Predation risk assessment by olfactory and visual cues in a coral reef fish

Assessment of predation risk is vital for the success of an individual. Primary cues for the assessment include visual and olfactory stimuli, but the relative importance of these sources of information for risk assessment has seldom been assessed for marine fishes. This study examined the importance of visual and chemical cues in assessing risk for the star goby, Asterropteryx semipunctatus. Visual and chemical cue intensities were used that were indicative of a high threat situation. The behavioural response elicited by both the visual cues of a predator (the rock cod, Cephalopholis boenak) and the chemical alarm cues from conspecifics were similar in magnitude, with responses including a decrease in feeding strikes and moves. A bobbing behaviour was exhibited when the predator was visible and not when only exposed to the chemical alarm cue. When visual and chemical cues were presented together they yielded a stronger antipredator response than when gobies were exposed solely to conspecific alarm cues. This suggests additivity of risk assessment information at the levels of threat used, however, the goby’s response is also likely to depend on the environmental and social context of the predator–prey encounter. This study highlights the importance of chemical cues in the assessment of predation risk for a coral reef fish.     

Predictable risk to native plants in weed biological control

Data on field host use of 112 insects, 3 fungi, 1 mite, and 1 nematode established for biological control of weeds in Hawaii, the continental United States, and the Caribbean indicate that the risk to native flora can be judged reliably before introduction. Virtually all risk is borne by native plant species that are closely related to target weeds. Fifteen species of insects introduced for biological control use 41 native plant species; 36 of which are congeneric with target weeds, while 4 others belong to two closely allied genera. Only 1 of 117 established biological organisms uses a native plant unrelated to the target weed. Thus the elements of protection for the native flora are the selection of weed targets that have few or no native congeners and the introduction of biological control organisms with suitably narrow diets.     

Parasite and predator risk assessment: nuanced use of olfactory cues

Foraging herbivores face twin threats of predation and parasite infection, but the risk of predation has received much more attention. We evaluated, experimentally, the role of olfactory cues in predator and parasite risk assessment on the foraging behaviour of a population of marked, free-ranging, red-necked wallabies (Macropus rufogriseus). The wallabies adjusted their behaviour according to these olfactory cues. They foraged less, were more vigilant and spent less time at feeders placed in the vicinity of faeces from dogs that had consumed wallaby or kangaroo meat compared with that of dogs feeding on sheep, rabbit or possum meat. Wallabies also showed a species-specific faecal aversion by consuming less food from feeders contaminated with wallaby faeces compared with sympatric kangaroo faeces, whose gastrointestinal parasite fauna differs from that of the wallabies. Combining both parasite and predation cues in a single field experiment revealed that these risks had an additive effect, rather than the wallabies compromising their response to one risk at the expense of the other.     

A two-phase open-field test to evaluate the host-specificity of candidate biological control agents for Heliotropium amplexicaule

This paper describes an open-field host-specificity test conducted to make a preliminary evaluation of the safety of four candidate agents for the biological control of Heliotropium amplexicaule, an invasive weed of South American origin. These agents were a leaf-eating beetle, Deuterocampta quadrijuga, a flea-beetle, Longitarsus sp., that feeds on leaves as an adult and roots as a larva, a shoot-feeding thrips, Haplothrips heliotropica, and a cell-sucking bug, Dictyla sp. During the first phase of the experiment, the four agents were given a choice between the target weed and six species of nontarget plant of varying degrees of phylogenetic relatedness. All four species were found to feed and reproduce on only H. amplexicaule and the closely related H. nicotianaefolium (a member of the same section of the genus Heliotropium). No plants outside this genus were attacked. For the second “no-choice” phase, the host-plants used in Phase 1 were cut, forcing the insects to use either other plant species within the plots, emigrate, or die of starvation. Heliothrips heliotropica disappeared rapidly from the plot, while D. quadrijuga persisted for several days on Heliotropium arborescens with some exploratory nibbling and then disappeared. Host-choice behavior of these species therefore remained unchanged, even in the absence of the primary host-plants. In contrast, adults of Longitarsus sp. rapidly colonized and fed on H. arborescens when the preferred hosts were removed. It therefore demonstrated a wider host-range under these more extreme conditions. In fact, on some plots, feeding commenced on H. arborescens before the “no-choice” phase, once the two preferred Heliotropium species were heavily damaged by these insects. The two-phase test is shown here to be a useful way of measuring host-choice behavior under “normal” conditions of choice and under more extreme conditions, such as it might occur were an agent to locally destroy the target weed following its release. It therefore provides a more refined assessment of the risk that would be posed to nontarget plants by the release of weed biological control agents.     

