Mycol. Res. 99 (7): 769-782 (1995) Printed in Great Britain 769 The mycobiota of the weed Lantana camara in Brazil, with particular reference to biological control ROBERT W. BARRETO l , HARRY C. EVANS 2 AND CAROL A. ELLISON 2 1 Departamento de Fitopatologia, Universidade Federal de Ｌ｡ｳｯｾｩｖ 36.570 Ｌ｡ｳｯｾｩｖ MG, Brazil Institute of Biological Control (lIBC), Silwood Park, Ascot, Berks, SL5 7TA, UK 2 International A survey of the fungi associated with the tropical weed Lantana camara was conducted during 1966 and 1969 in part of its native range in southern Brazil. The sampling activity was concentrated mainly in the state of Rio de Janeiro, covering 70 sites located in different climatic or ecological zones. The survey continued and was extended to the Amazon and northeast regions. Nine fungal species were identified as members of the mycobiota, including the previously undescribed fungi: Ceratobasidium lantanae-camarae sp. nov., Dendryphiella aspera sp. nov. and Micropustulomyces mucilaginosus gen. et sp. nov. Perisporiopsis lantanae, which has not been recorded previously in Brazil. has its anamorph described for the first time. The rusts Aecidium lantanae, Prospodium tuberculatum and Puccinia lantanae, as well as Mycovellosiella lantanae. Pseudocercospora guianensis and Anhellia lantanae were also recorded. Field observations suggest that several fungal pathogens have potential as classical biocontrol agents of the weed in the Old World tropics: Ceratobasidium lantanae-camarae and Puccinia lantanae in lowland humid climates; Mycovellosiella lantanae, Prospodium tuberculatum and Micropustulomyces mucilaginosus in more elevated or subtropical areas. In addition, databases and herbaria were consulted in order to compile a world list of fungal pathogens associated with L. camara. Lantana camara L. is a prickly shrub of the Verbenaceae native to the Americas. It was taken to Europe as early as the seventeenth century, where it was bred as an ornamental and the hybrids disseminated throughout the world, later becoming major pantropical weeds. Holm & Herberger (1969) listed it as one of the ten world's worst weeds. According to Holm et al. (1977), it has infested millions of hectares of natural grazing lands in Asia and Africa and in some areas of India its invasion of cultivated land has led to the shifting of entire villages. The same authors listed it among the worst weeds of a number of major subsistence and cash crops as well as of pasture and forested areas in many tropical countries in Africa, Asia, Australasia and the Pacific region. Lantana camara thickets are also said to be breeding places for feral pigs and insect vectors of disease, providing, in East Africa for example, a favourable environment for the tsetse fly (CIBC, 1982). The situation in the areas surveyed in Brazil is very different, and, although the species is a common component of the Brazilian weed communities (found in 60 % of the sites included in the survey in the state of Rio de Janeiro), it never occurs as an aggressive dominant species nor does it tolerate shade well. L. camara is included in all lists of Brazilian weeds, but this is due almost totally to the toxicity of its leaves and seeds to many domestic animals. Toxicity is also reported as an important cause of cattle losses throughout its pantropical range (Harley, 1974). Lantana camara was amongst the first weeds selected for biological control, with the pioneering work being undertaken in Hawaii at the beginning of the century (Perkins & Swezey, 1924). Emphasis was on insect biological control agents, as in all subsequent projects (Harley, 1974; Funasaki et al., 1988; Julien, 1992). Pemberton (1981) listed 21 projects, making L. camara easily the most targeted weed species for biological control. Results obtained have been unconvincing. Holm et al. (1977) commented that the introduced insect agents in Australia and Hawaii are effective in drier regions, but this is pOSSibly because L. camara is already under stress in these areas. Winder & Harley (1983) reported the results of work carried out in Brazil over a ten-year period on phytophagous insects, whilst Cilliers & Neser (1991) concluded that: 'A better understanding of the functional relationships and interactions between the various cultivars of L. camara and insect herbivores [in South Africa] may improve the chances of succeeding with biological control'. Evans (1987) considered the prospects for biological control of L. camara with fungal pathogens as being exceptionally good, based on his examination of material deposited at Herb. IMI. Recently, Queensland Department of Lands decided to renew the search for natural enemies of L. camara. As a result, a USA-based entomologist has been making collections of Lantana insects in Mexico, and the authors were asked to undertake preliminary surveys of fungal pathogens in South America. This paper reports on the mycobiota associated with L. camara in Brazil. MATERIALS AND METHODS The field survey was concentrated in the state of Rio de Janeiro (RJ) and the methodology is as described by Barreto The mycobiota of Lantana camara 770 & Evans (1994). In addition, shorter surveys were undertaken in the states of Amazonas, Espirito Santo, Bahia, Pani, Parana and Sao Paulo. For details of the laboratory studies and literature and herbarium surveys, see Barreto & Evans (1994). RESUL TS AND DISCUSSION Ceratobasidium lantanae-camarae Barreto & C. A. Ellison sp. nov. H. C. Evans, R. W. (Figs 1-8) Etym: reflecting its apparent restriction to Lantana camara. Laesiones in foliis vivis; maculae arnphigenae, irregulares, angulares, 1-20 mm latae, pallido-brunneae, fusco-marginatae, coalescentes. Fructificatio hypophylla, resupinata, tenuis, albida. Hyphae basales strietae, hyalinae, laeves, ramosae, 6-7'5 ｾｭ latae. Basidia subglobosa vel obovata, 12-20 x 7-10 lffil: 4 sterigmatibus, comutis, usque 7 x 3 ｾｭＮ Basidiosporae hyaJinae, laeves, aseptate, asymmetricae subglobosae vel late ellipsoideae, 6'5-8 x 4-7 ｾｭＬ valde apiculatae. Hyphae filamentosae steriles, petiolicola vel saepe call1icola, Lantana camara parasitica. Holotypus: !M! 360868, in Lantana camara, Rio Ariau-Rio Negro, Manacaparu, Amazonas, Brasilia, 31 July 1991, R. W. Barreto & H. C. Evans. Lesions initially absent, later appearing on both sides of leaf as irregular, angular, necrotic, occasionally water-soaked areas, 1-20 mm wide, pale brown to tan with a distinct dark brown border; producing shot-hole symptoms or coalescing to cover entire leaf surface, leading to disintegration and death of the