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Odour-mediated responses of a predatory mirid bug and its prey, the two-spotted spider mite.

It has been shown that many natural enemies of herbivorous arthropods use herbivore induced plant volatiles (HIPVs) to locate their prey. Herbivores can also exploit cues emitted by plants infested with heterospecifics or conspecifics. A study was conducted to test whether green bean HIPVs as well as odours emitted directly by spider mites influenced the orientation behaviour of the predatory mirid bug, Macrolophus caliginosus and its prey, Tetranychus urticae in a Y-tube olfactometer. Our results show that both spider mites and M. caliginosus preferred spider mite infested green bean plants to uninfested plants. For M. caliginosus this response was mediated by HIPVs whereas for T. urticae it was mediated through a composite response to both HIPVs and odours emitted directly by the conspecifics (and their associated products). The results may be of use in practical biocontrol situations, through e.g., plant breeding for improved HIPV production, conditioning of mass-reared predators to appropriate cues, and employment of "push-pull-strategies" by using HIPVs.

Animals↗

Influence of prey on developmental performance, reproduction and prey consumption of Neoseiulus californicus (Acari: Phytoseiidae).

The Spical strain of the predatory mite Neoseiulus californicus (McGregor) is used as a biological control agent, but little is known about its preferred prey and host plants in Japan. Here we studied the development, reproduction and prey consumption of the Spical strain when fed on eggs of five different spider mite species deposited on both their laboratory-rearing plant and cherry, on which all five spider mite species developed well. The developmental periods of immature N. californicus females and males were significantly affected by the prey species they fed on, but not by the plants. No difference was found between males and females. The developmental period was shorter on eggs of two Tetranychus species than on eggs of Panonychus ulmi. Immature females had a higher predation rate than immature males. Preoviposition period, oviposition period and the number of eggs laid per female were not significantly affected by either the plants or the type of prey eggs. The postoviposition period and total adult longevity were shorter on eggs of P. ulmi than of the other four prey species, but there was no effect of plant substrate. The postoviposition period of the Spical strain was much longer than that of other N. californicus strains or other predatory mite species: the postoviposition period of the Spical strain was more than three times longer than the oviposition period, accounting for more than 75% of the total adult longevity. This suggests that the females need multiple mating to reach full egg load, but this remains to be tested. Total consumption by N. californicus adults was lower for eggs of P. ulmi than for eggs of the other four species, apparently because of the shorter postoviposition period when fed on eggs of P. ulmi. The intrinsic rates of natural increase (r(m)) on the rearing plant did not differ among prey species, whereas those on cherry were significantly different: the value was higher on Tetranychus urticae eggs than on eggs of other species. Only when N. californicus fed on T. urticae eggs, the r(m)-values were significantly different between the rearing plant and cherry (higher on cherry). Thus, the Spical strain of N. californicus could feed on eggs of all five spider mite species, deposited on a variety of plants with similar r(m)-values, suggesting that it could be successfully used to control spider mites in orchards and various crop fields of Japan.

Animals↗

Phytoseiulus persimilis response to herbivore-induced plant volatiles as a function of mite-days.

The predatory mite, Phytoseiulus persimilis (Acari: Phytoseiidae), uses plant volatiles (i.e., airborne chemicals) triggered by feeding of their herbivorous prey, Tetranychus urticae (Acari: Tetranychidae), to help locate prey patches. The olfactory response of P. persimilis to prey-infested plants varies in direct relation to the population growth pattern of T. urticae on the plant; P. persimilis responds to plants until the spider mite population feeding on a plant collapses, after which infested plants do not attract predators. It has been suggested that this represents an early enemy-free period for T. urticae before the next generation of females is produced. We hypothesize that the mechanism behind the diminished response of predators is due to extensive leaf damage caused by T. urticae feeding, which reduces the production of volatiles irrespective of the collapse of T. urticae population on the plant. To test this hypothesis we investigated how the response of P. persimilis to prey-infested plants is affected by: 1) initial density of T. urticae, 2) duration of infestation, and 3) corresponding leaf damage due to T. urticae feeding. Specifically, we assessed the response of P. persimilis to plants infested with two T. urticae densities (20 or 40 per plant) after 2, 4, 6, 8, 10, 12 or 14 days. We also measured leaf damage on these plants. We found that predator response to T. urticae-infested plants can be quantified as a function of mite-days, which is a cumulative measure of the standing adult female mite population sampled and summed over time. That is, response to volatiles increased with increasing numbers of T. urticae per plant or with the length of time plant was infested by T. urticae, at least as long at the leaves were green. Predatory mites were significantly attracted to plants that were infested for 2 days with only 20 spider mites. This suggests that the enemy-free period might only provide a limited window of opportunity for T. urticae because relatively low numbers of T. urticae per plant can attract predators. Leaf damage also increased as a function of mite-days until the entire leaf was blanched. T. urticae populations decreased at this time, but predator response to volatiles dropped before the entire leaf was blanched and before the T. urticae population decreased. This result supports our hypothesis that predator response to plant volatiles is linked to and limited by the degree of leaf damage, and that the quantitative response to T. urticae populations occurs only within a range when plant quality has not been severely compromised.

