Plant Health Progress ¿ 2019 ¿ 20:255–260 https://doi.org/10.

1094/PHP-08-19-0052-DG

Diagnostic Guide

Black Rot of Sweetpotato: A Comprehensive Diagnostic Guide

Madison N. Stahr and Lina M. Quesada-Ocampo†
Department of Entomology and Plant Pathology, North Carolina State University, Raleigh, NC 27695-7613

Accepted for publication 24 October 2019.

Keywords: mycology, Ipomoea batatas, Ceratocystis fimbriata, pathogenicity testing

Disease: Black rot. wilting, leaf chlorosis, and leaf drop (Clark and Moyer 2013). If the
vine of an infected plant is pulled from the field prior to harvest, a
Hosts: Sweetpotato (Ipomoea batatas), morning glory (Ipomoea lesion on the vine below soil level may be visible (Fig. 1). Active
sp.) (de Beer et al. 2014), and carrot (Daucus carota subsp. sativus) lesions will appear brown and necrotic (Fig. 1A), and black, globose
(Stahr and Quesada-Ocampo, unpublished). perithecia with long necks should be observed protruding from the
lesion surface (Fig. 1B). As the lesion matures, the infected tissue
Pathogen: Black rot of sweetpotato is caused by the fungal will become increasingly more necrotic and can cause the sweet-
pathogen Ceratocystis fimbriata (Ellis and Halst.) (Halsted 1890). potato vine to split open (Fig. 1C). Although such splitting has been
Although this is the original name of the pathogen upon first observed in greenhouse trials (Stahr and Quesada-Ocampo, un-
publication, multiple synonyms were introduced over time before published), there has been no record of such advanced lesions in the
returning to the original name. Formerly used synonyms for C. field.
fimbriata include Ceratostomella fimbriata (J.A. Elliot), Endoco- Postharvest, black rot of sweetpotato causes a hard rot in the
nidiophora fimbriata (R.W. Davidson), Ophiostoma fimbriata sweetpotato storage roots (Clark and Moyer 2013). C. fimbriata is
(Nannf.), and Spaeronaema fimbriatum (Sacc.) (Clark et al. 2015; able to infect sweetpotatoes through healthy skin tissue, but in-
Webster and Butler 1967). fection preferentially takes place at lateral roots, lenticels, and
wound sites (Lauritzen 1926; Stahr and Quesada-Ocampo 2020).
Infected sweetpotatoes will exhibit shallow, circular lesions on the
Taxonomy
Kingdom fungi, phylum Ascomycota, subphylum Pezizomy-
cotina, class Sordariomycetes, subclass Hypocreomycetidae, order
Microascales, family Ceratocystidaceae, genus Ceratocystis, spe-
cies fimbriata. Current taxonomy can be found at MycoBank,
www.mycobank.org.
The name C. fimbriata has been used to describe fungi capable of
infecting many different plants in multiple families. However,
thorough research into the genetic relationship of fungi within the
Ceratocystis genus and closely related genera has called for the
name C. fimbriata to be restricted to isolates derived from
sweetpotato or from the same phylogenetic species (de Beer et al.
2014; Harrington 2013).

Symptoms and Signs

Due to the soilborne nature of causal agent C. fimbriata,
sweetpotato black rot starts in the field. However, during the
growing season symptoms of black rot are not often observed,
except intermittent lesions seen on storage roots at harvest if in-
fected seed is used and/or the field had a strong disease pressure.
According to the Compendium of Sweetpotato Diseases, Pests, and
Disorders, reported field symptoms (not pictured) include stunting,
†
Corresponding author: L. M. Quesada-Ocampo; lmquesad@ncsu.edu FIGURE 1
Appearance of stem lesions on sweetpotato vines. A, Active lesions will
Funding: This work was supported by the North Carolina Sweetpotato Commission appear brown to black in color on the sweetpotato vine. B, If black perithecia
and the United States Department of Agriculture (USDA) - Agricultural Research
Service under project number NC02628. are observed, the globose base will be partially embedded in the vine tissue
with the long necks of the perithecia protruding from the vine surface. C,
The author(s) declare no conflict of interest. Advanced lesions can cause the infected part of the vine to become
completely necrotic and ultimately split open, but this has not yet been
© 2019 The American Phytopathological Society documented outside of greenhouse trials.

