Melampsoridium on Alnus in Europe.

M. alni conspecific with M. betulinum

By F. ROLL-HANSEN and HELGA ROLL-HANSEN

Abstract

Melampsoridium betulinum from Betula verrueosa was found able in infeetion experiments to infect
Alnus barbata, A. eordata, A. glutinosa, A. ineana, A. orientalis, A. rhombifola, and A. viridis.
Melampsoridium etilleetions on Alnus spp. in Norway, Finland, Seotl.md, and Ireland were identified
with M. betulinum. Alnus species seem to be poorer hosts for M. betulinum than the Betula species.
M. alni is eonspeeifie with M. betulinum.

Introduction
A Melampsoridium sp. was found on leaves of plants of Alnus glutinosa (L.) Gaertn. in a
nursery at the Agricultural University of Norway at As on 2 October 1972 by the student
Peder GJERDRLFM. The plants were growing in a row beside a row of plants of Betula
verrueosa Ehrh. heavily infected by Melampsoridium betulinum Kleb. Uredinia and telia
were found on both hosts, but most abundantly on the birches. The alders might have been
infected from the birches.
Previous authors have based discrimination between Melampsoridium species on host
species, on length of ostiolar cells m the uredinial peridium, and on size and echinulation of
the urediniospores. In the present paper discussions on the morphology are based on the
same characters.
Literature data on Melampsoridium on Alnus, report on infection experiments, and
report on examination of lierbarium specimens are given.

Earlier descriptions and records

According to the literature, two Melampsoridium species are described from Alnus {M. alni
and M. hiratsukanum) and one frotn Betula (M. betulinum).

Melampsoridium alni Diet.

THUMEN (1878) described Melampsora alni on Alnus viridis (Chaix) DC. of the subgenus
Alnaster (Spach) Endl., from the Sayan mountains in Siberia; only uredinia were described;
the urediniospores were found to be cyltndrtcal-clavate, cylindrical, or oblong, subverrucu-
lous or nearly smooth, 40 (im long. TRANZSCHEL (1895) described both uredinia and telia
also on A. viridis from the Sayan mountains; for the urediniospores he gave the measures
28-42X 10-15 nm. DIETEL (1900), referring to THUMEN, transferred the fungus to the genus
Melampsoridium mentioning that it had been found on A. viridis from Ural to Japan.
In other papers, from 1936 and later, concerning M. alni in Asia similar measures for the
urediniospores were given: 27-47x9-18 |xm (HIRATSUKA 1936, 1958), 39.6x11.2 \im

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78 /•'. Roll-Hansen and Helga Roll-Hansen

(TRANZSCHEL 1939), 3 2 . 4 - 4 6 . 8 X 8 . 8 - 1 5 . 2 |im (KUPREVICZ and TRANZSCHEL 1957), 2 5 - ^ 5 X

8.7-17.5 \im (AzBUKtNA 1974). All the named authors stated that the urediniospores are
smooth at the upper end. Maximum length given of the ostiolar cells was 32 [im. The hosts
were restricted to the subgenus Alnaster, inter alia A. firma, A. rnaximowtczii, anci A.
pendula.
GAUMANN (1959) also gave similar measures for the urediniospores ( 2 7 ^ 7 x 8 - 1 8 \im),
mentioned the upper smooth end, and gave 32 [xm as maximum length of the ostiolar cells.
He indicated that A/, ^/wj reported on A. glutinosa and/I, ineana front Great Britain might be
M. betulinum.
Melampsoridium hiratsukanum S. Ito ex Hiratsuka
As described by ITO (HIRATSUKA 1927) this species can be distinguished from M. alniby the
long ostiolar cells (32.4-55.8 |im) of the uredinium peridium and by the relatively sinall
urediniospores (21.6-34.2x10.4-15.2 |xm) witli a uniformly echinulatc cpisporium. M.
hiratsukanum is found on A. hirsuta (HIRATSUKA 1927) and on ol\\er Alnus species within
the subgenus Gymnothyrsus (HIRATSUKA 1932, f936, 1958; TRANZSCI-IEL 1939; KUPKEVICZ
and TRANZSCHEL 1957; SATO 1966; AZBUKINA 1974). Infection experiments have given
positive results on species within the subgenus Gymnothyrsus, but negative within Alnaster
(HIRATSUKA 1932; SATO 1966).

