Peronospora On Compositae

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Peronospora complex on Compositae

O. CONSTANTINESCU
The Herbarium, Uppsala University, Box 541, S-751 21 Uppsala, Sweden
CONSTANTINESCU, O. (1989). Peronospora complex on Compositae. - SYDOWIA

41: 79-107.
Peronospora leptosperma and similar fungi parasitic almost exclusively on
plants belonging to the tribe Anthemideae are revised. A new genus, Para-
peronospora, is introduced to accommodate these fungi. It is mainly characterized by
conidiophores having branches broadening upward and by both dicho- and tri-
chotomic branching. Eight new combinations are proposed and lectotypes for 10 taxa
are selected. Detailed descriptions and illustrations of seven species of Para-
peronospora and of Peronospora radii are provided, and a key for their identification
is added. Five doubtful or excluded taxa are also treated.
Two species of Peronospora parasitic on different organs of var-

ious Compositae (Asteraceae) were described and illustrated by DE
BARY (1863). They were placed in the Section Pleuroblastae charac-
terized mainly by the dissemination organs, called conidia, having
continuous wall and germination by tube. The first species, Pe-
ronospora radii, was distinguished by its violaceous colour, and
sporulation in the inflorescence of its host. The second, Peronospora
leptosperma, was characterized by colourless organs and both
dichotomous and trichotomous type of conidiophore branching. In
addition, DE BARY mentioned the upward broadening of branches in
P. leptosperma.
Two taxa similar to P. radii, but parasitic on leaves, were later
described under P. radii var. epiphylla POIHAULT (1915) and P. danica
GÄUMANN (1923). Except for the question if the same fungus may
attack both flowers and leaves, Peronospora radii generated no
taxonomical problems for the subsequent authors.
On the contrary, because of its unusual morphology, P. lep-
tosperma was a source of controversial taxonomic decisions.
GÄUMANN (1923) restricted P. leptosperma to the fungus parasitic
on Chamomilla recutita, C. suaveolens and Matricaria perforata. He
introduced P anthemidis for the form parasitic on Anthemis spp., P.
tanaceti for the one on Tanacetum vulgaris, and added two more
species, P. sulphurea and P. artemisiae-biennis. Six more or less simi-
lar taxa were later described by SAVULESCU & RAYSS (1930), LING & TAI
(1946) and SAVULESCU & VANKY (1956).
79
Two P. leptosperma-like fungi were described in the genus Plas-

mopara by SAWADA (1931) and DUDKA & BURDJUKOVA (1977), and ten
such fungi were transferred to Plasmopara by SKALICKY (1966) and
TAO& QIN(1987).
Rather recently, three P. leptosperma-\ike fungi were placed in
the genus Bremiella, as one new species and two new combinations
(TAO & QIN, 1982).
The difficulty in finding the appropriate generic position of
these fungi is illustrated by the fact that each of the epithets arte-
misiae-annuae and chrysanthemi-coronarii have been combined
into Peronospora, Plasmopara and Bremiella, during the last
decades. Significantly enough, morphologically indistinguishable
collections of one and the same fungus were described by SAVULESCU
& VANKY (1956) under Plasmopara halstedii and four new taxa, Pe-
ronospora achilleae, P. dimorphothecae, P. ursiniae and P. buhrii.
During the preparation of a check list of Romanian Per-
onosporales (CONSTANTINESCU & NEGREAN, 1983), as well as of a mono-
graph of the genus Plasmopara, a number of these fungi parasitic on
Compositae were examined. It soon became apparent that, by their
morphological characters, they differ not only from typical Pe-
ronospora, but also from Plasmopara and Bremiella. Consequently,
a revision of all Peronospora-like fungi described on Compositae
was undertaken.
Material and methods

Some 355 herbarium specimens, including the types of 15 out of
the 24 taxa dealt in this paper were examined. Slides were prepared
by wetting the specimen with 95% ethyl alcohol and transferring
fungal material in lactic acid or lactofucsin. Except for the length of
the conidiophores, all other measurements were performed with the
aid of a oil-immersion lens and drawing tube at a 2000x final
enlargement. The nomenclature of the hosts is mainly in accordance
with CKONQUIST (1955), MOORE et al. (1976), SCOGGAN (1979) and SHI-
SHKIN & BOBROV (1961). Herbarium abbreviations are those used by
HOLMGREN et al. (1981). In some cases only selected specimens are
cited, but complete lists are deposited at the UPS herbarium. Except
where abbreviations may cause confusion, P. is used for Peronospora
and Pp. for Paraperonospora.
Results
Morphology. - With the exception of Peronospra radii, all
other fungi studied have a number of characters in common. The
branches, and in many cases even the trunk, are slightly but consis-
tently broadening toward the growing direction (Fig. 1 A). Although
80
the general pattern of the branching is dichotomous, the trichotomy

occurs at various points of branching (Fig. 1 A), and is present in a
number of conidiophores in almost every collection. When bran-
ching is dichotomous, the pair branches have appreciable different
length. Each branch commonly ends in (l-)2-3(-4) ultimate branch-
lets (Fig. 1 C). In some cases one ultimate branchlet is longer and
terminal, appearing as an extension of the branch, and two are
shorter and arise at the base of the terminal one. In other cases three,
more or less equally long ultimate branchlets emerge from a com-
mon, sometimes swollen, base. The tip of the ultimate branchlets
may be blunt, more or less rounded, or swollen in variable degree.
The dissemination organs are of conidium-type, i.e. with continuous
wall, no dehiscence apparatus and germination by tube. The conidia
are rather large, generally over 30 [im long.
In some conidia of P. leptosperma-like fungi a membrane-like
structure is visible below the conidium apex (Figs 1 D, E, 3 D-F, 4 D).
This structure superficially resembles a dehiscence apparatus like
the one present in Plasmopara or Bremiella. However, at a closer
examination, the wall of the conidium appears continuous and no
papilla is present. Moreover, in a few germinating conidia found in
the specimens examined (Fig. 6 C), as well as in one conidium
depicted by TAO & QIN (1987: 71), germination does not occur at, and
through the apex, but at randomly located points of the conidium
wall. Most probably, this membrane-like structure which, signifi-
cantly enough, is not present in all conidia, is only a vestigial dehis-
cence apparatus.
Conidium ontogeny. - The determinate growth of the ulti-
mate branchlets and synchronous production of solitary conidia/
sporangia are characteristic for Peronosporaceae. Peronospora-lep-
tosperma-like fungi fit this pattern. However, in a few herbarium
specimens, an apparently indeterminate growth associated with
basipetal production of conidia was noticed. This case is now under
study.
The r e l a t i o n to o t h e r g e n e r a of P e r o n o s p o r a l e s . -
The unique character of the conidiophores in P. leptosperma-like
fungi clearly differentiate them from Peronospora in which the
branching is consistently dichotomous, the branches have a constant
breadth, and each terminates in two ultimate branchlets with trun-
cate tips. By having conidia they are also distinct from Plasmopara
and Bremiella, genera with poroid dissemination organs, i.e. sporan-
gia. Peronospora leptosperma-like fungi are sufficiently distinct
from other genera of Peronosporales to be placed in a separate
genus. Since the introduction of the genus Peronospora by CORDA
(1837), nine genera of plant parasitic Peronosporales s.l. have been
erected. As in the case of P. leptosperma-like fungi, four of these
6 Svdowia, Vol. 41, 1989 81

Fig. 1: Paraperonospora spp. A-D. Pp. leptosperma. - E . Pp. chrysanthemi-coronarii.

A. conidiophores; arrows show the trifurcate branching. - B. oogonia and oospores. -
C. ultimate branchlets. - D & E. conidia; arrows show the vestigial dehiscence appa-
ratus.
show combined characters of different genera and this is also

reflected by their names: Peronosclerospora, Peronophythophthora,
Pseudoperonospora, Sclerophthora.
The value of the branching pattern of the sporangiophore and
conidiophore in the classification of Peronosporomycetidae was
recently emphasized by DICK et al. (1984). By its branching type and
the presence of conidia, P. leptosperma-like fungi show relationship
to both Plasmopara and Peronospora. The rudimentary, nonfunc-
tional dehiscence apparatus present in some specimens points out to
a possible link with Bremiella and, again, Plasmopara. In the dia-
gram of the classification of the Peronosporomycetidae presented by
DICK et al. (1984), P. leptosperma-like fungi should be placed between
Bremiella and Bremia, on the dichotomous lineage of the Pe-
ronosporaceae.
Taking into account the above mentioned distinctive characters,
a new genus, Paraperonospora, is introduced in this paper to accom-
modate Peronospora leptosperma and similar fungi parasitic on
Compositae.
S p e c i e s c o n c e p t . - The criteria used for delimitation of taxa
in P. leptosperma-like fungi followed the trend initiated in Pe-
ronospora by GÄUMANN (1923). It consists of a supposed affinity of a
certain fungus to a host or a limited number of hosts, and of bio-
metric studies of conidia, mostly based on a single collection. The
limitations of GÄUMANN'S concept have been pointed out, among
others, by GUSTAVSSON (1953, 1959b) and YERKES & SHAW (1959).
The species concept adopted in this paper is basically a mor-
phological one. Those specimens which could be placed reasonably
safely within a given morphological pattern are grouped under the
same species name, irrespective of their host. As a consequence it is
accepted that some species, like Pp. leptosperma, Pp. minor and Pp.
sulphurea have more than one host, whereas in other species, like
Pp. artemisiae-biennis, Pp. chrysanthemi-coronarii and Pp. tanaceti
the morphological features are consistently associated with one par-
ticular host.
Host r a n g e . - The hosts of Paraperonospora, and of Per-
onospora radii, belong almost exclusively to the tribe Anthemideae.
They should be divided into regular or „natural" hosts, i.e. plants
frequently attacked in nature by a particular fungus, and accidental
or occasional hosts, only rarely attacked. The last are either wild or,
more often, cultivated plants, particularly in botanical gardens.
The presence of Pp. leptosperma on cultivated Dimorphotheca,
a member of the tribe Calenduleae, as well as of Pp. sulphurea on
Helichrysum (tribe Inuleae) are in all probability due to accidental
infections. Both tribes, but particularly the Inuleae, are not closely
related to the Anthemideae (BREMER, 1987). As KUKKONEN (1972)
6« 83
pointed out in the case of Anthracoidea (Ustilaginales), „the excep-