Animals↗

Cannibalism and intraguild predation in Typhlodromus exhilaratus and T. phialatus (Acari: Phytoseiidae) under laboratory conditions.

Two species of Phytoseiidae are found in the same agroecosystem: Typhlodromus exhilaratus prevails in vine plots, while T. phialatus prevails in uncultivated surrounding areas. The objective of the present paper was to investigate whether the poor settlement of T. phialatus in vine plots can be explained by intraguild predation of these two species and/or cannibalism. Predatory abilities of the females on larvae and protonymphs were studied under laboratory conditions. A first experiment was conducted with only conspecific or heterospecific phytoseiid prey, in a second experiment Tetranychus urticae eggs were added to the phytoseiid prey. Oviposition, prey consumption, and escape rates of females were recorded. Oviposition and intraguild predation rates were higher for T. exhilaratus than for T. phialatus. Typhlodromus exhilaratus consumed fewer conspecifics than heterospecific phytoseiids, and oviposited when feeding on both diets. Typhlodromus phialatus consumed equal amounts of con- and heterospecifics. Although these two generalist predators belong to the type III defined by McMurtry and Croft (Annual Review of Entomology 42:291-321, 1997), our results suggest that they have different predation behaviour. However, because these results were obtained in experiments where no choice was given between the two phytoseiid species, they are difficult to link to previous studies conducted on the intraguild predation of the Phytoseiidae. The greater voracity and prolificacy of T. exhilaratus could partially explain the poor settlement of T. phialatus in vineyards and the predominance of T. exhilaratus. However, a full understanding of this phenomenon will require the study of other factors, such as susceptibility to pesticides and micro-climatic conditions, as well as the ability to cope with different food sources and host plants.

Animals↗

Influence of diet-related chemical cues from predators on the hatching of egg-carrying spiders.

Previous studies have shown that animals may make adaptive adjustments in response to chemical cues from predators, but hatching responses to diet-related chemical cues from predators have not been previously demonstrated. In the system studied here, the predator is an araneophagic jumping spider (Salticidae), Portia labiata, and the prey organism is a subsocial spitting spider, Scytodes pallida (Scytodidae). The spitting spider carries its eggs in its chelicerae, and carrying eggs is known to make it more vulnerable to predators. It is also known from an earlier study that the prior diet of the predator alters how dangerous the individual predator is to the spitting spider. In the experiments reported here, incubation time was shorter when volatile cues from the predator were present and longer in control tests when no chemical cues from the predator were present. The previous predator's diet also influenced incubation time: when in the presence of volatile cues from individuals of P. labiata that had previously fed on individuals of S. pallida, incubation time was shorter than when in the presence of volatile cues from individuals of P. labiata that had been feeding instead on house flies.

Animal Feed↗

Parallel arms races between garter snakes and newts involving tetrodotoxin as the phenotypic interface of coevolution.