leaf. Filaments of loosely parallel-growing, sterile hyphae, 4-6 I-Im diam, form webs on the stem, aggregating around the petioles as dense white clumps, inducing premature leaf fall. These stem webs, with prominent appressional pads, serve for internodal spread of the pathogen. Fructification hypophyllous, resupinate, thin, superficiaL forming a whitish, web-like bloom on young leaves. Basal hyphae hyaline, smooth, septate, without clamp connexions, branched, 6-7' 5 I-Im diam. These constitute the feeding mycelium which produce convoluted swollen 'haustoria' on and within the host stomata, as well as fertile hymenial branches which unite into a basidial-bearing mat. Basidia subglobose to obovate, 12-20 X 7-10 I-Im, with 4 horn-like sterigmata, up to 7 I-Irn long and 3!Jm wide at the base. Basidiospores hyaline, smooth, aseptate, asymmetrically subglobose to broadly ellipsoid, with a slightly flattened side, 6'5-8 x 4-7 fJIIl, strongly apiculate, apiculus 1'0-1'5 I-Im. Holotype: IMI 360868, on Lantana camara, Rio Ariau-Rio Negro, Manacaparu, Amazonas, Brazil, 31 July 1991, R. W. Barreto & H. C. Evans. Paratypes (all on L. camara): !M! 360870, Manaus-ltacoatiara, Amazonas, Brazil, 31 July 1991, R. W. Barreto & H. C. Evans; !M! 360869ltacoatiara, Amazonas, Brazil, 1 Aug. 1991, R. W. Barreto & H. C. Evans; IMI 360872, Arataca, Bahia, Brazil, 9 Aug. 1991, H. C. Evans; IMI 360871, Sao Jose da Vitoria, Bahia, Brazil, 9 Aug. 1991, H. C. Evans; IMI 360876, Una, Bahia, Brazil, 10 Aug. 1991, H. C. Evans; IMI 360875, Siquirres, Limon, Costa Rica, 2 June 1992, H. C. Evans & C. A. Ellison; IMI 360874, Cairo, Limon, Costa Rica, 2 June 1992, H. C. Evans & C. A. Ellison; IMI 360873, Puerto Bolivar, Cuyabeno, Napo, Ecuador, 23 July 1993, H. C. Evans & C. A. Ellison. Erkisson & Ryvarden (1973) re-described Ceratobasidium D. P. Rogers and separated it from other closely-related genera within the Corticiaceae on the basis of the secondary or repetitive basidiospores and the nature of the hymenium, since the basidia do not form a compact palisade layer but arise directly from the basal hyphae on short side branches. The fungus on L. camara clearly belongs to this genus (Figs 1, 6). Seven species are recognized (Eriksson & Ryvarden, 1973), including two plant parasitic species, C. anceps (Bres. & Syd.) H. S. Jacks. and C. bicorne J. Erikss. & Ryvarclen. On host, growth habit and morphological characteristics, principally basidiospore shape and size, C. lantanae-camarae can readily be separated from the other species assigned to the genus. The Brazilian records collected so far, and additional material from Ecuador and Costa Rica, indicate that C. lantanae-camarae has a wide distribution in the neotropics, but that it is favoured by lowland, humid tropical ecosystems. For example, it was not recorded during an intensive survey of L. camara in southern, subtropical Brazil (Barreto, 1991). In humid habitats, its affect on the host can be dramatic, with severe leaf fall and shoot die-back and, if it can be proven experimentally to be host specific (Evans & Tomley, 1994), it could be exploited as a classical biological control agent of L. camara in the palaeotropics. Field surveys already suggest that C. lantanae-camarae has an acceptably narrow host range, since other species of Lantana, such as L. lilacina L. growing in association with heavily infected L. camara, were found to be disease free. Finally, this appears to be the first report of the mode of nutrition of the parasitic Corticiaceae, with the finding of the substomatal, 'haustorial-like' structures (Figs 1, 4, 5), perhaps indicating some relationship with the Heterobasidiomycetes. Dendryphiella aspera R. W. Barreto & J. c. David sp. nov. (Figs 9-11) Etym: rough-walled conidia and conidiophores. Laesiones in foliis vivis; 1-10 mm, latae, maculae subrotundae atrobrunneae arnphigenae. Mycelium immersum intracellulosum, 1-3 IJm diametro, ramosum, septatum, pallidum brunneum. Mycelium superficiale absens. Stromata absentia. Conidiophora per stomata emergentia, hypophylla, plerumque singularia sed interdum fasciculata, usque 5 in fasciculo, cylindrica, recta nodosa, 136-544 x 4 lffil, 5-13 septata, non ramosa, brunnea, parietibus verruculosis. Cellulae conidiogenae terminales et intercalares, enteroblasticae, polyblasticae, cylindricae, parte fertilis inflatis terminanti, 13-66 x 4-7 ｾｭＬ brunneae. Loci conidiogeni minuti, usque 1 IJm lati, complanati, incrassati, fuscati. Conidia sicca, catenata (catenae ramosa), enteroblasticae, cylindrica vel elliptica, 10-22 x 4-6 1Jffi, apice et basi rotundata, 0-3 septata, eguttulata, brunnea, parietibus verruculosis. Holotypus: 1M! 345366, in Lantana camara Sao Paulo, Brasilia, 25 Aug. 1989, R. W. Barreto. Lesions on living leaves; 1-10 mm wide, forming subcircular dark-brown necrotic spots on both sides of the leaves. Internal mycelium intracellular, 1-3 I-Im diam., branched, septate, light brown. External mycelium absent. Stromata absent. Conidiophores arising through stomata, hypophyllous, mostly single but occasionally in loose groups of up to 5 conidiophores, cylindricaL 136-544 X 4 I-Im, straight, nodose, 5-13 septate, R. W. Barreto, H. C. Evans and Carol A. Ellison 771 A ( = 00 00 (J Fig. 1. Ceratobasidium lantanae-camarae, basal hyphae showing feeding mycelium and fertile hymenial branches with basidiospores on lower leaf surface. (A) Feeding mycelium showing stomata invaded by swoIlen, vesicle-like structures (smaIl arrows). Note convolution of hyphae near stomata (large arrows); (B) stages in development of basidia, sterigmata and basidiospores. Note young basidia (arrows); (C) basidiospores, showing apiculate condition (smaIl arrow), germination (large arrow); (D) baSidiospore showing repetitive germination, with secondary spore. Bars, 8 j..lm. unbranched, brown, rough-walled, becoming smooth towards the base. Conidiogenous cells terminal and intercalary, tretic (enteroblastic proliferation with simultaneous holoblastic conidial ontogeny), cylindricaL terminating in a swollen fertile portion with up to 3 conidiogenous loci, 13-66 x 4-7 !-lm, brown. Conidiogenous loci inconspicuous on the conidiogenous cells but conspicuous on the conidia (see Fig. 9), up to I j..lrn wide, raised, thickened, darkened, smooth. Conidia dry, formed in branched chains, holoblastic, cylindrical to ellipsoidal. 10-22 x 4-6 !