Invasive Species Compendium

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Datasheet

Phomopsis vexans
(Phomopsis blight of eggplant)

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Datasheet

Phomopsis vexans (Phomopsis blight of eggplant)

Summary

  • Last modified
  • 27 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Preferred Scientific Name
  • Phomopsis vexans
  • Preferred Common Name
  • Phomopsis blight of eggplant
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Fungi
  •     Phylum: Ascomycota
  •       Subphylum: Pezizomycotina
  •         Class: Sordariomycetes
  • Summary of Invasiveness
  • P. vexans is a pynicidial anamorph with a teleomorph in the genus Diaporthe. Easily seedborne and producing large numbers of conidia, it causes disease in Solanum melongena [aubergine/brinjal/eggp...

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Pictures

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PictureTitleCaptionCopyright
Pycnidia on Solanum melongena fruit. Original x7.5.
TitlePycnidia
CaptionPycnidia on Solanum melongena fruit. Original x7.5.
CopyrightUSDA-ARS/Systematic Mycology & Microbiology Laboratory
Pycnidia on Solanum melongena fruit. Original x7.5.
PycnidiaPycnidia on Solanum melongena fruit. Original x7.5. USDA-ARS/Systematic Mycology & Microbiology Laboratory
Pycnidia on Solanum melongena fruit. Original x10.
TitlePycnidia
CaptionPycnidia on Solanum melongena fruit. Original x10.
CopyrightUSDA-ARS/Systematic Mycology & Microbiology Laboratory
Pycnidia on Solanum melongena fruit. Original x10.
PycnidiaPycnidia on Solanum melongena fruit. Original x10. USDA-ARS/Systematic Mycology & Microbiology Laboratory
Cross section of pycnidium from lesion on Solanum melongena fruit. Original x200.
TitlePycnidium
CaptionCross section of pycnidium from lesion on Solanum melongena fruit. Original x200.
CopyrightUSDA-ARS/Systematic Mycology & Microbiology Laboratory
Cross section of pycnidium from lesion on Solanum melongena fruit. Original x200.
PycnidiumCross section of pycnidium from lesion on Solanum melongena fruit. Original x200. USDA-ARS/Systematic Mycology & Microbiology Laboratory
Beta-conidia from pycnidium. Original x400. Note scale bar.
TitleBeta-conidia
CaptionBeta-conidia from pycnidium. Original x400. Note scale bar.
CopyrightUSDA-ARS/Systematic Mycology & Microbiology Laboratory
Beta-conidia from pycnidium. Original x400. Note scale bar.
Beta-conidiaBeta-conidia from pycnidium. Original x400. Note scale bar.USDA-ARS/Systematic Mycology & Microbiology Laboratory
Beta-conidia from pycnidium. Original x1000. Note scale bar.
TitleBeta-conidia
CaptionBeta-conidia from pycnidium. Original x1000. Note scale bar.
CopyrightUSDA-ARS/Systematic Mycology & Microbiology Laboratory
Beta-conidia from pycnidium. Original x1000. Note scale bar.
Beta-conidiaBeta-conidia from pycnidium. Original x1000. Note scale bar.USDA-ARS/Systematic Mycology & Microbiology Laboratory

Identity

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Preferred Scientific Name

  • Phomopsis vexans (Sacc. & P. Syd.) Harter 1914

Preferred Common Name

  • Phomopsis blight of eggplant

Other Scientific Names

  • Ascochyta hortorum (Speg.) C.O. Sm. 1905
  • Diaporthe vexans (Sacc. & P. Syd.) Gratz 1942
  • Phoma solani Halst. 1892
  • Phoma vexans Sacc. & P. Syd. 1899
  • Phyllosticta hortorum Speg. 1881

International Common Names

  • English: brown spot of eggplant; fruit rot of eggplant; Phomopsis leaf blight; Phomopsis rot of eggplant; stem blight of eggplant; tipover of eggplant
  • Spanish: Phoma (berenjena)
  • French: pourriture de l'aubergine; pourriture des fruits de l'aubergine; taches foliaires de l'aubergine

Local Common Names

  • Germany: Blattfleckenkrankheit; Eierfrucht; Eierpflanze Fruchtfaeule

EPPO code

  • PHOPVE (Phomopsis vexans)

Summary of Invasiveness

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P. vexans is a pynicidial anamorph with a teleomorph in the genus Diaporthe. Easily seedborne and producing large numbers of conidia, it causes disease in Solanum melongena [aubergine/brinjal/eggplant], its only significant host. This ranges from poor seed germination and damping-off of seedlings, to leaf and stem lesions and to fruit rot, both in the field and after harvest. The fungus has been reported from widely distributed areas of most continents, but only a few of those are in Europe and Africa, even though the climates are favourable. Seed transmission may explain its broad historical distribution, but limitation of its host range to a non-staple vegetable crop can allow for its avoidance and eradication by cultural methods. As a result, perhaps, it does not appear often on lists of restricted pathogens, even though it may cause yield losses of more than 50%.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Fungi
  •         Phylum: Ascomycota
  •             Subphylum: Pezizomycotina
  •                 Class: Sordariomycetes
  •                     Subclass: Sordariomycetidae
  •                         Order: Diaporthales
  •                             Family: Diaporthaceae
  •                                 Genus: Phomopsis
  •                                     Species: Phomopsis vexans

