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Datasheet

Mycosphaerella fijiensis
(black Sigatoka)

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Datasheet

Mycosphaerella fijiensis (black Sigatoka)

Pictures

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PictureTitleCaptionCopyright
Range of symptoms from early streak stage to mature spot with grey centre on banana cultivar Williams (AAA 'Cavendish').
TitleRange of symptoms on leaf
CaptionRange of symptoms from early streak stage to mature spot with grey centre on banana cultivar Williams (AAA 'Cavendish').
CopyrightDavid Jones
Range of symptoms from early streak stage to mature spot with grey centre on banana cultivar Williams (AAA 'Cavendish').
Range of symptoms on leafRange of symptoms from early streak stage to mature spot with grey centre on banana cultivar Williams (AAA 'Cavendish').David Jones
Spots visible on a withered leaf of a 'Cavendish' cultivar.
TitleAdvanced symptoms on leaf
CaptionSpots visible on a withered leaf of a 'Cavendish' cultivar.
CopyrightDavid Jones
Spots visible on a withered leaf of a 'Cavendish' cultivar.
Advanced symptoms on leafSpots visible on a withered leaf of a 'Cavendish' cultivar.David Jones
'Cavendish' cultivar with no viable leaves at harvest.
TitleAdvanced symptoms on plant
Caption'Cavendish' cultivar with no viable leaves at harvest.
CopyrightDavid Jones
'Cavendish' cultivar with no viable leaves at harvest.
Advanced symptoms on plant'Cavendish' cultivar with no viable leaves at harvest.David Jones
M. fijiensis conidia and conidiophores. CMI Descriptions of Pathogenic Fungi and Bacteria No. 413. CAB International, Wallingford, UK.
TitleConidia and conidiophores - line drawing
CaptionM. fijiensis conidia and conidiophores. CMI Descriptions of Pathogenic Fungi and Bacteria No. 413. CAB International, Wallingford, UK.
Copyright©CABI BioScience
M. fijiensis conidia and conidiophores. CMI Descriptions of Pathogenic Fungi and Bacteria No. 413. CAB International, Wallingford, UK.
Conidia and conidiophores - line drawingM. fijiensis conidia and conidiophores. CMI Descriptions of Pathogenic Fungi and Bacteria No. 413. CAB International, Wallingford, UK.©CABI BioScience

Identity

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Preferred Scientific Name

  • Mycosphaerella fijiensis M. Morelet

Preferred Common Name

  • black Sigatoka

Other Scientific Names

  • Cercospora fijiensis M. Morelet
  • Cercospora fijiensis var. difformis J.L. Mulder & R.H. Stover
  • Mycosphaerella fijiensis var. difformis J.L. Mulder & R.H. Stover
  • Mycosphaerella fijiensis var. fijiensis
  • Paracercospora fijiensis (M. Morelet) Deighton
  • Pseudocercospora fijiensis (M. Morelet) Deighton

International Common Names

  • English: black leaf streak
  • Spanish: raya negra de la hoja; sigatoka negra
  • French: cercosporiose noire; maladie des raies noires de la feuille

Local Common Names

  • Germany: Schwarze Sigatoka: Banane

EPPO code

  • MYCOFI (Mycosphaerella fijiensis)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Fungi
  •         Phylum: Ascomycota
  •             Subphylum: Pezizomycotina
  •                 Class: Dothideomycetes
  •                     Subclass: Dothideomycetidae
  •                         Order: Capnodiales
  •                             Family: Mycosphaerellaceae
  •                                 Genus: Mycosphaerella
  •                                     Species: Mycosphaerella fijiensis

Notes on Taxonomy and Nomenclature

Top of page Leach (1964) found that black leaf streak of bananas was caused by a species of Mycosphaerella which had a Cercospora-like imperfect state. The fungus was examined by Deighton who considered it was a new species. The name M. fijiensis was suggested and was later validated by Morelet (1969).

Meredith and Lawrence (1969) made a detailed study of black leaf streak disease in Hawaii, comparing conidial states of M. fijiensis and M. musicola.

Stover (1974) assigned the name M. fijiensis var. difformis to the fungus causing black Sigatoka disease in Honduras. This name was validated by Mulder and Stover (1976) on the basis of specimens collected at La Lima, Honduras. The taxonomic criterion used to separate M. fijiensis var. difformis from M. fijiensis var. fijiensis was the presence of a sporadic stroma as the origin for dense or loose fascicles of conidiophores in the former in contrast to the absence of a stroma in the latter. However, the presence of the stroma in M. fijiensis var. difformis is an inconsistent feature. Pons (1987) examined specimens of M. fijiensis var. difformis and var. fijiensis and found that conidiophores could develop on a stroma in both. The two names are now considered synonyms (Pons, 1999).

The anamorph of M. fijiensis was first described as Cercospora fijiensis by Morelet (1969) and then transferred to Pseudocercospora by Deighton (1976) because of pigmented conidia. Deighton (1979) later placed the anamorph in Paracercospora due to minutely thickened spore scars.

Pseudocercospora fijiensis is again recommended as the anamorph because molecular data has proven that thickened spore scars, though useful in distinguishing the pathogen, are not phylogenetically important in the cercosporoids (Crous and Mourichon, 2002).

Description

Top of page (After Meredith and Lawrence, 1969; Mulder and Holliday, 1974.)

Conidiophores

Conidiophores first develop in the initial flecks or streaks on the lower surface of the leaf and continue to be produced until spots mature. They emerge singly or in diverging fascicles of 2-8 from stroma on the lower surface of the leaf within the boundary of the lesion; few arise on the upper surface. Conidiophores are pale to medium olivaceous-brown, becoming slightly paler towards the tip. They are straight or bent, often with geniculations and sometimes with a basal swelling up to 8 µm in diameter, 0- to 5-septate, 16.5-62.5 x 4-7 µm, usually slightly narrower, but occasionally wider, at the tip. One or more scars are present near the tip of the conidiophore, either flat against the apex or on the side, or on a slightly sloping shoulder.

Conidia

Conidia are formed singly at the apex of the conidiophore, later becoming lateral as the conidiophore develops. Up to four mature conidia may be attached to a single conidiophore. Conidia are not quite colourless, being pale-green or olivaceous. They are obclavate to cylindro-obclavate, 1- to 10-septate (commonly 5- to 7-septate), straight or curved, obtuse at the apex, truncate or rounded at the base with a visible and slightly thickened hilum, 30-132 x 2.5-5 µm, the broadest point being near the base.

Spermogonia

Spermogonia develop at the stage when streaks develop into spots and are more abundant on the lower surface of the leaf, being consistently associated with conidiophores. Spermogonia are hour-glass shaped, oval or almost globose and measure 55-88 x 35-50 µm. The ostiole is slightly prominent and protrudes through the stoma pore. Many hyaline, rod-shaped spermatia, 2.5-5.0 x 1.0-2.5 µm, are found in mature spermogonia.

Ascomata and Asci

Ascomata are perithecial, globose, 47-85 µm in diameter. They are immersed in the leaf tissue with protruding ostioles and are found on both leaf surfaces, although more abundant on the upper. Asci are numerous, obclavate, fisstunicate and 8-spored; paraphyses are lacking. Ascospores are unequally 1-septate and slightly constricted at the septum, the longer cell being uppermost in the ascus. They are hyaline, biseriate, fusiform, 11.5-16.5 x 2.5-5.0 µm.

Colony Morphology

Colonies on potato dextrose agar are slow growing, compact but with a velvety surface, prominently raised, grey to pale-buff or olive-green, black in reverse (Mulder and Holliday, 1974). On Mycophyl agar, colonies are dark-grey or grey-brown with a crenate edge or pale-grey to pink (Stover, 1976). Conidia can be produced in culture for use in inoculation experiments. Mourichon et al. (1987) used colonies growing on modified V8 juice agar at 25°C under continuous light as a source of conidia.

