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Taxonomic notes on Excipularia and Scolicosporium

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A recent collection of Excipularia fusispora, unknown from Britain for 121 years, is described and illustrated. Emended generic descriptions for Excipularia and Scolicosporium, both considered monotypic, are provided. Differences between the two genera, a source of confusion in the literature, are elucidated and their taxonomic positions discussed. Excipulariopsis gen. nov. is established for Excipularia narsapurensis, and a key to this and similar genera is offered. The relationship between Scolicosporium fagi and Asterosporium asterospermum is discussed, and notes on the remaining species of Excipularia and Scolicosporium are given. S. betulae and S. fusarioides are considered synonyms of E. fusispora.
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Trans. Br. mycol. Soc. 78 (2) 247-257
(1982).
[ 247 ]
Printed in Great Britain
TAXONOMIC
NOTES
ON
EXCIPULARIA
AND
SCOLICOSPORIUM
By
B. M.
SPOONER
The Herbarium,
Royal
Botanical Gardens,
Kew,
Richmond, Surrey,
U.K.,
TW93AE
AND
P. M.
KIRK
Commonwealth Mycological Institute, Ferry Lane,
Kew,
Richmond, Surrey,
U.K.,
TW9
3AF
Arecent collection of Excipularia fusispora, unknown from Britain for 121 years, is described
and illustrated. Emended generic descriptions for Excipularia and Scolicosporium, both
considered monotypic, are provided. Differences between the two genera, a source of con-
fusion in the literature, are elucidated and their taxonomic positions discussed. Excipulariopsis
gen. nov. is established for Excipularia narsapurensis, and a key to this and similar genera is
offered.
The
relationship between Scolicosporium fagi and Asterosporium asterospermum is
discussed, and notes on the remaining species of Excipularia and Scolicosporium are given.
S. betulae and S. fusarioides are considered synonyms of E. fusispora.
Arecent collection of Excipularia fusispora (Berk.
&Broome)
Sacco
(1884), received from Orpington,
Kent, has afforded an opportunity to redescribe
this neglected species and reassess its taxonomic
position. It has not been reported from Britain
since its description (Berkeley & Broome, 1859)
and has only rarely been collected elsewhere in
Europe.
The
inconspicuous habit of the tiny
conidiomata, usually hidden beneath or amongst
cracks in bark, is no doubt one of the reasons for
this.
From an examination of the present collection a
clearer concept of the delimitation of the genus
has emerged. During the process of initially
identifying the material the similarity of Scolico-
sporium fusaricides (Sacc.) B. Sutton (1975) was
noted which, on examination of the holotype, was
found to be conspecific.
This
prompted a re-
appraisal of the species described in Excipularia
and Scolicosporium.
It
was also found desirable to
clarify the distinction between similar, possibly
related genera.
Saccardo (1884) introduced Excipularia as a
monotypic genus for Excipula fusispora Berk. &
Broome with
the
following brief diagnosis:
'Perithecia excipuliformia, atra, setosa. Sporulae
fusoideae, z-pluri septatae, hyalino-fuscellae,
basidiis brevibus suffultae',
Four
species have
subsequently been described. Ellis (1976) gave an
expanded generic description and illustrated E.
narsapurensis Subram., a species of wood and
bark in India (Subramanian, 1971).
This
species,
rather than the holotype, was employed by both
Barnett &
Hunter
(1972) and Carmichael, Ken-
drick, Conners & Sigler (1980) to illustrate the
genus. Detailed examination of the Kentish
material and of the type collection, which is now
extremely scant, has revealed, however, that E.
fusispora and E.
narsapurensis
cannot be considered
congeneric. Current concepts of the genus, which
allow for inclusion of the latter species, must
therefore be revised.
Four
further collections of E. fusispora have
been available for study and the following emended
generic diagnosis and species description is
offered.
EXCIPULARIA
Sacc.,
Syll.
fungo 3: 689 (1884)·
Mycelium immersed. Conidiomata superficial,
cupulate, blackish brown to black, with a basal
aggregation of brown to dark brown textura
angularis. Sterile elements usually abundant
throughout the conidioma, usually proximally
branched, apex acuminate, pale brown, septate,
smooth, thin-walled. Marginal sterile elements
frequently setiform, with the apical cell modified,
becoming thick-walled and dark brown. Conidia-
phores pale brown, usually proximally branched,
arising from the basal aggregation of textura
angularis, septate, cylindrical, smooth-walled.
Conidiogenous cells integrated, monoblastic, deter-
minate, cylindrical, very pale brown to pale brown.
