Biological control of Ziziphus mauritiana (Rhamnaceae): feasibility,prospective agents and research gaps

The tropical fruit tree, Ziziphus mauritiana (Rhamnaceae), a native of the Indian subcontinent, is a pasture and environmental weed in northern Australia and Fiji. In their native range, Ziziphus spp., including commercially cultivated Z. mauritiana and Z. jujuba, are subjected to a wide range of pests and diseases. The feasibility of classical biological control of this weed has not been explored to date. Effective biological control could reduce plant vigour and seed output, thereby limiting the spread of Z. mauritiana in Australia. Two Ziziphus species are native to Australia, hence, any prospective biological control agent should be specific to Z. mauritiana. Opportunistic field surveys and literature searches identified 133 species of phytophagous insects, 9 species of phytophagous mites and 12 plant pathogens on Ziziphus spp. Host records suggest the following are possibly specific to Z. mauritiana and hence are prospective biological control agents in Australia: the seed‐feeding weevil Aubeus himalayanus; the leaf‐feeding gracillariid moth Phyllonorycter iochrysis; the leaf‐mining chrysomelid beetle Platypria erinaceus; the leaf‐folding crambid moth Synclera univocalis; the leaf‐galling midge Phyllodiplosis jujubae; and the gall‐mites Aceria cernuus and Larvacarus transitans. Host range of the rust Phakopsora zizyphi‐vulgaris includes many Ziziphus species, including the native Z. oenoplia and hence would not be a suitable biological control agent in Australia. The powdery mildew Pseudoidium ziziphi, with a host range restricted to Ziziphus species, has not been reported on Z. oenoplia. All available information on the pests and diseases of Z. mauritiana are from cultivated varieties. Hence, future surveys should focus on wild Z. mauritiana in the Indian subcontinent in areas that are climatically similar to the regions of northern Australia, where it is currently most abundant.     

Life cycle and host range of Phycita sp. rejected for biological control of prickly acacia in Australia

Prickly acacia (Vachellia nilotica subsp. indica), a native of the Indian subcontinent, is a serious weed of the grazing areas of northern Australia and is a target for classical biological control. Native range surveys in India identified a leaf webber, Phycita sp. (Lepidoptera: Pyralidae) as a prospective biological control agent for prickly acacia. In this study, we report the life cycle and host‐specificity test results Phycita sp. and highlight the contradictory results between the no‐choice tests in India and Australia and the field host range in India. In no‐choice tests in India and Australia, Phycita sp. completed development on two of 11 and 16 of 27 non‐target test plant species, respectively. Although Phycita sp. fed and completed development on two non‐target test plant species (Vachellia planifrons and V. leucophloea) in no‐choice tests in India, there was no evidence of the insect on the two non‐target test plant species in the field. Our contention is that oviposition behaviour could be the key mechanism in host selection of Phycita sp., resulting in its incidence only on prickly acacia in India. This is supported by paired oviposition choice tests involving three test plant species (Acacia baileyana, A. mearnsii and A. deanei) in quarantine in Australia, where eggs were laid only on prickly acacia. However, in paired oviposition choice trials, only few eggs were laid, making the results unreliable. Although oviposition choice tests suggest that prickly acacia is the most preferred and natural host, difficulties in conducting choice oviposition tests with fully grown trees under quarantine conditions in Australia and the logistic difficulties of conducting open‐field tests with fully grown native Australian plants in India have led to rejection of Phycita sp. as a potential biological control agent for prickly acacia in Australia.     

Making host specificity testing more efficient: Exploring the use of abridged test plant lists

Testing the specificity of candidate agents is a key component of risk analysis in weed biological control. This step is often time-consuming due to the numerous plant species that need to be tested under quarantine conditions in the invaded country of the weed species. Here, we examined whether an abridged phylogenetically based test list could be used in the weed's native range to quickly screen the host specificity of candidate agents. Ten plant species were used to test the host specificity of a promising candidate for the biological control of Sonchus oleraceus in Australia, the gall midge, Cystiphora sonchi. No-choice and choice tests were carried out in the native Mediterranean range of the midge. The results showed the midge has potential to threaten native Australian species, as those species showed high infestation levels in no-choice tests and produced significantly higher numbers of galls in choice tests. As a result of this approach, C. sonchi was rapidly discarded from the list of agents to be imported into Australian quarantines for further tests. This study demonstrates that testing a few key phylogenetically related species in the native range may save cost and effort in a weed biological control programme.     

