Cocoa canker caused by Phytophthora palmivora

Canker disease of Theobroma cacao L. caused by Phytophthora palmivora (Butler) Butler gives rise to serious damage in Fiji and is common in the S.W. Pacific area. Fusarium decemcellulareBiick (imperfect stage of Calonectria rigidiuscula (Berk. & Br.) Sacc.) was constantly associated with P. palmivora in disorders of cocoa but was not found to be a primary pathogen. Most Amelonado cocoa trees were resistant to canker but there were differences in the reactions of other Forastero and of Trinitario types. The F₁ generation of a local Trinitario selection, itself resistant, showed segregation resulting in a proportion of very susceptible plants. Isolates of P. palmivora from black pod, canker and green twigs were equally pathogenic, both to pods and seedling stems. Sporangial dimensions, particularly the length/breadth ratio, differed from those reported elsewhere. It is uncertain whether the reason that cocoa canker is more serious in some countries than others is because of differences in the pathogenicity of P. palmivora strains to bark or the genetic differences of the cocoa grown.     

Cocoa-based media for culturing Phytophthora palmivora (Butl.) Butl., causal agent of black pod disease of cocoa

Green cocoa pod husk agar (GCPA), ripe cocoa pod husk agar (RCPA), green cocoabean agar (GCBA), ripe cocoa bean agar (RCBA), green cocoa mucilage agar (GCMA)and ripe cocoa mucilage agar (RCMA) were prepared and assessd for their clarity andfor potential to support mycelial growth and sporulation of P. palmivora. Oatmeal agar (OMA), potato-dextrose agar (PDA), vegetable 8 juice agar (V8JA) and pineapple crown agar (PCA) were included for comparison. The highest radial growth rates of 8.3 and 7.2 mm/day were recorded, respectively, on OMA and GCPA but these were not significantly different (P ≤ 0 05) from each other. The two media also supported good aerial mycelial growth but were not clear. Radial mycelial growth rates of 6.5, 7.0 and 6.6 mm/day were obtained on GCMA, RCPA and V8JA, respectively, and these rates were also not significantly different from each other. Of the three media, only the GCMA was clear and supported the best aerial mycelial growth. In comparison, the RCMA supported a significantly lower radial growth (4.6 mm/day) of P. palmivora than the three media. Growth rates were least on RCBA, PCA and PDA but sporulation was poorest on PDA, PCA and V8JA. GCMA was found to be the best medium based on all the growth parameters and media characteristics. GCMA has been used effectively to isolate/detect P. palmivora from infected cocoa pod tissues. Apart from differences in radial growth rate, both the GCMA and RCMA were similar in all other respects and are recommended for culturing P. palmivora. This revised version was published online in June 2006 with corrections to the Cover Date.     

Sudden death of cocoa in Papua New Guinea associated with Phytophthora palmivora cankers invaded by bark beetles

An extensive survey in 1976-82 indicated that sudden death of cocoa was widespread in Papua New Guinea with losses up to 1% per month in trees over 10-yr-old, especially in neglected plantations. Over 95% trees affected had large Phytophthora palmivora canker lesions and over 90% of the cankers were invaded by bark beetles. In an intensive single year survey of one plot 18-8% of 69 healthy trees died. Ambrosiella spp., P. palmivora and Fusarium solani were isolated from wood surrounding beetle tunnels but only the last two caused extensive lesions when inoculated into the xylem of healthy trees. The evidence obtained indicated that the sudden death syndrome is initiated by P. palmivora cankers on trunks and main branches but bark beetles are a major contributory factor and fungi associated with their tunnels may contribute to the problem. There is a risk of further outbreaks of the disease as cocoa cultivars susceptible to P. palmivora continue to be planted.     

Electrophoretic protein patterns of three species ofPhytophthora associated with black pod disease of cocoa (Theobroma cacao L.)