PLANT HEALTH PROGRESS ¿ 2019, Vol. 20, No. 4 ¿ Page 255

surface of the root (Fig. 2A). Lesion color can range from brown, dark Although the host list is extensive, host specialization among
gray, greenish-black, to black depending on the sweetpotato cultivar isolates has been reported (Baker et al. 2003). As research into the
and surface moisture (Clark and Moyer 2013). When conditions are Ceratocystis genus continues and genomic tools advance, cryptic
favorable for sporulation, light gray mycelia will grow on the lesion species in the C. fimbriata complex are being reclassified as new
surface, typically originating in the lesion center or at a wound site. species. For example, previously classified C. fimbriata isolates
Perithecia development will follow mycelial growth, and light- from sycamore (Platanus spp.), cacao (Theobroma cacao), and
colored ascospore masses may be visible to the naked eye. How- almond (Prunus sp.) have now being reclassified as C. platani,
ever, diseased roots are often found after conditions are no longer C. cacaofunesta, and C. destructans, respectively (Engelbrecht and
favorable for sporulation or the sweetpotato has been washed on a Harrington 2005; Holland et al. 2019).
packing line, which will cause the gray mycelia to subside and leave
behind small black patches, which are remnants of sporulation
structures (Fig. 2B). If the storage root is cut open, the tissue below a Geographic Distribution
lesion will be black and necrotic (Fig. 2C), but the infection does not C. fimbriata has a large geographic range and has been docu-
often internalize beyond the vascular tissue of the storage root (Clark mented on all cultivated continents. More specifically, C. fimbriata
and Moyer 2013). Finally, the fungus will produce a fruity odor, can be found in all places where sweetpotatoes are cultivated,
which is observed in both infected roots and when the fungus is including Asia (Li et al. 2016), Africa (Roux et al. 2000), and the
grown in culture (Clark and Moyer 2013; Soares et al. 2000). Americas (Baker et al. 2003). Within the United States, C. fimbriata
is most prevalent within the Southeast, and has been reported in the
following states: North Carolina, Virginia, Georgia, Mississippi,
Host Range Alabama, Louisiana, Arkansas, Missouri, and California (Scruggs
C. fimbriata is known to infect many plant species in addition to et al. 2017). A summary of geographic distribution can be found in
sweetpotato, commonly producing canker and wilt diseases on the Fungal Databases at the Systematic Mycology and Microbi-
woody hosts. Hosts of C. fimbriata include, but are not limited to, ology Laboratory, USDA-ARS, which includes geographic distri-
coffee (Coffea sp.) (Baker et al. 2003; Johnson et al. 2005), Eu- bution by country (https://nt.ars-grin.gov/fungaldatabases/). A detailed
calyptus spp. (Roux et al. 2000), fig (Ficus carica) (Harrington et al. distribution map can also be found at https://www.cabi.org/dmpd,
2011), taro (Colocasia esculenta) (Johnson et al. 2005; Mizukami but it is not as current (1983).
1951), hickory (Carya sp.) (Johnson et al. 2005), mango (Man-
gifera indica) (Baker et al. 2003; Harrington et al. 2011), and
Pathogen Isolation
pomegranate (Punica granatum) (Li et al. 2016). Furthermore,
in greenhouse settings C. fimbriata has been observed to infect C. fimbriata is a slow-growing fungus that can take at least
tobacco (Nicotiana tabacum), soybean (Glycine max), and lettuce 2 weeks to grow large enough to fully cover a standard 100-mm-
(Lactuca sativa) when plants were inoculated after wounding or diameter agar plate. As such, steps to reduce the risk of contam-
at the seedling stage (Stahr and Quesada-Ocampo, unpublished). ination from faster growing organisms should be taken when
A summary of hosts can be found online in the Fungal Databases isolating C. fimbriata from any host tissue. Carrot baits and carrot
at the Systematic Mycology and Microbiology Laboratory, agar (described in Figure 3 and Table 1, respectively) are considered
USDA-ARS (https://nt.ars-grin.gov/fungaldatabases/fungushost/ selective for C. fimbriata (Moller and DeVay 1968) and as such are
fungushost.cfm). recommended for isolation, but they do allow for the growth of
other microorganisms. Currently, no other selective or differential
media for C. fimbriata exist, although potato dextrose agar and malt
yeast extract agar (Table 1) are also commonly used for C. fimbriata
propagation. Water agar should not be used for isolation or
propagation of C. fimbriata because the fungus will not grow on this
medium. Finally, C. fimbriata does not grow within sweetpotato