The host specificity of Anomalococcus indicus Ayyar (Hemiptera: Lecanodiaspididae), a potential biological control agent for prickly acacia (Vachellia nilotica ssp. indica) in Australia

ABSTRACT

Prickly acacia, Vachellia nilotica ssp. indica (Benth.) Kyal. & Boatwr, is a significant weed of northern Australia and has been a target of weed biological control in Australia since the 1980s. Following native range surveys in India, the scale insect Anomalococcus indicus Ayyar was identified as the most promising agent and was imported into Australia for further research. A. indicus is a major pest of prickly acacia on the Indian subcontinent, where it causes shoot tip dieback and plant death. Despite field observations suggesting the species was specific to V. nilotica, A. indicus completed development on 17 of the 84 non-target plant species tested during no-choice host specificity trials under quarantine conditions. Of these, Acacia falcata, V. bidwillii, V. sutherlandii and Neptunia major supported high numbers of mature females in all replicates. All of these species were utilised in choice trials. Combined risk scores indicate that V. sutherlandii, N. major and A. falcata may be attacked in the field. Due to the limited ability of scale insects to disperse, only those non-target species that occur on the Mitchell grass downs (i.e. V. sutherlandii) are considered to be at risk. Nevertheless, in view of the disparity between quarantine test results and the observed field host specificity of A. indicus in India, field trials are currently being conducted in India using Australian native species on which complete development has occurred. The future of A. indicus as a biological control agent for prickly acacia in Australia will be determined once results from these field trials are known.     

Density‐dependent reproduction and pollen limitation in an invasive milkweed,Calotropis procera (Ait.) R. Br. (Apocynaceae)

Calotropis procera (Ait.) R.Br. (Apocynaceae), an invasive woody milkweed, has expanded its range in northern Australia affecting rangeland and pastoral productivity. While self‐compatibility should enhance the species range expansion, spread of C. procera is limited by the availability of larger wasp and bee species that are able to vector its solid pollinia. Pollination efficiency is thus likely dependent on both pollinator abundance and plant density. Calotropis procera flowers year round in Australia but fruiting is limited to the warm months of the year when pollinators are most abundant, indicating that seasonal regulation of reproduction may be due to pollinator limitation. We examine the propositions that C. procera reproduction is regulated by the interaction between plant population density and pollinator pressure and that low pollinator pressure causes low per capita plant fecundity. All pollinators belonged to Order Hymenoptera and pollinator composition was similar at six of the seven sites. Fruit production per plant (fecundity) was lower above and below intermediate densities (350–550 plants ha^?1) of flowering plants with evidence of a weak Allee effect at lower plant density. Pollinator visitation rates per plant were low at high and low plant densities, and greatest at intermediate densities, while pollen supplementation experiments showed that C. procera is pollen limited (Pollen Limitation Index_fruit = 0.9) even at intermediate densities. Pollen limitation caused by low pollinator pressure at low plant densities and pollinator satiation at high plant densities may account for these fruit production trends. Management should be conducted in the colder months when pollinator pressure is low and plants are not reproducing. In addition, where stand eradication cannot be achieved in one attempt, management should reduce flowering plants to below intermediate densities where the fecundity per plant is low.     

