Mycoscience (2007) 48:135–144
DOI 10.1007/s10267-007-0347-7
© The Mycological Society of Japan and Springer 2007
REVIEW
Amy Y. Rossman · David F. Farr · Lisa A. Castlebury
A review of the phylogeny and biology of the Diaporthales
Received: November 21, 2006 / Accepted: February 11, 2007
Abstract The ascomycete order Diaporthales is reviewed
based on recent phylogenetic data that outline the families
and integrate related asexual fungi. The order now consists
of nine families, one of which is newly recognized as
Schizoparmeaceae fam. nov., and two families are recircumscribed. Schizoparmeaceae fam. nov., based on the genus
Schizoparme with its anamorphic state Pilidella and including the related Coniella, is distinguished by the threelayered ascomatal wall and the basal pad from which the
conidiogenous cells originate. Pseudovalsaceae is recognized in a restricted sense, and Sydowiellaceae is circumscribed more broadly than originally conceived. Many
species in the Diaporthales are saprobes, although some are
pathogenic on woody plants such as Cryphonectria parasitica, the cause of chestnut blight, and agricultural crops such
as canker diseases of soybean and sunflower caused by
species of Diaporthe-Phomopsis in both temperate and
tropical regions. Members of the Diaporthales such as
Apiognomonia-Discula and Diaporthe-Phomopsis are commonly encountered as endophytes of woody plants.
Key words Ascomycetes · Canker disease · Endophytes ·
Systematics
Introduction
The ascomycete order Diaporthales includes several plant
pathogenic fungi, of which the most notorious is the chestnut blight fungus [Cryphonectria parasitica (Murrill) M.E.
Barr] that altered the landscape of eastern North America
(Anagnostakis 1988). Numerous other tree diseases are
caused by members of the Diaporthales in either their ascomycetous or asexual states. These diseases include oak
A.Y. Rossman (*) · D.F. Farr · L.A. Castlebury
Systematic Botany and Mycology Laboratory, USDA Agricultural
Research Service, Beltsville, MD 20705, USA
Tel. +1-301-504-5264; Fax +1-301-504-5810
e-mail: arosman@nt.ars-grin.gov
dieback [Apiognomonia quercina (Kleb.) Höhn.], cherry
leaf scorch [A. erythrostoma (Pers.) Höhn.], sycamore canker [A. veneta (Sacc. & Speg.) Höhn.], and ash anthracnose
[Gnomoniella fraxinii Redlin & Stack, anamorph Discula
fraxinea (Peck) Redlin & Stack] in the Gnomoniaceae.
Diseases caused by anamorphic members of the Diaporthales include dogwood anthracnose (Discula destructiva
Redlin) and butternut canker (Sirococcus clavigignentijuglandacearum Nair et al.), both solely asexually reproducing species in the Gnomoniaceae. Species of Cytospora, the
anamorphic state of Valsa, in the Valsaceae cause diseases
on Eucalyptus (Adams et al. 2005), as do species of Chrysoporthe and its anamorphic state Chrysoporthella (Gryzenhout et al. 2004, 2005d). Many diseases of crop plants are
caused by members of the Diaporthales. For example, species of Diaporthe-Phomopsis attack sunflower and soybean,
causing stem cankers and seedling blights (MuntañolaCvetkovi, et al. 1991; Black et al. 1996). The asexually reproducing fungus Greeneria uvicola (Berk. & M.A. Curtis)
Punith., cause of bitter rot of grapes, belongs in the Diaporthales but is not affiliated with any family (Farr et al.
2001).
Molecular data support the Diaporthales as a distinct
order within the Sordariomycetes, the class including ascomycetous fungi that produce their asci primarily in perithecial fruiting bodies (Zhang and Blackwell 2001; Castlebury
et al. 2003; Zhang et al. 2006). In the latest multigene phylogeny of this class, the Diaporthales are a well-defined order in the subclass Sordariomycetidae, most closely related
to the Magnaporthaceae and Ophiostomatales and allied
with the Boliniales, Chaetosphaeriales, Coniochaetales, and
Sordariales (Zhang et al. 2006). Castlebury et al. (2002) and
Zhang et al. (2006) suggest that the Magnaporthaceae
is sister to the Diaporthales. However, recent work by
Mostert et al. (2006) places the Calosphaeriales even closer
to the Diaporthales.
