Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Diplodia seriata
(grapevine trunk disease)

Reeder R, 2020. Diplodia seriata (grapevine trunk disease). Invasive Species Compendium. Wallingford, UK: CABI. DOI:10.1079/ISC.9630.20210200697

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Diplodia seriata (grapevine trunk disease)

Pictures

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PictureTitleCaptionCopyright
Diplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus domestica).
TitleSymptoms on fruit
CaptionDiplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus domestica).
Copyright©Clemson University - USDA Cooperative Extension Slide Series/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus domestica).
Symptoms on fruitDiplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus domestica).©Clemson University - USDA Cooperative Extension Slide Series/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms on apple fruit and foliage (Malus spp.).
TitleSymptoms on fruit
CaptionDiplodia seriata (grapevine trunk disease); Symptoms on apple fruit and foliage (Malus spp.).
Copyright©University of Georgia Plant Pathology, University of Georgia/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms on apple fruit and foliage (Malus spp.).
Symptoms on fruitDiplodia seriata (grapevine trunk disease); Symptoms on apple fruit and foliage (Malus spp.).©University of Georgia Plant Pathology, University of Georgia/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of frog-eye spot on apple leaf (Malus spp.)
TitleSymptoms on leaf
CaptionDiplodia seriata (grapevine trunk disease); Symptoms of frog-eye spot on apple leaf (Malus spp.)
Copyright©Penn State Department of Plant Pathology & Environmental Microbiology Archives, Penn State University/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of frog-eye spot on apple leaf (Malus spp.)
Symptoms on leafDiplodia seriata (grapevine trunk disease); Symptoms of frog-eye spot on apple leaf (Malus spp.)©Penn State Department of Plant Pathology & Environmental Microbiology Archives, Penn State University/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of frog-eye spot on apple leaf (Malus spp.).
TitleSymptoms on leaf
CaptionDiplodia seriata (grapevine trunk disease); Symptoms of frog-eye spot on apple leaf (Malus spp.).
Copyright©University of Georgia Plant Pathology, University of Georgia/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of frog-eye spot on apple leaf (Malus spp.).
Symptoms on leafDiplodia seriata (grapevine trunk disease); Symptoms of frog-eye spot on apple leaf (Malus spp.).©University of Georgia Plant Pathology, University of Georgia/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus spp.).
TitleSymptoms on fruit
CaptionDiplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus spp.).
Copyright©University of Georgia Plant Pathology, University of Georgia/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus spp.).
Symptoms on fruitDiplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus spp.).©University of Georgia Plant Pathology, University of Georgia/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of black rot on Chinese pear fruit (Pyrus pyrifolia).
TitleSymptoms on fruit
CaptionDiplodia seriata (grapevine trunk disease); Symptoms of black rot on Chinese pear fruit (Pyrus pyrifolia).
Copyright©Mary Ann Hansen, Virginia Polytechnic Institute and State University/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of black rot on Chinese pear fruit (Pyrus pyrifolia).
Symptoms on fruitDiplodia seriata (grapevine trunk disease); Symptoms of black rot on Chinese pear fruit (Pyrus pyrifolia).©Mary Ann Hansen, Virginia Polytechnic Institute and State University/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus spp.).
TitleSymptoms on fruit
CaptionDiplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus spp.).
Copyright©University of Georgia Plant Pathology, University of Georgia/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus spp.).
Symptoms on fruitDiplodia seriata (grapevine trunk disease); Symptoms of black rot on apple fruit (Malus spp.).©University of Georgia Plant Pathology, University of Georgia/via Bugwood.org - CC BY 3.0
Diplodia seriata (grapevine trunk disease); Shrivelled black rot infected fruit. Hancock County, Maine, USA. March 2015.
TitleSymptoms on fruit
CaptionDiplodia seriata (grapevine trunk disease); Shrivelled black rot infected fruit. Hancock County, Maine, USA. March 2015.
Copyright©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Shrivelled black rot infected fruit. Hancock County, Maine, USA. March 2015.
Symptoms on fruitDiplodia seriata (grapevine trunk disease); Shrivelled black rot infected fruit. Hancock County, Maine, USA. March 2015.©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Wrinkled surface of fruit covered by a blanket of dark conidia from pycnidia embedded in tissue. Hancock County, Maine, USA. March 2015.
TitleSymptoms
CaptionDiplodia seriata (grapevine trunk disease); Wrinkled surface of fruit covered by a blanket of dark conidia from pycnidia embedded in tissue. Hancock County, Maine, USA. March 2015.
Copyright©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Wrinkled surface of fruit covered by a blanket of dark conidia from pycnidia embedded in tissue. Hancock County, Maine, USA. March 2015.
SymptomsDiplodia seriata (grapevine trunk disease); Wrinkled surface of fruit covered by a blanket of dark conidia from pycnidia embedded in tissue. Hancock County, Maine, USA. March 2015.©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Pycnidium embedded in fruit tissue, extruded dark conidia abound. Hancock County, Maine, USA. March 2015.
TitlePycnidium
CaptionDiplodia seriata (grapevine trunk disease); Pycnidium embedded in fruit tissue, extruded dark conidia abound. Hancock County, Maine, USA. March 2015.
Copyright©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Pycnidium embedded in fruit tissue, extruded dark conidia abound. Hancock County, Maine, USA. March 2015.
PycnidiumDiplodia seriata (grapevine trunk disease); Pycnidium embedded in fruit tissue, extruded dark conidia abound. Hancock County, Maine, USA. March 2015.©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Pycnidium embedded in fruit tissue, vertical section (Sphaeropsis anamorph). Hancock County, Maine, USA. March 2015.
TitlePycnidium
CaptionDiplodia seriata (grapevine trunk disease); Pycnidium embedded in fruit tissue, vertical section (Sphaeropsis anamorph). Hancock County, Maine, USA. March 2015.
Copyright©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Pycnidium embedded in fruit tissue, vertical section (Sphaeropsis anamorph). Hancock County, Maine, USA. March 2015.
PycnidiumDiplodia seriata (grapevine trunk disease); Pycnidium embedded in fruit tissue, vertical section (Sphaeropsis anamorph). Hancock County, Maine, USA. March 2015.©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Pycnidium, vertical median section showing conidiogenous wall and ostiolar neck (Sphaeropsis anamorph). Hancock County, Maine, USA. March 2015.
TitlePycnidium
CaptionDiplodia seriata (grapevine trunk disease); Pycnidium, vertical median section showing conidiogenous wall and ostiolar neck (Sphaeropsis anamorph). Hancock County, Maine, USA. March 2015.
Copyright©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Pycnidium, vertical median section showing conidiogenous wall and ostiolar neck (Sphaeropsis anamorph). Hancock County, Maine, USA. March 2015.
PycnidiumDiplodia seriata (grapevine trunk disease); Pycnidium, vertical median section showing conidiogenous wall and ostiolar neck (Sphaeropsis anamorph). Hancock County, Maine, USA. March 2015.©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Conidiogenous pycnidial wall and conidia on apple (Malus domestica). Hancock County, Maine. March 2015.
TitleConidiogenous pycnidial wall and conidia
CaptionDiplodia seriata (grapevine trunk disease); Conidiogenous pycnidial wall and conidia on apple (Malus domestica). Hancock County, Maine. March 2015.
Copyright©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0
Diplodia seriata (grapevine trunk disease); Conidiogenous pycnidial wall and conidia on apple (Malus domestica). Hancock County, Maine. March 2015.
Conidiogenous pycnidial wall and conidiaDiplodia seriata (grapevine trunk disease); Conidiogenous pycnidial wall and conidia on apple (Malus domestica). Hancock County, Maine. March 2015.©Bruce Watt, University of Maine/via Bugwood.org - CC BY-NC 3.0

Identity

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Preferred Scientific Name

  • Diplodia seriata De Not. (1845)

Preferred Common Name

  • grapevine trunk disease

Other Scientific Names

  • Botryosphaeria obtusa (Schwein.) Shoemaker (1964)
  • Diplodia profusa De Not. (1842)
  • Diplodia pseudodiplodia Fuckel. (1870)
  • Physalospora cydoniae G. Arnaud (1911)
  • Physalospora malorum Shear, N.E. Stevens & Wilcox. (1924)
  • Physalospora obtusa (Schwein.) Cooke (1892)
  • Sphaeria obtusa Schwein (1832)

International Common Names

  • English: apple black rot; bark: pome fruit necrosis; black rot canker: apple; black rot of apple; black: apple canker; black: grapevine dead-arm disease; Botryosphaeria dieback; canker: juniper; dieback: grapevine; dieback: oak; frogeye leaf spot: apple; loquat fruit rot; tree canker: apple
  • Spanish: black-rot del fresal; black-rot del manzano; black-rot del membrillo; chancro del manzano; falso black-rot del manzano; falso black-rot del peral; podredumbre negra del ciruelo; podredumbre negra del manzano
  • French: black-rot du cognassier; black-rot du fraisier; black-rot du pommier; chancre du pommier; dead arm noir de la vigne; faux black-rot du poirier; faux black-rot du pommier; pourriture noire du pommier; pourriture noire du prunier
  • German: Froschaugenkrankheit: Apfel; Rindenbrand: Obstgehoelze; Schwarzer: Obstgehoelze Krebs; Schwarzfaeule: Apfel

EPPO code

  • BOTSOB

Summary of Invasiveness

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Diplodia seriata is a cosmopolitan and plurivorous fungal species occurring on woody hosts belonging to many plant genera and families (Punithalingam and Waller, 1973; Phillips et al., 2007; Slippers et al., 2007). The fungus is encountered in many habitats, but has a primarily temperate distribution and is present on most continents.

D. seriata causes canker, dieback, fruit rot and leaf spot diseases on economically important forest and horticultural species (Farr and Rossman, 2020). Reports of the virulence of this pathogen vary depending upon the crop, varieties and hosts involved and it is often regarded as a stress-related pathogen taking advantage of weak or stressed plants. In common with other members of the Botryosphaeriaceae, D. seriata is capable of living endophytically inside plants (Crous et al., 2006; Slippers and Wingfield, 2007) and latent infections of fruits can result in storage rots. The pathogen is dispersed through both pycnidia and ascospores with conidia regarded as the most important inoculum source for short-distance spread. Infection is through wounds, natural openings, or direct penetration of the host tissue. There is no evidence that this species is seedborne although some members of the Botryosphaeriaceae have been shown to be present in seeds (Gure et al., 2005). The extensive host range of this species means that it is more likely to become established in new areas, as establishment will not depend on the presence of specific hosts. The widespread distribution of this species is presumably as a result of the word-wide movement agricultural, forestry and ornamental plants.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Fungi
  •         Phylum: Ascomycota
  •             Subphylum: Pezizomycotina
  •                 Class: Dothideomycetes
  •                     Order: Botryosphaeriales
  •                         Family: Botryosphaeriaceae
  •                             Genus: Diplodia
  •                                 Species: Diplodia seriata

Notes on Taxonomy and Nomenclature

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The genus Botryosphaeria was recently re-evaluated through a study of partial sequences of the LSU gene (Crous et al., 2006). This study determined that Botryosphaeria s. lat was composed of 10 phylogenetic lineages that represent individual genera. To avoid the introduction of new generic names, these authors chose to use existing asexual generic names for most of the lineages, and restricted the use of Botryosphaeria to B. dothidea (Moug. Fr.) Ces. & De Not. and B. corticis (Demaree & M.S. Wilcox) Arx & Müll. Consequently, the name Botryosphaeria is no longer acceptable for most of the species with Fusicoccum-like and Diplodia-like anamorphs including B. obtusa which has been named by its anamorph.

