molecule, whose individual molecular groups may undergo interallelic 

crossing-over. It differs essentially from the conversion hypothesis of 

WINKLER and of LINDE GREN in that the result of this new type of 

crossing-over is not the transformation of a dominant or a recessive 

into its allelomorph, but the creation of two new allelomorphs, and 

it is significant that one of its fundamental assumptions is that the 

enzymes synthesized by these closely-linked genes are probably chemically 

similar, since the difference between them does not exceed 

more than a portion of the whole molecular complex. 

LITERATURE 

Ephrussi, Boris: Nucleo-cytoplasmic relations in micro-organisms. - Oxford 

University Press, 1953. 

FRIESIA V 12

- 178- 

Gilliland, R. B.: A yeast hybrid heterozygotic in four fermentation characters. 

- Compt. Rend. d. Lab. Carlsberg, Ser. Physiol. 24: 347- 

356. 1949. 

: A study of a wild yeast - S a c c h a r o m y c e s d i a s t at 

i c u s. - Wallerstein Lab. Comm. 17: 165-175. 1954. 

LeupoldJ Urs & Hottinguer, Helene: Some data on segregation in S a cc 

h a r o m y c e s. - Heredity 8: 243-258. 1954. 

Lindegren, Carl C. & Lindegren, Gertrude: Selecting, inbreeding, recombining, 

and hybridizing commercial yeasts. - Journ. Bact. 46: 

405-419. 1943. 

: A new method for hybridizing yeast. - N atl. Acad. Sci. Proc. 

29: 306-308. 1943. 

Lindegren, Carl C., Spiegelman, S., & Lindegren, Gertrude: Mendelian inheritance 

of adaptive enzymes in yeast. - Natl. Acad. Sci. Proc. 

30: 346-352. 1944. 

Lindegren, Carl C. & Lindegren, Gertrude: The cytogene theory. - Cold 

Spring Harbor Symposia Quant. Biol. 11: 115-129. 1946. 

: Depletion mutation in S a c c h a r o m y c e s. - Natl. Acad. 

Sci. Proc. 33: 314-318. 1947. 

: Tetraploid S a c c h a r o m y c e s. - J ourn. Gen. Microb. 5: 

885-893. 1951. 

: Proximity of gene s controlling the fermentation of similar 

carbohydrates in S a c c h a r o m y c e s. - Nature 170: 965- 

968. 1952. 

Carl C.: Concepts of gene-structure and gene-action derived 

from tetrad analysis of S a c c h a r o m y c e s. - Experientia 9: 

75-80. 1953. 

Carl C. & Lindegren, Gertrude: Asci in S a c c h a r o m y c e s 

with more than four spores. - Genetics 38: 73-78. 1953. 

Mundkur, Balaji D. & Lindegren, C. C.: An analysis of the phenomenon of 

long-term adaptation to galactose by S a c c h a r o m y c es. 

- Arner. Jour. Bot. 36: 722-727. 1949. 

B. D.: Mitochondrial distribution in S a c c h a r o m y c e s. - 

Nature 171: 793-794. 1953. 

Pomper, Seymour & Burkholder, P. R.: Studies on the biochemical genetics 

of yeast. - Nat. Acad. Sci. Proc. 35: 456-464. 1949. 

S., Daniels, K. M. & McKee, D. W.: Genetic analysis of polyploid 

yeast. - Genetics 39: 343-355. 1954. 

Roman, Herschel, Hawthorne, Douglas C., & Douglas, Howard C.: Polyploidy 

in yeast and its bearing on the occurrence of irregular 

genetic ratios. - N atl. Acad. Sci. Proc. 37: 79-84. 1951. 

Phillips, Marcia M., & Sands, Stanley M.: Studies of polyploid 

S a c c h a r o m y c e s. L Tetraploid segregation. - Genetics 

40: 546-561. 1955. 

Spiegelman, S., Lindegren, Carl C., & Lindegren, Gertrude: Maintenance 

and increase of a genetic character by a substrate-cytoplasmic 

interaction in the absence of the specific gene. - N atl. Acad. 

Sci. Proc. 31: 95-102. 1945.

- 179- 

Spiegelman, S., Sussman, Racquel Rotman, & Pinska, E.: On the cytoplasmic 

nature of "long-term adaptation" in yeast. --:- Natl. Acad. Sci. 

Proc. 36: 591-606. 1950. 

Winge, O.: On haplophase and diplophase in som e S a c c h a r o m y c e t e s. 

- Compt. Rend. d. Lab. Carlsberg, Ser. Physiol. 21: 77-11l. 

1935. 

& Laustsen, O.: On two types of spore germination, and OD 

genetic segregations in S a c c h a r o m y c e s, demonstrated 

through single-spore cultures. - Ibid. 22: 99-116. 1937. 

& Laustsen, O.: Artificial species hybridization in yeast. - 

Ibid. 22: 235-244. 1938. 

& Laustsen, O.: On 14 new yeast types, produced by hybridi 

zation. - Ibid. 22: 337-352. 1939. 

& Roberts, Catherine: Inheritance of enzymatic characters in 

yeasts, and the phenomenon of long-term adaptation. - Ibid. 24: 

263-315. 1948. 

& Roberts, Catherine: The polymeric gene s for maltose fermen 

tation in yeasts, and their mutability. - Ibid. 25: 35-83. 1950. 

& Roberts, Catherine: Non-mendelian segregation from hetero 

zygotic yeast asci. - Nature 165: 157. 1950. 

& Roberts, Catherine: Causes of deviations from 2:2 segre 

gations in the tetrads of monohybrid yeasts. - Compt. Rend. d. 

Lab. Carlsberg, Ser. Physiol. 25: 285-329. 1954. 

: On interallelic crossing over. - Ibid. 25: 341-354. 1955. 

& Roberts, Catherine: Complementary action of melibiase and 

galactozymase on raffinose fermentation. - Nature 177: 383- 

384. 1956. 

Copenhagen, J anuary 1956.

- 181- 

("Buckthorn Brown", "Brussels Brown"), often with an olivaceous 

("Tawny Olive") and paler reddish tinge ("Morocco Red"), as older 

darker ("Auburn", "Reeds Brown"), on pressure immediately dark 

brown; cuticle not separable. 

T u b e s adnate to adnate sinuate, the walls of the tubes decurrent 

with a tooth on the stem, 0.5 to 2 cm long, yellow, lemon gold ("Strontium 

Yellow") with a tinge of olive, on injury becoming instantly 

very blue. 

p o r e s roundish to angular, in age edged and denticulate, lemon 

to gold, then pale greenish, at last rust y brown, immediately turning 

dark blue at touch. 

S t i p e 3.5-6 X 1-1.5 cm (sometimes broader at apex, to 3.5 cm), 

variable in stature, but mostly cylindraceous, at the base curved 

and radicating, dry, velutinous punctate, at the apex gold, red brown 

to dark brown at the base. 

C o n t e x t soft, gummous, citrin to gold, under the cuticle 

brownish to reddish, instantly turning blu e when cut, in the stem 

dark purpIe below. 

S p o r e s yellow, ellipsoid to fusiform, 10-12-14 (-16) X 4.5- 

5 (-6)1'. 

C Y s t i d i a clavate, fusiform, 40-75 X (5)-9-121'. 

T a x o n o m i c R e m a r k s. Owing to the strong inconstancy 

in colour in young and old stages the determination of B. pulverulentus 

often meets with difficulties. It is primarily characterized by 

the coarse, first yellow, mild context, immediately turning blue on 

injury and all parts being dark to almost black on pressure. 

The specimens found by the present writer remind us mainly of 

those described by SKOVSTED (1940-41) from Denmark. They agree 

best with BRESADOLA'S figure in leon. myc. (1931, vol. XIX, plate 

916, sub nom. radicans) and with USAK'S excellent pieture (Ceska 

mykologi e 1953, plate 11), but not with KONRAD & MAUBLANC'S 

figures (1924-37) which are too dark. Older stages have a strong 

likeness to OPATOWSKI'S (1836) original pieture. 

H i s t o r y. As regards the general historicaI survey of the 

species cf. KALLENBACH (1927, pag. 15-16). 

B. pulverulentus seems to be first reported from Sweden sub 

nom. B. radicans PERS. by FRIES (1836-38): "In quercetis (Naset 

ad Odensjo)". Although he refers to PERSOON (1801), it is evident 

from some parts of his description that he was not describing B. 

radicans sensu PERS. (see below) but B. pulverulentus: " .... stipite

-182 - 

attenuato-radicato laevi flavo e pruina rubente flocculoso, tactu nu do 

obscurato, tubulis adnatis amplis inaequalibus citrinis ...... odor et 

sapor ingratus. Caro flava, illico obscure caerulescit." These statements 

which are in agreement with OPATOWSKI'S diagnosis (l. c.), 

must be based on FRIES' own observations, because he as it seems 

had no access to OPATOWSKI'S thesis, when he wrote his diagnosis. 

First in Summa Veg. (1849) does he adopt B. pulverulentus as 

a synonym and records the species from Vastergotland, too, where 

it was found by LINDGREN (1845). In Hym. Eur. (1874) he still 

uses the name B. radicans but states "Sapor amaricans". This statement 

must be influenced by PERSOON'S diagnosis. 

The species was, then, not mentioned in Swedish literature before 

INGELSTROM (1940), but he does not communicate any stations. At 

present there are 13 localities known from Sweden. 

In Denmark it was discovered in 1924, but thi s find was not communicated 

before 1940 by CHRISTIANSEN (Friesia 1940-41). Earlier, 

however, it had been mentioned in excursion records from 1933 and 

1934 (Friesia 1934, 1936). Now 14 Danish localities are stated. 

MALMSTROM'S (1943) statement from Finland is due to a misinterpretation 

(MALMSTROM in litt. 30.4.1955). From Norway no records. 

N o r d i c D i s t r i b u t i o n. B. pulverulentus is a species, which 

reaches its northern limit in the southernmost Nordic. 

Denmark 

F Y n. District 29 - "Røverskov", 17.8.32 (SKOVSTED l. c.). 

Langesø Skov, J. LANGE (,sKOVSTED l. c.). D. 28 - Nørre-Søby st. 

(SKOVSTED 1. c.). 

S j æ Il a n d. D. 39 - Vintersbølle Skov, 21.9.41 (Friesia 2, p. 280). 

D. 40 - Jægersborg Dyrehave, 23.9.45 (Friesia 3, p. 232). - Bregentved, 

10.9.33 (Friesia 1, p. 196). - Charlottenlund Slotspark, 9.8.36 

(CHRISTIANSEN 1. c.). - Hæsede Skov, 4.10.36 (Friesia 2, p. 126). - 

Køg.e Aas, 20.9.24 (CHRISTIANSEN 1. c.). - Næsbyholm, 16.9.34 

(Friesia 1, p. 322). - Vallø Dyrehave, 1.8.36 (CHRISTIANSEN l. c.). 

D. 45 a - Ganløse Ore, 30.8.36 (Friesia 2, p. 123). - Gelskov, 9.10.38 

(Friesia 2, p. 192). D. 45 b - Lille Dyrehave at Hillerød (Friesia 2, 

p. 124). 

Sweden 

S k å n e. Bjaresjo, Bjaresjoholm, 27.8.52. - Fågeltofta, Kronovall, 

13.8.52. - Gladsax, ornaberga, 29.8.54. - Lovestad, "Lovestads 

åsar", 24.8.54. - Rorum, Strantemolla, 30.8.54. - ,skepparslOv, Kal-

- 183- 

lunda, 26.8.53. - Smedstorp, Listarum, 14.9.54. - S. Mellby, Stens 

huvud, 30.9.49. - Tosterup, near the church, 7.9.52. - Trane, Oves 

holm, 17.9.55. - o. SonnarslOv, Maltesholm, 2.10.52. 

S m å l a n d. Odensjo, ad Naset (FRIES L c.). 

Va s t e r g o t l a n d. Rackeby (LINDGREN 1. c.). 

E x t r a - N o r d i c D i s t r i b u t ion. The species has a cir 

cumpolar distribution in the northern hemisphere with a '8outhern 

tendency. In Europe it is known from Austria, Czechoslovakia, Eng 

land, France, Germany, Italy, Luxemburg, Switzerland (KALLENBACH 

L c.), Poland (SKIRGIELLO 1939) and Russia / Caucasia / (SINGER 

1947). MURRILL (1. c.) and SINGER (L c.) report it from North 

America. 

E n v i r o n m e n t a l C o n d i t i o n s. In the Nordic B. pulveru 

len tus occurs in deciduous woods, especiaIly those of beech and 

hornbeam both in the Allietum-ass. and the Galietum-ass., 

communities, corresponding to the Allietum-union resp. Galietum 

union, described by HÅKANSSON (1949, in MS). 

A few words will be said about the constitution of these communi 

ties which compose the field-Iayer ,especiaIly in the beech woods in 

southernmost Sweden. The Allietum-ass. occurs in clayey, calcareous 

moraine, alum slate moraine and sometimes in sandy soil, where the 

inclination of the ground favours the oozing out of the subsoil 

water, while the Galietum-ass. is correlated with more or less sandy 

soil. In the latter association Galium odoratum is dominating ofte n 

with a subdominance of Oxalis acetosella. The vernal aspect is cha 

racterized by Anemone nemorosa. The Galietum-ass. is richer in 

fungi than the Allietum-ass. In spring Allium ursinum and Anemone 

nemorosa are dominating in the Allietum-ass., the Allium ursinum 

soc. of which in summer is characterized by a few species with a 

low degree of cover. 

Outside the Nordic B. pulverulentus occurs in mixed woods but 

with partiality for beech woods in Europe. AIso in North America it 

prefers deciduous trees. SINGER (1. c.) states: "In high and meso 

phytic hammocks and in gardens, on lawns, farther north and in the 

mountains usually in mixed frondose woods or under shrubs near 

Fagales (in Florida Quercus laurijolia) and possibly other species of 

oak, maybe also hickory) on grassy, mostly sandy and argillaceous 

soil."

- 184- 

2. Boletus radicans PERS. ex FR. sensu KBcH. 

Syn.: B. candieans INZENGA (ex FR.) 1869. - Boletus albidus ss 

KONRAD 1929, non ROQUES 1841. - B. albidus ssp. eupachypus KONR. 1929. 

P i l e u s thick, hemisphericaI, then convex, with Iong incurved, 

une ven margin, 7-15- 20 cm, tomentose, then smooth, somewhat 

silky shining, dingy white-grey, paIe milky coffee, in age often 

shamoy ("Isabella eolor"). 

T u b e s first paIe yellow, soon lemon yellow, being bluish at 

touch, adnexed, up to 3 cm long. 

p o r e s citrin yellowish ("Reed Yellow") with a paIe greenish 

tinge, olive brownish, on injury and at touch blue, small, roundish 

angul ar. 

S t i p e 4-10 X 3-6 cm, ovate to bulbous, in age often elongated, 

radicating, tomentose to smooth, paIe yellow ("Primrose Yellow"), 

more vivid at the apex, with a concolorous net of fine meshes, which 

are Iarger against the middle, and an olivaceous tinge below, often 

brownish to reddish, being blue at touch. 

e o n t e x t very firm and hard, in age softer, in the cap paIe 

lemon yellow, at the apex of the tubes a more vivid yellow, under the 

cuticle somewhat greyish brownish, in the stem olive yellow above, 

olive brown below, turning blue on injury, then fading. 

S p o r e s subfusoid, 10-12-14 X 4.5-5-6,u. 

e y s t i d i a conicaI fusiform, hyalin to yellow, 30-60 X 5-11,u. 

H i s t o r y. The interpretations of this species have been very 

difficult and various. As pointed out above (pag. 181), it has been 

confused with B. pulverulentus. It was first described by PERSOON 

(1801) and then adopted by FRIES in Syst. Myc. (1821). 

That by ROQUES (1841, 3 ed., 1864) described and pictured B. al bi 

dus which by some authors, KONRAD (L c.) , KONRAD & MAUBLANC 

(1952) and KUHNER & ROMAGNESI (1953), has been regarded as ide n 

tical, does not belong here. According to ROQUES (L c.) the context of 

his ,albidus has a white colour and a sweet taste, while B. radicans has 

a paIe yellow to lemon context with a bitter taste. B. albidus sensu 

KONRAD (L c.) and B. albidus ssp. eupachypus KONRAD (1. c.) are both 

identical with B. radicans. 

B. radicans was first reported from Sweden by KALLENBACH (1. c.) 

but sine loco. His statement is based on an acceptable pieture of the 

species by ROMELL, who calls it B. jlavipes ad. int. It was next found 

by G. LYTTKENS in Gotland in 1946. At the present time it has been

- 185- 

collected in 13 stations. MØLLER (1940) first communicated it from 

Denmark, where it now has been stated from 18 localities. No records 

from Finland or N orway. 

N o r d i c D i s t r i b u t i o n. The species is limited to the 

southern provinces, where it aften grows in districts with limestone 

or loose calcareous layers. 

Denmark 

N ø r r e j y Il a n d. District 21 - Aarhus (MØLLER L c.). 

S ø n d e r j y Il a n d. D. 53 - Als Nørreskov, Taksensand, 21.8.48 

(F. TERKELSEN). - Løjt Sønderskov, 31.7.53 (F. TERKELSEN). 

L o Il a n d. D. 35 - Christianssæde (MØLLER L c.). - Pederstrup 

(MØLLER L c.). - Sæbyholm Skov at Nakskov (JENSEN). - Ryde 

Skov, 30.8.25 (MØLLER). - Ludvigshave, 20.8.20 (MØLLER). - Eriksvolde 

at Skåningshave, 16.8.24 (MØLLER). - D. 36 - GI. Fredskov, 

19.8.34 (MØLLER). - Knuthenborg Park, 15.8.37 (MØLLER). - Alholm 

Park, 11.9.37 (MØLLER). 

F a l s t e r. D. 37 - Ourupgaard Park (MØLLER L c.) - Korselitzeskovene 

(MØLLER L c.). 

M ø e n. D. 38 - Klinteskoven (JENSEN); 14.9.47 (Friesia 3, p. 

446). 

S j æ Il a n d. D. 39 a - Langebæk Skov (JENSEN). - D. 40. - 

Vallø Slotspark at Køge (JENSEN). - Marielyst Skov at Vordingborg, 

5.8.21 (MØLLER). 

Sweden 

S k å n e. Lund, Botanical Garden, amongst grass, on very hard 

soil, under beech, 30.9.50. 

O s t e r g ° t l a n d. V. Tollstad, Hoje, meadow with oak, 3.9.51 

(G. HAGLUND). 

U p P l a n d. Djuro, Runmaro, Våno brygga, 26.8.51 (G. HAGLUND) ; 

Sodersunda, about 150 m E of the church, under oak, 10.9.50 (R. 

RYDBERG) . - Ljustero, Kalvholmen, Siaro, under oak and hazel, 

19.8.51 (OLLE PERSSON) . - Lovo, Drottningholm, under oak, 25,.5.51 

(G. HAGLUND) ; 30.8.51 (MAJA ENGLUND). - osteråker, Fåglaro, 

Fåglaro brygga, 6.9.52 (OLLE PERSSON). 

G o t l a n d. Buttle, N of Ange, 30.8.48 (BENGT PETTERSSON) . - 

Endre, Allekvie, on hard clayey soil, under oak, 15.9.53; Svenskens, 

17.8.46 (G. LYTTKENS) ; 26.8.46 (BENGT PETTERSSON) ; 30.8.46 (E. TH. 

FRIES). - Eskelhem, Ekebys, 28.9.47 (GOSTA and ESTER LOFVEBERG). 

Kallunge, at the railway stn, 26.8.47 (E. TH. FRIES). - Vasterhejde, 

Stenstu (E. TH. FRIES).

-186 - 

E x t r a - N o r d i c D i s t r i b u t i o n. The are a lies within the 

limits of the deciduous woods in Europe. Within this are a it is recorded 

from Austria, England, France, Germany, Italy, Pol and, 

Switzerland, Russia (Latvia) and Yugoslavia. KALLENBACH (l. c.) 

reports it from North Africa, too. AIso found by SINGER (l. c.) in 

Maine, U. S. A. 

E n v i r o n m e n t a l C o n d i t i o n s. Owing to the faet that 

the species is rare, it is very difficult to get a general idea about its 

ecology. The finds hitherto made, however, indicate that it prefers 

light and dry woods and park meadows on hard lime rich soil. 

The small number of statements from other parts of Europe point 

in the same direction. Thus KALLENBACH (l. c.) states deciduous 

woods, even wood meadows, under oak and beech. In southern Europe 

it is known from oak woods and in North Africa under Quercu8 Suber. 

3. Phylloporus rhodoxanthus (SCHW.) BRES. 

Syn.: Agaricus rhodoxanthus SCHW. 1822. - Agaricus P elletieri LEV. 

apud CROUAN et CRaUA N 1867. - Agaricus paradoxus K ALCHBR. 1873. - 

Clitocybe Pelletieri GILL. 1878. - Gomphidius rhodoxanthus SACC. 1887. - 

Paxillus Pelletieri VEL. 1920. 

P i l e u s first pulvinate, then convex, with a somewhat uneven 

recurved margin, velutinous, 4.5-8 cm, reddish brown ("Auburn", 

"Burnels Brown"). 

G i Il s broad, rather distant, anastomosing, decurrent with ribs 

on the sti pe, golden ("Lemon Chrome", "Aniline Y ellow"), on injury 

reddish 'or fulvous. 

S t i p e 4--5 X 1.2-1.5 cm, tapering and somewhat curved at 

the base, yellow, with a tinge af rufous, fibrillase-striate and 

squamulose at the apex. 

C o n t e x t paIe yellow, under the cuticle somewhat reddish. 

S p o r e s ellipsoidic, 11-14 X 3.5-5J.l. 

C y s t i d i a fusoid, 45-70 X 5.5-10 J.l, yellowish-brownish. 

T a x o n o m i c R e m a r k s. P. rhodoxanthu8 has been placed 

both in Boletaceae and Agaricaceae. !ts natural position is in Boletaceae} 

which is indicated by the velutinous pileus, the spore shape 

and the characteristic chemical reaction of the pileus surfaee by a 

deep blue colour on ammonia. The latter character has not been found 

in any gill fungus (.SINGER 1. c.).

-187 - 

It is a rather variable species. SINGER (1. c.) has erected 4 sub 

species, which he distinguishes with regard to morphological dif 

ferences and different areas of distribution in ssp. europaeus J ameri 

canus J bogoriensis and foliiporus. Of these foliiporus and bogoriensis 

have been described as species before. SINGER'S interpretation seems 

to be right as regards the differentiating between foliiporus and 

bogoriensis and these against europaeus and americanus. From 

SINGER'S descriptions it se ems to the present writer that no marked 

morphological differences actually exist between these last mentioned 

taxa. They might be brought together into 'one taxon with the rank 

of a subspecies, in the present paper called the main type. 

H i s t o ry. The species was described by SCHWEINITZ (1822). 

It is uncertain if Agaricus Tammii ss FRIES, which by some authors 

(i. e. KONRAD & MAUBLANC 1924-37) is regarded as synonymous to 

P. rhodoxanthus J 

ean belong here. Neither FRIES' description nor 

illustration of A. Tammii in leon. sel. Hym. (1877) ean motive such 

a presumption. His diagnosis lacks some important characters, i. e. 

the broad anastomosing gills and the velutinous pileus. But 

PATOUILLARD'S pieture of A. Tammii (1885) is identicaI with the 

main type of P. rhodoxanthus. 

The first Swedish locality is reeorded by lNGELSTROM (1. c.). At 

the present time it is known from 8 stations. And in Denmark it is 

found in 7 loealities. No records from Norway and Finland. 

N o r d i e D i s t r i b u t i o n. P. rhodoxanthus (the main type) 

is restricted to the southern parts of the Nordie and occurs mainly 

within the beech region. Its northernmost station is Stockholm (?). 

Denmark 

F Y n. D. 32 - "Egeskov" at Kværndrup (JENSEN). 

L o Il a n d. D. 36 - Grænge skov (MØLLER). - Fuglsang Stor 

skov (JENSEN). 

S j æ Il a n d. D. 39 a - Vintersbølle skov at Vordingborg 

(JENSEN). - D. 40 - Skjoldnæsholm skov (JENSEN). - D. 45 a 

Jægersborg Dyrehave (JENSEN. - D. 45 b - Grib skov, 20.9.53 

(Friesia 5, p. 126). 

Sweden 

S k å n e. Borringe, Borringekloster, 2.9.45. - Genarp, Hacke 

berga, 18.8.45. - Hyby, Bokebergsslatt, 2.9.45. - Lovestad, "Love 

stads åsar" , 23.9.54. - Ravlunda, Brostorp, 26.8.51. - S. Mellby, 

Stenshuvud, 25.8.53. - Tolånga, Anklam, 1950-51.

- 189- 

Kalchbrenner, C. (1873): leones selectae Hymenomycetum Hungariae I. - 

Pestini. 

Kallenbach, J. (1926-38): Die Rohrlinge in "Die Pilze Mitteleuropas". - 

Leipzig. 

Kern, Hs. (1945): Die Rohrlinge. - Olten. 

Konrad, P. (1929): Notes critiques sur quelques champignons du Jura. 

- Bull. Soc. Myc. Fr. Tome XLV. Paris. 

et Maublanc, A. (1924-37): leones selectae Fungorum. Tome l 

VI. - Paris. 

(1952): Les AgaricaIes. - Encycl. Myc. XX. Paris. 

Kiihner, R. et Romagnesi, H. (1953): Flore analytique des champignons 

superieurs (Agaries, Bolets, Chantarelles). - Paris. 

Lindgren, S. J. (1845): Notiser från Wenern. - Bot. Not. Lund. 

Malmstrom, N. (1943): B o l e t u s P u l vel' u l e n t u s Opat. och B. 

c a s t a n e u s BuH., nya for Finland. - Mern. Soc. Fauna et 

Flora Fenn. 18. 1941-42. Helsingforsiae. 

Møller, F. H. (1940) : Svampeekskursion til Petersgaard-Skovene ved Kalvehave 

den 17.9.39. - Flora og Fauna. 46. Aarg. Aarhus. 

Murrill, W. A. (1943): More New Fungi from Florida. - Lloydia. Cincinnati. 

Ohio. 

Opatowski, G. (1836): De familia fungorum Boletoideorum. - Arch. Naturgesch. 

von Fr. Aug. Wiegmann Zweit. Jahrg. Berlin. 

Patouillard, N. (1885): Tabulae analyticae Fungorum. Fase. IV. - Paris. 

Pearson, A. A. (1950): British Boleti. - The Naturalist. Leeds. 

Persoon, D. C. H. (1801): Synopsis Methodica Fungorum. Par s I. - Gottingae. 

Pilåt, A. (1953): Suchobl'ib modracka - X e r o c o m u s p u l v e r u l e nt 

u s (Opat.) Gilb. - Ceskå myk. VII. 

Ridgway, R. (1912): Color Standards and Color Nomenclature. - Washington, 

D. C. 

Roques, J. (1864): Histoire de Champignons comestibles et veneneux. 3 ed. 

- Paris. 

Saccardo, P. A. (1887-1912): SyHoge fungorum. 

Schweinitz, L. D. (1822): Synopsis fungorum Carolinae superioris. - Schr. 

Naturf. Ges. Leipzig. 

Sin ger, R. (1938): Sur les gen re s I x o c o m u s, B o l e t i n u s, P h Y Il op 

o r u s, G y r o d o n et G o m p h i d i u s. - Rev. Myc., N. S. 

Paris. 

(1945-47): The Boletineae of Florida with notes on extralimital 

species. - Reprinted from Farlowia 1945-46 and The Arner. 

Midlands Nat. 1947. 

Skirgiello, Alina. (1939): Polskie nazi emme grzyby rurkowe (B o l e t ae 

e a e et partim P o l y p o r a c e a e t e r r e s t r e s Poloniae ). 

- Planta Pol. Vol. VIII. Warszawa. 

Skovsted, A. P. (1940-41): B o l e t u s P u l vel' u l e n t u s (Opat.) En 

ny Rørhat for Danmark. - Friesia 2. København. 

Velenovsky, J. (1920): teske Houby. Dil. II. - V Praze. 

Stockholm, March 1956.

THE NITROGEN REQUIREMENTS OF 

COPRINUS HEPTEMERUS M. LANGE & SMITH 

By ERIK BILLE-HANSEN 

Laboratory of Plant Physiology, University of Copenhagen 

Coprophilous species of the genus Coprinus have now been used for 

experimental work for nearly a century. Almost all investigators have 

used either natura l media e. g. sterilised horse dung or semisynthetic 

media as KAUFFMAN'S horse dung agar or potato-glucose agar. 

LEONIAN & LILLY (1938) were the first to grow a Coprinus on a 

truely synthetic medium. They studied the growth of Coprinus lag opus 

(i. e. C. fimetarius (L.) ex FR.) and 24 other fungi, combining thi amine 

with several nitrogen sources. KEYWORTH (1942) ob tai ned fruit bodies 

of Coprinus ephemerus on a synthetic medium. LISBETH FRIES (1945) 

showed thiamine-deficiency of eight species of Coprinus and the effects 

of pH on their growth. Later she studied the requirements of Coprinus 

fimetarius under high temperature conditions (L. FRIES 1953). 

I have obtained normal and re gul ar fructification of Coprinus sassii 

M. LANGE & SMITH, C. congregatus FR., C. heptemerus M. LANGE 

& SMITH (BILLE-HANSEN 1953 a, b) and C. bisporus J. LANGE and C. 

plagioporus Romagnesi (unpublished data) - all on the same synthetic 

medium. 

It has not been possible to reproduce the positive results obtained 

with C. heptemerus (1953 b). The error is probably due to the unnoticed 

presence of small fruit body primordia on the bits of mycelia 

used for inoculation. - The four other species, viz. Coprinus bisporus, 

C. congregatus, C. plagioporus and C. sassii fru it nicely on the synthetic 

medium employed. For experimental purposes they have the draw- 

-190 -

- 192- 

EXPERIMENT 2 

Y. M. agar with various amounts of yeast extract. Ca(N03)2 replaced 

by ammonium nitrate (2 g pr. liter); daylight; 20-22° C.; 

incubation-time 18 days; 3 flasks. 

Table 2 

Yeast extract g/ lOD g I Dry weight in mg 

1,25 

2,5 

5,0 

10,0 

362 

379 

379 

426 

Fruit bodies 

normal 

normal 

normal 

sterile 

Table 2 shows the effect of various amounts of yeast extract. The 

series "1,25 %", "2,5 %" and "5 %", produced normal spore shedding 

fruit bodies. The serie "10 %" made sterile, "etiolated" fruit bodies. 

EXPERIMENT 3 

As mentioned above C. heptemerus do es not grow on the synthetic 

medium, but addition of 1 per cent yeast extract gave good growth 

and fructification. This effect was tried imitated by adding to medium 

A folIowing mixture of vitamins: Thiamine, riboflavine, pyridoxine, 

nicotinamide, Ca-pantothenate, p-aminobenzoic acid (all 50 ,ug per 

liter), and biotin (5 ,u g per liter). No growth was obtained. 

The effect of various treatments of yeast extract was studied. To 

medium A (liquid cultures) was added: 1) 0.5 % yeast extract, 2) 

same treated with charcoal (1 g/100 ml), 3) same filtered 3 times 

through a kation filter (Amberlite IR-l00), 4) same dialysed 24 hours 

in tap water. - Four flasks in each series, grown 11 days at 25° C. 

Table 3 

Treatment of 

pH 

IDry weight 

0,5 % yeast extract Initial I final In mg 

Control 7,1 7,2 55,2 

Charcoal treated 6,9 7,1 3,5 

Kation filtered 5,8 6,0 3,5 

Dialysed 7,0 7,2 18,8

-193 - 

Table 3 shows that the various treatments have decreased the effect 

of the yeast extract much. The kation filter could e. g. remove NH 1 

and some amino acids. 

EXPERIMENT 4 

8 different amina acids were tested separately. To liquid medium 

A was added 10 pmol amina acid per flask. The cultures were grown 

2 months at room temperature. 3 flasks in each series. 

Control (Ca(NOa)2) 

L-glutamic acid 

L-tryptophane 

DL-alanine 

Table 4 

Nitrogen source added 

L-histidinemonochlor hydra t 

Glycine 

DL-phenylalanine 

L-cystine 

L( - ) -tyrosine 

Dry weight 

in mg 

2,4 

0,7 

2,2 

36,2 

1,2 

5,0 

12,0 

6,7 

39,2 

Table 4 shows the results. Only DL-alanine, L ( -) -tyrosine and 

to some extent DL-phenylalanine could support growth. Asparagine 

has been tested on the medium of HACSKA YLO et al. (1954) and could 

not be utilised at all. L. FRIES (1953) tried the effect of 21 different 

amina acids on Coprinus fimetarius at 44° C, but I do not think a 

comparison is justified as the Coprinus species and the experimental 

conditions are rather different. 

EXPERIMENT 5 

In this experiment the nitrogen sources potassium nitrate, ammo 

niumtartrate and L( -)-tyrosine were compared. Basal medium A, with 

out calcium nitrate. All combinations were made in liquid media 

and on agar media. The cultures were grown 16 days at 25° C. - Four 

flasks in each experiment. 

FRIESIA V 13

-194 - 

Table 5 

Nitrogen 20 ml liquid per flask 20 ml agar medium per flask 

source mg Initial 

I 

Final I D,: w,igh, Initial 

I Final per flask pH pH 10 mg. pH pH 

I D" w,igh, 

In mg 

No nitrogen 

added 6,3 6,0 0,5 7,1 6,9 4,9 

KN03: 10 mg 6,3 5,9 0,6 7,2 6,4 5,9 

NH 4-tartrate: 

20 mg 7,0 5,2 8,1 6,9 4,7 27,1 

L( - )-tyr osine: 

9,05 mg 6,9 6,3 14,8 7,1 6,4 23,0 

Table 5 demonstrates the inability of C. heptemerus to use nitrate. 

Ammoniumtartrate and tyrosine can both be utilised. In the liquid 

series tyrosine is twice as effective as ammoniumtartrate. In the agar 

series the ammoniumtartrate is the best but the difference is not very 

large. At present I have no explanation for this peculiar difference. 

When grown on the ammoniumtartrate medium, C. heptemerus produces 

a dark orange pigment which diffuses a few millimeter out in the 

agar. On the tyrosine medium a clear yellow pigment is produced 

diffusing so rapidly that the whole agar is coloured. 

The two synthetic media, viz. medium A with ammoniumtartrate 

and A with tyrosine as nitrogen source, permit a reasonable growth 

of C. heptemerus} but they are bad fructification media. Fruit bodies 

are allways obtained, but they are sterile and rather different from 

those produced on the Y. M. medium. An optimal synthetic fructification 

medium for C. heptemerus has not yet been found. The four 

species, Coprinus bisporus} C. congregatus} C. plagioporus and C. sas 

sii fruit all on medium A as mentioned above. According to LANGE & 

SMITH (1953) they are all closely related to C. heptemerus although 

not belonging in the same subsection, but the physiological difference 

seems to be noteworthy. 

SUMMARY 

1) The coprophilous species Coprinus heptemerus LANGE & SMITH 

is well suited for fructification experiments for the following reasons: 

a) it is homothallic, b) fruit bo dies are produced on 7 days on yeast 

extract maltose agar. Agreement in parallel experiments is very good. 

2) C. heptemerus is thiamine-deficient; it cannot utilise nitrate, 

but ammoniumtartrate, alanine or tyrosine can be used.

-195 - 

3) Reasonable growth is obtained on synthetic media, containing 

alanine, tyrosine or ammoniumtartrate as nitrogen source, but only 

sterile, somewhat atypical fruit bodies are developed. 

POSTSCRIPT 

N o t e a d d e d i n p r o o f: Since this paper was sent in print. I 

received the dissertation of Dr. LISBETH FRIES: "Studies in the physiology 

of Coprinus" (1956), in which the physiology of 19 species of Ooprinus have 

been investigated from various points of view. The thesis consists of the 

folIowing papers: 1) Studies in the physiology of Ooprinus. L Growth substance, 

nitrogen and carbon requirements. - Sv. Bot. Tidskr. 49: 475 

(1955); 2) Studies in the physiology of Ooprinus. II. Influence of pH, 

metal factors and temperature. - Sv. Bot. Tidskr. 50: 47 (1956); 3) 

Studies in the physiology of Ooprinus. III. Cultivation experiments with 

running media. - Bot. Not. 109: 12 (1956); 4) Mutations induced in 

Ooprinus fimetarius (L.) by nitrogen mustard. - Nature 162: 846 (1948); 

5) Factors promoting growth of Ooprinus fimetarius under high temperature 

conditions. - Physiol. Plantar. 6: 551 (1953). 

REFERENCES 

Bille-Hansen, E.: Fructification of a coprophilous C o p r i n u s on synthetic 

medium. - Phys. Plant. 6: 523-528. 1953. 

Fructification of three coprophilous species of C o p r i n u s using 

glucose, sucrose, and maltose as carbon source. - Bot. Tidsskr. 

50: 81-85. 1953. 

Fries, L.: tiber das Wachstum einiger C o p r i n u s-Arten bei verschiedener 

Wasserstoffionenkonzentrationen. - Arkiv f. Bot. 32: 1-8. 1945. 

Factors promoting growth of C o p r i n u s f i m e t a r i u s (L.) 

under high temperature conditions.-Phys. Plant. 6: 551-563.1953. 

Hacskaylo, J., V. G. Lilly) & H. L. Barnett: Growth of fungi on three sources 

of nitrogen. - Mycologia 46: 691-701. 1954. 

Hawker, L. E.: The effect of vitamin B, on the concentration of glucose 

optimal for the fruiting of certain fungi. - Ann. Bot. 6: 631 

-636. 1942. 

Keyworth, W. G.: Investigations on the physiology of some B a s i d i o m yc 

e t e s . . - Thesis, Univ. of London (cited by Hawker, 1942). 

Lange, M.: Species concept in the genus C o p r i n u s. - Dansk. Bot. Ark. 

14, Nr. 6: 1-161. 1952. 

Lange, M. & A. H. Smith: The Coprinus ephemerus group. - 

Mycologia 45: 747-780. 1953. 

Leonian, L. H. & V. G. Lilly: Studies on the nutrition of fungi. IV. Factors 

influencing the growth of some thiamine-requiring fungi. 

Phytopathology 28: 531-548. 1938. 

Lilly, V. G. & H. L. Barnett: Physiology of fungi. - New York 1951. 

Melin, E., & G. Lindeberg : tiber den Einfluss von Aneurin und Biotin auf 

das Wachstum einiger Mykorrhizapilze. - Bot. Not. 1939: 

241-245. 1939. 

Copenhagen, February 1956. 

13*

ON THE DIMORPHISM OF THE ASCOSPORES 

AND THEIR ARRANGEMENT IN THE ASCUS 

OF MONILINIA OXYCOCCI (WOR.) HONEY (SYN. 

SCLEROTINIA OXYCOCCI WOR.) 

By N. FABRITIUS BUCHWALD 

Contribution No. 46 from the Department of Plant Pathology, 

The Royal Veterinary and Agricultural College, Copenhagen. 

It is an outstanding feature of the morphology of Monilin'i,a 

(Sclerotinia) oxycocci (WOR.) HONEY that [our of the ascospores in an 

ascus are always much smaller than the other four spores, the proportion 

between the two sorts of spores being approximately 1 :2, both 

in respect of the length and the width (cf. the spore dimensions given 

at the bottom of this page) -)(-). This difference in the sizes of the 

ascospores within the same ascus - a difference which is also found 

in the most closely related species: Monilinia baccarum (SCHROT.) 

HONEY (cf. the annexed Fig. 1» '"* ) - was, of course, observed already 

by the author of the species, M. WORONIN (1885, 1888), and 

also by later authors (REHM 1893 (1896), SCHROTER 1893 (1908), 

SHEAR 1931). WORONIN writes on Sclerotinia oxycocci (1888, p. 29) : 

"In ihrer Vertheilung und gegenseitigen Lage im Ascus ist dabei 

immer eine gewisse Symmetrie vorhanden, die in alle moglichen 

* ) Measurements of 10 big and 10 small ascospores gave the folIowing 

results: 

Big ascospores: 15.2 X 7.2p (variation: 14.3-16.5X 6.0- 8.3!l). 

Small - : 8.1X 4.3 p (variation: 6.0- 9.8 X 3.8- 4.5p). 

* *) The author knows only of one instance among other fungi, viz.: 

Phaeobulgaria (Bulgaria) inquinans, where a similar difference is 

found between the eight spores. In this species the small spores are 

also more light-coloured than the big ones, which are dark brown. 

AIready TULASNE (1853, pp. 160- 164) made this observation. Cf. 

als o MOREAU'S treatise on Bulgaria (1914). 

-196 -

Fig. 1. Representation of 

WORONIN'S figures of four 

ripe asci of Monilinia (Sclerotinia) 

baccarum) with four 

big and four small spores, 

respectively, in each ascus 

(WORONIN 1888, Table VII 

Fig. 17). - Compare ascus 

types Nos. 1, 2, 3 and 5 in 

Fig. 2. 

- 197- 

Variationen auftritt, wie man sich durch 

die Betrachtung der hier beigegebenen 

Figuren 9-13 (Taf. VII) sofort iiberzeugen 

kann. Die vier kleineren Ascosporen 

erweisen sich als unentwickelte, 

sind dabei keimungsunfahig und gehen 

sehr bald zu Grunde." A similar observation 

is made by SHEAR (1931), who - 

unlike WORONIN - furthermore points 

out that the distribution of the big and 

the small spores within the same ascus 

is irregular and, in illustration of this 

faet, depicts four asci (SHEAR, 1931, 

Fig. 11 A, p. 10) three of which correspond 

to asci Nos. 4, 5 and 9, respectively, 

in the accompanying Fig. 2. 

A more thorough investigation of 

the spore arrangement in a larger number 

of asci shows, however, that the 

distribution is not so irregular by far as 

would appear at a cursory glance. 

The author of this paper has examined 

in detail a hundred asci picked at 

random from a single apothecium and 

thereby been able to demonstrate 19 

different types of arrangements of asco 

spores, of which t'he 12 occurring most frequently are depicted in Fig. 

2. Some of those types, again, were much commoner than the others; 

for example, 18 asci of Type No. 1 and 15 asci of Type No. 5 were 

found in which the distribution may be characterized as "regular" . The 

"irregular" types, on the other hand, were generally represented by 

a single ascus only; of Type No. 8 (Fig. 2 and 6) five asci were 

found, however. This distribution of the spores certainly tends to 

indicate that the development of big and small spores is determined 

genotypically, i. e. the size of the spores is determined by a pair of 

geneticaI factors, similarly to what has been demonstrated in respect 

of sex and colour in e. g. N eurospora) Pleurage and Bombardia 

(WILCOX 1928, DODGE 1930, AMES 1932, ZICKLER 1934). 

The types of distribution concerned may be referred to the three 

folIowing principal groups :

- 198- 

1st group: Asci Nos. 1-2 in Fig. 2; 30 asci in all. 

2nd 3-6 - - 2; 46 

3rd The other types of asci, of which some are depicted 

as Nos. 7-12 in Fig. 2; 24 asci in all. 

1 2 3 4 5 6 7 8 9 10 11 12 

Fig. 2. The 12 ascus types most frequently occurring in Monilinia (3clerotinia) 

oxycocci. Nos. 1- 6 show regular location of the ascospores, resulting 

from prereduction (N os. 1 and 2) and postreduction (N os. 3-6), respectively. 

Nos. 7-12 show some of the irregular types of location most 

frequently occurring. 

Let us, for simplicity's sake, term the pair of factors for the 

spore size Aa, A being the factor for the big as cos pores and a the 

factor for the small ascospores. The distribution of the spores in 

t h e f i r s t g r o u p may then be denoted in either of these 

ways:

- 201- 

Provided the foregoing assumptions are right, segregation of Aa 

would take place now in the first meiotic division (prereduction) and 

now in the second meiotic division (postreduction). If the attraction 

(or the repulsion, respectively) between two sister chromatids (chromosome 

halves) is equal to the attraction (or the repulsion, respectively) 

between two non-sister chromatids, the disjunction of the 

four chromatids may be expected to be purely accidental. Thus each 

individual chromatid would have the chance of pairing with the 

three others, one of which is a sister chromatid, the two others nonsister 

chromatids. If these possibilities are realised equally often, 

segregation of the pair Aa will take place half as frequently in the 

first division (prereduction) as in the second division (postreduction) ; 

Fig. 5. Monilinia ox ycocci. 

Diagram of postreduction. 

Location type No. 5 with 

two small spores at top and 

bottom and four big spores 

in the middle of the ascus. 

Fig. 6. Monilinia ox ycocci. 

An example of irregular 

location in the ripe ascus, 

probably resulting from 

postreduction and displacement 

of two spindles during 

the third division. - Compare 

location type No. 8 in 

Fig. 2.

- 202- 

in other words, the ratio between prereduction and postreduction 

will be 1 :2. Although the number of asci examined is very small 

(only 100), it will still be found that the ratio between the number 

of asci that may be referred to group 1, viz. 30%, and the number of 

asci that may be referred to group 2, viz.: 46% - i. e. approximately 

1 :2 - tends to indicate that the distribution of the spores in the former 

group is due to prereduction, whereas the distribution in the 

. latter group is due to postreduction. By his far more comprehensive 

examinations of Bombardia Zunata} ZICKLER (1934) arrived at a 

similar conclusion in respect of the segregation of the pair 'Df factors 

(viridis - rubiginosa) for the colour of the ascospores of this ascomycete. 

A study of t h e t h i r d g r o u p of ascus types, which represents 

24% of the material examined, shows, however, that the segregation 

of Aa is not always as regular as described above. The irregular 

location of the spores in these cases tends to indicate that a 

translocation of the nuclei must have taken place, the spindles 

having shot past each other during one of the three divisions resulting 

in the formation of spores, and this has happened whether 

the segregation has taken place in the form of prereduction or postreduction. 

An example of the mechanism leading to an irregular 

location of the ascospores is illustrated in Fig. 6. 

As far as is known, M oniZinia oxycocci has not been the subject 

of either cytological or geneticai examinations before, but it occurs 

to the author of this paper that this species of fungus, as well as 

M. baccarum} which is closely related to it, presents itself as an 

extremely favourable object of such examinations on account of the 

pronounced difference in the sizes of the a.scospores. 

ACKNOWLEDGMENT 

I am very grateful to Mr. J. H. WANSCHER, Amanuensis, M. Sc., for 

his excellent assistance in the preparation of thi s paper. 

LITERATURE 

Arnes, L. M.: An hermaphroditic self-sterile, but cross-fertile condition in 

p l e u r a g e a n s e r i n a. - BuH. Torrey Club 59: 341-345. 

1932.

- 203- 

Dodge, B. O.: Breeding albinistic strains of the M o n i l i a bread mold. - 

Mycologia 22: 9-38. 1930. 

Moreau, F.: Sur le dimorphisme des ascospores de B u.1 g a r i a i n q u in 

a n s (Pers.) Fr. - Bull. Soc. Mycol. France 30: 361-367. 

1914. 

Rehm, H.: Ascomyceten: H y s t e r i a c e e n und D i s c o m y c e t e n. In 

Rabenhorst's Kryptogamen-Flora. I. Die Pilze. 1893 (1896). 

Schroter, J.: Die Pilze Schlesiens. II. 1893 (1908). 

Shear, C. L., Stevens, N. E. & Bain, H. F.: Fungous Diseases of the cultivated 

Cranberry. - U. S. Dept. Agric., Techn. Bull. 258: 1-56. 

1931. 

Tulasne, L. R.: Nouvelles recherches sur l'appareil reproducteur des 

champignons. - Ann. Science nat., 3. Ser., 20: 129-182. 1853. 

Woronin, M.: Ueber P e z i z a b a c c a r u m. - Ber. Deutsch. Botan. Ges. 

3: LIX-LXII. 1885. 

: fiber die Sclerotienkrankheit der Vaccinieen-Beeren. - Mern. 

l'Acad. Imper. Science St.-Petersbourg, VII Ser; Tome XXXVI, 

No. 6. 1888. 

Wilcox, M. S.: The sexuality and arrangement of the spores in the ascus 

of N e u r o s p o r a s i t o p h i l a. - Mycologia 20: 3-17. 1928. 

Zickler, H.: Genetische Untersuchungen an einem heterothallischen Askomyzeten 

(B o m b a r d i a l u n a t a nov. spec.). - Planta 22: 

573-613. 1934. 

Copenhagen, J anuary 1956.

ORNAMENTATION OF 

SPORES OF RUSSULA LAUROCERACI MELZER 

AND RUSSULA FOETENS (PERS.) FR. 

By KJELD BULOW 

Russula lauroceraci MELZER was distinguished from Russula 

foetens (PERS.) FR. and set up as a particular species by MELZER 

(1927). According to SCHAFFER (1952) it must be regarded as a 

distinct sub-species, only distinguished from Russula foetens by two 

characteristics, viz. by having a smell like bitter almonds and by the 

spores being cristate with long crests. SCHAFFER is of the opinion that 

there are no intermediaries between the two types of spores. 

RICKEN (1915) does not mention that R. foetens may have a 

smell like bitter almonds. J. E. LANGE (1940) mentions that the 

smell of R. foetens is sometimes suggestive of cherry laurel (Prunus 

laurocerasus) and that "this form is sometimes segregated 

sub nom. RussuZa lauroceraci MELZ.", but he does not mention 

the difference in the spores. He states that the spores of R. foetens 

are coarsely warty-spinulose, spores from later finds having Y2- % p 

long spinules, generally without connecting lines. Therefore, he cannot 

have seen the typical Zauroceraci spores, which are markedly 

lamella te-crista te. 

KUHNER & ROMAGNESI (1953) distinguish between three species, 

viz.: R. foetens with warty-spinulose spores, adisgusting smell and 

a peppery taste; R. grata BRITZELMAYR with warty - partially reticulate 

- spores, a faint smell and a mild taste; and R. Zauroceraci 

MELZER smelling like bitter almonds and ha ving a peppery taste and 

cristate spores with long crests. PEARSON (1950) and MOSER (1953) 

distinguish between R. foetens and R. lauroceraci by spore ornamentation 

and smell. 

- 204-

- 205- 

In the autumn of 1952 I gathered in Riis Forest in Jutland some 

specimens of R. foetens which, apparently, were typical, but the 

smell of the fungi was quite pleasant, like that of bitter almonds. 

Microscopical examination by MELZER'S chloral-iodine mixture disc10sed 

the typically cristate lauroceraci spores. For comparison, I 

examined spores of some specimens of R. foetens. and was sometimes 

surprised at finding spores of types intermediate between lauroceraci 

spores and foetens spores. 

As may be known, the smell of R. foetens may vary considerably 

both in intensity and character. FERDINANDSEN & WINGE (1943) 

describe the smell as sweet and nauseous, and it is just this type of 

smell that is found in different mixtures. 

NO.of I 

specimen 

I 

II 

III 

IV 

----- 

V 

Smell 

Almost just like 

that of bitter almonds 

Faint, not characteristic 

Commonly mingled 

Strong, mingled 

Very disgusting 

TABLE 

Spore ornamentation 

Markedly warty-spinulose, almost without 

connecting lines (71h-9 fl incl. ornament) 

Like No. I (7%- 10 fl ) 

Most closely resembling lauroceraci spores, 

but with rather low crests « Ph fl ), a few 

of them warty-spinulose (8-10% fl) 

Pronounced lauroceraci spores with crests 

> 2 fl (- 4 fl ) ; (9-10 1 h - (12) fl ) 

Like No. I; some spores, however, like No. 

III, but with more conglomerate crests; 

(9 1 h - 10%- (12) fl ) 

In the autumn of 1954 I again gathered specimens of R. foetens 

and R. lauroceraci and compared spores and smells. The Table contains 

types of spores and smells of five specimens of ((foetens") gathered 

at different places in Slagelse Forest (Zealand) in the autumn 

of 1954 (Fig. 1). It is evident that widely different types of spores are 

represented, and that the pronounced smell of bitter almonds is not 

always found in specimens with cristate spores. Finally, it appears 

that the same specimen may produce widely different spores, though 

a clear-cut type is usually predominant in a given spore print.

-- 206 - 

I III IV V 

Fig. 1. Types of spores of four specimens of Russula »/oetens«, gathered 

at different places. 

The problem apparently calls for more detailed examination. 

Among other things, the taste ought to be accurately investigated 

(compare R. grata BRITZ.). 

At any rate, it is evident that the problem of species is not at all 

as simple as formerly assumed. 

BIBLIOGRAPHY 

Ferdinandsen, C. & Winge, ø.: Mykologisk Ekskursionsflora. 2nd. Ed. 1943. 

Kiihner, R. & Romagnesi, H.: Flore analytique des champignons superieurs. 

1953. 

Lange, J. E.: Flora Agaricina Danica, Vol. 5. 1940. 

Melzer, V. & Zvara, J.: Russulae bohemiae. 1927. 

Moser, M.: Die Blatter- und Bauchpilze. 1953. 

Pearson, A. A.: The Genus Russula. - The Naturalist 1948: 85-108, 

1948; 2nd Ed. 1950. 

Ricken, A.: Die Blatterpilze. I-II. 1915. 

Schaffer, J.: Russula-Monographie. Die Pilze Mitteleuropas. Vol. III. 1952. 

Slagelse, Denmark, January 1956.

TWO NEW SPECIES OF CORTICIACEAE 

FROM DEN MARK 

PENIOPHORA DANICA SP. N. AND CORTICIUM 

SALICICOLA SP. N. 

By M. P. CHRISTIANSEN 

In the following contribution two species, Peniophora danica and 

Corticium saZicicoZa) are described and illustrated and proposed as 

new species. Both have been known to me since 1948. 

1. Peniophora danica sp. n. 

Sjælland: Hareskoven Forest, in a copse of willows, on a rotten piece 

of trunk of Salix (einerea?) in a moist location, 27.9.1948 (!) and following 

years (!) (Nos. 259, 260, 261, 1296, 2264); in the same place on a 

decorticated but almost fresh piece of a trunk of Sambucus} 16.7.1950 

and 12.10.1950 (!) (1074, 1161); Sønderskov Forest, in a copse of willows, 

on Salix} 4.11.1955 (BREGNH0J LARSEN); Allerød, in a copse of willows 

close to the Bloustrød Path, on a rotten piece of wood, December 1955 

(BREGNH0J LARSEN). 

( !) = own finds. 

Fruit body resupinate, effused, adnate, at first thin and then 

lemon-greenish with a whitish to greenish-white, fringed margin, 

fully grown soft waxy-membranaceous, somewhat gelatinous, on the 

thickest portions ± densely covered by yellowish-light brownish, 

about 1 mm high and 1/4 to 1/2 mm wide warts or blunt-topped 

rounded upgrowths- X -); during drying-up, the fruit body shrinks 

heavily and assumes a much darker shade; the warts only become 

visible under a magnifying glass, and the thinnest portions of the 

fruit body can hardly be seen on the substrate. 

When dried up, the fruit body is 160-300 Il thick; the basal texture 

is very loosely built up of ± cylindrical, thin-walled hyphae, 

*) Colours from JAKOB E . LANGE'S colour plate: Margin 1 3-1 4-k3, 

inwardly k3- e3- b5-g2-h1-h2 ("Studies in the Agarics of Denmark", 

Dansk Botanisk Arkiv, Bd. 4, Nr. 12, 1926). 

