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Australasian Plant PathologyJournal of the Australasian PlantPathology Society ISSN 0815-3191 Australasian Plant Pathol.DOI 10.1007/s13313-013-0241-z

Jackfruit decline caused by Phytophthorapalmivora (Butler)

L. M. Borines, V. G. Palermo,G. A. Guadalquiver, C. Dwyer,A. Drenth, R. Daniel & D. I. Guest

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Jackfruit decline caused by Phytophthora palmivora (Butler)

L. M. Borines & V. G. Palermo & G. A. Guadalquiver & C. Dwyer &

A. Drenth & R. Daniel & D. I. Guest

Received: 30 May 2013 /Accepted: 20 June 2013# Australasian Plant Pathology Society Inc. 2013

Abstract In the late 1990’s a decline syndrome emerged injackfruit orchards in the Eastern Visayas region of thePhilippines. Symptoms included trunk cankers, wilting anddieback of the canopy and, in many cases, tree death. Thedecline resulted in significant yield losses for farmers. A surveywas conducted to assess disease incidence and to identify thecausal organism. Fifty two percent of farms surveyed had adisease incidence greater than 50 %. On some farms 100 % oftrees were affected. While Fusarium, Pythium, Colletotrichumand Phytophthora species were isolated during disease surveysonly Phytophthora was shown to consistently cause declinesymptoms in artificially inoculated jackfruit. Healthy jackfruitseedlings, detached leaves and fruit inoculated withPhytophthora isolates expressed similar symptoms to thoseobserved in the field. Based on morphological and molecularcharacteristics the pathogen was identified as Phytophthorapalmivora.

Keywords Nangka . Root rot . Dieback . Canker .

Oomycete

Introduction

Jackfruit (Artocarpus heterophyllus Lam.) is an importanttree crop in tropical and subtropical regions, grown in homegardens and commercially in orchards (Elevitch and Manner2006; Sangchote et al. 2003). The primary economic productis the fruit, but the tree is also used for timber, fodder, dyesand traditional medicines (Haq 2006). In the Eastern Visayasregion of the Philippines, jackfruit is a high-value fruitmarketed as a ‘flagship commodity’. It has the potential toprovide sustainable incomes for local farmers through thedomestic market, and export opportunities, as quality andproduction increase and infrastructure improves.

From 1996 until 2008, jackfruit was promoted in theEastern Visayas through a ‘plant now, pay later’ (PNPL)Government-run scheme resulting in large-scale plantingsnow covering more than 200 ha on the islands of Samar andLeyte alone. However, the jackfruit plantings have been af-fected by an unidentified decline with symptoms of leafchlorosis, wilting, defoliation, trunk cankers and, in manycases, tree death. The symptoms were particularly severe inareas prone to flooding or poor drainage, or in wet seasons. Insome orchards 100 % of jackfruit trees were affected andfarmers were re-planting with alternative crops. An assess-ment conducted between 2006 and 2008 of the distribution ofjackfruit planting material found that of all the jackfruit seed-lings planted, 63 % were still alive and 44 % were productive(Torres et al. 2011). Diagnosing the cause of jackfruit declinewas critical to the development and dissemination of diseasemanagement strategies and the survival of the industry.

Several diseases of jackfruit have been reported in theliterature, however, details about the incidence and severityof symptoms are scarce and crop losses appear to be minimalcompared to the newly emerged decline reported here (Haq2006; Sangchote et al. 2003). The most significant diseasesreported from major jackfruit growing regions (India, Sri

L. M. Borines :V. G. Palermo :G. A. GuadalquiverDepartment of Pest Management, Visayas State University,Visca, Baybay City 6521, Leyte, Philippines

C. Dwyer :A. DrenthCentre for Plant Science, Queensland Alliance for Agriculture andFood Innovation (QAAFI), The University of Queensland,GPO Box 267, Brisbane, Queensland 4001, Australia

