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44<br />

<strong>Plant</strong> <strong>Pathology</strong> & <strong>Quarantine</strong> — Doi 10.5943/ppq/2/1/7<br />

Comprehensive check list of Cercosporoid fungi from Iran<br />

Bakhshi M 3 , Arzanlou M 1* and Babai-Ahari A 2<br />

1<br />

Assistant Professor of <strong>Plant</strong> <strong>Pathology</strong> and Mycology, <strong>Plant</strong> Protection Department, Faculty of Agriculture, University<br />

of Tabriz, PO Box: 5166614766, Iran.<br />

2<br />

Professors of <strong>Plant</strong> <strong>Pathology</strong> and Mycology, <strong>Plant</strong> Protection Department, Faculty of Agriculture, University of<br />

Tabriz, PO Box: 5166614766, Iran.<br />

3<br />

PhD Student of <strong>Plant</strong> <strong>Pathology</strong>, <strong>Plant</strong> Protection Department, Faculty of Agriculture, University of Tabriz, PO Box:<br />

5166614766, Iran.<br />

Bakhshi M, Arzanlou M and Babai-Ahari A 2012 – Comprehensive check list of Cercosporoid<br />

fungi from Iran. <strong>Plant</strong> <strong>Pathology</strong> & <strong>Quarantine</strong> 2(1), 44-55, doi 10.5943/ppq/2/1/7<br />

Little is known about the biodiversity of cercosporoid fungi in Iran. We provide a comprehensive<br />

literature-based checklist for 89 cercosporoid species known to occur on different plant species in<br />

Iran. The complete annotated list covers 29 Cercospora species, 24 Ramularia, 21 Passalora and<br />