-207 -

- 208- 

Fig. 1. Peniophora danica sp. n. 

a. Hyphae; b. detail of hymenium; c. basidia; d. cystidia; e. basidiospores; 

f. detail of hymenium. - X 1250. a-d. drawn from Hareskov finds; 

f. drawn from Allerød find.

- 209- 

2-5 I' wide, with clamps, even and smooth closest to the substrate 

but intermingled with somewhat irregular, alittIe thick-walled, up to 

7 I' wide hyphae, which are ± heavily coated with a yellowish, finegrained, 

gritt y mass and then forming stellate crystal lumps; suhhymenial 

hyphae 0p3Jque, hyaline, thin-walled, closely interwowen; the 

hymenium consists of conglomerate basidia and few cystidia, though 

on the top of the warts there are often many. 

Cystidia somewhat fusiform, hyaline, thin-walled, blunt to ± 

pointed, 45-60 X 5 1', protruding up to 40 I' above the basidia. 

Basidia clavate, thin-walled, 25-35 X 3.5-51', with 4 sterigmata. 

Spore powder pallid to faintly cream-coloured. 

Spores ellipsoid, with a lateral tip, 4.5-6 X 2.8-3.21'; contents 

finely granulate or with a few droplets. 

L a t i n d i a g n o s i s: Fructificatio resupinata, late effusa, adhaerens, 

juvenilis tenuis, citrino-viridis, margine albo vel albidoviridio, 

leviter fimbriato, mature laxe membranacea vel ceracea, leviter 

gelatine a, dense et obtuse tuberculata, lutea - ± brunnea (k3e3 

- b5 - g2 - hl - h2, margine 13 -15 - k3) -;'- ), siccitate valde 

contraeta et obscure colorata; sicca 150-3001' crassa; subiculum 

hyphis basidum laxe intricatis, cylindraceis, tenuiter tunicatis, 

2-5 I' diam., nodoso-septatis, levibus, interdum usque 71' diam., ± 

crassiuscule tunicatis et valde luteo-incrustatis, hyphis subhymeniorum 

dense intricatis; cystidia sat sparsa, interdum numerosa, ± fusoidea, 

obtusa velleviter acutata, 45--60 X 5 1', usque 401' basidia 

superantes ; basidia densi, clavata, 25-35 X 3.5-51', 4 sterigmata gerentia 

; basidiosporae ellipsoide ae, oblique apiculatae, granulatae vel 

guttulatae, 4.5-6 X 2.8-3.21', tunicis tenuibus, levibus, non amyloideis. 

Hab. Ad ligna Salicis et Sambuci, Dania. 

2. Corticium salicicola sp. n. 

Sjælland, Ermelunden, on the bark on the underside of anearly horizontal, 

live stem of SaZix virens. 30.10.1948 (!) (No. 246) and in the same 

place 2.10.1949(!) (644). The tree was later felled. 

Fruit body resupinate, effused, membranaceous-waxy, fairly thick, 

about 3001', even or with a few warts, smooth, whitish-cream 

coloured, uniform out to the margin. 

*) Colours from JAKOB E. LANGE'S colour plate in »Studies in the Agaries 

of Denmark«, Dansk Botanisk Arkiv, Bd. 4, Nr. 12, 1926. 

FRIESIA V 14

- 210- 

Basal hyphae somewhat obscure, opaque, hyaline, thin-walled, 

smooth or slightly and somewhat irregularly thick-walled and ± incrusted, 

1.5-3.5/-l wide, often with granulate contents, few clamps ; 

hymenial hyphae thin-walled, 1.5-2/-l wide, as somewhat twisted. 

Gloeocystidia fe w, clavate or ending in a bulb-shaped enlargement 

at top, thin-walled, nidulant or projecting somewhat over the basidia, 

50-90 X 5-6.5/-l. Head slightly broader, contents even. 

Fig. 2. Corticium salicicola sp. n. 

a. Hyphae; b. detail of hymenium; c. gloeocystidia; d. basidiospores. - 

X 1350.

- 211- 

Paraphyses present, cylindrical, undulated, often branched at the 

end, 1-2,u wide. 

Basidia clavate, narrowing downwards, thin-walled, 20-32 X 

4.5-6,u, (2)- 4 thin sterigmata, 3-5,u long . 

.spores narrowly ellipsoid-oblong, obliquely acute towards the 

base, granulate, 6-8 X 2.4-2.8,u . 

L a t i n d i a g n o s i s: Fructificatio resupinata, late effusa, c. 

300,u crassa, membranacea vel ceracea, alba vel cremea, adhaerens, 

levis velleviter tuberculata; subiculum hyphis intricatis tenuiter 

tunicatis, hyalinis, 1.5-3.5,u diarn., nodoso-septatis, levibus vel 

leviter incrassatis et incrustatis; gloeocystidia clavata, interdum capitata, 

tenuiter tunicata, e subhymenio emergentia, 50-90 X 5- 6.5,u ; 

basidia clavata, 20-32 X 4.5-6,u; 4 gracilia sterigmata gerentia; 

basidiosporae subellipsoideae vel subcylindraceae, oblique apiculatae, 

6-8 X 2.4-2.8,u, granulosae, tunicis tenuibus, levibus, non amyloideis. 

Hab. Ad cortices Salicis virentis) Dania. 

Copenhagen, J anuary 1956. 

14*

A NEW SPECIES OF THE 

FORM-GENUS LICHENOCONIUM PETR. & SYD. 

(FUNGI IMPERFECTI), L. XANTHORIAE SP. N. 

By M. SKYTTE CHRISTIANSEN 

A parasitic fungus has on several occasions been noticed by the 

author on the apothecial disc of the lichen Xanthoria poZycarpa 

(EHRH.) RIEBER. It belongs to the Fungi Imperjecti (SphaeropsidaZes 

Phaeosporae)) and as it differs from the aIready known species of 

thi s group, it is described here under the name Lichenoconium xanthoriae. 

Lichenoconium xanthoriae sp. ll. 

L a t i n d i a g n o s i s: Pycnidia sparsa vel ±. dense aggregata, 

apotheciis ± denigratis hospitis immersa, initio clausa et globulosa, 

deinde conice erumpentia cupulariterque dehiscentia, nigra colore, 

diam. 100-200 p. Paries pycnidii 6-10 p crassus, textura paraplectenchymatica, 

cellulis ± angulosis, paene isodiametricis vel ± ellipsoideis, 

2-3-stratificatis, externis autem cellulis membrana crassa 

obscuraque instructis. Conidiophora hyalina, aseptata, subcylindrica, 

6-10p longa, 2-4p crassa, apicem versus attenuata, unde conidia 

formantur. Conidia sphaerico-subovoidea, fusca, simplicia, 2.5-4 X 

3-5p, cicatrice praedita. 

Habitatio: parasitice in apotheciis lichenis Xanthoriae poZycarpae 

per Daniam et Sueciam meridionalem. 

Locus classicus: Dania, Jutlandia borealis, in parochia Råbjærg 

(in Cimbrorum promontorio), parasitice in apotheciis Xanthoriae 

poZycarpae (habitantis ramulos emortuos Cytisi scoparii) in pineto 

loco arenoso posito). Legit M. SKYTTE CHRISTIANSEN 26.6.1942. Typus 

in Herbario Musei Botanici Hauniensis. 

- 212 -

- 213- 

Fig. 1. Lichenoconium xanthoriae (type collection). 

Pycnidia on the apothecial disc of Xanthoria polycarpa (EHRH.) RIEBER 

(magnification 19X, the white line at the top corresponds to 1 mm). 

Description of the new species (see Figs. 1 and 2) : 

Pycnidia scattered or ± densely aggregated, at first globose and 

immersed in the hymenium of the apothecia of the host (reaching 

down into the upper part of the hypothecium), later conically erumpent, 

opening by rupture of the pycnidium-wall and becoming cupshaped, 

diam. 100-200 p. The sides of the erumpent pycnidia remain 

covered by a thin layer of the epithecium of the lichen. 

The wall of the pycnidium is 6-10p thick, of paraplectenchymatic 

structure, composed of 2-3 layers of irregularly angular, almost 

isodiametric or oblong cells, 3-6 X 3-8p. In young pycnidia only the 

cells of the external part of the pycnidium-wall have thick, darkcoloured 

membranes, later all cells of the pycnidium-wall get thick, 

brown membranes. The inside of the pycnidium-wall is covered by 

closely standing, 1-celled, subcylindric conidiophores, which get narrower 

towards their tips, 6-10p long, 2-4p thick. The membrane 

of the conidiophore is thin, and in older pycnidia the conidiophores 

have almost disintegrated. 

The conidia are formed in succession at the tips of the conidiophores 

(sometimes 2 or 3 conidia remain attache d toeach other);

- 214- 

100 p. 

Fig. 2. Lichenoconium xanthoriae (type collection). 

a and b: sections through pycnidia in the hymenium of Xanthoria 

polycarpa (EHRH.) RIEBER; the lichen tissue (epithecium, paraphyses 

and hypothecium) is indicated schematically. - c: cells of the 

pycnidium-wall (from a tangential section of a pycnidium). - d: 

conidia. - e: part of a section through a pycnidium showing wallcells 

and conidiophores with young conidia. 

they are thick-walled and brown, sphaerical to broadly ovoid, with 

a distinct hilum, 2.5-4 X 3-5,u. The conidia to some degree agglutinate 

and form a mass which completely fills the cavity of the still 

closed pycnidia. The membrane of the conidia may sometimes have a 

somewhat scabrous appearance, pos si bly due to the substance which 

keeps the conidia together. The spores exude as a mass from the op en 

pycnidia when these are moistened. 

The mycelium of Lichenoconium xanthoriae is difficult to trace 

outside the pycnidia in sections of the host tissue, only a few sinuous, 

light-brown to almost hyaline, 1.5,u wide hyphae are seen. 

The parasitized apothecia of the lichen have a somewhat de generated 

hymenium. Only a few asci are still containing spores, most of the 

asci being empty and conglutinated with the paraphyses. No young 

asci are seen, nor any deeply staining ascogenous hyphae. The margin 

and the subhymenial tissue of the apothecia do still contain living 

algae. The colour of the apothecial disc may still be yellow as in 

unaffected apothecia (this coloration is due to a granular substance, 

parietin, excreted by the tips of the paraphyses and forming an

- 216- 

After having recognized Lichenoconium xanthoriae as a distinct 

species, I sent herbarium--specimens to Dr. ROLF SANTESSON. The 

species was then found by him in Sweden, and its occurrence here 

published without indication of locality (SANTESSON 1949, 142). Dr. 

SANTESSON has kind ly permitted me to publish these localities here, 

for which obligingness I am greatly indebted to him. 

Discussion of the systematic position of the new species: 

The form-genus Lichenoconium was established by PETRAK & 

SYDOW (1927, p. 432) to include species formerly referred to the 

form-genus Coniothyrium CDA., emend. SACC. (1880, p. 7), but having 

to be separated from this genus on account of the folIowing characters: 

1°. Pycnidia open irregularly, at last becoming erumpent and ± 

cup-shaped, while in Coniothyriu'Yfl' the pycnidia are opening with 

aminute ostiolum or a simple porus, and are as a rule immersed, 

except at the top. 

2°. The conidia are formed on distinct conidiophores, not on protruding 

parts of the cells of the pycnidium-wall, as in Coniothyrium. 

Owing to the insufficient observations concerning many species of 

Coniothyrium hitherto described it is difficult to ascertain, whether 

the above generic distinctions are really clear-cut and constant. The 

authors of the genus Lichenoconium have, however, formed their 

opinion after a careful study of the type specimens of a long series of 

species of the form-genus Coniothyrium. Until further studies have 

been made, I think it best to keep the two form-genera apart. No 

ascus-producing fungus connected with species of the form-genus 

Lichenoconium is known. 

Most species of Lichenoconium are parasitic on lichens. In the 

monograph on lichen-parasites by KErssLER (1930, p.558) 5 lichenicolous 

species of thi s form-genus are listed from North and Central 

Europe. 

The new species seems to have some resemblance in size and form 

of the pycnidia to L. lichenicolum (KARST.) PETR. & SYDOW, according 

to the descriptions of the type specimen made by KARSTEN (1887, p. 

104) and by PETRAK & SYDOW (1927, p. 432). I have not been able to 

study the original material, but a comparison of the descriptions of 

form and size of the spores and the conidiophores clearly shows that 

the two species cannot be identical:

- 217- 

Lichenoconium lichenicolum: spores oblong-ovoid or club-shaped, 

5-8 (-10) X 2.5-4.5 /-l. The length of the conidiophores does scarcely 

exceed that of the spores. 

L. xanthoriae: spores subsphaerical or broadly ovoid, 3-5 X 2.5- 

4 /-l. The length of the conidiophores 2-3 times that of the spores. 

A search of the available literature, especiaIly VOUAUX (1914, p. 

291), for lichenicolous species referred to the form-genus Coniothyrium) 

which might be identical with Lichenoconium xanthoriae) was 

without result. 

ACKNOWLEDGEMENT 

The author wishes to thank Mr. K. E. JENSEN, M. A ., for help with the 

Latin diagnosis of the new species. 

REFERENCES 

Karsten, P. A., 1887: Symbolae ad Mycologiam Fennicam. Pars XXI. - 

Meddel. Societas pro Fauna et Flora fennica 14: 103-110. 

Keissler, K. V., 1930: Die Flechtenparasiten. Rabenhorst's Kryptogamen 

Flora. 2. Aufl. Bd. VIII, 712 pp. 

Petrak, F. u. SydowJ H., 1926-1927: Die Gattungen der P y r e n o m y z et 

e n, S p h a e r o p s i d e e n und M e l a n c o n i e e n. 1. Teil: 

Die phaeosporen S p h a e r o p s i d e e n und die Gattung M ae 

r o p h o m a. - Fedde, Repert. spec. novar., Beih. Bd. XLII, 

551 pp. 

Saccardo, P. A., 1880: Conspectus generum fungorum Italiae inferiorum, 

nempe ad S p h a e r o p s i d e a s, M e l a n c o n i e a s et H yp 

h o m y c e t e s pertinentium, systemate sporologico dispositorum. 

- Michelia 2: 1-38. 

Santesson, R., 1949: Svampar som leva på lavar. - Sv. Bot. Tidskr. 43: 

141-143. 

Vouaux, l'abbe L., 1912-1914: Synopsis des Champignons parasites de 

Lichens. - BuH. Soc. Myc. France 28: 177-256; 29: 33-128, 

399--494; 30: 135--198, 281--329. 

Copenhagen, February 1956.

A NEW EUROPEAN CLAV ARlA: 

CLA VULINOPSIS SEPTENTRIONALIS SP. NOV. 

By E. J. H. CORNER 

Botany School, Cambridge. 

In December 1954, Mr. L. HJORTSBERG, Umeå, Sweden, sent me a 

painting and dried specimens of a clavaria collected by him from 

four localities in Sweden. It did not agree with any known European 

species, and I was much puzzled until I saw that the basidia had four 

to six sterigmata, as in the Malayan Clavulinopsis has tula. Its 

alliance lay clearly with the small phycophilous species which I put in 

section B of Clavulinopsis (CORNER, 1950, p. 346). Indeed, had the 

fungus been sent to me from the tropics, whence I have several more 

to describe, I should have been in no doubt, but the mind becomes 

"pigeon-holed" in its European dealings and has yet to learn that the 

European fungus-flora, though traditionally the fount of mycology, 

is in faet but a small part of the world-flora. 

Clavulinopsis septentrionalis is very close to the Malayan C. 

hastula) which has softly gelatinous, flattened and pinkish fruitbodies 

with unevenly inflated hyphae. The north temperate C. 

vernalis (SCHW.) has 2-4-spored basidia, longer spores, and smaller 

fruit-bodies. C. calocera (MARTIN), from Colombia, is 4-spored with 

larger spores and has rich yellow, very toughly gelatinous fruitbodies 

with stout hyphae, 20 fÆ wide. By analogy with these, I expect 

that the hyphae of C. septentrionalis may grow in intimate contact 

with soil-algae. 

Clavulinopsis septentrionalis sp. nov. 

R e c e p t a c u l a ad 20 mm. alta, ela vata, superne ad 2-4 mm. 

lata, simplicia vel plus minus subdigitata, compressa, levia vel subrugosa, 

gregaria, ochraceo-flavidula vel subaurantiaca, translucentia, 

-218 -

- 219- 

Fig. 1. Clavulinopsis seprentrionalis sp. nov. Figures (watercolour) executed 

from nature by Mr. L. HJORTSBERG. Nat. size. 

apicibus albidis: stipite 1-1.5 mm. lata, basi subcaesia: carne tenera, 

pallida, inodora, insipida: ad terram arenosam vel argillaceam et 

propter vias in pinetis, Sue cia. 

S p o r a e 5-7 X 2.5-3,u, hyalinae, leves, ovoideo-ellipticae vel 

subcylindricae, breviter apiculatae, ut videtur aguttatae. 

B a s i d i a 25-35 X 5,u , cla vata vel pyriformia, sterigmatibus 

4-5-6, 4,u longis, incurvatis: hymenium ad 100,u incrassatum: 

cystidia nulla: hyphae subhymeniales 2-3,u latae, tenuiter tunicatae, 

su bgela tinosae. 

li 

U

- 220- 

H Y P h a e 3-12,u latae, fibulatae, inflatae, tenuiter tunieatae, 

in stipite et propter subhymenium plus minus tenaeiter gelatinosae, 

eellulis regulariter inflatis, etiam fibulis inflatis. 

Fig. 2. Clavulinopsis septentrionalis sp. nov. Spores, basidia and hyphae. 

X IOOO. 

c o Il e e t i o n s: Stoeksjo, parish of Umeå, 24 Aug. 1949, on 

sandy earth in forest clearing among low moss and Calluna in pinewood. 

- StOeksjo, parish of Umeå, 9 km south of Umeå, 18 Aug. 

1950, Aug. 1951, 16 Aug. 1953, 20 Aug. 1954, 2 Sept. 1954, in gravelpit 

in pine-wood. - Håkmark, parish of Umeå, 14 Aug. 1950, 8 Sept. 

1953, by forest road in sandy pine-wood. - StOeke, parish 'of Umeå, 

Aug. 1951, 17 Sept. 1953, 6 Oet. 1953, by forest-road on bare soil, 

pine-wood. Leg. L. HJORTSBERG. 

T y P u s: Stoeke, parish of Umeå, 17 Sept. 1953. Institutionen for 

Systematisk Botanik, Uppsala, Sweden. 

REFERENCE 

Corner, E. J. H. (1950): A monograph of Clavaria and allied genera. - 

Ann. Bot. Mern. n. 1. 

Cambridge, February 1956.

COPRINUS COMATUS (SeHUM.) FR. 

SPRENGER ASFALT I BERGEN, NORGE 

Av ANDERS DANIELSEN 

I slutten av september 1955 kom det inn en pakke til Universitetets 

botaniske museum i Bergen. Innholdet, som ble fotografert av "Bergens 

Tidende" like etter ankomsten, er avbildet på tavle her (s. 222). 

Ved et besøk på voksestedet fikk jeg noen nærmere detaljer om 

forekomsten. Soppene vokste opp inne i et mørkt skur på Veivesenets 

hovedlager på Grønneviksøren i Bergen. Gulvet i skuret ble asfaltert 

på tomtens eldre fyllmasser for en del år tilbake. 

At noe var i gjære la man først merke til ved at asfaltdekket 

begynte å heve seg på forskjellige små og vel avgrensete partier - 

omtrent som en slags bobler under overfJaten. Lagerformannens 

nysgjerrighet var vakt, og han lot forhøyningene utvikle seg uten 

inngrep. I løpet av ca. 1 døgn hadde den avbildete gruppe vokset frem, 

og den ble raskt sendt til museet "forat botanikkstudentene skulle få 

se den merkelige og kraftige soppen". Det kan ellers nevnes at en 

annen sprengning ble forhindret ved harde slag med en slegge på 

asfaltbulken. 

Det var til en viss grad overraskende at fruktlegemene vokste opp 

inne i et skur, da blekksoppene trives best på lysåpne steder. Men 

dette skulle vel gi et visst håp om at de ikke kommer igjen inne i 

skuret. 

En slik ruptur i asfalt gjør et sterkt inntrykk første gang man 

ser den - ganske spesielt fordi den var laget av en bløt og lett forgjengelig 

Caprinus. Man tenker uvilkårlig på den kraft som må til 

for å sprenge asfalt med hakke eller trykklufthammer. Det skal imidlertid 

innrømmes at sammenligningen halter. Hva gatearbeideren 

klarer på få sekunder, har soppen brukt diverse timer - og tilsvarende 

mindre kraft - på. Jeg har ikke funnet noe tall for hvor 

- 221-

- 222- 

mange atmosfærers trykk voksende sopper kan utøve. I "Vaxternas 

Liv", bd. 1, p. 241, angir ROMELL opp til 12 atm. rent generelt på 

overflaten av voksende organer. 

Nå er tilfellet i Bergen på ingen måte enestående. Ifølge opplysninger 

fra amanuensis ECKBLAD bryter sjampinjonger årvisst gjennom 

asfalten på bestemte steder i Oslo. 

Fra "Vaxternas Liv" (ovennevte bd., p. 239) anføres: Voksende 

sopper har i sin tid sprengt asfalten et sted i Stockholm og en steinlegning 

i Koping. I et annet tilfelle skal en steinblokk på 160 kg være 

blitt løftet av voksende sopper. Følgende morsomme notis sto i den 

engelske avisen "Cheltenham Examiner" i 1841: "Forleden torsdag 

utførte tre eller fire store sopper en ny slags heltedåd da de på en av 

våre folkerikeste gater, i sin ivrige streben etter å bryte frem i den 

synlige verden, formelig løftet bort en stor gatestein." Sitatet er blitt 

klassisk fordi SCHOPENHAUER bruker det som bevis på den såkalte 

viljekraften eller livsviljen i naturen. 

Professor BUCHWALD har gjort meg oppmerksom på et godt bilde 

av en Coprinus (utvilsomt C. atramentarius)} "der den Steinbelag 

eines Kellers in die Hohe hebt" (Zeitschr. f. Pilzkunde, Bd. 18 (N. F.), 

Tafel 3, 1939). 

SUMMARY 

Coprinus comatus breaks through asphalt in Bergen, N orway. 

Fruiting bodies broke through an asphalt ground cover of c. 5 cm 

thickness in a dark store-house. EspeciaIly Psalliota spp. often break 

through asphalt, and are regularly found in Oslo. Breaking by Coprinus 

spp. might be considerably less frequent. 

LITTERATUR 

RomelI, Lars-Gunnar: De grona vaxternas kraftkalla. - Vaxternas Liv I: 

238-270, 1932. 

Bergen, januar 1956.

SOME OPERCULATE DISCOMYCETES NEW 

TO NORWAY 

By FINN-EGIL ECKBLAD 

The knowledge of N orwegian operculate Discomycetes is extremely 

scanty. The historicaI basis was formed by GUNNERUS (1766, 1772) 

and SOMMERFELT (1826). None of GUNNERuS' specimens has been 

left, but many of the species recorded by SOMMERFELT are still to be 

seen in the Herbarium of the Botanical Museum of Oslo. Some of his 

collections, however, contain but few specimens or only one, sometimes 

badly preserved, thus rendering a verification of his determinations 

in certain cases rather difficult. Later on A. BLYTT (1892) 

recorded Discomycetes from Dovre, Central Norway, and ROSTRUP 

(1904) determined a large number of gatherings made by BLYTT and 

his coUaborators. Since then the interest in these fungi died away, 

though records of species new to our flora have been published from 

time to time, viz. Caloscypha fulgens (ST0RMER 1946), Bulgaria globosa 

(NANNFELDT 1938), Anthopeziza protracta (GELIN 1938), Pithya 

cupressina (NANNFELDT 1949), and Helvella acetabulum (ECKBLAD 

1953). In 1949 NANNFELDT published a study of the Swedish species 

of Sarcoscyphaceae} including several records of Norwegian representatives. 

Although the exact number of operculate Discomycetes known to 

Norway cannot be given, it seems obvious that still several species 

rem ai n to be discovered. In this paper I have the pleasure to report 

10 species as new to Norway. 

My thanks are due to Professor J. A. NANNFELDT who has given 

me valuable hel p with several identifications and references to 

- 223-

- 224- 

literature. Thanks are also due to the many collectors who placed 

their collections at my disposal. 

The following abbreviations indicate the herbaria in which specimens 

are deposited: 

(O) Botanical Museum, Oslo 

(U) Institute of Systematic Botany, Uppsala. 

1. Ascobolus carbonarius KARST. 

Norwegian records: 

O s lo: Bygdøy, near the statue of Carl-Johan, Aug. 10. 1952, F.-E. 

ECKBLAD (O). 

A k e r s h u s: Bærum: At the Lysaker river near Jar, Sept. 3. and 

Sept. 17. 1955, F.-E. ECKBLAD (O). 

Seemingly not reported from N orway before. Confined to burned 

places. It seems to be the only species of Ascobolus growing on that 

substrate in our country. Easily recognized by the characteristically 

sculptured spores. 

2. Disciotis venosa (PERS. ex FR.) BOUD. - Syn.: Peziza venosa 

PERS. ex FR. 


O s l o: Bygdøy, under Larix at the cafe »Rodeløkken«, April 7. 1955, 

F.-E. ECKBLAD (O). 

The spring-fungi of N orway, as of most countries, are not well 

known. Among the lesser known Discomycetes occurring at that time 

of the year, are D. venosa and Diseina perlata (FR.) FR. They have 

ofte n been taken for one another. During the last two years I have 

studied Norwegian material which, judging from habit and time of 

collecting, might be either of them. Thus I have been unable to trace 

any specimen of D. venosa J whereas D. perlata is now known from at 

least 15 places in Southern N orway. 

In early spring of 1955, however, I collected several specimens 

of a rather large (4-7 cm in diam.), deeply cupulate discomycete. 

The hymenium was dark reddish brown, almost the same colour as 

in Gyromitra esculenta (PERS. ex FR.) FR. Excipulum dingy white,

- 225 

somewhat pruinose. Most specimens proved to be immature, but at 

least three of them contained mature ascospores, 20- 22.5 X 10.5-- 

11.5 Il. The microscopical analysis showed them to belong to Disciotis 

venosa. It is readily distinguished from Diseina perlata by the spores 

which are apiculate in the latter, but ellipsoid and non-apiculate in 

the former. 

3. Geopyxis carbonaria (A. & S. ex. FR.) SACC. - Syn.: Peziza 

carbonaria A. & S. ex. FR. 

N orwegian records: 

A k e r s h u s: Bærum: Eiksmarka Aug. 1953, F.-E. ECKBLAD (O); 

Near Ramsåsen, May 28. 1954, F .-E. ECKBLAD (O); At the Lysaker river 

near Jar, Sept. 3. 1955, F.-E. ECKBLAD (O). - Asker: Slottsberget, May 

28. 1955, S. G. SUNDBYE (O). 

Apparently this species has not been reported from Norway before. 

n grows exclusively in burned places, where it is one of the more 

conspicuous species. Easily recognized by its deeply cupulate, short· 

stipitate, dull yellowish apothecia which have crenated margins. It 

is probably rather common. 

4. Helvellella sphaerospora (PECK) IMA!. - Syn.: Helvella 

sphaerospora PECK, Gyromitra sphaerospora (PECK) SACC. 

Specimens preserved: 

A k e r s h u s: Bærum: Near the Sinober bridge in Sørkedalen, June 

1948, S. G. SUNDBYE (O), (U); July 10. 1949, S. G. SUNDBYE (O). 

In N orway this curious discomycete is known from only one 10cality. 

Mr. S. G. SUNDBYE, the collector of the species, has kindly 

informed me that he first discovered it in 1941, and has seen it fructificate 

almost every year since. It developed most abundantly in the 

dry and hot summer of 1947, the year in which the largest specimen 

occurred, measuring 28 cm high. 

n is easily distinguished from any Scandinavian species of Helvella 

by its globose spores. Macroscopically it may be recognized by the 

large size of the apothecia and the pinkish colour of the base of the 

stipe. Spores uniseriate, hyaline, 8.5-10 Il in diam., mostly with one 

large oil-drop. 

FRIESIA V 15

- 226- 

5. Helvella arctica NANNF. - Syn.: Paxina arctica (NANNF.) 

E. K. CASH. 


S ø r - T r ø n d e l a g: Oppdal: Kongsvoll, about 1 km W of the hotel, 

July 31. 1952, F.-E. ECKBLAD (O) . - T r o m s: Sørfjord: Breivikeidet, 

Aug. 29. 1954, F.-E. ECKBLAD (O). 

This small black and disciform, stipitate HelveZla was described 

by NANNFELDT (1937) on several Swedish collections. It is distinguished 

from H. corium (WEBERB.) MASS., its nearest relative, by the 

white tomentum covering the border and upper parts of the apothecium. 

Very probably it is an arctic-alpine species with more or less 

Northern circumpolar distribution. It was collected in Spitzbergen in 

1868 by TH. M. FRIES (NANNFELDT 1. c.), and was recently discovered 

in two localities in Alaska (CASH 1954). 

From Switzerland FAVRE (1955) describes a new variety, var. 

macrosperma FAVRE, distinguished by the larger spores (20-23 X 

13.5- 15.5/1). He collected it in three places in Engadine, one in Val 

Nliglia, and two at Murtarol, all specimens at rather high altitudes 

2400 to 2550 m a. s. 1.). 

6. Humarina aggregata (BERK. & BR.) SEAVER. - Syn. : Peziza 

aggregata BERK. & BR., Humaria aggregata (BERK. & BR.) SACC. 


A k e r s h u s: Skedsmo: Bråteskogen near Strømmen, May 1942, 

F. CHR. SØRLYE (U), and May 25. 1942, F. CHR. SØRLYE (O). 

H e d m a r k: Vå ler: Blakavegen, b etween Enberget and Gravberget, 

May 22. 1947, O. FURUSET (U). 

B u s k e r u d : Norderhov: Eikli SW of Hønefoss, May 31. 1954, F.-E. 

ECKBLAD (O). 

This bright orange discomycete occurs gregariously or in dense 

masses in spring on burned places or on needle beds in coniferous 

woods. The apothecia arise from a visible mycelial mat, and may 

reach a diameter of 5 mm. The spores are fusoid, 18-20 X 7-8.5/1 , 

smooth, hyaline, and mostly with two small oil-drops. Paraphyses 

septate below, strongly enlarged above, reaching a diameter of 5-6/1. 

The first Norwegian gathering of H. aggregata known to me, was 

made by the late Mr. F. CHR. SØRLYE, an interested amateur botanist, 

who collected it in Bråteskogen near Strømmen in the neighbourhood

- 227- 

of Oslo in May 1942. These specimens were sent to professor 

NANNFELDT for determination, and are kept in the Herbarium of the 

Botanical Museum, Uppsala. 

FUCK. 

7. Melastiza miniata (FUCK.) BOUD. - Syn.: Humaria miniata 


A k e r s h u s: Frogn: On a road nea r Drøbak south of the fortresses , 

Sept. 12. 1954, G. & F.-E. ECKBLAD (O). 

B u s k e r u d : Norderhov: Veienmoen, on a road, Aug. 23. 1953, F.-E. 

ECKBLAD (O) . 

This small and rather inconspicuous discomycete which so far 

has been found only twice in Norway, may at first sight be confounded 

with AZeuria aurantia (PERS.) FUCK. It differs from the latter i. a . 

in the smaller size, the colour of the hymenium which is slightly 

duller, and in having the excipulum furnished with hairs. At t he 

margin the hairs are bound together in fascicles. 

Asci cylindrical, 8-spored. Spores hyaline, ellipsoid, 14-17.5 X 

7- 9.5,u (without sculpturing), usually with two small oil-drops. The 

sculpturing soon forms a distinct reticulum, and mostly gives rise to 

a rather large apiculus at either end. Parafyses clavate above, granul 

ated. Our specimens agree well with the description by GRELET (1942, 

p. 22) who gave the spore-size as 16-20 X 9- 11/l. Those measurements, 

however, certainly included the spore-sculpturing. 

SEAVER (1928) united M. miniata with M. chateri (W. G. SM.) 

BOUD. (incorrectly speIling the specific epithet ))Charteri(( )' specific 

name after J. J. CHATER, the first collector of that species, see W. G. 

SMITH in Gard. Chron. 1872, p. 9). As clearly demonstrated and 

beautifully illustrated by Mme. LE GAL (1947) , M. miniata shows 

distinctly reticulated spores even before maturity, whereas the spore 

sculpturing of M. chateri consists of large warts with only few, obso 

lete anastomoses. It seems well founded to treat them as separate 

species. 

LEv. 

8. Sepultaria arenicola (LEv.) MASS. - Syn.: Peziza arenicoZa 


Ves t - A g d e r: Lista: The sand dunes at Kviljo, near the Soviet 

Russian war monument, Sept. 2. 1952, F.-E. ECKBLAD (O). 

15*

- 228- 

R o g a l a n d: Ogna: The sand dunes at Ogna, Aug. 30. 1952, F.-E. 

EKCBLAD (O). - Sola: The sand dunes at Sola, Aug. 29. 1952, F.-E. 

ECKBLAD (O). 

This species will probably prove to be common on sand dunes in 

Southern Norway. It may be difficult to detect as it is usually buried 

in the sand, and only appears as a small "hole" in the ground. 

ANDERSSON (1950) regards it as a facultative sand-fungus, but mentions 

that it even may grow on or in gravel-walks. 

The apothecia are subglobose, becoming cupulate, externally 

clothed with long, sinuous, septate hairs incrusted with sand. Asci 

cylindrical, 8-spored. Spores ellipsoid, 21-25 X 11.5-13.5,u, smooth, 

hyaline, usually containing one large oil-drop. Paraphyses slender, 

slightly enlarged above. 

9. Sowerbyella radiculata (Sow. ex FR.) NANNF. - For synonyms 

see NANNFELDT 1938. 

N orwegian specimen: 

O p P l a n d: Fåberg: Nordre Jørstad, Smiuhaugen, Aug. 28. 1950, 

OLAV S. JØRSTAD (O). 

The apothecia are cupulate and stipitate with a thick, hollow stipe 

which is mostly buried in the ground. The surface is first white, later 

brownish, and furnished with wavy, septate, blunt, hyaline hairs. The 

hymenium changes in colour from yellow to brownish. Spores ellipsoid, 

12-14.5 X 6.5-7.5,u, hyaline, verrucose, mostly biguttulate. 

The verrucae are often confluent, forming an incomplete network. 

A detailed description of thi s fungus is given by NANNFELDT 

(1938), who, discussing the delimitation of some difficult groups of 

operculate Discomycetes) established the new genus Sowerbyella 

NANNF., to comprise our species and a second one, viz. S. unicolor 

(GILL.) NANNF. (= Aleuria unicolor GILL.). The latter species, known 

from a few places in Sweden, ought to be searched for in N orway too. 

It differs i. a. in the brighter colour of the hymenium and the more 

elongated spores with smaller warts. 

10. Thecotheus pelletieri (CROUAN) BOUD. - Syn.: Ascobolus 

pelletieri CROUAN. 

N orwegian specimens : 

O s l o: Bygdøy Mars 16. 1955, on cow-dung, developed in moist chambers 

in ten days, leg. F.-E. ECKBLAD (O).

- 229- 

Apothecia about 1 mm in diameter, growing on dung of various 

kind. More or less cylindrical in shape, rare ly becoming discoid, white 

or whitish. Asci clavate, 300-350 X 40-60/1, 32-spored. Spores 

ellipsoid, smooth, hyaline, 23-27 X 13-14/1, with thick wall. Parafyses 

filiform. 

The very small apothecia of this species are easily overlooked, in 

faet, they were first thought to be young fructifications of a Coprinu8 

sp., which grew on the same substrate. T. pelletieri is probably not 

rare, but easily overlooked. It is readily distinguished by the microscopicaI 

characters mentioned above. 

LITERATURE 

Andersson, O., 1950: Larger fungi on sandy grass heaths and sand dunes 

in Scandinavia. - Bot. Notiser, Suppl. 2 (2): 1-89. 

Blytt, A., 1892: Bidrag til kundskaben om Norges sopparter. II. Ascomyceter 

fra Dovre, samlede af A. BLYTT, E. ROSTRUP. - Forh. 

Vidensk.-Selsk. Christiania 1891, No. 9, pp. 1-14. 

Cash, Edith K., 1954: Some D i s c o m y c e t e s new to Alaska. - Jour. 

Wash. Ac. Sci. 44: 44--46. 

Eckblad, F.-E., 1953: H e l v e Il a a c e t a b u l u m i Norge. - Blyttia 

11: 130-132. 

Favre, Jules, 1955: Les champignons superieurs de la zone alpine du parc 

national Suisse. - Ergebn. d. wissenschaftl. Unters. d. schweiz. 

Nationalparks 5: 33: 1-212. 

Gelin, Olov E. V., 1938: The distribution in Scandinavia of P l e c t a n i a 

p r o t r a c t a (Fries) Gelin, comb. nov. and P l e c t a n i a c o cc 

i n e a (Scop.) Fuckel. - D. Kgl. Norske Vidensk.-Selsk. Skr. 

10 (52): 194-197. 

Grelet, L.-J., 1942: Les Discomycetes de France d'apres la classification de 

Boudier. 9. fasc. - Rev. Myc. 7: 3-26. 

Gunnerus, J. E., 1766: Flora Norvegica. Pars Prior. - Nidrosiæ. 

1772: Flora N orvegica. Pars Posterior. - Hafniæ. 

Le Gal, Marcelle, 1947: Recherches sur les ornementations sporaIes des 

Discomycetes opercules. - Ann. d. Sci. Nat. 11. Ser. Bot. 

8: 73-297. 

Nannfeldt, J. A., 1937: Contributions to the mycoflora of Sweden. 4. On 

some species of H e l v e Il a, together with a discussion of 

the natural affinities within H e l v e Il a c e a e and P e z iz 

a c e a e trib. A c e t a b u l e a e. - Svensk Bot. Tidskr. 31: 

47-66.

- 230- 

Nannfeldt, J. A., 1938: Discomyceten B u l g a r i a g lob o s aSchmied. 

ex Fr. funnen i Norge. - Naturen 1938: 2-4. 

1938: Contributions to the mycoflora of Sweden. 5. On P e z iz 

a C a t i n u s Holmskj. ex Fr. and P. r a d i c u l a t a Sow. ex 

Fr. with a discussion of the genera P u s t u l a r i a Fuck. emend. 

Boud. and S o w e r b y e Il aNannf. n. gen. - Svensk Bot. 

Tidskr. 32: 108-120. 

1949: Contributions to the mycoflora of Sweden. 7. A new winter 

discomycete, U r n u l a h i e m a l i s Nannf. n. sp., and a short 

account of the Swedish species of S a r c o s c y p h a c e a e. 

- Svensk Bot. Tidskr. 43: 468-484. 

Rostrup, E., 1904: Norske Ascomyceter i Christiania Universitets botaniske 

Museum. - Vidensk.-Selsk. Skrifter. L Math.-naturv. Kl. 1904, 

No. 4, 1-44. 

Seaver, F. J.) 1928: The North American Cup-fungi (O p e r c u l a t e s) . 

- New York. 

Sommerfelt, S. Chr., 1826: Supplementum Florae Lapponicae. - Christiania. 

Størmer, Per, 1946: C a los c y p h a f u l g e n s ny for Norge. - Blyttia 

4: 81-82. 

Oslo, February 1956.

VARIATIONS IN RESISTANCE OF TRIFOLIUM 

PRATENSE TO ATTACKS OF KABATIELLA 

CAULIVORA (KIRCHN.) KARAK. 

By K. J. FRANDSEN 

D. L. F. and F. D. B. Plant Breeding station, 

0toftegaard, Denmark 

Attacks of Kabatiella caulivora (KrRCHN.) KARAK. on red clover 

are occasionally reported in Denmark and complete crop failures have 

been observed in red clover seed fields. In some cases it was necessary 

to abandon growing of clover for a time (STAPEL & BOVIEN 1943). 

Serious attacks have been reported both in Germany and the Netherlands. 

In the colder areas of North America, the so-called Northern 

anthracnose caused by Kabatiella 

caulivora) is a major disease 

of red clover (HANSON & KREITLOW 

1953, KREITLOW, GRAHAM & GARBER 

1953). 

Symptoms, dark brown or black 

spots, are mainly confined to stems 

and petioles. The lesions soon 

inhibit translooation tO' the superadjacent 

parts of stem and petiole 

which causes them to wilt and die. 

The petioles bend downward as a 

shepherd's crook (Fig. 2) at the 

site of the lesion and thus forms 

the characteristic picture of the di- 

- 231 - 

Fig. 1. 

Conidia of Kabatiella caulivora. 

X 1100.

- 232- 

Fig. 2. Plants of red clover (Trifolium pratense) . A. Sound plants. B. 

Plants infected by Kabatiella caulivora. Note the characteristically bended 

stems and petioles. 

sease in the fieId. Lesions on the stem develop as dark spots which 

lengthen and turn light brown with a dark margin. The stem s ofte n 

crack at the lesions as the plant grows. 

Attacks of Kabatiella caulivora on red elover have been observed 

several times in field experiments at øtoftegaard, Zealand. In 1948 

and 1954 our experiments with red elover families were infeeted with 

stem disease in the seeond harvest year. The severity of the attaek 

was judged and expressed in figures (O = no attaek; 10 = severe 

attaek). The figures showed that families differed in resistanee to 

the disease. The severest attaek was observed in 1948. Charaeters 

(figures) from a red elover test of that year are given. 45 families 

and 4 strains, one of whieh served as standard, were sown in 

8 series, eaeh with 6 families and the standard. The plot size 

was 10 sq. meters; the number of replicates 7. The attaek was severe 

in 4 of the series. It was rather evenly spread over the area. The average 

figure for attaek in the standard plots of the 4 series was 4.1, 3.3, 

4.0 and 3.7 respeetively. 

On August 20 characters (figures) were given for degree of attack 

in all 392 plots. 

Analysis of variation of the figures from series 5-8 appears in 

Table 1 where the 4 series have been treated as one. 

8

A NEW RHODOTORULA SPECIES, 

RHODOTORULA MACERANS SP. N., ISOLATED 

FROM FIELD-RETTED FLAX STRA W 

By P. SONNE FREDERIKSEN 

Report No. 23 from The Danish Flax Research Institute, 

Viby J ., Denmark. 

The processing of fibre plants comprises a microbiologioal proces;; 

called "retting" . By exposure to moisture the stem tissues are subjected 

to incipient decomposition, which gives the straw a brittleness 

enabling the fibres to be extracted mtchanically. 

In today's Danish flax industry the retting method most widely 

used is field retting. In connection with pulling (harvesting) , the flax 

is spread out on the ground in thin layers, thus being exposed to the 

weather conditions. Through moistening by precipitation and dew a 

condition is established which makes the activities of the microorganisms 

possible. Fermentation will take place first in the pectin 

substances and the water-soluble tissues of the straw and, next, in the 

actual component of value: the fibre bundle, which consists essentiaIly 

of cellulose. When rubbing of the straw by hand shows that it has 

obtained the brittleness desired, it should be removed from the 

ground and gathered in as soon as it has become sufficiently dry. 

LITERATURE 

Current literature on field retting and its biology is very sporadic 

and deals only with some rather commercial aspects and stray 

floristic studies, which, as far as the author can see, are often founded 

- 234-

- 236- 

connection between the retting flora and the quality of the rettingl). 

These investigations - spread over 6 years - demonstrated that the 

group of "red imperfect yeasts" amounts to about 30% of the mycolo 

gical flora 2 ). Isolation and determination of 38 isolates of these red 

yeast colonies revealed that 95% of all the isolates could be referred 

to the genus Rhodotorula. 

The occurrence of Rhodotorula species on the retted flax straw 

varies with the season as shown in Fig. 1. The yeasts are most 

frequent in the moist and cold season and may in certain spelIs of 

winter reign supreme within the mycological flora jointly with spe 

cies of Cryptococcus. While the mouIds are exterminated by the 

low temperatures of winter, the yeasts will spread in consequence of 

the higher humidity of the atmosphere, the soil and the flax straw. 

These microorganisms then give rise to the winter field retting the 

eminent quality of which is well known from practicaI and experi 

mental field retting in the cold season. 

Investigations of altogether 36 isolates of Rhodotorula according 

to LoDDER & VAN RIJ ("The Yeasts" 1952) proved that 9 isolates, or 

25%, could be referred to Rho 

dotorula glutinis) while 27 isola 

tes, or 75%, might be regarded 

as a new species. This Rhodo 

torula species, whose description 

is given below, consequently 

constitutes the major proportion 

of the red imperfect yeasts on 

field-retted flax straw and 

is therefore a microorganism 

which is ve ry valuable for flax 

retting. It represents 15-20% 

of the mycoflora of field retting 

in the six winter months. It is 

proposed to name this new spe 

cies Rhodotorula macerans) af- 

cP 

······, /O/7 O 

' ...... (// 

.. CJ 

Fig. 2. Cells of Rhodotorula macerans 

sp. n. after 3 days in malt extract. 

1) A detailed account of these investigations will be published later. 

2) Samples of the reUed flax straw were cut up by shaking with glass 

balls in sterile water. The author realizes that the mouids have thereby 

become more numerous because of the cuUing-up of the mycelium. 

Considering that the reUing organisms live in the flax straw, the method 

is still regarded as necessary and correct. Spreading of the shaking 

water af ter dilution was done on malt-agar.

- 237- 

ter the retting process. Probably, this species occurs commonly on 

dead plants in nature, and not exclusively on flax 1 ). 

Rhodotorula macerans sp. nov. 

C u l t u r a i n e x t r a c t o m a l t i: Post 3 dies ad 25° C cellulae 

sunt oblongo-oViales (3.3-5.5 X 7-12,u), singulae vel binae. (Fig. 2). 

Post unum mensem ad 17° C sedimentum atque annulus formati sunt. 

C u l t u r a i n m a l t o - a g a r: Post 3 dies ad 25° C cellulae 

sunt oblongo-ovales (3-5 X 7-12,u) , singulae vel binae. Cultura in 

striis post unum mensem ad 17° C laevis et nitens est vel ex parte 

rugosa et opaca, color roseus vel ruber. 

C u l t u r a i n l a m i n a v i t r e a: Nullum pseudomycelium. 

F e r m ,e n t a t i o: N ulla. 

A s s i m i l a t i o s a c c h a r i: Glucosis + 

Galactosis + ( exigua) 

Saccharosis + 

Maltosis + 

Lactosis + (saepe exigua). 

A s s i m i l a t i o k a l i u m n i t r a t i : Adest. 

E t h a n o l e u m u t u n i c a o r i g o c a r b o n i s: Nullum 

incrementum. 

D e c o m p o s i t i o a r b u t i n i: Variabilis. 

C u l t u r a i n p e c t i n o: Incrementum observatur. 

p r o d u c t i o c o m p o s i t o r u m a m y los i m i l i u m: Adest. 

Culturae huius speciei conservantur in "Centralbureau voor Schimmelcultures", 

Delft, Hollandia, item in collectionibus culturarum fungorum, 

quas Academia regia agriculturae, pars phytopathologica, in 

Hafnta sustentat. 

Standard description of Rhodotorula macerans sp. n. 

G r o w t h o n m a l t e x t r a c t: After 3 days at 25° C, cells 

are oblong-oval (3.3-5.5 X 7-12,u), single or in pairs (Fig. 2). 

Af ter one month at 17° C a sediment and a ring are formed. 

G r o w t h o n m a l t - a g a r: Af ter 3 days at 25° C, cells are 

oblong-oval (3-5 X 7-12,u ), single or in pairs. 

1) The specific name JJlini JJ (af ter the flax plant) ought not to be used 

for this saprophytic yeast but should be reserved to the pathogene 

fungi of the flax plant.

- 238- 

The streak culture after one month at 17° C is smooth and shiny 

or partially rugose and dull; the colour is pink to red. 

S l i d e c u l t u r e s: No pseudomycelium. 

F e r m e n t a t i o n: Absent. 

S u g a r a s s i m i l a t i o n: Glucose + 

Galactose + (weak) 

Saccharose + 

Maltose + 

Lactose + (often weak). 

p o t a s s i u m - n i t r a t e a s s i m i l a t i o n: Positive. 

E t h a n o l a s s o l e s o u r c e o f c a r b o n : No growth. 

S p l i t t i n g o f a r b u t i n: Variable. 

Growth on pectin: Positive. 

P r o d u c t i o n o f s t a r c h - l i k e c o m p o u n d s: Positive. 

Cultures of the abovenamed species are kept in "Centraalbureau 

voor Schimmelcultures", Delft, Holland, and in the colleotion of fungus 

cultures in the Plant Pathological Department of The Royal 

Veterinary and Agricultural College, Copenhagen. 

DISCUSSION 

Testing with LUGOL's iodine solution in accordance with LODDER 

Fig. 3. Slide cultures of Rhodotorula 

macerans sp. n. after 

10 days. 

& VAN RIJ's description gave a distinct 

blu e colour at all the cultures, which 

shows that the cells of Rhodotorula macerans 

con tai n compound s of a starchlike 

nature. This character is one of the 

most important character s of the species. 

Rh. macerans assimilates nitrate and 

thereby resembles Rh. glutinis) which is 

found on field-retted flax straw, too, 

and, according to LODDER & VAN RIJ 

(1952), also produces starch-like compounds. 

In relation to sugar species Rh. 

macerans differs from Rh. glutinis in the 

capacity for assimilation of lactose. The 

new species assimilates sugar species in 

the same way as Rh. jlava) but is easily 

discerned from the latter by the yellow 

colour of its streak culture and by its 

reaction as to nitrate and iodine. These

- 239- 

characters are sufficient to distinguish Rh. macerans from the Rhodo 

torula species described by LODDER & VAN RIJ in "The Yeasts" (1952). 

ACKNOWLEDGMENTS 

The author offers his best thanks to DRA. W. CH. SLOOT, "Centraalbureau 

voor Schimmelcultures", Delft, Holland, for investigations of two cultures of 

the abovenamed Rhodotorula species and for the opinion that it is a new 

species. Also to Dr. K. T. WIERINGA, Laboratorium voor Microbiologie de 

Landbouwhogeschool, Wageningen, Holland, the most sincere thanks are 

due for having been so kind as to forward the cultures to the "Centraal 

bureau voor Schimmelcultures" and, besides, assisted me in my work with 

the field-retting flora. 

In the preparation of the manuscript I have had valuable assistance 

from Professor N. FABRITIUS BUCHWALD. Mr. F:. BROCKMEYER has been so 

kind to prepare the Latin diagnosis. I acknowledge with gratitude the help 

rendered me by these two gentlemen. 

BIBLIOGRAPHY 

Lodder) J. & Kreger-van Rij, N. J. W., 1952: The Yeasts. 713 pages, 268 

figures. Amsterdam. 

Ruschmann, Gerhard, 1923: Grundlagen der Roste. 188 pages, 27 figures. 

Leipzig. 

Wieringa, K. T., 1955: Der Abbau der Pektine; der erste Angriff der 

organischen Pflanzensubstanz. Zeitschrift fur Pflanzenerniih 

rung, Dungung und Bodenkunde 69: 150-155. 

Viby J., Denmark, February 1956.

PARMELIA SUBAURIFERA NYL. AND 

P. FRAUDANS (NYL.) NYL. IN GREENLAND 

By PAUL GELTING 

An auto trip on the sea ice along the S. coast of Disko Is1., W. 

Greenland, gave me on February 23,1950 the opportunity of studying 

the lichens at Engelskmandens Havn. For the vegetation at the 

stenothermous springs here, keeping a temperature of ca. + 17° C, 

see PORSILD (Medd. om Grønl. 25, 1902, 181, & ibid. 60, 1921, 313). 

On the day of my visit the air temperature was ca. --;- 25° C, and 

the main stream of the springs ran unhampered into the sea. The 

easternmost spring, which in summer irrigates a meadowy stand of 

Juneus are ti eu s rich in Habenaria hyperborea) was frozen. Conse 

quently I could lay down with my back on the frozen spring and 

study the lichens growing on the lower side of the large gneiss 

blocks forming almost a cave over the spring. 

The roof of the cave was parti ally covered by rime and ice, evident 

ly formed when the spring was still flowing. In the crust of sterile 

and leprarioid lichens I could determine: Cetraria hepatizon) Parmelia 

saxatilis) P. subaurifera) Peltigera seutata and Physeia grisea. On the 

top of the blocks outside the cave grew: Cetraria eommixta) Parme 

lia disjuneta) P. fraudans and P. suleata. 

In the folIowing some data are given about Parmelia subaurifera 

and P. fraudans) the first never, the second rare ly recorded from the 

Arctic. Microtome sections studied, 5 and 10 p thick. Field numbers 

are cited in paranthesis after the collecting places. 

The material was partly worked up at the Arctic Station, God 

havn, W. Greenland, partly at the Institute of Systematic Botany of 

the University of Uppsala (UPS). To Professor J. A. NANNFELDT 

and his staff, especiaIly Docent ROLF SANTESSON, I express my cordial 

thanks. Fil. lic. T. E. HASSELROT, curator at the Riksmuseum, Stock 

holm (S), has kindly confirmed my determinations of Parmelia frau 

dans. 

-240 -

- 241- 

1. ParmeIia subaurifera NYL. 

in Flora, 56, 1873, 22. ZAHLBRUCKNER, Catal. 6, 1930, 117. TAVARES 1945, 

78. M AAS GEESTERANUS 1947, 126. 

leon.: GALL0E 1948, PI. 35 & 36, figs. 230-247. 

Exs.: CLAUD. 121; ELENK. 153; FLAG. Fr. C. 110; Krypt. Vind. 1872; 

MALME 135; NORRL. 31; OLlV. 12; ROUM. 405; Zw. 525, 865; KRAWIEC 82; 

SUZA 145. All seen in UPS. 

W. G re e n l a n d: Godhavn, Engelskmandens Havn, cave (cf. 

p. 241) , 3-4 m. s. m. (10689,12709). 

Specimens 1- 2 cm in diam., upper surface yellowish brown. Isidial 

soredia numerous, typical; as young yellowish, in age whitening. 

Lower surface black to brown, rhizinose.-Apothecia absent. 

Thallus 90-160/1 thick. Upper cortex 6-10 11. thick, brown, formed 

of one layer of firmly coherent cells (as in GALL0E L c., fig. 242). 

Medulla up to 120/1 thick, white, forming an open tissue, C + red. 

Hyphae 4-6(9) /1, membrane 1/1 thick, at intervals covere d by white 

granules, giving the hyphae a woolly appearance (as in GALL0E, fig. 

246). Lower cortex 6-8/1 thick, brown, formed of one cell-layer (as 

in GALL0E, fig. 243). Algae globoid, 8-14/1, light green, cystococcoid, 

forming a thin, ± open layer below upper cortex. Description af ter 

12709. 

The specimens recaU N. Fennoscandian ones and may be referred to 

the main species (cf. MAAS GEESTERANUS L c.). 

H a b i t a t : The lichen crust, in which Parmelia subaurifera 

takes part, is slightly dusty. The presence of Physcia grisea (interwoven 

with P. subaurifera) indicates the dust to be rich in nutritive 

matter. The dust may originate from droppings of the snow-buntings 

visiting the cave, or from the disintegrating "tufa" layers surrounding 

the gneiss area at Engelskmandens Havn. 

The spring breaks early and freezes late. It ameliorates in several 

respects the summer climate of the cave in diminishing the temperature 

oscillations and augmenting the aerial humidity. 