Distribution

Top of page Black leaf streak was first recognized in Fiji in 1963, but was present over a wide area of the Asia/Pacific region before its discovery. Stover (1978) believed the centre of origin of M. fijiensis may have been the Papua New Guinea/Solomon Islands area. Certainly, isolates of the pathogen have been found to be more genetically diverse in Papua New Guinea and also the Philippines than elsewhere(Carlier et al., 2000a), which supports Stover's argument. In the 'Distribution list' above, Irian Jaya/Papua New Guinea/Solomon Islands have been designated the locations where black leaf streak may have originated.

In Australia, black leaf streak is confined to the Torres Strait region of Queensland. Quarantine restrictions are in force to prevent its spread south in Cape York Peninsula and to other areas of Australia (Jones, 1990).

Black leaf streak was reported as present in Java (Indonesia) by Reddy (1969). However, a survey in 1988 showed the dominant leaf spot in Java to be Sigatoka (DR Jones, Brisbane, 1988, personal communication). In 1996, black leaf streak was again identified as present in Java, but Sigatoka was still the dominant leaf spot disease (X Mourichon, Montpellier, 1996, personal communication). Black leaf streak almost certainly exists in Irian Jaya (Indonesia), but has not been officially recorded.

Black leaf streak was first reported in peninsular Malaysia in 1965, but, if the record is correct, failed to become the dominant leaf spot as in many other tropical countries. More recently, the causal agent has been identified in Johore and Langkawi Island (X Mourichon, CIRAD, Montpellier, France, personal communication, 1996), but elsewhere eumusae leaf spot caused by the fungus Mycosphaerella eumusae is prevalent.

Although black leaf streak has been recorded in Thailand (Reddy, 1969), all leaf spot samples collected during a 1994 banana disease survey of the country were identified as eumusae leaf spot (DR Jones, INIBAP, personal communication, 1995).

Black leaf streak was reported in Bhutan in 1989, but the disease has not yet been recorded in India or Bangladesh. One would have expected black leaf streak to have spread from Bhutan to the Indian subcontinent before now if it was invasive in this region. Therefore, a similar situation may exist as in Thailand and peninsular Malaysia. Certainly, eumusae leaf spot has been identified has the predominant leaf spot in southern India and Sri lanka.

The first record of black leaf streak in Africa, which was in Zambia in 1973, is in doubt. Specimens sent to IMI for identification could not be confirmed as M. fijiensis (Tushemereirwe and Waller, 1993). A report of black leaf streak in Guinea (Jones and Mourichon, 1993) was erroneous and has been omitted. Recent reports from Guinea Bissau and Niger (EPPO, 2003) need confirmation.

Reports of black leaf streak in the Netherlands Antilles (EPPO, 2003) in the Caribbean can not be verified. Another report from Guyana has proved false and has been omitted (EPPO, 2009).

The disease may be present in the Northern Mariana Islands although the reports are unsubstantiated (EPPO, 2003).

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

BhutanPresentIntroduced1985 Not invasive Peregrine and, 1989; CABI/EPPO, 2011; EPPO, 2014
Brunei DarussalamPresentCABI/EPPO, 2011
ChinaRestricted distributionCABI/EPPO, 2011; EPPO, 2014
-GuangdongPresentIntroduced1990 Invasive Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
-HainanPresentIntroduced1980 Invasive Stover and Simmonds, 1987; CABI/EPPO, 2011; EPPO, 2014
-YunnanPresentIntroduced1993 Invasive Jones and Mourichon, 1993; CABI/EPPO, 2011; EPPO, 2014
IndonesiaRestricted distributionCABI/EPPO, 2011; EPPO, 2014
-Irian JayaPresentNative Invasive Davis et al., 2000; CABI/EPPO, 2011; EPPO, 2014
-JavaPresentIntroduced1969 Not invasive Reddy and, 1969; CABI/EPPO, 2011; EPPO, 2014
-KalimantanPresentIntroduced1996 Invasive Carlier et al., 2000a; CABI/EPPO, 2011; EPPO, 2014
-MoluccasPresentIntroduced Invasive Stover, 1978; CABI/EPPO, 2011; EPPO, 2014
-SumatraPresentIntroduced1993 Invasive Jones and Mourichon, 1993; CABI/EPPO, 2011; EPPO, 2014
MalaysiaRestricted distributionCABI/EPPO, 2011; EPPO, 2014
-Peninsular MalaysiaPresentIntroduced1965 Not invasive Graham, 1969; CABI/EPPO, 2011; EPPO, 2014
-SarawakPresentIntroduced1996 Invasive Carlier et al., 2003a; CABI/EPPO, 2011; EPPO, 2014
PhilippinesPresentIntroduced1964 Invasive Hapitan and Reyes, 1970; CABI/EPPO, 2011; EPPO, 2014
SingaporePresentIntroduced1964-1967 Invasive Graham, 1969; CABI/EPPO, 2011; EPPO, 2014
TaiwanPresentIntroduced1927 Invasive Stover, 1978; CABI/EPPO, 2011; EPPO, 2014
ThailandPresentIntroduced1969 Not invasive Reddy and, 1969; CABI/EPPO, 2011; EPPO, 2014
VietnamPresentIntroduced1993 Invasive Jones and Mourichon, 1993; CABI/EPPO, 2011; EPPO, 2014

Africa

BeninPresentIntroduced1993 Invasive Jones and Mourichon, 1993; CABI/EPPO, 2011; EPPO, 2014
BurundiPresentIntroduced1987 Invasive Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
CameroonPresentIntroduced1980 Invasive Tezenas, 1982; CABI/EPPO, 2011; EPPO, 2014
Central African RepublicPresentCarlier et al., 2003a; CABI/EPPO, 2011; EPPO, 2014
ComorosWidespreadIntroduced Invasive Jones and Mourichon, 1993; CABI/EPPO, 2011; EPPO, 2014
CongoPresentIntroduced1985 Invasive Mourichon, 1986; CABI/EPPO, 2011; EPPO, 2014
Congo Democratic RepublicPresentIntroduced Invasive Mobambo & Nakau, 1993; Sebasigari and Stover, 1988; Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
Côte d'IvoirePresentIntroduced1985 Invasive Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
GabonPresentIntroduced1978 Invasive Frossard, 1980; CABI/EPPO, 2011; EPPO, 2014
GhanaPresentIntroduced1986 Invasive Wilson, 1987; CABI/EPPO, 2011; EPPO, 2014
KenyaPresentIntroduced1988 Invasive Kung'u et al., 1992; IPPC-Secretariat, 2005; CABI/EPPO, 2011; EPPO, 2014
MadagascarPresentIntroduced2000 Invasive Jones, 2003; CABI/EPPO, 2011; EPPO, 2014
MalawiPresentIntroduced1990 Invasive Ploetz et al., 1992; CABI/EPPO, 2011; EPPO, 2014
MauritiusPresentCABI/EPPO, 2011; EPPO, 2014
MayottePresentIntroduced Invasive Carlier et al., 2003a; CABI/EPPO, 2011; EPPO, 2014
NigeriaPresentCABI/EPPO, 2011; EPPO, 2014
RwandaPresentIntroduced1986 Invasive Mourichon and Fullerton, 1990; Sebasigari, 1990; CABI/EPPO, 2011; EPPO, 2014
Sao Tome and PrincipePresentIntroduced1983 Invasive Frossard, 1980; CABI/EPPO, 2011; EPPO, 2014
TanzaniaPresentIntroduced1987 Invasive Sebasigari and Stover, 1988; CABI/EPPO, 2011; EPPO, 2014
-ZanzibarPresentIntroduced Invasive Sebasigari and Stover, 1988
TogoPresentIntroduced1988 Invasive Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
UgandaPresentIntroduced1990 Invasive Tushemereirwe and Waller, 1993; CABI/EPPO, 2011; EPPO, 2014
ZambiaAbsent, reported but not confirmedRaemakers, 1975; CABI/EPPO, 2011; EPPO, 2014