Conidia falcate to crescentic, smooth, euseptate,
apical cell rounded to corniform, basal cell
truncate with an indistinct unthickened scar, pale
brown to brown, terminal cells hyaline.
Type
species E. fusispora (Berk. &Broome)
Sacco
Excipularia and Scolicosporium
From
asurvey of the literature, it is apparent
that
no uniform concept of
the
position of
the
genus exists.
It
has, in fact, been variously
assigned to
both
Hyphomycetes and Coelomycetes.
Saccardo (1884) originally placed it in the Sphaer-
opsideae and Grove (1937) also included it in
the
Coelomycetes. Von Hohnel (1904), however,
considered it better placed in Tuberculariaceae
and
this position in
the
Hyphomycetes was
endorsed by Subramanian (1956, 1971), Barnett
&
Hunter
(1972), Ellis (1976) and Carmichael,
Kendrick, Connors &Sigler (1980). Wakefield &
Bisby (1941), in contrast, exclude any reference
to it in their check-list of British Hyphomycetes,
and
Clements &Shear (1931) inexplicably key it
out
in
both
Phomales and Moniliales. Sutton,
although including Scolicosporium fusarioides in
the
Coelomycetes (Sutton, 1975, 1980), excluded
Excipularia from his account of coelomycete
generic names (Sutton, 1977).
That
such disagreement should exist at all is
indicative of
the
unsatisfactory definition of the
terms 'coelomycete' and 'hyphomycete',
These
terms are too general and, whilst obviously
distinct
and
of useful application in extreme, and
perhaps even the majority of cases, would appear
to lack satisfactory distinguishing characters in
'borderline' cases where stromatic conidiomata
are involved.
Sutton
(1980) has
the
following
definition of Coelomycetes: 'anamorphic fungi
with pycnidial, acervular, stromatic or pycno-
thyrial conidiomata', An acervulus, by definition,
is never superficial in origin and, according to
Sutton
(1980), the 'conidia are initiated and
produced whilst still covered by the host tissue'.
The
development of
the
conidia has therefore
been regarded as
the
important character in
distinguishing
the
two groups, and on this basis
it is evident
that
Excipularia should be considered
as a hyphomycete. However, aconsideration of
'borderline' genera reveals that they exhibit a range
of structures for which
the
ontogeny of the coni-
dioma is imperfectly known or difficult to define,
and
in these cases it is questionable whether or
not
adistinction between Hyphomycetes and Coelo-
mycetes is desirable or even possible. Sutton
(1977) includes, for example, Bloxamia, Myco-
leptodiscus, Oramasia and Toxosporium as accept-
able coelomycete genera.
The
first three of these
are, however, clearly sporodochial in nature and,
having entirely superficial conidial development,
should, on
the
above definition, be regarded as
hyphomycetes, Toxosporium, having superficial
stromatic conidiomata, should be similarly inter-
preted.
It
is interesting
that
Ellis (1976) includes
Bloxamia, Mycoleptodiscus and Toxosporium in his
account of dematiaceous hyphomycete genera. A
further notable example is furnished by Hawks-
worth (1979) in his description of a new species of
Psammina.
This
genus was previously monotypic
and included by Sutton (1980) as an acervular
coelomycete,
The
type species, P. bommerae
Rouss. &Sacc., develops a rudimentary
sub-
epidermal acervulus on culms of Ammophila and
Juncus. Psammina stipitata D. Hawksw. is, in
contrast, a mainly superficial lichenicolous species
in which conidial development is initiated above
the host surface. Hawksworth has therefore inter-
preted
the
conidioma of P. bommerae as com-
prising merely a rudimentary stroma which may
appear acervular due to modification caused by a
subepidermal habit, and has suggested that
Psammina is better regarded as a hyphomycete.
We would suggest, on the basis of superficial
conidial development, that
Scolicosporium
should
also be so regarded.
Considering, however,
that
Deuteromycotina
(Ainsworth, 1973) comprises only form genera,
the
taxonomic arrangement of which, presently at
least, is for convenience only and does
not
neces-
sarily reflect natural relationships, it could be
argued that it is unnecessary to distinguish
betweenHyphomycetesand Coelomycetes. Indeed,
asuprageneric classification of Fungi Imperfecti
was proposed in 1973 by
Sutton
(in press;
recently summarized: Sutton, 1980), employing
conidiogenesis as a basis, and considering gross
morphology of
the
conidioma only at subordinal
level.
This
concept appears to be more satisfactory
than
that
traditionally employed.