FIGURE 2
Diagnostic symptoms of sweetpotato black rot caused by Ceratocystis fim-
briata. A, Dark and circular lesions form on the surface of the storage roots, FIGURE 3
usually centered around a wound, lenticel, or lateral root. B, Small black Use of carrot disc baiting for isolation of suspected Ceratocystis fimbriata
patches that are remnants of sporulation structures can often be observed in isolates. A, Infected tissue should be placed in an internal cavity cut into one
the lesion (arrows) and typically appear in the lesion center. C, Internally half of the carrot discs. B, The discs are reassembled and placed in a humidity
infected tissue will become black and necrotic, but necrosis does not typ- chamber with damp paper towels to prevent the discs from drying out. C and
ically extend beyond the vascular tissue. D, Fungal mycelia will grow out of the carrot disc after 4 to 5 days.

PLANT HEALTH PROGRESS ¿ 2019, Vol. 20, No. 4 ¿ Page 256

tissue when kept at temperatures below 18°C (Stahr and Quesada- free of soil, submerge the symptomatic area into a 0.6% sodium
Ocampo 2020). Therefore, isolation from plant material that has hypochlorite solution and surface sterilize as previously described.
been stored in a refrigerator or cold room for extended periods of If the stem is large and sturdy, cut a small, triangular tissue sample
time will have a lower success rate when compared with isolation from the edge of the lesion as previously described. If the stem is too
from a sample stored at room temperature. small or fragile to cut a triangular tissue sample, simply make a thin
Isolation from storage roots. Storage roots should be surfaced cross section of the stem near the outer edge of the lesion. Plate
sterilized by submersion in a 0.6% sodium hypochlorite solution for triangular tissue sample or stem cross section onto amended carrot
5 min. Roots should then be thoroughly rinsed with sterile distilled agar and proceed with the rest of the isolation as previously described.
water (SDW) and allowed to dry in a laminar flow hood. Using Isolation with carrot baits. If isolation onto agar is un-
sterile technique, small triangular tissue samples should be cut from successful, carrot baits can be used to attempt isolation. Carrot baits
the edge of the suspected black rot lesion using a flame-sterilized are also common when attempting isolation from non-sweetpotato
scalpel. Tissue samples are then to be plated onto carrot agar amended hosts and possible insect vectors (da Silva et al. 2017; Harrington
with ampicillin and rifampicin (20 mg/liter each), with the inner side of et al. 2011; Moller and DeVay 1968). To use a carrot bait, acquire a
the tissue facing down on the plate. Agar plates should be wrapped whole carrot and surface sterilize with sodium hypochlorite as done
with Parafilm and incubated at room temperature (;23°C), with an in prior isolation steps. Cut the carrot into 2-inch segments, and then
approximate 12-h photoperiod. Very limited growth will be visible half the segments into disks. On the interior surface of one disk, cut
within the first 3 to 4 days; if prolific growth is observed during this out a cavity large enough to contain a sample of infected tissue,
period, it is most likely due to other fungal contaminants. After 5 to insect, or whatever infected sample is being used (Fig. 3A). Under
7 days, the fungal colony should be large enough in size that the initial
sterile conditions, collect the infected tissue sample (using either of
hyaline mycelia will have turned greenish-black. Being careful to avoid
the methods described above), and then place the infected tissue
any other organisms that may be present, cut a 3 × 3-mm agar plug
into the carrot cavity. Reassemble the carrot discs and place in a
from the edge of the mycelial colony and transfer to a fresh plate.
humidity chamber or deep-dish Petri plate (25 × 100 mm) with a
Transfer of the colony, in this manner, should be repeated until no damp paper towel (Fig. 3B). The baits should then be placed into an
secondary organisms are present. Once free of secondary organisms, incubator set to 25°C. A 12-h photoperiod can be set, but C.
allow the fungi to grow until perithecia producing tan-orange asco- fimbriata can successfully grow in the absence of light (Webster
spore masses appear (;7 to 10 days). Because C. fimbriata is ho- and Butler 1967). Incubate the carrot discs 5 to 10 days, while
mothallic (Clark and Moyer 2013), singular ascospore masses should regularly checking moisture levels to ensure the carrot discs do not
be of a single genotype. To obtain an axenic, single genotype culture, dry out or become waterlogged. After 4 to 5 days, mycelial growth
use a flame-sterilized scalpel, inoculation loop, or inoculation needle to
grab a singular ascospore mass and transfer to a new plate. The as-
cospore masses are very sticky in nature and will adhere to a
transfer tool when lightly touched.
Isolation from plant stems. Stem isolations are generally less
successful than storage root isolations, because stem lesions in
sweetpotato are less common, can quickly become necrotic, and often
house saprophytic microbes. If an infection is suspected, gently
remove the whole plant, including roots, and rinse free of any soil.
Avoid cutting into or breaking the plant around the stem lesion,
because it provides an entry site for sanitizer during surface steril-
ization, which may result in senescence of the causal organism. Once