North America

MexicoRestricted distributionIntroduced1980 Invasive Stover and Simmonds, 1987; Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
USARestricted distributionCABI/EPPO, 2011; EPPO, 2014
-FloridaPresentIntroduced1998 Invasive Ploetz and Mourichon, 1999; CABI/EPPO, 2011; EPPO, 2014
-HawaiiPresentIntroduced1958 Invasive Meredith and Lawrence, 1969; CABI/EPPO, 2011; EPPO, 2014

Central America and Caribbean

BahamasRestricted distributionCABI/EPPO, 2011; EPPO, 2014
BelizeWidespreadIntroduced1975 Invasive Stover, 1980a; Stover, 1980b; Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
Costa RicaRestricted distributionIntroduced1979 Invasive Stover, 1980a; Stover, 1980b; Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
CubaPresentIntroduced1991 Invasive Vidal, 1992; Jones and Mourichon, 1993; CABI/EPPO, 2011; EPPO, 2014
Dominican RepublicPresentIntroduced1996 Invasive CABI/EPPO, 2011; EPPO, 2014
El SalvadorPresentIntroduced1979 Invasive Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
French West IndiesPresentBellaire et al., 2009
GrenadaRestricted distributionIPPC, 2007; IPPC, 2007; CABI/EPPO, 2011; EPPO, 2014
GuadeloupeAbsent, invalid recordCABI/EPPO, 2011; EPPO, 2014
GuatemalaRestricted distributionIntroduced1977 Invasive Stover, 1980a; Stover, 1980b; Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
HaitiPresentIntroduced1999 Invasive Jones, 2003; CABI/EPPO, 2011; EPPO, 2014
HondurasPresentIntroduced1972 Invasive Stover & Dickson, 1976; Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
JamaicaPresentIntroduced1995 Invasive Wilson, 1996; CABI/EPPO, 2011; EPPO, 2014
MartiniquePresentCABI/EPPO, 2011; Ioos et al., 2011; EPPO, 2014
NicaraguaRestricted distributionIntroduced1979 Invasive Stover, 1980a; Stover, 1980b; Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
PanamaPresentIntroduced1981 Invasive Stover, 1987; Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
Puerto RicoPresentIrish et al., 2006; CABI/EPPO, 2011; EPPO, 2014
Saint LuciaPresentIntroduced Invasive Compton, 2010; Lebourne, 2010; CABI/EPPO, 2011; EPPO, 2014
Saint Vincent and the GrenadinesWidespreadIPPC, 2009; CABI/EPPO, 2011; EPPO, 2014
Trinidad and TobagoWidespreadIPPC, 2012a; IPPC, 2012b; CABI/EPPO, 2011; EPPO, 2014

South America

BoliviaPresentIntroduced1996 Invasive Tejerina, 1997; CABI/EPPO, 2011; EPPO, 2014
BrazilPresentIntroduced1998 Invasive Cordeiro et al., 1998; CABI/EPPO, 2011; EPPO, 2014
-AcrePresentCavalcante et al., 2004; CABI/EPPO, 2011; EPPO, 2014
-AmapaPresentEPPO, 2014
-AmazonasPresentIntroduced Invasive Cordeiro et al., 1998; CABI/EPPO, 2011; EPPO, 2014
-BahiaAbsent, unreliable recordSenhor et al., 2009; EPPO, 2014
-Espirito SantoAbsent, unreliable recordEPPO, 2014
-Mato GrossoPresentSouza and Feguri, 2004; CABI/EPPO, 2011; EPPO, 2014
-Mato Grosso do SulPresentEPPO, 2014
-Minas GeraisAbsent, unreliable recordCastro et al., 2005; CABI/EPPO, 2011; EPPO, 2014
-ParaPresentIntroduced Invasive Trinidade et al., 2002; Benchimol et al., 2010; CABI/EPPO, 2011; EPPO, 2014
-ParanaPresentEPPO, 2014
-Rio Grande do SulPresentFavreto et al., 2007; CABI/EPPO, 2011; EPPO, 2014
-RondoniaPresentEPPO, 2014
-RoraimaPresentEPPO, 2014
-Santa CatarinaPresentEPPO, 2014
-Sao PauloRestricted distributionIntroduced Invasive Nogueira et al., 2000; Ferrari et al., 2005; CABI/EPPO, 2011; EPPO, 2014
-TocantinsPresentEPPO, 2014
ColombiaPresentIntroduced1991 Invasive Merchan, 1990; Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
EcuadorPresentIntroduced1986 Invasive Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
GuyanaPresentCABI/EPPO, 2011; EPPO, 2014
PeruPresentIntroduced1996 Invasive Jones, 2003; CABI/EPPO, 2011; EPPO, 2014
SurinameAbsent, unreliable recordCABI/EPPO, 2011; EPPO, 2014
VenezuelaPresentIntroduced1993 Invasive Jones and Mourichon, 1993; CABI/EPPO, 2011; EPPO, 2014

Oceania

American SamoaPresentIntroduced1975 Invasive Firman, 1975; CABI/EPPO, 2011; EPPO, 2014
AustraliaEradicatedIntroduced1981/2001CABI/EPPO, 2011; EPPO, 2014
-QueenslandEradicatedIntroduced Invasive Jones and Alcorn, 1982; CABI/EPPO, 2011; EPPO, 2014
Cook IslandsPresentIntroduced1976 Invasive Firman, 1975; CABI/EPPO, 2011; EPPO, 2014
FijiPresentIntroduced1963 Invasive Rhodes, 1964; CABI/EPPO, 2011; EPPO, 2014
French PolynesiaPresentIntroduced1964-1967 Invasive Graham, 1969; CABI/EPPO, 2011; EPPO, 2014
GuamPresentCABI/EPPO, 2011
Marshall IslandsPresentCABI/EPPO, 2011
Micronesia, Federated states ofPresentIntroduced1964-1967 Invasive Graham, 1969; Jones and Mourichon, 1993; CABI/EPPO, 2011; EPPO, 2014
New CaledoniaRestricted distributionIntroduced1964-1967 Invasive Graham, 1968; Mourichon and Fullerton, 1990; CABI/EPPO, 2011; EPPO, 2014
NiuePresentIntroduced1976 Invasive Stover, 1976; Dingley et al., 1981; CABI/EPPO, 2011; EPPO, 2014
Norfolk IslandPresentIntroduced1980 Invasive Jones, 1990; CABI/EPPO, 2011; EPPO, 2014
Northern Mariana IslandsPresentCABI/EPPO, 2011
PalauPresentCABI/EPPO, 2011
Papua New GuineaPresentNative Invasive Stover, 1976; CABI/EPPO, 2011; EPPO, 2014
SamoaPresentIntroduced1965 Invasive CABI/EPPO, 2011; EPPO, 2014
Solomon IslandsPresentNativeStover, 1976; CABI/EPPO, 2011; EPPO, 2014
TongaPresentIntroduced1965Johnston, 1965; CABI/EPPO, 2011; EPPO, 2014
VanuatuPresentIntroduced1964-1967 Invasive Graham, 1969; CABI/EPPO, 2011; EPPO, 2014
Wallis and Futuna IslandsPresentIntroduced1996 Invasive Carlier et al., 2003a; CABI/EPPO, 2011; EPPO, 2014

History of Introduction and Spread

Top of page First records

The first report of M. fijiensis causing damage was in the same Sigatoka valley on the island of Vitu Levu in Fiji where M. musicola was first recognised as a major pathogen of banana fifty years earlier. In February 1963, the disease caused by M. fijiensis was said to be spreading rapidly in the Sigatoka Valley (Rhodes 1964) and it was predicted that it would be island-wide by the end of 1964 (Leach 1964a). The causal agent was also described for the first time from material collected in Fiji (Leach 1964b). The disease caused by M. fijiensis was called 'black leaf streak' by Rhodes (1964). Leach (1964b) described the risk of spread of this new disease of banana as 'a grave threat' and that the abundance of airborne ascospores produced by the pathogen may lead to it being disseminated around the world faster than Sigatoka leaf spot (Jones, 2003).

