Systematics and diversity of the Phyllachoraceae associated with

Systematics and diversity of the Phyllachoraceae associated
with Rosaceae, with a monograph of Polysiigma
P.F.CANNON
International Mycological
Institute, Bakeham Lane, Egham, Surrey TW20 9TY, U.K.
Biotrophic members of the Phyllachoraceae are described and illustrated, and their nomenclature assessed. Isothea is considered to be
a monotypic genus with close affinities to Phyllachora. Four species of Phyllachora are accepted; some have distinctive features which
may provide evidence for subdivision of that genus. Two species currently referred to Plectosphaera are studied; neither is a typical
biotroph. Polysiigma is formally monographed. Five species are accepted, all on species of Prunus, including the previously
undescribed P. amygdalinum,
an economically important pathogen of almond (Prunus dulcis). An eastern Russian species originally
described in Polystigmella is treated as a subspecies of the widespread plum pathogen Polysiigma rubrum. A number of species names
is excluded from the Phyllachoraceae.
Biotrophic representatives o f the Phyllachoraceae associated
ISOTHEA
with species o f R o s a c e a e appear t o b e relatively small in
I s o t h e a Fr., Summa
number (12 species o f fungi k n o w n from 3100 plant species),
Isothea
but include a number o f e c o n o m i c a l l y important taxa. T h e
C. Bab. e x Berk, (non Corda). Five o r six further t a x a h a v e b e e n
important genus Glomerella
transferred t o Isothea
(anamorph Collelotrichum)
also has
Vegetabilium
Scandinaoiae:
421, 1849.
w a s erected for the single species Sphaeria
rhytismoides
b y different authors, but n o n e is well-
several important representatives associated with R o s a c e a e ,
known. It is n o t o b v i o u s from the original description w h y
but species o f this genus are necrotrophic or saprobic for at
Fries regarded the genus as distinct, but at that period m a n y
least m o s t o f their life cycle, and species concepts as currently
o f the g e n e r a n o w currently accepted w e r e either n o t y e t
recognized are n o t comparable (Cannon, in press). T h e y are,
recognized, or p o o r l y delimited. Isothea
therefore, not included in this survey.
related t o Phyllachora,
T h e apparent lack o f diversity o f Phyllachoraceae associated
with the R o s a c e a e can partly b e explained b y the
pre­
is clearly c l o s e l y
but seems t o differ primarily in having
peridial cells which are large and thick-walled. T h e ostiolar
structure with clavate p i g m e n t e d peridial cells outside o f true
dominantly temperate distribution o f the plant family; about
periphyses is n o t k n o w n in Phyllachora,
80 % o f k n o w n species o f Phyllachoraceae are tropical (Cannon,
b o t h Phyllachora
in press). A notable e x c e p t i o n is hothea
below) h a v e ostioles surrounded b y conic-papillate cells.
rhytismoides
(see
physocarpi
although interestingly
and Plectosphaera
fragariae
below), which extends at least to 83° N in the Canadian
O s t i o l a r structure has n o t b e e n studied in depth in
Arctic.
A n anamorph is n o t present in
The
transfer
o f Isothea
(see
Phyllachora.
Isothea.
rhytismoides
to
Phyllachora
by
Vasil'eva (1987) is. unjustified, and is formally incorrect as
MATERIALS AND
Isothea
METHODS
predates mmlachora.
b e found
Dried reference specimens o f Phyllachoraceae were e x a m i n e d
from a wide range o f herbaria. S o m e were collected b y various
collaborators at the request o f the author t o ensure
T h e t w o g e n e r a m a y ultimately
relatives, but m o r e studies in
the
variation within b$Q0pchora are required before this hypothesis
can b e properly tested.
the
availability o f material in the correct state o f maturity.
Stromata from the dried collections w e r e dissected in situ,
in a drop o f water. T h e c o n t e n t s were allowed t o swell before
b e i n g r e m o v e d from the specimen using the tip o f a scalpel
blade. Squash mounts were made and e x a m i n e d in water and
lactofuchsin, and sections cut using a freezing m i c r o t o m e .
Nomarski and phase optics were e m p l o y e d . A l l p h o t o g r a p h s
and measurements are o f material m o u n t e d in lactofuchsin
unless specified otherwise.
to b e H o s e
I s o t h e a r h y t i s m o i d e s Fr., Summa
421,
Vegetabilium
Scandinaoiae:
1849.
Sphaeria rhytismoides C. B a b . e x Berk., Ann. Mag. Nat. Hist. 6:
361, 1841, non Sphaeria
rhytismoides
Corda,
Icones
Fungorum
Sphaerella
4:-42,
1840.
rhytismoides
(Fr.) D e Not., Comm.
Soc. Critt.
Ital.
2 (3): 488, 1867.
Laestadia
1882.
rhytismoides
(Fr.) S a c c , Sylloge
Fungorum
1 : 424,
Hypospila rhytismoides (Fr.) Niessl, in Rabenhorst, Fungi
no. 3 2 6 1 , 1 8 8 5 , in sched.
Carlia
rhytismoides
(Fr.) Kuntze, Revisio
Generis
Europaei
Plantarum
2:
Guignardia
rhytismoides
(Fr.) Zahlbr., Krypt. exs. Mus.
Palat.
Vindoh. no. 6 1 8 , 1 9 0 1 , in sched.; Ann. k.k. Naturh.
Hofmus.
Wien 1 6 : 66, 1 9 0 1 .
rhytismoides
Lokuloaskomitsety
Sphaeria
(Fr.) Lar. N . Vassiljeva, Pirenomitsety
Severa
Dal'nego
dryadis Fuckel, Fungi Rhenani
Vostoka:
i
225, 1987.
no. 2 1 6 1 , 1 8 6 8 , in sched.
Host species:
Dryas
integrifolia,
Distribution:
Austria, Canada, Faeroe Islands, Germany, Greenland,
Iceland, Italy, Norway,
Anamorph:
United Kingdom (Scotland).
not known.
areas
sometimes
usually
1-2-5 m m
reddish
not
diam.,
brown.
visible from
the
Stromata
the
lower
included
leaf
tissue
hologenous
(though
surface
to
tomentum), the ostioles epigenous. Blackened
due
leaf
tissue sometimes
spreading b e t w e e n ascomata b e l o w the cuticle but much m o r e
heavily developed a b o v e and b e l o w ascomata (in roughly
circular regions 1 2 0 - 2 0 0 um diam.), restricted to an upper and
lower layer, subcuticular, c o m p o s e d o f heavily melanized
epidermal and some palisade cells, fungal tissue n o t clearly
visible, eventually c o v e r i n g almost the w h o l e o f the infected
area. Ascomata
ascospores,
Typification:
United Kingdom: Scotland: Sutherland, Assynt, Inchnadamph, on Dryas octopetala, Sep. 1838, C. Babington (K! - holotype
of Sphaeria rhytismoides
C. Bab. ex Berk.). Germany: in Bavariae
superioris alpibus, on Dryas octopetala, aestate, s. coll. (Fuckel, Fungi
Rhenani no. 2161, ?G - holotype, IMI! - isotype of Sphaeria
dryadis).
Description
Infected
to 7 um long, widely clavate,
mid brown, formed directly from germinating
occasionally while still in the ascoma.
846, 1 8 9 1 .
Phyllachora
smooth-walled, aseptate, often strongly guttulate, without a
gelatinous sheath. Appressoria
usually
strongly
aggregated,
clearly
dis­
Illustrations:
D.
octopetala.
Romania, Russia, Sweden, Switzerland,
Figs 1—2.
This arctic-alpine species is common throughout most of the
range of its host genus Dryas, though it appears not to extend
southwards along the Rocky Mountain range in Western
Canada and the U.S.A. Collections from central Europe appear
to have slightly longer ascospores than those from more
northerly latitudes, but the number of specimens examined in
detail was insufficient to confirm this impression. Canadian
collections seem more or less identical to those from northern
Europe.
tinguishable from the surface even in old colonies, from the
The
surface usually strongly domed, the ostiole inconspicuous;
rhytimoides
name on which Isothea
rhytismoides
is based,
Sphaeria
C. Bab. e x Berk., appears to b e a later h o m o n y m
2 0 0 - 3 2 0 um diam., usually slightly o b l a t e ; peridium 8 - 1 4 um
of S. rhytismoides
thick, c o m p o s e d o f 4 - 5 layers o f strongly compressed hyaline
publication b y B a b i n g t o n (in Abstr. Linn. Trans, p. 3 2 ) in his
Corda ( 1 8 4 0 ) . Berkeley ( 1 8 4 1 ) referred to a
pseudoparenchymatous cells with walls to nearly 1 um thick,
description o f Sphaeria
sometimes apparently m e r g i n g with compressed h o s t tissue at
b e e n traced, but if the n a m e was validly published there it
the sides o f the ascomata, in the ostiolar region
might predate Corda's h o m o n y m . T h e only effect would be a
forming
swollen and pigmented clavate periphysis-like elements with
cells to
1 8 x 4 um, m e r g i n g
into true hyaline
periphyses
within (Fig. 1). Centrum formed from a thin layer o f v e r y thinwalled pseudoparenchymatous tissue lining the l o w e r part o f
the wall. Paraphyses
inconspicuous, formed
(sometimes in
pairs) from + g l o b o s e cells and with at least the l o w e r septa
often constricted, to 8 um diam., gradually tapering, v e r y thinwalled, about as l o n g as the asci. Asci 6 7 - 9 5 x 1 6 - 2 2 - 5 um,
clavate, short- t o long-stalked (occasionally t o ca 4 0 um),
swelling
significantly just before
ascospore release,
thin-
walled at all stages, the apex obtuse to rounded, with a small
sometimes inconspicuous apical ring ca 4 um diam. and 0.5 um
thick, usually 8-spored but ascospores frequently
Ascospores
arranged
irregularly,
aborting.
( 9 - 5 - ) 1 2 - 5 - 1 6 x 5 - 6 - 5 um,
usually + cylindrical with round ends, s o m e t i m e s cylindricellipsoidal or ovoid, often slightly larger a n d / o r
shaped
Key
in asci with
to genera
irregularly
fewer than 8 a s c o s p o r e s ; thin-
and
rhytismoides.
This publication has n o t
change o f author citation.
Specimens examined: Austria: Tirol, Innsbruck, Hafelkar, 2 4 0 0 m, on
Dryas octopetala, July 1940, F. Petrak (WSP 3 1 5 6 3 ; IMI 22034, IMI
2 3 6 4 9 4 as Petrak, Reliquiae Petrakianae
no. 238). Canada: North
West Territories, Baffin Island, Inugsuin Fiord, Geog. Survey Site, on
dry lower slopes below cliffs 1 mile S of Inugsuin site, 6 9 ° 3 7 ' N,
70° 0 2 ' W, on Dryas integrifolia,
29 July 1967, /. A. Parmelee &
f. R. Seaborn (UPS, ex D A O M 117822); North West Territories,
Ellesmere Island, Franklin District, Hazen Camp: in small area on S
slope, 2 miles W N W of camp, 8 1 ° 4 9 ' N, 7 1 ° 2 1 ' W, 1200 m, on
Dryas integrifolia, 3 July 1962, D. B . O. Savile 4 5 3 4 (UPS, ex D A O M
92320); North West Territories, district of Franklin, Somerset Island,
Aston Bay, 73° 39'N, 94° 4 5 ' W, on Dryas integrifolia,
1 0 Aug.
1958, D. B . O. Savile 3767 (UPS, ex D A O M 6 0 0 7 5 ) ; North West
Territories, Mackenzie District, DEW Line site, pin 1, Clinton Point,
6 9 ° 3 5 ' N , 120° 4 4 ' W , on Dryas
integrifolia,
8 Aug. 1963,
/. A. Parmelee 3145a (IMI 137877, ex D A O M 115769); North West
Territories, Ellesmere Island, Hazen Camp, 81° 4 9 ' N, 71° 2 1 ' W, on
included
1. Stromata poorly developed; probably primarily saprobic
.
.
.
.
.
.
1 . Stromata well-developed; biotrophic for at least a large part of the life cycle
.
.
2. Stromata brightly coloured
.
.
.
.
.
.
.
.
.
.
2 ' . Stromata black .
.
.
.
.
.
.
.
.
.
.
.
.
3. Ascomatal walls composed of thin-walled compressed cells; ostioles±simple
.
.
3 . Ascomatal walls composed of large rather thick-walled cells; ostiole surrounded
from the periphyses
.
.
.
.
.
.
.
.
.
.
.
.
.
.
'Plectosphaera'
.
2
.
.
.
.
.
.Polystigma
.
.
.
.
.
.
.
3
.
.
.
.
.
Phyllachora
by clavate pigmented cells derived
.
.
.
.
.
.
Isothea
.
.
.
.
.
.
.
.
.
Figs 1 - 4 . Figs 1 - 2 . Isothea
rhytismoides.
Fig. 1 . Vertical section through ostiolar region, with clavate pigmented cells derived from
periphyses (arrow) (IMI 3 4 6 0 9 1 ; bar = 10 um). Fig. 2 . Ascus and ascospores (IMI 2 9 5 5 8 5 ; bar = 10 um). Fig. 3. Phyllachora
Asci and ascospores (IMI 3 6 7 5 6 3 ; bar = 10 um). Fig. 4 . Polysiigma
rubrum.
physocarpi.
Asci and ascospores. (Capron s.n., ex K; bar = 10 um).
is a small further contribution to understanding of the genus,
Dry as integrifolia, 16 July 1 9 6 2 , D. B. O. Savile 4 6 4 2 (IMI 1 0 5 1 6 9 , ex
D A O M 9 2 3 1 5 ) . Faeroe Islands: Oestero, in monte Kodlen prope
but all species studied in detail show unusual features.
Ejde, 300 m, on Dryns octopetaia, 17 Aug. 1895, H. G. Simmons
438Fung (UPS). Greenland: Ivigtut, on Dryas integrifolia, 22 June
P h y l l a c h o r a a c a e n a e Henn., Ofvers, K. Vet.-Akad.
Fork
1883, A. Berlin (UPS). Iceland: Sudur-Pingeyjarsysla, Mjvatnssveit,
Stockholm 1 9 0 0 ( 2 ) : 3 2 5 , 1 9 0 0 .
ca 500 m S of Haganes (at western shore of Lake Myvatn), on Dryas
octopetaia, 2 8 July 1 9 7 1 , K. Holm & L. Holm 4 3 c - 7 1 (UPS); Hof in
Catacauma
acaenae
(Henn.) Theiss. & Syd., Annls
Mycol.
Horgardal, on Dryas octopetaia, 2 4 June 1902, O. Davidsson (UPS); N
12: 3 8 7 , 1 9 1 5 .
Iceland, Husavik, on Dryas octopetaia, 2 5 June 1903, O. Paulsen (UPS).
Anamorph:
not known.
Italy: Sud-Tirol, Radein, E of Auer in Etschtal, 1600 m, on Dryas
Infected areas not differentiated. Blackened regions 3 5 0 — 1 0 0 0
octopetaia, 29 Oct. 1989, A. Nograsek (IMI 356695). Norway: SorTrondelag, Opdal h:d, Knudsho, on Dryas octopetaia, 28 July 1899,
pm diam., irregular in shape, very strongly raising the
A. T. Fredriksson (UPS); Sor-Trondelag, Opdal h:d, Kongsvoll on
substrate surface, 2 - to 10-loculate, the individual ascomata
Dryas octopetaia, 2 0 Aug. 1 8 8 4 , O.Juel (UPS); Opland, Lorn h:d,
easily distinguishable from the surface, shining black, appearing
Roisheim, on Dryas octopetaia, 8 July 1894, O.Juel (UPS); Svalbard,
slightly roughened due to the leaf epidermal architecture,
Bellsound, on Dryas octopetaia, 3 0 June 1883, A. G. Nathorst (UPS);
ostioles usually conspicuous. Ascomata
shallowly conical, to
Svalbard, Tempelbay, on Dryas octopetaia, 17 July 1882, A. G. Nathorst
2 5 0 pm diam. Upper wall composed of leaf epidermal tissue
(UPS); Svalbard, s. I o c , on Dryas octopetaia, 1861, s. coll. (UPS).
infiltrated by strongly melanized textura epidermoidea. Lower
Romania: Hunedoara, Muntii Retezatului, Piule, on Dryas octopetaia,
wall hardly developed. Paraphyses copious when immature but
30 Sep. 1974, G. Negrean (IMI 2 0 0 9 1 0 ; Herbarium
Mycologicum
many deliquescing at maturity, to 2-5 pm diam., thin-walled,
Romanicum no. 2487). Russia: Novaya Zemlya, Sinus Bessimanaja,
on Dryas octopetaia, July 1875, F. R. Kjellman & A. N. Lundstrom
gradually tapering, the apex swollen, about as long as the asci.
