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The Genera Cylindrocladium and Cylindrocladiella in South Africa, with Special Reference to Forest Nurseries P

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The Genera Cylindrocladium and Cylindrocladiella in South Africa, with Special Reference to Forest Nurseries P r.; Review paper The Genera Cylindrocladium and Cylindrocladiella in South Africa, with Special Reference to Forest Nurseries P. W. Crousl, A.J.L. Phillips I and M.J. Wingfield2 'Plant Protection Research Institute, Private Bag XI34, Pretoria 0001 and 'Department of Microbiology and Biochemistry, University of the Orange Free State, P. O. Box 339, Bloemfontein 9300 SYNOPSIS Species of Cylindrocladium and Cylindrocladiella are widely distributed throughout the world and are known to be important pathogens of numerous angiosperm and gymnosperm hosts. Several species are notorious pathogens in nurseries, and have frequently been found on tree genera such as Eucalyptus, Pinus and Acacia. This paper provides a review of current knowledge, identifies areas of future research, and outlines the importance of Cylindrocladium and Cylindrocladiella in South African forestry. INTRODUCTION has been reported from Eucalyptus in South Africa (Lundquist and Baxter, 1985). Additional local hosts The genus Cylindrocladium was erected in 1892 by for C. scoparium include Acacia spp. (Doidge, 1950; Morgan, with C. scaparium Morgan as the type. This Hagemann and Rose, 1988), Persea americana Mill. species was found on a dead pod of honey locust (Gle- (Darvas, 1978), Medicago truncatula Gaertn. (Lam- ditschia triacanthos L.) in Ohio (Morgan, 1892). Subse- precht, 1986) and Syncarpia glomulifera (Smith) quently, Graves (1915) isolated the same fungus from Niedz. (PREM 45419). Darvas et al. (1978) found C. roots of Pinus strobus L., P. resinosa Ai!. and Tsuga scoparium and Ca. parva (Anderson) Boesewinkel to canadensis L. but failed to induce symptoms through be pathogenic on pines. The latter fungus was subse- artificialinoculations. He, therefore, assumed it to be quently found on roots of avocado (Darvas, 1978), saprophytic. Massey (1917) and Anderson (1919) were Protea aurea (N.L. Burm.) Rourke (PREM 45440) the first to show that C. scoparium could cause disease. and has recently been isolated by us from roots of Jackson (1938) associated this fungus with damping- Acacia mearnsii de Wild, Pinus radiata D. Don. and off, root rot and crown canker of several conifer forest litter. Although Sharma and Mohanan (1982) species. The fungus has subsequently been found to be reported Ca. parva as pathogenic to Eucalyptus, we a devastating pathogen of numerous hosts (Cox, 1953; have found it growing saprophytically on Eucalyptus Bugbee and Anderson, 1963a, 1963b; Bertus, 1976a, leaf litter. These observations support similar findings 1976b). Gibson (1975) concluded that C. scaparium is that, under certain conditions, Cylindrocladium spp. probably one of the most serious nursery pathogens of can occur as saprophytes (Boedijn and Reitsma, 1950; eucalypts at all growth stages. At that time it had French and Menge, 1978). however not been noticed on African Eucalyptus The genera Cylmdrocladium and Cylindrocladiella stock. are hardly known in South Africa. Preliminary surveys Boesewinkel (1982a) selected several small-spored have, however, indicated that a number of species species in the genus Cylindrocladium (C.), and placed occur in this country, and that they are potentially them in a new genus, Cylindrocladiella Boesewinkel important pathogens of various plants, especially Euca- (Ca.) . This genus currently comprises five species lyptus spp. No previous review of information pertain- (Boesewinkel, 1982a, 1982b), three of which have been ing to the two genera has been published. Therefore, reported as pathogens of Eucalyptus (Batista et al., this paper reviews current knowledge of Cylindrocla- 1965; Sharma and Mohanan, 1982; Mohanan and dium and Cylindrocladiella and considers their import- Sharma, 1985a). ance in forest nurseries in South Africa. Future areas At present, approximately 33 Cylindrocladium spp. of research on the pathogenic species are also outlined. have been described (Boedijn and Reitsma, 1950; Al- fieri et al., 1970; Gill et al., 1971; Panwar and Bohra, 1974; Hunter and Barnett, 1978; Schoulties et al., TAXONOMY AND MORPHOLOGICAL 1982; EI-Gholl et al., 1986, 1989), 12 of which are CHARACTERISTICS pathogenic to Eucalyptus (Peerally, 1974a, 1974b; Al- meida and Bolkan, 1981a; Sharma and Mohanan, Cylindrocladium, together with other closely related 1982; Mohanan and Sharma, 1985a; EI-Gholl, et al., genera such as Cylindrocladiella, Gliocladium Corda, 1986). Only one Cylindrocladium sp., C. scoparium, Cylindrocarpon Wollemw. and Fusarium Link: Fr. are Suid-Afrikaanse Bosboutydskrif-nr. 157, funie 1991 69 grouped in the order Hypocreales. Cylindrocladium characteristic shape, called the vesicle. Chlamydo- has a Calonectria de Not. teleomorph, while the latter spores occur in clusters and form microclerotia in soil, genera have Nee/ria Fr. states (Booth, 1966; Rossman, debris, host tissue and culture media. 