Survey and prioritisation of potential biological control agents for prickly acacia (Acacia nilotica subsp. indica) in southern India

Prickly acacia, Acacia nilotica subsp. indica (Benth.) Brenan (Mimosaceae), a multi-purpose tree native to the Indian subcontinent, is a weed of national significance, widespread throughout the grazing areas of western Queensland and has the potential to spread throughout northern Australia. Biological control of prickly acacia has been in progress since the early 1980s, but with limited success to date. Based on genetic and climate matching studies, native surveys for potential biological control agents were conducted in 64 sites in Tamil Nadu state and eight sites in Karnataka state from November 2008 to December 2011. Surveys yielded 33 species of phytophagous insects (16 species of leaf-feeders, eight species of stem feeders, four species with leaf-feeding adults and root-feeding larvae, two stem-borers and bark-feeders and three flower-feeders) and two rust fungi. The number of species recorded at survey sites increased with the number of times the sites were surveyed. Using a scoring system based on field host range, geographic range, seasonal incidence and damage levels, we prioritised a scale insect (Anomalococcus indicus Ramakrishna Ayyar), two leaf-webbing caterpillars (Phycita sp. A and Phycita sp. B), a leaf weevil (Dereodus denticollis Boheman), a leaf beetle (Pachnephorus sp.), a gall-inducing rust (Ravenelia acacia-arabica Mundk. & Thirumalachari) and a leaf rust (Ravenelia evansii Syd. & P.) for detailed host specificity tests. The two rusts were sent to CABI-UK for preliminary host-specificity testing. Three insects (A. indicus, D. denticollis and Phycita sp. A) were imported into a quarantine facility in Brisbane, Australia where host-specificity tests are in progress.     

Influence of fruit traits on the infestation of Dacus persicus in two fruit morphs of Calotropis procera

The larvae of Dacus (Leptoxyda) persicus (aak fruit fly) are key predispersal seed predators in Calotropis procera (Asclepiadaceae). Based on fruit characteristics, two morphs are distinguishable in C. procera viz., the soft-fruited morph (SF morph) and the hard-fruited morph (HF morph). The work reported here examined whether the fruit characteristics influenced the infestation by the aak fruit fly and, if so, what mechanism(s) were operative. Fruits in the SF morph were significantly more acceptable to the aak fruit fly than those of the HF morph irrespective of their size class and availability or fly population density. A general ranking of fruit acceptability for oviposition by the aak fruit fly within the fruit size class was: size class III ≥ size class II > size class I and IV. The negative relationship between fruit infestation and pericarp toughness, which is suggestive of trade-offs between the fly’s oviposition obligation and energy/time (predation risk) constraint, was found to correlate with the requirement of greater force to puncture the pericarp in the hard fruits. Lower penetrability of the pericarp in the hard fruits appeared to be primarily due to the thickness of pericarp and secondarily on account of the thickened walls of endocarpic–mesocarpic cells in the inner pericarpic layer. The present data point to the existence of two fruit morphs in C. procera differing in the acceptability of fruits for oviposition by the aak fruit fly primarily on account of toughness and internal structure of the pericarp.     

Using molecular methods to determine the origin of weed populations of Pereskia aculeata in South Africa and its relevance to biological control

New biological control agents are required in order to reach and sustain an adequate level of control of the declared environmental weed Pereskia aculeata Miller (Cactaceae) in South Africa. Identifying the origin of weed genotypes has been important in a number of biological control programmes and is likely to be of importance for the control of P. aculeata due to its disjunct native distribution and morphological polymorphisms between plants from different regions of the native and introduced distribution. DNA sequencing of the trnL chloroplastic intron and the phyC nuclear gene indicate that the South African weed population’s origin was in the southern region of native distribution. Inter-Simple Sequence Repeats (ISSRs) confirmed this result and added resolution to the analysis indicating that the native plants with the closest genetic distance to the South African weed population were found in Rio de Janeiro Province, Brazil. The relationship between the South African weed population and garden variety plants as well as the large genetic distance between the South African plants and the native plants suggests that the South African population may be the progeny of escaped garden variety plants that have been cultivated and possibly hybridized. The low levels of genetic variation within the South African population and the monophyly of the South African plants indicates that these plants are the progeny of a single introduction or multiple introductions from the same source. Rio de Janeiro Province in Brazil is the most appropriate region in which to survey for new biological control agents.     