The anamorph of B. obtusa belongs in Diplodia Fr. because of its brown, aseptate conidia formed on phialides that line the inner wall of the pycnidial conidiomata and multiply via periclinal thickening, or annellations. However, the species differentiation within Diplodia has proven rather more difficult largely due to the lack of distinctive morphological features. This has resulted in Diplodia species being defined on the basis of host association and consequently a proliferation of species names. Unfortunately, the host is not a reliable means of species differentiation in the Botryosphaeriaceae and thus many of the names in Diplodia are likely to be synonyms (Slippers et al., 2004a).

As with the teleomorph the correct anamorph name for B. obtusa has also been the subject of much argument and confusion. In the past the debate has mainly revolved around the names Sphaeropsis malorum (Berk.) Berk. and S. malorum Peck. (Phillips et al., 2007). The anamorph S. malorum Peck. was introduced by Saccardo (1884) when he transferred S. malorum (Berk.) Berk to the genus Phoma, on the basis of its hyaline conidia. However, when Saccardo examined the S. malorum samples collected by Peck, he found them to be different from the Berkely collection, due to the production of brown conidia, and chose the name S. malorum Peck to represent them. Unfortunately, this name is an illegitimate, later homonym of S. malorum (Berk.) Berk. (1860) and is not recognized. The fungus S. malorum (Berk.) is itself a synonym of D. mutila, therefore neither of these names can be used for the anamorph of “Botryosphaeriaobtusa.  

The taxonomic position of the anamorph was recently clarified in a phylogenetic study of “B”. obtusa-type specimens conducted by Phillips et al. (2007). This study determined that D. seriata De Not. was the oldest name available for the asexual morph of what had been previously referred to as “B.obtusa and this is the currently accepted species name. The circumscription of this fungus is however, further complicated by the likely existence of cryptic species (Phillips et al., 2012).

The genome sequences of two D. seriata isolates are available (Morales-Cruz et al., 2015 ; Robert-Siegwald et al., 2017). They are highly similar and have a typical Ascomycota genome organisation.

Description

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Infection by D. seriata is thought to occur through wounds, however, it is not clear whether these wounds are simple entry points, or whether they provide chemical signals that enhance spore germination processes. In addition, pathogenicity studies performed on woody hosts such as apple, peach and pistachio have demonstrated that these pathogens can also infect through natural openings such as stomata and lenticels or even penetrate host tissue directly (Michailides, 1991; Pusey, 1989; Kim et al., 1999). Both conidia and ascospores are infective, although ascospores are rarely found in the natural environment. The release of conidia is triggered by rainfall or humidity levels of 70% or above. Dispersal by rain splash is then thought to be over relatively short distances (Úrbez‐Torres et al., 2010, Baskarathevan et al., 2013). Phillips et al. (2007) selected a specimen on Vitis vinifera collected in Portugal (CBS-H 19809) as the epitype when proposing the anamorph and preferred scientific name for this pathogen. He gave the following description of the morphological characteristics. The conidiomata are pycnidial, separate or aggregated and confluent, immersed in the host, partially emergent at maturity, dark brown to black, ostiolate, non papillate, thick-walled, outer layers composed of dark-brown textura angularis, inner layers of thin-walled hyaline textura angularis. Conidiogenous cells 3- 5.5 × 7-10(-15) µm, hyaline, thin-walled, smooth, cylindrical, swollen at the base, discrete, producing a single conidium at the tip, indeterminate, proliferating internally giving rise to periclinal thickenings or proliferating percurrently forming 2-3 annelations. Conidia (21.5-) 22-27(-28) × (11-)11.5- 14.5(-15.5) µm, 95% confidence limits = 24.3-25.4 × 12-6-13.2 µm ( x ± S.D. of 50 = 24.9 ± 1.9 × 12.9 ± 1.1 µm, L/W = 1.9 ± 0.1) initially hyaline, becoming dark brown, moderately thick-walled (ca. 0.5 µm thick), wall externally smooth, roughened on the inner surface, aseptate, ovoid, widest in the middle, apex obtuse, base truncate or rounded.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 10 Mar 2021
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

AlgeriaPresent
KenyaPresent
Sierra LeonePresent
South AfricaPresent
TanzaniaPresent
TunisiaPresent
ZambiaPresent
ZimbabwePresent

Asia

ChinaPresent
-HebeiPresent
-ShandongPresent
-XinjiangPresent
GeorgiaPresent
IndiaPresent, Localized
-BiharPresent
-ChhattisgarhPresent
-Himachal PradeshPresent
-Jammu and KashmirPresent
-KarnatakaPresent
-Madhya PradeshPresent
-MaharashtraPresent
-RajasthanPresent
-Uttar PradeshPresent
-UttarakhandPresent
-West BengalPresent
IranPresent
IraqPresent
JapanPresent
LebanonPresent
MalaysiaPresent
-SabahPresent
PakistanPresent
South KoreaPresent
Sri LankaPresent
TaiwanPresent
TurkeyPresent

Europe

BelgiumPresent
Bosnia and HerzegovinaPresent
BulgariaPresent
CroatiaPresent
CyprusPresent
CzechiaPresent
FrancePresent
GermanyPresent
GreecePresent
HungaryPresent
ItalyPresent
-SardiniaPresent
-SicilyPresent
LatviaPresent
NetherlandsPresent
PortugalPresent
RomaniaPresent
RussiaPresent
-Central RussiaPresent
-Southern RussiaPresent
Serbia and MontenegroPresent
SlovakiaPresent
SloveniaPresent
SpainPresent
UkrainePresent
United KingdomPresent

North America

CanadaPresent
-British ColumbiaPresent
-ManitobaPresent
-OntarioPresent
MexicoPresent
United StatesPresent, Widespread
-AlabamaPresent
-ArkansasPresent
-CaliforniaPresent
-ConnecticutPresent
-DelawarePresent
-GeorgiaPresent
-KansasPresent
-KentuckyPresent
-MarylandPresent
-MichiganPresent
-MinnesotaPresent
-MissouriPresent
-NebraskaPresent
-New YorkPresent
-North DakotaPresent
-OhioPresent
-OklahomaPresent
-PennsylvaniaPresent
-South DakotaPresent
-TexasPresent
-VirginiaPresent
-WashingtonPresent2014
-West VirginiaPresent

Oceania

AustraliaPresent, Localized
-New South WalesPresent
-TasmaniaPresent
-Western AustraliaPresent
New ZealandPresent
Papua New GuineaPresent

South America

ArgentinaPresent
BoliviaPresent
BrazilPresent, Localized
-Santa CatarinaPresent
ChilePresent
EcuadorPresent
UruguayPresent
VenezuelaPresent

History of Introduction and Spread

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D. seriata has been recorded worldwide on many different hosts of agriculture, forestry, or horticultural importance. The fungus was first described from Italy by Schweinitz (1832) as Sphaeria obtusa, and was found on dead stems of Jasminum. However, many past records relating to this pathogen are of questionable validity because of the previously confusing taxonomy and the unreliable nature of the morphological characters used in species identification. It is likely that much of the spread around the world was via the global trade of plants and plant products, but it is not possible to trace the historical routes of introduction.

Risk of Introduction

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Currently no regional plant protection organization consider D. seriata to be a quarantine pest, possibly due to its already extensive distribution. As with most canker fungi these organisms can survive in a reproductive state on woody host material. The prolonged latent infection, or endophytic phase on some hosts means that the fungus can pass undetected by quarantine systems in traded living plants, fruits, and other plant parts, illustrating the phytosanitary shortcomings related to the detection of endophytic plant pathogens. The colonization potential for D. seriata is high because of the wide host range and diversity of environments it could encounter upon entry. The pathogen is damaging to a wide variety of hosts especially if new more virulent strains are introduced that affect ornamental or high value plantings.

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial ManagedCultivated / agricultural land Principal habitat Harmful (pest or invasive)
Terrestrial ManagedProtected agriculture (e.g. glasshouse production) Secondary/tolerated habitat Harmful (pest or invasive)
Terrestrial ManagedManaged forests, plantations and orchards Principal habitat Harmful (pest or invasive)
Terrestrial ManagedDisturbed areas Present, no further details Natural
Terrestrial ManagedRail / roadsides Present, no further details Natural
Terrestrial ManagedUrban / peri-urban areas Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalNatural forests Present, no further details Natural
Terrestrial Natural / Semi-naturalRiverbanks Present, no further details Natural
Terrestrial Natural / Semi-naturalScrub / shrublands Present, no further details Natural
LittoralCoastal areas Present, no further details Natural

Hosts/Species Affected

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The fungus D. seriata, known for many years as Botryosphaeria obtusa, is an important pathogen of apples causing frog-eye spot, black rot, canker and shoot dieback. In addition to apples, B. obtusa has been isolated from at least 34 different hosts (Punithalingam and Waller, 1973). In recent years it has been recognized as a pathogen of Vitis vinifera in Europe (Phillips, 1998, 2002; Urbez-Torres, 2011), Australia (Castillo-Pando et al., 2001) and South Africa (van Niekerk et al., 2004).

Punithalingam and Waller (1973) reported this fungus to be isolated from 35 different plant genera, but the current number of known hosts is much greater. According to the Systematic Mycology and Microbiology Laboratory Fungal Database (Agricultural Research Service, United States Department of Agriculture) there are 209 named species representing 151 genera of herbaceous and woody hosts associated with this pathogen including synonyms (Farr and Rossman, 2020). The Herb IMI database (Herb IMI Database, Royal Botanic Gardens, Kew, UK) lists 47 species and 40 genera as being associated with Peyronellaea obtuse (an erroneous synonym for D. seriata). Combining the herbarium lists results in 241 named species from 163 genera that have been reported associated with this fungus. There are however some question marks over the authenticity of some of these associations given the complex taxonomy of this species and the unreliability of morphological characteristics for identification. Moreover, in some instances the fungus may have been growing saprophytically on dead material rather than acting as a primary pathogen.