The roof of the cave receives light reflected from the snow outside, 

which persists for a long time after the spring has broken 

through. It also receives light reflected from the spring itself running 

the whole vegetation period. Light reflected from the snow is of great 

importance in the Arctic and of particular influence in spots not 

reached by the sun. This may illustrate how P. subaurifera and other 

species can sustain life in a cave never receiving direct sunlight. The 

locality is far N. of the previously known distribution of the species 

F RI ESIA V 16

- 242- 

and represents so far the sole known Arctic record. Within its main 

area in more southern climes, P. subaurifera is mostly an epiphyte 

of trees. 

Geographical distrib.: Greenland (cf. p. 242). - Europe: 

From Finnmark, Swedish, Finnish and Russian Lapponia through all 

European eountries to the Mediterranean (Spain, Italy, Dalmatia, Greeee). 

Crimea: Jalta, leg. DEGELIUS (Herb.) ; Caucasus. From Ural: Molotov 

(Perm), pr. pag.Oborino, leg. A. OXNER (UPS), to Scotland and the islands 

W. hereof: lsI. of Mull, leg. P. O. LINDAHL (UPS). Ireland; Bretagne; 

Portugal. In S. E. Europe mainly montane. - N. A f r i e a: Moroeeo; 

Algeria. - A s i a: Middle Ural; Konda; Minusinsk. At the Ob: Kalinsk, 

610 N. lat., leg. M. BRENNER (S); at the Yenisej: Yenisejsk, Stolba and 

Uskij mys, 610 27' N. lat., leg. M. BRENNER (S). China: Kansu, Tsagaling, 

on Picea and Betula) 1900-2200 m. s. m., leg. D. HUMMEL (S). - N. A m er 

i e a: U. S. A.: Maine, Mt. Desert lsI., leg. G. LLANO (UPS); Conneetieut; 

New York; Tennessee; Miehigan; Wiseonsin: Lincoln eo., leg. J. W. 

THOMSON (2438, UPS); Minnesota. Canada: Nova Seotia, Cape Breton lsI.; 

Ontario: Bear lsI., leg. R. F . CAIN (21738, UPS); British Columbia: Vancouver 

lsI., leg. J. MACOUN (UPS). - I have seen no reeords from Asia 

Minor, E . Asia and western U. S. A. 

2. Parmelia fraudans (NYL.) NYL. 

in NORRLIN, Not. Sallsk. F. Fl. Fenn. Forh. 13, 1871, 324; ZAHLBRUCKNER, 

Catal. 6, 1930, 166; LYNGE 1921, 171; HASSELROT 1953, 80. - P. saxatilis 

* P. jraudans NYL., Lieh. Seand. 1861, 100 & Lich. Lapp. Orient. 1866, 119. 

leon.: Fig. 1, p . 244. 

Exs.: HAV. 557; Krypt Vind. 1970; MALME 556; NORRL. 25; RAS. 45, 46. 

W. G r e e n l a n d: Upernivik: På. Disko: Diskofjord, basalt 

block at trading post; Godhavn, Engelskmandens Havn, gneiss block 

4-5 m. s. m. (10688); Lyngmarken, gneiss block in Salix copse, 20 

m. s. m. (12719, 18573a). Egedesminde: Arfersiorfik Fjord: Serfartok 

Isl., low gneiss block in Betula nana heath, 15 m. s. m. (16538); Agpakarfik, 

N. exp. gneiss rocks with Lobaria scrobiculata} 30 m. s. m. 

(16236a-b); Ordlerfik, low gneiss blocks near bird place, 8 m. s. m. 

(16061b). All collected by GELTING. The first find of this species in 

Greenland was made by M. SKYTTE CHRISTIANSEN in 1946 at Holsteinsborg: 

Head of S. Strømfjord (kind verbal communication). 

Specimens up to 8 cm, occasionally 12 cm in diameter. Upper 

surface darker or lighter greyish, often with a yellow tinge. Soredia

- 243 - 

typical, frequently elevated on an isidioid base; as young limbiform 

(nu RIETZ 1924, 379), in age ± confluent (Fig. 1, p. 244). Lower sur 

face black to brown, rhizinose. - Apothecia absent. 

Thallus 150-260 p thick; upper surface covered by a 4-5 p thick, 

uncoloured, necral stratum. Upper cortex 20-36p thick, paraplect- 

Fig. 1. ParmeZia fraudans (NYL.) NYL. Specimen from Agpakarfik 

(16236 a). SVEN ERIKSSON (Vaxtbiol. Inst.) phot. X 2. 

enchymatous, uncoloured or greyish; at middle enlargements seem 

ingly divided in an upper darker and a lower lighter stratum. Cortical 

cells 4-6 in number, membrane not particularly thick. Medulla 40- 

150 p thick, whitish, forming an open tissue, C-, K + yellow-red; 

hyphae 2-3(4) p thick, leptodermatic; membrane 0.5-0.8p thick. 

Lower cortex 8-30 p thick, paraplectenchymatous, brown; cells 

3-4(5) in number, mesodermatic. In the apices of the lobes a brown 

ish stratum with ± coherent cells is found between lower cortex and 

medulla. It is found also in the interior of the rhizinae. Algae globoid, 

6-15 p in diam., cystococcoid, forming a layer, 30-90 p thick, below 

upper cortex. Description af ter 18573a. 

16*

- 244- 

NYLANDER (Flora 52, 1869, 292) doubted the value of his subspe 

cies, but raised it to a specific rank 1871. P.armelia fraudans devi 

ates from P. saxatilis in bearing soredia, from P. sulcata in the form 

and arrangement of the soredia. The Greenland specimens belong to 

the main species. 

p a r a s i t e s: P. fraudans is often overgrown by dark hyphae. In 

18573a from Lyngmarken the hyphae bear spinulose apothecia be 

longing to Echinothecium reticulatum ZOPF. 

H a b i t a t: In the interior of Upernivik and Egedesminde di 

stricts P. fraudans is mainly found in the vicinity of bird places 

(eiderducks, ptarmigans, snow-buntings), however, not on the more 

intense ly dunged rocks. At Ordlerfik the larger stones visited by 

ptarmigans were inhabited by: Physcia dubia) Umbilicaria decussata) 

Xanthoria candeZaria) Lecanora rubina) Rinodina cacuminum) R. 

oreina a. o., while P. fraudans was not found. In the vicinity, however, 

it occurred abundantly on low stones together with ± mesotrophic 

species as: Alectoria pubescens) Cetraria hepatizon) P. saxatilis a. o .. 

On a block slightly influenced by snow-buntings it occurred at 

Lyngmarken together with: P.armelia disjuneta) P. saxatilis) P. inte 

stiniformis) Sphaerophorus globosus) Umbilicaria hyperborea) Crocynia 

negleeta) Lecanora polytropa and Rhizocarpon geminatum)' in the 

main a mesotrophic to slightly eutrophic community. In Fennoscan 

dia P. fraudans is (HASSELROT, 1953, 82) in some degree favoured by 

human agency and becomes especiaIly towards S. an apophyte. 

As in Fennoscandia, Parmelia fraudans is also in Greenland most 

frequent in the inland. Godhavn and Diskofjord are the localities, 

where the species occurs nearest the open sea. 

G e o g r a p h i c a l d i s t r i b.: Greenland, W. coast 660-730 N. lat. - 

E u r o p e: Mainly Fennoscandian. From Finnmark, Swedish, Finnish and 

Russian Lapponia S. to Oslo, Småland, Gulf of Finland, Karelian Isthmus, 

Leningrad, Ladoga- and Olonetz Karelia. Recorded from Novaya Zemlya, 

Tallin and Tartu. Caucasus. - N. A m e r i c a: Miquelon. Canada: Keewa 

tin, Repulse Bay. Alaska: N. coast, Camden Bay. 

In addition an unlocalized record (ECKFELDT 1895, 243) from Labra 

dor or from New Foundland. 

RESUME 

Parmelia subaurijera rapporteres for første gang fra Grønland, 

hvor den er fundet i en grotte ved de "varme" kilder i Engelskman 

dens Havn ved Godhavn. Parmelia jraudans angives fra adskillige

- 245 

steder paa vestkysten indtil 73° n. br. Hos begge arter beskrives 

thallus-bygningen (apothecier fandtes ikke), ligesom der gives op 

lysninger om voksestedsbetingelser og geografisk udbredelse. Som 

parasit paa P. fraudans optræder i Godhavn Echinothecium reticula 

tum) der ikke tidligere er angivet fra Grønland. 

LITERATURE 

Ahlner, S.: Fleehten aus Nordfinnland. - Ann. Bot. Soe. Zool. Bot. Fenn. 

Vanamo, 9, 1. Helsinki 1937. 

Arnold, F.: Liehenologisehe Fragmente, 29, Miquelon. - Flora 71. Re 

gens burg 1888. 

Burnham, S. H.: Lichens of Lake George. - Bryologist 25. Pittsburgh 

1922. 

Degelius, G.: Contributions to the Lichen Flora of N. America I-II. - Ark. 

f. Bot. 30 A, 1 & 3. Uppsala 1940-41. 

Dubrovsky, G. 1.: Notulae ad floram lichenum reservati "Stolby" regionis 

Krasnojarsk. - Not. Syst. Crypt. Inst. Bot. Komarovii. Acad. 

Scient. URSS. 9. Moskva 1953. 

Du Rietz, G. E.: Die Soredien und Isidien der Flechten. - Sv. Bot. Tidskr. 

18. Uppsala 1924. 

Eckfeldt, J. W.: An Enumeration of the Lichens of New Foundland and 

Labrador. - BuH. Torrey Bot. Club. 22, 6. Lancaster, Pa. 1895. 

Evans, A. W. & Meyrowitz, R.: Catalogue of the Lichens of Connecticut. 

- Conn. State Geol. Nat. Hist. Surv. BuH. 37. Hartford 1926. 

Flagey, C.: Catalogue des Lichenes. - Battandier & Trabut: Flore de 

l'Algerie, 2. Alger 1896-1897. 

Galløe, O.: Natural History of the Danish Lichens. 7. Copenhagen 1948. 

Gattefosse, J. & Werner, R. G.: Catalogus Lichenum Marocanum. - BuH. 

Soe. Sei. Nat. Maroc. 11. Roehefort-sur-mer 1931. 

Hasselrot, T. E.: Nordliga lavar i Syd- oeh MeHansverige. - Aet. Phyto 

geogr. Sueciea 33. Uppsala 1953. 

Lamb, 1. M.: Lichens of Cape Breton Island, N ova Seotia. - Ann. Rept. 

Nat. Mus. Canada 1952-53, BuH. 132. Ottawa 1954. 

Lynge, B.: Studies on the Lichen Flora of N orway. - Vid. Selsk. Skr. r. 

Mat. Nat. KI. 1921, 7. Kristiania 1921. 

: Liehens from Novaya Zemlya. - Rep. Norw. Exp. N. Z. 1921, 

43. Oslo 1928. 

: A contribution to the Lichen Flora of Canadian Arctic. - Publ. 

Catholie Univ. America, Washington, D.C. 1939. 

Maas Geesteranus, R. A.: Revision of the Lichens of the N ether lands. 1. 

Parmeliaeeae. Leiden 1947 (Diss.). 

Magnusson, A. H.: Lichens from Torne Lappmark. - Ark. f. Bot. 2, 2. 

Uppsala 1952. 

Merrill, G. K.: Liehens CoHeeted by the Canadian Arctic Exp. 1913-18. - 

Rep. Can. Arct. Exp. 1913-18, 4. Ottawa 1924.

- 246- 

Nichols, G.: Lichens of Douglas Lake. - Bryologist 28. Pittsburgh 1925. 

Rasanen, V.: Die Flechten Estlands L - Ann. Acad. Sci. Fenn., Ser. A. 

34, 4. Helsinki 1931. 

Tavares, C. N.: Contribul

RUSTSOPPER OG RUSTVERTER 

NYE FOR FÆRØERNE 

Av ASBJØRN HAGEN 

Under studier av arktiske planter i Botanisk Museum i København 

så jeg også leilighetsvis over en del planter fra Færøene og gjorde 

herunder noen funn av mykogeografisk interesse. 

En historikk over den mykologiske utforskning av Færøene er gitt 

av MØLLER (1945, s. 11-13). MØLLER'S meget vakre avhandling samt 

ROSTRUP'S liste (ROSTRUP 1901, s. 304-316) er de to arbeider hvori 

vår viten om Færøenes rustsopper er samlet. 

Hva vertplanter for Puccinia hieracii angår, anfører ROSTRUP 

(1901, s. 307) bare ))On Hieracium(() og MØLLER (1945, s. 101-102) 

nevner Hieracium caesium FR. og H. vulgatum FR. Da både H. 

DAHLSTEDT og S. O. F. OMANG har revidert de færøiske hieracier i 

København-museet, ønsket jeg å se over disse, og jeg fant en del nytt 

materiale m. h. t. rustangrep på de forskjellige småarter. Det verdifulle 

rustsoppmateriale som ble tilveiebragt på denne måte, ble selvsagt 

til dels sparsomt, og jeg foretok derfor med en gang omhyggelige 

sporemålinger (i 50% meIkesyre), og har på grunn av materialknappheten 

oppbevart også preparatene. 

1. Chrysomyxa empetri (PERs.) SCHROET. 

På Empetrum hermaphroditum (LGE.) HAGERUP. Haugen ved 

Thorshavn, II, 12.6.1927; leg. FR. SVENDSEN (uredosporer 28.5-38 

( 46) X 23-28.5,u ). - Chrysomyxa empetri er ny for Færøene, men er 

kjent f. eks. fra Island (JØRSTAD 1952, s. 10-11, 85), Norge, Skottland, 

Vest- og øst-Grønland (HAGEN 1952, s. 97) og ø.- og V.-Nordamerika. 

- 2 47 -

- 248- 

2. Puccinia arenariae (SCHUM.) WINT. 

På Sagina procumbens L. Thorshavn: Høivikshauge, 23.7.1934; 

leg. LEVI RYGG (teleutosporer 30-52 X 14-17 p, apikal fortykkelse opp 

til 6.5 p, noen sporeskaft målte mellem 50 og 76.5 p). - Denne rust 

sopp iant jeg under studier av færøiske planter i Botanisk Museum, 

Oslo, og den oppbevares her. Puccinia arenariae er ny for Færøene, 

men er funnet en gang på Sagina procumbens (og bare på denne vert 

plante) på Island. J0RSTAD (1952, s. 25) skriver om denne rustsopps 

forekomst på Island: "Evidently this rust is ve ry rare in Iceland. 

S. procumbens is quite common, and although I regularly investigated 

this species carefully, I never succeeded in finding the rust." 

3. Puccinia hieracii MART. 

På .,.,Hieracia cerinthoidea((., det. H. DAHLSTEDT, av ROSTRUP be 

stemt til H. murorum L. Strømø: Sandegjærde, n+ III, 31.7.1867; leg. 

E. ROSTRUP (uredosporer 25.5-33 X 22-27 p; teleutosporer 26-37.5 X 

19-22.5 (26.5) p). 

På Hieracium constrictiforme DAHLST. (Vulgata genuina). "Insula 

Suderø, in rupibus ad Vaag, alt. m. 25, 10.8.1895, No. 757; leg. 

H. G. SIMMONS; det. H. DAHLSTEDT", II+III (uredosp. 25-27 X 20.5- 

24.5 p, teleutosp. 24-35.5 X 19-23 p). Strømø, "ved Bygden Kvalvik", 

(II) + III, 21.9.1925; leg. LEMCHE; det. S. O. F. OMANG (uredosp. 

ytterst sparsomme i prøven, gamle 26 X 25 p; teleutosp. rikelig, 22- 

34.5 X 18.5-24p ). 

På H. epileucoides DAHLST. (Rigida) ., DAHLSTEDT 1903, s. 639- 

640; av OMANG kalt H. simmonsianum DAHLST. var. epileucoides 

DAHLST. (OSTENFELD & GR0NTVED 1934, s. 154). Strømø: hammer ved 

Torsvig, II + In, 21.8.1897; leg. J. HARTZ & C. H. OSTENFELD (uredosp. 

24.5-30.5 X 19.5-27 p; teleutosp. 25-38 X 19-27 p). 

På H. faeroense DAHLST. (Rigida) . Suderø: Frodebø, n + (III), 

7.8.1867; leg. E. ROSTRUP (prøvene inneholdt så å si bare uredosporer : 

22-29.5 X 19-26 p, og ytterst få teleutosporer: 34 X 20 p). Frodebø, n + 

III, 19.7.1897; leg. J. HARTZ & C. H. OSTENFELD (uredosp. 22-28.5X 

21-25.5p; teleutosp. 27-40.5 X 19-22p). Thorshavn: skrænt ved Sande 

gjærde, n + III, 30.8.-5.9.1903; leg. C. H. OSTENFELD (uredosp. 22.5- 

29.5 X 22-26.5 p; teleutosp. 27-38 X 19-24.5 p). 

På H. gnophodes OM. (Cerinthoidea)., det. S. O. F. OMANG (av H. 

DAHLSTEDT bestemt som H. leucograptum DAHLST.). Strømø: Højvig, 

"in grassy hillslopes in the lowland", II + III, 29.8.1903; leg. C. H.

- 249- 

OSTENFELD (uredosp. 25.5-28.5 X 21.5-24.5 Il; mest av teleutosporer: 

26-38 (48) X16-261l). 

På H. kalsoense DAHLST. (Cerinthoidea). Kalsø: Mygledal, n + 

(III), 5.8.1897; leg. J. HARTZ & C. H. OSTENFELD (uredosp. 24-29.5 X 

18.5-27 Il; i prøven bare noen få teleutosporer: 28.5 X 19.5-24.5 Il). 

På H. peramplum DAHLST. (Cerinthoidea). 0sterø: hammer ved 

Troågjov nord for Svinå, n + nI, 12.8.1896; leg. C. H. OSTENFELD 

(uredosp. 26-30 X 19-24.5; teleutosp. 27-37 X 19-24.5 Il). Suderø: Tran 

gisvåg, oppskyllet floddelta, n + III, 18.7.1897; leg. J. HARTZ & C. H. 

OSTENFELD (uredosp. i flertall i prøven: 23.5-27.5 X 17.5-241l; te 

leutosp. 24.5-30.5 X 19-25.5 Il). Thorshavn, "in grassy, fairly dry pla 

ces near the rivulet," II + III, 29.8.1903; leg. C. H. OSTENFELD (i prø 

ven uredosp. i flertall: 24-36 X 18.5-241l; teleutosp. 27-30 X 19-241l). 

På H. sarcophylloides DAHLST. (Silvatica) . østerø: Kodlen nær 

Eide, 400 m o. h., II + III, 17.8.1895; leg. H. G. SIMMONS (uredosp. 

25.5-28.5 X 19-24.5 Il; teleutosp. 23-35.5 X 19-23 Il). 

På H. scoticiforme DAHLST.(Rigida). "Insula Suderø, in rupibus 

ad Vaag," II+III, 10.8.1895; leg. H. G. SIMMONS (uredosp. 23.5-27.5 X 

20.5-251l; teleutosp. (23) 29-35.5 X 18-21 (24.5)1l). Strømø: Kirkebø, 

II + III, 18.8.1897; leg. J. HARTZ & C. H. OSTENFELD (uredosp. 24.5-31.5 

X 19-24.5 Il; teleutosp. 27.5-33.5 X 19.5-24.5 Il). Strømø: Højvig, "in 

grassy, fairly dry slopes near the sea", (II) + III, 29.8.1903; leg. 

C. H. OSTENFELD (uredosp. i prøven sparsomt: 25-32 X 20.5-25.51l ; 

teleutosp. 28.5-41 X 16.5-241l). Strømø: Kirkebø, II + III, 10.8.1913; 

leg. J. JEPPESEN (uredosp. 23-25.5 X 21-241l; teleutosp. 22-34(38) X 

18.5-26.5 Il) . 

På H. subrubicundum DAHLST. (Cerinthoidea). "No. 755. Ad 

"Karagjov" prope Qualbø insulae Suderø. Alt. m. 100", II, 7.8.1895; 

leg. H. G. SIMMONS (sparsomt materiale, uredosp. 22-27.5 X 18-24.5 Il). 

Suderø: på en hammer ved Trangisvåg, II + III, 17.8.1895; leg. C. H. 

OSTENFELD (uredosp. 24.5-26 X 22-24.51l; teleutosp. dominerte i prø 

ven: 27.5-41.5 X 18-23 (31.5) Il). 0sterø: hammer ved Troågjov nord for 

Svinå, II, 12.8.1896; leg. C. H. OSTENFELD (bare uredosp. i prøven: 

23-29.5 X 19-26 Il). Bordø: Holgafjæld, 475 m, II + III, 16.8.1897; leg. 

J. HARTZ & C. H. OSTENFELD (overveiende uredo i prøvene, til dels 

gammel, uredosp. 26-30 X (16.5)18.5-26.51l; teleutosp. 27-35 X 19- 

221l ). 

På H. subvarians ZAHN, syn. H. heterophyllum DAHLST. (Cerinthoi 

dea). Kunø: Haugen ved Haraldssund, II + III, 4.8.1897; leg. J. HARTZ 

& C. H. OSTENFELD (uredosp. 22-32.5 X 19.5-24.51l; teleutosp. 24-33 X 

18.5-25 Il).

- 250- 

På H. veterascens DAHLST. (Cerinthoidea). "No. 475. Ad Mølen 

prope Eide insulae 0sterø", II + III, 19.8.1895, leg. H. G. SIMMONS 

(uredosp. 24.5-27.5 X 19-21.5 /-l; teleutosp. 30-39 X 18-24/-l). 0sterø: 

Næs- Reuk, på en hammer, II + III, 20.8.1897; leg. J. HARTZ & C. H. 

OSTENFELD (uredosp. 23-27.5 X 19.5-24.5/-l; teleutosp. 27.5-37 X 19- 

24/-l ). 

På H. veterascens DAHLST. var. eidense DAHLST. 0sterø: Mølen 

ved Eide, II + III, 19.8.1895; leg. H. G. SIMMONS (No. 11 og No. 461; 

uredosp. 22.5-28.5 X 19-27/-l; teleutosp. 25-31.5 X 18-22 (28) /-l). 

Sett under ett blir sporemålene for det her undersøkte materiale 

av Puccinia hieracii: 22-33 (36) X (16.5) 17.5-27 /-l for uredosporenes 

vedkommende og 22-41.5 ( 48) X 16-28(31.5) /-l for teleutosporene. 

Det fremgår av ovenstående at mange av de angrepne arter tilhører 

den vest-europæiske artsgruppe Cerinthoidea (OMANG 1949); av til 

sammen 25 prøver tilhørte de 14 Cerinthoidea) 8 Rigida) 2 Vulgata 

genuina og 1 Silvatica. 

SUMMARY 

Two rusts and several host plant s of Puccinia hieracii new to the Færoes. 

In thi s small annotated list of rust specimens from the Færoes 

Chrysomyxa empetri and Puccinia arenariae are new species to the 

rust flora of the islands. A number of Hieracium species (identified 

by the specialists H. DAHLSTEDT and S. O. F. OMANG) are new Færoese 

host plants of Puccinia hieracii. Accurate spore measurements are 

given for each specimen. 

LITTERATUR 

Hagen, A.: Neue Beitrage zur arktisehen Pilzflora. Ber. Deutsch. Bot. Ges. 

65: 97--100. 1952. 

Dahlstedt, H.: The Hieracia from the FærDes. Botany of the FærDes. Part 

II: 625--659 and Plate XI, XII. Copenhagen 1903. 

J ørstad, 1.: The Uredinales of Iceland. Skr. utgitt av Det Norske Vidensk. 

Akad. i Oslo, I, 1951, 2: 1--87. 1952. 

Møller, F. H.: Fungi of the FærDes. Part L Basidiomycetes. Copenhagen 1945. 

Omang, S. O. F.: Cerinthoidea, et nytt element i den norske Hieracium 

flora. Blyttia 7: 8--12. 1949. 

Ostenfeld, C. H. & J. Grøntved: The Flora of Iceland and the FærDes. Copen 

hagen 1934 (H i e r a c i u m by S. O. F. Omang, pp. 146--168). 

Rostrup, E.: Fungi from the FærDes. Botany of the FærDes. Part I: 304-- 

316. Copenhagen 1901. 

Oslo, Januar 1956.

TWO POLYPORACEOUS FUNGI WITH 

MERULIOID HYMENOPHORE 

PORIA TAXICOLA (PERS.) BRES. AND POLYPORUS 

DICHROUS FR. EX FR. 

By LISE HANSEN 

One of the distinguishing characters between M eruliaceae and 

Polyporaceae is that the hymenium in the Meruliaceae is formed before 

the tubes develope and thus cover the edges of the dissipiments. A 

small group in the genus Poria shows this merulioid character, combi 

ned with normal polyporoid features such as rather deep and regular 

pores. This group, which contains the species Poria taxicola (PERS.) 

BRES., Poria violacea FR., and Poria purpurea FR., has been placed 

by several authors in a merulioid group of Poria and recently ele 

vated to generic rank as Merulioporia by BONDARZEW & SINGER 

(1941). Merulioporia is characterized by a soft pellicular, resupinate 

fruit body and the hymenophore is at first folded reticulate, then 

tubular, with edges of the tubes covered by basidia. The hyphae have 

no ampullae and no clamp connections, and the spores are cylindrical. 

During anatomi cal investigations on Polyporus dichrous FR. ex FR. 

it was found that this fungus shows merulioid characters too, in 

having fertile edges on the dissipiments. It hence appears that this 

merulioid character can be found in different groups of Polypores) 

which are not normally classified together. The genus Merulius s.1. 

itself has been divided into two genera: Serpula (PERS.) S. F. GRAY 

with brown spores, and Merulius FR. with hyaline spores. Even this 

last group seems heterogenous, combining species with distinctly 

allantoid spores and coloured hymenium and species with ellipsoid or 

cylindric spores and paIe or white hymenium. 

- 251-

- 252- 

The Botanical Museum of the University of Copenhagen holds a 

few collections of Poria taxicola) and more collections are found in the 

Herbarium of the Department of Plant Pathology, Royal Vetr. and 

Agric. Coll., Copenhagen. There are one collection of Polyporus di 

chrous in the Botanical Museum, and two collections in the Department 

of Plant Pathology, Royal Vetr. and Agric. Coll. But none of these 

species have been reported in the Danish literature. 

1. Pori l taxier la (PERS.) BRES. 

Syn.: Xylomyzon taxicola PERS.; Merulioporia taxicoZa (PERS.) 

BOND. & SING.; Polyporus rufus SCHRAD. ex FR.?; Poria rufa 

(SCHRAD. ex FR.) COOIm; Polyporus haematodus Ro STK. ; Poria hae 

mat odes (ROSTK.) EGELAND. 

The fruit body resupinate, membranaceous or fleshy membranace 

ous, adherent, 5-6 X 5-6 cm., often confluent, forming rather big 

crusts; margin normally adpressed, but sometimes free and slightly 

reflexed, 1-3 mm bro ad, white, velvety-pubescent. Pore layer flesh 

colour to reddish-brown, in dried specimens black, less than 1 mm 

thick, fleshy-waxy. Tubes at first merulioid, then becoming deeper, 

somewhat irregular, with entire mouths and obtuse, fertile edges, 

ca. 5 pr. mm. 

Context white, up to 1 mm thick, monomitic, formed of hyaline, 

branche d, septate hyphae, 3-7 p broad, walls up to 1 p thick, several 

hyphae collapsed, H-connections very common, no clamp connections 

se en in the context or on the hyphae of the hymenium, but occasional 

ly on mycelium (in pure culture) on very wide and thinwalled hyphae. 

The context continues directly into the pore trama. Hyphae of 

dissipiments like those of the context, but all thinwalled and with large 

intercellular globules of purplish-brown pigment. Basidia 15-25 X 

3-5 p, forming a compact palisade; no cystidia. Spores hyaline, 

cylindric-allantoid, 4-6.5 X 1.5 p . 

Poria taxicola is not a common fungus in Denmark. It is known 

from all parts of the temperate zone on the northern hemisphere. 

It grows on coniferous trees and fallen branches and causes a rather 

intensive yellow rot. 

M a t e r i a l s t u d i e d: Sjælland: Tisvilde Hegn, July 1898, on Pinus 

silvestris) leg. E. ROSTRUP; Køge As, May 23. 1895, on Picea abies) leg. 

E. ROSTRUP (both labelled as Merulius serpens) corrected by EGELAND to 

Poria haematodes) ; Hundested, Dec. 4. 1951, on Pinus sp., leg. et det. L. 

HARMSEN; Asserbo, Oct. 26. 1954, on Picea sp., leg. et det. L. HARMS:SN;

- 253- 

Kregme, Jan. 6. 1956, on a pole of Picea sp., leg. et det. L. HARMSEN; all 

eolleetions from the Botanieal Museum of the University. 

Sjælland: Tisvilde, July 1. 1898, on Pop1tlus canescens, leg. E. ROSTRUP; 

Tisvilde, July 1898, on Pinus silvestris, leg. E. ROSTRUP; Hornbæk Plantage, 

Feb. 1935, on Pinus silvestris, leg. BOOTS; Søllerød, Sept. 5. 1936, on worked 

timber of Pinus, leg. N. F. BUCHWALD; Dyssekilde, Sept. 1937, on Pinus 

silvestris, leg. R. HESTEHAVE; Kongelunden, April 15. 1950, on Pinus, leg. 

M. P. CHRISTIANSEN; Asserbo Plantage, June 20. 1951, on Pinus silvestris, 

leg. E. JØRGENSEN. - Jylland: Lammehave Skov, Ry, Oet. 14. 1951, 

on Quercus, leg. N. F. BUCHWALD; 3 other finds without detailed reeords; 

all eolleetions det. N. F. BUCHWALD, and from the Herbarium of Dept. Plant. 

Pathol., Royal Vetr. and Agrie. Coll. 

Two collections, from Tisvilde and from Lammehave Skov, grew 

on deciduous wood; they seem nevertheless to belong here. 

2. Polyporus dichrous FR. ex FR. 

Syn.: Leptoporus dichrous (FR.) QUEL.; Bjerkandera dichroa (FR.) 

KARST. ; Gloeoporus dichrous (FR.) BRES.; Gloeoporus conchoides 

MONT. ? 

The fruit body is sessile, often imbricate, 1.5-3 cm broad, 2-5 

mm thick, gelatinous soft when fresh, rigid when dry. Surfaee of cap 

white, light grey, isabelline, near basis almost black, tomentose, in 

concentric zones loosely tufted.-hispid. Margin thick, wavy, obtuse, 

more or less involute as dry, white. Pore layer waxy, in dried specimens 

horny, reddish purpIe, on margin more paIe, near basis purplish-black 

to black, easily separable from the context. Pores small, 

polygonal, regular, with entire mouths, ca. 5 pr. mm, dissipiments 

thin with obtuse and fertil edges; marginal pores low and indistinct, 

merulioid. 

Context white, 2-4 mm thick, monomitic, composed of hyaline, 

septate hyphae with slightly thickened walls and numerous clamp 

connections, either very much branched and leading in all directions, 

binding the hyphae together, or longitudinal arranged in bunches, 

sparsely branche d with long intervals. Hyphae 2.3-5.111 wide, walls 

up to 111. Context slightly thickened under the tomentose surfaee 

cover and forming adistinet pseudoP3:renchymatic layer over the pore 

trama, ca. 5011 thick, penetrated of normal context hyphae. This layer 

is connected with the pore trama through a ca. 5011 thick absrission 

layer, compose d of thin-walled, branched, septate, clamped, very 100sely 

woven, 2-511 wide hyphae. Hyphae on surfaee of margin parallel, 

adpressed, breaking up into tufts as margin developes. Tube trama 

compact with strongly gelatinized hyp hal walls, still penetrated of 

context hyphae which colour strongly in lactophenol-cottonblue.

a b c d 

Polyporus dichrous. Section through the context and pore trama. a. context; b. pseudoparenchymatic layer; c. abscission 

layer; d. pore trama. X 1000. 

N 

Ul 

If:.

- 255 

Subhymenium pseudoparenchymatic, not gelatinized. Hymenium for 

ming a very compact palisade of cylindric, 12-15 X 2-4 p, 4-spo 

red basidia. Numerous large balls of intercellular, purplish-brown 

pigment present; no cystidia. Spores hyaline, smooth, allantoid, 4-5 

X 1-2 p. 

Polyporus dichrous se ems to be a rather rare fungus. It grows on 

fallen branches and trunks of deciduous trees, in Europe esp. on 

BetuZa) Fagus and Quercus) rarely reported on Conifers. It is a cos 

mopolitic species, known from temperate zones of both hemispheres. 

It is found rather frequently in U. S. A., but is rare in Europe. 

M a t e r i a l s t u d i e d: Sjælland: Horserød Hegn, March 28. 1943, on 

Betula J leg. 0. WINGE; Helsingør, Dec. 1. 1953, on Betula J leg. M. S. KROG; 

both collections det. N. F. BUCHWALD and from the Herbarium of Dept. 

Plant. Pathol., Royal Vetr. and Agric. Coll.; Tisvilde, Nov. 13. 1955, on 

Betula J leg. M . LANGE, det. L . HARMSEN ; in the Botanical Museum of the 

University. 

It is worth while noting that the two species mentioned above 

have some taxonomical very important characters in common. The 

spores are in both species hyaline, cylindric-curved or allantoid, and 

the colour of the spore layer and the pigment are exactly the same. 

Besides both of them have as mentioned merulioid pores. 

They differ in their anatorny, the one being imbricate, with a 

distinct stratified context, hyphae with clamp connections, the other 

resupinate, with homogenous context, hyphae without clamp connec 

tions. Poria taxicola was carefully examined for stratified context in 

the reflexed parts of the fruit body, but no such tissue was present. 

The gelatinous pore trama of Polyporus dichrous) which does not 

exist in Poria taxicola) places this species in the genus Gloeoporus 

BRES. 

ACKNOWLEDGMENT 

The author is indepted to Professor N. F. BUCHWALD, L. HARMSEN, M. 

Sc., and Dr. MORTEN LANGE, who have supplied me with the material of 

these two interesting and rare species. 

RESUME 

En af karaktererne hos slægten M erulius - at porerne har fertile 

rande - træffes også i en lille gruppe af Poria. BONDARZEW & SINGER

- 256- 

(1941) har oprettet slægten Merulioporia for de følgende tre arter: 

Poria taxicola) P. violacea og P. purpurea. Ved en anatomisk under 

søgelse af Polyporus dichrous har det vist sig, at denne art også har 

fertile porerande. Det ser således ud til, at den merulioide karakter 

kan træffes inden for forskellige grupper af poresvampe, der normalt 

ikke stilles systematisk nær hinanden. 

Der er undersøgt 13 danske kollektioner af Poria taxicola og 3 

kollektioner af Polyporus dichrous. 

LITERATURE 

Bondarzew, A. S., 1953: Polyporaceous fungi of the European part of the 

U. S. S. R. and the Caucasus. 

Bondarzew, A. S. & R. Sin ger, 1941: Zur Systematik der P o l y p o r a 

c e ae. - Ann. Myc. 39: 43-65. 

Bourdot, H. & A. Galzin, 1927: Hymenomycetes de France. 

Donk, M. A., 1932: Revision der NiederIandischen H o m o b a s i d i o m y c et 

a e, A p h y Il o p h o r a c e a e II. 

Egeland, J., 1914: Norske resupinate poresopper. - Nyt. Mag. Nat. 52:156. 

Overholts, L. O., 1953: P o l y p o r a c e a e of the United States, Alaska and 

Canada. - Univ. Mich. Stud. Sci. Ser. Vol. 19. 

Pilåt, A. 1942: P o l y p o r a c e a e, I-III, in Kavina, C. & A. Pilåt: Atlas 

des Champignons de l'Europe, Tome 3. 


ON MERULIUS SILVESTER FALCK AND 

M. HIMANTIOIDES FR. 

PRELIMINARY REMARKS 

By LOUIS HARMSEN 

Institute of Technology, Copenhagen. 

Everyone who has to do with investigations on fungi in buildings 

especially the Merulius species, will sooner or later meet the name 

Merulius silvester FALCK in literature and in connection with it, meet 

different opinions regarding the value of the species. 

Many years of investigations on wood destroying fungi have given 

the author the perception that FALCK in his thorough observations 

on the Merulius species has found characters which have given reason 

for proposing new species; though it seems that the material used 

has been heterogene. Merulius minor FALCK thus probably has been 

proposed on specimens partly of M. pinastri (FR.) BURT - compare 

FALCK'S remark (1912, p. 400) : "Die Art ist wahrscheinlich identisch 

mit Merulius umbrinus FR. und M. hydnoides HENNINGS" - partly of 

M. tignicola HARMSEN (HARMSEN 1952). 

This matter has presumably contributed to the mycologists' doubt 

of those of FALCK proposed species, the more so as the species have 

not been re ally described. 

C u l t u r e s of Merulius silvester and M. himantioides. The 

publications referring to M. silvester also indicate that the species 

partly include M. himantioides FR. (see ULBRICH 1941, p. 23). A photo 

graph published by FALCK (1912, tab. XIV) of a pure culture of 

M. silvester corresponds completely to a culture of M. himantioides 

isolated by the author from a Danish specimen (Fig. 1). The heavy 

exudation of colourless drops on the mycelium is very characteristic, 

but it is not se en in an other strain or in cultures neither German nor 

FRIESIA V - 257- 17

Fig. 1: M. himantioides) no. 

2704 pure culture. X 14 . 

- 258- 

Fig. 2: M. silvester) no. 3613, 

pure culture. X 14. 

Danish identified as M. silvester (Fig. 2). The present material is too 

sparse to decide whether thi s heavy ,exudation has any diagnostic 

value. In other ways the cultures of the two species are uniform with 

white mycelium, in older cultures sometimes with yellow patches, 

rather loose and with very fine strands radiating from the inoculum; 

their growth is faster than normal for M. lacrymans) which has a 

similar mycelium in pure culture though usually somewhat more 

dense. They have both an optimum temperature about 27° C. and 

maximum temperature a littIe above 34° C. while M. lacrymans has 

its optimum at 20-220 C. and maximum at 28° C. (table 1). An optimum 

temperature about 27° C. is found likewise for M. pinastri and 

M. molluscus FR., both occurring as well in the open as in buildings. 

The temperature relations must be ascribed some significance even 

if it may be possibIe to alter them in experiments (FALCK, MEZ). When 

Table 1. 

Daily increment in mm (in contemporary experiments) 

Culture 

Temperature -- o C. 

15 I 21 I 27 I 31 I 34 

M. silvester) mean of 4 German strains 3.3 5.3 6.5 2.5 0.3 

do. 3613 1.6 2.7 3.0 1.8 0.1 

M. himantioides) 2704 2.4 4.2 4.8 2.5 0.3 

M. lacrymans) 888 (fast growing strain) 3.5 4.6 0.5 O O 

do. German strain 1.5 2.6 0.7 O O

- 259- 

isolated and kept under uniform conditions the temperature relations 

of the cultures will be constant. 

The f r u i t b o d i e s of the two species have like M. lacryman.s 

the characteristic rugulose-porose surface and yellow to yellow brown 

spores of the same size and form. On the basis of the present material 

it seems that M. silvester takes a position between M. lacrymans and 

M. himantioides. The fruit body is not as thick as normal M . lacrymans 

but thicker than M. himantioides. It is easily detached from the 

substrate and when dry it is tough and leathery like that of M . 

lacrymans. The sterile margin is flat, indefinite and formed of white, 

dense mycelium. The fruit body of M. himantioides is thin, nearly 

membraneous, attached to the substrate and is loosened only in flakes. 

When dry it is brittle. The sterile margin is white, membraneous, 

formed of rather loosely woven hyphae, often arachnoid. 

The m y c e l i a l s t r a n d s are well developed in M. silvester. 

They are grey as in M. lacrymans} but not noticed equally vigorous; 

they are more smooth with a barklike outer layer which is rather 

easily loosened, and mainly consisting of fiber hyphae. The mycelium 

between the strands is rather sparse. In M. himantioides the 

superficial mycelium is poorly developed, the strands are most often 

brownish grey or very dark, rather fine, noticed up to 1 mm. 

Fig. 3: M. lacrymans) section 

through a thin fruit body; 

note the thick subhymenial 

layer; stained with aniline 

blue in lactic acid. X30. 

Fig. 4: M. silvester ) no. 3613, 

section through fru it body; 

stained as Fig. 3. X30. 

17*

- 260- 

As will be seen from the present material it might be possibIe 

to distinguish the two species frequently, but it is difficult to find 

exact characters for delimitating them. 

T h e m i c r o s c o p i c a l s t r u c t u r e of the fruit bodies 

(compare table 2) of the two species is distinctly different from 

M. lacrymans which between the hymenium and the context has a 

layer of densely interwoven hyaline hyphae, more or less gelatinized. 

Here also the context is rather dense and stratified with numerous 

brown thick-walled hyphae with septa and clamp connections (Fig. 3). 

In the other two species the context is loose and spongy, some brown 

coloured hyphae occur, but they are thin-walled (Fig. 4). A thin layer 

of hyaline thin-walled hyphae is found in M. himantioides between 

the hymenium and the context (Fig. 5), while no special subhymenial 

layer in M. silvester is seen (Fig. 6). 

In the strands the thickness of t h e f i b e r h y p h a e is of 

special interest. In M. lacrymans the mean value is 4p (3-5(-5.5p» 

as given by FALCK (1912) and in M. silvester 2.5p (2-3(-3.5p». 

In M. himantioides the author has found 1.5-2p but fiber hyphae 

up to 4p have been noticed. When stained in Benzo brilliant blue and 

Benzo brown (STARFINGER 1953) the fiber hyphae in M. lacrymans 

will stain blue, while in M. silvester and M. himantioides they will 

stain brown with a slight tinge of blue. 

Fig. 5: M. himantioides} no. 

3885, hymenium and subhymenial 

layer, stained as 

Fig. 3. X330. 

Fig. 6: M. silvester} no. 3613, 

hymenium and part of the 

context, stained as Fig. 3. 

X330.

fru it body 

superficial 

mycelium 

temperature 

relations 

losses in 

weight of 

wood blocks 

(spruce) exposed 

to attack 

during 

4 months 

M. lacrymans 

thick, fleshy; 

margin thick, 

sharp 

- 261- 

T a b l e 2. 

M. silvester 

rather thick; 

margin flat, 

indefinite, 

formed of dens e 

mycelium 

I M. himantioides 

thin; margin 

membraneous of 

loose woven 

hyphae, 

arachnoid 

easily detached from substrate; 

tough and leathery when dry I attached; 

brittle when dry 

subhymenial 

layer of densely 

interwoven. 

hyaline hyphae, 

± gelatinized 

context rather 

dense, with 

numerous brown, 

thick-walled 

hyphae with 

septa and clamp 

connections 

vigorous, often 

as greyish, 

skinlike flak es, 

also between 

the strands 

strands vigorous, 

up to 8 mm, 

grey, surface 

uneven, wolly 

fiber hyphae 

about 4 Il 

(3-5.5fl ), 

stain blue with 

B.B. 

optimum 

20-22° C; 

max. about 

28° C 

50-60 % 

no special 

subhymenial 

layer 

thin subhymenial 

layer of 

thin-walled, 

hyaline hyphae 

context loose, spongy, with some 

brown but thin-walled hyphae 

with septa and clamp connections 

rather well 

developed, 

spars e between 

the strands 

strands rather 

vigorous, 

up to 4-5 mm, 

grey to brown, 

smooth 

with "bark" 

of fiber hyphae 

sparse and 

passing 

strands thin, 

up to 1 mm, 

grey to brown 

or dark, wolly 

fiber hyphae about 2.5 fl (1.5-3.5 

(-4fl» I 

(2-3.5 fl ) (1.5-2.5( -4 fl », 

stain brown and partly slight blue 

with B. B. 

about 14 % 

(5-18 %) 

optimum 27-28° C; 

max. about 35 ° C 

about 7 % 

(4-10 %)

- 262- 

Some experiments have been done in respect of the ability of the 

fungi to d e c o m p o s e t h e wo o d. Pure cultures of M. lacrymans 

mayas known cause a loss in dry weight of the wood samples on 

50-60 per cent or more in 4 months. In experiments with M. silvester 

the loss in dry weight is found to be approx. 14 per cent and with 

M. himantioides approx. 7 per cent. 

c o n c l u s i o n s. As aresult of the investigations it must be 

concluded that M. silvester is specifically different from M. lacrymans 

and it seems to be specifically different also from M. himantioides. 

M. silvester and M. himantioides might as well be regarded as extreme 

forms of a variable species. The author though has seen no intermediate 

forms. 

WM. BRIDGE COOKE mentions in a letter (June 1955) to the author 

that he considers M. lacrymans) himantioides and some American 

species to be all one species, however, not including M. silvester. In 

the same letter he mentions M. tignicola as a synonym of M. pinastri) 

the former as a "domestic" form of the "wild" pinastri. The author 

does, however, not agree to these considerations. The faet of regarding 

M. lacrymans as a "domestic" or "dry rot" form of the "wild" M. 

silvester or M. tignicola of M. pinastri seems no solution of the 

species conception, as the "wild" species occur as well in buildings as 

in the open. Should thi s point of view be of any reality M. silvester 

might better be regarded as a "domestic" form of M. himantioides 

than M. lacrymans of M. silvester. 

At the same time COOKE unfortunately propose to re-introduce 

the generic name Serpula for the b r o w n - s p o r e d species. The 

author, mainly working with wood destroying fungi, would prefer 

to conserve the generic name Merulius which is well known - and 

fe are d - as everybody who has to do with timber knows. This 

because that group of fungi is of very great importance in world 

economy, and it is therefore a matter of interest, not only for the 

mycological nomenclature and taxonomy. 

M a t e r i a l u s e d. M. silvester. H e r b a r i u m m a t e r i a l: 

4 collections from Germany (2 with fruit bodies), 1 Danish with fruit 

bodies (3613, Asserbo 26/10 1954) and 1 sterile (3827, from cooling 

waggon, D.S.B. 6/10 1955). P u r e c u l t u r e s: 4 German and 2 

Danish cultures.

- 263- 

M. himantioides: 2 collections from Denmark (2704, Gribskov 

25/111950 and 3885, Kregme 6/11956) and 2 Danish cultures. 

M. lacrymans: Several Danish collections; 1 Danish and 1 German 

culture. 

SUMMAR y 

The author has taken up an investigation of M. silvester FALCK 

as compared with M. himantioides FR. Differences are seen in the 

anatomy of the fruit body (compare table 2). On basis of the rather 

sparse material at hand, it is difficult to decide if they are specifically 

different; they might be forms of a variable species, though no inter 

mediate forms are seen. M. silvester is considered to be specifically 

different from M. lacrymans. It is suggested to conserve the generic 

name Merulius for the brown-spored species of this group of fungi. 

LITERATURE 

Buchwald, N. F.: De danske Arter af Slægten M e r u l i u s (Hall.) Fr. med 

en særlig Omtale af Gruppen C o n i o p h o r i Fr. - Dansk Bot. 

Arkiv 5, Nr. 21. 1928. 

Falck, R.: Die M e r u l i u s - Faule des Bauholzes (in Moller: Haus 

sehwammforsehungen VI). 1912. 

Harmsen, L.: M e r u l i u s t i g n i e o l a sp. nov. og dens Forekomst i Dan 

mark. - Friesia 4. 1953. (Fortryk 1952). 

De danske M e r u l i u s-Arter. - Bot. Tidsskr. 50. 1954. 

Mez, C.: Der Hausschvamm. - Dresden 1908. 

Starfinger, Kathe: Farben der Strange von Bauholzpilzen zur mikro sko 

pisehen Artbestimmung. - Holz-Centralbl. nr. 137. 1953. 

Theden, Gerda: Holzzerstorende Pilze in Berliner Gebauden naeh dem 

Krieg. - Fortschritte und Forsch. im Bauwesen. Reihe D. Hf. 

11. 1953. 

Ulbrich, E.: Hausschwamm, Nassfaulen (Trockenfaulen) und andere Zersto 

rer unserer Hauser und Bauten. - 1941. 


DIE IN FINNLAND 

BISHER GEFUNDENEN W ASSERPILZE 

Von ERNST HAYREN 

Die Wasserpilze sind in Finniand bisher nur mehr oder weniger 

zufaUig untersucht worden. N achstehend folgt ein Verzeichnis der 

beobachteten Arten. Durch kiinftige Untersuchungen wird die Anzahl 

der Arten in dem an verschiedenartigen Gewassern sehr reichen Lande 

si cher erheblich steigen. Wenn im Folgenden nicht anders erwahnt 

wird, ist die Art in der Gegend von Helsingfors gefunden. 

Olpidiaceae 

Plasmophagus oedogoniorum DE WILD. var. ulothricis WILLE. 1m 

Friihjahr in H ormidium flaccidum in den Schmelzlochern im Eise. 

- Mern. 5, S. 137. 1929. 

Rhizidiaceae 

Phlyctidium laterale A. BRAUN var. minus HAYR. 1m Flusse bei der 

Stadt Borgå in · Siidfinnland. Auf Ophiocytium parvulum. - Bidrag 

88, No 5, S. 106. 1944. 

Rhizophidium sphaerocarpum (ZOPF) FISCHER. Sockenbacka in 

der Nahe von Helsingfors. Auf Tribonema viride. - Mern. 20, S. 6.1945. 

Phlyctochytrium vernale (ZOPF) SCHROTER. In Chlamydomonas. 

- Mern. 5, S. 138. 1929. 

Ancylistaceae 

Lagenidium Rabenhorstii ZOPF. In steriler umhertreibender Spiro 

gyra. - Bidrag 80, No 3, S. 110. 1921. 

Ancylistes Closterii PFITZER. In einem Scharentiimpel im Kirch 

spiel Korpo, Regio aboensis. - Mern. 30, S. 18. 1955. 

- 264-

- 265- 

Monoblepharidaeeae 

Monoblepharis sphaerica CORNU. Nykyrka auf dem lsthmus karelicus 

(ausserhalb der politischen Grenze Finniands). - Medd. 29, S. 

166.1904. 

Saprolegniaeeae 

Saprolegnia dioica DE BARY. - Medd. 29, S. 165. 1904. - Mem. 

18, S. 89-93.1943. - Comm. Biol. 15, No 3, S. 1-45. 1954. 

S. monoica (PRINGSH.) DE BARY. - Medd. 29, S. 165. 1904. 

Mern. 20, S. 149. 1945. 

S. Thuretii DE BARY. - Medd. 29, S. 165. 1904. 

S. torulosa DE BARY. - Medd. 29, S. 165. 1904. 

S. asterophora DE BARY. lsthmus karelicus (ausserhalb der politischen 

Grenze Finniands). Auf einem Brachsen. - Mern. 4, S. 50. 

1928. 

Leptolegnia caudata DE BARY. Auf toten Chironomidenlarven. - 

Mern. 31. 1956. 

Achlya prolifera (NEES) DE BARY. - Medd. 29, S. 166. 1904. 

A. polyandra (HILDEBRAND) DE BARY. - Medd. 29, S. 166. 1904. 

- Mern. 20, S. 149. 1945. 

A. racemosa (HILDEBR.) PRINGSH. - Medd. 29, S. 166. 1904. 

A. spin osa DE BARY. - Mern. 20, S. 149. 1945. 

Leptom i taeeae 

Gonapodya polymorpha THAXTER. 1m Kirchspiel Ilomantsi, Karelia 

borealis. - Mern. 5, S. 97. 1929. 

Leptomitus lacteus (ROTH) AGARDH. In verschmutzem Slisswasser 

und auf einem toten Stichling. - Medd. 29, S. 166. 1904. - Mern. 6, 

S. 121. 1929-1931. - Mern. 20, S. 7. 1945. 

VERKtJRZUNGEN 

Bidrag = Bidrag till kannedom af Finlands natur och folk, utgifna af Finska 

Vetenskaps-Societeten. 

Comm. Biol. = Commentationes Biologicae, Societas Scientiarum Fennica. 

Medd. = Meddelanden av Societas pro Fauna et Flora Fennica. 

Mem. = Memoranda Societatis pro Fauna et Flora Fennica. 

Helsingfors, Januar 1956.

OM FUND AF JUDASØRE 

(HIRNEOLA AURICULA JUDAE (L.) BERK.) 

I KORSØR OG NÆRMESTE OMEGN 

Af K. HADERSLEV 

Siden den 27. September 1953, da jeg første Gang fandt et lille 

Eksemplar af Judasøre (Hirneola auricula Judae) i Korsør Skov 

(Lokalitet Nr. 1, se Kortet), har jeg holdt en Del Hyldetræer (Sambucus 

nigra) under Observation for eventuelt at finde flere Eksemplarer. 

Tidligt paa Aaret 1954 fandtes derefter et enkelt Individ, 

ogsaa i Korsør Skov (Lokalitet Nr. 3). I Sommeren 1954 iagttoges 

yderligere flere Frugtlegemer paa Lokaliteterne Nr. 3 og 5. I Vjnteren 

1954 blev Antallet af Findesteder forøget · med Lokaliteterne 

Nr. 4, 6, 43 og 45, og en ret stor Posefuld indsamledes. Den varme 

og tørre Sommer 1955 frembragte ingen Eksemplarer af Judasøre, 

men sent paa Aaret begyndte Svampen atter at bryde frem, og i 

December fandtes den i Overflod mange Steder. I Juledagene 1955 

noteredes den saaledes paa ca. 50 Lokaliteter, og der indsamledes ca. 

400 Gram Svampe (tørret Vægt). Alle Fund var hidtil blevet gjort 

paa Hyld (Sambucus nigra)) Artens almindelige Værtplante, men 

iblandt de sidste Fund fra Juledagene stammede et Par fra P o p p e l 

(Populus sp.) og K i r s e b æ r (Cerasus vulgaris)) der begge er nye 

Værter for Svampen i Danmark. En Hyldegren i ca. 2 m's Højde 

(Lokalitet Nr. 20) var besat med et Dusin grumset-mælkehvide 

Frugtlegemer. De største og de fleste Eksemplarer fandtes paa Grene 

og Stammer af Hyld, der laa paa Jorden. Ingen Eksemplarer er set i 

over ca. 2% m's Højde. 

I Betragtning af, at Judasøre i Almindelighed anses for en her i 

Landet sjælden Svampeart, skal der nedenfor gives en samlet Fortegnelse 

over de ialt 48 Lokaliteter i Korsør 'Og nærmeste Omegn, 

hvor jeg i de sidste Par Aar har samlet den. Tallene i Fortegnelsen 

henviser til Findestedernes Nummer paa det vedføjede Kort (S. 269). 

- 266-

- 267 

BUCHWALD (1928) har tidligere henledt Opmærksomheden paa 

den ejendommelige geografiske Udbredelse af Judasøre her i Landet. 

Den synes nemlig at være en udpræget "oceanisk" Art, der kun kan 

vokse nær Kysterne af de større Landsdele eller paa Smaaøer, og 

kan paa saadanne Steder undertiden endog optræde i meget stor 

Mængde, f. Eks. paa Hesselø og langs Kysterne af Bornholm. Min 

Paavisning af Svampens hyppige Forekomst i Korsør og Omegn, 

altsaa langs Kysten af Store Bælt, stemmer saaledes ganske overens 

med tidligere Iagttagelser over dens Udbredelse. 

FORTEGNELSE OVER LOKALITETER 

Hyld (Sambucus nigra). 

1. Korsør Skov, Østrand : Gruppe paa et Dusin Træer; her er næ 

sten altid Svampe, især paa en væltet Stamme. 

2. Korsør Skov, Østrand : Gruppe paa en Snes Træer; kun faa 

Eksemplarer. 

