Genus Cercospora in Thailand: Taxonomy and Phylogeny (with a ...

Plant Pathology & Quarantine 

Genus Cercospora in Thailand: Taxonomy and Phylogeny (with a dichotomous 

key to species) 

To-Anun C 1* , Hidayat I 2 and Meeboon J 3 

1 

Department of Entomology and Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai, Thailand 

2 

Research Center for Biology, Indonesian Institute of Sciences (LIPI), JI. Raya Jakarta-bogor Km 46 Cibinong 16911, 

West Java, Indonesia 

3 

Faculty Of Agricultural Technology, Chiang Mai Rajabhat University, Sa Luang, Mae Rim District 50180, Chiang Mai, 

Thailand 

To-Anun C, Hidayat I, Meeboon J. 2011 – Genus Cercospora in Thailand: Taxonomy and 

Phylogeny (with a dichotomous key to species). Plant Pathology & Quarantine 1(1), 11–87. 

Cercospora Fresen. is one of the most importance genera of plant pathogenic fungi in agriculture 

and is commonly associated with leaf spots. The genus is a destructive plant pathogen and a major 

agent of crop losses worldwide as it is nearly universally pathogenic, occurring on a wide range of 

hosts in almost all major families of dicotyledonous, most monocotyledonous families, some 

gymnosperms and ferns. The information regarding Cercospora leaf spots in Thailand is scattered 

and mainly based on Chupp’s generic concepts. Therefore, this paper provides an update that 

includes synonyms, morphological descriptions, illustrations, host range, geographical distribution 

and literature related to the species. This will benefit mycologists, plant pathologists and quarantine 

officials who need to study this group of fungi. The present study represents a compilation of 52 

species of Cercospora s. str. associated with 29 families of host plants collected from several 

provinces in Thailand between 2004 and 2008. Twenty-four species represent C. apii s. lat. Plant 

families of Asteraceae, Amaranthaceae, Convolvulaceae, Euphorbiaceae, Fabaceae, Lamiaceae 

and Solanaceae, are commonly found infected with Cercospora s. str. Three species, Cercospora 

arecacearum, C. habenariicola and C. neobougainvilleae, have been validity published as new 

species from Thailand. 

Key words – diversity – hyphomycetes – leaf spot – taxonomy – Thailand. 

Article Information 

Received 24 November 2010 

Accepted 30 November 2010 

Published online 30 June 2011 

*Corresponding author: Chaiwat To-Anun – e-mail – agppi006@chiangmai.ac.th 

Introduction 

Overview of Genus Cercospora 

Cercospora, established by Fresenius 

(1863), is one of the most important genera of 

pathogenic fungi in agricultural field. The 

genus is a destructive plant pathogen throughout 

the world. Members of this genus are 

nearly universally pathogenic, occurring on a 

wide range of hosts in almost all major families 

of dicotyledonous, most monocotyledonous fa- 

milies, some gymnosperms and ferns (Pollack, 

1987). Cercospora species are commonly associated 

with leaf spots, but can also cause 

necrotic lesions on flowers, fruits, bracts, seeds 

and pedicels of numerous hosts in most climatic 

regions (Agrios 2005). In addition, several 

species of this genus are also known to be 

hyperparasites of other plant pathogenic fungi 

(Shin & Kim 2001), and are employed as 

biocontrol agents of alien weeds (Morris & 

Crous 1994). 

11

The genus Cercospora (type species: C. 

penicillata (Ces.) Fresen.) is one of the largest 

genera of hyphomycetes (Crous & Braun 2003). 

The name Cercospora, which is derived from a 

combination of the Greek “kerkok” (tail) and 

“sporos” (seed), designates the filiform conidia 

of the fungus (Crous & Braun 2003). The 

teleomorph state is Mycosphaerella Johanson 

(Dothidiomycetes, Capnodiales, Mycosphaerellaceae), 

a genus that has been linked with at 

least 30 different coelomycetes or hyphomycetes 

anamorph genera (Crous et al. 2007). 

Since Fresenius (1863) did not give the 

genus Cercospora a clear-cut definition, Saccardo 

(1880) defined it as having brown conidiophores 

and vermiform conidia which are 

brown, olivaceous or rarely subhyaline, but he 

did not mention the type species (C. apii) 

which has hyaline conidia. Saccardo considered 

C. ferruginea Fuckel as typical of Cercospora 

and repeated this definition in Sylloge 

Fungorum (1886). Since then, two anomalous 

groups of Cercospora are found to exist, i.e., 

those with colored conidia and those with 

hyaline conidia. 

Spegazzini (1910) was the first to split 

the genus Cercospora and published a new 

generic name Cercosporina Speg. to accommodate 

those species with hyaline conidia (i.e. 

with the characters of C. apii), but no type 

species was indicated for the new genus. 

Saccardo (1913) agreed with the establishment 

of Cercosporina, and transferred 89 species 

from Cercospora (including some with colored 

conidia as well as those with hyaline ones) to 

Cercosporina. This caused confusion among 

these species. Miura (1928) transferred C. apii 

to Cercosporina and also proposed the genus 

Cercosporiopsis Miura to accommodate certain 

Cercospora-like species with coloured cylindric 

conidia, but this genus is superfluous and 

illegitimate. Solheim (1930) proposed 21 sections 

of Cercospora by considering the presence 

or absence of external mycelium and 

prominent stromata, branching of conidiophores, 

as well as the shapes of conidia in order to 

make clear-cut morphology delimitation among 

species of Cercospora at that time. Later, 

Solheim & Stevens (1931) reconsidered their 

reclassification of Cercospora by adding the 

character of conidial scars, divided the genus 

into 38 sections and proposed the genus 

12 

Ragnhildiana Solheim for the species intermediate 

between Cladosporium Link and Cercospora 

based on these characters. 

Chupp (1954), in his monograph of genus 

Cercospora, made no attempt to subdivide the 

genus, however, the monograph provided a 

very valuable source of reference to almost all 

Cercospora species published up to 1954, but 

excluded those names other than Cercospora or 

Cercosporina. In Chupp’s monograph, the 

character of conidial scars are taken into 

account, either distinctly visible or obscured, 

and for those prominent scars, their sizes are 

noted as either large, medium or small. 

Deighton (1967, 1971, 1973, 1974, 1976, 

1979, 1983, 1987) continued study on Cercospora 

and allied genera, and reclassified 

numerous species. Deighton (1987) also stressed 

the characteristic of the conidial scars as an 

important character. Several genera of the cercosporoid 

fungi were redefined or newly proposed, 

which fall into two distinct taxonomic 

categories – those in which the conidial scars 

are conspicuously thickened (appearing as 

black rims when views under light microscopy) 

and those in which the conidial scars are 

unthickened (Deighton 1987). The hilum at the 

base of a conidium is thickened or unthickened 

in correspondence with the scars left on the 

conidiogenous cell (Deighton 1987). Thickened 

scars are present in Cercospora and allied 

genera such as Camptomeris Syd., Cercosporella 

Sacc., Cercosporidium Earle, Fusicladium 

Bonord., Mycovellosiella Rangel, Passalora 

Fr., Phaeoisariopsis Ferraris, Phaeoramularia 

Muntk.-Cvetk., Sirosporium Bubák & 

Serebrian., Stenella Syd., unthickened conidial 

scars occur in genera such as Cercoseptoria 

Petr., Mycocentrospora Deighton, Pseudocercospora 

Speg., and Stigmina Sacc. (Deighton 

1967, 1971, 1973, 1974, 1976, 1979, 1983, 

1987). 

The character of conidial scars, stressed 

by Deighton as an unambiguous taxonomic 

criterion, has been adopted by other mycologists 

in the classification of Cercospora and 

allied genera, e.g. Pons & Sutton (1988) and 

Braun (1988a,b, 1989, 1990, 1993). Braun 

(1993) insisted that the Cercospora generic 

conception adopted by Chupp (1954) was too 

wide, and this genus could be safely redefined 

into various additional genera to provide a

etter workable system. Braun (1993) also 

established generic separation of Cercospora 

on diverse criteria including ontogeny, pigmentation 

and ornamentation of conidia, conidiophores 

and conidiomata. Pons & Sutton (1988) 

described Distocercospora for Cercospora-like 

hyphomycetes with distoseptate, scolecosporous 

conidia. On the other hand, Braun (1993) 

separated Pseudocercospora-like species with 

percurrently proliferating conidiogenous cells 

and Mycosphaerella teleomorphs from Stigmina, 

and published the new genus Cercostigmina 

U. Braun. Although Deighton (1967) 

separated Passalora and Cercosporidium on 

account of the presence or absence of a 

substomatal stroma, Braun (1995) redefined 

Cercospora, Passalora and Phaeoisariopsis. 

Braun (1995) discussed the status of these 

genera and noted that small stromata were also 

developed in the type species of Passalora. 

Therefore, the degree of the development of 

stroma-like hyphal aggregations in the substomatal 

cavities should not be used for generic 

differentiations within Cercospora and allied 

genera (Braun 1995). 

Crous & Braun (2003) re-examined and 

presented a compilation of more than 3,000 

names published in the genus Cercospora and 

550 names in the genus Passalora. They 

separated the cercosporoid genera mainly based 

on a combination of characters, of which the 

structure of conidiogenous loci (scars) and hila, 

the presence and absence of pigmentation in 

conidiophores and conidia, and other characters 

described above (Crous & Braun 2003). A key 

to the true hyphomycetous cercosporoid fungi 

and related genera was also provided by Crous 

& Braun (2003). Based on these morphological 

categories, Crous & Braun (2003) retreated 

and re-examined 5,720 names that 

related to Cercospora s. str. (sensu stricto), and 

proposed 455 taxonomic novelties within 10 

genera including Cercospora, Dactylaria Sacc., 

Fusicladium, Mycosphaerella (teleomorph), 

Passalora, Scolecostigmina U. Braun, Semipseudocercospora 

J.M. Yen, Sirosporium, 

Sporidesmium Link and Stenella Syd. Crous & 

Braun (2003) recognized only 659 names in the 

genus Cercospora with a further 281 being 

referred to C. apii s. lat. They also stated the 

need of a “compound species”, a species that is 

formed when two or more species join together 


because of indistinguishable morphologically/genetically/physiologically, 

named C. apii s. 

lat. comprising all cercosporoid hyphomycetes 

indistinguishable from the Cercospora on 

Apium graveolens L. Introduction of new 

names for morphologically indistinguishable 

Cercospora collections detected on new host 

genera and families, should be avoided, and 

should simply be referred to C. apii s. lat. 

(Crous & Braun 2003). 

In Thailand, there have been only a few 

reports of cercosporoid fungi. These include 

Sontirat et al. (1980) who enumerated 21 

species of Cercospora; a list of 47 identified 

and 13 unidentified species of Cercospora in 

The Host Index of Plant Diseases in Thailand; 

by Giatgong (1980); and a report on 49 species 

by Petcharat & Kanjanamaneesathian (1989). 

These reports were based on the generic 

concept introduced by Chupp (1954). Further 

additional records of cercosporoid fungi in 

Thailand were also published by Ellis (1976), 

Manoch et al. (1986), Pons & Sutton (1988), 

Barreto & Evans (1994), Crous (1998), Crous 

& Braun (2003), Lumyong et al. (2003), Braun 

et al. (2006) and Hunter et al. (2006). As of this 

group of fungi in Thailand is little known, the 

information is scattered and based on Chupp’s 

generic concept, a survey on diversity based on 

more recent taxonomic concepts introduced by 

Deighton (1967, 1971, 1973, 1974, 1976, 1979, 

1983, 1987), Pons & Sutton (1988), Braun 

(1988a,b, 1989, 1990, 1993, 1994, 1995, 1996, 

1998, 1999), Crous & Braun (2003) and Crous 

et al. (2001, 2006, 2007) is for this group of 

fungi in this region. 

Morphology Characteristics 

Deighton (1967, 1971, 1973, 1974, 1976, 

1979, 1983), Pons & Sutton (1988), Braun 

(1993), Braun & Melnik (1997) and other 

authors divided Cercospora s. lat. into numerous 

smaller genera based on morphological 

characteristics. Later, a combination of morphology 

and molecular analysis were also 

carried out by Crous et al. (2000, 2001). From 

their intensive research on this group of fungi, 

Crous & Braun (2003) published the compilation 

of the names in Cercospora and Passalora, 

and re-defined the morphological characteristics 

of Cercospora s. lat. based on morphology 

and molecular analysis. The following 

13

description and illustration are the common 

items used to identify cercosporoid fungi, in 

particular the genus Cercospora. 

A. Symptoms 

Symptoms caused by Cercospora species 

are variable (Fig. 1). Leaf spots may be absent 

or present in every degree of distinctiveness 

from a faint discoloration on both leaf surfaces 

to well defined and conspicuous leaf spots with 

colored borders, eye-spot diseases or veinlimited 

lesions (Chupp 1954). Often effuse 

caespituli (or fruit bodies) are visible on the 

lower leaf surface when no leaf spots are 

visible, and the fungi may also be so minute 

that a hand lens is required to detect it (Chupp 

1954). When the disease reaches a certain stage 

of severity, the leaf may curl, dry and often 

drop from the plant (Chupp 1954). 

Many species of Cercospora also affect 

the blossoms, fruits, succulent petioles and 

young stems, for examples, C. agavicola 

Ayala-Escobar on Agave tequailana F.A.C. 

Weber and C. caricae-papayae P.K. Rajak & 

S.P. Gautam on Carica papaya L. Frequently, 

one to numerous spots may turn the entire leaf 

yellow or brown, after which it shrivels and 

dies (Chupp 1954). Only the leaf symptoms as 

they show in freshly collected (or herbarium) 

material are taken into account in describing 

the symptoms of the individual Cercospora 

species. 

B. Caespituli (Fruit Bodies) 

Caespituli of Cercospora, are turfs of 

conidiophores as seen under a microscope or 

hand lens (Fig. 2). Caespituli may be distributed 

on the upper surface (epiphyllous), lower 

surface (hypophyllous), both surfaces (amphigenous); 

evenly distributed on the spot or 

aggregated along the margin of the spot 

(Chupp 1954). Caespituli often appear velvety, 

floccose or arachnoid as effuse patches (Chupp 

1954). The colours are variable but mostly grey, 

olivaceous to whitish (Chupp 1954). 

C. Conidiophores, Conidiogenous Cells, and 

Conidiogenesis 

A conidiophore, defined as the entire system 

of fertile hyphae bearing conidia, may be 

either simple or branched, and includes the 

14 

conidiogenous cell (s) (Ulloa & Hanlin 1999). 

It can be reduced to a single fertile cell if the 

conidiophore and the conidiogenous cell are 

identical, or the conidiophore is composed of a 

single conidiogenous cell and a single or several 

supporting cells, or it consists of a system of 

conidiogenous cells with or without differentiated 

supporting structure (hyphal cells, stipe) 

(Gams et al. 1987). Some authors, for instance 

Hawksworth et al. (1995) and Pons & Sutton 

(1988), preferred to confine the term conidiophore 

to complex structures composed of two 

or more cells. In the present study, a wider 

concept of the term conidiophore is applied as 

a one-celled conidium-bearing structures can 

either be called conidiogenous cell or conidiophore, 

depending on the particular case. 

In the cercosporoid fungi, there are 

numerous species with tufts of mixed conidiophores. 

Some of them are continuous and onecelled 

(e.g. some species of Passalora), other 

conidiophores are septate, composed of two or 

more cells (e.g. species of Cercospora). 

Conidiophores may be colorless (hyaline) or 

variously pigmented, and the pigmentation is 

an important taxonomic feature (Crous & 

Braun 2003). Conidiophores may be formed 

singly, erumpent through the substratum or 

arising from hyphae as lateral branchlets, or 

they are caespitose, i.e. arranged in loose or 

dense fascicles (Gams et al. 1987) (Figs 3a–d). 

Conidiogenous cells can be formed as part of 

an undifferentiated hypha, and they also can 

form a unicellular conidiophore or they can 

mostly form part of a pluricellular conidiophores 

(Hennebert & Sutton 1994). In the latter 

case, they can be either terminal, intercalary or 

pleurogenous (Hennebert & Sutton 1994). If 

they are formed laterally or terminally but not 

in continuity with the main axis, they are called 

discrete (Hennebert & Sutton 1994). A coni- 

diogenous cell may be unilocal/single locus 

(Figs 3–b, d) or multilocal/more than two loci 

(Figs 3a, c). The loci can be apical, lateral or 

circumspersed (all around the conidiogenous 

cell) (Hennebert & Sutton 1994). 

A conidial scar, the minute structure at 

the end of a conidiogenous cell results from 

conidiogenesis, and is a recognizable portion 

where the conidium has been liberated 

(Hennebert & Sutton 1994). Conidia scars


Fig. 1 – Various types of cercospora leaf spot symptoms on leaves: a. Carica papaya, b. Lactuca 

sativa, c. Kopsia fruticosa and d. Lactuca sativa (cultivar head lettuce). (Meeboon et al. 2007c). 

may be conspicuous by thickened walls with 

dark coloration (Figs 3c–d) or inconspicuous 

(Hennebert & Sutton 1994). A scar on a 

conidium at the point of former attachment to 

the conidiophore is termed the hilum 

(Hennebert & Sutton 1994). The genus 

Cercospora is characterized by conspicuous, 

thickened and darkened scars and hila (Crous 

& Braun 2003). 

The development of a conidium from the 

conidiogenous cell or conidiophore is called 

conidiogenesis (Hennebert & Sutton 1994). 

Cercosporoid fungi conidiogenesis is characterized 

by blastic, sometimes determinate but 

often sympodial proliferation, mostly schizolytic 

conidial secession with single or conidia 

in chains. Blastic conidiogenesis is characterized 

by an elastic wall of the conidiogenous 

cells, bulging out to form a conspicuous, 

enlarged conidium initial (Hennebert & Sutton 

1994). It may be holoblastic [all wall layers of 

the conidiogenous cells contribute towards the 

formation of the conidium (blastoconidia)] or 

enteroblastic (only the inner wall of the conidiogenous 

cell contributes towards the formation 

of the conidium) (Hennebert & Sutton 

1994). Blastic conidiogenous cells may be 

monoblastic (only with a single conidiogenous 

15

Fig. 2 – Appearance of caespituli (as turfs of conidiophores) of genus Cercospora on the leaf spot 

of Coccinia grandis (L.) Voigt. (Meeboon 2009). 

Fig. 3 – Various types of conidiophores and conidiogenous cells of the cercosporoid fungi (40×). a. 

Fasciculate and non-divergent. b. Fasciculate, non-divergent and conidiogenous loci not darkened. c. 

Fasciculate, divergent, polyblastic, sympodial proliferation, with dark conidiogenous loci. d. 

Fasciculate, distinctly divergent, non-sympodial proliferation with dark conidiogenous loci. 

(Meeboon 2009). 

16


Fig. 4 – Various types of conidia of the true cercosporoid fungi sensu Crous & Braun (2003) found 

in this study (40×) (black arrows: conidia hila, white arrows: septation). a. Conidia of genus 

Cercospora s. str. b. Conidium of genus Passalora. c. Conidia of genus Pseudocercospora. 


locus or unilocal) or polyblastic (with two or 

more conidiogenous loci or multilocal), formed 

either synchronously or, mostly; in a sympodial 

succession (Hennebert & Sutton 1994). Conidiophores 

(or conidiogenous cells) can be 

determinate (growth ceasing with the production 

of a terminal conidium or conidial chain) 

or they can proliferate [indeterminate, proliferation 

being sympodial or percurrent (through 

the open end left when the first conidium 

becomes detached)] (Hennebert & Sutton 

1994). Cercospora is characterized by having 

blastic (monopolyblastic), schizolytic, and 

sympodial proliferation (Crous & Braun 2003). 

C. Conidia 

The important characters of conidia of 

the genus Cercospora are mostly related to the 

shape, septation, pigmentation and surface 

(Figs 4a–c), similar to the Saccardoan system 

(Crous & Braun 2003). The conidia of Cercospora 

species are either straight to curved, and 

acicular, filiform, obclavate or a combination 

of shapes (Crous & Braun 2003). There are two 

basic types of septation, viz, euseptate (septa 

formed by all existing wall layers) and 

distoseptate/pseudoseptate (septa formed only 

by the innermost layer) (Hennebert & Sutton 

1994). The term septum (septate) without 

specification is usually applied to euseptate 

(Hennebert & Sutton 1994). 

Hyaline and pigmented conidial structures 

are usually well separated in certain taxa 

(genera, species) of the cercosporoid fungi, but 

transitional phenomena are not uncommon 

(Crous & Braun 2003). However, taxa with 

subhyaline to pale (yellowish green, pale olivaceous, 

etc.) structures often cause serious 

taxonomic problems (Crous & Braun 2003). 

The conidia of Cercospora are characterized by 

hyaline or pale olivaceous pigmentation and 

euseptate conidial septation (Crous & Braun 

2003) 

Collection and Observation 

Specimen collection involved an observation 

of the presence/absence of the fruiting 

bodies/caespituli on the leaf. The observation 

was usually conducted using a 10× or 20× 

magnifying lens. Specimens that are positively 

showed the presence of Cercospora fruiting 

bodies/caespituli were placed in plastic bags. 

The collecting bags were sealed and labeled: 

Name of host plants, Collection site, Collector 

/s, and Collection date. 

Detailed observations of morphological 

characters were generally carried out by means 

of a dissecting microscope, followed by light 

compound microscope using oil immersion 

(1000×). Specimens for microscopic observation 

were prepared by hand sectioning or fine 

forceps. Water is very good as mounting medium. 

Shear’s solution or lactophenol was 

usually used as a media for permanent slides. 

Thirty conidia, hila, conidiophores, conidiogenous 

loci and 10 stromata were commonly 

measured for each specimen. Line drawings 

were prepared at a magnification of 400×, or 

17

1000×, if necessary. Dried herbarium specimens 

were deposited at herbarium at CMU 

Herbarium (CMU), Biology Department, 

Faculty of Science, Chiang Mai University, 

Chiang Mai, Thailand, and BIOTEC Herbarium 

(BBH), Bangkok, Thailand. Living 

cultures have been deposited at BIOTEC 

culture collection (BCC), Bangkok, and 

Molecular of Plant Pathology Laboratory, 

Department of Plant Pathology, Chiang Mai 

University. 

Single Spore Isolation 

Single spore isolation of each species 

followed the method outlined by Choi et al. 

(1999), with a modification (Fig. 5). 

A glass slide was sterilized with ethanol 

and wiped with a tissue on which ethanol (70%) 

had been sprayed. A sterilized pipette was then 

used to transfer 1–2 drops of sterilized water 

onto the glass slide. Flame sterilized fine forceps 

were used to take conidia from the 

specimen and to suspend the conidia with 

sterilized water on the glass slide. A triangle 

line was marked on the bottom of the water 

agar plate. The prepared homogenous spore 

suspension was then transferred with a sterilized 

loop, onto the surface of the water agar 

plate, and smeared following the drawn lines. 

The unsealed plate was incubated at room 

temperature for approximately 24 hours. The 

plate was not sealed as this allows some of the 

surface water to dry out, and thus reduce the 

chance of contamination. The conidia were 

usually checked within 24 hours to establish 

germination. Once the conidia had germinated, 

a sterilized glass needle was used to pick up a 

small piece of agar containing a conidium. In 

order to establish that the conidium is the one 

desired, and to maintain quality control, a slide 

was prepared and examined under the compound 

microscope. If the conidia do not 

germinate after 24 hours, the plate was then 

sealed with PARAFILM “M” ® (Chicago, USA) 

and examined periodically. Approximately 10- 

20 germinated conidia were transferred and 

distributed evenly onto two PDA plates and 

incubated at room temperature until their 

colony diameter was about 1 to 2 cm. A small 

piece of mycelium with agar could then be cut 

18 

and transferred to another PDA plate. The 

culture was checked after a few days. 

Preservation of Specimens 

Once fully examined, the host-plant 

tissue was spread out on folded drying paper or 

newspaper along with a collection-number tag 

and dried. 

Identification Procedures 

Identification of species was mainly 

based on the recent concepts of Deighton (1967, 

1971, 1973, 1974, 1976, 1979, 1983, 1987), 

Pons & Sutton (1988), Braun (1988a,b, 1989, 

1990, 1993, 1994, 1995, 1996, 1998, 1999), 

Crous & Braun (2003), and Crous et al. (2001, 

2006, 2007). In most cases the specimen could 

be identified to at least genus level with the 

above references, further identification required 

examination of the relevant literatures. Sources 

are often suggested in the above references and 

the Dictionary of Fungi (Kirk et al. 2008), the 

‘searchable’ Index Fungorum (http://www.in 

dexfungorum.org/Names/Names.asp) and Systematic 

Mycology and Microbiology Laboratory 

(SMML) Database (http://nt.ars-grin.gov/ 

fungaldatabases/fungushost/fungushost.cfm) on 

the internet are also invaluable. 

The following is dichotomous key to the 

four genera of the true cercosporoid fungi 

adopted from Crous & Braun (2003). 

1. Conidiogenous loci inconspicuous or subdenticulate, 

but always unthickened and not 

darkened or subconspicuous, i.e., unthickened, 

but somewhat refractive or rarely very 

slightly darkened, or only outer rim slightly 

darkened and refractive (visible as minute 

rings) .............................. Pseudocercospora 

1. Conidiogenous loci conspicuous, i.e., 

thickened and darkened throughout, only 

with a minute central pore.......................... 2 

2. With verruculose superficial secondary 

mycelium; conidia amero- to scolecosporous, 

mostly verruculose ..........................Stenella 

2. If superficial secondary mycelium present, 

hyphae smooth or almost so....................... 3


Fig. 5 – Diagram of single spore isolation employed in the isolation of Cercospora species. 