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Abies concolor (Rocky Mountain white fir)PinaceaeOther
    Acer negundo (box elder)AceraceaeOther
      Acer rubrum (red maple)AceraceaeOther
        Acer saccharinum (silver maple)AceraceaeOther
          Aesculus pavia (red buckeye)HippocastanaceaeOther
            Ailanthus altissima (tree-of-heaven)SimaroubaceaeOther
              Albizia julibrissin (silk tree)FabaceaeOther
                Alhagi maurorum (camelthorn)FabaceaeOther
                  Amorpha fruticosa (false indigo-bush)FabaceaeOther
                    Araucaria araucana (monkey puzzle)AraucariaceaeOther
                      Araucaria heterophylla (norfolk Island pine)AraucariaceaeOther
                        Aronia arbutifolia (red chokeberry)RosaceaeOther
                          Artemisia vulgaris (mugwort)AsteraceaeOther
                            Baccharis halimifolia (groundsel-bush)AsteraceaeOther
                              Betula nigra (river birch)BetulaceaeOther
                                Broussonetia papyrifera (paper mulberry)MoraceaeOther
                                  Callicarpa americana (American beautyberry)LamiaceaeOther
                                    Camellia sinensis (tea)TheaceaeOther
                                      Campsis radicans (trumpetcreeper)BignoniaceaeOther
                                        Canna glaucaCannaceaeOther
                                          Canna indica (canna lilly)CannaceaeOther
                                            Carpinus caroliniana (American hornbeam)BetulaceaeOther
                                              Carya cathayensis (Chinese hickory)JuglandaceaeOther
                                                Castanea dentata (American chestnut)FagaceaeOther
                                                  Castanea sativa (chestnut)FagaceaeOther
                                                    Ceanothus (white-thorn)RhamnaceaeOther
                                                      Cedrus atlantica (Atlas cedar)PinaceaeOther
                                                        Cedrus deodara (Himalayan cedar)PinaceaeOther
                                                          Celtis laevigata (Sugarberry)UlmaceaeOther
                                                            Celtis occidentalis (hackberry)UlmaceaeOther
                                                              Cephalanthus occidentalis (common buttonbush)RubiaceaeOther
                                                                Cercis canadensis (eastern redbud)FabaceaeOther
                                                                  Chamaecyparis lawsoniana (Port Orford cedar)CupressaceaeOther
                                                                    Chamaecyparis pisifera (sawara false cypress)CupressaceaeOther
                                                                      Chamaecyparis thyoides (Atlantic white cedar)CupressaceaeOther
                                                                        Citrus latifolia (tahiti lime)RutaceaeOther
                                                                          Citrus limon (lemon)RutaceaeOther
                                                                            Citrus nobilis (tangor)RutaceaeOther
                                                                              Citrus sinensis (navel orange)RutaceaeOther
                                                                                Cocculus hirsutusMenispermaceaeOther
                                                                                  Cornus florida (Flowering dogwood)CornaceaeOther
                                                                                    CorylusBetulaceaeUnknown
                                                                                      Corylus americana (American hazel)BetulaceaeOther
                                                                                        Corylus avellana (hazel)BetulaceaeOther
                                                                                          Corylus cornuta (beaked hazel)BetulaceaeOther
                                                                                            Cotinus coggygria (fustet)AnacardiaceaeOther
                                                                                              Cotoneaster bullatusRosaceaeOther
                                                                                                Cotoneaster salicifolius (willowleaf cotoneaster)RosaceaeOther
                                                                                                  Crataegus monogyna (hawthorn)RosaceaeOther
                                                                                                    Cupressus macrocarpa (Monterey cypress)CupressaceaeOther
                                                                                                      Cupressus sempervirens (Mediterranean cypress)CupressaceaeOther
                                                                                                        Cydonia oblonga (quince)RosaceaeOther
                                                                                                          Dalbergia sissooFabaceaeOther
                                                                                                            Diospyros kaki (persimmon)EbenaceaeOther
                                                                                                              Diospyros virginiana (persimmon (common))EbenaceaeOther
                                                                                                                Eriobotrya japonica (loquat)RosaceaeOther
                                                                                                                  Eucalyptus globulus (Tasmanian blue gum)MyrtaceaeUnknown
                                                                                                                    Ficus carica (common fig)MoraceaeOther
                                                                                                                      Fraxinus (ashes)OleaceaeOther
                                                                                                                        Fraxinus americana (white ash)OleaceaeOther
                                                                                                                          Fraxinus angustifolia (narrow-leaved ash)OleaceaeOther
                                                                                                                            Fraxinus excelsior (ash)OleaceaeOther
                                                                                                                              Fraxinus ornus (flowering ash)OleaceaeOther
                                                                                                                                Fraxinus pennsylvanica (downy ash)OleaceaeOther
                                                                                                                                  Gleditsia triacanthos (honey locust)FabaceaeOther
                                                                                                                                    Grevillea robusta (silky oak)ProteaceaeOther
                                                                                                                                      Hedera helix (ivy)AraliaceaeOther
                                                                                                                                        Humulus lupulus (hop)CannabaceaeOther
                                                                                                                                          Ilex opaca (American holly)AquifoliaceaeOther
                                                                                                                                            Juglans cinerea (butternut)JuglandaceaeOther
                                                                                                                                              Juglans hindsii (californian black walnut)JuglandaceaeOther
                                                                                                                                                Juglans nigra (black walnut)JuglandaceaeOther
                                                                                                                                                  Juglans regia (walnut)JuglandaceaeOther
                                                                                                                                                    Juniperus sabina (savin juniper)CupressaceaeOther
                                                                                                                                                      Juniperus virginiana (eastern redcedar)CupressaceaeOther
                                                                                                                                                        Lagerstroemia indica (Indian crape myrtle)LythraceaeOther
                                                                                                                                                          Leucaena leucocephala (leucaena)FabaceaeOther
                                                                                                                                                            Ligustrum vulgare (common privet)OleaceaeOther
                                                                                                                                                              Liquidambar styraciflua (Sweet gum)HamamelidaceaeOther
                                                                                                                                                                Liriodendron tulipifera (tuliptree)MagnoliaceaeOther
                                                                                                                                                                  Lonicera japonica (Japanese honeysuckle)CaprifoliaceaeOther
                                                                                                                                                                    Maclura pomifera (osage orange)MoraceaeOther
                                                                                                                                                                      Malus baccata (siberian crab apple)RosaceaeMain
                                                                                                                                                                        Malus coronaria (sweet crab-apple)RosaceaeMain
                                                                                                                                                                          Malus domestica (apple)RosaceaeMain
                                                                                                                                                                            Malus floribundaRosaceaeMain
                                                                                                                                                                              Malus ioensis (prairie crab-apple)RosaceaeMain
                                                                                                                                                                                Malus prunifolia (plum-leaved crab apple)RosaceaeMain
                                                                                                                                                                                  Malus pumilaMain
                                                                                                                                                                                    Malus sylvestris (crab-apple tree)RosaceaeMain
                                                                                                                                                                                      Melia azedarach (Chinaberry)MeliaceaeOther
                                                                                                                                                                                        Mespilus germanica (medlar)RosaceaeOther
                                                                                                                                                                                          Morus (mulberrytree)MoraceaeOther
                                                                                                                                                                                            Morus alba (mora)MoraceaeOther
                                                                                                                                                                                              Morus nigra (black mulberry)MoraceaeOther
                                                                                                                                                                                                Myrica cerifera (Southern waxmyrtle)MyricaceaeOther
                                                                                                                                                                                                  Nannorrhops ritchieanaArecaceaeOther
                                                                                                                                                                                                    Nerium oleander (oleander)ApocynaceaeOther
                                                                                                                                                                                                      Oenothera biennis (common evening primrose)OnagraceaeOther
                                                                                                                                                                                                        Olea europaeaOleaceaeOther
                                                                                                                                                                                                          Olea europaea subsp. europaea (European olive)OleaceaeOther
                                                                                                                                                                                                            Ostrya virginiana (American hophornbeam)BetulaceaeOther
                                                                                                                                                                                                              Oxydendrum arboreum (Sourwood)EricaceaeOther
                                                                                                                                                                                                                Parthenocissus quinquefolia (Virginia creeper)VitaceaeOther
                                                                                                                                                                                                                  Paulownia tomentosa (paulownia)ScrophulariaceaeOther
                                                                                                                                                                                                                    Pelargonium graveolens (Rose geranium)GeraniaceaeOther
                                                                                                                                                                                                                      Persea americana (avocado)LauraceaeOther
                                                                                                                                                                                                                        Picea glauca (white spruce)PinaceaeOther
                                                                                                                                                                                                                          Pinus nigra (black pine)PinaceaeOther
                                                                                                                                                                                                                            Pinus patula (Mexican weeping pine)PinaceaeOther
                                                                                                                                                                                                                              Pinus radiata (radiata pine)PinaceaeOther
                                                                                                                                                                                                                                Pinus strobus (eastern white pine)PinaceaeOther
                                                                                                                                                                                                                                  Pinus virginiana (scrub pine)PinaceaeOther
                                                                                                                                                                                                                                    Pistacia chinensis (chinese pistachio)AnacardiaceaeOther
                                                                                                                                                                                                                                      Pistacia lentiscus (mastic tree)AnacardiaceaeOther
                                                                                                                                                                                                                                        Pistacia vera (pistachio)AnacardiaceaeOther
                                                                                                                                                                                                                                          Platanus occidentalis (sycamore)PlatanaceaeOther
                                                                                                                                                                                                                                            Populus alba (silver-leaf poplar)SalicaceaeOther
                                                                                                                                                                                                                                              Populus deltoides (poplar)SalicaceaeOther
                                                                                                                                                                                                                                                Populus nigra (black poplar)SalicaceaeOther
                                                                                                                                                                                                                                                  Prunus armeniaca (apricot)RosaceaeOther
                                                                                                                                                                                                                                                    Prunus avium (sweet cherry)RosaceaeOther
                                                                                                                                                                                                                                                      Prunus cerasus (sour cherry)RosaceaeOther
                                                                                                                                                                                                                                                        Prunus domestica (plum)RosaceaeMain
                                                                                                                                                                                                                                                          Prunus dulcis (almond)RosaceaeOther
                                                                                                                                                                                                                                                            Prunus laurocerasus (cherry laurel)Other
                                                                                                                                                                                                                                                              Prunus munsoniana (wild goose plum)RosaceaeOther
                                                                                                                                                                                                                                                                Prunus persica (peach)RosaceaeMain
                                                                                                                                                                                                                                                                  Prunus salicina (Japanese plum)RosaceaeOther
                                                                                                                                                                                                                                                                    Prunus serotina (black cherry)RosaceaeOther
                                                                                                                                                                                                                                                                      Prunus spinosa (blackthorn)RosaceaeOther
                                                                                                                                                                                                                                                                        Prunus triloba (Rose tree of China)RosaceaeOther
                                                                                                                                                                                                                                                                          Prunus virginiana (common chokecherrytree)RosaceaeOther
                                                                                                                                                                                                                                                                            Psidium guajava (guava)MyrtaceaeOther
                                                                                                                                                                                                                                                                              Ptelea trifoliata (Hoptree)RutaceaeOther
                                                                                                                                                                                                                                                                                Punica granatum (pomegranate)PunicaceaeOther
                                                                                                                                                                                                                                                                                  Pyrus communis (European pear)RosaceaeMain
                                                                                                                                                                                                                                                                                    Pyrus pyrifolia (Oriental pear tree)RosaceaeMain
                                                                                                                                                                                                                                                                                      Quercus coccifera (kermes oak)FagaceaeOther
                                                                                                                                                                                                                                                                                        Quercus ilex (holm oak)FagaceaeOther
                                                                                                                                                                                                                                                                                          Quercus macrocarpa (mossy-cup oak)FagaceaeOther
                                                                                                                                                                                                                                                                                            Quercus nigra (water oak)FagaceaeOther
                                                                                                                                                                                                                                                                                              Quercus robur (common oak)FagaceaeOther
                                                                                                                                                                                                                                                                                                Quercus rubra (northern red oak)FagaceaeOther
                                                                                                                                                                                                                                                                                                  Quercus suber (cork oak)FagaceaeOther
                                                                                                                                                                                                                                                                                                    Quercus velutina (black oak)FagaceaeOther
                                                                                                                                                                                                                                                                                                      Quercus virginiana (Live oak)FagaceaeOther
                                                                                                                                                                                                                                                                                                        Rhamnus caroliniana (Carolina buckthorn)RhamnaceaeOther
                                                                                                                                                                                                                                                                                                          Rhododendron japonicum (Japanese azalea)EricaceaeOther
                                                                                                                                                                                                                                                                                                            Rhododendron maximum (Rosebay rhododendron)EricaceaeOther
                                                                                                                                                                                                                                                                                                              Rhus copallina (Shining sumac)AnacardiaceaeOther
                                                                                                                                                                                                                                                                                                                Rhus glabra (smooth sumac)AnacardiaceaeOther
                                                                                                                                                                                                                                                                                                                  Rhus typhina (staghorn sumac)AnacardiaceaeOther
                                                                                                                                                                                                                                                                                                                    Ribes aureum (golden currant)GrossulariaceaeOther
                                                                                                                                                                                                                                                                                                                      Ribes rubrum (red currant)GrossulariaceaeOther
                                                                                                                                                                                                                                                                                                                        Ricinus communis (castor bean)EuphorbiaceaeOther
                                                                                                                                                                                                                                                                                                                          Robinia pseudoacacia (black locust)FabaceaeOther
                                                                                                                                                                                                                                                                                                                            Rosa canina (Dog rose)RosaceaeOther
                                                                                                                                                                                                                                                                                                                              Rubus fruticosus (blackberry)RosaceaeOther
                                                                                                                                                                                                                                                                                                                                Rubus idaeus (raspberry)RosaceaeOther
                                                                                                                                                                                                                                                                                                                                  Rubus ursinus (boysenberry)RosaceaeOther
                                                                                                                                                                                                                                                                                                                                    Rumex crispus (curled dock)PolygonaceaeOther
                                                                                                                                                                                                                                                                                                                                      Rumex obtusifolius (broad-leaved dock)PolygonaceaeOther
                                                                                                                                                                                                                                                                                                                                        Ruta graveolens (common rue)RutaceaeOther
                                                                                                                                                                                                                                                                                                                                          Salix alba (white willow)SalicaceaeOther
                                                                                                                                                                                                                                                                                                                                            Salix babylonica (weeping willow)SalicaceaeOther
                                                                                                                                                                                                                                                                                                                                              Salix caprea (pussy willow)SalicaceaeOther
                                                                                                                                                                                                                                                                                                                                                Salix nigra (black willow)SalicaceaeOther
                                                                                                                                                                                                                                                                                                                                                  Sambucus canadensis (American black elderberry)CaprifoliaceaeOther
                                                                                                                                                                                                                                                                                                                                                    Sassafras albidum (common sassafras)LauraceaeOther
                                                                                                                                                                                                                                                                                                                                                      Simmondsia chinensis (jojoba)SimmondsiaceaeOther
                                                                                                                                                                                                                                                                                                                                                        Sorbus americana (American mountainash)RosaceaeOther
                                                                                                                                                                                                                                                                                                                                                          Sorbus aria (whitebeam)RosaceaeOther
                                                                                                                                                                                                                                                                                                                                                            Sorbus aucuparia (mountain ash)RosaceaeOther
                                                                                                                                                                                                                                                                                                                                                              Styphnolobium japonicum (pagoda tree)FabaceaeUnknown
                                                                                                                                                                                                                                                                                                                                                                Syringa vulgaris (lilac)OleaceaeOther
                                                                                                                                                                                                                                                                                                                                                                  Syzygium cumini (black plum)MyrtaceaeOther
                                                                                                                                                                                                                                                                                                                                                                    Tectona grandis (teak)LamiaceaeOther
                                                                                                                                                                                                                                                                                                                                                                      Thuja occidentalis (Eastern white cedar)CupressaceaeOther
                                                                                                                                                                                                                                                                                                                                                                        Thuja plicata (western redcedar)CupressaceaeOther
                                                                                                                                                                                                                                                                                                                                                                          Tilia americana (basswood)TiliaceaeOther
                                                                                                                                                                                                                                                                                                                                                                            Ulmus americana (American elm)UlmaceaeOther
                                                                                                                                                                                                                                                                                                                                                                              Ulmus rubra (slippery elm)UlmaceaeOther
                                                                                                                                                                                                                                                                                                                                                                                Ulmus thomasii (rock elm)UlmaceaeOther
                                                                                                                                                                                                                                                                                                                                                                                  Vaccinium arboreum (Tree huckleberry)EricaceaeOther
                                                                                                                                                                                                                                                                                                                                                                                    Vaccinium corymbosum (blueberry)EricaceaeOther
                                                                                                                                                                                                                                                                                                                                                                                      Vinca minor (common periwinkle)ApocynaceaeOther
                                                                                                                                                                                                                                                                                                                                                                                        Vitis labrusca (fox grape)VitaceaeMain
                                                                                                                                                                                                                                                                                                                                                                                          Vitis rotundifolia (Muscadine grape)VitaceaeMain
                                                                                                                                                                                                                                                                                                                                                                                            Vitis vinifera (grapevine)VitaceaeOther
                                                                                                                                                                                                                                                                                                                                                                                              Yucca glauca (great plains yucca)AgavaceaeOther
                                                                                                                                                                                                                                                                                                                                                                                                Zelkova carpinifolia (caucasian elm)UlmaceaeOther

