Pestalotiopsis—morphology, phylogeny, biochemistry and diversity

Abstract

The genus Pestalotiopsis has received consider-able attention in recent years, not only because of its role as a plant pathogen but also as a commonly isolated endophyte which has been shown to produce a wide range of chemically novel diverse metabolites. Classification in the genus has been previously based on morphology, with conidial characters being considered as important in distinguishing species and closely related genera. In this review, Pestalotia, Pestalotiopsis and some related genera are evaluated; it is concluded that the large number of described species has resulted from introductions based on host association. We suspect that many of these are probably not good biological species. Recent molecular data have shown that conidial characters can be used to distinguish taxa; however, host association and geograph-ical location is less informative. The taxonomy of the genera complex remains confused. There are only a few type cultures and, therefore, it is impossible to use gene sequences in GenBank to clarify species names reliably. It has not even been established whether Pestalotia and Pestalotiopsis are distinct genera, as no isolates of the type species of Pestalotia have been sequenced, and they are morphologically somewhat similar. When selected GenBank ITS accessions of Pestalotiopsis clavispora, P. disseminata, P. microspora, P. neglecta, P. photiniae, P. theae, P. virgatula and P. vismiae are aligned, most species cluster throughout any phylogram generated. Since there appears to be no living type strain for any of these species, it is unwise to use GenBank sequences to represent any of these names. Type cultures and sequences are available for the recently described species P. hainanensis, P. jesteri, P. kunmingensis and P. pallidotheae. It is clear that the important species in Pestalotia and Pestalotiopsis need to be epitypified so that we can begin to understand the genus/genera. There are numerous reports in the literature that various species produce taxol, while others produce newly discovered compounds with medicinal potential and still others cause disease. The names assigned to these novel compound-producing taxa lack an accurate taxo-nomic basis, since the taxonomy of the genus is markedly confused. Until the important species have been epitypi-fied with living strains that have been sequenced and deposited in public databases, researchers should refrain from providing the exact name of species.

Full text

Pestalotiopsis—morphology, phylogeny, biochemistry
and diversity
Sajeewa S. N. Maharachchikumbura &
Liang-Dong Guo &Ekachai Chukeatirote &
Ali H. Bahkali &Kevin D. Hyde
Received: 8 June 2011 / Accepted: 22 July 2011 / Published online: 31 August 2011
#Kevin D. Hyde 2011
Abstract The genus Pestalotiopsis has received consider-
able attention in recent years, not only because of its role as
a plant pathogen but also as a commonly isolated
endophyte which has been shown to produce a wide range
of chemically novel diverse metabolites. Classification in
the genus has been previously based on morphology, with
conidial characters being considered as important in
distinguishing species and closely related genera. In this
review, Pestalotia,Pestalotiopsis and some related genera
are evaluated; it is concluded that the large number of
described species has resulted from introductions based on
host association. We suspect that many of these are
probably not good biological species. Recent molecular
data have shown that conidial characters can be used to
distinguish taxa; however, host association and geograph-
ical location is less informative. The taxonomy of the
genera complex remains confused. There are only a few
type cultures and, therefore, it is impossible to use gene
sequences in GenBank to clarify species names reliably. It
has not even been established whether Pestalotia and
Pestalotiopsis are distinct genera, as no isolates of the
type species of Pestalotia have been sequenced, and they
are morphologically somewhat similar. When selected
GenBank ITS accessions of Pestalotiopsis clavispora, P.
disseminata, P. microspora,P. neglecta,P. photiniae, P.
theae, P. virgatula and P. v i s m i a e are aligned, most species
cluster throughout any phylogram generated. Since there
appears to be no living type strain for any of these species,
it is unwise to use GenBank sequences to represent any of
these names. Type cultures and sequences are available for
the recently described species P. hainanensis, P. jesteri,P.
kunmingensis and P. pallidotheae. It is clear that the
important species in Pestalotia and Pestalotiopsis need to
be epitypified so that we can begin to understand the
genus/genera. There are numerous reports in the literature
that various species produce taxol, while others produce
newly discovered compounds with medicinal potential and
still others cause disease. The names assigned to these
novel compound-producing taxa lack an accurate taxo-
nomic basis, since the taxonomy of the genus is markedly
confused. Until the important species have been epitypi-
fied with living strains that have been sequenced and
deposited in public databases, researchers should refrain
from providing the exact name of species.
Keywords Epitypify .Host occurrence .Pestalotia .
Pestalosphaeria .Pigmentation .Secondary metabolites .
Taxol
Introduction
Pestalotoiopsis Steyaert is an appendage-bearing conidial
anamorphic form (coelomycetes) in the family Amphi-
sphaeriaceae (Barr 1975,1990;Kangetal.1998,1999), and
molecular studies have shown that Pestalotiopsis is mono-
phyletic (Jeewon et al. 2002,2003,2004). Species of
Pestalotiopsis are common in tropical and temperate
S. S. N. Maharachchikumbura :L.-D. Guo (*)
Key Laboratory of Systematic Mycology & Lichenology,
Institute of Microbiology, Chinese Academy of Sciences,
Beijing 100190, People’s Republic of China
e-mail: guold@sun.im.ac.cn
S. S. N. Maharachchikumbura :E. Chukeatirote :K. D. Hyde (*)
School of Science, Mae Fah Luang University,
Thasud, Chiang Rai 57100, Thailand
e-mail: kdhyde3@gmail.com
A. H. Bahkali :K. D. Hyde
College of Science, Botany and Microbiology Department,
King Saud University,
P.O. Box: 2455, Riyadh 1145, Saudi Arabia
Fungal Diversity (2011) 50:167–187
DOI 10.1007/s13225-011-0125-x

ecosystems (Bate-Smith and Metcalfe 1957)andmaycause
plant disease (Das et al. 2010), are often isolated as
endophytes (Liu et al. 2006; Wei et al. 2007; Watanabe et
al. 2010), or occur as saprobes (Wu et al. 1982; Agarwal and
Chauhan 1988; Yanna et al. 2002;Huetal.2007; Liu et al.
2008a). The genus has received much attention from the
scientific community. However, this not because of its
pathogenic nature (Hyde and Fröhlich 1995; Rivera and
Wright 2000; Yasuda et al. 2003), but rather because its
species have been shown to produce many important
secondary metabolites (Strobel et al. 1996a,2002;Ding
et al. 2008a,b; Aly et al. 2010;Xuetal.2010). The aim of
the present paper on Pestalotia, Pestalotiopsis and similar
genera is to review (1) historical aspects, (2) morpholog-
ical and molecular studies, (3) life mode of taxa, (4)
species numbers and (5) biochemical production by selected
species. The problems of understanding the genus are
discussed and the work needed to resolve these problems
elaborated. In most cases problems arise due to misidentifica-
tion of taxa and the review illustrates the importance of the
correct identification of strains before they are used in
biochemical or other studies.
History
De Notaris (1839) introduced the genus Pestalotia De Not.
based on the generic type Pestalotia pezizoides De Not.,
which occurred on the leaves of Vitis vinifera in Italy. This
species is characterized by 6-celled conidia with four
deeply olivaceous central cells, distosepta, hyaline terminal
cells and simple or branched appendages arising from the
apex (Fig. 1.). Steyaert (1949) revised Pestalotia and
divided the genus into three main groups based on the
conidial forms. Steyaert (1949) also introduced two new
genera, Truncatella Steyaert for 4-celled conidial forms and
Pestalotiopsis Steyaert for the 5-celled forms, while the 6-
celled forms remained in Pestalotia.Pestalotia was consid-
ered to be a monophyletic genus and Steyaert (1949)
suggested that the type species could be distinguished from
Pestalotiopsis by it cupulate conidiomata and distoseptate
median cells. Steyaert (1949) further divided Pestalotiopsis
into additional sections based on the number of apical
appendages. These were the Monosetulatae, Bistulatae,
Trisetulatae and Multisetulatae, which were further divided
into subdivisions. Conidia with a single setulae (apical
appendage) were included in the Monosetulatae, which was
further divided into forms with simple and branched setulae.
Conidia with two setulae or on average two setulae were
included in the Bistulatae. Conidia with three setulae or on
average three setulae were included in the Trisetulatae,
which was further divided by concolorous or versicolorous
conidia, fusiform or claviform conidia and spatulate or non-
spatulate setulae. Conidia with more than three setulae were
included in the Multisetulatae. Steyaert (1949) reduced
Monochaetia (Sacc.) Allesch. from its generic state and
placed species with single setula in section Monosetulatae of
Pestalotiopsis and Trun cat ell a. Steyaert (1949)provided
descriptions of 46 species and Pestalotiopsis guepinii
(Desm.) Steyaert was considered to be the type species of
the newly introduced genus. Pestalotiopsis guepinii is
characterized by 4-euseptate and fusiform conidia with a
hyaline basal cell. Steyaert’s introduction of the genus
Pestalotiopsis was not supported by Moreau (1949),
Servazzi (1953) and Guba (1956,1961). Steyaert (1953a,
b,1961, 1963), however, published further evidence in
support of his new genus with answers to the criticisms
made by others.
Fig. 1 Pestalotia pezizoides
De Not. BPI0406483, aConidia
bconidiogenous cells. Scale
bars: a–b=20 μm
168 Fungal Diversity (2011) 50:167–187