TABLE 1
Recipes for media commonly used for Ceratocystis fimbriata
isolation and propagation
Mediuma Per liter
Carrot agar
Bolthouse Farms carrot juice 160 ml
Agar powder 16 g
Carrot broth
Bolthouse Farms carrot juice, clarified via 100 ml
centrifugation
Malt yeast extract agar
Malt extract (2%) 20 g
Yeast extract (0.2%) 2g
FIGURE 4
Agar (2%) 20 g
Morphological features used for identification of Ceratocystis fimbriata. A,
Potato dextrose agar
Appearance of culture in carrot agar. B, Sexual ascospores given charac-
Potato dextrose agar powder 39 g
teristic hat shape due to a gelatinous sheath forming a brim (arrow). C,
a
Antibiotics can be added to any listed medium to control for bacterial Asexual spore types include cylindrical endoconidia and brown, globose
contaminants. Rifampicin and ampicillin at 20 mg/liter each are chlamydospores. D, Black perithecia with globose base and long tapering
recommended. neck. Scale bars have been included for size reference.

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should be evident (Fig. 3C and D), and once perithecia have been (Fig. 4C) are also used for identification as subhyaline to hyaline,
produced the ascospore mass can be collected and plated to obtain a nonseptate, cylindrical endoconidia are typically produced in
pure culture as previously described. abundance on most substrates. Thick-walled, chlamydospores, also
known as aleurioconidia (Engelbrecht and Harrington 2005), can
also be produced. Chlamydospores are oval to globose in shape and
Pathogen Identification range from pale to olive brown in color (Morgan-Jones 1967).
Morphological characteristics are commonly used to confirm Certain strains of C. fimbriata are also known to produce doliform
pathogen identity of suspected C. fimbriata isolates. In agar me- endoconidia, but such a spore type has not been reported in
dium, cultures of C. fimbriata can range in color from black to sweetpotato-derived isolates (Webster and Butler 1967). Di-
brown to dark-olive green (Fig. 4A) (Morgan-Jones 1967; Webster mensions of perithecia and all spore types can be found in Table 2.
and Butler 1967). Perithecia readily form on infected plant tissue Molecular confirmation for C. fimbriata should be used when
and common agar media. These perithecia can be superficial or possible. DNA can be extracted from a pure sample, and the internal
partially embedded in the growing substrate, are dark brown or transcribed spacer region and the translation elongation factor 1a
black in color, and have a globose base with a long tapering neck region of the genome can be amplified and sequenced. Primer
from which hyaline, ostiolar hyphae protrude (Fig. 4D) (Clark and names and sequences for these regions can be found in Table 3. The