Melampsoridium betulinum (Fr.) Kleb.

Maximum length of the ostiolar cells was by KLEBAHN (1899) stated to be 35 [tin; the same
maximum length has been given later by Kut'REVicz and TRANZSCHEL (1957), GAUMANN
(1959), and AzBtJKiNA (1974). Measures of the urediniospores by the same authors and by
HIRATSUKA (1936, 1958), GJAERUM (1974), ZH.LER (1974), and MA.IEWSKI (1977) gave a
pt)oled total variation 20-40x8-17 [xm. The urediniospores have been stated to be smooth at
the upper end.

The European collections of Melampsoridium on Alnus spp.

Melampsoridium species on Alnus hosts have been reported from Finland (^|6RSTAD and
NANNEELDT 1958; RAUHALA 1959; GJAHRUM 1974), Ireland ( O ' G O N N O R 1936), Italy
(CiPERRi and CAMERA 1962), and Scotland (FARQUHARSON 19ff; GROVE 192f; WILSON
1924, 1934; HIRATSUKA 1927; WILSON and HENDERSON 1966; HENDERSON and BKNNEL
1979).
The reports frtini Finland, Ireland, and Italy were given without description of the
lungus, which was identified with M. alni, probably because of the host. But the I'lnnish
collection had originally been identified with M. betulinum by V. KtJ.|Ai.A.
In the Scottish reports, short descriptions and discussions were often given. The descrip-
tions were not uniform. The clifference from M. betulinum was in some papers stated to be
very small. But the urediniospores were sometimes reported to be smaller than those of M.
betulinum. Reports were given where tlie spores were stated to be echinulatc over the whole
surface, but aeeording to one of the papers this character did not appear to be constant. The
collections were in most cases identified with either M. alni, mainly because of the Alnus
hosts, orwith M. hiratsukanum because of the hosts within the subgenus Gymnothyrsus and
because of the urediniospores described as relatively small and echinulate over the wholc
surface. Only FAKQUMARSON (1911) and F1I-;NDI:RS(IN and BI'NNEL (1979) identified the
Scottish Melampsoridium on Alnus species with M. betulinum.
 Melampsoridium on Alnus in Europe 79

Material and methods

Infection rnaterial originating from the collection of Melampsoridium on A. glutinosa,

Norway, As, 2 October 1972 (the Alnus strain) and of M. betulinum on B. verrueosa,
Norway, As, 13 October 1972 (the Betula strain) were kept in separate greenhouses.
The uredinia infection material was kept vigorous by transferring urediniospores to pot
plants of Alnus und Betula species in the greenhouses where new leaves were fotnied all
winter.
Basidiospores of the two strains were obtained from the same two colleetions from leaves
with teliospores kept outdoors in separate places in wire boxes on the ground.
Aeciospores were obtained from Larix needles infected by the above-mentioned basidio-
spores. The Larix pot plants were grown in the greenhouses.
Application of urediniospores on Alnus or Betula leaves was carried out during the period
14 October to 18 June. The spores were applied with a hair brush to leaves pteviously
sprayed with tap water. Subsequently the plants were kept in moist ehambers for a eouple of
days. Urediniopustules could sometimes be observed 7 days after inoculation and ripe
urediniospores after 11 days, but in most cases more time was required.
During February-March teliospores germinated on the leaves after about 3 ^ days at 15 °C
in petri dishes lined with wet filter paper. The leaves with the getmin.iting teliospores
were kept on young needles of Larix plants in a nearly 100 % humidity chamber for about 4
days. In positive cases pyenia eould be observed on the Larix needles 11 days or more, and
aeeia 17 days or more after inoculation.
For infection experiments with aeciospores Larix needles with a few aecia were shaken in
5 ml of water and the spore suspension sprayed on both sides of the Alnus or Betula leaves. I n
positive cases ripe urediniospores eould be observed after 17 days.
Except for application of spores, eontrol plants were treated in the same way as the
inoculated plants. The control plants were never infected.
The moisture and the temperature in the greenhouses varied and so did the state of the
tested plants. Obviously the eonditions for infection were not always optimal. Therefore
only the positive results may be fully relied on.
Herbarium specimens were examined from field collections as well as from the infection
experiments. They are hsted in Table 2. Infection by both the Alnus sttain and the Betula
strain on each host species would have been ideal for comparison of the two strains, but we
aimed mainly at infecting Alnus species by the Betula strain and Betula species by the Alnus
strain.
Diameter of uredinia, maximum length of ostiolar cells, length and width of uredinio-
spores were measured; the urediniospore spines were included in the measures given. The
echinulation of the urediniospores was studied. Other objects of interest were the uredinium
peridium and, when present, telia and teliospores. Necrosis around the uredinia was noted
when present.
f'or the scanning electron micro.scopy (SEM), urediniospotes from dry herbarium
specimens were directly mounted and sputter-coated with gold/palladium. The micrographs
were taken with a Jeol-1SM-35C scanning electron microscope at an accelerating voltage of
8or lOkV.