The primary work on Pestalotia was carried out by Guba
(1961) in his “Monograph of Monochaetia and Pestalotia”.
Guba (1961) divided the genus into the sections quad-
riloculate, quinqueloculatae and sexloculatae for 4-celled
conidia, 5-celled conidia and 6-celled conidia respectively.
For his sections, Guba (1961) used a simple but very
effective system as proposed by Klebahn (1914), which was
based on the number conidial cells. Guba (1961) further
subdivided the sections into different categories, mainly on
the basis of conidial form, colour, and the position, and
character of the setulae. Monochaetia was retained as a
distinct genus based on its single apical appendage, while
Pestalotiopsis and Truncatella, the new genera proposed by
Steyaert (1949), were synonymised with Pestalotia. Guba
(1961) described 258 species of Pestalotia in his mono-
graph. Steyaert (1956) argued that the retention of Mono-
chaetia as a distinct genus based on a single character, a
single apical appendage was incorrect, while other genera
(Pestalotiopsis,Truncatella and Pestalotia) were differentiated
from each other based on a set of characters.
Sutton (1961,1980) gave more weight to conidiogenesis
when considering Pestalotia and Pestalotiopsis, and he
identified three major problems relating to their taxonomy.
According to the Steyaert system, Sutton (1980) concluded
that a large number of species that should be included in
Pestalotiopsis are still placed in Pestalotia by some authors.
In their studies, Guba (1961), Steyaert (1949,1953a,b,
1955,1956,1961) and most other workers used primarily
dried herbarium material. Sutton (1980) pointed out that
when species were grown in artificial culture, they show more
variability and species limits overlap. Therefore, identification
of species from culture and the application of names based on
herbarium taxonomy present a confusing situation.
Sutton (1980) used the investigation of Griffiths and
Swart (1974a,b), which showed the differentiation of
conidial wall development in two species of Pestalotiopsis,
P. funerea (Desm.) Steyaert and P. triseta (Moreau & V.
Moreau) Steyaert and in Pestalotia pezizoides to support
Steyaert’s opinions. Griffiths and Swart (1974a,b) electron
microscopic study was carried out to establish the relation-
ship among Pestalotia and Pestalotiopsis and other allied
generic members of Monochaetia and Seimatosporium
Corda. The minute zonation in conidial wall structure of
P. pezizoides was thought to separate it from Pestalotiopsis
(Griffiths and Swart 1974a,b). Until 1990, phylogenetic
understanding of the taxonomy associated with Pestalotiopsis
and allied genera was based mainly on conidial characters
(Steyaert 1949;Guba1961; Nag Rag 1993), conidiogenesis
(Sutton 1980) and teleomorph association (Barr 1975,1990;
Metz et al. 2000; Zhu et al. 1991).
Morphological characters used to differentiate species of
Pestalotiopsis and similar genera are limited (Hu et al. 2007);
the morphological characters are plastid and morphological
markers vary between host and environment (Egger 1995).
Hu et al. (2007) showed that colony morphology (colour,
growth rate and texture) is highly variable within single
isolates of Pestalotiopsis; this phenomenon can be easily
observed through repeated subculturing. Also within a single
species, conidial morphology (shape and colour of the
median cells), growth rate and fruiting structure, may vary
(Jeewon et al. 2003). Satya and Saksena (1984)observed
Pestalotiopsis glandicola (Castagne) Steyaert and P. versi-
color var. polygoni and found that the intensity of the median
cells varied with culture and host and concluded that colour
of median cells cannot be used to judge their taxonomic
position. Dube and Bilgrami (1965)observedPestalotiopsis
darjeelingensis Dube, Bilgrami & H.P. Srivast. and showed
morphological variation of conidia in culture (dimension,
length of the setulae, shape, number of cells and the colour
of the cells). Similar observations were made by Purohit and
Bilgrami (1969) when studying more than 100 pathogenic
strains. Conidiogenesis is also confusing when used for
species separation; Watanabe et al. (1998), showed that
Pestalotiopsis neglecta (Thüm.) Steyaert and P. guepinii
having similar acervuli development.
Jeewon et al. (2003) and Tejesvi et al. (2009) compared
morphology with sequence data and showed that species of
Pestalotiopsis display considerable diversity in morphology
and that isolates grouped together based on similarities in
conidial morphology. Hu et al. (2007) found that conidial
characters such as conidial length, median cell length,
conidial width and colour of median cells were stable
characters within Pestalotiopsis; however, the length of the
apical and basal appendages varied. Jeewon et al. (2003)
evaluated the morphological characters that could be used
to differentiate species of Pestalotiopsis. He suggested that
melanin granule deposition within the cell matrix providing
pigmentation to the median cells has taxonomic value; this
agreed with the findings of Griffiths and Swart (1974a,b).
He suggested that the colour of median cells was useful
for distinguishing species of Pestalotiopsis. Tejesvietal.
(2009) also agreed that species of Pestalotiopsis can be
distinguished on the basis of morphological characters
rather than host-specificity or geographical location. Liu et
al. (2010a) proposed that instead of using “concolorous”
and “versicolor”as proposed by Steyaert (1949) and Guba
(1961), “brown to olivaceous”and “umber to fuliginous”
median cells can be a key character in distinguishing
species in Pestalotiopsis. However the pigmentation can
be effected by environmental conditions, different stages
of spore maturity and the observer’sexpertise(Liuetal.
2010a), hosts, medium, and even different generations
through subculturing (Purohit and Bilgrami 1969;Satya
and Saksena 1984;Huetal.2007). The pigmentation of
the median cell however, can be stable even within a
successive subculture; when using standard conditions and
Fungal Diversity (2011) 50:167–187 169

culture on autoclaved carnation leaf segments (Liu et al.
2010a).
‘The teleomorph of a whole fungus has been traditionally
classified and named separately from their anamorphs. Each
of the morphs of anamorphosis was also given different
binomials as if they were different species. As a result, a whole
fungus finds itself in two classification and nomenclature
systems against the principle of natural classification’
(Shenoy et al. 2007). The gene responsible for the
expression of teleomorph and anamorph evolve at different
rates; anamorph characters tend to be morphologically
divergent even with the monophyletic groups while
teleomorph characters are highly conserved (Chaverri et al.
2003; Dodd et al 2003). The teleomorph characters can thus
be used as a precise taxonomic marker for Pestalotiopsis.
However the anamorph of Pestalotiopsis is Pestalosphaeria
M.E. Barr and only twelve species are known as compared
to the asexual state (235 species names). Pestalotiopsis has
been linked to Neobroomella Petr. one species and was
described by Petrak (1947)andPestalosphaeria (12
species), the genus being described by Barr (1975). As
such, the earliest name is Neobroomella, but this state has
rarely been recorded. Pestalotia De Not. has been linked to
Broomella Sacc. (1883) which has 20 species.
Since Pestalotiopsis is the most commonly used name,
we therefore suggest that this name be adopted for the
anamorph and teleomorph forms. However, if Pestalotia is
found to incorporate species of Pestalotiopsis in future
studies, then this name would be used to represent
Broomella, Neobroomella and Pestalosphaeria.
Morphological characters use in the differentiation
of species
Conidial morphology (Fig. 2.) is the most widely used
taxonomic character for the genus Pestalotiopsis. Most
species are divided into different groups based on the size
of the conidia. The length and width are good taxonomic
markers for the genus and stable within the different media
and the generations in most cases (Hu et al. 2007). Colour
of the median cells is still a widely used character, and all
species separate into three groups based on this- concolorous,
versicolorous umber olivaceous and versicolorous fulig-
inous olivaceous. Molecular evidence indicates that it is
more precise to group species according to concolorous
and versicolorous rather than the above three groups
(Jeewon et al. 2003). The length of the apical appendages
and the number of the apical appendages are also widely
used characters for species identification. Some species
can also be identified by the presence of knobbed apical
appendages. The apical appendages can arise from the top,
middle, bottom or different positions in the apical hyaline
cells and such characters are widely used in species
identification. Furthermore the apical appendages can be
divided into branches; in some species presence or
absence of the basal appendages is another character for
species diagnosis.
Recent molecular data
Hu et al. (2007) showed that the ITS gene is less
informative than the β-tubulin gene in differentiating
endophytic species of Pestalotiopsis in Pinus armandii
and Ribes spp. When gaps in the ITS region are treated as a
missing data, the total number of informative characters is
5% and this results in difficulty in separating taxa and low
statistical support. When β-tubulin gene data are used and
gaps are treated as missing data, the number of informative
characters is about 11%, and when gaps are treated as
newstate, it is more than 15%. Thus, Hu et al. (2007)
pointed out that the β-tubulin genes resolves Pestalotiopsis
phylogeny better than the ITS gene. A combination of both
the β-tubulin and ITS genes gave better phylogenetic
resolution, and they suggested that at least two genes
should be used to resolve the phylogeny of species of
Pestalotiopsis. However, Liu et al. (2010a) disagreed with
Hu et al. (2007) concerning the ITS region as being less
informative when compared to the β-tubulin region. They
indicated that proper analysis and alignment of the ITS
region can be a useful character in grouping Pestalotiopsis
to different types of pigmentation, which can be used as a
key character for the phylogeny of the species. Random
amplification of polymorphic DNA (RAPD) can also be
used to detect genetic diversity in species of Pestalotiopsis
(Tejesvi et al. 2007a). Tejesvi et al. (2009) showed that the
Fig. 2 Some commonly use conidial characters for Pestalotiopsis
species identification (1) colour of the median cells alight
concolorous bdark concolorous cversicolorous (2) size of the
conidia dsmall conidia elarge conidia frelatively long conidia g
relatively broad conidia (3) number of apical appendages htwo
apical appendages ithree apical appendages jfive apical appendages
(4) presence or absence of knobbed apical appendages kapical
appendages without knobbed apical appendages lapical appendages
with knobbed apical appendages (5) length of the apical appendages
mrelatively short apical appendages nrelatively large apical
appendages (6) branched or unbranched apical appendages o
branched apical appendages (7) position of the apical appendages
attached to the apical cell pattached to the top of the apical
appendages qattached to the middle of the apical appendages rsome
attached to the bottom of the apical cell (8) presence or absence of
basal appendages spresence of apical appendages tabsence of apical
appendages. Scale bars: a–b=20 μm
b
170 Fungal Diversity (2011) 50:167–187