tional host seems to be a species growing in the very same spot or in
the vicinity of a heavy infection". An occasional host may be recog-
nized by the fact that the presence of the parasite is discontinuous.
Such a case case is Plasmopara ammi CONSTANTINESCU (1968) parasi-
tic on Ammi majus, a plant originating from Egypt but cultivated in
Romania. Attempts to collect this species again failed, although the
host was cultivated in the same place for many years. I am now
convinced that Plasmopara ammi is only an accidental attack of the
widely distributed Plasmopara umbelliferarum s.l. on a new host.
During this study two specimens of Achillea clusiana, an „occasio-
nal" host for Paraperonospora, were examined. They were collected
at the Rostock Botanical Garden, by the same collector, in 1937 and
1939. The first specimens is parasitized by Pp. leptosperma (BUCM
3155) and the second by Pp. sulphurea (B).
The attack of an unusual host may be due to the presence of a
physiological race which may occur even in Peronosporaceae
attacking non-crop plants (CRUTE, 1981), or to the favourable envi-
ronmental and inoculum conditions (PALTI, 1974).
The only experimental study dealing with the host specializa-
tion of these fungi was made by DZHANUZAKOV (1962). He showed that
the fungus from Matricaria perforata can infect the same plant, as
well as M. suaveolens and Chamomilla recutita, but not Artemisia
vulgaris, Leucanthemum vulgäre and Tanacetum vulgäre. This is
hardly surprising as the last three plants are natural hosts for differ-
ent Paraperonospora species. Because the fungus from Matricaria
suaveolens could infect only this host but not M. perforata and
Chamomilla recutita DZHANUZAKOV concluded that the fungus para-
sitic on M. suaveolens represents a different physiological race.
Taxonomy
Paraperonospora O. CONST., gen. nov.
Etym.: from the Greek para = near/similar to.
Oomycotina. A Peronospora ramis sursum dilatatis et ramificatione dichotoma
vel trichotoma differt.
Specia typica: Paraperonospora leptosperma (DE BARY) O. CONST.
Oomycotina. - Plant parasitic. - C o n i d i o p h o r e s erect, col-
ourless or slightly yellowish, composed of a trunk and a branched
part; branching dichotomous, sometimes trichotomous; all branches
more or less broadening towards the distal part; dichotomous
branches of different length; each branch ends in (l-)2-3(-4) long-
conical ultimate branchlets, having blunt, rounded or inflated tip. -
C o n i d i a large, colourless to yellowish, ellipsoidal to broadly ellip-
soidal, base round, tip round or slightly apiculate; basal pedicel
84
short, slightly eccentric. - Oogonia irregular, thin-walled;

o o s p o r e s aplerotic, globose, with smooth, yellowish to brownish
wall.
Key to the species treated:

1 Branches of uniform width, conidia brownish jR radii
1* Branches broadening upward, conidia colourless, pale yellowish
or greyish-brown 2
2 Tip of ultimate branchlets swollen to 3-4 (ira 3
2* Tip of ultimate branchlets, blunt, rounded or only slightly
enlarged 5
3 Conidia greyish brown Pp. artemisiae-annuae
3* Conidia colourless or yellowish 4
4 Most conidia 35-40 |im long Pp. chrysanthemi-coronarii
4* Most conidia less than 35 (im long Pp. multiformis
5 Broadest part of conidia supramedian Pp. tanaceti
5* Broadest part of conidia median 6
6 Most conidia 35-45 urn long, length/width ratio
ca. 2 Pp. artemisiae-biennis
6* Conidia shorter, length/width ratio <2 7
7 Most conidia 20-26 |im broad, tip often
apiculate Pp. sulphurea
7* Most conidia 17-22 |jm broad, tip rounded 8
8 Conidia mostly 25-30 |j.m long Pp. minor
8* Conidia mostly 30-40 |0.m long Pp. leptosperma
1. Paraperonospora artemisiae-annuae (LEE & M. C. TAI) O. CONST,

comb. nov.
BAS.: Peronospora artemisiae-annuae LING & M. C. TAI - Lloydia 9: 144. 1946
= Plasmopara artemisiae-annuae (LING& M. C. TAI) SKALICKY - Preslia 38:
127. 1966
= Bremiella artemisiae-annuae (LING & M. C. TAI) TAO - Acta mycol. sin. 1:
64. 1982. Type on Artemisia annua L., CHINA: Chengtu, 21 April 1943.
No original specimen was available. According to the descrip-
tion and illustration there is no doubt that this is a Para-
peronospora. The swollen tip of the ultimate branchlets and the size
of conidia (24-46 x 14-24 yon, average 37.03 x 20.19) place this spe-
cies close to Pp. multiformis and Pp. chrysanthemi-coronarii. The
greyish brown colour of the conidia clearly distinguishes it not only
from the last two mentioned species but also from all other species of
Paraperonospora. However, such colour of conidia is quite unusual
in Paraperonospora.
Host. - Artemisia annua L.
Distribution. -CHINA.
85
2. Paraperonospora artemisiae-biennis (GÄUM.) O. CONST, comb. nov.

- Fig. 2.
BAS.: Peronospora artemisiae-biennis GÄUM. - Beitr. KryptogFl. Schweiz 5(4):
131. 1923
= Plasmopara artemisiae-biennis (GÄUM.) SKALICKY - Preslia 38: 127. 1966.
Type on Artemisia biennis WILLD , USA: Dakota, Kulm, 9 July 1912,
BRENCKLE, Fungi dakot. 197 (S - lectotype; S, NY - isolectotypes).
S p o t s epiphyllous, pale yellowish, margin diffuse. - Down
hypophyllous, yellowish, often covering the whole surface. - Con-
i d i o p h o r e s pale yellowish, erect, 320-600 um long, trunk 100-
280 x 7.5-18 urn; upper part dichotomously, rarely trichotomously
branched 3-4(-5) times; u l t i m a t e b r a n c h l e t s 2-4 at the tip of
Fig. 2: Paraperonospora artemisiae-biennis (lectotype); A. conidiophore. - B. ultimate

branchlets. - C. conidia.
86
each branch, long-conical, 8-16 urn long, base 2.5-5 um, gradually
tapering to 1-2 urn below the truncate or rarely almost rounded tip. -
C o n i d i a yellowish, ellipsoidal, (35-)38-45(-48) x (17-)19-21(-
24) um, broadest part median, wall ca 0.5 [im thick; pedicel
1 x 1-1.5 p.m. - S e x u a l o r g a n s unknown.
Host. - Artemisia biennis WILLD.
Distribution. - NORTH AMERICA.
A d d i t i o n a l s p e c i m e n s e x a m i n e d . - On Artemisia biennis. CANADA:
Alberta, Stettler District, Killan Station, 3 June 1926, coll. A.H. BRINKMAN, det. J. L.
CONNERS (FH). U.S.A.: Minn., Lake Minnetonka, Aug. 1883, FARLOW(NY); Iowa, Iowa
City, Oct. 1887, T. H. MACBRIDE (NY); S.Dakota, Brookings, 20 June 1895, WIIJ.IAMS & ?
(S); Wis., Racine, 23 July 1897, J. J. DAVIS (BPI).
N o t e s . - Paraperonospora artemisiae-biennis is easily recogni-
zable by its ellipsoidal conidia, mostly 40 urn or more long, and the
length/width ratio = ca 2.
3. Paraperonospora chrysanthemi-coronarii (SAWADA) O. CONST,