15 Pseudocercospora.<br />

Key words – hyphomycetes – taxonomy – mitosporic fungi<br />

Article Information<br />

Received 5 March 2012<br />

Accepted 6 March 2012<br />

Published online 10 April 2012<br />

*Corresponding author: Mahdi Arzanlou – e-mail – Arzanlou@hotmail.com<br />

Introduction<br />

Cercosporoid fungi are generally<br />

associated with leaf spots, but also can cause<br />

necrotic lesions on flowers, fruits, bracts, seeds<br />

and pedicels of numerous hosts in a wide range<br />

of climates worldwide (Agrios 2005, Crous et<br />

al. 2000, Goodwin et al. 2001), encompassing<br />

economically important pathogens of major<br />

agricultural crops such as cereals, vegetables,<br />

ornamentals, forest trees, grasses and many<br />

others (Goodwin et al. 2001). Furthermore, a<br />

number of cercosporoid fungi are considered as<br />

potential biocontrol agents of weeds (Morris &<br />

Crous 1994).<br />

The genus Cercospora was established<br />

by Fresenius in 1863. Since then many species<br />

have been assigned to this genus and there has<br />

been a significant expansion of the generic<br />

concept, including numerous Passalora-like<br />

fungi (Braun & Melnik 1997). The first<br />

monograph of Cercospora Fresen. s. lato. was<br />

published by Chupp (1954). Later, Deighton<br />

(e.g., 1967, 1976), Braun (1995), Braun &<br />

Melnik (1997) and then other authors narrowed<br />

the generic concept of Cercospora s. lato. and<br />

divided it into smaller units. Crous & Braun<br />

(2003) in their revision of cercosporoid fungi<br />

based on molecular sequence analyses and<br />

reassessment of morphological characters,<br />

recognized four true cercosporoid genera, viz.<br />

Cercospora, Pseudocercospora Speg., Passalora<br />

Fr. and Stenella Syd., and several other<br />

morphologically similar genera.<br />

Cercospora and allied genera have<br />

traditionally been treated as anamorphs of the<br />

ascomycetous genus Mycosphaerella Johanson,<br />

(e.g., Braun & Melnik 1997, Kim & Shin 1998,<br />

Crous & Braun 2003). However, a teleomorph<br />

connection for the majority of cercosporoid<br />

fungi still remains unknown, and a Mycosphaerella<br />

state has been proven for only a few<br />

species.<br />

With the advent of molecular<br />

techniques in recent years, sequence data from<br />

different protein coding and non-coding of<br />

genomic regions have widely been applied to


figure out phylogenetic relationships amongst<br />

wide arrays of fungal groups at different<br />

taxonomic levels of interest (Arzanlou 2007,<br />

Arzanlou 2008, Arzanlou 2010, Crous et al.<br />

2000, Crous et al 2009). Phylogenetic analysis<br />

based on concordance of multiple sequence<br />

data have placed Cercospora and allied genera<br />

in Mycosphaerella clade (Mycosphaerellaceae,<br />

Capnodiales, Dothideomycetidae) (Crous et al.<br />

2007, Crous et al. 2009). The genus Mycosphaerella<br />

is one of the largest genera in<br />

Ascomycetes and comprises several thousand<br />

species (Crous et al. 2001, Aptroot 2006, Crous<br />

et al. 2009).<br />

Contrary to earlier belief in monophyly<br />

of Mycosphaerella, recent work of Crous and<br />

co-workers on phylogeny of Capnodiales<br />

revealed Mycosphaerella to be polyphyletic,<br />

and it has been split into several families; those<br />

members of Mycosphaerellaceae, Teratosphaeriaceae<br />

and Schizothyriaceae have plant<br />

pathological relevance (Crous et al. 2007,<br />

2009). A bewildering diversity of anamorphs<br />

exists in these lineages and up to 30 anamorph<br />

genera have now been linked to<br />

Mycosphaerella (Crous et al. 2007, 2009,<br />

Arzanlou et al. 2007, 2008, Arzanlou &<br />

Bakhshi 2012). However, recent phylogenetic<br />

analysis based on multiple sequence data sets<br />

have shown this to be incorrect.<br />

Mycosphaerellaceae, in fact, consists of<br />

numerous genera with morphologically<br />

conserved Mycosphaerella-like teleomorphs,<br />

and distinct anamorphs (Crous et al. 2007,<br />

Crous et al. 2009). Species of Mycosphaerella<br />

have a worldwide distribution from tropical<br />

and subtropical to warm and cool regions and<br />

have adapted to different ecological niches as<br />

saprobes, plant pathogens or endophytes (Farr<br />

et al. 1995, Goodwin et al. 2001, Crous et al.<br />

2009).<br />

Cercosporoid fungi are common and<br />

widespread on a wide range of vascular plants,<br />

especially in tropical regions. The mainland of<br />

Iran covers diverse climatic zones with a great<br />

biodiversity of vascular plants and,<br />

accordingly, a high diversity of foliicolous<br />

fungi. The exploration of this fungal group is,<br />

however, far from being complete and the<br />

cercosporoid hyphomycetes from Iran are still<br />

insufficiently known. Therefore, a comprehensive<br />

examination of cercosporoid fungi in<br />

<strong>Plant</strong> <strong>Pathology</strong> & <strong>Quarantine</strong> — Doi 10.5943/ppq/2/1/7<br />