According to Kirk et al. (2001), the Diaporthales consists
of about 94 genera with 500 species. The anamorphic species derived from within the Diaporthales, including such
large genera as Cytospora (100+ species) and Phomopsis
(100+ species), may at least double the estimated number
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of species. The Diaporthales are characterized morphologically by brown to black perithecial fruiting bodies immersed
in a stroma or the substrata, lack of true paraphyses at maturity, and unitunicate asci that float free within the centrum at maturity and have a refractive ring in the apex (Barr
1978; Samuels and Blackwell 2001). The known asexual
states of members of the Diaporthales are generally coelomycetous, producing phialidic, often annellidic conidiogenous cells, and usually non- or one-septate conidia in
acervuli or pycnidia with or without a well-developed
stroma.
Generic concepts in the Diaporthales have not been reevaluated since they were established, primarily on the basis of the Saccardoan system, which placed high value on
ascospore characters. Characteristics of the stroma and the
anamorph were combined with ascospore characteristics in
the taxonomies proposed by Barr (1978, 1990), Kobayashi
(1970), and Monod (1983; Gnomoniaceae only) that represent the most recent monographic accounts of the Diaporthales. The few molecular studies in the Diaporthales
suggest that these generic concepts must be reevaluated
(Zhang and Blackwell 2001; Castlebury et al. 2002, 2003).
In these studies, many of the genera included were determined to be polyphyletic; thus, placement of the type species has been stressed.
Although for many years the genera Magnaporthe
and Gaeumannomyces were included in the Diaporthales
(Barr 1978; Yaegashi and Udagawa 1978; Monod 1983;
Cannon 1988), these genera are now placed in their own
family, Magnaporthaceae (Cannon 1994). In describing
this family with six genera, Cannon (1994) reviewed its
ordinal placement and concluded, based primarily on the
biology of these fungi, that it should be excluded from the
Diaporthales. Three recent studies of these taxa using molecular sequence data confirm the placement of Gaeumannomyces and Magnaporthe outside the Diaporthales (Zhang
and Blackwell 2001; Castlebury et al. 2002; Zhang et al.
2006).
Within the Diaporthales, up to eight families have been
recognized by various authors over the past 30 years. These
familial classifications of the Diaporthales were summarized by Zhang and Blackwell (2001), comparing Barr
(1978, 1990), Kirk et al. (2001), and Wehmeyer (1975). In
the most comprehensive molecular study to date, Castlebury et al. (2002) analyzed nLSU rDNA sequence data and
determined that there were six major lineages in the Diaporthales. Since then, an additional three lineages have
been added such that nine families are now included in the
order (Rossman et al. 2006). One of these families is named
herein, and two families are redefined. Despite this expanded phylogeny of the Diaporthales, many genera are
not affiliated with any of the lineages, and it is expected that
additional families will be uncovered with increased taxon
sampling. This was the case with the Sydowiellaceae. Initially, taxa now placed in the Sydowiellaceae were considered to be unaffiliated with any family but, with increased
sampling and the addition of more taxa, these species have
merged into a well-supported family (Rossman et al.
2006).
The phylogeny of each of the nine families currently
recognized in the Diaporthales is reviewed here based on
molecular data along with a summary of their biology. For
some families, extensive data exist, while for others very
little is known. Because of the polyphyletic nature of most
of the genera as defined based on morphology, emphasis is
placed on type species. At the end of the article, some genera in the Diaporthales are discussed for which data are insufficient to place them in any existing family.
Gnomoniaceae
Figs. 1– 4
Members of the Gnomoniaceae occur primarily on hardwood trees, although other hosts exist, such as conifers for
the anamorph genus Sirococcus. Several diseases are caused
by species in the Gnomoniaceae, as mentioned below, but
these species fruit primarily on overwintered leaves
(Stoykow 2005) and are also commonly isolated as endophytes (Kaneko and Kobayashi 1984; Barengo et al. 2000).
This family includes at least ten known teleomorphic
genera, namely, Apiognomonia-Discula, Apioplagiostoma,
Cryptodiaporthe, Cryptosporella-Disculina, Ditopella,
Gnomonia, Gnomoniella, Ophiovalsa, Phragmoporthe, and
Plagiostoma, and the anamorphic genus Sirococcus, for
which no teleomorph is known. Results of molecular studies
(Castlebury et al. 2002) generally agree with the concept of
the Gnomoniaceae as monographed by Monod (1983).
This finding differs significantly from other concepts of this
family based solely on morphology as proposed by Barr
(1978, 1990, 1991), Kobayashi (1970), and Vasilyeva
(1993).