EXCIPULARIA FUSISPORA (Berk. &Broome) Sacc.,
Syll.
fungo
3: 689 (1884). (Figs 1, 2)
Excipula fusispora Berk. &Broome, Ann. Mag.
nat. Hist. 1859
(III):
359 (1859).
Coryneum fusarioides Sacc., Michelia
2:
120
(1880).
Scolicosporium fusarioides (Sacc.) B. Sutton,
Mycol. Pap. 138: 111 (1975).
Scolicosporium betulae Rostrup, Vidensk, Selsk.
Skrlft. I. Math. Naturv. Kl. 1904: 39.
(1904)·
Conidiomata superficial, cupulate, 50-150
pm
diam, blackish-brown to black, with a basal
aggregation of brown to dark brown textura
angularis. Sterile elements abundant throughout
the conidioma, usually proximally branched, apex
acuminate, pale brown, septate, smooth,
thin-
walled,
45""90
pm
long,
2-3
pm
wide. Marginal
sterile elements frequently setiform, with
the
apical cell modified, becoming thick-walled and
dark brown, 20-65
pm
long,
2-3
pm
wide.
Conidiophores pale brown, proximally branched
B.
M.
Spooner
andP.
M.
Kirk
Fig. 1. Excipulariafusispora. Setiform sterile element, developing and mature conidia ( x 2000).
249
or simple, arismg from
the
basal cells, septate,
cylindrical, smooth-walled, 10-30
pm
long,
1'5-
2
pm
wide. Conidiogenous cells integrated, mono-
blastic, determinate, cylindrical, very pale brown
to pale brown. Conidia falcate to crescentic,
smooth-walled, 8(-10)-euseptate, apical cell
rounded
to comiform, basal cell truncate
with
an
indistinct unthickened scar, pale brown to brown,
terminal cells hyaline, (32-) 40-48
pm
long,
4-5'5
(-6) /lm wide.
Specimens examined: On inner surface of Clematis
uitalba
L.
bark, Batheaston, Somerset, U.K., Jan. 1859,
C. E. Broome 234,
IMI
249659,
slide ex herb. K,
holotype of Excipula fusispora, On bark of Populus
fastigiate Fouger., Rouen, France, undated, Letendre,
IMI
179078,
slideexherb. PAD, holotype ofCoryneum
[usarioides. On Acer saccharinum
L.
bark, Ottawa,
Canada, 14 Sept. 1897,J. M. Macoun, IMI 238243b.
On bark of Betula sp., Norway, c. 1900, E. Rostrup,
IMI
255640, slide ex herb. C, holotype of Scolico-
sporium betulae. On twigs of Berberis vulgaris L. f.
atropurpurea, Sweden, 21 July 1926, A. G. Eliasson,
IMI
31192.On Populussp. bark, Edenbridge, Orping-
ton, Kent, U.K., May 1980, Mrs J. Weightman,
IMI
24856Sdand K.
The
development
of
dark brown, thick-walled
setiforrn elements in Excipularia fusispora is
Excipularia and Scolicosporium
// DE
(
F
/
,
Fig. 2.
(A-G)
Excipularia fusispora. (A-C) 1M1 248565. (A) Setae and sterile elements in conidioma
( x 750). (B) Developing conidia. Arrow indicates position of septum delimiting conidium from conidio-
genous cell
(x
750). (C) Mature conidium ( x 750). (D and E) 1M1 249659 . (D) Conidioma ( x 300).
(E) Mature conidia ( x 750). (F and G) 1M1 179078. (F) Conidioma ( x 300). (G) Mature conidia ( x 750).
B. M.
Spooner
andP. M. Kirk
evidently
not
constant, varying, perhaps, in
relation to environmental conditions.
The
most
luxuriant development, as seen, for example, in
the
Kentish material, results in a dense fringing
of
the
conidioma so
that,
under
the
binocular
microscope, it resembles a tiny black apothecium
with
a well developed excipular structure.
The
thin-walled, pale brown sterile elements, which are
always present, may, on
the
other
hand, develop
no further, so
that
the
darker setiform elements
are lacking.
This
situation is seen in the Canadian
collection
and
in
the
holotype
of
Coryneum
fusarioides. Conidium morphology
and
conidio-
genesis in these
and
other
collections are, however,
indistinguishable
and
there can be no
doubt
as to
their
identity.
The
holotype of Scolicosporium betulae is
unfortunately arather poor collection.
The
conidiomata are poorly developed
and
neither
setiform elements
nor
conidiophores have been
observed.
There
are, however,
abundant
conidia,
measuring 32-37 x
5-6
pm,
which agree in size
and
also in form with those of E. fusispora.
Of
the
four additional species described in
Excipularia, E.
narsapurensis
is
the
best known.