The problem caused by M. fijiensis in Fiji became apparent when the mist-sprays of light mineral oil being used successfully to control M. musicola could no longer keep leaf spot in check. The recognition of yet another important banana pathogen in Fiji before anywhere else can probably be attributed to the fact that at this location there were sizeable plantations of susceptible dessert banana cultivars and an efficient plant protection service experienced in banana problems (Jones, 2003).

Surveys undertaken after black leaf streak was discovered in Fiji led to the conclusion that the pathogen had most likely been present in the Pacific and parts of the Pacific rim for many years previously (Meredith 1970, Stover 1976, Stover 1978, Long 1979). It was suggested that M. fijiensis may have been in the Hawaiian Islands in 1958 (Meredith and Lawrence 1969). An analysis of herbarium specimens by Stover (1976), showed that M. fijiensis was present in Papua New Guinea by at least 1957 and in Taiwan as early as 1927. The similarity of symptoms with those of Sigatoka most likely masked the arrival of this new disease in many countries. Because of this, the year that black leaf streak was first discovered in many countries in this region does not reflect the order of spread of the pathogen (Jones, 2003).

Global spread

When the black leaf streak pathogen was first found in Honduras in 1972, it was thought from its morphology to be a variant of M. fijiensis and was named M. fijiensis var. difformis (Mulder and Stover 1976). The disease the fungus caused was called black Sigatoka. However, it was later shown that M. fijiensis and M. fijiensis var. difformis were synonymous (Pons 1987).
A measure of the rate of spread of M. fijiensis between countries can be gained from an examination of the records from the Latin American/Caribbean region . Here, year of first report roughly corresponds with the year of introduction. Within three years of its detection in Honduras in 1972, M. fijiensis was reported in Belize to the north and by 1977 had arrived in Guatemala to the west. Local spread was quicker in the direction of prevailing winds from the east and northeast (Stover 1980). In 1979, it appeared in El Salvador, Nicaragua and Costa Rica and by 1981 had spread north to Mexico and south to Panama and northern Colombia (Carlier et al. 2000a). Spread was believed to have been accelerated in Central America by the movement of diseased banana leaves and leaf trash across international boundaries with road-transported banana and plantain fruit (Stover 1980). By 1986, commercial plantations in northern Ecuador were affected and plantains in western Venezuela succumbed in 1991. Spread to northern Peru occurred in 1994 and to Bolivia in 1996. The first report from western Brazil came in 1996 and since then M. fijiensis has been advancing in a southwesterly direction towards the Brazilian coast. In 2001, movements of banana fruit and associated banana leaves from inland areas where M. fijiensis was found to coastal cities were being controlled in an effort to delay spread (R. S. Moreira, Brazil, 2001, personal communication) (Jones, 2003).

In the Caribbean, black leaf streak was first found in Cuba in 1990. Jamaica followed in 1995 and then the Dominican Republic in 1996. The first authenticated report from Haiti, which has a dry climate, was made in 1999. Natural spread to other Caribbean countries is inevitable, but may be slowed considerably by the prevailing winds, which blow from the east. Recent investigations involving the analysis of isolates have revealed that the source of inoculum for the outbreak in Jamaica may have come from Central America and not windblown from Cuba as initially suspected (G. Rivas, Costa Rica, 2001, personal communication). The outbreak in the Dominican Republic has also been linked to Central America, though the evidence is more circumstantial. In both cases, the disease appeared shortly after banana fruit was shipped to the islands (Jones, 2003).

The first record of black leaf streak in Africa was from Zambia in 1973 (Raemaekers 1975). Although the paper published on this outbreak is convincing, the identity of the pathogen could not be confirmed from specimens sent to the UK for identification and therefore doubt remains as to the authenticity of the report (Dabek and Waller 1990). The next record was from Gabon in 1978. Frossard (1980) believed it may have been introduced on planting material from Asia. The disease then spread steadily through Central and West Africa reaching Côte d'Ivoire, Nigeria and Ghana in 1985-1986, and Uganda and Malawi in 1990 (Table 2). A second, separate introduction of M. fijiensis into Africa is thought to have occurred in 1987 on the island of Pemba. This outbreak is believed to have led to the pathogen spreading to the island of Zanzibar and coastal areas of Tanzania and Kenya (Carlier et al. 2000a). In 2000, M.fijiensis was recorded in Madagascar for the first time (Jones, 2003).

The Australian experience- a case study in local disease spread

Stover (1978) believed that M. fijiensis may have originated in the Papua New Guinea-Solomon Islands area and disseminated around the South Pacific with banana leaves or planting material. This possibility has received credence by the discovery that isolates of M. fijiensis are more diverse in the Papua New Guinea /Philippines region than elsewhere, an indication that this area may be the centre of origin of the pathogen (Carlier et al. 2000a). Therefore, it is likely that M. fijiensis may have been present on banana plants on islands in the Torres Strait and on the tip of Cape York Peninsula in Australia long before its discovery on the first plant pathological survey of the area in 1981 (Jones and Alcorn 1982). Airborne inoculum of the pathogen may not have spread the disease any further south in Australia because of the barrier presented by the Cape York Peninsula, which is a large, remote area of native bush with comparatively few communities and banana plants (Jones, 2003).

The discovery of the disease led to an attempt to eradicate the pathogen from the Bamaga area at the tip of Cape York and some Torres Strait islands by the destruction of all banana plants at these locations. This attempt failed for unknown reasons, the disease reappearing on reintroduced banana plants after a significant host-free period (Jones 1984). In hindsight, it seems likely that the close proximity of this area to Papua New Guinea, where the pathogen was endemic, meant that reintroduction by airborne inoculum was inevitable.

In the 1980s, better land and air communications in the far north of Queensland encouraged more tourists and people seeking an alternative lifestyle to visit the remote area of Australia where black leaf streak was present. This led to a higher risk of spread through movement of affected propagating material and possibly leaves with viable lesions. A programme of replacing susceptible cultivars with those that were known to have some resistance to M. fijiensis, such as 'Mysore' and 'Pisang Awak', was begun on some islands in the Torres Strait and on Cape York Peninsula in order to reduce disease incidence and local inoculum levels.

During the 1990s, isolated outbreaks of M. fijiensis were detected and eradicated from small banana plantings within Cape York Peninsula. These were at Pascoe River (1991), Bloomfield (1993), Weipa (1996) and Daintree (1997). In all cases, the origin of the inoculum could not be positively determined. The latter outbreak was the farthest south and the greatest cause of concern because of its close proximity to Cairns and the main commercial banana growing areas.