The Gnomoniaceae is characterized by ascomata that
are immersed, solitary without a stroma, or aggregated in a
reduced prosenchymatous stroma in herbaceous plant material, especially in leaves or stems but also in wood. The
ascomata are generally soft-textured, thin-walled, and prosenchymatous with either central or lateral beaks. The asci
usually have a distinct apical ring. This family includes species having ascospores that are generally small, less than
25 µm long, although some are longer as in Cryptosporella,
and range in septation from nonseptate to one-septate or
multiseptate. The asexual states of members of the Gnomoniaceae are acervular or pycnidial with a broad opening.
Conidiogenous cells are phialidic, and conidia are usually
pallid and nonseptate (Monod 1983).
The genus Gnomonia is represented by the type species
G. gnomon (Tode) J. Schröt. (Sogonov et al. 2005). Species
of Gnomonia usually have beaked, thin-walled, solitary
perithecia immersed in the substrata. Recent data show that
Gnomonia may not be monophyletic (Sogonov et al. 2006).
Numerous additional species of Gnomonia have been described but cannot yet be placed in well-defined genera. The
genus Apiognomonia is distinguished from Gnomonia by
unequally septate ascospores. Recent molecular data show
that Apiognomonia errabunda and Plagiostoma euphorbiae
(Fuckel) Fuckel, type of the genus Plagiostoma, are allied
with a group that includes two species of Cryptodiaporthe
including the type species C. aesculi (Fuckel) Petr. and C.
salicella (Fr.) Petr. Monod (1983) placed P. euphorbiae in
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Gnomonia. Gnomoniella fraxinii and its asexual state Discula fraxinea (Redlin and Stack 1988) is a non-type species
of a genus similar to Gnomonia but having unicellular ascospores. Gnomonia also includes two morphologically
similar species on Corylus, Phragmoporthe conformis (Berk.
& Broome) Petr. with polysporous asci and three-septate
ascospores and Ditopella ditopa (Fr.) J. Schröt. with eightspored asci and one-septate ascospores. Monod (1983) recognized Ditopella ditopa in the genus Gnomonia.
The genus Cryptosporella is a well-defined genus within
the Gnomoniaceae, represented by the type species C.
hypodermia (Fr.) Sacc. (Castlebury et al. 2002). The type
species of Ophiovalsa, O. suffusa (Fr.) Petr., and a second
species, O. betulae (Tul. & C. Tul.) Petr., are congeneric
with C. hypodermia; thus, Ophiovalsa is considered a
synonym of Cryptosporella. The genera Cryptosporella
and Ophiovalsa were not included in Gnomoniaceae by
Monod (1983) or any other authors. The genus Winterella
has been confused nomenclaturally with Ophiovalsa (Reid
and Booth 1987) and includes a number of species that were
previously placed in the genus Cryptospora (Reid and
Booth 1989). In contrast to members of the Gnomoniaceae,
Cryptosporella is characterized by a distinctly valsoid
arrangement of ascomata. Cryptosporella is similar to
other members of the Gnomoniaceae in having stromatal
tissues that are prosenchymatous, nearly lacking discs, or
forming small ectostromatic discs that may be reduced to a
brown disc between the erumpent ostiolar cluster and
perithecia.
At least two groups of species in the Gnomoniaceae reproduce asexually and lack any known sexual state, as is the
case for most plant-associated fungi (Rossman 1993). The
cause of dogwood anthracnose, Discula destructiva, is such
a species for which a sexual state is unknown. Despite use
of molecular data, neither Zhang and Blackwell (2001) nor
Castlebury et al. (2002) were able to infer the sexual state
of D. destructiva. Its affinities to the Gnomoniaceae in the
Diaporthales were hypothesized by Redlin (1991) and confirmed by Zhang and Blackwell (2001) and Castlebury et al.
(2002). The anamorphic genus Sirococcus, typified by S.
conigenus (DC.) P.F. Cannon & Minter, includes only species that occur on conifers. No teleomorph is known for
these species, nor is one known for the noncongeneric species S. clavigignenti-juglandacearum, cause of butternut
canker in North America (Ostry et al. 1996).
Melanconidaceae
Figs. 5–8
At present the only genus represented in this family is the
type, namely, Melanconis, including the type species, M.
stilbostoma (Fr.) Tul. & C. Tul., and two other species,
M. alni Tul. & C. Tul. and M. marginalis (Peck) Wehm.
(Castlebury et al. 2002). The anamorphs of Melanconis are
placed in the genus Melanconium. The three species of
Melanconis are restricted to hardwood trees in the Betulaceae, on which they appear to be endophytic and weakly
pathogenic. This family is sister to the Gnomoniaceae, both
of which have prosenchymatous stromatic tissues. The
genus Melanconis sensu Wehmeyer (1941) included many
additional species, some of which have been studied but are
not affiliated with any families in the Diaporthales (L.A.
