911
Mycol. Res. '96 (11):911-924 (1992) Printed in Great Britain
Phaeoseptoria eucalypti and similar fungi on Eucalyptus, with
description of Kiwamyces gen. nov. (Coelomycetes)
J. WALKER
5 Cook Street, Baulkham Hills, N S W 2153, Australia1
B. C. SUTTON
Internationlzl Mycological Institute, Ferry Lane, Kew,Surrey TW9 3AF, U.K.
I. G . PASCOE
Department of Food and Agriculture, Institute of Plant Sciences, Swan Street, Burnley, Victoria 3121, Australia
Phaeoseptoria eucalypti, with prominent, brown, rough-walled, percurrently proliferating conidiogenous cells, is shown to be distinct
from the type species of Phaeoseptoria, P. papayae, which has small, smooth-walled and apparently non-proliferating conidiogenous
cells. As rlo suitable genus exists for P. eucalypti, the new genus Kirramyces is erected for it. Study of various Eucalyptus leaf fungi has
shown that the earliest name applied to this fungus is Cercospora epicoccoides, transferred here as Kirramyces epicoccoides. Other
synonym!, include Hendersonia grandispora and Phaeoseptoria luwnensis. Two other species of Kirramyces, K. eucalypti, based on
Cercosporh eucalypti, and K. lilianiae n. sp., are recognized. The taxonomic position of Stagonospora delegatensis and of other collections
which may represent additional species of Kirramyces requires further investigation. The host ranges of these fungi are discussed in
relationship to the subgeneric classification of the genus Eucalyptus.
Several pycnidial fungi with brown septate conidia have been
described from Eucalyptus, and some formerly placed in
Hendersolia Berk. were revised by Swart & Walker (1988). In
that paper the fate of Hendersonia grandispora McAlpine (1903)
was left in abeyance, for it proved to be an earlier name for
the spezies commonly known as Phaeoseptoria eucalypti
Hansford (1957).
Quite separately, the search for a suitable name for a
Pseudocer~cospora causing widespread damage on Eucalyptus
species in South Africa (Crous, Wingfield, Marasas & Sutton,
1989) iwolved examination of the type collections of
Cercospora epicoccoides Cooke & Massee apud Cooke (1891)
and C. eucalypti Cooke & Massee apud Cooke (1889). It was
concludtzd that neither of these species were correctly placed
in the &.nus Cercospora Fres., but C . epicoccoides proved again
to be the species commonly known as Phaeoseptoria eucalypti.
The name antedated both P. eucalypti and H. grandispora.
Since the name Phaeoseptoria eucalypti was introduced for a
pycnidial fungus with brown, verruculose, septate conidia
occurring on Eucalyptus grandis in Australia, the species has
been extensively collected on Eucalyptus species, especially,
but not exclusively, in the Southern Hemisphere. Records at
Previous address: Plant Pathology Branch, Biological and Chemical
Research institute, N.S.W.Department of Agriculture, Private Mailbag No.10,
Rydalmere, N.S.W. 2116, Australia.
the International Mycological Institute give the fungus on 14
named and numerous unnamed Eucalyptus species from 12
countries in South America, Africa, Asia a i d Australia. In
South Africa alone, Crous, Knox-Davies & Wingfield (1988)
recorded it on 15 Eucalyptus species and many hybrids: it
occurred on most widely planted Eucalyptus species there.
The genus Phaeosepforia Spegazzini (1908), type species
P. papayae Spegazzini, was redescribed by Morgan-Jones
(1974), but so far P. eucalypti has not been compared and
contrasted with it. Such a study is reported in this paper,
together with the results of comparative work on similar
species. Herbarium abbreviations used are those given in
Holmgren, Keuken & Schofield (1981).
Phaeoseptoria papayae Speg.
The genus Phaeoseptoria Spegazzini (1908) was described with
one species, P. papayae Speg., on leaves of Carica papaya L.
from SHo Paulo, Brazil. White, angular to suborbicular, slightly
raised leaf spots, 0.5-3 mm diam. were described, bearing on
their upper surface loosely grouped, dark, glabrous, lenticular
pycnidia, 60-90 pm diam., which contained olivaceous,
continuous to I-heptate, scarcely clavate to somewhat
fusoid conidia, 30 x 3 pm, sharply rounded at each end
(Saccardo & Trotter, 1913). Conidiogenous cells were not
mentioned.
Phaeoseptoria eucalypti and similar fungi on Eucalyptus
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Fig. 1. Phaeoseptoria papayae. Three conidia from the type, LPS
912
said to be produced holoblastically, and no indication of
subsequent proliferation of conidiogenous cells was given. He
commented (in lift. 7 Feb. 1974) that, although phase-contrast
observations indicated holoblastic conidial production, in such
an old specimen and with rather small conidiogenous cells it
was difficult to be absolutely sure.
In view of the scarcity of pycnidia of P. papayae on the type
specimen, it is unlikely that the precise mode of conidiogenesis
of this fungus will be readily clarified until better material of
the species becomes available. What does seem certain from
the observations of both Morgan-Jones (1974) and J.W. is
that large, brown, cylindrical, rough-walled, percurrently
proliferating conidiogenous cells are not present. The fungi on
Eucalyptus dealt with in the present paper, some of which have
in the past been considered as species of Phaeoseptoria, are thus
best removed from this genus. In their prominent percurrently
proliferating conidiogenous cells, they also seem distinct from
several of the other described species of Phaeoseptoria on
Gramineae (Sprague, 1943) and other hosts, e.g. Batista &
Peres (1962), Bubak (1916), Parmelee & Hiratsuka (1970),
Viegas (1945), Wehmeyer (1946). These fungi will need reexamination to define their methods of conidiogenesis and
their relationship to Phaeoseptoria Speg. and they will not be
further here.
12555. Bar 10 urn.
Phaeoseptoria eucalypti Hansford
The type specimen (LPS 12555) has been examined (J.W.).
It consists of five pieces of papaya leaf on which several fungi
are present. The most abundant is Asperisporium caricae (Speg.)
Maubl. (noted on the packet in Spegazzini's writing as
'Cercospora caricae Speg.') causing many circular leaf spots,
1-3 mm diam., with a whitish centre and a thin pale brown
slightly raised margin, often fusing into larger irregular areas.
Abundant Asperisporium conidiomata are present on the lower
surface of the spots. These appear to be the spots attributed
to P. papayae in the original description, but they are quite
characteristic of Asperisporium infection.
O n areas killed by Asperisporium, other fungi are present.
Pycnidia of Phoma sp. (probably P. caricae (Pat.) Punithalingam;
noted on the packet by Spegazzini as 'Phyllosticta caricaepapayae All.') and Coniothyrium sp. were found. One pycnidium
containing conidia agreeing with those described for Phaeoseptoria papayae was seen. Conidia were cylindrical, straight,
slightly curved to sinuous, pale olivaceous brown, smooth,
(1,2)-3-(4)euseptate and 20-30 x 2-2.5 (3) Dm (Fig. 1). They
were usually widest at or below the middle, narrowing
gradually to the rounded apex and with a slightly rounded
truncated base on which no marginal frill was observed. These
agree well with the original description of the conidia.
Conidiogenous cells were not seen in the one pycnidium
found.