comb. nov. - Figs 1 E, 3.
BAS.: Plasmopara chrysanthemi coronarii SAWADA - Rep. Govt. Res. Inst. dep.
Agric. Formosa 51: 15. 1931
= Peronospora chrysanthemi-coronarii (SAWADA) S. ITO & TOKUNAGA -
Trans. Sapporo nat. Hist. Soc. 14: 29. 1935
= Bremiella chrysanthemi-coronarii (SAWADA) TAO - Acta mycol. sin. 1: 64.
1982. Type on Chrysanthemum coronarium L., CHINA (TAIWAN): Tai-
chung Prov., Yuanlin, 21 March 1930, E. KUROSAWA (TNS F-192975 - holo-
type).
S p o t s epiphyllous, yellowish, irregular, up to 15 mm diam,
margin diffuse.- Down hypophyllous, greyish to yellowish, felt-
like. - C o n i d i o p h o r e s colourless, erect, 350-600 |im long, trunk
180-430 x 11-17.5 um, base not or slightly swollen; upper part
dichotomously and often trichotomously branched 4-6 times; u l t i -
m a t e b r a n c h l e t s 2-3 at the tip of each branch, often arising from
a slightly swollen, common base, divergent, long-conical, 6-20 um
long, base 2.5-4.5 um, gradually tapering to 1.5-2 um below the
swollen up to 4 jam diam tip. - C o n i d i a colourless to slightly yel-
lowish, ellipsoidal to broadly ellipsoidal, (31)36-42(-50) x (18-)22-
25(-27) urn, broadest part median, tip elongate to round, wall ca
0.3 urn thick; pedicel 1-2 x 2-3 um; a vestigial, nonfunctional dehis-
cence apparatus, appearing as a membrane located under the apex is
present in many conidia. - S e x u a l o r g a n s unknown.
Host. - Chrysanthemum coronarium L.
Distribution. - CHINA (TAIWAN).
A d d i t i o n a l s p e c i m e n s e x a m i n e d . - CHINA (TAIWAN): Taipei, 14 Feb.
1931, K. SAWADA (TNS F-234562); ditto (TNS F-234560); ditto, 13 Feb. 1931 (TNS
F-234561); ditto, 4 Jan. 1946 (TNS F-234572).
87
Fig. 3: Paraperonospora chrysanthemi-coronarii (A-C, E, F from holotype; D from

TNS F-234561); A. conidiophore. - B. branches. - C. ultimate brancbjets. - D, E.
conidia. - F. tips of conidia showing variation of the vestigial papilla.
4. Paraperonospora leptosperma (DE BARY) O. CONST., comb. nov. -

Figs I A-D, 4.
BAS.: Peronospora leptosperma D E BAEY - Annls Sei. nat., Bot., ser. 4 , 2 0 : 48 a n d
121. 1863
= Plasmopara leptosperma ( D E BARY) SKALICKY - Preslia 38: 127. 1966.
Type on Matricaria perforata MKRAT (= Tripleurospermum inodorum (L.)
SCHULTZ BIP.), GERMANY: Freiburg, Autumn 1862, DE BARY in RABENIL,
Fungi europ. 574 (BPI - lectotype, L 910.237-1204 - isolectotype).
SYN.: Peronospora leptosperma GÄUM. - Beitr. KryptogFl. Schweiz 5(4): 129.
1923 as „(DE BARY) GÄUM." Nom. illegit. (Art. 64 ICBN).
Peronospora anthemidis GÄUM. - Beitr. KryptogFl. Schweiz 5(4): 126. 1923
= Plasmopara anthemidis (GÄUM.) SKALICKY - Preslia 38: 127. 1966. Type
on Anthemis austriaca JACQ., CZECHOSLOVAKIA: Teplitz, Frühling (for-
merly AUSTRIA), 1872, F. THÜMEN, Fungi austr. 745 (UPS - lectotype; NY
- isolectotype).
Peronospora achüleae SAVUL. & VAKKY - Arch. Freunde Natgesch.
Mecklenb. 2: 347. 1956 = Plasmopara achüleae (SAVUL. & VANKY) SKALICKY
- Preslia 38: 127. 1966. Type on Achillea atrata L., GERMANY: Rostock,
Neuer Botanischer Garten, 3 Aug. 1940, H. BUHR (BUCM 868 - lectotype);
on Achillea sp., ditto, 12 Aug. 1940 (BUCM 869 - paratype).
Peronospora dimorphothecae SAVUL. & VANKY - Arch. Freunde Natgesch.
Mecklenb. 2: 350. 1956. Type on Dimorphotheca aurantiaca DC, GER-
MANY: Rostock, Neuer Botanischer Garten, 16 July 1950, H. BUHR (BUCM
1564 - holotype; B - isotype).
Peronospora ursiniae SAVUL. & VAKKY - Arch. Freunde Natgesch.
Mecklenb. 2: 352.1956 = Plasmopara ursiniae. (SAVUL. & VAKKY) SKALICKY-
Preslia 38: 127. 1966. Type on Ursinia calenduliflora (DC) N. E. BH., GER-
MANY: Rostock, Neuer Botanischer Garten, 16 July 1950, H. BUHR (BUCM
2885 - lectotype); on U. anthemoides (L.) POIRET (= U. pulchra N.E. BR),
ditto, 9 July 1940 (BUCM 2884 - paratype).
Peronospora buhrii SAVUL. & VANKY- Arch. Freunde Natgesch. Mecklenb.
2: 355. 1956 = Plasmopara buhrii (SAVUL. & VANKY) SKALICKY- Preslia 38:
127. 1966. Type on Ursinia anthemoides (L.) Pom., GERMANY: Rostock,
Neuer Botanischer Garten, 27 July 1947, H. BUHR (BUCM 1233 - lectotype;
B - isolectotype); on U. speciosa DC, ditto, 16 July 1950, H. BUHR (BUCM
1232 - paratype).
S y m p t o m s hardly visible. - Down commonly on leaves but

sometimes on involucral bracts or petals, whitish, rarely yellowish,
scattered, in patches or covering rather restricted areas. - C o n -
i d i o p h o r e s colourless, erect, 290-800 urn long, trunk 120-500 x 6-
15 urn; upper part dichotomously or trichotomously branched 3-5
times; u l t i m a t e b r a n c h l e t s 2-4 at the tip of each branch, often
arising from a slightly swollen common base, long-conical, 8-17 um
long, base 3-6 um, gradually tapering to 1-1.5 urn below the trun-
cate, rounded or slightly swollen tip. - Conidia colourless or pale
yellowish, cylindrical with rounded ends to broadly ellipsoidal
(21-)30-40(-45) x (15-)17-22(-25) um, broadest part median, rarely
supramedian, base round, tip sometimes slightly flattened, wall ca
0.3 um thick; pedicel 1-2 x 1-1.5 urn; a membrane-like structure,
possibly a rudimentary dehiscence apparatus, is rarely visible under
89
the apex. - Oogonia yellowish, irregular, 33-53 (im diam, wall 1.5-
2 urn thick. - Oosporcs aplerotic, globose, (22-)25-27(-31) u.m
diam, wall yellowish to pale brown, smooth, 2-4 |im thick.
H o s t s . - (Cultivated plants marked with asterisk) Achillea
atrata L..*, A. clusiana TAUSCH.*, A. millefolium L., A. umbellata SIEB.
& SM.*, Anthemis altissima L., A. arvensis L., A. austriaca JACQ. and
A. cotula L., Chamomilla aurea (LOEFL.) GAY ex COSSON & KRALIK, C.
recutita (L.) RAUSCHERT, C. suaveolens (PURSH) RYDB., Cotula turbinata
L.*, Dimorphotheca aurantiaca DC*, Lasiospermum bipinnatum
(THUNB.) DRUCE.*, Matricaria caucasica (WILLD.) POIRET, M. perforata
MERAT, Ursinia anthemoides (L.) POIRET.*, U. calenduliflora (DC) N. E.
BR* and U. speciosa DC*.
D i s t r i b u t i o n . -EURASIA.
S e l e c t e d s p e c i m e n s e x a m i n e d o t h e r t h a n t y p e s . - On Achillea
clusiana. GERMANY: Rostock, Neuer Botanischer Garten, 31 Aug. 1937, coll. H. BUHR,
det. T. SAVULESCU (BUCM 3155).
On Achillea millefolium. SWEDEN: Uppland, Lunda, 2 Sept. 1954, A.
GUSTAVSSON 4877 (LD); Skäne, Räften. 14 Sept. 1882, coll. E. LJUNGSTRÖM, det. A.
GUSTAVSSON (LD).
On Achillea umbellata. GERMANY: Rostock, Neuer Botanischer Garten, 12
Aug. 1940, coll. H. BUHR, det. T. SAVULESCU (BUCM 3154); ditto, coll. H. BUHR, det. O.
CONSTANTINESCU (B).
On Anthemis altissima. FRANCE: l'Herault, near St. Martin-de-Loudres, 19
June 1953, E. MAYOR (BPI).
On Anthemis arvensis. FRANCE: Depart, du Tarn, Esperausses, 15-30 Aug.
1927, E. MAYOR (BPI). GERMANY: Bayern, Bayreuth, July 1879, THÜMEN (NY).
ROMANIA: Prahova Dist., Plopeni forest, 19 June 1977, O. CONSTANTINESCU & G.
NKGREAN in Herb, mycol. rom. 2699 (UPS). SWEDEN: Skäne, Perstorp, 2 Sept. 1953,
A. GUSTAVSSON 3493 (LD); ditto, Matteröd, 2 Sept. 1953, A. GUSTAVSSON 3516 (LD);
ditto, Bosjökloster, 8 Sept. 1953, A. GUSTAVSSON 3549 (LD).
On Anthemis austriaca. CZECHOSLOVAKIA: Bohemia, Teplitz (formerly AUS-
TRIA) May 1872, THÜMEN (NY). GERMANY: Bayern, Parsberg Dist., Pielenhofen, 27
May 1937, E. EICHORN in SYDOW, Myc. germ. 3278 (S; UPS); ditto, Riedenburg Dist.,
Prunn, 12 June 1938, E. EICHORN in SYDOW, Myc. germ. 3279 (S, UPS). ROMANIA: Ilfov
Dist., Greci, 25 June 1978, O. CONSTANTINESCU & G. NEOREAN in Herb, mycol. rom.
2923 (UPS).
On Anthemis cotula. GERMANY: Rheinprovinz, Altenkirchen Dist., near
Mühlenberg, 13 Aug. 1933, A. LUDWIG in SYDOW, Myc. germ. 2661 (BPI; S). ROMANIA:
Prahova Dist., Busteni, Valea Cerbului, 10 Oct. 1978, O. CONSTANTINESCU & G.
NEGREAN in Herb, mycol. rom. 2924 (BPI; UPS).
On Chamomilla aurea. IRAQ: Mosul, 9 Apr. 1977, coll. S. A. KHAN, det. G. LUNN
(IMI 223334).
On Chamomilla recutita. CZECHOSLOVAKIA: Bohemia, Tabor, 12 June 1909, F.
BUBAK in SYDOW, Phycom. et Protom. 254 (S, UPS); ditto,8 May 1903 in SYDOW, Phy-
com. et Protom. 211 (UPS). ITALY: Catania, Mascalucia, Mar. 1902, G. SCALIA in SACC,
Myc. ital. 886 (S). GERMANY: Sachsen, Moritzburg, 11 Aug. 1899, W. KRIEGER, Fungi
sax. 1487 in (S). HUNGARY: Ujszasz, May 1927, G. MOESZ in PETRAK. Myc. gen. 340 (S,
UPS); ditto in Flora hung. exs. 801 (S). ROMANIA: Craiova dist., near Isalnita, 10 May
1965, M. COSTESCU in Fl. oil. exs. 513 (UPS). SWEDEN: Uppland, Uppsala, 27 Sep.
1884, L. ROMKI.L (S, UPS); Södermanland, Tuna, July 1915, T. VESTERGREN (S); Skäne,
Mölleberga, Önsvala, 16 Sept. 1953, A. GUSTAVSSON 3683 (LD); Svedala, Aggarp, 5
90
Fig. 4: Paraperonospora leptosperma (A-C from lectotype; D from Finland, Rein-