The
emended generic description offered here,
however, does
not
allow for
the
inclusion
of
this
species,
and
hence a new genus is established to
accommodate it.
Excipulariopsis P. M. Kirk &Spooner gen.nov.
Mycelium plerumque immersum. Conidiomata super-
ficialia, pulvinata, fusca ad atra, in fundamento ex
cellulis, crassitunicatis, fuscis et aggregatis composita
quae setas producunt in peripheria. Setae subulatae,
fuscae, septatae, crassitunicatae, laeves.Conidiophorae
breves, cylindricae, pallide brunneae, non ramosae.
Cellulae conidiogenae holoblasticae, monoblasticae, in
conidiophoris incorporatae, terminales, determinatae.
Conidia acrogena, solitaria, sicca, late fusoidea, ad
basim truncata, multiseptata, laevia, crassitunicata,
fusca, cellulisterminalibus hyalinis ad pallidebrunneis,
Sp,
typo: Excipulariopsis narsapurensis
(Subramanian) Spooner &P.
M.
Kirk
comb.nov.
(basionym: Excipularia
narsapurensis
Subramanian,
J. Indian Bot. Soc. 35: 56, 1956).
Mycelium mostly immersed. Conidiomata super-
ficial, pulvinate, dark brown to black,
with
a basal
aggregation of thick-walled, dark brown cells.
Setae peripheral, arising directly from cells of
the
basal stroma, subulate, dark brown, septate,
thick-walled, smooth. Conidiophores short, cylin-
drical, pale brown, unbranched. Conidiogenous cells
holoblastic, monoblastic, integrated, terminal,
determinate. Conidia aerogenous, solitary, dry,
broadly fusoid, truncate at the base, multiseptate,
smooth, thick-walled,
dark
brown,
with
hyaline
to very pale brown terminal cells.
The
genus differs from Excipularia in producing
non-falcate, thick-walled conidia
and
in possessing
septate, consistently thick-walled, dark brown,
subulate setae at
the
periphery
of
anon-cupulate
conidioma
and
arising directly from
the
basal
stroma. On the basis of available evidence,
Excipulariopsis is considered to be monotypic,
Having established
that
S. fusarioides is synony-
mous with E. fusispora the possibility of
the
synonymy of
Scolicosporium
and
Excipularia was
considered. Examination of an isotype
of
S. fagi
Lib.,
the
type species of Scolicosporium, reveals,
however,
that
this is
not
the
case, as is evident
from
the
following description.
SCOLICOSPORIUM
Lib.
apud
Roum., Rev. Mycol.
2:
22 (1880).
Conidiomata
erumpent
or superficial,
brown
to
dark brown, stromatic, discoid to pulvinate,
composed of thin-walled angular cells. Setae
absent. Conidiophores simple or branched, hyaline,
septate. Conidiogenous cells holoblastic, mono-
blastic, integrated, terminal, percurrent. Conidia
aerogenous, solitary, dry, falcate to crescentic,
truncate at
the
base, corniform to distinctly
curved at
the
apex, multiseptate, smooth, brown
to dark brown, thinner-walled
and
paler towards
the
base
and
apex, terminal cells frequently
hyaline.
Type
species S. fagi
Lib.
SCOLICOSPORIUM
MACROSPORIUM
(Berk.) B.
Sut-
ton, Mycol. Pap.
141:
185 (1977). (Figs
3,4)
Coryneum macrosporium Berk. in Hooker's
English Flora 5 (2): 355 (1836).
Scolicosporium
.fagi
Lib.
apud
Roum.,
Rev.
Mycol.
2:
22 (1880).
?Sporidesmium uermiforme Riess in Fres.,
Beitrage zur
Myk.:
51 (1863). (fide
Sutton,
1980).
Teleomorphosis: Asteromassaria
macrospora
(Desm.)
Hohnel
(1917).
Specimens
examined:
(a) Anamorph only: On cortex
of
Fagus
syluatica L., Malmedy, Belgium, undated,
M. A. Libert,
Mycotheca
universalis
(Thumen),
Reliquiae Libertianae No. 1683, isotype of Scolico-
sporium
fagi, in herb. K, slide IMI 249657; sine loc.,
July 1939, F. Petrak,
IMI
31095.
(b) Anamorph socio
Asterosporium asterospermum (Pers, ex Gray) S.
Hughes: On twigs of ?
Fagus
sp., Apethorpe, North-
amptonshire, sine datum, M. J. Berkeley, neotype
(designated here) of
Coryneum
macrosporium,
in herb.
