Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Colletotrichum acutatum
(black spot of strawberry)


Colletotrichum acutatum (black spot of strawberry)


  • Last modified
  • 16 July 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Colletotrichum acutatum
  • Preferred Common Name
  • black spot of strawberry
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Fungi
  •     Phylum: Ascomycota
  •       Subphylum: Pezizomycotina
  •         Class: Sordariomycetes
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Preferred Scientific Name

  • Colletotrichum acutatum Simmonds ex Simmonds

Preferred Common Name

  • black spot of strawberry

Other Scientific Names

  • Colletotrichum xanthii Halsted

International Common Names

  • English: crown rot (of anemone and celery); leaf curl of anemone; post-bloom fruit drop of citrus; terminal crook disease (of pine)
  • Spanish: antracnosis del fresón; manchas negras del fresón
  • French: anthracnose du fraisier; taches noires du fraisier

EPPO code

  • COLLAC (Glomerella acutata)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Fungi
  •         Phylum: Ascomycota
  •             Subphylum: Pezizomycotina
  •                 Class: Sordariomycetes
  •                     Subclass: Sordariomycetidae
  •                         Family: Glomerellaceae
  •                             Genus: Colletotrichum
  •                                 Species: Colletotrichum acutatum

Notes on Taxonomy and Nomenclature

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The classification of the genus Colletotrichum is currently very unsatisfactory, and several species occur on the principal economic host (strawberry) which are regularly confused. As well as C. acutatum, these include the Glomerella cingulata anamorphs C. fragariae and C. gloeosporioides, all of which can be distinguished by isoenzyme analysis (Bonde et al., 1991). Studies are continuing. Colletotrichum xanthii appears to be an earlier name for C. acutatum, but more research is necessary before it is adopted in plant pathology circles.


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Colonies in culture are usually white, pale grey or pale orange, sometimes producing strong pinkish-purple pigments. Conidiomata are usually poorly developed, with few or no setae, especially in culture. Conidiogenous cells are roughly cylindrical, sometimes borne in weak clusters, and produce conidia successively from single loci. Conidia are 8-16 x 2.5-4 µm in size, fusiform, thin-walled, aseptate and hyaline. Appressoria are few in number, 6.5-11 x 4.5-7.5 µm in size, clavate to circular and light to dark brown.

Full descriptions are given by Dyko and Mordue (1979), Sutton (1980), Baxter et al. (1983) and Gunnell and Gubler (1992).


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Some country records may refer instead to the Glomerella cingulata-Colletotrichum fragariae aggregate.

A record of C. acutatum in Chile (EPPO, 2009; CABI/EPPO, 2010) published in previous versions of the Compendium has been removed as the pathogen in the original source (Peredo et al., 1979) is now confirmed as a separate species, Colletotrichum pseudoacutatum (Damm et al., 2012). C. acutatum is a quarantine pest for Chile (Servicio Agrícola y Ganadero, 2013).