(modified from Choi et al. 1999). 

3. Conidia hyaline or subhyaline, scolecosporous, 

acicular, obclavate-cylindrical, 

filiform, usually pluriseptate ..... Cercospora 

3. Conidia pigmented or, if subhyaline, conidia 

non-scolecosporous, ellipsoid-ovoid, short 

cylindrical, fusoid and only few septa 

......................................................Passalora 

Presentation of Results 

Cercospora species described and illustrated 

in this book are presented in alphabetical 

order according to plant families. All species 

are fully described and illustrated. The following 

data are provided: 

• Names of species with reference 

• Synonyms 

• Full description 

• Specimen examined 

• Habitat (host range) 

• Distribution with countries in alphabetical 

order 

• Notes (if necessary) 

Molecular Phylogenetic Analysis 

A. DNA Extraction, Polymerase Chain Reaction 

(PCR) and Sequencing 

Molecular characterization was carried 

out in order to elucidate the phylogenetic relationship 

of the members of genus Cercospora, 

and the relationship with other related genera 

within anamorphic taxa of Mycosphaerella the 

teleomorph. Total genomic DNA was extracted 

from fungal mycelia cultured on MEA (Difco, 

USA) following a 2× cetyltrimethylammoniumbromide 

(CTAB) protocol (Rogers & 

Bendich, 1994). DNA amplification of ITS 

region of nrDNA (Fig. 6) was performed by 

polymerase chain reaction (PCR) using ITS4 

(5′-TCCTCCGCTTATTGATATGC-3′) and 

ITS5 (5′-GGAA- GTAAAAGTCGTAACAA 

GG-3′) primers (White et al. 1990) to generate 

about 580 nucleotides from the complete ITS, 

including 5.8S rDNA region. The amplification 

condition was performed in a 50 mL reaction 

volume as follows: 1× PCR buffer, 0.2 mM 

each dNTP, 0.3 mM of each primer, 1.5 mM 

MgCl2, 0.8 units Amplitaq Taq Polymerase 

(Perkin-Elmer, Foster City, CA, USA), and 10 

ng DNA. PCR parameters for all the regions 

were performed as follows: initial denaturation 

at 94ºC for 3 min, followed by 30 cycles of 

94ºC for 1 min, 52ºC for 50 s, 72ºC for 1 min, 

and final extension of 72ºC for 10 min. 

The characterization of PCR products 

was performed via agarose gel electrophoresis 

on a TAE 1% agarose gel containing EB (EtBr) 

as the staining agent. The PCR product was 

purified using Qiaquick purification kit (Qiagen) 

and DNA concentration of the PCR 

products was subjected to automatic sequencing 

(ABI PRISM Dye Terminator Cycle 

Sequencing and ABI PRISM Sequencer model 

377, Perkin Elmer). 

19

Fig. 6 – Diagrammatic illustration of Internal Transcribed Spacer (ITS) region of ribosomal DNA 

(rDNA). Primers positions, ITS5 and ITS4, in highly conserved small subunit (SSU) 18S and large 

subunit (LSU) 28S ribosomal DNA gene sequences flanking the spacer regions, ITS I and ITS II, 

are shown. 

B. Sequence Alignment and Phylogenetic 

Analysis 

Sequences obtained from the respective 

primers (ITS5 and ITS4) were aligned in 

Clustal X (Thomson et al. 1997) and BioEdit 

(Hall 1999). The sequence alignments were 

also refined by direct examination. Regions 

designated as ambiguously aligned were 

excluded from the analysis. Gaps were treated 

as missing data. Phylogenetic analyses were 

performed in PAUP version 4.0b10 (Swofford 

2002). 

Unweighted Maximum Parsimony (UMP) 

analysis was performed in order to confirm the 

phylogenetic relationship with related taxa. 

Trees were inferred using the heuristic search 

option with 1000 random sequence additions. 

Maxtrees were unlimited, branches of zero 

length were collapsed and all multiple parsimonious 

trees were saved. Descriptive tree statistics 

(tree length [TL], consistency index [CI], 

retention index [RI], related consistency index 

[RC], homoplasy index [HI] and log likelihood 

[-ln L]) were calculated for trees generated 

under different optimality criteria. The Kishi 

no-Hasegawa (KH) likelihood test (Kishino & 

Hasegawa 1989) was carried out using PAUP 

to compare the best tree topology obtained by 

the nucleotide sequence data with a constrained 

tree. Clade stability was assessed in bootstrap 

analyses with 1000 replicates, each with 1000 

replicates of random stepwise addition of taxa. 

20 

Random sequence addition was used in the 

bootstrap analysis. Trees were figured in 

TreeView (Page 1996). 

Phylogenetic Affinities of Cercospora and 

Allied Genera Based on ITS nrDNA Sequence 

Analysis 

Current Understanding on Phylogeny of 

Cercospora and Allied Genera 

Every living organism contains DNA, 

RNA and proteins. Closely related organisms 

generally have a high degree of agreement in 

the molecular structure of these substances, 

while the molecules of organisms distantly 

related usually show a pattern of dissimilarity. 

With the advent of molecular technique, particularly 

since the finding of fungal ribosomal 

RNA genes amplification and direct sequencing 

technique by White et al. (1990), nucleotide 

sequences sampled from genome have 

been commonly employed in recent years by 

systematists to investigate the phylogeny of 

various groups of fungi, and consequently, the 

progress in molecular phylogenetic of Kingdom 

Fungi has accelerated rapidly. 

In Cercospora and allied genera, until the 

present time, only a few molecular phylogeny 

analyses have been published. One of the first 

significant phylogenetic analyses was published 

by Stewart et al. (1999) who reported the 

monophylogeny of Cercospora, Passalora and

Pseudocercospora based on ITS region of 

partial rDNA sequence analysis, and reaffirmed 

that Ramulispora Miura and Mycocentrospora 

Deighton are not related to Mycosphaerella 

teleomorph. Stewart et al. (1999) also reduced 

Paracercospora Deighton to a synonym of 

Pseudocercospora. However, because of limited 

taxa, no other species linked to Mycosphaerella 

teleomorph were included in their 

analysis, and it was not possible to determine 

the phylogenetic relationship of the Cercospora 

species to other anamorphs genera. 

Similar to the anamorphic state, the 

taxonomy and phylogenetic of Mycosphaerella 

(teleomorphic state) is also complicated (von 

Arx 1983, Crous et al. 2000). Due to the large 

number of associated anamorphs, Crous & 

Wingfield (1996) noted that Mycosphaerella 

was a polyphyletic assemblage of presumably 

monophyletic anamorphic genera. Goodwin et 

al. (2001), based on the analysis of a large 

number of anamorphs of Mycosphaerella using 

ITS region of rDNA sequence, also found that 

Mycosphaerella was not monophyletic. However, 

Goodwin et al. (2001) noted that Cercospora 

s. str. formed a highly supported monophyletic 

group, and the Cercospora species 

that produced the toxin cercosporin were suggested 

to have a single evolutionary origin. 

Crous et al. (2007), based on the analysis of 

Large Sub Unit (LSU) region of ribosomal 

DNA (28SrDNA), reaffirmed that Mycosphaerella 

was polyphyletic. 

Although the Mycosphaerella complex 

encompasses thousands of names, studies on 

the phylogenetic relationship among taxa in 

this group are still rare compared with other 

fungal groups. This is probably due to the fact 

that these organisms are relatively difficult to 

isolate on artificial medium (Crous et al. 2007). 

In fact, most taxa belonging to Mycosphaerella 

and anamorphs (the cercosporoid fungi) which 

are seen successfully cultivated on the artificial 

medium grow relatively slower than other 

fungi. 

Phylogeny of Cercospora Species from 

Thailand 

A phylogenetic tree of 42 representative 

sequences of Cercospora and allied genera 

from Thailand and other sequences obtained 

from NCBI (National Center for Biotechnology 


Information) GenBank database, obtained from 

unweighted maximum parsimony (UMP) analysis 

method are shown in Fig. 7. 

Based on this analysis, 6 genera of the 

cercosporoid fungi included in the analysis, 

namely, Cercospora, Septoria, Pseudocercospora, 

Stigmina, Ramularia and Passalora 

appear as monophyletic groups with 60%, 53%, 

95%, 100%, 100% and 79% bootstrap support, 

respectively (Fig. 7). Cladosporium was used 

as an out group. This result shows that morphological 

definitions of all these genera are also 

well-defined phylogenetically. A similar finding 

was also reported by Crous et al. (2001) 

with the exception of genus Stigmina. The 

general morphological characteristics among 

these six genera and other cercosporoid fungi 

are also briefly illustrated in Fig. 7. This 

diagram shows the differences among taxa 

within cercosporoid fungi which are mainly 

separated by distinct structures of conidia, 

conidiophores, conidiogenous cells, hila and 

scars. 

The Cercospora species formed a monophyletic 

clade with 60% bootstrap support (Fig. 

7). This clade appeared as a sister group to 

Septoria clade with 89% bootstrap support 

which indicates a close relationship between 

the two genera. Septoria, a coelomycetous 

fungus, shares similar morphology characteristics 

to Cercospora in having holoblastic 

and sympodial conidiophore proliferation, as 

well as hyaline, filiform to acicular and 

multiseptate conidia (Sutton 1980). However, 

the two genera are morphologically separated 

due to Septoria producing pycnidial conidiomata 

(Figs 7–8). These genera are maintained 

as separate taxa, although Verkley & Starink- 

Willemse (2004) noted that conidiomatal structure 

seems to have little predictive value for 

phylogenetic relatedness, but phylogenetically 

analysis showed that Cercospora and Septoria 

are not monophyletic within Mycosphaerella 

and its anamorphs. It was probably because of 

The presence of intermediate species between 

Cercospora and Septoria, therefore, more 

genes loci or taxa are required to analyze the 

relationship between the two genera. 

Passalora clade appeared as a basal 

group in the phylogenetic tree with 79% bootstrap 

support. This genus was introduced by 

Fries (1849) with Passalora bacilligera (Mont. 

21

Fig. 7 – Best parsimonious tree (300 steps) based on ITS nrDNA sequence data representing 

phylogenetic affinities of Cercospora with closely related genera representatives of the 

Mycosphaerella anamorphs. The analysis yielded 486 total characters of which 327 characters were 

constant, 31 characters were variable and parsimony-uninformative and 128 characters were 

parsimony-informative Bootstrap values (>50%) from 1000 replicates of parsimony analysis are 

shown above internodes (To-anun et al. 2009). 

22


Fig. 8 – Illustration of morphological structures between Cercospora and Septoria. A. Cercospora 

christellae (To-anun et al. 2009). B. Septoria violae-patrinii (Shin & Sameva 2002) (arrows show 

the location of conidiophores inside conidiomata). 

& Fr.) Mont. & Fr. (≡ Cladosporium bacilligerum) 

as type species. This species was 

characterized by having pigmented conidiophores, 

and ellipsoid-fusiform, obclavatesubcylindric, 

pigmented conidia with (0-)1(-3)septa 

(basal clade, Fig. 7). The basal position 

of Passalora clade indicated that species in 

Passalora hold more plesiomorphic characters 

or ancestral state characters than other 

cercosporoid fungi clades. This indication was 

also reported earlier by Stewart et al. (1999). 

On the other hand, species within Pseudocercospora 

s. str. form a monophyletic clade 

wth 95% bootstrap support and appear as a 

sister group to Stigmina clade with 60% bootstrap 

support (Fig. 7). Morphologically, these 

two genera are similar in having holoblastic 

and terminal conidia. proliferation, as well as 

obclavate to filiform-acicular with truncate 

base and multiseptate conidia, However, they 

differs due to the conidia of Stigmina being 

verrucose, dark brown and sometimes producing 

longitudinal septa, which is quite distinct 

to Pseudocercospora s. str. which has 

smooth and subhyaline conidia with only transverse 

septation, and unthickened conidial loci 

and hila. Pseudocercospora s. lat. is morphologically 

highly variable, and accommodates a 

wide range of cercosporoid hyphomycetes with 

pigmented conidiophores and inconspicuous, 

unthickened and not darkened conidiogenous 

loci. It includes Paracercospora, Phaeoisariopsis, 

Stigmina and Pseudophaeoramularia 

(Crous et al. 2001). Crous et al. (2000) showed 

Pseudocercospora s. lat. to be polyphyletic 

within Mycosphaerella anamorphs, having 

evolved more than once from different Mycosphaerella 

holomorphs, and in several occasions 

having lost the teleomorph (Crous & 

Braun 2003). This complex includes genera 

with single to synnematous (Phaeoisariopsis) 

conidiophores, solitary or catenate (Pseudophaeoramularia) 

conidia, eu- (Pseudocercospora) 

or distoseptate (Stigmina), and scars 

which are inconspicuous to denticle-like 

(Denticularia and Semipseudocercospora), unthickened 

to slightly pigmented conidia (Paracercospora) 


Within Cercospora species (Cercospora 

s. str.), Crous & Braun (2003) stated the need 

of a “compound species”, a species that is 

formed when two or more species join together 

because of indistinguishable in morphology or 

genetic or physiology characteristics, named C. 

apii s. lat., comprising all cercosporoid hyphomycetes 

indistinguishable from the Cercospora 

23

on Apium graveolens L. Introduction of new 

names for morphologically indistinguishable 

Cercospora collections detected on new host 

genera and families, respectively, should be 

avoided, and should simply be referred to C. 

apii s. lat. Crous & Braun (2003) also revised 

these species and redisposed many of them. A 

total of 659 Cercospora species were recognized, 

with a further 281 being referred to 

synonymy under C. apii s. lat. 

Unfortunately, only a few species belonging 

to C. apii s. lat. have been cultured and 

molecular data addressing the phylogenetic 

relationship within this complex and related 

species is still lacking. Thus, it is necessary to 

examine phylogenetically whether the species 

within this complex are monophyletic or not. If 

the C. apii s. lat. complex forms a monophyletic 

group, it is possible that a single species of 

Cercospora (C. apii) occurs on a wide host 

range. 

The phylogenetic tree generated from 

unweighted maximum parsimony analysis of 

ITS nrDNA region showed the monophyletic 

origin of Cercospora s. str. with 100% bootstrap 

support (Fig. 9). The short branch lengths 

among species within the Cercospora cluster 

indicate that they all shared a common ancestor 

relatively recently. The species belonging to C. 

apii s. lat. such as C. apii (on Apium), C. 

beticola (on Beta), C. hayi (on Musa), C. 

kikuchii (on Lycine), C. penzigii (on Citrus), C. 

physalidis (on Solanum) are polyphyletic (Fig. 

9). To date, only Ayala-Escobar et al. (2005) 

reported the monophyletic of C. apii s. lat. by 

combining five genes loci, namely, internal 

transcribed spacer (ITS) nrDNA, elongation 

factor 1-α (EF), actin (ACT), calmodulin (CAL) 

and histone H3 (HIS), even though with a few 

sequences included in their analysis (2 species 

of C. apii s. lat. and 2 species of Cercospora s. 

str. non C. apii s. lat.). Groenewald et al. (2006) 

found molecular evidence that C. apii has a 

wider host range than had been accepted by 

Chupp (1954) and Ellis (1976), but has 

narrower host range than that proposed by 

Crous & Braun (2003). The host range data of 

C. apii s. lat. obtained in Thailand illustrate 

that C. apii is not entirely host specific, and it 

is not possible to identify this species (and C. 

24 

apii complex) solely based on host, morphology 

and/or geographic location. Therefore, 

this has to be further investigated by conducting 

pathogenicity studies on all the hosts 

previously listed for these species. 

Although the monophylogeny of C. apii 

has been reported using a combination of five 

genes loci (Ayala-Escobar et al. 2005), often 

all the five genes loci sequenced are not 

congruent and can not be used in particular 

when a large number of sequences included. In 

order to overcome this problem, Groenewald et 

al. (2005) used sequence data of the same five 

genes as Ayala-Escobar et al. (2005) in combination 

with other features such as growth rate 

to establish species boundaries for C. apii, C. 

apiicola (as Cercospora sp.) and C. beticola. 

From these established species boundaries, 

species-specific primers were designed in 

polymorphic areas of the calmodulin gene for 

the three species. This combined approach 

probably represents the most reliable way to 

characterize and identify species within this 

complex. 

One possible explanation of C. apiicomplex 

occurrence on various host plants is 

probably due to host jumping events in which 

C. apii occurs on ‘‘atypical’’ hosts in the 

process of reaching their real hosts. However, 

the reason why host jumping by C. apii occurs 

remains unknown. Perhaps under stress (a 

shortage of host tissue or unsuitable weather) 

the new species might be able to jump from 

celery onto other hosts (Crous & Groenewald 

2005). Thus, it would be interesting to determine 

whether the C. apii that exist on 

‘‘atypical’’ hosts are able to cause disease on 

those hosts or not. At the moment, the information 

of Cercospora isolates from Thailand have 

shown that C. apii-complex from ‘‘atypical’’ 

hosts do not produce pigmentation (cercosporin) 

on agar medium. Therefore, the ability to 

produce cercosporin on medium seems to be 

one of important factors in revealing the 

phylogenetic relationships among the Cerco- 

spora s, str. as it was also previously found by 

Goodwin et al. (2003) that Cercospora 

producing pigment on agar medium are 

monophyletic in the phylogenetic tree 

generated from beta tubulin gene analysis.


Fig. 9 – Single parsimonious tree based on ITS nrDNA sequence data representing phylogenetic 

affinities of Cercospora s. str. The tree is obtained from heuristic search with 1000 random taxon 

addition of the sequences alignment. Bootstrap values (>50%) from 1000 replicates of Unweighted 

Maximum Parsimony (UMP) analysis are shown above internodes. 

25

* IVS = Intervening region sequence: the one or more segments of a split gene that are transcribed but not included in the 

final messenger ribonucleic acid; each is flanked by two exons. Also known as intron. 

Fig. 10 – Partial nucleotide sequence of the β-tubulin of a Cercospora lactucae-sativae Ben R strain 

between codon 159 to 264. 

26 

Fungus Strain Type Alignment 

195 196 197 198 199 200 201 202 203 204 205 

C. lactucae-sativae CCR-18 R AAC TCC GAC GCG ACT TTC TGT ATC GAC AAC GAG 

C. beticola C-3 S AAC TCC GAC GAG ACC TTC TGT ATC GAC AAC GAG 

C. beticola AD-762 R AAC TCC GAC GCG ACC TTC TGT ATC GAC AAC GAG 

M. fijiensis 020501 S AAC TCT GAC GAG ACC TTC TGT ATC GAC AAC GAG 

M. fijiensis 020301 R AAC TCT GAC GCG ACC TTC TGT ATC GAC AAC GAG 

C. kikuchii JC-203 R AAC TCC GAC GCG ACC TTC TGT ATC GAC AAC GAG 

Fig. 11 Sequence alignment for C. lactucae-sativae β-tubulin predicted codons 195-205 compared 

with sequences of Ben R of C. beticola, C. kikuchii, and M. fijiensis and Ben S of C. beticola and M. 

fijiensis. 

Taxonomy 

Cercospora Fresenius, in Fuckel, Hedwigia 1 

(15): 133 (1863) and Fungi Rhen., exs, Fasc. II, 

No. 117 (1863) s. str. 

Non-original description (Ellis 1971, Dematiaceous 

Hyphomycetes: 275). 

Colonies effuse, greyish, tufted. Mycelium 

mostly immersed. Stroma often present, 

but not large. Setae and hyphopodia absent. 

Conidiophores macronematous, mononematous, 

caespitose, straight or flexuous, sometimes 

geniculate, unbranched or rarely branch- 

ed, olivaceous brown or brown, paler towards 

the apex, smooth. Conidiogenous cells integrated, 

terminal, polyblastic, sympodial, cylindrical, 

cicatrized, scars usually conspicuous. Conidia 

solitary, acropleurogenous, simple, obclavate 

or subulate, colourless or pale, pluriseptate, 

smooth. 

Holotype species – Cercospora penicillata 

(Ces.) Fresen 

= Cercospora depazeoides (Desm.) Sacc. 

Lectotype species – Cercospora apii Fres. 

Teleomorph – Mycosphaerella Johanson

Key to the treated species in Thailand, 

arranged by host family 

Acanthaceae 

1. Stromata lacking or rudimentary; conidiophores 

relatively short, 16–60 × 3–6.5 µm, 

geniculate; on Andrographis ........................ 

...................................C. andrographidicola 

1. Stromata small, 20–24 μm diam.; conidiophores 

long, 85–209 × 3–4 μm, slightly 

geniculate near the apex; on Barleria ......... 

.............................................. C. barleriicola 

Adiantaceae 

A single species, on Doryopteris ................... 

..............................................C. adiantigena 

Amaranthaceae 

1. Caespituli hypohyllous; stromata often 

lacking, if present, small, up to 8 μm diam.; 

conidiophores up to 5 in loose fascicles, 

long, 90.5–192 × 3–4 μm; conidia acicular, 

8–316 × 3–4 μm, 10–22-septate; on Iresine 

and Celosia ............. Cercospora canescens 

1. Caespituli amphigenous ............................. 2 

2. Conidiophores up to 13 in moderately dense 

fascicles, 34–85 × 2.5–5 μm; conidia 

acicular, 12–67 × 2.5–3.5 μm, 4–7-septate; 

on Celosia....................Cercospora celosiae 

2. Stromata small, 12–26 μm diam.; conidiophores 

5–9 in loose and divergent fascicles, 

47–125 × 3–5 μm; conidia narrowly obclavate 

to subacicular, 29–168.5 × 2.5–3.5 μm, 

2–12-septate; on Ricinus .............................. 

.................................... Cercospora ricinella 

Apocynaceae 

A single species, on Pentalino ......................... 

.................................. Cercospora peregrina 

Arecaceae 

A single species, on Areca ............................... 

.............................Cercospora arecacearum 

Asteraceae 

1. Caespituli hypophyllous.............................2 



2. Conidiophores sometimes branched, 55–181 

× 4–5.5 μm; on Artemisia............................. 

................................. Cercospora artemisiae 

2. Conidiophores unbranched......................... 3 

3. Conidiophores 79–184 × 3–5 μm, geniculate; 

conidia 120–215 × 3–4 μm, narrowly 

obclavate to acicular, 8–20-septate, obconically 

truncate at the base; on Helianthus 

............................ Cercospora helianthicola 

3. Conidiophores 25–102 × 25–4 μm, not 

geniculate; conidia 46–87 × 2.5–3 μm, 

acicular, 7–10-septate, truncate at the base; 

on Dahlia..................Cercospora dahliicola 

4. On Zinnia ................................................... 5 

4. On other hosts ............................................ 6 

5. Conidiophores unbranched, 54–100 × 2.5–5 

μm; conidia 24.5–93.5 × 2.5–3.5 μm, 

filiform to narrowly obclavate, 7–18-septate; 

on Zinnia elegans .....Cercospora zinniicola 

5. Conidiophores sometimes branched, 40–152 

× 3–5.5 μm; conidia 24–175 × 2–3.5 μm, 

narrowly obclavate to subacicular, 4–13septate; 

on Zinnia grandiflora...................... 

...................................... Cercospora zinniae 

6. Conidiophores not geniculate or rarely 

geniculate, 32.5–220 × 3–8 μm; conidia 45– 

196 × 1.5–3 μm, acicular, 13–19-septate, 

truncate at the base; on Cynara.................... 

..................................... Cercospora cynarae 

6. Conidiophores distinctly geniculate........... 7 

7. Conidia obclavate or subacicular ............... 8 

7. Conidia acicular ......................................... 9 

8. Conidia obclavate, 80–96 × 3.5–4 μm; on 

Conyza...................Cercospora nilghirensis 

8. Conidia narrowly obclavate to subacicular, 

60–198 × 2–4 μm; on Gerbera..................... 

....................................Cercospora gerberae 

9. Conidia truncate at the base, 23–190 × 2–4 

μm, 3–10-septate; on Chrysanthemum 

............................Cercospora chrysanthemi 

9. Conidia obconically truncate at the base, 36- 

182 × 3–6.5 μm, 7–13-septate; on Lactuca 

........................Cercospora lactucae-sativae 

27

Balsaminaceae 

1. Conidiophores 52–129 × 2–3.5 μm, 

unbranched; conidia obclavate with obconically 

truncate at the base, 35–73 × 4–5 μm, 

3–11-septate; on Impatiens walleriana ........ 

............................Cercospora balsaminiana 

1. Conidiophores 49–112 × 4–6.5 μm, sometimes 

branched; conidia narrowly obclavate 

to subacicular with truncate to obconically 

truncate at the base, 60–120 × 2.5–5 μm, 5– 

18-septate; on Impatiens balsamina............. 

............................... Cercospora fukushiana 

Basellaceae 

A single species, on Basella............................. 

...........................Cercospora basellae-albae 

Brassicaceae 

A single species, on Brassica and Cichorium 

...............................Cercospora brassicicola 

Caricaceae 

A single species, on Carica.............................. 

.....................................Cercospora papayae 

Convolvulaceae 

1. Conidiophores sometimes branched, 13.5– 

134 × 3–5 μm; conidia 44.5–143 × 3–3.5 

μm, narrowly obclavate to subacicular, 6– 

15-septate, base obconically truncate; on 

Ipomoea and Argyreia.................................. 

.................................. Cercospora ipomoeae 

1. Conidiophores unbranched.........................2 

2. Conidia 80–240 × 3–4 μm, 9–14-septate, 

acicular to long obclavate, base obconically 

truncate; on Ipomoea.................................... 

...................................Cercospora citrullina 

2. Conidia 22.5–96 × 3–3.5 μm, 6–9-septate, 

acicular, sometimes obclavate, truncate at 

the base; on Operculina................................ 