R. DanielElizabeth Macarthur Agricultural Institute,NSW Department of Primary Industries, Private Bag 4008,Narellan 2567, Australia

D. I. Guest (*)Faculty of Agriculture and Environment, The University of Sydney,Biomedical Building C81, Sydney 2006, Australiae-mail: david.guest@sydney.edu.au

Australasian Plant Pathol.DOI 10.1007/s13313-013-0241-z

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Lanka, Bangladesh and Thailand) include leaf spots, dieback,fruit rots and pink disease (Table 1). Many of the diseases aremore severe during wet seasons, under high relative humidityand temperature, or in orchards with poor air circulation. Fruitand blossom rots caused by Rhizopus artocarpi are a signif-icant problem with crop losses as high as 32 % reported

(Soepadmo 1992). The male inflorescences and young fruitsare attacked and only a small proportion of them mature. Theinfected fruit rot slowly, mummify and fall from the tree(McMillan 1974; Roy 1983). Leaf spots are commonly causedbyColletotrichum species,Corynespora cassiicola,Phyllostictaartocarpina and Septoria artocarpi.

Table 1 Summary of known pathogens and diseases of jackfruit

Disease Pathogen Symptoms Reference

Anthracnose Colletotrichum gloeosporiodes Brown lesions on leaves, twigs andfruit; leaf and blossom blights.

Sirayoi 1993, Srivasta and Mehra 2004

Corynespora leaf spot Corynespora cassiicola Irregular spots on leaves and, less often,on stems, roots and flowers. Possibilityof shot-hole development and defoliation

Sangchote et al. 2003

Diplodia fruit and collar rot Diplodia theobromae Lesions on fruit, twigs and branches. Sangchote et al. 2003

Rhizopus rot or transit rot Rhizopus oryzae,R. artocarpi, R. stolonifer

Male inflorescences and young fruit arecovered in a mass of black spores andwhite mycelium; results in poor fruitdevelopment and premature fruit drop;can also affect overripe fruit that havenot been harvested.

Nelson 2005, Butani 1978,Pandey et al. 1979, McMillan 1974

Brown root and crown rot Phellinus noxius Roots and base of tree become encrustedas soil is held together by mycelium.Cracking of bark and gummosis. Woodbecomes discoloured, dry, friable andhoneycombed. Basidiocarps developand girdle base of tree causing wiltingand yellowing of foliage and tree death.

Putter 1998, Trujillo 1971

Phytophthora root, fruitand stem rot

Phytophthora palmivoraand P. citrophthora

Lesions on fruit surfaces. Can causeentire fruit to rot.

Haq 2006

Pink disease Erythricium salmonicolor White patches develop on trunk, branchand twigs and expand to encircle entirebranch appearing salmon pink in colour.Infected branches may wilt and defoliate

Holliday 1980

Pinglap disease Unknown Severe wilt or dieback syndrome resultingin partial or whole tree death

Morton 1987, Trujillo 1970, 1971

Miscellaneous leaf diseases Alternaria sp., Botryodiplodiatheobromae, Cercospora sp.Colletotrichum orbiculare,Nigrospora sphaerica,Pestalotiopsis versicolour,Pseudocercosporaartocarpi, Phyllostictaartocarpina, Septoriaartocarpi

Leaf spots Basak 1995, Butani 1978, Haq 2006,Shaw 1984

Curvularia sp. Infects young shoots and petioles ofleaves

Haq 2006

Phomopsis sp. Defoliation Haq 2006

Pestalotiopsis elasticola Grey leaf blight Vevai 1971, Zhang et al. 2003

Uredo artocarpii Rust Basak 1995, Singh 1980

Root rot Pythium spendens,Phytophthora sp., Fusariumsp., Rhizoctonia sp.,

Infect roots and stem base of seedlings Haq 2006

Nematodes Aphelenchoides sp., Helicotylenchus dihystera, Helicotylenchusmulticinctus, Hemicriconemoides cocophilus, Meloidogune sp.,Pratylenchus sp. and Ximphenema brevicolle