-lm, apex and base rounded, with thickened and darkened hilum, 0-3 septate, eguttulate, brown, rough-walled. In culture: slow-growing, 2'4-4'7 em after 30 d, dark grey, felty layer of aerial mycelium and conidiophores sometimes covered by a layer of cottony mycelium, with concentric diurnal zonation and prodUcing a ring of brown diffusate at periphery (when subjected to alternating exposure to black light/white fluorescent on PCA media), grey reverse, sporulation abundant. Holotype: IMI 345366, on Lantana camara, Sao Paulo, BraziL 25 Aug. 1989, R. W. Barreto. The suitable placement for this fungus posed some difficulties. Initially, it was considered that it could be accommodated in Cladosporium. C. oxysporum Berk. & M. A. Curtis had already been recorded in the literature on this host, but examination under SEM (Figs 10-11) showed that the fungus did not have conidial scars with an external raised ring, diagnostic for Cladosporium. Another possibility was the genus Stenella, but species in this genus do not have nodose conidiogenous cells and are known to have verruculose external mycelium and smooth conidiophores. The fungus on L. camara does not produce any external mycelium and has verruculose conidiophores and nodose conidiogenous cells, and, in most morphological aspects, it fits best in Dendryphiella. The other species in the genus are D. vinosa (Berk. & M. A. Curtis) Reisinger, D. infuscans (ThUrn.) M. B. Ellis, D. armaria Nicot (= Scolecobasidium armarium (Nicot) M. B. Ellis) and D. salina (G. K. Suther!.) G. J. F. Pugh & Nicot (= Scolecobasidium salinum (G. K. Suther!.) M. B. Ellis). These are cosmopolitan saprotrophs while the new species appears to be a specific pathogen of L. camara. The two last species mentioned are restricted to marine and estuarine habitats. Morphologically, D. aspera differs from D. vinosa by having shorter conidia and narrower nodes on the conidiogenous cells. It differs from D. infuscans in having distinctly verruculose conidiophores and conidia. Conidiogenesis in Dendryphiella is tretic. Observation under SEM has revealed the possible presence of a central pore (Fig. II). However, clarification of conidiogenesis requires TEM examination. Dendryphiella aspera was collected only at the type locality, a steep and humid roadside site, 580 m above sea leveL in the winter months. A single brief attempt to fulfil Koch's postulates was undertaken without disease being reproduced. However, the field damage observed, and the ability of the fungus to grow and sporulate abundantly in culture, indicate that this fungus warrants further study and evaluation either as a classical biocontrol agent of L. camara, or a mycoherbicide. Mycovellosiella lantanae (Chupp) Deighton var. lantanae, Mycological Papers 137: 33 (1974) Cercospora lantanae Chupp in Toro, Journal of the Department of Agriculture of Puerto Rico 15: 10 (1931); Cladosporium trichophilum Petro & Of., Annales Mycologici 30: 337 (1932); Chaetotrichum lantanae Petr., Sydowia 5: 38 (1951). Lesions on living leaves: variable in appearance; from indistinct, smaiL chlorotic spots to large, necrotic patches covering whole areas of the leaves and leading to leaf death, the most common appearance being that of powdery spots on abaxial leaf surfaces with a dark brown centre (1-4 mm wide) surrounded by a wider chlorotic zone with ill-defined limits, the adaxial surface having dark reddish-brown to black spots. Internal mycelium intracellular, 2-4 \.lm wide, branched, septate, hyaline, smooth. External mycelium arising from stomata and The mycobiota of Lantana camara 772 Figs 2-5. Ceratobasidium lantanae-camarae, macro- and micro-symptoms on lower leaf surface. Fig. 2. Shot-hole symptom on older, infected part of leaf with advancing, asymptomless, white sporulating area (arrow) (Bar = 3 mm). Fig. 3. Typical old necrotic, pale brown lesions with prominent dark brown border, smaller lesions beginning to disintegrate (Bar = 5 mm). Figs 4-5. Close-up of leaf surface after clearing and staining (Bar = 8 ｾｭＩＮ Fig. 4. Shows two stomata invaded by swollen, vesicle-like structures from central basal hypha, arrow denotes abnormal build-up of chitin-like material around guard cells. Fig. 5. Shows swollen structures within the substomatal chamber (arrow). R. W. ｂ｡ｲ ･ｾｯＬ 773 H. C. Evans and Carol A. Ellison Fig. 9. Dendryphiella aspera sp. nov., showing upper part of conidiophore with conidiogenous loci (A) and acropleurogenous, rough-walled conidia (B). In culture: slow-growing, 0'4-2'4 em after 23 d either brown and stromatic, composed of dense mass of spherical spermogonia and chlamydospores, covered with tufts of white aerial mycelium, dark grey to black .reverse; sporulation ｡｢ｳ･ｮｾ ｾｯ abundant. Material examined: IMI 345367, Itaperuna, RJ, 13 Sept. 1989; 1M! 345368, Resende, RJ, 21 June 1989. Figs 6-8. Ceratobasidium lantanae-camarae, SEM studies of feeding mycelium and basidia on lower leaf surface. Fig. 6. Side branch from basal mycelium penetTating stomatal aperture (Bar = 4 I-Im); Figs 7-8. Basidia with sterigmata in various stages of development (Bar =3I-1m). spreading over ｾｨ･ abaxial leaf surface, ｧｮｩｷｴｲ･ｾ ｯｾ form ropes ｡ｭｯｮｧｳｾ ｾｨ･ ｾｲｩ｣ｨｯｭ･ｳＬ 1-3 IJITl diam, sparsely branched, hyaline, ｳｭｯｾｨＭｷ｡ｬ･､Ｎ Stroma absent. Conidiophares hypophyllous, produced terminally or laterally on ｾｨ･ superficial mycelium, solitary or forming ｾｵｦｳ on the ｾｲｩ｣ｨｯｭ･ｳＬ cylindrical, 14-46 x 3-6 1JITl, ｬ｡ｮｩｭｲ･ｾ conidiophores being much longer, ｳ･ｰｾ｡ and ｯｦｾ･ｮ branched, while lateral conidiophores are ｳｨｯｲｾＬ often continuous and unbranched, straight or sinuose, hyaline, smooth-walled. Conidiogenous cells terminal or intercalary, ｩｮｾ･ｧｲ｡､Ｌ proliferating irregularly or sympodially to form several conidiogenous loci per cell ; cylindrical, 8-31 x 3-6 1JITl; straight becoming sinuose and geniculate towards the apices, hyaline. Conidiogenous loci numerous, ｭｩｮｵｾ･Ｌ ｰｲｯｴｵ｢･ ｡ｮｾＬ thickened, dark. Conidia dry, produced in dry chains, holoblastic, subcylindrical 1443 x 2-4 1JITl, apex rounded, base obconic ｷｩｾｨ thickened scar, 1-2 !-lm, 0-2 septate, guttulate, hyaline, ｳｭｯｾｨＭｷ｡ｬ･､Ｎ Two varieties of this fungus were recognized by ｄ･ｩｧｨｾｯｮ (1974): Mycovellosiella lantanae var. lantanae and M. lantanae var. cubensis Deighton. The observation of ｣ｯｮｴｲ｡ｳｾｩｮｧ disease infection symptoms on L. camara in the field associated ｷｩｾｨ by Mycovellosiella strongly suggested that two, or possibly more, distinct taxa were involved. Examination and comparison of the morphology of the fungi obtained from samples showing some of the ｭｯｳｾ contrasting symptoms (indistinct chlorotic zones, dark-brown extensive necrosis, black adaxial did not spots and leaves dry - suffused with ｡ｮｾｨｯ｣ｹ｡