...............................Cercospora operculinae 

Cucurbitaceae 

1. Ceaspituli amphigenous; conidiophores 52– 

106.5 × 2.5–5 μm, not geniculate; conidia 

28 

acicular, 63–296.5 × 2.5–4.5 μm, 8–26septate, 

truncate at the base; on various 

genera of Cucurbitaceae .............................. 

...................................Cercospora citrullina 

1. Caespituli epiphyllous; conidiophores 18– 

108.5 × 3–5.5 μm, strongly geniculate; 

conidia obclavate-cylindric, 41–102 × 2.5–5 

μm, 5–10-septate, obconically truncate at 

the base; on Coccinia ................................... 

...................................Cercospora cocciniae 

Euphorbiaceae 

1. Conidiophores sometimes branched, 48.5– 

83.5 × 4–6 μm, slightly geniculate; conidia 

44–256 × 1.5–3 μm, narrowly obclavate to 

subacicular, 4–18-septate, base obconically 

truncate; on Acalypha................................... 

..................................Cercospora acalyphae 

1. Conidiophores unbranched......................... 2 

2. Conidiophores 56–213 × 4–5.5 μm, slightly 

geniculate; conidia 29–160 × 3–4.5 μm, 

acicular, rarely obclavate, 3–10-septate, 

truncate at the base; on Codiaeum 

...................................... Cercospora codiaei 

2. Conidiophores 36–66 × 3–5 μm, sometimes 

constrict at the septate, rough wall, geniculate, 

mostly near the apex; conidia 51–133 

× 3–4.5 μm, obclavate to acicular, 5–11septate, 

obconically truncate at the base; on 

Phyllanthu ........ Cercospora phyllanthicola 

Fabaceae 

1. Caespituli hypophyllous; conidiophores 76– 

129 × 3.5–5 μm, rarely branched; conidia 

80–132 × 3–3.5 μm, narrowly obclavate to 

subacicular, 6–11-septate; on Glycine 

.....................................Cercospora kikuchii 


2. Stromata well-developed, 26.5–67 μm diam.; 

conidia 56–113.5 × 3–4.5 μm, narrowly 

obclavate to subacicular, 3–9-septate; on 

various genera of Fabaceae ......................... 

..................................Cercospora canescens 

2. Caespituli amphigenous; stromata small, 

25–30.75 μm diam.; conidia 39–206 × 2–4 

μm, narrowly obclavate to subacicular, 5– 

17-septate; on Crotalaria ............................. 

................................ Cercospora crotalariae

Hydrangeaceae 

A single species, on Hydrangea....................... 

...............................Cercospora hydrangeae 

Lamiaceae 

1. Caespituli epiphyllous; stromata small, 12– 

32 μm diam.; conidiophores 36–127.5 × 

2.5–4 μm, rarely branched, geniculate to 

sinuous; conidia 40–87 × 2–3 μm, narrowly 

obclavate to subacicular, 3–10-septate, base 

obconically truncate; on Clerodendrum 

fragrans ................Cercospora volkameriae 

1. Caespituli amphigenous .............................2 

2. Stromata lacking; conidiophores 78–185 × 

3–5 μm, rarely branched, geniculate; on 

Solenostemon............Cercospora kabatiana 

2. Stromata small to well developed, 14–30 

μm diam.; conidiophores 20–70 × 3–6 μm, 

unbranched, often not geniculate; on 

Clerodendrum paniculatum.......................... 

............................. Cercospora physostegiae 

Malvaceae 

A single species, on Alcea................................ 

...................................Cercospora althaeina 

Moraceae 

1. Caespituli hypophyllous; conidiophores 42- 

229 × 3–6 μm, branched; conidia 42.5–161 

× 2–4.5 μm, narrowly obclavate to 

subacicular, 7–14-septate, base obconically 

truncate; on Ficus religiosa.......................... 

.........................................Cercospora ficina 

1. Caespituli epiphyllous; conidiophores 63– 

139 × 3–4 μm, not branched; conidia 120– 

160 × 3 μm, acicular, 8–13-septate, truncate 

at the base; on Ficus carica.......................... 

....................................Cercospora elasticae 

Nyctaginaceae 

A single species, on Bougainvillea .................. 

....................Cercospora neobougainvilleae 

Orchidaceae 

A single species, on Habenaria ....................... 

........................... Cercospora habenariicola 

Oxalidaceae 


A single species, on Oxalis .............................. 

..................................... Cercospora oxalidis 

Polypodiaceae 

A single species, on Platycerium ..................... 

...................................Cercospora platycerii 

Pteridaceae 

A single species, on Pteris ............................... 

....................................Cercospora cyclosori 

Rosaceae 

A single species, on Rosa................................. 

.................................... Cercospora scharifii 

Rubiaceae 

A single species, on Coffea.............................. 

..................................Cercospora coffeicola 

Saururaceae 

A single species, on Houttuynia....................... 

.......................... Cercospora houttuyniicola 

Solanaceae 

1. Caespituli epiphyllous, conidiophores relatively 

short, 27–79.5 × 2–4.5 μm, not 

branched, not geniculate; conidia 30–71.5 × 

3–3.5 μm, narrowly obclavate, 3–6-septate; 

on Solanum torvum ..Cercospora solanacea 


2. Conidiophores 27.5–54 × 2.5–5.5 μm, unbranched, 

strongly geniculate; conidia 46.5– 

160 × 2–4 μm, obclavate to acicular, 7–15septate; 

on various genera of Solanaceae 

..................................Cercospora physalidis 

2. Caespituli amphigenous, chiefly 

hypophyllous; conidiophores 39.5–127 × 3– 

4 μm, branched, geniculate to sinuous; 

conidia 64–165 × 2–5 μm, long obclavate to 

subacicular, 6–19-septate; on Solanum 

indicum.........................Cercospora puyana 

Verbenaceae 

29

A single species, on Tectona........................... 

.................................... Cercospora tectonae 

Zingiberaceae 

A single species, on Alpinia............................. 

.................................Cercospora alpiniicola 

Acanthaceae 

Cercospora andrographidicola S.Q. Chen & 

P.K. Chi, J. South China Agric. Univ. 11: 61 

(1990). (Fig. 12) 

Leaf spots 2–5 mm diam., amphigenous, 

subcircular to irregular, distinct on the upper 

surface, brown with dark margin, without definite 

margin on the lower surface. Caespituli 

amphigenous. Stromata often lacking, rudi- 

30 

mentary to poorly developed, if present small, 

up to 29.5 µm diam., composed of a few 

subglobose and dark brown cells. Conidiophores 

16–74 × 3–6.5 µm, arranged in loose 

fascicles, 1–9-septate, arising from stromata, 

straight or flexuous, simple, thick walled, 

brown to dark brown or paler towards the apex, 

unbranched, geniculate near the apex. Conidiogenous 

cell integrated, holoblastic, monoblastic 

or polyblastic, sympodially proliferating. Conidiogenous 

loci 1.5–3 μm diam., conspicuous, 

thickened and darkened. Conidia 30.5–91 × 2– 

4 µm, obclavate to acicular, 3–15-septate, hyaline, 

straight to curved, truncate at the base 

with subacute apex, smooth, hila 1.5–2.5 μm 

diam., conspicuous, thickened and darkened. 

Specimen examined – THAILAND, 

Uttradit Province, Sak Yai National Park, on 

Fig. 12 – Line drawings of Cercospora andrographidicola on Andrographis paniculata. a. 

Stromata and conidiophores. b. Conidia. Bars = 50 µm. (Meeboon et al. 2007c). 

leaves of Andrographis paniculata Nees 

(Acanthaceae), 25 November 2004, Jamjan 

Meeboon (CMU 27924). 

Hosts – Andrographis paniculata 

(Acanthaceae) (Crous & Braun 2003, Meeboon 

et al. 2007c).

Distribution – China, Thailand (Crous & 

Braun 2003, Meeboon et al. 2007c). 

Notes – The first report of this species 

from Thailand was made by Meeboon et al. 

(2007c). 

Cercospora barleriicola Payak & Thirum., 

Indian Phytopath. 2: 191 (1949). 

= Cercospora barleriae-cristatae 

Govindu & Thirum., Sydowia 10: 273 (1957). 

(= C. apii s. lat.) (Fig. 13) 

Leaf spots 5 – 8 mm diam., amphigenous, 

dark to yellowish, only leaf decoloration. Caespituli 

amphigenous. Stromata 20–24 μm diam., 

small, substomatal, composed of a few subglobose, 

and dark brown cells. Conidiophores 

85–209 × 3–4 μm, 5–7 in loose fascicles, 5–8septate, 

arising from stromata, straight, unbranched, 

cylindrical, slightly geniculate near 


the apex, smooth, brown at the base, paler 

towards the apex. Conidiogenous cells integrated, 

holoblastic, polyblastic, sympodially 

proliferating. Conidiogenous loci 2–3 μm diam., 

conspicuous, thickened, and darkened. Conidia 

61–91 × 2–3 μm, solitary, acicular, straight, 

hyaline, 6–13-septate, smooth, truncate at the 

base, tapering toward a subacute apex, hila 

1.5–2 μm diam., conspicuous, thickened and 

darkened. 

Specimens examined – THAILAND, 

Chiang Mai Province, Chiang Dao District, 

Huay Luek Royal Project, on leaves of 

Barleria cristata L. (Acanthaceae), 6 February 

2008, Jamjan Meeboon and Iman Hidayat 

(BBH 23592); Uttradit Province, A. Nam Pad, 

Sak Yai National Park, same host, 25 

November 2004, Chiharu Nakashima and 

Jamjan Meeboon (CMU 27885). 

Fig. 13 – Line drawings of Cercospora barleriicola on Barleria cristata. a. Conidia. b. Stroma and 

conidiophores. Bars = 25 µm. (Meeboon et al. 2007b). 

31

Hosts – Barleria cristata, B. prionitis, 

Barleria sp. (Acanthaceae) (Crous & Braun 

2003, Meeboon et al. 2007b). 

Distribution – India, Jamaica, Thailand 

(Crous & Braun 2003, Meeboon et al. 2007b). 

Notes – This species belongs to C. apii s. 

lat. fide Crous & Braun (2003). The first report 

of this species from Thailand was carried out 

by Meeboon et al. (2007b). 

Literature – Chupp (1954, p. 22). 

Adiantaceae 

Cercospora adiantigena U. Braun & Crous, 

CBS Biodiversity Series 1: 44–45 (2003). 

(Fig. 14) 

Leaf spots – 5–15 mm diam., 

amphigenous, solitary, circular, brown to dark 

brown, with dark margin and grey at the center. 

Caespituli amphigenous. Stromata 9–43 μm 

32 

diam., substomatal to intraepidermal, small to 

well-developed, composed of few subglobose, 

brown to blackish brown cells. Conidiophores 

74–106 × 3–4 μm, 6–11 in loose fascicles, 1–3septate, 

arising from stromata, straight to decumbent, 

smooth, brown at the base, and paler 

toward the apex, cylindrical, unbranched, geniculate. 

Conidiogenous cells integrated, terminal, 

holoblastic, mostly polyblastic, sympodially 

proliferating. Conidiogenous loci 1.5–2.5 μm 


Conidia 53–60 × 2–3 μm, solitary, obclavate, 

straight to slightly curved, hyaline, 7–16septate, 

smooth, obconically truncate at the 


1.5–2 μm diam., thickened and darkened. 


Chiang Mai Province, A. Mae Rim, Nong Hoi 

Royal Project Foundation, on leaves of 

Doryopteris ludens J. Sm. (Adiantaceae), 12 

September 2007, Parin Noiruang (BBH 23634) 

Fig. 14 – Line drawings of Cercospora adiantigena on Doryopteris ludens. a. Conidia. b. Stroma 

and conidiophores. Bars = 50 µm. (Meeboon 2009).

Hosts – Adiantum philippense, Doryopteris 

ludens (Adiantaceae) (Crous & Braun 

2003, Meeboon 2009). 

Distribution – Tanzania (Crous & Braun 

2003), Thailand (Meeboon 2009). 

Notes – Dorypoteris ludens is a fern 

belonging to family Adiantaceae. Three species 

of Cercospora s. str. have been reported from 

Adiantum spp., viz, C. adianticola R.K. 

Srivast., A.K. Srivast. & Kamal (C. apii s. lat.), 

C. adiantigena U. Braun & Crous, and C. 

pteridigena M.K. Khan, R.K. Verm & Kamal. 

This specimen is identified as C. adian-tigena 

due to short and obclavate conidia (53–60 × 2– 

3 μm vs 40–90 × (4) 5–8 μm C. adian-tigena. 

It is distinct form for C. pteridigena due to the 

later species having very long conidio-phores 

and large conidiogenous loci (4-5 μm diam.). 

This specimen was first reported from Thailand 

by (Meeboon 2009). 

Amaranthaceae 


Cercospora canescens Ellis & G. Martin, 

Amer. Naturalist 16: 1003 (1882). 

≡ Cercosporiopsis canescens (Ellis & G. 

Martin) Miura, Flora of Manchuria and East 

Mongolia 3: 529 (1928). 

= Cercospora vignicaulis Tehon, 

Mycologia 29: 436 (1937). 

(= C. apii s. lat.) (Fig. 15) 

Leaf spots – 3–6 mm diam., amphigenous, 

dark to yellowish. Caespituli hypophyllous. 

Stromata mostly lacking, if present 

small, up to 8 μm diam., composed of 4–5 

globose to subglobose, brown to dark brown 

cells. Conidiophores 90.5–192 × 3–4 μm, up to 

5 in loose fascicles, 3–7-septate, arising from 

stomata, straight, smooth, brown at the base 

and paler toward the apex, unbranched, cylindrical, 

geniculate. Conidiogenous cells integrated, 

holoblastic, polyblastic, sympo-dially 


diam., conspicuous, thickened and darkened 

Fig. 15 – Line drawings of Cercospora canescens on Celosia argentea. a. Conidia. b. 

Conidiophores and stromata. Bars: = 50 μm. (Meeboon 2009). 

33

Conidia 8–316 × 3–4 μm, solitary, acicular, 

straight, hyaline, 10–22-septate, smooth, truncate 

at the base, tapering toward a subacute 

apex, hila 2.5–3 μm diam., conspicuous, 

thickened and darkened. 


Chiang Mai Province, Chiang Mai University, 

Multiple Cropping Centre, on leaves of Celosia 

argentea L. (Amaranthaceae), 14 August 2008, 

Jamjan Meeboon (BBH 23725); Chiang Rai 

Province, Mae Fah Luang, Mae Jan, Doi Tung 

Development, on leaves of Iresine herbstii 

Hook. (Amaranthaceae), 16 August 2008, 

Jamjan Meeboon (BBH 23586). 

Hosts – Amaranthus sp., Celosia argentea 

(Amaranthaceae), Annona odorata, A. 

squarrosa (Annonaceae), Rauvolfia serpentina 

(Apocynaceae), Verschaffeltia splendida (Arecaceae), 

Aster novibelgii (Asteraceae), Bixa 

orellana (Bixaceae), Raphanus sativus (Brassicaceae), 

Rhynchosia aurea, R. minima, Ricinus 

communis (Euphorbiaceae), Arachis hagenbeckii, 

A. hypogaea, Alysicarpus sp., Bauhinia 

alba, B. variegata, Cajanus cajan, Calopogonium 

mucunoides, Canavalia ensiformis, C. 

gladiata, C. maritima, Cassia alata, C. lathyroides, 

Cassia sp., Centrosema acutifolium, C. 

arenarium, C. brasilianum, C. macrocarpum, 

C. plumieri, C. pubescens, C. virginianum, 

Clitoria ternatea, Codariocalyx gyroides, Crotalaria 

juncea, C. mucronata, C. mysorensis, C. 

retusa, C. spectabilis, C. usaramoensis, C. 

verrucosa, C. zanzibarica, Crotalaria spp., 

Cyamopsis psoralioides, Desmodium canum, D. 

gyrans, D. gyroides, D. incanum, D. intortum, 

D. lycioides ssp. guerkei, D. repandum, D. 

turtuosum, D. uncinatum, Dolichos biflorus, D. 

daltonii, D. lablab, D. lignosus, D. trilobus, D. 

turtuosum, D. uniflorus, Erythrina addisoniae, 

E. suberosa, E. subumbrans, E. variegata, 

Flemingia macrophylla, Gliricidia sepium, 

Glycine max, G. soja, G. ussuriensis, G. wightii, 

Heylandia latebrosa, Indigofera astragalina, 

Kotschya sp., Lablab niger, L. purpureus, 

Lespedeza sp., Lathyrus odoratus, Leucaena 

leucocephala, Lotononis bainesii, Lupinus sp., 

Macroptilium atropurpureum, M. lathyroides, 

M. daltonii, M. uniflorum, Medicago sativa, 

Mimosa invisa, Mucuna pruriens, Neonotonia 

wightii, Phaseolus aconitifolius, P. angularis, 

P. atropurpureus, P. aureus, P. calcaratus, P. 

lathyroides, P. limensis, P. lunatus, P. minimus, 

34 

P. panduratus, P. radiatus, P. trilobus, P. vulgaris, 

Pistia stratiotes, Pisum sativum, Psophocarpus 

tetragonolobus, Psoralea bituminosa, P. 

drupacea, Pterocarpus marsupia, Pueraria 

hirsuta, P. lobata, P. phaseoloides, P. trilobam, 

Senna alata, S. tora, Shuteria involucrata, 

Stylosanthes guianensis, S. humilis, Vicia ungiculata, 

Vigna angularis, V. catjang, V. luteola, 

V. marina, V. mungo, V. parkeri, V. radiata, 

V. repens, V. reticulata, V. sesquipedalis, V. 

sinensis, V. umbellata, V. vexillata, Voandzeia 

subterranea (Fabaceae), Quercus sp. (Fagaceae), 

Vitis vinivera (Vitaceae), Coleus sp., 

Ocimum basilicum, Plectranthus sp. (Lamiaceae), 

Tetramnus labialis, T. uncinatus (Malphigiaceae), 

Artocarpus integrifolia (Moraceae), 

Boerhavia erecta, Commicarpus sp. 

(Nyctaginaceae), Lycopersicon esculentum, 

Solanum laciniatum (Solanaceae) (Crous & 

Braun 2003, Meeboon 2009). 

Distribution – Worldwide, including 

Australia, Bangladesh, Barbados, Brazil, 

Bolivia, Brunei, Cambodia, China, Colombia, 

Costa Rica, Cuba, Dominican Republic, 

Ecuador, Fiji, Georgia, Ghana, Guyana, Haiti, 

Hong Kong, India, Indonesia, Iran, Japan, 

Kenya, Korea, Malawi, Malaysia, Malawi, 

Mauritius, Myanmar, Nepal, New Caledonia, 

New Zealand, Nigeria, Pakistan, Panama, 

Papua New Guinea, Peru, Philippines, Puerto 

Rico, Russia, Senegal, Sierra Leone, Solomon 

Islands, Somalia, South Africa, Saint Vincent 

and the Grenadines, Sudan, Tadzhikistan, 

Taiwan, Tanzania, Thailand, Trinidad and 

Tobago, Togo, Uganda, USA, Uzbekistan, 

Vanuatu, Venezuela, Virgin Islands, Zambia 

and Zimbabwe (Crous & Braun 2003). 

Notes – This species was first reported 

from Thailand by Sontirat et al. (1980) who 

found C. canescens on Vigna radiata. Crous & 

Braun (2003) assigned this species to C. apii s. 

lat. Iresine herbstii was reported as a new host 

of C. canescens by Meeboon (2009). 

Cercospora celosiae Syd., Ann. Mycol. 27: 

430 (1929). (Fig. 16) 

Leaf spots – up to 3 mm diam., amphigenous, 

circular to subcircular, brown at central 

area (somewhat grey-brown in the centre of 

larger spots), with dark brown margin. Caespituli 

amphigenous, chiefly hypophyllous.

Stromata 19–29 µm diam., small, composed of 

a few globose to subglobose, dark brown cells. 

Conidiophores 34–85 × 2.5–5 μm, up to 13 

in moderately dense fascicles, 1–3-septate, 

straight to decumbent, unbranched, light brown 

to brown, paler and narrower towards the apex, 

plainly geniculate with 1–5 geniculation near 

the apex. Conidiogenous cells integrated, terminal, 

holoblastic, polyblastic, sympodially 


conspicuous, thickened and darkened. Conidia 

12–67 × 2.5–3.5 μm, 4–7-septate, hyaline, 

acicular, obconically truncate at the base, with 

acute apex, hila 2–3 μm in diam., thickened 

and darkened. 


Chiang Rai Province, Wiang Pa Pao, on leaves 

of Celosia argentea L., 25 November 2005, 

Jamjan Meeboon (CMU 27902 ); same locality, 

on C. argentea var. cristata (L.) Kuntze, 25 

November 2005, Jamjan Meeboon (CMU 

27893). 

Hosts – Celosia argentea, C. argentea 

var. cristata, C. aristata, C. laxa, C. plumosa, 

C. trigyna, Celosia spp. (Amaranthaceae) 

(Crous & Braun 2003, Meeboon 2009). 

Distribution – Bangladesh, Brazil, Brunei, 

Cambodia, China, Cuba, India, Indonesia, 

Japan, Malaysia, Myanmar, Nigeria, Pakistan, 

Papua New Guinea, Sabah, Sri Lanka, Sudan, 

Taiwan, Thailand, Uganda, USA, and 

Venezuela (Crous & Braun 2003). 


Notes – Cercospora celosiae on C. argentea 

and C. argentea var. cristata was 

reported from Thailand by Petcharat & 

Kanjanamaneesathian (1989), Sontirat et al. 

(1980) and Meeboon (2009). 

Cercospora ricinella Sacc. & Berl., Atti Reale 

1 st . Ven. Sci. Lett. Art. 6, Ser. 3: 721 (1885). 

≡ Cercosporina ricinella (Sacc. & Berl.) 

Speg., Anales Mus. Nac. Hist. Nat. Buenos 

Aires 20: 429 (1910). 

= Cercospora albido-maculans G. 

Winter, Hedwigia 24: 202 (1885); also in J. 

Mycol. 1: 124 (1885). (Fig. 17) 

Leaf spots – 2–9 mm diam., distinct, 

amphigenous, circular or subcircular, greyish 

brown, with reddish brown margins. Caespituli 

amphigenous. Stromata 12–26 μm diam., intraepidermal, 

small, composed of globular to 

angular, brown to blackish brown cells. Conidiophores 

47–125 × 3–5 μm, 5–9 in loose and 

divergent fascicles, 1–4-septate, arising from 

stromata, erect to decumbent, smooth, pale 

yellow to pale brown, unbranched, subcylindrical, 

strongly geniculate. Conidiogenous 

cells integrated, terminal to intercalary, holoblastic, 

polyblastic, sympodially proliferating. 

Conidiogenous loci 2.5–3 μm diam., conspicuous, 

thickened and darkened. Conidia 29- 

168.5 × 2.5–3.5 μm, solitary, narrowly obclavate 

to subacicular, 2–12-septate, straight to 

curved, hyaline, smooth, base obconically 

Fig. 16 – Line drawings of Cercospora celosiae on Celosia argentea. a. Conidiophores and stroma. 

b. Conidia. Bars = 50 µm. (Meeboon 2009). 

35

Fig. 17 – Line drawings of Cercospora ricinella on Ricinus communis. a. Conidiophores and 

stromata. b. Conidia. Bars = 50 µm. (Meeboon 2009). 

truncate, with subacute apex, hila 2–2.5 μm 

diam., thickened and darkened. 


Chiang Mai Province, San Sai, Mae Fag, on 

leaves of Ricinus communis L. (Euphorbiaceae), 

3 August 2008, Jamjan Meeboon 

(BBH 23755). 

Hosts – Ricinus communis (Euphorbiaceae) 



Angola, Argentina, Australia, Bangladesh, 

Barbados, Brazil, Bulgaria, Cambodia, China, 

Colombia, Cuba, Dominican Republic, Egypt, 

El Salvador, Ethiopia, French Polynesia, 

Georgia, Ghana, Guatemala, Haiti, India, 

Indonesia, Iran, Jamaica, Japan, Kazakhstan, 

Kenya, Korea, Malawi, Malaysia, Mauritius, 

Morocco, Mozambique, Myanmar, Nepal, New 

Caledonia, Nigeria, Pakistan, Panama, 

Philippines, Puerto Rico, Russia (European 

part), Sierra Leone, Somalia, South Africa, Sri 

Lanka, Sudan, Tahiti, Taiwan, Tanzania, 

Thailand, Togo, Trinidad and Tobago, Uganda, 

36 

Ukraine, USA, Vanuatu, Venezuela, Zimbabwe 



from Thailand by Sontirat et al. (1980). 

Literature – Chupp (1954, p. 229), 

Sontirat et al. (1980), Meeboon (2009). 

Apocynaceae 

Cercospora peregrina Chupp, Monograph of 

Cercospora: 49 (1954). 

(= C. apii s. lat) (Fig. 18) 

Leaf spots 2–5 mm diam., distinct, 

amphigenous, circular to subcircular, scattered, 

dull brown, often paler at the centre, with dark 

brown margins. Caespituli epiphyllous. Stromata 

25–37 μm diam., intraepidermal, welldeveloped, 

composed of globular to angular, 


38–139 × 3–4.5 μm, 7–12 in loose to dense 

fascicles, 2–4-septate, often divergent, arising 

from stromata, simple, erect to decumbent,

smooth, pale brown to brown, unbranched, 

subcylindrical, slightly geniculate. Conidiogenous 

cells integrated, terminal, holoblastic, 

monoblastic to polyblastic, sympodially proliferating. 