1979a, 1979b, 1983). Cylindrocladiella is, at present recognised by having Apart from cultural characteristics, the main criteria small, cylindrical 0-1 septate spores, borne on mono- on which identifications of Cylindrocladium and Cylin- phialides. Conidiophores can either be subverticillate drocladiella species are based include conidial dimen- or penicillate. In the latter case the branches are usu- sions and septation, shape and size of the vesicle, ally arranged around a central stipe, which " non-sep- characteristics of the stipe, phialides, branching habit tate, but separated from a basal cell by a septum. A and individual branch dimensions. Species differentia- thin-walled vesicle is formed at its apex, and this can tion is complicated because of plasticity of the fungus develop a septum with age. Chlamydospores are more on different media and under various environmental frequently arranged in chains than in clusters (Boese- conditions (Zumpetta, 1976; Hunter and Barnett, winkel, 1982a): 1978). Cylindrocladium is characterised by having species with hyaline, cylindrical conidia, 1-7 (or more) sep- COMMONL Y ACCEPTED SPECIES tate, with obtuse ends (Hunter and Barnett, 1978), mostly encased in an irregular mucilaginous matrix. Both C. scoparium and Ca. parva occur in forestry Conidia form on monophialides, which occur singly or regions of the Cape, Transvaal and Natal. Although in groups of up to five on penicillate branches of the various other species have been found in South Africa, conidiophore. These branches arise laterally from cen- they will be dealt with in a subsequent study. Species tral or lateral specialised hyphae, referred to as the described in Cylindrocladium and Cylindrocladiella stipe, being septate, branched or unbranched, giving have been found to have a wide geographic distri but ion rise to a fertile, terminal, thin-walled structure of and host range (Tables 1 and 2). TABLE 1. Host range, geographic distribution, symptoms and literature pertaining to Cylindrocladium spp. Geographic References on General Fungus lIosts Eucalyptus spp. Symptoms distribution control references Cylindrocladium avesi- Georgia, Flor- /lex spp., Leaf spot, Gill, Alfieri & Sobers, culatum ida, USA Rhododendron twig die-back, 1971; Schubert, EI- Gill, Alfieri & Sobers obtusum defoliation Gholl. Alfieri & Teleomorph: Calonec- Pyranchanta Schoulties, 1989; Sob. tria avesiculata coccinea ers & Alfieri, 1972 Schubert, EI-Gholl, Al- fieri & Schoulties Cylindrocladium brassi- India Brassica Not given Panwar & Bohra, 1974 cae camprestis Panwar & Bohra Cylindrocladium brazil- Brazil Eucalyptus spp. E. alba, Damping-off, Batista, 1951; Peerally, iensis E. citriadora, die-hack of 1974g (Batista & Ciferri) E. grandis, adult trees Peerally E. saligna Synonym: C. scopa- rium Morgan var. braziliensis Batiste & Ciferri Cylindrocladium cande- Brazil Annona sp., Leaf spot Viegas. 1946 labrum Luma sp. Viegas Cylindrocladium citri Florida, USA Citrus sinensis Fruit decay Boedijn & Reitsma, (Fawcett & Klotz) Boe- 1950; dijn & Reitsma Fawcett & Klotz, 1937; Synonym: Cande/- Schoulties, EI-Gholl & spora citri Fawcett & Alfieri, 1982 Klotz 70 South African Forestry Journal- No. 157, June 1991 TABLE 1. Host range, geographic distribution, symptoms and literature pertaining to CylindrocIadium spp. (Conld) Geographic References on General Fungus Hosts Eucalyptus spp. S)'mptoms distribution control references Cylindrocladium clava- Brazil, India, Arachis hypo- E. cloeziana, Root rot. seed- Hodges & May, Almeida & Bolkan turn Mauritius gueu E. grandis, ling and shoot 1971; 1981b; Bolkan, Di- Hodges & May Araucaria E. microcorys, blight, stem Rattan & anese, Ribeiro & Al- agllslifolia, E. paniculala, canker, leaf Dhanda, 1985; meida, 1980; Balkan, Capsinum E. mligna, spot Rattan, Dhanda Ribeiro & Almeida, frutescens, E. lereticornis & Rhandhawa, 1981; Dianese, Ribeiro Cicer arietinum, 1983 & Urben, 1986; Di. Eucalyptus spp., anese, Ribeiro, Glycine max, Sharma, Ferreira & Pinus caribaea, Urban, 1987; Hodges, P. elliottii Reis & May, 1976; P. imularis, Hodges, Reis & May, P. mochoacana, 1976; Lopes & Reif- P. montesumae, schneider, 1982; P. oocarpa, Mohanan & Sharma, P. paluslris, 1985a. 1986; Oak. & P. patula, Uchida, 1982; Peerally, Pisum salivum, 1974b. Solanum tubero- sum Cylindrocladium col. Australia, India Annona relicu- E. robusla Leaf spot, fruit Hutton & Sanewski, hauni; Peerally Mauritius lala, ,ot 1989; Peerally, 1973; Teleomorph: Colo- Callislemon Peerally 1974j; Siddera- neClria colhouni; lanceolatus, maiah, 1988 Peerally Canavalia ensiformis, Camellia sinen- sis, Eucalyplus sp. Cy/indrocladium crota- Bra z ii, Sri Acacia sp.. E. camaldulen- Root rot.
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