Use of visual and olfactory cues of flowers of two brassicaceous species by insect pollinators

1. Pollinating insects exploit visual and olfactory cues associated with flower traits indicative of flower location and reward quality. Pollination can induce changes in these flower-associated cues, thereby influencing the behaviour of flower visitors. 2. This study investigated the main cues exploited by the syrphid fly Episyrphus balteatus and the butterfly Pieris brassicae when visiting flowers of Brassica nigra and Raphanus sativus plants. Whether pollen is used as a cue and whether pollination-induced changes affect flower volatile emission and the behavioural responses of the two pollinator species were also studied. 3. Pollinator preference was investigated by offering visual and olfactory cues individually as well as simultaneously in two-choice bioassays. Plant treatments included emasculation, hand-pollination and untreated control plants. The composition of flower volatiles from pollinated and unpollinated control plants was analysed by gas chromatography-mass spectrometry. 4. Both pollinators exhibited a strong bias for visual cues over olfactory cues. Neither pollinator used pollen as a cue. However, E. balteatus discriminated between newly opened and long-open flowers at short distance only when pollen was available. Flower visits by pollinators were influenced by pollination-induced changes in B. nigra but not R. sativus flowers. Pieris brassicae only responded to pollination-induced changes when visual and olfactory cues were offered simultaneously. The blend of volatiles emitted by B. nigra, but not R. sativus inflorescences was affected by pollination. 5. Collectively, the findings of this study show that different pollinators exploit different visual and olfactory traits when searching for flowers of two brassicaceous plant species.     

Applying a weed risk assessment approach to GM crops

Current approaches to environmental risk assessment of genetically modified (GM) plants are modelled on chemical risk assessment methods, which have a strong focus on toxicity. There are additional types of harms posed by plants that have been extensively studied by weed scientists and incorporated into weed risk assessment methods. Weed risk assessment uses robust, validated methods that are widely applied to regulatory decision-making about potentially problematic plants. They are designed to encompass a broad variety of plant forms and traits in different environments, and can provide reliable conclusions even with limited data. The knowledge and experience that underpin weed risk assessment can be harnessed for environmental risk assessment of GM plants. A case study illustrates the application of the Australian post-border weed risk assessment approach to a representative GM plant. This approach is a valuable tool to identify potential risks from GM plants.     

Field catches of Oxythyrea cinctella using visual and olfactory cues

Diurnal flower‐visiting scarabs (Coleoptera: Scarabaeidae, Cetoniinae) use visual and olfactory stimuli when locating possible feeding sites. In field trapping experiments in Turkey, testing different colours and floral volatile compounds, a combination of fluorescent yellow colour and a blend of 2‐phenylethanol and (±)‐lavandulol leads to the highest number of Oxythyrea cinctella (Schaum) being caught. Trap catches of O. cinctella with respect to the individually presented visual or olfactory cues are significantly lower than to a combination of the two. The closely‐related O. funesta, a sibling species in the Oxythyrea genus, is also known to be caught by the same colour‐volatile combination. When applied together in a high capture‐capacity trap, the fluorescent yellow colour and the binary floral odour blend may provide a suitable means of monitoring population changes of O. cinctella, which indicates their potential use in agriculture.     

Assessing the biosecurity risk from pathogens and herbivores to indigenous plants: lessons from weed biological control

Some potentially invasive herbivores/pathogens in their home range may attack plants originating from another geographic area. Methods are required to assess the risk these herbivores/pathogens pose to these plants in their indigenous ecosystems. The processes and criteria used by weed biological control researchers to assess the impact of potential biological control agents on a plant species in its non-native range provide a possible framework for assessing risks to indigenous plants. While there are similarities between these criteria such as the need for clear objectives, studies in the native range of the herbivore/pathogen, good knowledge of the ecology of the target plant and taxonomy of the plant and herbivore/pathogen, and modelling of the interaction between the two organisms, there are some important differences in approach. These include the need to consider the threat classification of the plant, the likely greater risk from polyphagous herbivores/pathogens than oligophagous or monophagous species, and the need to consider the impact of an additional natural enemy in conjunction with a suite of existing natural enemies. The costs of conducting a risk assessment of a herbivore/pathogen in another country that damages plants indigenous to another geographic area means that criteria will be needed for deciding which foreign herbivore/pathogen species should be assessed. These criteria could include the threat classification of the plant, the amount of damage to the particular plant organs affected, and the importance in key ecosystems.     