Conidiogenous loci 2–3 μm diam., 


25–170 × 3–3.5 μm, solitary, narrowly 

obclavate to subacicular, 3–12-septate, straight, 

hyaline, smooth, base obconically truncate, 

with acute apex, hila 2–3 μm diam., thickened 



Chiang Mai Province, Royal Flora, on leaves 

of Pentalinon luteum (L.) B.F. Hansen & 

Wunderlin (Apocynaceae), 27 July 2008, 


Hosts – Tabernamontana coronaria, T. 

divaricata (Apocynaceae) (Crous & Braun 

2003), Pentalinon luteum (Meeboon 2009). 

Distribution – India, Mexico, Pakistan, 

Thailand, USA (Crous & Braun 2003, 

Meeboon 2009). 

Notes – This specimen is a typical of C. 

apii s. lat fide Crous & Braun (2003) due to 

long and slightly geniculate conidiophores, and 

long acicular conidia with truncate base and 

acute apex. Cercospora peregrina is the only 

one C. apii s. lat. reported from plants family 

Apocynaceae. This specimen was first reported 

from Thailand by Meeboon (2009). 

Arecaceae 

Cercospora arecacearum Hidayat & Meeboon, 

Mycol. Prog. 8: 115-121 (2009a). 

MycoBank 510616 (Fig. 19) 

Leaf spots – 1–10 cm diam., amphigenous, 

irregular, brown to dull greyish brown, 

finally pale greyish with a white center and 

dark margins, spots usually overlapping. Caespituli 

amphigenous, scattered, dark yellowish. 

Stromata 30–100 µm diam., substomatal to 

intraepidermal, well-developed, subglobular, 

brown to blackish brown. Conidiophores 68.5– 

310 × 4–5 µm, variable in length, in dense 

fascicles, 2–8-septate, arising from stromata, 

smooth, pale yellowish to brownish throughout, 

sometimes paler at the apex, cylindrical, but 

narrowed towards the apex, straight, branched, 

strongly geniculate. Conidiogenous cells 24.5– 

67 × 4–5 µm, integrated, terminal, sympodially 


Fig. 18 – Line drawings of Cercospora 

peregrina on Pentalinon luteum. a. Conidia. b. 

Stromata and conidiophores. Bars: a = 50 μm, 

b = 40 μm. (Meeboon 2009). 

proliferating. Conidiogenous loci 2.5–3 µm 


Conidia 140–320 × 4–5 µm, acicular, straight, 

often curved at the apex, hyaline, 9–25-septate, 

thin-walled, smooth, tapered towards a 

subacute apex, base truncate, hila 2–3 µm 



Chiang Mai province, Mae Taeng District, T. 

Pa Pae, Mushroom Research Centre, on leaf 

spots of Areca catechu L. (Arecaceae), 17 

November 2006, Iman Hidayat (CMU 27946: 

Holotype). 

Habitat – Areca catechu (Arecaceae) 

(To-anun et al. 2009). 

Distribution – Thailand (type locality) 

(To-anun et al. 2009). 

Notes – According to Crous & Braun 

(2003), this species belongs to Cercospora s. 

str., which is characterized by having pigmented 

conidiophores, thickened and darkened 

conidiogenous loci, and hyaline scolecoid 

conidia. This fungus is distinct from the 

37

Fig. 19 – Symptoms, conidiophores, stroma and conidia of Cercospora arecacearum (from 

holotype). a. Symptoms. b. Stroma and conidiophores. c. Conidia. Bars: a = 5 cm; b = 50 μm; c = 

150 μm. (To-anun et al. 2009). 

plurivorous C. apii s. lat. by having welldeveloped, 

large stromata, and strongly 

geniculate, branched conidiophores in deve 

fascicles (Crous & Braun 2003). 

Currently, only three species have been 

maintained in Cercospora s. str. on Arecaceae, 

viz, C. palmae-amazonensis Bat. & Cavalc., C. 

raphiae Deighton and C. nucifera R.K. Srivast., 

S. Narayan & A.K. Srivast. (Crous & Braun 

2003). The later species, however, is now 

classified as C. apii s. lat. (Crous & Braun 

2003). 

Cercospora arecacearum is distinct from 

C. raphiae by having amphigenous caespituli, 

branched and strongly geniculate conidiophores 

as well as much narrower acicular conidia. 

Deighton (1985) characterized C. raphiae by 

having hypophyllous caespituli, unbranched, 

non-geniculate conidiophores and obclavatecylindrical 

conidia with slightly thickened hila. 

Cercospora arecacearum is also easily distinguishable 

from C. palmae-amazonensis by 

its large stromata, branched and strongly geni- 

38 

culate conidiophores with hyaline acicular 

conidia. 

Asteraceae 

Cercospora artemisiae Y.L. Guo & Y. Jiang, 

Mycosystema 19: 445 (2000). Fig. 20 

Leaf spots – 15–30 mm diam., amphigenous, 

circular to subcircular, at first pale 

greenish to ochraceous, later becoming brown 

to dark brown, finally with greyish brown at 

the centre, surrounded by a dark margin or 

brown halo. Caespituli hypophyllous. Stromata 

18–25 μm diam., substomatal, small to welldeveloped, 

composed of a few globose to 

subglobose, brown to blackish brown cells. 

Conidiophores 55–181 × 4–5.5μm, 3–10 in a 

loose fascicle, 2–7-septate, arising from stromata, 

erect to decumbent, smooth, pale yellow 

to pale brown, simple, straight, sometimes 

branched, subcylindrical, geniculate to sinuous. 

Conidiogenous cells integrated, terminal,


Fig. 20 – Line drawings of Cercospora artemisiae on Artemisia indica. b. Conidiophores and 

stromata. a. Conidia. Bars = 50 μm. (Meeboon 2009). 

holoblastic, monoblastic or polyblastic, sympodially 

proliferating. Conidiogenous loci 2.5–3 

μm diam., conspicuous, thickened and darkened. 

Conidia 43.5–207.5 × 2–4 μm, solitary, 

narrowly obclavate to subacicular, straight, 

hyaline, 4–17-septate, smooth, base obconically 

truncate, with subacute apex, hila 2–2.5 

μm diam., thickened and darkened. 



Multiple Cropping Centre, on leaves of Artemisia 

indica Willd. (Asteraceae), 14 August 

2008, Jamjan Meeboon (BBH 23726). 

Hosts – Artemisia lactiflora (Asteraceae) 

(Guo & Jiang 2000), Artemisia indica 


Distribution – China, Thailand (Guo & 

Jiang 2000, Meeboon 2009). 

Notes – Crous & Braun (2003) noted this 

species is probably a synonym of C. apii s. lat., 

but further investigation is needed to justify this 

preliminary comment. This species was first 

reported from Thailand by Meeboon (2009). 

Cercospora chrysanthemi Heald & F.A. Wolf, 

Mycologia 3: 15 (1911). 

= Cercospora chrysanthemi Puttemans, 

Bull. Soc. Roy. Bot. Belgique 48: 244 (1912) 

(nom.illeg.), homonym C. chrysanthemi Heald 

& F.A. Wolf (1911). 

≡ Cercosporina chrysanthemi Sacc., Syll 

Fung. 25: 898 (1931) (nom. nov.), as 

‘(Puttemans) Sacc.’ 

= Cercospora chrysanthemi-coronarii 

Sawada, Rep. Dept. Agric. Gov. Res. Inst. 

Formosa 2: 147 (1922). 

(= C. apii s. lat.) Fig. 21 

39


chrysanthemi on Chrysanthemum sp. a. Conidiophores 

and stromata. b. Conidia. Bars = 50 

µm. (Meeboon 2009). 


irregular, greyish brown, with dark brown 

margin. Caespituli amphigenous. Stromata 34– 

40.5 μm diam., well-developed, substomatal, 

composed of a few subglobose, brown to dark 

brown cells. Conidiophores 35–212 × 3–4.5 

μm, 3–11 in fascicles arising from stromata, 

straight, smooth, brown at the base, paler 

towards the apex, 1–10-septate, unbranched, 

cylindrical, strongly geniculate. Conidiogenous 

cells integrated, holoblastic, polyblastic, sometimes 

monoblastic and terminal, sympodially 

proliferating. Conidiogenous loci 2–3.5 μm 








Chiang Mai Province, Chiang Dao District, 

Huay Luek Royal Project, on leaves of 

40 

Chrysanthemum sp. (Asteraceae), 6 February 


(BBH 23577). 

Hosts – Callistephus chinensis, Centratherum 

anthelminticum, Chrysanthemum balsamita, 

C. coronarium, C. hortorum, C. indicum, 

C. maximum, C. morifolium hybrid, C. segetum, 

C. sinense, Chrysanthemum sp. (Asteraceae) 


Distribution – Bermuda, Brazil, China, 

Georgia, Hong Kong, India, Jamaica, Japan 

Korea, Mauritius, Myanmar, New Zealand, 

Panama, Philippines, Taiwan, Thailand, USA 




Cercospora cynarae Y.L. Guo & Y. Jiang, 

Mycosystema 20: 26 (2001). Fig. 22 


distinct, circular to subcircular, pale brown to 

tan, centre greyish brown to greyish white, 

with dark brown margins. Caespituli amphigenous. 

Stromata often lacking, if present, up 

to 28 μm diam., composed of a few globose, 

brown to dark brown cells. Conidiophores 

32.5–220 × 3–8 μm, variable in length, loosely 

fasciculate, 3–12-septate, emerging from stromata 

through the cuticle or secondary mycelium, 

straight to slightly curved, pale brown or 

sometimes paler towards the apex, unbranched, 

rarely geniculate. Conidiogenous cells integrated, 

terminal, often monoblastic, sympodially 



Conidia 45–196 × 1.5–3 μm, solitary, acicular, 

curved, hyaline, 13–19-septate, smooth, 

truncate at the base, apex acute, hila ± 1 μm 



Chiang Mai Province, Mae-jam District, Maehae 

Royal Project Area, on leaves of Cynara 

scolymus L. (Asteraceae), 12 February 2008, 

Jamjan Meeboon and Iman Hidayat (BBH 

23674). 

Hosts – Cynara scolymus (Asteraceae) 

(Jiang & Guo 2001, Meeboon 2009). 

Distribution – China, Thailand (Jiang & 

Guo 2001, Meeboon 2009). 


from Thailand by Meeboon (2009).


Fig. 22 – Line drawings of Cercospora cynarae on Cynara scolymus. a. Conidiophores. b. Conidia. 

Bars = 50 μm. (Meeboon 2009). 

Fig. 23 – Line drawings of Cercospora dahliicola on Dahlia sp. a. Conidiophores and stromata. b. 

Conidia. Bars = 50 µm. (Meeboon 2009). 

Cercospora dahliicola M.A. Salam & P.N. Rao, 

J. Indian Bot. Soc. 36: 424 (1957). 



dark to yellowish, only leaf decoloration. 

Caespituli hypophyllous. Stromata 19–21 μm 

diam., small to well-developed, composed of a 

few globose to subglobose, brown to blackish 

brown cells. Conidiophores 25–102 × 25–4 μm, 

3–5 in a loose and divergent fascicle, 1–3septate, 

arising from stromata, straight, smooth, 

brown at the base, paler towards the apex, 

cylindrical, unbranched, not geniculate. Conidiogenous 

cells integrated, holoblastic, monoblastic, 

sympodially proliferating. Conidioge- 

41

Fig. 24 – Line drawings of Cercospora gerberae on Gerbera jamesonii. a. Conidiophores and 

stromata. b. Conidia. Bars = 50 μm. (Meeboon 2009). 

nous loci 2.5–3 μm diam., conspicuous, 

thickened and darkened. Conidia 46–87 × 2.5– 

3 μm, solitary, acicular, straight, hyaline, 7–10septate, 

smooth, truncate at the base, tapering 

toward a subacute apex, hila 2–2.5 μm diam., 

conspicuous, thickened and darkened. 


Chiang Rai Province, Mae Fah Luang, Mae Jan, 

Doi Tung Development, on leaves of Dahlia sp. 

(Asteraceae), 16 August 2008, Jamjan 

Meeboon (BBH 23587). 

Hosts – Dahlia variabilis, Dahlia sp. 

(Asteraceae) (Salam & Rao 1957, Meeboon 

2009). 

Distribution – India, Thailand (Salam & 

Rao 1957, Meeboon 2009). 

42 


from Thailand by Meeboon (2009). Crous & 

Braun (2003) assigned this species as C. apii s. 

lat. 

Cercospora gerberae Chupp & Viégas, Bol. 

Soc. Brasil. Agron. 8: 27 (1945). 



circular or subcircular, pale greenish to 

ochraceous when young, becoming brown to 

dark brown, finally greyish brown at the centre, 

surrounded by a dark margin. Caespituli amphigenous. 

Stromata 20.5–39 μm diam., welldeveloped, 

intraepidermal, composed of a few


helianthicola on Helianthus annuus. a. Conidia. 

b. Stromata and conidiophores. Bars = 50 μm. 



Conidiophores 36–163 × 3–6 μm, numerous, in 

loose to dense fascicles, 1–3-septate, arising 

from stromata, simple, straight, erect to decumbent, 

smooth, pale yellow to pale brown, 

unbranched, subcylindrical, strongly geniculate 

to sinuous. Conidiogenous cells integrated, 

terminal, polyblastic, sympodially proliferating. 

Conidiogenous loci 2–3 μm, conspicuous, 

thickened and darkened. Conidia 60–198 × 2–4 

μm, solitary, narrowly obclavate to subacicular, 

straight, hyaline, 5–12-septate, smooth, base 

obconically truncate, with subacute apex, hila 

2–2.5 μm, thickened and darkened. 


Chiang Rai Province, A. Wiang Pa Pao, on 

leaves of Gerbera jamesonii Adlam cultivar 

(Asteraceae), 9 March 2005, Jamjan Meeboon 

(CMU 28219); Chiang Mai Province, A. 

Muang, Suthep, Chang Khian, same host, 2 

August 2008, Jamjan Meeboon (BBH 23690); 

Chiang Mai Province, Mae Jo, San Sai Farming 


area, same host, 9 August 2008, Jamjan 


Hosts – Gerbera jamesonii, Gerbera sp. 

(Asteraceae) (Crous & Braun 2003). 

Distribution – Australia, Bangladesh, 

Bermuda, Brazil, Brunei, Cuba, Cambodia, 

Ghana, Hong Kong, India, Indonesia, Iran, 

Jamaica, Kenya, Malawi, Malaysia, Pakistan, 

Philippines, Puerto Rico, Sierra Leone, 

Singapore, Solomon Islands, Taiwan, Tanzania, 

Thailand, Uganda, USA, Virgin Islands (Crous 

& Braun 2003). 

Notes – The first report of this species in 

Thailand was by Sontirat et al. (1980). 


Cercospora helianthicola Chupp & Viégas, 

Bol. Soc. Brasil. Agron. 8: 29 (1945). 


Leaf spots 2–26 mm diam., distinct, 

amphigenous, variable in shapes, from minute 

spot to large necrosis on the leaves, dark brown, 

with indistinct margins. Caespituli hypophyllous. 

Stromata 12-15 μm diam., intraepidermal, 

small, composed of a few globose to subglobose, 


79–184 × 3–5 μm, 3–6 in loose and 

divergent fascicles, 2–4-septate, arising from 

stromata, erect to decumbent, smooth, pale 


geniculate to sinuous. Conidiogenous 

cells integrated, terminal to intercalary, holoblastic, 

monoblastic or polyblastic, sympodially 



Conidia 120–215 × 3–4 μm, solitary, 

narrowly obclavate to acicular, 8–20-septate, 

straight to curved, hyaline, smooth, base 


1.5–2.5 1.5–3 μm diam., thickened and 

darkened. 


Chiang Mai Province, Suthep-Pui National 

Park, on leaves of Helianthus annuus L. 

(Asteraceae), 30 November 2004, Jamjan 

Meeboon (CMU 27879); Chiang Mai Province, 

Chiang Mai University, Faculty of Agriculture, 

on same host, 14 August 2008, Jamjan 


Hosts – Helianthus annuus, H. doronicoides, 

H. hirsutus, H. maximiliani, H. occi- 

43

dentalis, H. rigidus, H. strumosus, H. tuberosus 

(Asteraceae) (Crous & Braun 2003). 

Distribution – Brazil, Cambodia, China, 

India, Mauritius, Pakistan, Panama, Thailand 


Notes – In Thailand, this fungus was first 

reported by Petcharat & Kanjanamaneesathian 

(1989). Crous & Braun (2003) noted C. 

helianthicola as C. apii s. lat. 


Cercospora lactucae-sativae Sawada, Report 

of the Department of Industry, Government 

Research Institute, Formosa 85: 111 (1943). 

= Cercospora longispora (Cugini) Trav., 

Malpighia 17: 217 (1902) (nom.illeg.), 

homonym of C. longispora Peck (1884). 

≡ Cercospora longissima Trav., 

Malpighia 17: correzione (correction slip) to p. 

217 (1903) (nom. illeg.), homonym of C. 

longissima Cooke & Ellis (1889). 

≡ Cercospora longisima (Cugini) Sacc., 

Syll. Fung. 18: 607 (1906) (nom.illeg.), 

homonym of C. longissima Cooke & Ellis 

(1889). 

= Cercospora lactucae J.A. Stev., J. 

Dept. Agric. Puerto Rico 1: 105 (1917) 

(nom.illeg.), homonym of C. lactucae Henn. 

(1902). 

= Cercospora lactucae Welles, 

Phytopathology 13: 289 (1923) (nom.illeg.), 

homonym of C. lactucae Henn. (1902). 

= Cercospora ixeridis-chinensis Sawada, 

Rep. Gov. Agric. Res. Inst. Taiwan 86: 171 

(1943) (nom. inval.). 

= Cercospora lactucae-indicae Sawada, 


(1943) (nom. inval.). Fig. 26 


circular or subcircular, brown to dark brown, 

greyish brown at the centre, surrounded by 

dark margins. Caespituli amphigenous. Stromata 

17–36 μm diam., intraepidermal, moderately 

small to well-developed, composed of 

subglobular, and brown to dark brown cells. 

Conidiophores 47–128 × 3–6.5 μm, 3–8 in 

loose fascicles, 1–4-septate, arising through 

stomata, straight to decumbent, smooth, brown 

at the base, paler towards the apex, unbranched, 

cylindrical, strongly geniculate near the apex. 

Conidiogenous cells 19–40 × 2–3.5 μm, 

44 


lactuca-sativae of Lactuca sativa cv. red leaf 

lettuce. a. Conidia. b. Conidiophores and 

stromata. Bars = 50 μm. (Meeboon 2009). 

integrated, terminal, monoblastic or polyblastic, 

sympodially proliferating. Conidiogenous loci 

2–3 μm diam., conspicuous, thickened and 

darkened. Conidia 36–182 × 3–6.5 μm, solitary, 

acicular to narrowly obclavate, straight (occasionally 

curved), hyaline, 7–13-septate, smooth, 

obconically truncate at the base, tapering 

towards a subacute apex, hila 1.5–3 μm diam., 




on leaves of Lactuca sativa L. (Asteraceae), 31 

October 2004, Jamjan Meeboon (CMU 27900); 

the same province, Amphur Samoeng, Pang Da 

Royal Project, on leaves of Lactuca sativa cv. 

butter head lettuce, 7 February 2008, Jamjan 

Meeboon and Iman Hidayat (BBH 23572); cv. 

red leaf lettuce, Jamjan Meeboon and Iman 

Hidayat (BBH 23573); and cv. green corol, 


23572); the same province Amphur Sanpatong, 

Tambol Mae Win, Ban Mae Sapok, Mae Sapok 

Royal Project, 8 February 2008, cv. red corol, 


23569) and cv. red oak leaf, Jamjan Meeboon


Fig. 27 – Line drawings of Cercospora nilghirensis on Conyza sumatrensis. a. Conidia. b. 

Conidiophores and stromata. Bars: a = 25 µm, b = 50 µm. (Meeboon 2009). 

and Iman Hidayat (BBH 23570); cv. ice berg, 


23633); cv. lettuce green oak leaf, Jamjan 

Meeboon and Iman Hidayat (BBH 23597); and 

cv. lettuce sweet chart, Jamjan Meeboon and 

Iman Hidayat (BBH 23631) 

Hosts – Cichorium endivia, C. intybus L., 

Lactuca chinensis, L. denticulata, L. indica, L. 

paradoxa, L. saligna, L. scariola, and L. sativa 

(Asteraceae) (Crous & Braun 2003, Meeboon 

et al. 2007c). 

Distribution – Worldwide, wherever the 

host is grow, including China, Japan, Korea, 

Taiwan, Thailand (Crous & Braun 2003, 

Meeboon et al. 2007c). 

Notes – The first report of this species 

from Thailand was made by Meeboon et al. 

(2007c). The first report of C. lactucae-sativae 

infecting Cichorium endivia, and various 

cultivars of L. sativa, viz, cv. butter head 

lettuce, cv. red leaf lettuce, cv. green corol, cv. 

red corol, cv. red oak leaf, cv. ice berg, cv. 

lettuce green oak leaf, and cv. lettuce sweet 

chart, in Thailand was by Meeboon (2009). 

Cercospora nilghirensis Govindu & Thirum., 

Sydowia 9: 224 (1955). Fig. 27 


distinct, circular to subcircular, pale to whitish 

at the center with dark margin. Caespituli 

amphigenous. Stromata up to 12 μm diam., 

small, often lacking, intraepidermal, composed 

of a few globose to subglobose, brown cells. 

Conidiophores 88–118 × 4–6 μm, 4–9 in loose 

to dense fascicles, 2–5-septate, arising from 

stromata, straight to decumbent, smooth, brown 

at the base, paler toward the apex, unbranched, 

cylindrical, geniculate, mostly near the apex. 

Conidiogenous cells integrated, terminal or 

intercalary, frequently monoblastic, sometimes 

polyblastic, sympodially proliferating. Conidiogenous 

loci 2–2.5 μm diam., conspicuous 

thickened and darkened. Conidia 40–96 × 3.5– 

45

4 μm, solitary, obclavate, straight, slightly 

curved, hyaline, 6–12-septate, smooth, obconically 

truncate at the base, tapering towards a 

subacute apex, hila 2–2.3 μm diam., conspicuous, 



Chiang Mai Province, Mae Jam District, Mae 

Hae Royal Project, on leaves of Conyza 

sumatrensis (Retz.) E. Walker (Asteraceae), 12 

February 2008, Jamjan Meeboon and Iman 

Hidayat (BBH 23775). 

Hosts – Conyza ambigua, C. stricta, C. 

sumatrensis (Asteraceae) (Crous & Braun 2003, 


Distribution – India, Thailand (Crous & 


Notes – This specimen is much close to 

C. nilghirensis than to C. bidentis Tharp. due to 

distinct and amphigenous leaf spot, amphigenous 

caespituli, stromata small to lacking, 

unbranched and geniculation of conidiophores 

near the apex, and the conidia frequently 

obclavate. In C. bidentis, the leaf spot is 

indefinite as the lower surfaces and caespituli 

epiphyllous. The first report of C. nilghirensis 

from Thailand was by Meeboon (2009). 

Cercospora zinniicola A. Pande, Kavaka 3: 55 

(1975). Fig. 28–29 


irregular, pale, with dark red margin, numerous 

and scattered through the leaf surface. Caespituli 

amphigenous. Stromata 32.5–46 μm 

diam., intraepidermal, well-developed, composed 

of globose to subglobose, brown to 

blackish brown cells. Conidiophores 54–100 × 

2.5–5 μm, 9–16 in dense fascicules, not 

divergent, 3–6-septate, arising from stromata, 

smooth, brown at the base, paler toward the 

apex, straight to decumbent, unbranched, cylindrical, 

geniculate to sinuous, mostly near the 

apex. Conidiogenous cells 7.5–20 × 2.5–5 μm, 

integrated, terminal, holoblastic, mostly polyblastic, 

sympodially proliferating. Conidiogenous 

loci 2–2.5 μm diam., conspicuous, 

thickened and darkened. Conidia 24.5–93.5 × 

2.5–3.5 μm, solitary, filiform to narrowly obclavate, 

straight, hyaline, 7–18-septate, smooth, 




46 


Chiang Rai Province, A. Wiang Pa Pao, T. 

Wiang Ga Long, Moo11, Bahn Tung Ruang 

Tong, on leaves of Zinnia elegans Jacq 

(Asteraceae), 31 July 2007, Jamjan Meeboon 

(BBH 23563); Chiang Mai Province, Mae Rim, 

Queen Sirikit Botanical Garden, on leaves of Z. 

elegans, 5 August 2008, Jamjan Meeboon, 

(BBH 23731). 

Hosts – Zinnia elegans (Asteraceae) 


Distribution – India, Thailand (Crous & 


Notes – Both of these specimens are 

apparently distinct from C. apii s. lat. due to 

well developed stromata, very densely fasciculate 

conidiophores and obclavate conidia 

with obconically truncate base. Both of them 

are characterized by having amphigenous 

caespituli as well as having relatively short 

conidiophores (up to 100 μm long) and conidia. 

The conidiophores of the first specimen are not 

divergent, but the second one is divergence and 

conidia of the second specimen are only up to 4 

septate. Since these collections are not C. apii s. 

lat., therefore, we assigned them to C. zinniicola 

due to the similarity of morphological 

characteristics, and being recorded from Zinnia 

elegans. Cercospora zinniicola was firstly 

found in Thailand by Meeboon (2009). 

Literature – Crous & Braun (2003, p. 

434). 

Cercospora zinniae Ellis & G. Martin, J. 

Mycol. 1: 20 (1885). 

= Cercospora atricincta Heald & F.A. 

Wolf, Mycologia 3: 14 (1911). 