Morgan-Jones (1974) has also examined the type specimen
of P. papayae and described and illustrated a pycnidial fungus
with conidia very similar to those above. He also mentioned
conidiogenous cells, describing them as 'small, undifferentiated, hyaline, smooth-walled'. His drawing shows them to
be hardly different from the cells of the inner pycnidial wall
layer, which they resemble in size and shape. Conidia were
Phaeoseptoria eucalypti was described by Hansford (1957) from
living leaves of Eucalyptus grandis Hill ex Maiden from
Sydney, N.S.W., collected about 1955-6 by N. H. White. It
has subsequently been collected on several other Eucalyptus
spp., and leaf symptoms developed depend on the reaction
to infection of the various species. On some hosts, no leaf spot
develops, on others small spots or more extensive necrotic
areas may be formed. Hansford (1957) described P. eucalypti
with subglobose, glabrous, black, embedded pycnidia 150 x
100 urn, containing brown, cylindric-fusoid, smooth-walled,
1-3 transversely septate conidia, borne on simple cylindrical
brown conidiogenous cells, each bearing a single apical
conidium. Walker (1962) examined the type and several other
collections of P. eucalypti, figured both conidia and conidiogenous cells and emended the description. Both the type and
other collections show brown, rough-walled conidia with (I)
3-5 (7) transverse septa, a truncate base with a marginal frill
and rough-walled brown cylindrical conidiogenous cells
showing 1-3 annellations from successive percurrent proliferation.
This fungus is common on Eucalyptus spp. and has
sometimes been found causing severe leaf damage to seedlings
of Eucalyptus spp. in forestry nurseries in N.S.W. As shown in
this paper, it has been described several times with various
epithets, the earliest of which derives from Cercospora
epicoccoides Cooke & Massee apud Cooke (1891).
Cercospora epicoccoides Cooke 6 Massee apud Cooke
Cooke & Massee (in Cooke, 1891) described Cercospora
epicoccoides from leaves of Eucalyptus sp. collected by Mrs
J. Walker, B. C. Sutton and I. G. Pascoe
Fig. 2. Cercospora epicoccoides. Above, four conidia and several percurrently proliferating conidiogenous cells; bar 10 pm. Below, section
of pycriidium; bar 50 um. From the type, in K.
Phaeoseptoria eucalypti and similar fungi on Eucalyptus
Fig. 3. Hendersonia grandispora. Above, three percurrently proliferating conidiogenous cells; bar 10 pm. Below, three conidia; bar
20 wm. From the type, VPRI 5930.
Martin (600) in Victoria. The fungus was reported as
epiphyllous, causing small or confluent spots. ' ~ ~ fwere
t ~ ' Fig 4. Phaeoseptoria luwnensis. Above, percurrently proliferating
described as lgregarious, sphaeriform, rather compact, black, conidiogenous cell; bar 10 wm. Below, three conidia; bar 20 pm.
resembling an Epicoccurn, threads very short, simple'. Conidia From the type, TFM 4869.
were reported as 'profuse, fasciculate, cylindrical, slightly
attenuated upwards, 3-5 septate (50 x 50 p), pale olive'. The to originate from spherical, immersed pycnidia, 115-120 pm
type specimen (K) has been examined by B.C.S. The conidia diam., with walls brown, up to 10 p thick, of 2-4 layers of
and conidial masses on the leaf surface were not attached to brown-walled cells 4-5 pm diam., and a non-protruding
conidiogenous cells or conidiophores. Sections showed them ostiole 10 urn diam. Conidiogenous cells are situated on the
J. Walker, B. C. Sutton and I. G. Pascoe
Fig. 5. Cercospora eucalypti. Above, eight conidia and four small percurrently proliferating conidiogenous cells; bar 10 vm. Below, left
partial section of pycnidium (bar 50 ~ m and
) right, portion of pycnidial wall showing textura epidermoidea (bar 10 ~ m ) From
.
the type,
in K.
Phaeoseptoria eucalypti and similar fungi on Eucalyptus
inner wall layer and are 6.5-11 pm long, 3-5 pm wide, pale
brown with finely roughened walls, proliferating percurrently,
with 1-2 distinct roughened annellations. Conidia are brown,
cylindrical to narrowly obclavate, tapered to the apex, base
truncate with a marginal frill, finely roughened, 1-4 euseptate
and measure 37-50.5 x 5-6 ym (Fig. 2). The fungus is identical
with that seen in the type of Phaeoseptoria eucalypti Hansford.
The first description of this species in the hyphomycete
genus Cercospora when it is in fact a coelomycete, though
strange, is not unusual. P. eucalypti is frequently received for
identification in herb. IMI with provisional identifications such
as 'hyphomycete' or 'sooty mould'. The copious conidial
production which is characteristic of the species results in
conidial masses exuding from the pycnidial ostioles and
spreading over the leaf surface, giving the appearance of a
superficial colony. When sporulation is particularly heavy the
whole of the leaf surface will be blackened, and in such cases
its description as a sooty mould is apt. In several collections,
it has been noticed that the abundant pycnidial ooze will often
push up a small flap of leaf cuticle and epidermis, sometimes
taking with it the top of the pycnidium. This leaves an open
cup-shaped structure with the spore mass on top. It is
interesting that Chupp (1953), in his description of C.
epicoccoides, mentioned a large stroma with dense fascicles of
short conidiophores, which would fit the picture of a ruptured
pycnidium with an exposed hymenium of brown conidiogenous cells.
916
from the innermost cells of the pycnidial wall, brown to olive
brown, long cylindrical, 3-4-septate, 32-60 x 4-5 pm, tapering to the tip and with a truncate base. They were said to be
powdery and black in mass. Conidiogenous cells were not
them as
described in detail, but Koba~ashi(1978) figured
simple, isodiametric, conical to turbinate cells bearing a single
apical conidium. Proliferation was not mentioned or figured.
Conidia were illustrated as smooth-walled. Kobayashi (1978)
mentioned P. eucalypti Hansford on Eucalyptus in Australia and
commented that it is different from P. luzonensis in its
conidiophores (details not given) and pycnidia produced on
the undersurface of leaves, which were not spotted.
Examination of the type specimen of P. luzonensis (TFM
4869) by J. W. has clarified some details of this species, and
shown that it is identical to P. eucalypti. The small embedded
pycnidia occur both on leaf spots and on unspotted areas, and
the abundant conidial ooze has dried to a black flaky coating
on the leaf surface. These features are common in many
collections of P. eucalypti in Australia. Conidia of P. luzonensis
are narrowly obclavate and have a finely roughened wall, and
the truncate base has a small marginal frill (Fig. 4).
Conidiogenous cells are brown, up to 8 pm long and 4 pm
wide, have finely roughened walls, and proliferate percurrently,
with up to three finely roughened annellations (Fig. 4). These
characteristics are identical to those seen in P. eucalypti
Hansford, from which P. luzonensis cannot be distinguished.
Kobayashi & Guzman (1988) examined material from the
Philippines, Thailand, Japan and Australia and concluded that
P. luzonensis could not be distinguished from P. eucalypti.
Hendersonia grandispora McAlp.
McAlpine (1903) described Hendersonia grandispora from
sapling leaves of Eucalyptus sp. collected at Wangaratta,
Victoria in September 1899. The type specimen (VPRI 5930)
examined by J.W. is four leaves showing dark irregular
patches from a few millimetres across to areas covering up to
one-third of the leaf surface. Abundant small embedded
pycnidia, 120-140 ym diam., are present in these areas. The
pycnidial wall is brown, 10-15 ym thick, of 3-4 layers of
brown-walled cells 5-7 pm diam. and is pierced by an apical
ostiole flush with the leaf surface. Conidiogenous cells arise
from the inner wall layer and are 4-10 ym long, 3-4 vm wide,
pale brown with finely roughened walls, proliferating
percurrently, with up to four (perhaps more) distinct,
roughened annellations (Fig. 3). Conidia are brown, narrowly
obclavate, apex narrowly rounded, base flattened with a
distinct marginal frill, finely roughened, 3-5 euseptate and
measure 30-60 x 4-5 ym (Fig. 3). The fungus is identical to
that seen in the type and many other collections of
Phaeoseptoria eucalypti Hansford.