(UPS). Sweden: Harjeladen, Tannas s:n, Hammafjaflet, 1000 m, on
Asci 4 2 - 5 9 ( - 6 7 ) X 1 4 - 5 - 1 9 pm, clavate, usually very shortDryas octopetaia, 3 0 July 1933, J. A. Nannfeldt 4802a (UPS); Jamtland,
stalked, very thin-walled at all stages, the apex obtuse to
Are s: n, Snasahogarna, below Taljbilfanen, 750 m, on Dryas octopetaia,
rounded, without discernible apical structures, 8-spored.
27 July 1951, J. A. Nannfeldt 11606b (UPS); Jamtland, A r e s:n,
Ascospores
arranged biseriately, ( 1 2 - ) 1 4 - 5 - 1 6 ( - 1 7 - 5 ) x 7 - 8 - 5
Renfjallet, on Dryas octopetaia, 13 July 1884, M. M. Floderus (UPS);
pm, ellipsoidal, sometimes slightly curved, thin-walled,
Jamtland, Frostviken s:n, Raudek, on Dryas octopetaia, 2 July 1914,
hyaline, aseptate, without a gelatinous sheath.
E. Borner (UPS); Lule Lappmark, Skaitetjakko, on Dryas octopetaia, July
1900, T. Vestergren (UPS, Vestergren, M. sel. exs. no. 4 0 5 ) ; Lule
Typification: Chile: ad Conception, on Acaena ovalifolia, s.d., F. Neger
Lappmark, Koratjokka, on Dryas octopetaia, 1 4 Aug. 1 9 1 2 , T. Lindfors
122
(B - holotype, lost and presumed destroyed).
(UPS); Tome Lappmark, Jukkasjarvi parish, Kartimvare, on Dryas
octopetaia, 11 July 1928, J. A. Nannfeldt 1203b (UPS); Tome Lappmark,
Host species: Acaena ovalifolia, A. ovalifolia var. insulae-exterioris,
Jukkasjarvi s.-n, Nuolja, towards Bjorkliden, on Dryas octopetaia, 6
A. stangii.
July 1927, J. A. Nannfeldt 833a (UPS); T o m e Lappmark, Jukkasjarvi
parish, M t Laktatjakko, NE slope, 900 m, on Dryas octopetaia, 7 July
Distribution: Chile, Tristan da Cunha.
1928, /. A. Nannfeldt 1617b (UPS); Tome Lappmark, Jukkasjarvi s:n,
Nualju, on Dryas octopetaia, 23 Aug. 1946, I. Holm 5 0 / a (UPS);
The type has been destroyed, and none of the specimens
Tome Lappmark, Jukkasjarvi s:n, Nuolja, on Dryas octopetaia, 10 July
examined is suitable as a neotype, as the only Chilean one
1927, R. Olsen (UPS). Switzerland: Graubiinden, prope Arosa, on
lacks mature spores. Further collections around the type
Dryas octopetala,s.d., P. Magnus (IMI 11252; Kryptog. exs. Mus. Palat.
Vindobon, no. 618). United Kingdom: Scotland: Sutherland, Loch
locality should be made in order that the application of the
Eriboll, Kempie, on Dryas octopetaia, 14 M a y 1985, S. M. Francis
name Phyllachora acaenae may be properly fixed, which will
(IMI 2 9 5 5 8 5 ) ; Sutherland, 3 km S of Inchnadamph, on Dryas
allow confirmation or otherwise of the identity of the
octopetaia, 13 Aug. 1990, P. F. Cannon (IMI 3 4 6 0 9 1 ) ; Sutherland,
specimens from Tristan da Cunha. The paraphyses with
Bettyhill, Strathnaver NNR, NC7060, on Dryas octopetaia, 12 May
distinctly swollen tips are unusual for the genus.
1985, P. F. Cannon (IMI 3 0 4 6 8 7 ) ; Sutherland, Loch Eriboll, Ard
Neackie, N C 4 4 5 9 , on Dryas octopetaia, 13 M a y 1985, P. F. Cannon
Other specimens examined: Chile: Masafuera, Q. de las Casas, on
(IMI 304696).
Acaena ovalifolia var. insulae-exterioris, 11 Feb. 1917, C. Skottsberg (S).
Tristan da Cunha: Hottentot Gulch, 6 5 0 m, on Acaena stangii, 23
Mar. 1938, Y. Mejland 1760 (O); Inaccessible Island, above Blenden
Hall, 400 m, on Acaena stangii, 19 Feb. 1938, £. Christopherson 1357
PHYLLACHORA
(O).
P h y l l a c h o r a Nitschke ex Fuckel, fahrb. Nassauischen
Vereins
Naturk. 2 3 - 2 4 : 2 1 6 , 1 8 7 0 , non Phyllachora Nitschke ex
P h y l l a c h o r a d a l i b a r d a e (Peck) Sacc, Sylloge Fungorum
2:
Fuckel, Fungi Rhenani no. 2 0 5 6 , 1 8 6 7 , in sched., nom. cons.
600, 1883.
Phyllachora is a large genus, with nearly 9 0 0 species currently
Dothidea dalibardae Peck, Ann. Rep. NY St. Mus. 27: 1 0 9 ,
recognized. The exclusively biotrophic nature of most of its
1875.
species means that host specificity has played a major role in
Anamorph:
not known.
species definition. Species on Gramineae and Leguminosae
have been the subjects of at least reasonably modern mono­
Infected
graphic treatment (Parbery, 1 9 6 7 , 1 9 7 1 ; Cannon, 1 9 9 1 ) , but
leaf tissue around effete ascomata is apparent.
for species associated with most other host families the most
regions
useful study remains that of Theissen & Sydow ( 1 9 1 5 ) . This
ostioles epigenous, inconspicuous, the surface + flat, shining
areas hardly delimited, though some discoloration of
Blackened
1 8 0 - 3 2 0 pm diam., roughly circular, hologenous, the
black but the pigmentation poorly developed except around
the ostioles until a late stage of development, 1- to 8-locuIate.
Upper and lower walls composed of several layers of rather
irregular mid brown thin-walled textura angularis with cells to
9 p;m diam., more strongly pigmented in the ostiolar region,
covered with more or less unmodified epidermal cells at the
periphery and sometimes between the ostiolar regions of the
ascomata; lateral walls similar in construction but less strongly
pigmented. Periphyses well-developed, to 15 um long, septate,
tapering, merging into the paraphysis layer. Paraphyses welldeveloped, to 4 um diam., thin-walled, rather irregular,
sometimes slightly swollen between the septa (possibly an
effect of secondary invaders), roughly the same length as the
asci. Asci 6 4 - 7 9 x 8-5-9-5 um, narrowly clavate, short-stalked,
thin-walled at all stages, the apex obtuse, with an in­
conspicuous apical ring ca 2-5 um diam. and 0-5 um thick,
8-spored. Ascospores
arranged biseriately, (14-5—)17'5—19
x 3-5-4-5 um, narrowly fusiform, usually slightly curved
(with one + flat face), thin-walled, aseptate, eventually be­
coming very slightly pigmented and verrucose (perhaps a
degradation effect), without a gelatinous sheath.
epidermis, slightly melanized but with little evidence of fungal
tissue. Lower wall to 15 um thick, composed of flattened host
cells pervaded by dark brown thick-walled textura
epidermoidea,
the cells 2—3 um diam. Fertile layer not clearly differentiated
from the lower wall. Conidiophores
usually present, to 35 um
long and ca 2 um diam., cylindrical, hyaline, thin-walled,
formed directly from the strongly contorted cells of the lower
wall.
Conidiogenous
cells
usually
formed
laterally
but
occasionally in small terminal clusters, 18—26 urn long,
2—2-5 um diam., gradually tapering, sometimes
irregular
in
form,
proliferating
slightly
percurrently; periclinal
thickening often obvious, collarettes lacking. Conidia
4—6 x ca
1-5 um, clavate, the base slightly attenuated, hyaline, aseptate,
apparently smooth-walled.
Teleomorph:
genous,
Blackened
sometimes
regions
500—2500 um diam., amphi-
coalescing,
often
irregularly shaped,
eventually strongly raising the substrate surface, containing
1—3 ascomata. Ascomata
conic-oblate, the ostiolar region
distinctly elongate in transverse section. Upper wall very welldeveloped, to 120 um thick in the ostiolar region, decreasing
in thickness gradually towards the edge of the blackened
region, with an outer layer of host cuticle and the outer part
Typification: U.S.A.: New York State, Sullivan Co., Forestburgh, on
Dalibarda repens, Sep. 1 8 7 3 , C. H. Peck (NYS!, holotype of Dothidea
dalibardae).
of the epidermal cells, and a wide inner layer of very strongly
melanized flattened fungal tissue, the lumina completely
occluded except for the very innermost layers, which merge
into periphyses in the ostiolar region. Lower
Host species: Dalibarda repens L.
wall 20—30 um
thick, composed of a number of layers of strongly melanized
Distribution: U.S.A.; only known from New York State.
flattened fungal cells. Periphyses
This is an inconspicuous species, and poorly known. It
well-developed,
apparently immersed in a mucous coating. Paraphyses
hyaline,
copious,
appears rarely to have been collected since Peck's original
to 6 um diam., very thin-walled, gradually tapering, septate,
work, and the ambiguity of its host identity no doubt has contri­
branching not seen, enveloped
buted to its relative obscurity. The species is easily distin­
1 1 6 - 1 4 6 x 2 8 - 3 7 um, clavate-ellipsoidal but often rather
guished from other Phyllachora
irregularly shaped, thin-walled at all stages of development,
taxa on Rosaceae using
ascospore characters. It is the only true species of
Phyllachora
on that host family known from North America.
Giant spores, which are presumably multinucleate, occur
in a mucous coat.
Asci
short-stalked, the apex acute to obtuse, slightly thicker-walled
but without obvious apical structures, 8-spored.
Ascospores
arranged
20—23-5
obliquely
uniseriately
to
biseriately,
occasionally in asci containing fewer than eight ascospores. A
X 1 2 - 1 3 um, ellipsoidal to cylindric-ellipsoidal, rarely slightly
small proportion of asci also contains ascospores which
ovoid, quite thick-walled, hyaline, aseptate, smooth-walled,
develop abnormally, becaoming almost globose rather than
with a thin to thick epispore-like layer (apparently not mucous)
fusiform. Neither of these phenomena is particularly unusual
which quickly degenerates, the spore eventually becoming pale
within the Phyllachoraceae.
brown and slightly rough-walled.
Other specimens examined: U.S.A.: New York State, Essex Co.,
Newcomb, on Dalibarda repens, 2 2 July 1 9 2 2 , H. D. House (NYS);
New York State, Mud Pond, on Dalibarda repens, July, C. H. Peck
(NYS).
Typification: Chile: Andenthal Los Condos, on Kageneckia oblonga,
s.d., F. Neger (B - holotype of Phyllachora negeriana, lost and presumed
destroyed; UPS (F-06052) 102945! - isotype, here designated
lectotype). Chile: Cordillera de Santiago, Salto de Agua, on
Kageneckia oblonga, June 1925, £ Werdermann 1745 (Sydow, Fungi
exotici exsiccati no. 9 3 2 ; B - holotype of Pleuroplacosphaeria negeriana,
lost and presumed destroyed; K!—isotype, here designated lect­
otype).
P h y l l a c h o r a n e g e r i a n a Henn. & Lindau, Hedwigia
1897.
Pleuroplacosphaeria
negeriana
1928 [anamorph].
Syd., Annls
Mycol.
3 6 : 225,
26: 114,
Host species: Kageneckia oblonga Ruiz & Pavon.
Infected areas to 10 mm diam., the included leaf tissue hardly
modified.
Anamorph:
Blackened
regions 2 5 0 - 1 0 0 0 um diam, irregularly
shaped and often coalescing, intraepidermal, hardly raising the
substrate surface, possibly ostiolate through the host stomata,
some of which have much more melanized fungal tissue
associated with them than others. Upper
wall composed of
conspicuously thick host cuticle and the upper part of the
Distribution: Chile.
As Sydow (1928) noted, the ascospores of Phyllachora
negeriana
may eventually become brown and slightly rough-walled,
prompting Sydow's comparison with Sphaerodothis.
It is not
uncommon for Phyllachora ascospores to become brown, but
this appears to be a degeneration of the ascospores rather than
part of the normal maturation process (Cannon, 1989).
Genuine Sphaerodothis
species have ascospores which b e c o m e
T h e anamorph o f Phyllachora
a m o n g s t the Phyllachora
developmental
negeriana
e x t e m e rather
physocarpi).
is s o m e w h a t unusual
group, but this perhaps represents a
branch. Distinctive features
Komarov (Fungi Rossiae Exsiccati no. 290, LE - holotype, not seen; K!
— isotype of Phyllachora
dark b r o w n before release from the ascus.
than a major
Host species: Physocarpus amurensis Maxim., P. opulifolia (L.) Maxim.
evolutionary
include the large size o f the
conidiomata, the v e r y large numbers o f conidiogenous cells
Distribution: Russia (Russian Far East).
This species d o e s n o t h a v e the shining black stromata o f a
which are usually formed laterally from conidiophores, and
typical Phyllachora,
the narrowly clavate rather than filiform conidia. It is n o t
hypogenous
unlike the anamorph o f Phyllachora
breakdown o f the c o v e r i n g cuticle. T h e absence o f paraphyses
pterocarpi,
for example, and
but this m a y b e at least partly due t o its
placement within the leaf and the
apparent
this fungus also has amphigenous ascomata. A l t h o u g h early
is unusual for the genus, but is n o t unparalleled
workers (e.g. T h e i s s e n & S y d o w , 1915) considered ascomatal
1991). Judging from the appearance o f the dried h o s t leaves,
(Cannon,
position within the leaf to b e an important generic character
the fungus
which separated Phyllachora
recent research­
insignificant as the host is deciduous, the b r o w n i n g o f the leaf
that
in the
ers (e.g. Cannon,
from Catacauma,
1991) h a v e
considered
ascomatal
position is m o r e to do with the physical characteristics o f the
has s o m e deleterious effect, but this m a y
vicinity o f the
ascomata coinciding with
be
natural
senescence.
h o s t than the relationships o f the fungus.
T h e ascospores with a distinct epispore layer are also
unusual for the genus, but the systematic significance o f this
structure
Other specimen examined: Russia: Primorskii Krai, Vladivostok, on
Physocarpus opulifolia, Sep. 1986, L. N. Vasil'eva (IMI 367563).
is unclear. This is because collections o f Phyl­
lachoraceae frequently degenerate rapidly in storage if t h e y are
PLECTOSPHAERA
n o t preserved v e r y carefully (Pascoe, 1990) and ascospore
sheaths and epispores m a y frequently break d o w n to a degree
b e y o n d recognition. T h e rough-walled
ascospores o f this
species m a y
appearance
b e a result
of
o f old
epispore
P l e c t o s p h a e r a Theiss., Annls Mycol.
Plectosphaera
a c c o m m o d a t e non- or weakly stromatic Phyllachora
type
species P. bersamae
(Lingelsh.) Theiss. is a
stromatic m e m b e r o f Phyllachora.
P h y l l a c h o r a p h y s o c a r p i Jacz., in Jaczewski, K o m a r o v &
Anamorph:
Infected
Exsiccati n o . 290, 1899, in sched.
0-7-4 m m
diam.,
the
included
leaf
surface
distinctly browned, usually roughly circular, often coalescing.
Blackened
regions
o f Plectosphaera
others
seems to include a rather disparate array o f
probably have
and its relatives,
closer links to the
Glomerella
a g g r e g a t e , or t o the fungi currently placed in the
Hypo-
nectriaceae.
1 5 0 - 3 5 0 um diam., 1 0 - 5 0 per infected area,
h y p o g e n o u s , sometimes coalescing (to 6-loculate), irregular
Plectosphaera
but usually roughly circular, usually n o t s t r o n g l y domed, m a t t
black especially w h e n immature, the ostiole relatively large,
Phomatospora
oblate, o c c u p y i n g the l o w e r epidermis and mesophyll. U p p e r
Physosporella
Mycol.
textura
(Willi Krieg. & R e h m ) A r x
fragariae
Schweiz
&
1 1 ( 1 ) : 2 0 8 , 1954.
Willi Krieg. & Rehm, Annls Mycol.
4:
39, 1906.
wall 15—20 um thick, slightly thicker in the vicinity o f the
ostiole, c o m p o s e d primarily o f s t r o n g l y melanized
fragariae
E. Mull., Beitr. Kryptogfl.
easily visible when wet. In section a s c o m a t a spherical t o
epidermoidea,
typical
V o n A r x & Muller's c o n c e p t
fungi. S o m e h a v e clear affinities to Phyllachora
but
unknown.
areas
to
species.