Fungal Diversity (2011) 50:167–187 171

ITS region is more informative than internal transcribed
spacer—restriction fragment length polymorphism (ITS-
RFLP). They used five restriction enzymes (Alu I, Hae III,
Ava II, Hpa II and Taq I) in their ITS-RFLP analysis and
showed that ITS-RFLP profiles were distinctly different in
P.virgatula (Kleb.) Steyaert and P.theae (Sawada) Steyaert
and intraspecific polymorphism highly variable in P. micro-
spora (Speg.) G.C. Zhao & N. Li. Based on the ITS
sequence, pathogenic and endophytic strains clustered into
distinct groups and these clusters were irrespective of the
host, parts of the host or location.
Life cycle in Pestalotiopsis
A disease cycle of a pathogen may be closely related to its
life cycle, and the former refers to the emergence,
development and maintenance of the disease (Agrios
2005) but is not discussed further here. Species of
Pestalotiopsis are not highly host-specific and taxa may
have the ability to infect a range of hosts (Hopkins and
McQuilken 2000; Keith et al 2006). Species of Pestalo-
tiopsis cause a variety of disease in plants, including canker
lesions, shoot dieback, leaf spots, needle blight, tip blight,
grey blight, scabby canker, severe chlorosis, fruit rots and
leaf spots (Pirone 1978; Kwee and Chong 1990; Xu et al.
1999; Tagne and Mathur 2001; Sousa et al. 2004; Espinoza
et al. 2008). Pirone (1978) considered that species of
Pestalotiopsis are weak or opportunistic pathogens and may
cause little damage to ornamental plants; however, Hopkins
and McQuilken (2000) pointed out that some species of
Pestalotiopsis may cause serious damage to pot grown
plants and the number of known infected plant species is
generally increasing.
Pathogenic species of Pestalotiopsis initially make
contact with the host where the infection occurs (inoculum),
probably by means of the conidia or fragmented spores
(Espinoza et al. 2008). These inocula may survive during
harsh weather conditions and may cause primary infections.
Secondary inoculum produced on diseased tissue may
cause secondary infections and increase the severity of the
disease. The source of the inoculum can be wild plantations
(Keith et al. 2006), flowers (Pandey 1990), crop debris,
disease stock plants, used growing media, soil and
contaminated nursery tools (McQuilken and Hopkins
2004), splashed water droplets (Hopkins and McQuilken
1997; Elliott et al. 2004) and also spores in the air (Xu et al.
1999). Species of Pestalotiopsis have constantly been
isolated as endophytes from plant tissues (Wei and Xu
2004;Liuetal.2006; Wei et al. 2005,2007; Tejesvi et al.
2009; Watanabe et al. 2010). We suspect that many
endophytic species remain as dormant symptomless
inhabitants of plants until the plant is stressed, and then
the endophytes become pathogens. This is thought to
occur in other pathogenic genera (Gehlot et al. 2008). The
pathogenic phase may be triggeredbyacombinationof
environmental factors, plant susceptibility and the viru-
lence of the pathogen. However, further research is needed
to prove the endophytic pathogenic relationship in the
genus. Pestalotiopsis is also considered to be a weak
pathogen (Madar et al. 1991), and most weak pathogens
penetrate the host through natural openings such as stoma,
lenticels and hydathodes (Agrios 2005). Wright et al. (1998)
stated that species of Pestalotiopsis only infect wounded or
stressed plants, so pruning wounds or other physical means
play important roles in disease development (Elliott et al.
2004; McQuilken and Hopkins 2004; Keith et al. 2006).
Plants may also be stressed due to insect, pesticide or sun
damage (Hopkins and McQuilken 2000). High temperature,
high rainfall and human activities may also trigger infections,
and this may lead to disease development (Tuset et al. 1999;
Hopkins and McQuilken 2000; Elliott et al. 2004). The
anamorph-teleomorph relationships and life cycles are not
well known for most species, as the sexual stage does not
often develop (Armstrong-Cho and Banniza 2006). There-
fore, conidia therefore appear to play a key role in providing
the inocula. A general disease cycle for Pestalotiopsis is
illustrated in Fig. 3.
The spore of Pestalotiopsis is considered to be a dry
spore. Watanabe et al. (2000) studied conidial adhesion and
germination of spores of P. neglecta and showed that
infection occurs in four stages. At the beginning, the lower
median cell germinates and becomes firmly attached to the
substrate. Future successive infections can be achieved by
two upper median cells. In the first stage, weak adhesion is
achieved by the mucilaginous matrix coating the conidia. A
second weak adhesion occurs at the bases of the pedicel.
The next two stages provide a strong attachment by release
of fibrillar adhesive substances. In the third stage, fibrillar
adhesive substances are produced along the length of the
pedicel to the apex of the basal cell and at times a smaller
amount of fibrillar material is released from the apical
appendages. The fourth stage involves the release of
fibrillar material at the point of germ tube emergence. Nag
Rag (1993) described conidiomata of the genus as variable,
ranging from acervuli to pycnidia. Conidiomata can be
immersed to erumpent, unilocular to irregularly pluriloc-
ular with the locules occasionally incompletely divided
and dehiscence by irregular splitting of the apical wall or
overlying host tissue (Nag Rag 1993). Conidiophores
partly or entirely develop inside the conidiomata, and they
can be reduced to conidiogenesis cells which are discrete or
integrated, cylindrical, smooth, colourless and invested in
mucus (Nag Rag 1993). Pycnidia can mostly be seen with
the unaided eye as a black or brown spore masses with
copious conidia.
172 Fungal Diversity (2011) 50:167–187

Control strategies are needed for serious Pestalotiopsis
disease, and therefore, knowledge of the causal agent and
the disease cycle is important. Precise knowledge of the
plant/ pathogen interaction and its functional variation
according to the environmental factors are important for
integrated disease management using cultural, biological
and chemical methods. Elliott et al. (2004) stated that
Pestalotiopsis may produce large numbers of spores which
Fig. 3 Disease cycle of the genus Pestalotiopsis (References: revised and redrawn; Von Arx 1974; Nag Rag 1993; Kobayashi et al. 2001)
Fungal Diversity (2011) 50:167–187 173