= Cercospora zinniae Takah. & Yosh., 

Pl. Protect. Tokyo 7: 17 (1953). 



circular or subcircular, at first pale 

greenish to ochraceous, later brown to dark 

brown, finally with greyish brown centre, 

surrounded by a dark margins. Caespituli 

amphigenous, effuse. Stromata 17–25 μm 

diam., intraepidermal, small to well-developed, 

composed of globose to subglobose, brown to 


3–5.5 μm, 9–13 in dense fascicles, often


Fig. 28 – Line drawings of Cercospora zinniicola on Zinnia elegans. a. Conidia. b. Conidiophores 

and stroma. Bars = 50 µm. (Meeboon 2009). 

Fig. 29 – Line drawings of Cercospora zinniicola on Zinnia elegans. a. Conidiophores and stromata. 

b. Conidia. Bars = 50 µm. (Meeboon 2009). 

47

Fig. 30 – Line drawings of Cercospora zinniae 

on Zinnia grandiflora. a. Conidia. b. 

Conidiophores and stromata. Bars = 50 µm. 


divergent, 1–4-septate, arising from stromata, 

simple, straight, erect to decumbent, smooth, 

pale yellow to pale brown, rarely branched, 

subcylindrical, geniculate to sinuous. Conidiogenous 


polyblastic, sympodially proliferating. Conidiogenous 

loci 1.5–3 μm diam., conspicuous, 

thickened and darkened. Conidia 24–175 × 2– 

3.5 μm, solitary, narrowly obclavate to subacicular, 

straight, hyaline, 4–13-septate, smooth, 

obconically truncate at the base, with subacute 

apex, hila 1.5–2.5 μm diam., thickened and 

darkened. 


Chiang Mai Province, Mae Rim, Queen Sirikit 

Botanical Garden, on leaves of Zinnia grandiflora 

Nutt. (Asteraceae), 5 August 2008, 


Hosts – Cosmos sp., Zinnia elegans, Z. 

multiflora, Z. pauciflora, Z. peruviana, Z. 

violacea, Zinnia sp. (Asteraceae) (Crous & 

Braun 2003). 

Distribution – American Samoa, 

Bangladesh, Bhutan Brazil, Brunei, China, 

48 


balsaminiana on Impatiens walleriana. a. 

Conidiophores and stromata. b. Conidia. Bars: 

a = 50 µm, b = 40 µm. (Meeboon 2009). 

Colombia, Cook Islands, Cuba, Dominican 

Republic, El Salvador, Fiji, Ghana, Guam, 

Guatemala, Haiti, Hong Kong, India, Indonesia, 

Jamaica, Japan, Korea, Lithuania, Malawi, 

Malaysia, Mauritius, Mexico, Micronesia, 

Myanmar, Nepal, New Caledonia, Nigeria, 

Pakistan, Panama, Papua New Guinea, 

Philippines, Puerto Rico, Samoa, Singapore, 

Solomon Islands, South Africa, Sudan, Taiwan, 

Tanzania, Thailand, Togo, Tonga, Trinidad and 

Tobago, Tuvalu, Uganda, USA, Vunatu, 

Venezuela, Virgin Islands, Zambia and 

Zimbabwe (Crous & Braun 2003, Meeboon 

2009). 

Notes – Cercospora zinniae was first 



Meeboon (2009). 

Balsaminaceae 

Cercospora balsaminiana J.M. Yen & Lim, 

Cah. Pacifique 14: 91 (1970). Fig. 31 


circular to subcircular, brown to dark 

brown, pale at the center, with dark margin. 

Caespituli amphigenous. Stromata 10 – 15 μm

diam., substomatal to intraepidermal, small, 

composed of a few globose to subglobose, 

brown cells. Conidiophores 52–129 × 2–3.5 

μm, 6–8 in loose fascicles, 2–4-septate, arising 

from stromata, straight to decumbent, smooth, 

brown at the base, paler toward the apex, 

unbranched, cylindrical, geniculate, mostly 

near the apex. Conidiogenous cells integrated, 

holoblastic, polyblastic, sometimes monoblastic, 

sympodially proliferating. Conidiogenous 

loci 1–2 μm, conspicuous, thickened and 

darkened. Conidia 35–73 × 4–5 μm, solitary, 

obclavate, slightly curved, hyaline, 3–11septate, 


base, with tapering toward a subacute apex, 

hila 1–2.3 μm diam., thickened and darkened. 


Chiang Mai Province, A. Mueang, Sri Pum, 

Chiang Mai Public Garden, on leaves of 

Impatiens walleriana Hook. f. (Balsaminaceae), 

15 August 2008, Jamjan Meeboon (BBH 

23582). 

Hosts – Impatiens balsamina (Balsaminaceae) 

(Yen & Lim 1980), Impatiens 

walleriana (Meeboon 2009). 

Distribution – Singapore, Thailand (Yen 

& Lim 1980, Meeboon 2009). 

Notes – The symptoms, stromata and 

conidiophores of this specimen are close to C. 

apii s. lat. fide Crous & Braun (2003), but the 

conidia are obclavate with an obconically 

truncate base. We decide to assign this 

specimen to C. balsaminiana. The first report 

of C. balsaminiana from Thailand was by 


Literature – Yen & Lim (1980, p. 155). 

Cercospora fukushiana (Matsuura) W. 

Yamam., J. Soc. Trop. Agric. 6: 601 (1934). 

≡ Cercospora fukushiana Matsuura, J. 

Pl. Prot. 14: 699 (1927). 

= Cercospora balsaminae Mend. Philipp. 

J. Sci. 75: 166 (1941). 

= Cercospora balsaminae Kellerm. & 

Swingle, fide Chupp (1954). 






surrounded by a dark margin. Caespituli 



fukushiana on Impatiens balsamina. a. Conidia. 

b. Conidiophores and stromata. Bars: a = 100 

µm, b = 40 µm. (Meeboon 2009). 

amphigenous. Stromata 25-32 μm diam., small 

to well-developed, composed of globose to 



loose to dense fascicles, arising from stromata, 

simple, straight, 1–4-septate, erect to decumbent, 

smooth, subcylindrical, pale yellow to 

pale brown, sometimes branched, geniculate to 

sinuous. Conidiogenous cells integrated, terminal, 

holoblastic, monoblastic or polyblastic, 


conspicuous, thickened, and darkened. Conidiogenous 

loci 2–3 μm diam., conspicuous, 

thickened, and darkened. Conidia 60–120 × 

2.5–5 μm, solitary, narrowly obclavate to 

subacicular, 5–18-septate, straight, hyaline, 

smooth, base truncate to obconically truncate, 

with subacute apex, hila 1.5–2.5 μm diam., 



Phetchabun Province, Nam Nao National Park, 

on leaves of Impatiens balsamina L. 

49

Fig. 33 – Line drawings of Cercospora basellae-albae on Basella alba. a. Conidiophores and 

stromata. b. Conidia. Bars = 50 µm. (Meeboon 2009). 

(Balsaminaceae), 24 November 2004, Jamjan 

Meeboon (CMU 27917); Chiang Mai Province, 

Samerng District, Pang Da Royal Project, on 

leaves of I. balsamina, 7 February 2008, 


Hosts – Impatiens acaulis, I. balsamina, 

I. biflora, I. chinensis, I. gigantea, I. hawkeri, 

hybrid, I. noli-tangere, Impatiens sp. (Balsaminaceae) 

(Crous & Braun 2003, Meeboon 

2006, 2009). 

Distribution – Bangladesh, Bhutan Brazil, 

Brunei, China, Cuba, Estonia, Hong Kong, 

India, Indonesia, Iran, Japan, Korea, Lithuania, 

Korea, India, Indonesia, Malaysia, Mauritius, 

Myanmar, Nepal, New Caledonia, Papua New 

Guinea, Philippines, Sierra Leone, South 

Africa, Sudan, Taiwan, Tanzania, Thailand, 

USA (Crous & Braun 2003, Meeboon 2006, 

2009). 

Notes – In Thailand, C. fukushiana on I. 

balsamina was first reported by Meeboon 

(2006, 2009). 


50 

Basellaceae 

Cercospora basellae-albae R.K. Srivast., S. 

Narayan & A.K. Srivast., Indian Phytopathol. 

47: 229 (1994). Fig. 33 


subcircular to irregular, pale brown to dingy 

grey in the center with reddish brown to 

purplish brown margin on the upper surface, 

pale brown to olivaceous brown on the lower 

surface. Caespituli amphigenous. Stromata 13– 

53 μm in diam., small to well-developed, rarely 

lacking, irregular, composed of a few globose 

to subglobose, dark brown cells. Conidiophores 

25–70 × 3–6 μm, 10–15 in divergent 

fascicles, 1–4-septate, emerging through stomata 

and the cuticle, light brown, paler towards 

the apex, straight to slightly curved, unbranched, 

plainly geniculate near the apex. Conidiogenous 

cells integrated, terminal or intercalary, 

holoblastic, monoblastic or polyblastic, sympodially 

proliferating. Conidiogenous loci 2–3 

μm, conspicuous, thickened and darkened.

Conidia 17–93 × 3–5 μm, solitary, acicular to 

obclavate, hyaline, 6–12-septate, non-constricted 

at the septa, acute to obtuse at the apex, 

truncate at the base, hila 1–3 µm in diam., 



Chiang Mai province, Chiang Mai University, 

on leaves of Basella alba L. (Basellaceae), 27 


28214). 

Hosts – Basella alba (Basellaceae) 

(Srivastava et al. 1994, Meeboon et al. 2007d). 

Distribution – India, Thailand (Srivastava 

et al. 1994, Meeboon et al. 2007d). 

Notes – Cercospora basellae-albae on 

Basella alba has been previously recorded in 

India. The first report of this species from 

Thailand was by Meeboon et al. (2007d). 

Crous & Braun (2003) noted that this species is 

a true Cercospora s. str., close to or identical 

with C. apii s. lat. 

Brassicaceae 

Cercospora brassicicola Henn., Bot. Jahrb. 

Syst. 37: 166 (1906). 

= Cercospora brassicae-campestris 

Rangel, Arq. Mus. Nac., Rio de Janeiro 18: 163 

(1917). 

≡ Cercosporina brassicae-campestris 

(Rangel) Sacc., Syll. Fung. 25: 899 (1931). 

= Cercospora brassicae-juncea Sawada 

(brassicae-yunciae), Special Publ. Coll. Agric. 

Natl. Taiwan Univ. 8: 212 (1959) (nom. nud.). 

= Cercospora bloxami auct. sensu E. 

Young, Mycologia 8: 43 (1916). Fig. 34 


irregular, brown to dark brown, pale at the 

center, with dark margin, and limited by vein 

of the leaf. Caespituli amphigenous. Stromata 

12.5–19.5 μm diam., substomatal to intraepidermal, 

small, composed of a few globose to 

subglobose, brown cells. Conidiophores 24.5– 

64 × 3.5–5 μm, 6–11 in loose to dense fascicles, 

1–3-septate, arising from stomata, straight to 

decumbent, smooth, unbranched, cylindrical, 

geniculate near the apex, brown at the base, 

paler toward the apex. Conidiogenous cells 

integrated, terminal or intercalary, phyloblastic, 





brassicicola on Raphanus sativus. a. 

Conidiophores and stromata. b. Conidia. Bars 

= 50 µm. (Meeboon 2009). 

darkened. Conidia 54–135 × 2.5–5 μm, solitary, 

obclavate to subacicular, straight, slightly 

curved, hyaline, 7–16-septate, smooth, obconically 

truncate at the base, tapering toward a 

subacute apex, hila 2.5–4.5 μm diam., 



Faculty of Agriculture, Chiang Mai University, 

Chiang Mai Province, on leaves of Brassica 

pekinensis Skeels (Brassicaceae), 31 October 

2004, Jamjan Meeboon (CMU 27891); on 

leaves of B. campestris L. (Brassicaceae), 15 


on leaves of B. rapa L. (Brassicaceae), 9 


27905); Suthep-Pui National Park, Chiang Mai 

Province, on leaves of B. juncea (L.) Czern. 

(Brassicaceae), 21 November 2004, Chiharu 

Nakashima and Jamjan Meeboon (CMU 

27901); Chiang Rai Province, A. Wiang Pa Pao, 

on leaves of B. oleracea L. (Brassicaceae), 19 



Royal Project, on leaves of Raphanus sativus L. 

(Brassicaceae), 12 September 2007, Jamjan 

Meeboon and Iman Hidayat (BBH 23639); 

51

same locality, on leaves of Cichorium endivia 

L. (Asteraceae), 6 June 2007, Jamjan Meeboon 

and Iman Hidayat (JMC 29). 

Hosts – Brassica alba, B. alboglabra, B. 

campestris, B. chinensis, integrifolia, juncea, B. 

kaber, B. napus, B. nigra, B. oleracea, B. 

pekinensis, B. petsai, B. rapa, Brassica spp., 

Matthiola incana, Raphanus sativus (Brassicaceae) 



Angola, Armenia, Australia, Belarus, Brazil, 

China, Colombia, Cuba, Dominican Republic, 

Estonia, Great Britain, India, Indonesia, 

Jamaica, Japan, Kazakhstan, Kenya, Korea, 

Latvia, Lithuania, Malaysia, Malawi, Mauritius, 

Myanmar, Nigeria, Niue, Papua, New Guinea, 

Peru, Philippines, Puerto Rico, Russia, Sierra 

Leone, South Africa, Solomon Islands, Somalia, 

Sri Lanka, Sudan, Taiwan, Tanzania, Thailand, 

Togo, Trinidad and Tobago, Togo, Uganda, 

Ukraine and USA (Crous & Braun, 2003). 

Notes – Reports of C. brassicicola on 

Brassica from Thailand were published by 

Puckdeedindan (1966), and Petcharat & Kanjanamaneesathian 

(1989). Raphanus sativus 

was first reported by Meeboon (2009) as a new 

host of C. brassicicola. 


Caricaceae 

Cercospora papayae Hansf., Proc. Linn. Soc. 

London 155: 58 (1943). 

= Cercospora papayae Chupp & Viégas, 

Arq. Mus. Nac. Rio de Janeiro 8: 42 (1945). 

≡ Cercospora mamaonis Viégas & 

Chupp, Monograph of Cercospora: 107 (1954). 

≡ Pseudocercospora mamaonis (Viégas 

& Chupp) Tak. Kobay. & Tokash., Ann. 

phytopath. Soc. Japan 61: 51 (1995). 

(= C. apii s. lat) Fig. 35 


scattered to confluent, distinct, circular to 

subcircular, pale brown, centre greyish, with 

dark brown margins. Caespituli amphigenous. 

Stromata 12–34 μm in diam., small to welldeveloped, 

often rudimentary to poorly developed, 

intraepidermal, composed of a few 

sub-globular to irregular, brown cells. Conidiophores 

46–202 × 3–5.5 μm, very variable in 

52 

Fig. 35 – Line drawings of C. papayae (C. apii 

s. lat.) on Carica papaya. a. Conidia. b. 

Conidiophores and stromata. Bars = 50 µm. 


length, 4–13 in divergent fascicles, 1–9-septate, 

emerging from stromata and through the cuticle, 

pale olivaceous brown or sometimes paler 

towards the apex, smooth, straight to slightly 

curved, mostly strongly geniculate. Conidiogenous 

cells integrated, terminal, sympodially 




straight to mildly curved, hyaline, 14–36septate, 


base, tapering toward a subacute apex, hila, 1– 

2.5 μm diam., thickened and darkened. 


Chiang Mai Province, A. Sarapee, on leaves of

Carica papaya L. (Caricaceae), 12 September 


(BBH 23650). 

Hosts – Carica monoica, C. papaya 

(Caricaceae) (Crous & Braun 2003). 

Distribution – Barbados, Brazil, 

Cambodia, China, Cuba, Guatemala, India, 

Indonesia, Malawi, Mauritius, Myanmar, Nepal, 

Pakistan, Panama, Papua New Guinea, 

Philippines, Seychelles, Somalia, Sudan, 

Thailand, Togo, Tonga, Uganda, USA, 

Venezuela, Zimbabwe (Crous & Braun 2003, 


Note – Cercospora papayae was first 


Literature – Chupp (1954, p. 107), Ellis 

(1976, p. 247). 

Convolvulaceae 

Cercospora citrullina Cooke Grevillea 12: 31 

(1883). 

= Cercospora cucurbitae Ellis & Everh., 

J. Mycol. 4: 3 (1883). 

= Cercospora sechii J.A. Stev., Puerto 

Rico Agric. Exp. Sta. Rep.: 137 (1919). 

= Cercospora momordicae McRae, Ann. 

Cryptog. Exot. 2: 267 (1929). 

= Cercospora trichosanthis McRae, Ann. 

Cryptog. Exot. 2: 270 (1929). 

= Cercospora luffae Hara, Diseases of 

cultivated plants: 228 (1928). 

= Cercospora chardoniana Chupp, 

Monogr. Univ. Puerto Rico, B, 2: 245 (1934). 

= Cercospora momordicae Mend., 

Philipp. J. Sci. 75: 173 (1941). (nom. illeg.), 

homonym of C. momordicae McRae (1929). 

= Cercospora momordicae Sawada, Rep. 

Gov. Agric. Res. Inst. Taiwan 86: 173 (1943), 

(nom. inval.), homonym of C. momordicae 

McRae 1929. 


Leaf spots 1–5 mm in diam., amphigenous, 

scattered to confluent, distinct, circular 

to subcircular, pale brown, greyish brown to 

greyish white at the center, with dark brown 

margins. Caespituli amphigenous. Stromata 7– 

41.5 μm in diam., small, to well-developed, 

irregular, composed of a few globose to 

subglobose, brown to dark brown cells. Conidiophores 

62–148 × 3–5 μm, 4–12 in divergent 


fascicles, 2–3-septate, emerging from stromata 

and through the plant cuticle, pale olivaceous 

brown or sometimes paler towards the apex, 

straight to slightly curved, unbranched, geniculate. 


holoblastic, polyblastic, sympodially proliferating. 

Conidiogenous loci 2.5–3.5 μm diam., 


very variable in length, 80–240 × 3–4 μm, 

solitary, acicular to long obclavate, straight to 

mildly curved, hyaline, 9–14-septate, smooth, 


toward a subacute apex, hila 2.5–3.5 μm diam., 

hila thickened and darkened. 



Wiang Ga Long, Bahn Tung Ruang Tong, 

Moo11, on leaves of Ipomoea nil (L.) Roth 

(Convolvulaceae), 8 September 2007, Jamjan 

Meeboon (BBH 23594); Chiang Mai Province, 

Mae Jo, Sansai, Farming area, on leaves of I. 

aquatica Forssk. (Convolvulaceae), 31 July 


Hosts – Apadanthera sagittifolia, Benincasa 

cerifera, B. hispida, Bryoania sp., 

Bryonopsis laciniata, Citrullus lanatus, C. 

vulgaris, Citrullus sp., Coccinia cordifolia, C. 

indica, Ctenolepsis cerasiformis, Ctenolepsis 

sp., Cucumis anguria, C. callosus, C. maxima, 

C. melo, C. sativa, Cucurbita foetidissima, C. 

maxima, C. moschata, C. pepo, C. perennis, C. 

sativus, Lagenaria leucantha, L. siceraria, L. 

vulgaris, Luffa acutangula, L. aegyptiaca, L. 

amara, L. cylindrica, L. vulgaris, Melothria 

pendula, Momordica charantia, M. cochinchinensis, 

M. cordifolia, M. dioica, M. foetida, M. 

schimperiana, Sechium edule, Sicana odorifera, 

Telfaria pedata, Trichosamthes anguina, T. 

japonica (Cucurbitatceae) (Crous & Braun 

2003). 

Distribution – Worldwide, where the host 

plants are cultivated or growing, including 

American Samoa, Argentina, Austria, Banggladesh, 

Barbados, Belize, Bolivia, Brazil, 

Brunei, Bulgaria, Cambodia, Canada, Chile, 

China, Cook Island, Costa Rica, Cuba, Czech 

Republic, Denmark, Dominican Republic, El 

Salvador, Ethiopia, Fiji, French Polynesia, 

Gabon, Georgia, Germany, Ghana, Great 

Britain, Greece, Guam, Hong Kong, India, 

Indonesia, Iran, Iraq, Ireland, Israel, Israel, 

Italy, Jamaica, Japan, Kenya, Korea, Laos, 

53

Fig. 36 – Line drawings of Cercospora citrullina on Ipomoea nil. a. Conidiophores and stromata. b. 

Conidia. Bars = 50 μm. (Meeboon 2009). 

Malawi, Malaysia, Mauritius, Mexico, 

Morocco, Myanmar, Nepal, Netherlands, New 

Caledonia, New Zealand, Nicaragua, Nigeria, 

Norway, Pakistan, Panama, Papua New Guinea, 

Peru, Philippines, Pitcairn Island, Poland, 

Puerto Rico, Romania, Russia, Samoa, Saudi 

Arabia, Solomon Islands, Somalia, South 

Africa, Sri Lanka, Sweden, Switzerland, Sudan, 

Taiwan, Tanzania, Thailand, Togo, Tonga, 

Trinidad and Tobago, Uganda, Ukraine, UK, 

USA, Vanuatu, Venezuela, Virgin Islands, 

Zambia, Zimbabwe (Crous & Braun 2003). 

Notes – Ipomoea nil is reported here as a 

new host of C. citrullina. Crous & Braun (2003) 

assigned this species to C. apii s. lat. 


(1976, p. 255). 

Cercospora ipomoeae G. Winter, Hedwigia 26: 

34 (1887). 

= Cercospora dichondrae Katsuki, Ann. 

Phytopathol. Soc. Japan 20: 72 (1955). 




54 



surrounded by a dark margin. Caespituli amphigenous. 

Stromata 24–40 μm diam., intraepidermal, 

well-developed, subglobose, brown 

to blackish brown. Conidiophores 13.5–134 × 

3–5 μm, in loose to dense fascicules, 2–3septate, 

numerous, arising from stromata, 


pale yellow to pale brown, sometimes branched, 



monoblastic or polyblastic, sympodially proliferating. 



44.5–143 × 3–3.5 μm, solitary, narrowly 

obclavate to subacicular, straight, hyaline, 6– 

15-septate, smooth, base obconically truncate, 

with subacute apex, hila 2–2.5 μm diam., 




leaves of Ipomoea aquatica Forssk. and I. nil 

(L.) Roth (Convolvulaceae), 9 March 2005, 

Jamjan Meeboon (CMU 28220 and CMU 

28221); Chiang Mai Province, A. Pa Pae, Bahn

Fig. 37 – Line drawings of Cercospora ipomoeae 

on Ipomoea obscura. a. Conidiophores 

and stromata. b. Conidia. Bars = 50 μm. 


Phadeng, Mushroom Research Centre, on 

leaves of Argyreia henryi Craib (Convolvulaceae), 

10 November 2006, Ikumitsu Araki 

(CMU 27911); Chiang Mai Province, Chiang 

Mai University, Faculty of Agriculture, on 

leaves of I. obscura (L.) Ker Gawl. (Convolvulaceae), 

21 August 2008, Jamjan Meeboon 

(BBH 23558). 

Hosts – Argyreia tiliaefolia, Convolvulus 

arvensis, Dichondra repens, Hewittia bicolor, 

Hewittia sp., Ipomoea acuminata, I. alba, I. 

aquatica, I. armata, I. asarifolia, I. batats, I. 

biloba, I. bonanox, I. cairica, I. carnea, I. 

clarensis, I. coccinea, I. cordofana, I. cymosa, I. 

eriocarpa, I. fistulosa, I. forsteri, I. hederacea, 

I. hildebrantii, I. indica, I. kentrocarpa, I. 

lacumosa, I. leari, I. longicuspis, I. nil, I. 

pandurata, I. pes-caprae, pestigridis, I. purpurea, 

I. quamoclit, I. ramonii, I. reptans, I. 

septaria, I. triloba,I. turpethum, I. villosa, 

Jacquemontia tammifolia, Merremia chrysoides, 

M. Emarginata, M. Umbellata, Operculina 

sp. (Convolvulaceae) (Crous & Braun 2003, 

Meeboon et al. 2007c). 



American Samoa, Antigua and Barbuda, 

Argentina, Australia, Barbados, Brazil, Brunei, 

China, Cook Islands, Costa Rica, Cuba, Fiji, 

Guam, Hong Kong, India, Indonesia, Italy, 

Ivory Coast, Jamaica, Japan, Kenya, Kiribati, 

Korea, Malaysia, Marshall Islands, Mauritius, 

Myanmar, New Caledonia, New Zealand, 

Pakistan, Panama, Papua New Guinea, Puerto 

Rico, Samoa, Sierra Leone, Solomon Island, 

Sudan, Venezuela, Taiwan, Tanzania, Thailand, 

USA, Vanuatu (Crous & Braun 2003, Meeboon 

et al. 2007c). 

Notes – The first report of C. ipomoeae 

on Argyreia henryi from Thailand was made by 

Meeboon et al. (2007c). Crous & Braun (2003) 

assigned this species to C. apii s.lat. 


Cercospora operculinae Mendoza, Phiipp. J. 

Sci. 75: 174 (1941). 

= Cercospora operculinicola Kamal, in 

herb. (IMI 367133). 



distinct, circular to subcircular, brown, with 

dark margin. Caespituli amphigenous. Stromata 

7–55 μm diam., substomatal, small to welldeveloped, 


subglobose, brown cells. Conidiophores 64– 

127.5 × 3–5 μm, 2–7 in loose fascicles, 2–8septate, 

arising from stromata, straight, unbranched, 

cylindrical, smooth, brown at the 

base, paler toward the apex, plainly geniculate. 