Phaeoseptoria luzonensis T. Kobayashi
Kobayashi (1978) described Phaeoseptoria luzonensis from small,
2-5 mm diam., angular to irregular, brown to greyish brown
leaf spots on Eucalyptus sp., collected on 23 Feb. 1977 at
Agoo, La Union, Luzon, Philippine Islands. Globose black
pycnidia, 75-95 ym diam. and 100-125 ym high, were present
on both surfaces of the spots. Conidia were described arising
All the Eucalyptus fungi described above are conspecific,
with prominent rough-walled percurrently proliferating conidiogenous cells producing brown, rough-walled conidia which
are (1)3-5 (7) septate. The earliest epithet that can be applied
to these collections is based on Cercospora epicoccoides Cooke
& Massee apud Cooke (1891). Another series of Eucalyptus
leaf specimens shows a related fungus, with very pale brown
finely roughened conidia, 1-3-septate, borne on small finely
roughened percurrently proliferating conidiogenous cells.
These collections appear to be congeneric with, but specifically
distinct from, the species based on C. epicoccoides. They are
considered below in chronological order of the names which
have been applied to them.
Cercospora eucalypti Cooke & Massee apud Cooke
Cooke & Massee (in Cooke, 1889) briefly described Cercospora
eucalypti from fading leaves of Eucalyptus sp., collected by Mrs
Martin (436) from Oakleigh, Victoria. Lesions were described
as subcircular or confluent, pale with a rose-coloured margin
bearing a fungus with short hyphae and curved, cylindrical
conidia, with both ends obtuse, barely septate, pale,
30-35 x 4 pm. The type specimen (K) has been examined by
B.C.S. The conidia and conidial masses on the leaf surface
s or conidiophores.
were not attached to ~ o n i d i o ~ e n o ucells
Sections showed them to originate from spherical, immersed
pycnidia, 95-115 ym diam., with walls brown, up to 10 ym
thick, of 2-4 layers of brown-walled cells 3-5 ym diam., and
a non-protruding ostiole up to 10 pm wide. In surface view
J. Walker, B. C. Sutton and I. G. Pascoe
the wi~llsconsist of a textura epidermoidea. The conidiogenous
cells arise from the inner wall layer and are 4 - 5 4 pm
long > 2-3 pm wide, very pale brown with minutely roughened walls, proliferating percurrently, with 1-2 indistinct
annellations. Conidia are very pale brown, cylindrical, straight
or slightly curved, tapered gradually towards the apex, base
truncate, very finely roughened, 0-2-euseptate and measure
25-48 Urn long x 2-3 ym wide (Fig. 5).
This species is distinct from Phaeosepforia eucalypti Hansford
in mo~phologyand gross appearance. Features in common are
the globose thin-walled pycnidia, non-protruding ostioles,
percurrently proliferating roughened conidiogenous cells and
holoblastic, euseptate, brown, rough-walled conidia. Differences ,Ire the texture type of the pycnidial walls and the paler
pigmentation, finer roughening and more delicate structure of
the conidiogenous apparatus and conidia.
Guc~& Liu (1989) transferred the name C. eucalypti to
Pseudoi:ercospora Speg. without comment. The species was
recorded from E. globulus, E. macarthurii and an unnamed
Eucaly17fus sp. from China and it is evident that they did not
examine the type. If they were dealing with a species of
Pseudo,-ercospora,in all probability it was P. eucalyptorum Crous
ef a]. (1989).
was first used in a thesis by Heather (1961), which contained
both Latin and English descriptions of 'Sepforia normae
Heather n.sp.' and mentioned collections from Bull's Head
and Lee's Creek near Canberra on E. dalrympleana Maiden and
E. viminalis Labill. No type specimen was designated, although
it was stated in the thesis that type material had been
deposited in IMI and at the Waite Agricultural Research
Institute, Adelaide. The name Sepforia normae Heather n.sp.
was also mentioned without description in a summary of the
above thesis given by Heather (1962). Neither of the above
constitutes either valid or effective publication of the name
under the International Code of Botanical Nomenclature (1988).
A search at the Waite Agricultural Research Institute has
located one collection (ADW 13559) labelled 'Septoria normae
sp. nov. ined.'. It was collected in the Bull's Head-Lee's Creek
area near Canberra on 20 Aug. 1962 on E. daiympleana by
W. A. Heather and identified by him. It is marked 'Authentic
specimen' and contains a typewritten copy of the Latin and
English descriptions of the fungus from Heather (1961).
Although collected after 1961, this specimen comes from the
locality mentioned in Heather's (1961) thesis and was
determined by him as S. normae. It can thus be considered as
authentic for the unpublished name Sepforia normae Heather.
The collection shows several leaves with large roughly
circular to irregular carmine to brown blotches, 5-10 mm
Septoria pulcherrima Gadgil 6r Dick
diam., often fusing into larger patches occupying most of the
Sepforia pulcherrima was described from leaf spots on E. nifens leaf area. The small, black, embedded pycnidia contain conidia
(Dean€,& Maiden) Maiden collected in New Zealand (Gadgil and conidiogenous cells identical to those described above for
& Dicl;, 1983). Leaf spots were described as first yellowish, Cercospora eucalypti and Septoria pulcherrima. Both Gadgil &
later b-ight carmine and finally brown. The small embedded Dick (1983) and Heather (1961) have commented on the
pycnidia contained hyaline to pale brown, smooth-walled bright carmine colour of the lesions produced on some
Eucalyptus spp. by this fungus. The rose-coloured margin to
slightly tapering conidia 30-60 x 3-4 pm, with (0)-I-(&
transvt-rse septa, and truncate base, borne on simple, hyaline leaf spots mentioned in the original description of C. eucalypti
to pale brown, doliiform to ampulliform conidiogenous cells, indicates a similar symptom.
Other collections in DAR under the name Sepforia normae
up to 5 wm long, arising from the inside of the pycnidial wall.
bore the same fungus. The carmine leaf lesions were
were given.
No details of ~onidio~enesis
Authentic material of the species has been studied by Swart particularly prominent on E. dalympleana, a host examined by
(1988), and he described two modes of conidiogenesis. The both Gadgil & Dick (1983) and Heather (1961), but the colour
predominant conidiogenous cells produced several (up to was much less prominent on a collection of E. viminalis from
three are figured) percurrent proliferations. A few conidio- Tasmania and absent from the brown leaf lesions on an
genous cells proliferated sympodially and bore one apical undetermined host species from Queensland. Similar variation
holoblastic conidium at each proliferation. Swart (1988) in symptoms on different Eucalyptus spp. produced by the
considered that S. pulcherrima was closely related to, and same fungus has been seen with Phaeoseptoria eucalypti.
The fungus referred to as Septoria normae is thus identical
congeneric with Stagonospora delegatensis Park & Keane (1984)
and that it was better placed in the genus Stagonospora (Sacc.) with the earlier-described Cercospora eucalypti.
Sacc. Me transferred it as Sfagonospora pulcherrima (Gadgil &
Dick) Swart. However, he commented that much revisionary
Septoria ceuthosporoides (Cooke 6 Harkness) Sacc.
work on these fungi was still needed.
From the original description and the emended account by
Cooke & Harkness (1881) described Cryptosporium ceufhoSwart (1988), S. pulcherrima is seen to be very similar to sporoides from dead leaves of Eucalyptus sp. collected by H. W.