Results
Infection experiments
Infection experiments were earried out during the winter season 1972-1973.
The tree species used in tlie infection experiments are listed in Table 1. For each tree
species are given the number of successfully inoculated plants, the number of plants used for
80 F. Roll-Hansen and Helga Roll-Hansen

Table I
Melampsoridium. Number of .successfully inoculated plants. Total number of inoculated plants in
brackets. Number of control plants of each tree species

Inoculum
Uredospores or ^cnninatinj^ Aeciospores from C'^ontrol
Inoculated tree species leleutospores from plants
A. j^lutinosa, Betula sp.. Betula sp., /^.anx sp., Larix .sp.,
Alnus strain AhiHi strain Bclula strain Alnus strain Bclula strain

Alnaster:
Alnus firma S.etZ. 0(4) 4
A. maximowiezii C.^\\. 0(4) 4
A. pendula'Matsum. 0(1) 0
A.vmdis{Ch^iK)VC. 3(10) 2(8) 0(2) 11
Gymnothyrsus:
Alnus harbata C. A. 'Mey. 4(4) 4
/I. cort/a(a (Loisl.) Desf. 4(4) 4
A. glutinosa {L.)G3crm. 3(3) 8(8) 4(8) 12
A. birsuta (Spaeh) R u p r . 0 (4) 4
/l.mcan^(I..)Moc-nch 9(11) 10(12) 20
A. orientalis Dene. 1(2) 2
/I. r^om^j/b/M Nutt. 2(4) 4
Betula oeeidentalis Hook. <> (<i) 4
B. pubeseens Ehrh. or
B.wrr«cos<j E h r h . 3(4) l.S(16) 22(22) 2(2) (,(11) 26
Larix X eurolepis Henry or
L.r«ss(Cd (Endl.) Sahine ex Tratitv. 1(2) 4 (.S) 2