Iran, based on the current taxonomic<br />

classification (Crous & Braun 2003), is<br />

urgently needed. In order to contribute to the<br />

knowledge of this group of fungi in Iran, and in<br />

the hope that what is reported here will<br />

stimulate other researchers to study the<br />

diversity of cercosporoid fungi, we have<br />

assembled a checklist of these fungi recorded<br />

to date from Iran.<br />

Materials and Methods<br />

List of species<br />

The list of cercosporoid fungi was<br />

compiled using reports available in the<br />

literature. Most of the quoted works are the<br />

result of field research by Iranian mycologists,<br />

although a small number of reports have been<br />

documented by foreign investigators. The list<br />

includes cercosporoid species together with<br />

their host species from which they have been<br />

collected. The fungal nomenclature and<br />

taxonomy follows Crous & Braun (2003).<br />

Species names used in the original<br />

publications, which deviate from those in the<br />

latter publication, are cited as synonyms. The<br />

checklist is organised alphabetically by genus<br />

and species name.<br />

Results<br />

A list containing 29 species of<br />

Cercospora, 21 species of Passalora, 22<br />

species of Ramularia and 15 species of<br />

Pseudocercospora is given (Table 1).<br />

Cercospora species have been collected on 50<br />

host plants belonging to 41 genera and 28<br />

families. The highest numbers of cercosporoid<br />

parasites were recorded on representatives of<br />

the Fabaceae (with nine species), Malvaceae<br />

(with four species) and Euphorbiaceae (with<br />

three species). Passalora species were<br />

recordered on 29 host plants belonging to 23<br />

genera and 14 families, including the<br />

Asteraceae (with four species) and Apiaceae<br />

(with three species), which represent the host<br />

plant families with most Passalora species.<br />

Pseudocercospora species were recorded on 18<br />

host plants belonging to 15 genera and 12<br />

families. The highest numbers of<br />

Pseudocercospora parasites were recorded on<br />

the Solanaceae (three species). Ramularia<br />

species were collected on 30 host plants<br />

45


46<br />

Table 1 Cercosporiod fungi known from Iran.<br />

<strong>Plant</strong> <strong>Pathology</strong> & <strong>Quarantine</strong> — Doi 10.5943/ppq/2/1/7<br />

Species Hosts References<br />

Cercospora<br />

- acnidae Ellis & Everh.<br />

Amaranthus chlorostachys var. chlorostachys Willd. & Thell. Pirnia et al. 2010<br />

- althaeina Sacc. Althaea rosea Cav., Althaea sp. Petrak 1956, Scharif & Ershad 1966, Ebrahimi & Minnasian 1973<br />

- apii Fresen. Abutilon theophrasti Medik., Euphorbia heterophylla L.,<br />

Pelargonium zonale, , Solanum lycopersicum L., Vigna<br />

sinensis (L.) Walp., Zanthedeschia aethiopica (L.) Spreng.<br />

Pirnia et al. 2010<br />

- beticola Sacc. Beta maritima L., B. vulgaris L., Spinacia oleracea L. Esfandiari 1946a, Esfandiari 1947, Khabiri 1952, 1958, Golato 1960,<br />

Eskandari 1964, Scharif & Ershad 1966, Eskandari et al. 1969, Vinnot-<br />

Bourgin et al. 1969, Altman et al. 1972, Ebrahimi & Minnasian 1973,<br />

Vaziri 1973, Alian et al. 2008<br />

- canescens Ellis & Martin Phaseolus vulgaris L., Vigna sinensis (L.) Endl Altman et al. 1972, Ershad 1995<br />

- cheiranthi Sacc. Cheiranthus cheiri L. Esfandiari 1948, Scharif & Ershad 1966<br />

- citrullina Cooke Citrullus vulgaris Schrad. ex Eckl. & Zeyh. Scharif & Ershad 1966<br />

- datiscicola Esfand. Datisca cannabina L. Esfandiari 1951<br />

- elaeagni Heald & Wolf Elaeagnus angustifolia L. Ershad 1995<br />

- fukushiana (Mat.) Yam. Impatiens balsamina L. Hedjaroude 1983<br />

- gerberae Chupp & Viegas Gerbera jamesonii Hook. Ershad 1995<br />

- hydrangea Ellis & Everh. Hydrangea hortensia Siebold. Hedjaroude 1983<br />

- iridis Chupp Iris sp. Pirnia et al. 2010<br />

-kikuchii Matsumoto & Tomoy Glycine max (L.) Merr. Zad 1979, Majidieh-Ghassemi 198<br />

-lactuca-sativae Sawada = C. longissima<br />

Cooke & Ellis<br />

Lactuca sativa L. Banihashemi 1985, Pirnia et al. 2010<br />

- malayensis Stev. & Solh. Hibiscus esculentus L. Petrak 1956, Scharif & Ershad 1966<br />

- medicaginis Ellis & Everh. Medicago sp. Scharif & Ershad 1966<br />

- mercurialis Pass. Mercurialis annua L. Pirnia et al. 2010<br />

-musae Zimm. Musa sapientum L. Amani et al. 2010<br />

- neriicola Ershad Nerium oleander L. Ershad 2002<br />

-ricinella Sacc. & Berl. Ricinus communis L. Petrak 1956, Khabiri 1958, Scharif & Ershad 1966<br />

-scharifii Petr. Rosa sp. Petrak 1956<br />

- sorghi Ellis & Everh. Sorghum bicolor (L.) Moench, S. halepense Pers. Mehrian & Rajoo 2004, Mehrian 2006, Pirnia et al. 2010<br />