Cercospora eucalypti. The authentic material of S. pulcherrima Harkness (2005) from California. The fruiting structures were
seen by Swart and several other collections have been described as flattened, brown, ceuthosporoid, then fissuring
examin2d by I.P. and found to be identical with C. eucalypti. above. Spores were reported as curved, hyaline, 0.0180.02 x 0.003 mm. The name was transferred as Septoria
ceuthosporoides (Cooke & Harkness) Saccardo (1884). The type
Septoria normae Heather ined.
specimen (K) has been examined by B. C. S. The conidiomata
In Australia, the name Sepforia normae Heather has been used are wrinkled, dark brown, unilocular, thick-walled, 180for a Eucalyptus leaf fungus similar to P. eucalypti but having 195 pm diam., with a thinner lower wall 2-3 cells thick and a
smaller, paler and more finely roughened conidia. The name . thicker, darker brown upper wall 3-5 cells thick. No distinct
918
Phaeosepforia eucalypti and similar fungi on Eucalyptus
ostiole is present, dehiscence occurring by radiating fissures.
Conidiophores are filiform, hyaline, smooth, septate, occasionally branched, with apical and lateral phialidic conidiogenous
loci, 10-15 pm long x 2 pm wide. Conidia are hyaline,
aseptate, falcate, 13-15 pm long, 2 pm wide. This species is
identical with Pilidium acerinum Kunze and the names have
therefore been added to the synonymy given by Sutton
(1991).
Septoria mortolensis Penz. 6 Sacc.
Penzig & Saccardo (in Saccardo, 1884) described Sepforia
rnortolensis from fallen leaves of E. amygdaloides and Acacia sp.
and from living leaves of Hedera algeriensis from north Italy
(Funghi Morfolenses no. 27). Lesions were described as absent,
and pycnidia as loosely gregarious, numerous, large, hypogenous, eventually erumpent, 150-200 pm diam., spherical,
depressed, with a somewhat prominent ostiole, parenchymatous, black, carbonaceous, and conidia which were long
cylindrical, straight, with both ends obtuse, aseptate or
spuriously 1-2-septate, filled with granular contents, 5055 x 3-3.5 Urn, and short, hyaline, simple conidiogenous cells.
The species was stated to be close to S. sicula Penz., which is
known from Citrus limonum. The type is reported to be in
herb. PAD (Gola, 1930), but the Curator (in lift. 1989) stated
that it was not filed in the collection. The Curator, Herb.
ROPV (in litf. 1991) reported that no collection of S.
mortolensis was filed in that herbarium and that, although
Penzig's herbarium of Citrus fungi is present in ROPV, the
general Penzig herbarium seems to have been destroyed
during World War 11. No information about S. mortolensis
could be obtained from herb. GE, where types of some Penzig
fungal collections are reported to be present (Vegter, 1983).
Since the species was described originally from three
separate host substrata, a lectotype must be selected before
firm conclusions can be made about the identity of the taxon.
It is possible that the fungus on E. amygdaloides is the same as
that which has been called Cercospora eucalypti, but there are
differences. The pigmentation of conidia was not mentioned
in the original account, but the number of septa and the length
and breadth measurements are consistent with this species.
However, conidia were also reported to be rotund (obtuse) at
each end, and this is not typical. Conidia in C. eucalypti are
tapered gradually towards the apices. In such circumstances, it
is not possible to state with any degree of certainty that S.
rnortolensis is the same as C. eucalypfi.
Septoria eucalypti Winter 6 Roum.
Winter & Roumeguilre (1887) described S. eucalypfi from
almost dead ('subvivis') leaves of E. amygdalina from Bone,
Algeria. Pycnidia were described as more or less gregarious,
amphigenous, sparse, globose to irregular, often formed
subangularly, sometimes irregularly concentric, without lesions, immersed, subglobose, only slightly attenuated in the
vertical axis, later opening widely, erumpent, black, membranaceous, ca 100 pm wide. Conidia filiform-acicular, not
very flexuous, both ends acute, chlorinous-hyaline, in the
middle finely septate, 14-18 pm long, 1.5 pm wide. The type
collection has not been examined but the conidial size alone,
especially the width, precludes this species from consideration
in the context of the particular species on Eucalyptus under
consideration.
These studies have shown that, on Eucalypfw leaves, there
are two related congeneric taxa which form pycnidia
containing brown, rough-walled, percurrently proliferating
conidiogenous cells producing brown, rough-walled, euseptate, long, narrowly obclavate to cylindrical conidia. They
differ in colour, degree of roughening and robustness of the
conidiogenous cells and conidia. Both have been placed in
various genera and a more detailed consideration of their
generic position is required.
GENERIC CONSIDERATIONS
Saccardo & Trotter (1913) categorized the genus Phaeosepforia
Spegazzini as 'est quasi Septoria sporulis olivaceis distincta'
and many modern authors have followed this lead, regarding
it merely as a coloured-spored Sepforia, e.g. Barnett & Hunter
(1972), Parmelee & Hiratsuka (1970), Roger (1953), Sprague
(1943) and Wehmeyer (1946). Hohnel (1924) regarded
Phaeosepforia as a synonym of the earlier-described Scoleciasis
Fautrey & Roumeguilre in Roumeguilre (1889), type species S.
aquatica Fautrey & Roum. on Scirpus. He described this species
with hyaline to pale brown conidia of very variable length,
with from 3 to several transverse septa, a wide, blunt base and
borne on long, thin (8-24 x 2 pm) conidiogenous cells. Sutton
(1977) mentioned Scoleciasis but did not give diagnostic
features, and it is not included in his later treatment of the
Coelomycetes (Sutton, 1980). A potential isotype for this
name (K) has been examined by B.C.S., but no fungus
corresponding to the original account was found on it. The
brown conidia, originally described as 120-160 x 4 pm, are
extremely long for a pycnidial species, and no fungus with
conidia even approaching this size range could be seen on the
specimen. This is in contrast to Hohnel (1924), who found
three spore forms in the material he examined (presumably
another potential isotype of the same exsiccatus -C.
RoumeguGre, Fungi selecti exsiccafi 5072). The long-spored
fungus he considered to be identical with Sepforia dolichospora
Trail, the medium-spored species identical with S. lacusfris
Sacc. & Thiim., and the small-spored species a variant within
the Septoria scirpi Sacc., S. scirpoides Pass., S. holoschoeni (Mont.)
Pass., S. nervispora Sacc. complex. With its long, thin
conidiogenous cells, it is unlikely that Scoleciasis would
provide a suitable place for the Eucalyptus leaf fungi considered
here, but further work to characterize the genus is needed,
especially since potential isotypes seem to bear several fungi.
Petrak (1941) considered fungi placed in Phaeoseptoria as
scolecospored forms of Hendersonia Berk. He did not study the
generic type specimen. From the description and from a study
of other species placed by other authors in Phaeosepforia, he
concluded that Phaeoseptoria could not be considered a
separate genus in its own right, but graded into fungi placed
in Hendersonia. The name Hendersonia Berk. is now a nomen
rejiciendum in favour of Stagonospora (Sacc.) Sacc. (Sutton,
1977), and several species on Eucalyptus placed formerly in
Hendersonia have been revised by Swart & Walker (1988). The
J. Walker, B. C. Sutton and I. G. Pascoe
Phaeoseptoria-like fungi on Eucalypfus are close to but distinct
from :hese species which have been placed in Sonderhenia
Swart & Walker, but the latter have distoseptate conidia bome
on smaller, smooth-walled annellidic conidiogenous cells.
Stagonospora is also unsuitable for these Eucalypfus fungi
because it has smooth, hyaline conidia bome on holoblastic,
occasionally annellidic, doliiform, smooth-walled conidiogenous cells. In the absence of any suitable genus, it is
necessary to introduce a new generic name for these fungi.
Kirrarnyces J. Walker, B. Sutton & I.Pascoe, gen. nov.