When electrophoretic profiles of native proteins from vegetative mycelia ofPhytophthora palmivora, Phytophthora capsici and Phytophthora citrophthora causing black pod disease of cocoa in India were compared on a single Polyacrylamide gel, the isolates of same species were readily distinguished both qualitatively by visual similarity in banding patterns and quantitatively by calculating similarity coefficients. Similarity coefficients were generally much higher between isolates within a species than between isolates of different species. The dendrograms obtained after unweighted pair grouping with arithmetic averaging cluster analysis, revealed that all the isolates ofPhytophthora capsici were highly homogenous and formed a single cluster. The isolates ofPhytophthora citrophthora were resolved into two electrophoretic types which were clustered into two distinct sub groups.Phytophthora palmivora formed a separate group. Thus, the results reveal that polyacrylamide gel electrophoresis can be used successfully in distinguishing species and sub groups within a species ofPhytophthora encountered on cocoa. CPCRl contribution No. 914.     

Separating effects of species identity and species richness on predation,pathogen dissemination and resistance to invasive species in tropical ant communities

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Phytophthora palmivora from Sulawesi and Java Islands,Indonesia, reveals high genotypic diversity and lack of population structure

Phytophthora palmivora is the causal agent of cocoa black pod disease, one of the primary diseases of cocoa in Indonesia. A better understanding of P. palmivora population genetics is needed to aid the development of relevant disease management strategies. This study is the first population genetic study of P. palmivora in Indonesia using microsatellite markers based on the alleles genotyping method. The microsatellite markers were used to determine the genotype of 44 P. palmivora isolates from Sulawesi (24) and Java (20) islands. The total number of observed multilocus genotypes (MLG) from both populations was 34. The genotypic diversity of P. palmivora from Sulawesi (2.90; 16.0; 0.938) and Java (2.76; 14.3; 0.930) islands was high as seen from Shannon's diversity index (H), Stoddart and Taylor's Index (G), and Simpson's Index (λ) respectively. Evenness and Nei's unbiased gene diversity exhibited similarly high levels from both populations. The linkage disequilibrium test indicated that sexual recombination occurred in the Java population (P = 0.312). Analysis of molecular variance (AMOVA) and Bayesian clustering revealed five genetic clusters, and isolates from both islands were evenly distributed across the five gene clusters. All genetic diversity was from within individuals. P. palmivora from Sulawesi and Java showed a high genotypic diversity but a lack of genetic differentiation among populations (Fst = 0.006). Both populations formed one highly diverse group. Minimum spanning network analysis showed no particular grouping of MLGs, and shared MLGs from both populations indicated long-distance migration of P. palmivora facilitated by human activities.     

Trichoderma from Aceh Sumatra reduce Phytophthora lesions on pods and cacao seedlings

The cocoa tree, Theobroma cacao L., suffers large yield losses in Aceh Indonesia due to the disease black pod rot, caused by Phytophthora spp. Despite having the largest area under cacao production in Sumatra, farmers in the Aceh region have low overall production because of losses to insect pests and black pod rot. Trichoderma spp. were isolated from the roots and leaves of cacao trees and screened as potential biological control agents. Isolates used in the study were Trichoderma asperellum isolates T2 and T4, Trichoderma longibrachiatum isolates T15 and T16, and Trichoderma virens isolates T1 and Tv. T1, T2, T4, and Tv completely colonized and destroyed Phytophthora tropicalis and Phytophthora palmivora mycelium in precolonized plate assays. All six isolates reduced P. tropicalis, but none reduced the growth of P. palmivora in dual plate assays. Phytophthora growth was suppressed on MIN media amended with sterile heat inactivated Trichoderma culture filtrates, with Tv best suppressing growth of both Phytophthora spp. T. virens isolate Tv was the only isolate observed coiling around P. tropicalis mycelium and disrupted the formation of P. palmivora sporangia. Of all six isolates, only Tv reduced P. palmivora lesion expansion in a detached pod assay, reducing severity by 71%. Tv also reduced P. palmivora infection on seedlings when applied aerially at 1 × 10⁶ and 1 × 10⁸ conidia/ml, by 19% and 59%, respectively. T. virens isolate Tv is a mycoparasite, antagonizes Phytophthora in a dual plate assay, and shows antibiosis against Phytophthora spp., suggesting that multiple modes of action contribute to its ability to limit Phytophthora lesion expansion on cacao pods and seedlings.     