Parallel "arms races" involving the same or similar phenotypic interfaces allow inference about selective forces driving coevolution, as well as the importance of phylogenetic and phenotypic constraints in coevolution. Here, we report the existence of apparent parallel arms races between species pairs of garter snakes and their toxic newt prey that indicate independent evolutionary origins of a key phenotype in the interface. In at least one area of sympatry, the aquatic garter snake, Thamnophis couchii, has evolved elevated resistance to the neurotoxin tetrodotoxin (TTX), present in the newt Taricha torosa. Previous studies have shown that a distantly related garter snake, Thamnophis sirtalis, has coevolved with another newt species that possesses TTX, Taricha granulosa. Patterns of within population variation and phenotypic tradeoffs between TTX resistance and sprint speed suggest that the mechanism of resistance is similar in both species of snake, yet phylogenetic evidence indicates the independent origins of elevated resistance to TTX.

Animals↗

Diet-related modification of cuticular hydrocarbon profiles of the Argentine ant, Linepithema humile, diminishes intercolony aggression.

Territorial boundaries between conspecific social insect colonies are maintained through a highly developed nestmate recognition system modulated by heritable and, in some instances, nonheritable cues. Argentine ants, Linepithema humile, use both genetic and environmentally derived cues to discriminate nestmates from nonnestmates. We explored the possibility that intraspecific aggression in the Argentine ant might diminish when colonies shared a common diet. After segregating recently field-collected colony pairs into high or moderate aggression categories, we examined the effect of one of three diets: two hydrocarbon-rich insect prey, Blattella germanica and Supella longipalpa, and an artificial (insect-free) diet, on the magnitude of aggression loss. Aggression diminished between colony pairs that were initially moderately aggressive. However, initially highly aggressive colony pairs maintained high levels of injurious aggression throughout the study, independent of diet type. Each diet altered the cuticular hydrocarbon profile by contributing unique, diet-specific cues. We suggest that acquisition of common exogenous nestmate recognition cues from shared food sources may diminish aggression and promote fusion in neighboring colonies of the Argentine ant.

Aggression↗

Increased sulfur precursors and volatiles production by the leek Allium porrum in response to specialist insect attack.

To defend themselves against herbivory, plants use a variety of direct and indirect strategies involving induced increases in secondary substances. Species of the Allium genus (Alliaceae), such as the leek Allium porrum (L.), produce nonprotein sulfur amino acids derived from cysteine, i.e., alk(en)yl-cysteine sulfoxides that are precursors of volatile thiosulfinates and disulfides. These defend most species including the specialist leek moth, Acrolepiopsis assectella. We determined by measuring the increase in the sulfur precursor propyl-cysteine sulfoxide (PCSO) if production of this precursor is induced in response to moth attack and mechanical wounding. The concentration of PCSO was determined by HPLC in 2- or 6-mo-old leeks after attacks of various intensity either by the specialist leek moth or by a generalist moth, Agrotis ipsilon. Injury-induced release of sulfur volatiles was measured by GC/MS after the attacks. Results showed an increase in the production of sulfur compounds in both the precursor and volatile form, occurring only in association with intensive attacks by leek moths. The increase in sulfur precursors also led to an increase in the release of sulfur volatiles. This induced response may provide an effective defense strategy against the plant's main natural enemy, both directly and indirectly by attracting entomophagous insects.

Animals↗

Methyl salicylate, a soybean aphid-induced plant volatile attractive to the predator Coccinella septempunctata.

Induced volatiles provide a signal to foraging predatory insects about the location of their prey. In Iowa, early in the growing season of soybean, Glycine max, many predacious seven-spotted lady beetles, Coccinella septempunctata, were observed on plants with heavy infestations of soybean aphid, Aphis glycines. We studied whether the attraction of this beetle is caused by the release of specific volatile compounds of soybean plants infested by aphids. Volatile compounds emitted by soybean plants infested by aphids were compared with those of undamaged, uninfested, and artificially damaged plants. Gas chromatography-mass spectrometry analyses revealed consistent differences in the profiles of volatile compounds between aphid-infested soybean plants and undamaged ones. Significantly more methyl salicylate was released from infested plants at both the V1 and V2 plant growth stages. However, release patterns of two other induced plant volatiles, (D)-limonene and (E,E)-alpha-farnesene, differed between the two plant growth stages. Gas chromatographic-electroantennographic detection of volatile extracts from infested soybean plants showed that methyl salicylate elicited significant electrophysiological responses in C. septempunctata. In field tests, traps baited with methyl salicylate were highly attractive to adult C. septempunctata, whereas 2-phenylethanol was most attractive to the lacewing Chrysoperla carnea and syrphid flies. Another common lady beetle, the multicolored Asian lady beetle, Harmonia axyridis, showed no preference for the compounds. These results indicate that C. septempunctata may use methyl salicylate as the olfactory cue for prey location. We also tested the attractiveness of some selected soybean volatiles to alate soybean aphids in the field, and results showed that traps baited with benzaldehyde caught significantly higher numbers of aphids.