<Emphasis Type="Italic">Nimbya alternantherae</Emphasis> a potential biocontrol agent for alligatorweed, <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis>

Alligatorweed, (Alternanthera philoxeroides (Mart.) Griseb.), an aquatic and wetland plant native to South America, is an aggressive weed in many parts of the world. Its ability to compete with other native plants and to impede waterways has made it a serious threat to aquatic ecosystems. Although biological control with insects has been fairly successful in aquatic habitats, there is a need for additional agents to manage the weed in upland sites. Accordingly, in a survey in Brazil in 1997 a fungus, Nimbya (=Alternaria) alternantherae (Holcomb and Antonopoulus) Simmons and Alcorn, was discovered and confirmed to be highly damaging to alligatorweed. Studies were conducted to determine the potential of this fungus for controlling this weed. Several isolates from Brazil, USA, and Puerto Rico were compared and no differences in virulence were observed, although a lower dew requirement was demonstrated for the Brazilian isolates. Conidia were more effective than mycelial suspension, and inoculum concentrations of 1×10⁵ and 1×10⁶ conidia per ml provided significant levels of control of the weed in greenhouse and field experiments, respectively. In a host-range study, N. alternantherae infected 6 plant species from a total of 42 species belonging to 23 families. N. alternantherae has the potential to be an effective mycoherbicide for alligatorweed.     

Potential for ants and vertebrate predators to shape seed-dispersal dynamics of the invasive thistles <Emphasis Type="Italic">Cirsium arvense</Emphasis> and <Emphasis Type="Italic">Carduus nutans</Emphasis> in their introduced range (North America)

The population dynamics of invasive plants are influenced by positive and negative associations formed with members of the fauna present in the introduced range. For example, mutualistic associations formed with pollinators or seed dispersers may facilitate invasion, but reduced fitness from attack by native herbivores can also suppress it. Since population expansion depends on effective seed dispersal, interactions with seed dispersers and predators in a plant species introduced range may be of particular importance. We explored the relative contributions of potential seed dispersers (ants) and vertebrate predators (rodents and birds) to seed removal of two diplochorous (i.e., wind- and ant-dispersed), invasive thistles, Cirsium arvense and Carduus nutans, in Colorado, USA. We also conducted behavior trials to explore the potential of different ant species to disperse seeds, and we quantified which potential ant dispersers were prevalent at our study locations. Both ants and vertebrate predators removed significant amounts of C. arvense and C. nutans seed, with the relative proportion of seed removed by each guild varying by location. The behavior trials revealed clear seed preferences among three ant species as well as differences in the foragers’ abilities to move seeds. In addition, two ant species that acted as potential dispersal agents were dominant at the study locations. Since local conditions in part determined whether dispersers or predators removed more seed, it is possible that some thistle populations benefit from a net dispersal effect, while others suffer proportionally more predation. Additionally, because the effectiveness of potential ant dispersers is taxon-specific, changes in ant community composition could affect the seed-dispersal dynamics of these thistles. Until now, most studies describing dispersal dynamics in C. arvense and C. nutans have focused on primary dispersal by wind or pre-dispersal seed predation by insects. Our findings suggest that animal-mediated dispersal and post-dispersal seed predation deserve further consideration.     

Biological control of alligator weed: Unsuccessful attempts to control terrestrial growth using the flea beetleDisonycha argentinensis [Col.: Chrysomelidae]

Biological control of alligator weed growing in aquatic habitats in Australia is successful but the agents, a flea beetleAgasicles hygrophila and a mothVogtia malloi, do not control terrestrial growth. Consequently another flea beetleDisonycha argentinensis was introduced into Australia specifically to control the terrestrial growth. Progeny of adults collected in Brazil from areas similar in climate and habitat to areas infested with alligator weed in Australia, were released but failed to become established. Eggs were laid by females released into a large field cage and some completed development, but the new adults failed to reproduce. Tentative conclusions are that microclimate or predation may have prevented establishment ofD. argentinensis but the results should not preclude attempts to establish this insect in North America, China or elsewhere.      