3. Korsør Skov, Vestrand nær Havet: Et enkelt Træ med mange 

Individer. 

4. Søskær Mose: Et Træ paa Roden med faa Eksemplarer og nogle 

fældede Træer med mange og store Frugtlegemer. 

5. Søskær Mose: 1/2 Dusin Træer, hvor der næsten til Stadighed 

har været nogle faa Individer. 

6. Søskær Mose: % Dusin Træer med kun faa og smaa Eksem- 

plarer. 

7. Søskær Mose: Et enkelt mindre Træ med et Par Individer. 

8. Søskær Mose: Et enkelt stort Træ med et Par Svampe. 

9. Forreste Markvej: Hegn af høje, klippede Træer med mange 

Frugtlegemer. 

10: Forreste Markvej: Ligeledes. 

11: Bageste Markvej: Lavt klippet Hyldehegn med et Par Haand 

fulde Eksemplarer. 

12. Bageste Markvej: Enkelte Træer i et klippet Hegn med faa 

Frugtlegemer. 

13. Fjordstien ved Grusgraven: Et Par Træer med faa Frugtlegemer. 

14. Fjordstien ved Kolonihaverne: Hegn paa Skrænten med faa Eks 

emplarer. 

15. Fjordstien ved Opgangen til Skoven: Et Par Hyldetræer i et 

Slaaenhegn med faa Individer.

Korsør 

o 

- 268- 

Store Bælt 

Store Bælt 

I ' 20000 

2 k m 

Korsør 

Kort over Korsør og nærmeste Omegn med Findesteder for Judasøre 

(Hirneola auricula Judae (L. ) BERK.) . 

Nor

- 269- 

16. Landevejen til Halseby, hvor der drejes af til Svenstrup: Et Hegn 

ved en Have med faa Frugtlegemer. 

17. Landevejen, Haneklint ind mod Vejen: En halv Snes Træer med 

tykke Stammer; her samledes en lille Kurvfuld Svampe. 

18. Landevejen, Høneklint ind mod Vejen: En halv Snes Træer og 

væltede Stammer; her samledes flere Haandfulde Frugtlegemer. 

19. Lunden ved Vandværket: Et fældet Træ med en Haandfuld store 

Svampe. 

20. Taarnborg Park: Et Par Træer ud mod Vejen med et Par Haandfulde 

Eksemplarer. 

21. Taarnborg Park: En gruppe paa et Dusin Træer med faa Individer. 

22. Revvej, vest for den nye Havn: Lavt Hegn og Buske i Grøften: 

Judasøre voksede mest paa Grene tæt ved Jorden; der indsamledes 

en lille Kurvfuld til Dels store Eksemplarer. 

23. Re vve j , Pilevænget : En enkelt Svamp paa en nedfalden Gren i 

Krattet. 

24. Revvej, Grosserer Næsteds Have: Faa Eksemplarer paa en 

Række nedskaarne Træer i et Hegn. 

25. Revvej, ved Privatvejen til Frugtplantagen: Paa Stammen af et 

stort Træ en Haandfuld Frugtlegemer. 

26. Revvej : Hegn ved Kolonihaverne med faa Svampe. 

27. Birkemosevej : Højt Hegn ved en Have med faa Individer. 

28. Birkemosevej : Mindre, fritstaaende Busk med faa Svampe. 

29. Slotsvej : Hegn ved en Have med faa Eksemplarer. 

30. Revvej Nr. 78: Et Træ i en Forhave med faa Eksemplarer. 

31. Hejsesvej, "Kærlighedsstien" : Hegn ved en Have med faa Frugtlegemer. 

32. Ved Viadukten: En nedskaaret Busk med en Haandfuld Individer. 

33. Magleøgade Nr. 22: Et Træ i Forhaven med faa Eksemplarer. 

34. Lilleøvej: Større Træ ved en Fiskerhytte med en Haandfuld 

Frugtlegemer. 

35. Lilleøvej: En Busk med faa Frugtlegemer. 

36. Lilleøvej: En Busk med faa Frugtlegemer. 

37. Fæstningsterrænet: Et Træstød samt flere spredte Træer eller 

Trægrupper; her samledes flere Haandfulde Frugtlegemer. 

38. Flasken Nr. 3: Træ i en Have med faa Eksemplarer. 

39. Badstuevej Nr. 16: Et Træ i en Have med faa Eksemplarer. 

40. Liitzensvej: Et Træ ved "De Gamles Hjem" med faa Individer. 

41. Nygade Nr. 25: En Busk med faa Svampe.

- 270- 

42. Møllebjergvej, ved Mathiesens Have: Et Hegn med faa Frugtlegemer. 

43. Sandvejsbanken, Fisehers Have: Faldefærdig Stamme med faa 

Eksem plarer. 

44. Kjærsvej i Quistgaards Have: Et Træ med en Haandfuld Eksemplarer. 

45. Jens Baggesensgade i Ejlerts Have: Et Træ med en Haandfuld 

Frugtlegemer. 

46. TejImanns Alle, ved Gasværket: Flere Træer, men kun eet Frugtlegeme. 

Kirsebær (Cerasus vulg,aris). 

47. Taarnborg Park: Ca. 7 cm. tyk Gren paa Jorden med nogle mindre 

Eksemplarer. 

Poppel (Populus sp.) 

48. Landevejen til Halseby: En Stribe af Frugtlegemer op ad den 

afbarkede Del af Stammen af et Vejtræ. 

SUMMARY 

A mass occurrence of Hirneola auricula Judae (L.) Berk. 

in Korsør, Sealand, in Denmark. 

After a single specimen of Hirneola auricula Judae (L.) BERK. 

found in 1953 near the town Korsør (Sealand, Denmark) an increasing 

number of finding-places has been discovered here. In the Christmas 

holidays ca. 400 gr. of the mushrooms (dried) was collected from ca. 

50 localities. 

LITTERATUR 

Buchwald, N. Fabritius: Om en Masseforekomst af Judasøre (Hirneola 

auricula Judae (L.) Berk.) paa Hesselø. - Meddelelser Foren. 

t. Svampekundsk.'s Fremme 4: 21-24. 1928. 


RECHERCHES SUR 

LE CYCLE DE COPRINUS DISSEMINATUS 

(PERS. EX FR.) S. F. GRAY 

Par J. P. JENSEN 

Les recherches sur le Coprinus disseminatus traitees ici ont eu 

lieu au pare "Jægersborg Dyrehave" a Klampenborg pres de Copenhague. 

Elles comprennent la periode allant du commencement de mai 

jusqu'a la fin de septembre 1948. 

Plusieurs localites ont ete etudiees, dont les trois indiquees sur 

le tableau, a savoir: 

A. Souche tres pourrie de Ulmus 

B. Souche assez fraiche de Acer campestre 

C. Base du tronc de Fraxinus sur l'ecorce. Arbre sur pied. 

Pendant la plus grande partie de la saison indiquee, la pluie est 

tombee avec des intervalles de secheresse de sorte qu'il parait possibIe 

d'etablir un lien entre certaines tombees de pluie et l'apparition du 

champignon et de constater que la croissance commence 4-5-6 jours 

apres une tombee de pluie suffisante. 

Elle a - d'une maniere constante - dure 5 jours, c. a. d. au 

quatrieme jour apres leur apparition les champignons s'affaissent en 

etat noirci. 

La petit Coprin a fleuri 7 fois au cours de la saison en question, 

revenant en meme temps dans les differentes localites, le plus vite 

sur la souche pourrie, un jour ou deux plus tard sur les autres h6tes, 

mais toujours de telle sorte que les memes endroits sont tout a fait 

envahis par les corpuscules. 

l re floraison: (date de l'apparition) le 4 juin, causee semble-t-il 

par les pluies 28/5 6 mm, 29/5 25 mm et 30/5 6 mm apres trois 

semaines assez seches. 

- 271-

- 273- 

2 me floraison: le 4 juillet. Pluies: 29/6 9 mm, 30/6 7mm et 1/7 3 mm. 

Dans l'intervalle 4-28 juin l'humidite ne mesurait que 10 mm en 

tout. 

3 me floraison: le 24 juillet. L'humidite tombee le 12-17/7 etait 

due il des averses d'orage fort limitees. Impossible d'etablir ici la 

relation faute de connaissance exacte. 

4 me floraison: le 11 aoilt. Pluies: 7/8 7 mm, 8/8 18 mm et 9/8 9 mm. 

La periode precedente 18/7-6/8 donnait seulement 2 mm. 

5 me floraison: le 24 aoilt. Pluies: 18/8 12 mm et 19/8 3 mm. 

Retardee ou non par la floraison precedente. 

6 me floraison: le 4 septembre. Pluies: 22-26/8 environ 30 m.m. 

L'apparition paralt ici retardee par la floraison precedente. 

7 me floraison: le 20 septembre. Pluies: 12-17/9 7 mm (?). Sans 

relation marquante. 

Les quatre dernieres floraisons sont indiquees en detail sur le 

tableau qui montre qu'une regeneration a eu lieu au bout de 11 il. 13 

jours. 

Quant il la temperature, elle a os cille d'une maniere stable entre 

15 0 et 30 0 le jour et 10 0-20 0 la nuit. 

Copenhague, fevrier 1956. 

FRIESIA V 18

MONASCUS RUBER VAN TIEGH. 

DEMONSTRATED IN DENMARK 

By HENRIK ALB. JbRGENSEN 

The Department of Plant Pathology, The Royal Veterinary 

and Agricultural College, Copenhagen. 

The Botanicai Department, The Danish State Experimental 

and Research Station for Plant Diseases and Pests, Lyngby. 

In September 1949 the Department of Plant Pathology of the 

Royal Veterinary and Agricultural College received from Mr. ALBERT 

HANSEN, Director of ALFRED JORGENSEN'S Zymological Laboratory, 

Copenhagen, for determination a culture of a fungus which had been 

isolated from the faeces of a child. It was stated that the mycelium 

assumed an intense red colour when cultivated on an acid nutrient 

medium. 

The fungus was transferred to the four following media: 2 % 

potato-dextrose-agar; 2% potato-dextrose-agar with an addition of 

0.1 % malie acid, sterilized in a current of water vapour (pH 4.2); 

autoclaved whole milk (pH 7.2); and autoclaved curdled milk (pH 

4.5). The cultures were allowed to stand at ordinary room temperature. 

In a few days, growth was ascertained on all media, and after ten 

days in both of the agar cultures a red mycelium had developed, which 

was proved by microscopy to contain perithecia as well as conidia. 

The perithecia, which were suggestive of Papulaspora-bulbils, 

were brownish red, roundish, had a mycelium appendage, and measured 

35-40/1 in diameter. The hyaline, ellipsoid ascospores measured 

4.5-6 X 3-4.5/1, on an average 5.25 X 3.75/1 (10 spores were 

measured). 

The conidia, which were present in large numbers, were brownish, 

pearshaped, and measured 10-13 X 8-11/1, on an average 11.75 X 

- 274-

- 275- 

9.25p (10 conidia were measured) ; they were constricted off terminally 

from short hyphae, ofte n 2 or 3 in a row, and may be designated 

as aleuriospores (VUILLEMIN 1910, 1931). 

Hence, there could be no doubt that the species of fungus concerned 

might be grouped under the genus M onascus VAN TIEGH., within 

the order Eurotiales (Plectascales). Fungi of this order form fruit 

bodies surrounded by a pseudoparenchymatous peridium within which 

asci are forme d irregularly in the loose parenchyma; on their maturit 

y the ascospores are released passive ly by the decay of the asci. 

Judging from the literature available on the subject (LIND 1913, 

O. ROSTRUP 1916 and 1935, KU)CKER 1924, and ALFR. JORGENSEN 

1940), none of the species of Monascus have ever been observed in 

Denmark before. 

According to YOUNG (1930), five well-defined species of Monascus 

have been described, viz. Monascus ruber VAN TIEGHEM (1884), the 

type species; M. mucoroides v. TIEGH (1884); M. purpureus WENT 

(1895); M. Barkeri DANGEARD (1907); and M. olei PIEDALLU (1910). 

It does not appear, either from PETRAK'S Verzeichnis (1938-39, 

1944) or the "Index of Fungi" (1940-55), that new species of this 

genus have been described later. 

In the determination of the fungus sent in, Monascus mucoroides 

could be left out of consideration directly because - unlike the 

present one - it will not form a red pigment in culture. 

All the four other species form a more or less intense red pigment 

and differ only by minor characters. 

M onascus olei is most easily distinguishable from the other three 

species by biological characters; for one thing, it is more thermophilic, 

it can growat temperatures right up to 46° C; and for this reason it 

could be ruled out. 

The last three species: Monascus ruber) M. purpureus and M. Barkeri) 

can be distinguished from each other by certain minute dissimilarities 

in shape and size of the perithecia, and, besides, there is a 

difference in the colour of the ascospores, both M. purpureus and 

M. Barkeri having reddish ascospores - purpie and brownish red, 

respectively - while the ascospores of M. ruber are hyaline. As the 

isolate received had hyaline ascospores, the species should thereby 

be determined as Monascus ruber. 

To support the determination, pure cultures of M onascus ruber 

and M. purpureus were imported from "Centraalbureau voor Schimmelcultures" 

at Baarn, in Holland, and for comparison these were 

18*

- 277- 

J ahresbericht 58 (1930), Abt. 1: 447-570, 1938; 60 (1931), Abt. 

1: 449-514, 1939; 63 (1935),2. Abt.: 805-1056, 1944. 

Peh·ak, F.: Index of Fungi 1936-1939. - Published by Commonwealth 

Mycological Institute, Kew. 116 pp., 1950. 

PiedalluJ A.: Sur une nouvelle moisissure du tannage a l'huile. Monascus 

oleL - Compt. Rend. Acad. Sci. Paris. 151: 397-399. 1910. 

Rostrup, O.: Bidrag til Danmarks Svampeflora. I. (Contributions to the 

fungus-flora of Denmark. L). - Dansk Bot. Arkiv 2, Nr. 5: 

1-56.1916. 

: Bidrag til Danmarks Svampeflora. II. (Contributions to the 

fungus-flora of Denmark. IL). - Dansk Bot. Arkiv 8, Nr. 8: 

1-74. 1935. 

van Tieghem, M.: Monascus, genre nouveau de l'ordre des Ascomycetes. 

Bull. Soc. Bot. France. 31: 226-230. 1884. 

Went, F. A. F. C.: Monascus purpureus, le champignon de l'Ang-Quac, une 

nouvelle Thelebolee. - Ann. ScL Nat. Bot., Ser. 8: 1-18. 1895. 

Y oung, E. M.: Physiological Studies in Relation to the Taxonomy of 

Monascus spp. - Trans. Wisc. Acad. ScL, Arts Lett. 25: 227- 

244. 1930. 

Index of Fungi. 1940-1954. - Publish. by Commonwealth Mycological 

Institute, Kew, 1940-1955. 

Lyngby, January 1956.

ON THE SONCHUS RUST 

PERISTEMMA PSEUDOSPHAERIA (MONT.) N. COMB. 

(SYN. PUCCINIA SONCHI ROB.) 

By IVAR JØRSTAD 

The Sonehus rust that in more than a hundred years has passed 

under the name of Puccinia sonehi) is certainly no true Puccinia) owing 

to the uredospores being produced in a peridium; nor is the specific 

epithet "sonehi" tenable, as the oldest valid name of the rust is 

Puccinia pseudosphaeria Mont. 

It seems that GROVE (1913, p. 156) was the first one to discover 

the peridial condition of the uredosori, stating that the "so-called 

paraphyses are really the upper part of adelicate, imperfect peridium, 

composed of hyaline pseudo-parenchyma (cells 5-10p diam.) ; at 

the top these cells become elongated, linear parallel, at first colourless, 

then brownish and more or les s clavate, and finally very dark brown, 

subopaque and irregular" . GROVE also expressed the opinion that "the 

species is widely different from a typical Puccinia)). LAMB (1934, p. 

185) agrees with GROVE, stating that "die Paraphysen seien in Wirklichkeit 

der obere Teil eines zarten Pseudoperidiums". 

H. SYDOW (1921, p. 174) places the rust near Miyagia) which like 

the Sonehus rust possesses Puccinia-like teleutospores, and in which 

the uredospores, and subsequently the teleutospores, develop in a funnel-shaped 

peridium consisting of elongated, paraphyse-like cells. But 

as the sole member of Miyagia then known, viz. M. anaphalidis Miyabe, 

is an aut-euform possessing also aecidia, H. SYDOW erected the new 

genus Peristemma for the brachyform Puccinia sonehi)· apparently 

he would otherwise have placed it in Miyagia. However, such difference 

is not considered to be of generic importance any more, and the genus 

Peristemma has, in faet, never been recognized; it has generally been 

looked upon as synonymous with Puccinia) what is rather surprising, 

considering that H. SYDOW placed it in the vicinity of Miyagia) which 

-278 -

- 279- 

is recognized as a valid genus by most authors; thi s is als o in accordance 

with the view of the present author.-l

- 280- 

sori; they are essentially of similar type, but in the long-covered teleutosori 

free paraphyse tips are less or not at all apparent. Instructive 

drawings of uredosori of the Sonehus rust are published by MAGNUS 

(1901, plate XIV, fig. 3) and LAMB (1934, p. 184, fig. 1), by the latter 

author also of teleutosori (with 1-celled teleutospores). 

Regarding the specific name of the Sonehus rust C. MONTAGNE 

described the species in 1840 under the name of Puccinia pseudosphaeria) 

while the name P. sonchi ROB. dates from 1849. However, in 1856 

MON TAGNE republished without alterations his diagnosis of P. pseudosphaeria) 

terming it a new species and without referring to his 

earlier description; as the latter was published in a work not well 

known by mycologists, it was generally overlooked. However, the 

oldest name, viz. from 1831, is Aecidium sonchi JOHNST., but as referring 

to the uredo-stage it is not valid; owing to the peridial paraphyses 

the uredosori have some faint resemblance to aecidia. - A full 

synonym list follows: 

Peristemma pseudosphaeria (MONT.) JØRST. n. comb. 

[Aeci.dium sonchi JOHNST., Flora of Berwick, 2, p. 205, 1831.] 

Puccinia pseudosphaeria MONT. apud BARKER-WEBB & BERTHELOT, 

Hist. Nat. Iles Canar., III, 2, p. 89, 1840. 

P. sonchi ROB. ex DESM., Ann. ScL Nat., Bot., III, 11, p. 274, 1849. 

P . pseudo-sphaeria MONT., Syll. PI. Crypt., p. 313, 1856. 

[Aecidium sonchi WEST., BuH. Acad. R. ScL Belg., 2. Ser., 11, p. 649, 

1861.] 

Uromyces sonchi OUDS., Rabh., Fungi Eur., No. 1952, 1872. 

U . puccinioides FAUTR. & ROLL. apud Roumeg., Rev. Mycol. 15, p. 15, 

1893. 

Puccinia tagananensis P. MAGN., Ber. D. Bot. Gesellsch. 19, p. 297,1901. 

Peristemma sonchi H. SYD., Annal. Mycol. 19, p. 175, 1921. 

Peristemma pseudosphaeria lives in the Eastern Hemisphere on 

many species of Sonehus) but is also reporled (ep. H. & P. SYDOW 

& BUTLER 1907, p. 494, and BUTLER & BISBY 1931, p. 74) from the 

Himalayas and from Mysore in South India on Launaea nudicaulis 

(L.) HOOK (syn. Zollikoferia nudicaulis BOISS.) , which is allied to 

Sonehus. 

Main hosts of the rust are in Europe the common weeds Sonehus 

a.rvensis L. s. 1. -)(' ), asper (L.) HILL, and oleraceus L. On these 3 hosts 

it has been found also in North Africa. On S. arvensis it has, further, 

*) Incl. Sonehus brachyotus DC., decorus CAST., mauritanicus BOlSS & 

REUT., and uliginosus M. B.

282 

Distribution of Peristemma pseudosphaeria. 

Russia and the Near Orient it extends eastward into West Siberia, 

Russian Central Asia and the Northwest Himalayas, but it has also 

an area of distribution in East Asia. It is als o reported from 

South India (on Launaea)) Ceylon (probably introduced) , New Zealand 

(introduced) , and from Uganda (here on a native host). It is 

thus seen, that the rust do es not extend as far towards north as some 

of its hosts, and als o avoids the more continental lowlands of central 

Europe. Apparently no other rust has a similar distribution, but Puccinia 

virgae-aureae (DC.) LIB., on Solidago) resembles it in so far as 

it inhabits the coastal lowlands of West Europe, but not the more 

continental lowlands of central Europe; it also occurs in the more 

southern European mountains, but differs from Peristemma pseudosphaeria 

in extending much farther towards north, chiefly in subalpine 

areas, while it is tot ally absent from the Mediterranean lowlands. 

However, in the subalpine areas of North Europe species of Sonchus 

are practically lacking, while on the other hand Solidago is often 

scarce in the Mediterranean lowlands and may be absent; in the Canary 

Islands it is lacking. The present author has elsewhere (JØRSTAD 

1948, p. 26) tried to explain the peculiar European distribution of 

P. virgae-aureae by assuming that infection in spring is possibie in

- 283- 

coastal are as with mild winters, and also in areas with a heavy snowcover 

that melts late, after warm weather having set in, but that 

infection is hampered in areas with cold winters without snow or with 

little snow melting early, as e. g. in the coastal areas of West Norway 

and in the continentallowlands of Germany. Possibly a similar reasoning 

is applicable also to Peristemma pseudosphaeria)' in this connection 

it should hardly make much difference that P. virgae-aureae is 

a non-leptosporic microform, but Peristemma pseudosphaeria a 

brachyform, consequently possessing a ure do-generation. 

REFERENCES 

Butler, E. J. & Bisby, G. R.: The fungi of India. - Imp. Counc. Agric. Res., 

Monogr. 1. Calcutta 1931 (237 pp.). 

Grove, W. B.: The British rust fungi (U r e d i n a l e s). - Cambridge 1913 

(412 pp.) 

Iwanow, K. S.: Parazitnye griby v okrestnostsjach S.-Peterburga letom 

1898 go da. - Trav. Soc. Imp. Natural. St. Petersb., 30, Fase. 3, 

Sect. Bot., pp. 1-20, 1900. 

Jørstad, 1.: Microcyclic Uredineae on Geranium and Solidago. - Nytt Mag. 

Naturv. 86: 1-30, 1948. 

Kuhnholtz-Lordat, G.: Les Uredinees corbiculees. - BuH. Acad. Sci. Lett. 

Montpellier, 1941 (Reprint, 2 pp.) 

Lagerheim, G.: Verzeichnis von parasitischen Pilzen aus Sodermanland und 

BohusHin. - Sv. Bot. Tidskr. 1: 18-40, 1909. 

Lamb, 1.: Entwicklungsgeschichtliche Untersuchung einer morphologisch 

abweichenden Puccinia-Art (P u c c. S o n c h i Rob.) - Hedwigia 

74: 181-194, 1934. 

Magnus, P.: tiber einige von J. Bornmiiller im Jahre 1900 auf den canarisehen 

Inseln gesammelte Uredineen. - Ber. D. Bot. Gesellsch. 

19: 292-300, 1901. 

Sydow, H.: Die Verwertung der Verwandtschaftverhaltnisse und des gegenwartigen 

Entwicklungsganges zur Umgrenzung der Gattungen 

bei den Uredineen. - Annal. Mycol. 19: 161-175, 1921. 

Sydow, H. & P. & Butler, E. J.: Fungi Indiae orientalis. Pars II. - Annal. 

Mycol. 5: 485-515, 1907. 

Sydow, P. & H.: Monographia Uredinearum. I. Genus P u c c i n i a. - Lipsiae 

1904 (972 pp.) 

Thirumalachar, M. J.: Some noteworthy rusts. II. - Mycologia 39: 231- 

248, 1947. 

Tranzschel, W.: Conspectus Uredinalium URSS. - Mosqua & Leningrad 

1939 (426 pp.). 

Trotter, A.: Basidiomycetes. - Saccardo, Sylloge Fungorum 23. - Abellini. 

1925 (1026 pp.) 

Oslo, January 1956.

BEITRAGE ZUR PILZFLORA JUTLANDS 

Von AXEL B. KLINGE 

1. Hirneola auricu)a Judae (L.) BERK. Diese Art ist von 

mehreren Standorten an der Ostkliste Djurslands bekannt, wo der 

Pilz recht zahlreich auf Sambucus nigra vorkommt, so z. B. bei Elsegaarde 

und Blushøj, 12.4.1953 (neuer Fundort), sowie in den Rugaard 

Waldern (Friesia 3: 69- 70,1944.). Wenn der Pilz wieder vom letzteren 

Fundort erwahnt wird, liegt es daran, dass ich ihn am 2.6.1952 

auf einem neuen Wirt fand, namlich auf abgefallenen Zweigen von 

Fagus silvatica (Friesia 4: 347, 1953). 

2. Cytidia floccuJenta (FR.) v. HOHN et LITSCH. (Fig. 1). In 

Spalten eines umgestlirzten, teilweise entrindeten Stammes von Popu 

lus alba. Gesellschaftlicher Wuchs. Der Pilz scheint an Populus 

Arten geknlipft zu sein, ist aber hier in Danemark selten. Am Strande 

bei Rugaard, Djursland, 16. 10. 1955. - Ein frliherer Fund aus Jlitland 

liegt vor, namlich Hammer Bakker bei Aalborg, 1935 auf Populus 

tre mul a} leg. POUL LARSEN. 

3. Hydnum coralloides SCOP. Auf Stammen von Fagus silvatica. 

Walder bei Fussingø bei Randers, im Troldeskov-Gebiet, 6.11. 

1955. 

4. Boletus versico)or ROSTK. Hut kirschrot. Die Fruchtkorper 

wachsen blindelweise im Grase an Waldwegen oder an Grabenrandern 

a u s s e r h a l b des Waldes. 

Wald bei østrup, Randers, 13. 8. 1949. 

Wald bei Hungstrup, Randers, 18.9. 1948. 

Am Grabenrand an der Landstrasse Randers-Grenaa in der Nahe 

der Løvenholm-Walder 31 km ostlich der Stadt Randers, 3.9.1951. 

Krabbesholm bei Skive am Grabenrand, 5. 9. 1955. 

- 284-

- 285- 

Fig. 1. Cytidia flocculenta (Fr.) v. Hohn et Litsch. 

Rugaard Strand, Djursland, auf Populus alba) 16. 10. 1955. 

Links: Fruchtkorper in feuchten Perioden. In der Mitte: Fruchtkorper 

von unten gesehen, so dass man das faltig-geaderte Hymenium sieht. 

Rechts: Zusammengerollte Fruchtkorper in trockenen Perioden. 

5. Collybia racemosa (PERS.) FR. Hut 0,3 cm, aschgrau, Stiel 

4-6 cm, Wurzelfasern treibend, von der Basis aufwarts mit zahlreichen 

abstehenden, zerstreuten Haaren versehen, die oft rudimentare 

Htite tragen. Wald bei Eskær, Tolne, 25.9.1955. In der modernden 

Laubschicht unter Fagus silvatica. 6 Fruchtkorper, von denen zwei 

mit einem schwarzen, rundlichen Sclerotium verkntipft waren. 

6. Dictyophora dnplicata (Basc.) ED. FISCR. Ein Individuum 

unter Fagus silvatica am Fusse des Htigels "Knøsen", Dronninglund 

Storskov, 10.9.1950. 

7. Geaster nanns PERS. Zwischen Flechten und Moos auf alteren 

Strandwallen mit einer Pflanzendecke aus Hieracium pilosella) 

Trifolium arvense) Veronica spicata) Saxifraga grwnulata und Festuca 

ovina bestehend. Sporen rund, 4,5-6!-l im Diameter. Capillitienfaden 

5-7!-l breit. Rugaard Strand beim Kalkbruch Glatved stidlich von 

Grenaa, 25.11.1951.

- 286- 

8. Ciboria Sydowiana (REHM) REHM. Auf Blattnerven und 

Blattstielen vorjahriger Bliitter von Quercus robur. Ascosporen 11,7 X 

6,5 p. Asci 115-130 X 8,5-9,0 p. Htigellandschaft, Læsten bei Randers, 

1.8.1954. Die Art ist ein Mal frtiher in Danemark beobachtet, 

namlich auf Seeland 1914, leg. O. ROSTRUP (Friesia 3: 277-278, 1947). 

9. Sclerotinia ficariae REHM. Auf sclerotisierten Wurzelknollen 

von Ficaria verna. Apothecien hellbraun. Stiel dunkler braun, haarfein. 

Fruchtscheibe 2-5 mm breit; der Sti el 1- 15 mm lang, je nach der 

Tiefenlage des Sclerotiums in der Erde. Ascosporen 12,5-15 X 4,5 

- 6,5p. Laut N. FABRITIUS BUCHWALD: Sclerotiniaceae Daniae (Friesia 

3: 312, 1947) ist Sclerotinia ficariae nicht in Danemark nachgewiesen; 

aber seit 1951 habe ich den Pilz an verschiedenen Standorten der Randersgegend 

sowie bei Mariager gefunden. Nicht allzu dichte Ficaria 

Bestande auf feuchtem, lockerem Boden bieten dem Pilze die besten 

Bedingungen. 1st der Ficaria-Teppich zu dicht, suche man am Rande 

des Bestandes oder wo Baume oder andere Vegetation den T'eppich unterbrechen. 

Die Apothecien erreichen ihre Vollreife wahrend der Bltitezeit 

der Ficaria-Pflanzen und etwa acht Tage spater. Die Fruchtkorper 

konnen dann so zahlreich sein, dass sie in Form von Sporenwolken 

ihre Anwesenheit verraten, wenn man die Finger durch den Teppich 

gleiten lasst. Sclerotien, die in der Erdoberflache liegen, konnen von 

fast sitzenden Apothecien ganz bedeckt sein. Die Sclerotien sind sehr 

verschieden geformt. Auf steifem Lehmboden sind sie langlich, oft 

schwach gekrtimmt, zugespitzt 3-5 X 1- 2 mm, wahrend sie auf lockerem, 

feuchtem Boden ofte r langlich-knollenformig sind und eben die 

schwammige Konsistenz haben, die man bei Sclerotinia tuberosa 

kennt, aber die Grosse tibersteigt wohl kaum 6 mm. Die sclerotisierten 

Wurzelknollen der Ficaria verna findet man Ende Juni, und sie sind zu 

dem Zeitpunkt im lnnern rosafarbig unter einer ausseren schwarzen 

Rindenschicht. 

Wald bei Haslund, Randers, 22.4.1951. - 28.4.1951. 

Wald bei Mygind, Randers, 29.4.1951. 

Walder bei Fussingø, Randers, 3.5.1951. - 7.5.1955 Massenvorkommen. 

Marenmølle bei Mariager, 13.5.1951 Massenvorkommen. - 29.7. 

1951 Sclerotien. - 26. 4. 1952. - 9. 5. 1952. - 29.6. 1952 Sclerotien. 

Wald bei Ulstrup, Randers, 6.5.1951. 


Walder bei Bidstrup, Laurbjerg, 8. 5. 1955.

- 287- 

10. Myriosc1erotinia Duriaeana (TUL.) BUCHW. Auf vorjahrigen, 

fast vom Wasser aufgelOsten Stange In von Carex acutiformis. 

Ascosporen 13 X 5,2p. Asci 130-140 X 7-9 p. Sclerotien spindelformig, 

schwach gesaumt, dicht und deutlich gestreift, mit 1-2 Apothecien. 

Wald bei Toholt, Langaa, im Sumpfwald auf Carex acutiformis} 

22.5.1955. 

In einem Carex-Sumpf auf vorjahrigen Stangeln einer Carex sp., 

der H omostachyae-Vigneae-Gruppe angehorig (C. disticha ?). Sclerotien 

mit 1-3 Apothecien. Moor bei Læsten, Randers, 28.5.1955. 

Die giirtelweise gestellten Mikrosporodochien, die man Ende Juni 

sollte beobachten konnen, habe ich nicht nachgewiesen. 

11. Rhizina inflata (SCHAEFF.) KARST. Auf Brandstatten zusammen 

mit Flammula carbonaria} Geopyxis carbonaria sowie zahlreichen 

Collybia ambusta. Wald bei Ulstrup, Randers, 25.9.1955. 

12. Hypocrea alutacea (PERS.) TUL. Wenige Individuen auf 

einem Erdwall (Waldeinfriedigungswall) unter Corylus avellana zu- 

Fig. 2. Hypocreopsis lichenoides (Tode) Seaver auf Corylus avellana .. 


Man bemerke, wie der handformige Pilz die Unterlage mit seinen »Fingern« 

umfasst.

- 288- 

sammen mit zahlreichen SpathuZaria flavida. Wald bei Gjerrild, Djursland, 

26.7.1953. 

13. Hypoereopsis liehenoides (TODE) SEAVER (Fig. 2). Nach 

J. LIND, Danish Fungi (1913) solI E. ROSTRUP am 31. 12. 1864 den Pilz 

auf Corylus gesammelt haben. In den spateren Jahren habe ich den 

Pilz an verschiedenen Standorten notiert, aber in keinem Falle waren 

Ascosporen gebildet. Der kanelartige Belag auf den Fruchtkorpern 

besteht aus Nebensporen. (PETCH in Trans. Brit. Myc. Soc. 21: 288, 

1938). 

Hiigellandschaft bei Læsten, Randers, 1.8.1954 auf Zweigen von 

Sorbus aucuparia. 

Moor bei Læsten, Randers, 11. 4. 1955 auf Zweigen von Salix aurita 

und vorjahrigen Stangeln von Chamaenerium. 

Klørup bei Randers, 3. 4. 1955 auf Salix caprea und Picea canadensis. 

Kaas, im westlichen Salling, 27.7.1955 auf Corylus avellana und 

Lonicera periclymenum. 

Wald bei østrup, Randers, 13.11.1955 auf Corylus avellana. 

Weiter habe ich 1955 Material aus der Gegend bei Ribe (leg. LEN 

NART EDELBERG) erhalten. 

14. Daldinia tuberosa (ScoP.) SCHROET. Auf abgefallenen 

Zweigen eines Laubbaumes (Fagus ? )} Rugaard Sønderskov, Djursland, 

6. 6. 1943. 

Ein Massenvorkommen in einem Erlensumpf auf abgefallenen 

Zweigen von Alnus glutinosa. Walder bei Fussingø, Randers, 6.11. 

1955. 

Randers, Danemark, Februar 1956.

PYRENOMYCETES PARASITIC ON 

HYPOGEOUS FUNGI 

By MORTEN LANGE 

Two members of Clavicipitales) viz. Cordyceps capitata and C. 

ophioglossoides range with Nyetalis among the most well known 

fungi parasitic on other fungi. They inhabit various species of the 

Elaphomyces granulatus group. C. ophioglossoides is not uncommon 

in Denmark ; I have noted it on Elaphomyces muricatus) E. variegatus) 

and E. granulatus coll. C. capitata is much less frequent and have 

been found but a few times in this country. I have collected it in 

Grib Skov (Elaphomyces variegatus) Sept. 27, 1945), Tisvilde Hegn 

(E. granulatus coll., Oct. 14, 1945 and later years), Frederiksdal 

Skov (E. variegatus) Aug. 10, 1952), and Rude Skov (E. variegatus) 

Aug. 18, 1952 and later years) :>:-) 

There are, however, at least three other "pyrenomycetes" which 

are known to occur on hypogeous· fungi. During my study of Danish 

hypogeas (LANGE, 1956), I have encountered these three species; 

two of them have not previously been found in Denmark, while the 

third has only been met with on epigeous hosts. Hypomyces chryso 

spel'mus was found on Arcangeliella asterosperma) Battarrina inclusa 

grew in the gleba of Tuber puberulum) while Melanospora ornat a inhabited 

the gleba of Elaphomyces asperulus. All three species belong 

in Hypocreales) although the one of them, Battarrina) is very reduced, 

and fin ally may be placed somewhere else. 

:;:) A third species with the same biology, Cordyceps intermedia IMAI, 

has not yet been found in Denmark. It is sometimes considered a 

hybrid, as it in all characters is intermediary between C. ophioglossoides 

and C. capitata. 

FHIESTA V - 289- IC)

- 291- 

isodiametric cells, inner part of 2-3 rows of more narrow cells; asci 

forming cluster in bottom of perithecium; paraphyses not seen; setae 

pointed, 25-40 p long, base more or less inflated, up to 10 p broad; 

asci clavate, short, stipitate, very thinwalled, 50 X 12-25 p , 7-8 

spored, spores irregularly arranged; ascus wall bursting before the 

spores are fully mature, no special apical structure, not coloured by 

Iodine ; spores 17.5-20.5 X 9.5-12.5p, lemon-shaped, snout-like protracted 

with germ pore in both ends, flattened, almost cigar-shaped 

in side view; distinctly reticulate, reticulum with wide, irregular, low 

meshes; dark brown when fully mature, but long remaining hyalin. 

Mycelium hyaline, a few hyphae springing from peridium. 

Numerous perithecia scattered in half mature gleba of Elaphomyces 

asperulus) Jylland, Munkebjerg by Vejle, Aug. 22, 1940, leg. 

J. E. & M. LANGE. 

The genus Melanospora has recently been monographed by DOGUET 

(1955), who divides the genus in several groups, using spore characters 

as main criterion. A small group (four species) is characterized 

by reticulate spores, but these species seem to be rare. None of them 

are mentioned by VON ARX & MULLER (1954), and PETCR (1938) reports 

but one of them, M. episphaeria PRILL. & PLOWR. from England. 

It differs from the present species through much larger spores and 

lack of setae. ZUKAL found Melanospora ornata growing on Polyporus 

zonatus) whHe DOGUET reports it from Elaphomyces (France) and 

from soH with Fusarium (Holland, Madagascar), DOGUET'S description 

covers my plant exactly, while ZUKAL gives somewhat smaller 

spores (14- 15 X 10-11p). 

DOGUET mentions no differences between his specimens from 

Elaphomyces and from soH. It should, however, be further studied, 

whether the plant occurring on Elaphomyces represents a distinct 

taxon. 

Another species of MeZanospora) M. zobeZii (CORDA) FUCK. is recorded 

from Tuber sp. (cp. PETCR, 1938), but is more frequently 

found on fruit bodies of Sepultaria (cp. LANGE & HANSEN, 1950). 

3. Hypomyces chrysospermus TUL. 

The imperfect stage of this fungus, Sepedonium chrysospermum 

FR., is frequently found on BoZetus spp. and Paxillus invoZutus. 

TULASNE (1860) gives als o Melanogaster and ArcangelieZZa (Octaviania) 

as hosts, and HAWKER (1955) has recently found it on 

Melanogaster and on Rhizopogon) while records from ArcangelieZla 

19*

- 292- 

are very few (SCHROETER, 1908). alder records from Scleroderma 

needs confirmation, especially as Boletus parasiticus could have been 

the true host, without thi s being noticed. 

The perfect stage, Hypomyces chrysospermus TUL., is rare ly 

found, and has, as far as I know, only been encountered on Boletus 

and Paxillus. The Danish finds of the perfect stage are all from 

Boletus. 

All Danish collections of Rhizopogon) Melanogaster and Arcangeliella 

asterosperma have been examined for Hypomyces-attacks. The 

result was negative for Melanogaster and Rhizopogon) but three colleetions 

of Arcangeliella asterosperma had distinct yellow crusts with 

chlamydospores of the Sepedonium-stage. ane collection (ML4601, 

Sjælland, Jægersborg Dyrehave, aet. 3, 1955, very strongly decayed) 

had only chlamydospores; another collection (ML4032, Sjælland, 

Pramskov, Vallø, Sept. 9, 1951) had young chlamydospores, and very 

well developed, hyaline, elliptic conidia. The third specimen (ML4199, 

Sjælland, Rude Skov, Aug. 20, 1952) was very strongly infected, the 

whole fruit body covered by yellow chlamydospores, which also 

developed in the surrounding soil. The gleba was, however, normally 

developed, with mature spores, and the peridium only slightly collapsed. 

Numerous perithecia were imbedded in the peridium; they 

were globose, up to 300 p broad, paIe yellowish brown, and hyaline, 

with a wall of 3 layers of thinwalled cells. The ascogonial hyphae 

were developed in the central part of the larger perithecia, while no 

asci were formed as yet. 

LITERATURE 

von Arx, J. A. & E. Miiller, 1954: Die Gattungen der amerosporen Pyrenomyceten. 

- Beitr. Kryptogamenflora Schweiz 11, 1. 

Doguet, G., 1955: Le genre M e l a n o s p o r a; biologie, morphologie, developpement, 

systematique. - Le Botaniste 39: 1-313. 

Hawker, L. E., 1955: Hypogeous fungi V. Two fungal parasites of hypogeous 

fruit bodies. - Trans. Brit. Myc. Soc. 38: 75-77. 

Lange, M., 1956: Danish hypogeous Macromycetes. - D. Bot. Ark. 16, 1. 

& E. Bille Hansen, 1950: Notes on Danish Fungi. - Friesia 4: 

61-65. 

Petch, T., 1938: British Hypocreaies. - Trans. Brit. Myc. Soc. 21: 243-305. 

Schroeter J J., 1908: Die Pilze Schlesiens. Zweite Halfte. 

Tulasne, L. R. & C., 1851: Fungi hypogei. 

1860: De quelques spheries fongieoles. - Ann. ScL Nat. (Bot.) 

IV, 13: 5-19. 


QUELQUES LEPIOTES NOUVELLES 

OU CRITIQUES 

LEPIOTA HELVEOLA BRES. SS. JOSSERAND, L. SERICIFERA 

(LocQ.) LocQ. et L. KUHNERIANA sP. NOV. 

par MARCEL LOCQUIN 

Des trois especes decrites ci-apres, l'une est nouvelle, l'autre rare, 

la troisieme mal connue bien que toxique. Elles appartiennent toutes 

les trois au genre Lepiota) florissant au Danemark, puisque c'est dans 

la F lo r a A g a r i c i n a D a n i c a, de J. E. LANGE que se trouve 

la source de toutes les etudes modernes sur ce genre. 

1. Lepiota helveola BRES. ss. JOSSERAND. 

Nous redonnons une description detaillee de cette espece toxique 

souvent meconnue car nous la connaissons bien de nombreuses stations 

differentes. 

Nombreuses recoltes dans les regions: Lyonnaise, AIsacienne et 

Parisienne, sous feuillus ou au bord des haies. Septembre-Octobre. 

C h a p e a u: (D = 20-35 mm) arrondi, puis convexe, puis 

etale, plus ou moins regulier, non mamelonne; revetement separable, 

densement veloute-hispide au disque chez le jeune, puis rompu diffracte 

en meches de taille et de forme tres irreguliere, ocre briquete, 

ocraee lie de vin, sur le fond rose ocre ou rose incarnat de la chair; 

marge incurvee puis droite, reguliere, iL peine plus paJe et moins ornee; 

chair cassante, peu epaisse, blanche, rosissant iL peine; odeur suave 

puis de lepiote; saveur fongique. 

p i e d: (H = 35-40 mm; d = 4-5 mm) non separable, assez 

trapu, cylindrace; chair iL cortex miel hyalin, puis pelure oignon, 

blanche en profondeur; moelle blanche; revetement supere blanc 

- 293-

SPOROBOLOMYCES AND OTHER YEASTS 

ON GRAINS OF BARLEY 

By AAGE LUND 

Tuborg Breweries, Copenhagen. 

The genus Sporobolomyces) together with Bullera) holds a special 

position among yeasts, due to its formation of the so-called ballistospores. 

These arise on sterigmata and, when ripe, are discharged 

by a drop-excretion mechanism. After KLUYVER & VAN NIEL (1925) 

calle d attention to the similarity between the spore-discharging 

mechanism in Sporobolomyces and in many Basidiomycetes) pointing 

out that Sporobolomyces might belong to the Basidiomycetes) many 

mycologists have associated themselves with this view. Others, however, 

regard the ballistospores as conidia and consequently in cl ude 

Sporobolomyces under Fungi imperfecti. However, it would seem more 

reasonable to regard the genus as belonging to the Basidiomycetes. 

According to LoDDER & . KREGER-VAN RIJ (1952), 7 species of 

Sporobolomyces are known which have been isolated from leaves, 

flower nectar, soil, air, yeast, beer, etc. in various parts of the world. 

As far as the writer knows, this yeast has not so far been mentioned 

in the literature as having been found in either Denmark or, for 

that matter, in the rest of Scandinavia. However, it should be noted 

that one of the red yeasts pictured by E. CHR. HANSEN (1879, Tab. II, 

Figs. 1-37) must doubtless be regarded as a Sporobolomyces· X ). This 

yeast, to which HANSEN gave no systematic designation, came from 

rose-coloured stains on some paste which he had left standing for a 

few days in the laboratory. 

The discharge of spores in Sporobolomyces salmonicolor (F. et BR.) 

KL. et v. N. was examined by MULLER (1954) through a very instruc- 

* ) P resumably being the oldest drawing of Spor obolom yces. 

- 297 -

- 298 - 

tive film he had made. However, the culture which MULLER used for 

his investigations was of Dutch, not of Danish origin. Mr. E. BILLE 

HANSEN, M. Sc., has informed the writer that in air analyses on agar 

plates he has found red yeasts which produced "mirror images" characteristic 

of spores dis charge d by Sporobolomyces. 

Recently, the writer found Sporobolomyces to occur commonly 

on grains of barley, and the idea readily suggests itself that it may 

occur, just as commonly on other vegetable matter in Denmark. 

DERX (1930) in the vicinity of Prague found Sporobolomyces on 

grains of barley and other cereals and also on straws and old leaves. 

Recently, LAST (1955) has shown Sporobolomyces roseus KL. et v. N. 

to occur commonly on old and dead leaves of barley and wheat in 

England. From these, moreover, he isolated a species of Bullera) the 

other ballistospore-forming yeast genus. 

It may seem strange that Sporobolomyces was not observed in the 

writer's former investigations on yeasts on natural substrates (LUND 

1954). This presumably is due to the faet that - as will be mentioned 

below - Sporobolomyces forms exceedingly small colonies with 

the method used and hence might be easily overlooked. The faet that 

it was found on barley was probably due to its being a dominant 

element of the yeast flora on such grains. 

COUNTING AND ISOLATING YEASTS 

In the very hot and dry summer of 1955, the yeast flora on grains 

of barley was examined. The investigations comprised barley-grain 

samples taken aseptically in sterile Petri dishes from the plants in 

two fieIds immediately before the harvest in August as well as samples 

of threshed barley taken immediately on its arrival at the Tuborg 

Breweries, Copenhagen. 

From each sample, 10 healthy undamaged grains were transferred 

to 10 cc sterile water in which they stood for 30 minutes during which 

time they were subjected to repeated vigorous shaking. The shakings 

were then plated out on hoppe d wort agar to which 0,25 % sodium 

propionate had been added. On thi s medium, the growth of mouIds 

is inhibited while the development of yeasts as a rule is unaffected. 

The agar plates were permitted to stand at about 20 0 C for 3-4 days, 

whereupon the yeast colonies that were found were counted and converted 

into number of yeast cells per grain. The figures found give 

the amount of yeast cells present on the outside of the grains. Some

- 299- 

A B 

Fig. 1. Sporobolomyces on agar plates at about 20° C. 

A. Wort agar 3 days old. - B. Wort agar with 0,25 % sodium propionate 

3 days old (colonies hardly visible to the naked eye). 

of the yeast colonies were transferred to wort agar slopes, and pure 

cultures were secured from these by repeated platings. 

Sporobolomyces} unlike other yeasts examined by the writer, 

forme d very small colonies on wort agar with sodium propionate, so 

that they could be counted only through a magnifying glass. In 

A B 

Fig. 2. Sporobolomyces on agar plates at about 20° C. 

A . Wort agar 5 days old. - B . Wort agar with 0,25 % sodium propionate 

5 days old (colonies readily visible to the naked eye) .

- 300- 

experiments with a pure culture, the yeast after 3 days at about 20° C 

formed readily visible colonies, frequently 1 mm in diameter, on wort 

agar (without sodium propionate). On the other hand, its colonies 

on wort agar + 0.25% sodium propionate under the same conditions 

were barely visible to the naked eye, measuring only 100- 250/1 in 

diameter (Fig. 1). Only after 5 days' growth the colonies were readily 

visible on that medium (Fig. 2). However, in the examination of 

grains for yeasts the agar plates should be counted af ter 3-4 days, 

longer standing frequently resulting in extensive mould growth. 

YEASTS ON BARLEY GRAINS 

The results of yeast-cell counts on grains of barley are given in 

Table 1. It will be se en that yeasts were found in 14 out of the 17 

samples (Nos. 1-17) taken on barley plants in two fields. They occurred 

in greatly varying numbers, all the way from ve ry few to 

3400 yeast cells per grain. Some of the samples contained so large 

amounts of yeast cells that the yeasts no doubt grew on the grains 

in question. 

Table 1, moreover, shows that the samples of threshed barley 

behaved somewhat differently. Out of the 15 samples (Nos. 18-32) 

from 3 lots of barley that wera examined, the presence of yeasts 

Field 

Table 1. 

Barley grains from fieIds Barley grains on arrival at Tuborg 

I 

Sample INumber of yeast Barley I Sample I Number of yeast 

no. cells per grain. lot no. cells per grain. 

A 1 

2 

3 

8 

13 

30 

I 18 

19 

20 

O 

O 

1 

4 41 21 3 

5 60 22 28 

6 160 

7 750 

II 23 O 

B 8 

9 

O 

O 

24 

25 

26 

O 

1 

1 

10 O 

27 2 

11 1 

12 1 

13 80 III 28 O 

14 110 29 1 

15 900 30 1 

16 2600 31 3 

17 3400 32 7

- 301- 

was demonstrated in 10 samples but they occurred in small amounts 

only, from 1 to 28 yeast cells per grain. 

Incidentally, somewhat similar findings were also made in regard 

to moulds. These occurred in large amounts in platings of the field 

samples, rendering the examination of yeast colonies exceedingly difficult. 

On the other hand, mouIds occurred in much smaller amounts 

in platings of threshed barley, so that yeast analyses presented no 

difficulties. It would seem that large amounts of mould spores and 

yeast cells were removed in the threshing process. 

A total of 36 isolates of yeasts from the barley samples were 

pure-cultured. 17 cultures of RhodotoruZa and 11 of SporoboZomyces 

were isolated, the rest being distributed on HansenuZa (3), ToruZopsis 

(4), and Candida (1). 

SporoboZomyces and the red RhodotoruZa yeasts are very similar. 

The difference between them is that SporoboZomyces forms basidiospores 

while no perfect stage is known in RhodotoruZa. According to 

MULLER (1954), the spores in Sp. saZmonicoZor are discharged to a 

distance of up to 0.4 mm in both the horizontal and vertical directions. 

To determine which of the red-coloured isolations belonged to 

Fig. 3. "Mirror images" consisting of discharged 

spores from Sporobolomyces colonies 

on the lid of an inverted Petri dish. 

Sporobolomyces) the isolates were inoculated on wort agar in Petri 

dishes and on wort agar slopes in tubes, both of which were placed 

with the layer of agar and the culture facing downwards. After 2-3

- 302- 

days at about 20° C "mirror images" consisting of discharged Sporobolomyccs 

spores were formed in 11 of the red yeasts. No discharge 

of spores occurred in the other 17 red isolations, which were consequently 

regarded as belonging to Rhodotorula. 

Future investigations, however, may show this distinction to be 

unjustified. As expressed by LoDDER & KREGER-VAN RIJ (1952), it 

seems very probable that at any rate certain Rhodotorula species must 

be regarded as asporogenous Sporobolomyces species. 

SUMMARY 

Sporobolomyces) together with Rhodotorula) was found to be a 

dominant ,element in the yeast flora on grains of barley in Denmark 

examined after the very hot and dry summer of 1955. On grains 

taken from barley plants immediately before the harvest, considerable 

amounts of yeast cells were often found. On the other hand, few 

yeast cells occurred in samples of threshed barley. 

REFERENCES 

Derx, H. G.: Etude sur les S p o r o b o lom y c e t e s. - Ann. Myc. 28: 

1-23. 1930. 

Hansen, E. Chr.: Organismer i 01 og 01urt. Kjøbenhavn 1879. 

Kluyver, A. J. & C. B. van Niel: uber Spiegelbilder erzeugende Hefenarten 

und die neue Hefengattung S p o r o b o lom y c e s. - 

Centralbl. Bakteriologie II, 63: 1-20. 1925. 

Last, F. T.: Seasonal incidence of S p o r o b o lom y c e s on cerealleaves. 

- Trans. Brit. Myc. Soc. 38: 221-239. 1955. 

Lodder, J. & N. J. W. Kreger-van Rij: The Yeasts. A taxonomic study. 

Amsterdam 1952. 

Lund, A.: Studies on the ecology of yeasts. Copenhagen 1954. 

Muller, D.: Die Abschleuderung der Sporen von S p o r o b o lom y c e s - 

Spiegelhefe - gefilmt. - Friesia 5: 65-74. 1954. 


ON METASPHAERIA COCCODES (KARST.) SACC. 

AND OTHER FUNGI 

PROBABLY RELATED TO MASSARINA SACC. 

(MASSARINACEAE N . FAM.) 

By ANDERS MUNK 

In a previous paper (MUNK 1953) I with some doubt placed the 

genus Massarina SACC. together with Massaria DE NOT. and Massar';'ella 

SPEG. Further investigations haverevealed that Massarina is 

more related to other genera which I (1953) placed elsewhere in the 

system. The present paper is an attempt to clarify the problem of the 

relationship of Massarina. As a conclusion of the report the new 

family Massarinaeeae is established. 

The most elucidating investigation was the microscopic examination 

of a Leptosphaeria-like fungus with hyaline spores, growing on 

sheaths of grass. It is preserved in the herbarium of the Department 

af Plant Pathology of The Royal Veterinary and Agricultural College, 

Copenhagen. The specimen was collected at Dronninggaard Skov 

(Sealand) by OVE ROSTRUP and was labelled Leptosphaeria anarithma 

(BERK. & BR.); it seems to me that abetter name for the fungus 

would be Metasphaeria coccodes (KARSTEN) SACC. (= Leptosphaeria 

coccodes KARSTEN). The fungus is distinctly related to Massarina 

and not to any gro up of Leptosphaeria. I shaH make no attempt to 

discuss the generic name of this fungus; all other species of M etasphaeria 

examined by me, however, have appeared to belong to more 

or less well-defined genera with other names, and I think it is reasonable 

to use the name Metasphaeria for it. (I am not aware that the 

genus Metasphaeria is so strictly typified as to forbid thi s procedure). 

- 303-

- 305- 

ecology which reminds one of Pseudotrichia; its spores are somewhat 

atypical for the group; it seems probable that it belongs to the same 

taxon as the other genera mentioned here. 

In my opinion the facts presented above justify the erection of 

the group to a new family, Massarinaceae: 

Massarinaeeae fam. n. (Genus typicum: Massarina SACC.). 

Pseudotheciis non-stromaticis vel saepius clypeatis. Peridio textura prismatica, 

atro-brunneo, ad basim saepius subhyalino, ad porum obscuriore, 

parvicellulare (conum internum nigrum, vel pilos rigidos, atros, acutos 

saepe formante ). - Ascis basalibus, bitunicatis; textura interasciculari 

paraphysiformi. - Ascosporis hyalinis, transversaliter septatis (2-cellularibus 

vel saepius 4-cellularibus vel ad 6-7-cellularibus), saepius strato tenui, 

mucoso, persistente obductis, ±. distincte guttulatis, typice valde refringentibus. 

e 

Massarinaceae. Pseudothecia (X 95) and spores (a X 765, f X 930, 

b-e X 585). - a. Metasphaeria coccodes. - b. Trichometasphaeria culmi/ida. 