Sangchote et al. 2003

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Pink disease, caused by the fungusErythricium salmonicolor,is widespread, causing wilting and defoliation of infectedtrunks, branches and twigs. The disease is more severe in therainy season, and in trees with a dense canopy. Dieback hasbeen reported to be caused by Botryodiplodia theobromae, afungus that affects growing shoots, causing discolouration ofthe bark as it spreads downwards, ultimately killing the tree(Haq 2006). The bacterium Erwinia carotovora also causesdieback resulting in leaf yellowing and gummy exudates fromthe stems and branches (Ton et al. 1990).

Phytophthora has not previously been confirmed as thecause of dieback and decline of jackfruit. Phytophthora spe-cies, including P. palmivora, have been associated with fruit,stem and root rot of the related species breadfruit (A. integer)and chempedek (A. altilis) (Drenth and Guest 2004; Lim1990). A disease of unknown cause referred to as ‘Pingelap’resulted in severe devastation of breadfruit crops in the Pacificin the 1970s and again in the 1990s (Sangchote et al. 2003;Trujillo 1970). While several fungi have been isolated, includ-ing Rhizopus, Phomopsis and Pythium, none have been shownto be pathogenic. Trujillo (1970) noted that the symptomswere similar to those caused by P. palmivora but the associ-ation has not been confirmed by isolation and reinoculation(Koch’s Postulates).

In the southern Philippines, the newly emerged decline hasaffected the roots, trunks, branches and fruit of jackfruit. Rootand collar rot and stem lesions developed on infected trees,followed by wilting and loss of canopy cover. In many cases,the entire tree died and some orchards had 100 % tree losses.Under high relative humidity or wet conditions, water soakedlesions also developed on leaves and fruit. The decline causeda significant decline in tree health and productivity. In thenursery, affected seedlings wilted and died. A summary ofknown diseases of jackfruit are presented in Table 1. Moredetailed descriptions of jackfruit diseases and disorders aregiven by Haq (2006) and Sangchote et al. (2003).

In order to develop management strategies to reduce theimpact of jackfruit decline it was paramount that the cause ofthe decline was identified so that science-based improve-ments could be developed for the jackfruit orchards anddisseminated to the industry. This study aimed to determinethe cause of jackfruit decline in the Philippines and morespecifically to:

(1) describe the symptoms of the decline of the jackfruit trees(2) determine if the symptoms and cause of the decline

were the same for different regions in the southernPhilippines (Leyte and Samar)

(3) identify the causal agent and confirm Koch’s postulates(4) determine if there was a correlation between disease

incidence and the age of the trees

Materials and methods

Distribution of jackfruit decline

In order to determine the range and extent of symptoms and theimpact of the decline forty-two jackfruit orchards on Leyte andSamar Islands in the Eastern Visayas Region of the Philippineswere surveyed. The number of trees, age, disease symptomsand disease incidence (% of trees showing symptoms) wererecorded. Plant and soil samples were collected from selectedorchards for pathogen isolation.

Pathogen isolation

Tissue samples from canker lesions, roots, leaves and samplesof soil near the rhizosphere of affected trees were collected andtaken back to the laboratory for isolation. Samples were plateddirectly onto onion agar (200 g red onion, 17 g agar/L distilledwater), carrot agar (200 g carrots, 17 g agar/L distilled water)and water agar (17 g agar/L distilled water), supplemented withStreptomycin (0.1 mg/L), Benomyl (0.05 mg/L) and RoseBengal (500 mg/L). Samples were also baited using disinfectedKuomintang (Catharanthus roseus) or eggplant (Solanummelongena) fruit then isolated onto onion agar. Cultures wereincubated at room temperature (23 °C) and examined micro-scopically for growth and characterisation of the organisms.