Specificity and host specialization of Puccinia chondrillina

Various strains of Puccinia chondrillina, the Chondrilla rust, were collected during surveys in the Mediterranean, undertaken to discover biological control agents suitable for use in Australia against skeleton weed, Chondrilla juncea. Only the strain collected at Vieste (S.E. Italy) was highly virulent against the common Australian form of C. juncea. The specificity of this strain of the rust was tested by inoculation of many unrelated cultivated plants, related cultivated plants and closely related wild plants; all were immune to the rust. These tests satisfied the Australian plant quarantine authorities and this strain of the rust has now been introduced into Australia as a biological control agent for C. juncea.     

Role of molecular genetics in identifying ‘fine tuned’ natural enemies of the invasive Brazilian peppertree, <Emphasis Type="Italic">Schinus terebinthifolius</Emphasis>: a review

Brazilian peppertree, Schinus terebinthifolius Raddi (Sapindales: Anacardiaceae), is a highly successful invasive species in the continental United States, Hawaiian archipelago, several Caribbean Islands, Australia, Bermuda, and a number of other countries worldwide. It also is one of only a few invasive intraspecific hybrids that has been well characterized genetically. The natural enemy complex of Brazilian peppertree includes two thrips and two psyllids that appear to be highly adapted to specific haplotypes or their hybrids. Successful biological control of Brazilian peppertree will require careful matching of the appropriate natural enemies with their host plant genotypes. The Brazilian peppertree model reviewed here could provide a useful framework for studying biological control agents on other invasive weed species that have exhibited intraspecific hybridization.     

Prioritisation of potential agents for the biological control of the invasive alien weed,Pereskia aculeata (Cactaceae), in South Africa

Pereskia aculeata Miller (Cactaceae) is an invasive alien species in South Africa that is native in Central and South America. In South Africa, P. aculeata outcompetes native plant species leading to a reduction in biodiversity at infested sites. Herbicidal and mechanical control of the plant is ineffective and unsustainable, so biological control is considered the only potential solution. Climatic matching and genotype matching indicated that the most appropriate regions in which to collect biological control agents were Santa Catarina and Rio de Janeiro provinces in Southern Brazil. Surveys throughout the native distribution resulted in 15 natural enemy species that were associated with the plant. Field host range data, as well as previous host plant records, were used to prioritise which of the species were most likely to be suitably host specific for release in South Africa. The mode of damage was used to determine which species were most likely to be damaging and effective if released. The most promising species prioritised for further study, including host specificity and impact studies, were the stem-wilter Catorhintha schaffneri Brailovsky & Garcia (Coreidae); the stem boring species Acanthodoxus machacalis Martins & Monné (Cerambycidae), Cryptorhynchus sp. (Curculionidae) and Maracayia chlorisalis (Walker) (Crambidae) and the fruit galler Asphondylia sp. (Cecidomyiidae). By prioritising the potential biological control agents that are most likely to be host-specific and damaging, the risk of conducting host specificity testing on unsuitable or ineffective biological control agents is reduced.     

Ecological niche modelling of an invasive alien plant and its potential biological control agents

Invasive alien plants are of concern in South Africa. Pompom weed (Campuloclinium macrocephalum) is currently invading the Grassland and Savannah biomes of South Africa and is likely to continue spreading in the southern African sub- region. Two possible biological control agents (Liothrips tractabilis and Cochylis campuloclinium) have been identified for control of pompom weed. We used ecological niche modelling to predict which areas in southern Africa are likely to be suitable for pompom weed and the two potential biological control agents. The overlap between areas predicted to be highly suitable for pompom weed and areas suitable for the biological control agents was assessed. Methods of reducing sampling bias in a data set used for calibrating models were also compared. Finally, the performance of models calibrated using only native range data, only invaded range data and both were also compared. Models indicate that pompom weed is likely to spread across a greater region of southern Africa than it currently occupies, with the Savannah and Grassland biomes being at greatest risk of invasion. Poor overlap was found between the areas predicted to be highly suitable for pompom weed and those areas predicted to be suitable for the biological control agents. However, models of the potential distribution of the biological control agents are interpreted with caution due to the very small sample size of the data set used to calibrate the models. Models calibrated using both native range and invaded range data were found to perform best whilst models calibrated using only native range data performed the worst. There was little difference found between models that were calibrated using spatially reduced (selecting only one record per 30 min grid cell) and randomly reduced (randomly selecting 50% of available records) biased data sets.     