K, slide
IMI
249658; On
Fagus
sylvatica, Ranmore
Common, Surrey, U.K., 6 Jan. 1947, S. J. Hughes,
Excipularia and Scolicosporium
Fig. 3. Scolicosporium macrosporium, (A)
IMI
249658. Developing and mature conidia
(x
1200).
(B)
IMI
249657. Mature conidium
(x
1200).
Fig. 4.
(A-F)
Scolicosporium macrosporum. (A-D)
IMI
249658. Conidioma. Arrow indicates disinte-
grating conidium of Asterosporium asterospermum
(x
200).
(B) Conidiogenous cells showing annellations
(arrows) and developing conidia
(x
750). (C) Developing conidia
(x
600).
(D) Mature conidium
(x
500).
(E and F)
IMI
249657. Mature conidia (E, x
300;
F, x 500).
B. M.
Spooner
andP. M. Kirk
253
F
E
c
The
generic name
Scoliciosporum
Massal. 1852
(Lichens) is probably of the same etymological
derivation as
Scolicosporium.
They
are, however,
sufficiently distinct, and should not be regarded
as homonyms.
There
has been confusion in the literature
concerning the synonymy and possible anamorphs
of Asteromassaria macrospora. Tulasne &Tulasne
(1863: 221) listed Sporidesmium vermiforme as a
synonym of Coryneum macrosporium, which was
correctly indicated to be the conidiophorous state
of Cucurbitaria macrospora (Desm.) Ces, &de
Not. (syn. Asteromassaria macrospora). An excel-
lent illustration (Plate XXVI, fig. 10) was provided
to substantiate this connexion, although Currey
(1859: 233) was apparently the first author to
suggest the link. Tulasne &Tulasne (1863: 229)
also described
Massaria
loricata
Tul.
&C.
Tul.
suggesting
that
the anamorph of this species is
Stilbospora
kickxii
Westend. (syn, Neohendersonia
kickxii
(Westend.) B. Sutton &Pollack). Cucur-
bitaria macrospora and
M.
loricata are illustrated
together on Plate XXVI, where they are clearly
seen to be distinct taxa. Grove (1937: 342),
however, for an obscure reason, suggested
that
M. loricata may simply be an early state (sic)
of C.
macrospora.
He also suggested (p. 326)
that
Stilbospora
pyriformis
(Otth) Grove is the pyc-
nidial state of
M.
loricata. Stilbospora
kickxii
and
Hendersonia loricata
Sacco
&Roum. are listed as
synonyms. Scolicosporium macrosporium (as S.
fagi),
with Sporidesmium vermiforme as a synonym,
was given (p. 340) as the anamorph of
Massaria
macrospora.
Following Grove, this species would,
consequently, exhibit more than one generically
distinct anamorph.
Later
(p. 342), Grove concluded
that
Astero-
sporium
asterospermum
(as A. hoffmanniiKunze ex
Wallr.) represents another conidial state of this
species, based on their frequent association.
The
association of these fungi is, however, better
interpreted as the result of a specialized sapro-
254 Excipularia andScolicosporium
IMI
98906b.
(c) Anamorph socio teleomorph: On phytic or hyperparasitic habit by Scolicosporium
Fagus sp., Wiltshire,Dec. 1850,ex herb. C. E. Broome; macrosporium. Tulasne &Tulasne (1863: 142)
sine
loco
et datem, on wood and bark, ex herb.
Bloxam
state
that
this species occurs surrounding areas
336. (d) Anamorph socio teleomorph socio
A.
astero- formerly occupied by A. asterospermum (as A.
spermum: Dyrham Park, 12 Oct.
1850,
C. E. Broome;
On Fagus syloatica, Kings Lynn, 1872, C. B. Plow- hoffmannii) and further state (1863: 222)
that
right; ex herb. M. C. Cooke,sine
loco
et datum; On
Massaria
macrospora '
...
arises most often round
Fagus sp., ex herb.
Berkeley
607, sine
loco
et datum; perithecia of Asterosporium hoffmannii,
and
indeed
On Fagus sp., Batheaston,Somerset. Collections from seems to grow out of it, exactly as if it were truly
groups (c) and (d) are disposed sub teleomorph in parasitic upon it'.
This
is apparently the first
herb. K. It should be added that a total of 15 British published suggestion of hyperparasitism involving
collections of Asteromassaria macrosporaare preserved these species.
in herb. K. Eight collections, therefore, lack the Von Hohnel (1917: 368) also discussed the
:;=~:h
and are not associated with the Astero- association of these fungi.
In
the many examples
he studied, he found that conidia of S. macro-
sporium always developed eventually on conidio-
mata of A.
asterospermum.
