Description of Gibberella sacchari and neotypification of ... - Mycologia

Mycologia, 97(3), 2005, pp. 718–724.
2005 by The Mycological Society of America, Lawrence, KS 66044-8897
Description of Gibberella sacchari and neotypification of its anamorph
Fusarium sacchari
John F. Leslie 1
Department of Plant Pathology, Throckmorton Plant
Sciences Center, Kansas State University, Manhattan,
Kansas 66506-5502
Brett A. Summerell
Suzanne Bullock
Royal Botanic Gardens and Domain Trust, Mrs.
Macquaries Road, Sydney, New South Wales 2000,
Australia
Frank J. Doe
Department of Biology, University of Dallas, Irving,
Texas 75061
Abstract: We described the teleomorph of Fusarium
sacchari as Gibberella sacchari, sp. nov. This species
can be separated from other species of Gibberella on
the basis of the longer, narrower ascospores found in
G. sacchari and by sexual cross fertility. Female-fertile
mating type tester strains were developed that can be
used for making sexual crosses with this heterothallic
fungus under laboratory conditions. The anamorph,
Fusarium sacchari, was neotypified.
Key words: biological species, Fusarium neoceras,
Fusarium subglutinans sensu lato, Gibberella fujikuroi
mating population B, maize, sorghum, sugarcane
INTRODUCTION
Nine mating populations or biological species, denoted
by the letters A–I, have been identified within
the Gibberella fujikuroi species complex (Britz et al
2002, Samuels et al 2001, Zeller et al 2003). The anamorphs
of all nine have unique Latin binomials as
do eight of the nine teleomorphs. Our objective in
this report is to provide a name and formal description
for the teleomorph of the ninth biological species
and to describe the mating-type tester strains that
are commonly used for identification purposes. To
clarify the nomenclature and taxonomy of this taxon
we neotypify the anamorph, Fusarium sacchari.
Accepted for publication 6 Jan 2005.
1 Corresponding author. Department of Plant Pathology, 4002
Throckmorton Plant Sciences Center, Kansas State University, Manhattan,
Kansas 66506-5502; Phone: 785-532-1363; Fax: 785-532-
2414; E-mail: jfl@plantpath.ksu.edu
718
MATERIALS AND METHODS
Cultures and culturing conditions.—Cultures examined were
recovered from sugarcane and sorghum and either provided
to us or collected by us as indicated in the list below.
Strain numbers from other collections are given where
known, and are: ATCC—American Type Culture Collection,
Manassas, Virginia; BBA—Biologische Bundesanstalt für
Land- und Forstwirtschaft, Berlin, Germany; FGSC—Fungal
Genetics Stock Center, University of Missouri-Kansas City,
Kansas City, Missouri; FRC—Pennsylvania State University,
University Park, Pennsylvania; SNS—University of California,
Davis, California; KSU—Kansas State University, Manhattan,
Kansas; PTS—University of California, Berkeley, California.
FGSC and KSU numbers are used preferentially.
When isolates were recovered from plant material, they
were plated onto semiselective peptone-PCNB media (Nash
and Snyder 1962). After 4–7 d incubation at 25 C, we selected
Fusarium colonies that emerged from this material.
We purified all cultures, including those received from other
investigators, by subculturing single conidia separated by
micromanipulation. Isolates were cultured routinely on carnation
leaf agar (CLA) (Fisher et al 1982), or on modified
Czapek’s complete medium (Correll et al 1987).
All strains are maintained as spore suspensions in 15%
glycerol at 70 C. We have deposited a dried culture of
FGSC 7610 FGSC 7611, representing G. sacchari with the
National Fungus Collection, Beltsville, Maryland, as BPI
843974. Female-fertile mating-type tester strains (strain
numbers KSU B-03853 [ FGSC 7611] and KSU B-03852
[ FGSC 7610]) were deposited as isotypes with the Fungal
Genetics Stock Center.
