Plant hosts of the phytoplasmas and rickettsia-like-organisms associated with strawberry lethal yellows and green petal diseases

Candidatus Phytoplasma australiense (Ca. P. australiense) is associated with the plant diseases strawberry lethal yellows (SLY), strawberry green petal (SGP), papaya dieback (PDB), Australian grapevine yellows (AGY) and Phormium yellow leaf (PYL; New Zealand). Strawberry lethal yellows disease is also associated with a rickettsia-like-organism (RLO) or infrequently with the tomato big bud (TBB) phytoplasma, the latter being associated with a wide range of plant diseases throughout Australia. In contrast, the RLO has been identified only in association with SLY disease, and Ca. P. australiense has been detected only in a limited number of plant host species. The aim of this study was to identify plant hosts that are possible reservoirs of Ca. P. australiense and the SLY RLO. Thirty-one plant species from south-east Queensland were observed with disease between 2001 and 2003 and, of these, 18 species tested positive using phytoplasma-specific primers. The RLO was detected in diseased Jacksonia scoparia and Modiola caroliniana samples collected at Stanthorpe. The TBB phytoplasma was detected in 16 different plant species and Ca. P. australiense Australian grapevine yellows strain was detected in six species. The TBB phytoplasma was detected in plants collected at Nambour, Stanthorpe, Warwick and Brisbane. Ca. P. australiense was detected in plants collected at Nambour, Stanthorpe, Gatton and Allora. All four phytoplasmas were detected in diseased Gomphocarpus physocarpus plants collected at Toowoomba, Allora, Nambour and Gatton. These results indicated that the vector(s) of Ca. P. australiense are distributed throughout south-east Queensland and the diversity of phytoplasmas detected in G. physocarpus suggests it is a feeding source for phytoplasma insect vectors or it has a broad susceptibility to a range of phytoplasmas.

Agonosoma trilineatum (Heteroptera: Scutelleridae) a biological control agent of the weed bellyache bush, Jatropha gossypiifolia (Euphorbiaceae)

Bellyache bush, Jatropha gossypiifolia L., is a serious weed of northern Australia. Agonosoma trilineatum (F.) is an insect from tropical America released in Australia in 2003 as a biological control agent against bellyache bush. It feeds on seeds and has the potential to reduce seed production, thereby potentially reducing the rate of spread and recruitment. To test the host specificity of A. trilineatum, four biological responses to host plant species were determined: development of nymphs, oviposition preferences, adult feeding and frequency of mating. Development of nymphs to adults and adult feeding only occurred on three Jatropha spp. These species also supported mating and oogenesis but only J. gossypiifolia was accepted for oviposition. Mating did not occur in the presence of other plant species. The evidence indicates that there is little risk associated with the release of this insect species in Australia and probably other countries where this weed is a problem. The probability of this insect expanding its host range is low because multiple aspects of the biology would need to change simultaneously. A. trilineatum was released in Australia between 2003 and 2007. A Climex model indicated that coastal areas of Queensland and the Northern Territory would be climatically most suitable for this insect.

Will biological control of Lantana camara ever succeed? Patterns, processes & prospects

Despite biocontrol research spanning over 100 years, the hybrid weed, commonly referred to as Lantana camara, is not under adequate control. Host specificity and varietal preference of released agents, climatic suitability of a region for released agents, number of agents introduced and range or area of infestation appear to play a role in limiting biocontrol success. At least one of 41 species of mainly leaf- or flower-feeding insects has been introduced, or spread, to 41 of the 70 countries or regions where lantana occurs. Over half (26) of these species have established, achieving varying levels of herbivory and presumably some degree of control. Accurate taxonomy of the plant and adaptation of potential agents to the host plant are some of the better predictors of at least establishment success. Retrospective analysis of the hosts of introduced biocontrol agents for L. camara show that a greater proportion of agents that were collected from L. camara or Lantana urticifolia established, than agents that were collected from other species of Lantana. Of the introduced agents that had established and were oligophagous, 18 out of 22 established. The proportion of species establishing, declined with the number of species introduced. However, there was no trend when oceanic islands were treated separately from mainland areas and the result is likely an artefact of how introductions have changed over time. A calculated index of the degree of herbivory due to agents known to have caused some damage per country, was not related to land area infested with lantana for mainlands nor for oceanic islands. However, the degree of herbivory is much higher on islands than mainlands. This difference between island and mainland situations may reflect population dynamics in patchy or metapopulation landscapes. Basic systematic studies of the host remain crucial to successful biocontrol, especially of hybrid weeds like L. camara. Potential biocontrol agents should be monophages collected from the most closely related species to the target weed or be phytophages that attack several species of lantana. Suitable agents should be released in the most ideal ecoclimatic area. Since collection of biocontrol agents has been limited to a fraction of the known number of phytophagous species available, biocontrol may be improved by targeting insects that feed on stems and roots, as well as the agents that feed on leaves and flowers.