Moyer 2013; Engelbrecht and Harrington 2005). Asci are not resulting sequences can then be compared against the full C.
visible, but single-celled hyaline ascospores are exuded in an ex- fimbriata genome (GenBank accession APWK00000000) using
tracellular matrix and form tan (Stahr and Quesada-Ocampo 2020), BLAST (Scruggs et al. 2017; Wilken et al. 2013). At this time,
pink (Clark and Moyer 2013), cream (Clark and Moyer 2013; species-specific primers are not available.
Engelbrecht and Harrington 2005), or orange (Stahr and Quesada-
Ocampo 2020) colored masses depending on isolate and substrate.
These smooth, elliptical ascospores (Fig. 4B) are key for identi- Pathogen Storage
fication because they possess a gelatinous sheath adhered to one There are multiple options for maintaining cultures of C. fim-
side of the spore, leading to their characteristic hat shape (Clark briata for long periods of time. First, because the fungus is slow
and Moyer 2013; Webster and Butler 1967). Asexual spore types growing (Webster and Butler 1967) and does not appear to lose

TABLE 2
Description of key morphological features used for identification of Ceratocystis fimbriata
Structure Colora Component Dimensionsa
Perithecia Dark brown to black Base Width: 110 to 250 mm
Height: 120 to 250 mm
Neck Base width: 28 to 40 mm
Top width: 16 to 24 mm
Length: Up to 900 mm
Hyaline to light brown Ostiolar hyphae Width: 2.2 mm
Number present: 7 to 16
Ascospores Hyaline Spore Width: 3.5 to 5 mm
Height: 3.0 to 4.5 mm
Length: 5 to 7.5 mm
Brimb Tapering width: 0.5 to 0.1 mm
Endoconidia Hyaline to pale brown Endoconidiophore Width: 3 to 8 mm
Length: 35 to 172 mm
Hyaline to subhyaline Spore Width: 3 to 7 mm
Length: 7 to 37 mm
Chlamydospores (aleurioconidiac) Pale to olive brown Conidiophore Width: 4 to 7 mm
(aleurioconidiophorec) Length: 10 to 76 mm
Spore Width: 6 to 16 mm
Length: 8 to 20 mm
a
All color and dimensions have been compiled from multiple publications (Clark and Moyer 2013; Engelbrecht and Harrington 2005; Morgan-Jones 1967;
Webster and Butler 1967). The largest published range in size for each structure component is shown.
b
Measurements taken with an electron microscope (Webster and Butler 1967).
c
Alternate nomenclature as used in Engelbrecht and Harrington (2005).

TABLE 3
Primer sequences to be used for molecular identification of Ceratocystis fimbriata
Region amplified Reference Primer name Primer sequence
Internal transcribed spacer region White et al. (1990) ITS1-F 59-CTTGGTCATTTAGAGGAAGTAA-39
ITS4 59-TCCTCCGCTTATTGATATGC-39
Translation elongation factor 1-a Geiser et al. (2004) ef1 59-ATGGGTAAGGARGACAAGAC-39
ef2 59-GGARGTACCAGTSATCATGTT-39