We used this standard Gibberella fujikuroi mating population
mating-type tester strains in all mating tests (G. fujikuroi
mating population, Gibberella teleomorph, FGSC strain
numbers and mating types): G. fujikuroi mating population
A, Gibberella moniliformis, FGSC 7600 (MATA-1), FGSC
7603 (MATA-2); G. fujikuroi mating population B, Gibberella
sacchari, FGSC 7611 (MATB-1), FGSC 7610 (MATB-2);
G. fujikuroi mating population C, Gibberella fujikuroi, FGSC
8931 (MATC-1), FGSC 8932 (MATC-2); G. fujikuroi mating
population D, Gibberella intermedia, FGSC 7615 (MATD-1),
FGSC 7614 (MATD-2); G. fujikuroi mating population E,
Gibberella subglutinans, FGSC 7616 (MATE-1), FGSC 7617
(MATE-2); G. fujikuroi mating population F, Gibberella thapsina,
FGSC 7057 (MATF-1), FGSC 7056 (MATF-2); G. fujikuroi
mating population G, Gibberella nygamai, FGSC 8934
(MATG-1), FGSC 8933 (MATG-2); G. fujikuroi mating population
H, Gibberella circinata, FGSC 9022 (MATH-1), FGSC
9023 (MATH-2); G. fujikuroi mating population I, Gibberella
konza, FGSC 8910 (MATI-1), and FGSC 8911 (MATI-2).

Microscopy.—The description of Gibberella sacchari is based
on crosses (FGSC 7610 FGSC 7611) produced on carrot
agar. Perithecia were treated with 3% KOH and 100% lactic
acid to observe any color reaction, and measured in situ.
Asci and ascospores were mounted in water for measurement
and photography. Measurements were taken of 20
each of perithecia, asci and ascospores. Whole perithecia
were fixed in 6.5% glutaraldehyde in 100 mM sodium cacodylate
buffer at pH 7.6 for 4 h at room temperature (Bullock
et al 1980), dehydrated in a graded ethanol series and
infiltrated and embedded in LR White resin. Sections 1.5
m thick were cut with a Reichert ultramicrotome, dried
onto poly-L-lysine-coated glass slides and stained with 0.5%
toluidine blue O for 10 s (Feder and O’Brien 1968). Sections
were mounted in immersion oil. Morphological features
of the anamorph (F. sacchari) were photographed in
situ on carnation leaf agar. Macro- and microconidia were
mounted in water on glass slides and photographed with
DIC optics.
Mating-type specific PCR and crossing procedures.—We identified
mating-type idiomorphs (MAT-1 or MAT-2) for the F.
sacchari isolates with PCR-based assays, as described in either
Kerényi et al (1999) or Steenkamp et al (2000), and
in crosses to the female-fertile tester isolates (FGSC 7610
MAT-2 or FGSC 7611 MAT-1) developed as a part of this
study. We made crosses on carrot agar as described in Klittich
and Leslie (1988).
RESULTS
The initial mating-type testers used for this species
were the field isolates FGSC 7608 and FGSC 7609.
These strains are both highly fertile but are subject
to strain degeneration, in which a tough, dense, mycelium
with fingerlike projections is produced, with
a thinner, wispier mycelium between the denser mycelial
fingers. If these colonies are subcultured, then
the amount of dense mycelia continues to increase
at the expense of the thinner mycelium. The thinner
mycelium eventually disappears and only the denser
mycelium can be found. Spores from such colonies
have an apparently normal ability to serve as a male
parent in a sexual cross. When used as a female parent,
however, the only portion of the colony on which
perithecia form is that with the thinner mycelium.
Once the strain produces only the denser mycelium,
the colony no longer is useable as a female parent.
The frequency with which the dense sectors appear
is irregular, but once the process has begun we have
not been able to recover a colony that has only the
thin phenotype. Instead we begin a new culture from
the frozen stock and discard the degenerating culture.