- c. Massarina eburnea. - d. Massarina rubi. - e. Pseudotrichia 

minor. - f. Keissleriella sambuci. 

FRIESIA V 20

- 307- 

cylindric-clavate, sessile, double-walled; interascicular tis sue paraphy 

soid. Spores 1-2-seriate, transversally septate and constricted, hyaline, 

with strongly refractive contents, generally with a "gelatinous" 

outer wall of the epispore. - On herbaceous stems. 

Species examined: 

Trichometasphaeria dianthi (ROSTR.) MUNK (type species). 

pentamera sp. n. (MUNK 1956, in press). 

culmifida (KARSTEN) comb. n. (Leptosphaeria 

culmif'ida KARSTEN). 

sp., cfr. Leptosphaeria setulosa SACC. & 

ROUM. 

Massarina SACC. 

With a + distinct clypeus-stroma coalescent with the upper por 

tion of the peridium. Pseudothecia subspheric to flattened, in M. ebur 

nea hyaline in the bottom, uniform in the other species; towards the 

porus the inner portion of the peridium is differentiated into an 

opaque blackish, small-celled tis sue in the shape of a cone (in M. 

eburnea a flat disc). Asci clavate; interascicular tissue paraphysoid. 

Spores biseriate, transversally septate, hyaline, strongly refractive, 

constricted at the septa, outer epispore "gelatinous". - Corticolous. 


Massarina eburnea (TUL.) SACC. (type species). 

salicinicola REHM. 

micacea (KZE.) SACC. 

polymorpha (REHM) SACC. 

alni (OTTH) SACC. 

rubi (FUCK.) SACC. 

Pseudotrichia KIRSCHSTEIN 

Pseudothecia small, gregarious, immersed, vertically elongate. 

Peridium thin and soft; towards the porus the inner portion is 

differentiated as in Massarina) but the black tissue forms a cylinder 

instead of a cone. Asci cylindric-clavate; interascicular tissue paraphy 

soid. Spores biseriate, 2-, then 4-celled, strongly constricted at the 

primary septum, hyaline. - Hypersaprophytes, probably approach 

ing real mycoparasitism. 


Pseudotricha minor MUNK. 

20*

- 308 

Keissleriella v. H. (Syn.: Coenosphaeria MUNK). 

Pseudothecia immersed, but prominent, clypeate, Peridium smalIcelIed. 

Papilla outside and in the porus covered with dark brown bristles. 

Asci clavate; interascicular tissue paraphysoid. Spores 1-2-seriate, 

clavate, bicellular, hyaline. - Hypersaprophytes on rotten wood. 

(Dr. V. ARX has called my attention to the fact that my Coenosphaeria 

diaporthides (MUNK 1953 : 134) seems to be identical with 

Didymella sambuci REHM, which is the type species of Keissleriella 

v. H . My thanks are due to Dr. v. ARX for this valuable information). 


Keissleriella sambuci (REHM) v. H. 

REFERENCES 

Munk, A. (1953): The System of the Pyrenomycetes. 163 pp. 

Munk, A. (1956): Danish Pyrenomycetes. A preliminary Flora. About 550 

pp. (in press). 


L YCOGALA FLA VO·FUSCUM IN DENMARK 

By D. MULLER 

Until now the big but rather rare myxomycet Lycogala flavofuscum 

(Ehrenb.) Rost. has not been recorded from Denmark. It is 

neither mentioned in the monograph by RAUNKIÆR (1888) nor 

by ELLIOTT (1927) in his review of the Danish myxomycetes in the 

Botanical Museum of Copenhagen. 

In Sweden it is mentioned from 4 localities by FRIES (1899); from 

Norway no finds have been published. 

Fig. 1. Lycogala flavo-fuscum. Cluster of aethalia 

on A esculus hippocastanum. Elsinore, 18. 

July 1955. Phot. by D. MULLER. 

- 309-

- 311- 

The hypothallus had a yellow-ochraceus colour, a character men 

tioned by EHRENBERG (1818) but not since. 

CEJP (1952) who mentions that Lycogala flavo-fuscum has been 

found in 6 localities in Czechoslovakia, says that thi s myxomycet nor 

mally occurs on Acer. But the Danish finds, and also some of those 

stated by CEJP, are not from Acer. Re also mentions, that the preva 

lence is at the end of the summer or in the autumn. But in Denmark the 

aethalia ripen in July. The fruit bodies, which I gathered on the 18. 

July 1955, had ripened during the days from the 13. to the 18. July. 

The Danish finds are the following four, all from Zealand: 

1) Frederiksværk, the Arresodal Forest, on a stump. Spores 5.5- 

6.2 fÆ. July 1890. Leg. et det. E. ROSTRUP. At Laboratory of Plant 

Pathology, Royal Vetr. and Agric. College, Copenhagen. 

2) Tisvilde, on Aesculus hippocast,anum in the neighbourhood of 

the Sand-flight Monument. Spores 6.2-6.9 fÆ. 9. Sep. 1954. Leg. M. 

SKYTTE CHRISTIANSEN; det. M. P. CHRISTIANSEN. At Botanical Mu 

seum of Copenhagen. 

3) Copenhagen, Assistens Cemetery, on Fraxinus excelsior. Spores 

5.5 fÆ. 6. Aug. 1954. Leg. et det. M. P. CHRISTIANSEN. 

4) Elsinore, Marienlyst Alle, on Aesculus hippocastanum. Spores 

5.5 fÆ. 12. July 1953 and 18. July 1955. Leg. D. MULLER. At Labora 

tory of Plant Pathology, Royal Vetr. and Agric. College, Copenhagen.*) 

REFERENCES 

Cejp, K.: New finds of the rare Myxomycet, L y c o g a l a f l a v o - f u se 

u m, in this country. (Orig. in Czech.). - Ceska Mykologie 6: 

156-159, 1952. 

Ehrenberg, G.: Sylvae mycologicae Berolinensis. Dissert. Berlin 1818. 

Elliott, W.: Danish Myxomycetes. - Botan. Tidsskr. 39: 357-367, 1927. 

Fries, R.: Sveriges Myxomyceter. - Kungl. Vetensk.-Akad. Forhandl., 

Stockholm 1899: 215-246, 1899. 

Raunkiær, C.: Myxomycetes Daniae eller Danmarks Slim svampe. - Botan. 

Tidsskr. 17: 20-105, 1888. 

Rostafinski, J.: Sluzowce (Mycetozoa). Monografia. Paris 1874-76. 

Copenhagen, J anuary 1956. 

*) In 1956 the following two finds are recorded: 23. July on Acer pseudo 

platanus at the Vigeland Museum in Oslo, Norway, the first find in 

Norway. Leg. et det. M. P. CHRISTIANSEN. 31. July on Ulmus glabra 

at a height of 1,5 m by Kronborg, Elsinore. Leg. D. MULLER.

TWO LITTLE-KNOWN DANISH MUSHROOMS: 

BOLETUS FRAGRANS VITT. AND BOLETUS EDULIS VAR. 

CITRINUS PELTEREAU 

By F. H. MaLLER 

. The occurrences of our less common Danish Boletus species in 

Lolland, Falster and Mon, as well as in the vicinity of Vor 

dingborg (Zealand) are now fairly well established in detail. Never 

theless, the following three species have not yet been found in that 

area: Boletus cavipes OPAT., B. lignicola KALLENB., and B. placidus 

BoN., all of which are associated with particular trees or substrates 

and have been observed only once or a couple of times in Denmark. 

A great number of the other species are rare, or fairly rare, in the 

abovenamed southern region of Denmark, viz.: 

Boletus aeruginascens SECR. (syn. B. viscidus FR.); Lolland: 2 

localities; Falster: 3 localities. 

B. castaneus BULL. ex. FR.; Mon: 2 localities. 

B. cramesinus SECR. (syn. B. gentiZis QUEL., B. aurisporus sensu 

KALLENB.) ; Falster: 1 locality, 1 specimen. 

B. crocipodius LET. (syn. B. nigrescens RICH. & ROZE, B. rimosus 

sensu KALLENB.) ; Lolland: 2 localities. 

ties. 

B. cyanescens BULL. ex. FR.; Lolland: 1 locality; Falster 2 locali 

B. duriusculus KALCHBR. & SCHULZ.; Lolland: 1 locality. 

B. fechtneri VEL. (syn. B. aestivalis sensu KALLENB.); Falster: 3 

localities. 

B. flavidus FR. var. juranus QUEL.; Falster: 1 locality. 

- 312-

- 313- 

B. impolitus FR.; Lolland: 2 localities; Falster: 1 locality. 

B. lividus BULL. ex. FR.; Lolland: 2 localities; Falster: 1 locality. 

B. parasiticus BULL. ex. FR.; Lolland: 4 localities. 

B. pseudoscaber SECR., non KALLENB. (syn. B. porphyrosporus 

FR.) ; Falster: 1 locality. 

B. pulverulentus OPAT.; Lolland: 2 localities; Vintersbølle Forest 

at Vordingborg: 1 locality. 

B. purpureus FR. (syn. B. rhodoxanthus KROMBH.) ; Lolland: 2 

localities; Falster: 2 localities. 

Phylloporus rhodoxanthus (SCHW.) BRES.; Lolland: 2 localities; 

Vintersbolle Forest: 1 locality. 

Besides, two other species or varieties may deserve the designation 

"rare", viz. Boletus fragrans VITT. and B. edulis BULL. ex. FR. 

var. citrinus PeItereau. These two species are mentioned in greater 

detail in the folIowing. 

1. Boletus fragrans VITT. 1835. 

Syn.: Boletus aereus SECR. 1833; B. xanthoporus KROMBH. 1846. 

H a b i t a t: Lolland: Skejten at Fuglsang 6/VII 1920 under Quer 

cus. Falster: Hovedskov Forest at Tromnæs 9/IX 1934 under Fagu8 

and Quercus; Vålse Vesterskov Forest 15IVII 1937 under Quercus J' 

Nykobing Falster at the Vesterskov Pavilion under old, preserved 

Quercus 13/VIII 1952, 17 lIX 1953. 

C a p: 6-12 cm wide, pulvinate, repand, highly tomentose, umber 

brown or umber-olive, becoming purple-brown-spotted where touched 

or rubbed. 

T u b e s: lemon-yellow, sinuate-free, in the last stage fairly long; 

pore orifices sulphur-yellow, small, round. 

S t e m: 4-6 cm X 4--6 cm, ventricose, attenuated at the base, 

tomentose, apex not reticulate but more or less roughly warty-to 

mentose with yellow granules, everywhere yellow or, below the yellow 

apex, olive and usually spotte d with a purple-brown colour. 

F l e s h: compact, parti ally white in the young cap and stem, gene 

rally deep lemon-yellow above the tubes and near the cuticle of the

- 315- 

It is a familiar fact that Boletus impolitus is described with flesh 

which does not turn blu e when cut, but in Falster this species is 

occasionally found with thi s flesh reaction, and KUHNER & ROMAGNESI 

(Flore anal., p. 38, 1953) distinguish between B. jragrans and B. impolitus 

in the following way: 

Boletus jragrans. Chair et pores bleuissant legerement .... 

Spores 9.5-11 X 3.5--4 fÆ. 

Boletus impolitus. Chair bleuissant tres rarement un peu iL la 

coupe au moment de la recolte, en general immuable 

.... Spores (9)10-14(16) X 4-6 fÆ. 

The very dark coloured figures of B. impolitus in MICHAEL Fiihr. 

f. Pilzfr. nI. No. 282, 1927, are probably B. jragrans. BRESADOLA has 

given a bette r, though a little too paIe picture (lcon. Myc., pI. 926) of 

B. jragrans) which KONRAD & MAUBLANC (Agaricales II, p. 120, 1952), 

however, call a very dubious, too paIe figure. On the other hand, they 

refer to PELTEREAU's colour plate of B. jragrans in BULL. Soc. MYc. 

FR. 47, Atlas Pl. 45, as "tres bonne", but no doubt thi s one represents 

rather B. appendiculatus) because the stem above is reticulate. 

FRIES, who has not seen B. jragrans in nature, describes it very 

well, but he states the colour of the cap to be darker ("fusco-umbrino") 

than does the author of this paper, and according to FRIES the 

stem is "laevi", which, however, probably means "not reticulate", for 

the stem of B. impolitus is also described as "laevi". 

2. Boletus edulis BULL. ex FR. var. citrinus PELTEREAU 1931. 

Only a brief, incomplete description can be given, because the 

numerous (about 20) specimens had all expanded and were rather 

old. These fruit bodies had an appearance quite different from that of 

the typical Boletus edulis) but the reticulate stem and the spore size, 

together with the qualities of the flesh (mild, not turning blue) agreed 

only with those of the latter species. 

H a b i t a t: Zealand: Vintersbølle Forest, east of Vordingborg, 

under young beeches, in old leaves, immediately west of the gamekeeper's 

lodge at Blegen, 28/VlII 1921. 

C a p: 8-12 cm wide, pulvinate, in the last stage with acute, 

rising margin, beautifully lemon-coloured with fulvous parts, especiaIly 

at the centre of the capo

- 316- 

T u b e s: Sinuate-free, long, lemon-coloured (the first colour not 

seen) , in the last stage with an olive tinge. The pore orifices paIe 

yellow, later yellowish-green, small and round. 

S t e m: 8-11 X 2-3 cm, more or less fusiform, rather slender, 

downwards white, upwards Isabel coloured and white-reticulate like 

that of Boletus edulis. 

F l e s h: white, later yellow in the cap and sometimes also at 

the base of the stem; also a reddish tinge is seen in the thick portion 

of the cap, not turning blue, mild. 

S p o r e s: 13-15 X 3.75-4.5,u. 

Two water-colours executed in 1921 were sent to KALLENBACH, 

who did not recognize the fungus and wrote on the tables : "Bitte 

weiter beobachten" ! Unfortunately, the variety was not found again, 

and if KURNER & ROMAGNESI (1. c., p. 38) had not given the name 

of the fungus, the Danish find of it had not been published here. 

As I have not seen PELTEREAU'S description (in GILBERT: Les Bolets, 

1931), I cannot compare it with my notes of 1921. It would therefore 

be a good thing if the variety could be found again. 

Nykøbing Falster, February 9th, 1956.

POLYPORUS HISPIDUS (BULL.) FR. 

FUNNEN PÅ OLAND 

Av J. A. NANNFELDT 

Under en sommarvistelse på Oland 1953 antraffade jag den 3. aug. 

på en relativt ung ask i en tradgård vid vagen från Kalla odekyrka 

till Kalla hamn några ung a just utvaxande fruktkroppar av en stor 

Polyporus-art, som jag misstankte vara P. hispidus (BULL.) FR. (= 

Inonotus hispidus KARST.) . Vid återkomsten till Uppsala var Dr. 

SETH LUNDELL vanlig bekrafta bestamningens riktighet. Svampen 

sågs .ej på något mera trad i omgivningen. Tiden medgav dock ej en 

grundligare inventering. Ej heller såg jag den någon ytterligare gång 

under de 10 dagar jag var kvar på Oland och hade arten i tankarna. 

Fyndet har sitt givna intresse, i det att arten i Sverige tidigare 

antraffats blott på Gotland, dar den emellertid ar ganska utbredd och 

vaxer på bl. a. ask, alm och oxel (Sorbus intermedia). Det kan darfor 

måhanda fort jana påpekas, att Kalla liksom ovriga hamnar på norra 

Olands ostkust i gångna tider var en viktig overfartsort till och från 

Gotland. 

Det foreligger en litteraturuppgift från Skåne (Hackeberga, på 

bok, N. F. BUCHWALD; Friesia 1, p. 146, 1933) men denna uppgift 

avser P. cuticularis (BULL.) FR. (= Inonotus cuticularis KARST.), 

som ar en typisk bok-svamp och som bland danska mykologer gått 

under namnet hispidus) anda tills detta rattats av BUCHWALD (Naturhist. 

Tid. 5, p. 64, 1941). Xkta hispidus ar annu ej kand från Sveriges 

fastland. 

I Mellaneuropa ar arten en allman och skadlig parasit, bl. a. på 

appeltrad. Från Danmark synes efter BUCHWALD'S korrigering foreligga 

blott en uppgift, namligen om ett fynd från Falster på bjork 

(F. H. MØLLER, Friesia 2, p. 186, 1941). Från Norge foreligger en 

uppgift, naml. från Oslo på asp, och från Finland likaledes en, naml. 

från Runsala på ek. Jag har emellertid icke forsokt kontrollera om 

dessa uppgifter från Norge och Finland ara riktiga. 

- 31 7 -

- 318- 

SUMMARY 

Polyporus hispidus (BuH.) Fr. found on the Island of Oland (Sweden). 

The author reports a find of Polyporus hispidus (BULL.) FR. on a 

planted ash (Fraxinus excelsior) between "Kalla odekyrka" and 

"Kalla hamn" . The species was previously known in Sweden only 

from the Island of Gotland, where it is not rare. It is still unknown 

from the mainland of Sweden. 

Uppsala, February 1956.

ZUR VERBREITUNG 

DER AGARICALES IN DEN WALDERN DES 

SUDWESTLICHEN SCHWEDENS 

Von T. NATHORST-WINDAHL 

In den -' letzten zwanzig Jahren habe ich in den Landschaften 

BohusUin und Vastergotland im sUdwestlichen Schweden zahlreiche 

Exkursionen gemacht, um besonders die AgaricaZes-Arten zu studieren. 

Aber auch im nordlichen Halland, in DaIsIand und im mittleren 

Varmland bis an die norwegische Grenze hinauf habe ich an verschiedenen 

Orten Aufzeichnungen der Pilzflora gemacht und mir 

unbekannte Arten gesammeIt, eingehend beschrieben, photographiert 

oder gemalt, um sie spater zu bestimmen. Das sehr unterschiedliche 

Auftreten der Pilzarlen in den klimatisch stark voneinander abweichenden 

Beobachtungsjahren machte es notwendig, fortlaufende 

Untersuchungen im Gebiet zu machen, um ein getreues Bild dieser 

Flora zu erhalten. Ich habe keine Probeflachen verwendet, sondern 

mich mit Notierung aller gefundenen Arten und der begleitenden 

Flora begnUgt. 

Der Felsengrund dieses Gebietes besteht grosstenteils aus Gneis, 

welcher teilweise mit Moranensand, Kies und Geroll bedeckt ist. Die 

Humusschicht ist oft mit Sand und Lehm untermischt. An mehreren 

Stellen kommt das Urgebirge zum Vorschein. Besonders in Vastergotland 

kommt Sand- und Kalkstein vor, und in gewissen Teilen dieser 

Landschaft dominiert das bebaute Flachland. Bohuslan ist an der 

KUste reich an Felsen, Inseln, Scharen, tiefen Meerbusen, Mooren 

und Strandwiesen. Auf diesen Wiesen wachsen sehr oft PsaZZiota 

campestris und Lepiota excoriata. Die Berge dieser Landschaft sind 

nicht hoch (bis 225 m) und oft teilweise mit CaZZuna bedeckt. In den 

Landschaften DaIsIand und Varmland sind sie dagegen hoher (in 

- 319-

- 320- 

Varmland bis 700 m) und die Walder grosser. Die Seen sind zahlreich, 

und der Wanersee, der grosste See Schwedens, bedeckt eine Flache 

von 5568 qkm. Das Klima des sUdwestlichen Gebietes ist subatlantisch, 

die Bodenflliche erhebt sich gegen die norwegische Grenze, und 

das Klima ist dort mehr kontinental. Die mittlere Temperatur ist in 

Gotenburg + 7,7 0 C, in Karlstad am Wanersee + 5,9 0 C. Die jahrliche 

durchschnittliche Niederschlagsmenge variiert im Gebiet zwischen 

625 und 800 mm. 

Grosse Walder von einheitlichen Charakters sind in Bohuslan und 

Vastergotland kaum vorhanden, und deshalb wurde dort die Feststellung 

der okologischen Bindung der Agaricales an bestimmte Wald 

Gesellschaften sehr erschwert. Die Fruchtkorper vieler Pilze treten 

nur in einzelnen Jahren auf, das Myzel aber kann manchmal jahrelang 

steril im Boden verbleiben. 1m Herbst 1955 erschien Clitocybe 

(Hygrophoropsis) aurantiaca var. pallida Uberall in den Nadelwaldern 

in der Nahe von Gotenburg. Dieser Pilz ist dort frUher sehr selten 

beobachtet worden. In den Jahren 1941-1943 wurde Tricholoma 

co los sus an verschiedenen Orten im Gebiet beobachtet. Seitdem ist 

sie meines Wissens nur einmal dort wiedergefunden. FUr die Erscheinung 

der Fruchtkorper dUrfte wohl die Warme und Feuchtigkeit 

des Bodens und die Lage des Grundwasserstands massgebend 

sein. 

Die unterschiedliche Verbreitung vieler Agaricales entspricht in 

mancher Beziehung der Verbreitung bestimmter Waldverbande. 

Folgende Baume kommen im Gebiet Uberall vor: Pinus silvestris) 

Picea abies) Populus tremula) Betula pubescens) B. verrucosa und Sorbus 

aucuparia. Die Larix-Arten werden zuweilen kultiviert und Picea 

abies kommt an der KUste spontan nicht vor. Ulmus glabra) Malus 

silvestris) Sorbus intermedia) S. suecica) Tilia cordata und Acer platanoides 

kommen in Varmland zerstreut vor, sind aber sonst haufig. 

Fraxinus excelsior) Quercus robur und Corylus avellana haben ihre 

Nordgrenze im mittleren Varmland. Fagus silvatica ist sUdlich 

orientiert und in Varmland nicht spontan. Alnus incana ist dagegen 

nordlich orientiert und kommt im Gebiet hauptsachlich in Varmland 

vor. Es gibt grossere Bllitterpilze, die nur in den Waldern leben, ohne, 

wie es scheint, mit den Baumen dort Mykorrhiza zu bilden, z. B. Stropharia 

(Geophila) Hornemannii) Lepiota rhacodes) Clitocybe clavipes) 

Tricholoma (Rhodopaxillus) nudum und die meisten Cortinarien. Collybia 

maculata) die sonst in den Kiefernwaldern wachst, kann zufallig 

im Laubwald ganz Uberraschend erscheinen. Es lasst sich ja denken,

- 321- 

das s diese Art nicht Mykorrhizabildner ist. Nach verschiedenen Floren 

solI Boletus badius im N adelwald wachsen, im slidwestlichen 

Schweden kommt diese Art auch unter Laubbaumen, besonders 

Eichen, vor. Professor FABRITIUS BUCHWALD teilt mir mit, dass sie 

auch in Danemark unter Laubbaumen vorkommen kann. In Bohuslan 

und im nordlichen Halland habe ich niemals Hygrophorus erubescens 

und Stropharia (Geophi1a) Hornemannii gesehen, obwohl sie in den 

librigen Landschaften des Gebietes nicht selten sind. Dagegen scheint 

Russula sardonia viel haufiger an der Westkliste zu sein als in den 

librigen Teilen Schwedens. Einige Agarica1es} die ich im Gebiet nie 

beobachtet habe, wachsen im slidlichen Schweden, z. B. Boletus /loccopus} 

Lepiota Bucknal1ii} L. hystrix} Mycena crocata} P1uteus leoninus} 

Inocybe jurana} I. Patouillardii und Coprinus picaceus. 

Die folgenden Listen sind keineswegs volIstandig. Manchmal dauert 

es eine geraume Zeit, bis z. B. eine Cortinarius-Art in ihrer ganzen 

Variationsbreite erfasst wird, und ganz gewiss erlibrigt es, noch 

weitere Arten im Gebiet zu finden und manche Gegenden, besonders 

in Daisiand und Varmland, zu untersuchen. In den Listen, die alphabetisch 

angeordnet sind, habe ich nicht die Moglichkeit die Pilze umgebende 

Flora zu erIautern. Die Listen wlirden dann viel zu umfangreich 

werden. Die Autornamen mussten wegbleiben. Nur im Falle, das s 

cine Art nicht eindeutig ist, folgt hinter dem Artnamen "ss.", das 

heisst sensu oder im Sinne von, z. B. ss. Lge. Die Zahl hinter den 

Namen gibt die Haufigkeit an, wobei 4 die grosste Haufigkeit, 3 

ziemlich haufig, 2 ziemlich selten und 1 selten angibt. Der Buchstabe 

"s." hinter dem Artnamen bedeutet saurer Boden, "S." Sphagnum 

und "m.", dass der Pilz auch im Mischwald wachst; "a." bedeutet 

Alnus} "b." Betu1a} "c." Cory1us} "f." Fagus} "p." Popu1us tremu1a und 

"q." Quercus} unter welchen Baumen diese Pilze das Optimum ihrer 

Verbreitung haben. 

Kiefernwalder 

Boletus bovinus 4, edulis ssp. pinicola 

3, flavidus S. 1, luteus 4, 

piperatus 3, variegatus 4, 

Oollybia maculata 2, tenaeella ss. 

FAVRE 2, 

Oortinarius mucosus 3, obtusus 4, 

Gomphidius rutilus (viscidus) 1, 

Hygrophorus hypothejus 4, 

Lactarius deliciosus 3, helvus s. 4, 

mammosus 3, rufus 4, 

FRIESIA V 

Russula caerulea 3, sardonia 4, 

Tricholoma colossus 1, flavovirens 

(equestre) 2, imbricatum 3, 

portentosum 3. 

Nadelwalder 

A manita citrina 3, fulva 2, muscaria 

ssp. regalis 2, porphyria 3, 

spissa 2, vaginata 1, 

Armillaria (Olitocybe) mellea 3, 

A. (Biannularia) imperialis 1,

Boletus badius 2, calopus 2, felleus 

3, subtomentosus 3, 

Clitocybe Alexandri 2, C. (Hygrophoropsis) 

aurantiaca 4, brumalis 

ss. LGE 2, clavipes 3, 

diatreta 2 , dicolor ss. LGE 2, 

ditopoda 3, gilva ffi. 3, inversa 

ss. RICK. ffi. 3, Langei ffi. 2, 

metachroa 2, odora 3, pithyophila 

ss. LGE 3, squamulosa 

1, C. (Hygrophoropsis) umbonata 

s. 2, vibecina ss. RICK. 2, 

Collybia (Marasmius) acervata ss. 

F AVRE 2, C. butyracea 3, distorta 

1, esculenta ss. FAVRE 3, 

Cortinarius armeniacus 2, brunneus 

l , callisteus 4, camphoratus 

3, cinnamomeo-lutescens 3, 

cinnamomeus 4, collinitus ffi. 3, 

cumatilis 1, erythrinus ss. LGE 

2, fasciatus 3, gentilis 2, glaucopus 

2, malachius ss. LGE 2. 

obtusus 4, rapaceus ss. LGE 

1, sanguineus 3, scaurus ss. 

RICK. S . 2, semisanguineus 4, 

traganus 3, turmalis 2, violaceus 

2, 

Cystoderma amianthinum 3, carcharias 

3, 

Flammula (Dryophila) lenta 3, F. 

(Gymnopilus) penetrans 3, F. 

(Gymnopilus) picrea 2, F. 

(Dryophila) astragalina 2, F. 

(DryophiZa) scamba 2, F. 

(DryophiZa) spumosa 3, 

Gomphidius gZutinosus 4, roseus l , 

H ebeZoma mesophaeum 2, sinuosum 

ss. K. & M. 1, spoZiatum 2, 

strophosum 2, 

Hygrophorus agathosmus 3, camarophyllus 

2, erubescens 3, 

pustuZatus 4, olivaceoalbus ss. 

K. & R. 3, tephroZeucus 2, 

Hypholoma (Geophila) capnoides 

4, H. (Geophila) dispersum ss. 

K. & M. 2, H. (Geophila) radicosum 

ss. LGE l, 

Inocybe calamistrata 2, dulcamara 

ss. HElM l, lacera 4, sambucina 

1, 

Laccaria laccata 3, 

Lactarius camphoratus 3, deliciosus 

2, flexuosus 4, fuliginosus 

2, helvus b. s. 4, hepaticus l, 

hysginus 1, lignyotus 2, mitissimus 

2, repraesentaneus s. 2, 

rufus 4, scrobiculatus 2, trivialis 

3, voZemus 1, 

322 - 

Lentinus (Lentinellus) bisus 1, 

Lepiota rhacodes 3, 

Marasmius androsaceus 3, M. (Xeromp 

halina) fulvo bulbillosus 

2, impudicus l, perforans 4, 

putilZus 1, 

Mycena aZcaZina 3, amicta 2, cyanorrhiza 

1, elegans ss. SMITH 

1, epipterygia 4, flavoalba 2, 

galop oda 3, lactea 2, metata 

ss. ROMELL, LUNDELL, RICK., 

LGE, SMITH, 3, rosella 3, rubromarginata 

ss. SMITH 2, sanguinolenta 

4, vulgaris 3, zephirus 

3, 

Naucoria (Galerina) sideroides 1, 

Omphalia bisphaerigera 1, cyathiformis 

3, chrysophylla 2, O. 

(Xeromphalina) camp anella 3, 

Paxillus atrotomentosus 3, involutus 

4, panuoides 2, 

PhoZiota (Rozites) caperata 3, P. 

fZammans 3, P. (Galerina) 

marginata 3, 

Pleurotus (Panellus) mitis 4, P. 

(PZeurotellus) porrigens 3, 

Pluteus atromarginatus 1, 

Psalliota silvatica 2, 

Rhodophyllus hirtipes ss. K. & M. 

1, madidus 2, nitidus 2, vernus J 

non Ent. pascuum ss. BRES. 1, 

Russula aurata 1, azurea 2, consobrina 

1, decolorans 4, emetica 

3, integra 3, Mariae ss. J. 

SCHAFF. 1, obscura (vinosa) 

3, ochroleuca 4, paludosa (elatior) 

s. 3, puellaris 2, Queletii 

2, sanguinea 2, 

Stropharia (Psathyrella) Caput 

Medusae 1, S. (Geophila) Hornemannii 

(depilata) 2, 

Tricholoma aestuans 2, albobrunneum 

3, T. (M elanoleuca) cognatum 

1, T. decorum 3, T. inamoenum 

2, T. (Leucopaxillus) 

lentum 1, T. (Rhodopaxillus) 

nudum 3, T. rutiZans 3, saponaceum 

3, sejunctum 2, sudum 

1, vaccinum 3. 

Larchenbaume 

Boletus cavipes 1, Grevillei 3, viscidus 

1, 

Lactarius porninsis 1, 

Tricholoma psammopus 2.

Laub- und Mischwalder 

A manita citrina m. 4, excelsa 1, 

julva m. 4, inaurata (strangulata) 

m. 2, muscaria m. 4, 

pantherina m. 2, phalloides m. 

1, rubescens m. 4, vaginata m. 

2, virosa f . 2, 

Armillaria (Clitocybe) mellea m. 

3, A. (Collybia) mucida f. 2, 

Asterophora lycoperdoides m. 3, 

parasitica 1, 

Boletus appendiculatus 1, aurantiacus 

p. 3, badius m. 2, calopus 

f. m. 2, carpini c. p. 1, chrysent 

eron 4, cyanescens 2, edulis 

4, erythropus q. 3, lividus 

a. 1, luridus 1, parasiticus 2, 

reticulatus q. 2, rubellus (versicolor) 

s. a. q. 2, scaber b. 3, 

subtomentosus m. 3, B. (L eccinum) 

testaceo-scaber b. 3, 

Claudopus (Phyllotopsis) nidulans 

1, 

Clitocybe candicans ss. RICK. 2, 

C. cerussata 1, connata 2, 

fragrans 3, C. (Leucopaxillus) 

gigant ea 2, C. injundibulijormis 

m. 3, nebula1"is m. 3, odora 

m. 3, phyllophila ss. LGE 2, 

Clitopilus cretatus 1, hirneolus m. 

2, prunulus m. 3, 

Collybia butyracea m. 3, butyr. 

var. asema 4, cirrhata 2, cirrh. 

var. Cookei 2, dryophila 4, palustris 

(leucomyosotis) S. 3, 

platyphylla 3, plexipes 1, racemosa 

1) radicata f. 3, C. (Lyophyllum) 

1"ancida m. 2, C. 

tuberosa m. 3, velutipes 3, 

Coprinus atramentarius 3, disseminatus 

3, dilectus 1, extinctorius 

ss. RICK. 1, impatiens 

3, lagopus 2, micaceus 3, 

niveus 2, plicatilis 2, silvaticus 

(tardus KARST. ss. LGE) 2, 

Cortinarius acutus m. 2, alboviolaceus 

m . 2, anomalus b. m. 3, 

armillatus b. m. 4, balteatus 

ss. K. & M. 2, bibulus s. a. 1, 

bolaris q. 3, cinnamomeus m. 

4, cinnamomeo-lutescens m. 3, 

concinnus s. 1, croceocaeruleus 

ss. LGE 1, cyanopus m. 2, delibutus 

m. 2, elatior 3, emollitus 

f. 2, evernius m. s. 3, jlexipes 

m. 3, hemitrichus m. 3, hinnu 

Zeus q. 2, helvelloides a. 2, 

holophaeus f. 1, injractus m. 3, 

lucorum 1, multijormis ss. K. & 

- 323- 

M . m. 2, nemorensis ss. LGE 

f. 1, paleaceus m. s. 3, phoen'lceus 

ss. K. & M. 2, pholideus 

m. 4, psammocephalus ss. LGE 

1, rigidus ss. LGE (rigidus 

forma) 3, rubicundulus 1, sciophyllus 

2, scutulattts ss. K. & 

M. s. 2, subpurpurascens ss. 

RICK. 2, torvus f. 3, triumphans 

b . 2, 

Crepidotus Cesati m. 3, jragilis 

(autochtonus ss. LGE) 2, haustellaris 

2, Lundellii 3, mollis 2, 

variabilis 3, 

Cystoderma g1"anulosum m. 2, 

Flammula (Dryophila) alnicola 4, 

F. gummosa m. 2, 

H ebeloma longicaudum 2, sacchariolens 

3, 

Hygrophorus chrysodon 1, citrinus 

ss. LGE 1 , cossus 2, eburneus f. 

2, leucophaeus f. 1, subradiatus 

1, turundus s. 2, unguinosus 

2, 

Hypholoma (Drosophila) appendiculatum 

(hydrophilum auct. 

plur.) 3, H. (Drosophila) Candolleanum 

3, H. (Geophila) 

jasciculare 4, H. (Geophila) 

sublateritium 3, 

Inocybe asterospora q. 2, calospora 

s. 2, cincinnata 2, Cookei ss. 

LGE 2, decipientoides 3, descissa 

1, eutheles 2, jastigiata 4, 

geophylla 4, grammata ss. LGE 

2, hystrix f. 1, lacera 4, lanuginosa 

q. c. 2, maculata 2, microspora 

1, napipes q. 3, paludinella 

(trechispora ss. LGE) 

S . 1, perlata 1, petiginosa f . 2, 

pusio ss. LGE 1, umbrina q. 2, 

xanthomelaena 1, 

Laccaria laccata 4, tortilis 2, 

Lactarius aspideus 2 (Salix) ) blennius 

f. 4, camphoratus m. 3, 

chrysorrheus q . 2, juliginosu8 

m. 3, glyciosmus 3, insulsus 

ss. NEUH. q. 1, lilacinus a. 1, 

obscuratus a. 4, pallidus f. 2, 

piperatus 2, pyrogalus c. 3, 

quietus q. 4, serijluus (cimicarius) 

3, spinosulus b. 2, subdulcis 

f. 3, thejogalus 3, tormino8us 

b. 4, turpis s. b. ffi. 

4, uvidus s. m. 2, vellereus 2, 

vietus s. 3, violascens 1, volemus 

f. m. 2, 

Lentinus (Lentinellus) cochleatus 

2, suavissimus 3, 

Lepiota acutesquamosa 2, cristata 

3, 

21*

Marasmius alliaceus f. 2, confluens 

3, epiphyllus 3, foetidus c. 1, 

fuscopurpureus ss. LGE f. 3, 

languidus (candidus LGE) 3, 

peronatus 3, prasiosmus 1, ramealis 

4, recubans f. 1, ro tula 

4, scorodonius 3, splachnoides 

2, 

Mycena acicula 2, adoni:l 1, alcalina 

3, capillaris f. 3, citrinomarginata 

m. 1, corticola 3, fag etorum 

f. 2, fibula (Omphalia) 

4, M. filopes ss. K. & M. 2, 

flavescens m. 2, galericulata 

4, galopoda 4, haematopoda 3, 

inclinata q. 3, leptocephala ss. 

RICK. m. 3, niveipes 2, polygramma 

3, pterigena 3, pura 

3, rorida 2, sanguinolenta 4, 

M. (Omphalia) speirea 2, M. 

stylobates 2, vitilis ss. LGE 2, 

Naucoria (Alnicola) alnetorum a. 

2, N. (Agrocybe) arvalis 2, N. 

bohemica 2, carpophila f. 3, 

N. (Phaeocollybia) Christinae 

1, N. (Geophila) crobula 1, N. 

(Macrocystidia) cucumis m. 2, 

N. erinacea 3, escharoides ss. 

LGE a. 4, granulosa 1, reducta 

a. 1, scolecina FR. a. 2, subme 

Zinoides a. 2, N. (Gymnopilus) 

subsphaerospora 1, N. triscopoda 

2, 

Omphalia (Delicatula) integrella 

s. 3, O. oniscus ss. FAVRE S. 

1, picta 2, 

Panus conchatus b. 3, ringens b. 1, 

stipticus q. 4, 

Paxillus involutus m. 4, 

Pholiota (Dryophila) aurivella 2, 

P. (Rozites) caperata m. 3, 

P. destruens 2, P. (Agrocybe) 

erebia 2, P. (Conocybe) filaris 

1, P. (Dryophila) flammans 

m. 1, P. (Naucoria) muricata 

f. 1, P. (Dryophila) mutabilis 

4, P. (Agrocybe) praecox 3, 

P. (Hebeloma) radicosa f. 1, 

P. (Gymnopilus) spectabilis 

q. 3, P. (Dryophila) squarrosa 

3, 

Pleurotus (Geopetalum) applicatus 

2, P. (Geopetalum) atrocaeruleus 

3, P. dryinus (corticatus) 

1, P. Zignatilis 1, P. 

(Geopetalum) rhacodium 1. P. 

(Panellus) serotinus 3, P. 

(Lyophyllum) ulmarius 2, 

324 - 

Pluteolus (Bolbitius) aleuriatus 2, 

Pluteus cervinus 3, chrysophaeus 

1, Godeyi 1, lutescens 2, nanus 

3, pellitus f. 1, phlebophorus 

2) plautus 1, saZicinus 2, 

Psalliota arvensis 3, augusta 2, 

Langei 1, macrospora 2, silvatica 

m. 2, silvicola 1, vaporaria 

ss. MOLL. 1, 

Psathyrella (Drosophila) fibrillosa 

ss. RICK. 2, gracilis 3, microrrhiza 

ss. RICK. 1, multipedata 

2, noZi tangere a. 1, obtusata 

ss. LGE 2, papyracea ss. LGE 1, 

spadicea ss. RICK. 2, sphagnicola 

S. 1, 

Rhodophyllus apriZis 1, byssisedus 

2, carneoalbus s. 1, cetratus 

2, clypeatus 3, dysthales 1, 

euchrous 2, icterinus 3, minutus 

2, nidorosus S. 3, porphyrophaeus 

1, rhodocyZix s. 1, 

rhodopoZius 2, staurosporus 4, 

undatus 1, versatilis 1, 

Russula adusta m. 3, aeruginea b. 

4, albonigra m. 1, alutacea m. 

2, claroflava s., b. 3, cyanoxantha 

3, delica 3, emetica f. 

fagicola 3, exalbicans b. 2, 

farinipes 1, fellea f. 3, foetens 

3, fragilis s. 3, grisea f. 1, 

heterophylla 1, integra m. 3, 

lauroceraci 2, lepida f. 2, lilacea 

3, Lundellii b. 1, lutea 2, 

nigricans 4, nitida s. 3, ochroleuca 

4, pectinata 2, pseudointegra 

q. 1, solaris f. 1, sororia 

q. 2, Velenovskyi b. 2, versicolor 

b. 2, vesca 3, virescens 

1, xerampelina m. 3, 

Schizophyllum commune 2, 

Stropharia (Geophila) squamosa 2, 

Tricholoma album 2, T. (Melanoleuca) 

cognatum 1, T. columbetta 

l, flavobrunneum b. 3, 

T. (Melanoleuca) grammopodium 

1, T. (Rhodopaxillus) 

irinum 1, T. (Rhodopaxillus) 

nudum 3, T. (Rhodopaxillus) 

personatum 1, T. scalpturatum 

2, sulphureum 2, terreum 2, 

ustale f. 2, virgatum f. 3, 

Tubaria (Naucoria) autochthona 1, 

conspersa ss. K. & R. 3, furfuracea 

ss. ROMAGN. 1, pellucida 

ss. RICK. 2. 

Gotenburg, Februar 1956.

YELLOW PHYLLODE SPOT 

AND DROP OF ACACIA ARMATA R. BR. 

(SEPTORIA ACACIAE SP. NOV.) 

By PAUL NEERGAARD 

Phytopathological Service of Denmark 

In October 1954 a serious phyllode spot and drop was encountered 

in a consignment of Acacia armata R. BR. (A. paradoxa DC.), subjected 

to inspection for export at the Phytopathological Service of Denmark. 

The disease seems not previously to have been reported in literature. 

The following description of the disease and its probable causal agent 

is based upon examination of plants kindly placed at my disposal by 

the exporter. 

ECONOMIC SIGNIFICANCE 

In Denmark the host, Acacia armata has been cultured for some 

years in a few nurseries as a pot culture under glass. During 1954 and 

1955 a considerable number of plant s in the nursery of the exporter 

was so severely attacked, that they had to be discarded, and the rest 

part of the culture, comprising several thousand plants, were affected 

to such a degree that their commercial value was reduced by 

about 50 per cent. The losses due to the disease during these years in 

the said nursery are estimated at several thousand kr. Essentially on 

account of the disease it has been decided to abandon the culture. 

SYMPTOMS 

Small chlorotic spots appear, scattered irregularly on the phyllode. 

At first they are almost circular, later more irregular, sometimes finally 

necrotic at the center. The spots are frequently less than 1 mm 

in diameter, but the chlorotic areas commonly coalesce to affect almost 

the entire phylIode (Fig. 1). 

- 325-

- 326 -

Fig. 2. Plants of Acacia armata, attacked by the yellow phyllode spot and drop. The two plants at left severely, 

the two plants at right slightly affected. 

., 

w 

t--:) 

-J

FRIESIA V, 1956 PLATE III 

Figs. 3-8. Septoria acaciae sp. nov. 

a Fig. 3 b 

Fig. 4 Fig. 5 

Fig. 6 Fig. 7 

a Fig. 8 b 

Fig. 3, a . Epidermis of a phyllode; b. Texture of hyphae developed below 

the epidermis shown in a. - Fig. 4. Pycnidium forme d around and below 

a stoma. - Fig. 5. Transverse section of a pycnidium. - Fig. 6. Sporulating 

pycnidial structures on m a lt extract agar. - Fig. 7. Sporulating tissues 

on malt extract agar. - Fig. 8. Conidia; a) from a phyllode of A cacia 

anl1,ata,' b) from culture on malt extract agar. - Figs. 3- 5 and 7- 8 

about X 250; Fig. 6 about X 100.

- 329- 

Fig. 9. Primordium of a pycnidium of Septoria acaciae) developed 

around a stoma. 

c o n i d i a (Fig. 8) filiform to subcylindrical, rounded at both 

ends, straight, seldom slightly curved, frequently with one indistinct 

medial septum, hyaline, 8-36 X 1-2.5 f1 (see table 1), issuing in 

white to rosy masses. 

On phyllodes of Acacia armata R. BR. (A. paradoxa DC.), October 

1954, Sept. 1955, January 1956, in North Zealand, Denmark. 

C u l t u r e on malt extract agar. Aerial mycelium dense, velvety, 

white to slightly grey, submerged mycelium dark olive green to almost 

black, producing pycnidial structures and sporulating profusely. Diameter 

of 5 days old culture at room temperature : 8-9 mm. 

TABLE 1 

Spore measurements in f1 for Septoria acaciae sp. nov. 

from Acacia armata. 

Medium Length Width lAVerage length 

(25 conidia) 

Leaf 10- 22 1- 1.5 17.9 

Malt extract agar 8- 36.5 1.5- 2.5 19.2

330 - 

Fig. 10. Pyenidium of L eptothY1'ium sp. on Acacia armata. 

DIAGNOSE LATINE 

Septoria acacia sp. nov. 

M a e u l i s amphigenis, subrotundatis irregularibusque, eonfluentibus, 

chlorinis. 

p y c n i d i i s amphigenis, immersis, dispersis vel subgregariis, subglobosis 

vel lentieularibus, glabris, atro-brunneis, 60- 110 Il diam., eontextu 

pseudoparenehymatieo. 

C o n i d i o P h o r i s vix ullIs. 

C o n i d i i s filiformibus vel subeylindraeeis, reetis vel leviter eurvulis, 

utrinque rotundatis, eontinuis vel indistinete-1-septatis, hyalinis, 8- 36X 

1-2,5fl. 

Hab. in phyllodiis vivis vel emortuis-dejeetis Acaciae armatae R. BR., 

Oet. 1954, Sept. 1955, Jan. 1956. Selandia, Daniae. 

Typus in Kgl. Veterinær- & Landbohøjskole, Plantepatologisk Laboratorium, 

Hafnia, Dania, eultura ex typo in Centraalbureau voor Sehimmeleultures, 

Baarn, Hollandia, depos. 

CYCLE OF TRE DISEASE AND CONTROL 

The fungus seems not to develop pycnidia in the affected, but 

still living tissues of the phyllodes on the plant. In the dead tissues 

of fallen phyllodes, however, the pycnidia are formed profusely, and 

from these the conidia have ample opportunities of reaching and

- 331 - 

infecting healthy phyllodes. Consequently, in order to control the 

disease, it is imperative to remove all phyllodes as soon as they have 

been dropped, and still better, to remove cautiously all infected plants. 

Furthermore, it is suggested repeatedly to spray the plants with fungicides. 

This has been practised in the said nursery by spraying with 

% % Captan, but the treatment caused a heavy phylIode drop and 

no further experiments were carried out. The whole lot of plants, 

however, were here entirely affected by the fungus so that no or liUle 

protection from the spraying could be obtained anymore. Only in the 

case of healthy plants, protection with fungicides can be expected to 

work efficiently. 

SUMMARY 

A serious phylIode spot and drop on Acacia armata R. BR. was 

detected in October 1954. The disease caused a loss of several 

thousand kr. in one Danish nursery. Septoria acaciae sp. nov. was 

isolated, partly from the diseased tissues of phyllodes taken from the 

plants, partly from pycnidia developed in dead, fallen phyllodes. 

Description and Latin diagnosis are given and measures for control 

suggested. 


- 333- 

experiment found that more than one-third of the isolates (mouids 

and bacteria) had inhibitory effects on the annosus mycelium in 

vitrO J one of them being Trichoderma viride J which was inhibitory 

at 22° C, but was itself overgrown by P. annosus at 12° C. 

On the other hand, as far as the author knows, no investigations 

have been made of the antagonistic effect of soil-borne actinomycetes 

on Polyporus annosus. In view of the great importance of these 

microorganisms as producers of antibiotics it was therefore natural 

to go into this question in greater detail, and in the folIowing a brief 

account is given of som e preliminary investigations made by the 

author on actinomycetes isolated from woodland soil. 

EXPERIMENTAL METHODS AND RESULTS 

1. Isolation of actinomycetes from woodland soil. 

In the period May-October 1954 a number of soil samples were 

taken from two regions in Jægersborg Hegn Forest, Zealand. 

a. Older stands of Norwegian spruce (Picea abies) (forestal 

sample plot, former name of the region: Husarsletten) . The soH 

samples were taken under sterile conditions at a depth of 5-20 cm 

underneath the unde compose d layer of needles. pH of the samples 

(measured electrometrically): 4.6-4.8. In the region were found 

numerous fruit bodies of Polyporus annosus FR., which according to 

the Danish Forest Experiment Station caused considerable damage. 

b. Deciduous forest, consisting preponderantly of beech (Fagus 

silvatioa) J immediately west of the abovenamed Norwegian spruce 

stand. The soil samples were taken under sterile conditions at a depth 

of 5-20 cm underneath the unde compose d layer of leaves. pH of the 

samples (measured electrometrically): 5.9-6.2. Attacks by Polyporus 

annosus could not be demonstrated. 

Spreading on glucose-casein-agar (pH 6.8) was done according 

to the conventional microbiological technique immediately af ter the 

samples were taken. All the colonies produced were investigated, 

but only actinomycetes were isolated. Many of these isolates proved 

to be replicates of the same species. Altogether, 18 different species 

were found, out of which 15 had to be referred to Streptomyces J 2 to 

Micromonospora and 1 to Nocardia. 

2. I nteraction experiments in agar media. 

The isolated actinomycetes were then tef.;ted for antagonistic 

effect in vitro on the mycelium of Polyporus anno sus FR. Petri

- 334- 

dishes with 2% potato-glucose-agar (pH 6.9) were first inoculated 

with Polyporus annosus in one side, after which they were left to 

stand for 3 to 5 days at 25° C, because the fungus grows rather 

slowly. Only when the growth of the fungus was in good progress 

were the dishes inoculated with the actinomycete - at a fair distance 

from Polyporus annosus. 

Fig. 1. Interaction between Polyporus annosus FR. and Streptomyces sp. ; 

some colonies (clear) of Nocardia sp. are seen in the left-hand side of the 

left-hand Petri dish. The antagonistic activity of the actinomycetes takes 

place in a distinct zone around the antagonist, into which the hyphae of 

the fungus do not penetrate. 

Of the 18 isolated actinomycetes, altogether 8 species, including 

Nocardia sp., displayed a pronounced antagonistic activity, distinct 

inhibitory zones being produced af ter 3 to 7 days (Fig. 1). Five of the 

antagonists originated from deciduous forest soil, the other three 

from coniferous forest soil. 

3. Interaction experiments in sterilized soil. 

On one of the abovenamed antagonistic actinomycetes which had 

shown the strongest activity (Streptomyces sp.) a number of experiments 

were made in 1954-55 in order to investigate whether the 

inhibitory activity ascertained in agar media (in vi tro) could also be 

demonstrated in soil experiments. 

In the tests, larger glass dishes with close-fitting lids were used. 

To each dish 75 g air-dried garden soil (pH 7.3) and 20% water were 

added, after which the dishes were autoclaved at 120° C for 3 hours. 

They were then inoculated with suspensions of spores of the above-

- 335- 

mentioned Streptomyces in sterile water. Some dishes, the soil of 

which was not inoculated with Streptomyces spores, were used as 

controis. 

The following tests on Polyporus annosus were then made on two 

- slightly different - methods. 

1. After 7 days, a slide covered on one side with nutrient agar 

and overgrown with mycelium of Polyporus annosus was placed on 

the soil surface under sterile conditions, with the covered side down, 

in direct contact with the soil. 

The slide cultures were produced in the following way: A sterile 

slide was placed on the bottom of a Petri dish and then covered with 

a thin layer of nutrient agar; next, the dish was inoculated with 

Polyporus annosus. When the agar was overgrown with the mycelium 

the slide was cautiously cut out of the culture and placed in a dish 

with soil, as described above. Also in each of the control dishes, 

which were not inoculated with Streptomyces) a slide with a culture 

of Polyporus annosus was placed. All the tests were made at 25° C. 

Af ter 7 days, the results of the tests were made up. Polyporu8 

annosus in the control dishes then appeared to have spread from the 

edges of the slides over the soil surface as a fine cobweb-like mycelium, 

while in the dishes which had been inoculated with Streptomyces 

no growth of the mycelium from the slides on to the soil could be 

a b 

Fig. 2. Interaction experiments in soil between Polyporus annosus FR. and 

Streptomyces sp. a. Growth of Polyporus annosus from 3 inocula (agar 

dises) in sterile soil. b. Antagonistic activity. No growth of Polyporus 

annosus in soil previously inoculated with a spore suspension of Streptomyces 

sp. Around the three agar discs the growth of Streptomyces sp. 

is seen as tiny white dots.

- 336- 

ascertained. The growth of the actinomycetes could be demonstrated 

as numerous, tiny white dots all over the soil surface. 

2. Instead of slide cultures, agar discs overgrown with Polyporus 

annosus were used as inocula. The discs were 10 mm in diameter and 

had been cut from young cultures in Petri dishes. Otherwise, the 

experiment was performed in a manner similar to that described 

under 1. 

AIso in thi s test series, a distinct inhibition of the mycelium of 

Polyporus annosus could be demonstrated in the dishes which had 

been inoculated with spores of Streptomyces sp. 7 days before, while 

there was no inhibition at all in the control dishes (Fig. 2 a & b). 

DISCUSSION 

Great importance is attache d to the question whether the antagonistic 

microorganisms of which it is known that they form strong 

antifungal antibiotics in synthetic media, are also capable of producing 

such substances in unsterilized soil. WRIGHT (1954) has investigated 

this question with a view to Trichoderma viride) which produces in 

vitro two antifungal substances, gliotoxin and viridin. She found that 

gliotoxin was also produced in acid heath soil, when this soil was 

amended with accessibIe carbon compounds (e. g. 1 % dried clover), 

but not in garden soil, unless its pH was changed at the same time 

towards higher acidity by the addition of acid. Furthermore, she was 

of the opinion that the increase in the production of antibiotics which 

was demonstrable after autoclaving of the green-manured soil must 

be ascribed to the autoclaving having increased the quantity of 

accessibIe N and C, rather than having removed competitive microorganisms. 

The possibility of controIling soil-borne plant diseases by inoculation 

of the soil with suitable antagonistic microorganisms has been 

discussed, for instance, by NISSEN (1953) and BRIAN (1954), and 

they both point out that present experience tends to indicate that 

control by thi s means would give positive results only when large 

quantities of a suitable medium (green manure) are added together 

with the antagonist, seeing that it is very difficult for a foreign 

microorganism to gain a foothold in the well-balance d environment 

constituted by the soil microflora. On the other hand, there is much to 

indicate that it will be possibIe by suitable green-manuring alone to 

stimulate the antagonistic fraction of the natural micropopulation

- 337- 

or to give the latter a favourable trend so that infestations originating 

from soil-borne microorganisms are thereby controlled or entirely 

prevented. 

It would be natural to correlate WRIGHT'S demonstration of antibiotic-production 

in natural soil (1954) with the studies on fungistasis 

carried out by DOBBS & HrNSON (1953) and JEFFERYS & HEMMING 

(1953). According to these investigations, fungistasis having the 

effect that fungal spores do not germinate, has a universal occurrence 

in soil. The fact that it is very difficult to demonstrate antibiotics in 

the soil is, in the first place, presumably due to the considerable 

adsorption power of the soil, but mayaIso be ascribed to instability 

of the substances and to their microbial decomposition (JEFFERYS 

1952, NISSEN 1954). 