Castlebury, unpublished data). One of these has been segregated from Melanconis as Melanconiella spodiaea (Tul. &
C. Tul.) Sacc., type of the genus Melanconiella, whereas
Melanconis desmazieri Petr. appears to be affiliated close
to the Sydowiellaceae (L.A. Castlebury, unpublished
data).
Schizoparmeaceae Rossman, fam. nov.
Figs. 9–11
Ascomata fusca vel nigra, collapsa, erumpentia, superficialentia. Asci annulo apicali distincto praediti, ad maturitatem
separati. Paraphyses nullae. Ascosporae non septatae. Anomorphi pycnidiales in Coniella Höhn. & Pilidiella Petr. &
Syd.
Genus typicum: Schizoparme Shear
Ascomata brown to black, collapsed collabent, erumpent, becoming superficial. Asci with distinct apical ring,
floating free at maturity. Paraphyses lacking. Ascospores
nonseptate. Pycnidial anamorphs in Coniella Höhn. and
Pilidiella Petr. & Syd.
Type genus: Schizoparme Shear, Mycologia 15: 121,
1923.
This new family includes the distinctive teleomorph
genus Schizoparme and its asexual state Pilidiella and the
closely related anamorph genus Coniella. Samuels et al.
(1993) were the first to recognize the unique nature of
Schizoparme and its relationship to Coniella and Pilidiella.
These fungi were initially placed in the Melanconidaceae,
but Castlebury et al. (2002) determined that these species
constitute a distinct lineage within the Diaporthales, which
they referred to as the Schizoparme complex, as did van
Niekerk et al. (2004). Members of the Schizoparmeaceae
have fruiting bodies that are often erumpent through the
host epidermis, becoming superficial on living or decaying
herbaceous and woody plants. Members of this family produce diseases such as white rot of grapes caused by Pilidiella
diplodiella (Speg.) Crous & van Niekerk (Sutton and
Waterston 1966, as Coniella diplodiella) and leaf and fruit
diseases of strawberry caused by P. castaneicola (Ellis &
Everh.) Arx (Maas 1998). Although known in temperate
regions, species of the Schizoparmeaceae are most common
in tropical areas.
Both the ascomata of Schizoparme and conidiomata of
Pilidiella and Coniella are brown or black, occasionally
pallid yellowish brown or gray. They have a distinctive wall
layer that is irregularly thickened, often with plate-like ornamentation. They become collapsed cupulate upon drying,
erumpent through the substromatal surface at maturity.
The asci, which arise from the base of the fruiting body
(Samuels et al. 1993), are typically diaporthalean with a
distinct apical ring and float free in the centrum.
The ascospores are one-celled, initially hyaline, but may
become pale to dark brown, smooth, rarely with gelatinous
appendages. The type species of Schizoparme is S. straminea Shear having the anamorph Pilidiella castaneicola.
Seven species are included in Schizoparme (Samuels et al.
1993).
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Until recently, Pilidiella was considered a synonym of
Coniella (Sutton 1980; Nag Raj 1993); however, van Niekerk
et al. (2004) demonstrated that these two anamorph genera
should be distinct. Both produce one-celled conidia rarely
with a longitudinal slit. Pilidiella includes species with hyaline to pale brown conidia having a length:width ratio of
greater than 1.5, whereas conidia of Coniella are dark brown
and much narrower. The pycnidia of both Coniella and Pilidiella are relatively thick-walled, often hyaline. They produce pale brown to dark brown conidial masses in which
the conidiogenous cells develop from a basal pad described
as a “pulvinate basal parenchyma” by Sutton (1980) and
described in detail by Muthumary and Vaneja (1986) for
Coniella fragariae (Oudem.) B. Sutton. This structure is
morphologically similar to the basal pad from which the
ascogenous hyphae develop in the ascomatal state. Species
of Schizoparme have Pilidiella anamorphs; no teleomorph
is known for Coniella.