He
concluded them to
be 'metagenetically related', the Asterosporium
representing afurther anamorph of
Massaria
macrospora.
The
development of the fungus, as
he understood it, began with the formation,
beneathbark, of conidia ofthe Asterosporium. These
germinated to form a much-branched mycelium,
on which conidia of S. macrosporium appeared,
functioning for dispersal. At the edge of the older
specimens perithecia of the
Massaria
developed,
at which time the Asterosporium conidia broke
down and disappeared, followed by those of the
Scolicosporium. On the basis of this apparently
unique mode of development, he established the
genus Asteromassaria for the perfect state.
The
situation may be more complex than is so
far indicated. Currey (1856), discussing the fact
that
various distinct conidial states belong,
apparently, to a single holomorph, gave an account
of Myriocephalum botryosum (Mont.) Fres. sporu-
lating on mature conidiomata of Asterosporium
asterospermum and concluded them to be synana-
morphs, Tulasne &Tulasne (1863: 242-243) also
discuss these taxa,
but
describe the Myriocephalum
(as
Hyperomyxa
sp.) as growing on 'old worn-out
areas occupied by Asterosporium hoffmannii':
They
concluded
that
the relationship is better inter-
preted as being 'either parasitic or secondary'.
This
would indeed appear to be the case and, if so,
represents a second example of a fungus utilising
Asterosporium.
Further,
there is evidence
that
yet a
third
fungus may do so. Von Hohnel (1902)
discussed the association, first reported by Briard
(in Roumeguere, 1886), between A.
asterospermum
(as
A.
hoffmannii) and Fusicoccum macrosporium
Sacco
&Briard. He stated that the latter species is
always found with the former and referred also to
the account by Fuckel (1870: 351) which evidently
also describes the association of these fungi, the
Fusicoccum being described
but
not
identified.
The
large hyaline conidia of F. macrosporium
were considered by Von Hohnel to be immature
B. M.
Spooner
and P. M. Kirk
255
asci, so
that
the fungus was thought, therefore, to
represent the perfect state of A. asterospermum.
Furthermore,
Tulasne
&
Tulasne
(1863: 241)
described the occurrence
of
the Asterosporium on
both
Massaria
eburnea
and M. loricata.
They
concluded it to be
the
anamorph of one or
other
of these species, an interpretation which, however,
is
not
necessarily justified.
The
Asterosporium is
not
likely to represent the conidial stage of
both
these taxa and, therefore, if the anamorph of one,
it
must
have a different, unexplained relationship
with the other. Massaria loricatahas already been
shown to have N
eohendersonia
kickxii as a conidial
state.
Unfortunately,
further
confusion is liable to
arise on two counts. Firstly, Melanconis macro-
sperma
Tu!.
(1856 :
110)
is, by coincidence,
described on the previous page to the similar-
sounding Melanconis
macrospora
Tul.
Moreover,
the
reference cited
under
that name in
the
Carpo-
logia (Tulasne &Tulasne, 1863: 132) is incorrect
and
refers, rather, to Melanconis macrospermat
Secondly, Melanconis macrosperma has Stilbospora
macrosperma
Pers. ex
Merat
as an anamorph for
which
the
Tulasnes have, incorrectly, suggested
that
Stilbospora pyriformis Hoffm. ex
Fr.
non
Otth
(syn. Stegonsporium pyriforme (Hoffm. ex
Fr.)
Corda) is a synonym.
A
number
of anamorph genera are similar to
Excipularia
and
Scolicosporium in being charac-
terized by acervular or simple stromatic,
erum-
pent
or superficial conidiomata,
and
non-sym-
podial holoblastic conidiogenous cells forming
euseptate, versicoloured, phragmoseptate conidia.
These
conidia
mayor
may
not
possess cellular
appendages,
but
lack either agelatinous sheath or
setiform appendages. Genera of this type treated
by
Sutton
(1980) are placed in his new suborder
Blastostromatineae. However, despite
the
fact
that
conidiogenesis in the type species of S
colicosporium
is annellidic,
that
genus is keyed
out
by Sutton in
the monoblastic group.
It
has
not
proved asimple task to distinguish
satisfactorily genera exhibiting these characters.
In
an
attempt
to
understand
possible relationships
of
the
remaining species described in Scolico-
sporium
and
Excipularia, the following tentative
key has, therefore, been compiled:
2
3
7
Pe
sta/ot
ia
5
Seimatosporium
6
1.
Conidiogenous cells determinate. Conidiomatafrequently setose, Conidialackingdistinct
appendages
1. Conidiogenous cellsannellidic. Setaeabsent. Conidia frequentlywith terminalappendages
2.