ikainen, 12 Sept. 1963, H. ROIVAINEN in UPS; D from PETRAK, Myc. gen. 340 in UPS); A.
conidiophore. - B. branch and ultimate branchlets. - C, D, conidia. - E. oogonium and
oospore.
91
Aug. 1953, A. GUSTAVSSON 3004 (LD); Svalöv, 22 Sept. 1928, coll. C. HAMMARLUND, det.
O. CONSTANTINESCU (LD); ditto, 16 Oct. 1929 (LD). U.S.S.R.: Litov SSR, Kaunas,
Botanical Garden, 14 June 1934, A. MINKEVICIUS (S).
On Chamomilla suaveolens. FINLAND: Alandia, Hammarland, Drygsböle, 1
Aug. 1919, T. PUTKONEN in Myc. fenn. 23 (UPS); Regio aboensis, Kakskerta, Monnonen,
29 June 1936, L. E. KARI in Fungi exs. fenn. 738 (UPS). GERMANY: Bayern, distr.
Regensburg, Mosham, 21 June 1936 and 26 June 1937, E. EICHHORN in SYDOW, Myc.
germ. 3079 (S); Sachsen, Moritzburg, 11 Aug. 1899, W. KRIEGER in Fungi sax. 1488 (S).
ROMANIA: Harghita Dist., Gheorgheni, 23 July 1977, O. CONSTANTINESCU & G.
NEGREAN in Herb, mycol. rom. 2700 (UPS). SWEDEN: Gästrikland, Gävle, Lövudden,
24 Aug. 1942, J. A. NANNFELDT in Fungi exs. suec. 1677 (S, UPS); Skäne, Snärestad, 28
July 1953, A. GUSTAVSSON 2864 (LD). U.S.S.R.: Latvij SSR, Latgale Dist., Vidsmuiza,
12 June 1934, K. STARKS 1621 (S); ditto, Vidzeme Prov., Rizh Dist., Maryupe, 18 July
1955, YU. SMARODS in Griby SSSR 114 (L, UPS).
On Cotula turbinata. GERMANY: Rostock, Neuer Botanischer Garten, 14 July
1948, coll. H. BUHR, det. T. SAVULESCU (BUCM 3152); ditto, coll. H. BUHR, det. O.
CONSTANTINESCU (B).
On Lasiospermum bipinnatum. GERMANY: Rostock, Neuer Botanischer Gar-
ten, 16 July 1950, coll. H. BUHR, det. T. SAVULESCU (BUCM 3140).
On Matricaria caucasica. GERMANY: Rostock, Neuer Botanischer Garten, 12
Aug. 1940, coll. H. BUHR, det. T. SAVULESCU (BUCM 3153).
On Matricaria perforata. ENGLAND: Exeter, Stoke Woods, 4 Sept. 1978, G. M.
WATERHOUSE (IMI 232657). FINLAND: Regio aboensis, Korppoo, Bonäs, 11 July 1948,
L. A. KARI in Fungi exs. fenn. 981 (UPS). GERMANY: Bayern, Bayreuth, July 1874,
THÜMEN. Myc. univ. 50 (L, UPS); Brandenburg, Lichtenrade near Berlin, 10 May 1918,
H. SYDOW. Myc. gen. 1516 (S, UPS); Freiburg, summer 1862, DE BARY in RABENH., Fungi
europ. 573 (UPS). POLAND: Schottwitz near Wroclaw (formerly GERMANY:
Breslau), SCHRÖTER, Pilze schles. 385 (S). ROMANIA: Ialomita Dist., Fierbinti railway
station, 21 May 1982, G. NEGREAN (BUCM 70340); Satu-Mare Dist, Dindesti, 16 Aug.
1974, G. NEGREAN in Herb, mycol. rom. 2425 (UPS). SWEDEN: Södermanland, Sock-
holm, Oct. 1915, G. LAGERHEIM (S); Skäne, Bösarp, Markiedal, 3 Aug. 1953, A.
GUSTAVSSON 2959 (LD). U.S.S.R.: Latvian SSR, prov. Zemgale, distr. Tervete, Uplejas,
30 June 1963, E. VIMBA in SMARODS, Fungi latv. exs. 1357 (UPS).
On Ursinia speciosa. GERMANY: Rostock, Neuer Botanischer Garten, 12 July
1947, coll. H. BUHR, det. O. CONSTANTINESCU (B).
N o t e s . - It seems that the original material of this fungus,

distributed in Rabenh. Fungi europ. 574, was not carefully checked.
The copies present at S and UPS contain no fungus, whereas of the
two specimens preserved at L one (910.237-1221) shows Pp. lep-
tosperma and another (910.237-1204) no fungus at all.
The fungus parasitic on Anthemis spp. was traditionally treated
as a separate species, P. anthemidis. Besides the different host, P.
anthemidis was stated to have yellowish conidia vs. colourless ones
in P. leptosperma. However, many specimens exhibiting intergrading
characters were found during this study. Although the conidia of the
fungus on Anthemis are slightly broader, no clear distinction could
be made on the basis of morphological characters.
The pigmentation of the down and conidia in Pp. leptosperma is
variable. The down is almost always white in properly dried speci-
mens, but it may be greyish or yellowish, especially when the drying
92
was poor. The young conidia are commonly colourless but in most
specimens they become yellowish with age.
5. Paraperonospora minor (SAVUL. & RAYSS) O. CONST., comb. nov. -
Fig. 5.
BAS.: Peronospora sulphured GÄUM. forma minor SAVUL. & RAYSS. in SAVUL. -
Herb, mycol. rom. 146. 1930. Type on Artemisia vulgaris L., USSR: Distr.
Cetatea Alba, Crocmaz (formerly in ROMANIA), 22 June 1926, T. SA-
VULESCU & T. RAYSS (BUCM 60146 - lectotype; L - isolectotype).
S p o t s epiphyllous, pale, brown when tissues become necrotic,
polyangular. - Down hypophyllous, whitish or yellowish, in
patches, sometimes powdery, covering large, vein-limited areas or
the entire lower surface, but often obscured by the trichoma. -
C o n i d i o p h o r e s colourless, erect, 160-730 urn long, trunk 90-
400 x 6-15 urn; upper part dichotomously or very rarely tri-
chotomously branched 3-6 times; u l t i m a t e b r a n c h l e t s 2-3 at
the tip of each branch, long-conical, up to 27 urn long when termi-
nal, 6-15 urn when lateral, base 3-5 um, gradually tapering to 1.5-
2 um below the truncate or rounded tip. - Conidia yellowish,
broadly-ellipsoidal, (21-)25-30(-35) x (15-)19-22(-24) um, broadest
part median or, rarely, submedian, wall ca 0.3 urn thick. Oogonia
pale, globose to irregular, 34-45 urn diam, wall 1-3 um thick. -
Oospores aplerotic, globose, 22-26 um diam, wall yellowish,
smooth, 2.5-3 urn thick.
H o s t s . - (Cultivated plants marked with asterisk) Achillea
clusiana TAUSCH.*, Artemisia sieversiana WILLD., A. tilesii LEDEB., A.
vulgaris L. and Leucanthemum nipponicum FRANCH.* (= Chrysan-
themum nipponicum MATSUM).
Distribution. -EURASIA.
A d d i t i o n a l s p e c i m e n s e x a m i n e d . - On Achillea clusiana. GERMANY:
Mecklenburg, Rostock Neuer Botanischer Garten, 20 Sept. 1939, coll. H. BUHK, det. O.
CONSTANTINKSCU (B).
On Artemisia sieversiana. U.S.S.R.: Siberia occid., Omsk Prov., near Omsk, 3
June 1922, MURASHKINSKY (BPI).
On Artemisia tilesii. JAPAN: Ishikari Prov., Hokkaido, Nopporo, 8 May 1920, K.
TOGASHI (BPI, NY); Shiribeshi Prov., Hariusu, 22 July 1930, K. TOGASHI (NY).
On Artemisia vulgaris. DENMARK: Jylland, Skive, 12 July 1909, J. LIND in
SYDOW, Phycom. et Protom. 253 (S, UPS). SWEDEN: Skäne, Oppomanna, Bokenäset,
24 June 1954, A. GUSTAVSSON 4272 (LD); Iistorp, 23 Sept. 1953, A. GUSTAVSSON 3777
(LD); Genarp, Häckeberga, 20 Sept. 1953, A. GUSTAVSSON 3724 (LD); Hörröd, Rib-
betuaröd, 24 Sept. 1953, A. GUSTAVSSON 3798 (LD); Vanstad, Äsperöd, 22 Sept. 1953, A.
GUSTAVSSON 3743 (LD); Eslöv, 15 Sept. 1953, A. GUSTAVSSON 3658 (LD).
On Leucanthemum nipponicum. ROMANIA: Bucuresti, Botanical Garden, Oct.
- Nov. 1970, E. ELIADE (BUCM 1487).
N o t e s . - The size of conidia in Pp. minor is remarkably uni-
form. Besides SAVULESCU & RAYSS (1930), GUSTAVSSON (1953, 1959a)
also noticed that the fungus on A. vulgaris has smaller conidia.
93
Fig. 5: Paraperonospora minor (A, B, E & F from lectotype; C, D, G from various