-taurica Tranzsch. Heliotropium sp. Petrak 1939<br />

-traversiana Sacc. Trigonella foenum-graceum L. Scharif & Ershad 1966 as Cercospora sp., Vinnot- Bourgin et al. 1970,<br />

Ershad 1995<br />

- viola Sacc. Viola odorata L., V. sintenisii Becker, V. sylvestris Lam., Esfandiari 1948, Khabiri 1952, Khabiri 1958, Scharif & Ershad 1966,<br />

Viola sp.<br />

Vinnot-Bourgin et al. 1970, Ershad 1995<br />

- zebrina Pass. Medicago sativa L. Ebrahimi & Minassian 1973


<strong>Plant</strong> <strong>Pathology</strong> & <strong>Quarantine</strong> — Doi 10.5943/ppq/2/1/7<br />

Species Hosts References<br />

-zonata Wint. = C. fabae Fautr. Vicia faba L. Esfandiari 1948, Scharif & Ershad 1966, Ebrahimi & Minnasian 1973,<br />

- Cercospora spp. Arachis hypogaea L., Capparis spinosa L., Citrus aurantium<br />

L., C. sinensis (L.) Osbeck., Citrus sp., Ficus carica L.,<br />

Gladiolus segetum Ker-Gawl., Glycine max (L.) Merr.,<br />

Hibiscus cannabinus L., Hydrangea hortensia Siebold.,<br />

Lactuca sativa L., Malus pumila Mill., <strong>Plant</strong>ago major L.,<br />

Hedjaroude 1976, Pirnia et al. 2010<br />

Afzali et al. 2010, Scharif 1953, Ebrahimi 1964, Scharif & Ershad 1966,<br />

Scharif & Akhavizadegan 1967, Daftari & Behdad 1968, Vaziri 1973,<br />

Ebrahimi & Minassian 1973, Roohibakhsh & Ershad 1997<br />

Punica granatum L.<br />

Passalora<br />

- bolleana (Thum.) Braun = Cercospora Ficus carica L. Petrak 1956, Scharif & Ershad 1966, Vinnot- Bourgin et al. 1970, Ershad<br />

bolleana (Thum.) Speg.<br />

1995<br />

- calotropidis (Ellis & Everh.) Braun =<br />

Cercospora calotropidis Ellis & Everh. = C.<br />

patouillardi Sacc.<br />

Calotropis procera Dryand Petrak 1953, Petrak 1956, Vinnot- Bourgin et al. 1970, Ershad 1995<br />

- calystegiae (Speg.) Braun = Cercospora<br />

calystegiae Speg.<br />

Calystegia sp. Hedjaroude 1983<br />

- carlinae (Sacc.) Braun = Cercospora<br />

carlinae Sacc.<br />

Calina sp. Petrak 1941<br />

- circumscissa (Sacc.) Braun = Cercospora Amygdalus communis L., Cerasus avium (L.) Moench, C. Petrak & Esfandiari 1941, Esfandiari 1946a, Khabiri 1958, Vinnotcircumscissa<br />

Sacc.<br />

vulgaris Miller, Prunus domestica L.<br />

Bourgin 1958, Scharif & Ershad 1966<br />

- cousiniae Petr. Cousinia nekarmanica Rech. f., Cousinia sp. Petrak 1949, Esfandiari 1951, Jorstad 1960, Ershad 1995<br />

- dubia (Riess) Braun = Cercospora dubia<br />

(Riess) Wint.<br />

Chenopodium brumale L. Hedjaroude 1976<br />

- fraxini (DC.) Arx Fraxinus excelsior L. Kalte et al. 2008<br />

-fraxinicola (Ershad) Braun & Crous =<br />

Cercospora fraxinicola Ershad<br />

Fraxinus rotundifolia Miller Ershad 2000<br />

- fulva (Cooke) Braun & Crous = Fulvia<br />

fulva (Cooke) Ciferri = Cladosporium<br />

fulvum Cooke.<br />

Lycopersicum esculentum Mill. Scharif & Ershad 1966, Ershad 1995<br />

- graminis (Fuckel) Hohn. Glyceria fluitans R.Br. Esfandiari 1948<br />

- kirchneri (Hegyi) Petr.<br />

Anethum graveolens L., Foeniculum vulgare Mill. Esfandiari & Petrak 1950, Esfandiari 1951, Petrak 1956, Scharif &<br />