We crossed FGSC 7608 and FGSC 7609 and collected
78 random ascospore progeny. We used these
progeny as the female parents in crosses with the pa-
LESLIE ET AL: DESCRIPTION OF G. SACCHARI
719
rental strains and identified strains that repeatedly
produced, at least qualitatively, a large number of mature
perithecia. Mating type in all of the progeny was
identified on the basis of these crosses. We also identified
progeny that retained exclusively the thin phenotype
and did not produce sectors with the dense
mycelial phenotype in at least three successive subcultures.
We selected two progeny (FGSC 7610 and
FGSC 7611) based on these criteria and have been
using them as the standard testers for this mating
population since 1990. We have never seen these testers
develop the dense, female-sterile mycelial sectors
that can occur in their parents and have not noticed
any appreciable reduction in female fertility of these
strains over this time.
Members of the B mating population are distinct
from other mating populations in the G. fujikuroi
species complex in that they do not cross with the
testers from any of the other eight described mating
populations. Strains listed are cross-fertile with either
FGSC 7610 or FGSC 7611 but not with testers for the
other known biological species in the G. fujikuroi species
complex. In most cases no perithecia are made
in the crosses between strains from the different mating
populations. On rare occasions a few perithecia
may be produced, presumably due to homothallism
such as that observed with these testers and a few
other strains of mating population B (Britz et al
1999). Thus any crosses in which only a few perithecia
are produced on a plate must be viewed as suspect
until the progeny produced are shown to have a biparental
origin.
TAXONOMY
Gibberella sacchari Summerell et Leslie sp. nov.
FIGS. 1–7
Perithecia superficialia, livida, 270–390 m alta, 250–390
m diam. Asci fusiformes, dehiscentes, octospori. Ascosporae
exudatae in cirrhis, laeves, hyalinae, ellipsoideae vel
obovoideae, 0–1 septatae, plerumque 1-septatae et ad septum
leviter constrictae, 3–4.5 7.0–8.0 m. Anamorphosis:
Fusarium sacchari (E.J. Butler) W. Gams.
HOLOTY PUS. Cultura exsiccata in agaro ex FGSC 7610
FGSC 7611.
Etymology.—Taken from the anamorph, referring to
the original host plant, sugarcane.
Teleomorph.—Perithecia superficial, solitary to aggregated
in groups of a few and seated on a minute
stromatic base, obovoidal and warty (FIGS. 1–2); 270–
390 (mean 325) m, 250–390 (mean 310) m
diam; blue-black, color not changing in 3% KOH,
turning red in 100% lactic acid. Perithecial wall 23–
39 (mean 30.6) m thick laterally, formed of two

720 MYCOLOGIA
FIGS. 1–7. Gibberella sacchari, characteristics of perithecia, asci and ascospores produced from a cross of strains FGSC
7610 FGSC 7611 on carrot agar. 1–2. Perithecia. 3. Transverse section of a perithecium stained with toluidine blue O. 4.
Detail of transverse section of perithecial wall. 5–6. Asci. 7. Ascospores. Bars: 1 and 2 200 m; 3 and 5 50 m; 4, 6 and
7 25 m.
obvious regions (FIGS. 3–4). Outer wall region 18–33
(mean 25.6) m thick; outer cells angular to
elliptic in transverse section, 5.3–12 (mean 8.4)
m length 3.1–5.9 (mean 4.3) m thick, with
the largest cells at the exterior and the smallest cells
toward the interior of the wall, walls of cells 0.5–1.2
(mean 0.9) m thick and pigmented. Inner wall
region 4.0–5.9 (mean 4.9) m thick; cells angular
to elliptic, 7.2–16 (mean 10.6) m length
1.1–3.2 (mean 2.2) m thick. Inner wall cells fu-

siform, thin-walled and irregular, walls of inner cells
0.3–0.6 (mean 0.4) m thick and pigmented (FIG.