Cryphonectriaceae
Figs. 12–15
The Cryphonectriaceae was recently described to accommodate Cryphonectria, Endothia, and several segregate genera including Amphilogia, Chrysoporthe, and
Rostraureum (Gryzenhout et al. 2004, 2005a–c, 2006), a
group first recognized as the Cryphonectria-Endothia complex by Castlebury et al. (2002). All members of this family
have microscopic characteristics typical of the Diaporthales
with the addition of fruiting bodies with orange-reddish
stromatic tissues that turn purple in 3% KOH. The genus
Cryphonectria was recently conserved with a new type species, C. parasitica, thus ensuring that the name of this important fungus remains stable and serves as the basis for
this family (Gryzenhout et al. 2005a). The anamorph states
of members of the Cryphonectriaceae are pycnidial, often
with the same orange-reddish KOH+ purple pigments, phialidic conidiogenous cells, and small, hyaline, nonseptate
conidia, placed in the anamorph genera Endothiella and the
recently described Chrysoporthella. Additional species in
this family include Wuestneia xanthostroma (Mont.) J. Reid
& C. Booth, type of the genus Wuestneia (L.A. Castlebury,
unpublished data), and Cryptodiaporthe corni (Wehm.)
Petr., a non-type species (Redlin and Rossman 1991). Both
of these species have orange-reddish pigments in the
stroma.
The Cryphonectriaceae includes virulent pathogens such
as the well-characterized Cryphonectria parasitica, cause of
chestnut blight (Anagnostakis 1988; Milgroom and Cortesis
1999; Breullin et al. 2006). In addition, Chrysoporthe cubensis (Bruner) Gryzenh. & M.J. Wingf. and several related
species threaten eucalyptus trees throughout the world
Figs. 1–15. Representative species of four families in the Diaporthales.
1–4 Gnomoniaceae: Gnomonium gnomon. 1 Ascomata. 2 Section
through ascomata. 3 Asci. 4 Ascospores. 5–8 Melanconidaceae:
Melanconis stilbostoma. 5 Ascomata. 6 Section through ascomata. 7
Asci. 8 Ascospores. 9–11 Schizoparmeaceae: Schizoparme straminea.
9, 10 Ascomata. 11 Section through ascomata. 12–15 Cryphonectria-
(Gryzenhout et al. 2004, 2005d). Many members of this
family occur primarily on woody plants as saprobes, e.g.,
Endothia gyrosa (Schwein.) Fr., Cryphonectria nitschkei
(G.H. Otth) M.E. Barr, C. macrospora (Tak. Kobay. &
Kaz.-Ito) M.E. Barr, and C. radicalis (Schwein.) M.E. Barr
(Hoegger et al. 2002; Myburg et al. 2004) and are often observed in their Endothiella pycnidial states on the exposed
roots of hardwood trees.
Valsaceae
Figs. 16–19
The Valsaceae is restricted to the genus Valsa and its segregates, including Valsella and Leucostoma, based on Castlebury et al. (2002). Previously, Barr (1978) had recognized
the Valsaceae to include members of the Diaporthaceae
such as Diaporthe as well as Plagiostoma and Cryptodiaporthe, now placed in the Gnomoniaceae, and Cryphonectria, now placed in the Cryphonectriaceae.
The genera Leucostoma, Valsa, and Valsella have been
long recognized as closely related genera (Spielman 1985;
Vasilyeva 1998; Castlebury et al. 2002), and the distinction
between them is not clear. Traditionally they are separated
based on the characteristics of having eight-spored asci
(Leucostoma and Valsa) or polysporous asci (Valsella), with
Leucostoma having a white to grayish-brown, ectostromatic
disc, but this latter character appears to vary with specimen
age. All three genera have anamorphs referred to as Cytospora. In Castlebury et al. (2002), the genus Valsa was represented by its type species V. ambiens (Pers.) Fr., which
occurs on woody angiosperms in temperate regions throughout the world (Spielman 1985), and several other species.
In the phylogenetic tree presented by Castlebury et al.
(2002), the two species of Valsella, including the type species V. salicis Fuckel and V. adherans Fuckel, were intermingled with species of Valsa and Leucostoma such as L.
cincta (Fr.) Höhn. and L. nivea (Hoffm.) Höhn. However,
the type species L. massarina (De Not.) Höhn. was not included in this study. The genus Leucostroma is considered
a synonym of Valsa by Vasilyeva (1998).
Members of Leucostoma, Valsa, and Valsella occur on
hardwoods and occasionally on conifers throughout the
world, often in their Cytospora anamorphic states. Species
of Valsa on hardwoods in North America were delineated
by Spielman (1985); three species of Valsa were reported
from the eastern Himalayas by Dargan and Sharma (1991).