Notes on Taxonomy and Nomenclature

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The existence of several pycnidial fungi causing leaf spots on Solanum melongena [eggplant/aubergine] resulted in difficulties with the identification of each one. Spegazzini (1881) described a fungus occurring on leaves of S. melongena in Italy as Phyllosticta hortorum. Halsted (1892) described the same fungus on leaves and fruits of eggplants in New Jersey, USA, as Phoma solani. However, the name P. solani was already applied by Cooke and Harkness to another fungus on another host, and therefore Saccardo and Sydow (1899) substituted Phoma vexans. In 1905, Smith, observing septate conidia in the USA, proposed the name Ascochyta hortorum instead of P. hortorum. In Italy, Voglino (1907) studied a fungus on aubergine and agreed with Smith, concluding that the fungus described by Spegazzini as P. hortorum was an Ascochyta.

Cross-inoculation tests and morphological studies indicated to Harter (1914) that Phoma solani and Phyllosticta hortorum were the same species. He also concluded that the genus to which the fungus belonged was not Phoma, Phyllosticta or Ascochyta, but Phomopsis. Unlike the previous workers, Harter observed and described the beta conidia (“stylospores”) characteristic of the genus. He proposed the name Phomopsis vexans for the fungus, and Spegazzini agreed that the American isolates were different from P. hortorum (Harter, 1914).

Whereas the anamorph on eggplant that produces both alpha and beta conidia is a true Phomopsis (Uecker, 1988), the species Phoma hortorum Speg. and Ascochyta hortorum (Speg.) C.O. Sm. have recently been synonymized with Phoma exigua Desm. var. exigua; a weak pathogen of many plants that may be found in older lesions caused by other fungi (Boerema et al., 2004). Smith and Voglino were apparently observing yet another species, Ascochyta lycospersici Brunaud (Harter, 1914).

The teleomorph of the fungus has not yet been encountered in nature. Gratz (1942) observed perithecia on 2% potato dextrose agar in culture, and assigned the name Diaporthe vexans. The current view is that D. vexans is the teleomorph of P. vexans (Rehner and Uecker, 1994). Nevertheless, although the known connections of some Phomopsis species are to teleomorphs in the genus Diaporthe, the name D. vexans (Sacc. & P. Syd.) Gratz, although previously regarded as illegitimate, is now considered to apply only to the anamorph (Punithalingam and Holliday, 1972).

Furthermore, species concepts in Phomopsis have often been based on host specificity, but the phylogeny based on molecular data obtained so far indicates that either species have broader host ranges, or significant changes (“jumps”) between hosts have occurred in species evolution (Rehner and Uecker, 1994). Therefore, additional molecular evidence might connect the eggplant pathogen to an older species in either the anamorph genus or the teleomorph genus.

Description

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Conidiomata pycnidial, subepidermal, erumpent, dark, thick-walled, flattened to globose, varying in size, often 100-300 µm diameter, with or without a beak; beak to 76 µm. Phialides hyaline, simple or branched, sometimes septate, 10-16 µm long, arising from the innermost layer of cells lining the cavity. Alpha conidia hyaline, aseptate, sub-cylindrical, 5-8 x 2-3 µm. Beta conidia filiform, curved, hyaline, septate, 18-32 x 0.5-2.0 µm, non-germinating. Hyphae hyaline, septate, 2.5-4.0 µm diameter (see Edgerton and Moreland, 1921; Sherf and MacNab, 1986; Singh, 1987).

Ascomata perithecial, in culture usually in clusters, 130-350 µm diameter, beaked; beaks sinuous, carbonaceous, irregular, 80-500 µm long. Asci clavate, sessile, 24-44 x 5-12 µm, eight-spored. Ascospores biseriate, hyaline, narrowly ellipsoid to bluntly fusoid, one-septate, constricted at the septum, 9-12 x 3.0-4.5 µm (see Gratz, 1942).

A number of workers have studied the factors affecting growth and sporulation of the fungus in culture (Gratz, 1942; Pawar and Patel, 1957; Lapis and Deangkinay, 1967; Panwar and Chand, 1968; Hasija and Chowdhury, 1980; Singh and Chand, 1986; Islam and Pan, 1990a).