Ershad 1966, Ebrahimi & Minassian 1973, Ershad 1995<br />

- microsora (Sacc.) Braun = Cercospora Tilia begonifolis Stev., Tilia sp. Petrak 1941, Esfandiari 1946b, Khabiri 1958, Scharif & Ershad 1966,<br />

microsora Sacc.<br />

Ershad 1995<br />

- miser (Karst.) Redhead, Vilgalys &<br />

Hopple = Coprinus miser Karst.<br />

Salix sp. Saber & Esmaeili Taheri 2002<br />

- phaeopappi Petr. Phaeopappus aucheri (DC.) Boiss; P. kotschyi (Boiss. &<br />

Heldr) Boiss<br />

Petrak 1939, Esfandiari 1948, Golato 1960<br />

47


Species Hosts References<br />

- punctum (Delacr.) Petzoldt = Cercospora<br />

Petroselini Sacc.<br />

Petroselinum sativum Hoffm. Ebrahimi & Minassian 1973<br />

- rhamni (Fuckel) Braun = Cercospora<br />

rhamni Fuckel<br />

Rhamnus frangula L. Vinnot-Bourgin 1958<br />

- rosae (Fuckel) Hohn. Rosa sp. Esfandiari 1948<br />

- scandicearum (Magnus) Braun = Anthriscus sylvestris Hoffm. Vinnot-Bourgin et al. 1970<br />

Cercospora scandicearum Magnus<br />

- scariola Syd.<br />

Scariola orientalis (Boiss) Sojak Petrak 1949, Esfandiari 1951<br />

- smilacis (Thum.) Braun = Cercospora<br />

smilacis Thum = C. smilacina Sacc.<br />

Pseudocercospora<br />

- abelmoschi (Ellis & Everh.) Deighton =<br />

Cercospora abelmoschi Ellis & Everh. = C.<br />

hibisci Tracy & Earle<br />

- atromarginalis (Atk.) Deighton =<br />

Cercospora atromarginalis Atk.<br />

- cavarae (Sacc. & D.Sacc.) Deighton =<br />

Cercospora cavarae Sacc. & D.Sacc.<br />

- cruenta (Sacc.) Deighton = C. cruenta<br />

Sacc.<br />

- fici (Heald & Wolf) Liu & Guo =<br />

Cercospora fici Heald & Wolf<br />

- fuligena (Roldan) Deighton<br />

- gomphrenae Sawada ex Goh & Hsieh =<br />

Cercospora gomphrenae<br />

- jujubae (Chowdhury) Khan & Shamsi =<br />

Cercospora jujubae Chowdhury<br />

-kaki Goh & Hsieh = Cercospora kaki Ellis<br />

& Everh.<br />

- phyllitidis (Hume) Braun & Crous =<br />

Cercospora petunia (Saito) Muller & Chupp<br />

- punicae (Henn.) Deighton = Cercospora<br />

punicae Henn.<br />

- rubi (Sacc.) Deighton = Cercospora rubi<br />

Sacc.<br />

- salvadorae (Maire) Deighton =<br />

Cercospora salvadorae Maire<br />

48<br />

<strong>Plant</strong> <strong>Pathology</strong> & <strong>Quarantine</strong> — Doi 10.5943/ppq/2/1/7<br />

Smilax aspera L., S. excels L. Petrak & Esfandiari 1941, Esfandiari 1946b, Khabiri 1958<br />

Hibiscus cannabinus L.<br />

Esfandiari 1947, 1948, Scharif & Ershad 1966, Scharif & Akhavizadegan<br />

1967 as Cercospora sp., Vinnot-Bourgin et al. 1970<br />

Solanum nigrum L. Vinnot-Bourgin et al. 1970, Ershad 1995<br />

Glycyrrhiza glabra L. Hedjaroude 1983<br />

Vigna sinensis (L.) Endl. Kaiser et al. 1968 as Cercospora sp., Vaziri 1973<br />