Members of the Valsaceae cause canker diseases such as
Cytospora canker or gummosis of peach and stone fruits
caused by Leucostoma persoonii (Nitschke) Höhn. (Hayova
and Minter 1998a) and other species of Leucostoma as described by Adams et al. (2002). Species of Valsa including
V. ambiens, V. ceratosperma (Tode) Maire, V. cypri (Tul.)
ceae: Cryphonectria parasitica. 12 Ascomata. 13 Section through ascomata. 14 Asci. 15 Ascospores. 1–4 BPI 844237; 5, 6 BPI 748234; 8 BPI
872036; 9, 11 BPI 600008; 10 BPI 600023; 12, 13 BPI 749121; 14, 15 BPI
748234. Bars 1, 2, 5, 6, 9, 10, 11, 13 100 µm; 12 400 µm; 3, 4, 7, 8, 14, 15
10 µm
139
140
Figs. 16–27. Representative species of three families in the Diaporthales. 16–19 Valsaceae: Valsa ambiens. 16 Ascomata. 17 Section
through ascomata. 18 Asci. 19 Ascospores. 20–23 Diaporthaceae:
Diaporthe eres. 20 Ascomata. 21 Section through ascomata. 22 Asci.
23 Ascospores. 24–27 Sydowiellaceae: Sydowiella fenestrans. 24 Ascomata. 25 Section through ascoma. 26 Asci. 27 Ascospores. 16–19 BPI
843602; 20, 21 BPI 841335; 22, 23 BPI 872076; 24–27 BPI 843503. Bars
16 200 µm; 17, 20, 21, 24, 25 100 µm; 18, 19, 22, 23, 26, 27 10 µm
Tul. & C. Tul., V. eugeniae Nutman & F.M. Roberts, V.
malicola Z. Urb., V. salicina (Pers.) Fr., and V. sordida
Nitschke cause canker diseases of hardwood trees, especially those that are stressed or damaged (Sivanesan and
Holliday 1970; Hayova and Minter 1998b–h). Adams et al.
(2005) described 27 species of Valsa and Cytospora causing
diseases on Eucalyptus in Australia and South Africa.
Diaporthaceae
Figs. 20–23
The Diaporthaceae consists of the very large genus
Diaporthe with 801 named taxa and its Phomopsis anamorph with more than 900 described species (Uecker 1988).
Based on Castlebury et al. (2002), this family includes only
Diaporthe-Phomopsis and the genus Mazzantia based on
141
the non-type species M. napelli (Ces.) Sacc. Wehmeyer
(1933) recognized the relationship of Mazzantia to Diaporthe, especially as the anamorph Mazzantiella having “a
stroma similar to the ascospore state with pycnidial locule
containing comma-shaped conidia on filiform conidiophores . . . similar to the Phomopsis state of the genus Diaporthe.” At least one segregate genus, Diaporthopsis, on the
basis of the presence of nonseptate ascospores, was determined to belong within Diaporthe based on a study of the
type species, Diaporthopsis angelicae (Berk.) Wehm. (Castlebury et al. 2003). The genus Allantoporthe, based on the
type species A. tessella (Pers.) Petr., was established for
species similar to Diaporthe but having a short narrow appendage at the ends of the ascospores. Wehmeyer (1933)
suggested that the presence of faint hyaline appendages was
not a constant character. Barr (1978) segregated Allantoporthe from Diaporthe based on the prosenchymatous stromatic tissues in Allantoporthe. Molecular data suggest that
both A. tessella and A. decedens (Fr.) M.E. Barr belong in
Diaporthe sensu lato (L.A. Castlebury, unpublished data).
Relationships within this genus and family have yet to be
determined.
The proliferation of species names, especially in Phomopsis, has resulted from the assumption that these fungi
are host specific. Some species of Phomopsis appear to be
host-specific pathogens such as Phomopsis amygdali
(Delacr.) J.J. Tuset & M.T. Portilla, cause of peach and almond canker (Farr et al. 1999; Kanematsu et al. 1999, 2000),
although this species has recently been isolated from grape
(van Niekerk et al. 2005). Other recently described, hostspecific species of Phomopsis include P. gossypii (Sacc.)