Distribution

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P. vexans has been reported from many areas in the warmer parts of most continents, but is unknown in Europe, except in Romania (Smith et al., 1988) and known in only a few African countries. It is probably native to southern Asia, the area of origin of the host Solanum melongena (eggplant/aubergine) (Prance and Nesbitt, 2005), where it is also reported to infect some wild Solanum species (Datar and Ashtaputre, 1988). It is readily transmitted in and on the seed (Porter, 1943; Vishunavat and Kumar, 1993) of a crop that is only grown in limited areas and this may explain its lack of a continuous distribution in the tropics and subtropics. The fungus could be introduced to a region within a seed lot, but then die out if its presence discouraged continuous local cultivation of S. melongena.

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

BangladeshPresentCABI/EPPO , 2008; Masuduzzaman et al., 2008
Brunei DarussalamPresentPeregrine and Kassim, 1982; CABI/EPPO , 2008Reported as causing a minor leaf spot
ChinaPresentTai, 1979; Teng ShuChün, 1996; CABI/EPPO , 2008
-FujianRestricted distributionTeng, 1932; CABI/EPPO , 2008
-GansuPresentGuo, 2005; CABI/EPPO , 2008
-GuangdongPresentBPI, US National Fungus Collections; Luo et al., 2004; CABI/EPPO , 2008
-GuangxiPresentTeng ShuChün, 1996; CABI/EPPO , 2008
-HebeiPresentCABI/EPPO , 2008
-HeilongjiangPresentCABI/EPPO , 2008
-Hong KongPresentLu et al., 2000; CABI/EPPO , 2008
-HubeiPresentCABI/EPPO , 2008
-HunanPresentCABI/EPPO , 2008
-JiangsuPresentBPI, US National Fungus Collections; Teng ShuChün, 1996; CABI/EPPO , 2008
-JiangxiPresentCABI/EPPO , 2008
-JilinPresentCABI/EPPO , 2008
-LiaoningPresentCABI/EPPO , 2008
-Nei MengguPresentCABI/EPPO , 2008
-NingxiaPresentGuo, 2005
-ShaanxiPresentCABI/EPPO , 2008
-ShandongPresentCABI/EPPO , 2008
-ShanxiPresentCABI/EPPO , 2008
-SichuanPresentCABI/EPPO , 2008
-XinjiangPresentCABI/EPPO , 2008
-YunnanPresentCABI/EPPO , 2008
-ZhejiangPresentCABI/EPPO , 2008
IndiaPresentMathur, 1979; CABI/EPPO , 2008
-Andaman and Nicobar IslandsPresentCABI/EPPO , 2008
-Andhra PradeshPresentCABI/EPPO , 2008
-AssamPresentAli and Saikia, 1993; CABI/EPPO , 2008
-BiharPresentMathur, 1979On Acacia
-ChandigarhPresentCABI/EPPO , 2008
-DelhiPresentPanwar et al., 1970; CABI/EPPO , 2008
-HaryanaPresentCABI/EPPO , 2008
-Himachal PradeshPresentPaul et al., 1990; Sugha and Kumar, 2003; CABI/EPPO , 2008
-Indian PunjabPresentPanwar et al., 1970; Singh and Chand, 1986; CABI/EPPO , 2008
-Jammu and KashmirPresentCABI/EPPO , 2008
-KarnatakaPresentThippeswamy et al., 2006; CABI/EPPO , 2008
-KeralaPresentCABI/EPPO , 2008
-Madhya PradeshPresentChaudhary and Hasija, 1980; CABI/EPPO , 2008
-MaharashtraPresentUppal et al., 1935; Pawar and Patel, 1957; Mathur, 1979; CABI/EPPO , 2008
-OdishaPresentDas, 1995; Beura et al., 2008; CABI/EPPO , 2008
-Uttar PradeshPresentVerma and Khan, 1965; Vishunavat and Kumar, 1994; CABI/EPPO , 2008On brinjal seed
-UttarakhandPresentCABI/EPPO , 2008
-West BengalPresentManna et al., 2004; CABI/EPPO , 2008
IranPresentCABI/EPPO , 2008
IraqPresentCABI/EPPO , 2008
JapanPresentShishido et al., 2006; CABI/EPPO , 2008
-HonshuPresentShishido et al., 2006In Mie prefecture
JordanPresentEPPO, 2000On phytosanitary list as A2 organism
Korea, Republic ofPresentCho and Shin, 2004; CABI/EPPO , 2008
LaosPresentCABI/EPPO , 2008
MalaysiaPresentCABI/EPPO , 2008
-Peninsular MalaysiaPresentThompson and Johnston, 1953; CABI/EPPO , 2008One record
-SabahPresentCABI/EPPO , 2008
-SarawakPresentTeo, 1984; CABI/EPPO , 2008
MaldivesPresentCABI/EPPO , 2008
MyanmarPresentCABI/EPPO , 2008; Maung Mya Thaung, 2008
PakistanPresentCABI/EPPO , 2008
PhilippinesPresentPalo, 1938; Lapis and Deangkinay, 1967; CABI/EPPO , 2008
Saudi ArabiaPresentAbu-Yaman and Abu, 1972; CABI/EPPO , 2008
TaiwanPresentSawada, 1959; CABI/EPPO , 2008