the separate kinds of inoculation, and the nutnber of eontrol plants. Positive results were
recorded if spores were formed in uredinia, spermogonia, or aeeia.
f n the subgenus y4/ni«5£er positive results were obtained in A. viridis when inoeulated with
urediniospores of thej4/««s strain as well as of the Betula strain. A. firma, A. maximowiezii,
and A. pendula gave negative results (Table 1).
In the subgenus Gymnothyrsus, A. hirsuta was the only species tested that always gave
negative result. A. eordata, A. glutinosa, A. ineana, A. orientalis, and A. rhombifolia all gave
positive results in at least half of the plants inoculated with urediniospores of the Betula
strain, fn addition, positive results were obtainecf when A. glutinosa was inoculated with
urediniospores o{ the. Alnus sirain and when A. glutinosa or A. ineana were inoeulated with
aeciospores of the Betula strain (Table 1).
B. pubeseens and B. verrueosa were easily infected by both strains. B. occidentalis was
tested only with the Betula strain to which it was very susceptible (Table 1).
Infection of Larix needles by basidiospores from germinating teliospores was obtained
with both strains. TwopIantsofL.r«s«Cit were inoeulated with the/l/w«i strain; one got ripe
aeeia, the other had rust hyphae in the needles, but no pyenia or aecia were observed, and the
plant was registered as negative (Table 1). Five Larix plants were inoculated with the Betula
strain; one L. Xeurolepis and two L. russiea got ripe aecia, one L. russica got only spermogonia,
and one L. russiea got no infection (Table I).
It is of special interest that in the infection experiments in the greenhouses, the uredinia
formed on the Alnus species were generally fewer and smaller than those formed on the
Betula speeies. On Alnus species there were often formed dark neerotic tissue around the
uredinia or necrotic spots without uredinia. Thus in the greenhouses the Alnus leaves might
get a characteristic, dark-spotted appearance not seen on the Betula plants (Figs. 1,2, and 3).
In the field collections, the necrotie spots were not distinct or not observed.
Figs. I'-3. Melampsoridium betulinum from the infection experinients in greenhouses, fig. /. Necrotic
spots on leaves of Alnus glutinosa. Fig. 2. Uredinia and necrotic spots on leaf of A. ineana x l . 7 .
Fig. 3. Uredinia on lc-.if of Betula pubescens X2.7
82 F. Roll-Hansen and Helga Roll-Hansen

We found that the susceptibility of B. verrueosa to M. betulinum varies in forests as well

as in nurseries. Immune or nearly immune individuals have been observed both in infection
experiments and in the field.

Comparison of the Norwegian Alnus strain with the Norwegian Betula strain
The colleetions examined were the original Norwegian field collection on A. glutinosa, the
field collection on the neighbouring 5. verrueosa, and samples from the greenhouse infection
experiments (Table 2).
The diameter of the uredinia seemed to be smaller on the Alnus speeies than on the Betula
species, but no difference between the strains was found within eaeh host genus.
The maximum length found of ostiolar cells varied from collection to collection,
probably dependent on the general development of the uredinia and on the relatively small
number of ostiolar cells measured; there might seem to be some difference between the two
strains, but the differenee is not significant. No other difference was found in the peridium.
The echinulation of the urediniospores was the same in both strains, fairly even on the
whole surface, exeept for a bald area at the top of the spores. The size of the bald area varied;
on some spores it was not seen at all, possibly beeause of the orientation of the spores in the
slides.
The size of the uridiniospores did not vary mueh from eollection to eollection; no
significant difference was found between urediniospores on the two host genera or between
Table 2
Diameter of uredinia, maximum length observed of ostiolar celKs, and length and width of
urediniospores in the examined collections of Melampsoridium

Uredinia O.stiolar Uredini ospores

cells maxi-
Host pl.int Orif^in diameter num lengdi length width
observed
mm nm |lni Uni