T h i s c o n c e p t o f the genus is unfortunately untenable, as the
degeneration rather than wall ornamentation.
Tranzschel, Fungi Rossiae
1 4 : 413, 1916.
was used b y v o n A r x & Miiller (1954)
Stromata
fragariae
(Willi Krieg. & R e h m ) Hohn.,
Annls
16: 61, 1918.
absent. Ascomata
immersed in leaf tissue, 120—200 um
the epidermis s t r o n g l y degraded and the cuticle
diam., + g l o b o s e , with a slightly papillate ostiole surrounded
apparently missing. L o w e r wall p o o r l y developed, c o m p o s e d
b y a patch ca 50 um diam. o f almost black thick-walled, often
o f compressed h o s t tissue filled with
strongly
contorted
curved cells to 15 um in length and 4—5 um wide, g i v i n g the
fungal cells, the palisade layer s o m e t i m e s slightly melanized
appearance o f blunt spines. Peridium
but n o t disrupted. Ostiole
outer layers o f pale b r o w n textura globulosa-angularis
papillate
cells. Periphyses
surrounded b y melanized conicwell-developed,
t o 35 um
hyaline, thin-walled, gradually tapering. Paraphyses
long,
absent, at
thin, membranous, with
with cells
up to 12 um diam., and inner layers o f thin-walled hyaline
flattened cells. Paraphyses
n o t seen; perhaps e v a n e s c e n t at
least at maturity. Asci 6 0 - 7 7 x 13-5-16 um, clavate-cylindrical,
maturity. Periphyses
v e r y well-developed, t o 2-5 um
the stalk rather variable in length, thin-walled at all stages, the
rather thick-walled. Asci 6 2 - 7 4 x 1 2 - 1 5 um, clavate, short-
diam.,
a p e x obtuse, with a small apical ring 2-5—3 um diam. and ca
stalked, v e r y thin-walled, the a p e x obtuse, apical structures
0-5 um
not
thick,
8-spored.
Ascospores
arranged
biseriately,
clearly
visible,
8-spored.
slightly curved (fabiform), occasionally slightly ovoid, fairly
slightly papillate, hyaline, aseptate, thin- and smooth-walled,
thin-walled, hyaline, aseptate, without a gelatinous sheath.
apparently without a gelatinous sheath.
Typification: Russia: Russian Far East, 'in saxosis montibus Londoko
Typification: Germany: Konigstein, on dry leaves of Fragaria vesca,
Apr.-May 1 8 9 8 - 1 9 0 0 , W. Krieger (Krieger, Fungi Saxonici no. 1929,
IMI! — isotype of Phomatospora
fragariae).
Burejensium', on Physocarpus
amurensis,
in regione montium
2 0 A u g . - l Sep. 1895,
to
ovoid,
10-5—12(-14)
x 5-5-6-5 um,
ad ripas fl. Kirma, fl. Amur influentis,
ellipsoidal-fusiform
Ascospores
12-5-14 x 4-5-5-5 um, cylindrical with rounded ends, often
occasionally
Host: Fragaria vesca L.
Von Arx & Muller did not specify the source of the material
of this species that they examined, so although the specimen
Distribution: Germany.
from Padova is authentic material, it is not clear whether it is
This may not be a member of the Phyllachoraceae, as von Arx
the holotype. The sample examined is very small, and further
& Muller ( 1 9 5 4 ) implied when they transferred Krieger &
collections are needed in order that a complete examination of
Rehm's name into Plectosphaera.
The apparently saprobic
this species may take place. Von Arx & Muller apparently
nutritional status and the blackened tissue which is confined to
relied upon the opinion of Petrak ( 1 9 2 9 ) as to the identity of
blunt spine-like processes in Phomatospora
Sphaeria
placement in Phyllachora.
fragariae
The genus name
precludes
Physosporella
rosicola;
Pringsheimia
he suggested that it was a synonym of
sepincola
(Fr.) Hohn. ( = Saccothecium
sepincola
Hohn. may need to be resurrected for this species, and
(Fr.) Fr.). Judging from Fuckel's original description, this was
perhaps also other saprobic taxa formerly placed in
Pkcto­
not the case, and his species name may in fact be an earlier and
sphaera
with
priorable synonym of Plectosphaera
such
Glomerella,
as
P. spartii
E. Mull.
Relationships
as well as the Hyponectriaceae, need exploration.
The apparent similarity of the ostiolar processes to the spines
Rhenani
surrounding the ostiole in Acanthorhynchus
been located.
Physalospora
vaccinii Shear (syn.
vaccinii (Shear) Arx & E. Mull.; Hyponectriaceae)
The material examined has degenerated, and some structures
Unusually, Fuckel
in the original publication, and no type material has
Plectosphaera
cingulata,
needs further investigation.
rosae.
did not cite type material from his exsiccatum series Fungi
rosae
appears very similar to
Glomerella
but appears to have at least primarily biotrophic
nutrition. Glomerella
species often have a period of biotrophic
were difficult to intepret. Modern collections are needed. As
nutrition immediately following infection of the host, but
the various examples of the exsiccatum set were gathered at
become necrotrophs well before ascoma formation. The links
different times (to judge from the label), the specimen
between generic placement and mode of nutrition need further
examined may not technically constitute isotype material.
study, especially because of the considerable significance of
Glomerella
P l e c t o s p h a e r a r o s a e Arx & E. Mull., Beitr. Kryptogfl.
11 ( 1 ) : 2 0 6 , 1 9 5 4 .
ISphaeria rosicola Fuckel, Jahrb.
2 3 - 2 4 : 114, 1870.
IPhysalospora
1882.
Stromata
Nassauischen
Vereins
rosicola (Fuckel) Sacc, Sylloge Fungorum
absent. Ascomata
aggregated, usually
Schw.
POLYSTIGMA
Naturk.
1: 4 3 5 ,
300—400 pm diam., strongly
+ circular when viewed from above,
with a poorly-defined outer ring of blackened tissue, a central
black dot where the ostiole penetrates the plant epidermis, and
the intervening tissue appearing hardly differentiated from the
surrounding substratum; inserted beneath the epidermal layer,
oblate to shallowly conical. Upper wall composed of brown
thick-walled textura angularis
with cells to 9 pm diam. around
the ostiole and at the edge of the ascoma, merging into largercelled textura globuiosa
to plant pathologists.
with cells to 1 7 pm diam. in the middle
P o l y s t i g m a Pers. ex D C , in Lamarck & de Candolle, Flore
Frangaise 6 : 1 6 4 , 1 8 1 5 . Lectotype: Polystigma
rubrum
(Pers.) DC. (Clements & Shear, 1 9 3 1 ) .
Polystigma Pers., in Mougeot & Nestler, Stirpes
Cryptogamae
Vogeso-Romanae
nos 270—1, 1 8 1 2 , in sched. Nom. inval.
(Art. 3 2 ) .
Polystigmella
}acz. & Natalyina, in Natal'ina, Mater.
Mikol.
Fitopatol. Leningrad 8 : 1 6 3 , 1 9 2 9 . Holotype: P. ussuriensis
Jacz. & Natalyina.
Polystigmina Sacc, Sylloge Fungorum 3: 6 2 2 , 1 8 8 4 [anamorph].
Holotype: Polystigmina
rubra (Pers.) Sacc.
Rhodoseptoria
Naumov, Bull, trimest. Soc. mycol. Fr. 29: 2 7 9 ,
1 9 1 3 [anamorph]. Holotype: Rhodoseptoria
ussuriensis
Naumov.
portion, and in the lower wall. Ostiole well-developed, not
Polystigma
papillate, lined with hyaline periphyses to ca 1 0 pm long and
of systematic mycology, but this contribution appears to be
2 pm diam. Paraphyses
has been recognized as a genus since the early days
well-developed, to 4 pm diam., very
the first attempt at a formal monograph. The genus is
thin-walled, gradually tapering, ? deliquescing at maturity.
characterized principally by its brightly coloured stromata,
Asci 8 4 - 9 4 x 1 1 - 5 - 1 4 pm, cylindric-clavate, very thin-walled
which occur on living leaves of Prunus spp. in the Euro-Asiatic
at all stages, + short-stalked but tending to break down near
region. The genus is probably a close relative of
the base making measurement difficult, the apex rounded, ?
as there are few distinguishing characteristics apart from the
Phyllachora,
with a small apical ring 2 - 3 pm diam. and ca 0-5 pm thick,
stromatal pigmentation. A possible further delimiting character
8-spored.
is that Polystigma
Ascospores
arranged
+
biseriately,
17'5-21-5
anamorphs have conidiogenous cells which
X 6-5—7-5 pm, fusiform-ellipsoidal, often slightly curved (fabi-
at least mostly proliferate sympodially rather than percur­
form), thin-walled, hyaline, aseptate, mucous sheath not seen.
rently, as is usual in the Phyllachoraceae. Observation is
Typification: Italy: Treviso, Cansiglio, on dead stems of Rosa canina,
Aug. 1898, s. coll. (D. Saccardo, Mycotheca Italica no. 287, as
Physalospora rosicola (Fuckel) Sacc.; PAD! — syntype of Plectosphaera
rosae).
distinction does not appear to be absolute.
Host species: Rosa canina. .
and overwinter on the ground after leaf fall. Ascomata are
difficult due to the small size of the conidial structures, and the
Polystigma
species are temperate in distribution, and their
host leaves are deciduous. In most species therefore conidial
stromata are formed on living leaves throughout the summer,
formed within the stromata during early spring, and ascospores
Distribution: Italy; perhaps also from Germany (see below).
are released to infect emerging leaves. Polystigma
is one of the
few
genera
o f Phyllachoraceae which
appears
to
cause
species b e l o n g s to y e t a further
assemblage. Link ( 1 8 3 3 )
e c o n o m i c damage to its host plants, causing a m o n g o t h e r
accepted a greatly expanded c o n c e p t o f Polystigma,
s y m p t o m s premature leaf fall (e.g. Banihashemi, 1 9 9 0 ; Kranz,
rather nebulous boundaries. His w o r k on the genus appears
1962).
n o t to have been accepted either b y his contemporaries o r b y
The
deleterious
effect
apparently
has
not
been
with
subsequent authors.
quantified.
Muller ( 1 9 8 6 ) n o t e d that at least in s o m e instances the
stromata o f Polysiigma
species accumulate starch, presumably
as e n e r g y reserves for ascomatal formation, in c o m m o n with
species o f Diachora.
This feature m a y well serve as a further
distinguishing characteristic from Phyllachora,
staining in iodine o f Polysiigma
but the blue
stromata does not occur
Article 5 9 o f the International
Code of Botanical
rears its ugly head again for Polystigma.
Nomenclature
T h e names currently
used for the anamorph and teleomorph o f the t y p e species P.
rubrum
are linked t o the same basionym, Sphaeria
There
seems
little point
in
w h e t h e r the name Polystigma
expending
rubra Pers.
energy
deciding
should refer t o the teleomorph
reliably. It is possible that significant starch reserves also build
or the anamorph when there is ample evidence linking the t w o
up within Phyllachora
morphs. A p p l y i n g the rules strictly would probably necessitate
stromata, but the blueing might be
masked b y melanin deposits. Starch accumulation is likely t o
the use o f the p o o r l y - k n o w n
be correlated with the deciduous nature o f the host leaves o f
teleomorph. T h e n a m e Polystigma
Polysiigma
duced b y P e r s o o n on an exsiccatum label in 1 8 1 2 , but n o
and Diachora
species. T h e coloration o f the stromata
has in the past resulted in Polysiigma
ordinal level from Phyllachora
being separated
at
and its relatives (e.g. Chadefaud,
1 9 6 0 ) , but the almost complete lack o f melanin pigments does
description
o f the genus
name Polystigmella
was
for
the
was apparently first intro­
provided.
T h e name
was
validated three years later b y de Candolle (in Lamarck & de
Candolle, 1 8 1 5 ) .
not in itself preclude placement in the Phyllachoraceae.
Polysiigma
is a small genus, apparently restricted to
Species such as P. astragali
to Stigmatula
Prunus.
on Leguminosae are here referred
(Sacc.) S y d . & P. S y d . (Cannon, 1 9 9 1 ) ; t h e y differ
in anamorph characteristics and stromatal characters as well as
in h o s t preference. A number o f other species has been
referred to the genus in the past which probably do n o t
b e l o n g ; for example Polysiigma
Hyponeclria
adenostomatis
(referred
in this publication), and P. montserratis
believed to be a species o f Phyllachora
to
which is
with reduced melanin
pigmentation and which causes death o f host cells immediately
surrounding
the
stromata
(Cannon,
probably does not b e l o n g in Polysiigma,
1991).
P.
sonneratiae
though its affinities
are n o t y e t fully established (Hyde & Cannon, 1 9 9 2 ) , and P.
pusillum
again does n o t b e l o n g to the genus sensu stricto
but
P o l y s t i g m a a m y g d a l i n u m P. F. Cannon, sp. n o v .
Septoria
rubra |3 amygdali
(Fig. 7)
Desm., Ann. Sci. Nat., B o t . ser. 2, 1 9 :
343, 1843.
Polystigmina rubra var. amygdali (Desm.) S a c c , Sylloge
3 : 6 2 2 , 1 8 8 4 , as
"amygdalinum".
Polystigma
rubrum
var. amygdali
Rehm, Annls. Mycol.
Fungorum
4 : 70,
1 9 0 6 , nom. nud.
Slromata primo ochracea deinde atropurpurascentia, crassa; conidio­
mata sparsa; cellulae conidiogenae
1 2 - 2 2 x 2—2-5 um, laterales vel
terminales, conidia sympodialia efficientia; conidia 22—34 x 0-75-1 um,
filiformia, sigmoidea, ad apicem angustata; ascomata numerosa,
hologena, leniter papillata; paraphyses sparsae, tenuitunicatae; asci
74—97 x 1 8 - 2 3 um, clavati, brevistipitati, tenuitunicati, cum apparatu
apicali ca 4 um diam., octospori; ascosporae
(13-)14—16-5 x 5-5—
waits for relatives to be identified (Cannon, 1 9 9 1 ) . A c o m p l e x
6-5 um, cylindricae, ad centrum minutus constrictae, aspetatae,
of species on leafy and phyllodinous Australasian
hyalinae, Iaeves, sina vagina gelatinosa.
Key
1.
I'.
Acacia
to species of P o l y s t i g m a
Anamorph present .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
2
Teleomorph present
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
6
2. Stromata yellowish, at least when young
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
3
2'. Stromata reddish brown at all stages .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
5
3. Stromata at first pale yellow, becoming orange; conidia 5 5 - 1 2 0 um long; on Prunus cornuta, Himalayas .
.Polystigmina
pallescens
3'. Conidia shorter
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
4
4. Stromata becoming yellowish brown, causing inrolling of leaves but not significant distortion; conidiomata sparse; conidia
2 2 - 3 4 um long; on Prunus dulcis, widely distributed
.
.
.
.
.
.
.
.
.
Polystigma
amygdalinum
4'. Stromata initially pale yellow, becoming orange-red, causing significant distortion of young shoots; conidiomata abundant;
conidia 38—53 um long; on Prunus armeniaca,
China
.
.
.
.
.
.
.
.
.
.
Polystigma
deformans
5. Conidia 2 2 - 4 2 um long; on Prunus domestica and P. spinosa, widespread
.
.
.
.
.
Polystigma rubrum subsp. rubrum
5'. Conidia 3 8 - 6 3 um long; on various Prunus spp., eastern Russia .
.
.
.
.
.
Polystigma rubrum subsp. ussuriense
6. Stromata yellowish, mature in autumn or spring without a preceding anamorphic stage; ascospores 13-5-15-5 x 5-5-7 um; on
Prunus padus, widespread .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Polystigma
fulvum
6'. Stromata reddish or brown to black, at least when mature; anamorphic stage always present
.
.
.
.
.
.
7
7. Stromata brown to purplish black when mature, thick, causing inrolling of the leaf; ascospores 13-16-5 x 5-5-6-5 um, + cylindrical;
on Prunus dulcis, widespread
Polystigma
amygdalinum
7'. Stromata usually reddish brown to black .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
8
8. Stromata reddish brown, causing deformation of shoots and leaves; ascospores 1 2 - 1 5 x 5 - 6 um, ovoid-cylindrical to
ellipsoidal; on Prunus armeniaca,
China
.
.
.
.
.
.
.
.
.
.
.
.
Polystigma
deformans
8'. Stromata dark reddish brown to black, causing some hypertrophy but not inrolling .
.
.
.
.
.
.
.