ｮｩＩ reveal any significant differences. Deighton (1974) separated M. lantanae var. cubensis from M. lantanae var. lantanae on the basis of the fonner having narrower conidiophores with shorter and narrower, and mostly non-septate conidia. Samples examined from Brazil clearly belonged to M. lantanae var. lantanae. With respect to pathological symptoms, the material of the two varieties examined by Deighton differed little: M. lantanae var. lantanae having 'Leaf spots indefinite, merely a yellowish-brown discoloration of the upper surface ... ' and M. lantanae var. cubensis with 'Leaf spot none' (Deighton, 1974). The contrast of Deighton's observations with those made in Brazil is great. The mycobiota of Lantana camara 774 This variation could be due to environmental factors, but, if inherent variation is present, either polymorphism or aggregate taxon status could be involved. M. lantanae var. lantanae was extremely common throughout the year in the state of Rio de Janeiro (77% of the sites where its host was present) and was also common in other states (Sao Paulo, Parana, Espirito Santo). L. camara appeared to be less affected by this pathogen at higher altitudes (900 m and above). Nevertheless, this is a fungus with a broad environmental tolerance, occurring in all the main climatic types in the state of Rio de Janeiro. A mycoparasite, Verticillium lecanii (H. Zimm.) Viegas was found in association with 14'5 % of the samples of M. lantanae var. lantanae in sites from sea level up to 1140 m. The frequency of M. lantanae var. lantanae in a range of environmental conditions, coupled with the severe defoliation it can cause to its host and its absence in the Old World, places this fungus among the most promising classical biocontrol agents of L. camara. Mycovellosiella lantanae var. lantanae will grow in artificial media and sporulate, however, the chances of mycoherbicidal development with this fungus appear to be remote because all isolates grew very slowly and sporulated irregularly. Figs 10-11. Dendryphiella aspera, IMI 345366, SEM detail of conidial chains and scars (arrows). Bar = 2'5 Ilm (Fig. 10); 0'8 Ilm (Fig. 11). Perisporiopsis lantanae (F. Stevens) R. W. Barreto comb. nov. (Figs 12-15) Perisporina lantanae F. Stevens, Transactions of the Illinois Academy of Science 10: 170 (1917). 12 15 Figs 12-15. Perisporiopsis lantanae comb. nov., IMI 345369. Fig. 12. T.S. through ascoma (Bar = 2'5 Ilm); Fig. 13. Asci showing detail of ascus tip (Bar = 10 Ilffi); Fig. 14. Ascospore (Bar = 8 Ilm); Fig. 15. Conidium (Bar = 8 Ilm). Lesions not produced (the fungus forms a sooty coating on the leaves of its host). Internal mycelium absent. External mycelium 3-4 I-lm diam., branched, septate, brown, smooth-walled. Teleomorph Ascomata pseudothecioid, amphigenous, superficial, abundant, intermixed with anamorph, subspherical, 122-190 x 95163 I-lm; wall 1-3 cells thick, 3-10 I-lm, composed of dark brown textura angularis, ornamented with abundant brown septate setae (8-10 I-lm wide and up to 286 I-lm long), often with a group of shorter setae arranged as a crown surrounding the ostiole. Dehiscence ostiolate, one ostiole in the centre of each pseudothecium, circular, 3 I-52 I-lm diam., papillate. Interthecial filaments pseudoparaphyses, thin, branched, hyaline. Asci bitunicate, fasciculate, subclavate, 66-98 x 17-22 I-lm, 8spored. Ascospores fusiform, 35-49 x 8-10 I-lm, rounded in one extremity subacute in the other, 2-3 septate, guttulate, greyish-brown, smooth-walled. Anamorph Conidiomata pycnidial, amphigenous, superficial, abundant, intermixed with ascomata, indistinguishable from ascomata in external appearance, wall thickness, composition and colour, 136-170 x 109-150 I-lm. Dehiscence as for teleomorph. Conidiophores absent. Conidiogenous cells arising from lower half of pycnidial wall, subcylindrical, 4-10 x 4--6 I-lm, hyaline. Conidia holoblastic, cylindrical, 60-80 x 6-8 I-lm, straight to slightly curved, apex round, base truncate, 3 I-lm wide, 4--6 septate, guttulate, greyish, smooth-walled. R. W. Barreto, H. C. Evans and Carol A. Ellison 775 Figs 18-19. Puccinia lantanae, IMI 345374. Fig. 18. Light micrograph of unicellular and bicellular teliospores (Bar = 20 ｾＩ［ Fig. 19. SEM of telium (Bar = 14 ＮＩｭｾ Figs 16-17. SEM of Prospodium tuberculatum, IMI 345371. Fig. 16. Teliospore (Bar = 8'5 ｾＩ［ Fig. 17. Teliospore and urediniospores (below) (Bar = 5 ｾＩＮ Material examined: IMI 345369, Cachoeiras do Macaw, RJ, 27 July 1989; IMI 345370, Angra dos Reis, RJ, 2 Aug. 1989. Stevens (1917) ereded the species Perisporina lantanae based on material from Puerto Rico, which, until now, has remained the sole record. This is, therefore, the first record from Brazil. Muller & Arx (1962) synonymized Perisporina under Perisporiopsis but did not propose a new combination for Perisporina lantanae. No reference was made by Stevens to an anamorph. This was found in the two samples colleded in Rio, but it is not named because it did not appear to occur separately from its teleomorph. In addition to the two above colledions, the fungus was also colleded in Macae (RJ) and Manacapuru (Amazonas). The fungus seems to be adapted to very warm and humid sites. Perisporiopsis lantanae grows as an epiphyte rather than a pathogen, and no penetration of the tissue of L. camara by the fungus was noted. It is, therefore, of no significance for weed biocontrol. Prospodium tuberculatum (Speg.) Arthur, North American Flora 7; 161 (1912) (Figs 16-17) Uredo tuberculata Speg., Anales de la Sociedad Cientifica Argentina 9: 172 (1880) Uredinial and teliallesions on living leaves; extensive, irregular, without clear limits, coalescing, orange to purplish brown. Mycelium intercellular, 1-4 11m diam., branched, septate, hyaline. Uredinia mainly hypophyllous, but also occurring on adaxial leaf surface, sub-epidermal, scattered initially, coalescing as disease progresses, 190-544 IJffi wide. Paraphyses densely-packed below the layer of urediniospores within the sori, clavate, 15-52 x 7-15 11m, on a short pedicel, 1121 x 6-7 11m, thin-walled, hyaline. Urediniospores abundantly produced, globose, 17-24 11m diam., unicellular, thin-walled, minutely aculeate, 2-pored, pores in opposite equatorial position. Urediniospore pedicels cylindrical, 42-63 x 3-4 11m, hyaline, not ornamented. Teliospores produced in epiphyllous uredinia, few in each sori, ellipsoidal, 34-46 x 24-31 11m, bicellular, slightly constrided at septa, wall 3-8 IJffi thick, dark brown, sparsely verrucose, verrucae distant 1-6 11m apart, one distind pore at each cell (at the apex in the upper cell and adjacent to the septum in the lower cell). Teliospore