Conidiogenous cells integrated, holoblastic, 

polyblastic, rarely monoblastic, terminal, sympodially 



Conidia 22.5–96 × 3–3.5 μm, solitary, 

acicular, sometimes obclavate, straight, hyaline, 

6–9-septate, smooth, truncate at the base, 

tapering toward a subacute apex, hila 1–2.3 μm 



Chiang Mai Province, A. Mae Taeng, on leaves 

of Operculina turpethum (L.) Silva Manso 

(Convolvulaceae), 6 February 2008, Jamjan 

Meeboon and Iman Hidayat (BBH 23768). 

Hosts – Operculina bufalina, O. riedeliana, 

O. turpethum (Convolvulaceae) (Crous 

& Braun 2003). 

55

Fig. 38 – Line drawings of Cercospora operculinae 

on Operculina turpethum. a. Conidia. b. 

Conidiophores and stromata. Bars = 50 μm. 


Distribution – India, Papua New Guinea 

and Philippines (Crous & Braun 2003). 

Notes – This specimen is the first report 

of C. operculinae from Thailand. Crous & 

Braun (2003) assigned this species to C. apii 

s.lat. 


Cucurbitaceae 

Cercospora citrullina Cooke Grevillea 12: 31 

(1883). 

= Cercospora cucurbitae Ellis & Everh., 

J. Mycol. 4: 3 (1883). 

= Cercospora sechii J.A. Stev., Puerto 

Rico Agric. exp. Sta. Rep. 1917–1918: 137 

(1919). 

= Cercospora momordicae McRae, Ann. 

Cryptog. Exot. 2: 267 (1929). 

= Cercospora trichosanthis McRae, Ann. 

Cryptog. Exot. 2: 270 (1929). 

= Cercospora luffae Hara, Diseases of 

cultivated plants: 228 (1928). 

56 

= Cercospora chardoniana Chupp, 

Monogr. Univ. Puerto Rico, B, 2: 245 (1934). 

= Cercospora momordicae Mend., 

Philipp. J. Sci. 75: 173 (1941). (nom. illeg.), 

homonym of C. momordicae McRae (1929). 

= Cercospora momordicae Sawada, 


(1943), (nom. inval.), homonym of C. 

momordicae McRae 1929. 



irregular, greyish brown, with dark brown 

margin. Caespituli amphigenous. Stromata 14- 

30 μm diam., substomatal, small to welldeveloped, 


subglobose, brown to dark brown cells. 

Conidiophores 52–106.5 × 2.5–5 μm, 6-11 in 

loose fascicules, 2-5-septate, arising from 

stomata, straight, smooth, brown at the base, 

and paler toward the apex, unbranched, 

cylindrical, not geniculate. Conidiogenous cells 

integrated, holoblastic, monoblastic, terminal, 


1.5–2.5 μm diam., conspicuous, thickened and 

darkened. Conidia 63–296.5 × 2.5–4.5 μm, 

solitary, acicular, straight, hyaline, 8-26-septate, 

smooth, truncate at the base, tapering toward a 

subacute apex, hila 1.5–2 μm diam., 




leaves of Sechium edule (Jacq.) Sw. 

(Cucurbitaceae), 19 December 2005, Jamjan 

Meeboon (CMU 28215); the same host, Chiang 

Mai Province, Faculty of Chiang Mai 

University, Multiple Cropping Centre, 1 


same locality, Faculty of Agriculture, on leaves 

of Coccinia grandis (L.) Voigt (Cucurbitaceae), 

19 January 2005, Jamjan Meeboon (CMU 

27903); ibid, 12 June 2007, Jamjan Meeboon 

(BBH 23652); Chiang Rai Province, A. Wiang 

Pa Pao, T. Wiang Ga Long, Moo 11, Bahn 

Tung Ruang Tong, on leaves of Cucumis 

sativus L. (Cucurbitaceae), 30 July 2007, 

Jamjan Meeboon (BBH 23623); Chiang Mai 

Province, Hang Dong, Num Phrae, Farming 

area, on leaves of Lagenaria siceraria (Molina) 

Standl. (Cucurbitaceae), 7 August 2008, 


Province, Sansai, Mae Fag, on leaves of


citrullina on Citrullus vulgaris. a. Conidiophores 



Momordica charantia L. (Cucurbitaceae), 3 

August 2008, Jamjan Meeboon (BBH 23754).; 

Chiang Mai Province, Mae Jo University, 

Farming area, on leaves of Citrullus vulgaris 

Schrad. (Cucurbitaceae), 9 August 2008, 


Hosts – Apadanthera sagittifolia, Benincasa 

cerifera, B. hispida, Bryoania sp., 

Bryonopsis laciniata, Citrullus lanatus, C. 

vulgaris, Citrullus sp., Coccinia cordifolia, C. 

indica, Ctenolepsis cerasiformis, Ctenolepsis 

sp., Cucumis anguria, C. callosus, C. maxima, 

C. melo, C. sativa, Cucurbita foetidissima, C. 

maxima, C. moschata, C. pepo, C. perennis, C. 

sativus, Lagenaria leucantha, L. siceraria, L. 

vulgaris, Luffa acutangula, L. aegyptiaca, L. 

amara, L. cylindrica, L. vulgaris, Melothria 

pendula, Momordica charantia, M. cochinchinensis, 

M. cordifolia, M. dioica, M. foetida, 

M. schimperiana, Sechium edule, Sicana 

odorifera, Telfaria pedata, Trichosamthes 

anguina, T. japonica (Cucurbitatceae) (Crous 

& Braun 2003). 

Distribution – Worldwide, where the host 

plants are cultivated or growing, including 


American Samoa, Argentina, Austria, 

Bangladesh, Barbados, Belize, Bolivia, Brazil, 

Brunei, Bulgaria, Cambodia, Canada, Chile, 

China, Cook Islands, Costa Rica, Cuba, Czech 

Republic, Denmark, Dominican Republic, El 

Salvador, Ethiopia, Fiji, French Polynesia, 

Gabon, Georgia, Germany, Ghana, Great 

Britain, Greece, Guam, Hong Kong, India, 

Indonesia, Iran, Iraq, Ireland, Israel, Israel, 

Italy, Jamaica, Japan, Kenya, Korea, Laos, 

Malawi, Malaysia, Mauritius, Mexico, 

Morocco, Myanmar, Nepal, Netherlands, New 

Caledonia, New Zealand, Nicaragua, Nigeria, 

Norway, Pakistan, Panama, Papua New Guinea, 

Peru, Philippines, Pitcairn Island, Poland, 

Puerto Rico, Romania, Russia, Samoa, Saudi 

Arabia, Solomon Islands, Somalia, South 

Africa, Sri Lanka, Sweden, Switzerland, Sudan, 

Taiwan, Tanzania, Thailand, Togo, Tonga, 

Trinidad and Tobago, Uganda, Ukraine, UK, 

USA, Vanuatu, Venezuela, Virgin Islands, 


Notes – Sechium edule is reported here as 

a new host of C. citrullina. Cercospora 

citrullina was previously reported from 

Thailand by Petcharat & Kanjanamaneesathian 

(1989), and Meeboon et al. (2007b) on 

Coccinia grandis. Crous & Braun (2003) 

considered this species as C. apii s. lat. 


(1976, p. 255). 

Cercospora cocciniae Munjal, Lall & Chona, 

Indian Phytopathol. 12: 86 (1959). Fig. 40 


scattered to confluent, distinct, circular to 

subcircular, pale to pale brown, greyish brown 

to greyish white at the center, with distinct, 

yellowish brown to dark brown margins. 

Caespituli epiphyllous, rarely amphigenous. 

Stromata 18–29.5 μm diam., small, sometimes 

rudimentary to poorly developed, composed of 

a few sub-globular to irregular, brown to dark 

brown cells. Conidiophores 18–108.5 × 3–5.5 

μm, very variable in length, 4–15 in divergent 

fascicles, 1–9-septate, arising from stromata 

through the cuticle, pale olivaceous brown, 

sometimes paler at the apex, straight to slightly 

curved, unbranched, strongly geniculate. Coni- 

diogenous cells integrated, terminal, holoblastic, 

mostly polyblastic, sympodially proli- 

57


cocciniae on Coccinia grandis. a. Conidia. b. 



ferating. Conidiogenous loci 1.5–2.5 μm diam., 


41–102 × 2.5–5 μm, solitary, obclavatecylindric, 

straight to mildly curved, hyaline, 5– 

10-septate, very variable in length, smooth 


toward a subacute apex, hila 1.5–2.5 μm diam., 




Faculty of Agriculture, on leaves of Coccinia 

grandis (L.) Voigt (Cucurbitaceae), 29 

February 2008, Jamjan Meeboon (BBH 23564). 

Hosts – Coccinia indica, Momordica 

charantia (Cucurbitaceae) (Crous & Braun 

2003), Coccinia grandis (Meeboon 2009). 

Distribution – Brunei, India, Pakistan, 

Thailand (Crous & Braun 2003, Meeboon 

2009). 

Notes – This specimens is typical of C. 

cocciniae is having obclavate conidia. 

Cercospora cocciniae was first reported from 

Thailand by Meeboon (2009). 

58 

Euphorbiaceae 

Cercospora acalyphae Peck, Rep. (Annual) 

New York State Mus. Nat. Hist. 34: 48 (1881). 

= Cercospora acalypharum Tharp, 

Mycologia 9: 106 (1917). 

≡ Cercosporina acalypharum (Tharp) 

Sacc., Syll. Fung. 25: 902 (1931). Fig. 41 


circular or subcircular, symptoms at first 

pale greenish to ochraceous, later brown to 

dark brown, finally with grayish brown at the 

centre, surrounded by a dark margin. Caespituli 

amphigenous. Stromata 25–47 μm diam., 

intraepidermal, well-developed, composed of 

globose to subglobose, brown to blackish 

brown cells. Conidiophores 48.5–83.5 × 4–6 

μm, 2–8 in loose fascicules, 1–3-septate, 

arising from stomata, simple, straight, erect to 

decumbent, smooth, pale yellow to pale brown, 

sometimes branched, subcylindrical, slightly 


terminal, holoblastic, mostly monoblastic, 

sometimes polyblastic, sympodially proliferating. 



44–256 × 1.5–3 μm, solitary, narrowly obclavate 

to subacicular, straight, hyaline, 4–18septate, 

smooth, base obconically truncate, 

with subacute apex, hila 2–3 μm diam., 



Phetchabun Province, Nam Nao National Park, 

on leaves of Acalypha wilkesiana Mull. Arg. 

(Euphorbiaceae), 24 November 2004, Chiharu 


27898); Chiang Mai Province, A. Mae Jo, 

Sansai, Farming area, on leaves of A. 

wilkesiana, 31 July 2008, Jamjan Meeboon 

(BBH 23694). 

Hosts – Acalypha alopecurioides, A. 

australis, A. ciliata, A. gracilens, A. grandis, A. 

hispida, A. indica, A. marginata, A. marvorata, 

A. ostryifolia, A. rhomboidea, A. virginica, A. 

wilkesiana, Acalypha sp. (Euphorbiaceae) 

(Crous & Braun 2003, Meeboon et al. 2007c). 

Distribution – Brazil, China, Cuba, India, 

Jamaica, Japan, Solomon Islands, Thailand, 

USA, Venezuela (Crous & Braun 2003, 

Meeboon et al. 2007c).


Fig. 41 – Line drawings of Cercospora acalyphae on Acalypha wilkesiana. a. Conidiophores and 


Fig. 42 – Line drawings of Cercospora codiaei on Codiaeum variegatum. a. Conidiophores and 

stromata. b. Conidia. Bars = 50 μm. (Meeboon 2009) 

Notes – Meeboon et al. (2007c) were the 

first to report this species from Thailand. 

Literature – Chupp (1954, p. 200-201). 

Cercospora codiaei Gonz. Frag. & Cif., Boln 

de la Real Soc. Españ. Hist. Nat., Madrid 26: 

199 (1926). 



subcircular, solitary, pale brown, with reddish 

brown margin. Caespituli amphigenous. Stromata 

38–44 μm diam., small, substomatal, 

composed of a few globose, dark brown cells. 


loose fascicles, 4–7-septate, arise through 

stromata, straight, smooth, brown at the base, 

paler toward the apex, unbranched, cylindrical, 

slightly geniculate. Conidiogenous cells integrated, 

holoblastic, polyblastic, sometimes 

monoblastic, terminal, sympodially proliferating. 


59

Fig. 43 – Line drawings of Cercospora phyllanthicola on Phyllanthus acidus. a. Conidia. b. 

Conidiophores and stromata. Bars = 50 μm. (Meeboon 2009). 


29–160 × 3–4.5 μm, solitary, acicular, rarely 

obclavate, straight, hyaline, 3–10-septate, 

smooth, truncate at the base, with tapering 

toward a subacute apex, hila 1.5–2 μm diam., 



Chiang Mai Province, T. Suthep, A. Muang, 

Suthep-Pui National Park, Medicinal Plant 

Garden, on leaves of Codiaeum variegatum (L.) 

A. Juss. (Euphorbiaceae), 25 July 2008, 


23666). 

Hosts – Codiaeum sp., Codiaeum 

variegatum (Euphorbiaceae) (Crous & Braun 

2003, Meeboon 2009). 

Distribution – Cuba, Dominican Republic, 

Nigeria, Thailand (Crous & Braun 2003, 


Notes – Cercospora codiaei was first 


Literature – Chupp (1954, p. 200–215). 

Cercospora phyllanthicola S.A. Khan & 

Kamal, Indian Phytopathol. 15: 296 (1962) 

[1963]. 


60 


irregular, brown to dark brown, pale at the 

center, with dark margin, and limited by vein 

of the leaf. Caespituli amphigenous. Stromata 

15–32 μm diam., small to well-developed, 

substomatal to intraepidermal, composed of a 

few globose to subglobose, dark brown cells. 


to dense fascicles, 1–3-septate, arising from 

stomata, straight to decumbent, smooth, brown 

at the base, paler toward the apex, cylindrical, 

sometime constrict at the septate, rough wall, 

unbranched, geniculate, mostly near the apex. 


intercalary, holoblastic, polyblastic, sympodially 



Conidia 51–133 × 3–4.5 μm, solitary, 

obclavate to acicular, straight, slightly curved, 

hyaline, 5–11-septate, smooth, obconically 

truncate at the base, tapering toward a subacute 

apex, hila 1–2.3 μm diam., thickened and 

darkened. 


Uttradit Province, A. Muang, Sak Yai National 

Park, on leaves of Phyllanthus sp. (Euphorbiaceae), 

25 November 2004, Chiharu Nakashima 

and Jamjan Meeboon (CMU 27876); Chiang 

Rai Province, Wiang Pa Pao, on leaves of 

Phyllanthus acidus (L.) Skeels, 15 February 

2008, Jamjan Meeboon (BBH 23670).

Hosts – Phyllanthus niruri, Sauropus 

androgynus (Euphorbiaceae) (Crous & Braun 

2003, Meeboon et al. 2007c). 

Distribution – Brunei, India, Malaysia, 

Myanmar, Pakistan, Singapore, Thailand 


Notes – Three species of Cercospora s. 

str., viz, C. kirganeliicola R.K. Srivast., S. 

Narayan & A.K. Srivast. (C. apii s. lat.), C. 

phyllanthicola (C. apii s. lat.), and C. tarii 

Deighton have been recorded associated with 

plant genus Phyllanthus. This specimen is very 

close to C. phyllanthicola in having simple 

conidiophores and acicular conidia. The first 

report of C. phyllanthicola from Thailand was 

by Meeboon et al. (2007c). Crous & Braun 

(2003) assigned this species to C. apii s.lat. 

Fabaceae 

Cercospora canescens Ellis & G. Martin, 

Amer. Naturalist 16: 1003 (1882). 

≡ Cercosporiopsis canescens (Ellis & G. 

Martin) Miura, Flora of Manchuria and East 

Mongolia 3: 529 (1928). 

= Cercospora vignicaulis Tehon, 

Mycologia 29: 436 (1937). 




irregular, brown to dark brown, limited 

by vein of the leaf. Caespituli amphigenous. 

Stromata 26.5–67 μm diam., well-developed, 

intraepidermal, and composed of globose to 


Conidiophores 60.5–118 × 3–5 μm, 12–20 in 

loose to dense fascicles, 1–3-septate, straight to 

decumbent, cylindrical, smooth, brown at the 

base, and paler toward the apex, unbranched, 

geniculate to sinuous. Conidiogenous cells 12– 

35.5 × 3–5 μm, integrated, terminal, holoblastic, 

polyblastic, sometimes monoblastic, sympodially 


μm diam., conspicuous, thickened and 

darkened. Conidia 56–113.5 × 3–4.5 μm, 

solitary, narrowly obclavate to subacicular, 

straight, hyaline, 3–9-septate, smooth, obconically 

truncate at the base, tapering toward a 

subacute apex, hila 2–3 μm diam., thickened 




Wiang Ga Long, Moo11, Bahn Tung Ruang 

Fig. 44 – Line drawings of Cercospora canescens on Dolichos lablab. a. Conidiophores and stroma. 

b. Conidia. Bars = 50 μm. (Meeboon 2009). 

61

Tong, on leaves of Vigna radiata (L.) R. 

Wilczek, 25 October 2005, Jamjan Meeboon 

(CMU 27888); the same locality, on leaves of 

V. unguiculata var. sesquipedalis (Fabaceae), 

18 November 2005, Jamjan Meeboon (CMU 

27894); same locality, on leaves of Lablab 

purpureus (L.) Sweet (Fabaceae), 16 July 2007, 


Province, San Sai, Mae Fag, 3 August 2008, 

Jamjan Meeboon (BBH 23749); on leaves of 

Psophocarpus tetragonolobus DC. (Fabaceae), 

31 July 2007, Jamjan Meeboon (JM 104); on 

leaves of Dolichos lablab L. (Fabaceae), 25 

July 2008, Jamjan Meeboon (BBH 23773); 


Royal Project, on leaves of V. unguiculata (L.) 

Walp. (Fabaceae), 27 September 2007, Jamjan 

Meeboon and Iman Hidayat (BBH 23678); 


Multiple Cropping Centre, on leaves of V. 

unguiculata, 1 August 2008, Jamjan Meeboon 

(BBH 23722). 

Hosts – Amaranthus sp., Celosia 

argentea (Amaranthaceae), Annona odorata, A. 

squarrosa (Annonaceae), Rauvolfia serpentina 

(Apocynaceae), Verschaffeltia splendida (Arecaceae), 

Aster novibelgii (Asteraceae), Bixa 

orellana (Bixaceae), Raphanus sativus (Brassicaceae), 

Rhynchosia aurea, R. minima, Ricinus 

communis (Euphorbiaceae), Arachis hagenbeckii, 

A. hypogaea, Alysicarpus sp., Bauhinia 

alba, B. variegata, Cajanus cajan, Calopogonium 

mucunoides, Canavalia ensiformis, C. 

gladiata, C. maritima, Cassia alata, C. lathyroides, 

Cassia sp., Centrosema acutifolium, C. 

arenarium, C. brasilianum, C. macrocarpum, 

C. plumieri, C. pubescens, C. virginianum, 

Clitoria ternatea, Codariocalyx gyroides, Crotalaria 

juncea, C. mucronata, C. mysorensis, C. 

retusa, C. spectabilis, C. usaramoensis, C. 

verrucosa, C. zanzibarica, Crotalaria spp., 

Cyamopsis psoralioides, Desmodium canum, D. 

gyrans, D. gyroides, D. incanum, D. intortum, 

D. lycioides subspecies guerkei, D. repandum, 

D. turtuosum, D. uncinatum, Dolichos biflorus, 

D. daltonii, D. lablab, D. lignosus, D. trilobus, 

D. turtuosum, D. uniflorus, Erythrina addisoniae, 

E. suberosa, E. subumbrans, E. variegata, 

Flemingia macrophylla, Gliricidia sepium, 

Glycine max, G. soja, G. ussuriensis, G. wightii, 

Heylandia latebrosa, Indigofera astragalina, 

62 

Kotschya sp., Lablab niger, L. purpureus, 

Lespedeza sp., Lathyrus odoratus, Leucaena 

leucocephala, Lotononis bainesii, Lupinus sp., 

Macroptilium atropurpureum, M. lathyroides, 

M. daltonii, M. uniflorum, Medicago sativa, 

Mimosa invisa, Mucuna pruriens, Neonotonia 

wightii, Phaseolus aconitifolius, P. angularis, 

P. atropurpureus, P. aureus, P. calcaratus, P. 

lathyroides, P. limensis, P. lunatus, P. minimus, 

P. panduratus, P. radiatus, P. trilobus, P. 

vulgaris, Pistia stratiotes, Pisum sativum, 

Psophocarpus tetragonolobus, Psoralea bituminosa, 

P. drupacea, Pterocarpus marsupia, 

Pueraria hirsuta, P. lobata, P. phaseoloides, P. 

trilobam, Quercus sp., Senna alata, S. tora, 

Shuteria involucrata, Stylosanthes guianensis, 

S. humilis, Vicia ungiculata, Vigna angularis, 

V. catjang, V. luteola, V. marina, V. mungo, V. 

parkeri, V. radiata, V. repens, V. reticulata, V. 

sesquipedalis, V. sinensis, V. umbellata, V. 

vexillata, Vitis vinivera, Voandzeia subterranea 

(Fabaceae), Coleus sp., Ocimum basilicum, 

Plectranthus sp. (Lamiaceae), Tetramnus 

labialis, T. uncinatus (Malphigiaceae), Artocarpus 

integrifolia (Moraceae), Boerhavia 

erecta, Commicarpus sp. (Nyctaginaceae), 

Lycopersicon esculentum, Solanum laciniatum 

(Solanaceae) (Crous & Braun 2003). 

Distribution – Worldwide, wherever the 

crop is cultivated, including Australia, 

Bangladesh, Barbados, Brazil, Bolivia, Brunei, 

Cambodia, China, Colombia, Costa Rica, Cuba, 

Dominican Republic, Ecuador, Fiji, Georgia, 

Ghana, Guyana, Haiti, Hong Kong, India, 

Indonesia, Iran, Japan, Kenya, Korea, Malawi, 

Malaysia, Malawi, Mauritius, Myanmar, Nepal, 

New Caledonia, New Zealand, Nigeria, 

Pakistan, Panama, Papua New Guinea, Peru, 

Philippines, Puerto Rico, Russia, Senegal, 

Sierra Leone, Solomon Islands, Somalia, South 

Africa, Saint Vincent and the Grenadines, 

Sudan, Tadzhikistan, Taiwan, Tanzania, 

Trinidad and Tobago, Togo, Uganda, USA, 

Uzbekistan, Vanuatu, Venezuela, Virgin 

Islands, Zambia, Zimbabwe (Crous & Braun 

2003). 


from Thailand by Sontirat et al. (1980) on V. 

radiata. Crous & Braun (2003) assigned this 

species to C. apii s. lat. 


Meeboon (2009).


crotalariae on Crotalaria montana. a. Conidia. 

b. Conidiophores and stroma. Bars: a = 40 μm, 

b = 50 μm. (Meeboon 2009). 

Cercospora crotalariae Sacc., Syll. Fung. 22: 

129 (1913). 

= Cercospora crotalariae-junceae 

Sawada, J. Taihoku Soc. Agric. 7: 27 (1942) 

(nom. inval.). 



scattered to confluent, subcircular to angular, 

pale brown at the young symptoms, later 

becoming greyish brown, greyish to pale at the 

centre, with reddish brown or purplish brown 

margins. Caespituli amphigenous. Stromata 

25–30.75 μm diam., intraepidermal, small to 

well-developed, composed of globose to 


Conidiophores 35–231 × 3.5–5.5 μm, numerous 

in dense fascicules, 2–6-septate, arising 

from stromata, simple, straight, unbranched, 

erect to decumbent, smooth, pale yellow to 

pale brown, subcylindrical, geniculate to 

sinuous. Conidiogenous cells integrated, termi- 


nal, holoblastic, mostly polyblastic, sympodially 

proliferating. Conidiogenous loci 2.5–3.5 

μm diam., conspicuous, thickened, and darkened. 

Conidia 39–206 × 2–4 μm, solitary, 



truncate, with subacute apex, hila 2.5–3.5 



Chiang Mai Province, Mae Jo University, 

Farming area, on leaves of Crotalaria montana 

Heyne ex Roth (Fabaceae), 9 August 2008, 


Hosts – Crotalaria incana, C. juncea, C. 

montana, C. mucronata, C. retusa, C. sericeum, 

C. spectabilis, C. striata, C. stricta, C. Usaramoensis, 

Crotalaria sp. (Fabaceae) (Crous & 


Distribution – Bangladesh, China, Cuba, 

Ethiopia, India, Indonesia, Pakistan, Papua 

New Guinea, Puerto Rico, Sri Lanka, Taiwan, 

Thailand, Venezuela (Crous & Braun 2003, 


Notes – Cercospora crotalariae was first 

recorded from Thailand by Meeboon (2009). 

Crous & Braun (2003) assigned this species as 

C. apii s. lat. 

Cercospora kikuchii T. Matsumoto & Tomoy., 

Ann. Phytopathol. Soc. Japan 1: 1 (1925). 

≡ Cercospora kikuchii T. Matsumoto & 

Tomoy., l.c.: 10. 

≡ Cercospora kikuchii (T. Matsumoto & 

Tomoy.) M.W. Gardner, Proc. Indian Acad. Sci. 