Animals↗

Chemical defense in harvestmen (arachnida, opiliones): do benzoquinone secretions deter invertebrate and vertebrate predators?

Two alkylated 1,4-benzoquinones were identified from the defensive secretion produced by the neotropical harvestman Goniosoma longipes (Gonyleptidae). They were characterized as 2,3-dimethyl-1,4-benzoquinone and 2-ethyl-3-methyl-1,4-benzoquinone. We tested the effectiveness of these benzoquinone secretions against several predator types, including invertebrates and vertebrates. Different predators were exposed to the harvestmen's gland secretion or to distilled water in laboratory bioassays. Our results indicate that secretions containing the 1,4-benzoquinones released by G. longipes can be an effective defense against predation, and that the effectiveness of the secretion is dependent on the predator type. The scent gland secretion repelled seven ant species, two species of large wandering spiders, and one frog species, but was not an effective defense against an opossum. Our study also demonstrates that the scent gland secretion of G. longipes can work as a chemical shield preventing the approach of three large predatory ants for at least 10 min. The chemical shield may protect the harvestman against successive attacks of the same ant worker and also allow the harvestman to flee before massive ant recruitment. Our data support the suggestion that chemical defenses may increase survival with some but not all potential predators. This variation in defense effectiveness may result from many interacting factors, including the attack strategy, size, learning ability, and physiology of the predators, as well as the chemical nature of the defensive compounds, type of emission, and amount of effluent released by the prey.

Adaptation, Physiological↗

Similarity of cuticular lipids between a caterpillar and its host plant: a way to make prey undetectable for predatory ants?

Ithomiine butterflies (Nymphalidae) have long-lived, aposematic, chemically protected adults. However, little is known about the defense mechanisms in larvae and other juvenile stages. We showed that larvae Mechanitis polymnia are defended from ants by a chemical similarity between their cuticular lipids and those of the host plant, Solanum tabacifolium (Solanaceae). This is a novel defense mechanism in phytophagous insects. A field survey during one season showed that larval survivorship was up to 80%, which is high when compared with other juvenile stages. In a laboratory bioassay, live larvae on their host plant were not attacked by the predatory ant Camponotus crassus (Formicidae). Two experiments showed that the similarity between the cuticular lipids of M. polymnia and S. tabacifolium protected the larvae from C. crassus: (a) when the caterpillar was switched from a host plant to a non-host plant, the predation rate increased, and (b) when a palatable larva (Spodoptera frugiperda, Noctuidae) was coated with the cuticular lipids of M. polymnia and placed on S. tabacifolium leaves, it no longer experienced a high predation rate. This defensive mechanism can be defined as chemical camouflage, and may have a double adaptive advantage, namely, protection against predation and a reduction in the cost of sequestering toxic compounds from the host plant.

Adaptation, Physiological↗

Predator-labeling effect on byssus production in marine mussels Perna viridis (L.) and Brachidontes variabilis (Krauss).

Mussels Perna viridis and Brachidontes variabilis were exposed to chemical cues from the predatory crab Thalamita danae maintained on different diets, and byssal thread production of the mussels was studied. P. viridis produced the highest number as well as the thickest and longest byssal threads when they were exposed to crabs maintained on a diet of P. viridis as compared with those exposed to crabs maintained on a diet of the top shell Monodonta labio, the rock oyster Saccostrea cucullata, or crabs that were starved. For B. variabilis, results were similar, in that a diet containing B. variabilis elicited the greatest response as compared with other treatments. This indicates that the mussels were able to discriminate chemical cues released from predators maintained on different diets, and respond accordingly to the level of predation risk. By increasing the strength of byssal attachment as a defensive trait, the chance of being dislodged and consumed by crabs is reduced. As energy cost involved in the induction of an antipredatory response is considerable, this defensive trait seems to be an advantage to the mussels in enhancing efficiency. The short response time in byssal thread production allows the mussels to increase resistance against predation by crabs at the time when predation pressure is the highest in a tidal cycle.