9
9. Stromata almost black when mature, the ostioles inconspicuous; on Prunus domestica and P. spinosa, widespread
Polystigma rubrum subsp. rubrum
9'. Stromata reddish brown when mature, with black pigmentation around the ostioles; on various Prunus spp., eastern Russia
Polystigma rubrum subsp. ussuriense
Figs 5 - 8 . Figs 5 - 6 . Polystigma
7. Polystigma
amygdalinum.
fulvum
(IMI 163055). Fig. 5. Paraphyses; bar = 10 um. Fig. 6 . Asci and ascospores; bar = 10 um. Fig.
Asci and ascospores (IMI 3 4 3 4 1 5 ; bar = 10 um). Fig. 8. Polystigma
rubrum.
Periphyses. (Capron s.n., ex K;
bar = 10 um).
Stromata-.
developing through late spring and summer
on
Conidiomaial
wall p o o r l y differentiated, c o m p o s e d o f a narrow
living leaves, producing conidia in summer and autumn, and
layer o f small hyaline thick-walled textura
ascospores from fallen overwintered
which the conidiogenous cells are directly derived.
leaves the
following
genous
spring.
Conidial
stromata
to 4 0 x 15 m m diam., irregularly shaped,
angularis,
from
Conidio­
cells arising as lateral or terminal branches from short
-
relatively undifferentiated conidiophores, 12—22 x 2—2 5 um,
at first yellow-brown to orange-brown but soon darkening in
almost cylindrical but usually gradually tapering from
the central portion, h o l o g e n o u s , crinkling the leaf but n o t
base, the upper portion (up to 5 um) often slightly irregular
causing noticeable h y p e r t r o p h y ;
due to successive conidial scars, proliferating
composed
of two
layers o f epidermal tissue and strongly tanned
outer
parenchyma
cells, and a hyaline inner layer.
Conidiomata
sparse, v e r y inconspicuous, 80—120 pm diam.,
+ g l o b o s e , the ostiolar region slightly papillate, h y p o g e n o u s .
Conidia
the
sympodially.
2 2 - 3 4 x O ' 7 5 - l pm, the widest point usually almost
at the base, usually tapering s m o o t h l y towards the sigmoidally
curved filiform a p e x ; hyaline, aseptate, apparently
walled.
smooth-
extensive, s o m e t i m e s c o v e r i n g large portions
Prunus dulcis, 1 Jan. 1993, A. Sztejnberg (IMI 3 5 8 7 9 7 ) ; same locality
o f the leaf, apparently causing leaf curl, purplish black (dark
and host, 19 Feb. 1993, s. coll. (IMI 3 5 8 8 1 6 ) ; Lower Galilee, Gazit, on
Ascostromata
red w h e n fresh; Ghazanfari & Banihashemi, 1 9 7 6 ) , 1 - 5 - 2 m m
thick. Ascomata
v e r y numerous, shallowly domed, h o l o g e n o u s ,
the ostioles epigenous, inconspicuous, v e r y slightly papillate.
Paraphyses
rather sparse, c o m p o s e d o f chains o f v e r y thin-
walled textura globulosa,
at
the
base,
branched at the base, t o 1 8 pm wide
tapering
towards
the
acute
apex.
Asci
74—97 x 18—23 pm, clavate, short-stalked, v e r y thin-walled at
Prunus dulcis, 1 Jan. 1993, A. Sztejnberg (IMI 358796);
same locality
and host, 19 Feb. 1993, s. coll. (IMI 3 5 8 8 1 7 ) ; Golan Heights, Kefar
Haruv, on Prunus dulcis, 1 Jan. 1993, A. Sztejnberg (IMI 358798).
Italy: dintorni di Bologna, on Amygdalus
communis, estate 1889,
G. £ Mattel (IMI 16905; Briosi & Cavara, Funghi Parassiti delle Piante
Coltivate
ed Utili no. 145). M o r o c c o : El Hadjeb, on
Amygdalus
communis, 15 Sep. 1937, R. Delecluse (IMI 16908, UPS;
Mycotheca
generalis; exs. Rev. Mycol. no. 19). Portugal: Azores, unlocalized, on
all stages, the a p e x obtuse t o truncate, with an apical ring ca
almond, comm. Nov. 1924, Bensanda (IMI 16908). Romania: reg.
4 pm diam. and 1 pm deep, usually 8-spored but occasionally
Dobrogea-Ostrov,
4-spored.
Herbarium Mycologicum
Ascospores
arranged
biseriately,
(13—)14-16'5
x 5-5—6-5 pm, + cylindrical, o n e end rounded and the other
1957, M. Olangiu
(IMI 1 5 0 5 7 9 ; Savulescu,
Romanicum no. 1765). Turkey. Ankara, on
Amygdalus webbii, 22 Oct. 1944, collector indecipherable (WSP 30529).
obtuse t o acute, usually slightly constricted in the central
region, quite thin-walled, s m o o t h , hyaline, aseptate, without a
gelatinous sheath. A s c o s p o r e s in s o m e o n l y o f the 4-spored
P o l y s t i g m a d e f o r m a n s Syd., Annls Mycol
asci are larger (16—18 x 6-5—7 pm), and similarly shaped.
Stromata
3 4 : 417, 1936.
Typification: Israel: Kibbutz Givat Oz, on Prunus dulcis, 28 Feb. 1992,
o c c u p y i n g almost the w h o l e o f the leaf, the surface pale ochre
t o 3 0 m m diam., restricted to e m e r g i n g leaves and
apparently arresting d e v e l o p m e n t o f y o u n g shoots, s o m e t i m e s
A. tin (IMI 353415, holotype of Polystigma
amygdalinum).
to reddish b r o w n (dried material), often causing the leaf t o
incurl, to ca 1 m m thick, inner tissues hyaline to pale yellow,
Host species: Prunus dulcis (syn. P. amygdalus, Amygdalus communis, A.
webbii), Amygdalus nana, A. ledebouriana.
Distribution: Afghanistan (Gattani, 1962 as P. rubrum), Cyprus,
Greece (Alexopoulos, 1940 as P. rubrum), India, Iran, Israel, Italy,
Kazakhstan (Byzova & Vasyagina, 1 9 8 1 ; Byzova, 1983 as P. rubrum),
Lebanon (Khatib et al, 1969 as P. ochraceum), Libya (Kranz, 1962 as
P. rubrum), Morocco, Portugal, Romania, Spain (Gonzales Fragoso,
1927), Tadzhikistan (Stepanov, 1966 as P. rubrum), Turkey, Ukraine.
This species is a serious p a t h o g e n o f almonds in the M i d d l e
c o m p o s e d o f hypertrophied leaf cells filled with crystalline
material interspersed with ramifying hyphae.
Anamorph:
conidiomata
1 5 0 - 2 5 0 pm diam., + spherical t o
flattened, the wall p o o r l y developed, difficult t o distinguish
from the stromatal tissue. C o n i d i o g e n o u s cells arranged o v e r
+ the w h o l e o f the inner surface, 1 5 - 2 1 X 2-5—3-5 pm, rather
irregular, + cylindrical, s o m e t i m e s swollen at the base, often
curved,
apparently
proliferating
clinally. Conidia 38-53
at least s o m e t i m e s
x ca 1 pm,
peri-
+ cylindrical, the basal
East, often causing premature defoliation (e.g. Kranz, 1 9 6 2 ) . It
portion usually slightly swollen, fusiform, often sigmoidally
has been referred to in the past b o t h as Polystigma
curved, hyaline, aseptate.
(a s y n o n y m o f P. fuluum)
and P. rubrum,
ochraceum
but is distinct from
Teleomorph:
ascomata
immersed within
the stroma,
the
these species in stromatal m o r p h o l o g y and ascospore shape.
ostioles epigenous, visible from the surface o n l y as slightly
T h e development and a s c o m a formation o f P. amygdalinum
sunken dark b r o w n t o black spots ca 25 pm diam., + spherical,
P. ochraceum)
was
studied
by
Ghazanfari
(as
& Banihashemi
the wall fairly thin, + hyaline, difficult t o distinguish from the
( 1 9 7 6 ) . T h e y found that mature ascomata w e r e formed m o s t
stromatal
plentifully in leaves collected from affected trees in O c t o b e r ,
parenchymatous cells, the ostioles periphysate.
and their formation was dependent on a sufficiently
low
tissue, c o m p o s e d
of very
thin-walled
tapering gradually, v e r y thin-walled. A s c i cylindric-clavate,
v e r y thin-walled at all o b s e r v e d stages, without
incubation temperature.
T h e fungus appears to b e susceptible to various fungicides,
pseudo-
Paraphyses
apical
structures.
Ascospores
not
seen
developed
[ 1 2 - 1 5 X 5 - 6 pm,
although these are o n l y efficaceous w h e n applied at petal fall,
ovoid-cylindrical to ellipsoidal, narrower at the base, s o m e ­
w h e n the ascospores are being transmitted from overwintered
times slightly curved; S y d o w , 1 9 3 6 ] .
stromata t o y o u n g leaves (Banihashemi, 1 9 9 0 ) . C o n t r o l can
also b e effected b y burning o f fallen leaves in the autumn and
winter.
This t a x o n is described as a n e w species due t o
difficulties
Septoria
Typification:
China: Beijing, Prince park, on Prunus armeniaca,
15
Sept. 1935, Y. C. Wang (B - holotype, lost and presumed destroyed);
o f typifying
rubra [3 amygdali,
unequivocally D e s m a z i e r e s '
the
same locality and host, 1932, Y. C. Wang 233 (B, lost and presumed
destroyed; HMAS 13282!), anamorphic stage cited by Sydow.
name
and confusion o v e r the application
Host species: Prunus armeniaca, P. davidiana.
o f Article 5 9 .
Other specimens examined: Cyprus: unlocalized, on almond, comm. 19
Nov. 1930, H. M. Morris (IMI 16910); Agros, on Prunus amygdalus,
Sept. 1937, R.M.Nattrass
Prunus amygdalus,
869 (IMI 16898). India: Shalimar, on
12 Aug. 1987, V. K. Razdan (IMI 321486). Iran:
40 km SE of Shiraz, Maharlou, on Prunus dulcis, 28 M a y 1993, Z.
Distribution: China.
The
material
immature
examined
contains
ascomata. Its identity,
only
conidiomata
therefore,
could n o t
and
be
unequivocally established, n o r could its relationships. T h e
Banihashemi (IMI 358821). Israel: Kibbutz Gazit, on Prunus dulcis, 20
species as interpreted
Mar. 1992, A. Lin (IMI 3 5 3 4 1 6 ) ; near Tel Aviv, Ben Sherman, 6
amygdalinum,
Aug. ????, I. Reichert (IMI 16897); Givat Oz, on Prunus dulcis, 4
ostioles and the v e r y l o n g conidia serve as
Dec. 1992, A. Sztejnberg (IMI 3 5 5 4 5 6 ) ; Givat Oz, Emek Yizre'el, on
features.
but
the
here might b e close t o
blackish pigmentation
Polystigma
around
the
distinguishing
The
restriction
of
stromata
to
young
leaves
seems
t o the immature stromata, but conspicuously thickened (ca
diagnostic, a feature n o t e d in the original description, and
1 mm), at first bulging around d e v e l o p i n g
their at least almost c o m p l e t e maturation while still attached
eventually b e c o m i n g m o r e uniformly thickened, at first reddish
to the tree is also n o t e w o r t h y .
o r a n g e but eventually b e c o m i n g dark orange-brown, the leaf
cuticle eventually
Other specimens examined: China: prov. Shansi austr., Yiian-ch'ii distr.,
ad pag. Wu-li-Tsun, ca 900 m, on Prunus davidiana,
H.Smith
12 July 1924,
(Plantae sinenses no. 8 2 9 8 ; UPS); Hebei Province, Ping-
shang county, on Prunus armeniaca, M a y 1990, S. M. Tian (IMI);
Liaoning, Xingcheng, on Prunus armeniaca, 12 July 1955,
Guang-zheng
Jiang 6036 (HMAS 13334); Shandong, Muping, on Prunus armeniaca,
peeling
away,
with
ascomata
30—200
but
ascomata,
visible as black patches 80—100 um diam. on the
adaxial
surface, and sunken ostioles on the abaxial one. O u t e r layer o f
stroma 30—40 um thick at the abaxial surface,
somewhat
thinner at the adaxial one, dark orange-brown,
composed
primarily o f strong tanned leaf tissue. Inner layer at first white
28 May 1930, T. N. liou 6 6 6 9 (HMAS 161); Beijing, Fangshan, on
but b e c o m i n g pale brown, c o m p o s e d almost entirely o f fungal
Prunus armeniaca,
tissue.
26 June 1935, Fong-yan Huang
(HMAS 1615);
Liaoning, on Prunus armeniaca, 19 M a y 1950, T. N. Liou 2 5 9 (HMAS
Conidiomata
1614).
Ascomata
absent.
roughly spherical but s o m e t i m e s slightly flattened,
the walls n o t well differentiated from stromatal tissue, the
ostioles n o t conspicuous, at first clearly sunken but eventually
P o l y s t i g m a f u l v u m D C , in Lamarck & de Candolle,
Francaise
6: 1 6 4 , 1 8 1 5 .
Flore
(Figs 5 - 6 )
Dothidea
fulva
(DC.) Fr., Systema
Hypocrea
fulva
(DC.) D e Not., Erb. Critt. Ital. n o . 1 4 7 3 , 1 8 6 5 ,
in
Mycologicum
2: 5 5 4 , 1 8 2 3 .
fulvum
Pers.,
Cryptogamae
in
Mougeot
Vogeso-Romanae
& Nestler,
no. 2 7 1 , 1 8 1 2 , in
inflated b e t w e e n the septa, e x t r e m e l y thin-walled. Periphyses
2 0 um long, 3—4 um wide at the base, gradually
98-128
x 1 4 - 5 - 1 6 ( - I 8 - 5 ) um,
nar­
sched.
thin-walled at all stages, with a subapical ring 3—3-5 um diam.
and ca 1 um thick, 8-spored. Ascospores
1 3 - 5 - 1 5 - 5 x 5-5-&5(—
arranged biseriately,
7) um, cylindric-ellipsoidal t o o b o v o i d ,
thin- and smooth-walled, hyaline, aseptate, often
guttulate,
without a gelatinous sheath.
des
Typification: Finland: Lapponia: Komenji ad Sodankyla, on Padus, 5
n o . 3 2 2 , 1 8 0 1 ; in sched. N o m . inval. (Art.
Sept. 1802, s. coll. (UPS; ex herb Wahlenberg, authentic and probably
|3
Fichtelgebirg's
the apex. Asci
to
tapering
Stirpes
Polystigma fulvum (Fr.) Tomilin, Botanicheskii
Materialy
16:
137, 1 9 6 3 . Based on Dothidea fulva (DC.) Fr. Norn, inval.,
Art. 3 3 . 2 ; full reference t o b a s i o n y m n o t cited, and n o w
unnecessary under the m o d e r n nomenclatural rules.
rubrum
to 1 2 um
r o w l y clavate, long-stalked (to ca 50 um) the a p e x obtuse,
Norn, inval. (Art. 3 2 ) .
Xyloma
Paraphyses
diam., tapering gradually t o an acute tip, septate, s o m e t i m e s
towards
sched.
Polystigma
b e c o m i n g v e r y slightly protuberant.
padi
Funck,
Cryptog.
Gewachse
type material of Sphaeria ochracea Wahlenb., here designated neotype) ;
32).
Lapponia, Komenji, host not specified, s.d., Wahlenberg (K; authentic
Xyloma
aurantiacum
Schleich., Plantae
Cryptogamae
Helveticae
no. 4 8 7 , 1 8 0 7 . N o m . inval. (Art. 3 2 ) .
Sphaeria
ochracea
Dothidea
Polysiigma
Prunus padus, Sept. [year unknown], s. coll. (K; Funck, Cryptog.
Wahlenb., Flora Lapponica:
ochracea
dinaviae
(Wahlenb.) Fr., Summa
518, 1812.
Vegetabilium
Gewachse des Fichtelgebirg's no. 322, authentic material o f Xyloma
Scan-
rubrum (3 padi Funck). Germany: unlocalized, host not specified, s.d.,
s. coll. (K; Schmidt & Kunze, Deutschlands Schwamme
2: 3 8 7 , 1 8 4 9 .
ochraceum
material of Sphaeria ochracea Wahlenb.). Germany: unlocalized, on
(Wahlenb.) S a c c , Nuovo
Giorn.
Bot. Ital.
8: 1 8 1 , 1 8 7 6 .
no. 1, authentic
material of Sphaeria padi Schmidt & Kunze).
Host species: Prunus padus L. (syn. Cerasus padus, Padus
racemosa),
Sphaeria padi J . C. Schmidt & Kunze, in Holl, Schmidt &
Kunze, Deutschland
Schwamme:
1, 1 8 1 5 , in sched. N o m .
inval. (Art. 3 2 ) .