                                                                                                                                                                                                                                                                                                                                                                                                  Growth Stages

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                                                                                                                                                                                                                                                                                                                                                                                                  Fruiting stage, Post-harvest, Vegetative growing stage

                                                                                                                                                                                                                                                                                                                                                                                                  Symptoms

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                                                                                                                                                                                                                                                                                                                                                                                                  D. seriata has been associated with diseases such as fruit rot, dieback and cankers on a wide range of economically and environmentally important plants. There are too many hosts to discuss them all, so only a couple of economically important hosts are provided below, although it is likely that the symptoms of cankers and die back will be similar across many of the reported hosts.

                                                                                                                                                                                                                                                                                                                                                                                                  On apple, the fungus affects a variety of plant parts including leaves, fruit and branches. One of the most damaging is the fruit rot phase known as black rot which causes the fruit of apples and pears to rot before harvest and in storage. The disease can cause latent infections of these fruits which do not become apparent until after harvest. The first visible symptoms of latent fruit infection are small black lesions (2-4 mm diam.) which are slightly sunken with a corky texture. These black lesions do not enlarge further and only give rise to a rapidly progressing pale brown rot 2-3 weeks preceding harvest. The active stage of the fruit rot can be seen in the orchard and is characterised by rot that has concentric zones of lighter and darker brown colours, later the rotted areas turn black. Fruits affected by this kind of brown rot are rapidly colonised within 3-5 days. The fungus also causes a distinctive leaf spot, known as frogeye spot. Leaf lesions are initially small, purple specks that enlarge to form spots 3 to 6 mm in diameter, these spots have light brown-to-grey centres which are surrounded by one or more darker rings of tissue and a purple border. Dark pycnidia of the fungus may develop in the centre of older leaf spots. Stem symptoms of D. seriata begin as slightly sunken, reddish-brown patches within the bark. These areas enlarge and darken to form cankers with sunken centres and raised margins. Cankers may also develop as a superficial roughening or cracking of the bark, especially at the margins, where the cankers girdle the twigs or branches a blight and dieback is seen. D. seriata is regarded as an important pathogen of apple in the USA (Stevens, 1933; Brown and Britton, 1986; Brown-Rytlewski and McManus, 2000) but as a weak secondary pathogen on the same host in the UK and New Zealand (Laundon, 1973).

                                                                                                                                                                                                                                                                                                                                                                                                  On grapevines D. seriata is known to cause the death of spring buds, leaf chlorosis, fruit rot and trunk dieback, with brown, hard necrosis of the wood that appears as wedge-shaped necrosis in cross sections of the affected plant parts (van Niekerk et al., 2006; Urbez-Torres, 2011). Other symptoms include internal streaking and pith necrosis of wood, failure of graft union in young vines and cane bleaching (Urbez-Torres, 2011). D. seriata is one of the most cited Botryosphaeriaceae species occurring on grapevines worldwide and is frequently associated with the ‘black dead arm’ disease of grapevine (Larignon et al., 2001; Urbez-Torres, 2011). Recently Urbez-Torres (2011) proposed the name ‘Botryosphaeria dieback’ to include the increasing number of Botryosphaeriaceous species besides D. seriata that have been associated with most of the symptoms and diseases above. Reports of the virulence of this pathogen on grapes varies with some artificial inoculation studies (Spagnolo et al., 2017; Pinto et al., 2018; Reis et al., 2019) suggesting that it is a weak pathogen to grapevine and possibly takes advantage of weak or stressed plants (Qiu et al., 2016). These differences may be due to variations in virulence between strains, or they may be a result of the incomplete knowledge of the taxonomy of the genus, which in turn hampers accurate species recognition and identification. It is also possible that in species with a broad host range, such as D. seriata, virulence of any given isolate may vary according to the host that is being attacked.