Africa

AlgeriaPresentCABI/EPPO , 2008
EgyptPresentHaggag, 2014
KenyaPresentCABI/EPPO , 2008
MauritiusPresentIntroduced Invasive Orian, 1951; Felix et al., 1965; CABI/EPPO , 2008
SenegalPresentCABI/EPPO , 2008
SeychellesPresentCABI/EPPO , 2008
South AfricaPresentCrous et al., 2000; CABI/EPPO , 2008
TanzaniaPresentRiley, 1960; CABI/EPPO , 2008
-ZanzibarPresentCABI/EPPO , 2008
ZambiaPresentCABI/EPPO , 2008
ZimbabwePresentCABI/EPPO , 2008

North America

BermudaPresentOgilvie, 1924; CABI/EPPO , 2008
CanadaPresentCABI/EPPO , 2008
-British ColumbiaPresentCABI/EPPO , 2008
-OntarioPresentConners, 1939; CABI/EPPO , 2008
-QuebecPresentCABI/EPPO , 2008
MexicoPresentBPI, US National Fungus Collections; McGuire and Crandall, 1967; CABI/EPPO , 2008Reported on tomato only
USAPresentCABI/EPPO , 2008
-AlabamaPresentBlain, 1931; CABI/EPPO , 2008
-DelawarePresentCABI/EPPO , 2008
-FloridaPresentGratz, 1942; Harrison and Kelbert, 1944; CABI/EPPO , 2008
-HawaiiPresentIntroduced Invasive Raabe et al., 1981; CABI/EPPO , 2008
-IowaPresentGilman Archer, 1929; CABI/EPPO , 2008
-LouisianaPresentEdgerton and Moreland, 1921; CABI/EPPO , 2008
-MississippiPresentParris, 1959; CABI/EPPO , 2008
-New JerseyPresentHalsted, 1892; Harter, 1914; Martin, 1928; CABI/EPPO , 2008
-North CarolinaPresentGrand, 1985; CABI/EPPO , 2008
-OklahomaPresentPreston, 1945; CABI/EPPO , 2008
-TexasPresentBPI, US National Fungus Collections; CABI/EPPO , 20081936. Hidalgo county
-VirginiaPresentBPI, US National Fungus Collections; CABI/EPPO , 2008
-WashingtonPresentCABI/EPPO , 2008
-West VirginiaPresentBPI, US National Fungus Collections; CABI/EPPO , 20081920
-WisconsinPresentHarter, 1914

Central America and Caribbean

Antigua and BarbudaPresentCABI/EPPO , 2008
BarbadosPresentBourne, 1923; CABI/EPPO , 2008
Costa RicaPresentCABI/EPPO , 2008
CubaPresentArnold, 1986; CABI/EPPO , 2008
Dominican RepublicPresentCiferri, 1961; CABI/EPPO , 2008
El SalvadorPresentCABI/EPPO , 2008
GuadeloupePresentJacqua and Gerion, 1988; CABI/EPPO , 2008
GuatemalaPresentMuller, 1950; CABI/EPPO , 2008
HaitiPresentBenjamin and Slot, 1969; CABI/EPPO , 2008
JamaicaPresentSmith, 1937; CABI/EPPO , 2008
PanamaPresentPiepenbring, 2006; CABI/EPPO , 2008
Puerto RicoPresentNollo, 1929; Stevenson, 1975; CABI/EPPO , 2008Considered endemic; limiting to production
United States Virgin IslandsPresentCABI/EPPO , 2008

South America

ArgentinaPresentIntroduced2000 Invasive Dal and Sisterna, 2000; CABI/EPPO , 2008On Prunus armeniaca
BrazilPresentCABI/EPPO , 2008
-CearaPresentCABI/EPPO , 2008
-ParanaPresentCABI/EPPO , 2008
-PernambucoPresentCABI/EPPO , 2008
-Rio de JaneiroPresentCABI/EPPO , 2008
-Sao PauloPresentDefigueiredo and Pereira, 1944; CABI/EPPO , 2008
ColombiaPresentChardon and Toro, 1930; CABI/EPPO , 2008
VenezuelaPresentChardon and Tora, 1934; CABI/EPPO , 2008

Europe

RomaniaPresentIntroduced Invasive Smith et al., 1988; CABI/EPPO , 2008
RomaniaPresentSmith et al., 1988; CABI/EPPO , 2008