Impact of habitat modification on the distribution and abundance of fruit flies (Diptera: Tephritidae) in Southeast Queensland

Loss of rainforest because of agricultural and urban development may impact the abundance and diversity of species that are rainforest natives. Tropical fruit flies are one group of such organisms indigenous to rainforests. In southeast Queensland, a region subject to rapid urbanization, we assessed the impact of habitat disturbance on the distribution and abundance of native fruit flies. Data on four species (Bactrocera tryoni, Bactrocera neohumeralis, Bactrocera chorista, and Dacus aequalis) were gathered and analyzed over 6 months in three habitat types: suburbia, open sclerophyll forest, and rainforest. We also analyzed the data at a combined "dacine fruit fly" level incorporating all fruit fly species trapped over the period of study (as might occur in a biodiversity assessment): these included the four species already named and Bactrocera melas, Bactrocera bryoniae, Bactrocera newmani, and Dacus absonifacies. Analysis at the species level showed that the polyphagous pest species responded differently to the monophagous species. Bactrocera tryoni, which has more exotic than native hosts, was positively affected by transformation of natural habitat into suburbia whereas B. neohumeralis, which has nearly identical numbers of native and exotic hosts, was found equally across habitat types. Bactrocera chorista and Dacus aequalis, each monophagous on a species-specific rainforest host plant, were most abundant in rainforest. The analysis based on the combined data suggests that replacing rainforest with suburbia has a neutral, or even positive, effect on the abundance of fruit flies as a whole. At the species level, however, it can be seen that this is an erroneous conclusion biased by the abundance of a single pest species. Our discussion raises the issue of analyses at supraspecific levels in biodiversity and impact assessment studies. Received: March 6, 2000 / Accepted: June 19, 2000     

Fruit production in Sorbus aucuparia L. (Rosaceae) and pre-dispersal seed predation by the apple fruit moth (Argyresthia conjugella Zell.)

Variation in fruit production and pre-dispersal seed predation by Argyresthia conjugella was studied in␣four populations of Sorbus aucuparia in northern Sweden.␣The number of infructescences, fruits per infructescence, consumed seeds and developed unattacked seeds per fruit were scored in marked trees from 1984 to 1990. The results showed that the number of fruits produced in each population determined the number of seed predators occurring in the host population, as the yearly number of seed predators was significantly and positively correlated with yearly number of fruits, in all but one population. The seed predators showed a delay in response to variation in number of fruits produced. This lag in response resulted in a large proportion of fruits being attacked and seeds consumed in a bad fruiting year that followed a good fruiting year, and vice versa. The proportion of fruits attacked and seeds consumed was largest in the population showing the greatest between-year variation in fruit production and lowest in the population showing the lowest between-year variation in fruit production. Furthermore, the individuals within the former population were synchronised, while they were not in the latter population. These results contradict one of the possible explanations of mast-seeding, where large synchronised between-year variation is supposed to reduce pre-dispersal seed predation. Instead, differences in attraction of the seed predator to differences in fruit crop size could explain the observed difference in seed predation between the two populations with opposite fruiting patterns. Within each population, irrespective of year, the proportion of fruits attacked and seeds consumed was independent of a tree's fruiting display. Therefore, trees with high fruit production, despite harbouring the largest number of seed predators, produced the largest number of developed seeds in absolute numbers, compared to trees that produced few fruits. Received: 25 February 1996 / Accepted: 30 November 1996     