Pathogenicity studies

Pure cultures of each isolate grown on onion agar (OA) for 4-5 days at 23 °C were used to inoculate detached leaves, fruitand healthy jackfruit seedlings. Jackfruit cultivar EVIARCSweet was used in all experiments. Young, fully emerged leaveswere detached then disinfected with 1 % NaOCl, rinsed threetimes with sterile water and dried with paper tissue. Cut petioleswere wrapped with moist cotton wool and the leaves placedinside sterile Petri dishes lined with sterile moistened papertissue. A mycelial plug (9 mm diameter) was placed mycelialside down onto the surface of the detached leaves. The plug wasremoved after 48 h and the inoculated leaves were observeddaily for the development of lesions. Uninoculated leaves wereincluded as a control. Leaves were incubated at 23 °C undernatural light regimes. The pathogen was re-isolated from leaflesions 5-7 days after inoculation by surface sterilising theleaves in 70% ethanol and plating sections on to onion agar asdescribed above.

To assess the development of symptoms on stems, one-month-old jackfruit seedlings were inoculated by making a V-cut halfway up the stem and inserting an agar plug, mycelialside down, into the slit. The inoculation point was sealed withparafilm. The mycelial plug and parafilm were removed after

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48 h and the stem observed for the development of lesions.The pathogen was re-isolated 24 days after inoculation byplating on to onion agar.

To confirm the infection of roots, one-month-old jackfruitseedlings grown in 25 cm pots filled with 3.5 L potting medium(1:1 w/w garden soil and fine river sand, sterilised in a pressurecooker twice for 1 h at 103 kPa) were inoculated by adding 100 gof mungbean inoculum (autoclaved mungbean seeds inoculated,then incubated in Erlenmeyer flasks at 23 °C for 14 days undernatural light regimes) around the roots. Control plants wereinoculated in the same way using sterile mungbeans. The seed-lings were examined daily for the development of symptoms.After 9 days the seedlings were harvested, roots washed and thepathogen reisolated from soil and roots by baiting with eggplantfruit for 3 days and then plating onto onion agar.

Detached immature jackfruit, 2.5 months from flowering,was sprayed with 70 % ethanol and allowed to air dry. A cut(~10 mm) was made on the fruit using a sterile scalpel and anagar disc of the culture was inserted into the wound and sealedwith transparent tape. The inoculated fruit was placed in indi-vidual crates and wrappedwith nylon tulle bag to prevent entryof insects. The fruit was maintained at room temperature(23 °C) and observed daily for lesion development. The path-ogen was reisolated from lesions by plating onto onion agar.

Pathogen identification

The pathogen was reisolated from inoculated plant tissuesshowing distinctive disease symptoms by surface sterilisingand plating onto modified onion agar. Isolated cultures werecompared to the originally isolated cultures. Colony morphol-ogy was examined after 4-5 days growth at 23 °C under 12 hlight. Small pieces of agar with mycelium were also suspendedin sterile water in Petri dishes to induce sporulation. Wetmounts of each isolate were examined under the microscopeto observe hyphal characteristics and reproductive structures.

Nine isolates were selected for DNA-based identification(Table 2) according to the method of Drenth et al. (2006).These isolates are stored in the Phytophthora culture collec-tion at the University of Queensland.

Results

Disease symptoms

Jackfruit decline affected all parts of the jackfruit tree includingthe roots, trunks, branches, leaves and fruit of jackfruit resultingin a significant reduction in tree health and productivity. Severe

Table 2 Response of jackfruit to inoculation with isolates from jackfruit orchards and the identity of isolates

IsolateNo.

Sample Source Location Symptoms development on inoculatedjackfruit (S = stem canker; L = leaflesion; R = root lesion; W = wilting,F = fruit lesion)

Identification based on cultural,morphological and DNA Characteristics

S L R Wa F

VSU001 Lower trunk canker Abuyog, Leyte X Xb X Pythium sp.

VSU003 Leaf Mahaplag, Leyte X X X X P. palmivora

VSU004 Soil Calbayog, Samar X X X X P. palmivora

VSU005 Durian leaf Calbayog, Samar X X X X P. palmivora

VSU006 Soil Sogod, Leyte Xb X Pythium sp.