specimens); A. conidiophores. - B, C & D. branches and ultimate branchlets. - E, G.
conidia. - F. oogonium and oospore.
94
6. Paraperonospora multiformis (TAO & Y. QIN) O. CONST., comb. nov.

- Fig. 6.
BAS.: Bremiella multiformis TAO & Y. QIN - Acta mycol. sin. 1: 62. 1982. Type on
Chrysanthemum coronarium L. var. spatiosum BAILEY, CHINA: Tian-quan
Region, Sichuan Prov., 12 May 1980, Y. QIN & J. Z. QIN 0666 (Herb. Sichuan
Agric. College - holotype).
SYN.: Peronospora crossostephii SAWADA - Trans, nat. Hist. Soc. Formosa 31:
266. 1941. Nom. invalid. (Art. 35.1 ICBN). Type on Crossostephium arte-
misioides LEES (= C. chinense (MAXIM.) MAKINO, CHINA (TAIWAN): Tai-
peh, 8 May 1936, K. SAWADA (BPI - holotype).
S p o t s epiphyllous pale, irregular with diffuse margin. -
Down hypophyllous, whitish to yellowish, scattered. - Co-
n i d i o p h o r e s colourless, erect 250-700 urn long, trunk 180-
460 x 7.5-15 |im, base bulbous, swollen up to 18 urn; upper part
dichotomously, rarely trichotomously branched 3-6 times; u l t i -
m a t e b r a n c h l e t s 2-5 at the tip of each branch, sometimes arising
from a slightly swollen common base, divergent, long-conical, 15-
20 um long when terminal, 7-13 urn when lateral, base 3-5 urn, grad-
ually tapering to 2 urn below the swollen, up to c. 3 urn tip. - Co-
n i d i a colourless or slightly yellowish, broadly ellipsoidal to ovoidal
rarely ellipsoidal, 26-31(-39) x (19-)20-23(-25) um, broadest part
median, wall c. 0.25 um thick; pedicel 0.5-1 x 2-2.5um. - S e x u a l
o r g a n s unknown.
H o s t s . - Artemisia annua L., Crossostephium artemisioid.es
LESS, and Chrysanthemum coronarium L.
Distribution. -CHINA.
A d d i t i o n a l s p e c i m e n e x a m i n e d . - On Artemisia annua. CHINA:
Sichuan, Yaan, 19 Apr. 1982, coll. Y. QIN, det. TAO (Herb. Sichuan Agric. Coll.).
7. Paraperonospora sulphurea (GÄUM.) O. CONST, comb. nov. - Fig. 7.

BAS.: Peronospora sulphurea GÄUM. -Beitr. KryptogFl. Schweiz 5(4): 128. 1923 =
Plasmopara sulphurea (GÄUM.) SKALICKY- Preslia 38: 127. 1966. Type on
Artemisia absinthium L., USSR: near Nowotserkassk, 27 Apr. 1911, O.
TREBOUX in SYDOW, Phycom. et Protom. 280 (UPS - lectotype; BPI, S -
isolectotypes).
SYN.: Peronospora helichrysi TOGASHI & EGAMI in S. ITO & TOKUNAGA - Trans.
Sapporo nat. Hist. Soc. 14: 30. 1935 = Plasmopara helichrysi (TOGASHI &
EGAMI) TAO - Acta mycol. sin. 6: 72. 1987. Type on Helichrysum brac-
teatum (WENT.) ANDREWS, JAPAN: Hokkaido, Ishikari Prov., Sapporo, 26
July 1932, Y. IMAI (n.v.).
Plasmopara pyrethri DUDKA & BURDYUKOVA - Novit. syst. PI. vase. & non
vase. (Kiev) 1976: 240. 1977. Type on Tanacetum corymbosum (L.) SCHULTZ
BIE, U.S.S.R.: Ukr. SSR, Odessa, distr. Baltuanus, Pesniczovka, 24 June
1974 (KW - holotype).
S p o t s epiphyllous, pale, vein limited or with diffuse margin;
attacked tissues become necrotic. - D o w n hypophyllous, sometimes
also epiphyllous, whitish to yellowish to golden-yellow, scattered
95
Fig. 6: Paraperonospora multiformis (A, B, C from holotype; D, H from Artemisia

annua 19 Apr. 1982, QIN & TAO in Sichuan Agric. Coll.; E, F, G from the holotype of
Peronospora crossostephi); A. conidiophore. - B, D. branches. - E, F. ultimate branch-
lets. - C, G, H. conidia.
96
Fig. 7: Paraperonospora sulphurea (A-C from lectotype; D, E from the holotype of

Plasmopara pyrethri; F, G & I from Herb. Mycol. Rom. no. 347 in BPI; H from Heli-
chrysum Orientale in VLA). A. conidiophores. - B, D, F. branches and ultimate
branchlets. - C, E, G-I. conidia.
7 Sydowia, Vol. 41, 1989 97

and obscured by trichoma, to dense, felt-like and above trichoma,

often covering large areas. - C o n i d i o p h o r e s colourless, erect 240-
450 urn long, trunk 110-320 x 5-15 um; upper part dichotomously or
very rarely trichotomously branched 2-3 times; u l t i m a t e
b r a n c h l e t s 2-3 at the tip of each branch, long-conical up to 22 urn
long when terminal, 7-13 urn when lateral, base 3.5-6 um, gradually
tapering to 1-1.5 urn below the truncate or rarely rounded tip. -
C o n i d i a yellowish, broadly ellipsoidal when young but ellipsoidal
when mature, (26-)32-37(-44) x (18-)20-26(-29) um, broadest part
median tip round in young conidia but slightly apiculate in mature
ones, wall ca 0.3 urn thick; pedicel 1.2-2 x 1.5 urn. - S e x u a l
o r g a n s unknown.
H o s t s . -Artemisia absinthium L., A. annua L., A. ludoviciana
NUTT., A. serrata NUTT., A. suksdorfii PIPER, Helichrysum bracteatum
(WENT.) ANDREWS, H. orientate (L.) GAERTNER and Tanacetum cor-
ymbosum (L.) SCHULTZ BIP. (= Pyrethrum corymbosum (L.) SCOP).
D i s t r i b u t i o n . - EURASIA and NORTH AMERICA.
S e l e c t e d s p e c i m e n s e x a m i n e d o t h e r t h a n t y p e s . - On Artemisia
absinthium. ROMANIA: Ilfov Dist., Moldoveni, 8 May 1977, G. NEGREAN (BUCM
48784); Ialomita Dist., Lehliu, 18 June 1933, T. SAVULESCU & T. RAYSS (LD). U.S.S.R.:
Moldav SSR, Lapusna dist., Cornesti (formerly in ROMANIA), 18 June 1931, T. SAV-
ULESCU & T. RAYSS- Herb, mycol. rom. 347 (BPI, L).
On Artemisia annua. ROMANIA: Ialomita Dist., Fierbinti Railway Station, 21
May 1982, G. NEGREAN (BUCM 70338); Ilfov Dist., Gradistea, Sitaru, 21 May 1982, G.
NEGREAN (BUCM 70334); Constanta dist., Deleni, 25 May 1971, O. CONSTANTINESCU &
G. NEGREAN - Herb, mycol. rom. 1980 (UPS).
On Artemisia ludoviciana. U.S.A.: Cal., Santa Barbara, Apr. 1895, W. G. FARLOW
(BPI); ditto, Santa Cruz, May 1885, W. G. FARLOW (BPI); Dakota, Dickey Co., Hollans'
farm, 4 July 1915, J. F. BRENCKLE, Fungi Dakot. 336 (BERN; BPI; IMI 26460; NY);
Kansas, Riley Co., Manhattan, 20 June 1889, KELLERMAN & SWINGLE 1644 (NY); Wis-
consin, Taylor, 11 July 1916, J. J. DAVIS in Fungi Wise. exs. 30 (BPI, K, NY).
On Artemisia serrata. U.S.A.: Iowa, Decorah, July 1885, E. W. D. HOLWAY in ELL.
& Ev, N. Am. Fungi 1804 (BPI; NY, UPS).
On Artemisia suksdorfii. U.S.A.: Cal., Oxnard, 10 May 1939, Coll. C. L. SHEAR,
det. J. A. STEVENSON (BPI 187922).
On Helichrysum bracteatum. CHINA: Sichuan Prov., Yaan, 6 June 1988, TAO JIA-
FENG 1387 (Herb. Sichuan Agric. Univ.); U.S.S.R.: near Vladivostok, 13 Sept. 1968, E.
S. NELEN (VLA).
On Helichrysum Orientale. U.S.S.R.: Sakhalin, Novo-Alexandrovskoe, 16 Aug.
1960, E. S. NELEN (VLA).
N o t e s . - The holotype of Peronospora helichrysi was not avail-