Oceania

AustraliaPresentCABI/EPPO , 2008
-QueenslandPresentIntroduced Invasive Simmonds, 1966; Meurant et al., 1999; CABI/EPPO , 2008
FijiIntroduced Invasive CABI/EPPO , 2008
French PolynesiaPresentCABI/EPPO , 2008
GuamPresentIntroduced Invasive Schlub and Yudin, 2002; CABI/EPPO , 2008
New CaledoniaPresentIntroduced Invasive Bugnicourt et al., 1951; CABI/EPPO , 2008

Risk of Introduction

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P. vexans may be introduced readily in the seed, as well as in or on harvested fruit. Phytosanitary regulation of imported seed and fruit, as well as a grower’s selection of clean seed, will readily prevent most introductions. If introduction occurs, destruction of crop debris and crop rotation for several years will reduce or eliminate the fungus from a specific area.

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Present, no further details Harmful (pest or invasive)

Hosts/Species Affected

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P. vexans has been considered to be restricted to Solanum melongena [eggplant/aubergine] (Edgerton and Moreland, 1921; Pawar and Patel, 1957; Sherf and McNab, 1986), but there are reports of pathogenicity to Capsicum annuum (pepper) and Lycopersicon esculentum [Solanum lycopersicum] (tomato) (Sawada, 1959; Tai, 1979) as well as of isolation from Acacia arcuaefolia (Mathur, 1979), Prunus armeniaca (apricot) (Dal Bello and Sisterna, 2000; Cho and Shin, 2004), and seeds of Sorghum bicolor (Mathur, 1979) and interception on imported Capsicumfrutescens (BPI, 2009 [1945]). In India, it has been reported to infect some wild Solanum species in inoculation trials (Datar and Ashtaputre, 1988), and Solanumincanum (Dubey et al., 1987). Edgerton and Moreland (1921), nevertheless, were unable to obtain infection of tomato, pepper, potato [Solanum tuberosum] or wild Solanum species, and Pawar and Patel (1957) report identical results for tomato, pepper and potato, as well as finding no infection of Solanum nigrum. Those reports did not specify the plant parts inoculated, but uninjured tomato and pepper fruits were found to be unaffected by the fungus in parallel trials with brinjal [S. melongena] in India (Chaudhary and Hasija, 1979). Both young and fruiting pepper and tomato plants sprayed with suspensions of conidia were not infected (Harter, 1914).

Growth Stages

Top of page Flowering stage, Fruiting stage, Post-harvest, Pre-emergence, Seedling stage, Vegetative growing stage

Symptoms

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The symptoms range from poor germination and seedling blight to fruit rot. Post-emergence damping-off of seedlings results from infection of the stem just above the soil surface. The symptoms on leaves are more prominent during the early stages of plant growth. At first the lesions are small, more or less circular, and buff to olive, later becoming cinnamon buff, with an irregular blackish margin (Pawar and Patel, 1957). Irregular spots result from coalescence. After transplanting, leaves coming into contact with the soil may become infected directly or develop leaf spot due to infection by conidia. Lesions on the petiole or the lower part of the midrib can result in death of the entire leaf. Affected leaves may drop prematurely, and the blighted areas become covered with numerous black pycnidia.

On stems and branches, elongated, blackish-brown lesions are formed, eventually containing pycnidia. The diseased plant bears smaller leaves and the axillary buds are often killed. When stem girdling occurs, the shoot above the infected area wilts and dries up and the plant may be toppled by the wind (Edgerton and Moreland, 1921; Pawar and Patel, 1957; Sherf and MacNab, 1986). Pycnidia develop readily in lesions on young stems, but rarely on older ones (Harter, 1914).

On the fruits the symptoms appear first as minute sunken greyish spots with a brownish halo, which later enlarge and coalesce, producing concentric rings of yellow and brown zones. These spots increase in size and form large rotten areas on which conidiomata often develop concentrically, covering most of the rotten fruit surface. Pycnidia on fruit are larger than those on stems and leaves (Harter, 1914). If the infection enters the fruits through the calyx, the whole fruit may become mummified due to dry rot (Pawar and Patel, 1957).

Rot may appear in fruit, in transit after harvest (Sherf and MacNab, 1986).

List of Symptoms/Signs

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SignLife StagesType
Fruit / lesions: black or brown
Fruit / mummification
Fruit / premature drop
Leaves / abnormal leaf fall
Leaves / necrotic areas
Leaves / wilting
Leaves / yellowed or dead
Seeds / discolorations
Stems / canker on woody stem
Stems / internal discoloration
Stems / lodging; broken stems
Stems / necrosis
Whole plant / damping off
Whole plant / dwarfing
Whole plant / plant dead; dieback
Whole plant / seedling blight
Whole plant / uprooted or toppled

Biology and Ecology

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Life Cycle

Conidia germinate after 6 hours and penetration occurs after 12 hours. In tissue, the spread of the fungus is both intercellular and intracellular. Seedlings and young stems are highly susceptible. Mature tissue exhibits hypertrophy and hyperplasia below the infected region, preventing further spread of the fungus (Divinagracia, 1968).