pedicels cylindrical, up to 46 11m long, with branched ornaments, collapsing after discharge and becoming coiled. Material examined: IMI 345371, Pirai, RJ, 13 Aug. 1989; IMI 361065, Tijuca, RJ, Brazil, 15 July 1989 (with Aecidium lantanae);IMI 361067, Pico do Frade, Macae, RJ, 15 Oct. 1993; IMI 361066, Terras Frias, RJ, 20 Jan. 1994; IMI 361068, Trajano de Morais, Macae, RJ, 29 Jan. 1994. P. tuberculatum was never particularly common, although a wide geographical distribution was noted (the fungus being found outside the main sampling range in Sao Paulo, Parana and Bahia). On several occasions, the uredinial state was heavily parasitized by a Sphaerellopsis sp. The rust can cause severe leaf necrosis alone, as noted in the field (Parana) and in the greenhouse, or in association with M. lantanae var. lantanae. Based on field observations and damage assessment, Evans (1989, unpub. report) suggested that P. tuberculatum The mycobiota of Lantana camara 776 surrounded by aecia, flask-shaped, 94-170 x 70-164 I-lm. Aecia hypophyllous, deep-seated, in groups surrounding the spermogonial cluster, cup-shaped, 150-354 I-lm diam. Peridia well-developed; membrane cells 13-24 x 11-22 I-lm, polygonate, hyaline. Material examined: IMI 345365, Saquarema, RJ, 17 Aug. 1989; IMI 361065 (see Prospodium tuberculatum). The sole previous record of Aecidium lantanae on L. camara was from Brazil (Hennen et al., 1982) and only the aecia of this fungus were described. Here spermogonia of type 4 (group 5) were found (Hiratsuka & Cummins, 1963). Aecia were freguently parasitized by Tuberculina sp. Greenhouse inoculations with aeciospores have not been successful but this was probably due to low spore viability rather than to failure to infect. This rust has been collected from a number of sites together with Prospodium tuberculatum (1M! 361065-361068), often on the same leaf. Whilst it is tempting to link these two species, there is no experimental proof of a connexion and the relatively few Prospodium species in which all spore stages have been discovered, possess subcuticular spermogonia with a flat hymenium (group 6, type 7) and uredinioid aecia (Cummins, 1940; Hiratsuka & Cummins, 1963). Fig. 20. Aecidium lantanae, IMI 345365, showing aecial cups surrounding cluster of spermogonia (arrow) (Bar = 1 mm). Fig. 21. MicroplIStulomyces mucilaginosus gen. et sp. nov., IMI 345375, t.s. through acervulus (Bar = 20 IJ.ID). merits further study as a biocontrol agent, as a result, greenhouse trials are now being carried out with this fungus. Initial results are promising. Thirty plant species have so far been screened, belonging to the Verbenaceae and related families, following the centrifugal, phylogenetic testing sequence (Wapshere, 1974). None of these species has shown any macroscopic symptoms of infection. Despite the fad that P. tuberculatum has a wide host range within Lantana (Cummins, 1940; Herb. IMI records), the isolate being tested from Rio de Janeiro state appears to be specific to a single cultivar of L. camara (common pink) which is one of the most important weedy types in Queensland, Australia. Further surveys are underway to collect additional strains of the rust which are pathogenic to all the main weedy cultivars. Elucidation of the life-cycle for P. tuberculatum is now required using artificial inoculation with basidiospores. This would be a prerequisite for its use as a classical weed biocontrol agent, because, although Prospodium is an autoecious genus (Cummins, 1959), it would remove any uncertainties about an alternate host of ecological or economical significance. Aecidium Iantanae Mayor, Memoires de la Societe neuchateloise des Sciences naturelles 5: 567 (1913) (Fig. 20) Aecial and Spermogonial lesions on living leaves; 4-6 mm wide, circular to irregular, vein-delimited, initially pale green, becoming necrotic with a chlorotic halo. Spermogonia type 4, hypophyllous, subepidermal, grouped in the centre of lesions, Pseudocercospora guianensis (p. Stevens & Solheim) Deighton, Mycological Papers 140: 145 (1976). Cercospora guianensis F. Stevens & Solheim, Mycologia 23: 375 (1931) Lesions on living leaves; scattered over the surfaces and along the margins, up to 7 mm wide, irregular, vein-delimited. dark brown. Internal mycelium intracellular, 1-2 I-lm diam. branched, septate, light brown. Superficial mycelium absent. Stromata erumpent; 24-45 x 14-24 I-lm; composed of dark brown textura angularis. Conidiophores bursting through cuticle, arnphigenous, densely fasciculate, 6-29 conidiophores per fascicle, cylindrical, 66-175 x 3 I-lm, becoming slightly sinuose towards the apices, 2-5 septate, unbranched, brown. Conidiogenous cells terminal, integrated, holoblastic, proliferating holoblastically and sympodially, cylindrical, 25-66 x 4-6 I-lm, becoming sinuose and geniculate towards the apices, pale brown. Conidiogenous loci inconspicuous, up to 1 I-lm, flat, unthickened, not darkened. Conidia dry, holoblastic, cylindrical 32-69 x 4--6 I-lm, apex rounded, base truncate, 1 I-lm wide; 0-5 septate, guttulate, light brown, smooth-walled. In culture: colony slow-growing, 1'0-1'5 ern after 28 d, either stromatic, covered with felty aerial mycelium and dark grey (occasionally with white sectoring), black reverse; no or little sporulation. Material examined: IMI 345372, Pirai, RJ, 7 Mar. 1989; IMI 345373, Pirai, RJ, 13 June 1989. Evans (1987) was of the opinIOn that a reVISIOn of the Cercospora-like complex on L. camara was necessary. L. camara is the host of five different species of Cercospora and also other Cercospora-Iike fungi: M. lantanae, Ramularia sp. and two species of Pseudocercospora (Table 2). In addition to P. guianensis, Pseudocercospora formosana (W. Yamam.) Deighton has been described from L. camara. The R. W. Barreto, H. C. Evans and Carol A. Ellison 777 Fig. 22. Micropustulomyces mucilaginosus, section of acervulus and conidia (Bar = 20 ｾｭＩＮ latter species differs from P. guianensis in having narrower conidia arising from non-fasciculate conidiophores as indicated by Chupp (1954). P. guianensis was rare in the state of Rio de Janeiro (obtained once from a single site), and it was suggested (Barreto, 1991) that this might be an indication of a preference for lower altitudes, considering the numerous records from typically tropical countries (see Table I). This hypothesis was confirmed by the ubiquity of this fungus on L. camara noted during a survey in the Brazilian Amazon Guly - Aug. 1991). The effect of P. guianensis on its host was not particularly pronounced, even in the Amazon, and its