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 ..... ... .'3 <,. :, ..... ..: ... .>. :.,,. ..::, I. ;;.. . ..... ... ... :: i.. , :. ' " . .:,r .." ,~: .... ::.. ..... ..- ...... ........ ............. .. ... . .::: . . L : .::. .,....... .. . . :,.. .,. ,... . .. ...12...-... :. ....... .. : .. ... . .'! .., ,. ,....,,: .. ,:.,, . ,?. > ^ I..: , ! . , . :.?. ...... 1 .'. .'.., .:$ ...; '.I, .L .- , 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. REFERENCES Alcorn, J. L. 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