Finalizing host range determination of a weed biological control pathogen with best linear unbiased predictors and damage assessment

Colletotrichum gloeosporioides f. sp. salsolae (Penz.) Penz. & Sacc. in Penz. (CGS) is a facultative parasitic fungus being evaluated as a classical biological control agent of Russian thistle or tumbleweed (Salsola tragus L.). In initial host range determination tests, Henderson’s mixed model equations (MME) were used to generate best linear unbiased predictors (BLUPs) of disease severity reaction to CGS among 89 species of plants related to S. tragus. The MME provided: (1) disease assessments for rare and difficult or impossible to grow species, (2) environmentally independent measures of disease severity, (3) measures of disease severity for species versus a sample of material tested in a greenhouse, (4) objective indicators of susceptible and non-susceptible species, (5) a means to objectively compare disease on targets versus non-targets. Of the 89 species evaluated by the MME, eight native N. American species were predicted to be susceptible. As a result of these predictions, these eight species were further evaluated to determine the amount of actual damage caused by CGS. This was done by comparing root and shoot areas and weights between non-inoculated plants and plants inoculated with CGS. Results showed that several of the species exhibited some minor reduction in root weight and root area, but none of the species had any damage to above-ground plant parts. This supports the BLUP output in the initial host range determination tests. As a result of both analyses, there is no evidence that CGS would cause any non-target effects in nature.     

无翅桃蚜对龟纹瓢虫的视觉和嗅觉识别反应

为探明桃蚜Myzus persicae是否可以识别来自其捕食性天敌的嗅觉和视觉信息,本研究采用桃蚜和龟纹瓢虫Propylaea japonica这一猎物-捕食者系统,在室内测定了无翅桃蚜对龟纹瓢虫成虫体色和挥发物的识别反应。选择反应试验结果表明,无翅桃蚜既能通过视觉信息识别瓢虫的存在,也能通过嗅觉信息识别瓢虫的存在,说明桃蚜对龟纹瓢虫体色和气味信息的识别存在一定的冗余现象。剂量反应试验结果表明,龟纹瓢虫粗提物的剂量与其对桃蚜的驱避率呈现良好的线性关系,识别反应剂量的临界值在0.3~0.6个瓢虫当量之间,1.2~1.5个瓢虫当量的粗提物的驱避作用与20头活体龟纹瓢虫的驱避作用相当。结论认为,桃蚜可以识别龟纹瓢虫的存在,并对其作出一定的躲避反应,从而降低被捕食的风险。     

Trapping of European buprestid beetles in oak forests using visual and olfactory cues

Trapping approaches developed for the emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), were adapted for trapping several European oak buprestid species. These approaches included the use of natural leaf surfaces as well as green and purple plastic in sticky trap designs. Plastic surfaces were incorporated into novel ‘branch‐trap’ designs that each presented two 5 × 9‐cm² rectangular surfaces on a cardboard structure wrapped around the leaves of a branch. We used visual adult Agrilus decoys in an attempt to evoke male mating approaches toward the traps. Our first experiment compared the attractiveness of visual characteristics of the surfaces of branch‐traps. The second looked at the effect on trap captures of adding semiochemical lures, including manuka oil, (Z)‐3‐hexen‐1‐ol, and (Z)‐9‐tricosene. In total, 1 962 buprestid specimens including 14 species from the genus Agrilus were caught on 178 traps in a 22‐day time‐span. Overall, the green plastic‐covered branch‐traps significantly out‐performed the other trap designs. We further found that the presence of an EAB visual decoy placed on the trap surface often increased captures on these green traps, but this effect was stronger for certain Agrilus species than for others. The visual decoy was particularly important for the most serious pest detected, Agrilus biguttatus Fabricius, which was captured 13 times on traps with decoys, but only once without a decoy. There were some small but significant effects of odor treatment on the capture of buprestids of two common species, Agrilus angustulus Illiger and Agrilus sulcicollis Lacordaire. There were also 141 Elateridae specimens on these traps, which were not influenced by trap type or decoys. The results suggest that small branch‐traps of this nature can provide a useful new tool for monitoring of buprestids, which have the potential to be further optimized with respect to visual and olfactory cues.     

Divergent preferences of Rhagoletis pomonella host races for olfactory and visual fruit cues

The quality of different species of aphids as food for aphidiphagous ladybird beetles varies greatly. The optimal oviposition theory predicts that a female should lay eggs preferentially in patches of suitable prey and should be reluctant to oviposit in patches of less suitable prey. A no‐choice experiment was used to test whether aphids (Homoptera: Aphididae) of different suitability influence the oviposition behaviour of the two‐spot ladybird beetle, Adalia bipunctata (L.) (Coleoptera: Coccinellidae). The results indicate that A. bipunctata females are not more reluctant to lay eggs in the presence of moderately suitable compared to highly suitable aphids. However, females laid fewer eggs in the presence of toxic aphids.     