Etym.: kirra, folium, in lingua una aboriginum Australiae et
myces (Gk), fungus
M y c e l i ~ w irnrnersurn. Conidiomata pycnidialia, irnrnersa, brunnea,
solitaria, unilocularia; parietes 2-5 cellulis crassitunicati, ex textura
angulari vel epidermoidea brunnea cornpositi; ostiolurn central,
circulars:, non papillaturn. Conidiophora absentia. Cellulae conidiogenae
discretae, arnpulliforrnes, lageniformes vel brevi-cylindricae, bnmneae,
verruculosae, uno vel aliquot proliferationibus enteroblasticis, percurrentibus, venuculosis vel laevibus, ex cellulis interioribus parieturn
pycnidiorurn forrnatae. Conidia holoblastica, euseptata, cylindrica vel
longi-ol~clavata, apicem obtusurn versus leniter deminuta, basirn
truncata cum segrnenti rnargine, verruculosa.
Typus generis : K. epicoccoides (Cooke & Massee) J. Walker, B. Sutton &
I. Pascoe
Mycelii-tm immersed. Conidiomafa pycnidial, immersed, brown,
solitary, unilocular; wall 2-5 cells thick, of brown textura
angulaiis to epidermoidea; ostiole central, circular, not
papillale. Conidiophores absent. Conidiogenous cells discrete,
ampulhform, lageniform or short cylindrical, brown, verruculose, with I-several proliferations, formed from the inner
cells of the pycnidial wall. Conidia holoblastic, euseptate,
cylindr~calto long obclavate, tapered t o obtuse apices, bases
truncatl with a marginal frill, verruculose.
Kirramyces epicoccoides (Cooke & Massee) J. Walker, B.
Sutton & I. Pascoe, comb. nov.
Cercosp2ra epicoccoides Cooke & Massee apud Cooke, Grevillea
19 : 9 1 (1891).
Hendersonia grandispora McAlp., Proc. Linn. Soc. N.S. W. 28 : 9 9
(15103).
Phaeosejltoria eucalypti Hansf., Proc. Linn. Soc. N.S.W. 82 : 225
(15'5 7).
Phaeoseptoria luzonensis T. Kobayashi, Trans. Mycol. Soc. lapan
19 : 377 (1978).
In Australia, this is a common leaf spot fungus on several
species of Eucalyptus in eastern states and can cause severe
damage t o seedlings of some species when cultivated
intensively in nurseries. There are n o records of this species
under m y of its synonyms in recent check lists of plant
diseases for South Australia (Cook & Dubd, 1989) and
Western Australia (Shivas, 1989). It is also recorded from
several ']verseas countries. The Eucalyptus species on which K.
epicoccoides has been reported (based on specimens) and the
localities from which the records came are summarized in
Table 1. Of the 25 named species of Eucalyptus, 2 1 fall into the
919
Table 1. Named species of Eucalyptw reported as hosts of Kirramyces
epicoccoides, their subgeneric classification and their distribution, based on
specimens
Host
Subgenus' Distribution
Eucalyptus amplifolia
E camaldulensis
S
S
N.S.W. (DAR 41900)
Q. (BRIP 5442) (Alcom, 1976),
E. citriodora
E. nebra
E. dealbata
E. delegatensis
E. drepanophylla
E. exserta
E. globulw
C
Q. (BRIP 5443)'
Q. (BRIP 5444) (Alcorn, 1976)
Q. (BRIP 5445) (Alcorn, 1976)
Tas. (DAR 24899 b)
E. globulus ssp. bicostata
S
S
Vic. (VPRI 15150)""
E. grandis
E. longifolia
E. macarthurii
E. maculata
E. major
E. microcoys
E. platypus
E. punctata
E. radiata ssp. robertsonii
E. resinifera
E. robusta
E. saligna
S
S
M
S
S
S
S
S
C
S
Q. (BRIP 5446) (Alcom, 1976)
Q. (IMI 161742b)
N.S.W. (DAR 5833a),
Q. (IMI 161745)"
N.S.W. (DAR 32409)
N.S.W. (DAR 41899).
Q. (BRIP 5463) (Alcom, 1976)*
N.S.W. (DAR 6476)
N.S.W. (DAR 6339)
N.S.W. (DAR 6338)
Q. (BRIP 5451) (Alcom, 1976)
S
S
S
N.S.W. (DAR 13879)
Q. (BRIP 5452) (Alcom, 1976)
M
Vic. (Chambers, 1982)
S
S
S
E.
sideroxylon
S
E.
tereticornis
E. urophylla
S
S
E. viminalis
S
N.S.W. (DAR 38821,
Q. (BRIP 5454) (Alcom, 1976)'
N.S.W. (DAR 6340).
Q. (BRIP 5455)
Q. (BRIP 5457) (Alcom, 1976)"
Vic. (VPRI 2418)
" Subgeneric classification from Pryor &Johnson(1971).C, Corymbia; M,
Monocalyptus; S, Symphyomyrtus.
" An asterisk (*) indicates that extra-Australian specimens have also been
seen (see Specimens examined).
subgenus Symphyomyrtus of Pryor & Johnson (1971), with 2
in the subgenus Corymbia and 2 in the subgenus Monocalyptus. Further collecting could well extend both the host
and geographic ranges of this species.
Spec~mens examined: locations are in Australia unless otherwise
mentioned. On Eucalyptw amplifolia Naudin, Chipping Norton,
N.S.W., 28 Apr. 1982, C. Whitternore, DAR 41990. On E.
camaldulensis Dehnh., Ivanhoe, Victoria, 25 Oct. 1986, H.-Y. Yip,
VPRI 15150, 15852; Bahia, Brazil, 24 Sept. 1983, C. S. Hodges, IMI
283115; Formosa Prov., Argentina, 16 July 1985, A. Rornero, IMI
296606 (as E. rostrata Schlecht.);on E. citriodora Hook., Zambia, 1986,
N. Sikornbwa, IMI 311080; Queensland, 1974, J. L. Alcom, BRIP
5443. On E. delegatensis R. T . Baker, at forestry camp, Deloraine,
Tasmania, 27 Mar. 1974, C. Palzer GI, DAR 24899b. On E. exserta
F. Muell., Kenmore, Queensland, 15 Aug. 1971, J. L. Alcom, IMI
161742 b. On E. globulus Labill., Balgowlah, N.S.W., 21 Mar. 1960,
L. R. Fraser, DAR 5833a (originally as Cercospora epicoccoides);
Mooloolabah, Queensland, 11 Oct. 1965, A. Tange 16755, IMI
161745; Kamataka, India, Oct. 1975, K. M. Ponnappa 5, IMI 198912;
Gora, Ethiopia, Jan. 1977, Guillaurna, IMI 210737~; Gumuro,
Ethiopia, 3 Dec. 1976, E. Nlernann 2079, IMI 225573; Jabalpur,India,
Nov. 1986, N. D. Sharma, IMI 311973; Irnlikhera, 23 Nov. 1990,
M. K. Rai 14, IMI 337563. On E. globullls doubtfully (host given
as E. globulosw St.-Lag.), Calcutta, India, 24 Jan. 1979, B. Das PCC
Phaeoseptoria eucalypti and similar fungi on Eucalyptus
Table 2. Named species of Eucalyptus reported as hosts of Kirramyces
eucalypti, their subgeneric classification and their distribution based on
specimens
Host
Subgenus,, Distribution
Eucalyptus cephalocarpa
S
E. dalrympleana
S
E. gardneri
E. globulrrs
S
S
E. goniantha
E. glrnnii
S
S
E. melliodora
E. nitens
E. nutans
E. ouaia
E. platypus
E. rubida
E. vitninalis
S
S
S
S
S
S
S
New Zealand (NZFRI 101)
(Gadgil & Dick, 1983)
N.S.W. (DAR 12841).