Setae brown, septate, thick-walled, arising directly from the stroma.
Th
in-walled
sterile elements absent. Conidia not falcate or crescentic
Excipulariopsis
2.
Thin-walled sterile element typically present throughout the conidioma with apices
frequently becoming setiform at the margin. Conidia falcate to crescentic
Excipularia
3. Conidia with at least one distinct, often branched apical and/or basal appendage. 4
3. Conidia lacking distinct appendages; apical cell at most acuminate, but then basal
appendagesabsent
4. Conidia with median cells distoseptate
4. Conidia with median cells euseptate .
5. Exogenousbasal appendage(s) present .
5. Basal appendages endogenousor absent
6. Apical cell of conidia with more than one, branched or unbranched appendage
Pestalotiopsis and Truncatella
6. Apical cell of conidia without, or with a single, sometimes branched appendage
Seiridium and
Mono
ch
aetia
7. Conidia 5-septate, the central septum appearing deeply pigmented and thickened Toxosporium
7. Number of septa variable. Central septum not modified . 8
8. Conidia curved, falcate, usually with 7 or more septa . Scolicosporium
8. Conidia ellipsoid to obovoid, straight, usually with 5 or fewer septa Seimatosporium
It
should be emphasized
that
the appearance
of
genera in this key does
not
necessarily reflect a
close natural relationship between them.
Further,
it will be noted
that
PestalotiopsisfTruncatella
and
Seiridium/Monochaetia key
out
in pairs. Distinc-
tion between
the
genera in each of these pairs is
based solely on the
number
of
septa in the conidia.
Thus
Truncatella has conidia
with
3septa,
whereas in Pestalotiopsis they are 4-septate.
Similarly, Seiridium has 5-septate conidia whilst in
Monochaetia they may be 3 or more frequently
4-septate.
Sutton
(1969) has further discussed the
distinction between Monochaetia
and
Seiridium,
but
we have been unable to discover any more
positive distinguishing character
than
the
number
of septa in the conidia.
It
should be noted
that
a
number
of
species
of
Stigmina (Ellis, 1971, 1976) will key
here
to
Scolicosporium.
This
does not, however, necessarily
reflect their
true
relationships.
The
current
concept
of Stigmina includes a large
and
certainly hetero-
geneous assemblage of species, not all of which
Excipularia and Scolicosporium
have versicoloured or even phragmoseptate
conidia.
Ten
species have been described in Scolico-
sporium. Sutton (1975) has pointed
out
that
the
genus is heterogeneous
but
that
no attempt has
been made at a taxonomic treatment. Such is
certainly beyond the scope of the present study,
although some comment on the possible relation-
ships of the species involved can be made from
published descriptions.
Synopsis of remaining
species
of
Excipularia and Scolicosporium
EXCIPULARIA
EPIDENDRI
P. Henn., Hedwigia 44:
174 (1905).
This
species, from Brazil, on dead flower stalks of
Epidendron sp., is described as being densely clad
with rigid black setae.
The
conidia are given as
fusoid, subhyaline, 3- to 4-septate, 12-30 x
4-5
{lm.
The
species does not appear to belong in
Excipularia,
but
no definite conclusion as to its
correct systematic position can be reached on the
strength of the description alone.
EXCIPULARIA
LIGNICOLA
P. Karsten &Malbr.,
Symb. Myc. Fenn. 24: 18 (1888).
This
species, on branches of Syringa vulgaris L.
from Finland, is described as having curved,
fusoid-bacillar, 1- to 3-septate conidia 12-15 x2
{lm.
Setae are not mentioned, and the description
is very brief. Beyond doubting it to be a good
species of Excipularia, no conclusion as to its
true
affinities can be made.
EXCIPULARIA
NIGROSEPTATA
Dev Rao &P. R. Rao,
Sydowia 22: 171 (1969).
Described from Pakistan, on unidentified wood.
From
the description, this species appears re-
markably similar to E.
narsapurensis,
differing in
having somewhat narrower conidia and possibly
smaller conidiomata.
It
is almost certainly con-
generic,
but
no formal transfer of the name is
proposed here as type material has not been
examined.
SCOLICOSPORIUM
CORYLI
Dearn. &House, N. Y.
St. Mus. Bull. 188: 39 (1916).
Described from New York, causing spots on
fading leaves of Corylus americana Watt.
Though
the small 3-septate conidia are reported as having
atapered hyaline apical cell, the remaining cells
are described as brown, with the basal cell
rounded.