On the part played by the actinomycetes as soil-borne antagonists 

to plant diseases a few positive accounts are available. As early as in 

1927 MILLARD & TAYLOR demonstrated that potato scab (caused by 

Streptomyces scabies) could be effectively controlled by the application 

of green manure, and they assumed that this effect of green 

manure was due to microorganisms. LoCHHEAD & LANDERKIN have 

rather recently (1949) established the rightness of this assumption, 

having managed to isolate from the potato rhizosphere in greenmanured 

soil actinomycetes which were antagonistic to the potato 

scab parasite. Promising are also the results of MEREDITH'S experiments 

(1946) with the control of a banana disease (Fusarium oxysporum 

cubense) by inoculation of the soil with actinomycetes. Quite 

recently (1955) it has been demonstrated at Rothamsted Experimental 

Station that the activity of the actinomycetes on H elminthosporium 

sativum) which is the cause of foot-rot of barley and wheat, is antibiotic 

and not mere ly competitive. 

In view of the wide-spread occurrence of antagonistic actinomycetes 

(LANDERKIN et al. 1950, ROUATT et al. 1951) and the ever 

increasing number of antibiotics which it has been possibIe to isolate 

from actinomycetes, it is certainly justifiable to suppose that these 

microorganisms are able to make their antagonistic activities felt in a 

high degree in nature. As, moreover, the actinomycetes belong to 

the zymogenic fraction of the soil micro-flora, i. e. the fraction which, 

in particular, enters into function when organic matter is added to 

the soH, there seem to be rather good chances of shifting, by means 

of green manuring, the microbial balance in a direction adverse to 

soil-borne pathogens. 

FRIESIA V 22

- 338- 

CONCLUSIONS 

Previous investigations, made particularly by Swedish workers, 

have proved that cerlain mould fungi, especially Penicillium species, 

exert an antibiotic activity on Polyporus annosus FR. By the experiments 

made here it has been demonstrated - and, as far as the 

author knows, for the first time - that also actinomycetes, not only 

in vitro but also in sterilized soil, may have a strong antibiotic effect 

on the mycelium of Polyporus annosus. Indeed, such antagonistic 

actinomycetes seem to be rather common in woodland soils. These 

results are bound to encourage a closer investigation to find out 

whether it will be possible, by the addition of suitable green manures, 

to stimulate the natural actinomycete flora in forests infested with 

Polyporus annosus) thereby possibly inhibiting the spread of this 

highly injurious pathogenic fungus. 

SUMMARY 

Of 18 actinomycetes species isolated from Danish woodland soil, 

8 proved to be antagonistic to Polyporus annosus FR. in vitro. One 

species (Streptomyces sp.) showed a pronounced inhibitory effect 

on this fungus in experiments with sterilized soil. It is discussed 

whether it will be possibie to control Polyporus annosus biologically 

by green manuring, so that the actinomycete flora of the soil is 

stimulated or shifted in a direction adverse to the fungus. As such 

possibility appears to be reasonable, it is proposed to proceed with 

experiments of thi s kind. 


This investigation has received financial support by a grant from 

The Danish State General Research Foundation to Professor N. FABRITIUS 

BUCHWALD, The Royal Veterinary and Agricultural College, Copenhagen. 

I am indebted to Professor N. FABRITIUS BUCHWALD and to Dr. H. L. 

JENSEN, the Stat e Laboratory of Plant Culture, Lyngby, for valuable 

discussions of these problems. 

REFERENCES 

Benedict, R. G.: Antibiotics produced by actinomycetes. - Botan. Review 

19: 229-320. 1953. 

Bjorkman, E.: Soil Antibiotics acting against the root-rot fungus (P o l yp 

o r u s a n n o s u s Fr.). - Physiol. Plant. 1: 1-10. 1949.

- 339- 

Brian, P. W.: The use of antibiotics for control of plant diseases caused by 

bacteria and fungi. - Jour. Appl. Bacteriology 17: 142-151. 

1954. 

Dobbs, C. G. & W. H. Hinson: A widespread fungistasis in soils. - Nature 

172: 197-199. 1953. 

Enebo, L.: Experiments with Claviformin as an antibiotic against P o l y 

p o r u s a n n o s u s Fr. - Physiol. Plant. 2: 56-60. 1949. 

Jefferys, E. G.: The stability of antibiotics in soils. - Journ. Gen. Microbiol. 

7: 295-312. 1952. 

Jefferys, E. G. & H. G. Hemming: Fungistasis in soils. - Nature 172: 

872-873. 1953. 

Landerkin, G. B. et al.: A study of the antibiotic activity of actinomycetes 

from soils of N orther Canada. - Canad. Jour. Res. C. 28: 690- 

698. 1950. 

Lochhead, A. G. & G. B. Landerkin: Aspects of antagonisms between 

microorganisms in soil. - Plant and Soil 1: 271-276. 1949. 

Meredith, C. H.: Soil actinomycetes applied to banana plants in the fieid. 

- Phytopathology 36: 983-987. 1946. 

Millard, W. A. & C. B. Taylor: Antagonisms of microorganisms as the 

controIling factor in the inhibition of scab by green manuring. 

- Ann. Appl. Biol. 14: 202-216. 1927. 

Nissen, T. Vincents: Om mikrobiel antagonisme og om antibiotica i jord 

bunden og mulighederne for praktisk anvendelse af disse fak 

torer. - Tidsskr. Planteavl 56: 633-659. 1953. 

: Effects of antibiotics on carbon dioxide produetion in soil. - 

Nature 174: 226-227. 1954. 

: Actinomycetes antagonistic to P o l y p o r u s a n n o s u s Fr. 

- Experientia 12: 229-230. 1956. 

Rennerfelt, E.: The effect of soil organisms on the development of 

P o l y p o r u s a n n o s u s Fr., the root rot fungus. - Oikos 1: 

65-78. 1949. 

Rennerfelt, E. & Sheila K. Paris: Some physiological and ecological experi 

ments with P o l y p o r u s a n n o s u s Fr. - Oikos 4: 58-76. 

1952-53. 

Rouatt, J. W., M. Lechevalier & S. A. Waksman: Distribution . of anta 

gonistic properties among actinomycetes isolated from different 

soils. - Antibiotics & Chemotherapy 1: 185-192. 1951. 

Wright) Joyce M.: The production of antibiotics in soil. I. Produetion of 

gliotoxin by T r i c h o d e r m a v i r i d e. - Ann. Appl. Biol. 41: 

280-289. 1954. 

Rothamsted Experimental Station: Report for 1954. - Nature 176: 1253- 

1254. 1955. 

Copenhagen, February 1956. 

22*

BOLETINUS ASIATICUS SING. I FINLAND 

A v WOLMAR NYBERG 

Bland de svampar som jag for flere år sedan overHimnade till 

Helsingfors universitets botaniske museum i exsiccat fanns aven 

Boletinus cavipes Opat. De voro tagna i en skogsdunge av någon 

gång i tiden planterade Iarktrad, som uppenbarligan spritt sig ut 

over sitt egentliga område, i vilket åter talriKa representanter for 

våra vanliga skogstrad trangt in. I dess centrum voro traden gamla, 

många redan doda. De forsta f ynden gjordes i aug. 1930 och under 

åren 1931, 1932 och 1933 hade jag tillfalle att fortsattningsvis finna 

svampen dar. Jag hade tyckt mig att det fanns två typer av arten. 

Den ena av dem, som jag for mig sjalv kallade var. karmin skilde 

genom vakkert karminrod hattfarg, morkare sporer och något storra 

dimensioner på dem från den mera i brunt gående formen. I "Karstenia", 

vår inhemska svampjournal, konstaterar numera professor 

R. TUOMIKOSKI (rattare redan 1953 i "Karstenia" II) att denna mera 

karminfargade svamp ar Boletinus asiaticus Sing. och att en så sall· 

synt framling - får man val saga - funnit vagen till oss i Finland 

har ju ett visst interesse. 

SUMMARY 

Boletinus asiaticus Sing. found in Finland. 

A short note is given about finds of the rare Boletinus asiaticus 

Sing. in Finland. 

REFERENCES 

Nyberg, w.: Några i Borgå och dess omnejd funna sallsyntare svamp-arter. 

- Memor. Soc. F. Fl. Fenn. 10: 20-23. 1934. 

Tuomikoski, R.: Boletinus asiaticus Sing. in FinnIand gefunden.- Karstenia 

2:42. 1953. 

Helsinki, Januar 1956. 

- 340-

INFECTIO;N OF BARLEY BY LOOSE SMUT, 

USTILAGO NUDA (JENS.) ROSTR. 

By P. NORUP PEDERSEN 

Department of Plant Culture, Royal Veterinary and 

Agricultural College, Copenhagen. 

INTRODUCTION 

Since MADDOX (1895, 1897) discovered that barley could only be 

infected by Ustilago nuda through the flowers, other workers have 

observed mycelium of this fungus in the barley seed at various stages 

of its development. Thus HECKE (1905) found hyphae in the scutellum 

and hypocotyl of germinating seeds, and BREFELD (1905) noted that 

almost all the loose smut spores located on the stigma had germinated 

and the hyphae penetrated to the style tissue, but he was unable to 

follow the course of the fine mycelium filaments further. Later 

BREFELD (1907) reported hyphae to be present in both embryo and 

endosperm. FALCK (1908) like-wise discovered the presence of hyphae 

in the scuteHum, the embryo and the tissues immediately surrounding. 

BROILI & SCHIKORRA (1913) examined germinated as well as dormant 

seeds and found an abundance of hyphae in the scutellum, and observed 

hyphae also in various parts of the embryo and hypocotyl as well 

as in leaf and bud primordia. A few years later LANG (1917) published 

the results of his investigations on the stages of infection by U. nuda 

in barley. His results which will be discussed later are briefly as follows: 

The spores germinate on the styles and follow the path of the 

pollen tube to the integuments which they follow to the embryo, ar 

they germinate on the surface of the ovary, penetrate the ovary wall 

and the integuments and continue between the latter and the nucellar 

epidermis till they reach the embryo. 

- 341-

- 342- 

In seeds studied 12 days after inoculation RUTTLE (1934) found 

hyphae in the pericarp, in the crushed nucellus, in the aleuron and 

endosperm as well as in the scutellum and hypocotyl. The pericarp 

was invaded at various points along the ovary wall, especiaIly in the 

brus h end. In latter years, recognition of the faet that hyphae are 

present in the scutellum has been useful in the development of 

methods to demonstrate loose smut infection in eertified seed barley. 

It appears from this short review that most investigations coineide 

with regard to most points of the infection biology mentioned, but that 

there is some disagreement as to the aetual place from which the 

penetration hyphae start. The present investigation is centered on 

this' problem. 

MATERlALS AND METHODS 

In the summer of 1950, 400 heads of Rigel barley were inoeulated 

according to the method of SHANDS (1940). Inoculation was performed 

immediately af ter flowering. From the date of inoculation till shortly 

before ripening, 10 heads were fixed daily. NAVASHIN'S fluid (Cromie 

acid and formalin) was used for fixation, and the specimens were kept 

in 70 pct alcohol. During the winter of 1950-51 the fixed material 

was imbedded in paraffin wax, sectioned by microtome (15 f.l), stained 

with safranin-lightgreen, mounted in Canada balsam, and examined. 

From this treatment very good slides resulted, the plasma-filled 

hyphae appearing a vivi d red, which showed up well against the green 

cells of the host. 

RESULTS 

As early as 24 hours after inoculation many spores have fully 

developed promycelia, and some have developed even further, having 

long filaments of mycelium twisted around the spores (Fig. 1). The 

spores do not seem to germinate better on the styles than on the 

ovary. A count made on flowers with many germinated spores, 

shows a germination percentage of about 60 in both areas. Although 

there are large numbers of germinating spores on the styles, only one 

instance is noted in whieh the promycelium has entered the style tissue, 

whereas several cases of infection are observed on the surface of 

the ovary. In these latter cases appressoria-like swellings ean be observed, 

from which vigorous pointe d hyphae have entered the cells 

immediately below (Figs. 2-4). Af ter 48 hours, the first hyphae have 

penetrated the epidermal eell layer and a few the subepidermal one

- 343 - 

Fig. 4. Spore with promycelium, appressorium, and penetration hypha 24 

hours after inoculation. - X 900. 

Fig. 5. Infection hypha which has penetrated the epidermis and entered 

the subepidermis. Note smaller diameter of hypha at point of penetration. 

48 hours af ter inoculation. - X 1300. 

Fig. 6. Infection hypha running straight from Uie epidermis to the integuments. 

5 days af ter inoculation. - X 225. 

Fig. 7. Infection hyphae running parallel to each other. - X 400. 

Fig. 8. Hyphae in the collapsed integuments. - X 1300. 

Fig. 9. Hyphae in the integuments near the base of the embryo. - X 330 . 

Fig. 10. Hyphae in the scutellum and hypocotyl. 24 days af ter 

inoculation. - X 90. 

. i 

i 

j

- 344- 

(Fig. 5), and new infections, which seem to have been established 

within the preceding 24 hours, are observed. In the next few days the 

infection develops further, and additional new infections are noted. 

After 5 days the first hyphae have reached the integuments (Fig. 6), 

but the greater part of the hyphae, however, have not penetrated this 

far. Having reached the integumental layers, the hyp ha e branch 

off and proceed through these toward the embryo (F'igs. 8-9). On the 

eighteenth day the first hyphae are found in the scutellum, and a few 

days later hyphae are plentiful in the embryo (Fig. 10). 

The promycelium and the hyphae located outside the pericarp, with 

exception of the appressoria, have a much smaller diameter than the 

hyphae found within the pericarp, and they are only stained very 

lightly by the method used. Partly for thi s reason it was impossible 

to ascertain whether the appressoria had been formed by swelling of 

promycelia or by swelling of hyphae originating from the latter. In 

every instance of infection observed, the infection hyphae have penetrated 

the cell wall near the centre of the wall. The pore thus made 

by the infection hyphae is always smaller than the average diameter 

of the hyphae af ter penetration (Figs. 3-4). The oldest parts of the 

hyphae from early infections are often emptied of protoplasm. It 

is highly characteristic of hyphae from early infections that they 

gro w directly towards the integuments. In several cases hyphae were 

seen to run parallel to one another toward the integuments from 

two closely situated foci of infection (Fig. 7). Having entered the 

integuments, the hyphae bend rather sharply and grow toward the 

embryo. The hyphae running on a straight course from the surface 

of the pericarp to the integuments have very few branches, or none 

at all. They seem to have no difficulties in penetrating cell walls, 

and they develop no swellings, or very small ones only, at the points 

of penetration (Figs. 6-7). 

After a few days, the situation changes somewhat. Along with the 

hyphae travelling a straight course, as described above, several hyphae 

are now observed growing irregularly, with several branches and 

bulbous swellings. On penetrating cell walls, these hyphae form large, 

irregular swellings, often shaped like a dumb-bell (Figs. 13-14). Swollen 

U-shaped hyphae located next to the cell wall are frequently note d 

within the cells. At a later stage, large are as are seen in which all 

cells are invaded by hyphae. Such areas are found in all parts of the 

pericarp from the brush end to the chalaza region. Ha ving reached 

the region where the scutellum adjoins the integuments, the hyphae

- 345- 

Fig. 11. Infection in the upper part of the scutellum. 24 days af ter inoculation. 

- X I80. 

Fig. 12. Heavily infected scutellum. - X 120. 

Fig. 13. Hyphae in the upper part of the pericarp 18 days after inoculation. 

At this time, large swellings of the hyphae appear upon penetration 

of the cell walls. - X 340. 

Fig. 14. Detail from ;Fig. 13. - X 1700. 

branch off and ente r the scutellum. At this time the diameter of the 

hyphae tends to increase. Many hyphae are now present in the layer 

of collapsed cells lying between endosperm and the outer cell layer 

of scutellum. This epidermal cell layer consists of long, epitheliallike 

cells whose distal ends are adjacent to the endosperm. Here 

too, the hyphae form a vigorous network, and frequently large 

areas are seen to be completely interwoven with hyphae. The hyphae

- 346- 

then proceed further into the epithelial layer (Fig. 12) and continue 

up through the hypocotyl (Fig. 10). In the week prior to ripening, 

hyphae are present in all parts of the embryo. Most are observed in 

the hypocotyl and leaf primordia; some, however, are also found in 

root primordia. At thi s stage, hyphae are furthermore not uncommon 

in the endosperm, especially at the base. 

DISCUSSION 

As mentioned in the introduction, the mycelium of U. nu da has 

often been found in barley seed at various stages of its development. 

The observations published by various authors coincide in most respects. 

There is, however, as mentioned before some disagreement 

regarding the place of the primary infection and the route of the 

hyphae to the young embryo. It is the object of this investigation to 

shed some light on these points. It has not been possibIe to confirm 

LANG'S assertion that infection takes place through the styles as well 

as through the pericarp; in the material used here infection only takes 

place through the pericarp. It is highly unlikely that, in the material 

examined, a possibIe path of infection through the styles could remain 

unobserved, as the contrast staining used was extremely effective as 

regard all other parts of the material, and there is no reason why the 

same should not apply to the styles. However, to exclude the possibility 

that the staining method should have failed in the styles, a large 

part of the material was also examined by phase contrast microscop 

but nothing was found. LANG'S statement that pericarpial cells are 

frequently destroyed by the fungus could not be confirmed, and 

this applies also to his allegation that very few hyphae are present 

in the embryo, an allegation which also is inconsistent with the findings 

of HECKE. Furthermore, LANG'S theory to the effect that the thin 

epidermis surrounding the young embryo should protect the latter 

against the early phases of hyphal invasion, making it impossible for 

the fungus to reach the embryo except through the epithelial cells of 

the scutellum at a late stage of embryonal development, must be 

doubted, as in several instances hyphae were observed coming from the 

integument, penetrating the scutellum "from behind" and fin ally 

reaching the epithelial cells of that body. 

According to the results presented in this paper, the following 

hypothesis seems more probable. During its early growth, the embryo 

secretes substances (growth-hormones?) which chemotactically attract 

the fungus hyphae. Via the epitheliallayer of the scutellum and

- 347- 

the integuments, these substances by diffusion enter the pericarp. 

About a week af ter pollination the chemotactic action is much lessened 

and the substances probably no longer reach the pericarp. 

This hypothesis would explain several of the observed peculiar 

features. Due to chemotactic attraction, the early infection hyphae will 

strive to reach the integuments, hence their straight course of trave!. 

The hyphae of later infections, which are not in the same degree subject 

to this chemotactic influence develop irregularly and erratically, 

with large swellings upon penetration of cell walls. These swellings are 

strongly reminiscent of the appressoria-like swellings found outside 

the pericarp which appear at a time when they cannot be .affected by 

any substance seereted by the embryo. 

Consistent with this hypothesis is furthermore the faet that, having 

reached the integuments, the hyphae branch off and follow the 

integuments toward the embryo instead 'of going through the endosperm, 

frequently a much shorter route. The hypothesis also explains 

why hyphae are often seen in the endosperm at a later stage, since by 

this time there is no longer any chemotactic attraction. 

SUMMARY 

The previously held theory that barley may be infected with U. 

nuda through the styles should be discarded. The result of the present 

investigation shows that the hyphae penetrate the pericarp and enter 

the integuments which they follow to the scutellum. Through the scutellum 

they finally reach the embryo. 

REFERENCES 

Brefeld, O. (1905): Untersuchungen aus dem Gesamtgebiete der Mycologie 

XIII. 

(1907): tiber die Brandkrankheiten des Getreides. - Jahrb. d. 

Deutch. Landw.-Ges. 22: 75-84. 

Broili, J. & Schikorra, W. (1913): Beitrage zur Biologie des Gerstenflugbrandes. 

- Ber. Deutsch. Bot. Ges. 31: 336-339. 

Falck, R. (1908): Die Flugbrandarten des Getreides, ihre Verbreitung und 

Bekampfung. - Journ. f. Landw. 56: 173-182. 

Hecke, L. (1905): Zur Theorie der Blliteninfektion des Getreides durch 

Flugbrand. - Ber. Deutsch. Bot. Ges. 23: 248-250.

- 348- 

Lang, W. (1917): Zur Ansteckung der Gerste durch Ustilago nuda. - Ber. 

d. Deutsch. Bot. Ges. 35: 4-20. 

Maddox, F. (1895): Experiments at Eastfield. Dept. Agric., Tasmania. 

(1897): Notes and results on agricultural experiments carried on 

under the auspices of the council of Agriculture of Tasmania at 

Eastfield, Newhavn. Tasmania. 

Popp, W. (1951): Infektion in seeds and seedlings of wheat and barley in 

relation to development of loose smut. - Phytopath. 41: 261 

-275. 

Russell, R. C. (1950): The whole embryo method of testing barley for loose 

smut as a routine test. - Scient. Agr. 30: 361-366. 

Russell, R. C. & Popp, W. (1951): The embryo test as a method of forecasting 

loose-smut infection in barley. - Scient. Agr. 31: 559-565. 

Ruttle, M. L. (1934): Studies on barley smuts and on loose smut of wheat. 

- N. Y. Agr. Exp. Sta. Tech. BuH. 221. 

Shands, H. L. (1940): Wisconsin Agr. Exp. St. BuH. 449, p. 74. (Description 

of inoculation method). 

Simmonds, P. M. (1946): Detection of the loose smut fungi in embryos of 

barley and wheat. - Scient. Agr. 26: 51-58. 


YIELD EXPERIMENTS WITH MUSHROOMS 

CULTIV ATED ON SYNTHETIC COMPOST OR 

HORSE AND PIG MANURE SUPPLEMENTED 

WITH VARIOUS SUBSTANCES 

By C. RIBER RASMUSSEN 

The Mushroom Laboratory, The Royal Veterinary and Agricultural College, 

Copenhagen 

From June 1954 to January 1955 and from February 1955 to October 

1955 two identicaI experiments were carried out with the cultivation 

of mushrooms in the experimental houses of the Mushroom 

Laboratory under the Royal Veterinary and Agricultural College, 

Copenhagen. The object of these experiments was to establish the 

difference between the yields from three synthetic composts, two 

horse manure composts and one pig manure compost. Furthermore, 

in both experiments two casing soil types were tested, one of them 

a meadow soil, rich in humus, which had given the highest yield in 

previous experiments, and the other, used for comparison, a very 

stiff clayey soil (potter's clay). Finally, an investigation was made 

of the difference in yields obtained by growing on shelves and by 

growing in trays, as well as of the effect of pasteurization on the yield. 

PLAN OF EXPERIMENTS 

A. - Horse manure (control). 

Addition per ton of fresh manure : 

4 kilos ammonium sulphate (at stacking) 

15 fodder lime (at 1st turn) 

4 superphosphate plus 15 kilos gypsum (at 2nd turn) 

B. - Horse manure (+ 50 %). 

Addition per ton of fresh manure : 

6 kilos ammonium sulphate (at stacking) 

22.5 fodder lime (at 1st turn) 

6 superphosphate plus 22.5 kilos gypsum (at 2nd turn) 

- 349-

- 350- 

O. - M. R. A. (100 % blood meal) 1 ) 

Per 1000 kilos of dry wheat straw: 

10 kilos ammonium sulphate (preparation of straw) 

Aet. 2) X 

(at staeking) { 

Aet. Y 

(at 1st turn) 

Aet. Z 

(at 2nd turn) 

{ 

150.0 kilos 

6.5 

16.0 

6.5 

22.5 

blood meal 

superphosphate 

gypsum 

potassium sulphate 

fodder lime 

340 grammes magnesium sulphate 

340 

iron sulphate 

42 

aluminium sulphate 

42 

eopper sulphate 

32 

zine sulphate 

32 

boric aeid 

32 

ammonium molybdate 

14 

ehromium sulphate 

7 

potassium bromide 

7 

potassium iodide 

6.5 kilos superphosphate 

32.0 gypsum 

D . - M. R. A. (50 % blood m eal) 3) 

Per 1000 kilos of dry wheat straw: 


E. 

75.0 kilos blood meal 

20.0 urea 

J 4 Aet. Xa 

(at staeking) 

) 

6.5 superphosphate 

16.0 gypsum 

6.5 potassium sulphate 

22.5 fodder lime 

l 

Aet. Y { 6.5 kilos superphosphate 

(omitted) 5) 32.0 gypsum 

- Pig manure 6) 

Addition per ton of manure : 

4 hectolitres brewers grains plus 4 kilos ammonium sulphate 

(at staeking) 

15 kilos fodder lime (at 1st turn) 

4 superphosphate plus 15 kilos gypsum (at 2nd turn) 

(30 gypsum (at 4th turn)) 

1) A detailed description of M. R. A. (Mushroom Research Association's 

formula) compost is given in "Gartnertidende", Copenhagen, No. 4/1953. 

2 ) Aet. = Activator. 

3) In the 2nd experiment a further amount of 75 kilos blood meal was 

added at the 4th turn. 

4 ) According to English statements, urea gives, in general, a lower yield 

than does blood meal. 

5) Omission of Activator Y (the micronutrients) should, theoretically, enhance 

the chance of tracing a decrease in the yield, thus proving the 

indispensability of these substance. 

6) In the 2nd experiment neither brewers grains nor ammonium sulphate 

was added at the set-up. Furthermore, 30 kg gypsum were omiUed at 

the 4th turn.

F. - Brewers grains compost 

- 351 

Per 1000 kilos of wheat straw: 


Aet. Xb 

(at staeking) 

Aet. Ya 

(at 1st turn) 

Aet. Za 

(at 2nd turn) 

{ 

{ 

15 heetolitres brewers grains plus 5 kilos 

ammonium sulphate 

40 kilos fodder lime 

340 grammes magnesium sulphate 

340 iron sulphate 

42 aluminium sulphate 

42 eopper sulphate 

32 zinc sulphate 

32 boric acid 

32 ammonium molybdate 

14 chromium sulphate 

7 potassium iodide 

7 potassium bromide 

12 kilos superphosphate 

40 gypsum 

According to the plan of experiments, the object was to investigate 

(1) whether it would be possibIe to increase the yield by an extra ad 

dition of chemicals to horse manure than had been used so far; (2) 

whether any difference in yields from the two M. R. A. composts could 

be traced when 50 % of the blood meal was replaced with urea while 

the micronutrients were omitted; (3) whether, after all, the M. R. A. 

composts were able to give the same yields as the tried-out horse 

manure composts (according to the British Mushroom Laboratory, 

the two M. R. A. composts give the same yield - at any rate when 

micronutrients are added - and they are on a level with horse ma 

nure composts); (4) whether it is possibIe to produce a synthetic 

compost, but at a considerably lower cost of materials ; and, finally 

(5) whether pig manure lends itself to be used as a culture medium 

at all. 

COMPOSTING 

Composting of the culture medium for the 1st experiment was 

carried out under favourable climatic conditions in the summer of 

1954, and everything turne d out approximately as expected. In the 

2nd experiment, however, composting took place under unfavourable 

climatic conditions (in the winter of 1955 with very severe frost), 

and in particular it proved difficult to make the M. R. A. composts 

absorb sufficient heat, which involved adisplacement of the dates 

prefixed for the stackings for simultaneous termination of the fer 

mentation; the treatments which were completed first therefore had

- 352- 

to be "stored" for a shorter or longer period, until filling and pasteu 

rization could be carried out simultaneously. 

Table 1 gives the numbers of days and turns required before the 

individual composts could be considered ready for filling. 

TABLE 1 

1st experiment 2nd experiment 

NO.of 

\ 

No.of NO.of 

I 

NO. of 

days turns days turns 

A: Horse manure (control) 14 4 24 5 

B : - - + 50 % 14 4 24 5 

C: M. R. A. (100 % blood meal) 17 4 49 7 

D: M. R. A. ( 50 % - - ) 19 4 49 7 

E: Pig manure 17 5 28 5 

F: Brewers grains compost 18 4 30 5 

For each of the three synthetic composts (C, D and F), 1000 kilos 

fresh wheat straw were used, which were pre-fermented by addition 

of the ammonium sulphate and large quantities of water before the 

actual stacking with activators (vide the Plan of Experiments). For 

the two sorts of horse manure, 2000 kilos fresh racing stable manure 

were used, while 3000 kilos pig manure wereemployed. 

The compost heaps were stacking as normal for the substratum 

concerned and covered with straw mats on the sides and at the ends. 

All turns were performed meticulously but otherwise as is done in 

practice. The additives were spread as evenlyas possible, and a turn 

ing machine was used at all turns. The dates for the turns were, as a 

rule, determined by the changes in temperature, the formation of the 

actinomycetes and the relative humidity. Great importance was at 

tached to the proper supply of water during the turns. 

Shelf house: 

At filling 

At spawning 

Tray house: 

At filling 

At spawning 

TABLE 2 

Water percentages (averages of 3 samples) 

A I B I C I D E I F 

1 * ) 70 70 79 77 66 78 

2*) 72 71 79 77 72 79 

1 64 63 74 74 46 75 

2 70 . 68 79 74 70 78 

1 70 69 79 78 64 77 

2 72 71 80 77 73 79 

1 67 66 77 76 48 76 

2 69 68 78 75 71 79 

:1: ) 1 = 1st experiment. 2 = 2nd expenment.

Shelf house: 

At filling 

At spawning 

Tray house: 

At filling 

At spawning 

- 353- 

TABLE 3 

pH readings (averages of 3 samples) 

A I B I C I D E I F 

1 *} 7.5 7.4 7.4 7.4 8.2 7.1 

2*} 6.9 6.8 6.7 6.6 6.6 6.6 

1 7.3 7.2 7.1 7.4 7.8 7.3 

2 7.0 7.0 6.9 7.4 7.4 7.0 

1 7.5 7.4 7.4 7.3 8.1 7.2 

2 6.9 6.8 6.5 6.6 6.8 6.8 

1 7.5 7.3 7.3 7.4 7.8 7.2 

2 7.3 7.5 7.6 7.5 7.5 7.3 

Tables 2 and 3 indicate the water percentages and the pH-values, 

respectively, at the filling and at the spawning. The six treatments 

were filled partly on shelves and partly in trays after having been 

intimately mixed shortly before. All the while, compost was tak en out 

alternately from each heap both for shelves and trays. The experiments 

were laid out with systematic plot distribution and with six 

repetitions in the shelf house (72 m 2 bed surface) and 8 repetitions 

in the tray house (56 m 2 bed surface). The area of each plot in the 

shelf house was 1 m 2 , while the area of each frame was only V2 m 2 • 

On an average, 80 kilos compost were filled per square metre of bed 

surface. 

After filling, the trays were stacked in a specially designed pasteurizing 

room, and immediately thereafter the so-called "peak-heating n 

(pasteurization) was commenced, steam being piped into the shelf 

house and the tray room from a steam locomobile, while the central 

heating lines were fully open. 

In the 1st experiment, it took 10-12 hours for the temperature 

to rise to the desired 60-61° C, and pasteurization was terminated 

after some 50 hours at thi s temperature. In the 2nd experiment, it 

took 20-22 hours to reach the temperature of about 60° C, and the 

steam was turned off after 75 hours. 

The mushroom spawn was planted when the temperature had dropped 

to 30° C in the composts. After another 21 days, the plots were 

cased with the two types of soil. At the same time the trays were 

stacked for growing in an adjacent room. In both rooms the tempera- 

* } 1 = 1st experiment. 2 = 2nd experiment. 

FRIE SIA V 23

TABLE 4 

A = Horse manure (control) 

B = Horse manure (+ 50 %) 

o = clayey soil 

No of days 

Com bi· from planting 

nation to galhering 

H 

I 

l{ 

1 

Ao 2 

37 

41 

38 

42 

1 

Bo 2 

37 

39 

37 

41 

1 

Co 2 

38 

41 

37 

43 

1 

Do 2 

39 

44 

39 

44 

1 

Eo 2 

(60) 

45 

(60) 

45 

1 

Fo 2 

38 

41 

38 

42 

Ab 1 

2 

35 

39 

38 

40 

Bb 1 

2 

35 

37 

37 

41 

Cb 1 

2 

38 

38 

36 

41 

Db 1 

2 

38 

40 

39 

45 

Eb 1 

2 

(60) 

42 

(60) 

44 

Fb 1 

2 

36 

37 

38 

40 

YIELDS 

C = M. R. A. (100 % blood meal) E = Pig manure 1 = 1st experiment 

D = M. R. A. (50 % blood meal) F = Brewers grains compost 2 = 2nd experiment 

b = meadow soil H = shelves K = trays 

1 st period 2 nd period 3 rd period 

Normal picking period 

1: abt. 90 days 

2: abt. 90 days H 

28 days kg/m 2 28 days kg/m 2 34 days kg/m2 Shelf house Tray house less 

K 

I Grames I crease In· I Grames I crease In- 

kg m 2 mush in yield kg ll\21 JlIush in yield 

H J{ 

I 

H J( 

I 

H I 

!{ Irooms I m 2 rooms I m 2 

kgm2 8.3 

8.3 

5.6 

8.2 

4.9 

2.8 

5.0 

4.8 

4.2 

4.7 

3.7 

6.9 

17.4 12.6 O 

15.8 12.2 O 

14.3 

19.9 

12.2 

12.3 

O 

O 

+3.1 

- 4.1 

9.2 

8.7 

6.0 

8.2 

7.5 

5.1 

5.1 

3.9 

4.5 

5.2 

5.1 

4.2 

4.5 

4.6 

4.5 

4.3 

7.3 

4.5 

18.8 

17.2 

15.0 

13.1 

12.2 

15.1 

+1.4 

+1.4 

- 2.4 

17.7 

19.9 

13.8 

13.3 

12.4 

14.1 

+3.4 

O 

-0.5 

+1.1 

- 2.7 

+1.2 

4.0 4.1 1.8 2.6 3.5 4.5 9.3 13.1 - 6.5 11.2 12.9 - 8.7 -1.9 

4.7 

2.7 

4.1 

1.6 

4.7 

1.7 

4.2 

2.3 

5.8 

3.5 

5.6 

3.8 

15.2 

7.9 

14.8 

12.5 

- 

- 

2.2 

7.9 

13.9 

7.7 

15.1 -0.4 

10.2 - 12.2 

+1.3 

+0.2 

0.4 0.4 1.1 0.6 1.3 0.6 2.8 12.2 - 14.6 1.6 12.1 - 12.7 +1.2 

5.9 5.8 3.8 5.4 3.5 5.5 13.2 12.3 - 2.6 16.7 12.5 -3.2 -3.5 

4.4 4.5 3.6 3.3 4.1 3.5 12.1 15.1 -5.3 11.3 12.1 -3.0 +0.8 

3.5 4.1 2.6 3.1 4.7 5.4 10.8 13.6 - 5.0 12.6 13.3 - 7.3 -1.8 

9.6 7.8 5.7 5.5 4.0 4.1 19.3 11.8 O 17.4 11.7 O +1.9 

8.8 9.2 3.4 5.4 6.4 7.4 18.6 11.3 O 22.0 11.3 O -3.4 

11.2 8.7 6.1 5.4 4.9 4.2 22.2 11.8 +2.9 18.3 10.9 +0.9 +3.9 

9.4 7.6 3.7 5.1 5.5 7.4 18.6 11.4 O 20.1 11.2 - 1.9 -1.5 

6.0 4.8 4.3 4.0 5.9 4.5 16.2 13.5 - 3.1 13.3 13.3 -4.1 +2.9 

4.5 3.9 2.6 2.4 4.2 5.4 11.3 11.9 -7.3 11.7 11.4 - 10.3 -0.4 

5.0 4.4 5.8 4.6 7.0 5.6 17.8 13.6 -1.5 14.6 13.0 - 2.8 +3.2 

3.0 1.5 2.9 2.5 4.9 4.1 10.8 11.5 -7.8 8.1 10.3 - 13.9 +2.7 

0.7 1.4 1.4 1.5 1.3 1.5 3.4 10.6 - 15.9 4.4 10.8 - 13.0 -1.0 

6.3 6.4 3.1 5.2 5.4 6.3 14.8 10.9 -3.8 17.9 11.2 -4.1 - 3.1 

5.3 4.5 3.9 3.4 3.9 3.8 13.1 11.7 -6.2 11.7 13.7 - 5.7 +1.4 

3.7 3.7 2.8 3.4 4.9 6.6 11.4 12.9 -7.2 1 13.7 11.4 - 8.3 -2.3 

w 

C/l 

tf:.

- 355- 

ture was lowered gradually to about 16° C and the ventilation was 

increased gradually so as to correspond to complete air renewal 2 or 

3 times an hour. In the 1st experiment, all the composts, with the exception 

of pig manure, were almost transpierced by spawn. In the 2nd 

experiment, the two M. R. A. composts had a considerably poorer 

growth than the other treatments. 

The mushrooms ripe ,for picking from the individual plots were 

counted, weighed and recorded daily. The normal picking period extended 

over about 90 days. 

Table 4 represents the results of the pickings. In the first column 

a combination of compost treatment and casing soil is indicated. The 

second column contains the number of days which passed from the 

spawning to the picking of the first rip e mushrooms. The following 

three column show the results from shelves and trays in the 1st picking 

period (28 days), the 2nd period (the next 28 days) , and the 3rd 

period (about 34 days). Finally, the total yield, the average weight 

per mushroom, and the increase (decrease) in yield in comparison 

with the control (horse manure with normal addition). In the last 

column the difference between the yields from shelves and trays is 

given. 

NORMAL PICKING PERIOD 

Experiment 1. 

(a) Shelf house 

The highest total yield was noted for the B manure (+ 50 %) 

cased with meadow soil, viz. 22.2 kilos per square metre, and in comparison 

with the control manure (A) cased with the same soil the 

increase in yield was 2.9 kilos per square metre, and 1.4 kilos when 

clayey soil is used for the casing. The best synthetic compost was D: 

M. R. A. (50 %), though the difference is but slight. A comparison of 

the best synthetic com post and the best horse manure (B + 50 %) 

cased with meadow soil shows a difference of 4.4 kilos per sq. metre. 

The F compost cased with meadow soil gave 9.1 kilos less per sq. 

metre than the best horse manure. Altogether, the meadow soil has: 

given a somewhat larger increase in yield than the clayey soil - in 

a single case up to over 3 kilos per sq. metre. Both soils seem, 

however, to be very suitable. Considering the column for grammes 

per mushroom, it is evident that, irrespective of the casing soil, the 

two horse manure composts give the lowest weight and thus the 

poorest quality. All mushrooms picked from clayey soil plots give 

heavier weights than those from the meadow soil. 

23*

(b) Tray house 

- 356- 

Here, almost the same conditions prevailed as in the shelf house. 

The two horse manures were best, and the B manure (though case d 

with clay soil) gave an increase in yield of 3.4 kilos per sq. metre. 

AIso D: M. R. A. (50 % blood meal) gave slightly higher yields than 

C: "(100 % blood meal). F: Brewers grains compost gave a comparatively 

low yield. Generally, the difference between the test treatments 

was lower in the tray house than in the shelf house. 

(a) Shelf house 

Experiment 2. 

In the total picking period the yields were, on an average, somewhat 

lower than in the 1st experiment. The two horse manures gave 

the highest yields (A: control, 18.6 kilos per sq. metre cased with 

meadow soil) ; and they proved equally good when cased with meadow 

soil, whereas B: + 50 % gave 1.4 kilos more per sq. metre with 

meadow soil. The two M. R. A. composts gave a considerable decrease, 

as was expected. C. (100 % blood meal) was somewhat better than 

D (50 % bio od meal) with 11.3 kilos per sq. metre cased with meadow 

soil. The pig manure proved surprisingly good, with a yield only 3-4 

kilos lower than that obtained with the horse manures. Of the synthetic 

composts, F: brewers grains compost was best 11-12 kilos per sq. 

metre, but the difference between brewers grains compost and the 

M. R. A. composts was comparatively small, considering that the brewers 

grains compost fermented satisfactorily, whereas the M. R. A. 

composts, and particularly the D compost, were all but abandoned. In 

this experiment, the meadow soil gave about the same yield as in the 

1st experiment. The average weight per mushroom was considerably 

lower in this experiment, and it was also obvious that the quality was 

poorer, but it was once more established that clayey soil gives abetter 

quality than does meadow soil. 

(b) Tray house 

The variation within the five treatments was almost the same as 

in the shelf house; however, there was no increase in yield for B: 

+ 50 % - on the contrary, the control A cased with meadow soil 

gave 1.9 kilos more per sq. metre (22.0 kilos altogether).

- 357- 

In the last column in the left-hand main section (H less K) of 

Table 4 the difference between the yields from shelves and trays is 

stated. In the 1st experiment the shelves gave a somewhat higher 

yield than the trays; but in the 2nd experiment the reverse is the 

case, the trays generally giving surprisingly high increases in yields. 

It will probably be difficult to account for this faet, but the difference 

may be due to a more or less optimal fresh-air supply during peakheating, 

even though it was impossible in any of the experiments to 

established this by comparison of the pH readings, the water percentages 

and the growth of the spawn at eas in g time. It is quite likely, 

however, that the difference is caused by other factors. 

In addition to the normal number of trays used in the experiments. 

several trays were left unpasteurized, and Tables 5 and 6 show the 

increases in yield which were obtained by pasteurization. In most 

cases the increases in yield are quite appreciable, proving that the 

extra work and expenditure pay many times over. 

DISCUSSION 

The two experiments did not agree very well, which, in the first 

place, must be ascribed to the abnormal fermentation in the 2nd 

experiment. Altogether, it may be said that the horse manures 

suffered no deterioration by the long storage time before the filling. 

Whereas in the 1st experiment a fairly constant increase in yield 

was obtained with an extra 50% of the additive which was used in 

control A, this was not pronounced in the 2nd experiment. However, 

it is impossible to say whether this is due to the abnormal conditions 

or to other things, or whether the increased yield in the 1st experiment 

is accidental; but for the present it cannot be recommended to 

add 50% of chemicals in relation to the control with "normaF' 

addition, in the hope of obtaining a higher yield. Nor is it possible 

to say whether the extra addition may give greater security in 

cultivation. In the 1st experiment it is interesting to note that 

although half of the nitrogen has been applied in the form of urea, 

and the micronutrients have been left out, the M. R. A. (50 % blood 

meal) has nevertheless given the highest yield in the total picking 

period. In the 2nd experiment there is upwards of 3 kilos more per 

sq. metre for M. R. A. (100%) as compared with M. R. A. ("50%") 

less micronutrients, but, considering the comparatively low yields 

alto gethe r and the very abnormal course of the fermentation, the

- 359- 

2nd experiment actually shows nothing concrete beyond the faet that 

M. R. A. compost requires favourable conditions to be a complete 

success, and that M. R. A. (100 % blood meal) fares slightly better 

under unfavourable climatie conditions than when half blood meal 

and half urea are used. 

F: Brewers grain compost, has not given particularly high, though 

fairly equal, yields in the two experiments. It should be observed, 

however, that the brewers grains compost fermented excellently, even 

under the unfavourable climatie conditions, and apparently there is 

good reason to use brewers grains as kind of a "starter" and then 

add suitable nitrogen sources. The two experiments demonstrate 

clearly that the brewers grains compost lacks one or more "substanees" 

to give yields comparable to those obtained with horse 

manure - but what substances, and in what quantities, ean only be 

found by future experiments. 

Whereas the pig manure in the 1st experiment failed to give good 

results owing to various causes, the 2nd experiment showed that 

good yields ean be obtained if manure from pig herds with normal 

feeding and wheat-straw litter is used - and with a likelihood of 

more constant yields than with the use of the synthetic compost used 

in Europe today. 

In both experiments the interesting observation is made for the 

synthetic compost, particularly D and C in the 1st experiment, that 

the yields are increasing from period to period. This certainly tends 

to indicate that: (1) some sort of fermentation takes place during 

growing, (2) or the composting / "peak heating" has been inadequate, 

(3) or the mushroom spawn ean somehow "decompose" the compost 

(say, by enzymes) during cultivation. 

SUMMARY 

1. The horse manures proved superior to all other compost. 

2. An extra addition of 50 % of the inorganic substances used in the 

control had no consistent effect on the yield. 

3. Under favourable climatie conditions the M. R. A. compost gave 

fairly good yields, but they are uncertain when composting is 

done during spelIs of severe frost.

- 360- 

4. The brewers grains compost fermented excellently under all climatic 

conditions, but yields were too low in comparison with those 

obtained with horse manures. 

5. Fairly good yields are obtainable if a good wheat-straw littered 

pig manure is used. 

6. It is generally alleged that lower yields are obtained by growing 

in trays, but the experiments demonstrated that this need not 

always be so. 

7. The meadow soil gave the highest yield, but the poorest quality of 

mushrooms. 

8. Pasteurizing of the composts gave considerably increased yields 

in most cases. 


THE NATURAL RESISTANCE TO DECAY OF 

CERTAIN CONIFERS 

By ERIK RENNERFELT 

Forest Research Institute of Sweden, Stockholm 51. 

A number of conifers contain heartwood which is strongly resistant 

to insects, termites and fungi while in others the heartwood 

can be destroyed almost as easily as the sapwood. It is now generally 

accepted that this natural resistance depends to very great extent on 

the occurrence of specific constituents which are laid down during the 

formation of the heartwood. Through the extensive investigations of, 

in the first place, ERDTMAN and his co-workers (ERDTMAN 1953,1955) 

many of these substances have been isolated and their chemical constitution 

determined. In some cases it has also been shown that these 

compound s have strong fungicidal properties; pinosylvin, nootkatin 

and the thujaplicins inhibit decay fungi on an agar substrate in 0.01 

-0.001 % concentrations (RENNERFELT & NACHT 1955). 

In the present investigation the resistance to decay of the heartwoods 

of a number of conifers has been tested. The wood samples 

were kindly provided by Professor H. ERDTMAN, Stockholm. The 

rotting tests have been carried out with three commonly used fungi, 

namely Coniophora puteana) Lentinus lepideus and Poria vaporaria; 

four different samples of each kind of wood were tested over a period 

of four months by the LEUTRITZ' soil method. 

From the results in Table 1 it may be seen that different species of 

conifers have very different resistances to decay. The following species 

show g r e a t d u r a b i l i t y: Actinostrobus pyramidalis) Cedrus 

deodara) Chamaecyparis nootkatensis) Ch. pisifera) Cupressus Law 

soniana) Libocedrus spp., Podocarpus totara) Sequoia sempervirens) 

Thuja plicata and Widdringtonia whytei. In one or two cases one of 

the three fungi, principally Coniophora puteana) has been able to 

aUack the wood although neither of the other fungi had any appreci- 

- 361-

- 362- 

Table 1 

Durability of sorne conifer heartwoods 

Per cent los s in Per cent 

Density weight caused by moisture 

No. Species of the content 

dry wood 

Con. Lent. Poria af ter 

put. lep. vap. decay 

I I 

1 Actinostrobus pyramidalis 0.68 2.8 0.1 31.1 

2 Agathis alba 0.58 19.4 23.3 24.6 47.3 

3 Araucaria imbricata 0.54 30.4 23.9 30.2 51.7 

4 Cedrus deodara 0.40 9.5 2.4 2.5 28.6 

5 Chamaecyparis nootkatensis 0.48 18.1 1.5 1.2 40.0 

6 - pisifera 0.36 20.5 1.7 6.3 28.4 

7 Cryptomeria japonica 0.54 22.1 25.1 46.6 

8 Cupressus Lawsoniana 0.52 4.2 4.2 6.5 61.7 

9 Fitzroya patagonica 0.34 2.5 16.5 30.3 

10 Juniperus communis 0.52 30.7 12.0 21.0 63.2 

11 Larix decidua 0.55 24.4 29.3 32.6 58.9 

12 Libocedrus Bidwillii 0.35 21.5 0.7 3.6 37.5 

13 - chilensis 0.51 1.6 0.7 0.8 31.4 

14 - decurrens 0.38 3.3 0.7 0.6 23.1 

15 Pilgerodendron uviferum 0.45 1.8 2.7 31.5 

16 Podocarpus totara 0.51 1.3 31.2 

17 Sequoia sempervirens 0.42 2.8 2.9 3.0 75.2 

18 Sequoiodendron giganteum 0.29 23.6 1.5 12.1 42.1 

19 Taxus baccata 0.59 22.7 13.7 23.2 39.8 

20 Thuja plicata 0.27 5.7 1.9 0.8 33.4 

21 Widdringtonia whytei 0.42 0.4 1.9 1.8 28.4 

22 Pinus silvestris 0.63 25.9 30.5 34.2 54.4 

23 - - sapwood 0.62 67.6 42.0 50.3 64.7 

able effeet. The folIowing species ean be elassed as of l o w o r m od 

e r a t e r e s i s t a n e e: Agathis alba) Araucaria imbricata) Juni 

perus communis) Larix decidua and Pinus silvestris. Taxus baccata 

also suffered eonsiderable attaek. Field tests by other investigators 

(SMITH 1949, GRAHAM 1954) have given results whieh agree well 

with these laboratory tests. 

As already mentioned resistanee to deeay depends on the oeeurrenee 

of e h e m i e a l e o m p o u n d s in the heartwood. As ean be 

seen from Fig. 2 the deeay resistant woods, with the exeeption of 

Actinostrobus pyramidalis) do not have a partieularly high density; 

the very durable Thuja plicata has the lowest density.

- 363- 

Table 2 

Resinous appearence and heartwood constituents of the conifers 

No. Species 

1 Actinostrobus pyramidalis 

2 Agathis alba 

3 Araucaria imbricata 

4 Cedrus deodara 

5 Chamaecyparis nootkatensis 

6 Ch. pisifera 

7 Cryptomeria japonica 

8 Cupressus Lawsoniana 

9 Fitzroya patagonica 

10 Juniperus communis 

11 Larix decidua 

12 Libocedrus Bidwillii 

13 chilensis 

14 decurrens 

15 Pilgerodendron uvijerum 

16 Podocarpus totara 

17 Sequoia sempervirens 

18 Sequoiodendron giganteum 

19 Taxus baccata 

20 Thuja plicata 

21 Widdringtonia whytei 

22 Pinus silvestris 

*) toxic to fungi. 

Resinous 

appearance 

+ 

+ 

(+ ) 

? 

+ 

+ 

Heartwood eonstituents 

not investigated 

sesquiterpenes, e. g. atlantone 

nootkatin *), earvaerol *), ehamie 

acid * ), ehaminie acid * ) , 

sesquiterpenes 

sesquiterpenes, no tropolones 

diterpenes, sesquiterpenes 

little investigated; no fungicidal 

eompounds known 

unknown; eonsiderable amounts 

of »tannins« 

diterpenes, sesquiterpenes; no 

fungicidal compounds known 

lariciresinol, taxifolin 

nothing known 

one per cent fJ -thujapliein*) 

carvacrol *), hydrothymoquinone, 

thymoquinone*), libocedrol 

sesquiterpenes; no tropolones; 

not yet possibie to explain 

decay resistance 

totarol (probably not active) 

eonsiderable »tannins«, pinitol, 

sequoyin etc. 

unknown 

isotaxiresinol (not active) 

a-,fJ-,y- thujaplicin*), thujic ae id 

sesquiterpenes, sesquiterpene 

aleohols 

pinosylvin *), pinobanksin, pinoeembrin 

The summary in Table 2 gives a list of the compounds known to 

occur (ERDTMAN 1955) in the conifer heartwoods examined in the 

present investigation. Nootkatin, carvacrol, chamic acid, a-, {Jo and 

y-thujaplicin, thymoquinone, pinosylvin etc. have been shown to have 

fungicidal properties (Fig. 1). In a number of cases e. g. sesquiterpenes, 

which sometimes occur in relatively large amounts, fungicidal 

activity could not be detected with the method used. In other cases 

e. g. with Fitzroya patagonica and Sequoia sempervirens the resistance

- 364- 

Fig. 1. Effect of pinosyIvin (left) and nootkatin (right) on the growth of 

the bIue stain fungus Pullularia pullulans. 

to decay must be connected at least partly with the occurrence of 

tannins. 

Certain physical properties should assis t thi s resistance in some 

types of heartwood. In a number of trees the wood is rich in o i l 

which thus renders it water-repellent. Four of the species examined, 

namely Cedrus deodara) Chamaecyparis pisifera) Libocedrus chilensis 

and Widdringtonia whytei) are noted for their oiliness. The curves for 

water content (Fig. 2) show that the water uptake of the wood of 

these species was small as estimated in the pieces of wood after exposure 

to rotting with Lentinus lepideus for four months. 

There appear to be two methods for the protection of heartwood. 

In one, poisonous, somewhat water-soluble substances (nootkatin, 

pinosylvin, thujaplicins etc.) have been deposited - as in Chamaecyparis 

nootkatensis) Libocedrus chilensis) L. decurrens and Thuja plicata 

- in the other the wood is impregnated with oily, water-repellent 

substances. In certain cases heartwood is impregnated in both ways 

as in Libocedrus chilensis which has been found in this investigation 

to be the most resistant type of wood. It is both resinous and contains 

not less than 1 % ,8-thujaplicin (ca 5 kg/m 3 ).

SOME FUNGI OCCURRING ON DIED-BACK 

TOPS AND BRANCHES OF PICEA ABIES AND 

ABIES SPP. IN WESTERN NORWAY 

By HAKON ROBAK 

Forest Research Institute of Western Norway, Bergen. 

With many interruptions, the author has for some years made 

studies on an apical die-back 'Df Norway spruce (Picea abies (L.) 

KARST.). In its typical appearance it is characterized by the leader 

and the youngest branch whorl and the upper part of the last year's 

internodium being dead, just as in the top necrosis once described 

from Sweden by LAGERBERG (1913). By less advanced attack the 

leader and branch whorl still carry healthy bark and green needles 

while a more or less broad belt on the stem, most often 'Dn the last 

year's internodium, exhibits dead and resin-soaked bark and yellow 

or wilted needles. Thus the die-back is initiated by a girdling of the 

main axis, just as stated by LAGERBERG for the corresponding disease 

described by that author. 

Tops showing the initial stage of attack carried no external signs 

of fungal growth. Culture tests demonstrated that the bark had already 

been invaded by fungi, however. On the tops already dead a 

more or less abundant vegetation of fungal fructifications flourished. 

From causes brie fly referred to in two earlier pamphlets (ROBAK 

1952 a and b) none among the fungi met with can be considered the 

causal organism of the disease. 

It is not my intention now to deal with the etiology of the latter. 

I should only think it of interest to mycologists to get a brief account 

on the species of fungi encountered during my work with the problem 

in question because our knowledge of the distribution of Micromycetes 

on conifers in Fenno-Scandia is still rather incomplete (for closer 

information on that subject see f. i. JØRSTAD 1945 and KUJALA 1950). 

- 366-

- 367- 

For the same reason I inc]ude in my list some findings made in connection 

with adieback of branch ends observed on several species 

of Abies. 

Unfortunately, my list can give only a very deficient knowledge 

of the fungus flora really occuring. Partly, thi s is due to the fact 

that a great number of mycelia isolated in culture have been unidentifiable. 

Further one must have in mind that in the beginning taxonomical 

studies were of only secondary interest to me compared with 

the etiological problem. Therefore, some of the species which I observed 

during my first examinations 

of died-back tops 

and which, owing to tlieir 

obvious unimportance as 

possibIe causal organisms, 

N 

I studied only superficially, 

could not be recovered by 

renewed examination of the 

top samples. Among the mycelia 

forming pycnidia in culture 

some lost their fertility 

so quickly that specialists to 

which I sent them for identification 

were not able to get 

much out of them any longer. 

Others formed pycnidia 

of such an abnormal structure 

that identification had 

in any case to be rather 

hypothetical. 

Fig. 1. Map of Western Norway. 

Localities mentioned in text numbered 

in succession. 

In the following account 

a quotient in parentheses 

tells on how many of the 

examined samples the species 

has been observed. In cases 

of fungi littIe known in artificial 

culture I have given a 

brief description of their 

cultural characters. When 

nothing particular is said 

about the medium used, the

- 368- 

latter has been malt agar with 3 % malt extract and 2 % agar 

agar. 