are easily dispersed in air or by water splash, thus sanitation
and disease management are critical. They suggested that
water management strategies, such as elimination of
overhead irrigation, decreasing wetness of leaves, increas-
ing the spacing of plants and increasing the air circulation,
can reduce disease in palm plantations. Different harvesting
factors also directly affected disease development in tea
plantations. Sanjay et al. (2008) showed that highest disease
incidence occurred in continuously shear-harvested fields
and least in hand-plucked plantations, and they evaluated
systemic fungicide and biocontrol agents such as a
Trichoderma, Gliocladium and Pseudomonas for use in
controlling grey blight disease in tea.
Mode of life
Species of Pestalotiopsis commonly cause disease in a
variety of plants (Hyde and Fröhlich 1995; Hopkins and
McQuilken 2000; Tagne and Mathur 2001), are commonly
isolated as endophytes (Kumar and Hyde 2004; Wei and Xu
2004; Wei et al. 2005,2007; Liu et al. 2006; Tejesvi et al.
2009; Watanabe et al. 2010) and some species likely have
endophytic and pathogenic stages in their life cycle (Wei et
al. 2007: Tejesvi et al. 2009). Species have also been
recorded as saprobes (Guba 1961; Wu et al. 1982; Agarwal
and Chauhan 1988; Yanna et al. 2002; Liu et al. 2008a)
where they are recyclers of dead plant material (Okane et al.
1998; Osono and Takeda 1999; Tokumasu and Aoiki 2002)
and even rarely cause disease in humans (Sutton 1999)
Sexual and asexual forms
One fifth of all known anamorphic fungi lack known sexual
states (Shearer et al. 2007), and out of 2,873 anamorphic
genera names, 699 genera and 94 anamorph-like genera are
linked to a sexual state (Hyde et al. 2011). The links
between sexual and asexual stage are mostly from indirect
evidence, with some links known through experimental
or molecular data (Kendrick 1979;Reynolds1993;
Shenoy et al. 2007;Hydeetal.2011). Pestalotiopsis is
a species-rich anamorphic genus with species mostly
lacking sexual morphogenesis, unlike the coelomycetous
genera Colletotrichum and Phyllosticta (Armstrong-Cho
and Banniza 2006; Wulandari et al. 2009)andPenicillium
(Cannon and Kirk 2000). The sexual states or teleo-
morphs of Pestalotiopsis species have been identified as
Pestalosphaeria (Barr 1975)andNeobroomella (Kirk et
al. 2008).
The asexual Pestalotiopsis state and ascomycetous
sexual state have rarely been recorded in the same host
plant (Barr 1975; Nag Raj 1985; Hyde 1996). However, it is
not always clear that the two stages found are definitely the
same biological species and therefore molecular evidence is
needed to link them. In the laboratory species of Pestalo-
tiopsis rarely develop sexual forms (Metz et al. 2000). Zhu
et al. (1991) induced Pestalosphaeria accidenta P.L. Zhu,
Q.X. Ge & T. Xu and P. jinggangensis P.L. Zhu, Q.X. Ge &
T. Xu to form on potato dextrose agar (PDA). However, this
took 5 to 6 months of incubation. Metz et al. (2000)
obtained the sexual state of P. microspora, an endophytic
isolate that produced taxol. The asexual stage formed after
3–6 weeks on water agar with dried yew needles when
incubated at 16–20 C with 12 h of light per day and was
identified as Pestalosphaeria hansenii Shoemaker & J.A.
Simpson. The twelve sexual states known for species of
Pestalotiopsis are listed in Table 1.
Pestalotiopsis Steyaert as a plant pathogen
Pestalotiopsis is a relatively important plant pathogenic
genus known mostly from the tropics, where it causing leaf
blights (Guba 1961) in many plant species (Hyde and
Fröhlich 1995;Xuetal.1999; Das et al. 2010). Species
may also cause rots of fruit and other post harvest disease
(Ullasa and Rawal 1989; Korsten et al. 1995; Xu et al.
1999). It has been estimated that in southern India grey
blight disease of tea (Camellia sinensis)causedby
Pestalotiopsis has resulted in 17% production loss (Joshi
et al. 2009) and 10–20% yield loss in Japan (Horikawa
1986). Five species of Pestalotiopsis - have been recorded
from tea (Agnihothrudu 1964), although P. longiseta
(Speg.) H.T. Sun & R.B. Cao and P. theae are considered
to be the major species causing grey blight (Joshi et al. 2009).
Pestalotiopsis sydowiana (Bres.) B. Sutton causes foliage,
root and stem-base browning disease in container-grown
ericaceous plants, resulting in plant losses and reduced plant
quality (McQuilken and Hopkins 2004). Antheraea assa-
mensis, a silkworm endemic to the north eastern part of India
that depends on Perseabombycina as the primary food plant,
is endangered due to grey blight disease cause by Pestalo-
tiopsis disseminata (Thüm.) Steyaert (Das et al. 2010).
Pestalotiopsis funerea was found to cause leaf spots of
Hakea sericea, a plant that is considered as an invader of
natural habitats in northern Portugal, and this may allow
its use in biological control (Sousa et al. 2004). P.
menezesiana (Bres. & Torrend) Bissett and P. uvicola
(Speg.) Bissett causes postharvest disease of grape (Xu et
al. 1999)andP. clavispora (G.F. Atk.) Steyaert, P.
disseminata and P. microspora cause scab in Guava in
Hawaii (Keith et al. 2006). The economically important
blueberry fruit from Chile is infected by pathogenic P.
clavispora and P. n e g l e c t a , which cause canker and twig
dieback (Espinoza et al. 2008).
174 Fungal Diversity (2011) 50:167–187

In Sicily, the economically important plant Laurus nobilis is
infected by P. uvicola, which causes causing leaf spots and
stem blights (Vitale and Polizzi 2005). Chlorosis and
reduction of growth were recorded in maize fields in the
Cameroons when the plants were infected by P. n e g l ec t a
(Tagne and Mathur 2001). The medicinally important
ornamental shrub Lindera obtusiloba, which grows wild in
the mountain areas of the Korean Peninsula, is infected by P.
microspora, and the affected leaves initially have grey or dark
brown lesions, surrounded by yellowish halos; these enlarge,
coalesce and become entire at a later stage, finally causing
full leaf blight (Jeon et al. 2007). Affected leaves of
Hymenaea courbaril show symptoms of leaf spots and the
pathogen was identified as a P. subcuticularis (Guba) J.G Wei
& T. Xu (Fail and Langenheim 1990). Pathogenic P. funerea
infects conifer species and causes necrosis on infected tissues
and sometimes death of the plants involved (Bajo et al.
2008). The medicinal and ornamental Carapa guianensis is
infected by P. macrochaeta (Speg.) J. Xiang Zhang & T. Xu,
and foliar blight has been observed in the lower canopy of the
plants (Halfeld-Vieira and Nechet 2006). Species of Pestalo-
tiopsis also have the potential to cause leaf and/or fruit spots
on ginger, rambutan, lychee and orchid (Keith and Zee 2010)
Pestalotiopsis glandicola is a postharvest pathogen on
mango in Bangalore; the disease can be observed on the
leaves throughout the year and it provides the inoculum for
mature fruits, which develop postharvest decay during
storage (Ullasa and Rawal 1989). Fruit rot of grapevine is
caused by P. menezesiana and P. uvicola, and the pathogens
were not only isolated from diseased and healthy fruits but
also from the airspora in grape orchards; thus, the authors
pointed out that latent infection or conidial attachment to
the barriers in the field will lead to postharvest disease in
grapes (Xu et al. 1999). Pestalotiopsis fruit rot is one of the
serious postharvest diseases of rambutan fruit in Thailand
(Sangchote et al 1998). Pestalotiopsis psidii (Pat.) Mordue
is considered to be the causal agent of scabby fruit canker
of guava in India and infection results in rapid yield loss
and affects the postharvest quality of the fruits (Kaushik
et al. 1972).
Pestalotiopsis as an endophyte
Most resent Pestalotiopsis research is based on endophytic
isolates (Liu et al. 2006; Wei et al. 2007; Watanabe et al.
2010; Aly et al. 2010) and has resulted in a four new species
being described. These are P. hainanensis A.R. Liu, T. Xu &
L.D. Guo, P. jesteri Strobel, J. Yi Li, E.J. Ford & W.M. Hess,
P. kunmingensis J.G. Wei & T. Xu and P. pallidotheae Kyoko
Watanabe & Yas. Ono. Most endophytic studies have used
morphological characters and either gene sequence data (Hu
et al. 2007; Liu et al. 2007; Wei et al 2007)orRFLP
technique (Tejesvi et al. 2007a) or a combination of gene
sequence and RFLP techniques (Tejesvi et al. 2009)to
distinguish species. The distribution of the endophytic
species of Pestalotiopsis is ubiquitous and is not largely
influenced by geographical factors (Wei et al. 2007; Tejesvi
et al. 2009). Tejesvi et al. (2005) stated that the endophytic
species of Pestalotiopsis dominant in the winter season and
their colonization are comparatively low in the monsoon
season. The colonization frequency of species of Pestalo-
tiopsis increased with the increasing the age of the host plant
and colonization frequency was variable (Wei et al. 2007).
Some endophyte studies in which species of Pestalo-
tiopsis have been recovered are listed in Table 2.
Pestalotiopsis as a saprobe
Species of Pestalotiopsis have been repeatedly isolated as
saprobes from dead leaves, bark and twigs (Guba 1961).
Table 1 List of anamorphs with known teleomorphs
Asexual form Sexual form
Pestalotiopsis baarnensis Steyaert Pestalosphaeria accidenta
Pestalotiopsis sp. Pestalosphaeria alpiniae P.K. Chi & S.Q. Chen
Pestalotiopsis sp. Pestalosphaeria austroamericana Nag Raj & DiCosmo
Pestalotiopsis guepinii var macrotricha (Kleb.) B. Sutton Pestalosphaeria concentrica M.E. Barr
Pestalotiopsis sp. Pestalosphaeria elaeidis (C. Booth & J.S. Robertson) Aa
Pestalotiopsis eugeniae (Thüm.) S. Kaneko Pestalosphaeria eugeniae P.K. Chi & S.M. Lin
Pestalotiopsis neglecta Pestalosphaeria gubae Tak. Kobay., Ishihara & Yas. Ono
Pestalotiopsis microspora Pestalosphaeria hansenii
Pestalotiopsis podocarpi (Dennis) X.A. Sun & Q.X. Ge Pestalosphaeria jinggangensis
Pestalotiopsis sp. Pestalosphaeria leucospermi Samuels, E. Müll. & Petrini
Pestalotiopsis maculiformans (Guba & Zeller) Steyaert Pestalosphaeria maculiformans Marinc., M.J. Wingf. & Crous
Pestalotiopsis besseyi (Guba) Nag Raj Pestalosphaeria varia Nag Raj
Fungal Diversity (2011) 50:167–187 175