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


ChinaPresentCABI/EPPO, 2010; EPPO, 2014
-ChongqingPresentChen et al., 2016
-FujianPresentCABI/EPPO, 2010
-HainanPresentCABI/EPPO, 2010; EPPO, 2014
-HenanPresentCABI/EPPO, 2010
-Hong KongPresentCABI/EPPO, 2010; EPPO, 2014
-HubeiPresentCABI/EPPO, 2010; EPPO, 2014
-HunanPresentXia et al., 2011
-JiangsuPresentCABI/EPPO, 2010; EPPO, 2014
-LiaoningPresentXu et al., 2013; EPPO, 2014
-NingxiaPresentCABI/EPPO, 2010
-ShaanxiPresentCABI/EPPO, 2010
-YunnanPresentCABI/EPPO, 2010
IndiaPresentCABI/EPPO, 2010; EPPO, 2014
-Andhra PradeshPresentCABI/EPPO, 2010
-AssamPresentCABI/EPPO, 2010; EPPO, 2014
-ChhattisgarhPresentCABI/EPPO, 2010
-Indian PunjabPresentCABI/EPPO, 2010; EPPO, 2014
-KeralaPresentCABI/EPPO, 2010
-Tamil NaduPresentCABI/EPPO, 2010
-Uttar PradeshPresentCABI/EPPO, 2010
-UttarakhandPresentCABI/EPPO, 2010
IndonesiaPresentCABI/EPPO, 2010; EPPO, 2014
-JavaPresentCABI/EPPO, 2010
-SumatraPresentCABI/EPPO, 2010
IranPresentZafari and Hamadani, 2009; Mousakhah and Khodaparast, 2012
IsraelPresentCABI/EPPO, 2010; EPPO, 2014
JapanPresentCABI/EPPO, 2010; EPPO, 2014
-HokkaidoPresentCABI/EPPO, 2010
-HonshuPresentCABI/EPPO, 2010; EPPO, 2014
-KyushuPresentCABI/EPPO, 2010
-ShikokuPresentCABI/EPPO, 2010
Korea, Republic ofPresentCABI/EPPO, 2010; EPPO, 2014
MalaysiaPresentCABI/EPPO, 2010; EPPO, 2014
-Peninsular MalaysiaPresentCABI/EPPO, 2010
-SabahPresentCABI/EPPO, 2010
NepalPresentCABI/EPPO, 2010; EPPO, 2014
Saudi ArabiaPresentAmmar and El-Naggar, 2011
Sri LankaPresentCABI/EPPO, 2010; EPPO, 2014
TaiwanPresentCABI/EPPO, 2010; EPPO, 2014
ThailandPresentCABI/EPPO, 2010; EPPO, 2014
TurkeyPresentCABI/EPPO, 2010; EPPO, 2014


CongoPresentCABI/EPPO, 2010
EgyptPresentCABI/EPPO, 2010; EPPO, 2014
EthiopiaPresentCABI/EPPO, 2010; EPPO, 2014
KenyaPresentCABI/EPPO, 2010; EPPO, 2014
MauritiusPresentCABI/EPPO, 2010
NigeriaPresentCABI/EPPO, 2010; EPPO, 2014
South AfricaPresentCABI/EPPO, 2010; EPPO, 2014
TanzaniaPresentCABI/EPPO, 2010; EPPO, 2014
TunisiaPresentSayeh et al., 2016
UgandaPresentCABI/EPPO, 2010
ZimbabwePresentCABI/EPPO, 2010; EPPO, 2014

North America

CanadaRestricted distributionCABI/EPPO, 2010; EPPO, 2014
-British ColumbiaPresentCABI/EPPO, 2010; EPPO, 2014
-ManitobaPresentCABI/EPPO, 2010; EPPO, 2014
-New BrunswickPresentCABI/EPPO, 2010; EPPO, 2014
-Nova ScotiaPresentCABI/EPPO, 2010; EPPO, 2014
-OntarioPresentCABI/EPPO, 2010; EPPO, 2014
-QuebecPresentCABI/EPPO, 2010; EPPO, 2014
MexicoPresentCABI/EPPO, 2010
USAPresentCABI/EPPO, 2010; EPPO, 2014
-AlabamaPresentCABI/EPPO, 2010
-ArkansasPresentCABI/EPPO, 2010; EPPO, 2014
-CaliforniaPresentCABI/EPPO, 2010; Swain et al., 2012; EPPO, 2014
-ConnecticutPresentCABI/EPPO, 2010; EPPO, 2014
-FloridaPresentCABI/EPPO, 2010; EPPO, 2014
-GeorgiaPresentCABI/EPPO, 2010
-KentuckyPresentCABI/EPPO, 2010
-LouisianaPresentCABI/EPPO, 2010; EPPO, 2014
-MarylandPresentCABI/EPPO, 2010
-MassachusettsPresentCABI/EPPO, 2010
-MichiganPresentCABI/EPPO, 2010; Rodriguez-Salamanca et al., 2012
-MississippiPresentCABI/EPPO, 2010; EPPO, 2014
-MissouriPresentCABI/EPPO, 2010; EPPO, 2014
-New MexicoPresentFrench et al., 2013; EPPO, 2014
-New YorkPresentCABI/EPPO, 2010; EPPO, 2014
-North CarolinaPresentCABI/EPPO, 2010; EPPO, 2014
-OhioPresentCABI/EPPO, 2010; EPPO, 2014
-OklahomaPresentCABI/EPPO, 2010; EPPO, 2014
-PennsylvaniaPresentPollok et al., 2012
-Rhode IslandPresentCABI/EPPO, 2010
-South CarolinaPresentCABI/EPPO, 2010
-TennesseePresentCABI/EPPO, 2010
-VirginiaPresentCABI/EPPO, 2010
-WashingtonPresentCABI/EPPO, 2010