Interactions between two biological control agents and their target weed: a beetle,a bug and a cactus weed

ABSTRACT

Pereskia aculeata Miller (Cactaceae) is an invasive alien shrub introduced into South Africa from Brazil. The leaf-feeding beetle, Phenrica guerini Bechyne (Chrysomelidae), was released as a biological control agent in South Africa in 1991 followed by the stem-wilting bug, Catorhintha schaffneri Brailovsky & Garcia (Coreidae), in 2014. This study investigated the interactions between the two agents under laboratory conditions. Potted plants were exposed to one of four treatments: control (no agents), P. guerini only, C. schaffneri only and both species together. Four densities, ranging from 2 to 12 insects per plant were used. Cathorhitha schaffneri alone at low to moderate densities resulted in the same reduction in number of leaves and shoot length as when combine with P. guerini. At the highest density, C. schaffneri reduced the number of leaves significantly more than any treatment. Mortality of P. guerini was significantly higher than C. schaffneri at the highest density when in combination. The antagonistic interaction between P. guerini and C. schaffneri suggests that these agents should not be released together because this would impact negatively on the overall biocontrol programme against P. aculeata. It is recommended that C. schaffneri should be released at sites where P. guerini is not present. Extrapolation of laboratory-based studies into the field is often challenging, so mass-rearing and releases of P. guerini should continue until there is convincing proof that C. schaffneri alone is more effective than P. guerini in the field.     

The role of pre-release efficacy assessment in selecting classical biological control agents for weeds—applying the Anna Karenina principle

The goals in selecting classical biological control agents for weeds are to identify agents that will be both safe for release and effective in controlling their target plants. The release of ineffective agents should be avoided, as these add to the costs and risks of biological control without contributing to its benefits. While the principles of host-specificity testing and risk assessment for weed biological control agents have been extensively debated and refined, there has been less attention given to assessing the probable efficacy of agents prior to release. This reluctance to undertake pre-release efficacy assessment (PREA) is probably based on concerns that it will both add to the cost of screening biological control agents and introduce a risk of wrongly rejecting effective agents. We used a project simulation model to investigate the implications of using PREA as an additional filter in the agent selection process. The results suggest that, if it can be done at a lower cost than host-specificity testing, the use of PREA as the first filter can make agent selection more cost-effective than screening based on host-specificity alone. We discuss examples of PREA and potential approaches. The impact of biocontrol agents is a function of their range, abundance, and per-capita damage. While it will always be difficult to predict the post-release abundance of biological control agents from pre-release studies, some estimates of potential range can be obtained from studies of climatic adaptation. For agents that affect the vegetative growth or survival of their target weeds, experimental measurement of per-capita damage is feasible and can contribute to a reduction in the numbers of ineffective agents released. The Anna Karenina principle states that success in complex undertakings does not depend on a single factor but requires avoiding many separate causes of failure. We suggest that, in biological control of weeds, the use of agents that are not sufficiently damaging is one such cause that can be partially avoided by the use of pre-release efficacy assessment.     

A methodology for risk analysis of plurivorous fungi in biological weed control: Sclerotinia sclerotiorum as a model

In New Zealand, research is being performed on the use of the ubiquitous plurivorous ascomycete Sclerotinia sclerotiorum, as a biocontrol agent for Cirsium arvense in pasture. As a consequence of the wide host range of this fungus, the proposed biocontrol may pose a risk to non-target arable plants. Crop disease risk is primarily due to the formation of sclerotia. These perennating bodies may directly infect susceptible crops sown at the biocontrol site in years after the biocontrol event. Additionally they may cause infection in neighbouring crops through the formation of apothecia from which airborne ascospores are dispersed. A methodology for a risk analysis is proposed, embodying both temporal and spatial processes, where the relative disease risk from biocontrol is defined as the ratio of added to naturally occurring inoculum.Exponential decay is proposed as a suitable model for the survival of sclerotia in soils beneath pastures, and data from an ongoing study are used in a preliminary parameterisation. The model is applied to published data on initial sclerotium population size and unpublished data on natural levels of soilborne sclerotia, to reveal how a withholding period, defined as the interval of time that must elapse for the ratio of added to natural inoculum to equal unity, may be calculated.For quantifying escape and aerial dispersal of ascospores from a treated pasture, three mechanistic models are proposed. Firstly, a simple one-layer model of escape, giving a provisional estimation of 81% of released ascospores escaping the pasture canopy, is discussed. Secondly, ascospore dispersal within and close to the biocontrol site (<30 m) is described by a model revealing abrupt decline in the atmospheric concentration of ascospores within short distances from the biocontrol site. Thirdly, for predicting ascospore concentrations at greater distances (>100 m), a Gaussian plume model is proposed. In a first approximation, these models are used in an example calculation to show how a hypothetical safety zone might be determined. A map of the province of Canterbury showing the locations of different agricultural activities and infestations of dense populations of C. arvense, revealed that potential biocontrol sites mostly do not border arable land and that a safety zone would therefore be feasible.The study presented here provides conceptual and mathematical frameworks for risk analysis of the use of plurivorous fungi in biological weed control, using S. sclerotiorum as an example. However, before a judgement may be made about the phytosanitory risks associated with the use of S. sclerotiorum to control C. arvense in pastures in Canterbury, the models developed in this paper must be parameterized using data collected under a wide range of conditions in the field.     

