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Phoma - Chap 00 Pre 13/4/04 10:49 Page I Phoma - Chap 00 Pre 13/4/04 10:49 Page i PHOMA IDENTIFICATION MANUAL Differentiation of Specific and Infra-specific Taxa in Culture [Result of many years’ experience at the diagnostic mycological section of the Dutch Plant Protection Service (PD)] G.H. Boerema, J. de Gruyter, M.E. Noordeloos and Maria E.C. Hamers Phoma - Chap 00 Pre 13/4/04 10:49 Page ii Phoma - Chap 00 Pre 13/4/04 10:49 Page iii PHOMA IDENTIFICATION MANUAL Differentiation of Specific and Infra-specific Taxa in Culture G.H. Boerema Zandvoort The Netherlands J. de Gruyter Plant Protection Service Wageningen The Netherlands M.E. Noordeloos National Herbarium of The Netherlands Leiden The Netherlands and Maria E.C. Hamers Plant Protection Service Wageningen The Netherlands CABI Publishing Phoma - Chap 00 Pre 13/4/04 10:49 Page iv CABI Publishing is a division of CAB International CABI Publishing CABI Publishing CAB International 875 Massachusetts Avenue Wallingford 7th Floor Oxfordshire OX10 8DE Cambridge, MA 02139 UK USA Tel: +44 (0)1491 832111 Tel: +1 617 395 4056 Fax: +44 (0)1491 833508 Fax: +1 617 354 6875 E-mail: [email protected] E-mail: [email protected] Website: www.cabi-publishing.org ©G.H. Boerema, J. de Gruyter, M.E. Noordeloos and M.E.C. Hamers 2004. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Phoma identification manual : differentiation of specific and infra-specific taxa in culture / G.H. Boerema ... [et al.]. p. cm. Includes bibliographical references and index. ISBN 0-85199-743-0 (alk. paper) 1. Phoma--Identification. 2. Phoma--Pictorial works. I. Boerema, G. H. (Gerhard H.) II. Title. QK625.S5P56 2004 579.5Ј5--dc21 2003012556 ISBN 0 85199 743 0 Typeset in 10pt Souvenir by Columns Design Ltd, Reading. Printed and bound in the UK by Biddles Ltd, King’s Lynn. 00Chap 00 Pre 19/4/04 14:34 Page v Contents Preface vii 1 Introduction 1 (with Figs 1–3) 2 Nomenclator of the Genus and its Sections 8 3 The Generic Characters 11 4 The Methods Used for Differentiation and Identification 14 5 Entries to the Differentiation of the Sections 19 (with Fig. 4) Key to the Sections Based on Characteristics In Vitro 20 Table of the Collective and Differential Features In Vitro 22 Synopsis of the Collective and Differential Characteristics of the Sections In Vivo 23 6 Notes on ‘Adjacent’ or ‘Convergent’ Genera 24 (with Fig. 5) 7 Entries to the Species per Section 30 8APhoma sect. Phoma (68 taxa) 32 (with Figs 6–13) 9BPhoma sect. Heterospora (17 taxa) 119 (with Figs 14–20) 10 C Phoma sect. Paraphoma (12 taxa) 158 (with Figs 21–23) 11 D Phoma sect. Peyronellaea (16 taxa) 179 (with Figs 24–28) 12 E Phoma sect. Phyllostictoides (41 taxa) 214 (with Figs 29–32) v Phoma - Chap 00 Pre 13/4/04 10:49 Page vi vi Contents 13 F Phoma sect. Sclerophomella (12 taxa) 297 (with Figs 33–35) 14 G Phoma sect. Plenodomus (32 taxa) 320 (with Figs 36–46) 15 H Phoma sect. Macrospora (9 taxa) 387 (with Figs 47–50) 16 I Phoma sect. Pilosa (2 taxa) 404 (with Fig. 51) 17 Miscellaneous (14 taxa) 411 References 426 Index of Fungus Names 446 Index of Substrata 467 Phoma - Chap 00 Pre 13/4/04 10:49 Page vii Preface This monographic study involves the cultural descriptions of 223 specific and infraspecific taxa of Phoma Sacc. emend. Boerema & G.J. Bollen, with 1146 syn- onyms in various Coelomycetes genera. They are divided into nine sections, which assists in their identification. Three of these sections include species with a Didymella teleomorph and one section includes two species with a Mycosphaerella teleomorph; another section shows a specific metagenetic relation with the genus Leptosphaeria and another with the genus Pleospora. Most of the species are described from isolates of European origin and refer to parasites as well as ubiquitous saprophytes. A smaller number of species is described from isolates made in other continents and refers to both parasitic and saprophytic species. Descriptions in vivo are only added when important for identification; species with a teleomorph are always described in vivo. References on diagnos- tic literature and short notes on ecology and distribution are given for each species. With one exception the keys refer only to the characteristics in vitro. Various tables on host relations or connections are given as well as indices to all fungal taxa and hosts mentioned. All well-known plant pathogenic and economically important Phoma species come up. Some examples: Ph. andigena (black potato blight) and Ph. foveata (potato gangrene), both included in the lists of quarantine organisms of the European Plant Protection Organization (EPPO), Ph. betae (black leg of beet and spinach), a common seed-borne pathogen, Ph. apiicola (phoma root and crown rot of celeriac and celery), Ph. terrestris (pink root of onion) and