Many species have been isolated from soil, polluted stream
water or are associated with the deterioration of wood,
paper, fabrics and decay of wool (Guba 1961). For an
example, P. bicolor (Ellis & Everh.) A.R. Liu, T. Xu & L.D.
Guo, P. funerea,P. monochaetioides (Doyer) Steyaert, P.
montellica (Sacc. & Voglino) Tak. Kobay., P. disseminata,P.
foedans (Sacc. & Ellis) Steyaert, P. versicolor and P.
virgatula are common species recorded either from
decaying leaves or bark. Several saprobic species of
Pestalotiopsis arelistedinTable3.
Pestalotiopsis as a parasymbiont
Lichen symbiosis is an association between a fungus (the
mycobiont) and an alga or a cyanobacterium (the photo-
biont) (Schwendener 1868). Most lichens associate with
only one fungal species, while some have additional
species. In most cases these additional fungal species are
parasitic while few are parasymbiont. A parasymbiont is a
secondary fungus present in the lichen thallus, growing in
intimate association with the primary symbionts without
causing them any apparent harm (Sun et al. 2002).
Pestalotiopsis maculans (Corda) Nag Raj is considered to
be the dominant parasymbiont in the North American
lichen species Cladonia rangiferina, C. subtenuis, C. mitis,
C. leporina, Parmotrema perforatum and Usnea strigosa
(Sun et al. 2002).
Pestalotiopsis as potential human and animal pathogens
Species of Pestalotiopsis are also known to cause human
and animal disease. Pestalotiopsis has been isolated from
Table 2 List of endophytes and associated host
Species Host References
P.clavispora Camellia oleifera, C.sinensis, Terminalia arjuna,
Podocarpus macrophyllus
Liu et al. 2007; Tejesvi et al. 2007a,2009;
Weietal.2007
P. conigena (Lév.) G.C. Zhao & N. Li Lithocarpus glabra, C.nitidissima Wei et al. 2005,2007
P. funerea Catharanthus roseus Srinivasan and Muthumary 2009
P. hainanensis Podocarpus macrophyllus Liu et al. 2007
P.heterocornis (Guba) Y.X.Chen Camellia japonica, C.oleifera, Castanopsis
sclerophylla, Cephalotaxus fortunei, Podocarpus
macrophyllus, Lithocarpus glabra,
Wei et al. 2005,2007; Liu et al. 2007
P. jesteri Fragraea bodenii Strobel et al. 2000
P.karstenii (Sacc. & P. Syd.) Steyaert Camellia japonica, C.sasanqua Liu et al. 2007; Wei et al. 2007
P.kunmingensis Podocarpus macrophyllus Wei et al. 2007
P.mangifolia (Guba) J. Xiang Zhang
&T.Xu
Camellia japonica, C.reticulate, C.sasanqua,
Podocarpus nagi
Liu et al. 2007; Wei et al. 2007
P.microspora Azadirachta indica, Camellia sinensis, Maytenus
ilicifolia, Podocarpus macrophyllus Terminalia
arjuna, T. chebula, Taxus wallichiana,
Taxodium distichum,
Li et al. 1996; Strobel et al. 1996a,b;
Weietal.2005,2007; Gomes-Figueiredo
et al. 2007; Liu et al. 2007; Tejesvi et al.
2007a,2009
P.neglecta Camellia sinensis, C.nitidissima, Podocarpus
macrophyllus, P.nagi, Taxus chinensis,
T.yunnanensis
Liu et al. 2007; Wei et al. 2007
P.olivacea (Guba) G.C. Zhao & J. He Camellia sasanqua, Podocarpus macrophyllus, Liu et al. 2007; Wei et al. 2007
P.oxyanthi (Thüm.) Steyaert Camellia nitidissima, Podocarpus macrophyllus Liu et al. 2007; Wei et al. 2007
P.paeoniae (Servazzi) Steyaert Camellia sasanqua, Cephalotaxus fortune,
Ginkgo biloba, Podocarpus macrophyllus,
Taxus yunnanensis
Wei et al. 2005,2007 Liu et al. 2007
P. palliditheae Pieris japonica Watanabe et al. 2010
P.photiniae (Thüm.) Y.X. Chen Camellia japonica, C.sasanqua, Podocarpus
macrophyllus P.nagi, Taxus chinensis,
Acer palmatum
Wei et al. 2005,2007; Liu et al. 2007
P.subcuticularis Camellia sasanqua, Taxus yunnanensis, T.chinensis, Liu et al. 2007; Wei et al. 2007
P. submersa Sati & N. Tiwari Equisetum sp., Lyonia ovalifolia Sati and Belwal 2005
P.theae Camellia nitidissima, C.sinensis, Holarrhena
antidysenterica, Podocarpus macrophyllus,
Terminalia arjuna
Liu et al. 2007; Tejesvi et al. 2007a,2009;
Weietal.2007
P.versicolor (Speg.) Steyaert Tamarindus indica Liu et al. 2007,2010a
176 Fungal Diversity (2011) 50:167–187

the human sinuses, fingernails, a bronchial biopsy, eyes,
scalp and feet with corneal abrasions (Sutton 1999). One
isolated from cotton was tested in a toxicity bioassay,
which indicated that it caused reduction in weight,
pathological abnormalities and even mortality in rats
(Diener et al. 1976)
Pestalotiopsis in extreme environments
Some species of Pestalotiopsis have also been isolated from
extreme environments and these isolates have been shown
to produce bioactive metabolites (Tejesvi et al. 2007b).
Pestalotiopsis microspora isolated from Taxu s sp. from the
foothills of Himalayas produced taxol (Strobel et al. 1996a),
P. microspora isolated from Sepik River drainage system in
Papua New Guinea produced isopestacin (Strobel et al.
2002)andPestalotiopsis sp. obtained from the gut of a grass
hopper (Chondracris rosee) produced two new phytotoxic
g-lactones, pestalotines A and B (Zhang et al. 2008).
Endophyte-pathogen relationships
Lee et al. (1995) was able to show that P. microspora has an
endophyte-pathogen relationship with the North American
endangered tree Torreya taxifolia.Theydemonstrated
that P. microspora inhabits the inner bark of the tree
without causing symptoms. However, physiological or
environmental factors trigger the fungus to become
pathogenic. Typical symptoms include needle spots,
needle death and stem cankers. The pathogenic ability
of the fungus depends upon it producing phytotoxins,
pestalopyrones, hydroxypestalopyrones and pestalosides.
At the same time antifungal activity by the fungus
produces exudates of pestaloside; this competes with
other fungi. Pestalotiopsis subcuticularis naturally
inhabits Hymenaea courbaril (Leguminosae) and remains
dormant until leaves become mature. Fail and Langenheim
(1990) stated that when leaves become mature the fungal
hyphae spread and enter in to the intracellular spaces of the
leaves. When the plant tissues are damaged due to
mechanical injury such as insect feeding, active infection
by the fungus occurs. The typical symptoms of infected
leaves included serious leaf blight.
Phylogenetic analysis of existing data in GenBank
ITS sequences of 48 species of Pestalotiopsis were down-
loaded from GenBank and aligned using Clustal X. The
alignment was optimized manually to allow maximum
alignment and maximum sequence similarity. Gaps were
treated as missing data. Phylogenetic analysis was carried
out based on the aligned dataset using PAUP* 4.0b10
(Swofford 2002). Ambiguously aligned regions were
excluded from all analyses. Trees were inferred using the
heuristic search option with TBR branch swapping and
1,000 random sequence additions. Maxtrees were unlimited,
branches of zero length were collapsed and all multiple
parsimonious trees were saved. Trees are figured in Treeview
(Page 1996).
An example of the confusion which results from
molecular data is shown in Fig. 4. In this phylogram we
downloaded 44 selected strains of eight species which have
high number of ITS sequences in GenBank plus 4
sequences from ex-type cultures available in GenBank
(Table 4).
According to Jeewon et al. (2003) and Liu et al. (2010a),
pigmentation is a highly weighted character in the lineage of
species of Pestalotiopsis and which can be differentiated into
two main groups based on the colour of the median cells.
This recent finding was previously supported in the
separation of species by Guba (1961) and Steyaert (1949),
based on versicolorous median cells as well as those species
characterized by concolorous median cells. Jeewon et al.
(2003) showed that species such as P. theae with dark
colored concolorous median cells with knobbed apical appen-
dages should be included in the versicolorous group. Jeewon et
al. (2003) argued that the arrangement of Guba (1961)that
groups the versicolorous assemblages of species into umber
olivaceous and fuliginous olivaceous depends on the color
intensity of the median cells. This statements was followed by
Liu et al. (2010a) and they proposed the use of “brown to
Table 3 List of recently recorded saprobes with their host/substrata
Species Host/ substrate References
Pestalotiopsis sydowiana Dead leaves of Calluna vulgaris, Erica sp., Rhododendron ponticum,
R. hybridum,Prunus laurocerasus
Dennis 1995; Ellis and Ellis 1997
P. funerea Dead leaves of Rhododendron sp, Chamaecyparis sp., Cupressus sp.,
Pinus sp., Juniperus sp.
Dennis 1995; Ellis and Ellis 1997
P. theae Seeds of Diospyros crassiflora Douanla-Meli and Langer 2009
P. guepinii Decaying leaves of Dracaena loureiri Thongkantha et al. 2008
P. palmarum Dead culms of Schoenoplectus triqueter Wu et al. 1982
Fungal Diversity (2011) 50:167–187 177