Norwegian specimens: Herb. NFRI

Alnus glutinosa As, 2. 10. 1972 0.1-0.2 38 26^0 10-17
A. glutinosa Inoc. from Alnus 0.1 40 26^2 13-17
A. viridis Inoc. frtini Alnus 0.1-0.3 28 22-34 13-16
Betula verrueosa As, 13. 10. 1972 0.2-0.5 49 30-42 12-15
A. barbata Intic. from Betula 0.1-0.2 38 25-36 10-15
A. eordata Inoc. from Betula 0.1-0.2 46 24-38 12-15
A. glutinosa Inoc. from Betula 0.1-0.2 46 30-40 13-19
A. ineana Inoe. from Betula 0.1 30-38 13-17
A. orientalis Inoc. from Betula 0.1-0.2 47 26-37 12-15
A. rhombifolia Inoc. from Betula O.I 22-35 11-16
A. viridis Inoc. from Betula 0.1-0.2 32 24-38 12-15
B. verrueosa Inoc. from Betula 0.2-0.4 43 26^0 11-16
B. oeeidentalis Inoc. from Betula 0.1-0.4 41 24-38 11-14
Finnish specimen: Herb. V. Kujala
A. glutinosa Helsinki,Mustavuori, 18.10.1953 0.2-0.3 33 26-38 13-16
Scottish specimens: Herb., Royal Bot. Garden, Edinburgh
A. eordata Oet. 1931 M.WiLson 0.1-0.3 30^3 13-15
A. glutinosa 10 collections 0.1-0.4 32-45 25^6 12-16
A. ineana 3 collections 0.1-0.4 35^3 28-40 12-16
Irish specimens: Herb., Nat. Bot. Garden, Dublin
A. glutinosa 4 collections 0.1-0.25 33-40 24^4 11.5-16.5
Siberian specimens: Herb., Bot. Inst. Acad. Sci. USSR, Leningrad
A.frutieosa {A. viridis) Collection I 0.1-0.25 40 35-47 12.5-16.5
Collection 11 0.1-0.25 35.5 28-t3 12-16.5
 Melumpsoridiurn on Alnus in Europe 83

urediniospores of the two strains. Neither was there any difference between telia and
teliospores in the field collections of the two hosts. All the examined characters of the
Norwegian herbarium material thus indicate that the two strains are identical. As also the
infection experiment gave no basis for separation, the two strains will be pooled in our
further discussion and named Melampsoridium betulinum.

Comparison of the Finnish, Scottish, and Irish collections on Alnus species with
Norwegian M. betulinum
No significant difference was found between the Finnish, Seottish, Irish, and Norwegian
material as to diameter of uredinia, peridium with ostiolar eells, echinulation and size of
the urediniospores (Table 2, Figs. 4, 5, and 6), neither as to telia and teliospores.
It must be eoneluded that there is no reason not to identify the examined Finnish,
Scottish, and frish material on Alnus species with M. betulinum.
We were not able to examine any Italian speeimens.

The conspccifity of
M. alni with M. betulinum
The M. alni descriptions given
of ostiolar cells and size and
echinulation of the uredinio-
spores by THUMEN (1878),
TRANZSCHEL (1895), DIETI-L
(f900), HIRATSUKA (1936,
1958), KupREVicz and TRANZ-
SCHEL (1957), and AZDUKINA
(1974) fit M. betulinum. The
hosts were said to be speeies
within the subgenus Alnaster.
Maximum length given for the
ostiolar cells was 32 [xm, the
urediniospores were described
as echinulate except for a bald
spot at the top of the spores,
and measures of the uredinio-
spores were given within the
boundaries 25^7 X 8.8-18 [im.
Department of Ciyptoga-
mic Plants, Botanical Institute
of Aeademy of .Sciences of
USSIl in Leningrad kinifly lent
us two eolleetions of "Melamp-
sora Alni Thum. n. sp.", both
from A. fruticosa Rupr. {A. vi-
nt^w [Chaix] DC.) in the Sayan
Mountains in Siberia. Collec-
tion I was labelled: "f'ungi Mi-
nusiensis exsiecati. Melamp-
sora Alni Thum. n. s. in syl-
vinis subalpinis prope monteni l-igs. 4-6. Melampsoridium betulinum, urediniospores. SEM
X1320. Fig. 4. From Betula verrueosa, Norway. Fig. 5. From
Borus in alpibus sajanensibus Alnus glutinosa, Finland. Fig. 6. From A. glutinosa, Scotland
84 F. Roll-Hansen and Helga Roll-Hansen

ad Alni viridis D. C. folia viva, ex

herb. N. Martianoff". Collection II
was labelled (translated from Rus-
sian): "Fungi, Minusinsk flora.
Melampsora Alni Thum n sp, on
leaves of Alnus fruticosa {viridis),
taiga at the foot of Borus mountain.
N. Martianoff". The two eolleetions
did not differ from M. betulinum
as to size of uredinia, maximum
length of ostiolar cells, or size of
urediniospores (Table 2). The echi-
nulation and the bald spot of the
urediniospores were also the same
(Figs. 7, 8).
M. betulinum {M. alni) is clearly
different from M. hiratsukanum,
which has urediniospores echinulate
over the whole surface (Fig. 9).
We conclude that M. alni is eon-
specific with M. betulinum from:
1. earlier deseriptions of M. alni, 2.
our investigations of the Siberian
material on A. fruticosa {A. viridis),
and 3. the faet that M. betulinum
infects Alnus speeies also within the
subgenus Alnaster.