Palmateer et al., cause of boll rot of cotton (Palmateer et
al. 2003), P. vaccinii Shear et al., cause of blueberry and
cranberry canker (Farr et al. 2002a), P. viticola (Sacc.) Sacc.
on grapes (Schilder et al. 2005), and P. columnaris Farr. &
Castl. on lingonberry (Farr et al. 2002b). Recent studies
suggest that many species of Phomopsis occur on a variety
of plant host genera (Rehner and Uecker 1994) or, conversely, the same plant host may harbor several species of
Phomopsis as exemplified by the six different species of
Diaporthe-Phomopsis on Vitis vinifera (Phillips 1999; Kajitani and Kanematsu 2000; Mostert et al. 2001; van Niekerk
et al. 2005).
Many plant diseases are caused by species of Diaporthe
(Wehmeyer 1933) and Phomopsis (Uecker 1988), some of
which have already been mentioned. The taxa causing diseases on soybean are difficult to separate, with at least four
different taxa involved (Fernandez and Hanlin 1996).
Northern soybean stem canker caused by D. phaseolorum
(Cooke & Ellis) Sacc. var. caulivora Athow & Cauldwell
occurs on hosts in addition to soybean (Black et al. 1996),
as does D. ambigua Nitschke, reported to cause a disease
on rootstocks of rosaceous fruit trees (Smit et al. 1996) but
also known on grapes (van Niekerk et al. 2005). For many
disease-causing species of Phomopsis, host specificity is not
known, as, for example, P. limonii I.C. Harv. et al. 2000
(non Vegh 1994), cause of stem canker on the cut-flower
hybrid Limonium sp. (Harvey et al. 2000), Diaporthe foeniculacea Niessl on fennel (Phillips 2003), and D. actinidiae
N.F. Sommer & Beraha, cause of stem-end rot of kiwifruit
(Lee et al. 2001).
Phomopsis is the most prevalent endophytic fungus isolated from both tropical and temperate woody plants
(Fisher et al. 1994, 1995; Shamoun and Sieber 2000; Tomita
2003) and was present in the sapwood of almost all angiospermous trees examined (Boddy and Griffith 1989). Their
potential role in protecting plants from fungal diseases such
as Dutch elm disease has been explored (Brayford 1990).
At least one endophytic Phomopsis from a woody tropical
tree is known to produce toxins that affect the central nervous system in vertebrates (Bills et al. 1992), suggesting an
adaptive advantage to plants that harbor these fungi.
Little is known about the mating systems in the
Diaporthe-Phomopsis complex; however, Linders and van
der Aa (1995) demonstrated that D. adunca (Roberge ex
Desm.) Niessl was heterothallic with two mating types.
Fertilization occurs in the fall when the alpha-conidia that
function as both fertilizing agents and infective agents
were splashed onto adjacent substrata, resulting in crossfertilization and development of the Diaporthe sexual state
the following spring (Linders and van der Aa 1995).
Pseudovalsaceae
The Pseudovalsaceae was established by Barr (1978) in a
broad sense to include members of the Diaporthales having
upright, erumpent perithecia with central beaks. Based on
Castlebury et al. (2002), many members of Pseudovalsaceae
sensu Barr belong elsewhere, such as Ditopella and Phragmoporthe, now placed in the Gnomoniaceae, Allantoporthe,
now placed in the Diaporthaceae, and Chapeckia and Sydowiella, now placed in the Sydowiellaceae (L.A. Castlebury, unpublished data). In fact, the only genus included in
the Pseudovalsaceae at present is Pseudovalsa based on the
type species P. lanciformis (Fr.) Ces. & de Not. and related
species P. longipes (Tul.) Sacc., P. modonia (Tul. & C. Tul.)
Höhn., and P. umbonata (Tul. & C. Tul.) Sacc., all species
that have a Coryneum anamorph and occur on temperate
hardwood trees in the Fagales.
Sydowiellaceae
Figs. 24–27
The Sydowiellaceae is based on Sydowiella, type species S.
fenestrans (Duby) Petr., with an assortment of genera and
species that do not have any clear features in common (L.A.
Castlebury, unpublished data). These taxa occur on herbaceous, dicotyledonous plants as well as hardwood trees.
Genera in this family include the type species of Chapeckia,
C. nigrospora (Peck) M.E. Barr, Hapalocystis, H. berkeleyi
Fuckel with the closely related H. occidentalis Jakl. &
Voglmayr (Jaklitsch and Voglmayr 2004), Rossmania, R.
ukurunduense Lar. N. Vassiljeva (Vasilyeva 2001), Stegophora, S. ulmea (Fr.) Syd. & P. Syd., and Sillia, S. ferruginea
(Pers.) P. Karst. In addition, the Sydowiellaceae includes
species previously placed in Gnomonia, such as G. rostellata
(Fr.) Brefeld on herbaceous plants, a species potentially bet-
142
ter placed in Sydowiella. The biology of the Sydowiellaceae
is also quite diverse, ranging from parasites of living leaves
such as Stegophora ulmea on Ulmus americana and saprobes
on herbaceous plants to Hapalocystis berkeleyi forming stromatic fruiting bodies on decaying woody plants.