4). Outer and the inner wall regions merging imperceptibly;
cells of the outer region more pigmented.
Perithecial apex continuous with the outer and inner
wall layers; cells at the apex smaller appearing as a
reticulum; cells forming the ostiolar opening clavate
and thick-walled at the cell tips; nonpigmented
merging periphyses. Cells of the apex attaining the
same length to form an apical disk.
Asci fusiform (FIGS. 5–6), regularly dehiscing during
slide preparation and 8-spored. Ascospores exuded
in a cirrus (FIGS. 1–2), ellipsoidal to obovoid
with both ends rounded, 0–1-septate and slightly constricted
at the septum, 7.0–8.0 3–4.5 (mean 7.6
4.2) m (FIGS. 5–7). Heterothallic species, reproductively
isolated from previously described species of
Gibberella.
Isolates examined.—BRAZIL: culture isolated from sorghum
by Shirley Nash Smith and received from J. Puhalla and P.T.
Spieth 1984, MAT-2, KSU B-00205 (PTS F-1178, SNS
Br19a). CHINA: Taiwan, Hsingying; cultures isolated from
sugarcane plants by Shirley Nash Smith in 1981 and received
from J. Puhalla and P.T. Spieth 1984, MAT-1, KSU
B-00281 (ATCC 201263, FGSC 7609, FRC M3128, SNS HY-
7, PTS F-1254); MAT-2, KSU B-00278 (ATCC 201262, FGSC
7608, FRC M3127, SNS HY-4, PTS F-1251). EL SALVADOR:
La Libertad; cultures isolated from diseased (stalk rot) sorghum
plants by L.E. Claflin 2002; all MAT-1, KSU B-12751,
KSU B-12752, KSU B-12753, KSU B-12754, KSU B-12755 &
KSU B-12756. GERMANY: from Cattleya and received from
H. Nirenberg via P.E. Nelson; MAT-2 KSU B-03828 (FRC
1217, BBA 62260). INDIA: all from sugarcane and received
from H. Nirenberg via P.E. Nelson; MAT-1, KSU B-03819
(FRC M941, BBA 63340), KSU B-03820 (FRC M942, BBA
63342); MAT-2, KSU B-03821 (FRC M943, BBA 63448).
MEXICO: Guadalajara; cultures isolated from diseased
(stalk rot) maize plants by L.E. Claflin 2002; MAT-1, KSU
B-12747; MAT-2, KSU B-12748, KSU B-12749 & KSU B-
12750. PHILIPPINES: Pioneer Overseas Corporation Research
Station, Banbic, Calruyao, Laguna; all cultures isolated
by J.F. Leslie 1988; from diseased sorghum leaf, MAT-
1, KSU B-01722 (FRC M5476), KSU B-01724 (FRC M5478),
KSU B-01725 (FRC M5479), KSU B-01726 (FRC M5480);
from diseased sorghum stalk, MAT-2, KSU B-01721 (FRC
M5467); from sorghum seed, MAT-1, KSU B-01730 (FRC
M5484), KSU B-01732 (FRC M5486), KSU B-01735 (FRC
M5489); MAT-2, KSU B-01727 (FRC M5481), KSU B-01728
(FRC M5482). USA: Kansas, Manhattan; cultures recovered
from a laboratory cross of FGSC 7608 (KSU B-00278) and
FGSC 7609 (KSU B-00281) by J.F. Leslie 1989; MAT-1 KSU
B-03853 (ISOTY PE, ATCC 201265, FRC M6866, FGSC
7611); MAT-2 KSU B-03852 (ISOTY PE, ATCC 201264, FRC
M6865, FGSC 7610).