During banana disease surveys in 1998-1999, the presence of the pathogen was reported at Bamaga and Pascoe river (Wattle Hills) on Cape York Peninsula and on islands in the Torres Strait (Davis et al., 2000). In 2000, an outbreak occurred on a commercial banana planting for the first time. This was at Daintree, where an earlier outbreak on a much smaller planting in 1997 had been eradicated. The grower concerned was compensated for the destruction of his crop by the Australian banana industry. The pathogen was again eradicated (Jones, 2003).

Towards the end of the wet season in April 2001, M. fijiensis was detected on unmanaged (feral) and adjacent cultivated banana plants in the Tully Valley, which is in the heart of the commercial banana-growing area in North Queensland centred south of Cairns. Subsequently, the pathogen was reported from other locations in the same area. An eradication campaign was immediately mounted. This campaign gathered momentum when the governments of banana-growing states and the Commonwealth Government pledged funds. Measures employed included: - (1) the establishment of a special banana quarantine area, (2) a ban on the movement of fruit from this area to other banana-growing areas in Australia, (3) the close monitoring of crops and the diagnosis of any leaf spots detected, (4) the destruction of fields where affected plants were found (5) the drastic pruning of all banana plants in the growing area, (5) the regular application of systemic fungicides and (6) zero tolerance for leaf spot. This campaign was conducted during the 2001 dry winter season, which also significantly reduced chances of spore release and infection, with the co-operation of most growers. A total of 25 plants were found infected with M. fijiensis in the Tully area in 2001. The last seven plants found were either growing in private gardens or were unmanaged (Jones, 2003).

At the time of writing (January 2004), M. fijiensis has not been detected on banana in the Tully Valley for over two years. It seems likely that M. fijiensis has been successful eradicated from a growing area, which is the first time that this has been accomplished anywhere in the world. This was only achieved because (1) the pathogen was recognised very soon after introduction because of routine leaf spot monitoring operations, (2) the banana growers' association immediately lobbied the government for action to be taken to eradicate the pathogen, (3) the government provided funds necessary to eradicate the pathogen and (4) the local growers worked in unison for the benefit of the entire industry and undertook the control measures necessary for the eradication campaign to be successful.

The future for the banana industry in North Queensland as regards black leaf streak is uncertain especially as the sources of inoculum for the numerous outbreaks remains undetermined. It seems unlikely that airborne inoculum originating in Papua New Guinea/Torres Strait could be responsible for outbreaks as far south as Daintree and the Tully Valley, which is a distance of over 700 km. It is also unlikely that inoculum originates from undiscovered small pockets of disease further south on Cape York Peninsula, though this theory cannot be entirely discarded. It seems probable that inoculum is being introduced by another pathway, perhaps on illegally introduced propagating material or by deliberate mischievousness.

Risk of Introduction

Top of page Black leaf streak disease has not yet been reported in the islands in the eastern Caribbean where smallholders produce fruit for export. It has also not been recorded in India, Sri Lanka, South Africa and commercial production areas in Australia where bananas are mostly grown for local consumption. Quarantine measures are in place in Australia to prevent the spread of black leaf streak from the Torres Strait region to other areas (Jones, 1990). The disease can be disseminated long distances by the movement of conventional planting material, such as infected sword suckers or as diseased leaf tissue. Banana plantlets in tissue culture pose no risk of contamination by fungal diseases if maintained in a sterile environment.

Hosts/Species Affected

Top of page Advanced symptoms of black leaf streak caused by M. fijiensis have only been recorded on cultivated banana and plantain and the wild banana species Musa acuminata (subsp. banksii and subsp. zebrina). Banana cultivars differ in their reaction to the pathogen. Many are susceptible, but some show varying degrees of resistance from slow lesion development to a hypersensitive-like response to infection (see Biology and Ecology). Most wild species that have been tested are infected, but invasion is usually halted at a very early stage in a hypersensitive-like response (Carlier et al., 2000a).

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Heliconia psittacorumHeliconiaceaeOther
Musa (banana)MusaceaeMain

Growth Stages

Top of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage

Symptoms

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Symptoms are first visible as faint, minute, reddish-brown specks on the lower surface of the leaf. Specks elongate, becoming slightly wider, to form a characteristic narrow, reddish-brown streak with dimensions of 20 x 2 mm with the long axis parallel to leaf veins. Streaks frequently overlap to form compound streaks. The colour of streaks, which are now clearly visible on the upper leaf surface, changes to dark brown, almost black. The entire leaf can blacken at this stage if streaks are numerous. If less densely congregated, streaks broaden and become fusiform or elliptical spots. Water-soaked borders appear around spots and surrounding leaf tissue yellows slightly. The centres of spots become slightly depressed and dry out, becoming light grey or buff. Each spot has a well-defined, narrow dark brown or black border and surrounding tissue is often yellow. Whole sections of leaves can become necrotic as spots coalesce. After the leaf has withered, spots remain visible because of their light-coloured centres. Different stages of disease development have been identified (Meredith and Lawrence 1969; Fouré, 1987). Often, all stages of disease development can be seen on one leaf.

If inoculum pressure is high, leaves are rapidly destroyed. Often, fewer than six living leaves may be seen on a susceptible plant that is growing vegetatively. On resistant cultivars, symptoms are only usually seen on the older, lower leaves. The disease is more severe on plants with bunches because new leaves are no longer being produced to replace those lost due to disease. If disease pressure is great, it is not uncommon for a susceptible cultivar to have no viable leaves at harvest.

List of Symptoms/Signs

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SignLife StagesType
Leaves / abnormal colours
Leaves / abnormal leaf fall
Leaves / necrotic areas

Biology and Ecology

Top of page Disease development.

The period between infection and the formation of mature lesions depends on the resistance or susceptibility of the cultivar, intensity of infection and environmental conditions. Infection is believed to occur as a new leaf emerges from the pseudostem and unfurls. If the cultivar is susceptible, initial specks may appear on the second and third open leaves of a growing plant, streaks on the third and fourth leaves and both spots and streaks on older leaves. If a cultivar has resistance, streaks and spots may only be seen on the very oldest leaves. In some highly resistant cultivars, specks develop quite rapidly in response to infection, but there is no further disease development. Some authors believe that in these cases, the speck may represent a hypersensitive-like reaction (Carlier et al., 2000a).

Reproductive biology.

Conidia are formed first in lesions and spread the infection to other leaves on the same plant or to adjacent plants. They are dislodged from conidiophores by wind and water (Stover, 1980b). Germination occurs in water and the leaf is penetrated through stomata. Ascospores are thought to contribute to most of the inoculum and can spread the disease further distances than conidia. They are forcibly discharged when the leaf surface is wet and can be carried many kilometres in air currents (Stover, 1980b). However, recent studies suggest that ascospores are susceptible to UV radiation which may prevent spread over very long distances (Parnell et al., 1998). Ascospores also germinate in moisture and infect leaves through stomata. Both conidia and ascospores can germinate within 2-3 hours, but stomata are not usually penetrated until after 48-72 hours of humidity at or near saturation, and at temperatures above 20°C. After infection, hyphae emerge from the stomata and either develop into conidiophores or grow across the surface and infect adjacent stomata. Streaks usually appear first near the leaf apex and along the leaf margin, which is indicative of infection by ascospores (Meredith, 1970). Spotting can develop on the third or fourth fully opened leaf of a susceptible, vegetatively growing plant and sometimes on the second (Stover, 1980b).

Genetics

Populations of M. fijiensis maintain a high level of genetic diversity and it is speculated that pathogenic variability is, therefore, also likely to exist (Carlier et al., 2000a). Isolates of the pathogen from different locations in Papua New Guinea and elsewhere have in fact been found to vary in their pathogenicity in glasshouse screening tests using differential Musa genotypes (Fullerton and Olsen, 1995). Isolates have also been shown to vary in aggressiveness (Jacome and Schuh, 1993; Romero and Sutton, 1997).