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virulence after serial transfer through artificial media compared recollected from the plate, the concentration determined using a
with other fungi such as Botrytis spp. (Breen et al. 2016; Butt et al. hemocytometer, and the suspension diluted with SDW to a
2006), C. fimbriata can be continually transferred through agar working concentration of 5 × 103 spores/ml (Stahr and Quesada-
every 10 to 14 days for many months. However, if keeping and Ocampo 2020). If mixed spore suspensions are undesirable due
repeatedly transferring plate cultures is undesirable, mycelial plugs to the difficulty of separating ascospore masses, primarily
can be prepared for short- and long-term storage. endoconidial suspensions can be prepared by flooding 7-day-old
To prepare storage cultures, first procure carrot broth (Table 1), a C. fimbriata cultures, grown on malt yeast extract or potato
50% glycerol solution, and a mature C. fimbriata plate culture. dextrose agar, with 10 ml of SDW and scraping with a rubber
Using sterile technique, cut five small (<5 mm) mycelial plugs from spatula. The spore suspension can then be filtered through four
the edge of the C. fimbriata culture. Place the mycelial plugs into a layers of sterile cheesecloth and rinsed with an additional 5 ml
1.5-ml tube along with 500 ml of carrot broth. Loosely close the tube of SDW. The spore suspension concentration should then be
to maintain limited airflow and allow an adjustment period of 3 to determined using a hemocytometer and diluted to a working
5 days. If contamination is a concern, the adjustment period can be concentration of 2 × 105 spores/ml (Baker et al. 2003).
shorter, but recovery of storage isolates may be less successful. Currently, there are no known resistant sweetpotato cultivars to C.
After the adjustment period, add 500 ml of 50% glycerol solution to fimbriata (Scruggs et al. 2017), so commercially available varieties
the tube and gently shake or vortex to mix. Place the tube on ice for such as Covington in North Carolina or Beauregard in Louisiana can be
1 h before transitioning to a 4°C fridge for short-term storage (<1 used for pathogenicity testing. If unfamiliar with disease signs and
year) or a –80°C freezer for long-term storage (>1 year). When symptoms, varieties with lighter skin colors are recommended over
recovering stored isolates, take steps to gradually warm the isolate purple-skinned varieties, because lesions will be easier to observe with
before attempting plating. First, remove the tube from the 4 or the naked eye. Pathogenicity should be tested on the storage roots,
–80°C and thaw on ice (for –80°C). Once thawed, hold the tube at which can be acquired from a local market or packing house. Roots
room temperature for 1 to 2 h, or until the room temperature is should be surface sterilized as described previously, to prevent other
reached. Finally, place the tube into a warm water bath, no hotter common and faster growing pathogens such as Rhizopus or Fusarium
than 40°C, for an additional 1 to 2 h. Plate mycelial plugs onto from skewing the results. Common to many postharvest pathogens, C.
room-temperature amended carrot agar plates. Growth should be fimbriata will preferentially infect at wound sites (Stahr and Quesada-
observed within 3 to 5 days. Ocampo 2020); thus, storage roots can be wounded to promote
infection. If not wounding roots, it is recommended to perform
Pathogenicity Tests pathogenicity tests with a larger sample size because infection through
Inoculum for pathogenicity tests can be prepared in two ways, healthy skin tissue is less common (Lauritzen 1926; Stahr and
depending on desired spore type. Mixed spore suspensions Quesada-Ocampo 2020). If wounding, wounds can be made with
containing endoconidia, chlamydospores, and the highly in- either a toothpick to produce a small puncture wound or a handheld
fectious ascospores can be obtained from 10-day-old C. fim- wounding tool (Collins et al. 2018) to produce three evenly spaced
briata cultures grown on carrot agar by adding a 5% Tween-20 punctures on the sweetpotato (Fig. 5). Roots, wounded or not, should
solution to the plate. Once 1 to 2 ml of the Tween solution is then be soaked in either the mixed spore suspension or the endoconidia
added, spores can be gently dislodged using a rubber scraper suspension for 20 min with periodic gentle agitation. After soaking,
or spatula and carefully massaged against the agar to separate place roots into sterile plastic containers and maintain a relative hu-
the ascospore masses. The spore suspension should then be midity of 80% or greater. Methods to maintain relative humidity
within the plastic containers will vary depending on container di-
mensions and number of roots per container but can typically be
managed by adding small Petri plates containing SDW or damp paper
towels to the containers (Scruggs and Quesada-Ocampo 2016). Store
the containers in a dark growth chamber or incubator at 29°C for a
maximum 3-week period. If the isolate tested is pathogenic, lesion
development and/or sporulation will be observed within 14 days of
inoculation, with lesion development occurring as early as 4 days
post-inoculation and sporulation as early as 7 days postinoculation. A
variety of measurements can be taken to determine the virulence and
aggressiveness of the isolate, including incidence of lesion devel-
opment and sporulation, and lesion and sporulation area diameters
taken at the end of a 3-week period or taken periodically every 2 to
4 days (Stahr and Quesada-Ocampo 2020).