Invertebrate vectors of Phytophthora palmivora, causing black pod disease of cocoa in Ghana

Evidence is presented for the involvement of a range of invertebrates in the epidemiology of black pod disease of cocoa. Tent-building ant species, notably Crematogaster striatula, Camponotus acvapimensis and Pheidole megacephala , are considered to contribute to vertical spread of the pathogen within the cocoa tree. The building material used in tent construction is a potential source of inoculum. Certain phytophagous invertebrates and sugar-feeding Diptera colonize black pods in the field and may carry viable inoculum externally, on their bodies and mouthparts, or internally, in their faeces. Two species, Brachypeplus pilosellus (Coleoptera) and Chaetonerius latifemur (Diptera), stand out as important vectors because of their abundance, their habit of visiting wounded pods and their proven efficiency in transmission experiments. These vector species quickly colonize the lesions of pod-wounding insects (mirids, pod borers) and are considered to be the main means of local and long-distance horizontal spread of the pathogen. Radioactive tracers were extensively used to determine the movements of potential vectors.     

Association of Phytophthora palmivora bark canker and insect damage in cocoa in Papua New Guinea

Bark canker caused by the fungus Phytophthora palmivora has caused the death of many cocoa trees over 15 years old in Papua New Guinea in recent years. Cankers are significantly associated with channels made by the larvae of bark-damaging insects, which provide the major entry point for the pathogen. The combination of canker and insect damage may limit the economic life of cocoa to less than 15 years in affected areas.     

Seasonal Changes in Total Free Sugar Content of Field Cocoa Plants

As part of a study of factors affecting the vegetative and reproductive characteristics of cocoa, Theobroma cacao L., (viz. leaf flushing, flowering, cherelle wilt, pod setting and pod growth), seasonal changes in total free sugar levels of the bark and wood of field cocoa trees growing under medium shade provided by Gliricidia sepium Steud. trees were studied over a period of 13 months. On dry weight basis, sugar content of the bark (13 to 63 mg/g) was higher than that of the wood (9 to 40 mg/g). Free sugar content was significantly correlated with sunshine hours for the whole period, except October-November 1975, and March 1976 when pod load, and flowering and leaf flushing respectively, caused considerable reduction in free sugar levels. The peaks of flowering and flushing coincided with those of sunshine hours and free sugar levels. The results, analysed in terms of the prevailing microclimatic conditions, suggest that light is a limiting factor for photsynthesis of shaded cocoa and that the reported stimulation of flowering and leaf flushing, which accompany shade removal, may be attributed to increased sugar status of the trees due to increased photosynthetic activity.     

Roosting Ecology of the Tent‐Roosting Bat Artibeus watsoni (Chiroptera: Phyllostomidae) in Southwestern Costa Rica1

We described the plants used as roost resources by Artibeus watsoni in southwestern Costa Rica, assessed roost fidelity, and compared roosting ecology between two sites, Golfito and Corcovado, which vary in the degree of human influence. A total of 349 tents from 25 different plant species were used by A. watsoni as roosts; some plant species (e.g., Carludovica palmata, Asplundia alata, Heliconia imbricata and Calathea lutea) were modified into tents with significantly higher frequency than others. The highest tents above the ground were observed in Philodendron popenoei and Rhodospatha wendlandii, whereas tents in Philodendron grandipes and A. alata were significantly lower than any other species. Asplundia alata and R. wendlandii also had the highest frequency of leaves modified per plant. Fidelity of bats to tents was low, although bats used several tents intermittently within a restricted area. Males generally were more faithful to tents than females, although not significantly so. This observation, along with indirect evidence of leaf modification, suggests that males are primarily responsible for tent construction. The two study sites differed in the plants used for roosting and in tent fidelity. Bats in Corcovado used a greater variety of plant species for tent roosting, whereas bats in Golfito were more faithful, suggesting that roosting resources were scarcer at the latter site.     