Central America and Caribbean

BelizePresentCABI/EPPO, 2010; EPPO, 2014
Costa RicaPresentCABI/EPPO, 2010; EPPO, 2014
DominicaPresentCABI/EPPO, 2010; EPPO, 2014
Dominican RepublicPresentCABI/EPPO, 2010; EPPO, 2014
JamaicaPresentCABI/EPPO, 2010
Saint LuciaPresentCABI/EPPO, 2010

South America

ArgentinaRestricted distributionCABI/EPPO, 2010; Sir et al., 2012; EPPO, 2014
BrazilRestricted distributionCABI/EPPO, 2010; EPPO, 2014
-GoiasPresentCABI/EPPO, 2010
-Minas GeraisPresentCABI/EPPO, 2010; EPPO, 2014
-Rio Grande do SulPresentCABI/EPPO, 2010
-Santa CatarinaPresentCABI/EPPO, 2010
-Sao PauloPresentCABI/EPPO, 2010; EPPO, 2014
ChileAbsent, invalid recordCABI/EPPO, 2010; EPPO, 2014
ColombiaPresentCABI/EPPO, 2010; EPPO, 2014
EcuadorPresentCABI/EPPO, 2010; EPPO, 2014
GuyanaPresentCABI/EPPO, 2010
UruguayPresentCABI/EPPO, 2010; Alaniz et al., 2012; EPPO, 2014
VenezuelaPresentCedeño et al., 2007; CABI/EPPO, 2010; EPPO, 2014


AustriaPresent, few occurrencesCABI/EPPO, 2010; EPPO, 2014
BelgiumPresentCABI/EPPO, 2010; EPPO, 2014
Bosnia-HercegovinaPresentCABI/EPPO, 2010
BulgariaRestricted distributionCABI/EPPO, 2010; EPPO, 2014
CyprusPresentCABI/EPPO, 2010
Czech RepublicRestricted distributionCABI/EPPO, 2010; Víchová et al., 2013; EPPO, 2014
DenmarkPresentCABI/EPPO, 2010; EPPO, 2014
EstoniaAbsent, confirmed by surveyEPPO, 2014
FinlandRestricted distributionCABI/EPPO, 2010; EPPO, 2014
FrancePresentCABI/EPPO, 2010; EPPO, 2014
-France (mainland)PresentCABI/EPPO, 2010
GermanyPresent, few occurrencesCABI/EPPO, 2010; EPPO, 2014
GreecePresentIliadi et al., 2018
HungaryPresent, few occurrencesIrinyi and Kövics, 2008; CABI/EPPO, 2010; EPPO, 2014
IrelandPresent, few occurrencesEPPO, 2014
ItalyPresentCABI/EPPO, 2010; Mari et al., 2012; EPPO, 2014; Vitale and Infantino, 2014; Vitale et al., 2015; Frisullo et al., 2016
-Italy (mainland)PresentCABI/EPPO, 2010
-SicilyPresentPolizzi et al., 2011
LatviaPresentCABI/EPPO, 2010
LithuaniaPresent, few occurrencesCABI/EPPO, 2010; EPPO, 2014
MaltaRestricted distributionCABI/EPPO, 2010; EPPO, 2014
MontenegroPresentLatinovic et al., 2012; EPPO, 2014
NetherlandsPresentNPPO of the Netherlands, 2013; CABI/EPPO, 2010; EPPO, 2014
NorwayPresentCABI/EPPO, 2010; EPPO, 2014
PolandPresentCABI/EPPO, 2010
PortugalPresentCABI/EPPO, 2010; EPPO, 2014
SerbiaPresentCABI/EPPO, 2010; EPPO, 2014
SloveniaPresent, few occurrencesCABI/EPPO, 2010; EPPO, 2014
SpainRestricted distributionCABI/EPPO, 2010; EPPO, 2014
-Spain (mainland)PresentCABI/EPPO, 2010
SwedenPresent, few occurrencesCABI/EPPO, 2010; EPPO, 2014
SwitzerlandPresentCABI/EPPO, 2010; Michel et al., 2011; Michel et al., 2013; EPPO, 2014
UKRestricted distributionCABI/EPPO, 2010; Baroncelli et al., 2014; EPPO, 2014
-Channel IslandsPresentCABI/EPPO, 2010; EPPO, 2014
-England and WalesRestricted distributionCABI/EPPO, 2010; EPPO, 2014
-Northern IrelandPresentCABI/EPPO, 2010
-ScotlandAbsent, intercepted onlyCABI/EPPO, 2010