Acknowledgments
The authors thank all the members of the Quesada lab for their valuable help.

FIGURE 5 Literature Cited

Optional wound sites to be used for pathogenicity testing. A, Small singular
Baker, C. J., Harrington, T. C., Krauss, U., and Alfenas, A. C. 2003. Genetic
puncture wound produced by a sterile toothpick. B, Slightly larger but more
variability and host specialization in the Latin American clade of Ceratocystis
shallow set of puncture wounds produced by C, a handheld wounding tool. fimbriata. Phytopathology 93:1274-1284.
This tool was assembled from a piece of stainless steel, a drawer handle, and Breen, J., Mur, L., Sivakumaran, A., Akinyemi, A., Wilkinson, M., and
three 4-mm self-drilling screws spaced 3.8 cm apart. Rodriguez Lopez, C. M. 2016. Botrytis cinerea loss and restoration of virulence

PLANT HEALTH PROGRESS ¿ 2019, Vol. 20, No. 4 ¿ Page 259

 during in vitro culture follows flux in global DNA methylation. bioRxiv. doi: Lauritzen, J. I. 1926. Infection and temperature relations of black rot of sweet
10.1101/059477 potatoes in storage. J. Agric. Res. 33:663-676.
Butt, T. M., Wang, C., Shah, F. A., and Hall, R. 2006. Degeneration of Li, Q., Harrington, T. C., McNew, D., Li, J., Huang, Q., and Somasekhara,
entomogenous fungi. Pages 213-226 in: An Ecological and Societal Approach Y. M. 2016. Genetic bottlenecks for two populations of Ceratocystis
to Biological Control. J. Eilenberg and H. M. T. Hokkanen, eds. Springer, fimbriata on sweet potato and pomegranate in China. Plant Dis. 100:
Dordrecht, the Netherlands. 2266-2274.
Clark, C. A., Ferrin, D. M., Smith, T. P., and Holmes, G. J. 2015. Diseases of Mizukami, T. 1951. Comparison of the pathogenicity of Ceratostomella
sweetpotato. Common Names of Plant Diseases. American Phytopatho- fimbriata and Endoconidiophora sp. causal agent of taro-blackrot,
logical Society, St. Paul, MN. https://www.apsnet.org/edcenter/resources/ on sweetpotatoes and taroes. Sci. Bull. Faculty Agric. Kyushu Univ.
commonnames/Pages/Sweetpotato.aspx 12:5-9.
Clark, C. A., and Moyer, J. W. 2013. Black rot. Pages 29-33 in: Compendium of Moller, W. J., and DeVay, J. E. 1968. Carrot as a species-selective isolation
Sweetpotato Diseases, Pests, and Disorders. C. A.Clark, D. M. Ferrin, T. P. medium for Ceratocystis fimbriata. Phytopathology 58:123-124.
Smith, and G. J. Holmes, eds. APS Press, St. Paul, MN. Morgan-Jones, G. 1967. Ceratocystis fimbriata. IMI Descriptions of Fungi
Collins, H., Adams, M. L., and Quesada-Ocampo, L. M. 2018. Evaluation of and Bacteria. No. 15, Sheet 141. CABI Publishing, Wallingford, U.K.
fungicides for postharvest control of black rot in sweetpotato, 2017. Plant Dis. Roux, J., Wingfield, M. J., Bouillet, J.-P., Wingfield, B. D., and Alfenas, A. C.
Manage. Rep. 12:V150. 2000. A serious new wilt disease of Eucalyptus caused by Ceratocystis
da Silva, A. C., Cândido, T. S., Sales, N. L. P., Harrington, T. C., and Alfenas, fimbriata in Central Africa. For. Pathol. 30:175-184.