Prunus asiatica (syn. Padus asialica), Prunus maximowiczii.
Sphaeria
Distribution: Austria, Belgium [?], China (unverified record cited by
Eriksson & Yue, 1988), Czechoslovakia, Finland, France, Germany,
Hungary, India, Italy, Japan (Anon., 1965, unverified), Latvia, Norway,
Romania, Russia, Sweden, Switzerland, Ukraine, United Kingdom.
xantha
Fr., Observationes
Mycologici
1: 1 7 2 , 1 8 1 5 ,
nom. illeg. (Art. 6 3 ) .
Polystigma
aurantiacum
Pers. e x Fr., Systema
5 5 4 , 1 8 2 3 , nom. inval.
Mycologicum
2:
(Art. 3 4 ) ; apparently erroneously
attributed in s y n o n y m y t o Persoon, Mycologia
Europaea
t.
5, f . l , 1 8 2 2 .
ILibertella
in
Immature
fulva
Prunus grayana (e.g. Anon., 1965) have not been verified.
S t r o m a t a are formed on living leaves in the spring and
summer, a s c o m a t a developing during the summer and autumn.
Thiim., Mycotheca
Universalis
no. 1 1 9 5 , 1 8 7 8 ,
sched.
stromata
Records on
A n anamorph is apparently n o t produced. Immature asci are
present in material collected in S e p t e m b e r ( I M I 1 2 7 6 0 , 3 0 1 8 4 ) ,
(in summer and autumn) without a delimited
and apparently mature o n e s in an autumn Italian collection (de
infected area, at first hardly causing h y p e r t r o p h y o f the leaf,
Notaris, Erb. Crittog.
t o ca 1 0 m m diam., roughly circular, h o l o g e n o u s , the adaxial
p r o b a b l y delayed until the following spring, at least in m o s t
Ital.
n o . 1 4 7 3 ) . A s c o s p o r e release is
surface y e l l o w t o pale brown, quickly b e c o m i n g
orange-
regions. Relatively few collections h a v e b e e n seen which bear
b r o w n and rather thicker due to the development o f fungal
the mature teleomorph. T h e described variation o f features
tissue, the adaxial surface sometimes appearing
such as ascus and ascospore size and shape m a y n o t therefore
minutely
punctate (perhaps due t o leaf glands), and the abaxial surface
strongly punctate due t o the development o f sunken ostioles.
Mature
stromata
(in late autumn and spring) similar in size
adequately reflect their true e x t e n t .
Sphaeria
ochracea
W a h l e n b . appears t o h a v e b e e n described
as a c o m p o s i t e o f Polystigma
fulvum
and P. rubrum.
It is here
neotypified as a s y n o n y m o f P. fulvum,
in accordance with
(K;
Mougeot
&
Nestler,
Stirp.
Crypt.
Voges.
no.
271).
long-established practice; Fries ( 1 8 1 5 ) appears t o have b e e n
Czechoslovakia: am Sattler, in montibus sudeticis, host not specified,
the first to implicitly restrict application o f the name t o the
s.d., T. Bail (K; Rabenhorst, Herb. Mycol. edn 2, no. 579) [mature asci].
fungus on Prunus padus.
T h e heterogeneity of Wahlenberg's
species c o n c e p t was apparently o n e reason w h y Fries adopted
the n a m e Sphaeria
xantha
(here interpreted as superfluous and
thus illegitimate) instead o f W a h l e n b e r g ' s name, but h e also
s e e m e d t o consider the first-published n a m e
inappropriate
bearing in mind the coloration o f the fungus.
fulvum
that n a m e is therefore sanctioned
and takes precedence o v e r the earlier n a m e Sphaeria
fulva,
22 July 1959, E. R. Julin (UPS); Ostrobotnia borealis: Ii, Maakrunni,
on Prunus padus, 6 Aug. 1933, /. /. Liro & H. Roivanen (IMI 60898,
ochracea
Jalonvaara, Tupinmylly, on Prunus padus, 13 Aug. 1937, ] . I. Liro &
H. Roivanen (UPS; Liro, Mycotheca
Fennica no. 4 7 7 ) ; Ostrobotnia
australis: Maalahti, Storskar, on Prunus padus, July 1962, L. Roivanen
in
& H. Roivanen (UPS); Ks Posio Pemu, riverside grove by Kurttajoki
giving a figure in P e r s o o n
(2 km upstream of Koivukongas falls), on Prunus padus, 23 Aug. 1964,
W a h l e n b . Fries cited the n a m e Polystigma
s y n o n y m y with Dothidea
locality and host, 27 Sep. 1886, W. Krieger (K; Krieger, Fungi Saxonici
no. 229b) [mature asci]. Finland: Ob, Sdeni, Ajos, on Prunus padus,
UPS; Liro, Mycotheca Fennica no. 870); Karelia ladogensis: Suistamo,
Fries ( 1 8 2 3 ) adopted de Candolle's n a m e Polystigma
and transferred it to Dothidea;
Czechoslovakia [?]: Sachsen, Erzgebirge, bei Bockau, on Prunus padus,
28 July 1 8 8 5 , W. Krieger (K; Krieger, Fungi Saxonici no. 229a); same
aurantiacum
( 1 8 2 2 ) as a source. A l t h o u g h the fungus depicted appears t o
T. Ulvinen (UPS; ex herb Univ. Oulu); Regio Aboensis: Inio,
be Polystigma
the name is apparently never referred t o
Lokholm, on Prunus padus, 22 July 1932, L. E. Kari (IMI 16896).
in P e r s o o n ( 1 8 2 2 ) , and is n o t included in the index t o that
France: Jura, on Prunus padus, s.d., Morthier 8 8 9 (UPS; ex herb Fuckel
fulvum,
w o r k (Rogers, R o g e r s & Seeler, 1 9 4 2 ) . T h e epithet presumably
originated from Schleicher's nom. nud. Sphaeria
aurantiaca
above). Material has n o t b e e n seen o f Libertella
fulva
(see
Thiirn.
Saccardo ( 1 8 8 4 ) stated that the exsiccatum he saw contained
asci, and placed the n a m e into s y n o n y m y with
fulvum.
Polystigma
If the t a x o n is conidial as T h t i m e n intimated b y his
choice o f name, s y n o n y m y with P. rubrum
is m o r e likely.
ussuriense,
and found
ochraceum)
that
stromata on Padus asiatica,
than that o f Polystigma
P. fulvum
form
rubrum
subsp. ussuriense.
Prunus
dulcis
(syn. Amygdalus
X P. ussuriensis,
communis),
but little
fruticosa,
or P.
Limin referred t o an anamorphic stage forming on
asiatica,
Dresden, host not specified, s.d., Rabenhorst
(K; ex herb Cooke);
Badens. no. 6 5 ) ; Gottingen, host not specified, 1835, Wissmann (K);
ca. Ostrich, host not specified, s.d., s. coll.. (K; Fuckel, Fungi Rhenani no.
Yellowish-
Cerasus
Prunus americana
Rabenhorst, Fungi Europaei no. 1036); Reichenhall, on Prunus padus,
Aug. 1869, A. Braun (K; Rabenhorst, Fungi Europaei no. 1358);
2 6 6 4 , originally identified as Polystigma rubrum) [mature ascomata];
development o f the fungus occurred after inoculation o n t o
Prunus domestica
edn 2, no. 6 9 ) ;
mature
though d e v e l o p m e n t was slower
o r a n g e stromata were also formed on Padus maackii,
s.d., Lasch (K; Rabenhorst, Herb. Mycol.
Franconia, Eschenau, on Prunus padus, autumn 1864, P. Reinsch (K;
subsp.
and P. rubrum
would
padus,
unlocalized, on Prunus padus, s.d., s. coll. (K; Knieff & Hartm., PI. Crypt.
Limin ( 1 9 8 6 ) investigated the h o s t range o f this species
(referred to as Polystigma
1894; ex herb Barbey-Boissier); Jura, on Prunus padus, s.d., Morthier
(K; Fuckel, Fungi Rhenani no. 1826). Germany: Driesen, on Prunus
nigra.
Eisleben, on Prunus domestica,
1875, /. Kunze (K; ex herb Kunze)
[mature ascomata; the host is probably misidentified]. Hungary:
Comit. Szepes: Iglofiired, on Prunus padus, Sept. 1905, F. Filarsky (IMI
30184, UPS, WSP 1 8 1 5 1 ; Petrak, Mycothecageneralis
no. 284). India:
Kanawar, Punang Forest, 8500', on Prunus padus, 8 Oct. 1890,
A. Barclay (K). Italy: Val Anzasca, Macugnaga, on Cerasus sp. ('un
ciliegio'), autumn 1865, de Notaris (K; Erbar. Critlogam, Hal. no. 1473)
Padus
[mature teleomorph]; Sudtyrol, Predazzo, on Prunus padus, 1 8 8 5 - 6 ,
but n o n e was found on the o n l y collection seen on
Arnold (K; Rehm, Ascomyceten no. 872). Latvia: Zemgale: distr.
that host from eastern Russia ( I M I 3 6 7 5 6 1 ) . T h i s is o f stromata
Dobele, Telrvete, in silvam ad cementario, on Prunus racemosa, 27
on senescing leaves collected in September, and
contains
July 1956, E. Vimba (UPS; Fungi Latvici no. 6). Norway: Troms:
mature asci. It is possible that there is a further eastern Asiatic
Nordreisa, Bilto, ca 200m, on Prunus padus, 6 Aug. 1960, L. Roivanen
form o f that species which has an anamorph, so m o r e research
& H. Roivanen (UPS; ex H); Vesteralen: Hadseloy, Ongstad, vid
is needed.
stigen till Storvand, ca 100m, on Prunus padus, 21 Aug. 1966,
T h e collection seen from Prunus maximowiczii
is
immature,
containing
only
asci
in
early
(LE 1 4 6 5 8 2 )
stages
of
development. O t h e r features are consistent with identification
as Polystigma
fulvum.
ochraceum
Porsmyr, on Prunus padus, 17 Aug. 1975, F.-E. Eckblad (UPS: ex herb
Univ. Bergen); Gtta. [?], on Prunus padus, 28 July 1947, F. J.
Newhook
(IMI 16833); Kongsvoll, near Oppdal, on Prunus padus, Aug. 1985,
S. M. Francis (IMI 314478). Romania: Moldova: distr. Neamt, Piatra
A widely distributed collection on Pyrus communis
as Polystigma
H. Smith 4 8 2 6 : 2 (UPS); Hordaland: Kvam hd., Karpedammen ved
var. aurantiacum
labelled
Neamt, pe muntele Cozla, on Prunus padus, 3 0 July 1954, T. Savulescu
W e s t e n d , , the n a m e
(IMI 2 8 9 3 4 ; Herbarium Mycologicum
Pers. e x Fr. (Herbier
Siberia, Krasnoyarsk, Minusinsk, on Prunus padus, s.d., Martianoff
Barbey-Boissier no. 8 9 0 ; e x herb Muller A r g . 1 8 9 6 , K!, U P S ! ) ,
(UPS; ex herb Thumen); European Russia, Novgorod, ad fluv.
is actually Venturia pyrina
Tigoda, on Prunus padus, 1894, V. Komarov (K; Jaczewski, Komarov
perhaps based on Polystigma
aurantiacum
Aderh.
&
Tranzschel,
Fungi
Rossiae
Romanicum no. 1301). Russia:
Exsiccati
no.
82); Khabarovsk,
Other specimens examined: Austria: Carinthia, pr. Greifenburg, on
Bolshekhekhzirski Reservation, on Padus asiatica,
Prunus padus, Sept. ????, F. von Hdhnel (IMI 1 2 6 7 0 ; Crypt. Exs. ed.
L. N. Vasil'eva (IMI 3 6 7 5 6 1 ) ; Dal'nevostochnyi Krai, Vladivostokskii
16 Sep. 1982,
Mus. Hist. Nat. Vindob. no. 3 2 7 2 ) ; same locality and host, Sept. 1901,
okrug, Poluostov Murav'eva Amurskogo, on Prunus maximowiczii, 19
von Hdhnel (IMI 2 2 2 2 2 ) ; Niederosterreich, Purkersdorf, on Prunus
Aug. 1929, V. TransheV (LE 146582). Sweden: Are, on Prunus padus,
padus, July 1940, F. Petrak (IMI 3 3 0 0 6 6 ; Reliquiae Petrakianae
23 July 1950, G. M. Waterhouse (IMI 4 2 3 3 4 ) ; unlocalized, host not
no.
716); Austria inferior, Krems, on Prunus padus, June 1871, Thumen
specified, s.d., s. coll. (K; Fries, Scleromyci Sueciae no. 2 4 1 , as Sphaeria
(K; Thumen, Fungi Austriaci no. 178); Carinthia, Bleiburg, on Prunus
xantha);
padus, s.d., Kristof (K, W S P ; Flora Exsiccata Austro-Hungarica
L. Romell (K; Romell, Fungi Exsiccati Praesertim Scandinavici no. 77);
no.
ad Jonsered Vestrog, on Prunus padus,
28 Aug. 1889,
1577); Austria inf., Senftonberg, host not specified, s.d., s. coll. (K; ex
Hosjo, DLr Vika, Karlslund, on Prunus padus,
herb Thumen). Belgium [?]: unlocalized, on Prunus padus, s.d., s. coll.,
R. Morander 3 2 1 5 (UPS); Gastrikland: Hille parish, Iggon, near the
5 Aug. 1977,
village on the central part of the island, on Prunus padus, 3 0 July
W. Gardiner (K); Scotland: near Aberdeen, host not specified, 1869,
1972, R. Santesson 24173 (UPS); Smaland: Nobbele s:n, Orraryd,
s. coll. (K; Cooke, Fungi Britannici Exsiccati no. 4 6 4 ) ; unlocalized, host
host not specified, 1881, C.J.Johanson
not specified, s.d., s. coll. (IMI 16899, K; Cooke, Fungi Britannici
(UPS); Smaland: Granna,
Gyllene, Uttern, on Prunus padus, 14 July 1945, K. V. O. Dahlgren
Exsiccati edn 2, no. 578).
(UPS); Smaland: Vrigstad, on Prunus padus, July 1887, R. Tolf (UPS);
VG Skopde, host not specified, Aug. 1880, J . Carlson (UPS); VG
Henneberg: Barsled, on Prunus padus, 23 July 1889, A G. Eliasson
(UPS); Mossekeys [?], host not specified, 13 Aug. 1869, J. £. Zetterstedt
(UPS); V G Finnerodja, on Prunus padus, s.d., G. Neander (UPS); ad
Tjufordei in paroecia Astra Karup, Hallendil, on Prunus padus, 11
Aug. 1927, A. G. Eliasson (UPS); Halland: Getinge, host not specified,
s.d., s. coll. (UPS, ex herb Fries); Varmland: Langserud par., Norra
Botten, on
Prunus padus,
2 9 July 1889, H. Dahlstedt
Sodermanland: Soderstelje, Hall, on Prunus padus,
(UPS);
July 1895,
T. Vestergren (UPS); Sdm Allhelgona s:n, Bullersta, on Prunus padus,
Aug. 1892 G. V. Schotte (UPS); Sodermanland: Strangnas parish,
Hospitaler, on Prunus padus, 12 Aug. 1931, ] . A. Nannfeldt
P o l y s t i g m i n a p a l l e s c e n s Petr., in Petrak & Ahmad,
8: 1 8 3 , 1 9 5 4 .
Stromata
Sydowia
formed on living leaves in the spring and summer,
conidiomata developing
during the summer
A s c o m a t a formed in mature
and
autumn.
stromata; immature
asci are
present in material collected late in the season. A s c o s p o r e
release is p r o b a b l y delayed until the following spring.
Stromata
without a delimited infected area, at first hardly
causing h y p e r t r o p h y o f the leaf, t o ca 1 7 m m diam., r o u g h l y
4288
circular, s o m e t i m e s coalescing, h o l o g e n o u s , pale y e l l o w t o
(UPS); Uppland: Hanas par., Fageln, on Prunus padus, 2 Sept. 1975,
pale brown, quickly b e c o m i n g bright o r a n g e t o o r a n g e - b r o w n
S. Ryman 3629 (UPS); Uppland: Fuubo S:n, Marichued [?] gard, on
and rather thicker due t o the d e v e l o p m e n t o f fungal tissue, the
Prunus padus, 8 Oct. 1928, N. Hyalnder (UPS); Uppland: Vendels s:n,
Tomsarbo, on Prunus padus, 19 Sept. 1970, K. Holm & I . Holm (UPS);
Uppland: Uppsala, Slattobacken, on Prunus padus, 2 0 Sept. 1892,
A. G. Eliasson (UPS); Uppland; Alvkarleby par., Billudden Peninsula,
Billskaten, on Prunus padus, 3 0 Aug. 1974, N. Lundqvist 9466 (UPS);
Uppland: Stockholm, Nacka, on Prunus padus, 2 Aug. 1882, /. Eriksson
(UPS; Eriksson, Fungi parasitici
Scandinavian
no. 198); Uppland:
adaxial surface with minute b r o w n spots to 5 0 um
where
the conidiomatal ostioles e m e r g e and
the
diam.
abaxial
surface with ill-defined g r e y i s h - b r o w n b l o t c h e s t o 2 5 0 um
diam. w h e r e the ascomata are developing, with darker sunken
ostiolar regions t o 7 5 um diam.