able. Nevertheless, both the trichotomic condition in same ramifica-
tions and the size of conidia mentioned in the description are rele-
vant for Paraperonospora. Helichrysum is also parasitized by a
Plasmopara in North America.
98
8. Paraperonospora tanaceti (GÄUM.) O. CONST., comb. nov. - Fig. 8.

BAS.: Peronospora tanaceti GÄUM. - Beitr. KryptogFl. Schweiz 5 (4): 130. 1923
= Plasmopara tanaceti (GÄUM.) SKALICKY - Preslia 38: 127. 1966. Type on
Tanacetum vulgäre L., CZECHOSLOVAKIA: Bohemia, near Teplitz (for-
merly in AUSTRIA), summer 1873, THÜMEN in Fungi austr. 1235 (UPS -
lectotype).
S y m p t o m s hardly visible on the upper leaf surface. - Down

hypophyllous, yellowish, rarely whitish, dense, felt-like, often cov-
ering the whole leaf. - C o n i d i o p h o r e s colourless, erect, 350-
800 urn long, trunk 160-400 x 7-15 urn; upper part dichotomously,
rarely trichotomously branched 3-5 times; u l t i m a t e b r a n c h l e t s
2-3 at the tip of each branch, when three often arising from a
slightly swollen common base, long-conical, 8-22 urn long, base 3-
5 urn, gradually tapering to 1-2 urn below the rounded or rarely
slightly swollen tip. - C o n i d i a colourless when young, pale yellow-
ish when mature, broadly ellipsoidal, (33-)37-40(-52) x (21-)23-28
(-29) um, broadest part median or supramedian, base round or grad-
ually narrowing, tip round to slightly flattened, wall ca 0.3 urn thick;
pedicel 1-1.5 x 1-2 urn, longer in young conidia. - O o g o n i a irregu-
lar, 34-55 um diam, wall colourless or pale yellowish, 1-3 urn thick. -
Oospore aplerotic, (22-)27-36(-42) (im diam, yellowish, globose,
wall smooth, 3-7 urn thick.
Host. - Tanacetum. vulgäre L.
Distribution. -EUROPE.
S e l e c t e d s p e c i m e n s e x a m i n e d o t h e r t h a n t y p e . - CZECHOSLO-
VAKIA: Sternberg, June 1923, PETRAK, Flora Bohem. et Morav. exs. 1514 (S); Moravia,
Rüdersdorf (formerly GERMANY), June 1889, P. SYDOW, Myc. march. 2654 (S; UPS).
DENMARK: Fyn, Skärup, 7 June 1874, E. ROSTRUP (UPS); ditto, 11 June 1883 (S).
FINLAND: Nyl., Helsinki, Munkkiniemi, 27 July 1935, V. LETIIOLA in Myc. fenn. 327
(UPS); Regio aboensis, Turku, Ruissalo, 19 July 1959, L. A. KARI. in Fungi exs. fenn.
994 (UPS); Karelia australis, Haapasaari, 8 July 1953, L. E. KARI in Fungi exs. fenn.
996 (UPS). FRANCE: Seine-et-Marne, Fontainbleau, May 1896, FEUILLEAUBOIS (S);
sine loc. & dat., FAUTREY (herb. Fries in UPS). GERMANY: Bayern, Bayreuth, June
1879, THÜMEN (UPS); ditto, June 1875, A. WALTHER (S); ditto, 1880 (UPS); ditto, sum-
mer 1875, THÜMEN Myc. univ. 424 (UPS); Bayern, Hassfurt, June 1898, ALLESCHER &
SCHNABLI, Fungi bavarici 646 (S); Brandenburg, Hönow near Berlin, 5 June 1910, H.
SYDOW, Myc. germ. 884 (S; UPS); Hessen, Ostrich, FUCKEL, Fungi rhen. 1606 (S). NOR-
WAY: Dovrefjeld, N. Knutshö, 9 July 1925, A. G. ELIASSON (S, UPS). POLAND:
Wroclaw (formerly Breslau, GERMANY), sine dat, J. SCHRÖTER, Pilze schles. 385 (S).
SWEDEN: Helsingland, Söderhamn, 7 Aug. 1926, T. VESTERGREN (S); Jämtland, Äs, 17
July 1920, K. FALCK (LD); Norrbotten, Luleä, Aug. 1915, J. VLEUGEL (S); ditto, Aug.
1913 (S, UPS); Södermanland, Fjäderholmen, spring 1907, T. VESTERGREN (S). Öland,
Vickleby, 2 Aug. 1957, A. GUSTAVSSON 5727 (LD). USSR: Akmolensk prov., Albasarsk
dist. sine dat., coll. S. GANESEININ, det. A. JACZEWSKI (S).
9. Peronospora radii DE BAEY. - Fig. 9.

Annls Sei. nat., Bot., ser. 4, 20: 48-49 and 121. 1863. Type on Matricaria perforata
MERAT (= Tripleurospermum inodorum SCHULTZ BIP), GERMANY: Freiburg, Summer
1862 in RABENH., Fungi europ. 573 (see notes).
7. 99
Fig. 8: Paraperonospora tanaceti (A-D from the lectotype; E from Fungi exs. fenn. 994
in UPS; F from Fungi exs. fenn. 997 in UPS; G from Fungi exs. fenn. 996 in UPS); A.
conidiophores. - B, F. branches and ultimate branchlets. - C-E. conidia. - G.
oogonium and oospore.
100
SYN : Peronospora radii DK BARY var. epiphylla POIKAUI.T - Bull. Assoc. Nat.
Nice. 2(3): 9. 1915. Type on Coleostephus myconis (L.) REICIIENB. fil. (=
Chrysanthemum myconis L.), FRANCE: Cap d'Antibes. Collection not
indicated.
Peronospora danica GAUM. - Beitr. KryptogFl. Schweiz 5(4): 128. 1923.
Type on Chrysanthemum segetum L., DENMARK: near Lingby, 7 Oct.
1913, J. LIND in SYDOW, Phycom. et Protom. 309 (UPS - lectotype; BPI, S -
isolectotypes).
Down commonly on florets, rarely on leaves and more rarely
on stems, greyish, ochreous, greyish-brown to greyish-violaceous,
scattered or densely agglomerated, sometimes completely covering
all the florets within the inflorescence or large areas of the leaves. -
C o n i d i o p h o r e s colourless, the branched part rarely yellowish,
erect, rarely flexuous, 280-650 |im long, trunk 120-450 x 8-18 urn,
base swollen up to 25 urn; upper part dichotomously branched 4-6
times, very rarely three branches may arise from the trunk, branches
straight or rarely flexuous; u l t i m a t e b r a n c h l e t s 1-3 at the tip of
each branch, long-conical to almost cylindric, up to 20 urn long
when terminal, 5-12 um when lateral, base 2.5-4 um, gradually
tapering to c. 1.5-2 urn below the truncate tip. - Conidi a brownish,
obovoidal to broadly ellipsoidal or ellipsoidal, (24-)30-35(-42) x
(18-)20-23(-32) urn, broadest part median or slightly supramedian,
base gradually narrowing, tip rounded, elongate or variously apicu-
late, wall ca 0.5 urn thick, smooth to finely verruculose; pedicel 1-
2 x 1.5-2 jam or only slightly protruding. - O o g o n i a in florets, yel-
lowish, globose to irregular, 38-58 urn diam, wall 1-2 Jim thick. -
O o s p o r e s aplerotic, golden-yellowish, globose, 24-34 urn diam,
wall smooth 1-4 um thick.
H o s t s . - Achillea ptarmica L., Anthemis arvensis L., A.
ruthenica BIEB., Artemisia mexicana WILLD. ex SPRENG., Chamomilla
recutita (L.) RAUSCHERT, Chrysanthemum segetum L., Leucanthemum
vulgäre LAM. (= Chrysanthemum leucanthemum L.), Matricaria ma-
ritima L. (= Tripleurospermum maritimum (L.) KOCH), M. perforata
MERAT.
D i s t r i b u t i o n . - EUROPE, ISRAEL, MEXICO.
S e l e c t e d s p e c i m e n s e x a m i n e d o t h e r t h a n t y p e s . - On Achillea
ptarmica. SWEDEN: Södermanland, Strengnäs, July 1913, G. LAGERIIEIM in Vesterg,,
Microm. rar. sei. 1669 (S, UPS).
On Anthemis arvensis. GERMANY: Hessen-Nassau, Manderbach, 24 June 1934,
A. LUDWIG (BPI, S); Zahlendorf near Berlin, Aug. 1895, P. SYDOW, Myc. march. 4329 (S,
UPS); Bayern, Unterfranken, Brünnstadt, Aug. 1904, A. VILL, Fungi bavarici 767 (S);
ditto, July 1912 in SYDOW, Myc. march. 1084 (S, UPS). POLAND: Zedlitz near Wroclaw
(formerly GERMANY, Breslau), June, July 18H6, SCHNEIDER in RABENH., Fungi eur.
1369 (S, UPS); Wilanöw near Warszawa, 1 June 1958, J. KOCHMAN, Myc. pol. 14 (UPS).
SWEDEN: Öland, Stora Rör, 15 May 1929, A. G. ELIASSON (S);
On Anthemis ruthenica. ROMANIA: Satu-Mare Dist., Sanislau, 30 May 1983, G.
NEGREAN (BUCM 76638; UPS); Dolj Dist., Cernele, 18 May 1977, O. CONSTANTINF.SCU &
G. NEGREAN in Herb, mycol. rom. 2690 (UPS).
101
Fig. 9. Peronospora radii (from various specimens); A. conidiophores. - B & C.