A. C. 2017. First report of Ceratocystis wilt caused by Ceratocystis fimbriata Scruggs, A. C., Basaiah, T., Adams, M. L., and Quesada-Ocampo, L. M. 2017.
on Caryocar brasiliense trees in Brazil. Plant Dis. 101:1822. Genetic diversity, fungicide sensitivity, and host resistance to Ceratocystis
de Beer, Z. W., Duong, T. A., Barnes, I., Wingfield, B. D., and Wingfield, M. J. fimbriata infecting sweetpotato in North Carolina. Plant Dis. 101:
2014. Redefining Ceratocystis and allied genera. Stud. Mycol. 79:187-219. 994-1001.
Engelbrecht, C. J. B., and Harrington, T. C. 2005. Intersterility, morphology and Scruggs, A. C., and Quesada-Ocampo, L. M. 2016. Cultural, chemical, and
taxonomy of Ceratocystis fimbriata on sweet potato, cacao and sycamore. alternative control strategies for Rhizopus soft rot of sweetpotato. Plant Dis.
Mycologia 97:57-69. 100:1532-1540.
Geiser, D. M., Ward, T. J., Zhang, N., Kuldau, G. A., and O’Donnell, K. 2004. Soares, M., Christen, P., Pandey, A., and Soccol, C. R. 2000. Fruity flavour
FUSARIUM-ID v. 1.0: A DNA sequence database for identifying Fusarium. production by Ceratocystis fimbriata grown on coffee husk in solid-state
Eur. J. Plant Pathol. 110:473-479. fermentation. Process Biochem. 35:857-861.
Halsted, B. D. 1890. Some fungous diseases of the sweet potato. Bull. N.J. Agric. Stahr, M., and Quesada-Ocampo, L. M. 2020. Assessing the role of temperature,
Exp. Sta. 76:7-14. inoculum density, and wounding on disease progression of the fungal
Harrington, T. C. 2013. Ceratocystis diseases. Pages 230-255 in: Infectious pathogen Ceratocystis fimbriata causing black rot in sweetpotato. Plant Dis.
Forest Diseases. CABI, Wallingford, U.K. 104. doi: 10.1094/PDIS-12-18-2224-RE.
Harrington, T. C., Thorpe, D. J., and Alfenas, A. C. 2011. Genetic variation and Webster, R. K., and Butler, E. E. 1967. A morphological and biological concept
variation in aggressiveness to native and exotic hosts among Brazilian of the species Ceratocystis fimbriata. Can. J. Bot. 45:1457-1468.
populations of Ceratocystis fimbriata. Phytopathology 101:555-566. White, T. J., Bruns, T., Lee, S., and Taylor, J. 1990. Amplification and direct
Holland, L. A., Lawrence, D. P., Nouri, M. T., Harrington, T. C., and Trouillas, sequencing of fungal ribosomal RNA genes for phylogenetics. Pages
F. P. 2019. Taxonomic revision and multi-locus phylogeny of the North 315-322 in PCR Protocols: A Guide to Methods and Applications. Aca-
American clade of Ceratocystis. Fungal Syst. Evol. 3:135-156. demic Press, San Diego, CA.
Johnson, J. A., Harrington, T. C., and Engelbrecht, C. J. B. 2005. Phylogeny and Wilken, P. M., Steenkamp, E. T., Wingfield, M. J., de Beer, Z. W.,
taxonomy of the North American clade of the Ceratocystis fimbriata com- and Wingfield, B. D. 2013. IMA draft nuclear genome sequence for the plant
plex. Mycologia 97:1067-1092. pathogen, Ceratocystis fimbriata. IMA Fungus 4:357-358.

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