Anamorph:
Conidiogenous
cells arising from a p o o r l y defined
Elfkarelby, Tanunsberg, on Prunus padus, 3 1 July 1886, E. Henning
layer o f small-celled hyaline textura
(UPS); Uppsala, on Prunus padus, 2 0 Sept. 1884, A. Nilsson (UPS);
fascicles but
Uppland: Uppsala (Bondyrka) at the river Hagaan ca 1 km S of
veloped;
Norby, on Prunus padus, 2 6 Aug. 1927, /. A. Nannfeldt 15790 (UPS);
acute, proliferation unclear. Conidia
Holmiae,
Sept. 1882,
filiform (ca 0-5 um wide), v e r y flexible, often with a v e r y
(UPS); Uppland: Alsihe I, Kungshann, on Prunus
inconspicuous g l o b o s e k n o b at the base, hyaline, apparently
ad
Bellevue, host
K. ] . Haeggblom
not
specified,
23
padus, 18 Sept. 1898, R. Sernand (UPS); Uppland: Bondkyrka s:n,
Fabordarna, on Prunus padus, 20 Sept. 1928, ] . A. Nannfeldt
e v e n short
angularis,
conidiophores
2 4 - 3 2 x 2 - 5 - 3 um,
almost
s o m e t i m e s in
almost n e v e r
cylindrical, the
55—120 um in
de­
apex
length,
s m o o t h and aseptate.
3762
(UPS); Dalarna: Husby parish, N of Flinsberget, on Prunus padus, 1
Sept. 1973, L. Junell 2179 (UPS); Dalarna: Husby par., E of Lovnas
( = 6 km SE of Husby) at the edge of the wood, on Prunus padus, 1
Sept. 1973, N. Lundqvist 8714 (UPS); Gastrikland: Hamrange s:n,
Axmar bruk, Overhammar, host not specified, 19 Aug. 1942,
G. Lohammar (UPS); Gastrikland: Valbo s:n, Turuvik, Garrmarkuna
Typification:
Pakistan: Kagan Valley, on Prunus cornuta, 2 6 July
1951, S. Ahmad 4427 (GZU - holotype of Polystigmina pallescens, not
seen).
Host species: Cerasus sp., Prunus cornuta.
W om jainvagstationen, on Prunus padus, 29 July 1 9 3 8 , / . A. Nannfeldt
5591 (UPS), Gastrikland: Hille S:n, Galgrund, on Prunus padus, 10
Aug. 1972, S. Ahlner (UPS); Dalama: Transtrand, on Prunus padus,
Sept. 1911, N. Sylven (UPS; Lagerberg & Sylven, Skogens skadesv. no.
23); Medelpad: Torp s:n, Torpshammar, on Prunus padus, 9 Aug.
1933, /. A. Nannfeldt 4 9 2 2 (UPS); Angermanland: Ytterlannas socken,
Distribution: India, Nepal, Pakistan.
This species w a s described as an anamorph, but is clearly
refrable t o Polystigma,
and s o m e specimens e x a m i n e d (e.g. I M I
7 1 4 3 6 ) h a v e immature asci. T h e coloration o f the
young
stromata suggests affinities with Polystigma
fulvum,
Angermanland: Vitryggeia s:n, Skuleberget, O-siden, on Prunus
species does n o t produce
T h e conidia o f
padus, 14 July 1932, G. B. E. Hasselberg(UPS);
Polystigmina
Sel, on Prunus padus,
13 July 1970, N. Lundqvist
6 7 2 0 (UPS);
Jamtland: ad cataractam
Rista in paroecia Undersaker, on Prunus padus,
12 Aug. 1925,
A G. Eliasson (UPS); Jamtland: Froso, Valla, strandsnar vid Storsjostranden, on Prunus padus,
22 Sept. 1923, R. Sernander
Jamtland: Are, on Prunus padus, 7 Sept. 1885, C.f.fohanson
pallescens
an anamorph.
but that
are significantly l o n g e r than t h o s e o f
other k n o w n species o f Polystigma.
T h e h o l o t y p e has n o t b e e n
seen, but several o f A h m a d ' s collections from similar localities
(UPS);
are in I M I , and Petrak's description is comprehensive. T h e r e is
(UPS);
little doubt as t o the identity o f the species.
Norrbotten: Nedertornea, Laivandden i Vuono, on Prunus padus, 6
Sept. 1942, £. R. fulin (UPS); Norrbotten: Karl Gustav par., Kukkola,
Other specimens examined: India: unlocalized, on Prunus cornuta, 8
on Prunus padus,
July 1973, R. D. Ram 2 (IMI 185885); Garwhal Himalaya, on Betula
11 Aug. 1960, E. R. fulin (UPS). Switzerland:
Graubfinden: Klosters, Novai, on Prunus padus,
17 July 1970,
alnoides (probably actually Prunus sp.), Aug.-Sept. 1983, R. D. Gaur
E. Muller (IMI 163055a, WSP 89027) [mature asci]; Bemer Oberland:
R-76 (IMI 2 8 0 9 5 2 ) ; same data, R. D. Gaur R-80 (IMI 280956).
Kiental, on Prunus padus, 15 Aug. 1926, W. Ryby (IMI 22223, ex hort.
Nepal: below Mugu, Mugu Khola, 3 5 0 0 m, on Cerasus sp., 2 4 Aug.
bot. Bemensis); Zurich, host not specified, s.d., Muller (K; ex herb
1952, O. Polunin, W. R. Sykes & L. H. ] . Williams 3 0 1 9 (K). Pakistan:
Shuttleworth); in montibus prope Neuchatel, on Cerasus padus, Aug.
Shogran, on Prunus cornuta, 27 July 1956, S. Ahmad
1869; P.Morthier
81154); same locality and host, 25 July 1956, S. Ahmad 1 4 1 4 8 (IMI
(K). United Kingdom: England: near Ilkley,
1 4 1 4 9 (IMI
Middleton, on Prunus padus, Sep. 1893, H. T. Soppitt (K; ex herb
81153); Swat, Kalam, on Prunus cornuta, 2 1 Aug. 1952, S. Ahmad
Crossland); Scotland: banks of the Clunie, on Prunus padus, July 1895,
(IMI 2 3 6 6 1 5 ; Reliquiae Petrakianae no. 3 6 0 ) ; Kamal Ban Forest, on
Prunus cornuta, 20 Oct. 1 9 5 4 , A. H. Khan (IMI 7 1 4 3 6 , ex herb.
Pakistan Forest Institute no. 4 0 0 ; immature asci).
2 8 - 4 2 x 0 - 5 - 0 - 7 5 pm, the l o w e r part (to approximately the
mid point) v e r y narrowly lanceolate to fusiform, the upper
part filiform
P o l y s t i g m a r u b r u m (Pers.) D C , in Lamarck & de Candolle,
Flore Francaise 6: 1 6 4 , 1 8 1 5 .
(Fig. 8 )
Pers., Observationes
rubrum
Cryptogamae
Mycologiae
Pers., in M o u g e o t
Vogeso-Romanae
2: 101, 1799.
& Nestler,
Stirpes
n o . 2 7 0 , 1 8 1 2 , nom.
inval.
rubra
(Pers.) Fr., Observationes
Mycologicae
1: 172,
Teleomorphic
stromata
1—5 m m diam., irregularly shaped but
hologenous,
reddish
brown
to
black, s o m e t i m e s
faintly
verrucose (a feature o f the leaf epidermal architecture), the
ostioles sometimes inconspicuous but appearing as small black
pigmented outer layers 20—40 pm thick and a hyaline inner
layer containing the ascomata.
1815.
Dothidea
the
dots on paler stromata, often s o m e w h a t sunken; c o m p o s e d o f
(Art. 3 2 ) .
Sphaeria
curved,
the leaf, flat o r slightly c o n c a v e on the abaxial surface,
rubrum
Polystigma
wide), sigmoidally
usually r o u g h l y circular, s t r o n g l y raising the adaxial surface o f
subsp. r u b r u m
Xyloma
{ca 0-25 um
b a s e + truncate, hyaline, aseptate, apparently smooth-walled.
rubra (Pers.) Fr., Systema
Mycologicum
2: 553, 1823.
Septoria rubra (Pers.) Desm., Ann. Sci. Nat., B o t . ser. 2 , 1 9 : 3 4 3 ,
1 8 4 3 [anamorph].
Ascomata
+ spherical. Paraphyses
gradually
tapering
towards
rather sparse, t o 7 pm diam.,
the
apex,
s t r o n g l y inflated b e t w e e n the septa.- Asci
very
thin-walled,
94—118 X 10-5—
1 2 pm, narrowly clavate, v e r y long-stalked (to ca 6 0 pm),
Libertella
rubra
Mykologie
Polystigmina
(Pers.) Bonord.,
Handbuch
der
Allgemeine
1 : 5 5 , 1 8 5 1 [anamorph].
ring 2—3 pm diam. and ca 0-5 um thick, 8-spored.
rubra (Pers.) S a c c , Sylloge Fungorum
3 : 622, 1884
[anamorph].
Polystigma rubrum forma pruni-spinosae S a c c , Mycotheca
n o . 6 3 7 , 1 8 7 6 ; in sched. N o m . inval. (Art. 3 2 ) .
Veneta
Polystigma
rubrum forma pruni-domesikae
Sacc,
Mycotheca
Veneta n o . 6 3 8 , 1 8 7 6 ; in sched. N o m . inval. (Art. 3 2 ) .
rubra forma ramipetiolicola
S a c c , Annls Mycol.
1;
2 6 , 1 9 0 3 [anamorph].
Stromata:
developing throughout late spring and summer on
living leaves, producing conidia in summer and autumn, and
ascospores from fallen overwintered
leaves the
following
spring; the conidiomata s o m e t i m e s persisting in an apparently
viable condition until the ascomata are mature.
Conidial
stromata
t o 10(—35) m m diam., irregularly shaped
but usually roughly circular, yellowish-brown in v e r y y o u n g
lesions but
becoming
arranged
Ascospores
biseriately, 1 0 - 5 - 1 4 x 3-5—4-5(—5-5) pm, cylindric-
ellipsoidal, occasionally o b o v o i d , occasionally slightly curved
Sphaeria hyetospilus Mart., Flore Cryptogamica
Erlangensis: 4 7 8 ,
1 8 1 7 , nom. illeg., Art. 6 3 . Xyloma
rubrum Pers. and
Sphaeria padi D u v . cited in s y n o n y m y ; apparently a
c o m p o s i t e o f the t w o taxa.
Polystigmina
v e r y thin-walled at all stages, the a p e x obtuse, with an apical
quickly b e c o m i n g
darker
in the
orange
t o reddish
brown,
central region, h o l o g e n o u s ,
the
surrounding leaf tissue hardly affected, containing a large
(fabiform), hyaline, aseptate, thin- and smooth-walled, without
a gelatinous sheath.
Typification: 'Utplurimum automno, in foliis Pruni domeslkae, rarius Pr.
spinosae' (typification statement of Xyloma rubrum). Belgium [?],
unlocalized, 'in foliis Pruni domeslkae et spinosae, autumno', s. coll.
(IMI!, K!; Mougeot & Nestler, Stirp. Crypt. Vogesomm no. 270,
authentic material of Polystigma rubrum Pers., nom inval.). Italy:
Belluno, Cadore, on Prunus spinosa, 187'4, S. Venzo (Kl; Saccardo,
Mycotheca Veneta no. 637, authentic material for Polystigma rubrum
forma pruni-spinosae). Italy: Treviso, Vittorio, on Prunus domestica,
1875, s. coll. (K!; Saccardo, Mycotheca Veneta no. 638, authentic
material for Polystigma rubrum forma pruni-domesticae). Italy: Treviso,
Volpago, Montello, in ramulis tenellis Pruni spinosae, Sept. 1902, s.
coll. (?PAD — holotype of Polystigmina rubra forma ramipetiolicola, not
seen).
Host species:
Prunus domestica
L., Prunus institia L. (syn. Prunus
domestica subsp. institia), Prunus spinosa L. Records on Prunus salicina
(e.g. Anon., 1965, 1972) have not been verified, and may refer to
subsp. ussuriense. The host cited as Malus
baccata (see below) is
probably misidentified.
number o f conidiomata; c o m p o s e d o f an upper layer o f plant
tissue 4 0 - 5 0 pm thick w h o s e
cells are filled with
bright
orange-brown material, an intermediate layer 3 0 0 - 5 0 0 pm
thick o f almost c o m p l e t e l y occluded angular to vertically
elongated fungal cells, and a l o w e r layer similar t o the upper
one.
Anamorph:
conidiomata
150—250 pm diam., roughly spheri­
cal, epigenous o r h o l o g e n o u s , the ostiole epigenous, v e r y
inconspicuous. Conidiomatal wall v e r y p o o r l y developed, n o t
clearly distinguishable from the stromatal tissue.
Conidiogenous
cells developing o v e r the entire inner surface o f the wall,
often laterally from sequential cells o f short conidiophores t o
1 0 um l o n g and ca 2 pm w i d e ; derived from a thin layer o f
textura
angularis
Conidiogenous
with hyaline thin-walled cells 3—5 pm diam.
cells 13—24 x 2—3 pm, at first usually cylindrical
Distribution: Armenia, Austria, Belgium (Grove, 1937), Bulgaria
(Yossifovitch, 1937), Channel Islands, China, Cyprus, Czechoslovakia,
Denmark, Finland, France, Germany, Greece (Pantidou, 1973),
Hungary, India (Bilgrami et al, 1979), Iran, Iraq, Ireland, Italy, Japan
(Anon., 1965), Kazakhstan (Byzova & Vasyagina, 1981), Kirghizia
(Byzova & Vasyagina, 1981, unverified), Korea (Anon, 1972,
unverified), Lebanon (Khatib et al, 1969), Lithuania (Brundza, 1961),
Moldova (Fatina, 1989), Netherlands (Oudemans, 1897), Poland,
Portugal (Sousa de Camara, 1916), Romania, Russia, Serbia, Spain
(Gonzales Fragoso, 1927), Sweden, Switzerland, Tadzhikistan (Stepanov, 1966), Turkey, Ukraine, United Kingdom, Uzbekistan (Byzova
& Vasyagina, 1981, unverified). The records cited by Byzova &
Vasyagina (1981) from countries other than Kazakhstan do not cite
host names; they may thus refer to Polystigma amygdalinum instead.
Their statement that the species occurs in North America has not
been substantiated.
but gradually tapering towards the upper region, which is
slightly irregular in appearance due t o successive conidial
scars;
usually
proliferating
sympodially.
Conidia
(22-)
T h e typification o f Polystigma
rubrum
presents problems if
application o f the n a m e is t o continue with current practice,
not least because the teleomorph and anamorph names h a v e
Sphaeria ochracea Wahlenb.). France: Var, Le Luc, on Prunus spinosa,
the same basionym, as discussed a b o v e . T h e r e are only t w o
s.d., s. coll (K; ex herb Shuttleworth); Puy de Dome, Royat, on Prunus
collections identified as Xyloma
spinosa, Aug. 1890, A. He'ribaud (K); unlocalized, on Prunus spinosa,
rubrum in Persoon's herbarium
in L, neither being definite type material as minimal collection
data are given. L 9 1 0 2 6 3 — 7 6 0 contains t w o portions o f leaves,
containing w h a t m a y b e developing ascomata o f
rubrum.