NEOTY PIFICATION OF FUSARIUM SACCHARI
Fusarium sacchari (E.J. Butler) W. Gams first was described
as Cephalosporium sacchari Butler from sug-
LESLIE ET AL: DESCRIPTION OF G. SACCHARI
721
arcane in India in 1913 (Butler and Khan 1913) but
did not include a description of macroconidia. Cultures
with macroconidia were described by Wollenweber
and Reinking (1925) as Fusarium neoceras Wollenweber
& Reinking. Gams (1971) synonymized the
two names to the present form. Because the original
type specimen no longer is available we have chosen
isolate KSU B-03852 (ATCC 201264, FGSC 7610, FRC
M6865) to neotypify the anamorph. Because this isolate
is a single ascospore isolate from two parents isolated
from sugarcane, has been shown to be conspecific
with isolates KSU B-03819 (FRC M941, BBA
63340), KSU B-03820 (FRC M942, BBA 63342); KSU
B-03821 (FRC M943, BBA 63448) from sugarcane in
India by sequencing and AFLPs (data not shown)
and is one of the two parental strains used to create
the teleomorph we have chosen it to neotypify.
Neotype, designated here Fusarium sacchari (E.J.
Butler) W. Gams. KSU B-03852 (ATCC 201264, FRC
M6865, FGSC 7610). A dried culture of this strain is
deposited as DAR 76835 at the Plant Pathology Herbarium,
Orange Agricultural Institute, Department
of Primary Industries, New South Wales, Australia.
Anamorph features (FIGS. 8–13) are similar to
those described by Nirenberg (1976) and Gerlach
and Nirenberg (1982). Key features are 3–4-septate
macroconidia (FIGS. 8–9) and oval microconidia
(FIGS. 10–11) borne in false heads on monophialides
and polyphialides (FIGS. 12–13).
DISCUSSION
Fusarium sacchari (E.J. Butler) W. Gams first was described
as Cephalosporium sacchari Butler from sugarcane
in India (Butler and Khan 1913). The original
description did not include a description of macroconidia,
which is not surprising given the scarcity of
macroconidia formed by this species in in vitro culture.
Wollenweber and Reinking (1925) described
cultures with macroconidia as Fusarium neoceras Wollenweber
& Reinking. Gams (1971) synonymized the
two names to the present form. Kuhlman (1982) described
four varieties of G. fujikuroi—moniliformis,
subglutinans, fujikuroi and intermedia. He used the
name G. fujikuroi var. subglutinans for the teleomorph
of F. sacchari because these strains at that time
were identified as F. subglutinans. The name Gibberella
subglutinans now is associated with the former
Gibberella fujikuroi mating population E (Samuels et
al 2001), which reflects its initial usage for a fungal
pathogen of maize (Edwards 1935). Thus we selected
sacchari as the species epithet to maintain similarity
between the anamorphic and teleomorphic names.
Relative to other Gibberella species, the ascospores
of G. sacchari are large. In this study the sizes that

722 MYCOLOGIA
FIGS. 8–13. Characteristics of Fusarium sacchari. 8–9. Macroconidia. 10–11. Oval to obovoid microconidia. 12–13. Microconidia
on monophialides and polyphialides produced in carnation leaf agar cultures. Bars 25 m.
we measured were 28–32 3.0–4.5 (mean 30.4
4.2) m, which is somewhat shorter and narrower
than the values observed by Kuhlman (1982) (16–32
4–8 [mean 22.4 5.6] m) wide. Kuhlman (1982)
found the mean values for the ascospore sizes to be
significantly different from the means for G. moniliformis
(17.5 4.8 m), G. fujikuroi (12.5 4.7 m),
and G. intermedia (14.6 4.8 m). Our observations
are consistent with his conclusions. In addition G.
sacchari probably can be distinguished from G. nygamai
(13.9 5.3 m), G. subglutinans (19.6 5.9
m), G. thapsina (17.0 6.0 m) and G. circinata
(12.7 5.2 m) on the basis of the generally longer
and narrower ascospores found in G. sacchari. This
trait could be especially useful for distinguishing G.
sacchari from other species in Fusarium subglutinans
sensu lato, which generally are similar morphologically.