New Zealand (NZFRI 103)
Q. (BRIP 13248)
New Zealand (NZFRI 104)
(Gadgil & Dick, 1983)
Q. (BRIP 13249)
New Zealand (NZFRI 105)
(Gadgil & Dick, 1983)
Q. (BRIP 11345)
New Zealand (NZFRI 2524)
Q. (BRIP 5464)
New Zealand (NZFRI 106)
Q. (BRIP 5465)
Vic. (DAR 65646)
Tas. (DAR 30156).
Vic. (VPRI 2419)
" Subgeneric classification from Pryor & Johnson (1971).C, Corymbia; M,
Monocalyptus; S, Symphyomyrtus.
2841, IMI 242968; West Bengal, India, 1987, A. K. Kar 6011,
IMI 316948. O n E. globulus ssp. bicostata (Maiden, Blakely & J. Simm.)
Kirkpatr., West Ryde, N.S.W., 23 June 1978, S. Stanton, DAR
32409 b. On E. grandis W. Hill ex Maiden, Sydney, N.S.W., about
1955, N. H. White, ADW 7137 (holotype of Phaeosepforia eucalypti
Hansford, slides as DAR 44448); Mullumbimby, N.S.W., 31 May
1982, K. Richens, DAR 41899; Jiggi, near Lismore, N.S.W., 8 Aug.
1988, P. Faulk, DAR 63318b; Natal, South Africa, 5 May 1988,
M. J. Wingfield, IM1 325555; E. Transvaal, South Africa, 11 Apr.
1988, M. J. Wingfield, IMI 323975. O n E. longifolia Link, West
Pennant Hills, N.S.W., 18 Nov. 1960, J. Walker, DAR 6476. On E.
macarthurii Deane & Maiden, West Pennant Hills, 30 June 1960, J.
Walker, DAR 6339. O n E. maculata Hook., West Pennant Hills, 30
June 1960, J. Walker, DAR 6338. On E. microcoys F. Muell., Zomba,
Malawi, 29 Apr. 1981, N. Chipompha PCNIO, IMI 257674. On E.
piafypw Hook., Greenwich, N.S.W., 23 Mar. 1965, Mr Baltpurvins,
DAR 13879. O n E. resinifera Smith, Mazabuka, Zambia, 4 Apr. 1982,
G. D. Piearce FP. 117/A, IMI 267755. On E. robusta Smith, Dai Lung,
Hong Kong, 3 Feb. 1966, R. I. Leather CB43, IMI 118467; Dai Lung,
Hong Kong, 29 June 1966, R. I. Leather, IMI 121997a. On E. saligna
Smith, Canberra, A.C.T., June 1959, W. Stahl, ADW 8633 (duplicate
as DAR 6337); Glenbervie State Forest, New Zealand, 13 Apr. 1983,
D. Bartram, NZFRI 2457 (duplicate as VPRI 15501); North Plymouth,
New Zealand, 31 May 1983, M. Stoodley, NZFRI 2458 (duplicate as
VPRI 15500); Gora, Ethiopia, Feb. 1977, L. S. Foster, IMI 211589.
O n E. sideroxylon A. Cunn. ex Woolls, West Pennant Hills, N.S.W., 30
June 1960, J. Walker, DAR 6340; Dai Lung, Hong Kong, 3 Feb. 1966,
R. I. Leather CB45, IMI 118469. O n E. tereticornis Smith, Kerala, India,
13 Mar. 1980, J. K. Shama 15, IMI 246483; Hyderabad, India, Feb.
1981, B. S. Reddy SFE-I, IMI 262292. O n E. urophylla S. T. Blake,
Bahia, Brazil, 25 Sept. 1983, C. S. Hodges, IMI 283118. O n E.
viminalis Labill., Victoria, no date, Mrs Martin, VPRI 2418 (perhaps
isotype of Cercospora eprcoccoides Cooke & Massee). O n Eucalyptus
hybrid, Choma, Zambia, 30 Sept. 1986, N. Sikombwa FP809/2, IMI
311075. O n Eucalyptus spp. indet., Victoria, before 1891, Mrs Martin
600, K (holotype of Cercospora epicoccoides); Wangaratta, Victoria,
Sept. 1899, D. McAlpine, VPRI 5930 (holotype of Hendersonia
grandispora McAlp.; slides as DAR 59811); Agoo, La Union, Luzon,
920
Philippines, 23 Feb. 1977, T. Kobayashi, TFM 4869 (holotype of
Phaeoseptoria luwnensis T. Kobayashi; slides as DAR 37804); East
Parramatta, N.S.W., 27 Aug. 1976, J. Walker 761126, DAR 27983;
Bangalore, India, 31 Jan. 1973, B. C. Sutton 85, IMI 174780;
Maymyo, Burma, 20 Jan. 1974, M. Thaung 25, IMI 185241;
Bangalore, India, 4 Apr. 1978, B. A. Ullasa 2, IMI 226824; Assam,
India, 9 May 1980, M. D. Mehrotra LA28, IMI 248469; Jabalpur,
India, 24 Dec. 1981, R. C. Rajak I, IMI 264372; Uttar Pradesh, India,
1 Jan. 1984, P. Narayan PN64, IMI 282875; Nainital, India, 12 Nov.
1984, P. Narayan PN166, IMI 291635; Bhubaneswar, India, 11 Jan.
1985, Narain OUAPPl9, IMI 293321; Bhutan, 21 Feb. 1985, W.
Peregrine SIB 223, IMI 294224; Ugunga Island, Tanzania, Mar. 1985,
J. Waller W2311, IMI 297200; Kath Godam, India, 31 Oct. 1985,
Verma KK117, IMI 299170; Sagar, India, 30 Nov. 1988, P. Chauksey
RC70, IMI 332512; Uttar Pradesh, India, Aug. 1987, S. Chauhan 6,
IMI 332670; Madhya Pradesh, India, Dec., S. A. Firdausi FV-54, IMI
332674.
Kirramyces eucalypti (Cooke & Massee) J. Walker, B. Sutton
& I. Pascoe, comb. nov.
Cercospora eucalypti Cooke & Massee apud Cooke, Grevillea
18: 7 (1889).
Pseudocercospora eucalypti (Cooke & Massee) Guo & Liu,
Mycosystema 2 : 234 (1989).
Septoria pulcherrima Gadgil & Dick, N e w Zealand Journal of
Botany 21: 49 (1983).
Stagonospora pulcherrima (Gadgil & Dick) Swart, Trans. Brit.
Mycol. soc. 9 0 : 285 (1988).
The names Septoria n o m a e Heather (1961) ined. and
Phaeoseptoria sp. have been used commonly in eastern Australia
for this species. It is not as well known as K. epicoccoides but
has now been recognized on 13 species of Eucalyptw (Table 2),
all in the subgenus Symphyomyrtus. Like K. epicoccoides, it can
cause severe leaf damage, but with both species much remains
to be learned of the pattern of susceptibility and symptom
expression across the genus Eucalyptus.
Specimens examined: locations are in Australia unless otherwise
mentioned. On Eucalyptw dalympleana Maiden, Bull's Head-Lee's
Creek area, near Canberra, A.C.T., 20 Aug. 1962, W. A. Heather,
ADW 13559 (authentic for the unpublished name Septoria nomae
Heather, dupl. as DAR 65724); Canberra, 28 Feb. 1964, W. Stahl,
DAR 12841 (as S. normae, dupl. as VPRI 15453); Rotoehu State
Forest, New Zealand, 26 Mar. 1982, A. Zandvoort, NZFRI 103 (as
Septoria pulcherrima Gadgil & Dick, dupl. as VPRI 15504). O n E.
gardneri Maiden, Lawes, Queensland, Jan. 1981. A. H. Wearing, BRIP
13248 (as Phaeoseptoria sp.). O n E. goniantha Turcz., Lawes,
Queensland, Jan. 1981, A. H. Wearing, BRIP 13249 (as Phaeosepforia
sp.). O n E. melliodora Cunn. ex Schauer, Wynyabbie Nursery,
Jindalee, Queensland, 9 Dec. 1971, J. L. Alcom, BRIP 11345. O n E.
nitens (Deane & Maiden Maiden), Tawhero Road, New Zealand Forest
Products Ltd, Tokoroa, New Zealand, 13 Mar. 1981, R. M. J.