UQ7177d Roots Casilda, Biliran, Leyte X Xc X X P. palmivora

UQ7178d Roots Ormoc City, Leyte X X X P. palmivora

UQ7179d Nursery potting mix Abuyog, Leyte X Xc X X P. palmivora

UQ7180d Roots Baybay, Leyte X X X X P. palmivora

UQ7181d Soil DA-RIARC, Abuyog, Leyte X Xc X X X P. palmivora

UQ7182d Roots DA-RIARC, Abuyog, Leyte X X X X P. palmivora

UQ7183d Root with soil Catarman, Northern Samar X X P. palmivora

UQ7184d Roots with soil Salcedo, Eastern Samar X X P. palmivora

UQ7185d Soil DA-RIARC, Abuyog, Leyte X X X P. palmivora

DA-RIARC Department of Agriculturea Following root inoculationb Pin-size, water-soakedc Blightd Selected for DNA/PCR based identification; UQ codes indicate isolates lodged in the Phytophthora collection at the University of Queensland;VSU codes indicate isolates from the collection at VSU

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cankers developed at the collar (Fig. 1a) and on the trunk(Fig. 1b) of infected trees. Cankers were particularly severe inorchards with poor drainage and those that were subjected toperiodic flooding. As the infection progressed, leaves wiltedand became yellow and there was a loss of canopy cover(Fig. 1c). In many cases, the entire tree died. Under highrelative humidity or wet conditions, water soaked lesions wereobserved on leaves and fruit (Fig. 1d).

Disease incidence

Forty-two farms were surveyed on Leyte and Samar Islands.The number of trees per farm ranged from 30 to 4,000 andtheir age ranged from 2 months to 17 years old. Disease

incidence ranged from 0% to 100 % (Fig. 2). Twenty-two ofthe 42 (52.4 %) farms surveyed had a disease incidence of50 % or higher and the median disease incidence was 50 %.There was a weak correlation between disease incidence andorchard age (R2=0.4075; Fig. 3).

Pathogen isolation and characterisation

While several organisms were isolated from diseased tissue,only Phytophthora palmivora and Pythium spp. were foundto be pathogenic on jackfruit under experimental conditions(Table 2). The Phytophthora species was positively identi-fied as P. palmivora based on cultural, morphological andmolecular characteristics (Fig. 4). Phytophthora palmivorainfected all jackfruit tissues that were inoculated and wasreisolated to confirm Koch’s Postulates. Cankers and brownlesions developed on the stem within 4 days of inoculation(Fig. 5a). Infected seedlings that were root-inoculated beganto wilt within 7 days after inoculation (Fig. 5b) and subse-quently died; some as early as 9 days after inoculation(Fig. 5c). Water-soaked, dark lesions, more than 2 cm indiameter developed on inoculated leaves within 4 days ofinoculation (Fig. 5d). Dark lesions also began to develop oninoculated fruit 2 days after inoculation and mycelial growthwas visible after 7 days (Fig. 5e).

Fig. 1 Symptoms of jackfruit decline in the field. a Canker lesions atthe collar of an infected tree developing at the soil line; b Canker lesionson the trunk of an infected tree; c loss of canopy cover as a result ofinfection; d lesions on jackfruit seedlings in a commercial nursery

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ease

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Fig. 2 Disease incidence(percentage infected trees) and injackfruit orchards surveyed onLeyte and Samar Islands. Thedashed line indicates the mediandisease severity

y = 7.3598x

R2 = 0.4075

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Fig. 3 Correlation between disease incidence and age of trees in thejackfruit orchard surveyed on Leyte and Samar Islands. R2=0.4075

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Inoculation of leaves with the Pythium sp. isolates(VSU001 and VSU006) lead to the development of pin-sized, water soaked lesions within 4 days. No canker lesionswere observed on inoculated stems. Yellowing and wilting,attributed to root infection, was observed on inoculatedseedlings, but this did not lead to plant death. No symptomsdeveloped in jackfruit inoculated with Fusarium sp. isolates.