36: 12 (1927) (comb. superfl.). 




initially appearing pale brown, later becoming 

tan to dingy grey, greyish white at the centre, 

with reddish brown or purplish brown margins. 


diam., intraepidermal, well-developed, composed 

of globose to subglobose, brown to 


3.5–5 μm, 9 to numerous in dense and 

divergent fascicules, 2–4-septate, arising from 

stromata, simple, straight, erect to decumbent, 

smooth, pale yellow to pale brown, sometimes 


Conidiogenous cells integrated, terminal, holo- 

63

Fig. 46 Line drawings of Cercospora kikuchii on Glycine max. a. Conidia. b. Conidiophores and 

stromata. Bars = 50 μm. (Meeboon 2009). 

blastic, monoblastic or polyblastic, sympodially 






truncate, with subacute apex, hila 2–2.5 




on leaves of Glycine max Merr. (Fabaceae), 21 


27878); Chiang Mai Province, Mae Jo 

University, Farming area, on same host, 9 

August 2008, Jamjan Meeboon (BBH 23707). 

Hosts – Cassia obtusifolia, Cyamopsis 

tetragonoloba, Dolichos biflorus, Glycine 

hispida, G. max, G. soja, G. tabacina, G. 

ussuriensis, Phaseolus aureus, P. lunatus, P. 

mungo, Senna sp., Vigna prainiana (Fabaceae) 


Distribution – Worldwide where the host 

is cultivated, including Argentina, Bangladesh, 

Bolivia, Brazil Brunei, Burkina Faso, 

64 

Cameroon, Canada, China, Colombia, Cuba, 

Egypt, Ethiopia, Fiji, France, Gabon, Ghana, 

Guinea, India, Indonesia, Iran, Jamaica, Japan, 

Korea, Liberia, Malaysia, Mexico, 

Mozambique, Nepal, New Caledonia, Nigeria, 

Pakistan, Panama, Papua New Guinea, Peru, 

Puerto Rico, Russia, Sierra Leone, Somalia, 

South Africa, Sri Lanka, Taiwan, Tanzania, 

Thailand, Togo, Trinidad and Tobago, Uganda, 


Note – The first record of this species 

from Thailand was by Sontirat et al. (1980). 

Crous & Braun (2003) assigned this species to 

C. apii s.lat. 


Hydrangeaceae 

Cercospora hydrangeae Ellis & Everh., J. 

Elisha Mitch. Sci. Soc. 8: 52 (1892). 

= Cercosporina hydrangeicola Speg., 

Anales Mus. Nac. Buenos Aires 20: 426 (1910). 

≡ Cercospora hydrangeicola (Speg.) 

Vassiljevsky, in Vassiljevsky & Karakulin,

Fungi imperfecti parasitici 1. Hyphomycetes: 

339 (1937). 

= Cercospora hydrangeana Tharp, 

Mycologia 9: 110 (1917). 

≡ Cercosporina hydrangeana (Tharp) 

Sacc., Syll. Fung. 25: 915 (1931). 

= Cercospora arborscentis Tehon & E. 

Daniels, Mycologia 17: 246 (1925). 



irregular, pale olivaceous to light 

brown, centre greyish brown to greyish white 

with purplish brown to dark brown margins. 

Caespituli amphigenous. Stromata lacking. 

Conidiophores 111–227 × 3.5–5 μm, 3–11 in 

loose fascicles, 1–3-septate, straight to decumbent, 

unbranched, smooth, very dark brown at 

the base, and paler toward the apex, cylindrical, 

strongly geniculate. Conidiogenous cells integrated, 

holoblastic, polyblastic, sometimes 

monoblastic when young, sympodially proliferating. 



113–278 × 2.5–3.5 μm, acicular, hyaline, 13– 

27-septate, solitary, slightly curved at the apex, 

base obconically truncate, hila 1.5–2.5 μm 



Chiang Mai Province, Suthep-Pui National 

Park, on leaves of Hydrangea macrophylla 

(Thunb.) Ser. (Hydrangeaceae), 21 November 

2004, Jamjan Meeboon (CMU 27921); Chiang 

Mai Province, Sanpatong District, Mae Wang 

Sub-district, T. Mae Win, Bahn Mae Sapok, 

Mae Sapok Royal Project, on same host, 8 



Hosts – Hydrangea angustisepala, H. 

arborescens, H. hortensis, H. macrophylla, H. 

opuloides, H. paniculata, H. serrata, Hydrangea 

sp. (Hydrangeaceae) (Crous & Braun 

2003). 

Distribution – Argentina, Borneo, Brazil, 

Brunei, China, Hong Kong, India, Iran, Japan, 

Korea, Malawi, Malaysia, Myanmar, Nigeria, 

Philippines, Puerto Rico, Romania, Russia 

(Asian part), Sabah, Sierra, Leone, Singapore, 

Taiwan, Thailand, USA, Virgin Islands, 

Zimbabwe (Crous & Braun 2003). 

Notes – The first record of this species 

from Thailand was by Petcharat & 



hydrangeae on Hydrangea macrophylla. a. 

Conidiophores and stromata. b. Conidia. Bars 

= 50 μm. (Meeboon 2009). 

Kanjanamaneesathian (1989), but they did not 

give any detailed morphological description 

and illustration for this fungus. Braun (2000) 

assigned this species to C. apii s.lat. 


Lamiaceae 

Cercospora kabatiana Allesch. ex Lindau, 

Rabenh. Krypt.- Fl. ed. 2, 9: 130 (1910). 

≡ Cercospora kabatiana (Allesch. ex 

Lindau) Moescz, Magyar Biol. Kutatόint. 

Munkái 3: 115 (1930). Fig.48 



greenish to ochraceous, later become dull 

brown, finally with pale to greyish white at the 

centre, surrounded by a dark margin. Caespituli 

amphigenous. Stromata lacking. Conidiophores 

78–185 × 3–5 μm, numerous in loose 

fascicules, 2–5-septate, arising from stromata, 


pale yellow to pale brown, rarely branched, 

subcylindrical, geniculate. Conidiogenous cells 

65

integrated, terminal, holoblastic, polyblastic, 



darkened. Conidia 95–144 × 2.5–4 μm, solitary, 


hyaline, 7–12-septate, smooth, base truncate, 

with subacute apex, hila 2–2.5 μm diam., 




on leaves of Solenostemon scutellarioides (L.) 

Codd (Lamiaceae), 1 August 2008, Jamjan 


Hosts – Lamium amplexicaule, L. 

galeobdolon, L. maculatum, L.montanum, L. 

nepetaefolia, Leonotis sp., Solenostemon 

scutellarioides (Lamiaceae) (Crous & Braun 

2003, Meeboon 2009). 

Distribution – Armenia, Austria, Czech 

Republic, Lesotho, USA, Thailand (Crous & 


Notes – This specimen is a typical of C. 

apii s. lat. by having slight or lacking stromata, 

long conidiophores, and hyaline and long 

acicular conidia, with truncate base. 

Cercospora kabatiana was assigned to C. apii s. 

lat. by Crous & Braun (2003). This species was 

first reported from Thailand by Meeboon 

(2009). 


Cercospora physostegiae Jenkins, Phytopathology 

35: 329 (1945) Fig. 49 


solitary, sometimes clustered to form larger 

spots, circular to subcircular, sometimes irregular, 

greyish brown, with dark brown margin, 

limited by leaf veins. Caespituli amphigenous. 

Stromata 14–30 μm diam., small to welldeveloped, 

substomatal, composed of a few 

globose to subglobose, brown-walled cells. 


fascicles, 1–3-septate, arising through stromata, 

straight, smooth, brown at the base, and paler 

toward the apex, unbranched, cylindrical, often 

not geniculate, very rarely geniculate. 

Conidiogenous cells integrated, terminal, holoblastic, 

mostly monoblastic, sympodially proliferating. 



20– 129 × 2– 4 μm, solitary, obclavate-filiform 

66 

to acicular, straight, hyaline, 12–19-septate, 

smooth, truncate at the base, tapering toward a 

subacute apex, hila 1.5–2 μm diam., conspicuous, 

thickened, and darkened. 



of Clerodendrum paniculatum L. (Lamiaceae), 

13 February 2008, Jamjan Meeboon and Iman 


Hosts – Clerodendrum paniculatum, 

Physostegia virginiana (Lamiaceae) (Chupp 

1954, Meeboon 2009). 

Distribution – Thailand, USA (Meeboon 

2009, Chupp 1954). 

Notes – Four species of Cercospora s. str. 

non C. apii s. lat. have been reported from the 

plant family Lamiaceae, viz, C. isanthi Ellis & 

Kellerm., C. physostegiae, C. scorodoniae 

Unamuno, and C. teucrii Ellis & Kellerm. All 

these species are characterized by relatively 

short conidiophores and amphigenous caespituli 

(Chupp 1954). This specimen is similar to 

C. physostegiae due to the conidiophores often 

not geniculate and obclavate conidia with an 

obconically truncate base. The other three 

species are characterized by acicular conidia 

and geniculate conidiophores. Cercospora physostegiae 

was firstly reported from Thailand by 



Cercospora volkameriae Speg., Revista del 

Museo de La Plata 15: 47 (1908). 



circular or subcircular, limited by vein, often 

greyish at the centre, brown with a dark reddish 

margin. Caespituli epiphyllous. Stromata 12– 

32 μm diam., intraepidermal, small to welldeveloped, 

composed of globose to subglobose, 


36–127.5 × 2.5–4 μm, 8–10 in loose and 

divergent fascicules, 2–4-septate, arising from 

stomata, erect to decumbent, smooth, pale 

yellow to pale brown, straight, rarely branched, 


cells integrated, terminal to intercalary, 

holoblastic, polyblastic, sympodially proliferating. 

Conidiogenous loci 2.5–3 μm diam., 


40–87 × 2–3 μm, solitary, narrowly obclavate


Fig. 48 – Line drawings of Cercospora kabatiana on Solenostemon scutellarioides. a. Apical part 

of conidiophores. b. Conidia. Bars = 50 μm. (Meeboon 2009). 

Fig. 49 – Line drawings of Cercospora physostegiae on Clerodendrum paniculatum. a. Conidia. b. 

Conidiophores and stromata. Bars: a, b = 50 μm. (Meeboon 2009). 

to subacicular, 3–10-septate, straight, hyaline, 

smooth, base obconically truncate, with subacute 


darkened. 



of Clerodendrum fragrans Willd. (Lamiaceae), 

27 July 2008, Jamjan Meeboon (BBH 23763). 

Hosts – Clerodendrum cordifolium, C. 

fragrans, C. indicum, C. infortunatum, C. 

paniculatum, C. scandens, C. schweinfurthii, C. 

siphonatus, C. speciosissimum, C. speciosum, 

67

C. splendens, C. thomsonae, C. trichotomum, C. 

volubile, Gmelina arborea (Lamiaceae) (Crous 

& Braun 2003, Meeboon et al. 2007c). 

Distribution – Barbados, Brazil, Brunei, 

Cuba, Ghana, Guinea, India, Indonesia, 

Jamaica, Korea, Malawi, Malaysia, Nepal, 

Nigeria, Sierra Leone, Singapore, Sudan, 

Taiwan, Tanzania, Thailand, Togo (Crous & 


Notes – The morphological characteristicss 

of this specimen are close to C. volkameriae. 

The first report of C. volkameriae from 

Thailand was by Meeboon et al. (2007c). Crous 

& Braun (2003) considered this species as C. 

apii s.lat. 


Malvaceae 

Cercospora althaeina Sacc. Michelia 1: 269 

(1878). 

= Cercospora kellermanii Bubák, J. 

Mycol. 9: 3 (1903). 

68 

= Cercospora ramularia Siemaszko, Izv. 

Kavkazsk. Muz. 12: 28 (1919) and Arch. Nauk 

Biol. Towarz. Nauk. Warszawsk. 1: 49 (1923). 

≡ Cercopsorina ramularia (Siemaszko) 

Sacc., Syll. Fung. 25: 910 (1931). 

= Cercospora altheina var. praecincta 

Davis, Trans. Wisconsin. Acad. Sci. 18: 260 

(1915). 

≡ Cercospora praecincta (Davis) Chupp, 

Monograph of Cercospora: 376 (1954). 

= Cercospora althaeina var. althaeaeofficinalis 

Săvul. & Sandu, Hedwigia 73: 127 

(1933). 

= Cercospora althaeicola J.M. Yen & 

S.K. Sun, Cryptog. Mycol. 4: 189 (1983). 

Fig. 51 


dark to yellowish, only leaf decoloration. 


diam., substomatal, small, composed of a few 


brown 

Fig. 50 – Line drawings of Cercospora volkameriae on Clerodendrum fragrans. a. Conidiophores 

and stromata. b. Conidia. Bars = 50 μm. (Meeboon 2009).


Fig. 51 – Line drawings of Cercospora althaeina on Alcea rosea. a. Conidiophores and stromata. b. 


cells. Conidiophores 41–186 × 3–5 μm, up to 8 

in dense fascicles, 3–7-septate, arising from 

stomata, straight, sometime slightly constricted 

at the septate, smooth, brown at the base, paler 

toward the apex, cylindrical, unbranched, not 


holoblastic, often monoblastic, sometimes 

polyblastic and sympodially proliferating. 

Conidiogenous loci 2–3 μm diam., conspicuous, 


μm, solitary, acicular, straight, hyaline, 14–25septate, 

smooth, truncate at the base, tapering 




Chiang Rai Province, Mae Fah Luang, a. Mae 

Jan, Doi Tung Development, on leaves of 

Alcea rosea L. (Malvaceae), 16 August 2008, 


Hosts – Abutilon avicennae, A. incanum, 

A. theophrastii, Alcea flavovirens, A. froloviana, 

A. kusariensis, A. litwinowii, A. nudiflora, 

A. pallida, A. rosea, A. rugosa, A. tabrisiana, 

Alcea sp., Althaea ficifolia, A. hirsuta, 

A. officinalis, A. rosea, Callirhoë involucrata, 

C. triangulata, Gossypium hirsutum, Hibiscus 

trionum, Hibiscus sp., Kydia calycina, Kydia 

sp., Lavatera thuringiaca, Malva neglecta, M. 

pusilla, M. rotundifolia, Malva sp., Modiola 

caroliniana, Napaea dioica (Malvaceae) 

(Crous & Braun, 2003). 

Distribution – Worldwide, Argentina, 

Armenia, Australia, Azerbaijan, Bangladesh, 

Brazil, Bulgaria, Canada, China, Cuba, Georgia, 

Germany, Guatemala, India, Iran, Italy, 

Jamaica, Japan, Kazakhstan, Kenya, Kirghizia, 

Korea, Lithuania, Malawi, Malaysia, Mauritius, 

Moldova, Myanmar, New Zealand, Pakistan, 

Romania, Russia (European part), Tadzhikistan, 

Taiwan, Thailand, Ukraine, USA, Zambia, 

Zimbabwe (Crous & Braun 2003, Meeboon 

2009). 

Notes – Cercospora althaeina was first 


Literature – Chupp (1954, p. 369, 376). 

Moraceae 

Cercospora ficina Tharp, Mycologia 9: 109 

(1917). 

≡ Cercospora ficina (Tharp) Sacc., Syll. 

Fung. 25: 911 (1931). Fig. 52 





surrounded by a dark margin or brown halo. 


diam., intraepidermal, well-developed, subglobose, 

brown to blackish brown. Conidiophores 

42–229 × 3–6 μm, numerous, in loose to 

69

Fig. 52 – Line drawings of Cercospora ficina 

on Ficus religiosa. a. Conidia. b. Conidiophores 

and stroma. Bars = 50 μm. (Meeboon 

2009). 

densely fascicules, arising from stomata, branched, 

subcylindrical, 2–9-septate, geniculate to 

sinuous, erect to decumbent, smooth, pale 

yellow to pale brown. Conidiogenous cells 

integrated, terminal, monoblastic to polyblastic, 


2–2.5 μm diam, conspicuous, thickened and 

darkened. Conidia 42.5–161 × 2–4.5 μm, 

solitary, narrowly obclavate to subacicular, 

straight, hyaline, 7–14-septate, smooth, apex 

subacute, base obconically truncate, hilum 1.5– 

2.5 μm diam., thickened and darkened. 



Faculty of Agriculture, on leaves of Ficus 

religiosa L. (Moraceae), 18 August 2008, 


Hosts – Ficus carica, F. hispida, F. 

religiosa, F. uliginusa, F. urceolaria, Streblus 

asper (Moraceae) (Crous & Braun 2003). 

Distribution – India, Indonesia, Nigeria, 

Pakistan, Sudan, Uganda, USA (Crous & 

Braun 2003). 

70 

Notes – The first report of C. ficina from 

Thailand was by Meeboon (2009). 

Cercospora elasticae A. Zimm., Bull. Inst. Bot. 

Buitenzorg 10: 17 (1901). 


Leaf spots 5–8 mm diam., distinct, amphigenous, 

scattered, circular or subcircular to 

angular, sometimes forming large symptoms, 

up to 30 mm diam., greyish brown, with dark 

margins. Caespituli epiphyllous. Stromata 18- 

24 μm diam., intraepidermal, small, composed 

of globose to subglobose, brown to blackish 

brown cells. Conidiophores 63–139 × 3–4 μm, 

5–8 in loose and divergent fascicules, 2–4septate, 

arising from stromata, erect to decumbent, 

smooth, pale yellow to pale brown, 

unbranched, subcylindrical, geniculate to sinuous. 


holoblastic, monoblastic to polyblastic, sympodially 



Conidia 120–160 × 3 μm, solitary, acicular, 

8–13-septate, hyaline, smooth, truncate at the 

base, with acute to subacute apex, hila 2–2.5 



Chiang Mai Province, Pang Da Royal project, 

on leaves of Ficus carica L. (Moraceae), 5 

August 2008, Jamjan Meeboon (BBH 23728). 

Hosts – Ficus carica, F. elastica 

(Moraceae) (Crous & Braun 2003). 

Distribution – India, Indonesia, USA, 

Venezuela (Crous & Braun 2003). 

Notes – The first report of C. elasticae 



Nyctaginaceae 

Cercospora neobougainvilleae Meeboon, 

Hidayat & C. Nakash., Sydowia 60: 254 (2008). 

Fig. 54) 


orbicular, center pale brown, with dark brown 

margin. Caespituli epiphyllous. Stromata 11.5– 

71.5 μm diam., intraepidermal, well-developed, 

composed of globose to subglobose, dark 

brown cells. Conidiophores 13.5–165 × 1–9 

μm, 4–20 in loose to dense fascicules, 1–3-


elasticae on Ficus carica. a. Conidiophores 




septate, arising from stromata, narrower toward 

the apex, unbranched, geniculate 1–2 times 

near the apex, thin-walled to slightly thickened, 

smooth, brown at the base, paler towards the 

apex. Conidiogenous cells 2–30 × 1–9, 

integrated, terminal, sympodial proliferation. 



μm, solitary, obclavate, straight to mildly curve, 

truncate to obconically truncate at base, acute 

to subobtuse at the apex, 4–5-septate, hyaline, 

thin-walled, smooth, hila 1–4 μm diam., 




on leaves of Bougainvillea spectabilis Willd. 

(Nyctaginaceae), 30 November 2005, Jamjan 

Meeboon (CMU 27930: Holotype); Chiang 

Mai Province, A. Muang, RAMA IX Garden, 

on leaves of B. spectabilis, 26 August 2008, 


Hosts – Bougainvillea spectabilis 

(Nyctaginaceae) (Meeboon et al. 2008). 


(Meeboon et al. 2008). 

Fig. 54 – Line drawings of Cercospora neobougainvilleae on Bougainvillea spectabilis (from 

holotype). a. Conidiophores and stroma. b. Conidia. Bars = 20 μm. (Meeboon et al. 2008). 

Notes – Four species of Cercospora are 

hitherto known associated with Nyctaginaceae, 

viz, Cercospora canescens Ellis & G. Martin, 

C. furfurella Speg., C. mirabilis Tharp & C. 

salpianthi Chupp & A.S. Mull. (Crous & Braun 

2003). Two species, C. canescens and 

71

C. salpianthi belong to the species complex 

C. apii s. lat. (Crous & Braun 2003). Cercospora 

neobougainvilleae differs from the 

plurivorous C. apii s. lat. by having obclavate 

conidia and well-developed stromata (11.5– 

71.5 μm diam.). 

Cercospora neobougainvilleae differs 

from C. furfurella in appearance of leaf spots, 

stromata and septation characteristics. The 

symptoms of C. neobougainvilleae are pale at 

the center with dark brown margin, but C. 

furfurella symptoms are almost lacking or dark 

purple to almost black with grey center. The 

stromata of C. neobougainvilleae is welldeveloped 

but C. furfurella stromata are small 

or sometimes lacking. The conidia septation in 

C. neobougainvilleae are distinct with 3–6septa, 

but C. furfurella is characterized by 4–5indistinct 

septa. Moreover, the conidia sizes of 

C. neobougainvilleae are different (4–112 × 4– 

8 μm vs 30–120 × 2–4.5 µm for C. furfurella). 

Cercospora mirabilis Tharp, described 

from Mirabilis jalapa, is characterized by 

having amphigenous caespituli, small or lacking 

stromata, short branches conidiophores, 

and acicular conidia with indistinct septation 

(Chupp 1954). Cercospora neobougainvilleae 

differs from C. mirabilis by having epiphyllous 

caespituli, well-developed stromata, unbranched 

conidiophores, and obclavate conidia with 

distinct septation. 

Orchidaceae 

72 

Cercospora habenariicola Meeboon, Hidayat 

& C. Nakash., Mycotaxon 99: 118 (2007). 

(Fig. 55) 





surrounded by a dark margin or brown halo. 

Caespituli amphigenous, ochre yellow, velvety. 

Stromata 25–75 μm diam., intraepidermal, 

well-developed, subglobose, brown to blackish 

brown. Conidiophores 50–285 × 7.3–7.5 μm, 

occasionally up to 952 μm long, loose to dense 

fascicules, 2–9-septate, numerous, simple, 

straight, erect to decumbent, smooth, pale 

yellow to pale brown, rarely branched, subcylindrical, 

geniculate to sinuous. Conidiogenous 

cells integrated, terminal, sympodially 



Conidia 75–110 × 4.9–5 μm, solitary, narrowly 

obclavate to subacicular, straight, hyaline, 6– 

10-septate, smooth, apex subacute, base 

obconically truncate, hila 1.2–2.9 μm diam., 



Chiang Mai Province, Queen Sirikit Botanic 

Garden, on leaves of Habenaria susannae (L.) 

R. Br. (Orchidaceae), 14 July 2006, Jamjan 

Meeboon (CMUMH 155: Holotype). 

Hosts – Habenaria susannae (Orchidaceae) 

(Meeboon et al. 2007a). 

Fig. 55 – Line drawings of Cercospora habenariicola on Habenaria susannae (from holotype). a. 

Conidiophores and stromata. b. Conidia. Bars: a, b = 40 μm. (Meeboon et al. 2007a).


(Meeboon et al. 2007a). 

Notes – This species belongs to Cercospora 

s. str. because of pigmented conidiophores, 

thickened and darkened conidiogenous 

loci, and hyaline scolecoid conidia (Crous & 

Braun 2003). It is easily distinguishable from 

the plurivorous C. apii s. lat. by having welldeveloped 

stromata and obclavate conidia with 

an obconically truncate base (Crous & Braun 

2003). 

On orchids numerous species of Cercospora 

s. lat. are known, including C. cypripedii 

Ellis & Dearn., C. dendrobii H.C. Burnett, 

C. odontoglossii Prill. & Delacr. and C. peristeriae 

H.C. Burnett, which have been excluded 

and reallocated to the genus Pseudocercospora 

Speg. (Crous & Braun 2003). Cercospora 

angraeci Feuilleaub. & Roum., described from 

orchids, is an insufficiently known species of 

unclear generic affinity (Crous & Braun 2003), 

but based on the original description C. 

habenariicola differs from C. an-graeci in 

having much longer, occasionally branched 

conidiophores (Chupp 1954). Cercospora 

cephalantherae Ondřej & Zavřel, a genuine 

species of Cercospora s. str., is characterized 

by having very short, narrow conidiophores 

(10–25 × 3.5 μm) and relatively short, narrow 

conidia (40–100 × 2–3.5 μm). Cercospora 

habenariicola is morphologically close to C. 

epipactidis C. Massal. However, the latter 

species has consistently unbranched, small 

conidiophores (10–45 × 4–6 μm), and short, 

narrow conidia (30–130 × 3.5–5 μm) (Chupp 

1954). Cercospora eulophiae M.S. Patil is 

another cercosporoid fungus on an orchid 

(Eulophia sp.), but this species was described 

having straight to flexuous, smaller 

conidiophores and pigmented conidia (75–250 

× 3.2–4 μm). 

Oxalidaceae 

Cercospora oxalidis (A.S. Mull. & Chupp) U. 

Braun & Crous, CBS Biodiversity Series 1: 

300 (2003). 

≡ Cercospora oxalidis A.S. Mull. & 

Chupp, Arq. Inst. Biol. Veget. Rio de Janeiro 1: 

218 (1935) (nom. inval.) Fig. 56. 


Leaf spots 1–5 mm in diam., amphigenous, 

scattered to confluent, distinct, circular 

to subcircular, pale brown to tan, centre greyish 

brown to greyish white, dark brown margin. 


in diam., small, brown to dark brown, irregular, 

composed of a few brown hyphal cells. 

Conidiophores 14–122 × 2.5–4.5 μm, loose 

fascicules, 1–4-septate, emerging through the 

cuticle, or sometimes from stromata, straight to 

slightly curved, pale olivaceous brown or 

sometimes paler towards the apex, geniculate. 


intercalary, holoblastic, polyblastic, sympodially 

proliferating. Conidiogenous loci 2.5–3.5 



acicular, straight to mildly curved, hyaline, 5– 

10-septate, smooth, obconically truncate at the 

base, tapering toward a subacute apex, hila ± 1 



Chiang Mai province, Mae Jam District, Mae 

Hae Royal Project Area, on leaves of Oxalis 

debilis Kunth var. corymbosa (DC.) Lourteig 

(Oxalidaceae), 12 February 2008, Jamjan 


Hosts – Oxalis sp., Oxalis debilis Kunth 

var. corymbosa (Oxalidaceae) (Crous & Braun 

2003, Meeboon 2009). 