Figs. 7-8. "Melampsora Alni Thum.",

urediniospores. SEM. Kg. 7. Collection I
X 1320. Fig. 8. Collection II X600. Fig. 9.
Melampsoridium hiratsukanum from Al-
nus ineana (Danish provenance), Japan,
urediniospores. SEM X600

Acknowledgements

Dr. D. M. HENDERSON, The Royal Botanic Garden, Edinhurgh, tent us the Scottish herbarium material
of Melampsoridium on Alnus spp. Professor Dr. Vnjo KUJALA gave us part of his Finnish collection.
Miss MAURA SCANNELL lent us four Irish specimens from the Herbarium at the National Bot.mic
Gardens, Dublin. Department of Cryptogainic Plants, Botanical Institute of Academy of Sciences of
USSR, Leningrad, lent us the two collections of "Melampsora Alni Thum. n. sp.". Dr. HAKUYOSHI
SAHO sent us specimens of Melampsoridium hiratsukanum from Japan. The scanning electron micro-
graphy has been carried out by the Instrument Service of the Norwegian Agricultural Research
Council. Mr. HAt^voR B. GJAERUM has carefully read the manuscript and made valuable suggestions.

Summary
A Norwegian isolate of Melampsoridium from Alnus glutinosa was eompared in infeetion experiments
with an isolate of M. betulinum from Betula verrueosa. The isolate from Betula was able to infeet
A. barbata, A. eordata, A. glutinosa, A. ineana, A. orientalis, A. rhombifolia, and A. viridis, \. e. spec
within the subgenus Gymnotbyrsus as well as Alnaster. Studies on the morphology of the fungi in tbe
original collections from Alnus and Betula and of the isolated strains gave no basis for discrimination
 Melampsoridium on Alnus in Europe 85

between tbe collections. The Norwegian collection on Alnus was therefore identified with M. betu-
linum.
A Finnish collection of Melampsoridium sp. on Alnus glutinosa and Scottish collections of Melam-
psoridium on A. eordata, A. glutinosa, and A. ineana were eompared witb the above-mentioned
Norwegian material. Irish collections on A. glutinosa were .also examined. All eolleetions were
identified with M. betulinum. They cannot be identified with M. hiratsukanum, which, according to
the diagnosis, has somewhat longer ostiolar cells .md urediniospores without a bald area.
In the infection experiments Betula speeies generally seemed to be better hosts for M. betulinum than
were the Alnus speeies. But resistant Betula individuals have been found both in infeetion experiments
and in tbe field. On the Alnus speeies the uredinia were relatively small; in the greenhouse experiments
the uredinia were often surrounded by a neerotie zone and sterile neerotie spots were common.
There may be the different mces within the .species M. betulinum, some of which may not be able
to infeet Gymnotbyrsus or Alnaster, or perhaps not any Alnus speeies at .ill; but we think it is right
to name all of them M. betulinum as long as clear morphological differences are not found.
The diagnosis and other descriptions given of M. alni Diet, .agree well with our descriptions of
characters of M. betulinum. Study of "Melampsora Alni Thum. n. sp." on A. frutieosa {A. viridis) from
tbe Sayan Mountains settled the question of conspeeificity.