Ph. lingam (dry rot and canker or black leg of brassicas). Pathogens of pulse and forage crops such as Ph. sub- boltshauseri (blotch or leaf spot of beans and cowpea), Ph. medicaginis (black stem of lucerne) and Ph. pinodella (foot rot and black stem of pea and clover). vii Phoma - Chap 00 Pre 13/4/04 10:49 Page viii viii Preface On tomato plants Ph. destructiva (fruit rot, leaf and stem blight) and Ph. lycop- ersici (canker, stem and fruit rot). On florists’ chrysanthemums Ph. chrysan- themicola (root rot and basal stem rot) and Ph. ligulicola (ray blight, leaf and stem spot). On Amaryllidaceae Ph. narcissi (red spot diseases, leaf scorch). Important plurivorous opportunistic plant pathogens are also treated, e.g. Ph. exigua, Ph. macrostoma, Ph. multirostrata and Ph. sorghina (common in the tropics and subtropics). Phoma - Chap 01 13/4/04 10:49 Page 1 1 Introduction This study had an unplanned start at the diagnostic mycological section of the Dutch Plant Protection Service at Wageningen (PD) in the early 1960s with the examination of a sample of Brassica seed suspected of being infected with Phoma lingam (Tode: Fr.) Desm., the anamorph of the causal fungus of dry rot and canker or black leg. However, it appeared that the seed coat was contami- nated with two other species of Phoma (Boerema, 1962). This was established with a comparative study of isolates on agar media. The dominating species proved to be Phoma herbarum Westend. nomi- nated as type species of Phoma sensu Saccardo (nomen genericum conservan- dum) at the 8th International Botanical Congress at Paris in 1954. According to Saccardo’s taxonomic system, the genus Phoma refers to ‘pycnidia with one- celled hyaline conidia occurring on herbaceous stems’ (Saccardo, 1884). However, our study showed Ph. herbarum to be a ubiquitous saprophyte, occurring on very diverse substrata, not only on all parts of dead and dying herbaceous plants, but also on woody plants, in soil, water, milk, butter, paint, paper, etc. (Boerema, 1964). Since the experimental study by Hughes (1953) on Hyphomycetes, the conidiogenesis has been recognized as the most impor- tant taxonomic criterion in all conidial fungi. In mature pycnidia of Ph. herbarum and other similar Phoma species the hyaline unicellular conidia arise from less differentiated cells lining the pycnidial cavity in a kind of repetitive monopolar budding. Apart from some wall-thickening at the top of the conidio- genous cells, the light microscopy gave few details. In cooperation with the Agricultural Highschool at Wageningen (LH, now Wageningen University and Research, WUR) it was possible to study the conidiogenesis in Phoma spp. with transmission electron microscopy (Brewer & Boerema, 1965; Boerema & Bollen, 1975). The conidiogenesis appeared to begin with the formation of a papilla and breakage of the conidiogenous cell to leave a ‘collarette’ (periclinal thickening). The double layered wall of the first conidium arises as new, its mat- uration synchronous with the ontogeny; the wall building of successively pro- duced conidia starts with the basal layer of the septum remaining after © G.H. Boerema, J. de Gruyter, M.E. Noordeloos and M.E.C. Hamers 2004. Phoma Identification Manual (G.H. Boerema et al.) 1 01Chap 01 19/4/04 14:35 Page 2 2 Chapter 1 secession of the previous conidium, maturation still being synchronous with ontogeny (for details see Figs 1–3). The conidia are produced at about the same level by percurrent proliferation. This conidiogenesis, associated with abundant production of mucilage, is analogous with that in Hyphomycetes now recognized as phialidic (Kendrick, 1971) with blastic conidial ontogeny. This is more precisely termed false- or no-chain phialidic (Minter et al., 1982) as the conidia are produced in mucilaginous masses or chains (Boerema, 1964) that are not held together by wall material as in true phialides. On agar media Ph. herbarum shows specific cultural characteristics which very much simplify its identification. This appeared to be true also for other species of Phoma. In addition to characteristics of pycnidia and conidia, cul- tures give information about mycelial characters such as development of chlamydospores, and biochemical properties such as production of pigments and crystals. The importance of cultural characteristics for identifying and dif- ferentiating Phoma species was already noted by Wollenweber & Hochapfel (1936) in their study on fruit-rotting, and by Dennis (1946) in a study of Phoma isolates from various sources in comparison with those found in associ- ation with rots of potato tubers. Their in vitro studies provided valuable infor- mation about various plurivorous species of Phoma, described under many different names; see Boerema & Dorenbosch (1973) and Boerema (1976).
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