Distribution – Brazil, Thailand and USA 


Notes – The first report of C. oxalidis 


Polypodiaceae 

Cercospora platycerii Chupp, Monograph of 

Cercospora: 456 (1954). Fig 57 


subcircular to irregular, grey to pale brown, 

with dark brown margin, numerous and scattered 

through the leaf surface. Caespituli 

amphigenous. Stromata 16–58 μm diam., small 

to well-developed, substomatal and composed 

of a few, globose to subglobose, brown to 


3–5.5 μm, densely fasciculate, 3–10-septate, 

straight to decumbent, smooth, brown at the 

73

Fig. 56 – Line drawings of Cercospora oxalidis on Oxalis debilis. a. Conidiophores and stromata. b. 


Fig. 57 – Line drawings of Cercospora platycerii on Platycerium wallichii. a. Conidia. b. 


74

ase, paler toward the apex, unbranched, 

cylindrical, geniculate, sinuous at the apex. 





μm, solitary, obclavate to acicular, straight, 

hyaline, 5–24-septate, smooth, truncate at the 



darkened. 


Doi Sa Ket, Chiang Mai Province, on leaves of 

P. bifurcatum (Cav.) C. Chr., 5 July 2006, 

Jamjan Meeboon (CMU 27904); Chiang Mai 

Province, T. Sansai, on leaves of P. wallichii 

Hook. (Polypodiaceae), 12 September 2007, 

Parin Noiruang (BBH 23741); Chiang Mai 

Province, Pang Da Royal Project, on leaves of 

P. wallichii, 5 August 2008, Jamjan Meeboon 

(BBH 23733). 

Hosts – Platycerium bifurcatum, Platycerium 

sp. (Polypodiaceae) (Crous & Braun, 

2003, Meeboon et al. 2007b,c). 

Distribution – Thailand, USA (Crous & 

Braun 2003, Meeboon et al. 2007b,c). 

Notes – The first record of C. platycerii 

from Thailand was by Meeboon et al. (2007b, c) 

on P. wallichii and P. bifurcatum. 

Pteridaceae 

Cercospora cyclosori Goh & W.H. Hsieh, 

Trans. Mycol. Soc. R.O.C. 4: 26 (1989). 

≡ Cercospora cyclosori Sarbajna & 

Chattopadh., J. Mycopathol. Res. 28: 14 (1990) 

(nom. illeg.), homonym of C. cyclosori Goh & 

W.H. Hsieh (1989). 

(= C. apii s. lat.) Fig 58 


irregular, white to pale at the center, with 

brown margin, sometimes limited by vein. 


diam., substomatal, small to well-developed, 

composed of a few subglobose, brown-walled 

cells. Conidiophores 111–190 × 3–5 μm, 9–17 

in loose fascicules, 5–7-septate, arising through 

stromata, straight, smooth, brown at the base, 

paler toward the apex, unbranched, cylindrical, 

not geniculate. Conidiogenous cells integrated, 

holoblastic, monoblastic, terminal, sympodially 




cyclosori on Pteris biaurita. a. Conidia. b. 




179–283 × 2.5–3.5 μm, solitary, acicular, 






Chiang Mai Province, Mae Jam District, Mae- 

Hae Royal Project Area, on leaves of Pteris 

biaurita L. (Pteridaceae), 12 February 2008, 


23617). 

Hosts – Cyclosorus acuminatus, 

Cyclosorus sp. (Thelypteridaceae) (Crous & 

Braun 2003), Pteris biaurita L. (Pteridaceae) 


Distribution – India, Taiwan, Thailand 


Notes – This specimen is close to C. 

cyclosori in having dark brown symptoms, 

amphigenous caespituli, conidiophores in 

divergent fascicles and long acicular conidia 

with truncate base (Hsieh & Goh 1990). Crous 

& Braun (2003) assigned this species to 

75


scharifii on Rosa hybrida. a. Conidia. b. 

Conidiophores. Bars: a = 25 μm, b = 50 μm. 


C. apii s. lat. This species was first reported 


Literature – Hsieh & Goh (1990, p. 327– 

329). 

Rosaceae 

Cercospora scharifii Petr., Sydowia 10: 14 

(1957) [1956]. Fig. 59 


circular to subcircular, brown, with dark brown 

to blackish margin. Caespituli amphigenous. 

Stromata lacking. Conidiophores 30.5–141 × 

3–5 μm, 4–9 in loose fascicules, 1–6-septate, 

arising from stomata, straight to decumbent, 

unbranched, cylindrical, smooth, brown at the 

base, paler toward the apex, geniculate, mostly 

near the apex. Conidiogenous cells integrated, 

terminal or intercalary, sympodially proliferating. 



25–38.5 × 3.5 μm, solitary, obclavate, straight, 

hyaline, 4–5-septate, smooth, obconically truncate 


apex, hila 1.5–2 μm diam., thickened and 

darkened. 



76 

Hae Royal Project Area, on leaves of Rosa 

hybrida E.H. L. Krause (Rosaceae), 12 



Hosts – Rosa sp., Rosa hybrida 

(Rosaceae) (Crous & Braun 2003, Meeboon 

2009). 

Distribution – Iran, Thailand (Crous & 


Notes – Cercospora scharifii was first 


Rubiaceae 

Cercospora coffeicola Berk. & M.A. Curtis, 

Grevillea 9: 99 (1881). 

= Cercospora coffeae Zimm., Ber. 

Land-Forstw. Deutch-Oatafr. 2: 35 (1904). 

= Cercospora herrerana Farneti, Atti Ist. 

Bot. Univ. Pavia, Ser. 2, 9: 37 (1911). Fig. 60 


circular to subcircular, brown to dark brown, 

pale at the center, with dark margin. Caespituli 

amphigenous. Stromata 16.5–31 μm diam., 

substomatal to intraepidermal, small, composed 

of a few globose and brown-walled cells. 

Conidiophores 20–140 × 2.5–5 μm, 9–23 in 

loose to dense fascicules, divergent, 2–7-septate, 

arising from stomata, straight, mostly near 

the apex, smooth, brown at the base, paler 

toward the apex, unbranched, cylindrical, 


holoblastic, polyblastic, sometimes monoblastic, 

terminal or intercalary, sympodially 

proliferating. Conidiogenous loci 2–2.5 μm 


Conidia 35–178 × 3–4 μm, solitary, obclavate, 

straight, slightly curved, hyaline, 4–21-septate, 

smooth, obconically truncate at the base, tapering 

toward a subacute apex, hila 2–2.3 μm 




Hae Royal Project Area, on leaves of Coffea 

arabica L. (Rubiaceae), 12 February 2008, 


23600). 

Hosts – Coffea arabica, C. canephora, C. 

excelsa, C. laurina, C. liberica, C. robusta, C. 

stenophylla, Coffea spp. (Rubiaceae) (Crous & 

Braun 2003).


Fig. 60 – Line drawings of Cercospora coffeicola on Coffea arabica. a. Conidia. b. Conidiophores 

and stromata. Bars = 50 μm. (Meeboon 2009). 

Distribution – Widely distributed, including 

American Samoa, Angola, Australia, Brazil, 

Brunei, Cambodia, China, Colombia, Congo, 

Costa Rica, Cuba, Dominican Republic, E1 

Salvador, Ethiopia, Fiji, French Guiana, French 

Polynesia, Guiana, French Polynesia, Gabon, 

Ghana, Guadeloupe, Guatemala, Guyana, Haiti, 

India, Indonesia, Ivory Coast, Jamaica, Japan, 

Kenya, Laos,n Madagascar, Malawi, Martinique, 

Mauritius, Micronesia, Mosambique, 


Panama, Papua New Guinea, Peru, Philippines, 

Puerto Rico, Samoa, Sierra Leone, Somalia, 

South Africa, Sudan, Suriname, Taiwan, 

Tanzania, Thailand, Togo, Trinidad and 

Tobago, Uganda, USA, Vanuatu, Venezuela, 

Yemen, Zimbabwe (Crous & Braun 2003). 

Notes – Cercospora coffeicola was first 

reported from Thailand by Sontirat et al. (1980). 

Literature – Chupp (1954, p. 493-494). 

Saururaceae 

Cercospora houttuyniicola Goh & W.H. Hsieh, 

Bot. Bull. Acad. Sin. Taipei 30: 118 (1989). 



clustered, irregular, dark brown to blackish, 

with dark reddish margins. Caespituli amphigenous. 

Stromata 13–43 μm diam., substomatal, 

small to well-developed, and composed of 5–6, 


brown cells. Conidiophores 47.5–176 × 3–4.5 

μm, 5–11 in loose fascicules, 1–3-septate, 

arising through stomata, straight, cylindrical, 

smooth, brown at the base, paler toward the 

apex, unbranched, mostly not geniculate, sometimes 

slightly geniculate. Conidiogenous cells 

integrated, terminal, holoblastic, monoblastic, 


2–3.5 μm diam., conspicuous, thickened and 

darkened. Conidia 27–99 × 2–5 μm, solitary, 

acicular, straight to curve at the apex, hyaline, 

7–12-septate, smooth, obconically truncate at 

the base, tapering toward a acute apex, hila 2–3 




on leaves of Houttuynia cordata Thunb. 

(Saururaceae), 6 December 2006, Ikumitsu 

Araki (CMU 27907); ibid 19 July 2007, Jamjan 


77

Fig. 61 – Line drawings of Cercospora houttuyniicola on Houttuynia cordata. a. Conidiophores 

and stromata. b. Conidia. Bars = 50 μm. (Meeboon 2009). 

Hosts – Houttuynia cordata 

(Saururaceae) (Goh & Hsieh 1989, Meeboon 

et al. 2007c). 

Distribution – Taiwan, Thailand (Goh & 

Hsieh 1989, Meeboon et al. 2007c). 

Notes – The first report of C. 

houttuyniicola from Thailand was carried out 

by Meeboon et al. (2007c). 

Solanaceae 

Cercospora physalidis Ellis, Amer. Naturalist 

16: 810 (1882), emend. Braun & Melnik, Trudy 

Bot. Inst. im V. L. Komarova 20: 79 (1997). 

≡ Cercosporina physalidis (Ellis) Miura, 

South Manch. Railway Co. Agric. Rept. 27: 

525 (1928). 

= Cercospora solanicola G.F. Atk., J. 

Elisha Mitchell Sci. Soc. 8: 53 (1892). 

= Cercospora nicotianae Ellis & Everh., 

Proc. Acad. Sci. Philadelphia 45: 170 (1893). 

= Cercospora phyalidicola Ellis & 

Barthol., Erythea 4: 28 (1896). 

= Cercospora physalidicola Speg., 

Anales Mus. Nac. Buenos Aires 3: 342 (1899). 

(nom. illeg.). 

= Cercospora raciborskii Sacc. & Syd., 

Syll. Fung. 16: 1070 (1902). 

= Cercosporina physalidicola Speg., 

Anales Mus. Nac. Hist. Nat. Buenos Aires 20: 

426 (1910). 

78 

= Cercosporina daturicola Speg., 

Anales Mus. Nac. Hist. Nat. Buenos Aires 20: 

425 (1910). 

≡ Cercospora daturicola (Speg.) 

Vassiljevsky, Fungi imperfecti parasitici 1. 

Hyphomycetes: 247 (1937). 

≡ Cercospora daturicola (Speg.) W.W. 

Ray, Mycologia 36: 175 (1944). 

= Cercospora capsici Heald & W.A. 

Wolf, Mycologia 3: 15 (1911). 

= Cercospora abchasica Siemaszko, Izv. 

Severo-Kavkazsk. Muz. 12: 26 (1919). 

= Cercospora melongenae Welles, 

Phytopathology 12: 63 (1922). 

= Cercospora atropae Kvashn., Izv. 

Severo-Kavkazsk. Kraev. Stantsii Zashch. Rast. 

4: 37 (1928). 

= Cercosporina petuniae Saito, Trans. 

Tottori Soc. Agric. Sci. 3: 271 (1931). 

≡ Cercospora petuniae (Saito) Chupp & 

A.S. Mull., Ceiba 1: 176 (1950) (nom. illeg.). 

= Cercospora petuniae A.S. Mull. & 

Chupp, Arq. Inst. Biol. Veg. Rio de Janeiro 3: 

96 (1936) (nom. inval.). 

= Cercospora petuniae Sandu & Sarea, 

Lucr. Sti. Inst. Agron. 1962: 94 (1962) (nom. 

illeg.). 

= Cercospora petuniae var. Brevipedicellata 

Chidd., Indian Phytopathol. 12: 120 

(1960) (nom. inval.). 

(= C. apii s. lat.) Fig. 62


irregular, brown to dark brown, pale at 

the center, with dark margin, limited by leaves 

vein. Caespituli amphigenous. Stromata 10.5- 

19 μm diam., substomatal to intraepidermal, 

small, composed of few globose to subglobose, 

brown-walled cells. Conidiophores 27.5–54 × 

2.5–5.5 μm, 3–8 in loose to dense fascicules, 

1–3-septate, straight to decumbent, smooth, 

brown at the base, paler toward the apex, 

unbranched, cylindrical, strongly geniculate. 



Conidiogenous loci 1.5–2.5 μm diam., conspicuous, 

thickened and darkened. Conidia 46.5– 

160 × 2–4 μm, solitary, obclavate to acicular, 

straight, slightly curved, hyaline, 7–15-septate, 

smooth, obconically truncate at the base, 

tapering toward a subacute apex, hila 1–2.3 μm 



Phetchabun Province, Num Nao National Park, 

on leaves of Capsicum frutescens L. 


(Solanaceae), 24 November 2004, Chiharu 


27965); the same host, Chiang Mai Province, A. 

Sarapee, 28 November 2006, Jamjan Meeboon 

(CMU 28065); Chiang Mai Province, Doi 

Suthep-Pui National Park, on leaves of C. 

annuum L., 2 October 2005, Jamjan Meeboon 

(CMU 27938); Chiang Rai Province, Wiang Pa 

Pao, on leaves of C. annuum var. acuminatum 

Fingerh (Solanaceae), 2 February 2008, Jamjan 


A. Mae Taeng, on leaves of Nicotiana tabacum 

L. (Solanaceae), 6 February 2008, Jamjan 


San Sai, Mae Fag, on leaves of C. annuum, 3 



Multiple Cropping Centre, on leaves of 

Solanum nigrum L. (Solanaceae), 1 August 

2008, Jamjan Meeboon (BBH 23612); Chiang 

Mai Province, Suthep-Pui National Park, on 

leaves of S. verbascifolium L., 25 July 2008, 


Fig. 62 – Line drawings of Cercospora physalidis on Solanum nigrum. a. Conidiophores and 


79

Hosts – Atropa bella-donna, Capsicum 

annuum, C. baccatum, C. frutescens, C. 

grossum, Datura alba, D. arborea, D. fastuosa, 

D. metel, D. stramonium, D. suaveolens, 

Hyoscyamus agrestis, H. niger, Lycopersicon 

esculentum, Nicandra physalodes, Nicotiana 

repanda, N. rustica, N. tabacum, Petunia 

axillaris, P. hybrida, P. variabilis, P. violacea, 

Petunia sp., Physalis alkekengi, P. angualata, 

P. franchetii, P. heterophylla, P. hybrida, P. 

lanceolata, P. lobata, P. longifolia, P. minima, 

P. mollis, P. parviflora, P. pubescens, P. 

subglabrata, P. variabilis, P. violacea, P. 

virginica, P. viscosa, Physalis sp., Quincula 

lobata, Solanum aculeatum, S. aethiopicum, S. 

incanum, S. laciniatum, S. luteum, S. 

melongena, S. nigrum, S. torvum, S. tuberosum, 

S. xanthocarpum (Solanaceae) (Crous & Braun 

2003). 

Distribution – Worldwide where the host 

is cultivated, including including Afghanistan, 

American Samoa, Argentina, Armenia, 

Australia, Bangladesh, Barbados, Bhutan, 

Bolivia, Brazil, Brunei, Bulgaria, Cambodia, 

China, Colombia, Congo, Cuba, Cyprus, 

Dominican Republic, Egypt, El Salvador, 

Ethiopia, Fiji, French Antilles, Gabon, Gambia, 

Georgia, Germany Ghana, Guam, Guatemala, 

Guinea, Guyana, Haiti, Hong Kong, India, 

Indonesia, Iraq, Jamaica, Japan, Jordan, Kenya, 

Korea, Laos, Libya, Malawi, Malaysia, 

Mauritius, Mexico, Micronesia, Morocco, 


New Zealand, Pakistan, Palau, Panama, Papua 

New Guinea, Philippines, Puerto Rico, 

Romania, Russia (European part), Samoa, 

Seychelles, Sierra Leone, Singapore, Solomon 

Islands, Somalia, South Africa, Spain, Sri 

Lanka, Sudan, Suriname, Swaziland, Taiwan, 

Tanzania, Thailand, Tonga, Trinidad and 

Tobago, Uganda, Ukraine, USA, Vanuatu, 

Venezuela, Virgin Islands, Wallis and Futuna 

Islands, Yemen, Zambia, Zimbabwe (Crous & 

Braun 2003). 

Notes – The first record of this species 

from Thailand was by Sontirat et al. (1980) as 

‘C. capsici Heald & F.A. Wolf’. Crous & 

Braun (2003) considered this species as C. apii 

s.lat. bird chili (Capsicum frutescens) is an 

important crop in Thailand, and its leaf spot 

disease caused by C. capsici is recognized as 

an important disease. Solanum verbascifolium 

80 

Fig. 63 – Line drawings of Cercospora puyana 

on Solanum indicum. a. Conidiophores and 

stromata. b. Conidia. Bars = 50 μm. (Meeboon 

2009). 

was reported as a new host of this pathogen by 


Cercospora puyana Sydow, Ann. Mycol. 37: 

431 (1939). Fig. 63 



initially appearing pale brown, later becoming 

greyish at the centre, with reddish brown or 

purplish brown margins. Caespituli amphigenous, 

chiefly hypophyllous. Stromata 24–40.5 

μm diam., intraepidermal, well-developed, 

composed of globose to subglobose, brown to 

blackish brown cells. Conidiophores 39.5–127 

× 3–4 μm, numerous in dense fascicules, 

slightly divergent, 1–3-septate, straight, erect to 

decumbent, smooth, pale yellow to pale brown, 






μm, solitary, long obclavate to subacicular, 6– 

19-septate, straight, hyaline, smooth, base


2.5–3 2–3 μm diam., thickened and darkened. 




Solanum indicum L. (Solanaceae), 1 August 

2008, Jamjan Meeboon (JM 108). 

Hosts – Solanum indicum, S. 

trachycypum (Solanaceae) (Meeboon 2009, 

Chupp 1954). 

Distribution – Ecuador, Thailand (Chupp 

1954, Meeboon 2009). 

Notes – This species is distinct from the 

plurivorous C. apii s. lat. in having welldeveloped 

stromata, branched conidiophores 

and being obconically truncate at the base of 

conidia (Chupp 1954). Ten species of 

Cercospora (non C. apii s. lat.) have been 

recorded on plant genus Solanum, viz, C. 

lanugiflori Chupp & A.S. Mull., C. nigri var. 

microsporae L.N. Bhardwaj & Y.S. Paul, C. 

puyana, C. sciadophila (Speg.) Chupp, C. 

solanacea Sacc. & Berl., C. solani Thüm., C. 

solanigena Bhartiya, R. Dubey & S.K. Singh, 

C. solani-nigri Chidd., C. solani-tuberosi 

Thirum. and C. venezuelae var. indica Govindu 

& Thirum. (Crous & Braun 2003). This 

specimen is close to C. puyana in having 

amphigenous caespituli, branched conidiophores 

and long obclavate conidia with an 

obconically truncate base. This species was 

first reported from Thailand by Meeboon 

(2008). 


Cercospora solanacea Sacc. & Berl., Atti 

Reale Ist. Veneto Sci. Lett. Arti VI, 3: 721 

(1885). Fig. 64 


angular, at first pale greenish to ochraceous, 

later brown to dark brown, finally with 

grayish brown centre, surrounded by a dark 

margin. Caespituli epiphyllous. Stromata 19– 

24 μm diam., intraepidermal, small to welldeveloped, 

composed of globose to subglobose, 


27–79.5 × 2–4.5 μm, 5–7 in dense fascicules, 

1–3-septate, arising from stomata, simple, 

straight, erect to decumbent, smooth, pale 


not geniculate. Conidiogenous cells 



solanacea on Solanum torvum. a. Conidiophores 



integrated, terminal, holoblastic, mostly monoblastic. 



30–71.5 × 3–3.5 μm, solitary, narrowly obclavate, 

3–6-septate, straight, hyaline, smooth, 

base obconically truncate, with subacute apex, 

hila 2–2.5 2–3 μm diam., thickened and 

darkened. 




Solanum torvum Sw. (Solanaceae), 1 August 


Hosts – Solanum melongena, S. Nigrum, 

S. Torvum, S. verbascifolium (Solanaceae) 


Distribution – Australia, China, India, 

Taiwan, Thailand, Venezuela (Crous & Braun 

2003, Meeboon 2009). 

Notes – This specimen is close to C. 

lanugiflori and C. solanacea based on epiphyllous 

caespituli, relatively short and not 

geniculate conidiophores, and narrowly obcla- 

81

vate and short conidia with a few septate 

(Chupp 1954). However, the status of C. 

lanugiflori is unclear (Crous & Braun 2003), 

therefore, this specimen is assigned to C. 

solanacea. Cercospora solanacea was first 



Verbenaceae 

Cercospora tectonae F. Stevens (tectoniae), 

Bernice P. Bishop Mus. Bull. 19: 155 (1925). 


Leaf spots 2–14 mm in diam., angular 

to suborbicular, limited by leaf veins, confluent, 

brown to greyish brown or white at the centre, 

with a dark margin. Caespituli amphigenous, 

chiefly epiphyllous. Stromata 8–41 µm diam., 

small to well–developed, composed of a few 

globose to subglobose, brown to dark brown 

cells. Conidiophores 33.5–76 × 3–5 µm, in 

loose fascicules, 1–5–septate, straight to decumbent, 

light brown to medium brown, paler 

towards the apex, geniculate at the apex. 



Conidiogenous loci 2–3 µm diam., conspicuous, 

thickened and darkened. Conidia 31–96.5 × 2– 

3 µm, hyaline, acicular to obclavate–cylindric, 

4–13–septate, straight to curved, truncate or 

obconically truncate at the base, with subacute 

82 

apex, hila 2–2.5 µm diam., thickened and 

darkened. 




Tectona grandis L.f. (Verbenaceae), 1 

December 2005, Jamjan Meeboon (CMU 

27928). 

Hosts – Tectona grandis (Verbenaceae) 


Distribution – China, Indonesia, Taiwan, 

Thailand, Trinidad and Tobago, USA (Crous & 


Notes – Crous & Braun (2003) assigned 

this species to C. apii s.lat. The first report of C. 

tectonae from Thailand was by Meeboon et al. 

(2007c). 

Literature – Chupp (1954, p. 595) 

Zingiberaceae 

Cercospora alpiniicola S.Q. Chen & P.K. Chi 

(alpinicola), Journal of South China 

Agricultural University 11: 57 (1990b); also in 

Chi, Fungal Diseases of Cultivated Medicinal 

Plants in Guangdong Province: 33 (1994). 

Fig. 66 


distinct, circular to irregular, pale olivaceous 

brown at the center, sometimes discoloration 

forming surrounding the margin. Caespituli 

amphigenous. Stromata 12.5–19 μm diam., 

Fig. 65 – Line drawings of Cercospora tectonae on Tectona grandis. a. Conidia. b. Conidiophores 

and stromata. Bars = 50 μm. (Meeboon 2009).


Fig. 66 – Line drawings of Cercospora alpiniicola on Alpinia purpurata. a. Conidia. b. 


small, substomatal to intraepidermal, composed 

of a few globose to subglobose, brown–walled 

cells. Conidiophores 48.5–100 × 4.5–6.5 μm, 

3–8 in loose to dense fascicules, 1–3–septate, 

arising from stromata, straight, smooth, brown 

at the base, paler toward the apex, unbranched, 

cylindrical, strongly geniculate. Conidiogenous 

cells integrated, holoblastic, terminal, polyblastic, 

sometimes monoblastic, sympodially 



Conidia 39.5–162 × 3.5–5.5 μm, solitary, 

obclavate to acicular, straight, slightly curved, 

hyaline, 4–11–septate, smooth, obconically 

truncate at the base, tapering toward a subacute 


darkened. 


Chiang Mai Province, Sanpatong District, Mae 

Wang Sub-district, Tambol Mae Win, Bahn 

Mae Sapok, Mae Sapok Royal Project, on 

leaves of Alpinia purpurata K. Schum, 

(Zingiberaceae), 8 February 2008, Jamjan 


Hosts – Alpinia oxyphylla, A. purpurata 

(Zingiberaceae) (Chen & Chi 1990b, Meeboon 

2009). 

Distribution – China, Thailand (Chen & 

Chi 1990b, Meeboon 2009). 

Notes – The first report of C. alpiniicola 


Acknowledgements 

This work was financially supported by 

the Thailand Research Fund (DBG5380011) 

and the Hitachi Scholarship Foundation. The 

authors would like to thank Dr. Eric McKenzie 

for critical reading and providing suggestions 

to improve the manuscript. 

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