Biology and host range of Ophiomyia camarae Spencer (Diptera: Agromyzidae), a potential biocontrol agent for Lantana spp. (Verbenaceae) in Australia

The life history and host range of the herringbone leaf-mining fly Ophiomyia camarae, a potential biological control agent for Lantana spp., were investigated. Eggs were deposited singly on the underside of leaves. Although several eggs can be laid on a single leaf and a maximum of three individual mines were seen on a single leaf, only one pupa per leaf ever developed. The generation time (egg to adult) was about 38 days. Females (mean 14 days) lived longer than males (mean 9 days) and produced about 61 mines. Oviposition and larval development occurred on all five lantana phenotypes tested. Eleven plant species representing six families were tested to determine the host range. Oviposition and larval development occurred on only lantana and another nonnative plant Lippia alba (Verbenaceae), with both species supporting populations over several generations. A CLIMEX model showed that most of the coastal areas of eastern Australia south to 30°16' S (Coffs Harbour) would be suitable for O. camarae. O. camarae was approved for release in Australia in October 2007 and mines have been observed on plants at numerous field sites along the coast following releases.

Predicting risk and benefit a priori in biological control of invasive plant species: a systems modelling approach

While the method using specialist herbivores in managing invasive plants (classical biological control) is regarded as relatively safe and cost-effective in comparison to other methods of management, the rarity of strict monophagy among insect herbivores illustrates that, like any management option, biological control is not risk-free. The challenge for classical biological control is therefore to predict risks and benefits a priori. In this study we develop a simulation model that may aid in this process. We use this model to predict the risks and benefits of introducing the chrysomelid beetle Charidotis auroguttata to manage the invasive liana Macfadyena unguis-cati in Australia. Preliminary host-specificity testing of this herbivore indicated that there was limited feeding on a non-target plant, although the non-target was only able to sustain some transitions of the life cycle of the herbivore. The model includes herbivore, target and non-target life history and incorporates spillover dynamics of populations of this herbivore from the target to the non-target under a variety of scenarios. Data from studies of this herbivore in the native range and under quarantine were used to parameterize the model and predict the relative risks and benefits of this herbivore when the target and non-target plants co-occur. Key model outputs include population dynamics on target (apparent benefit) and non-target (apparent risk) and fitness consequences to the target (actual benefit) and non-target plant (actual risk) of herbivore damage. The model predicted that risk to the non-target became unacceptable (i.e. significant negative effects on fitness) when the ratio of target to non-target in a given patch ranged from 1:1 to 3:2. By comparing the current known distribution of the non-target and the predicted distribution of the target we were able to identify regions in Australia where the agent may be pose an unacceptable risk. By considering risk and benefit simultaneously, we highlight how such a simulation modelling approach can assist scientists and regulators in making more objective decisions a priori, on the value of releasing specialist herbivores as biological control agents.

Specificity of Carvalhotingis visenda (Hemiptera: Tingidae) as a biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae) in Australia

Biological control is considered the most suitable management option for cat's claw creeper, Macfadyena unguis-cati, a major environmental weed in coastal and sub-coastal areas of Queensland and New South Wales, Australia. The potential host range of the leaf-sucking bug, Carvalhotingis visenda (Hemiptera: Tingidae) was evaluated on the basis of nymphal survival and development, adult feeding and survival, and oviposition preference using choice and no-choice tests involving 38 plant species in 10 families. In no-choice tests, although adults survived on a few of the non-target plants, no eggs were laid on any of the non-target plants. In no-choice condition, the tingid oviposits and completes nymphal development only on M. unguis-cati. There was also no visible feeding damage on any of the non-target plants. In choice tests, adults showed distinct preference for M. unguis-cati, and the preference level increased over time as the tingids moved away from the non-target plants. At the end of the trial no adults were evident on any of the non-target plants. Host specificity tests confirm that the tingid is a highly host specific biocontrol agent, and does not pose risk to any non-target plants in Australia. This agent has been approved for field release by the relevant regulatory authorities in Australia.

The host range of Isturgia deeraria, an insect considered for the biological control of Acacia nilotica in Australia

The geometrid caterpillar Isturgia deerraria was imported from Kenya into quarantine facilities in Australia as a potential biological control agent for prickly acacia, Acacia nilotica subsp. indica (Benth.) Brenan (family Mimosaceae). The insect was then tested on 30 plant species presented to neonate larvae as a no-choice cut foliage test and 13 plant species presented as a no-choice potted plant test. In these tests the insect was able to complete its life cycle on 13 native Acacia spp. and also on Acacia farnesiana and the exotic ornamental Delonix regia (family Caesalpiniaceae). The tests supported field observations that the insect has a host range spanning many leguminous species and as such the insect could not be considered for release in Australia.