Resume
Melampsoridium sur Alnus en Europe. M. alni identique a M. betulinum

A. orientalis, A. rbombifolia clA. viridis, e'est-a-dire des espeees appartenant au .sous-genre Gymno-
thyrsus aussi bien qu'au sous-genre / l / f
d port.int sur lla morphoiogie du champignon dans les colleetions originales provenant
Des etudes
d'Alnus et Betula et a partir de diverses souehes isolees n'ont t'ourni aueune base de differeneiation entre
les collections. La rc-colte norvegienne sur Alnus a c'tc dc's lors identifiee comme appartenant .a l'espeee
M. hetulinum.
Une reeolte finlandaise de Melampsorodium sp. sur Alnus glutinosa et des rc-eoltes effeetuees en
Ecosse de Melampsoridium sur/1. eordata, A. glutinosa et /I. ineana ont ete companies avec le materiel
norvegien dont il est question plus haut. Des reeoltes irlandaises sur A. glutinosa ont c-te egalement
examinees. Toutes les rc-eoltes ont c-tc- identifiees comme c-tant M. betulinum. Elles ne peuvent c-tre
rattachees a M. hiratsukanum, qui, d'apres la diagnose, possede des cellules ostiolaires quelque peu plus
longues et des urc-diospores sans zone dcnudee.
Lors des expc-riences d'infection, les espeees du genre Betula semblent en gc-neral eonstituer des hotes
plus favorables pour M. betulinum que les Alnus. Mais des individus resistants .ippartenant au genre
Betula se sont revt-lcs, tant ati cours des expe-riences d'infection que dans la nature. Sur les Alnus, les
uredies sont relativements petites; dans les experienees en serre les uredies sont frequemment entourees
d'une zone necrosee et des t.iclies nc-erosees steriles sont conimunement notees.
Il peut y avoir des raees differentes a l'interieur de l'espeee M. betulinum, dont les unes ne peuvent
avoir d'aptitude .i infecler le sous-genre Gymnothyrsus ou Alnaster, ou peut-c-tre aucune espece d'Alnus;
inais nous pensons qu'il est legitime de toutes les nommer M. betulinum, tant qu'on aura pas deeouvert
de differences morphologiques notables entre elles.
La di.ignose et d'autres deseriptions fournies sur M. alni Diet, sont en bon accord avec nos des-
criptions des earaetc-res de M. betulinum. L'evude de "Melampsora alni Thum. n. sp." sur A.frutieosa
{A. viridis) provenant des montagnes de Sayan elarifie le problbme de la eo-spe-cificitt- (identite).

Zusammenfas.sung
Melampsoridium an Alnus in Europa. M. alni identiseb mit M. betulinum
Ein norwegisches Melampsoridium-hoht aus Alnus glutinosa wurde in Infektionsversuehen mit einem
M. betulinum-lsohi aus Betula verrueosa vergliehen. Das BetuLi-lsohi infizierte die Arten Alnus
barbata, A. eordata, A. glutinosa, A. ineana, A. orientalis, A. rhombifolia und A. viridis, d. h. Arten
irinerhalh der Subgenera Gymnothyrsus wie Alnaster. Die Untersuchungen iiber die Morphologie des
Pilzes anhand von Originalkottektionen von Alnus und Betula sowie der isotierten Stamme fiihrten zu
keiner Untenscheidung zwischen den Kollektionen. Die norwegische KoUektion an Alnus wurde daher
als M. betulinum ideiuifiziert.
Eine finnisehe Koltektion von Melampsoridium sp. zn Alnus glutinosa und sehottisehe Kollektionen
von Melampsoridium an A. eordata, A. glutinosa und A. ineana wurden mit dem oben erwalinten
86 F. Roll-Hansen and Helga Roll-Hansen