use in biocontrol does not appear to be good. are usually fractured in microscopic mounts, wall irregularly light thickened (often thicker at the apical portion), 2-7 ｾｭＬ reddish-brown, smooth, pores discrete. Pedicels cylindricaL up to 45 ｾ ｭ long, pale brown, smooth. Material examined: IMI 345374, Cachoeiras do Macaai, RJ, 14 June 1989. Evans (1989, unpub. report) expressed doubts about the distinction of P. lantanae from P. natalensis Dietel & Sydow (also recorded on L. camara). In the state of Rio de Janeiro, P. lantanae was common and damaging on Lantana lilacina but very rare on L. camara (only one collection was made on this host). Material resulting from the field survey and that from Herb. IMI were compared. The results are presented in Table 2. Puccinia lantanae Far!., Proceedings of the American Academy of Arts and Sciences 18: 83 (183) (Figs 18-19) Lesions on living leaves few; up to 4 mm wide, angular, veindelimited, chlorotic becoming dark brown and necrotic. Aecia and uredinia unknown. Telia hypophyllous, subepidermal, scattered over the leaf surface, rarely confluent, 1-2 mm wide. Teliospores tightly packed within the sori, of two kinds: unicellular, obovate, 21-31 x 14-20 ｾ ｭ and bicellular, clavthe unicellular kind being much more ate, 22-46 x 14-22 ｾｭＬ common, the bicellular teliospores slightly constricted at the septum and occasionally showing abnormalities such as asymmetry and an additional cell. The apices of the teliospores The data suggest that P. lantanae and P. natalensis are distinct and that P. lantanae var. erythraeae is a synonym of P. natalensis. Examination of the material confirmed the occurrence of P. lantanae in Ghana, an isolated occurrence outside the Americas on L. camara. Records of P. natalensis on L. camara should be considered dubious, perhaps resulting from misidentification of the host since most records of P. natalensis are on species of Lantana native in Africa, whilst P. lantanae is common on neotropical endemic species in South America Gackson, 1932). Puccinia lantanae was collected only once on L. camara during the survey in Rio but was found to be extremely 778 The mycobiota of Lantana camara Table 1. Fungal pathogens recorded on Lantana camara (ex CABI Database, Herb. 1M!) Distribution Ascomycotina &< Deuteromycotina Acanthosligma (= Nematostoma) lantanae (Henn.) Theiss. (N) Anhellia lantanae (Viegas) Arx (N) Alternaria sp. (0) Capnodium sp. (0) Cercospora sp. (0) Cercospora canescens Ellis & G. Martin (0) Cercospora lantanae-aculeatae J. M. Yen (0) Cercospora lantanae-camarae ]. M. Yen & Gilles (0) Cercospora lantanae-camarae R. C. Rajak & R. K. Rajak (homonym of above) (0) Cercospora lantanicola Chupp (N) Cladosporium oxysporum Berk. & M. A. Curtis (N, 0) Colletotrichum capsici (Syd.) E. ]. Butler & Bisby (0) Corynespora cassiicola (Berk. & M. A. Curtis) C. T. Wei (0) Diaporthe sp. (N) Epiphyma (= Botryosphaeria) nervisequens (Chardon) J. H Mill. & Burton Dominican Rep. (Ciferri, 1961), Puerto Rico (Stevenson, 1975) Brazil (Arx, 1963) India (IMI 228072) India (Sarbhoy et al., 1986), Malaysia (Johnston, 1960) Mauritius (Orieux & Felix, 1968) India (lMI 91342, 288610) Singapore (Yen, 1966) Gabon (Yen, 1971) India (Rajak & Rajak, 1981) Colombia (Chupp, 1954) Costa Rica (IMI 275388), India (lMI 276361) India (Sarbhoy et al., 1986) India (1M! 288610) U.S.A. (Hawaii) (Parris, 1940) Venezuela (Chardon, 1939) (N) Eutypa aspera (Nitschke) Fuckel (0) Gloeosporium sp. (N) Godronia lantanae (E. K. Cash) Seaver (N) Meliola sp. (N) Meliola ambigua Pat. & Gaillard (N, 0) Meliola cookeana Speg. (N, 0) Meliola durantae Gaillard (N) Mycovellosiella lantanae (Chupp) Deighton var. cubensis Deighton (N) Mycovellosiella lantanae (Chupp) Deighton var. lantanae (N) Myrothecium roridum Tode (0) Necfria wegeliniana (Rehm) Rehm ex Strasser (0) Oidium sp. (0) Oidium erysiphoides Fr. (N) Perisporina lantanae F. Stevens & Solheim (N) Phoma sp. (0) Phoma lantanae Verwoerd & du Plessis (0) Phomopsis lantanae (M. E. A. Costa & Sousa da Camara) B. Sutton (0) Phyllosticta sp. (0) Phyllosticfa lantanae-verae T ratter (N) Phylloslicta lantanae Pass. (N) Physalospora abdita (Berk. & M. A. Curtis) Voorhees (N) Pseudocercospora formosana (W. Yamam.) Deighton (N, 0) Pseudocercospora guianensis (F. Stevens & Solheim) Deighton (N) Ramularia sp. (N) Rosenscheldia paraguaya Speg. (N) Scolecobasidium sp. (0) Scolecopeltidium lantanae Bat. & I. H. Lima (N) Septoria lantanae Garman (N) Septoria lantanifolii Bat. & J. L. Bezerra (N) Sphaerulina sp. (0) Stictis radiata (L.) Pers. (0) New Caledonia (Huguenin, 1966) Barbados (Norse, 1974) U.S.A. (Hawaii) (Farr et al., 1989) Brazil (lACM) Dominican Rep. (Ciferri, 1961), Ghana (Piening, 1962), Puerto Rico (Stevenson, 1975), Trinidad & Tobago (Baker & Dale, 1951), Venezuela (Farr et aI., 1989) Sierra Leone (Farr et aI., 1989), USA (Farr et al., 1989) Dominican Rep. (Ciferri, 1961) Cuba (Deighton, 1974) Barbados (Norse, 1974), Bermuda (Waterson, 1947), Brazil (Chupp, 1954), Panama (Rios, 1982), Puerto Rico (Stevenson, 1975), U.S.A. (Farr et al., 1989), Venezuela (Chupp, 1954) Malaysia (Johnston, 1960) Zambia (IMI 96627) India (Butler & Bisby, 1960) South America (Viegas, 1961) Puerto Rico (Stevens, 1917) Burma (IMI 182584), India (lMI 150937), Zambia (IMI 100131) India (Mathur, 1979), South Africa (Gorter, 1981) India (Evans, 1987), Portugal (Costa & Camara, 1955), Singapore, Zambia (Evans, 1987) Solomon Islands (IMI 77259) Brazil (Batista & Vital, 1952), Puerto Rico (Stevenson, 1975) (as P. lantanicola) Barbados (Norse, 1974) Cuba (Farr et al., 1989), U.s.A. (Hawaii) (Farr et al.. 1989) Brazil (Evans, 1987), Brunei (Peregrine & Ahmad, 1982), China (Tai, 1979), Hong Kong (Leather & Hor, 1969), India (Evans, 1987), Japan (Yamamoto, 1934), Philippines (Evans, 1987), Singapore (Yen & Lim, 1980), Taiwan (Sawada, 1942) Bermuda (Chupp, 1954), Brazil (Hino & Tokeshi, 1978), Cuba (Urtiaga, 1986), Guyana (Chupp, 1954), Venezuela (Chupp, 1954) Trinidad & Tobago (Evans, 1987) South America (Viegas, 1961) Australia (IMI 151647) Brazil (Batista, 1959) Dominican Rep. (Ciferri, 1961), Puerto Rico (Stevenson, 1975) Brazil (Batista et al., 1959) Zambia (lMI 100131) New Caledonia (Huguenin, 1966) Basidiomycotina Aecidium lantanae Mayor (N) Aleurodiscus sp. (0) Prospodium tuberculatum (Speg.) Arthur (N) Brazil (Hennen et al., 1982) Zambia (IMI 96628) Argentina (Lindquist. 