Polystigma
It is inscribed ' H b . Pers. ded. Letellier' and (in a
different hand) 'Dothidea
condition,
and
its
rubra Fr.'). T h e material is in p o o r
identity
has
not
been
s.d., Fautrey 1627 (UPS); Colmar, on Prunus domestica, 1869, Kaufmann
(UPS); unlocalized, on Prunus spinosa, Fautrey 1068 UPS: unlocalized,
on Prunus sp„ s.d., Ripart (UPS); Germany: Westfalen: Siegen,
Gaiargriind [?], bei Hainiken, on Prunus spinosa, 8 Oct. 1922, A. Ludwig
(IMI 16911, ex herb Ludwig); Westfalen: Siegen, Hainiken, on Prunus
spinosa,
16 July 1922, A. Ludwig (IMI 16912, ex herb Ludwig);
unequivocally
unlocalized, on various Prunus species, s.d., s. coll. (K; Fuckel, Fungi
established. L 9 1 0 2 6 3 — 7 7 9 contains several leaves with well-
Rhenani no. 1003); Berlin, on Prunus domestica, Aug. 1882, P. Sydow
developed stromata, but the h o s t is g i v e n as Prunus padus
two subsequent annotations (in unknown hands)
the collection as Polystigma
fulvum.
and
(K; Sydow, Mycotheca Marchica no. 9 5 8 ) ; Sachsen, Halle, bei Ober-
re-identify
roblingen, on Prunus domestica, Aug. 1877, R. Staritz (UPS; ex herb
Clearly, neither o f these
Staritz); Mittelfranken: Hersbruck, Aschbach, on Prunus
4 July 1948, K. Stares
Polystigma
Ellenbach, on Prunus spinosa, 8 Sep. 1948, K. Stares 9337 (UPS);
rubrum.
Neotypification
with a collection from
another source is n o t carried out here, pending rationalization
(UPS); Mittelfranken: Hersbruck,
bei Dresden, on Prunus domestica, Oct. 1882, P. Magnus (UPS); Haut
Rhin, Websheim, 210m, sur pruniers, 2 0 Aug. 1869, ] : P. Becker
of Art. 5 9 with its implications for the generic name.
From evidence provided b y serial collections in Serbia,
mature ascospores are present in stromata b e t w e e n M a r c h and
M a y . In collections from Prunus domestica,
6997
spinosa,
collections is suitable either as a l e c t o t y p e o r a n e o t y p e o f
conidia tend to b e
(UPS; ex herb Becker). Hungary: Transilvania australis, ad pagam
Giregrace, on Prunus sp., 25 Sep. 1875, F. Michael
20 Feb. 1989, V. Minassian
Prunus domestica,
those from P. spinosa,
locality and host, 14 Mar. 1990, V. Minassian
variation
between specimens, and the effect if it is statistically significant
T h e r e are m a n y literature records o f Polystigma
rubrum
here t o the n e w l y described species Polystigma
Polystigma
Laestadia
rubrum is in fact P.
circumscissa
on
T h e s e are referred
The material distributed in Fuckel's Fungi Rhenani
amygdalinum.
no. 2 6 6 4 as
fulvum.
rubrum
(IMI 3 5 8 8 2 4 ) ; same
(IMI 3 5 8 8 2 5 ) ; same
locality and host, 10 Apr. 1991, V. Minassian (IMI 3 5 8 8 2 6 ) ; Lorestan,
3 5 8 8 2 3 ) ; Lorestan, Hamedan, on Prunus domestica,
V. Minassian
10 May 1991,
(IMI 358827, mature ascospores); same locality and
host, 10 June 1991, V. Minassian (IMI 358828). Iraq: Shaklaurh, on
Prunus Idomestica
A. A. Attrackchi
(misidentified as Morus
sp.), 22 Oct. 1976,
(IMI 2 1 4 6 0 2 ; ex herb. Dep. PI. Prot., Hammam
Al-Alil no. 28). Ireland: Co. Tipperary, shore of Lough Derg, near
Sacc. (Saccardo, 1 9 0 3 ) , was placed into
s y n o n y m y with Polystigma
4 Mar. 1991, V. Minassian
Tooysarkan, on Prunus domestica, 4 May 1991, V. Minassian (IMI
might b e geographical rather than host-related.
almond (Prunus dulcis, syn. P. amygdalinum).
domestica,
(IMI 3 5 8 8 2 2 ) ; Lorestan, Hamedan, on
shorter and often h a v e shorter apical filiform regions than
but there is considerable
(UPS; ex herb
Richter); Iran: Lorestan, Khorram Abad, on Prunus
Gortmore Point, on Prunus spinosa, 10 July 1967, M. P. Scannell (K);
b y v o n A r x & Muller
Co. Mayo, Lough Carra, on Prunus spinosa, 2 0 July 1966, M. Scanned
( 1 9 5 4 ) . N o material has b e e n seen, but the fact that ascomata
(K); Co. Clare, Burren, N of Leamanah Castle, by Lough Aleenaun, on
mature in autumn and the stromata form shot-holes in the h o s t
Prunus spinosa, 12 July 1966, M. ] . P. Scannell; 1 mile W of Sligo
leaves suggests that the affinities o f this species lie elsewhere.
town, 7 July 1962, M. Scannell (K). Italy: Serra-Pamparato, on Prunus
It might b e y e t another s y n o n y m o f Glomerella
domestica,
cingulata,
indeed a c o m p o s i t e o f that species and Polystigma
or
rubrum;
a
1888, Mondovi
(IMI 16900; Briosi & Cavara, I Funghi
Parassiti delle Piante Coltivate ed Utili no. 12); ad Oldenico, presso
collection examined from Serbia ( I M I 3 6 8 4 7 2 ) includes b o t h
Vercelli, on Prunus spinosa, 1858, Malinvemi
species and exhibits apparently similar s y m p t o m s .
Ital. no. 240). Romania: Oltenia, Segarcea, prope pagum Timburesti,
(K; de Notaris, Erb. Critt.
90m, on Prunus domestica, 10 Sep. 1 9 6 3 , 1 . Comes, I. Ene, M. Costescu
Other specimens examined: Armenia: near Erevan, on Prunus domestica,
& I. Terbea (IMI 120527; Flora Oltenia Exsiccata no. 132); Muntenia,
Sep. 1956, K.Abakyan
distr. Ilfov, Buceresti, on Prunus domestica, 22 Oct. 1924, T. Savulescu
& D. Babayan
(IMI 7 5 9 8 4 ; Griby
SSSR
(Moskovskoe Obshchetvo Ispytatelei Prirody) no. 120). Austria:
& C. Sandu (IMI 16902, 16913; Herbarium Mycologicum
Carinthia, prope Bleiburg, on Prunus domestica, s.d., Kristof (UPS; Flora
no. 556). Russia: Chuvashskaya ASSR, Il'inka, Zelenaya Stoyanka,
exsiccata Austro-Hungarica
banks of river Volga, 4 Aug. 1978, V.A.Mel'nik
no. 1981); Mahr.-Weisskirchen, on Prunus
domestica, Aug. 1929, F. Petrak (IMI 30800, WSP 18577; Petrak,
Mycotheca
Generalis no. 700); Austria inf., Dross, on Prunus spinosa,
Romanicum
(IMI 2 4 9 3 4 5 ) ;
Primorskii Krai, environs of Vladivostok, on cultivated plum, 6 Apr.
1993, I . N . Vasil'eva (IMI 3 6 7 5 6 4 ; teleomorph); prov. Kursk, distr.
s.d., s. coll. (K, ex herb Thiimen, Plowright). Channel Islands:
Dmitriewsk., Derjutschino, on Prunus domestica,
Guernsey: Petit Port, on Prunus spinosa, 16 Sep. 1948, M. B. Ellis &
A. Bondarzew (WSP 14193; Bucholtz, Fungi Rossiae exsiccati ser A., no
July—Aug. 1911,
f. P. Ellis (IMI 31818, K); Guernsey, unlocalized, on Prunus spinosa,
92). Serbia: Valjevo, on Prunus domestica, June 1986, M. Arsenijevic
1847, s. coll. (K). China: Shansi austr., Yuan-ch'iii district, Hsiung-shan-
(IMI 368472); Valjevo, Jovanja, on Prunus domestica,
ting, ca 800 m, on ' Malus baccata' [probably actually Prunus sp.], 10
M. Arsenijevic (IMI 3 6 8 4 7 3 ) ; same locality and host, 17 Feb. 1995, S.
Mar. 1994,
July 1924, H. Smith (Plantae sinenses no. 8299, UPS). Cyprus:
Siojanovic
Trikoukhia, on Prunus domestica, comm. 3 Nov. 1929, H. M. Morris
S. Stojanovic (IMI 368475); same locality and host, 8 Apr. 1995,
(IMI 368474); same locality and host, 10 Mar. 1995,
(IMI 16909; very immature, identification doubtful). Czechoslovakia:
S. Stojanovic (IMI 3 6 8 4 7 6 ) ; same locality and host, 8 May 1995,
Bohemia, Teplitz Schonau, on Prunus domestica, July 1901, Eichler
S. Stojanovic (IMI 368477). Sweden: Goteborg, on Prunus spinosa, 17
[?] (IMI 16901); Brno, on Prunus domestica, Sep. 1924, R. Picbauer
Aug. 1895, K. L. Larsen (IMI 2 0 7 7 9 4 ; ex herb LD); Bergiiland,
(IMI 16904; Baudys & Picbauer, Phytopathologicky
unlocalized, host not specified, 5 Aug. 1889, collector
Herbdr Pestovanych
indecipherable
Rostlin no. 30). Denmark: Fyen Skaarup, on Prunus institia, 7 Sep.
(UPS); Skane: Kullen, host not specified, 18 July 1909, H. Du Rietz
1882, C. /. fohanson (UPS); Micus Klink [?], host not specified, 3 Aug.
(UPS); Skane: ad Apelryd, prope Banad, on Prunus spinosa, 14 Sep.
1873, O. Thoen (UPS). Finland: ad Panckoco, on Prunus domestica, 27
1925, A. G. Eliasson (UPS); Skane: Dalby, on Prunus spinosa, 15 Oct.
Aug. 1811, s. coll., (UPS; ex herb Wahlenberg, authentic material of
1890, I . Romell (UPS; Eriksson, Fungi parasitici scandinaviae no. 3 4 5 ) ;
Blek, Karlehamn, on Prunus spinosa, Aug. 1883, C. J. Johanson (UPS);
Oland, ad Farjestaden, on Prunus spinosa, 3 0 Aug. 1914, A. G. Eliasson
(UPS); Oland, Torslunda socken, Arontorp, on Prunus spinosa, 6 Sep.
1963, L.Junell
1622 (UPS); Gotland: Visby, Galybeigu, on Prunus
subsp. u s s u r i e n s e (Jacz. & Natalyina) P. F. Cannon, c o m b , et
stat. n o v .
Polystigmella
ussuriensis Jacz. & Natalyina, in Natal'ina,
Mikol. Fitopatol. Leningrad 8 : 1 6 2 , 1 9 2 9 .
Mater.
spinosa, 14 July 1895, A. G. Eliasson (UPS); Gotland, norr om Visby,
on Prunus spinosa, July 1895, T. Vestergren (UPS); Halland, unlocalized,
host not specified, s.d., s. coll. (UPS; ex herb Fries); Halland, ad
Godhem in paroecia Oustala, on Prunus spinosa,
17 June 1914,
A. G. Eliasson (UPS); Getange, host not specified, s.d., s. coll. (UPS; ex
Polystigma
ussuriense (Jacz. & Natalyina) Prots., Sad i Ogorod
7:
35, 1957.
Rhodoseptoria
ussuriensis N a u m o v , Bull, trimest.
29: 2 7 8 , 1 9 1 3 [anamorph].
Soc. mycol
Fr.
herb Fries); Halland, host not specified, s.d., s. coll. (UPS; ex herb
Differs from subsp. rubrum
Fries); Bohuslan, Ljungs s:n, Aufacterod, on Prunus spinosa, 2 8 July
rather than dark brown, with the ostioles distinctly visible and
1887, A. G. Eliasson (UPS); Bohuslan, Storon ex Gaderoa, on Prunus
spinosa, 30 Aug. 1911, A. G. Eliasson (UPS); Bohuslan, in Lindholen
ad Skafioland, on Prunus spinosa, 2 Aug. 1911, A. G. Eliasson (UPS);
Bohuslan, Spekerod s:n, 0-5 km N O om Rod, on Prunus spinosa, 8
Sep. 1946, P. Karlvall (UPS); Bohuslan, Spekerod s:n, 0-5 km N O om
Rora gard, on Prunus spinosa, 10 Sep. 1947, F. Karlvall 3 2 4 9 (UPS);
Bohuslan, rod prope Hallevikstrand, on Prunus spinosa, 16 Aug. 1882,
O. Nordstedt (UPS); Goteborg, on Prunus spinosa, 17 Aug. 1895,
A. L. Larsen (UPS); Osterg. Taby, Beateborg, host not specified, 11
Aug. 1913, P. A. Issen (UPS); Uppland: Overgran parish, Arno, on
Prunus spinosa, 14 July 1936, S. Junell (UPS). Switzerland: Vaud,
Valleyres, on Prunus spinosa, 15 Sep. 1886, ] . Veller (K; Barbey, Socie'te
Dauphinoise no. 4385 bis). Turkey: Kutahya, on Prunus domestica, 13
Aug. 1968, H. K. Wagnan
2 9 (IMI 135768). United Kingdom:
England: Cornwall, near Perranporth, Lamboume, on Prunus spinosa,
16 Oct. 1926, F. Rilstone (K, LIV); England: Porlock Beach, on Prunus
b y : ascostromata reddish b r o w n
surrounded b y a small melanized r e g i o n ; and the conidia
which are 38—63 um long.
Typification: Russia: Russian Far East: in prov. Austro-Ussuriensi, on
Prunus sp. (a Manchurian
Rhodoseptoria
species), s.d., s. coll.
below). Russia: Russian Far East: Shkotovskogo Raiona, near
Petrovka, on Prunus triflora var. koreana, M a y 1928, s. coll. (holotype
of Polystigmella ussuriensis, not seen; see possible type material cited
below).
Host species: Prunus mandschurica, P. maximowiczii, P. triflora, P. triflora
var. koreana, Prunus ussuriensis.
Distribution: Russia (Dalny Vostok region).
spinosa, 25 M a y 1957, P. K. Austwick (IMI 6 9 4 3 3 ) ; England: Bungay,
Polystigma
host not specified, s.d., Stock
the o n l y representative o f the separate genus
(K); Northamptonshire,
host not
specified, July 1848, s. coll. (K; ex herb Broome); England: North
Wootton, on Prunus spinosa,
s.d., C. B. Plowright
(K;
Sphaeriacei
Britannici no. 15); England: near Ryde, host not specified, Sep. 1843,
s.d., s. coll. (K; ex herb Broome); England: Bungay, host not specified,
Sep. 1865, M. C. Cooke (K; Cooke, Fungi Britannici Exsiccati no. 1 8 2 ) ;
England: Weymouth, near Portisham, 15 Aug. 1890, s. coll. (K; ex
(holotype of
ussuriensis, not seen; see probable type material cited
rubrum subsp. ussuriense was originally described as
Polystigmella.
T h e stated difference b e t w e e n it and Polystigma
Polystigmella
was
that
had ascospores which were septate. T h e fungus
otherwise appears v e r y similar t o Polystigma
rubrum.
The
o b s e r v e d septum within the ascospores was presumably an
observational error for the juxtaposition o f oil droplets, a n o t
herb Grove); England: Brixham, on Prunus spinosa, Aug. 1889,
u n c o m m o n p h e n o m e n o n when spores are examined in certain
W. R. Hughes (K); England: Scarborough, Hayburn Wyke, on Prunus
mounting media. S o m e o f the specimens e x a m i n e d have p o o r
spinosa, 14 Aug. 1892, s. coll. (K; ex herb Grove); England: near
documentation;
Weymouth, on Prunus spinosa, 10 Aug. 1890, s. coll. (K; ex herb
possible t y p e material, but it is n o t easy t o reconcile them with
two
o f the
collections cited
below
are
Grove); England: Cornwall, Saltash, on Prunus spinosa, 8 Sep. 1932,
the collection data g i v e n in the original publications. O n l y
s. coll. (K; ex herb Grove); England: North Wootton, Lex's [?] Wood,
o n e collection, itself rather sparse, s h o w e d mature ascomata.
host not specified, Sep. 1871, C. B. Plowright (K); England: Isleworth,
host not specified, s.d., Murray (K); England: Oxfordshire, Crowell
Hill, on Prunus spinosa, July 1844, s. coll. (K; ex herb Ayres); England:
East Bergholt, host not specified, Sep. 1855, s. coll. (K; ex herb
Currey); England: Weymouth, on Prunus spinosa, 15 Aug. 1890, s.
coll. (K; ex herb Grove); England: Devon, cliffs W of Ilfracombe, on
Polystigma
rubrum
subsp. ussuriense
is p r o b a b l y an eastern
Asian variant o f subsp. rubrum, restricted t o Prunus triflora
and
immediate relatives. It is possible that there is clinal variation
at least in conidium length b e t w e e n the subspecies in northern
Europe and eastern Asia, but suitable collections from localities
Prunus spinosa, Aug. 1930, R. W. G. Dennis (K); England: Ilfracombe,
in central Russia h a v e n o t been seen in order t o test this
Cairn Top, on Prunus spinosa, 3 0 Aug. 1908, R. H. Philip 1438 (K);
hypothesis. T h e collection on Prunus mandschurica
England: Shere, host not specified, s.d., Capron (K; ex herb Cooke)
conidia which are slightly shorter on average than t h o s e from
[mature ascomata]; England: Hampshire, Basingstoke, near Oakley,
collections on Prunus triflora, but there is insufficient evidence
on Prunus spinosa, July 1871, R. S.Hill
(K); Wales: Anglesey, near
Bryn Siencyn, on Prunus spinosa, 19 July 1927, P. G. M. Rhodes 3 0 0 9
(K) Wales: Lampeter, host not specified, 1841, S. Ralfs (K); Wales:
;
Aberystwyth, on Prunus spinosa, 3 Sep. 1905, s. coll. (K; ex herb
Grove); Wales: Abergele, on Prunus spinosa, 2 6 Aug. 1882, s. coll (K;
ex herb Grove); Wales: Tenby, on Prunus spinosa, Aug. 1925,
E. M. Wakefield (IMI 12792); Wales: Aberystwyth, on Prunus spinosa,
to support the erection o f a further
taxon. T h a t
as affected leaves. Subsp. rubrum
also occurs in eastern Asia,
presumably introduced with cultivated plums.