Selecting non-target species for risk assessment of entomophagous biological control agents: Evaluation of the PRONTI decision-support tool

The release of entomophagous biological control agents can pose risks to non-target invertebrate species in the release area and beyond. Pre-release risk assessment of these agents often involves tests with non-target species; however, selecting appropriate test species can be difficult when there is a large number to choose from. The PRONTI (priority ranking of non-target invertebrates) tool has been developed to aid this selection process. This automated tool prioritises species for testing using five criteria: (1) direct and indirect hazards posed by the agent, (2) likelihood of exposure to the hazards, (3) ecological impacts that may result from that exposure, (4) species’ anthropocentric value and (5) testability. Criteria (1) and (2) produce a risk estimate that drives the ranking process. In a test of PRONTI’s ability to identify non-target species at most risk from a proposed biological control agent, we used a generalist predator already present in New Zealand, the Asian paper wasp Polistes chinensis, as if it were the agent in a hypothetical biocontrol programme aimed at lepidopteran pests in New Zealand kiwifruit orchards. A ranked list of 340 invertebrate taxa known to occur in kiwifruit orchards was produced. To validate the risk estimates for a direct attack by P. chinensis on each taxon, wasps were introduced to kiwifruit orchards and prey taxa identified. Risk estimates were accurate except where identified prey taxa had not previously been recorded from kiwifruit orchards.     

Responses to olfactory and visual cues by over-wintered and summer generations of the pollen beetle, Meligethes aeneus

Abstract.  Behavioural responses to odours of oilseed rape in bud and flower stage and to green and yellow colours are studied for the two main phenological stages of pollen beetles, Meligethes aeneus , a major pest of oilseed rape, Brassica napus . Over-wintered individuals oviposit in buds of oilseed rape. Adults of the new generation (i.e. the summer generation) feed on flowers of different plant species before over wintering. In olfactometer experiments, the over-wintered beetles display a higher preference for odour of oilseed rape at the bud stage compared with the summer generation, both with and without colour stimuli. Flower odours are preferred in combination with yellow colour. Colour stimuli presented alone do not affect the behaviour. The summer generation beetles respond to both bud and flower odour. Adding colour stimuli changes the summer generations preference towards yellow and flower odour.     

Oviposition preferences in Grapholita molesta: the relative importance of visual and olfactory cues

The oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae), is a major pest of fruit trees worldwide. Females oviposit on the young leaves of peach [Prunus persica (L.) Batsch (Rosaceae)] shoots at twilight, when light intensity changes markedly. Previous researches have shown that the oviposition response of this moth is guided particularly by visual and olfactory cues from host plants. However, the relative importance of visual and olfactory cues in their oviposition preference is largely unknown. As a crepuscular moth, how do choices change as light intensity drops rapidly from day to night? In the present study, through two‐choice behavioural experiments, the oviposition responses of G. molesta to visual and olfactory cues (alone or in combination) from peach shoots were tested at four light intensities ranging from daylight to starlight. Grapholita molesta showed similar oviposition responses to plant cues under 1 000, 100, 1 and 0.01 mW m^?2 illumination. Olfactory cues from peach shoots attracted significantly more oviposition than the negative control, whereas visual cues alone did not. Furthermore, olfactory cues were more attractive than visual cues, and no significant interaction was observed between the responses to the two cues. Our findings indicate that G. molesta females mainly rely on olfactory cues to recognize oviposition sites, regardless of differences in light intensity. These results do not provide evidence for attraction to visual cues, but the possibility that the brightness of leaves might be used to guide oviposition is discussed.