The
species may consequently not be-
long in any of the genera considered here, or,
alternatively could be an aberrant species of
Seimatosporium or Monochaetia.
SCOLICOSPORIUM
PEDICELLATUM
Dearn. &Overh.,
Mycologia 16: 170 (1924).
The
conidia of this species, described from bark
of living Malus, measure 18-25 x
7'5-10
{lm,
and are 5-septate with the basal and apical cells
both appendaged. As suggested by Sutton (1975)
it is almost certainly referable to Seiridium.
SCOLICOSPORIUM
PHOEBES
T.
S. Ramakr., Proc.
Ind. Acad. Sci., Sec. B 34: 70 (1951).
Described from living leaves of
Phoebe
paniculata
Nees.
The
curved, falcate conidia are 8- to 10-
septate and measure 42-85 x8-11
{lm.
There
is
no reference to their being versicoloured which,
if they are not, would probably exclude the
species from Scolicosporium, and, similarly, from
any of the genera considered here. Otherwise, the
species could be correctly referred to Scolico-
sporium,
SCOLICOSPORIUM
TRANSVERSUM
Fairman, Proc.
RochesterAcad. Sci. 6: 124 (1922).
This
species, on bark of roots of Pyrus from New
York, is described as having oblong-fusoid, 3- to
5-septate conidia 20-30 x10
{lm,
with paler or
hyaline terminal cells.
The
conidia are described
as being apiculate at one end, which
mayor
may
not indicate the presence of an appendage. No
reference is made to the presence of setae. From
this description, it is not possible to say with
certainty to which genus this species belongs,
though, on the basis of conidium shape and
septation, it appears unlikely to belong in Exci-
pularia or Scolicosporium. Sutton (1975) states
that
the species is almost certainly referable to
Seiridium.
If
the conidia are interpreted as having
either an apical or endogenous basal appendage
this may be so, although the apparently variable
septation is not a character of
that
genus. Alter-
natively, as seems likely, it may be referable to
Seimatosporium.
SCOLICOSPORIUM
TYPHAE
(Oud.) Hohnel, Sber.
Akad. Wiss. Wien 118: 405 (1909).
This
species is currently referred to Scolecosporiella
(Petrak, 1921).
SCOLICOSPORIUM
SYZIGII
Ciccarone, Mycopath.
mycol. appl. 5: 230 (1951).
This
species, described from leaves of Syzigium
guineense
(?)
from Africa has verruculose conidia
and is unlikely to belong in Scolicosporium.
It
is
not clear from the description where its
true
affinities lie.
The
authors are indebted to Mrs J. Weightman
for collecting additional material of Excipularia
B. M.
Spooner
andP. M. Kirh
257
fusispora,
Dr
D. L. Hawksworth for drawing our
attention to
Psammina,
Dr
D. W. Minter for
correcting the Latin diagnosis, and
The
Keeper,
herb. C for loan of material of
Scolicosporium
betulae.
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Note
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Whilst this manuscript was in press a collection of
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9
MYC
78
... During this study, an Excipularialike taxon was gathered, but comprehensive morphological analysis showed that it is distinct from Excipularia s. str. since it lacks a thin-walled sterile element of conidiomata and the crescent to falcate conidia (Spooner & Kirk 1982, Seifert & Gams 2011. As a result, this unique strain was accommodated in Excipulariopsis (Spooner & Kirk 1982). ...
... since it lacks a thin-walled sterile element of conidiomata and the crescent to falcate conidia (Spooner & Kirk 1982, Seifert & Gams 2011. As a result, this unique strain was accommodated in Excipulariopsis (Spooner & Kirk 1982). The phylogeny of this unusual asexual fungus has not yet been investigated. ...
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Four new combinations Neoantrodiella gypsea, Neoantrodiella thujae (Neoantrodiellaceae), Punctulariopsis cremeoalbida, Punctulariopsis efibulata (Punctulariaceae) are also introduced here for the division Basidiomycota. Furthermore Absidia caatinguensis, Absidia koreana and Gongronella koreana (Cunninghamellaceae), Mortierella pisiformis and Mortierella formosana (Mortierellaceae) are newly introduced in the Zygomycota, while Neocallimastix cameroonii and Piromyces irregularis (Neocallimastigaceae) are introduced in the Neocallimastigomycota. Reference specimens or changes in classification and notes are provided for Alternaria ethzedia, Cucurbitaria ephedricola, Austropleospora, Austropleospora archidendri, Byssosphaeria rhodomphala, Lophiostoma caulium, Pseudopithomyces maydicus, Massariosphaeria, Neomassariosphaeria and Pestalotiopsis montellica.
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