olivaceous”and “umber to fuliginous”colour median cells as
valid for the taxonomy of the genus instead of the use of the
“concolorous”and “versicolor”median cells grouping system
proposed by Steyaert (1949)andGuba(1961).
Pestalotiopsis clavispora, P. disseminata, P. microspora,
P. neglecta,P. photiniae, P. theae,P. virgatula and P. vismiae
can be divided into two groups depending mainly on the
colour of the median cells. One group is the versicolorous
group, consisting of P. clavispora, P. photiniae and P.
virgatula, and dark concolorous median cells with knobbed
apical appendages containing the P. theae group. The other
group consists of species with concolorous median cells (i.e.,
P. disseminata, P. microspora,P. neglecta and P. vismiae.
Almost all strains that separate into two main clades depend
on the concolorous and versicolor system, and only P.
microspora strains AY924295 and FJ478120 cluster in the
wrong clade. However, within the two main groups, the
respective species distributions are scattered and most
species overlap with each other. Because of the limitation
of characters used to differentiate species (Hu et al. 2007)
and many overlapping characters (Sutton 1980), identifica-
tion to species in Pestalotiopsis is presently difficult. For an
example according to Guba (1961), P. disseminata, P.
microspora,P. neglecta and P. vismiae within the concolo-
Fig. 4 Maximum parsimony
phylogram generated from ITS
sequence analysis of selected
sequences from selected species
of Pestalotiopsis including
P. clavispora, P. disseminata,
P. microspora,P. neglecta,
P. photiniae, P. theae,P.virgatula
and P. vismiae downloaded from
GenBank with other
related taxa. Data were analyzed
with random addition sequence,
unweighted parsimony and
treating gaps as missing data.
Type sequences of Pestalotiopsis
pallidotheae,P. hainanensis,
P. jesteri and P. kunmingensis are
in black and bold
178 Fungal Diversity (2011) 50:167–187

rous group have the same conidia size (18–26×5–8μm).
Pestalotiopsis vismiae can be differentiated as it has two
apical appendages, while Pestalotiopsis microspora is
differentiated from P. neglecta and P. dissementa by the
length of the apical appendages. Pestalotiopsis neglecta and
P. dissementa can be distinguished from each other only by
the shape of the conidia. Most of above characters vary
when in culture and following successive subculturing (Hu
et al. 2007). Within the versicolorous group, P. clavispora
and P. photiniae are morphologically very similar (conidia
size 19–26×6–8.5 μm), while P. virgatula can be differen-
tiated from P. clavispora and P. photiniae by its relatively
small conidia (17–23×6–8μm). However, these characters
overlap and thus identification to these species is rather
difficult. For this reason, naming of species is difficult and
highly subjective and many sequences for Pestalotiopsis
deposited in GenBank are likely to be wrongly named.
Species numbers
According to Index Fungorum (http://www.indexfungorum.
org/names/names.asp; accession date, 2010.10.21) there are
235 Pestalotiopsis names, while in MycoBank (www.
mycobank.org/mycotaxo.aspx; accession date, 2010.10.21)
there are 232 names. The reason for the large number of
names is historical and may not reflect the actual number of
species (Jeewon et al. 2004). As with other pathogenic
genera such as Colletotrichum (Cai et al. 2009), species of
Pestalotiopsis were historically named according to the host
fromwhichtheywerefirstobserved.Ifanewhost
occurrence was found a new species was described. For
example, Venkatasubbaiah et al. (1991) isolated a species of
Pestalotiopsis from leaves of Oenothera laciniata and
described the new species P. oenotherae Venkatas., Grand
& Van Dyke. The new species was justified because no
species of Pestalotiopsis had been described previously
from Oenothera and its morphological characters clearly
distinguished it from other species found on any member of
the family Onagraceae (Venkatasubbaiah et al. 1991).
Kohlmeyer and Kohlmeyer (2001) described Pestalotiopsis
juncestris Kohlm & Volkm.-Kohlm which was isolated from
the host Juncus roemerianus; the taxon is morphologically
similar to P.versicolor and several other species of
Pestalotiopsis, but the taxon was described as a new species
based on the host occurrence. Similarly, Pal and Purkayastha
(1992)andSingh(1981) described the new species P.
agallochae A.K. Pal bis and Purkay and P. arborei N.I.
Singh, respectively based on host occurrence. As recently as
2002, Chen et al. (2002) described P. afinis Y.X. Chen & G.
Species GenBank accession numbers Species GenBank accession numbers
P. clavispora AY682928 P. neglecta EU342212
P. clavispora AY924263 P. neglecta FJ037759
P. clavispora DQ812921 P. neglecta GU595050
P. clavispora GU362540 P. pallidotheae AB482220
P. disseminata AY687870 P. photiniae AY682937
P. disseminata DQ001000 P. photiniae AY682943
P. disseminata DQ195782 P. photiniae AY682946
P. disseminata EF055196 P. photiniae DQ812939
P. disseminata HM535728 P. photiniae EU030345
P. disseminata HM535738 P. virgatula AY924281
P. disseminata HM535752 P. virgatula DQ812936
P. disseminata HM535759 P. virgatula DQ813436
P. hainanensis GQ869902 P. virgatula HM535725
P. jesteri AF377282 P. vismiae EF055220
P. kunmingensis AY373376 P. vismiae EF055221
P. microspora AY924278 P. vismiae EF055222
P. microspora AY924285 P. vismiae EU273510
P. microspora DQ000996 P. vismiae EU326213
P. microspora FJ459945 P. vismiae HM535710
P. microspora FJ478120 P. vismiae HM535751
P. microspora FJ487936 P. theae AY924265
P. neglecta AY682930 P. theae DQ812917
P. neglecta DQ812935 P. theae EF423551
P. neglecta EF055209 Truncatella angustata DQ093715
Table 4 Isolates and GenBank
accession numbers of taxa
used to generate the phylogram.
Type species are marked
in bold
Fungal Diversity (2011) 50:167–187 179