Animals↗

Kairomonal response of predators to three pine bast scale sex pheromones.

The kairomonal activity of the sex pheromones of three pine bast scales, Matsucoccus feytaudi, Matsucoccus josephi, and Matsucoccus matsumurae, as well as a new analog of the M. feytaudi sex pheromone, were investigated in pine forests of France, Portugal, and Italy. The response of the maritime pine bast scale predators, Elatophilus spp. and Hemerobius stigma, was used to test the influence of trapping methods, kairomone composition, and dose. Both predators showed significant attraction to all compounds except to the sex pheromone of M. josephi. Significant increase in captures was observed as a function of dose, and within the studied dose range, up to 2200 mug, no threshold saturation limits were observed for any of the attractive compounds. Trap design and size did not significantly influence predator captures, except for high population levels of Elatophilus crassicornis, when plate traps were more efficient than delta traps. Geographic variations were found in the kairomonal responses patterns of both predators, with the M. matsumurae sex pheromone being more attractive to the oriental populations from Corsica and Italy, whereas the western populations in Aquitaine and Portugal were more attracted to the M. feytaudi sex pheromone.

Animals↗

Chemically mediated intraguild predator avoidance by aphid parasitoids: interspecific variability in sensitivity to semiochemical trails of ladybird predators.

The avoidance responses of aphid parasitoids with varying host ranges (Aphidius eadyi, Aphidius ervi, and Praon volucre) to chemical trails deposited by intraguild predatory ladybirds, Coccinella septempunctata and Adalia bipunctata, were investigated. Females of all three parasitoid species avoided leaves previously visited by C. septempunctata or A. bipunctata adults. The avoidance responses shown by the two Aphidius species were stronger to trails of C. septempunctata than to those of A. bipunctata. However, P. volucre avoided trails of both ladybird species to a similar degree. Dose responses of these three parasitoid species to the hydrocarbons n-tricosane (C23H48), n-pentacosane (C25H52), and n-heptacosane (C27H56), which are components of the trails of both C. septempunctata and A. bipunctata, were evaluated. Dual-choice bioassays indicated the following: (1) A. eadyi showed more sensitive avoidance responses to n-tricosane than did the other two parasitoid species, (2) all three species showed similar responses to n-pentacosane across a range of doses, and (3) only P. volucre showed avoidance responses to n-heptacosane. Quantitative analyses of each hydrocarbon in the trails of the two ladybird species showed that n-pentacosane and n-heptacosane occur in significantly greater amounts in C. septempunctata trails than in those of A. bipunctata. The trails of the two species also differ qualitatively in the other hydrocarbons present.

Alkanes↗

Production and predator-induced release of volatile chemicals by the plant bug Lygus hesperus.

Both sexes of adult western tarnished plant bug, Lygus hesperus Knight (Heteroptera: Miridae), released three volatile chemicals in relatively large amounts when attacked by ants (Pogonomyrmex rugosus and Solenopsis xyloni) or when grabbed by forceps, as determined by solid-phase microextraction (SPME) and gas chromatography-mass spectrometry (GC-MS). The relative amounts of the volatile compounds, hexyl butyrate, (E)-4-oxo-2-hexenal, and (E)-2-hexenyl butyrate, absorbed by SPME as a percentage of the largest were 100%, 44%, and 4%, respectively, from females, and 83%, 37%, and 3% from males. Both ant species were repelled by the defensive discharges (confirmed by SPME) when the ants attacked L. hesperus adults. Sexually mature L. hesperus were individually extracted in pentane to quantify the mean amounts of hexyl butyrate (14.9 microg/female; 10.3 microg/male), (E)-4-oxo-2-hexenal (2.7 microg/female; 3.1 microg/male), and (E)-2-hexenyl butyrate (1.2 microg/female; 0.6 microg/male). (E)-4-Oxo-2-hexenal was unstable in solvent when in contact with a macerated adult, but relatively stable when the solution was decanted within minutes. The production of the three major volatile components began soon after the emergence of the adult and amounts increased for about 5-10 d with little or no increase thereafter. Minor additional constituents were cross-correlated in many cases with the three major ones. A cost of defensive secretion is suggested for females but not for males, because heavier females produced more volatile compounds than lighter females. The initial discharge percentage, defined as the proportion of volatile compounds initially present that is discharged to defend against predation was estimated at about 50% in males and 70% in females. Newly eclosed adults did not produce volatile chemicals until 2 d after molting.