Of synonyms I have quoted only the most important ones. 

Parish: 

Bolsøy 

Eid i Romsdal 

Evanger 

Fana 

" 

Fjaler 

" 

Førde 

Gaular 

" 

Hyllestad 

Jondal 

Jostedal 

Kyrkjebø 

" 

Luster 

" 

Os 

Surnadal 

Tingvoll 

Vik 

Voss 

" 

List of collection localities mentioned in text. 

Locality: 

Bårdalen 

Torviken 

Evanger Priest yard-Forest 

Kismul 

Stend 

Dale 

Hellebust State Forest 

Svinevik 

Førde Priest-yard Forest 

Døskeland 

Senneset 

Sy stad 

Hallaråkershagan 

Jostedal Priest-yard Forest 

Osland in Bjordal 

Tangen at Vadheim 

AIsmo 

Berteig 

Sørheim 

Ekhaug at Syfteland 

Heggset 

Gyl 

Nessedalen 

Bjørke 

Møen 

HYMENOMYCETES 

No. on the 

map: 

3 

4 

19 

23 

22 

6 

7 

8 

5 

9 

10 

11 

25 

15 

13 

12 

16 

18 

17 

24 

2 

1 

14 

20 

21 

Only in one case hymenomycetous fructifications have been observed 

on the samples. At Dale in Fjaler parish, on one among 20 examined 

spruce tops collected 31.3.1948, were observed small, sterile fructifications 

closely resembling those of Corticium laeve PERS. (Syn. C. 

evolvens FR.). From the border of necrosis of the same top was isolated 

a mycelium which I am very much inclined to identify with that 

of the species in question the culture characters of which I have 

studied quite closely before (ROBAK 1942). There is but little doubt 

that our fungus is C. laeve. 

From another sample collected in the same locality 8.11.1947 the 

same mycelium has been isolated together with another hymenomycete 

which remained unidentified. 

Finally, from one among three tops received from Gyl in Tingvoll 

parish 13.6.1949 a mycelium was isolated which had very much in

- 369- 

common with that of Stereum sanguinolentum (A. & SCHW.) FR. but 

which carried its clamp connections singly at the septae and not in 

pairs or whorls as is obviously always the case in S. sanguinolentum 

(ROBAK 1942, NOBLES 1948). 

These ,findings might indicate that the die-back may in some cases 

serve as a way of entrance to apical decay in the stem. 

BY)? 

DISCOMYCETES 

1. Cenangium ferruginosum FR. (Syn. C. abietis (PERS.) Du 

On Picea abies: 

Os parish, Ekhaug at Syfteland 14.4.1948 (3:3). 

As far as I know, C. ferruginosum has never been reported from 

spruce before. Therefore, I hesitated in identifying' the ·examined specimens 

with that species, the more so as their colour (exterior almost 

yellow-grey !) was much lighter than normally observed in the lastnamed 

one when growing on pines. Actually, however, I think thi s colour 

(which may to some extent be due to their somewhat immature 

state) was the only characteristic distinguishing our apothecia from 

those of C. ferruginosum. The distinguished expert on arboricolous 

fungi in Norway, Dr. 1. JØRSTAD, thought their microscopical features 

showed good agreement with that species. The apothecia occurred in 

small groups near the base of the dead part of the tops. Bark cultures 

from the same areas yielded mycelia different from that of C. 

ferruginosum} however (cultural characteristics see GREMMEN 1952). 

PYRENOMYCETES 

2. Diaporlhe eres NKE. sens. WEHMEYER 

Isolated from Picea abies: 

Fjaler parish, Dale 8.11.1947 (1:1), fructifications obtained in culture 

identified by L. E. WEHMEYER. 

As is well known, WEHMEYER operates with very wide species 

limits within the genus Diaporthe (WEHMEYER 1933). His D. eres 

covers 12 among the Diaporthe species noted by WINTER (1887). 

Obviously, however, WEHMEYER thinks those species to be more or 

FRIESIA V 24

- 373- 

exuding black drops of conidia. The brown-olive-coloured spores 

measured 4.5-9.2 X 2.8-4.61-'. Size and shape extremely variable. 

(Fig. 3 c-d).) 

6. Cytispora abietis (SACC.) ? 


Hyllestad parish, Systad 

Fana '" Kismul 

Gaular '" Senneset school 

Isolated from the bark of Picea abies : 

Gaular parish, Senneset school 

7. C. Curreyi (SACC.) ? 

Isolated from the bark af Picea abies : 

Gaular parish, Senneset school 

8. C. Mougeotii LEv. 


Fjaler parish, Dale 

Gaular '" Senneset school 

February 1948 (1:2) 

26.5.1948 (1:1) 

3.4.1948 (2 :4) 

3.4.1948 (1 :4) 

3.4.1948 (1:4) 

31.3.1948 (1 :20) 

31.3 and 3.4 1948 (2:6) 

Isolated from the bark of Picea abies from the same localities on the 

same occasions. 

Partly, these identifications must be considered rather uncertain. 

Df what is here calle d Cytispora abietis the natural material was very 

scanty, so that only culture material (partly on sterilized twigs) was 

sent Dr. PETRAK for identifications. His opinion regarding the Systad 

material of this fungus was as follows: uC. abietis SACC., identification 

very doubtful". Df the Senneset material he writes: uC. abietis 

SACC. Development entirely abnormous". 

Df our C. Mougeotii Dr. PETRAK received natural material from 

both localities quoted above. Re identified both samples as C. Mougeotii. 

Df the Senneset fungus he writes: «Cytospora Mougeotii LEv. 

Stimmt am besten mit dieser Art iiberein, die aber zweifelhaft und 

wahrscheinlich mit C. abietis oder C. Curreyi identisch ist". (C. M.) 

shows the best agreement with this species which seems to be dubious 

and most probably identicaI to C. abietis or C. Curreyi) however). 

Dur supposed C. Curreyi (culture material) Dr. PETRA K refers to 

C. Mougeotii. 

I think it justified, however, to keep the name C. Mougeotii only 

for the material quoted under that name above. For the same material

- 374- 

Dr. PETRAK'S identification has confirmed my own. Among the Cytispora 

species described from coniferous hosts none is note d to have 

conidia as long and narrow as observed in our fungus, with the only 

exception of C. Mougeotii. The conidia (Fig. 4 a) have measured 

5.5-7 X 0.6-0.8 (1.0) p. They are exuded in slimy, white masses. 

Mycelium on malt agar appressed, with a low, felt y, white to creamcoloured 

aerial growth. Daily radial increment at room tempo 3- 

4.5 (5) mm. 

In shape of the conidia and in cultural characteristics thi s fungus 

has been distinctly different from the remaining two forms. 

Of these, the one for which (following PETRAK'S suggestion) I 

have chosen the name C. abietis exhibits dingy-yellow spore masses 

just as noted by ALLESCHER (1901) 

and GROVE (1935) for that species. 

The conidia (Fig. 4 b- c) measure 

4.5 - 6 X 1-1.4 p (abnormally 

Fig. 4. Pycnospores and sporophores 

of Oytispora from culture pycnidia. 

shaped individuals from malt agar 

cultures up to 8-9 p) . Cultures: 

Mat appressed, margin even, with 

very low, felt y to mealy, greybrown 

(Ostw. ge2-ig2) air mycelium. 

Daily radial increment at 

room tempo 6-7.5 mm. 

a. O. M ougeotii LEV. - b. O. abietis 

(SACC.) (?) from Systad. - c. 

do. from Senneset. d.-e. O. 

Ourreyi SACC. (? ). - ( X 1330) . 

For the third fungus I have 

proposed C. Curreyi SACC. mainly 

because of its brick-red spore masses 

(LINn says purplish, acc. to 

GROVE 1935) and its entirely simple conidiophores (Fig. 4 e). In my 

other two Cytispora forms the conidiophores have been very variable 

and most ofte n abnormal in shape and thus of very little help to the 

identification work. Individually, the conidia (Fig. 4 d) resemble 

quite closely those of our C. abietis. Dimensions 4.1-6.3 X 1-1.3p. 

Cultures: Mat appressed, with deeply lobed margin, brown-black with 

little or no aerial mycelium, daily radial increment at room tempo 

normally 10-15 mm. ,self-staling is ofte n pronounced, however, and 

may lead to early suppression of the growth. 

Pycnidia have developed rather abundantly in all three species 

(or varieties?) on malt agar and on sterilized twigs. Most often 

they have appeared as large black stromata, more or less like gigantic 

magnifications of natural Cytispora pycnidia.

- 375- 

9. Dothichiza pithyophila (CDA.) PETR. (Syn. Sclerophoma pithyophila 

(CDA.) HOEHN.) 



Kyrkjebø" ,Tangen at Vadheim 

Luster '" Berteig 

" '" Sørheim 

Gaular " ,Senneset school 

Bolsoy '" Bårdalen state forest 

On Abies Nordmanniana: 

Os parish, Ekhaug at Syfteland 26.5.1948 and 24.1.1949. 

Isolated from the bark of Picea abies: 


" " 

Gaular " ,Døskeland saw-mill 

Kyrkjebø" ,Tangen at Vadheim 

Luster '" Berteig 

Bolsoy '" Bårdalen state forest 

Tingvoll " ,Sandvika at Gyl 

31.3.1948 (3:20) 

30.3.1948 (2:5) 

April 1948 (2:8) 

21.2.1948 (2:2) 

31.3.1948 (1:2) 

May 1948 (1:3) 

12.2.1948 (1:2) 

31.3.1948 (3 :20) 

16.6.1948 (1:2) 

31.3.1948 (1:6) 

30.3.1948 (1:2) 

April 1948 (5:8) 

May 1948 (1:3) 

8.6.1948 (2:4) 

The mycelium of D. pithyophila is of the Pullularia pullulans type 

and certainly identical with Hormonema dematioides LAGER BERG et 

MELIN (ROBAK 1952 a). 

!ts cultural characteristics are well known from the comprehensive 

description given by LAGERBERG, LUNDBERG & MELIN (1927). 

I have never succeeded in obtaining pycnidia in culture. From 

USSR NAZAROWA (1936, p. 1208) describes cultures of Sclerophoma 

pithyophila developing pycnidia and open acervuli. She does not mention 

the formation of conidia by direct budding from the hyphal walls 

which is so characteristical to the P. pullulans type of mycelia. Consequently, 

I doubt that her identification is correct. By me numerous 

cultures have been made from spores exuded by pycnidia within 

material the identification of which has been confirmed by Dr. PETRAK 

(the Senneset and Sørheim samples). 

However, pycnidia resembling very much D. pithyophila occurred 

abundantly in one culture isolated from the bark of a die d-back spruce 

leader from Bjørke at Voss 20.4.1948 and, in faet, Dr. PETRAK was 

inclined to identify those culture pycnidia as D. pithyophila. However, 

the mycelium was of a type entirely different from that of the pithyophila 

mycelia otherwise isolated by me. It was very slow-growing, 

daily radial increment at ro om tempo 0.5-0.8 mm. against nearly 

5 mm. in fresh isolates of D. pithyophila) and exhibited no formation 

of conidia from the hyphae. A pycnidiferous mycelium of almost

- 376- 

identicaI appearance was isolated from spores of pycnidia observed 

on one sample among the tops sent me from Berteig in Luster April 

1948. The natural pycnidia differed from those of D. pithyophilc(, 

mainly in their decidedly smaller size, their somewhat more narrow 

conidia and their superficial position. Unfortunately, the fertility of 

the culture strain rapidly declined, and Dr. PETRAK could not find 

any fertile pycnidia when he received cultures for identification. 

Owing to other work I could not spare time for persecuting the 

problem of the identity of those two strains. I have mentioned them 

here only to show that pycnidiferous fungi exist which resemble 

D. pithyophila closely enough to be mistaken for that species and 

which are, nevertheless, distinct. Most probably, NAZAROWA has 

happened to meet with such a fungus. 

10. Sclerophoma entoxylina LAGER BERG et MELIN 



Vik '" N essedalen valley 

16.6.1948 (1:2) 

24.4.1948 (2:2) 

This fungus which has been described on culture material only 

has been so comprehensively dealt with by its authors that identification 

of cultures of the same can be made without difficulty. I refer 

to the original description (LAGERBERG, LUNDBERG & MEL IN 1927). 

11. Phomopsis conorum (SACC.) DIED. 


Evanger 

Fjaler 

Gaular 

Jondal 

Os 

Voss 

parish, Evanger priest-yard forest 

June 1948 (1:3) 

" 

" 

" 

,Dale 

,Døskeland saw-mill 

,Hallaråkershagan 

31.3.1948 ( 4:20) 

31.3.1948 (2:6) 

April-May 1948 (2:2) 

" 

" 

,Ekhaug at Syfteland 

,Bjørke 

26.5.1948 and 24.1.1949 (2:7) 

April 1948 (1:3) 


Fjaler parish, Dale 12.2.1948 

31.3.1948 

" " , " 16.6.1948 

Gaular 

, Døskeland saw-mill 31.3.1948 

Os " , Ekhaug 14.4.1948 

" 

" " , " 26.5.1948 

Luster 

, Berteig April 1948 

Bolsoy " , Bårdalen state forest May 1948 

" 

(1:2) 

(7 :20) 

(1:2) 

(1:6) 

(2:3) 

(4:6) 

(1:8) 

(1:3)

- 378- 

13. Phomopsis pseudotsugae WILS. (Syn. Discula pinicola 

(NAUMOW) PETR.) 



Gaular " ,Døskeland saw-mill 

Os '" Ekhaug at Syfteland 

Tingvoll " ,Gyl 

Bolsøy ", Bårdalen state forest 

12.2. and 31.3. 1948 (5:22) 

31.3.1948 (2:6) 

26.5.1948 (1:6) 

8.6.1948 and 13.6.1949 (3 :7) 

May 1948 (1:3) 

Apart from an earlier communication by the author (ROBAK 

1952 b) Phomopsis pseudotsugae has - under that name - never 

been reported from spruce. Under the name Discula pinicola it is well 

known however, as a blueing fungus in pine and spruce wood and in 

ground-wood pulp of spruce as well (LAGERBERG, LUNDBERG & MELIN 

1927, ROBAK 1932). That the two names are synonyms has been confirmed 

by HAHN (ROBAK 1952 b). Thus, my isolations demonstrate 

that P. pseudotsugae must be relatively wide-spread as asaprophyte 

on spruce although this host does not form a favourable medium for 

its fructification. 

The culture characteristics of the fungus have been described by 

HAHN (1930) and - very comprehensively - by LAGER BERG & MELIN 

(1. c.). 

14. Zytostroma pinastri (KARST.) HOEHN. (Syn. Zythia pinastri 

KARST.) 


Gaular parish, Døskeland saw-mill 

Jostedal " ,Jostedal priest-yard forest 


Fjaler parish, Svinevik 

Gaular '" Døskeland saw-mill 

" '" Senneset school 

Jondal '" Hallaråkershagan 

Voss '" Bjørke 

Bolsøy '" Bårdalen state forest 

31.3.1948 (1: 6) 

June 1948 (1 :4) 

1.4.1948 (1:2) 

31.3.1948 (4:6) 

31.3. and 3.4.1948 (3:6) 

April-May 1948 (1:2) 

March-April 1948 (1:3) 

May 1948 (1:3) 

Identification confirmed by Dr. PETRAK for the Døskeland material. 

Cultures: Mat at first appressed, glabrous, later gradually covered 

by a low, white, cottony air mycelium of a loose texture. Daily 

radial increment at room tempo 3 mm. Pycnidia not observed. Conidia 

forming directly from the mycelium. AIso this fungus seems to be 

more widely distributed than expected from the sparse occurrence of 

its pycnidia.

- 379 - 

15. Hormococcus conorum (SACC. et ROUM.) n. comb. (Syn. 

Sirococcus conorum SACC. et ROUM.) 


Hyllestad parish, Sy stad near Leirvik 13.12.1947 

The cultures carried superficial, globular, less than 0.5 mm. 

broad, black, pore-Iess pycnidia the almost globular, hyaline spores of 

which were formed in chains from bottle-shaped to cylindrical, hyaline 

sterigmata and measured ab. 2 X 2p, each spore having one oil guttul 

e (Fig. 6 a). These observations agree so well with ALLESCHER'S 

description of SirococcU8 conorum (ALLESCHER 1901, p. 445) that 

I do not hesitate in identifying our fungus with that species. 

The mycelium grows very slowly on malt agar. Daily radial increment 

at room tempo ab. 0.4 mm. Mat dense, almost glabrous and 

colourless, with even margin. Pycnidia on the fresh isolate numerous, 

crowded. 

There is no doubt that this species and the folIowing are far from 

being nearly related to each other. Sirococcus conorum is not the 

type species of the genus and ought to be given another generic name. 

I have here proposed Hormococcu8 n. g. 

16. Sirococcus strobilinus (DESM.) PETR. (Syn. Sporonema 

strobilinum DESM., Discella strobilina (DESM.) DIED., S. strobilinus 

PREUSS (?), Ascochyta parasitica (HART.) ROSTR. (?) , A. piniperda 

LINDAU (?) a. o.) 



" '" Hellebust state forest*) 

Gaular '" Senneset school*) 

?Luster " , Berteig 

Vik '" N essedalen valley 

Voss '" Bjørke 

Isolated from the bark of Picea abies : 


Luster Berteig 

Os , Ekhaug at Syfteland 

Vik N essedalen 

31.3.1948 (1:20) 

1.4.1948 (1:1) 

31.3. and 3.4.1948 (2:6) 

April 1948 (1:8) 

24.4.1948 (2:2) 

March- April 1948 (1:3) 

31.3.1948 (1:20) 

April 1948 (1:8) 

26.5.1948 (1:6) 

24.4.1948 (1:2) 

JØRSTAD (1945) considers Discella strobilina (DESM.) DIED. idencal 

to Ascochyta piniperda LINDAU and A. parasitica (RART.) ROSTR. 

*) Material examined by Dr. PETRAK, who confirmed the identification.

- 380- 

In my opinion, the material resting in the Herbarium of the Botanical 

Museum of Oslo university supports this view. What separates the 

specimens given the species names in question is only the different 

habitat from which they have been collected (i. e. cones or twigs !). 

HARTIG'S illustrations of his Septoria parasitica (HARTIG 1890) 

show spores which - according to the magnification noted - measure 

ab. 12.5-14.5 X ab. 2%-3p. For Sporonema strobilinum DESM. 

ALLESCHER (1903) notes: 10-15X2.5-3p. For the Norwegian ma- 

Fig. 6. a. Hormococcus conorum (SACC. 

et ROUM. ) n. comb. Conidia and conidiophore 

( X 1000). 

b.-f. Eirococcus strobilinus (DESM.) 

PETR. Main form; b. Conidia from natural 

pycnidia; c.- f. Conidia from culture 

pycnidia of different isolations. 

( X 500). 

g. Eirococcus strobilinus (?) deviating 

type from natural pycnidium. Berteig in 

Luster, with exclusively quadricellular 

conidia. ( X 500) . 

h.-i. Eirococcus strobilinus var. acutisporus. 

Conidia and conidiophore with 

young conidia. ( X 500) . 

terial (referred to both species 

names) J 0RSTAD quotes : 

9.5-15 X2-2.5 (-3) p . 

Within my own material 

(Fig. 6 b) the Bjørke specimens 

had spore dimensions 

like 9.8-17.2X3-4p and 

the N essedalen specimens 

9.8-19.8 X 3-4p (where 

two-celled spores are concerned) 

. For Danish material 

of A. parasitica (HART.) 

ROSTR. FERDIN ANDSEN & J 0R 

GENSEN (1938) note 13-15 X 

5-6p, i. e. obviously broader 

conidia then ever described 

for Sirococcus strobilinus. In 

this respect their figures differ 

just as much from HAR 

TIG'S original illustrations of 

S. parasitica) however. Thus, 

thi s slight difference in spore 

width which seems to be the 

only noticeable difference between 

the shoot-inhabiting 

fungus of the Danish authors 

and the cone-inhabiting S. 

strobilinus is most probably 

due to inter-regional variabilit 

Y only. Obviously, some of 

my own specimens are in the 

said respect intermediate.

- 381- 

A peculiarity of my material is, however, that many pycnidia 

contained a · small proportion of 3-4-celled conidia, the length of 

which was correspondingly greater, i. e. up to 30/1. (Fig. 6 c-f). In 

pycnidia developed in culture such big and otherwise strange-looking 

conidia often dominated entirely, particularly in mycelia having been 

cultivated for a more extended period. 

The mycelium of Sirococcus strobilinus on malt agar develops a 

slow-growing mat with even margin and low, felty-woolly, yellowgreen 

(Ostw. ne2-pg2 or in sectors ig2) aerial mycelium. Daily 

radial increment at room tempo 0.7-1.6 mm. Pycnidia sometimes 

occur on malt agar, abundantly when a sterilized spruce twig is partly 

embedded in the agar slant. 

Two forms differing more or less markedly from the rest have 

been met with. 

One form was isolated from spores exuded by a singly occurring 

pycnidium on spruce from Berteig in Luster parish April 1948. This 

pycnidium had almost exclusively 4-celled spores, closely resembling 

the 4-celled individuals amongst the S. strobilinus conidia (Fig. 6 g). 

By germination they developed a mycelium which differed from that 

of S. strobilinus by its somewhat darker colour and slightly faster 

growth. The identity of this fungus with S. strobilinus may be doubtful. 

Material is too scanty for giving conelusive information in thi s 

respect, however. 

The other form, occurring as pycnidia on one of the original 

Senneset samples (besides typical specimens) was distinguished by 

its distinetly pointed spore extremities (resembling those of Darlucal). 

Dr. PETRAK identified this fungus as Sirococcus strobilinus) 

and the pycnidia could hard ly if at all be distinguished from those of 

that species. In culture, however, it behaved entirely different from 

the last-named one. In faet, it developed an appressed, glabrous mat, 

almost without aerial hyphae, but with distinct zones forming concentricaI 

waves 7-8 cm. in width on the mat surface. At first colourless, 

later more or less black. Daily radial increment 1.1-1.4 mm. Growth 

rate about three times as great when a sterilized spruce twig has been 

partly embedded in the slant. Pycnidia then abundant. Pycnospores 

exclusively two-celled, 11.8-17.8 X 2.5-3/1, shape more variable in 

culture than in the original material (Fig. 6 h). I should like to propose 

the provisional name Sirococcus strobilinus var. acutisporus for this 

fungus.

- 382- 

17. Brunchorstia pinea (KARST.) HOEHN. (Syn. B. destruens 

ERIKSS.) 


Gaular parish, Døskeland saw-mill 31.31.1948 (3:6) 

Jondal '" Hallaråkershagan May 1948 (1:2) 

Voss '" Bjørke March- April 1948 (1:3) 

Isolated from the bark of Picea abies from one sample of each of the 

three collections noted above. 

Although the ontogeneticaI connection between Brunchorstia 

pinea and CrumenuZa abietina LAGERB. (ScZeroderris abietina 

(LAGERB.) GREMMEN 1953) must be considered definitely prooved and 

seems now to be generally accepted, I mention the fungus here under 

the first name because (with one, somewhat doubtful exception) only 

pycnidia have been met with during my investigation. 

The fungus has been encountered by me in relatively few cases. 

It has neither been observed on nor isolated from samples showing 

only initial symptoms of the disease. As a possibIe causal agency 

obviously no importance can be attached to it. Its culture characteristics 

are well known (cmp. f. i. VAN VLOTEN & GREMMEN 1953) for 

which reason I do not mention them here. Within my material, pycnospores 

have measured 30-47 (-51) X3-4p (cmp. ETTLINGER 

1945). 

18. Hendersonia abietis ROUM. et FAUTR. 


Luster parish, Berteig 

Det. Dr. PETRAK. 

April 1948 (2:8) 

Spores (Fig. 7 a) 12-16.2X5-6p (rarely up to 18 X7p) against 

ALLESCHER (1903): 12 X 4p. 

Cultures on malt agar glabrous, almost without air mycelium, 

colourless, daily radial increment at room tempo 2-2.3 mm. Pycnidia 

having extremely delicate walls develop in the presence of sterilized 

spruce twigs in the slant. 

19. Camarosporium strobilinum BONN. 

Gaular parish, Senneset school 31.3.1948 (1:2) 

Førde '" Førde priest-yard forest 21.2.1948 (1: 1) 

Jondal '" Hallaråkershagan April- May 1948 (1:2) 

The Senneset material examined and identification confirmed by Dr. 

PETRAK. 

The conidia (Fig. 7 b) are very much like the Camarosporiurn 

conidia attributed to Cucurbitaria piceae BORTHW. but our fungus has

- 384- 

20. Myxoeyelus eenangioides (ELL. & ROTH) PETR. (Syn. Camarosporium 

abietis WILS. & AND.) 

On Abies concolor Fana parish, Kismul 26.5.1948 

A. nobilis Os Ekhaug 24.1.1949 

" A. nordmanniana: " " 

A. grandis Fana Stend 19.1.1949 

The material from A. concolor was identified by r. JØRSTAD. 

Spores of Myxocyclus cenangioides germinate extremely slowly on 

malt agar. When the mycelia have attained macroscopicaI dimensions, 

growth may for some days be moderately rapid (daily radial increment 

max. 1.4 mm.), the low, dark-coloured mat very unevenly outlined 

owing to self-staling which quickly prevents further growth. 

Staling les s pronounced on a douglas fir decoction agar on whieh 

pycnidia with mature spores (Fig. 7 c) developed. 

21. Robakia aretiea PETR. n. g. et n. sp. 


Fjaler parish, Svinevik 1.4.1948 

This species was discovered by Dr. PETRAK on a top sample sent 

him for examination and has been given a very full description by him 

(PETRAK 1952). I have not succeeded in recovering the fungus on any 

of the samples otherwise examined and, consequently, I am unable to 

yield any informations about its culture characteristics. 

MELANCONIALES 

22. Coryneum juniperi ALLESCHER (?) 


Gaular parish, Senneset school 3.4.1948 (1:4) 

Of his Coryneum juniperi ALLESCHER (1903) notes: "Spores with 5 

septae, the 4 central cells of almost even size, dark-brown, terminal 

cells hyaline, conicaI. Spores at first n o t, later f a i n t l Y constricted 

at the septae, 25-30 X 6-91-'." The spores of our fungus (Fig. 7 d) 

agree fairly well with that description, their dimensions being 20- 

30 X 6- 10 1-', the short terminal spines not included. Cultures on malt 

agar: Mat low, covered by a finelyand irregularly curIe d air mycelium, 

margin white and finely fringed, otherwise blueish olive-grey 

(near Ostw. gc17). Daily radial increment at room tempo 1.4 mm. 

Acervuli develop af ter a few weeks.

- 385- 

23. Pestalolia sabinae FAUTR. (?) 

On Pseudotsuga taxifolia f. viridis: 

Førde parish, Førde 1.5.1948 

Among the Pestalotia species described from conifers P. sabinae 

FAUTR. is certainly the species with which our fungus has most in 

common. In faet, ALLESCHER'S description of P. sabinae agrees very 

well with my observations on the Førde fungus. P. sabinae: spores 

20-22 X 7-8 ,li, our fungus: 16-24 X 6.5-7.5 ,li. Important seems 

the folIowing characteristic which both have in common : The only 

existing terminal projection divided into two or three stiffly diverging 

branches (cfr. ALLESCHER 1903). Certainly, within the original material 

of our fungus the spores were slightly smalle r (max. 20 ,li ?) and 

entirely without any projections. However, in acervuli developed in 

culture on douglas fir decoction agar a certain proportion among the 

spores carried the projections described above (Fig. 7 e) . 

24. Cryptosporiopsis diversispora ROBAK 


Os parish, Ekhaug 

Isolated from A bies grandis : 

Fana parish, Stend 

25. Cryptosporiopsis balsameae ROBAK 

Isolated from A bies balsamea: 

Os parish, Ekhaug 

Isolated from Abies grandis: 

Fana parish, Stend 

14.4.1948 (1:3) 

19.1.1949 (2:2) 

24.1.1949 (1:3) 

19.1.1949 (1:2) 

These two fungi have been comprehensively dealt with by the author 

in earlier publications (ROBAK 1949 and 1950), for the last-named 

one als o its Pezicula stage. 

HVPHOMVCETALES 

26. Tubercularia vulgaris TODE 



Luster , Berteig 

" '" Sørheim 

Eid in Romsdal, Torviken 

Surnadal par., Heggset 

FRIESIA V 

31.3.1948 (5 :20) 

Apr.1948 (1:8) 

21.2.1948 (1:2) 

17.6.1948 (1:3) 

10.6.1948 (1:22) 

25

- 387- 

The excellent student of Alternaria) StemphyZium and morphologically 

related genera Dr. P AVL NEERGAARD, Copenhagen, has prop ose d 

to place the Senneset fungus within the genus Sirodesmium DE NOT., 

preliminarily as a new species. The fungus forms its spores in chains, 

in basipetal succession (contrary to Alternaria) . The chains radiate 

from a center, thus forming a dense, semiglobular mass of max. 2 mm. 

width and of a jet-black colour. Mature spores (Fig. 7 f) are dark 

brown, 20-33 X 12-14 f.Æ (when still longer apparently formed by 

incomplete separation of two individuals). Cultures: On malt agar 

mat fur-like to woolly, few mm. high, of a light-greyolive colour 

(Ostw. ec2 to ge2). Daily radial increment at room tempo 1.3-1.9 mm. 

Spores form in fresh isolates, fertility then quickly declining. 

CONCLUSIONS 

In this paper I have mentioned and, partly, described the fungi 

which could with some degree of certainty be identified, amongst the 

total of species met with during my work regarding the diseases mentioned 

in the introduction. 

A greater number, however, could not be identified at all, although 

in some cases suggestions could be made as to the genera in which 

they had to be placed. Both sphaeriaceous and shaeropsidaceous fungi 

as well as Hyphomycetes and sterile mycelia have been represented 

within this incompletely known material of fungi, the greater number 

of them amongst the species encountered in cultures made from the 

bark tissue. This fact indicates that the dead bark of conifers is invaded 

by a far greater number of fungi than those known to fructify on 

the different species in Iquestion. The repeated isolations of Phomopsis 

pseudotsugae Wils. from Norway spruce yield a striking example of a 

fungus the host list of which includes most probably, a far greater 

number of tre e species than might be concluded merely from the occurrence 

of its fructifications in nature. 


Specialists to whom I am much indebted for their kind and valuable 

help are Dr. F. PETRAK, Vienna, Dr. G. G. HAHN, New Haven, Conn., Dr. 

L. E. WEHMEYER, Ann Arbor, Mich., Dr. P. NEERGAARD, Copenhagen, and 

Dr. I. JØRSTAD, Oslo. 

25*

- 388- 

LITERATURE 

1901. AllescherJ H.: Fungi imperfecti. I. In Rabenhorst's Kryptogamenflora 

V. 1, Abt. VI. Leipzig 1901. 

1903. AllescherJ H.: Fungi imperfecti. II. Ibid. V. 1, Abt. VII. Leipzig 1903. 

1945. Ettlinger, L.: fiber die Gattung C r u m e n u l a sensu Rehm etc. Dissertation. 

Bern 1945. 

1938. Ferdinandsen, C. & Jørgensen, C. A.: Skovtræernes Sygdomme. I. Copenhagen 

1938. 

1952. Gremmen, J.: A preliminary study on the culture of D i s c o m y c et 

e s, especially the perfect stage. - Antonie v. Leeuwenhoek V. 

18: 153-164, 1952. 

1935. Grove, W. B.: British Stem- and Leaf-Fungi. S p a e r o p s i d a l e s I. 

Cambridge 1935. 

1930. Hahn, G. G.: Life-history studies of the species of P h o m o p s i s occurring 

on Conifers. I. - Trans. Brit. Mycol. Soc. 15: 32-93, 

1930. 

1890. Hartig, R.: Eine Krankheit der Fichtentriebe. - Zeitschr. f. Forst-u. 

J agdwesen 22: 667-670, 1890. 

1952. Jørgensen, H. A.: Studies on N e c t r i a c i n n a b a r i n a. Hosts 

and variations. - Contrib. No. 35. Dept. Plant Pathology of the 

Royal Veterinary and Agricult. CoU., Copenhagen. Den kg!. 

Veterinær- og Landbohøjskoles Årsskrift 1952: 57-120, 1952. 

1945. J ørstad, I.: Parasittsoppene på kultur- og nyttevekster i Norge, I. 

Tillegg til Meld. fra Landbruksdirektøren, Oslo 1948. 

1950. Kujala, V.: fiber die Kleinpilze der Koniferen in FinnIand. - Commun. 

Instit. Forestali Fenniae V. 38.4.1950. 

1913. Lagerberg, T.: Granens topptorka. - Meddel. Statens Skogsfors.anst. 

10: 9-44, 1913. 

1927. Lagerberg, T., Lundberg, G. & Melin, E.: Biological and Practical 

Researches into Blueing in Pine and Spruce. I. - Sv. Skogsvårdsf. 

Tidskr. 1927: 145-272, 1927. 

1936. Nazarowa, E. S.: Pine disease caused by S c l e r o p h o m a p i t h y op 

h i l a v. H. - BuU. Acad. Sci. USSR Ser. bio!. 6: 1191-1208, 

1936. 

1948. Nobles, M. K.: Studies in forest pathology. VI. Identification of cultures 

of wood-rotting fungi. - Canad. Journ. Res. C. V. 26: 

281-431, 1948. 

1930. Ostwald, W.: Die kleine Farbmesstafel, Edit. A. Muster-Schmidt, 

Gottingen 1930. 

1952. Petrak, F.: Zwei neue Gattungen der P a r a s p h a e r o p s i d e e n. 

- Sydowia VI: 372-377, 1952. 

1896. Rehm, H.: H y s t e r i a c e e n und D i s c o m'Y c e t e n. In Rabenhorst's 

Kryptogamenflora V. 1, Abt. III. Leipzig 1896. 

1932. Robak, H.: Investigations regarding fungi in Norwegian groundwood 

pulp and fungal infection at wood-pulp mills. - Nyt Mag. 

f. Natvsk. 71: 185-330, 1932.

- 389- 

1942. Robak, B.: Cultural studies in some Norwegian wood-destroying 

fungi. - Meddel. VestI. Forstl. Fors.st. No. 25, 1942. 

1949. - : A heterosporous D i p l o i d i a sp. Isolated from necrotic 

bark tissue of Conifers in Western Norway. - Sv. Bot. Tidskr. 

43: 534-539, 1949. 

1950. - : On the identity of D i p lod i a d i ve r s i s p o r a Robak 

n. sp. ad int. and of a related fungus likewise isolated from dead 

coniferous bark tissue. - Ibid 44: 465-472, 1950. 

1951. - : Noen iakttakelser til belysning av forholdet mellom klimatiske 

skader og soppangrep på nåletrær. - Meddel. VestI. Forstl. 

Fors.st. No. 27, 1951. 

1952 a. - : Dothichiza pithyophila (Cda.) Petr., the pycnidial 

stage of a mycelium of the type P u Il u l a r i a p u Il u l a n s 

(De B.) Berkhout. - Sydowia VI: 361-362, 1952. 

1952 b. - : P h o m o p s i s p s e u d o t s u g a e Wilson D i s c u l a 

p i n i c o l a (Naumov) Petro as asaprophyte on coniferous 

woods. - Ibid. VI: 378-382, 1952. 

1953. Van Vloten, B. & Gremmen, J.: Studies in the Discomycete genera 

C r u m e n u l a de Not and C e n a n g i u m Fr. - Acta Botan. 

N eerlandica 2: 226-246, 1953. 

1933. Wehmeyer, L. E.: The genus D i a p o r t h e and its segregates. 

Ann Arbor, Mich. 1952. 

1887. Winter, G.: Gymnoascaceen und Pyrenomyceten. In 

Rabenhor.st's Kryptogamenflora V. 1, Abt. II. Leipzig 1887. 

Bergen, February 1956.

CAPILLIPES CAVORUM G. NOV., SP. NOV., A NEW 

TERRICOLOUS INOPERCULATE DISCOMYCETE 

FROM SWEDISH LAPPLAND 

By ROLF SANTESSON 

In 1945 I collected in Torne Lappmark a small Discomycete with 

a remarkable appearance and a characteristic anatomy of its fruit 

bodies. For many years I have now sought for it in the literature but 

in vaino Apparently it represents a new genus. 

The fungus was fouild on plant debris in the entrance of a small 

rodent's cave, most probably a lemming's, Lemmus lemmus. This may 

seem to be a strange habitat for a plant. However, on such places the 

lichen Coniocybe furfuracea is not very rare. The locality is situated 

in Abisko National Park in the subalpine Betula pubescens (B. tortuosa) 

forest between Abisko Tourist Station and the southern shore 

of the lake Torne Trask, at an altitude of somewhat more than 350 m. 

Capillipes nov. gen. 

Ascomata stipitata, ± ceracea, sti pit e longo, tenui. Excipulum cellulis 

prismaticis vel subglobosis formatum. Hymenium valde convexum, iodo 

non mutatum. Asci cylindrici. Sporae ellipsoide ae, continuae, incoloratae. 

Paraphyses sim pli ces apice subsphaeroidaliter incrassatae, verrucosae, 

ascos superantes. Planta saprophytica. 

Species typica (et adhuc unica): 

C. cavorum nov. spec. 

Ascomata pusilla, gracilia. Pileus ± globosus, (0,2- ) 0,3- 0,4 (- 0,65) 

mm diarn., pallide griseo-fulvus (fere avellaneus). Stipes (8- ) 11- 18 mm 

longus, 0,05- 0,08 (- 0,12) mm crassus, parte apicali pallida, parte basali 

nigra. Excipulum tenue, parte externa solum leviter fulvescente, ceterum 

incoloratum. Hymenium 50- 70 fl altum; apices paraphysium epithecium 

pallide fulvescens formantes; asci ut paraphyses non agglutinatae. Asci 

- 390-

- 391- 

octospori. Sporae monostiehae, oblongo-ellipsoideae, 8-10X 2-3 fl. Paraphyses 

basaliter 1,5-21-'_' apiee inerassato 6-8 fl erassae, sat dense 

verrueosae. 

Hab.: Suecia, Lapponia tornensis, Abisko, alt. cire. 350 m, in terra 

(plantis valde destruetis) in introitu eavi animalis (Lemmus lemmus ?). 

9.VIII.1945, leg. ROLF SANTESSON. (Holotypus in herb. Instit. Botan., Uppsala.) 

Fig. 1. Capillipes cavorum. Two specimens on a pieee of plant debris. 

About X 20. (Delin. M. INDUSS.) 

Ascomata solitary or in loose groups, stipitate, with an almost 

globose pileus on a very long and slender stipe. Pileus (0.2-) 0.3- 

0.4 (wet 0.35-0.65)mm diam., ± waxy, paIe greyish brown (RIDGWAY: 

Tawny-Olive to Dark Olive-Buff), sometimes as with a thin grey 

pruina. Stipe (8-) 11-18 mm long, 0.05-0.08 (wet 0.09-0.12) mm 

thick, abruptly and repeatedly bent in various directions, when wet 

± cylindrical, when dry angularly and irregularly flattened, its upper 

(114-115) part grey to paIe brownish, its lower part black, ± nitidous. 

Stipe a bundle of hyphae forming a regular prosoplectenchymatous 

tissue, cells 15-30p long, 5-7 (-9) p diam., their walls dark 

brown (except for the upper part of the stipe) and thick in the 

peripheral, thinner in the central part of the stipe. 

Pileus exposing the hymenium at the surface of the globe which 

it forms. The stipe reachs the centre of this globe, its hyphae there 

bending 'outwards and downwards, ending in the excipular tissue. 

Excipulum thin (15-40 p) formed by leptodermatous cells which 

are mainly prismaticai and uncoloured, the outermost ones are ±

- 392- 

Fig. 2 Oapillipes cavorum. a. Section through pileus and upper part of the 

stipe. - X 100. b. Detail of a, showing excipulum, marginal part of hymenium, 

etc. - X 450. c. Upper part of paraphyses. - X 1000. 

globose and with paIe brown walls; the margin of the excipulum not 

prominent. Between the hyphae of the top of the stipe and the hypothecium 

there is a loose medullary tissue, 40-120p thick, formed by 

leptodermatous, 2-4p thick hyphae. Hypothecium uncoloured, nubilous, 

10-20p thick. Hymenium I-, 50-70p thick, its elements not 

glued together, with relatively few paraphyses the swollen ends of 

which form a distinct but not very dense, 7-10p thick, light brown 

epithecium. Asci 8-spored, cylindrical, 45-60 X 3.5-4.5p, their wall 

thickened at the apex but otherwise very thin; the apical apparatus 

totally I-; the spores obliquously arranged in one row. Spores 

(oblong-) ellipsoidal, continuous, uncoloured, smooth, 8-10 X 2-3p; 

many spores I + paIe bluish after a long time in iodine solution. Paraphyses 

simple, not septate, basally 1.5-2p thick, apically broadly 

clavate to almost globose, 6-8p thick, the uppermost part with a 

thick, paIe brownish wall furnished with rather numerous small 

warts. 

c

- 394 - 

matica") but has a very well developed stipe with a "textura porrecta". 

In the family Hyaloscyphaceae the apothecia are often distinctly 

stipitate, the excipulum with a "textura prismatica" is usually furnished 

with characteristic hairs, the asci are usually ± clavate and 

with a positive iodine reaction. Capillipes has no hairs, its asci are 

cylindrical and I - . I know no representative of the Hyaloscyphaceae 

with a strongly convex hymenium, and none with a stipe comparable 

with that of Capillipes. 

The family HeZotiaceae) as delimited by NANNFELDT, has an 

excipulum with a "textura intricata" or/ and "textura oblita". Thus 

the structure of the excipulum of Capillipes is clearly differing. 

In the Helotiaceae there are many species with apothecia having a 

long stipe and also with convex hymenium. However, none of the species 

studied by me, from specimens or descriptions, showed any characteristics 

revealing close affinity of Capillipes. 

The most remarkable character of Capillipes is the verrucose, 

head-like ends of its paraphyses. This type of paraphyses seem to 

be extremely rare in the whole order Helotiales. 

In spite of the comprehensive and careful studies on the taxonomy 

of Helotiales performed by NANNFELDT there are still a great number 

of imperfectly known members. Future studies may probably reveal 

the exsistence of fungi related to Capillipes) making it possibly to 

give this genus a correct place within the families of Helotiales. Now 

I shaH not try to force it into any of the families recognized. 

Capillipes cavorum is known only from the type collection. Most 

probably the fungus is not so rare as could be supposed by our present 

knowledge. The reason why it has not been found previously, 

and now only once, certainly lies in its inconspicuousness and its rather 

peculiar habitat ecology. Ii the small rodent's caves in alpine - 

and perhaps other - regions are carefully studied it will perhaps 

be found in many places. 


I am very much indebted to Prof. J. A. N ANNFELDT for valuable advise 

during my studies of thi s fungus, and to Docent H ARRY SMITH for his 

kind help at the preparation of the manuscript.

- 395- 

LIT E R A T U R E C I T E D. 

Dm'and, E. J. (1908): The G e o g los s a c e a e of North America. - 

Ann. Myc. 6: 387-477. 

Nannfeldt, J. A. (1932): Studien iiber die Morphologie und Systematik der 

nicht-lichenisierten inoperculaten Discomyceten. - Nov. Aet. 

Reg. Soe. Sei. Ups. ser. 4, 8(2): 1-368. 

(1942): The G e o g los s a c e a e of Sweden. - Ark. f. Bot. 

30 A(4): 1-67. 

Rizzini, C. T. (1952): On a new Brazilian Hemiliehen. - Arq. Jard. Bot. 

Rio de Janeiro 12: 139-144. 

Uppsala, January 1956.

A SHORT SURVEY 

OF DANISH NEMATOPHAGOUS FUNGI 

By AUDREY M. SHEPHERD 

School of Agriculture, University of Cambridge 

The first discovery of a fungus which captured, killed and consumed 

nematodes took place in 1888, when ZOPF recorded that Arthrobotrys 

oligospora) earlier described by FRESENIUS (1850) as asaprophytic 

species, was in fact also parasitic upon nematodes. Since then 

DRECHSLER, DUDDINGTON and others have described some 80 species 

and the morphological diversity amongst these fungi is now well 

established. There are two main groups, t h e e n d o z o i c, o b l ig 

a t e p a r a s i t e s which only exist independently of the host in the 

form of spores, and t h e n e m a t o d e - t r a p p i n g f u n g i, existing 

as saprophytes in the absence of nematodes but stimulated by 

their presence to form trap s for their capture. These traps, which 

are specialised branches of the mycelium, are of four main types; the 

adhesive network, the adhesive knob, the non-constricting ring, and 

the constricting ring, all of which have been fully described by 

various authors. 

In Denmark, 19 species of nematophagous fungi were recorded 

during a brief survey carried out in 1952-54, which comprised mainly 

an investigation into the incidence of predacious species in soH. An 

analysis of the records from 130 collections of various kinds from 

widely distributed parts of the country is shown in Table L 

One species, Harposporium crassum) was described for the first 

time (SHEPHERD 1955) and another, Haptoglossa heterosporr.t 

DRECHSLER, was recorded for the first time in Europe. As far as the 

Danish mycoflora is concerned, all the species listed in Table 1, with 

the exception of Arthrobotrys oligospora, are almost certainly first 

records. O. ROSTRUP (1916, 1935) reported several species of Arthro- 

- 396-

- 397- 

TABLE 1. 

Occurrence of predacious fungi in various substrata in Denmark 

Analysis of records from 130 collections 

Species 

Leaf 

mouid. 

Rotting 

wood 

Decayed 

plant 

remain 

s 

Moss Soil l\Iisc. Total 

Acrostalagmus obovatus - - - 1 - - 1 

Arthrobotrys conoides ...... 

A . cladodes var. macroides 

- 

- 

I 

- 

- 

- 

- 

-- 

2 

2 

1 

- 

- 

2 

3 

A . dactyloides .............. .. - - - 1 1 - 2 

A . musiformis .... ... .... .... . - - 1 .. - 4 - 5 

A . oligospora .............. ... 2 - 1 1 35 - 39 

A. robusta .. .... ............... 2 - 1 - 2 - 5 

Dactylaria brochopaga ... 1 - - - - - 1 

D. psychrophila .......... .. .. - - - l 1 - 2 

D. thaumasia var. longa - - - - 7 - 7 

Dactylella cionopaga .... .. ·1 - - 1 1 - 6 

D. doedycoides .... .... ...... . 1 1 - - - - 2 

D. ellipsospora ...... ......... - 1 - - - - 1 

Haptoglossa heterospora - - - - - 1 1 

H arposporium anguillulae 3 - - - 1 -- 4 

H . crassum .......... ... ....... - - - - - 1 1 

H. lilliputanum ............... 1 - - - - - 1 

N ematoctonus tylosporus 1 - - - - - 1 

Stylopage hadra ... ... ... .. .. 1 - - - - - 1 

Total records 16 2 3 7 55 2 85 

No. of collections 19 11 5 17 76 2 130 

botrys) Dactylaria and Monacrosporium in Denmark, including the 

folIowing: 

Arthrobotrys oligospora FRES. 

A. longispora PREUSS 

A. superba CORDA 

Dactylaria echinophila MASSAL. 

D. acicularia ROSTRUP 

Monacrosporium elegans OUD. 

M. oxysporum SACC. 

Of these, only Arthrobotrys oligospora FRES. corresponds definitely 

to any of the present records, although A. longispora PREUSS may 

perhaps be synonymous with A. conoides DRECHSLER and Monacro-

- 398 - 

sporium elegans OUD. with Dactylella ellipsospora (PREUSS) GROVE. 

A. superba CORDA is now known to be nematode-trapping but none of 

the others can with certainty be regarded as synonymous with any 

of the predacious species. O. ROSTRUP does not mention predacious 

activity in any other than A. oligospora J for which he refers to a 

publication by EMIL CHR. HANSEN (1890), where ZOPF'S description 

of A. oligospora is quoted. Some of ROSTRUP'S records are again listed 

by LIND (1913). 

Whilst the primary aim of the survey was to establish as far as 

possible which species were to be found in the country, account was 

also taken of the type of location and the substratum from which 

collections were made. The results indicate that leafmould from rather 

damp situations was the richest source of predacious fungi. From 

nineteen collections, sixteen records were obtained comprising nine 

species. Beech litte r, however, seemed rather less suitable than that 

from other deciduous trees, possibly on account of its much slower 

rate of decomposition. Soil gave a high yield, sixty-two records being 

obtained from se vent y collections, although only ten different species 

occurred. Moss and rotting wood, which other workers have found to 

yield predacious fungi in abundance, were not very productive in 

this survey. 

Considering the various locations from which collections were 

made, the reed-swamp association appeared to be a very favourable 

habitat, yielding both quantity and variety. Beechwood was less 

favourable, and boggy situations proved to be ve ry unsatisfactory, 

eleven collections yielding only one record. The incidence of predacious 

fungi in soil had not been investigated until DUDDINGTON 

(1954) published an introductory survey from British agricultural 

soils. The outstanding point which emerged from a survey of Danish 

agricultural soils from widely distributed districts was the frequency 

of appearance of Arthrobotrys oligospora FRES. This fungus, apparently, 

is not noticeably affected by any variations of hydrogen ion 

concentration, organic matter content or soil structure, since it was 

found over almost the whole range of each of these factors. From 

the results obtained, which corroborate DUDDINGTON'S results in 

Britain, Arthrobotrys oligospora e m e r g e s a s a n i m p o r t a n t 

element in the soil mycoflora and can almost 

c e r t a i n l y b e r e g a r d e d a s a n e n d e m i c s o i l s p e c i e s.

- 399- 

The folIowing is a description of the generic characteristics and 

specific details of those nematophagous fungi found in Denmark. 

Further investigations would undoubtedly yield many more species. 

ACROSTALAGMUS 

Mycelium branched, septate, endozoic, giving rise to fertile branches 

emerging from the host. Whorls of flask-shaped phialides borne at intervals 

on the fertile branches, from which aseptate conidia are produced. 

Hyphomycetes. 

Acrostalagmus obovatus DRECHSLER, Phytopathology 31, p. 

784,1941. 

In moss (DicraneZZa sp.), Sorgenfri, Sept. 1952. 

Mycelium 2Ji wide, fertile branches 50-400Ji long. Phialides 

2-3 per whorl; conidia very small, 2 Ji across, slightly egg-shaped. 

HARPOSPORIUM 

Mycelium branched, septate, endozoic, giving off fertile hyphae emerging 

from the host, these bearing spherical or subspherical branches with 

short sterigmata, from which arcuate conidia are produced. Hyphomycetes. 

Harposporium anguilIulae LOHDE, Tageblatt 47. Versamml. 

deutsch. Naturforscher u. Arzte in Breslau, p. 203, 1874. 

In rotting leaves (UZmus and Sambucus)) Sorgenfri, Slotsparken, 

Sept. 1952 and leafmould, Rungsted, Dec. 1952. 

Very common parasite of nematodes. Conidia 13-17 Ji around the 

arc. Chlamydospores produced on the endozoic mycelium. 

Harposporium lilIiputanum DIXON, Trans. Brit. mycol. Soc. 

35, p. 146, 1952. 

In leafmould (beech), Folehave, Dec. 1952. 

Endozoic mycelium 2-3 Ji wide. Fertile hyphae up to 80 Ji long. 

Conidia very small, 4-8Ji long around the arc, 0.5-1Ji wide. 

Harposporium crassum SHEPHERD, Trans. Brit. mycol. Soc. 

38, p. 47, 1955. 

In horse dung, Agård, Oct. 1952. 

Endozoic mycelium 5.5-6.5 Ji wide, cells rather short and broad,

- 400- 

somewhat thickwalled. Fertile hyphae 50-150p long, 1.5-2.5p 

wide. Subspherical branches 2.5-4.5 p in diameter, bearing conidia 

18-22 p long around the arc, 2-3 p wide. 

HAPTOGLOSSA 

Haptoglossa heterospora DRECHSLER, J. Wash. Acad. Sci. 

30, p. 246, 1940. 

This fungus appeared in nematodes (Ditylenchus dipsaci) isolated 

from infected celeriac. It consisted of an oval-shaped thallus which 

broke up into spores, these being released through a papilla penetrating 

the cuticle of the nematode. On release, the spores were of a 

rather irregular shape, 5.0-6.5 p wide. They then encysted, and on 

emerging from the cyst the empty membrane remained attached to the 

spore, which had now developed a tongue-shaped lobe. These infective 

bo dies were 5-10 p in width. 

b 

O·05mm. 

Fig. 1. Haptoglossa heterospora DRECHSLER. 

a, b, eelworms containing a thallus ; c, thallus, now a sporangium, showing 

cleavage lines; d, release of spores through a papilla penetrating the cuticle 

of a nematode; e, release of sporangiospores from a sporangium producing 

only a few spores; f, infective bodies with the cyst membrane still attached.

- 403 - 

Arthrobotrys cladodes var. macroides DRECHSLER, Myeologia 

36, p. 144, 1944. 

In moss, Sorgenfri, Sept. 1952 and Rungsted Dee. 1952. 

Trap meehanism adhesive networks, often extensive. In pure eulture, 

large storage hyphae, 8-12,.u wide, were observed. Conidia 

15-22 X 6.5-8.5,.u. Tips of eonidiophores ofte n widened and lobed. 

Arthrobotrys dactyloides DRECHSLER, Myeologia 29, p. 486, 

1937. 

In moss and soil, Sorgenfri, Sept. 1952 and Oet. 1953. 

Trap meehanism eonstrieting rings, mostly 20,.u in diameter. Conidia 

43-51 X 8.5-9.6,.u, in loose heads eontaining up to 12 eonidia. 

Terminal eell of trophie hyphae within the nematode swollen. 

I I 

0 '02.""" 

Fig. 4. Arthrobotrys dactyloides DRECHSLER. 

a, conidia; b, constricting rings, showing the three-celled ring and the 

two-celled stalk; c, constricting rings as they appear after stimulation; 

d, a captured nematode with trophic hyphae, showing the enlarged ultimate 

cell of the branch. 

Arthrobotrys musiforrnis DRECHSLER, Myeologia 29, p. 481., 

1937. 

In rotting corn stubble and soil, Sorgenfri, Sept. 1952. 

Trap mechanism simple adhesive networks, characteristic for the 

species. Conidia 32- 44 X 10-14,.u, in loose heads. 

b 

26*

I o-OZmrn I 

404 - 

Fig. 5. Arthrobotrys musiformis DRECHSLER. 

a, conidia; b, c, e, networks; d, f, captured nematodes showing the infection 

bulb and trophic hyphae. 

Arthrobotrys oligospora FRESENIUS, Beitdige zur Mykologie, 

p. 18, 1850. 

In soil from many parts of Denmark ; in leafmould, decaying 

plant remains and moss. 

Trap mechanism adhesive networks. Conidial ranges within those 

given by DRECHSLER, 22-32 X 12-20 p (slightly lower in pure culture), 

varying from strain to strain. Conidia in heads, or later in 

whorls owing to proliferation.

- 405- 

Arthrobotrys robusta DUDDlNGTON, Trans. Brit. mycol., Soc. 

34, p. 598, 1951. 

In leafmould, Sorgenfri, Slotsparken, Sept. 1952 and Rungsted, 

Dec. 1952; in decaying plant remains, Valby, March, 1953. 

Trap mechanism adhesive networks. Conidia 18-24 X 8-12p, in 

heads. 

DACTYLELLA 

Mycelium branched, septate. Conidia hyaline, 3-6-celled, borne singly 

at the apex of aerial conidiophores. Hyphomycetes) Moniliales) Moniliaceae) 

Phragmosporae. 