Epidemiology

P. vexans requires hot and humid conditions for infection and disease development. Spore germination is optimal at 27°C, and pycnidial formation is greatest between 30 and 35°C (Pawar and Patel, 1957). The optimum relative humidity for disease development is 55% RH and above (Chaudhary and Hasija, 1979), and the optimum temperature for fungal growth is 28°C (Pawar and Patel, 1957). Fruit rot was maximal at 30°C and 50% RH in the growth chamber (Islam and Pan, 1990b); temperatures of 5, 10 and 40°C were unfavourable for disease development in inoculated, detached fruit.

Physiology and Phenology

Isolates from various locations and different parts of the plant varied in some characteristics in culture, but the differences in source could not be related to differences in virulence (Islam and Pan, 1990a). Differences in colony morphology and growth rate, in production of the two forms of conidia, and in virulence on different plant parts were also observed among isolates by Edgerton and Moreland (1921).

Climate

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ClimateStatusDescriptionRemark
A - Tropical/Megathermal climate Preferred Average temp. of coolest month > 18°C, > 1500mm precipitation annually
Af - Tropical rainforest climate Preferred > 60mm precipitation per month
Am - Tropical monsoon climate Preferred Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
As - Tropical savanna climate with dry summer Preferred < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Df - Continental climate, wet all year Tolerated Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)

Means of Movement and Dispersal

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Natural Dispersal

Conidia are disseminated locally by wind and rain (Edgerton and Moreland, 1921). The fungus also survives in crop debris (Ogilvie, 1924; Panwar et al., 1970).

Vector Transmission

Edgerton and Moreland (1921) stated that insects may carry the conidia, but no particular genera or species were reported.

Accidental Introduction

The fungus can be transmitted in and on seed (Porter, 1943; Vishunavat and Kumar, 1993) and on tools (Edgerton and Moreland, 1921). Infected seedlings may be transplanted from the nursery (Nolla, 1929).

Seedborne Aspects

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Incidence

The fungus is seedborne at significant levels (Edgerton and Moreland, 1921; Ogilvie, 1924; Martin, 1934; Toole et al., 1941; Porter, 1943; Singh and Chakrabarti, 1982; Pan and Acharya, 1995), although certain varieties are more likely to be infected (Porter, 1943). Infected seeds contain profuse branched septate mycelium aggregated in the seed coat, between the seed coat and endosperm and in the embryo region of the seeds. Pycnidia are produced in the seed coat, between the seed coat and endosperm, and in the endosperm tissue (Vishunavat and Kumar, 1994).

Effect on Seed Quality

Infection in seed adversely affects the seed quality, causing seed discolouration, reduced seed weight and density, poor germinability and reduced viability (Toole et al., 1941; Porter, 1943; Panwar et al., 1970; Vishunavat and Kumar, 1993).

Pathogen Transmission

The fungus is seed transmissible (Nolla, 1929; Martin, 1930; Vishunavat and Kumar, 1993; Ogilvie, 1994; Pan and Acharya, 1995). Seedborne infection leads to pre-emergence and post-emergence damping-off of seedlings (Kaushal and Sugha, 1995). Infected seedlings bear conidiomata on the first true leaves, which serve as sources of primary inoculum. Conidia are disseminated by rain splash to other plants.

The fungus also survives on infected crop debris, but seedborne inoculum is of great concern when the seeds are exported or imported to areas where the fungus is not already present.

Seed Treatment

Hot-water seed treatment has been recommended to reduce the incidence of infection in seed without adversely affecting seed viability (Martin, 1930; Felix et al., 1965). Seed treatment with formaldehyde is also effective (Edgerton and Moreland, 1921). Chemical seed treatment with captan, carbendazim, carboxin, metasulfovax, thiram and triadimenol was found to increase germination and to reduce the incidence of damping-off of seedlings in artificially infested soil (Kaushal and Sugha, 1995). In the Republic of Georgia, extracts of garlic and celery were found effective as seed treatments for the control of P. vexans (Kuprashvili, 1996). Treatment with captan, carbendazim, carboxin, dithane and mancozeb reduced the incidence of seed-borne fungi, including P. vexans, in local farmer seed lots, but not without reducing seed germination in some cases (Thippeswamy et al., 2006).

Seed Health Tests

Dry seed examination: examining dry seed with a magnifying lens or under a stereobinocular microscope reveals the presence of black pycnidia on the seed surface. However, this test may only give a partial measure of the presence of P. vexans; the absence of conidiomata on the seed surface does not indicate the absence of the fungus on or in seeds. Infected seeds are often discoloured, appearing rusty-brown to black (Vishunavat and Kumar, 1993).