                                                                                                                                                                                                                                                                                                                                                                                                  List of Symptoms/Signs

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                                                                                                                                                                                                                                                                                                                                                                                                  SignLife StagesType
                                                                                                                                                                                                                                                                                                                                                                                                  Fruit / abnormal shape
                                                                                                                                                                                                                                                                                                                                                                                                  Fruit / discoloration
                                                                                                                                                                                                                                                                                                                                                                                                  Fruit / lesions: black or brown
                                                                                                                                                                                                                                                                                                                                                                                                  Fruit / mummification
                                                                                                                                                                                                                                                                                                                                                                                                  Growing point / dieback
                                                                                                                                                                                                                                                                                                                                                                                                  Growing point / lesions
                                                                                                                                                                                                                                                                                                                                                                                                  Growing point / rot
                                                                                                                                                                                                                                                                                                                                                                                                  Growing point / wilt
                                                                                                                                                                                                                                                                                                                                                                                                  Leaves / abnormal colours
                                                                                                                                                                                                                                                                                                                                                                                                  Leaves / abnormal leaf fall
                                                                                                                                                                                                                                                                                                                                                                                                  Leaves / necrotic areas
                                                                                                                                                                                                                                                                                                                                                                                                  Leaves / rot
                                                                                                                                                                                                                                                                                                                                                                                                  Leaves / wilting
                                                                                                                                                                                                                                                                                                                                                                                                  Leaves / yellowed or dead
                                                                                                                                                                                                                                                                                                                                                                                                  Stems / canker on woody stem
                                                                                                                                                                                                                                                                                                                                                                                                  Stems / dieback
                                                                                                                                                                                                                                                                                                                                                                                                  Stems / discoloration
                                                                                                                                                                                                                                                                                                                                                                                                  Stems / gummosis or resinosis
                                                                                                                                                                                                                                                                                                                                                                                                  Stems / internal discoloration
                                                                                                                                                                                                                                                                                                                                                                                                  Stems / necrosis
                                                                                                                                                                                                                                                                                                                                                                                                  Stems / ooze
                                                                                                                                                                                                                                                                                                                                                                                                  Whole plant / discoloration
                                                                                                                                                                                                                                                                                                                                                                                                  Whole plant / early senescence
                                                                                                                                                                                                                                                                                                                                                                                                  Whole plant / plant dead; dieback

                                                                                                                                                                                                                                                                                                                                                                                                  Biology and Ecology

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                                                                                                                                                                                                                                                                                                                                                                                                  Genetics

                                                                                                                                                                                                                                                                                                                                                                                                  The genetic diversity of a species is highly influenced by the relative contribution of its asexual and sexual reproduction forms. Reproduction of Botryosphaeriaceae species is believed to be mainly by asexual means (Phillips, 2002; Úrbez‐Torres, 2011) and sexual fruiting bodies are rarely found in the field (van Niekerk et al., 2004). Nevertheless, it has been reported that compatible genotypes can exchange genetic material through parasexual recombination (Leslie, 1993). The genetic diversity of D. seriata populations on grapevines in Spain was investigated using the inter‐simple sequence repeat (ISSR) technique (Elena et al., 2015). This study found that isolates from different geographic origins or from different hosts were not grouped in genetically distinct clusters. This suggests that the isolates are genetically similar regardless of their geographic and host origin. A similar conclusion was reported by Phillips et al. (2007), who found no correlation between the host origin of different D. seriata isolates and the clustering structure obtained from a phylogenetic study of this species based on ITS sequence data. The pathogenicity and virulence of D. seriata is usually evaluated through its ability to cause brown necrotic lesions in the wood and the length of these lesions, respectively, there are conflicting reports as to the virulence of D. seriata on hosts form different geographic locations. These differences may be due to variations in virulence between strains, or they may be a result of the incomplete knowledge of the taxonomy of the genus, which in turn hampers accurate species recognition and identification.

                                                                                                                                                                                                                                                                                                                                                                                                  Life-cycle

                                                                                                                                                                                                                                                                                                                                                                                                  The fungus overwinters in fruiting bodies (pycnidia and perithecia) on dead bark, dead twigs or mummified fruit. It has also been demonstrated to survive endophytically inside some hosts, where it can invade almost any dead, woody tissues.

                                                                                                                                                                                                                                                                                                                                                                                                  In the spring, pycnidia and perithecia are trigged to release conidia and ascospores, under high humidity and during wet periods throughout the growing season. The spores are dispersed by splashing rains, wind and insects. The pathogen invades the tissue primarily through wounds, although in some hosts entry through natural openings such as lenticels and stomata is possible as well as direct penetration. Depending upon the host, the conidia can infect a variety of organs including leaves, the calyxes of blossoms, tiny fruit, and wounds in twigs and limbs. Infections of fruit and wood may not become visible for several weeks. The spores germinate at temperatures between 15 and 37°C and grow between 5 and 37°C. Infection is favoured by conditions that can stress the plant such as drought, frost damage, hail damage, poor nutrition and poor pruning practices. 

                                                                                                                                                                                                                                                                                                                                                                                                  Climate

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                                                                                                                                                                                                                                                                                                                                                                                                  ClimateStatusDescriptionRemark
                                                                                                                                                                                                                                                                                                                                                                                                  C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C
                                                                                                                                                                                                                                                                                                                                                                                                  Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
                                                                                                                                                                                                                                                                                                                                                                                                  Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)
                                                                                                                                                                                                                                                                                                                                                                                                  Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
                                                                                                                                                                                                                                                                                                                                                                                                  Ds - Continental climate with dry summer Preferred Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)
                                                                                                                                                                                                                                                                                                                                                                                                  Dw - Continental climate with dry winter Preferred Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)
                                                                                                                                                                                                                                                                                                                                                                                                  Df - Continental climate, wet all year Preferred Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)

                                                                                                                                                                                                                                                                                                                                                                                                  Air Temperature

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                                                                                                                                                                                                                                                                                                                                                                                                  Parameter Lower limit Upper limit
                                                                                                                                                                                                                                                                                                                                                                                                  Mean annual temperature (ºC) 5 37

                                                                                                                                                                                                                                                                                                                                                                                                  Natural enemies

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                                                                                                                                                                                                                                                                                                                                                                                                  Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
                                                                                                                                                                                                                                                                                                                                                                                                  Aureobasidium pullulans Antagonist Pinto et al.; 2018
                                                                                                                                                                                                                                                                                                                                                                                                  Bacillus subtilis Antagonist not specific
                                                                                                                                                                                                                                                                                                                                                                                                  Fusarium proliferatum Antagonist Mondello et al.; 2019
                                                                                                                                                                                                                                                                                                                                                                                                  Gibberella baccata Antagonist
                                                                                                                                                                                                                                                                                                                                                                                                  Trichoderma asperellum Antagonist not specific Vineyards, nursery

                                                                                                                                                                                                                                                                                                                                                                                                  Notes on Natural Enemies

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                                                                                                                                                                                                                                                                                                                                                                                                  Many of the host plants attacked by D. seriata are infected through wounds, therefore much of the research into natural enemies has concentrated on wound protection products. The majority of the commercial products available make use of species of Trichoderma which are antagonistic to D. seriata. Despite extensive research and increased availability, there has been limited adoption of biocontrol agents in commercial agriculture, mainly due to inconsistent and unpredictable performance. For grapevine pruning wounds the physiological state (dormant or active) of the vines at pruning can affect how the Trichoderma spp. colonises the wound. Delayed pruning may result in excessive sap bleeding which may dislodge any wound protectants applied immediately after pruning.

                                                                                                                                                                                                                                                                                                                                                                                                  Means of Movement and Dispersal

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                                                                                                                                                                                                                                                                                                                                                                                                  Natural dispersal

                                                                                                                                                                                                                                                                                                                                                                                                  Ascospores (teleomorph state) are spread by both air or water, whereas the conidia (anamorph state) are mainly splash-dispersed. Conidia are thought to be the primary propagule responsible for the short-distance spread between woody hosts.

                                                                                                                                                                                                                                                                                                                                                                                                  Vector transmission

                                                                                                                                                                                                                                                                                                                                                                                                  Insect vector transmission of conidia has been reported by Holmes and Rich (1970) who found that the convergent lady beetle (Hippodamia convergens), a common inhabitant of north American fruit orchards during the period of flower pollination, could move viable conidia of Physalospora obtuse (syn = Diplodia seriata) around the orchards. Similarly, Epstein et al. (2008) recorded conidia of D. seriata on rove beetles (Staphilinidae) collected from pruning wounds in California vineyards and Panzavolta et al. (2018) also found conidia of D. seriata on the bodies of wood boring beetles (Coraebus fasciatus [C. florentinus], Cerambyx welensii and Purpuricenus kaehleri) in oak woodlands in Italy. It is quite likely that other insects are also capable of passively transporting spores between hosts on their legs and bodies.

                                                                                                                                                                                                                                                                                                                                                                                                  Accidental introduction

                                                                                                                                                                                                                                                                                                                                                                                                  Members of the Botryosphaeriaceae including D. seriata have undoubtedly been spread around the world on traded agricultural plants and ornamentals (Burgess et al., 2016; Crous et al., 2016). The latent infection, or endophytic phase on some hosts implies that the fungus can easily pass undetected by quarantine systems in traded living plants, fruits and other plant parts associated with trade and transport. In a study undertaken in New South Wales, Australia, D. seriata infection occurred in the rootstock source plant canes of 95% of the canes sampled (Whitelaw-Weckert et al., 2013). Similarly, D. seriata was also reported in the basal and central parts of grapevine rootstock canes in New Zealand (Billones et al., 2010).

                                                                                                                                                                                                                                                                                                                                                                                                  Seedborne Aspects

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                                                                                                                                                                                                                                                                                                                                                                                                  There are no references in the literature of D. seriata being seed transmitted, however this species, in common with other members of the Botryosphaeriaceae, can live endophytically inside plants and latent infection of fruits is commonly reported for D. seriata. Despite there being no records of seed transmission of D. seriata, there is evidence that at least some members of the Botryosphaeriaceae family can be transmitted in seed (Gure et al., 2005), however, there is little evidence that these seed infections result in systemic infections in the plants as they develop (Slippers and Wingfield, 2007).