1982), Brazil (Evans, 1987), Dominican Rep. (Ciferri, 1961), Mexico (Leon-Gallegos & Cummins, 1981), Puerto Rico (Stevenson, 1975), Jamaica (Dale, 1955) 779 R. W. Barreto, H. C. Evans and Carol A. Ellison Table 1. (coni.) Distribution Puccinia lanlanae Farl. (N, 0) Antigua (Evans, 1987), Argentina (Lindquist, 1982), Barbados (Norse, 1974), Brazil (Lindquist & Costa Neto, 1963), Cuba (Rosenada, 1973), Dominican Rep. (Gferri, 1961), Ghana (Dade, 1940), Jamaica (Baker & Dale, 1951), Panama (Rios, 1982), Puerto Rico (Stevenson, 1975), Trinidad & Tobago (Baker & Dale, 1951), Venezuela (Evans, 1987), U.SA (Farr el al., 1989) Ethiopia (Baccarini, 1906), Ghana (Piening, 1962): Sudan (Evans, 1987" Philippines (Orillo & Valdez, 1959), U.SA (Farr el al., 1989) U.SA (Farr el al., 1989) Puccinia naialmsis Dietel & Syd. (0) Rhizoctonia sp. (N, 0) Rhizoctonia solani J. G. Kiihn (N) N, Recorded only from the New World; 0, recorded only from the Old World; ., records require confirmation. Table 2. Biometric data of microcyclic Puccinia spp. on Lantana (in 1IDl) Bicellular teliospores Unicellular teliospores Host (Length x width) L. odorata 26-38 x 19-26 23-27 x 15'5-20 Lantana sp. 20-42 x 15-21 22-28 x 15-21 L. lilacina 24-32 x 13-22 28 x 15 18-32 x 15-20 25 x 16 L. lilacina 22-35 x 14--18 26 x 15 14-35 x 11-28 23 x 17 L. camara 22-46 x 14-22 30 x 17 21-31 x 14-20 25 x 17 L. camara 35-49 x 17-28 38 x 21 28-35 x 14-24 32 x 20 L. camara 36-42 x 20-28 31 x 20 28-42 x 15-22 26x 16 (x) (Length x width) (x) Puccinia Iantanae Farlow (1883) (ex U.SA) Lindquist (1982) (ex Argentina) IMI361062 (ex RJ) !M! 361063 (ex RJ) 1M! 345374 (ex RJ) 1M! 38054" (ex Ghana) 1M! 361064 (ex Colombia) Puccinia Iantanae var. erythraeae L. camara Baccami (1906) 45 x 28 (ex Ethiopia) Puccinia natalensi. Sydow (1899) (ex South Africa) !M! 67571 (ex Sudan) L. salvaifolia 43-55 x 26-36 L. trifolia 35-59 x 17-24 50 x 23 " Deposited as P. nalalensis in Herb. IMI. common and damaging on this host in more tropical regions of Brazil (Para., Bahia). These observations suggest that P. lantanae is not a significant pathogen of L. camara in subtropical situations, but is of potential interest for classical weed biocontrol in warmer, more humid regions. Micropustulomyces R. W. Barreto gen. nov. Etym: derived from the discrete leaf pustules. Conidiomata acervularia, hypophylla, subepidermalia, in calervis densis rolundalis aggregata, irregulariler dehiscenlia, parielibus hyalinis. Conidiophora e pseudoparenchyma basali formala, cylindrica, erecla, septata, interdum ramosa, hyalina, laevia. Cellulae conidiogenae terminales, inlegralae, holoblasticae, cydindricae, hyalinae. Conidia mucosa, holoblaslica, cylindrica vel fusiformia, apice obtuso, base obconico vel truncato, 0-3 seplala, guttulata, hyalina, laevia. Paraphyses filiformes, hyalinae, aseplalae, a apices inflalae. Sp. type.: Micropustulomyces mucilaginosus R. W. Barrelo (hololypus) Conidiomata acervular, hypophyllous, subepidermal, in dense circular groups, wall hyaline; dehiscence irregular. Conidiophores formed from the basal pseudoparenchyma, cylindrical, erect, occasionally branched, hyaline, smooth-walled. Conidiogenous cells terminal, integrated holoblastic, cylindrical, hyaline. Conidia slimy holoblastic, cylindrical to fusiform, apex obtuse, base obconic to truncate, 0-3 septate, guttulate, hyaline, smooth. Paraphyses filiform, hyaline, aseptate, swollen at apices. Micropustulomyces mucilaginosus R. W. Barreto sp. nov. (Figs 21-22) Etym: referring to the mucilaginous appearance of the conidiomata Laesiones in foliis vivis; chlorolicae, brunneae - emorienles, subrolundatae, usque 25 nun latae, coalescentes, conidiomalibus aggregalis sicul numerosis puslulalis aperientibus. Mycelium immersum inlracellulosum, 1--6 IJm diametro, ramosum, seplatum, The mycobiota of Lanfana camara hyalinum vel pallidum brunneum. Mycelium superficiale absens. Conidiomata acervularia, hypophylla, subepidermalia, in catervis irregulariter densis rotundatis aggregata, 41-218 x 27-136 ｾｭ［ crassis, hyalinis. Conidiophora e dehiscentia; parietibus 3-7 ｾ ｭ pseudoparenchyma basali orientia, cylindrica, ereda 9-14 x 1-7 ｾｭＬ 1-4 septata, ramosa, hyalina, laevia. Cellulae conidiogenae terminales, integratae, holoblasticae, cylindricae, hyalinae. Loci conidiogeni minuti. Conidia mucosa, holoblastica, a1iquando acropetala catenae brevis (usque 3 conidia) formantia, cylindrica vel fusifonnia, 15-28 x 2-5 ｾｭＬ apice obtuso, base obconico vel truncato, 1-2 ｾ ｭ lato, 0-3 septata, guttulata, hyalina, laevia. Paraphyses filifonnes, hyalinae, aseptatae, ad apices inflatae. HoJotypus: IMI 345375 in Lantana camara, lacupiranguinha, Sao Paulo, Brasilia, 26 Aug. 1989, R. W. Barreto & H. C. Evans. Lesions on living leaves; up to 25 mm wide, subcircular, chlorotic becoming brown necrotic, coalescing, the groups of acervular conidiomata opening as numerous blisters breaking open to liberate slimy masses of hyaline conidia. Infemal mycelium intracellular, 1-6 ｾ diam., branched, septate, hyaline to light brown. Exfernal mycelium absent. Conidiomafa acervular, hypophyllous, subepidermal, in dense roundish groups, 41-218 x 27-136 lAm in section; wall 2 cells thick, 3-7 lAm, hyaline. Dehiscence irregular. Conidiophores formed from the basal pseudoparenchyma, cylindrical, 9-14 x 1-7 lAm, erect, 1-4 septate, branched, hyaline, smooth-walled. Conidiogenous cells terminal, integrated, holoblastic, cylindrical, hyaline. Conidiogenous loci inconspicuous, minute. Conidia slimy, holoblastic, occasionally forming short, acropetalous chains of up to 3 conidia, cylindrical to fusiform, 15-28 x 2-5 lAm, apex obtuse, base obconic to truncate, wide, 0--3 septate, guttulate, hyaline, smooth. Paraphyses filiform, hyaline, aseptate, swollen at apices. Holofype: IMI 345375, on Lanfana camara, Jacupiranguinha, Sao Paulo, 26 Aug. 1989, R. W. Barreto & H. C. Evans. Parafype: IMI 363648, on L. camara, PinheiraL RJ, 29 Aug. 1994, R. W. Barreto. The original descriptions of Sepforia lanfanijolii Bat. & J. L. Bezerra (Batista ef a/., 1959) and Sepforia lanfanae Garman (1915) were compared with the present fungus. Both descriptions mention the presence of well-defined pycnidial conidiomata. This feature, together with considerable differences in spore morphology, indicated that the species was distinct and that it did not fit well into the concept of Sepforia. Similarities are shown with Sepfogloeum Sacco and Phloeosporella Hahn., but the conidiogenous cells in the former genus are enteroblastic phialides with prominent periclinal thickening (Sutton, 1980). In addition, conidia in this genus carry away the upper part of the conidiogenous cell as a basal, open-ended frill or appendage. In Phloeosporella, conidiophores and paraphyses