Blotter test: a 9.5 cm diameter Petri dish, made of glass or clear plastic, should be used to allow light to penetrate. Three layers of blotting paper, moistened with sterile water, are placed in the dish. Seeds from working samples are placed at a rate of 25 seeds per plate, equidistantly. Petri dishes are incubated at 25 ± 1°C for 7 days under artificial daylight or NUV light with alternating periods of 12 hours light and 12 hours darkness (Vishunavat and Kumar, 1993). The seeds are examined under a microscope. Infection is measured by the appearance of black conidiomata on the seed surface.

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Flowers/Inflorescences/Cones/Calyx hyphae Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Fruits (inc. pods) fruiting bodies; hyphae; spores Yes Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Leaves fruiting bodies; hyphae; spores Yes Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Seedlings/Micropropagated plants hyphae Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Stems (above ground)/Shoots/Trunks/Branches fruiting bodies; hyphae; spores Yes Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
True seeds (inc. grain) fruiting bodies; hyphae; spores Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Growing medium accompanying plants
Roots
Wood

Impact Summary

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CategoryImpact
Economic/livelihood Negative

Economic Impact

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Fruit rot is the most destructive stage of the disease, as it damages the fruits partially or completely in the fields or during transit. The disease on stems and leaves results in reduction of fruit size and weight as well as loss of plants.

In Louisiana, USA, in 1921, at least 50% yield reduction was observed in eggplant [Solanum melongena] crops due to infection in the field (Edgerton and Moreland, 1921). Later, Martin (1930) in the USA and Nolla (1929) in Puerto Rica also reported losses of 50% or more due to Phomopsis blight in aubergines [Solanum melongena]. In Brazil, in 1944, P. vexans caused such devastating losses that all control measures were impractical (De Figueiredo and Pereira, 1944). In India, the yield losses due to fruit rot ranged from 10 to 20% in the Punjab and Delhi (Panwar et al., 1970). In an advanced stage of disease, seed quality is also adversely affected, and infected seed becomes discoloured, with poor germinability and reduced seed viability (Toole et al., 1941; Porter, 1943; Vishunavat and Kumar, 1993). Seed infection results in pre-emergence and post-emergence damping-off of seedlings; approximately one-third of the plants were lost at each stage (Kaushel and Sugha, 1995).

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • Proved invasive outside its native range
  • Has a broad native range
  • Has high reproductive potential
  • Reproduces asexually
Impact outcomes
  • Host damage
  • Negatively impacts agriculture
  • Negatively impacts livelihoods
Impact mechanisms
  • Pathogenic
Likelihood of entry/control
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field

Diagnosis

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A pure culture can be isolated from pieces of infected tissues on agar plates (Islam and Pan, 1990a). P. vexans produces abundant conidiomata on 4-7% oat meal agar medium at 30°C under light (Divinagracia, 1969). Pawar and Patel (1957) reported good production of pycnidial conidiomata on agar made with an extract of the host.

The blotter method can be used to confirm infection on seeds, as described under 'Seedborne Aspects' (Seed Health Tests).

Sequences of ITS and LSU regions of rDNA for two isolates identified as P. vexans are available in GenBank for comparison (NCBI, 2009).

Detection and Inspection

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Infection is easily visible in the field on close examination of leaves, stems and fruits; characteristic conidiomata appear as black pinhead-sized structures, which are often concentrically arranged on fruits. Infected fruits are soft and mushy or mummified and black. Infection of seed may be confirmed using the methods described for Seed Health Tests in 'Seedborne Aspects'.

Similarities to Other Species/Conditions

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Species of Phomopsis often have similar or overlapping ranges of morphological measurements, and the actual host specificity of each reported species is usually unknown (Uecker, 1988). At least nine other species are reported on Solanum hosts, including those on Lycopersicon esculentum [Solanum lycopersicum] (tomato), now considered to belong in Solanum (USDA-ARS, 2009). Comparative studies of morphology and pathogenicity under identical conditions may be needed to provide a basis for the accurate separation of these Phomopsis species.

Similar post-emergence damping-off of seedlings may be caused by Rhizoctoniasolani, which does not produce pycnidia; its distinctive broad hyphae may be observed with a microscope (Edgerton and Moreland, 1921).

Other fungi cause spots on leaves and fruits of eggplant (Solanum melongena) (Schlub and Yudin, 2002), but the large, dark Phomopsis conidiomata produced in the lesions are distinctive (Chupp and Sherf, 1960). Phoma exigua, which may colonize the lesions as well, produces only small ellipsoid conidia, some of which may be septate (Boerema et al., 2004).