                                                                                                                                                                                                                                                                                                                                                                                                  Pathway Causes

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                                                                                                                                                                                                                                                                                                                                                                                                  CauseNotesLong DistanceLocalReferences
                                                                                                                                                                                                                                                                                                                                                                                                  Breeding and propagation Yes Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Crop production Yes Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Escape from confinement or garden escape Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Forestry Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Hitchhiker Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Nursery trade Yes Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Ornamental purposes Yes Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Timber trade Yes Yes

                                                                                                                                                                                                                                                                                                                                                                                                  Pathway Vectors

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                                                                                                                                                                                                                                                                                                                                                                                                  VectorNotesLong DistanceLocalReferences
                                                                                                                                                                                                                                                                                                                                                                                                  Germplasm Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Host and vector organisms Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Plants or parts of plants Yes Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Wind Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Water Yes

                                                                                                                                                                                                                                                                                                                                                                                                  Plant Trade

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                                                                                                                                                                                                                                                                                                                                                                                                  Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
                                                                                                                                                                                                                                                                                                                                                                                                  Bark Yes Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
                                                                                                                                                                                                                                                                                                                                                                                                  Flowers/Inflorescences/Cones/Calyx Yes
                                                                                                                                                                                                                                                                                                                                                                                                  Fruits (inc. pods) Yes Pest or symptoms usually invisible
                                                                                                                                                                                                                                                                                                                                                                                                  Leaves Yes Yes Pest or symptoms usually visible to the naked eye
                                                                                                                                                                                                                                                                                                                                                                                                  Roots Yes Pest or symptoms usually invisible
                                                                                                                                                                                                                                                                                                                                                                                                  Stems (above ground)/Shoots/Trunks/Branches Yes Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
                                                                                                                                                                                                                                                                                                                                                                                                  Wood Yes Yes Pest or symptoms usually invisible

                                                                                                                                                                                                                                                                                                                                                                                                  Wood Packaging

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                                                                                                                                                                                                                                                                                                                                                                                                  Wood Packaging liable to carry the pest in trade/transportTimber typeUsed as packing
                                                                                                                                                                                                                                                                                                                                                                                                  Solid wood packing material with bark No
                                                                                                                                                                                                                                                                                                                                                                                                  Solid wood packing material without bark No
                                                                                                                                                                                                                                                                                                                                                                                                  Wood Packaging not known to carry the pest in trade/transport
                                                                                                                                                                                                                                                                                                                                                                                                  Loose wood packing material
                                                                                                                                                                                                                                                                                                                                                                                                  Processed or treated wood

                                                                                                                                                                                                                                                                                                                                                                                                  Impact Summary

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                                                                                                                                                                                                                                                                                                                                                                                                  CategoryImpact
                                                                                                                                                                                                                                                                                                                                                                                                  Cultural/amenity Negative
                                                                                                                                                                                                                                                                                                                                                                                                  Economic/livelihood Negative
                                                                                                                                                                                                                                                                                                                                                                                                  Environment (generally) Negative

                                                                                                                                                                                                                                                                                                                                                                                                  Economic Impact

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                                                                                                                                                                                                                                                                                                                                                                                                  D. seriata has been implicated in causing economic damage to fruit crops, forestry and ornamental plants around the world. It is recognised as being one of the most prominent pathogens involved in grapevine trunk disease (GTD, or grapevine decline). This pathology can result in the death of adult plants and therefore it produces severe economic losses all around the world. The worldwide economic cost for the replacement of dead grapevines is roughly estimated to be more than 1.5 billion dollars per year (Hofstetter et al., 2012). In recent years, GTD has been increasing in importance as it is increasingly found affecting plants at a younger age being commonly reported in vineyards that are over 7-year-old (Díaz and LaTorre, 2013). According to a survey led by the French Directorate General for Food (DGAL) in 2012, nearly 13% of French vineyards were affected by trunk diseases (Grosman and Doublet, 2012). In 2014 these diseases lowered the French potential wine production by 13%, according to the agriculture ministry and French Wine Institute (IFV). The IFV estimate that GTD is costing France the equivalent of 1bn euros ($1.14bn) annually in lost wine production, and more than 100,000 hectares of vineyard was lost in 2014. In California USA, Eutypa dieback and Botryosphaeria canker were estimated to have caused over 260 million dollars of damage in reduced yields and increase production costs (Siebert, 2001).

                                                                                                                                                                                                                                                                                                                                                                                                  D. seriata is also damaging to the production of apples and pears. In south-eastern USA, fruit losses of between 25 and 50% have been reported due to black rot (Brown and Britton, 1986). Similarly, since 2007 when D. seriata was first recorded on apples in the Lower Elbe region (northern Germany), annual crop losses of over 5% at harvest have been reported (Brockamp and Weber, 2014).

                                                                                                                                                                                                                                                                                                                                                                                                  Risk and Impact Factors

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                                                                                                                                                                                                                                                                                                                                                                                                  Invasiveness
                                                                                                                                                                                                                                                                                                                                                                                                  • Invasive in its native range
                                                                                                                                                                                                                                                                                                                                                                                                  • Proved invasive outside its native range
                                                                                                                                                                                                                                                                                                                                                                                                  • Has a broad native range
                                                                                                                                                                                                                                                                                                                                                                                                  • Abundant in its native range
                                                                                                                                                                                                                                                                                                                                                                                                  • Highly adaptable to different environments
                                                                                                                                                                                                                                                                                                                                                                                                  • Is a habitat generalist
                                                                                                                                                                                                                                                                                                                                                                                                  • Capable of securing and ingesting a wide range of food
                                                                                                                                                                                                                                                                                                                                                                                                  • Has propagules that can remain viable for more than one year
                                                                                                                                                                                                                                                                                                                                                                                                  • Reproduces asexually
                                                                                                                                                                                                                                                                                                                                                                                                  Impact outcomes
                                                                                                                                                                                                                                                                                                                                                                                                  • Host damage
                                                                                                                                                                                                                                                                                                                                                                                                  • Negatively impacts agriculture
                                                                                                                                                                                                                                                                                                                                                                                                  • Negatively impacts forestry
                                                                                                                                                                                                                                                                                                                                                                                                  • Negatively impacts livelihoods
                                                                                                                                                                                                                                                                                                                                                                                                  Impact mechanisms
                                                                                                                                                                                                                                                                                                                                                                                                  • Competition - monopolizing resources
                                                                                                                                                                                                                                                                                                                                                                                                  • Pathogenic
                                                                                                                                                                                                                                                                                                                                                                                                  Likelihood of entry/control
                                                                                                                                                                                                                                                                                                                                                                                                  • Highly likely to be transported internationally accidentally
                                                                                                                                                                                                                                                                                                                                                                                                  • Highly likely to be transported internationally deliberately
                                                                                                                                                                                                                                                                                                                                                                                                  • Difficult to identify/detect as a commodity contaminant
                                                                                                                                                                                                                                                                                                                                                                                                  • Difficult to identify/detect in the field
                                                                                                                                                                                                                                                                                                                                                                                                  • Difficult/costly to control

                                                                                                                                                                                                                                                                                                                                                                                                  Diagnosis

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                                                                                                                                                                                                                                                                                                                                                                                                  D. seriata can be identified using classic and molecular biology methods. In classic identification, the fungus is first isolated into pure culture using aseptic techniques and grown on standard agar, such as half strength Potato Dextrose Agar (PDA). Colony growth, colour, conidiophore, and conidial morphology are used to distinguish species (Crous et al., 2006; Urbez-Torres et al., 2006; Pitt et al., 2010). Classic identification of D. seriata relies on morphological features, however, recent taxonomic evaluations by several researchers have shown that these characteristics are variable and often overlap between species (Phillips et al., 2013).

                                                                                                                                                                                                                                                                                                                                                                                                  The accurate identification of Botryosphaeriaceae is therefore best achieved by DNA sequence data rather than relying on morphological descriptions. Phillips et al. (2013) recommended that at least two loci, the internal transcribed spacer (ITS) region, and the translation elongation factor 1-alpha (tef1α), be used for species separation. However, Slippers et al. (2013) recommended the use of four loci, including the ITS region, tef1α, beta-tubulin (tub), and the RNA polymerase II (rpb2), as these loci provide better resolution to distinguish cryptic species. Unfortunately, the amplification of rpb2 is challenging and subsequently there is lack of data for comparisons (Slippers et al., 2013). Procedures and protocols for DNA isolation and sequencing are explained in detail by Alves et al. (2004).

                                                                                                                                                                                                                                                                                                                                                                                                  Detection and Inspection

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                                                                                                                                                                                                                                                                                                                                                                                                  D. seriata is difficult to diagnose definitively based on symptomology alone as the symptoms it produces vary depending upon the host and environmental conditions and are often not unique. Moreover, D. seriata may be present in apparently healthy-looking plants as a latent infection, or colonise dead woody parts damaged by other fungal pathogens, insects or abiotic agents. Where present, the fruiting bodies of the fungus on mummified fruit, cankers or dead wood are a reasonable indication of its presence, however other closely related fungi can produce similar structures so this is not conclusive unless backed up by microscopic analysis, or better still, molecular testing.

                                                                                                                                                                                                                                                                                                                                                                                                  Similarities to Other Species/Conditions

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                                                                                                                                                                                                                                                                                                                                                                                                  This pathogen can induce cankers, diebacks, and fruit rots on a wide variety of hosts and these symptoms can easily be confused with those caused by many different fungal pathogens and abiotic disorders. Given the enormous host range of this pathogen, it is not within the scope of this document to attempt to describe all the possible organisms with which it could be confused. Therefore, two of the most important economic hosts, grapes and apples have been selected for further discussion.

                                                                                                                                                                                                                                                                                                                                                                                                  D. seriata has been implicated along with several other fungi in the condition known as ‘grape vine trunk disease’ (GTD). Affected vines show external symptoms that include a general and progressive decline (delayed budburst, dead buds, dieback, cankers, stunted development, chlorosis, apoplexy) as well as internal symptoms of brown streaking of vascular tissues and sectorial, or central necrosis. Based on the predominant organism responsible, GTD currently includes black foot, Esca, Eutypa dieback, Phomopsis dieback, Petri disease and Botryosphaeria dieback (Pascoe, 1998; Mugnai et al., 1999; Úrbez-Torres et al., 2006; Úrbez-Torres and Gubler, 2011; Lecomte et al., 2012Mondello et al., 2018).

                                                                                                                                                                                                                                                                                                                                                                                                  D. seriata is one of the most cited Botryosphaeriaceae species occurring on grapevines worldwide and is frequently associated with the ‘black dead arm’ disease of grapevine (Larignon et al., 2001; Úrbez‐Torres, 2011). However, it is just one of at least 20 different species in the Botryosphaeriaceae occurring in grapevines (Úrbez-Torres, 2011), the most common other species being Neofusicoccum parvum, Lasiodiplodia theobromae and D. mutila [Botryosphaeria stevensii]. Therefore, what is commonly referred to as Botryosphaeria dieback of grapes is a complex of different fungi that cannot be distinguished in the field based on symptomology. Moreover, symptoms such as dieback, dead spurs, stunted shoots, and bud mortality are shared with multiple trunk diseases that often occur in mixed infection within the vineyard and even within an individual vine. It is sometimes possible to distinguish Eutypa dieback from D. seriata dieback by the presence of foliar symptoms, but these are not always present.

                                                                                                                                                                                                                                                                                                                                                                                                  Differentiation of the different members of the Botryosphaeriaceae involved in GTD is possible using isolation and morphological examination using the light microscope, but this is complicated by the overlapping characteristics of these pathogens and molecular-based techniques are therefore the preferred method for accurately ascribing species.