MacKenzie, NZFRI 2524 (authentic for the name Septoria pulcherrima
Gadgil & Dick; duplicate as VPRI 15503). O n E. nutans F. Muell.,
Everton Park, Queensland, 17 Oct. 1973, F. D. Hockings, BRIP 5464;
Dalby, Queensland, 20 Feb. 1985, D. Cummins, BRIP 14589. O n E.
ovata Labill., Kinleith, New Zealand, 6 May 1981, R. M. J. MacKenzie,
NZFRI 106 (as 5. pulcherrima, dupl. as VPRI 15502). O n E. platypus
Hook., Dalby Forestry Nursery, Queensland, 26 July 1972, L.
Bolland, BRIP 5465. On E. rubida H. Deane & Maiden, Errinundra
Plateau, Victoria, 6 Sept. 1982, R. Park, DAR 65646. O n E. viminalis
Labill., Pittwater, Tasmania, June 1977, G. A. Kile, DAR 30156;
J. Walker, B. C. Sutton and I. G. Pascoe
Fig. 6 . Kirramyces lilianiae. Left, four conidia; bar 20 urn. Right,
percurreitly proliferating conidiogenous cells; bar 10 urn. From the
type, DAR 3832.
Victoria, 1900, Mrs Martin, VPRI 2419. On E. spp. undetermined,
Oakleigl~,Victoria, before 1889, Mrs Martin 436, K (holotype of
Cercospo~.a eucalypti); Eden, N.S.W., April 1975, W. Stahl, DAR
25300, i.5306; Macalister, Queensland, Feb. 1970, B. Brown, DAR
19581; Jindalee, Queensland, 9 Dec. 1971, 1. L. Alcom, BRIP 8734
(dupls a: BRIP 2574 and IMI 167047).
In orrler to avoid confusion in use of the above names, it
should be mentioned that the fungus formerly known as
Phaeose~lforia eucalypti Hansford is now named Kirramyces
epicoccoides. The name Kirramyces eucalypti (whose epithet
derives from Cooke & Massee, not Hansford) refers to a
differenl- fungus.
Other species of Kirramyces
Several other collections of Eucalypfus leaf spots examined
have associated with them a variety of fungi showing many
features in common with those described above. Some of
these deserve special mention.
Kirrarnl~ceslilianiae J. Walker, B. Sutton & I. Pascoe, sp.
no\.
(Fig. 6)
92 1
which occurs naturally only in the coast and adjacent tableland
districts of New South Wales about 200 krn north and south
of Sydney, mainly on poorer sandstone soils (Chippendale,
1988). The two collections come from locations about 10 krn
apart. They are similar but differ slightly in that the conidia in
DAR 3832 are more finely roughened than those in DAR
3833, which show more prominent verruculations. A similar
variation in wall roughness is seen in a range of collections of
K. epicoccoides, to which K. lilianiae appears closely related.
Both have conidia that are a quite definite brown in colour and
obviously roughened, in contrast to the paler more finely
roughened conidia of K eucalypti. The pycnidia and conidiogenous cells of K. epicoccoides and K. lilianiae are virtually
identical. The two species differ only in their conidia, which
are cylindrical with up to 3 eusepta in K. lilianiae and narrowly
obclavate with up to 7 eusepta in K. epicoccoides. Conidia of K.
lilianiae are generally slightly wider than those of K. epicoccoides.
To date, the only known host of K. lilianiae is E. eximia, a
member of the subgenus Corymbia of Pryor & Johnson
(1971). Most of the hosts shown in Table 1 for K. epicoccoides
and in the list of specimens examined are placed in subgenus
S y m p h y ~ m ~ r t ubut
s , three collections on subgenus Corymbia
have been seen. These are on E. maculafa (DAR 6338) and E.
citriodora (Alcorn, 1976; BRIP 5443) from Australia and on E.
citriodora from Zambia (IMI 311080). All have been examined
in detail and confirmed as K. epicoccoides. However, DAR 6338
is a collection on seedlings from a forestry nursery where an
epidemic of K. epicoccoides was damaging seedlings of several
species, mainly of the subgenus Symphyomyrtus. Whether
the seedling reaction to infection under these conditions of
overwhelming inoculum density gives a true indication of the
innate reaction of this host species to infection in nature is
open to question. BRIP 5443 was from a sapling in a home
garden. Similar details for the Zambian collection are not
available. Savile (1979) has indicated the need for care in cases
such as this when using parasites as indicators of host
relationships. In this context it is interesting that to date there
is no field collection in Australia of K. epicoccoides on E.
maculafa or E. citriodom. The only Field collection on E.
maculafa available (DAR 22092 b) is of a pale small-spored
species more closely related to K. eucalypti. Many more field
collections are needed before the true host ranges of these
fungi and their relationship to the subgeneric classification of
the genus Eucalyptus are known.
Etym. Lilian Ross Fraser, beata; collector et fungorum discipula
Conidiom,zta et cellulae conidiogenae Kirramyceti epicoccoidi similes.
Conidia c:qlindrica, brunnea, venuculosa, 1-3 euseptata, 35-50 x 4-6
(7) um.
Habitat n~aculisin foliis Eucalypti eximiae Schauer, Wiseman's Ferry,
Nova W.dlia Australis, Australia, mensis Julius 1949, L. R. Fraser,
DAR 3832 (Holotypus);Maroota, Nova Wallia Australis, Sept. 1949,
L. R. Fraser, DAR 3833.
Conid~omata and conidiogenous cells similar to those of
Kirramyces epicoccoides. Conidia cylindrical, brown, verruculose,
1-3 euseptate, 35-50
X
4-6 (7) pm.
Kirranryces lilianiae is known to date only from the above
two collections on yellow bloodwood, Eucalyptus eximia,
Ofher possible Kirramyces faxa
A smaller number of other collections bear similar fungi, but
these require further study and more extensive collections
before their relationships to the above species can be
determined. Most are pale-spored species resembling
K. eucalypti but differing from this species in conidial shape and
wall roughening. Three collections (DAR 19894 on Eucalyptus
sp., BRIP 13623 on E. ptychocarpa F. Muell. and BRIP 13059
on E. fiifolia F. Muell.) show a variable fungus with very pale
finely roughened cylindrical conidia, which could be a palespored analogue of the darker-spored K. lilianiae. The known
Eucalyptus host both of this fungus and of K. lilianiae are in the
subgenus Corymbia.
Phaeoseptoria eucalypti and similar fungi on Eucalyptus
922
(Swart, 1982, 1986, 1988; Swart & Walker, 1988; Swart &
Williamson, 1983) have led to the discovery of many
reassessed. This species is known only from the type collection previously unrecognized Coelomycete taxa. Some of these
on E. delegatensis R. Baker, on which it causes circular grey leaf have been placed in new genera such as Dauisoniella Swart
spots, 5-10 mm diam., with a darker margin. It is the pycnidial (1988), Macrohilum Swart (1988), Sonderhenia Swart & J.
anamorph of Mycosphaerella delegatensis Park & Keane (1984). Walker (1988) and Vemisporium Swart & Williamson (1983),
Conidia were described as cylindrical, straight, hyaline, which so far are known only from Eucalyptus, whilst others
smooth-walled, 21-51 x 3-5 ym, with a truncate base on represent undescribed species in genera with a wider host
which a minute basal frill was figured. Conidia were borne on range, for example Leptomelanconium ausfraliense Sutton (1974)
holoblastic, annellidic, hyaline conidiogenous cells up to and the various Coniothyrium species on Eucalyptus (Sutton,
31 x 13 ym. Swart (1988) compared S. delegatensis and Septoria 1975, 1980; Swart, 1986).