Impact of an introduced ant on native rain forest invertebrates: Pheidole megacephala in monsoonal Australia

Pheidole megacephala is an exotic ant species that has severely affected native invertebrate biodiversity throughout the tropics. Its impacts have been documented extensively in relatively depauperate invertebrate communities, but not in species-rich habitats such as tropical rain forests. Here we describe the local distribution of P. megacephala and its impacts on native invertebrate assemblages in and around a rain forest patch at Howard Springs, in Australia's monsoonal tropics. P. megacephala was found to be confined to a single area of approximately 25 ha, with its distribution centered on drainage lines and the rain forest. Significant but weak correlations were found between its abundance and vegetative canopy cover (positive) and distance from the rain forest (negative). In the most heavily infested area within the rain forest, the abundance of P. megacephala was 37–110 times that of total native ant abundance found within uninfested plots, as measured by pitfall traps. The abundance and richness of native ants and other invertebrates were significantly reduced in litter samples, pitfall catches and foliage beats where P. megacephala was present, inversely relative to the abundance of P. megacephala. Only two individuals of a single native ant species were found within the most infested plot, with native ant richness being reduced to about half in the least infested plot. The most persistent functional groups of native ants in infested plots were Cryptic species, which forage primarily within soil and leaf litter, and Opportunists, which exhibit highly generalised foraging behaviour. The highest abundance of P. megacephala corresponded with a 42–85% decrease in the abundance of other native invertebrates. Insect larvae were totally absent from foliage beats collected at the most heavily infested plot. P. megacephala was found overall to be expanding its range, averaging 12 m range expansion in the dry season and contracting 7 m in the wet season. It is able to spread into surrounding savanna habitats by occupying relatively sheltered microsites, such as beneath logs and at the bases of trees. However, it is unlikely to attain high population densities in open savanna habitats because of its relative intolerance of desiccation, and the prevalence of behaviourally dominant native ant species. Howard Springs is currently the only rain forest patch in monsoonal Australia known to be infested by P. megacephala, but clearly this ant is a serious potential threat to the region's rain forest invertebrate fauna. Received: 19 August 1998 / Accepted: 12 May 1999     

Tent Construction by the Short-nosed Fruit Bat Cynopterus sphinx (Chiroptera: Pteropodidae) in Southern India

The short-nosed fruit bat Cynopterus sphinx (Chiroptera: Pteropodidae) constructs shelters by severing stems of the curtain creeper, Vernonia scandens, and stems and leaves of the mast tree, Polyalthia longifolia, creating partially enclosed cavities (stem tents) in which to roost. Our observations indicate that the construction and maintenance of stem tents are primarily, if not exclusively, the behaviour of single males. A stem tent is formed in V. scandens when a single male C. sphinx severs up to 300 small- to medium-sized stems creating a partially flattened, bell-shaped cavity, and in P. longifolia when a male severs a few medium- to small-sized branches and many leaf petioles, creating an entry/exit portal and space in which to roost. A tent constructed in V. scandens is completed in approximately 30 d, whereas one in P. longifolia is completed in about 50 d. Stem-tent construction takes place mostly at night, but some stem chewing occurs in late afternoon. At night a stem tent is occupied by a single male, whereas females are usually absent. During the day the number of bats occupying completed tents is highly variable, ranging from two to 19 females (and their pups) and a single adult male. Tent construction is annually bimodal, which corresponds to a biannual breeding season. A dominant male sometimes deposits saliva on branches inside his tent cavity and actively defends this space from intrusions by other males. Both behaviours suggest forms of scent marking and territorial display. Our observations indicate that dominant males construct tents, recruit females and then defend the tents (and their female occupants) for the purpose of gaining reproductive access. The variance in harem group size indicates that some tent-making males are more successful than others in recruiting females.     