                                                                                                                                                                                                                                                                                                                                                                                                  D. seriata is also an important disease of pome fruits, where it attacks the leaves, stems and fruits. Leaf lesions are referred to as frogeye spot due to the characteristic dark-brown concentric rings surrounded by a purple margin that develop around a light brown-to-grey centre, giving it a ‘frogeye’ appearance. These spots start off as small, purple specks that enlarge to form spots 3 to 6 mm in diameter. Black pycnidia, may develop on the upper surface in the centres of the older leaf spots which help to distinguish frogeye leaf spots from similar spots caused by spray injury. Stem symptoms of D. seriata begin as small, slightly sunken, reddish-brown areas that develop in the bark. These areas slowly enlarge and darken to form cankers with depressed centres and slightly raised and lobed margins. Cankers may also appear as a superficial roughening of the bark; or the bark may be killed and conspicuously cracked, especially at the margins. There are many potential causes of cankers in pome fruits, including other fungi, bacteria and mechanical injury, the development of black, pimple-like pycnidia and perithecia in older cankers is an indication that the canker may be caused by D. seriata, however cankers caused by another common apple pathogen Botryosphaeria dothidia cannot be told apart.

                                                                                                                                                                                                                                                                                                                                                                                                  D. seriata also produces a fruit rot known as black rot of apples and pears. Initially the fungus infects the fruit through wounds caused by insects, hail or growth cracks, particularly at the calyx end of the fruit. At first, a light brown spot forms on the fruit which enlarges and is surrounded by a concentric zonation of lighter and darker brown colours. The rotted fruit finally turns black. The fruit symptoms are difficult to tell apart from rots caused by Colletotrichum gloeosporioides [Glomerella cingualata] and C. acutatum (bitter rot), however the development of ‘pimple-like’ fruiting bodies (pycnidia) on the surface of rotted fruit can help to distinguish between these pathogens. The fungus B. dothidia also causes a rot of apples that is known by the common name ‘white rot’, to distinguish it from the black rot caused by D. seriata. In practice these two diseases on fruits can be very difficult to tell apart. However, with black rot of apple, the flesh in the decayed portion of the fruit remains firm and somewhat leathery and the surface of the spot is not sunken. Conversely, the decay caused by the white rot pathogen is soft and forms a slightly sunken lesion.

                                                                                                                                                                                                                                                                                                                                                                                                  Phillips et al. (2012) used molecular techniques to examine in detail the Botryosphaeriaceae attacking apples. This study revealed that D. seriata is a complex of species, two of which are associated with fruit rot and canker of apples and other Rosaceae, namely D. seriata and a species that they named D. intermedia. Both species are virtually indistinguishable based on morphology, which raises questions over previous reports regarding the black rot pathogen of apple (Phillips et al., 2012).

                                                                                                                                                                                                                                                                                                                                                                                                  Prevention and Control

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                                                                                                                                                                                                                                                                                                                                                                                                  Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

                                                                                                                                                                                                                                                                                                                                                                                                  Cultural and chemical management options for control of Botryosphaeria diseases are similar in many cropping systems including apple, blueberry, grape, peach and pistachio. Benzimidazoles, quinone outside inhibitors (QoI), and sterol biosynthesis inhibitors (DMI) are extensively used to treat the external symptoms of Botryosphaeria blight in apple (Brown and Britton, 1986), grape (Bester et al., 2007) and pistachio cropping systems (Ma et al., 2001Ma et al., 2002). These products are applied either prophylactically, or as treatments applied to pruning wounds, as these serve as important entry points for infection. In addition to the synthetic chemical pruning treatments there are also several commercially available biological and botanical wound treatments. The biological products have already been outlined in the section ‘Notes on natural enemies’ and mostly make use of Trichoderma fungal antagonists that are painted on to the wounds. In addition to these several botanical products have been tested for their ability to manage D. seriata infections in grapevines, these products include chitosan oligosaccharide, garlic extract and vanillin. In field experiments all three were able to significantly reduce infection in pruning wounds by D. seriata and P. chlamydospora, with the most effective treatment being a mix of all three (Cobos et al., 2015).

                                                                                                                                                                                                                                                                                                                                                                                                  Cultural control mostly relies on sanitation by reducing inoculum sources such as cankers, blighted shoots, mummified fruit, and pruning. In Californian vineyards delayed pruning is recommended as the current timing coincides with the highest periods of spore dispersal by fungi in the Botryosphaeriaceae.

                                                                                                                                                                                                                                                                                                                                                                                                  Host resistance

                                                                                                                                                                                                                                                                                                                                                                                                  Work is continuing to determine grape varieties with enhanced resistance to D. seriata and other members of the Botryosphaeriaceae. A study conducted by Guan et al. (2016) into the of genetic resistance of Vitaceae found differential susceptibility to wood necrosis caused by Neofusicoccum parvum and D. seriata. Several accessions of V. vinifera subsp. sylvestris, the ancestor of V. vinifera, were found to be more resistant to artificial inoculation than cultivars such as Chardonnay and Gewürztraminer. These findings suggest that creating new grapevine varieties with enhanced resistance to trunk pathogens is a realistic possibility.

                                                                                                                                                                                                                                                                                                                                                                                                  Similarly, the host resistance of apples to black rot has been investigated by several authors experimentally and in the field. Biggs et al. (2004) tested 23 apple varieties for resistance and was able to classify the cultivars into three relative susceptibility groups - most susceptible: ʻOrinʼ, ʻPristineʼ and Sunriseʼ; moderately  susceptible: ʻSun-crispʼ, ʻGinger Goldʼ, ʻSenshuʼ, ʻHoneycrispʼ, ʻPioneerMacʼ, ʻFortuneʼ, ʻNY 75414ʼ, ʻArletʼ, ʻGolden  Supremeʼ, ʻShizukaʼ, ʻCameoʼ, ʻSansaʼ and ʻYatakaʼ; and least susceptible: ʻCrestonʼ, ʻGolden  Deliciousʼ, ʻEnterpriseʼ, ʻGala  Supremeʼ, ʻBraeburnʼ, ʻGoldRushʼ and ʻFujiʼ. 

                                                                                                                                                                                                                                                                                                                                                                                                  For further information on the management of grapevine trunk disease, see Gramaje et al. (2018) and Mondello et al. (2018, 2019).

                                                                                                                                                                                                                                                                                                                                                                                                  References

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                                                                                                                                                                                                                                                                                                                                                                                                  Abreo, E., Martinez, S., Bettucci, L., Lupo, S., 2013. Characterization of Botryosphaeriaceae species associated with grapevines in Uruguay. Australasian Plant Pathology, 42(3), 241-249. doi: 10.1007/s13313-013-0200-8

                                                                                                                                                                                                                                                                                                                                                                                                  Akgul DS, Savas NG, Eskalen A, 2014. First report of wood canker caused by Botryosphaeria dothidea, Diplodia seriata, Neofusicoccum parvum, and Lasiodiplodia theobromae on grapevine in Turkey. Plant Disease, 98(4):568. http://apsjournals.apsnet.org/loi/pdis

                                                                                                                                                                                                                                                                                                                                                                                                  Alves, A., Correia, A., Luque, J., Phillips, A., 2004. Botryosphaeria corticola, sp. nov. on Quercus species, with notes and description of Botryosphaeria stevensii and its anamorph, Diplodia mutila. Mycologia, 96(3), 598-613. doi: 10.2307/3762177

                                                                                                                                                                                                                                                                                                                                                                                                  Ammad, F., Benchabane, M., Toumi, M., Belkacem, N., Guesmi, A., Ameur, C., Lecomte, P., Merah, O., 2014. Occurrence of Botryosphaeriaceae species associated with grapevine dieback in Algeria. Turkish Journal of Agriculture and Forestry, 38(6), 865-876. http://journals.tubitak.gov.tr/agriculture/

                                                                                                                                                                                                                                                                                                                                                                                                  Arzanlou M, Dokhanchi H, 2013. Morphological and molecular characterization of Diplodia seriata, the causal agent of canker and twig dieback disease on mulberry in Iran. Archives of Phytopathology and Plant Protection, 46(6):682-694. http://www.tandfonline.com/loi/gapp20

                                                                                                                                                                                                                                                                                                                                                                                                  Auger, J., Esterio, M., Ricke, G., Pérez, I., 2004. Black dead arm and basal canker of Vitis vinifera cv. Red Globe caused by Botryosphaeria obtusa in Chile. Plant Disease, 88(11), 1286. doi: 10.1094/PDIS.2004.88.11.1286A

                                                                                                                                                                                                                                                                                                                                                                                                  Barradas C, Phillips AJL, Correia A, Diogo E, Bragança H, Alves A, 2016. Diversity and potential impact of Botryosphaeriaceae species associated with Eucalyptus globulus plantations in Portugal. European Journal of Plant Pathology, 146(2):245-257. http://rd.springer.com/journal/10658

                                                                                                                                                                                                                                                                                                                                                                                                  Baskarathevan, J., Jaspers, M. V., Jones, E. E., Ridgway, H. J., 2013. Development of isolate-specific markers for Neofusicoccum parvum and N. luteum and their use to study rainwater splash dispersal in the vineyard. Plant Pathology, 62(3), 501-509. doi: 10.1111/j.1365-3059.2012.02675.x

                                                                                                                                                                                                                                                                                                                                                                                                  Beer, M., Brockamp, L., Weber, R. W. S., 2015. Control of sooty blotch and black rot of apple through removal of fruit mummies. Folia Horticulturae, 27(1), 43-51. https://www.degruyter.com/downloadpdf/j/fhort.2015.27.issue-1/fhort-2015-0013/fhort-2015-0013.pdf

                                                                                                                                                                                                                                                                                                                                                                                                  Ben Ghnaya-Chakroun A, Rezgui A, Vallance J, Kharoubi I, Dridi M, Hajlaoui MR, Rey P, Sadfi-Zouaoui N, 2014. First molecular and biochemical characterization of Phomopsis viticola and Diploidia seriata two pathogens of esca and black dead arm diseases of grapevine in the northern region of the Tunisia. International Journal of Current Microbiology and Applied Sciences, 3(8):977-987. http://www.ijcmas.com/vol-3-8/Asma%20Ben%20Ghnaya-Chakroun,%20et%20al.pdf

                                                                                                                                                                                                                                                                                                                                                                                                  Bester, W., Crous, P. W., Fourie, P. H., 2007. Evaluation of fungicides as potential grapevine pruning wound protectants against Botryosphaeria species. Australasian Plant Pathology, 36(1), 73-77. doi: 10.1071/AP06086

                                                                                                                                                                                                                                                                                                                                                                                                  Biggs, A. R., Miller, S. S., 2004. Relative susceptibility of selected apple cultivars to fruit rot caused by Botryosphaeria obtusa. HortScience, 39(2), 303-306.

                                                                                                                                                                                                                                                                                                                                                                                                  Billones R, Jones EE, Ridgway HJ, Jaspers MV, 2010. Botryosphaeriaceae infection in New Zealand grapevine nursery plant materials. Phytopathologia Mediterranea, 49, 115.