branches and ultimate branchlets; D-H. conidia. - I. oogonium and oospore.
102
On Artemisia mexicana. MEXICO: Chihuahua, 17 Aug. 1980, coll. R. M. ALVAREZ,

det. D. WEAST (BPI).
On Chamomilla recutita. BELGIUM: Groenendael, 1 June 1890, E. BOMMEH & M.
ROUSSEAU in ROUMEG, Fungi sei. exs. 5615 (UPS). FRANCE: Lille, 4 June 1918, A.
LUDWIG (S). POLAND: West Beskiden Mt, Wsetin, June 1923, F. PKTRAK, Myc. carpat.
319 (S). SWEDEN: Skäne, Svalöv, 21 Sept. 1928; Oct. 1928; 16 Oct. 1929, C. HAM-
MARLUND (LD);
On Chrysanthemum segetum. DENMARK: Lyngby, 7 Oct. 1913, J. LIND (S); ditto,
30 Sept. 1913 (BPI, LD). SWEDEN: Hallan, Äled, Enslöv, 4 Sept. 1952, H. ANDERSON
1068 (LD).
On Leucanthemum vulgäre. CZECHOSLOVAKIA: Bohemia, Kamärover Teich
at Dymokur, 7 June 1900, F. BUBÄK in VESTERG., Microm. rar. sei. 347 (UPS). FIN-
LAND: Nylandia, Kerava, Alikerava, 11 July 1965, H. ROIVAINEN (LD). SWEDEN:
Södermanland, Strengnäs, July 1913, G. LAGERHEIM in VESTERG., Microm. rar. sei. 1670
(S, UPS); Stockholm, Frescati, 1923, H. DAHLSTEDT (S); Uppland, Norrtälje, Aug. 1915,
G. LAGERHEIM in Fungi exs. suec. 1684 (S, UPS); Ljusterö, Äpplerö Isl., 5 July 1927, T.
VESTERGREN (S).
On Matricaria maritima. ISLAND: Reykjavik, 16 July 1962, J. A. NANNFELDT
(UPS). SWEDEN: Gotland, Västerhejde, 30 June 1972, M. THULIN 2010 (UPS); Upp-
land, Storvreta, Vretalund, 11 July 1981, N. LUNDQVIST 13260 (UPS); ditto, 2 July 1967,
N. LUNDQVIST 5325 (UPS); Uppsala, 29 June 1959, R. SANTESSON 12832 (UPS).
On Matricaria perforata. CZECHOSLOVAKIA: Bohemia, Laschin, Schlössles,
June 1917, R. STEPPAN in PETRAK, Flora Boch. & Morav. exs. 1345 (S). DENMARK:
Jylland, Frijsenborg, 26 June 1915, J. LIND (UPS); Nebsager, 16 July 1892, O. ROSTRUP
(S); Fölle, 25 July 1874, E. ROSTRUP (UPS); Fyn, Hindsgavl, 7 Aug. 1928, R. SERNANDER
(UPS); Falster, Stubbeköbing, 1 Aug. 1876, E. ROSTRUP (UPS). ENGLAND: Herts.,
Sandridge, Aug. 1941, A. SMITH (S). FINLAND: Alandia, Saltvik, Hjortö, 8 July 1919,
T. PUTKONEK in Myc. fenn. 22 (UPS); Regio aboensis, Lohja, Karkalinniemi, Lin-
naniemi, 29 July 1955, L. E. KARI in Fungi exs. fenn. 986 (UPS); Masku, Taipale, 8 July
1956, L. E. KARI in Fungi exs. fenn. 987 (UPS); Turku, Ravattula, 5 July 1937, L. E.
KARI in Fungi exs. fenn. 988 (UPS). GERMANY: Bayern, Bayreuth, Summer 1877,
WALTHER (UPS); ditto in THÜMEN, Myc. univ. 135 (UPS); Hessen-Nassau, near Lan-
genaubach, 14 July 1935, A. LUDWIG in SYDOW, Myc. germ. 2898 (S, UPS). NORWAY:
Hordaland, Hardanger, Jondal, Herandsbygden, 10 July 1908, S. K. SELLAND (S).
SWEDEN: Västmanland, Hamre, 19 July 1924, J. A. NANNFELDT in Fungi exs. suec. 697
(S, UPS); Södermanland, Strengnäs, July 1913, G. LAGERHEIM in VESTERG., Microm.
rar. sei. 1671 (S, UPS); Uppland; Nortälje, Aug. 1915, G. LAGERHEIM (S); ditto, in Fungi
exs. suec. 1684 (UPS); Vaksala par., Skölsta, 11 July 1931, H. SMITH (UPS); ditto in
Fungi exs. suec. 199 (S); Uppsala, Norbyskogen, 4 July 1926, J. A. NANNFELDT (UPS);
Norrbotten, Rokliden, Aug. 1908, N. SYLVEN in VESTERG., Microm. rar. sei. 1418 (UPS).
U.S.S.R.: Latvij SSR, Zemgale Prov., Dobele Dist, Tervete, 30 June 1963, E. VIMBA in
Fungi latv. exs. 1357 (UPS).
N o t e s . - Original specimens of P. radii have been distributed in

RABENH., Fungi europ. 573. As in the case of P. leptosperma, no. 574 of
the same exsiccatum, it seems that the material was carelessly
selected. The copies in S and UPS contain no P. radii but few con-
idiophores and conidia of Pp. leptosperma on the involucral bracts
and/or the leaves. Although no original specimens was examined,
there is little doubt about the identity of this rather common species.
A lectotytpe should be selected from among the copies of this exsic-
catum, if a representative such specimen exists.
Peronospora radii is a variable species. When a limited number
103
of specimens are examined, it appears that the form on the leaves

has more slender conidiophores and branches, longer ultimate
branchlets and larger conidia, compared to the form on florets.
However, intermediates or forms contradicting this pattern are
found when more specimens are seen.
Peronospora radii is commonly found on the ligulate florets of
its host. Following the description of P. danica by GÄUMANN (1923), on
materials collected by J. LIND and distributed in SYDOWS Phycom. et
Protom, no. 309, it was considered that the morphologically similar
fungus attacking the leaves is a different species. However,
GUSTAVSSON (1959: 214) found among the original specimens of LIND
preserved at C and CP one specimen with the fungus present on both
florets and leaves. Actually it was POIRAULT (1915) who first men-
tioned an attack on the leaves of Chrysanthemum myconis L., now
known as Coleostephus myconis (L.) REICHEND, fil., in South of
France. He even found plants showing the fungus on both leaves and
corolla, but his paper, as well as the new variety epiphylla intro-
duced for the form attacking the leaves, have been overlooked by
subsequent authors. Recently BEN-ZE'EV et al. (1987) found plants of
cultivated Argyranthemum frutescens SCHULTZ BIP. (= Chrysan-
themum frutescens L.) and wild Chrysanthemum coronarium L.
showing attack of P radii on both leaves and petals. By isozyme
analysis they demonstated that the same fungus is responsible for
these two forms of attack. During this study additional specimens
showing attack on flowers and leaves were found, particularly on
Anthemis. In one specimen (KOCHMAN, Myc. pol. 14 in UPS) the
fungus was present on florets, leaves and stems.
Peronospora radii is sometimes associated with Pp. lep-
tosperma. Several specimens of Anthemis spp., Chamomilla recutita
and Matricaria perforata having florets attacked by P. radii and the
involucral bracts and/or leaves attacked by Pp. leptosperma were
found during this study.
Important damages produced by Peronospora radii to culti-
vated Chrysanthemum spp. and/or Argyranthemum frutescens, were
reported from England (WILCOX, 1961), Yugoslavia (MIJUSKOVIC, 1968)
and Israel (KENNETH & BEN-ZE'EV, 1985; BEN-ZE'EV & al., 1987).
Doubtful and excluded taxa