A gregarine parasite ofLeptothorax ants from North America

Summary A gregarine parasite (Neogregarinida, Ophryocystidae) ofLeptothorax ants was found near West-Yellowstone, Montana, USA. Infested pupae are characterized by grey to black coloration, irregular pigmentation of compound eyes, and toothless mandibles. The pathogenic effects closely resemble those in the fire ant,Solenopsis geminata, when infested byMattesia geminata. Infested pupae die. The infection can be transmitted to healthy colonies and to differentLeptothorax species by feeding them with infested pupae.     

An in vitro investigation of Malaysian Phytophthora palmivora isolates and pathogenicity study on oil palm

Bud rot disease affecting oil palm in South American countries is reported to be caused by Phytophthora palmivora. P. palmivora is a local pathogen affecting various crops in Malaysia, and this finding caused an alarm, which prompted an investigation of pathogenicity using Malaysian P. palmivora to assess the potentials of this Oomycete to infect oil palm in Malaysia. A total of 11 P. palmivora isolates were obtained from cocoa and durian for the study. Leaf bioassays via artificial inoculation using 50,000 zoospores/ml and mycelial agar disc showed severe necrotic lesions on the infection spot of oil palm (DxP) spear leaves. Mild infection was observed in oil palm spear leaves of OxG hybrids indicating lower susceptibility against P. palmivora infection. Phylogenetic analysis using internal transcribed spacer (ITS) data revealed that Malaysian isolates were genetically similar to Colombian isolates supported by significant bootstrap values. The leaf bioassay results revealed that Malaysian oil palm materials are susceptible towards local P. palmivora infection. The Colombian P. palmivora isolates causing bud rot incidence may have evolved over a long period of time, undergone sequential genetic shift to become more virulent towards Colombian oil palm planting materials.     

Contribution of cocoa plantations to the conservation of native ants (Insecta: Hymenoptera: Formicidae) with a special emphasis on the Atlantic Forest fauna of southern Bahia,Brazil

By maintaining a forest-like structure, shaded cocoa plantations contribute to the conservation of ants that usually live in the soil, leaf litter or canopy of tropical forests. Here we synthesize the available information on the diversity and community structure of ants in shaded cocoa plantations in the Atlantic forest region of Brazil, compare ant assemblages in cocoa agroforests with forests and other forms of agriculture, and discuss how these shaded plantations contribute to the conservation of the ants in the Atlantic Forest region. We also discuss ants of economical importance and of special interest, including Camponotus, Dolichoderus, Gnamptogenys, Pachycondyla, Pseudomyrmex and other litter dwelling genera. We discuss the situation of the tramp ant Wasmannia auropunctata in the Bahian cocoa-producing region where it is considered as native, and that of the two cryptobiotic genera Thaumatomyrmex and Typhlomyrmex, as well as that of proven and possible endangered army ant and Ponerini species. A total of 192 ant species from four strata were found in extensive sampling of a cocoa plantation with a relatively simple shade canopy (comprised primarily of Erythrina). Species richness in the cocoa plantations corresponded roughly to that of low diversity native forests, and species composition of cocoa plantations was most similar to native habitats (forest and mangroves) while ant composition in other agricultural habitats was most similar to that of urban areas. Although occurrences of Wasmannia auropunctata were similar in cocoa plantations and forests, abundance of Thaumatomyrmex and Typhlomyrmex, generally thought to be rare ants, was relatively high in cocoa plantations. These results, from cocoa plantations with relatively simple shade, demonstrate the importance of cocoa for ant conservation in the Atlantic forest region of Brazil. It is likely that cocoa plantations with a greater number of vegetation strata and higher tree species richness (such as traditional cabruca plantations) provide even more important habitat for ants generally and for ant species of conservation concern.     