AustraliaPresentCABI/EPPO, 2010; EPPO, 2014
-New South WalesPresentCABI/EPPO, 2010; EPPO, 2014
-QueenslandPresentCABI/EPPO, 2010; EPPO, 2014
-South AustraliaPresentCABI/EPPO, 2010
-TasmaniaPresentCABI/EPPO, 2010
-VictoriaPresentCABI/EPPO, 2010; EPPO, 2014
-Western AustraliaPresentCABI/EPPO, 2010; EPPO, 2014
GuamPresentCABI/EPPO, 2010
New ZealandPresentCABI/EPPO, 2010; EPPO, 2014
Papua New GuineaPresentCABI/EPPO, 2010
VanuatuPresentCABI/EPPO, 2010; EPPO, 2014

Risk of Introduction

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C. acutatum has not been considered to be a quarantine pest by EPPO or any other regional plant protection organization. A certain ambiguity remains on its geographical distribution and impact on the strawberry crop, due to confusion with other Colletotrichum spp. In several countries of mainland Europe, the names C. fragariae or C. gloeosporioides have been used for all fungi causing anthracnose on strawberry. C. acutatum was only described on strawberry in the 1960s (Simmonds, 1966) and it is not clear whether its subsequent appearance as a strawberry pathogen in the literature is due to geographical spread of a pathogen which previously had a restricted distribution, to the rise in importance of a pathogen which was previously insignificant, or simply to the clarification of a taxonomic situation which was previously confused. As C. acutatum attacks several other crops without being a serious cause of concern, and indeed many other plant species, it does not appear logical to attempt to control it by international phytosanitary measures. In addition, identification in imported consignments presents difficulties because of the confusion with related species. Pathogen-free certification of strawberry planting material seems the best approach.

Phytosanitary Measures

The inclusion of C. acutatum (and other Colletotrichum spp.) among the species covered by a strawberry certification scheme would ensure that healthy planting material is traded nationally and internationally. A suitable scheme has been recommended by EPPO (OEPP/EPPO, 1994).

Hosts/Species Affected

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The species has a very wide host range, but is economically most important on strawberries.

C. acutatum can apparently affect almost any flowering plant, especially in warm temperate or tropical regions, although its host range needs further clarification. It has rarely been noted on other than agricultural or forestry land.