Aging↗

Individual variation in alkaloid content of poison frogs of Madagascar (Mantella; Mantellidae).

Brightly colored Malagasy poison frogs, Mantella spp., sequester lipophilic, basic alkaloids from arthropod prey for their own chemical defense. Consequently, microsympatric prey diversity is expected to influence alkaloid diversity observed in poison frogs. Twenty-two specimens of three Mantella species from four localities in moist forests of southeastern Madagascar were analyzed individually via gas chromatography-mass spectrometry, revealing that they contain over 80 known alkaloids. Frogs within a locality possessed significantly similar alkaloid content and diversity, while frogs from areas that varied in disturbance, elevation, and/or species showed greater differences. Based on dietary data, the larger frog species Mantella baroni consumed more and larger prey, and showed greater diversity in skin alkaloids than significantly smaller Mantella bernhardi. Additionally, frogs from the most pristine locality had the greatest number of alkaloids, whereas individuals from the most disturbed localities had the least. In a comparison of frog alkaloid profiles over a 10- to 14-yr period, alkaloid turnover, and thus presumably alkaloid-source arthropod turnover, was high in a disturbed locality and low in the pristine primary forest locality. We demonstrate that the nonlethal transcutaneous amphibian stimulator (TAS) is effective for harvesting alkaloids from poison frogs; future studies using this device could obtain larger sample sizes without harming local frog populations.

Alkaloids↗

Effects of Bt maize-fed prey on the generalist predator Poecilus cupreus L. (Coleoptera: Carabidae).

We investigated the effects of transgenic maize (Zea mays) expressing Bacillus thuringienses toxin (Bt maize) on larval and adult Poecilus cupreus carabid beetles in laboratory studies. In no-choice trials, neonate P. cupreus larvae were fed exclusively with Spodoptera littoralis caterpillars, which had been raised on Bt maize. S. littoralis raised on conventional maize or "high quality" Calliphora sp. pupae were fed to the beetle larvae in two control treatments. Bt-maize-fed caterpillar prey increased mortality to 100% within 40 days. The experiment was repeated with 10-day-old beetle larvae. Bt treatment resulted in fewer pupae than in both controls, and in a higher mortality than in the Calliphora control. S. littoralis was suitable as exclusive prey in no-choice tests, at least for 40 days, although prey quality seemed to be low compared to Calliphora pupae. The observed effects are most likely indirect effects due to further reduced nutritional prey quality. However, direct effects cannot be excluded. In the second part of the study, exposure of P. cupreus to Bt intoxicated prey was examined in paired-choice tests. Adult beetles were offered a choice between different prey conditions (frozen and thawed, freshly killed or living), prey types (S. littoralis caterpillars, Calliphora sp. pupae, cereal aphids) and prey treatments (raised on Bt or conventional maize). Living prey was preferred to frozen and dead prey. Caterpillars were only preferred to fly pupae and aphids when living. Prey treatment seemed to be least important for prey selection. The tests showed that P. cupreus ingested caterpillars readily and there was no evidence of them avoiding Bt containing prey, which means exposure in the field could occur. The presented protocols are a first step towards ecological risk assessment for carabid beetles.

Animals↗

Modeling the role of diffusion coefficients on Turing instability in a reaction-diffusion prey-predator system.

The paper is concerned with the effect of variable dispersal rates on Turing instability of a non-Lotka-Volterra reaction-diffusion system. In ecological applications, the dispersal rates of different species tends to oscillate in time. This oscillation is modeled by temporal variation in the diffusion coefficient with large as well as small periodicity. The case of large periodicity is analyzed using the theory of Floquet multipliers and that of the small periodicity by using Hill's equation. The effect of such variation on the resulting Turing space is studied. A comparative analysis of the Turing spaces with constant diffusivity and variable diffusivities is performed. Numerical simulations are carried out to support analytical findings.

Algorithms↗