Environmental requirements

Maximum germination of conidia and ascopsores occurs in water and decreases as the relative humidity (RH) lowers. No conidia have been observed germinating below 95% RH and no ascospores below 98% RH (Jacome et al., 1991). The minimum, optimum and maximum temperatures for the development of ascospore germ tubes of M. fijiensis is 12°C, 27°C and 36°C respectively, with no development taking place at 11°C and 38°C (Stover, 1983; Jacome et al., 1991; Porras and Pérez, 1997). However, ascospore germ-tube growth at 20°C is half the rate it is at 27°C. A film of water on the leaf surface is required for ascospore infection under controlled conditions. Water is not required for conidial infection provided the relative humidity is high. Epiphytic growth of hyphae from one stoma to another is most likely encouraged by leaf surface moisture as disease severity increases the longer the period of leaf wetness (Jacome and Schuh, 1992). Maximum disease development probably takes place at around 25-27°C under wet conditions.

Conidiophores develop under conditions of high humidity. They were seen in streak symptoms in Cameroon in the rainy season, but not in the dry (Fouré and Moreau, 1992). Condia are dislodged by wind and water (Stover, 1980). Mature perithecial ascocarps need to be impregnated with water before ascospores can be discharged (Stover, 1976). As UV radiation has been shown to affect the viability of ascospores (Parnell et al., 1998), maximum survival could be expected during periods of heavy cloud cover as is experienced during thunderstorms or cyclonic disturbances. Strong winds that occur during these natural phenomena would be expected to carry viable ascospores the maximum distances.

Records showing that black leaf streak is gradually becoming dominant at higher and higher altitudes suggest that M. fijiensis may be slowly adapting to cooler temperatures (Carlier et al., 2000a).


Associations

M. fijienis is well suited for the environmental conditions prevailing in the tropical coastal areas and has now virtually replaced M. musicola, which is the pathogen causing Sigatoka, in most of these locations. This process took 2-3 years in coastal Honduras and less than 5 years in coastal Costa Rica. Lesion expansion and ascospore production is greater for M. fijiensis in wet tropical environments and this has probably given it a competitive advantage over M. musicola. However, the situation is different at altitude as M. musicola seems more suited to cooler environments.The optimum growth of ascospore germ tubes of M. musicola is at 25°C, which is 2°C lower than that the optimum for M. fijiensis (Porras and Pérez, 1997). This physiological difference may explain the dominance of M. musicola in upland areas.

In South-East Asia, M. fijensis was recorded as present in Indonesia, peninsular Malaysia and Thailand in the 1960s, but it has not become the dominant leaf spot pathogen. Eumusae leaf spot caused by M. eumusae appears to be common and widespread in peninsular Malaysia and Thailand and in these countries this pathogen may be outcompeting M. fijiensis. In Java, its lack of dominance over M. musicola is harder to explain, but may be related to environmental factors and the great genetic diversity of banana cultivars which are mainly grown in mixed plantings.

Means of Movement and Dispersal

Top of page Natural dispersal

Rain splash and wind are believed to carry conidia from leaf to leaf and plant to plant thus initiating new infections. Ascopsores, which are forcibly ejected from perithecial ascocarps and are the most common form of inoculum, have the potential to be carried for longer distances. During overcast and windy conditions, such as occurs during tropical disturbances, ascospores may remain viable for extended periods and spread the pathogen for considerable distances. However, dispersal over a distance of a few hundred kilometres is considered unlikely (Parnell et al., 1998).

Agricultural practices

Sword suckers, which are an important form of planting material, especially in developing countries, usually have some attached leaf material. This material could be infected by the pathogen. Therefore, if affected sword suckers are moved long distances prior to planting, the pathogen could be disseminated to new areas. The intercontinental spread of M. fijiensis is believed to have occurred in this way.
The chances of spread of M. fijiensis may be much less if corm pieces, with no leaf material, are used as planting material. However, there is a possibility that viable spores could be carried passively on the surface of such material and later be in a position, perhaps by rain splash, to infect leaves.

The risk of spread is eliminated if planting material is moved in tissue culture. This is recommended for the international movement of banana germplasm (Jones and Diekmann, 2000; Jones 2002)

Movement in trade

In some societies, banana leaves are used as ornaments on special occasions and for wrapping foods. In most banana producing countries, banana leaves are used to cushion and protect banana fruit from the sun during transportation from places of production to market. The long distance movement of leaves that could be infected by pathogens is viewed as a quarantine risk. The spread of black leaf streak in Central America may have been accelerated when plantains and green reject export bananas cushioned by infected leaves were trucked across international borders (Stover, 1980). Imported banana leaves and leaf trash should be destroyed. Leaf trash may be found in boxes of commercially packed banana fruit (Jones, 2002).

A recent paper from Venezuela suggests that small sporulating lesions of M. fijiensis can form on fruit of 'Harton' (AAB, Plantain subgroup) (Cedeno et al., 2000). If this report is accurate, then it is possible for the pathogen to be disseminated with plantain fruit. However, Sigatoka lesions have never been reported on fruit of Cavendish cultivars, which is the main commodity of the banana trades (Jones, 2002).

The risk from spores of banana pathogens carried passively on or with fruit is also a quarantine consideration. Research in Brazil has found that large numbers of conidia of M. fijiensis are present on the surfaces of fruit of 'Prata Anã' (AAB, Pome subgroup) from areas where black leaf streak is present. These spores retain their viability for 18 days. On cardboard and polyethylene surfaces, such as is used in packaging banana fruit, viability of spores has been found to extend to 30 days (L. Gasparotto, EMBRAPA, 2002, personal communication). However, it is difficult to show that spores in these situations would initiate infections. One would suppose that there is a chance that conidia on fruit and packaging discarded in banana plots could be splashed onto leaves during rainstorms (Jones, 2002).

Plant Trade

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

Impact Summary

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CategoryImpact
Animal/plant collections None
Animal/plant products None
Biodiversity (generally) Negative
Crop production Negative
Environment (generally) Negative
Fisheries / aquaculture Negative
Forestry production None
Human health Negative
Livestock production None
Native fauna None
Native flora None
Rare/protected species None
Tourism None
Trade/international relations None
Transport/travel None

Impact

Top of page Black leaf streak disease of banana has spread to all major banana-growing regions of the world since it was first recognised in Fiji in 1963. In the Pacific, it has been recorded in American Samoa, Australia (Torres Strait and Cape York Peninsula), the Cook Islands, Fiji, French Polynesia, Hawaii (USA), Micronesia, New Caledonia, Niue, Norfolk Island, Papua New Guinea, the Solomon Islands, Tonga, Vanuatu, Wallis and Fortuna Islands and Samoa. The disease has been found in many countries in Latin America namely Belize, Bolivia, Brazil (Amazonia), Colombia, Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panama and Venezuela. In the Caribbean region, black leaf streak has been identified in Florida (USA), Cuba, Jamaica, the Dominican Republic and now poses a threat to the export industry based in the Windward Islands. West African countries with the disease are Benin, Cameroon, the Central African Republic, Congo, Côte d'Ivoire, the Democratic Republic of Congo, Gabon, Ghana, Nigeria, S¦o Tomé and Togo. In the East African region, black leaf streak has been found in Burundi, Comoros, Kenya, Madagascar, Malawi, Mayotte, Rwanda, Tanzania (including Zanzibar), Uganda and possibly Zambia. Bhutan, China (Hainan, Guangdong and Yunnan), Indonesia (Halmahera, Java, Kalimantan and Sumatra), Malaysia (Johore, Langkawi Island and Sarawak), the Philippines, Singapore, Taiwan, Thailand and Vietnam have been found with the disease in Asia. However, the situation is more confused in this region because M. fijiensis is not common at all locations. Competition from other leaf spots and the diversity of banana germplasm with varying degrees of disease resistance may account for its erratic distribution (Carlier et al., 2000; Jones, 2003).