With respect to perithecial diameter, the values we
obtained (250–390 m [mean 310 m]) are somewhat
narrower in the range but similar in the mean
(240–420 m [mean 307 m]) to those reported by
Kuhlman (1982). Perithecial diameter could be used
to differentiate perithecia of G. sacchari from those
of G. fujikuroi (mean perithecial diameter 231 m),
G. thapsina (250 m), and G. intermedia (389 m)
but not from those of G. moniliformis (321 m) or
G. nygamai (309 m) and probably not from those
of G. subglutinans (336 m) or G. circinata (337 m)
(Klassen and Nelson 1996, Klittich et al 1997, Kuhlman
1982). The comparative measurements of these
different species must be evaluated with caution however
because the perithecia and ascospores they contain
were grown on several different substrates (e.g.
carrot agar, V-8 juice agar and carnation leaf agar)
and differences in nutrition could alter the values
observed. The degree of maturity of both the peri-
thecia and the spores that they contain similarly
could alter the observed sizes of the asci, ascospores
and perithecia. Other described Fusarium species to
which F. sacchari shows morphological similarity and
is closely related (e.g. F. acutatum, F. bulbicola, F. concentricum,
F. denticulatum, F. guttiforme, F. pseudocircinatum,
F. pseudonygamai and F. ramigenum [Nirenberg
and O’Donnell 1998]), have as yet no reported
sexual stage, and the utility of ascospore size or perithecial
diameter as distinguishing characters for
these species remains unknown.
Representative isolates now assigned to G. sacchari
often have been included in studies of variation in
physiological traits or molecular characters. Differences
that separate G. sacchari from the other members
of the G. fujikuroi species complex include sensitivity
to hygromycin and benomyl (Yan et al 1993),
polymorphism in isozyme banding patterns (Huss et
al 1996), chromosome length (Xu et al 1995), AFLP
banding pattern (Marasas et al 2001) and DNA sequences
of representative genes (O’Donnell et al
1998). The only unique isozyme band for G. saccahara
is a pattern observed for esterase (Huss et al
1996). However the pattern for glucose-4-phosphate
isomerase is found only in G. moniliformis and
G. sacchari and these two species can be distinguished
easily on the basis of the pattern for triose
phosphate isomerase, which is unique to G. moniliformis.
O’Donnell et al (1998) concluded that G. sacchari
was related most closely to F. fujikuroi, F. proliferatum,
F. globosum and F. concentricum based on the
sequence of the -tubulin, mitochondrial small subunit
rDNA, and 28S rDNA genes. Species to which
G. sacchari is more similar morphologically (e.g. G.
subglutinans and G. circinatum) were not closely related
to G. sacchari when these DNA sequences were
used to estimate relatedness. Marasas et al (2001)

showed that G. sacchari was distinct from F. andiyazi,
F. nygamai, F. pseudocircinatum, F. ramigenum, F. thapsinum
and F. verticillioides on the basis of AFLP fingerprinting
patterns. Strains of G. sacchari are not
known to produce fumonisins (Leslie et al 1992).
The strains examined for this study were from Taiwan,
the Philippines, Germany (greenhouse), India,
Mexico and El Salvador, but other strains have been
reported from Brazil, India, Malaysia, South Africa
and Thailand (Leslie 1995). In crosses with the tester
strains, however, many of the isolates not included in
this study were reported to produce relatively few
perithecia, and thus their identity as G. sacchari
should be regarded as tentative due to the potential
problems associated with selfing by the testers. The
host range similarly should be considered limited to
maize, sugarcane, orchids and sorghum until the isolates
reported from banana, peanut and rice have
been examined more closely.