Fire damage on seeds of Calliandra parviflora Benth. (Fabaceae), a facultative seeder in a Brazilian flooding savanna

Optimal sexual reproduction in relation to fire effects varies in Fabaceae species. Calliandra species have a large investment in reproduction. We investigated the consequences of fire during the fruiting period of Calliandra parviflora Benth., by checking fruit exposure to fire, pre-dispersal seed predator infestation, and the effect of fruit burning on germination. We conducted this study in a floodable savanna in central Brazil, where we collected burnt and unburnt fruits. We measured the fruit and seed mass, and counted the number of damaged and undamaged seeds and live larvae per fruit. We analyzed the seed germination percentage from burnt and unburnt fruits. The burnt fruits presented greater mass than the unburnt fruits, despite their seed mass being similar. The number of damaged seeds per fruit was only slightly higher in burnt compared to unburnt fruits (p = 0.047). The number of larvae on pre-dispersal seeds per fruit varied from 0 to 4 and did not differ between burnt and unburnt fruits. The germination percentage of unburnt fruit seeds (mean = 22 ± 17%), was significantly higher than that of burnt fruit (mean = 3.0 ± 2.0%, p < 0.001). Fire during fruiting or pre-dispersion decreases seed germination from 22 to 3%, but it does not hurt vegetative regeneration or resprout capacity of C. parviflora, which is a facultative seeder. Hence, we suggest that C. parviflora has potential for post-fire restoration in floodable open grassy savannas, in the ecotone between Cerrado and Pantanal, because this species may sprout quickly after first post-fire rains.     

Host-specificity of Uromyces heliotropii, a fungal agent for the biological control of common heliotrope (Heliotropium europaeum) in Australia

Common heliotrope, Heliotropium europaeum (Boraginaceae), a summer annual, is a serious weed of pastures in Australia causing poisoning in sheep and other grazing animals. The weed is native to Mediterranean and Middle Eastern regions of Eurasia where it is attacked by several arthropods and pathogens including the rust fungus Uromyces heliotropii. The host-specificity of a virulent and effective strain of the rust from Turkey has been tested by inoculating 96 plants of importance to the Australasian region using both microscopic and macroscopic observations of the reaction of host and non-host plants. The test plants included several members of the Boraginaceae and related families as well as species of Heliotropium native to Australia. The infection was mostly limited to European species of Heliotropium. U. heliotropii was thus found to represent a safe introduction and has recently been introduced to Australia for the biological control of common heliotrope.     

Predicting the host range of Nystalea ebalea: Secondary plant chemistry and host selection by a surrogate biological control agent of Schinus terebinthifolia

The safety of weed biological control depends upon the selection and utilization of the target weed by the agent while causing minimal harm to non-target species. Selection of weed species by biological control agents is determined by the presence of behavioral cues, generally host secondary plant compounds that elicit oviposition and feeding responses. Non-target species that possess the same behavioral cues as found in the target weed may be at risk of damage by classical biological control agents. Here we conducted host range tests and examined secondary plant compounds of several test plant species. We studied the specialist herbivore Nystalea ebalea (Lepidoptera: Notodontidae) a Neotropical species, present in Florida as a surrogate biological control agent of the weed, Brazilian peppertree Schinus terebinthifolia, invasive in Florida and Hawaii. We found that the larvae had the greatest survival when fed the target weed, the Neotropical species Spondias purpurea, the Florida native species Rhus copallinum, and the ornamental Pistacia chinensis. Reduced survival and general larval performance were found on the native species Metopium toxiferum and Toxicodendron radicans. Both the volatiles and the allergen urushiols were chemically characterized for all species but urushiol diversity and concentration best predicted host range of this herbivore species. These results provide insight into host selection and utilization by one oligophagous Schinus herbivore. Other potential biological control agents may also be sensitive to plants that contain urushiols and if so, they may pose minimal risk to these native species.