Black leaf streak is a major constraint to banana production in most countries where it occurs (Stover, 1983, 1986; Fouré, 1985; Fullerton, 1987; Stover and Simmonds, 1987). After the first occurrence of black leaf streak in an area, the disease usually builds up and often reaches an epidemic level in a few years (Fullerton and Stover, 1990; Belalcazar, 1991). Chemical control costs and crop losses are well documented for industrially produced bananas (Stover, 1986, 1990). Losses to smallholders' crops that are consumed locally are harder to estimate.

Black leaf streak does not kill plants immediately, but crop losses increase gradually with the age of plantings. The decrease in functional leaf area caused by the disease results in a reduction in the quality and quantity of fruit (Stover, 1983; Stover and Simmonds, 1987; Pasberg-Gauhl, 1989; Mobambo et al., 1993, 1996b). Fruit from infected plants ripens prematurely and does not properly fill. Bananas for export are sometimes harvested at a lower grade (younger age) in order to reduce the risks of premature ripening in transit to overseas markets (Stover and Simmonds, 1987).

Until the 1970s, the common leaf diseases of plantain were not considered economically important. This changed when black leaf streak spread to areas where the crop was extensively grown. All over the tropics, plantain is cultivated and fruit consumed by smallholders. In many areas, black leaf streak has caused a considerable decrease in the availability of fruit for local consumption and this has resulted in a substantial increase in their market price. Smallholders growing plantain in the Americas either go out of business, because they cannot cover the high costs of chemical control, or form cooperatives so that their limited resources can be pooled to fight the disease.

Black leaf streak is endangering the food security of resource-poor people. Africa alone contributes about 50% of the world plantain production and the demand for plantain is steadily increasing (Wilson, 1987). All known plantain cultivars (Fouré, 1985; Mobambo et al., 1996a) are susceptible to black leaf streak and are severely defoliated by the disease. Plants in the ratoon crop are weaker than in the first cycle and thus more affected by wind damage. On poor sandy soils in West Africa, Mobambo et al. (1996b) estimated that yield losses due to black leaf streak are 33% and 76% during the first and second cropping cycle respectively. However, in intensively cropped backyard or home garden systems, cultivation is not so seriously affected (Mobambo et al., 1994). Under marginal conditions, plantain production is often abandoned due to low yields.

Plantain is not the only smallholder banana to be affected in Africa. The disease also causes serious damage to East African highland cultivars in the Lujugira-Mutika subgroup (AAA). In Uganda, Tushemereirwe (1996) reported yield losses of 37% due to the effects of a leaf spot complex consisting mainly of black leaf streak and Cladosporium leaf speckle.

Black leaf streak has had a devastating effect on the production of export bananas in the South Pacific. Firman (1972) noted that only 49% of unsprayed Cavendish cultivars produced fruit that reached the export quality standard. Fiji ceased exporting bananas in 1974 and Samoa in 1984. Exports also dropped in Tonga and the Cook Islands, because producers had problems maintaining fruit quality standards for their markets in New Zealand. Black leaf streak control has become the single largest production cost (Fullerton, 1987).

In 1974, the production of dessert bananas and plantains in Central America was seriously affected by hurricane Fifi, which was also thought to be responsible for the wind borne spread of black leaf streak to new areas. In many countries, production subsequently dropped substantially. Before 1974, Honduras exported 500,000 boxes of plantain each year, but afterwards exports dropped to below 1,000 boxes (Stover, 1983). In 1978, the export of plantain from Honduras to the USA was curtailed because of the shortage of fruit with the required quality (Bustamente, 1983). Plantain exports only resumed in 1985, when black leaf streak was controlled by the aerial application of fungicides (Stover, 1987).

After the identification of black leaf streak in Costa Rica in 1979, the government initiated a quarantine programme (Woods, 1980). This programme consisted of the destruction of host plants in the affected area and the establishment of roadside quarantine stations strategically located to stop movements of banana and plantain leaves which were used for padding and shading fruit. About 3,000 hectares of plantain were destroyed in 1979 and early 1980. The Costa Rican government paid about US$ 3 million for the eradication program, but the spread of the disease could not be stopped (Woods, 1980). By 1982, the Ministry of Agriculture in Costa Rica estimated that black leaf streak alone reduced plantain production by 40% (Romero, 1986). In general, yields of plantains in well maintained fields on rich fertile soils in Central America may have fallen by 20-50% (Stover, 1983a; Pasberg-Gauhl, 1989).

Black leaf streak was first detected in Panama in 1980. Bureau (1990) estimated that plantain production in Panama decreased by 69% between 1979 (100,910 t) and 1984 (31,134 t). During this period, the price of plantains rose by up to 50% in local markets. Jaramillo (1987) reported that between 1982 and 1985, the area planted with plantain decreased by 22% from 7432 ha to 5800 ha. About 34% of growers were believed to have abandoned their holdings leading to a decrease in production of 47%.

Colombia is one of the largest plantain producers in Latin America with 400,000 ha under cultivation and an estimated yearly production of 2.5 million t. About 96% of plantains are consumed locally, the remainder being exported (Belalcazar, 1991). Smallholders grow about 88% of the plantain in association with coffee. Only 12% of the crop is grown in monoculture in larger plantations. After the introduction of black leaf streak, this staple food became scarce and much higher prices were demanded in the market (Belalcazar, 1991). Due to the high cost of plantain, consumers changed to other, cheaper food crops. This in turn had a negative effect on plantain production. Black leaf streak has thus had a significant impact on Colombian agriculture and the eating habits of a nation.

Black leaf streak can be chemically controlled on plantations, but the cost is substantial. Up to 36 spray cycles per year may be required for plantations growing dessert bananas for export and up to 19 cycles for commercial plantings of plantain (Fouré, 1983, 1988a, b; Stover, 1980b, 1990; Belalcazar, 1991; Gauhl, 1994; Romero and Sutton, 1997b). Stover and Simmonds (1987) reported that 27% of production costs in dessert banana plantations was spent controlling black leaf streak. From 1972 until 1985, the estimated cost of black leaf streak control in Central America, Colombia and Mexico was more than US$ 350 million (Stover and Simmonds, 1987). The cost of chemical control measures has been calculated to be US$ 400-1,400 per ha per year (Anon., 1993). During the 1980s, the cost of black leaf streak control in export banana crops in Costa Rica was estimated at approximately US$ 17.5 million per year (Stover and Simmonds, 1987). Between 1985 and 1994, the area under banana cultivation increased from approximately 21,000 ha to 52,737 ha (Serrano and Marín, 1998). As a consequence, the cost of black leaf streak control in 1995 was estimated to have increased to US$ 49 million per year (Romero and Sutton, 1997b). Recent information from Venezuela indicates that the cost of black leaf streak control in plantain farms south of Lake Maracaibo amounts to almost 50% of production costs. However, even with control measures being taken, yields were still 25% below those before the appearance of the disease in that country (Zabala and Bermudez, 1999).

Environmental Impact

Top of page The extensive use of chemicals is believed to be having a detrimental effect on the environment of countries where inoculum pressures are high and aerial spraying is practised on a large scale, such as Costa Rica. The health of plantation workers as a result of exposure to chemicals has also been raised. Numerous websites are devoted to this topic.