Wei, P. alpiniae Y.X. Chen & G. Wei,P.antiarisY.X. Chen
and G. Wei,P.dilleniaeY.X. Chen & G. Wei,P.kuwang-
siensis Y.X. Chen and G. Wei,P.nelumbinisY.X. Chen & G.
Wei,P.schimaeY.X. Chen & G. Wei and P. synsepali Y. X .
Chen & G. Wei based on the host association.
More recently, some new species have been introduced
based on host occurrence, plus morphological and mole-
cular data. Wei and Xu (2004) isolated an endophytic
species of Pestalotiopsis (P. kunmingensis J.G. Wei & T.
Xu) from Podocarpus macrophyllus (Thunb.) Sweet and
described it as a new species, supported by both morpholo-
gical and molecular evidence. An endophytic species isolated
from the Japanese plant Pieris japonica Thunb. L. was
named as Pestalotiopsis pallidotheae Kyoko Watanabe and
Yas. Ono;its conidial morphology is quite similar to P. theae
but molecular data showed it to be distinct (Watanabe et al.
2010). Similarly, Strobel et al. (2000) and Liu et al. (2007)
described P. jesteri Strobel, J. Yi Li, E.J. Ford & W.M. Hess
and P. hainanensis A.R. Liu, T. Xu & L.D. Guo, respectively,
using the same considerations.
Species status and host-specificity within the genus
Pestalotiopsis has been questioned previously or investi-
gated (Zhu 1989; Jeewon et al. 2004; Wei et al. 2005,2007;
Hu et al. 2007). These authors showed that different species
isolated from the same host may not be phylogenetically
closely related (Jeewon et al. 2004; Wei et al. 2007).Weietal.
2007 investigated endophytic species of Pestalotiopsis
associated with plant species in the families Podocarpaceae,
Theaceae and Taxaceae. The endophytic species of Pestalo-
tiopsis associated with these host families were not generally
host-specific, occurring on a range of hosts. For example, P.
neglecta (Thüm.) Steyaert and P.photiniae were isolated
from all the host plants in three plant families. Tejesvi et al.
(2007a) isolated endophytic species of Pestalotiopsis asso-
ciated with the medicinal plants Azadirachta indica, Holar-
rhena antidysenterica, Terminalia arjuna and T. chebula.
They showed that isolates obtained from a single plant were
genetically diverse, while the same species occurred in most
plants. According to Guba (1961), most species of the
Pestalotia were listed from a range of hosts. For example,
Pestalotia microspora was listed from several different host
plants (i.e., Ananas comosus, Araucaria sp., Carya sp.,
Hedera helix, Juniperus bermudiana and Platanus occidenta-
lis). Hu et al. (2007) tested the relationships of endophytic
Pestalotiopsis strains from two tissues of Pinus armandii
and found that even strains isolated from the same tissue
type were not phylogenetically related. Zhu (1989)used
artificial cross inoculation studies to show that pathogenic
species of Pestalotiopsis may not be specific to the single
host. Jeewon et al. (2004) pointed out that host-specificity of
Pestalotiopsis is not supported by the large number of
species recorded on one host. They also argued that many
taxa used in literature can be misinterpretations or synonyms
of species with wide host ranges. Jeewon et al. (2004)used
analysis of ITS and 5.8S rDNA to show that isolates taken
from the same host were not phylogenetically related and
that taxa with similar morphological characters were
phylogenetically related.
Up to this time, most phylogenetic research on Pestalo-
tiopsis has shown that Pestalotiopsis is not highly host-
specific and that species are found on a range of hosts
(Jeewon et al. 2004; Wei et al. 2005,2007;Huetal.2007).
The diseases caused by species of Pestalotiopsis have been
recorded in different ecosystems and infect a diverse range
of unrelated plant taxa. Isolation of endophytic Pestalo-
tiopsis strains for bioprospecting for new biochemical
compounds have shown that the same species can be found
in a range of hosts. Therefore, most of the species recorded
in checklists and the literature may not reflect what actually
occurs. As in other related plant pathogenic genera such as
Colletotrichum, the Pestalotiopsis species concept depends
mostly on the conidial characteristics. It has been shown
that most of the key conidial characters used in species
level separation are not stable and vary with host range,
generation, culture and other environmental conditions (Hu
et al. 2007). The arrangement of species by Steyaert (1949)
and Guba (1961) in various coloured groupings is prob-
lematic because this character has been shown to be
variable within a species (Liu et al. 2010a). Thus, most
species in the above arrangements may be confused and
many species are probably synonyms. Due to the fact that
(1) species of Pestalotiopsis are generally not host-specific,
(2) conidial characters vary and species limits overlap, and
(3) species arrangements in Steyaert (1949) and Guba
(1961) are problematic, then the actual number of species in
Pestalotiopsis is likely to be much lower than presently
recorded in databases (e.g., Index Fungorum, MycoBank)
and the literature (Kirk et al. 2008).
For example, according to Guba (1961), Pestalotiopsis
breviseta (Sacc.) Steyaert, P. e u g e n i a e ,P. ilicicola T. , P.
microspora,P. podocarpi and P. sinensis (C.I. Chen) P.L. Zhu,
Q.X. Ge & T. Xu have very similar, overlapping morpholog-
ical characters and these species were justified mainly
according to the host association. Also the above six species
vary from P. carissae Guba, P. d i s s e m i n a t a ,P. neglecta,andP.
olivacea by the length of the apical appendages. We question
whether these names are synonyms of a single biological
species. Furthermore, the versicolorous umber olivaceous
group which comprises 40 species and versicolorous fuligi-
nous olivaceous group comprising 56 species. These groups
are differentiated depending on the intensities of the median
cells, while most species have similar conidial measurements
and thus are likely to be synonyms. We suspect that the actual
number of biological species may be fewer than 50. The
scientific community, however, uses many more names when
diagnosing disease and in phylogenetic studies and biochemical
180 Fungal Diversity (2011) 50:167–187

studies. Therefore, modern research approaches are needed
for species of Pestalotiopsis in order to establish the
acceptable names.
Species numbers and accepted species
When species are morphologically distinct and molecular
evidence shows they are monophyletic, then such species
can be considered as a distinct and valid species in a
particular genus. Based on their distinct morphological
characters, we suggest that the 20 species listed in Table 5
can be considered as good species in the genus at this time.
Furthermore some other species (Table 6) which have
considerable value because of their economic roles (in
bioactive metabolites production, frequent pathogens, or
frequently isolated endophytes) are possibly good species.
We suggest that type material of these species should be
reexamine and epitypified with fresh collections. With the help
of ex-type living cultures and sequence data, a robust species
concept can be developed for the genus Pestalotiopsis.
Novel Pestalotiopsis biochemistry
Species of Pestalotiopsis have been well-studied because of
the diverse array of novel compounds that they have been
shown to produce. As such, they are thought to be a rich
source for bioprospecting when compared to those of other
fungal genera (Aly et al. 2010; Xu et al. 2010). Strobel and
Long (1998) described Pestalotiopsis as the ‘E.coli of the
temperate and tropical rainforest systems’.Speciesof
Pestalotiopsis may have an important role in forest
ecosystems; they have a cosmopolitan geographical distri-
bution and are found almost everywhere (Tejesvi et al.
2007a). Moreover, species of Pestalotiopsis have been
found to produce an enormous number of secondary
metabolites that may have medicinal, agricultural and
industrial applications. The majority of compounds have
been discovered from endophytic strains of Pestalotiopsis
(Lee et al. 1996; Strobel et al. 1996a,b;LiandStrobel
2001) plus some pathogenic strains (Kwon et al. 1996).
Species of Pestalotiopsis have been shown to produce
bioactive alkaloids, terpenoids, isocoumarin derivatives,
coumarins, chromones, quinones, semiquinones, peptides,
xanthones, xanthone derivatives, phenols, phenolic acids,
and lactones with a range of antifungal, antimicrobial, and
antitumor activities (Xu et al. 2010). Xu et al. (2010)
reviewed 130 different compounds isolated from species of
Pestalotiopsis. In the present review, we discuss some
selected species and their bioactive potential.
Pestalotiopsis microspora is a common species present
in tropical and subtropical plants and is a widespread
saprobe of bark and decaying plant material (Metz et al.
Table 5 Morphologically distinct Pestalotiopsis species with their host and location
Species with distinct morphological characters Host and location
P. gaurae Guba On stem of Gaura parviflora in Hays, Kansas, United States
P. multiseta (Speg.) Guba On fallen leaves of Iris germanica in Conegliano, Italy
P. trevoae Speg. On dead decaying branches of Trevoa trinervia in Santiago, Chile
Pestalotiopsis bicolor Isolated from the dead leaves of Salix sp. in Tuskegee, Alabama, United States
P. distincta (Guba) K. Yokoy. On leaves of Castanopsis cuspidate in Japan
P. funerea On dead leaves of Thuja sp. in Paris, France
P. guepinii On stem and leaves of Camellia japonica in France.
P. hughesii Steyaert On stems of Cyperus articulate in Gold Coasts in West Africa
P. karstenii On leaves of Camellia japonica in United States
P. leucopogonis Nag Raj On leaves of Leucopogan lanceolatus in Australia
P. macrospora (Ces.) Steyaert On fronds of Pteridium aquilinum in Italy
P. maculans On leaves of Camellia japonica and Camellia sp. in Czechoslovakia, France, Germany
and United States
P. monochaetioides On dead twig of Chamaecyparis lawsoniana in Naarden, Holland
P. montellica On dead leaves of Quercus rubra in Canada
P. palustris Nag Raj On Euphorbia palustris in Italy
P. perseae Nag Raj On leaves of Persea borbonea in United States
P. pseudomontellica Nag Raj On leaves of Lithocarpus densiflora in United States
P. smilacis (Schwein.) B. Sutton On stem of Smilax rotundifolia in United Sates
P. tecomicola Nag Raj On Tecoma radicans in United States
P. trichocladi (Laughton) Steyaert On leaves of Trichocladus crinitus in South Africa
Fungal Diversity (2011) 50:167–187 181