Studies on the rust Prospodium tuberculatum, a new classical biological control agent released against the invasive alien weed Lantana camara in Australia. 2. Host range

A strain of the rust Prospodium tuberculatum from Brazil was screened as a potential biocontrol agent against 40 Australian Lantana camara forms and 52 closely related, non-target plant species. Results under glasshouse conditions showed that the Brazilian rust strain is pathogenic to only two flower colour forms: pink and pink-edged red. Macro- and microsymptoms were recorded using 11 assessment categories and four susceptibility ratings. No macrosymptoms were observed on any of the non-target plants.

How widespread is Parthenium hysterophorus and its biological control agent Zygogramma bicolorata in South Asia?

Parthenium hysterophorus is a weed of global significance causing severe economic, environmental, human and animal health problems in Asia, Africa, Australia and the Pacific. In South Asia, P. hysterophorus occurs in India, Pakistan, Sri Lanka, Bangladesh and Nepal. A host-specific leaf-feeding beetle Zygogramma bicolorata from Mexico was introduced into India in 1984, as a biological control agent for P. hysterophorus. In this study, a GIS-based distribution map of P. hysterophorus and its biological control agent Z. bicolorata in South Asia based on meta-analysis is presented. The map highlights the limited published information on P. hysterophorus incidence in many of the states and territories in India, as well as in neighbouring Bangladesh, Bhutan, Nepal and Pakistan. Incidence of Z. bicolorata was recorded as three distinct clusters, covering many states in India. In Pakistan, Z. bicolorata was recorded in the Punjab region bordering India. A CLIMEX model based on the current distribution of Z. bicolorata in India suggests that the geographic range of this agent in India and Pakistan can extend to other P. hysterophorus-infested areas in the region. The CLIMEX model also suggests that all of Bangladesh and Sri Lanka, and parts of Nepal are climatically suitable for Z. bicolorata.

The infectivity and host range of Orgyia anartoides nucleopolyhedrovirus.

The painted apple moth (PAM), Teia anartoides (Walker) (Lepidoptera: Lymantriidae) made a recent incursion into New Zealand. A nucleopolyhedrovirus (NPV), Orgyia anartoides NPV (OranNPV), originally isolated from PAM in Australia, was tested for its pathogenicity to PAM and a range of non-target insect species found in New Zealand, to evaluate its suitability as a microbial control for this insect invader. Dosage-mortality tests showed that OranNPV was highly pathogenic to PAM larvae; mean LT50 values for third instars ranged from 17.9 to 8.1 days for doses from 102 to 105 polyhedral inclusion bodies/larva, respectively. The cause of death in infected insects was confirmed as OranNPV. Molecular analysis established that OranNPV can be identified by PCR and restriction digestion, and this process complemented microscopic examination of infected larvae. No lymantriid species occur in New Zealand; however, the virus had no significant effects on species from five other lepidopteran families (Noctuidae, Tortricidae, Geometridae, Nymphalidae and Plutellidae) or on adult honeybees. Thus, all indications from this initial investigation are that OranNPV would be an important tool in the control of PAM in a future incursion of this species into New Zealand.

Austropleospora osteospermi gen. et sp nov and its host specificity and distribution on Chrysanthemoides monilifera ssp rotundata in Australia.

Hendersonia osteospermi was found for the first time in Australia on leaf spots of the introduced invasive plant Chrysanthemoides monilifera ssp. rotundata (bitou bush) in coastal regions of New South Wales. Pathogenicity tests on species from 11 tribes in the family Asteraceae, demonstrated that H. osteospermi caused severe necrosis on leaves and stems of C. monilifera ssp. rotundata and its congener C. monilifera ssp. monilifera (boneseed). Small necrotic spots also developed on Osteospermum fruticosum and Dimorphotheca cuneata in the Calenduleae and on Helianthus annuus (sunflower) in the Heliantheae. None of the other plant species tested developed leaf spots, although H. osteospermi was re-isolated from senescent leaves of Cynara scolymus (globe artichoke) in the Cynareae and Vernonia cinerea in the Vernonieae. Single ascospores from ascomata of a Pleospora-like fungus found on diseased stems of bitou bush produced H. osteospermi in culture, which proved the anamorph/teleomorph connection. The ITS region of both a single-ascospore isolate and a single-conidium isolate were sequenced and found to be identical. The taxonomic status of H. osteospermi is re-examined and Austropleospora osteospermi gen. et sp. nov. is described as its teleomorph based on morphology, host range tests and DNA sequence analysis. The potential of A. osteospermi for the biological control of bitou bush and boneseed in Australia is discussed.