Growth Stages

Top of page Post-harvest


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The spread of the disease is often so rapid that by the time symptoms are noticed, the crop is in serious danger. For strawberry, fruit and occasionally petiole rots may be noticed, with sunken, water-soaked spots enlarging to cover the whole fruit within 2-3 days, with dark-brown fruit bodies producing pink spore masses. For other crops such as anemone and celery, crown rots and leaf curl may be the principal symptoms. In pine seedlings, the developing leaves around the apical bud are affected, with small, brown lesions appearing and rapidly extending. Severe stunting is eventually caused as the uninfected tissue beneath the apex continues to develop.

List of Symptoms/Signs

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SignLife StagesType
Fruit / lesions: scab or pitting
Leaves / leaves rolled or folded
Stems / rot
Stems / stunting or rosetting

Biology and Ecology

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The conidia germinate to form appressoria on plant surfaces, from which penetration hyphae develop into plant cells. Infection may occur through almost any plant surface, but for the particularly susceptible herbaceous species such as strawberry and anemone, the crown with its relatively humid microclimate is often favoured. In suitable conditions, the fungus can grow rapidly inside the plant and cause severe symptoms very quickly, but in other circumstances the fungus may be quiescent inside host tissues for a period, in some cases only becoming apparent after harvest. Once the fungus has developed sufficiently inside the plant, dark fruit-bodies are produced, causing typical anthracnose symptoms. Conidia are formed liberally, and are normally dispersed by watersplash (Yang et al., 1992). They may lie dormant in the soil for some time, often overwintering in this fashion. Survival is longest under relatively cool, dry conditions (Eastburn and Gubler, 1992). The fungus can also remain dangerous for long periods in dead plant material on the surface or buried in the soil.

Although the disease in strawberry crops tends to be more virulent in warm climates, where damage can be devastating, it frequently has its origins in cooler conditions where propagating material is grown (Opgenorth et al., 1989; Wilson et al., 1990; Sutton, 1992). The disease may possibly occur in all countries where strawberries are cultivated. However, it is reported to be absent from the premises of most major strawberry propagators in the UK, and it may be possible to exclude the fungus from these sites despite its presence elsewhere in the areas concerned. There is little information on the biology of C. acutatum other than for strawberry crops.

In some crops, notably mango (Liu et al., 1986) and tamarillo (Yearsley et al., 1988), C. acutatum causes postharvest diseases of fruits.

Means of Movement and Dispersal

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Most natural transmission is probably by conidia, although appressoria, hyphal fragments and appressorium-like thick-walled cells may also play a part (Nair et al., 1983). Local dispersal seems to be at least mostly by water-splash (Yang et al., 1990), with propagules sometimes overwintering in soil to affect strawberry crops planted in subsequent years (Eastburn and Gubler, 1990).

Long-distance transmission due to human influence is probably widespread, and has contributed to the rapid spread of the fungus in recent years. The disease is frequently intercepted on strawberry material imported into the UK.

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Bark hyphae; spores Yes Yes Pest or symptoms usually invisible
Flowers/Inflorescences/Cones/Calyx hyphae; spores Yes Yes Pest or symptoms usually visible to the naked eye
Fruits (inc. pods) hyphae; spores Yes Yes Pest or symptoms usually invisible
Leaves hyphae; spores Yes Yes Pest or symptoms usually visible to the naked eye
Roots hyphae Yes Pest or symptoms usually invisible
Stems (above ground)/Shoots/Trunks/Branches hyphae; spores Yes Yes Pest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Growing medium accompanying plants
Seedlings/Micropropagated plants


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The disease is significant worldwide on strawberry (on which it is considered the second most important pathogen after Botrytis cinerea), and also on a few other crops such as anemones. The disease on pine may not now be so severe as in recent years, judging from the decline in research papers. Little detailed information on economic losses is available. In France, the disease has caused up to 80% losses of unsprayed strawberry crops, especially of ever-bearing cultivars (Denoyes and Baudry, 1991). Crops sprayed for B. cinerea control have suffered much less. In the UK, where the disease is statutorily notifiable, presence forces the burning of crops and fumigation of the soil.