Ficus carica L. Scharif & Ershad 1966<br />

Solanum nigrum L. Physalis alkekengi L. Aghajani & Ahmadi 2010<br />

Gomphrena globosa L. Hedjaroude 1983<br />

Ziziphus spina-christi (L.) Willd. Ershad & Khosravi 1996<br />

Diospyros kaki L., D. lotus L. Petrak & Esfandiari 1941, Esfandiari 1946b, Khabiri 1958, Golato 1960,<br />

Scharif & Ershad 1966, Vinnot-Bourgin et al. 1970<br />

Petunia var. Hort. Ershad 1995<br />

Punica granatum L. Petrak & Esfandiari 1941, Esfandiari 1946b, Golato 1960, Ershad 1995<br />

Rubus sp. Hedjaroude 1976<br />

Salvadora persica L. Ershad 1990


<strong>Plant</strong> <strong>Pathology</strong> & <strong>Quarantine</strong> — Doi 10.5943/ppq/2/1/7<br />

Species Hosts References<br />

- sphaerella-eugeniae (Sacc.) Crous, Eugenia jambos L. Ershad 1990<br />

Alfenas & Barreto = Cercospora eugeniae<br />

(Rangel) Chupp-Tis<br />

- vitis (Lev.) Speg. = Cercospora vitis (Lev.) Vitis sylvestris Gmel, V. vinifera L. Esfandiari 1947, Vinnot-Bourgin et al. 1970, Hedjaroude 1976, Ershad<br />

Sacc.<br />

Ramularia<br />

1990<br />

-alpine (Massal.) Nannf.= Ovularia alpine<br />

Massal.<br />

Alchemilla sp. Petrak 1949, Esfandiari 1951<br />

-anchusae Massal. Anchusa italica Retz., A. ovate Lehm. Petrak 1953, Khabiri 1958, Vinnot-Bourgin et al. 1970, Moaven et al.<br />

2003<br />

- beccabungae Fautr. Veronica beccabunga L. Petrak & Esfandiari 1941<br />

- bornmulleriana (Magnus) Braun = Onobrychis sintenisii Bornm. Petrak 1939<br />

Ovularia bornmulleriana Magnus<br />

- cynarae Sacc.= R. carthami Zaprom. Carthamus oxyacantha M. Bieb., C. tinctorius L. Minassian 1971, Vinnot-Bourgin et al. 1969 as Cercospora carthami<br />

Sundar & Ramakr., Altman et al. 1972, Ebrahimi & Minassian 1973 as<br />

Ramularia sp.<br />

- geranii Fuckel var. geranii Geranium pyrenaicum L. Hedjaroude & Abbasi 2000<br />

- heraclei (Oud.) Sacc. Heracleum persicum Desf. ex Fischer, Heracleum sp. Esfandiari 1948, Ershad 1995<br />

- iranica Petr. Acantholimon sp. Petrak 1949, Esfandiari 1951<br />

- lamii Fuckel = R. leonuri Sacc. & Penz. = Leonurus cardiac L., Mentha arvensis L. Vinnot-Bourgin et al. 1969 as R. menthicola Sacc., Hedjaroude 1983,<br />

R. menthae Sacc.<br />

Ershad 1995, Anonymous 2005 as R. menthicola Sacc.<br />

- macrospora Fres. Campanula rapunculus L. Petrak & Esfandiari 1941, Esfandiari 1946b, Scharif & Ershad 1966<br />

- marrubii Massal.= R. sideritis Hollos. Sideritis sp. Petrak 1939<br />

- primulae Thum. Primula aqualis L. Aghapour et al. 2010<br />

- rhabdospora (Berk. & Broome) Nannf. =<br />

R. plantaginea Sacc.<br />

<strong>Plant</strong>ago lanceolata L. Esfandiari 1948<br />

- rubella (Bonord) Nannf. = Ovularia<br />

obliqua (Cooke) Oud.<br />

Rumex crispus L., Rumex sp. Esfandiari 1948, Vinnot- Bourgin et al. 1969, Ershad 1995<br />

- rubicola Ershad Rubus caesius L. Ershad 2000<br />

- rufomaculans Peck Polygonum sp. Khabiri 1958<br />

- rumicis Kalchbr. & Cooke = R. decipiens<br />

Ellis & Everh.<br />

Rumex crispus L., Rumex sp. Vinnot-Bourgin et al. 1970, Ershad 1995, Fotouhifar et al. 2003<br />