In the present work, the new genus Kirramyces has been
pulcherrima Gadgil & Dick and regarded them as congeneric.
Septoria pulcherrima is here reduced to synonymy under established for a group of taxa centred on the fungus most
commonly known as Phaeoseptoria eucalypti Hansford (1957).
Kirramyces eucalypti.
Examination of the holotype of Stagonospora delegatensis With their combination of pycnidial conidiomata, brown,
(DAR 45718b) generally confirms the above description. euseptate, cylindrical to narrowly obclavate rough-walled
However, conidia are faintly tinted yellowish brown rather conidia and brown roughened annellidic conidiogenous cells,
than hyaline and the conidiogenous cells are smaller (up to these taxa do not fall readily into any known genus. The
15 x 4-5 vm) than originally described, and very similar to collections placed here in Kirramyces form two series - those
the illustration in Swart (1988). This species is very similar to with brown, rough-walled conidia and conidiogenous cells (K.
K. eucalypti in spore size, but differs in having paler, slightly epicoccoides and K. lilianiae) and those with paler, more
less tapered conidia, which are apparently smooth. Whilst it is delicately roughened conidia and conidiogenous cells (K.
not correctly placed in Stagonospora, its correct taxonomic eucalypti and a group of at present ill-defined taxa represented
position and relationship to the pale-spored Kirramyces taxa by only a few collections). Of the three named species, K.
lilianiae is known only on E. eximia (subgenus Corymbia),
must await study of further collections from E. delegatensis.
The above collections show considerable intergradation in which has not been reported as a host of either of the other
conidial shape, size, colour and degree of wall roughening. two species. All known collections of K. eucalypti occur on
Conidiogenous cells are also generally smaller than in the hosts in the subgenus Symphyomyrtus (13 species) and, with
three species of Kirramyces recognized here, and precise details the exception of E. citriodora and E. maculata (subgenus
of conidiogenesis remain to be determined. Resolution of the Corymbia) and E. delegatensis and E. radiata (subgenus
taxonomic problems they pose must await further work and Monocalyptus), all other hosts of K. epicoccoides also fall into
especially a larger number of collections from a range of subgenus Symphyomyrtus (21 species) (Pryor & Johnson,
locations and Eucalyptus species. A division of available 1971). Three species, E. globulus, E. platypus and E. viminalis,
collections into two tentative taxa is given in the list of are hosts to both K. epicoccoides and K. eucalypti (Tables 1, 2).
The genus Kirramyces as defined here would thus seem to
specimens examined.
show a decided preference for hosts in the subgenus
Specimens examined: Kirramyces tax. sp. 1 (pale cylindrical conidia): on
Symphyomyrtus, although it is expected that examination of
Eucalypfw ficifolia F. Muell., Toowoomba, Queensland, 22 Jan. 1980,
further collections will reveal further species like K. lilianiae on
W. Mills, BRIP 13059. On E. pfychocarpa F. Muell., Mitchelton,
other
subgenera.
Queensland, 23 Mar. 1982, E. G. Phillips, BRIP 13623. On E. sp.,
An
analysis of the subgeneric distribution within Eucalypfus
Bouddi National Park, N.S.W., 5 Feb. 1967, 0.M. Williams 67/62,
of the leaf-attacking Coelomycetes (and other pathogens)
DAR 19894~.
Kirramyces tax. sp. 2 (pale short obclavate conidia): on E. maculata, would be of considerable interest, but the sample sizes for
10 krn S of Naroorna, N.S.W., 9 Dec. 1971, J. Walker, DAR 22092 b. each pathogen are at present too small to draw definite
Stagonospora delegatensis Park & Keane; on E. deiegafensis, conclusions. Care should also be taken to allow for the soNarbethong, Victoria, 28 June 1983, R. F. Park, DAR 45718b called adventitious infections that may result on a normally
(holotype).
non-host species in nature if it is exposed to large doses of
inoculum or growing under artificial conditions (in a
glasshouse, nursery or in cultivation). In nurseries, seedlings of
DISCUSSION
Eucalyptus spp. are often damaged by conidial powdery
The genus Eucalyptus is host to a wide range of leaf-attacking mildews (Oidium spp.), but powdery mildew has not been
fungi, and amongst these are many Coelomycetes with brown found on Eucalyptus in the field in Australia. Savile (1979) has
conidia borne on percurrently proliferating conidiogenous stressed that listing plants with adventitious infections as
cells. In the past, some of these have been placed in classical hosts may give a false suggestion of relationships when using
genera such as Coniothyrium Corda, Diplodia Fr., Hendersonia parasites as aids in the study of higher plant classification.
auctt., Phaeoseptoria Speg. and Septoria Sacc. However, studies Several of the Kirramyces specimens listed here were made
in recent years which have elucidated the structure of from seedlings in nurseries or from cultivated garden plants,
conidiomata and methods of conidiogenesis (Sutton, 1980) of and many more collections from plants in the bush are needed
many classical genera on the one hand, and applied this work to check the natural host ranges of these fungi.
Although these fungi are not recorded from Western
to a detailed consideration of Eucalyptus leaf fungi on the other
In considering these pale-spored taxa, the status of
Stagonospora delegatensis Park & Keane (1984) must be
J. Walker, B. C. Sutton and I. G. Pascoe
Australia (Shivas, 1989), at least three Eucalypfus species
native tc Western Australia have been found to be infected
when grown in eastern states. Of the species listed in Tables
1 and 2 . E. platypus is attacked by K. epicoccoides and E.
goniantha, E. nutans and E. plafypm are infected by K.eucalypti.
Further searches for these fungi in central and western
Australia are needed.
Overseas, there are numerous reports of these fungi on
Eucalyptus. Phaeoseptoria eucalypti was first reported from India
by Padaganur & Hiremath (1973) and subsequently it was
stated to cause leaf blighting of seedlings of E. camaldulensis
and E. f~reficomisin nurseries (Jamaluddin, Soni & Dadwal,
1985). In Italy, a fungus reported as Cercospora eucalypti was
recorded on 43 species of Eucalyptus seedlings in a nursery
(Magnani, 1965). Damage varied from light to severe,
depending on the species. It is of some interest that, of the 43
species 1i:jtedas hosts, 40 are in the subgenus Symphyomyrtus
and three in the subgenus Monocalyptus of Pryor & Johnson
(1971). Three of the species listed (E. gomphocephala DC., E.
occidenfalis Endl. and E. rudis Endl.) are native to Western
Australia. Literature reports on the host range and geographic
distribution of these fungi will need to be confirmed by a reexamination of the specimens on which they are based before
accurate host and distributional data can be compiled.
The authors gratefully acknowledge assistance from Dr Eileen
Scott (AIIW), Dr J. L. Alcorn (BRIP), Dr D. Pegler (K),Dr Irma
Gamundi de Amos (LPS)and Dr T. Kobayashi (TFM) who lent
specimens in their care, and from Margaret Dick, New
Zealand Forest Research Institute, who sent duplicates of New
Zealand collections. The assistance of the Curators of herb.
PAD anc herb. ROPV in searching for collections of Sepforia
morfolens:~is also acknowledged. They also thank Professor G.
Morgan-lones, Auburn University, Alabama, U.S.A. for his
comments on the conidiogenous cells of Phaeoseptoria papayae.
The senior author is also grateful to the Plant Pathology
Branch Herbarium (DAR), N.S.W. Department of Agriculture,
for providing working facilities.
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