Peronospora brachycomes ENKINA - Mikol. i Fitopatol. 3: 482.
1969. Holotype on Brachycome iberidifolia BENTH., U.S.S.R.:
Novosibirsk, Botanical Garden, 3 Aug. 1966, T. V. ENKINA The holo-
type is preserved at LE, but was not sent upon request. The mor-
phological characters are reminiscent of Paraperonospora lep-
tosperma or Pp. sulphurea. Brachycome belongs to the tribe
104
Astereae and is a very improbable host for a Peronospora. This spe-

cies was not included in the monograph of U.S.S.R. Peronosporales
(NOVOTEL'NOVA & PYSTINA, 1985).
Peronospora leptosperma DE BARY var. siegesbeckiae LAGERH. -
Bull. Herb. Boissier 3: 61. 1895 = Plasmopara siegesbeckiae (LAGERH.)
TAO - Acta mycol. sin. 6: 69. 1987 as ,(LAGERH. in SACC.)'. Type on
Siegesbeckia, ECUADOR: Pichincha. The authentic material was
not available but specimens of Siegesbeckia collected from Peru,
China and Taiwan contain Plasmopara.
Peronospora pospelovii GAPONENKO - Semejstvo Per-
onosporaceae Srednej Azii i Yuzhnogo Kazakhstana p. 316. 1972.
Holotype on Aster dahuricus BENTH. ex BAKER (= Galatella dahurica
De), U.S.S.R.: Kirgiz SSRN, Tianshan region, Koczar distr.,
Czapyldyk, 26 July 1946, A. G. POSPELOV. The holotype is preserved at
TASM, but was not sent upon request. This taxon is apparently
based on conidia of a Peronosporaceous fungus attached to the
leaves of Galatella. It is very unlikely that the host, a member of the
tribe Astereae, may bear a Peronospora.
Peronospora senecionis FUCKEL - Symb. mycol. p. 69. 1870. Type
on Senecio cordatus KOCH, GERMANY: Oberbayern, near Hohensch-
wangau, summer. No authentic material is available. The host
belong to the tribe Senecioneae on which the occurrence of Per-
onospora is highly improbable. According to GÄUMANN (1923) this is
Bremia lactucae REGEL.
Peronospora sonchi GAPONENKO - Not. syst. Sect, crypt. Inst. bot.
Acad. Sei. U.S.S.R. 8: 87. 1952 (as ,Napon.'). Holotype on Sonchus
oleraceus L., U.S.S.R.: Uzbekistan SSR, Margelan distr., Frunze Col-
lectiv Farm, 25 May 1949. Authentic material was requested from
TASM but not received. Judging from the description and illustra-
tion it may be Paraperonospora leptosperma. However, the host is a
member of the tribe Lactuceae and thus very improbable of being
parasitized by this fungus.
Acknowledgments
I am indebted to the directors and curators of the herbaria B, BERN, BPI, FH,
IMI, KW, L, LD, NY, PC, S, TNS, UPS, VLA and Sichuan Agric. College for loan of
material, to Dr. Kare BREMER for informations concerning the taxonomy of the hosts
and to Mrs Ulla-Britt SAHLSTRÖM for photography. This work was supported by
grants from the Swedish Natural Science Research Council and Swedish Council for
Forestry and Agricultural Research.
Literature cited
BARY, A. DE (1863). Recherches sur le developpemcnt de quelques champignons para-
sites. - Annls Sei. Nat., Bot., ser. 4, 20: 5-148.
BEN-ZE'EV, I. S., R. G. KENNETH & M. R. BONDE (1987). Peronospora radii DE BY., a
105
causal agent of downy mildew of Anthemideae: complementary description

and new hosts recorded in Israel. - Phytoparasitica 15: 51-67.
BREMER, K. (1987). Tribal interrelationships of the Asteraceae. - Cladistics 3: 210-253.
CONSTANTINESCU, O. (1968). Plasmopara ammii sp. nov. - Sydowia 20: 268-270.
— & G. NEGKEAN (1983). Check-list of Romanian Peronosporales. - Mycotaxon 16:
537-556.
CORDA, A. C. I. (1837). Icones Fungorum hucusque cognitorum. 1. - Pragae.
CRONQUIST, A. (1955). Compositae. In: HITCHCOCK C. L. et al. (eds). Vascular Plants of
the Pacific Northwest. Part 5. - University of Washington Press, Seattle.
CRUTE, I. R. (1981). The host specificity of peronosporaceous fungi and the genetics of
the relationship between host and parasite. In: SPENCER, D. M. (ed.). The
Downy Mildews. Pp. 237-253. - Academic Press, New York & London.
DICK. M. W., P. T. W. WONG & G. CLARK (1984). The identity of the oomycete causing
,Kikuyu Yellows', with a reclassification of the downy mildews. - Bot. J. Linn.
Soc. 89: 171-197.
DUDKA. I. & L. BURDYUKOVA (1977). A new species of the genus Plasmopara SCHROETER
from USSR. - Novit. syst. PI. vase. & non. vase. (Kiev) 1967: 240-241. (In
Russian).
DZHANUZAKOV, A. (1962). The specialization and variability of Peronosporaceous
fungi. - Bot. Zh. SSSR 47: 862-866. (In Russian).
GÄUMANN, E. (1923). Beiträge zu einer Monographie der Gattung Peronospora CORDA.
- Beitr. KryptogFl. Schweiz 5(4): 1-360.
GUSTAVSSON, A. (1953). Some species of the genera Peronospora and Pseu-
doperonospora new to Sweden. - Bot. Notiser 1953: 48-64.
— (1959 a). Studies on Nordic Peronosporas. I. Taxonomic part. - Opera bot. 3(1)
1-271.
— (1959 b). Studies on Nordic Peronosporas. II. General account. - Opera bot. 3(2)
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HOLMGREN, P. K., W. KEUKEN & E. K. SCHOFIELD (1981). Index Herbariorum. Part I. The
Herbaria of the World, 7th ed. - Regnum veg. 106: 1-452.
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downy mildew of marguerite daisy in Israel. - Phytoparasitica 13: 151-152.
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parasites. - Ann. Bot. 36: 1029-1040.
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107

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Peronospora complex on Compositae

CONSTANTINESCU, O. (1989). Peronospora complex on Compositae. - SYDOWIA

Two species of Peronospora parasitic on different organs of var-

Two P. leptosperma-like fungi were described in the genus Plas-

Material and methods

the general pattern of the branching is dichotomous, the trichotomy

6 Svdowia, Vol. 41, 1989 81

Fig. 1: Paraperonospora spp. A-D. Pp. leptosperma. - E . Pp. chrysanthemi-coronarii.

show combined characters of different genera and this is also

pointed out in the case of Anthracoidea (Ustilaginales), „the excep-

short, slightly eccentric. - Oogonia irregular, thin-walled;

Key to the species treated:

1. Paraperonospora artemisiae-annuae (LEE & M. C. TAI) O. CONST,

2. Paraperonospora artemisiae-biennis (GÄUM.) O. CONST, comb. nov.

Fig. 2: Paraperonospora artemisiae-biennis (lectotype); A. conidiophore. - B. ultimate

3. Paraperonospora chrysanthemi-coronarii (SAWADA) O. CONST,

Fig. 3: Paraperonospora chrysanthemi-coronarii (A-C, E, F from holotype; D from

4. Paraperonospora leptosperma (DE BARY) O. CONST., comb. nov. -

S y m p t o m s hardly visible. - Down commonly on leaves but

Fig. 4: Paraperonospora leptosperma (A-C from lectotype; D from Finland, Rein-

N o t e s . - It seems that the original material of this fungus,

Fig. 5: Paraperonospora minor (A, B, E & F from lectotype; C, D, G from various

6. Paraperonospora multiformis (TAO & Y. QIN) O. CONST., comb. nov.

7. Paraperonospora sulphurea (GÄUM.) O. CONST, comb. nov. - Fig. 7.

Fig. 6: Paraperonospora multiformis (A, B, C from holotype; D, H from Artemisia

Fig. 7: Paraperonospora sulphurea (A-C from lectotype; D, E from the holotype of

7 Sydowia, Vol. 41, 1989 97

and obscured by trichoma, to dense, felt-like and above trichoma,

N o t e s . - The holotype of Peronospora helichrysi was not avail-

8. Paraperonospora tanaceti (GÄUM.) O. CONST., comb. nov. - Fig. 8.

S y m p t o m s hardly visible on the upper leaf surface. - Down

9. Peronospora radii DE BAEY. - Fig. 9.

Fig. 9. Peronospora radii (from various specimens); A. conidiophores. - B & C.

On Artemisia mexicana. MEXICO: Chihuahua, 17 Aug. 1980, coll. R. M. ALVAREZ,

N o t e s . - Original specimens of P. radii have been distributed in

of specimens are examined, it appears that the form on the leaves

Doubtful and excluded taxa

Astereae and is a very improbable host for a Peronospora. This spe-

causal agent of downy mildew of Anthemideae: complementary description

SKALICKY, V. (1966). Taxonomie der Gattungen der Familie Peronosporaceae. - Preslia

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