Impact: Biodiversity

Top of page In some countries, the affect of black leaf streak on the yield of some cultivars has been so great that they are being replaced with cultivars that are more resistant to the disease (Watson, 1993). If this trend continues, it could mean that the diversity of cultivated banana germplasm will be eroded.

Diagnosis

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Overnight incubation of lesions at the streak stage under 100% RH and at 25°C should produce abundant conidia and conidiophores for identification. However, these closely resemble those caused by M. musicola, the cause of Sigatoka disease, and M. eumusae, the cause of eumusae leaf spot. A thickened basal hilum on the (on average) longer conidia and scars on the conidiophores where conidia have detached, distinguish M. fijiensis from M. musicola and M. eumusae.

Cultures can be initiated from ascospores. Necrotic leaf tissue with abundant advanced spots should be incubated in a humid chamber for 48 hours. The tissue should then be cut into small squares no bigger than 5 x 5 cm and soaked in 2% sodium hypochlorite solution for 5 minutes. After rinsing in tap water, the squares of tissue are stapled to a piece of paper towel and immersed in water for 3 minutes. The paper towel is then pressed against the lid of a Petri dish with the leaf surface facing the agar. With the paper stuck to the inside, the lid is placed on a Petri dish containing water agar. The surface of the agar is regularly checked for ascospores which discharge from ascomata. Individual, germinating ascospores can be transferred to nutrient agar for further growth and development. Conidia produced on colonies can be used for diagnostic purposes and in inoculation studies.

Zapater et al. (2008) detail a protocol for diagnosing M. fijiensis based on observation of anamorphs.

Mycosphaerella fijiensis and M. musicola can also be distinguished in culture and leaf tissue by a polymerase chain reaction technique (Johanson and Jeger, 1993; Carlier et al., 2000). Molina and Kahl (2004) developed locus-specific simple sequence repeat (SSR) markers for M. fijiensis and M. musicola, for use in PCR methods. Real-time PCR assays were successfully used to diagnose M. fijiensis, and differentiate it from related species, in an Australian banana plantation (Henderson et al., 1996, 2006; Ioos et al., 2011). TaqMan real-time quantitative PCR assays have also been developed (Arzanlou et al., 2007).

Phylogenetical analysis based on sequences of the ITS of ribosomal DNA from M. fijiensis, M. eumusae, M. musicola and P. musae has confirmed that all are different species (Carlier et al., 2000).

Vázquez-Euán et al. (2012) report on the use of direct colony-polymerase chain reaction (DC-PCR) approach to rapidly distinguish M. fijiensis and M. musicola strains in multiplex PCR reactions.

Detection and Inspection

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Mycosphaerella fijiensis can be detected by direct observation of conidiophores and conidia on banana leaves (Zapater et al., 2008). The disease may be difficult to identify during the early stages of disease development because streak symptoms are caused by many fungal diseases of banana (Carlier et al., 2000). Advanced symptoms of black leaf streak disease may also be confused with those of Sigatoka and Sigatoka-like leaf spots. Diagnosis needs to be undertaken using a light microscope or by the polymerase chain reaction method (Souza and Feguri, 2004). See Similarities to Other Pests and Diagnostic Methods.

Similarities to Other Species/Conditions

Top of page Black leaf streak is difficult to diagnose as symptoms resemble those of Sigatoka, eumusae leaf spot and Phaeoseptoria leaf spot. The causal agent of Phaeoseptoria leaf spot can be distinguished from those causing eumusae leaf spot, Sigatoka and black leaf streak because Phaeoseptoria musae forms conidia in pycnidial conidiomata whereas M. eumusae, M. musicola and M. fijiensis form conidia borne on conidiophores on sporodochia. However, M. eumusae can give the impression that conidia form in pycnidia (Carlier et al., 2000a, b; Crous and Mourichon, 2002) so care must be taken in the identification. The presence of scars on conidia and conidiophores is a feature that can be used to separate M. fijiensis from M. eumusae and M. musicola.

Prevention and Control

Top of page Cultural Control

Removal and destruction of diseased leaves will reduce inoculum levels. If diseased leaves cannot be removed from the plot and burnt, they should be cut from plants and stacked on top of one another. This will prevent ascospores discharging effectively from the lower leaves in the pile. Overhead irrigation encourages disease and under-canopy micro-irrigation is preferable. Plants are also more prone to black leaf streak in sheltered areas where humidity levels are high. Good drainage systems that take surface water rapidly out of plantations can reduce humidity levels.

Host-Plant Resistance

Genetic resistance to black leaf streak is clearly the best long-term goal for disease control, especially for smallholders who cannot afford to purchase chemicals. Cultivars with high levels of resistance include 'Yangambi Km 5' (AAA), 'Mysore' (AAB), 'Pelipita' (ABB), 'Saba' (ABB) and 'Pisang Awak' (ABB). However, these do not suit all local tastes and some are susceptible to Fusarium wilt (Fusarium oxysporum f.sp. cubense). Conventional banana breeding programmes utilize resistance to black leaf streak found in wild species of Musa, notably M. acuminata subsp. burmanicca, subsp. malaccensis and subsp. siamea, and in diploid cultivars such as 'Paka' (AA) and 'Pisang Lilin' (AA). Although a black leaf streak-resistant Cavendish-type dessert banana useful for the export trades has still to be developed, progress has been made towards breeding banana hybrids for local consumption. Plantain hybrids with good resistance have been bred by the International Institute for Tropical Agriculture (IITA), Nigeria, and by the Fundación Hondureña de Investigación Agricola (FHIA), Honduras. FHIA has also bred resistant dessert bananas hybrids which have fruit of a subacid or apple flavour and resistant robust cooking banana hybrids. Other breeding programmes include the Centre de coopération internationale en recherche agronomique pour le développement (CIRAD) in Guadeloupe and the Centre de recherches sur bananiers et plantains (CARBAP) in Cameroon Biotechnological techniques, such as exploiting somaclonal variation, gamma irradiation and genetic engineering, are also being utilised in attempts to produce banana cultivars resistant to black leaf streak (Jones et al., 2000).

Chemical Control

The disease can be controlled by the application of systemic fungicides such as benzimidazoles (benomyl, carbendazim), morpholines (tridemorph), triazoles (difenoconazole, propiconazole, triadimenol, flusilazole, fenbuconazole, bitertanol, tebuconazole, hexaconazole and cyproconazole) and strobulins (azoxystrobin, trifloxystrobin). Protectant fungicides, such as dithiocarbamates (mancozeb in oil or oil-water emulsions) and chlorothalonil in water, can also control the disease during periods of low inoculum pressure. (Carlier et al., 2000; Marin et al., 2003). In countries where disease pressure is high, fungicide sprays need to be applied at very regular intervals for control to be effective. The cultivation of banana for export in Central America has become a chemical intensive industry and the cost of black leaf streak control is high (see Economic Impact).

Resistance to benomyl (Stover 1980a; Fullerton and Tracy, 1984), propiconazole (Romero and Sutton, 1997) and oxystrobin has been reported. Strategies to combat this problem, such as alternating triazoles and morpholines, which have different modes of action, and the application of systemics in mixtures with protectants, have been developed (Carlier et al., 2000a; Marin et al., 2003).

There are concerns that the high use of fungicides in export industries is polluting the environment. Although the number of sprays can be significantly reduced in some countries when used in conjunction with disease prediction systems, this method has not been successful in Central America (Carlier et al., 2000a; Marin et al., 2003).


Biological Control

No biological control methodshave been adopted in commercial plantations. Some bacteria have shown promise, but the challenge of controlling this polycyclic fungus by biological methods is immense (Marin et al., 2003).

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