Identification of G. sacchari can be accomplished
in many ways. The size of the ascospores and the fertility
of the strain in crosses with standard tester isolates
are the technically simplest means of distinguishing
G. sacchari from other sibling species in the
former F. subglutinans sensu lato. The species also can
be distinguished based on AFLP patterns, isozymes
and by differences in the DNA sequence of several
commonly studied genes.
Gibberella sacchari has yet to be reported under
field conditions, but the anamorph, F. sacchari, has
been recovered from a diverse, although small, set of
host plants. The fungus often has been associated
with Fusarium sett and stem rot of sugarcane (Egan
et al 1997); however no comprehensive studies of
host range or pathogenicity have been conducted.
Given the recent increase in our knowledge of the
taxonomy of this fungus and its close relatives, such
studies would be both interesting and important.
ACKNOWLEDGMENTS
This research was supported in part by the Kansas Agricultural
Experiment Station and by the Sorghum and Millet
Collaborative Research Support Program (INTSORMIL)
AID/DAN-1254-G-00-0021-00 from the U.S. Agency for International
Development. We thank Larry E. Claflin, John
Puhalla, Philip T. Spieth and the late Paul E. Nelson for
providing some of the strains used in this study. JFL thanks
the Australian-American Fulbright Commission and The
Royal Botanic Gardens and Domain Trust, Sydney, for the
award of fellowships to support his sabbatical leave in Australia.
Contribution No. 03-20-J from the Kansas Agricultural
Experiment Station, Manhattan.
LESLIE ET AL: DESCRIPTION OF G. SACCHARI
LITERATURE CITED
723
Britz H, Coutinho TA, Wingfield MJ, Marasas WFO. 2002.
Emended description of Gibberella circinata, the cause
of pitch canker of pines. Sydowia 54:9–22.
, , , , Gordon TA, Leslie JF. 1999.
Fusarium subglutinans f. sp. pini represents a distinct
mating population in the Gibberella fujikuroi species
complex. Appl Environ Microbiol 65:1198–1201.
Bullock S, Willetts HJ, Ashford AE. 1980. The structure and
histochemistry of sclerotia of Sclerotinia minor Jagger 1.
Light and electron microscope studies on sclerotial development.
Protoplasma 104:315–331.
Butler EJ, Khan AH. 1913. Some new sugarcane diseases.
Mem Dept Ag in India, Bot Ser 6:185–190.
Correll JC, Klittich CJR, Leslie JF. 1987. Nitrate non-utilizing
mutants of Fusarium oxysporum and their use in vegetative
compatibility tests. Phytopathology 77:1640–
1646.
Edwards ET. 1935. Studies on Gibberella fujikuroi var. subglutinans
the hitherto undescribed ascigerous stage of
Fusarium moniliforme var. subglutinans and its pathogenicity
on maize in New South Wales. Dept. Agric.
New South Wales Sci Bull 49:1–68.
Egan BT, Magarey RC, Croft BJ. 1997. Sugarcane. In: Hillocks
RJ, Walker JM, eds. Soilborne diseases of tropical
crops. CAB International: Wallingford, Oxon, U.K. p
277–302.
Feder N, O’Brien TP. 1968. Plant microtechnique: some
principles and new methods. Am J Bot 55:123–142.
Fisher NL, Burgess LW, Toussoun TA, Nelson PE. 1982. Carnation
leaves as a substrate and for preserving cultures
of Fusarium species. Phytopathology 72:151–153.
Gams W. 1971. Cephalosporium-artige Schimmelpilze (Hyphomycetes).
Stuttgart, Germany: Gustav Fischer Verlag.
Gerlach W, Nirenberg HI. 1982. The genus Fusarium—a
pictorial atlas. Mitt Bio Bundesanstalt für Land- u
Forstw (Berlin-Dahlem) 209:1–406.
Huss MJ, Campbell CL, Jennings DB, Leslie JF. 1996. Isozyme
variation among biological species in the Gibberella
fujikuroi species complex (Fusarium section Liseola).