Recent studies in Australia showed that C. acutatum caused losses of 25-50% in celery crops in Queensland (Wright and Heaton, 1991).

Detection and Inspection

Top of page No rapid methods exist, although early results from a detection system using monoclonal antibodies are promising. Current tests involve either inoculation of apples with strawberry petioles or paraquat treatment of petioles to stimulate sporulation of the pathogen (Cook, 1993). These tests are time-consuming and labour-intensive.

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.

The only serious research on control has been in connection with strawberry crops. Some success was reported in New Zealand by spraying with dichlofluanid and a captan-benomyl mixture (Cheah and Soteros, 1984), with various chemicals in Australia (Washington et al., 1992), and in South Africa with captan (van Zyl, 1985). Recently in the USA, studies showed that no acceptable fungicide is effective (Milholland, 1989). Fungicide-resistant strains of related species have been reported in the USA and Japan (Chikuo and Kobayashi, 1991; McInnes et al., 1992). There have been considerable efforts in the USA to develop resistant strawberry cultivars, but limited success has been achieved due to the presence of varied races within the species (Delp and Milholland, 1981; Smith, 1985; Smith and Black, 1990; McInnes et al., 1992). Gupton and Smith (1991) have suggested some potentially useful directions for further research.

In the UK, the disease is rare owing to strict quarantine controls and a policy of destroying affected crops and fumigating soil. McInnes et al. (1992) found that nursery material derived from tissue culture which was free from the related species C. fragariae and planted in isolated fields remained healthy, suggesting that careful selection of disease-free stock and soil sterilization in affected beds might be at least as effective as attempting chemical control.

In celery crops, Wright and Heaton (1991) found both a variation in cultivar susceptibility and amenability to chemical control of the disease. For anemone, disease incidence decreased with storage of corms (Doornik and Booden, 1990), and treatment by soaking with hot water proved effective (Doornik, 1990). Yearsley et al. (1988) found that dipping of tamarillos in imazalil and prochloraz reduced the incidence of postharvest disease caused by C. acutatum. However, dipping strawberry plants in hot water or fungicides did not eliminate the disease.

For pine, regular applications of prochloraz have been found to be effective, as has dichlofluanid (Vanner, 1990).


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Alaniz S, Hernández L, Damasco D, Mondino P, 2012. First report of Colletotrichum acutatum and C. fragariae causing bitter rot of apple in Uruguay. Plant Disease, 96(3):458.

Ammar MI, El-Naggar MA, 2011. Date palm (Phoenix dactylifera L.) fungal diseases in Najran, Saudi Arabia. International Journal of Plant Pathology, 2(3):126-135.

Baroncelli R, Sreenivasaprasad S, Lane CR, Thon MR, Sukno SA, 2014. First report of Colletotrichum acutatum sensu lato (Colletotrichum godetiae) causing anthracnose on grapevine (Vitis vinifera) in the United Kingdom. New Disease Reports, 29:26.

Barquero Quirós M, Peres NA, Arauz LF, 2013. Presence of Colletotrichum acutatum and Colletotrichum gloeosporioides on leatherleaf fern, key lime, papaya, star fruit and mango in Costa Rica and Florida (United States). (Presencia de Colletotrichum acutatum y Colletotrichum gloeosporioides en helecho hoja de cuero, limón criollo, papaya, carambola y mango en Costa Rica y Florida (Estados Unidos).) Agronomía Costarricense, 37(1):23-38.

Baxter AP, Westhuizen GCAVan der, Eicker A, 1983. Morphology and taxonomy of South African isolates of Colletotrichum. South African Journal of Botany, 2(4):259-289.

Bonde MR, Peterson GC, Maas GL, 1991. Isozyme comparisons for identification of Colletotrichum spp. pathogenic to strawberry. Phytopathology, 81:1523-1528.