Discussion

This study confirmsP. palmivora as the cause of jackfruit declinedisease in the Eastern Visayas region of the Philippines. The

organism was consistently isolated from jackfruit trees showingsymptoms including trunk cankers, chlorosis and wilting of thefoliage, root lesions and tree death. A description of thesymptomology has been provided above.

When healthy seedlings, leaves and fruit were inoculatedusing the P. palmivora isolates the symptoms that developedwere similar to those observed in the field. Phytophthorapalmivora has previously been isolated from jackfruit in thePhilippines but its association with the complex of symptomsnow referred to as ‘jackfruit decline’ was never confirmed byre-inoculation (Tsao et al. 1994). While other organisms in-cluding Pythium sp. and Fusarium sp. were isolated during thesurveys, they are not considered to be the primary pathogen.

Fig. 4 Cultural andmorphological characteristics ofrepresentative isolates ofPhytophthora palmivora. aCulture grown on onion agar at23 °C for 5 days (UQ7177); bhyphae; c chlamydospores and dpapillate, caducuous sporangiaof representative isolates

Inoc

ulat

edC

ontr

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a b c d e

f g h i j

Fig. 5 Symptoms on jackfruitseedlings, leaves and fruitartificially inoculated withPhytophthora palmivora(VSU005). a lesions on stems ofjackfruit seedlings, b rootlesions, and c wilting ofseedlings 24 days afterinoculation; d leaf lesions(UQ7177) 5 days afterinoculation and e fruit 8 daysafter inoculation compared withf–j control inoculations on theplant parts (sterile water alone)

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Inoculation of jackfruit seedlings with Pythium resulted inroot damage, and the organism may contribute to root diseasein the field. The other organisms did not lead to the develop-ment of symptoms and may be more likely to be involved assecondary inhabitants of damaged tissues.

Phytophthora diseases are among the most economicallysignificant diseases of fruit tree crops in the tropics (Drenthand Guest 2004). The survey of jackfruit orchards in Leyteand Samar indicated that 85 % of orchards were affected bythe decline. Jackfruit decline consists of a range of differentsymptoms all caused by the one pathogen, so it was expectedthat the disease incidence would be greater in older orchardsas the different plant parts from the same and different plantsbecome infected if left untreated and population levels in-crease. While disease was more severe in older orchards, thecorrelation between age and disease severity was not highalthough disease incidence was generally lower on farms withyounger plantings. This may be due either to better manage-ment, following the extension of disease management strate-gies, or insufficient time for disease to become established.

The results of our study confirm P. palmivora as the cause ofjackfruit decline in the southern Philippines. The survey resultsclearly indicate that the decline has a significant impact onyields and the long term viability of plantations, and conse-quently, farmer incomes. The confirmation of the causal organ-ism enables the development of appropriate and effective man-agement strategies.

Acknowledgments This project was conducted as part of the Austra-lian Centre for International Agricultural Research (ACIAR) projectHORT2007/067/2. We acknowledge the Philippine Council for Agri-culture and Aquatic Resources Research and Development, the Depart-ment of Agriculture, Leyte, Visayas State University and the jackfruitfarmers of Leyte and Samar for their support.

References

Basak AB (1995) Fruit rot disease of jackfruit caused by Colletotrichumgloeosporioides Penz in Chittagong. Bangladesh J Bot 24:197–199

Butani DK (1978) Pests and diseases of jackfruit in India and theircontrol. Fruits 33:351–357

Drenth A, Guest DI (2004) Diversity and management of phytophthorain southeast Asia. ACIAR Monograph series no. 114. AustralianCentre for International Agricultural Research, Canberra. http://aciar.gov.au/publication/MN114