norwegisehen Material vergliehen. Auch irisehe Kollektionen an A. glutinosa wurden iiberpriift. Alle
Kollektionen wurden als M. betulinum identifiziert. Sie konnten nicht als M. hiratsukanum bestimmt
werden, weil diese Art - den Diagnosen zufolge - etwas langere ostioiare Zellen und Uredosporen ohne
eine kahle Flaebe auf weist.
Den Infektionsversuehen zufolge seheinenBc/«/d-Arteni. a. hesser geeignete Wirte fiir/W.Z'ete/mMm
darzustellen als y4/n«s-Arten. Resistente Individuen wurden aher bei Betula sowohl in den Infektions-
versuehen als im Freiland gefunden. An Alnus-Arten waren die Uredosporen relativ klein; in Versuchen
unter Glas waren die Uredosporen oft von einer nekrotisehen Zone umgeben, haufig traten sterile
nekrotisehe Stellen auf.
Moglieherweise existieren innerhalb der Art M. betulinum verschiedene R.issen, von denen einige
nicbt imstande sind, Gymnothyrsus oder Alnaster, vielleieht iiberbaupt jedwede Alnus-An zu iiifi-
zieren. Wir halten es aber fiir korrekt, alle M. betulinum zu nennen, solange keine klaren morpholo-
gisehen Differenzen gefunden werden.
Die Diagnose und andere Besebreibungen von M. alni Diet, stimmen gut mit unseren Merkmals-
besehreibungen fiir M. betulinum uberein. Untersuebungen an Melampsora Alni Thiiiii. n. sp. an
A.frutieosa aus den Sayan-Bergen festigten die Frage der Identitat.

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Authors'address: Professor V. ROLL-HANSEN and Forest Research Officer HELGA ROLL-HANSEN,

Norwegian Forest Research Institute, P.O.Box 62, 1432 As-NLH, Norway
Receipt of Ms. 19.yi9S0

Lehrstuhl fiir Forstbotanik der Universitat Miinehen

Rhizographische Untersuchungen an Feinwurzelsystemen

aus verschiedenen Abies alba-Bestanden Bayerns
Von H . BLASCHKE

Abstract
Rfiizographie investigation of fine root systems of silver fir stands in Bavaria. Fine root samples
were taken in topsoil and subsoil of silver fir stands affected by diebaek. The results of the rhizographie
investigation show th.it the myeorrhizal rootlet formation and the fine root distribution was markedly
dependent on soil structural characteristics. The features of myeorrhizal rootlet development stages
and decomposing stages and fine-root regeneration in affeeted trees were different from those in tieatthy
trees, wtiieli showed an abundanee ot living fine-root mmifications.

1 Einleitung
Als Ursache fur einen Riickgang der Vitalitat von Waldbaumen werden vielfach auch Seha-
den im Wurzelbereich als Folge ungiinstiger Umwelteinfliisse vermutet (ULRICH et al.
1979). Die Kenntnisse iiber Wurzelsysteme von Waldbaumen, insbesondere (iber Beziehun-
gen zwischen den Leistungen des Wurzelsystems und den Bodenfaktoren, sind jedoch
immer noch recht unvollstandig (KREUTZER 1961; REYNOLDS 1974).
Die genauc Lokalisierung von Wurzelschaden 1st nur moglich, wenn detailliette Wurzel-
untersuchungen zur Analyse gehoren. Dazu konnen verschiedene rhizographische Me-
thoden herangezogen werden, die, ergiinzt durch wurzelphysiologische Untersuchungen, zu
der gcwiinschten Aussage iiber Wurzelfunktionen und die im Wurzelbereicb auftretenden
Scbadigungen fiihren (BLASCHKE f980b). Im Rahmen von Untersuchungen iiber das Tan-
nensterben war es das Ziel, anhand der Bewurzelungsverhaltnisse mogliche Zusammen-
hange zwischen dem Riickgang der Vitalitat von Tannen und deten Wurzelentwicklung
aufzuzeigen.

2 Material und Methoden

Fur wurzelmorphologische Untersuchungen stand Material aus mehreren bayerischen

Forstamtern zur Verfijgung (Tab. 1). Neben erkrankten wurden auch gesunde Alttannen
sowie Tannen aus Naturverji.ingungen in die Untersuchung einbezogen.

U.S. Copyright Cleamnce Center Code Statement: 0300-1237/81 / 1101 /0087 $ 02.50/0
Eur.J. For. Path. 11 (1981) 87-97
© 1981 Verlag Paul Parey, Hamburg und Berlin
ISSN 0300-1237/ InterCode: EJF'PA9