Butin H, Peredo HL, 1986. Bibliotheca Mycologica, 101. Berlin, Stuttgart, Germany: J. Cramer, 100 pp.

CABI/EPPO, 2010. Colletotrichum acutatum. [Distribution map]. Distribution Maps of Plant Diseases, No.October. Wallingford, UK: CABI, Map 705 (Edition 2).

Cedeño L, Briceño A, Fermín G, Domínguez I, Pino H, Quintero K, 2007. First record of Colletotrichum acutatum on lisianthus (Eustoma grandiflorum). Fitopatología Venezolana, 20(2):41-43.

Cheah LH, Soteros JJ, 1984. Control of black fruit rot of strawberry. In: Proceedings of the thirty-seventh New Zealand weed and pest control conference. Hastings, New Zealand: NZ Weed Pest Control Soc., Inc., 160-162.

Chen YJ, Tong HR, Wei X, Yuan LY, 2016. First report of brown blight disease on Camellia sinensis caused by Colletotrichum acutatum in China. Plant Disease, 100(1):227-228.

Chikuo Y, Kobayashi N, 1991. [A study of strawberry anthracnose. Perfect state of the pathogen which originated from the northern area of Kyushu Island and its benomyl resistance]. Proceedings of the Association for Plant Protection of Kyushu, 37:23-26.

Cook RTA, 1993. Strawberry black spot caused by Colletotrichum acutatum. In: Plant health and the European single market. BCPC Monograph, No. 54 [ed. by Ebbels, D.]. Farnham, UK: BCPC, 301-304.

Damm U, Cannon PF, Woudenberg JHC, Crous PW, 2012. The Colletotrichum acutatum species complex. Studies in Mycology, No.73:37-113.

Delp BR, Milholland RD, 1981. Susceptibility of strawberry cultivars and related species to Colletotrichum fragariae. Plant Disease, 65(5):421-423.

Denoyes B, Baudry A, 1991. Characterization of species of Colletotrichum isolated from strawberry in France, taxonomy and pathogenicity abstract. Strawberry Diseases and Breeding for Varietal Resistance International Workshop, Bordeaux 1991.

Doornik AW, 1990. Hot-water treatment to control Colletotrichum acutatum on corms of Anemone coronaria. In: Acta Horticulturae, No. 266. 491-494.

Doornik AW, Booden EMC, 1990. Decrease in viability of Colletotrichum acutatum in corms of Anemone coronaria during storage. In: Acta Horticulturae, No. 266. 505-507.

Dyko BJ, Mordue JEM, 1979. Colletotrichum acutatum. C.M.I. Descriptions of Pathogenic Fungi and Bacteria. Kew, UK: Commonwealth Mycological Institute, No. 630.

Eastburn DM, Gubler WD, 1990. Strawberry anthracnose: detection and survival of Colletotrichum acutatum in soil. Plant Disease, 74(2):161-163.

Eastburn DM, Gubler WD, 1992. Effects of soil moisture and temperature on the survival of Colletotrichum acutatum.. Plant Disease, 76(8):841-842.

EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization.

Freeman S, Horowitz-Brown S, Afanador-Kafuri L, Maymon M, Minz D, 2013. Colletotrichum: host specificity and pathogenicity on selected tropical and subtropical crops. Acta Horticulturae [IV International Symposium on Tropical and Subtropical Fruits, Bogor, Indonesia.], No.975:209-216.

French JM, Randall JJ, Stamler RA, Segura AC, Goldberg NP, 2013. First report of anthracnose of sunflower sprouts caused by Colletotrichum acutatum in New Mexico. Plant Disease, 97(6):838-839.

Frisullo S, Mang SM, Elshafie HS, Camele I, 2016. First report of anthracnose disease caused by Colletotrichum acutatum on Lupinus albus in Italy. Plant Disease, 100(8):1789.

Gunnell PS, Gubler WD, 1992. Taxonomy and morphology of Colletotrichum species pathogenic to strawberry. Mycologist, 84(2):157-165

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