are absent (Sutton, 1980). According to B. C. Sutton (pers. comm.), the closest related genera are Sepforiella Oudem. and Alloneoffiospora Nag Raj, but both are pycnidial and the spores have apical, gelatinous appendages; the best solution, therefore, was to introduce a new generic name. M. mucilaginosus is difficult to detect in the field since it is frequently in association and masked by Mycovellosiella lanfanae and Prospodium fuberculafum. However, it was associated with severe defoliation in the states of Sao Paulo and Rio de Janeiro at higher altitudes and thus appears to 780 show potential as a classical biological control agent of L. camara in subtropical climates. Anhellia lantanae (Henn.) Arx, Persoonia 2: 433 (1963). Trabutia lanfanae Henn., Hedwigia 34: 108 (1895). Agostaea lanfanae (Henn.) Theiss. & Syd., Annales mycologici 13: 359 (1915). This fungus was collected only once at a site at Saquarema, RJ, and did not cause any Significant damage. Although the specimen was sterile, the presence of the distinctive stroma suggested its identity as Anhellia lanfanae (Arx, 1963). This fungus is known only from the original description based on material from the state of Sao Paulo on an unidentified Lanfana sp. Subsequent collections have since been made of fertile ascomata associated with lesions of Prospodium fuberculatum. Its habit appears to be that of a mycoparasite rather than a primary pathogen. GENERAL DISCUSSION The discipline of biological control of weeds with fungal pathogens, with the two main approaches, classical and mycoherbicide, is beginning to prove its potential and several successful projects have already been implemented and others are in progress. These recent advances in the field of weed biocontrol with fungi have been documented for both the classical (Watson, 1991) and the mycoherbicidal approach (Charudattan, 1991). No indication is given in any of these references of attempted uses of pathogenic fungi against Lanfana camara in biocontrol programmes. This is surprising considering the number of projects already undertaken against this target weed (Julien, 1992). The fungi described here represent only a limited portion of the pathogenic mycoflora of L. camara. A literature and herbarium accessions search (Table 1) yielded a total of 53 suspected fungal pathogens recorded on L. camara. Only nine fungal pathogens were found in the field during the Brazilian survey, six of which were already known and three are newly described. The aim of this work was not the preparation of a complete picture of the mycobiota of L. camara in Brazil. Hence, most fungi which were either weak, opportunistic or non-specific pathogens, and therefore of no interest for weed biocontroL were excluded from this work. Such criteria could not be adopted for the preparation of Table 1, because of the difficulties involved in deciding on pathogenic status based only on the information (often scanty and imprecise) given in the literature. For example, Mukerji & Juneja (1975) give records of nearly 30 species of fungi associated with L. camara in India, most of which were obtained from dead or moribund tissues. Such obvious saprotrophs or opportunistic pathogens are of no interest for biocontrol and hence were excluded from Table 1. Even so, 26 entries were from countries or regions of the world where L. camara is an alien. Twenty-one species (plus one variety) were exclusive to the native range, whilst only six were common to both situations. The Old World fungi clearly form an assemblage of non-specific, opportunistic 781 R. W. Barreto, H. C. Evans and Carol A. Ellison pathogens. The small number of fungal pathogens present in both native and exotic range of the weed is an important indication of a continuing isolation of the New World's pathogenic mycoflora of L. camara. This represents a clear encouragement for classical introductions of fungal pathogens and several of the fungi discussed here appear to be suitable candidates. Ceratobasidium lantanae-camarae, Mycovellosiella lantanae var. lantanae, Prospodium tuberculatum and Micropustulomyces mucilaginosus are certainly worthy of investigation for their potential as biocontrol agents of L. camara, either individually or in combination, or in association with arthropod natural enemies. Puccinia lantanae could be of interest for introduction into warmer regions of the exotic range of L. camara. Many other fungal pathogens are likely to be found if the survey is extended to cover a wider part of the native range of L. camara. Almost a century of effort by entomologists on biocontrol has not solved the L. camara problem on a global scale, but it is possible that the exploitation of fungal pathogens may yield more positive results. ClBC (1982). Biological control of Lantana camara. Commonwealth Institute of Biological Control, Status Paper 3, 1-4. Ciferri, R. (1961). MycoRora Domingensis Integrata. Quaderno Instituto Botanico della Universita Pavia 19, 1-537. Giliers, C. J, &: Neser, S. (1991). Biological control of Lantana camara 01erbenaceae) in South Africa. Agriculture, Ecosystems &: Environment 37, 57-75. Costa, M. E. P. &: Camara, E. S. (1955). Species a1iquae Mycologicae Lusitaniae. Agronomia Lusitana 17. 153-165. Cummins, G. B. (1940). The genus Prospodium (Uredinales). Llnydia 3, 1-78. Cummins, G. B. (1959). Illustrated Genera of Rust Fungi. Burgess Publ. Co.: Minneapolis, MN, U.S.A. Dade, H. A. (1940). A revised list of Gold Coast fungi and plant diseases. Kew Bulletin 6, 205-247. Dale, W. T. (1955). A preliminary list of Jamaican Uredinales. Mycological Papers 60, 1-21. Deighton, F. C. (1974). Studies on Cercospora and allied genera. 5. Mycovellosieila Rangel and new species of Ramuiariopsis. Mycological Papers 137, 1-75. Eriksson, J. &: Ryvarden, L (1973). The Corticiaceae of North Europe, Vol. 2. FungiRora: Oslo, Norway. Evans, H. C (1987). Fungal pathogens of some subtropical and tropical weeds and the possibilities for biological control. Biocontrol News and Information 8,7-30. &: Tomley, A J. (1994). Studies on the rust Maravalia cryptostegiae, a potential biological control agent of rubber-vine weed, Cryptostegia grandif/ora (Asclepiadaceae: Periplocoideae), in Australia. III. Host range. Mycopatholngia 126, 93-108. Evans, H. C This work forms part of a research project submitted as a Ph.D. thesis to the University of Reading by R. W. B., who would like to thank: Dr M. W. Dick for guidance; Drs B. C. Sutton, J. David and E. Mordue for their suggestions; Ms G. 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