Blossom end rot of eggplant, due to a physiological condition, occurs only on the bottom part of the fruit (Meurant et al., 1999); fruit rot due to Phomopsis is more likely to begin at the top from infection of the calyx (Edgerton and Moreland, 1921).

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.

Prevention

Early Warning Systems

A linear model, based on environmental factors, for predicting Phomopsis blight in aubergines [Solanum melongena] has been developed in India (Islam and Pan, 1992), but is not yet in use. Leaf blight severity was correlated with maximum and minimum temperatures and the number of rainy days.

Control

Cultural Control and Sanitary Measures

Burning of crop debris and burying it by deep ploughing are some of the cultural practices that may help to reduce disease incidence (Singh, 1987). The fungus is also capable of growing well on sterile vegetative structures of a number of other field and garden crops, such as cauliflower petioles, and carrot and beet roots, some of which could then serve to perpetuate the fungus indefinitely (Howard and Desrosiers, 1941). Therefore, the efficacy of crop rotation as a control measure may vary, although a three-year rotation can be useful in reducing initial inoculum (Sherf and MacNab, 1986).

Use of an appropriate nitrogen source at a reduced level with higher rates of phosphorus and potassium fertilizer may increase yield without increasing disease (Sugha and Kumar, 2003).

The pathogen also survives on and in seeds, therefore seeds should be collected from healthy plants and only disease-free seeds should be used.

Chemical Control

Chemical control, especially the use of fungicides, is largely practised for Phomopsis blight control in aubergines [Solanum melongena] crops throughout the world where the disease is prevalent (De Figueiredo and Pereira, 1941; Felix et al., 1965; Teo, 1982, 1984; Singh and Chakrabarti, 1982; Grewal and Jhooty, 1987; Jacqua and Gerion, 1988; Islam and Pan, 1989; 1993; Mohanty et al., 1994; Manna et al., 2004). The more common fungicides applied as foliar sprays are Bordeaux mixture, captan, carbendazim, carboxin, chlorothalonil, copper oxychloride, dithiocarbamates, maneb, mancozeb, thiophanate-methyl, tolclofos-methyl, ferbam and zineb.

Confirming the results of other workers, Beura et al. (2008) found that carbendazim provided the best control of Phomopsis under their test conditions in Orissa state (India); its use also allowed for the maximum increase in yield. In the laboratory, carbendazim completely inhibits culture growth (Mohanty et al., 1994); sensitivity of spore germination to the fungicide is high, though not as high as sensitivity to prochloraz (Sugha and Kumar, 2004). The newer systemic fungicide tebuconazole also provides a high level of control at a low concentration (Manna et al., 2004).

Tests of some natural plant extracts and homeopathic drugs showed that thuja, teucrium and extracts from Allamanda cathartica and Aegle marmelos could prevent or reduce growth of the fungus in vitro as did an effective fungicide, although higher concentrations of active ingredient were required (Panda et al., 1996). Some unidentified compounds extracted from A. cathartica, using organic solvents, prevented the growth of P. vexans in culture at unspecified concentrations (Masuduzzaman et al., 2008).

Seed treatment with mancozeb, carbendazim and thiophanatemethyl has also produced a reduction in disease incidence (Singh and Agarwal, 1999).

Recently, fungicides such as cacrio, quadris, and endura have been registered for use on brinjal, but their efficacy against phomopsis fruit rot is unknown. Fungicides are most effective when combined with cultural control strategies (Howard and David, 2007).

Biological control

Antagonistic Pseudomonas fluorescens and Trichoderma harzianum seed treatment and spray treatment were found to be effective against P. vexans (Srinivas et al., 2005).

Host Resistance

The use of resistant varieties can be one of the most effective methods of control (De Figueiredo and Pereira, 1944). Extensive work in breeding for resistance to Phomopsis blight in aubergines has been carried out with some success in Florida, USA (Decker, 1946; 1947; 1948; 1949), India (Kalda et al., 1976; Datar and Ashtaputre, 1988; Pandey et al., 2002), China (Ren and Zhang, 1993; Liu, 1998) and Brazil (Reifschneider et al., 1993). In India, other Solanum species have been identified as sources of genes for resistance (Sherf and MacNab, 1986; Datar and Ashtaputre, 1988).

Nevertheless, Pandey et al. (2002) found no variety tested to be immune from stem blight or fruit rot. Some were moderately resistant and one escaped severe disease due to early maturity.

Resistance to P. vexans is probably due to chemical and protoplasmic factors rather than structural and mechanical processes (Howard and Desrosiers, 1941).

Gaps in Knowledge/Research Needs

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The frequency of occurrence of the sexual (Diaporthe) form in nature and its possible role in the epidemiology and biology of the pathogen remain undetermined. Additional molecular examination of Phomopsis species on Solanum hosts could clarify their identities and host ranges. Continued breeding for resistance may yield better cultivars for areas where the pathogen is endemic.

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