- rumicis-scutati Allesch. Rumex scutatus L. Petrak & Esfandiari 1941, Esfandiari 1946a, Esfandiari 1946b<br />

- sambucina Sacc. Sambucus ebulus L. Petrak & Esfandiari 1941, Esfandiari 1946b<br />

- simplex Pass. Ranynchulus oxyspermus Willd. Moaven et al. 2003<br />

- uredinicola Khodap. & Braun Melampsora sp. Khodaparast & Braun 2005<br />

- urticae Ces. Urtica dioica L., U. urens L. Esfandiari 1948, Vinnot-Bourgin 1958<br />

- valeriana (Speg.) Sacc. Valeriana sisymbrifolia Vahl. Petrak 1939<br />

Ramularia sp. Foeniculum vulgare Mill., Potentila sp., Rumex sp. Khabiri 1952, Vaziri 1973, Hedjaroude 1976<br />

49


elonging to 22 genera and 17 families,<br />

including the Polygonaceae (with five species)<br />

and Rosaceae (with three species), which<br />

represent the host plant families with most<br />

Ramularia species.<br />

Discussion<br />

Despite their widespread distribution,<br />

current knowledge on the cercosporoid fungi of<br />

Iran is scant. Hopefully, the information<br />

gathered here will provide both a reference<br />

work and be an incentive for further work<br />

aimed at disclosing the diversity of<br />

cercosporoid fungi in Iran.<br />

Since the description of the genus<br />

Cercospora, the taxonomy of this genus as well<br />

as the description of individual species within<br />

this group has proven burdensome. While<br />

Cercospora was defined at genus level by<br />

morphology, species definition was based<br />

largely on host association. A significant<br />

problem pertaining to the taxonomy of<br />

Cercospora is the degree of host-specificity of<br />

the various species. Most species are still<br />

defined based on host, and they are assumed to<br />

be host-specific or restricted at least to a family<br />

of plants (Chupp 1954). However, the<br />

tenability of many species may be called into<br />

question because some taxa, including C. apii,<br />

the type species of Cercospora, have been<br />

shown to be non host-specific (Crous & Braun<br />

2003).<br />

Most hyphomycetous genera linked to<br />

Mycosphaerella have conventionally been dealt<br />

with as part of the cercosporoid fungi (Crous &<br />

Braun 2003). In most cases cercosporoid fungi<br />

have been treated as asexual fungi, and<br />

teleomorphs have been confirmed for only a<br />

few species. Groenewald et al. (2006) detected<br />

the two mating type genes in approximately<br />

even proportions in C. beticola, C. zeae-maydis<br />

and C. zeina populations, and speculated that a<br />

sexual cycle may occur regularly in these<br />

species. However, the actual sexual stage was<br />

not observed. Hence, the application of the<br />

criterion of intersterility is also mainly limited<br />

in cercosporoid fungi.<br />

Iran is one of the richest biodiversity<br />

hotspots in the world. This is due to the variety<br />

of climatic zones ranging from subtropical and<br />

tropical rain forests to wetlands. Despite this<br />

wealth, however, little information is available,<br />

50<br />

<strong>Plant</strong> <strong>Pathology</strong> & <strong>Quarantine</strong> — Doi 10.5943/ppq/2/1/7<br />

regarding cercosporoid fungi and their<br />

distribution as well as host range in Iran. Most<br />

publications describe the species from<br />

herbarium material and no cultures of<br />

cercosporoid fungi are present in the country.<br />

It is difficult to rely solely on morphology<br />

for the species identification in this group.<br />

Since the combination of morphological and<br />

phylogenetic analyses of new proposed taxa in<br />

Cercospora complex is very important in order<br />

to avoid misidentification, in our opinion,<br />

future work should be focused on phylogenetic<br />

analysis of cercosporoid fungi. Hence, we are<br />

currently busy with phenotypic and molecular<br />

characterization of cercosporoid fungi of Iran<br />

based on single spore cultures.<br />

Acknowledgements<br />

The authors would like to thank the<br />

Research Deputy of the University of Tabriz,<br />

Iran for financial support.<br />

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53

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