Dactylella cionopaga DRECHSLER, Mycologia 42, p. 30, 1950. 

In leafmould (beech), Sorgenfri, Sept. 1952 and Folehave, Dec. 

1952; in Sphagnum, Lyngby Mose, Sept. 1952. 

Trap mechanism adhesive networks, characteristic for the species, 

the cells being short and constricted at the septa. Conidia spindleshaped, 

4-5-celled, 40-50 X 18-24p . 

Daclylella doedycoides DRECHSLER, Mycologia 32, p. 454, 

1940. 

In leafmould (beech) and rotting wood, Sorgenfri, Sept. 1952. 

Trap mechanism constricting rings, 20-22p across. Conidia topshaped, 

28-38 X 16.5-24p, 3-celled. Conidiophores slightly swollen 

at the tip. 

Dactylella ellipsospora (PREUSS) GROVE, J. Bot. 34, p. 200, 

1886. 

In rotting wood, Sorgenfri, Sept. 1952. 

Trap mechanism adhesive knobs, sometimes proliferating to form 

simple networks. Conidia spindle-shaped, 43-50 X 10-13 p, mostly 

3-celled. 

DACTYLARIA 

Mycelium branched, septate, Conidia hyaline, 3-7-celled, in heads on 

aerial conidiophores. Hyphomycetes) Moniliales) Moniliaceae) Phragmosporae.

- 406- 

Dactylaria brochopaga DRECHSLER, Mycologia 29, p. 517, 

1937. 

In leafmould, Rungsted, Dec. 1952. 

Trap mechanism constricting rings, 25-30 p in diameter. Conidia 

cylindricaI, 4-celled, 33-41 X 6.6-8.1p. 

Dactylaria psychrophila DRECHSLER, Mycologia 36, p. 161, 

1944. 

In moss and soil, Sorgenfri, Sept. 1952 and Aug. 1953. 

Trap mechanism adhesive networks. Conidia ellipsoidal, 4--5-celled, 

48-63 X 21-27 p. 

Fig. 6. Dactylaria psychrophila DRECHSLER. 

a, b, eonidia; c, a eaptured nematode showing trophie hyphae; d, networks. 

Dactylaria thaumasia DRECHSLER var. longa DIXON var. nov. 

ined. 

In soil, Horsens, Amager, Maarsø, Næstved, Lyngby, Aug. 1952. 

Trap mechanism adhesive networks. Conidia 3-4-celled, 40.8- 

54.0 X 17.6-25.2p, central cell rather swollen. Conidia frequently 

germinating i n s i t u.

- 407- 

STYLOPAGE 

Stylopage hadra DRECHSLER, Mycologia 27, p. 206, 1935. 

In rotting leaves (Ulmus and Sambucus)) Sorgenfri, Slotsparken, 

Sept. 1952. 

Mycelium branched, aseptate, hyphae 3.5-5.0 p wide. Conidia 

aseptate, obovoid, 33-37 X 15-21p, containing rather granular cytoplasm 

and borne singly on aerial conidiophores. Nematodes captured 

by adhesion to the ordinary vegetative mycelium. Zoopagaceae. 


This work was carried out under the tenure of a Danish Government 

scholarship, awarded through The British Council. I am indebted to both 

these authorities and also to the Director and staff of Statens plantepatologiske 

Forsøg, Lyngby, for so kindly providing the necessary facilities. 

I should especiaIly like to thank Dr. C. L. DUDDINGTON for his continual 

help and advice and Dr. P. BOVlEN, Mr. KNUD LINDHARDT and Dr. R. A. 

DUNNING for their interest and criticism. 

The material here set out has formed part of a thesis for the Ph. D. 

degree in the University of London (SHEPHERD, 1955). 

REFERENCES 

Dixon, S. M. (1952): Predacious fungi from rotten wood. - Trans. Brit. 

mycol. Soc. 35: 144-148. 

Drechsler, C. (1935): A new species of conidial phycomycete preying on 

nematodes. - Mycologia 27: 206-215. 

(1937): Some Hyphomycetes that prey on free-living terricolous 


(1940): Three fungi destructive to free-living terricolous nematodes. 

- J. Wash. Acad. ScL 30: 240-254. 

(1940): Three new Hyphomycetes preying on free-living terricolous 


(1941): Some Hyphomycetes parasitic on free-living terricolous 

nematodes. - Phytopathology 31: 773-802. 

(1944) : Three Hyphomycetes that capture nematodes in adhesive 

networks. - Mycologia 36: 138-171. 

(1950): Several species of Dactylella and Dactylaria that capture 

free-living nematodes. - Mycologia 42: 1-79. 

Duddington, C. L. (1951): Two new predacious Hyphomycetes. - Trans. 

Brit. mycol. Soc. 34: 598-603. 

(1954): Nematode-destroying fungi in agricultural soils. - 

Nature, Lond. 173; 500.

- 408- 

Fresenius, G. (1850): Beitrage zur Mykologie. Heft 1, p. 18. 

Grove, W. B. (1886): New or noteworthy fungi. Part 3. - J. Bot., Lond. 

34: 129-137, 197-206. 

Hansen, E. C. (1890): Nye biologiske Undersøgelser hos Svampe. - Naturen 

og Mennesket 3: 129-132. 

Lind) J. (1913): Danish fungi as represented in the herbarium of E. Rostrup. 

Copenhagen. 

Lohde, G. (1874): Einige neue parasitische Pilze. - Tageblatt der 47. VersammIung 

deutscher Naturforscher u. Arzte in Breslau: 203- 

206. 

Rostrup, O. (1916): Bidrag til Danmarks Svampeflora. I. - Dansk bot. 

Arkiv 2, Nr. 5: 1-56. 

(1935): Bidrag til Danmarks Svampeflora. II. - Dansk bot. 

Arkiv 8, Nr. 8: 1-74. 

Shepherd, A. M. (1955): Harposporium crassum sp. nov. - Trans. Brit. 

mycol. Soc. 38: 47-48. 

(1955): Some observations on the distribution and biology of 

fungi predacious on nematodes. - Thesis for degree of Ph. D. 

The Library of Univel'sity of London. 

Zopf, W. (1888): Zur Kenntnis der Infections-Krankheiten niederer Thiere 

und Pflanzen. - N ova Acta Leop. Carol. Deut. Acad. naturf. 52: 

314-376. 

Cambridge, J anuary 1956.

DISTRIBUTION OF TRICHOLOMA GAMBOSUM 

(FR.) GILL. AND BOLETUS GRANULATUS L. EX FR. 

IN NORWAY 

By JENS STORDAL 

It is a well-known faet that several higher plants prefer calcareous 

soH, requiring at the same time particular climatie conditions, e. g. 

in respect of temperature. This is an important factor in the distribution 

of the plants (ep. HULTEN 1950). 

However, in the case of Basidiomycetes only ve ry little is known 

in this matter. All the same, in several references it is suggested 

that Tricholoma gambosum} Boletus granulatus} and others are particularly 

fond of lime. This induced me to procure information about 

Norwegian finds to find out whether their distribution might verify 

these suppositions. The two species mentioned above are well known 

to most of us who are engaged in the study of fungi. The faet that 

they are delicious mushrooms as well, has contributed to the comparatively 

great attention that has been paid to them. 

Everything that we know about the occurrence of these species 

in N orway before 1905 is collected in "Norges Hymenomyceter" 

(BLYTT & ROSTRUP 19,05). The information given here is based on 

collections and records by A. BLYTT, E. HENNING (1885), K. O. 

BJORLYKKE, J. OLSEN (dr. O. Sopp), and P. HENNINGS (1904). In the 

last twenty years especially much information has been gained 

from Eastern Norway by CHR. FR. BOHME (1935-52, 1938). Also 

in Trondelag the species are known, mentioned by O. A. HOEG (1942, 

1943, 1946) in his reports. 

With a view to procure further information I arranged for sending 

out -inquiry forms to the people interested all over the country. In 

-409 -

- 410- 

this way answers have been submitted, stating new localities or veri 

fying the non-existence of the species in certains parts of the country. 

The existing herbarium material is available in the Botanic Mu 

seum of Oslo, being marked in the lists with (O). 

1. Tricholoma gambosum (FR.) GILL. 

In N orway this species has only been found in spring-time, in 

May and June. The earliest date recorded, to my knowledge, is May 

10, 1937. However, it is only in the latter half of May it can be said 

to appear in larger numbers. Then it usually keeps for a month's time. 

In July it is only seldom to be seen, but in the Oslo area it may some 

times keep till well into the middle of J uly (1938, 1951). 

Fig. 1. The distribution of Tricholoma gambo8um 

in Norway.

- 411- 

T. gambosum is recorded by BLYTT (1905: 26) to grow in mixed 

spruce forest. This has also been verified by several others later. At 

Grav in Bærum it was found growing on a dry crest under pine trees. 

On Kråkeroy it occurred among grass both in thick brush-wood and 

on more open grass plains among frondage trees and scrubs. 

Even BLYTT stated that it occurred in several places in the lower 

areas near Oslo. A closer look at the particular localities will show 

that they can all be located in Oslo or on the western side of the 

town. Reports made by BORME also verify these facts. S. G. SUNDBYE, 

who has been collecting mushrooms in Oslo for years, has been 

aoquainted with a great many localities which have given very good 

yield. 

On the western side of the Oslofjord T. gambosum has been recorded 

in the lowland down to Sandefjord. In ostfold the Fredrikstad 

area has ,been subjected to thorough investigations by W. RAMM. 

He states that T. gambosum in that area has only been found on 

Kråkeroy, where it was found growing in several localities. The 

ground in which it has been found, has always contained shell fragments. 

In the interior of Eastern Norway it has so far been recorded up 

to Toten and Vardal. The interior of the country with its broad 

valleys stretching along Eastern N orway has not been subjected to 

closer investigations when larger fungi are concerned, and this is 

particularly the case with spring fungi. - In Trondelag it reappears 

in the lower areas around the Trondheim fjord, but here it is very 

rare. 

LOCALITIES 

Ostfold 

K r å k e r o y: Enhus, Holte, Strålsund and several places along 

the road between Enhus and the church 1951/55 (W. RAMM). 

Oslo 

Several places in Oslo (A. BLYTT 1905: 26); Malmoya June 17, 

1945 (J. STORDAL) ; Hauketo (S. G. SUNDBYE) ; V. Aker June 2-July 

17, 1950, May 18 - June 23, 1952; o. Sogn June 20 and 29, 1951; 

Ullevålsalleen June 29, 1951; Bergsalleen July 4, 13 and 16, 1951 

(A. BRATSBERG) ; Dronningberget on Bygdoy May 24, 1885 (K. O. 

BJORLYKKE) (O); Gaustad, Frognerparken, V. Gravlund, Slottsparken,

- 412- 

Parkveien, Skoyen, Hofsbyen, Vækero, Bjornsletta, Valler-Gjettum 

and Fleskum (S. G. SUNDBYE). 

Akershus 

Many places around Oslo (A. BLYTT, CRR. FR. BORME). S k e d sm 

o: Leirsund (I. A. LYSEBRAATE). B æ r u m: Grav (l. A. LYSE 

BRAATE, J. STOKKE); Kalvoya May 31, 1953 (F.-E. ECKBLAD) (O); 

V. Sandvika June 7, 1953 (F.-E. ECKBLAD) (O). A s k e r: Gullhella 

1944 (H. ROED). 

Oppland 

T o t e n (J. OLSEN). Var d a l (J. OLSEN). 

Buskerud 

H u r u m: Tofteholmen (J. EGELAND 1921). 

Vestfold 

B o r r e: Falkensten (J. OLSEN). S a n d e f j o r d and the surrounding 

country (The mushroom controller in Sandefjord) . 

Nord -Trondelag 

.s t j o r d a l: Vikan 1942 (O. A. HOEG). Ve r d a l: Svinhamarmarka 

near the sea-side (K. BRAARUD). 

2. Boletus granulatus L. ex FR. 

This is one of the earliest larger fungi in the autumn season. If 

the weather is not too dry, it appears in the latter half of July. The 

earliest find ever recorded is dr. O. Sopp's material from Hovedoya 

near Oslo, July 13, 1882. It usually is at its height in August, but It 

mayaIso keep till late in September, as for instance in Eidanger, 

Sept. 23, 1942. Last autumn a sample was even found as late as Oet. 

11 on Bygdoy. However, the drought in the summer of 1955 caused 

several anomalies in the normal occurrences. 

According to BLYTT (1905: 115) B. granulatus grows in spruce 

and pine forest. The same kind of localities is noted by RAMM on 

Kråkeroy, but here it als o appears among grass in glades. In Tronfjell 

it was common in spruce forest and by road-sides (E. HENNING 

1885: 67). Pines are always to be found near the localities, and most 

likely it lives in mycorrhiza with this kind of trees.

- 413- 

Fig. 2. The distribution of Boletus granulatus 

in Norway. 

BLYTT noted it as being frequent in Vestre Aker, Bærum and 

Asker, and P. HENNINGS (1904) found it "in vast quantities" June 

1902, in Kongsskogen on Bygdoy (Oslo). AIso later it has been 

ascertained that thi s fungus is not rare in Oslo and in the districts 

west of the town. It appears in Vestfold and Buskerud, and reappears 

in the south in the lime area near Brevik in Telemark. 

In Ostfold the same factors come into play as in the case of T. 

gambosum. It has only been seen on Kråkeroy in three localities where 

the soi! was rich in shell fragments. 

Up in the valleys of Eastern Norway a few more localities are 

known for this species than for T. gambosum. This is probably due 

to the fact that there are a greater number of botanists out practising 

at the time when B. granuZatus fructifies.

- 414- 

Then this species, too, reappears in Trondelag along the Trondheim 

fjord, but also thi s species is rare here. 

OsUold 

LOCALITIES 

K r å k e r o y: Fuglevik Aug. 11, 1951, July 24 and 29, 1953, July 

17, Aug. 17 and 20, 1954, Aug. 22 and 30, 1955 (W. RAMM); Sandbukta 

July 16, 1953 (W. RAMM); Barkedalen Aug. 1953 (W.RAMM). 

Oslo 

Rustadsaga in ostmarka (R. RUl); Ulvoya 1942 (B. BULL) ; Gaustad-Sognsvatn 

Aug. 18, 1940 (CHR. FR. BOHME); y. Aker Sept. 9, 

1950 (A. BRATSBERG) ; V. Aker (A. BLYTT 1905: 115); Sogn Sept. 4, 

1951 (A. BRATSBERG) ; Bygdoy June 1902 (P. HENNINGS 1904: 17); 

Kongsskogen on Bygdoy Oct. 11, 1955 (L A. LYSEBRAATE) (O); 

Hovedoya July 13, 1882 (J. OLSEN) (O). 

Akershus 

E i d s von: SW. of Eidsvoll st. Sept. 3, 1942 (CHR. FR. BOHME) ; 

B æ r u m and A s k e r: Common in the lower areas (A. BLYTT 

1905: 115). 

Hedmark 

R a m a r (J. OLSEN); A l v d a l: The crest E. of Alvdal st. Aug. 

20, 1943 (CHR. FR. BOHME); Around Tronfjell July 30, 1884 (E. 

HENNING 1884: 67); T Y n s e t: Around Tronfjell July 30, 1884 

(E. HENNING 1885: 67); F o Il d a l: Enunda-Arlet (A. BLYTT 1905: 

115). 

Oppland 

L u n n e r: Roa, near the old road to Bergen (H. RUl). 

Buskerud 

K o n g s b e r g: W. of the town Aug. 28, 1940 (CHR. FR. BOHME). 

Vestfold 

S k o g e r (A. BLYTT 1905: 115).

Telemark 

- 415- 

E i d a n g e r: The areas around Heistad Sept. 9, 1942 (CRR. FR. 

BORME). 

Sor -Trondelag 

Strinda: W. of Lianvatn (O. A. HOEG 1946: 22). 

Nord-Trondelag 

V e r d a l: Vera, about 400 m above sea level (K. BRA AR UD ) ; 

S n å s a: F'insåsskogen Sept. 22, 1944 (O. A. HOEG). 

SUMMARY 

Although we have not nearly complete knowledge of the distribution 

of Tricholoma gambosum and Boletus granulatus in Norway 

as yet, certain interesting tendencies seem to reveal themselves which 

might induce us to closer investigations. Apparently both species 

show a rather similar distribution, a fact that is reminiscent of 

several heat and lime loving phanerogamia, e. g. Polygala amarella 

( NORD HAGEN 1940: 426). 

In the Oslo area, which must be said to be well investigated, they 

undoubtedly prefer the Silurian in Oslo and on the western side of the 

Oslofjord. East of the Oslofjord this calcareous rock is on the whole 

lacking. Here the localities are to be found near the fjord, an occasiGnal 

occurrence of lime being left in the soil, e. g. in form of shell. 

An exception to this are finds from Rustadsaga in bstmarka. 

On the western side of the Oslofjord they extend over Vestfold. 

B. granulatus also reappears in the numerous calcareous are as around 

Brevik. 

Up in the country they follow our warm valleys in the interior of 

the east. The interesting plant mountains around Tronfjell seem to 

offer favourable conditions for B. granulatus at least (E. HENNING 

1885), and here it gets high into the pine forest. - Then we again 

rediscover both species around the Trondheim fjord, but here they 

are rarer. 

; In Southern, Western and Northern Norway neither has been 

noted even though they have been searched for in several places both 

in outer and inner districts.

- 416- 

LITERATURE 

Blytt, A. & Rostrup, E.: Norges Hymenomyceter. - Vidensk.-Selsk. 

Skrifter L Math.-N aturv. Kl. No. 6. Christiania 1905. 

Bøhme, Chr. Fr.: Våre Nyttevekster. Heggedal 1935/ 52. 

: Oslo Helseråds kontroll av torvfort sopp gjennom 6 år. - 

Nytt Mag. for Naturv. B. 78. Oslo 1938. 

Ferdinandsen, C. & Winge, ø.: Mykologisk Ekskursionsflora. 2. Udg. København 

1943. 

Henning, E.: Bidrag till svampfloran i Norges sydligare fjelltrakter. - 

ofvs. Kongl. Vetensk.-Akad. Forh. Stockholm 1885. 

Hennings, P.: Beitrag zur Pilzflora der Umgebung Christiania (Halbinsel 

Bygdo). - Nyt Mag. for Naturv. Christiania 1904. 

Hulten, E.: Atlas over vaxternas utbredning i Norden. Stockholm 1950. 

Høeg, O. A.: Det Kongelige Norske Videnskabers Selskab. Museet. Årsberetning. 

Trondheim 1942. 

: Notater om sopper i Trondelag og Nordmore. - Våre Nyttevekster. 

Heggedal 1943. 

: Norsk Botanisk Forening. Trondelagsavdelingen. Årsmelding 

for 1944. - Blyttia. Oslo 1946. 

Nordhagen, R.: Norsk Flora. Oslo 1940. 

Ski, Norway, February 1956.

LACTARIUS GROENLANDICU5 SP. NOV. 

By FREDE TERKELS EN 

Thanks to a grant from The Danish National Scientific Foundation 

in August-September, 1955, I was able to make a journey to West 

Greenland for the purpose of collecting Macromycetes) especially 

BoZetaceae and Agaricaceae) in the regions north of the Arctic Circle 

which in that respect had so far been rather unexplored. 

Of course, it was a revelation for me to make investigations of 

the fungus flora of areas which, practically speaking, were entirely 

virgin. I came across a large number of strange fungi, and many 

species from these regions will probably appear to be undescribed, 

especially species within the genera RussuZa and Lactarius. 

The fungi of Greenland may in many cases be so closely related 

to well-known European and American species that it would be right 

to consider them just as varieties of these. But in other cases it is no 

doubt possibIe to find species of fungi which have never been described 

before. Such species may, of course, resemble some species 

hitherto known, but may nevertheless possess certain characteristics 

of their own which show that they are separate species, rather than 

varieties of other species. In my opinion, thi s applies to the species 

of the genus Lactarius which I am going to describe in the following. 

- As this fungus seems to have been found only in Greenland, I find 

it appropriate to name it: Lactarius groenZandicus. 

Lactarius groenlandicus sp. nov. 

F i n d s: Col!. No. 16 (F. T.), Hassells Fjeld, Sondre Stromfjord, 13/VIII, 

1955. - Co l!. No. 49 (F. T.), Godhavn, between the town and the arctic station, 

17/VIII, 1955, (colour-photos were taken, see Fig. 1). - Also found on 

the heath between the arctic station and the Rodeelv stream, 19/VIII, 1955, 

22/VIII, 1955, and 27/VIII. 1955. 

FRIESIA V - 417- 27

- 418- 

Fig. 1. Lactarius groenZandicus sp. n. - Photo. FREDE TERKELSEN. 

H a b i t a t: Always found in shrubs of willow (Salix ) or near such 

shrubs. 

D e s c r i p t i o n: Cap: 7- 11 cm broad, convex-expanded, often with 

ovaloutlines and then alittie concave at the centre between two flat "sadd 

Ies" ; whitish, turning yellowish; slightly viscid; later dry and somewhat 

shining; smooth, but the incurved margin at first very shaggy. - Gills: 

rather narrow; moderately crowded; whitish, at last straw-coloured, stained 

rust y or brownish when cut or bruised. - Milk: white, unchangeable, copious, 

mild. - Stem: sometimes a little eecentric, 2- 5 X 1--1.5 cm, whitish 

or yellowish, sometimes pitted with lemon-yellow spots, cylindrical or ae uminate, 

hollow. - Flesh: white, firm, with apleasant, sweetish smell of 

Lactarius helvus or curry, which gets stronger when dried. 

Gregarious, often almost hidden by earth and foliage sticking to the 

viscid surfaee of the capo 

S p o r e s: Ellipsoidic or subglobose, minutely echinulate or reticulate, 

(6.5-) 7-8 X 5-5.5 fl. See Fig. 2. 

V e i l made up of hyaline fibrils, 3-4 fl broad; similar hyphae densely 

matted on surfaee of pileus, not gelatinous. 

C y s t i d i a on face of lamellae hardly protruding over mature basidia, 

about 30- 50 X 6- 7 fl. 

DIAGNOSE LATINE 

Pileus 7-11 cm latus, e convexo expansus, centro plus minusve 

concavo, albidus, dein lutescens, e viscido siccus, glaber, margine 

primitus inflexo et floccoso-barbato. 

LamelIae subangustae, subconfertae, albidae, 

postremo stramineae, vulneratis ferrugineo-maculatis. 

Lac album, invariabile, copiosum, mite. 

Stipes 2-5 cm X 1-1.5 cm, interdum 

subexcentricus, albidus vel lutescens, saepe 

scrobiculis citrinis ornatus, cylindricus vel 

sursum attenuatus, cavus. 

Caro alba, firma, odore subdulci fere Coci 

nuciferae} sicca Lactarium helvum vel "curry" 

redolente. 

Fig. 2. Spores of Lactarius 

groenZandicus 

Del. F. H. MØLLER 

X 2000.

- 419- 

Sporae (6.5-)7-8 X 5-5.5 1-', ellipticae vel subglobosae, minute 

echinulatae vel minutissime et parce reticulatae. 

Gregarius, in salicetis, saepe sub sabulone et foliis fere occultus. 

Typus die 16' Augusti 1955 sub Salice ad sinum Sdr. Strømfjord 

Groenlandiae occidentalis sub numero F. T. 16 lectus, in Museo Botanico 

Hauniensi depositus. 

This big and beautiful species, the most prominent Lactarius I 

saw in Greenland, should be classed in the section EuZactarius) subsection 

L CZaricoZorini teste NEUHOFF. It does not seem to be found 

anywhere outside Greenland. 

Lactarius groenZandicus is very much like L. controversus (PERS.) 

FR., but is not acrid and never zoned or stained with pink or red; it 

also differs in habitat. From L. resimus FR. it differs in the milk not 

turning sulphur-yellow. - The two American species which seem to 

be most closely related to L. groenlandicus} viz. L. deceptivus Peck 

and L. subveZZereus Peck, are both characterized as "very acrid", and 

also their spore dimensions are different. 


In the preparation of this paper I have had valuable assistance from 

Mr. F. H. Moller) Senior Schoolmaster, who also has made the drawings of 

the spores, as well as from Mr. Morten Lange) Ph. D. Mr. E. Brockmeyer 

has been so kind as to go over the Latin diagnosis with a critical eye. I 

acknowledge with gratitude the help rendered me by these three gentlemen. 

Danebod Folk-Highschool, Als, Denmark, April 1956. 

27*

TRICHOLOMA VERRUCIPES FR. ELLER 

CLITOCYBE VERRUCIPES (FR.) MAIRE FUNNEN 

I HALSINGBORGSTRAKTEN, SYDSVERIGE 

Av HERVID VALLIN 

Forst några litteraturuppgifter om denna sallsynta, sporadiskt 

upptradande, vita, vackra svamp. 

E. FRIES: Hymenomycetes Europaei, Upsaliae 1874. - S. 43. Agaricus 

(Armillaria) verrucipes. Upptackt av v. POST 4.5.1870 nara 

Uppsala. "Odor farin ae recentis, gratus; facile esculentus, ut plurimi 

fungi stricte vernales. " 

P. A. KARSTEN: Rysslands, Finlands och den skandinaviska halfons 

hattsvampar. 1. Helsingfors 1879 (Bidr. t. kannedom af Finlands 

natur och folk 32). - S. 65: Clitocybe puellula KARST. Vit, hatten 

snovit, n. pucklig ; foten brun- eller svartfjallig, lamellerna fastvaxta. 

Karrjord, dikeskanter, r, 6, 9. Finland (Mustiala) . S. 20 upptages 

Armillaria verrucipes) lokal Uppsala. 

L. QUELET: Flore Mycologique de la France et des pays limitrophes. 

Paris 1888. - S. 272-273: Gyrophila verrucipes: Chair 

ferme, blanche, tres parfume, odeur de farine et de fruits. - 

Printemps. - En cercle dans les pres ou bois sablonneux. Jura. 

Comestible. 

G. BRESADOLA: lconographia Mycologica. Vol. II. Mediolani 1927. 

Tab. 68. Tricholoma verrucipes (FR.) QUEL.; QUEL., Fl. Myc., p. 272; 

Sacc., Fl. It. Crypt., Hym., p. 141. Armillaria verrucipes FR. in QUEL., 

Champs du Jura II, p. 304, t. 11, f. 1; FR., Hym. Eur., p. 43; Sacc., 

Syll. V. p. 78. Clitocybe puellula KARST., leon II, t. I, f. 31. .... odore 

pirino demum farinaceo, sapore grato; ..... 

-420 -

- 421- 

Hab.: in herbidis juxta vias, saepe caespitosus, autumno, raro. 

Area distr. : Europa. 

P. MAGNUS: Die Pilze (Fungi) von Tirol, Vorarlberg und Lichtenstein. 

Innsbruck 1905. - S. 296: Tricholoma verrucipes (Fr.) 

Bres. Italienisches Tirol: auf grasigen PHitzen an Wegrandern, meist 

rasig (Bres. II, p. 6.) 

A. RICKEN: Die BIatterpilze (Agaricaceae) Deutschlands und der 

angrenzenden Lander, besonders Oesterreichs und der Schweiz. I. Leipzig 

1915. - S. 330: Tricholoma verrucipes (Fr. 1873.*) - Bres.). Warzenftissiger 

Ritterling .... Fleisch weiss, riecht birnen-schl. mehlartig. 

An grasigen Orten, neben Wegen, meist rasig 8-11. Selten. 

P. KONRAD & A. MAUBLANC: Icones selectae Fungorum, VI. Paris 

1924-1937. - S. 333: Clitocybe verrucipes (FRIEs) R. MAIRE (1911). 

Syn .. Armillaria verrucipes FRIES ap. QUELET (1872); Agaricus ver 

rucipes FRIES (1874); Gyrophila verrucipes QUELET (1886); Tricholoma 

verrucipes BRESADOLA (1892); Agaricus puellula KARSTEN 

(1876); Clitocybe puellula KARSTEN (1879). 

Jura (QUELET), Grande-Chartreuse (R. MAIRE), Trentin (BRESA 

DOLA), Finland (KARSTEN), etc. 

E. BILLE HANSEN & F. H. MØLLER: Clitocybe verrucipes (FR.) 

MAIRE (Rufodet Tragthat) i Danmark. Friesia III (1944-48), Hefte 

2 (1945). - S. 101: "Paa Botanisk Forenings Svampeexkursion til 

Lillerød d. 8.10.1944 fandtes (E. BILLE HANSEN) ved en Sti paa fugtig 

Bund i Allerød Mose mellem Græs (Molinia coerulea) nær Eg (Quercus 

robur) og Rødel (Alnus glutinosa) en Flok paa 5-6 Individer af en 

særpræget, hvid Tragthat, som senere bestemtes (F. H. MØLLER) til 

ovenstaaende Art." 

E. INGELSTROM: Svampflora, Stockholm 1940. - S. 184: Tr. verrucipes 

FR., parlmusseron ..... uppgiven for Uppsala- och Goteborgstrakten. 

5 km oster om centrum av Halsingborgs stad ligger Vasatorps 

skog, i vilken finns ett tamligen oppet, grasbevuxet område, en rest av 

en gammal faladsmark med glest stående enbuskar och bjorkar och 

en sparsam vegetation av ljung (Calluna vulgaris) och odon (Vaccinium 

uliginosum) J blåtåtel (Molinia coerulea) m. fl. gras och orter. 

I utkanten av faladsmarken ligger en liten damm, kring vilken vaxer 

klibbal (Alnus glutinosa) med kuddar av Leucobryum glaucum och 

Sphagnum i sankorna. 7-8 m vaster om dammen fann jag på gras- 

* ) i. e. "Hym. Eur.", 1874!

- 422 - 

Fig. 1. Tricholoma verrucipes FR. 

Vasatorps skog, Halsingborg, 28.9.1941. 

mark med Galium saxatile och Rhytiadelphus squarrosus nara en 

enbuske den 28.9.1941 5 st. glansande, vita, vackra parlmusseroner, 

Tricholoma verrucipes FR., med sina synnerligen karakteristiska 

svartbruna flockprickar (darav namnet "parlmusseron") på foten, 

vilka på oversta delen av foten, %-1 cm under hatten, sitta tatare 

och darigenom gora en antyd an om ett velum. 

Enar parlmusseronen utforligt beskrivits av BILLE HANSEN & 

F. H. MØLLER (1945), skall jag endast i korthet namna några ord 

om den. 

De 5 ex. stodo nara varandra, hattarna vore plana, mattgIansande, 

vita till mycket svagt ljusbruna med en karakteristisk något morkare 

liten puckel i mitten. Hattarnas diametrar vore 6,5 X 6,5, 5,5 X 5,5, 

4 X 3,5, 6 X 6 och 7 X 7 cm; fotterna voro resp. 7, 6, 5, 7.5, 7.5 cm långa 

samt något tjockare nedtill ; alltså av betydligt mindre dimensioner 

an hos de i Allerøds mosse (Danmark) funna exemplaren. Jag återfann 

några individ av samma art på samma lokal den 1.11.1942, 13.9. 

1946, 2.10. och 16.10.1949 samt 6.10 och 12.10.1950. Den 12.10.1950 

var hattdiametern hos ett ex. 6,5 X 5,5 cm, foten 5 cm och upptill med 

en diam. av 7 X 5 mm. Ett annat ex. (det storsta) hade en hatt av

- 423 - 

Fig. 2. TTicholoma ven'ucipes FR. 

Vasatorps skog, Halsingborg, 12.10.1950. 

8 X 7,5 cm, foten var 7 cm lång. Sedan dess har jag tyvarr ej återfunnit 

dem. Enbusken, under vilken svamparna vaxte, ar borthuggen, 

men detta har troligen icke med svampens forsvinnande att gora. 

Aldre ex. hade hatten svagt inbuktad. Lukten hos svampen var forst 

angenam, men ett pioekat ex. hade efter några timmar en sotaktig, 

acklig lukt. Ett torkat ex., som jag haft i ett kuvert sedan den 

2.10.1949 har en fran kryddlukt, svår att definiera. 

Av ett visst intresse ar, att den enda kanda danska lokalen i 

Allerøds mosse och den i Vasatorp vid Halsingborg blott ligger på 

c:a 30 km:s avstånd från varandra. For ovrigt ar ju den sallsynta 

svampen kand endast från ett fåtal lokaler i Europa, belagna på

- 424- 

stora avstånd från varandra. Dess sporadiska upptradande ar ocksa 

en gåta. Intressant vore, om man genom mykologers medverkan i 

olika lander exakt kunde få reda på, nar och var svampen funnits 

under t. ex. de sista 10 åren. 

SUMMARY 

Tricholoma verrucipes Fr. is found in Scania, Sweden. 

Tricholoma verrucipes FR. (Clitocybe verrucipes (FR.) MAIRE) is 

only known from a few localities in Europe. In Sweden the species 

was found near Uppsala in the year 1870, in Gothenburg(?) and in 

the neighbourhood of the town of Halsingborg during the years 

1941-1950. In Denmark it was found 1944 (good description of the 

fungus), and in the 19th century it was found in Finland, France and 

Italy. It would be of great interest to known the exact localities of 

this rare, beautiful and peculiar fungus, f. i. during the latest 10yearperiod 

in the different countries of Europe. 

Halsingborg, February 1956.

NÅGRA VASTSVENSKA SVAMPFYND 

Av S. WOLDMAR 

Svampfloran på Sveriges vastkust har behandlats i arbeten av 

O. ROB. FRIES, T. NATHORST-WINDAHL m. fl. Dess sardrag, praglade 

framfor allt av ett maritimt klimat, kunnaemellertid annu knappast 

skonjas. I den mån uppgifter om fynd och fyndorter bli kanda, klarnar 

dock bilden, och sammanstallda med de mykofloristiska undersokningar, 

som gjorts i skilda delar av Sverige och i de ovriga 

nordiska landerna, vidgas den vaxtgeografiska overblicken, och hittills 

outredda problem betraffande svamp arnas forekomst kunna måhanda 

harigenom lOsas. 

Foreliggande anteckningar avse fynd av anmarkningsvarda arter 

inom ett mindre område på den svenska vastkusten, omfattande mellersta 

BohusIan, speciellt Uddevallatrakten, och angransande del av 

Vastergotland (Vanersborg och V. Tunhem). Samtliga arter, som 

antecknats från området, ha ej kunnat medtagas, vilket hade varit 

onskvart, då harigenom en battre bild av floran hade kunnat erhållas. 

Den av LUNDELL & NANNFELDT i "Fungi Exsiccati Suecici" anvanda 

nomenklaturen har jag sokt fOlja, så långt det varit mojligt. 

Hymenomycetes 

Amanita muscaria L. ex FR. f. aureola KALCHBR. - Uddevalla, 

terrangen S om sjon Sarven, under bjork och Salix) 5.10.1947. 

Denna form stammer val med såval text som avbildning (PI. 3 C) 

hos J. E. LANGE. 

-425 -

- 426- 

A. phaZZoides (VAILL. ex FR.) SECR. - Skafto, Vagerod, under ek, 

27.7.1953. Åven antecknad från S. Bohuslan (L. ÅKERBLOM in Htt.) : 

Marstrand, Koon, 1935 och flera foljande år; Lycke, 1941. 

A. virosa FR. - Uddevalla, Rosserod sept. 1950; terr. S om Sarven, 

sept. 1952. 

Amanitopsis adnata (W. G. SM. ap. SAUND. & SM.) SACC. - Uddevalla, 

Bjorback, i gras under bjork, sept. 1948 (det. A. MELDERIS), i 

grus intill bjork (fig. 1), aug. 1953 (det. S. LUNDELL). - Artens 

taxonomi ar omtvistad. Enl. LUNDELL (in litt.) bor den av mig 

funna arten t. v. beteeknas som 

A manitopsis adnata SMITH sensu 

COOKE, då den val overensstammer 

med COOKE'S avbildning 

men icke med SMITH'S ursprungliga 

beskrivning. Den 

stammer aven val med LANGE'S 

bild a v den ringlosa formen 

(var. exannuZata) av Amanita 

junquiZZea QUEL. (PI. 2 D), som 

emelIertid vaxer i barrskog. 

Kategoriskt anmarker LANGE: 

"This is Amanitopsis adnata of 

W. G. SMITH. But even when the 

ring is entirely absent it cannot 

be mistaken for a true A manitopsis((. 

Fig. 1. A manitopsis adnata SMITH 

sensu COOKE. 

Uddevalla, Bjorback, Aug. 1953. Note 

the habitat! - Phot. by GUSTAF 

CARLSSON. 

Conocybe striaepes (CKE) 

LUNDELL. - U ddevalla, Porsen, 

14. 9. 1949 och flera foljande år. 

- Det forsta svenska f yndet av 

denna calcifila art, som har 

vaxer i starkt snackskalsbemangd jord. - Fung. exs. suec., n. 2049. 

Mycena quisquiZZiaris (Joss.) KUHNER. - Uddevalla, Sarven, på 

Phragmites} sedan 1950 (det. R. RYDBERG) (Fig. 2). - Denna art har 

av RYDBERG observerats på Carex paradoxa m. fl. arter i karr på Runmaro, 

Uppl., sedan 1948. Av JOSSERAND fordes den till OmphaZia medan 

CEIP inrangerat den under DeZicatuZa. En art som latt forbises (hatt:

- 427- 

2- 4 mm; fot 3- 5 mm) och som sakerligen ar allrnannare an de fåtaliga 

f ynden utvisa. Från Danmark ar den noterad av MORTEN LANGE 

(Sjælland: Maglemose och Buresø, på Molinia och Eriophorum vaginatum). 

M. buZbosa (CEIP) KUHNER. - Vg. Vanersborg, på Carex graciZis i 

sjon Vassbotten sept. 1953 (det. R. RYDBERG). - Denna art, som ar 

ny for Sverige, ar mindre och har kortare fot an den narstående allmanna 

M. styZobates. 

EnI. RYD BERG (in liU.) 

igenkannes den på sin 

gelatinosa hatthud. Å ven 

denna art forbises la tt 

(storlek ung. som foregående). 

PZeurotus chioneus 

PERS. U ddevalla, 

Bjorback, på forarbetat 

virke, okt. 1947, sept. 

1948. 

Fig. 2. Mycena quisquilliaris (JOSS.) KOHNER. 

Uddevalla, the lake Sarven, on Phragmites, 

Sept. 1955. X 3. Phot. by GUSTAF CARLSSON. 

P. atrocoeruZeus FR. 

ex FR. - Lane-Ryr. terr. NO om St. Koperodsjon, juli 1947, okt. 

1948. 

P. porrigens PERS. ex FR. - Ett flertal lokaler i Uddevallatrakten 

antecknade for denna art, som vissa år forekommer tamligen rikligt. 

Marasmius Zimosus BouD. & QUEL. - Uddevalla, Sarven, på Phragmites} 

juli-okt. 1951-1955. - En liten art (hatt: 2-3 mm; fot: 

10-15 mm, hårfin), latt igenkand på den for RotuZa-gruppen karakteristiska 

lamellringen. Jfr. ø. WINGE'S vackra teckning av M. rotuZa 

i FERDINANDSEN & WINGE: Mykologisk Ekskursionsflora, 1943, p. 218! 

Artnamnet ar missvisande, eftersom den ej vaxer i dy jord uta n på 

fuktiga, vissna blad, strån o.d. J. E. LANGE har sett den på Deschampsia 

caespitosa. 

Lentinus suavissimus FR. - Uddevalla, Graskarr, sept. 1946; terr. 

S om Sarven, sept. 1947; Bjorback, flera lokaler 1947 och de foljande

- 428- 

åren; Grindhult, skagen O om Grindsjon, 2. 12. 1949; Lane-Ryr, terre 

V om Hukeberg, sept.-nov. 1947. På samtliga lokaler på Salix. - Av 

lokaluppgifterna framgår, att denna svamp ej ar ovanlig i Uddevallatrakten. 

I ovriga delar av Sverige torde den doek vara sallsynt, vilket 

framgår av HARRY G. SVENSSON'S sammanstallning av tidigare fynd 

(SVENSSON 1944). Dessa uppgå till ett lO-tal, Troligen ar den forbisedd 

p. g. a. sin forekomst i ± tata Salix-snår oeh allmannare i vastra 

Sverige an i ovriga delar av landet. NATHORST-WINDAHL (1949) har 

sett den på flera stallen i Goteborgstrakten. - Fung. exs. suee., n. 

1774. 

Panus torulosus (PERS. ex FR.) FR. - Uddevalla, terre S om Sarven, 

sept.-okt. 1947 oeh flera foljande år. - Fung. exe. suee., n. 1771. 

SchizophyllurYl, commune FR. - Uddevalla, skogen O om Bjorbaeks 

gård, på asp, 1. 12. 1948; Salekarr, på asp, 5. 5. 1953. 

Trogia crispa (PERS. ex FR.) FR. - Uddevalla, skogskant e:a 300 

m O om Bjorbaeks skola, 19.11.1948. 

Asterophora parasitica (BuLL. ex FR.) SING. - Skaf to, Vagerod, 

på Russula delica, 17.8.1953. S. å. rikligt i V. Tunhem, dar den tidigare 

(1944) antraffats av NATHORST-WINDAHL. Denna saregna svamp, 

som ej antraffats utanfor ekens nordgrans (WOLDMAR 1954), synes 

vara beroende av hog luftfuktighet. Under den torra hosten 1955 stod 

ej något exemplar att finna vare sig i V. Tunhem eller vid Vagerod. 

A. lycoperdoides FR. ex FR. - Forshalla, Stenshult, på Russula 

nigricans, 21.9.1946 oeh de fOljande åren t. O. m. 1954. En storvuxen 

form (LANGE'S f. major?) har jag anteeknat från denna lokal 29.9. 

1949. 

Cantharellus umbonatus PERS. ex FR. - Liksom C. aurantiacus 

synes denna svamp "ersatta" vanlig kantarell under torra hostar. Så 

var fallet i Uddevallatrakten 1947 oeh 1955. 

Entoloma bahusiense LUNDELL. - Uddevalla, Bjorbaek. - En intressant 

E. clypeatum narstående art, som årligen 1948-1953 fore-

- 429- 

kom rikligt på ett begriinsat område. 1954 och 1955 syntes den ej till. 

Marklig ar den korta fruktifikationstiden, c:a 10 dage r, i slutet av 

maj. - Fung. exs. suec., n. 2007 (originalkoll.). 

Stropharia Hornemannii FR. - Uddevalla, S och SV Sarven, i enstaka 

ex. under åren 1947-1953 och Uimligen rikligt hosten 1955; 

Lane-Ryr, teIT. V om St. Koperodsjon, okt. 1948. 

Coprinus Friesii var. (C. Friesii QUEL.?). - Uddevalla, Sarven, 

på Phragmites 26.6.1950, aven 1951-1954. - EnI. LUNDELL (in 

litt.) ar det ovisst om detta ar QUELET'S art. 

Hydnum corrugatum FR. - Uddevalla, S om Sarven, på bjork, 

juli 1948; Lane-Ryr, Skallerod, på bjork, sept. 1948. 

Clavaria abietina PERS. var. flaccida. - Lane-Ryr, Koperod, sept. 

1950. (Det. S. LUNDELL, som aven bestamt de fOljande Clavaria-arterna 

utom C. botrytis). 

C. botrytis PERS. - Uddevalla, skogskant S om Bjorbacks gård, 

sept. 1946 (det. T. NATHORST-WINDAHL); Kurod, sept. 1948. 

C. crocea FR. - Uddevalla, Bjorback, 1. 9. 1953. - Det andra 

svenska f yndet. Tidigare har den antraffats av NATHORST-WINDAHL 

i Bot. Tradgården i Goteborg. 

C. epichnoa FR. - Uddevalla, Graskarr, sept.-nov. 1947-1949. 

C. inaequalis MULL. - V. Tunhem, 20.8.1953. 

C. rugosa BULL. - Forshalla, i barrskog V om oresjon, 18. 9. 1940. 

C. corniculata SCHAEFF. ex FR. - Uddevalla, Kurod, på skalgrus, 

31. 8.1953. 

C. corniculata forma (?). - V. Tunhem, aug. 1953. - En intressant 

form, habituellt lik C. corniculata men gråblek och med 2-sporiga 

basidier. 

C. subtilis PERS. ex FR. - Uddevalla, Bjorback, i mossa vid ormbunksrotter, 

1.9.1953. 

C. pulchra PECK. - V. Tunhem, 1.9.1953.

-- 430 - 

Fig. 3. Clavaria cineroides ATKINSON or a related species. 

Viisterg. V. Tunhem, 20.8.1953. X 4. Phot. by GUSTAF CARLSSON. 

Clavaria Sp. - V. Tunhem, 20.8.1953. - EnI. LUNDELL (in litt.) 

ny art eller C. cineroides Atkinson. (Fig. 3). 

Solenia crocea KARST. - En forbisedd svamp, som synes forekomma 

Himligen allmant på aldre ex. av Matteuccia struthiopteris 

(WOLDMAR 1954). Jag har aven funnit den på Matteuccia vid Farum, 

Sjælland, 5. 8. 1954, vilket ar det forsta danska f yndet. 

Peniophora subcalcea Litsch. - Uddevalla, Graskarr, 20.5.1950. - 

En intressant art, som ej tidigare uppmarksammats i Sverige. Det. 

E. RUTH DEARDEN och H. S. JACKSON, Toronto, som aven haft tillgång 

till LITSCHAUER'S typmaterial. Bestamningen har också bekraftats av 

JOHN ERIKSSON. 

Discomycetes 

Cudonia con/usa BRES. - Lane-Ryr, i skogen S om St. Abborrvattnet, 

5. 9. 1950 (det. S. LUNDELL). En sammanstallning av tidigare

skandina viska fynd har gjorts 

av J. A. NANNFELDT (1942). 

Cyathicula coronata 

(BULL.) REHM. - Uddevalla, 

St. Uggelhult, på Matteuccia) 

23.9.1954 (det. J. A. NANN 

FELDT); Ramserod, på Dry 

opteris filix mas 15.10.1954. 

Rhizopodella melastoma 

(Sow. ex FR.) RICHON. - Uddevalla, 

Bjorback, på ljungrotter, 

30.4.1949, i enstaka 

ex. aven flera foljande år 

(det. J. A. NANNFELDT). - 

Artens taxonomi och fore 

431 - 

Fig. 4. Rhizopodella melastoma (SOW. ex 

FR.) RICHON. 

UddevalIa, Bjorback, April 1953. X 4. 

Phot. by GUSTAF CARLSSON. 

komst i Skandinavien behandlad av NANNFELDT (1949), som anser 

att den intar en mellanstallning mellan Urnula och Bulgaria. (Fig. 4). 

P I ectascal es 

Onygena equina WILLD. - Uddevalla, Graskarr, terr. SV om St. 

Koperodsjon, på ruttnande horn, 13.12.1948. 

Professor J. A. NANNFELDT, Uppsala, har hjalpt mig med artbestamningar 

och lamnat mig vardefulla uppgifter och goda råd under flera år. 

Hjalp med examinering har jag också fått av fil. dr. SETH LUNDELL, Uppsala, 

tradgårdsdirektOr T. NATHORST-WINDAHL, GOteborg, lektor JOHN 

ERIKSSON, Varnamo, och disponent R. RYDBERG, Stockholm. Min van, 

folkskolIarare GUSTAF CARLSSON, UddevalIa, har stallt de valtagna fotografierna 

till mitt forfogande. Till alIa ett varmt tack. 

SUMMARY 

Notes Oll Macromycetes from Western Sweden. 

The author reports about fort y finds of Hymenomycetes) Discomycetes) 

and Plectascales (one species) from the Swedish West Coast, 

especiaIly from the district ro und Uddevalla. The folIowing species 

are new to Sweden: Conocybe striaepes (CKE.) LUNDELL, Mycena bul 

bosa (CEIP) KUHNER, Entoloma bahusiense LUNDELL, Peniophora

432 - 

subcalcea LITSCHAUER, and one Clavaria sp., probably C. cineroides 

ATKINSON. A number of species, for instance Mycena quisquilliaris 

(Joss.) KUHNER and M. bulbosa (CEIP) KUHNER, Marasmius limosus 

BOUD. & QUEL., Rhizopodella melastoma (Sow. ex FR.) RICHON, 

Solenia crocea KAR ST. , and Cyathicula coronata (BULL.) REHM seem 

to have been overlooked, owing to their smallness and/or their special 

way of growing. A find of Solenia crocea KARST. is also reported from 

Denmark, where it has not been met with previously. 

LITTERATUR 

Ferdinandsen, C. & Winge, ø., (1943): Mykologisk Ekskursionsflora. - 

København. 

Fries, O. Rob., (1907): Anteckningar om Svenska H y m e n o m y c e t e r. 

- Ark. f. bot., Bd. 6, N:o 15. Stockholm. 

Lange, Jakob E., (1935-1940): Flora Agaricina Danica. Vol. I-V. - 

København. 

Lange, Morten, (1946): Mykologiske Iagttagelser i Danmark 1943-1945. 

- Friesia. Bd. III, H. 3. København. 

Lundell, S. & Nannfeldt, J. A., (1934-1953): Fungi exsiccati suecici, .... 

Fase. I-XLII. (pro parte). - Uppsala. 

Nannfeldt, J. A., (1942): The G e o g los s a c e a e of Sweden (with regard 

also to the surrounding Countries). - Ark. f. bot., Bd. 30 A., 

N:o 4. Stockholm. 

, (1949): Contributions to the Mycoflora of Sweden 7. A new 

winter Discomycete, U r n u l a h i e m a l i s N annfeldt n. sp., 

and a short account of the Swedish species of S a r c o s c yp 

h a c e a e. - Sv. Bot. Tidskr. Bd. 43. Stockholm. 

Nathorst-Windahl, T., (1945): Anmarkningsvarda fynd av H y m e n om 

y c e t e r i Bohuslan och Vastergotland. - Meddel. fr. Goteborgs 

Bot. tradgård. Bd. 16. Goteborg. 

, (1949): Anmarkningsvarda fynd av H y m e n o m y c e t e r 

i Bohuslan, Vastergotland och Daisiand. - Bot. Not. h. 3. Lund. 

Svensson, Harry G., (1944): Anteckningar om Karlstadtraktens skivlingflora 

II (Broskskivlingar: M a r a s m i a e och rodsporingar: 

R h o d o s p o r a e). - Medd. fr. Varmlands Naturhist. For. 15: 

1. Karlstad. 

Woldmar, S., (1954 a): Om utbredningen i Norden av gråkremling, 

A s t e r o p h o r a p a r a s i t i c a (BuH. ex Fr.) Sing. - Sv. 

Bot. Tidskr. Bd. 48. Stockholm. 

, (1954 b): S o l e n i a c r o c e a Karst. - en forbisedd svampart. 

- Friesia. Bd. V, H. 1. København. 

Bjorback, UddevaHa, januari 1956.

Indhold (fortsat) 

Side 

Marcel Locquin: Quelques Lepiotes nouvelles ou critiques. Lepiota 

helveola BRES. ss. JOSSERAND, L. sericifera (LocQ.) 

LocQ. et L. Kuhneriana sp. nov. ................................. 293 

Aage Lund: Sporobolomyces and other yeasts on grains of 

barley .................................................... . ............. 297 

Anders Munk: On Metasphaeria coccodes (KARST.) SACC. and 

other fungi probably related to Massarina SACC. (Massarinaeeae 

n. fam.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303 

D. Miiller: Lycogala flavo-fuscum in Denmark .. ................... 309 

F. H. MiHIer: Two little-known Danish mushrooms: Boletus 

fragrans VITT. and Boletus edulis var. citrinus PELTEREAU 312 

J. A. Nannfeldt: Polyporus hispidus (BULL.) FR. funnen på 

bland. (Summary: Polyporus hispidus (BULL.) FR. found 

on the Island of bland (Sweden». .. ...... ................... 317 

T. Nathorst-Windahl: Zur Verbreitung der Agaricales in den 

Waldern des siidwestlichen Schwedens ........................ 319 

Paul Neergaard: Yellow phyllode spot and drop of Acacia 

armata R. BR. (Septoria acaciae sp. nov.) .................. 325 

T. Vincents Nissen: Soil actinomycetes antagonistic to Polyporus 

annosus FR. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 332 

Wolmar Nyberg: Boletinus asiaticus SING. i Finland. (Summary: 

Boletinus asiaticus SING. found in Finland) .................. 340 

P. Norup Pedersen: Infectioll of barley by loose smut Ustilago 

nu da (JENS.) ROSTR. ....................... ......................... 341 

C. Riber Rasmussen: Yield experiments with mushrooms cultivated 

on synthetic compost or horse and pig manure 

supplemented with various substances .... .... ... ............. 349 

Erik Rennerfelt: The natural resistance to decay of certain 

conifers .. '" .................. " . .. .... ... ..... .... ... .. . ... .. . ... ... . 361 

Håkon Robak: Some fungi occurring on died-back tops and 

branches of Picea abies and Abies spp. in Western Norway 

..................................................................... 366 

Rolf Santesson: Capillipes cavorum g. nov., sp. nov., a new terricolous 

inoperculate discomycete from Swedish Lappland 390 

Audrey M. Shepherd: A short survey of Danish nematophagous 

fungi .................................................................. 396 

Jens Stordal: Distribution of Tricholoma gambosum (FR.) GILL. 

and Boletus granulatus L. ex FR. in Norway .. ... .......... 409 

Frede Terkelsen: Lactarius groenlandicus sp. nov. .................. 417 

Hervid Vallin: Tricholoma verrucipes FR. eller Clitocybe verrucipes 

(FR.) MAIRE funnen i Halsingborgstrakten, Sydsverige. 

(Summary : Tricholoma verrucipes FR. is found 

in Scania, Sweden) ........ ........................ ... ... ...... .... 420 

S. Woldmar: Några vastsvenska svampfynd. (Summary: Notes 

on Macromycetes from Western Sweden) ................ ..... 425 

REDAKTION: 

N. F. BUCHWALD F. H. MØLLER 

* 

Udgivet af Foreningen til Svampekundskabens Fremme 

Rolighedsvej 23, København V. 

PRIS: 50 KR. 

Trykningen afsluttet Maj 1956.

A New Standard Iconography of Agarics. 

JAKOB E. LANGE, the famous Danish mycologist, during his life 

time made a series of excellent water-colour figures with short de 

scriptions of about 1200 species of Agarics. The original paintings 

were purchased by the Botanical Museum of the Copenhagen Uni 

versity. 

A joint grant of 70.000 Danish Kroner from the Carlsberg Founda 

tion and the Rask-Ørsted Foundation made it possibIe to publish 

these during 1935-1940 and to offer the work at a very low price 

considering its first rate quality and the high cost of colour printing. 

The entire work consists of five volumes in folio (about 550 pages) 

and 200 plates in chromo-lithography, the process involving up to 

ten printings. Altogether about 1200 species are figured. The text 

includes a complete set of keys and a description of all the species 

including spores, basidia, cystidia etc. 

This iconography is of great importance to mycologists all over the 

world, many species of agarics being cosmopolitan. 

The price has been fixed at 700 Danish Kroner for the complete 

work. Single volumes are not sold. 

We shall be glad to send a specimen plate and a page of the text 

in order that an opinion may be formed of the high standard of 

the work. 

PIe ase apply to: 

Flora Agaricina Danica, 

The Society for the Advancement of Mycology, 

The Royal Veterinary and Agricultural College, 

23, Rolighedsvej, Copenhagen V 

Denmark. 

Hen. - BOgfrykkergaarden. KJlbenhavn

BIND V

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