2000). The species has most commonly been isolated as an
endophyte associated with rainforest plants (Strobel et al.
2002) or as a pathogen (Keith et al. 2006). Pathogen
associations include scab disease on Psidium guajava
(Keith et al. 2006), leaf blight of Lindera obtusiloba (Jeon
et al. 2007) and as an endophyte on Terminalia morobensis
(Womersley 1995). Pestalotiopsis microspora has the
potential to be a model organism for biological and
biochemical studies in the laboratory (Metz et al. 2000).
Isolates of this species (or possibly species complex) show
diverse genetic variation and thus each individual isolate is
generally unique in the substances that it produces (Harper
et al. 2003). Long et al. (1998) have shown that under
laboratory conditions it can take up heterologous DNA, add
telomeric DNA, express heterologous DNA and can
replicate independently of chromosomal DNA.
Such genetic diversity would be useful to the species in
nature, helping it adapt to a new plant by incorporating
plant DNA into its own genome (Strobel et al. 1996a;Li
et al. 1996). Bioactive compounds such as the anti-cancer
drug taxol, jesterone, ambuic acid, torreyanic acid,
pestaloside, pestalotiopsins and 2-a hydroxydimeniol
(Strobel et al. 2002), hetero-polysaccharides (Kai et al.
2003) have been obtained from P. microspora.The
multimillion dollar anti-cancer drug, taxol was obtained
from an endophytic strain of P. microspora isolated from
Taxus wallachiana (Strobel et al. 1996a)andTaxodium
distichurn (Strobel et al. 1996b).Kaietal.(2003) found
that P. microspora can metabolize various monosacchar-
ides and the composition of hetero-polysaccharides
depends on the type of monosaccharide in the media.
Harper et al. (2003) investigated the production of
pestacin, a 1,3-dihydro isobenzofuran with moderate
anti-fungal properties and high anti-oxidant activity when
compared with the vitamin E derivative trolox from
endophytic strains of P. m i c r o s p o r a . The anti-oxidant
activity works mainly by cleavage of an unusually reactive
C–H bond. Lee et al. (1995) obtained several anti-fungal
compounds such as pestaloside, an aromatic glucoside,
and two pyrones (pestalopyrone and hydroxypestalopyrone)
Table 6 Economically important Pestalotiopsis species with their host and location
Economically
important species
Host and location Economically importance
Pestalotiopsis adusta
(Ellis & Everh.) Steyaert
On leaves of Prunus cerasus in
Newfield, New Jersey, United States
Bioactive metabolites Li et al. 2008b
P. clavispora On leaves of Quercus sp. in Auburn,
Alabama, United States
Plant pathogen,
Common endophyte
Keith et al. 2006; Espinoza et al. 2008;
Weietal.2007; Liu et al. 2007
P. disseminata On dead leaves of Eucalyptus globules
in Coimbra, Portugal
Plant pathogen,
Bioactive metabolites
Das et al. 2010; Keith et al. 2006;
Deyrup et al. 2006
P. fici Steyaert On Ficus sp. in Kiagwe, Uganda Bio active metabolites Liu et al. 2008a,b,2009b
P. foedan (Sacc. & Ellis)
Steyaert
On decaying bark of Thuja occidentalis
in Newfield, New Jersey, United States
Bio active metabolites Ding et al. 2008a
P. heterocornis On leaves of Anarcardium occidentale
in Cantanduva, São Paulo, Brazil
Common endophyte Wei et al. 2007; Liu et al. 2007
P. longiseta On leaves of Rubus caesius in Susegana,
Conegliano, Italy
Plant pathogen,
Bioactive metabolites
Joshi et al. 2009; Nagata and Ando 1989;
Nagata et al. 1992; Xu et al. 2010
P. microspora On leaves of Hedera helix in Botanical
garden, College of Argentina, Buenos
Aires, Argentina
Plant pathogen,
Common endophyte,
Bioactive metabolites
Strobel et al. 1996a,b,2000;Metzetal.2000;
Keith et al. 2006;Jeonetal.2007;
Wom e rsl e y (1995); Harper et al. 2003;
Lee et al. 1995; Kai et al. 2003
P. neglecta On leaves of Euonymus japonicas in
Coimbra, Portugal
Plant pathogen, Endophyte Tagne and Mathur 2001; Espinoza et al. 2008;
Weietal.2007;Liuetal.2007
P. pauciseta (Sacc.)
Y.X. Chen
On leaves of Litsea glutinosa in Mount
Makiling, near Los Banos, Laguna
province, Philippine
Bioactive metabolites Gangadevi et al. 2008
P. photiniae On leaves of Photinia serrulata in Istria,
Australia
Bioactive metabolites Ding et al. 2009
P. theae On leaves of Camellia sinensis in Japan Plant pathogen, Endopyte,
Bioactive metabolites
Li et al 2008a; Nagata et al. 1992;
Shimada et al. 2001; Tuset et al. 1999;
Worapong et al. 2003; Joshi et al. 2009;
Muraleedharan and Chen 1997;
Ding et al. 2008b; Shimada et al. 2001
P. uvicola On Gaura parviflora and Vitis vinifera
in Italy
Plant pathogen Vitale and Polizzi 2005; Xu et al. 1999
182 Fungal Diversity (2011) 50:167–187

from a strain of P. microspora isolated from the endangered
North American tree Torreya taxifolia.WhenPestalotiopsis
microspora is cultured on media containing various mono-
saccharides as a carbon source, different polysaccharides are
produced and this mainly depends on the monosaccharide
used as the carbon source (Kai et al. 2003). Whether all these
strains were in fact P. microspora is yet to be determined,
since the identifications were based on morphology or
comparison with GenBank sequence data, which itself
may be erroneously named. This species is in need of
epitypification.
Pestalotiopsis theae is an economically important spe-
cies that has been reported from all major tea growing
countries of the world (Muraleedharan and Chen 1997) and
also as an endophyte (Worapong et al. 2003). Pestalotheols
A–D, four new metabolites isolated from endophytic
Pestalotiopsis theae, and pestalotheol C showed an inhib-
itory effect against HIV-1
LAI
replication in C8166 cells (Li
et al. 2008a). Three new compounds, pestalamides A–C
and two known metabolites, aspernigrin A and carbonarone
A, were obtained from the same fungus isolated from the
branches of tea (Ding et al. 2008b). The newly isolated
pestalamide B inhibited HIV-1 replication in C8166 cells
with EC
50
of 64.2 μM and antifungal activity against
Aspergillus fumigatus. Chloroisosulochrin and chloroisosu-
lochrin dehydrate were obtained from the culture filtrate of
Pestalotiopsis theae, and these compounds can be used as
plant growth regulators (Shimada et al. 2001). This species
is obviously important as a producer of novel medicinal
metabolites.
The generic type of Pestalotiopsis is P. g u e p i n i i ,a
plant pathogen that causes disease in important crop plants
(Karaca and Erper 2001). Strains of Pestalotiopsis
guepinii isolated as an endophyte from the plant families
Anacardiaceae, Apocynaceae, Leguminosae and Palmae
were tested for their in vitro acetylcholinesterase (AChE)
and butyrylcholinesterase (BuChE) inhibitory activity,
using Ellman’s colorimetric method adapted for thin layer
chromatography (Rodrigues et al. 2005). Pestalotiopsis
guepinii from Anacardium giganteum inhibited both
enzymes in the TLC polar region and a strain isolated
from Myracroduon urundeuva and Spondias mombin
showed selective inhibition of AChE. Parshikov et al.
(2001) suggested that P. guepinii may be a useful model
for the mammalian transformation of fluoroquinolones.
They obtained the metabolites N-acetylciprofloxacin
(52%), desethylene- N-acetylciprofloxacin (9.2%), N-
formylciprofloxacin (4.2%), and 7-amino-1-cyclopropyl-
6-fluoro- 4-oxo-1,4-dihydroquinoline-3-carboxylic acid
(2.3%) by specific culture of P. guepinii dosed with
ciprofloxacin (300 μM). In addition, by dosing with
norfloxacin (313 μM) and the metabolites N-acetylnor-
floxacin (55.4%), desethylene-N-acetylnorfloxacin (8.8%),
N-formylnorfloxacin (3.6%), and 7-amino-1-ethyl-6-
fluoro- 4-oxo-1,4-dihydroquinoline-3-carboxylic acid
(2.1%) were obtained.
Liu et al. (2008b) isolated five new cyclohexanone
derivatives, pestalofones A–E, with the known compounds
isosulochrin, isosulochrin dehydrate, and iso-A82775C,
from cultures of the plant endophytic fungus Pestalotiopsis
fici. Pestalofones A and B were inhibitory against HIV-1
replication in C8166 cells, pestalofones C showed anti-
fungal activity against Aspergillus fumigatus while pestalo-
fones E showed both the above effects. Chloropestolide A
extracted from the scale-up fermentation extract of Pesta-
lotiopsis fici showed significant inhibitory effects on
growth of two human cancer cell lines, HeLa and HT29
(Liu et al. 2009). Liu et al. (2010b) obtained chloropupu-
keanolides A and B (unprecedented spiroketal peroxide)
and chloropupukeanone A (three highly functionalized
metabolites featuring a chlorinated pupukeanane core) from
an endophytic strain of Pestalotiopsis fici. The compound
chloropupukeanolide A showed significant anti-HIV-1 and
cytotoxic effects.
These findings will most likely trigger further studies on
total synthesis. Whether Pestalotiopsis is unique amongst
endophytes or coelomycetes in producing large numbers of
secondary metabolites with medicinal and pathogenic
control significance has yet to be established.
Taxonomic confusion and way forward
Pestalotiopsis is taxonomically poorly understood both at
the inter- as well as the intraspecific level. It is not clear
whether Pestalotia is really distinct from Pestalotiopsis,
since stains of the type of the former have not been
sequenced. Nomenclature of the genus is confusing and
most host based names in databases may be synonyms.
Molecular data have still not been successfully applied for
species-level differentiation and names applied to data in
GenBank are doubtful, as they are not linked to any type
materials. Epitypification with molecular work is therefore
needed to understand the species and what distinguishes
them. Re-examination of type materials and establishment
of epitypes with living cultures is essential for real progress
(Hyde and Zhang 2008), and sequence data are needed to
develop a strong species-based taxonomic system for the
genus Pestalotiopsis. It is only then that plant pathologists
can confidently name disease causal agents, quarantine can
put in effective measures to prevent entry of unwanted
species of Pestalotiopsis, plant breeders can breed resis-
tance against pathogenic species and biochemists can
confidently put names to species producing novel chem-
icals and use an understanding of species relationships to
aid in bioprospecting.
Fungal Diversity (2011) 50:167–187 183

Acknowledgments This project was supported by the Global
Research Network for Fungal Biology, King Saud University and the
Key Lab of Systematic Mycology and Lichenology, Institute of
Microbiology, Chinese Academy of Sciences. Sajeewa Maharachchi-
kumbura thanks the Key Lab of Systematic Mycology and
Lichenology, Institute of Microbiology, Chinese Academy of
Sciences, Beijing and the Mushroom Research Foundation, Chiang
Mai, Thailand, for a postgraduate scholarship.
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