                                                                                                                                                                                                                                                                                                                                                                                                  Billones-Baaijens, R., Savocchia, S., 2019. A review of Botryosphaeriaceae species associated with grapevine trunk diseases in Australia and New Zealand. Australasian Plant Pathology, 48(1), 3-18. doi: 10.1007/s13313-018-0585-5

                                                                                                                                                                                                                                                                                                                                                                                                  Bobev SG, Lopes J, Phillips AJL, 2008. First report of Diplodia seriata causing shoot blight and cankers of Cotoneaster salicifolius in Bulgaria. Plant Disease, 92(6):976. HTTP://www.apsnet.org

                                                                                                                                                                                                                                                                                                                                                                                                  Britton, K. O., Hendrix, F. F., 1982. Three species of Botryosphaeria cause peach tree gummosis in Georgia. Plant Disease, 66(12), 1120-1121. doi: 10.1094/PD-66-1120

                                                                                                                                                                                                                                                                                                                                                                                                  Britton, K. O., Hendrix, F. F., 1986. Population dynamics of Botryosphaeria spp. in peach gummosis cankers. Plant Disease, 70(2), 134-136. doi: 10.1094/PD-70-134

                                                                                                                                                                                                                                                                                                                                                                                                  Brockamp, L., Weber, R. W. S., 2014. Black rot (Diplodia seriata) in organic apple production - infection biology and disease control strategies. In: Ecofruit. 16th International Conference on Organic-Fruit Growing: Proceedings, 17-19 February 2014, Hohenheim, Germany [Ecofruit. 16th International Conference on Organic-Fruit Growing: Proceedings, 17-19 February 2014, Hohenheim, Germany], Weinsberg, Germany: Fördergemeinschaft Ökologischer Obstbau e.V. (FÖKO). 77-82.

                                                                                                                                                                                                                                                                                                                                                                                                  Brown, E. A., II, Britton, K. O., 1986. Botryosphaeria diseases of apple and peach in the southeastern United States. Plant Disease, 70(5), 480-484. doi: 10.1094/PD-70-480

                                                                                                                                                                                                                                                                                                                                                                                                  Brown-Rytlewski, D. E., McManus, P. S., 2000. Virulence of Botryosphaeria dothidea and Botryosphaeria obtusa on apple and management of stem cankers with fungicides. Plant Disease, 84(9), 1031-1037. doi: 10.1094/PDIS.2000.84.9.1031

                                                                                                                                                                                                                                                                                                                                                                                                  Burgess, T. I., Crous, C. J., Slippers, B., Hantula, J., Wingfield, M. J., 2016. Tree invasions and biosecurity: eco-evolutionary dynamics of hitchhiking fungi. AoB Plants, 8, plw076. doi: 10.1093/aobpla/plw076

                                                                                                                                                                                                                                                                                                                                                                                                  CABI/EPPO, 2005. Botryosphaeria obtusa. Distribution Maps of Plant Diseases, No. 945. Wallingford, UK: CAB International

                                                                                                                                                                                                                                                                                                                                                                                                  Castillo-Pando, M., Somers, A., Green, C. D., Priest, M., Sriskanthades, M., 2001. Fungi associated with dieback of Semillon grapevines in the Hunter Valley of New South Wales. Australasian Plant Pathology, 30(1), 59-63. doi: 10.1071/AP00068

                                                                                                                                                                                                                                                                                                                                                                                                  Chattaoui M, Rhouma A, Msallem M, Pérez M, Moral J, Trapero A, 2012. First report of Botryosphaeria obtusa as causal agent of olive tree branch dieback in Tunisia. Plant Disease, 96(6):905. http://apsjournals.apsnet.org/loi/pdis

                                                                                                                                                                                                                                                                                                                                                                                                  Choudhury R, Modi P, Hanstad J, Elkins R, Gubler WD, 2014. First report of Diplodia seriata causing pear branch canker dieback in California. Plant Disease, 98(5):688-689. http://apsjournals.apsnet.org/loi/pdis

                                                                                                                                                                                                                                                                                                                                                                                                  Cobos, R., Mateos, R. M., Álvarez-Pérez, J. M., Olego, M. A., Sevillano, S., González-García, S., Garzón-Jimeno, E., Coque, J. J. R., 2015. Effectiveness of natural antifungal compounds in controlling infection by grapevine trunk disease pathogens through pruning wounds. Applied and Environmental Microbiology, 81(18), 6474-6483. doi: 10.1128/AEM.01818-15

                                                                                                                                                                                                                                                                                                                                                                                                  Crespo, M., Moral, J., Michailides, T. J., Trouillas, F. P., 2018. First report of black rot on apple fruit caused by Diplodia seriata in California. Plant Disease, 102(4), 824-825. http://apsjournals.apsnet.org/loi/pdis doi: 10.1094/PDIS-07-17-1023-PDN

                                                                                                                                                                                                                                                                                                                                                                                                  Crous PW, Slippers B, Wingfield MJ, Rheeder J, Marasas WFO, Phillips AJL, Alves A, Burgess T, Barber P, Groenewald JZ, 2006. Phylogenetic lineages in the Botryosphaeriaceae. Stud. Mycol, 55, 235-253.

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                                                                                                                                                                                                                                                                                                                                                                                                  Taylor A, St J Hardy G E, Wood P, Burgess T, 2005. Identification and pathogenicity of Botryosphaeria species associated with grapevine decline in Western Australia. Australasian Plant Pathology. 34 (2), 187-195. DOI:10.1071/AP05018

                                                                                                                                                                                                                                                                                                                                                                                                  Trapman M, Maxin P, Weber R W S, 2008. Diplodia seriata, cause of black fruit rot in organically grown apples in Holland, Belgium and Northern Germany. In: Ecofruit. 13th International Conference on Cultivation Technique and Phytopathological Problems in Organic Fruit-Growing. Proceedings of the conference, Weinsberg, Germany, 18-20 February 2008. [ed. by Fördergemeinschaft Ökologischer Obstbau e V]. Weinsberg, Germany: Fördergemeinschaft Ökologischer Obstbau e.V. (FÖKO). 177-181.

                                                                                                                                                                                                                                                                                                                                                                                                  Twomey M C, Stone J K, Gent D H, 2016. Black Wilt of hop (Humulus lupulus) caused by Diplodia seriata in New York state. Plant Disease. 100 (4), 861. DOI:10.1094/PDIS-10-15-1140-PDN

                                                                                                                                                                                                                                                                                                                                                                                                  Úrbez-Torres J R, Boulé J, O'Gorman D T, 2016. First report of Diplodia seriata and D. mutila causing apple dieback in British Columbia. Plant Disease. 100 (6), 1243-1244. DOI:10.1094/PDIS-11-15-1358-PDN

                                                                                                                                                                                                                                                                                                                                                                                                  Úrbez-Torres J R, Leavitt G M, Guerrero J C, Guevara J, Gubler W D, 2008. Identification and pathogenicity of Lasiodiplodia theobromae and Diplodia seriata, the causal agents of bot canker disease of grapevines in Mexico. Plant Disease. 92 (4), 519-529. DOI:10.1094/PDIS-92-4-0519

                                                                                                                                                                                                                                                                                                                                                                                                  Úrbez-Torres J R, Leavitt G M, Voegel T M, Gubler W D, 2006. Identification and distribution of Botryosphaeria spp. associated with grapevine cankers in California. Plant Disease. 90 (12), 1490-1503. HTTP://www.apsnet.org DOI:10.1094/PD-90-1490

                                                                                                                                                                                                                                                                                                                                                                                                  Urbez-Torres J R, Peduto F, Striegler R K, Urrea-Romero K E, Rupe J C, Cartwright R D, Gubler W D, 2012. Characterization of fungal pathogens associated with grapevine trunk diseases in Arkansas and Missouri. Fungal Diversity. 52 (1), 169-189. http://www.springerlink.com/content/r04710p5uv65j716/ DOI:10.1007/s13225-011-0110-4

                                                                                                                                                                                                                                                                                                                                                                                                  Valencia D, Torres C, Camps R, López E, Celis-Diez J L, Besoain X, 2015. Dissemination of Botryosphaeriaceae conidia in vineyards in the semiarid Mediterranean climate of the Valparaíso region of Chile. Phytopathologia Mediterranea. 54 (2), 394-402. http://www.fupress.net/index.php/pm/article/view/16055/15761

                                                                                                                                                                                                                                                                                                                                                                                                  Vico I, Žebeljan A, Vučković N, Vasić M, Duduk N, 2017. First report of Diplodia seriata causing postharvest rot of quince fruit in Serbia. Plant Disease. 101 (10), 1823. DOI:10.1094/pdis-04-17-0484-pdn

                                                                                                                                                                                                                                                                                                                                                                                                  White C L, Halleen F, Fischer M, Mostert L, 2011. Characterisation of the fungi associated with esca diseased grapevines in South Africa. Phytopathologia Mediterranea. 50 (Supplement), 204-223. http://www.fupress.net/index.php/pm/article/view/8983

                                                                                                                                                                                                                                                                                                                                                                                                  Zhang M, Zhang Y K, Geng Y H, Zang R, Wu H Y, 2017. First report of Diplodia seriata causing twig dieback of english walnut in China. Plant Disease. 101 (6), 1036. DOI:10.1094/pdis-04-16-0458-pdn

                                                                                                                                                                                                                                                                                                                                                                                                  Zhu ZongCai, Zhang WangBin, Yue Juan, Yi ZiBo, 2020. Identification of Diplodia seriata causing black spot on branches and trunks of apple trees in Xinjiang. Acta Phytopathologica Sinica. 50 (3), 373-376. DOI:10.13926/j.cnki.apps.000325

                                                                                                                                                                                                                                                                                                                                                                                                  Zlatković M, Keča N, Wingfield M J, Jami F, Slippers B, 2016. Botryosphaeriaceae associated with the die-back of ornamental trees in the Western Balkans. Antonie van Leeuwenhoek. 109 (4), 543-564. http://link.springer.com/article/10.1007%2Fs10482-016-0659-8

                                                                                                                                                                                                                                                                                                                                                                                                  Zlatkovic M, Wingfield M, Keca N, Jami F, Slipper B, 2014. Diplodia seriata on Ligustrum vulgare L. in Serbia. In: Proceedings of the VII Congress on plant protection, 24-28 November 2014, Zlatibor, Serbia [VII Congress on plant protection, 24-28 November 2014, Zlatibor, Serbia],

                                                                                                                                                                                                                                                                                                                                                                                                  Contributors

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                                                                                                                                                                                                                                                                                                                                                                                                  20/05/20 Original text:

                                                                                                                                                                                                                                                                                                                                                                                                  Robert Reeder, CABI E-UK, Bakeham Lane, Egham, Surrey,  TW20 9TY, UK

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