Growth Enhancement,Amino Acid Synthesis and Reduction in Susceptibility Towards Phytophthora megakarya by Arbuscular Mycorrhizal Fungi Inoculation in Cocoa Plants

The effects of some selected arbuscular mycorrhizal (AM) fungi, Gigaspora margarita and Glomus mossae on the growth and the role of soluble amino acids of two contrasting cocoa cultivars (ICS84 tolerant and SNK10 sensitive) against black pod disease caused by Phytophthora megakarya were investigated. Root colonization by AM fungi is between 50 and 70% 18 weeks after planting. Tested AM fungi significantly increased all the plant growth parameters (height, number of leaves, shoot and root matter) and P uptake as compared to non‐inoculated plants in pot experiments. AM fungi inoculated cocoa reduced the disease severity. Compared to the control, the soluble amino acid levels increased with inoculation of the AM fungi strains in the necrotic stems of disease on inoculated cocoa plants. Significant relationships between amino acids and disease severity observed for two cocoa cultivars imply that the induction of specific amino acids synthesized by leaves, such as arginine, cysteine and glutamic acid, may represent potential candidate molecules for adaptation of such cultivars to P. megakarya disease. Inoculating seedlings with AMF in nurseries could enhance the development of cocoa plants protected against P. megakarya.     

Impact of an invasive alien ant,Wasmannia auropunctata Roger., on a specialised plant–ant mutualism,Barteria fistulosa Mast. and Tetraponera aethiops F. Smith., in a Gabon forest

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Geographic and taxonomic distribution of a positive interaction: ant-tended homopterans indirectly benefit figs across southern Africa

Although species pairs and assemblages often occur across geographic regions, ecologists know very little about the outcome of their interactions on such large spatial scales. Here, we assess the geographic distribution and taxonomic diversity of a positive interaction involving ant-tended homopterans and fig trees in the genus Ficus. Previous experimental studies at a few locations in South Africa indicated that Ficus sur indirectly benefited from the presence of a homopteran (Hilda patruelis) because it attracted ants (primarily Pheidole megacephala) that reduced the effects of both pre-dispersal ovule gallers and parasitoids of pollinating wasps. Based on this work, we evaluated three conditions that must be met in order to support the hypothesis that this indirect interaction involves many fig species and occurs throughout much of southern Africa and Madagascar. Data on 429 trees distributed among five countries indicated that 20 of 38 Ficus species, and 46% of all trees sampled, had ants on their figs. Members of the Sycomorus subgenus were significantly more likely to attract ants than those in the Urostigma subgenus, and ant-colonization levels on these species were significantly greater than for Urostigma species. On average, each ant-occupied F.sur tree had 37% of its fig crop colonized by ants, whereas the value was 24% for other Ficus species. H. patruelis was the most common source for attracting ants, although figs were also attacked by a range of other ant-tended homopterans. P. megacephala was significantly more common on figs than other ant species, being present on 58% of sampled trees. Ant densities commonly exceeded 4.5 per fig, which a field experiment indicated was sufficient to provide protection from ovule gallers and parasitoids of pollinators. Forty-nine percent of all colonized F. sur trees sampled had ant densities equal to or greater than 4.5 per fig, whereas this value was 23% for other Ficus species. We conclude that there is considerable evidence to suggest that this indirect interaction occurs across four southern African countries and Madagascar, and involves many Ficus species. Received: 11 December 1997 / Accepted: 6 April 1998     

The host range and vector species of viruses from Cola chlamydantha K. Schum., Adansonia digitata L. and Theobroma cacao L

Seventeen species in the Tiliaceae, Sterculiaceae and Bombacaceae were tested for susceptibility to three virus isolates from naturally infected Cola chlamydantha trees and three from Adansonia digitata trees in Ghana. Seven species of Pseudococcidae were tested as vectors of the six isolates. These studies indicate that the Cola isolates should be classified with cocoa swollen shoot virus and those from Adansonia with cocoa mottle leaf virus.