Limin ( 1 9 8 6 ) studied the host specificity o f this subspecies
(under the s y n o n y m Polystigma
ussuriense),
though it is n o t
3 Sep. 1905, s. coll. (K; ex herb Grove); Wales: Glamorgan, Gower,
Mewslade, 17 July 1953, s. coll. 61 (K); Scotland: Dunbartonshire, on
subsp.
rubrum.
He
Prunus spinosa, s.d., s. coll. (K); Scotland: Midlothian, Auchendinny
infections o f Prunus
Woods, s.d., s. coll. (K); unlocalized, host not specified, s.d., s. coll. (IMI
americana,
16977;
calycina
doubtful, immature).
Exsiccati no. 577, identification
sample
contains semi-mummified fruit infected b y the fungus, as well
certain that he w a s working with subsp. ussuriense
Cooke, Fungi Britannici
seen has
P. nigra,
found
that
besseyi,
the
P. domestica
P. salicina,
P. salicina
x P. nigra and P. ussuriensis.
sections Armeniaca
fungus
and Cerasus
rather than
would
cause
x P. ussuriensis,
P.
x P. americana,
P.
H o s t species from Prunus
s h o w e d s o m e evidence o f
parasitism, but the lesions did not develop fully, although
comparatively large yellowish
stromata
Cerasus tomentosa,
and P. cerasifera.
Prunus spinosa
were
pathogenic effect was o b s e r v e d on Spiraea
asiatica,
and none at all on Padus
communis.
formed
Negligible
salicifolia
maackii
on
or
and
Padus
Amygdalus
T h e s e experiments are in general supportive o f
subspecific status for Polystigma
rubrum
subsp.
T h e transfer to Hyponectria
relative to the Phyllachoraceae reassessed. Nevertheless, t h e
much-reduced
ascomatal wall with melanized material re­
stricted to o n l y a small clypeus is n o t typical o f
Phyllachora,
the other obvious placement for the species.
ussuriense.
There is considerable diversity a m o n g Prunus species from
Phyllachora
eastern Asia, and their classification needs reinterpretation. It
? = Auerswaldiella
puccinioides
Mycol. 1 2 : 2 7 8 , 1 9 1 4 :
is not improbable that further Polysiigma
is made with reservations, as
that genus is in n e e d o f revision, and its t a x o n o m i c position
taxa remain t o b e
discovered in this region.
beaumontii
Auerswaldia
C o o k e , Grevillea
puccinioides
1 3 : 63, 1885.
(Speg.) Theiss. & Syd.,
Annls
Speg., An. Soc. Cient. Argentina
19:
247, 1885.
Specimens examined:
Russia-. Dal'nevostochnyi Krai, locality in­
decipherable, on Prunus triflora, 17 May, O.Natal'ina
(LE 1 4 6 8 5 3 ;
possible type material of Polystigmella ussuriensis); Dal'nevostochnyi
Krai, Vladivostokskii okrug., okrestnosti g. Vladivostoka, on Prunus
triflora, 23 July 1927, V. Transhel'
Krai,
Primorskaya
oblast',
(LE 146588); Dal'nevostochnyi
Dolina
r.
Surgan,
c
Vladimiro
Aleksandrov'ske, on Prunus triflora, 13 Sep. 1938, V. I. Protsenko (LE
146581); locality indeciperable, on 'sliva usyrskaya', 5 Aug. 1932, s.
coll. (LE 146580); Amur, Blagovestshenk, on Prunus mandschurica, 27
July 1924, Aleksahine
(LEP); Primorskii Krai, Spasskii Raion, near
Nakhimovskii, on Prunus ussuriensis, 23 Sep. 1981, V. A. Limin (LEP);
Yuzhno-ussuriskii Krai, Spasskoe, host not specified, 11 June 1912,
Naumov (LEP; probable type material of Rhodoseptoria
ussuriensis).
Typification: U.S.A.: Alabama, on IPrunus caroliniana
[given as in
Ceraso carolia], Beaumont, ex herb. Berkeley no. 4893 (K! - probable
syntype
of Phyllachora
beaumontii,
here
designated
neotype).
Unlocalized, ex herb. Berkeley no. 4635 (K! - probable syntype of
Phyllochora beaumontii). Unlocalized, ex herb. Berkeley no. 4 8 9 1 (K!
- probable syntype of Phyllachora beaumontii). Unlocalized, ex herb.
Cooke (K! — probable syntype of Phyllachora beaumontii). Paraguay:
Mbocaiate, prope Villa Rica, on leaves of IPrunus sp., 15 Jan. 1882,
B. Balansa {Plantes du Paraguay no. 3443, IMI 1 5 8 8 1 3 ! - isotype of
Auerswaldia
puccinioides).
T h e t y p e material studied o f Auerswaldiella
immature, but appears
Phyllachora
beaumontii,
puccinioides
is
t o b e l o n g t o the same species as
as was suspected b y T h e i s s e n & S y d o w
( 1 9 1 5 ) . C o o k e ' s name w o u l d appear t o b e published a few
EXCLUDED AND DOUBTFUL
SPECIES
weeks previous to that o f Spegazzini; the issue o f
Grevillea
Everh.,
Fungi
containing C o o k e ' s paper is dated M a r c h 1 8 8 5 , and according
= H y p o n e c t r i a a d e n o s t o m a t i s (Farl.) P. F. Cannon,
comb.
o f that year. Bearing in mind the lack o f c o m p l e t e certainty o f
Polystigma
adenoslomatis
Columbiani
in
Ellis
no. 2 0 4 9 , 1 9 0 5 , in
Farl.,
sched.
&
to Farr ( 1 9 7 3 ) Spegazzini's w o r k was n o t published until June
nov.
publication dates, the immaturity o f o n e o f the t y p e specimens
Infected areas n o t clearly delimited. Blackened
regions t o 1 m m
diam., rather irregular, the individual ascomata n o t c o m p l e t e l y
coalescing, containing 1—5 ascomata. Ascomata
hologenous,
the ostioles epigenous, 2 5 0 - 3 5 0 um diam., spherical t o slightly
oblate, the ostiolar region shallowly conical. Ascomatal
poorly
developed.
100—140 Mm diam.,
Upper
wall
composed
reduced
to
o f several
a
epidermoidea.
Lower
and
side
layers o f
walls
almost
very
Paraphyses
n o t seen.
well-developed, copious, to 4 Mm diam., fairly thin-
walled, slightly tapering, septate, branching n o t seen.
Asci
formed from a crozier system, 58—66 x 1 8 - 2 2 um, saccate,
short-stalked,
thin-walled
at
all
stages
of
development,
the apex rounded, without clearly-defined apical structures,
8-spored. Ascospores
arranged biseriately, 1 3 - 5 - 1 5 - 5 ( - 1 8 ) X 8 -
9-5 Mm, ellipsoidal to ovoid-ellipsoidal, thin-walled, hyaline,
smooth, aseptate, without a mucous sheath.
Dothidea fructigena
Fungorum
2:
Schwein., Trans. Amer. Phil. Soc. Philadelphia
= B o t r y o s p h a e r i a o b t u s a (Schwein.) Shoemaker, Can.
four.
Bot. 4 2 : 1 2 9 8 , 1 9 6 4 .
Typification: 'in pomis putridissimis in campo dejectis prope Camden'
[see below).
None
of
the
collections
cited
below
is
adequately
documented, and it is possible that t h e y are all authentic
material. T h e only o n e found to contain spores is the o n e from
Bloxam's
herbarium,
which
could
have
originated
from
Schweinitz's collections via Berkeley. T h i s is in p o o r condition,
but almost certainly represents the Sphaeropsis
Botryosphaeria
obtusa.
anamorph o f
S e e S h o e m a k e r ( 1 9 6 4 ) and Sivanesan
( 1 9 8 4 ) for further information. T h e Schweinitz n a m e p r o b a b l y
Adenosioma
fasciculatum, 12 April 1903, C. H. Thompson (FH - holotype, K!, WSP
5 9 3 5 ! — isotypes of Polystigma
(Schwein.) S a c c , Sylloge
4: 232, 1832.
represents
Typification: U.S.A.: California: San Mateo County, on
fructigena
622, 1883.
absent,
composed o f a narrow layer o f almost hyaline fungal tissue
surrounded b y strongly tanned host cells. Periphyses
Phyllachora
walls
clypeus,
degenerate plant cells + filled with fairly strongly melanized
textura
and lack o f research into species concepts, a n e w c o m b i n a t i o n
is n o t appropriate.
adenostomatis).
the earliest available n a m e for the
anamorph.
Bearing in mind the p o o r condition o f the material examined
and its dubious
type
status, not
to mention
the
highly
questionable need for a separate n a m e for the anamorph, n o
n e w combination is made.
Host species: Adenosioma fasciculatum Hook. & Am.
Materials examined; U.S.A.: Massachusetts, on 'decaying apples', ex
Distribution: U.S.A. (California); only known from the type collection.
herb. Currey (K); Massachusetts, on 'decaying apple', ex herb. Cooke
(K). Unlocalized, on 'decaying apples', ex herb. Bloxam (K);
Phyllachora
unlocalized, on 'old apples', misit Curtis, ex herb. Ravenel (K);
1883.
unlocalized, 'in poma', M. A. Curtis, ex herb. Berkeley (K). No
information, ex herb Schweinitz, ex herb. Berkeley ['no fruit'] (K).
Phyllachora
impressa
(Fr.) S a c c , Sylloge Fungorum
Dothidea
impressa
Fr., Systema
Discosia
impressa
(Fr.) Fr., Summa
423,
Mycologicum
Vegetabilium
(Schwein.) S a c c , Sylloge
2: 6 1 1 ,
The Schweinitz fungus appears t o represent the anamorphic
state o f Diplocarpon
Scandinaviae:
Fungorum
Dothidea rosae Schwein., Trans. Amer. Philos. Soc. Philadelphia,
n.s., 4 : 2 3 5 , 1 8 3 2 .
= D i p l o c a r p o n r o s a e F. A . W o l f , Bot. Gaz. 5 4 : 2 3 1 , 1 9 1 2 .
2: 609, 1883.
2: 5 6 4 , 1 8 2 3 .
rosae
rosae, s o the n a m e does n o t threaten that
of W o l f under the current nomenclatural rules. T h e fungus is
the c o m m o n black spot o f cultivated roses.
1849.
= D i p l o c a r p o n i m p r e s s u r n (Fr.) L. H o l m & K. Holm, Kew
Bull. 3 1 : 5 6 8 , 1 9 7 7 .
I have received m o s t valuable assistance from a number o f
Typification:
Sweden: Femsjo [?], on decaying leaves of Rubus
colleagues in the
chamaemorus,
s.d., E. M. Fries (Fries, Scleromyci Sueciae no. 152, UPS
Polystigma
- holotype of Dothidea
impressa).
collection o f overwintered
material o f
species, with mature ascomata. A l m o s t all collec­
tions in herbaria are o f the conspicuous anamorphic stromata
on living leaves. M y thanks are t o : Drs Z. Banihashemi and
Phyllachora
pomigena
(Schwein.) S a c c , Sylloge Fungorum
2: 622,
DOTHIDEA POMIGENA Schwein., Trans. Amer. Phil. Soc. n.s., 4 : 2 3 2 ,
L. N . Vasil'eva
= G l o e o d e s p o m i g e n a (Schwein.) C o l b y , Trans.
Illin.
St.
Sci. 1 3 : 1 5 7 , 1 9 2 0 .
(Vladivostok,
Russia)
and
Drs
M . Arsenijevic and S. S t o j a n o v i c (Serbia).
I
1832.
Acad.
V. Minassian (Iran); Prof. A . Sztejnberg and D r A . Lin (Israel);
Dr
1883.
have
following
also received hospitality
during
herbaria,
valuable
and
access
to
visits
to
the
collections:
R o y a l B o t a n i c Gardens, K e w , U.K. ( K ) ; M e r s e y s i d e C o u n t y
Typification: 'frequens in maturis pomis dictis "Newton Pippins",
M u s e u m s , Liverpool, U.K. (LIV); Swedish M u s e u m o f Natural
History, S t o c k h o l m (S); University o f Uppsala, S w e d e n ( U P S ) ;
Pennsylv.', not seen.
D e p a r t m e n t o f Plant P a t h o l o g y , W a s h i n g t o n S t a t e University,
(of which this is the type) appears n o t
U.S.A. ( W S P ) . T h e following herbaria kindly sent me material
to h a v e been examined recently. T h e fungus as interpreted b y
on loan; A c a d e m i a Sinica, Beijing, China ( H M A S ) ; K o m a r o v
The
Colby
genus Gloeodes
(1920)
may
represent
an
immature
species
of
Botanical Institute, St Petersburg,
Russia (LE); All-Union
( A s c o m y c o t a ) , rather than a c o e l o m y c e t e , the
Institute for Plant Protection, St Petersburg, Russia (LEP);
structures s h o w n as conidia actually being immature asci.
N e w Y o r k S t a t e M u s e u m , A l b a n y , U.S.A. ( N Y S ) ; Botanical
Schizothyrium
However,
there
is a c o m p l e x o f fungal
species on
this
Garden
and M u s e u m ,
University o f O s l o , N o r w a y
(O);
substratum which needs systematic study (Johnson, Sutton &
Erbario Patavinum, Padova, Italy ( P A D ) ; and the A c a d e m y o f
Hodges, 1 9 9 6 ) .
Natural Sciences o f Philadelphia, U.S.A. (PH).
Phyllachora
and M s G . G o d w i n
I M I staff are also gratefully acknowledged. M i s s T . S. Caine
potentillae
(Schwein.) P e c k e x Seym., Host Index of
the Fungi of North America:
Professor
388, 1929.
Sphaeria
potentillae
Schwein., Schriften
Gesellschaft zu Leipzig 1 : 4 6 , 1 8 2 2 .
der
naturforschenden
Drs
D. W . M i n t e r and P. M . Kirk made valuable
B . C. Sutton,
contributions
during discussions o f species concepts, nomenclature etc., and
U.S.A.: Pennsylvania, Bethlehem, host not cited
[Potentilla sp. presumed], s.d., s. coll. (Collins Collection no. 378, PHI,
possible type material of Sphaeria potentillae); same locality, host not
cited, 1826, s. coll. (PH! —authentic material of Sphaeria
T h e n a m e Phyllachora
potentillae
potentillae).
w a s cited in the h o s t index
published b y S e y m o u r ( 1 9 2 9 ) , and n o o t h e r reference has b e e n
found (J. H. Haines, pers. comm.
1 2 M a y 1 9 9 5 ) . Examination o f
authentic material o f Sphaeria
potentillae
shows a m e m b e r o f
the Dothideales, possibly referable t o Nodulosphaeria
(Shoemaker,
for technical assistance.
and
in their c o m m e n t s on the manuscript.
? = N o d u l o s p h a e r i a sp.
Typification:
are thanked
D . L. Hawksworth
1 9 8 4 ; Shoemaker
&
Babcock,
Rabenh.
1987).
The
collections studied are unfortunately slightly immature, w h i c h
makes their disposition uncertain. Unusual features o f this
species include a distinctly setose a s c o m a which is almost
c o m p l e t e l y superficial (presumably m o r e or less unavoidable
w h e n developing in a leaf), well-developed cellular pseudoparaphyses, and ascospores which appear t o h a v e a distinctly
swollen section. M a t u r e ascospores were n o t seen; immature
ones measured 2 6 - 2 9 x 3 - 5 - 4 pm, w e r e c o m p l e t e l y hyaline
and appeared t o be b e c o m i n g at least 3 - s e p t a t e .
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The
authors
gratefully
acknowledge
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R. A .
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concentrations induce
Department.
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