Drenth A, Wagels G, Smith B, Sendall B, O'Dwyer C, Irvine G, IrwinJAG (2006)Development of a DNA-basedmethod for detection andidentification of Phytophthora species. Australasian Plant Pathol35:147–159

Elevitch CR, Manner HI (2006) Artocarpus heterophyllus (jackfruit)ver. I.IV. In: Elevitch CR (ed) Species profiles for Pacific IslandAgroforestry. Permanent Agriculture Resouces (PAR) HõlualoaHawai'i. www.traditionaltree.org

Haq N (2006) Jackfruit: Artocarpus heterophyllus. Southampton Centrefor Underutilised Crops, Southampton

Holliday P (1980) Fungus diseases of tropical crops. Cambridge Uni-versity Press, Cambridge

Lim TK (1990) Durian diseases and disorders. Tropical Press Sdn Bhd,Malaysia

McMillan JR (1974) Rhizopus artocarpi rot of jackfruit (Artocarpusheterophyllus)

Morton J (1987) Breadfruit. In: Fruits of warm climates. Morton Col-lectanea. University of Miami, Coral Gables, Florida

Nelson S (2005) Rhizopus rot of jackfruit. www.ctahr.hawaii.edu.au/oc/freepubs/pdf/PD-29.pdf

Pandey RS, Bhargava SN, Shukla DN, Khati DVS (1979) Control ofRhizopus rot of jack fruit. Indian Phytopathol 32:479–480

Putter CAJ (1998) Phellinus noxius. Global Plant Protection Informa-tion System, FAO, Rome

Roy AK (1983) Perpetuation of Rhizopus artocarpi - the incitant ofRhizopus fruit rot of jack fruit (Artocarpus heterophyllus) Indian.Phytopathology 36:344–345

Sangchote S, Wright JG, Johnson GI (2003) Diseases of breadfruit,jackfruit and related fruit crops. In: Ploetz RC (ed) Diseases oftropical fruit crops CAB International Oxon UK

Shaw DE (1984) Microorganisms in Papua New Guinea. ResearchBulletin No. 33. Department of Primary Industry, Port Moresby

Singh (1980) A check list of hosts and diseases in Malaysia. Ministry ofAgriculture Bulletin No. 154:14–15

Sirayoi A (1993) Diseases of fruit crops, spices and control. TakansakFoundation, Kasetsart University

Soepadmo E (1992) Artocarpus heterophyllus Lam. In: Verheij EWM,Coronel RE (eds) Plant resources of South East Asia 2. Ediblefruits and nuts. PROSEA, Bogor

Srivasta MP, Mehra R (2004) Diseases of minor tropical and sub-tropical fruits and their management. In: Naqvi SAMH (ed) Dis-eases of fruits and vegetables, vol II. Springer, Netherlands, pp559–632

Ton HT, Rok MK, Champada K (1990) Die–back: a new bacterialdisease of champedak jack-fruit (Artocarpus sp. (Merr.) Thumb.).J Thai Phytopathol Soc 10:19–29

Torres EC,Garcia PV, Londina LA (2011)Development of regional baselineinformation for jackfruit. Terminal report. DA-RFU-8, Tacloban City

Trujillo EE (1970) Preliminary report on the causes of breadfruit dis-eases in the Trust Territory. Office of the High Commissioner,Trust Territory of the Pacific Islands, Saipan

Trujillo EE (1971) A list of diseases of economic plants in the TrustTerritory of the Pacific Islands. Trust Territory of the PacificIslands Department of Resources and Development, Division ofAgriculture

Tsao PH, Gruber LC, Portales LA, Gochangoco AM, Luzaran PB, Delos Santos AB, Pag H (1994) Some new records of Phytophthoracrown and root rots in the Philippines and in world literature.Phytopathology 84:871

Vevai EJ (1971) Know your crop its pest problems and control – minortropical fruits. Pesticides 5:33–54

Zhang J, Tong X, Qixin G (2003) Notes on Pestalotiopsis from southernChina. Mycotaxon 85:91–99

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