Species and Hybrids: Chemical and Physical Foliar Attributes and Implications for Herbivory.

Hybridization is an important biological phenomenon that can be used to understand the evolutionary process of speciation of plants and their associated pests and diseases. Interactions between hybrid plants and the herbivores of the parental taxa may be used to elucidate the various cues being used by the pests for host location or other processes. The chemical composition of plants, and their physical foliar attributes, including leaf thickness, trichome density, moisture content and specific leaf weight were compared between allopatric pure and commercial hybrid species of Corymbia, an important subtropical hardwood taxon. The leaf-eating beetle Paropsis atomaria, to which the pure taxa represented host (C. citriodora subsp. variegata) and non-host (C. torelliana) plants, was used to examine patterns of herbivory in relation to these traits. Hybrid physical foliar traits, chemical profiles, and field and laboratory beetle feeding preference, while showing some variability, were generally intermediate to those exhibited by parent taxa, thus suggesting an additive inheritance pattern. The hybrid susceptibility hypothesis was not supported by our field or laboratory studies, and there was no strong relationship between adult preference and larval performance. The most-preferred adult host was the sympatric taxon, although this species supported the lowest larval survival, while the hybrid produced significantly smaller pupae than the pure species. The results are discussed in relation to plant chemistry and physical characteristics. The findings suggest a chemical basis for host selection behavior and indicate that it may be possible to select for resistance to this insect pest in these commercially important hardwood trees.

Characterising insect plant host relationships facilitates understanding multiple host use

Weed biocontrol relies on host specificity testing, usually carried out under quarantine conditions to predict the future host range of candidate control agents. The predictive power of host testing can be scrutinised directly with Aconophora compressa, previously released against the weed Lantana camara L. (lantana) because its ecology in its new range (Australia) is known and includes the unanticipated use of several host species. Glasshouse based predictions of field host use from experiments designed a posteriori can therefore be compared against known field host use. Adult survival, reproductive output and egg maturation were quantified. Adult survival did not differ statistically across the four verbenaceous hosts used in Australia. Oviposition was significantly highest on fiddlewood (Citharexylum spinosum L.), followed by lantana, on which oviposition was significantly higher than on two varieties of Duranta erecta (‘‘geisha girl’’ and ‘‘Sheena’s gold’’; all Verbenaceae). Oviposition rates across Duranta varieties were not significantly different from each other but were significantly higher than on the two non-verbenaceous hosts (Jacaranda mimosifolia D. Don: Bignoneaceae (jacaranda) and Myoporum acuminatum R. Br.: Myoporaceae (Myoporum)). Production of adult A. compressa was modelled across the hosts tested. The only major discrepancy between model output and their relative abundance across hosts in the field was that densities on lantana in the field were much lower than predicted by the model. The adults may, therefore, not locate lantana under field conditions and/or adults may find lantana but leave after laying relatively few eggs. Fiddlewood is the only primary host plant of A. compressa in Australia, whereas lantana and the others are used secondarily or incidentally. The distinction between primary, secondary and incidental hosts of a herbivore species helps to predict the intensity and regularity of host use by that herbivore. Populations of the primary host plants of a released biological control agent are most likely to be consistently impacted by the herbivore, whereas secondary and incidental host plant species are unlikely to be impacted consistently. As a consequence, potential biocontrol agents should be released only against hosts to which they have been shown to be primarily adapted.

Quambalaria species: increasing threat to eucalypt plantations in Australia

Spotted gum (Corymbia citriodora subsp. variegata and C. maculata) is a valuable source of commercial timber and suitable for a wide range of different soil types in eastern Australia. The main biological constraint to further expansion of spotted gum plantations is Quambalaria shoot blight caused by the fungus Quambalaria pitereka. Surveys conducted to evaluate the impact of Quambalaria shoot blight have shown that the disease is present in all spotted gum plantations and on a range of Corymbia species and hybrids in subtropical and tropical regions surveyed in eastern Australia. More recently, Q. eucalypti has also been identified from a range of Eucalyptus species in these regions. Both pathogens have also been found associated with foliage blight and die-back of amenity trees and Q. pitereka in native stands of Corymbia species, which is the probable initial infection source for plantations. Infection by Q. pitereka commonly results in the repeated destruction of the growing tips and the subsequent formation of a bushy crown or death of trees in severe cases. In comparison, Q. eucalypti causes small, limited lesions and has in some cases been associated with insect feeding. It has not been recorded as causing severe shoot and stem blight. A better understanding of factors influencing disease development and host-pathogen interactions is essential in the development of a disease management strategy for these poorly understood but important pathogens in the rapidly expanding eucalypt (Corymbia and Eucalyptus spp.) plantation industry in subtropical and tropical eastern Australia.