Togniniaceae
The Togniniaceae was established by Reblova et al. (2004)
for species of Togninia and their Phaeoacremonium anamorphs that were removed from the Calosphaeriales and
recognized in the Diaporthales along with the genus Jobellisia (Reblova et al. 2004; Mostert et al. 2006). This conclusion was based on small subunit (SSU) rDNA; however,
analyses based on large subunit (LSU) rDNA are ambiguous about placement of this family (Mostert et al. 2006).
The Togniniaceae and the Calosphaeriales are unique in
having asci that develop in fascicles from short proliferating
ascogenous hyphae and are morphologically quite unlike
the Diaporthales. The phylogenetic data presented by Mostert et al. (2006) also suggest that Jobellisia may be allied
with the Diaporthales. Except for the presence of a conspicuous ring in the ascal apex, this genus does not have
characteristics of the Diaporthales, i.e., asci that float free
and paraphyses lacking at maturity; rather, the long-stalked
asci of Jobellisia remain attached among abundant straight
paraphyses.
The genus Togninia, especially as the hyphomycetous
anamorph Phaeoacremonium, is involved in two serious
diseases of grapevine (esca and Petri disease) and other
wilting and dieback diseases of plants (Mostert et al. 2003).
These fungi also are opportunists that infect humans, causing phaeohyphomycosis. Ten species of Togninia and 22
species of Phaeoacremonium were monographed by Mostert et al. (2006) in a publication that also includes a review
of the diseases caused by these fungi.
Diaporthalean fungi of unknown affiliation
Several genera in the Diaporthales either are not affiliated
with any family or have not been examined using molecular
sequences. Only a few of the most important taxa are mentioned here.
The anamorphic genus Harknessia belongs in the Diaporthales with several species grouping together but outside
of any recognized family (Castlebury et al. 2002). The teleomorph genus Wuestneia is linked to species of Harknessia
(Crous et al. 1993; Crous and Rogers 2001); however, the
type species of Wuestneia, W. xanthostroma (Mont.) J. Reid
& C. Booth, with distinctive orange-red pigments, has affinities with members of the Cryphonectriaceae, whereas
the remaining species of Wuestneia and species of Harknessia form a group outside this family (Lee et al. 2004). The
similar-looking anamorph genus Apoharknessia lacks a
known teleomorph and is phylogenetically distinct from
Harknessia (Lee et al. 2004) but also is not allied with any
known family in the Diaporthales. Species of Harknessia
and Apoharknessia are associated with diseases of primarily
tropical plants, especially Eucalyptus (Yuan and Mohammed 1997; Yuan et al. 2000) and Proteaceae (Crous et al.
1993; Lee et al. 2004). The anamorph genus Dwiroopa with
the non-type species D. lythri (D.F. Farr & Rossman) D.F.
Farr & Rossman, initially described as Harknessia lythri
D.F. Farr & Rossman, also could not be placed in any
known family within the Diaporthales (Farr and Rossman
2001, 2003). The cause of bitter rot of grape, Greeneria
uvicola, was determined to belong in the Diaporthales, but
it is not affiliated with any recognized family, nor is any teleomorph known for this anamorph species (Farr et al.
2001).
Hercospora is a distinct genus in that the ostioles from
individual fruiting bodies converge within the stroma and
emerge as one ostiole. Hercospora tiliae (Pers.) Tul. & C.
Tul., with its unusual anamorph, Rabenhorstia, groups with
Melanconis desmazieri, also on Tilia, and is allied with the
Sydowiellaceae but falls outside this family (L.A. Castlebury, unpublished data).
The genera Lollipopaia and Phruensis on Licuala (Arecaceae) are described as members of the Diaporthales with
unknown affinities, both having elongate ascospores and
the latter producing a phialophora-like anamorph (Inderbitzin and Berbee 2001; Pinruan et al. 2004). Phruensis appears morphologically similar to Melogramma in the
Melogrammataceae, Phyllachorales. The genus Vismaya
was described from rotting wood in Hong Kong and appears to be a member of the Gnomoniaceae, but it lacks a
distinct apical ring (Sarma and Hyde 2001). Without a culture, it will be difficult to determine the affinities of this
genus.
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