Appl Environ Microbiol 62:3750–3756.
Kerényi Z, Zeller KA, Hornok L, Leslie JF. 1999. Molecular
standardization of mating type terminology in the Gibberella
fujikuroi species complex. Appl Environ Microbiol
65:4071–4076.
Klaasen JA, Nelson PE. 1996. Identification of a mating population,
Gibberella nygamai sp. nov., within the Fusarium
nygamai anamorph. Mycologia 88:965–969.
Klittich CJR, Leslie JF. 1988. Nitrate reduction mutants of
Fusarium moniliforme (Gibberella fujikuroi). Genetics
118:417–423.
, , Nelson PE, Marasas WFO. 1997. Fusarium
thapsinum (Gibberella thapsina): a new species in section
Liseola from sorghum. Mycologia 89:643–652.
Kuhlman EG. 1982. Varieties of Gibberella fujikuroi with anamorphs
in Fusarium section Liseola. Mycologia 74:
759–768.
Leslie JF. 1995. Gibberella fujikuroi: Available populations

724 MYCOLOGIA
and variable traits. Can J Bot 73(Supplement 1):S282–
S291.
, Plattner RD, Desjardins AE, Klittich CJR. 1992. Fumonisin
B 1 production by strains from different mating
populations of Gibberella fujikuroi (Fusarium section
Liseola). Phytopathology 82:341–345.
Marasas WFO, Rheeder JP, Lamprecht SC, Zeller KA, Leslie
JF. 2001. Fusarium andiyazi sp. nov., a new species from
sorghum. Mycologia 93:1203–1210.
Nash SM, Snyder WC. 1962. Quantitative estimations by
plate counts of propagules of the bean root rot Fusarium
in field soils. Phytopathology 52:567–572.
Nirenberg HI. 1976. Untersuchungen über die morphologische
und biologische Differenzierung in der Fusarium-Sektion
Liseola. Mitt Biolog Bundesanstalt für
Land-u Forstw (Berlin-Dahlem) 169:1–117.
, O’Donnell K. 1998. New Fusarium species and combinations
within the G. fujikuroi species complex. Mycologia
90:434–458.
O’Donnell K, Cigelnik E, Nirenberg HI. 1998. Molecular
systematics and phylogeography of the Gibberella fujikuroi
species complex. Mycologia 90:465–493.
Samuels GJ, Nirenberg HI, Seifert KA. 2001. Perithecial spe-
cies of Fusarium. In: Summerell BA, Leslie JF, Backhouse
D, Bryden WL, Burgess LW, eds. Fusarium: Paul
E. Nelson memorial symposium. St Paul, Minnesota:
APS Press. p 1–14.
Steenkamp ET, Wingfield BD, Coutinho TA, Zeller KA,
Wingfield MJ, Marasas WFO, Leslie JF. 2000. PCR-based
identification of MAT-1 and MAT-2 in the Gibberella
fujikuroi species complex. Appl Environ Microb 66:
4378–4382.
Wollenweber HW, Reinking AO. (1925). Aliquot Fusaria tropicalia,
nova vel revisa. Phytopathology 15:155–169.
Xu J-R, Yan K, Dickman MB, Leslie JF. 1995. Electrophoretic
karyotypes distinguish the biological species of Gibberella
fujikuroi (Fusarium section Liseola). Molec Plant-
Microbe Interact 8:74–84.
Yan K, Dickman MB, Xu J-R, Leslie JF. 1993. Sensitivity of
field strains of Gibberella fujikuroi (Fusarium section
Liseola) to benomyl and hygromycin B. Mycologia 85:
206–213.
Zeller KA, Summerell BA, Bullock S, Leslie JF. 2003. Gibberella
konza (Fusarium konzum) sp. nov. from prairie
grasses, a new species in the Gibberella fujikuroi species
complex. Mycologia 95:943–954.