Phonotactic behaviour and vertical sound source localisation of the parasitoid fly Emblemasoma auditrix (Diptera: Sarcophagidae)

1. Acoustically guided movement in a three‐dimensional space is a complex behavioural task performed notably by birds, bats, and some insect species. The precision of acoustic orientation depends on the directionality of the hearing system as well as on auditory behaviour. 2. The fly Emblemasoma auditrix Diptera (Sarcophagidae) is a parasitoid of the cicada Okanagana rimosa Auchenorrhyncha (Cicadidae) and locates its host in the complex habitat of a forest. The phonotactic behaviour of the fly was analysed experimentally with emphasis on the vertical domain in the field. Different experimental setups allowed discriminating subsequent steps in the phonotactic behaviour of E. auditrix. 3. During the phonotactic flight, flies first landed on landmarks, which were used to re‐adjust to the elevation of the sound source. Acoustic targets were located from these resting positions. The sound source elevation was detected at the start of the flight as the longitudinal body axis was adjusted to the inclination of the target sound source. 4. Flies usually did not land directly upon the sound source, but landed nearby, and most often above the target. Within the target area, types of movement for the final approach differed in respect to target position; flies walked predominantly if the final target was located above or below, but for horizontally located targets much of the distance was covered by flight. 5. In conclusion, E. auditrix can locate the acoustic target in complex habitats and uses a flexible multi‐step approach for short‐range phonotaxis.     

Death comes suddenly to the unprepared: singing crickets, call fragmentation, and parasitoid flies

Male field crickets are subject to a delicate dilemma becausetheir songs simultaneously attract mates and acoustic predators.It has been suggested that in response, crickets have modifiedvarious temporal song parameters to become less attractiveto acoustic predators. We investigated whether crickets withchirping (versus trilling) song structures are less likely toattract acoustically orienting parasitoid flies. Experimentally,we evaluated the phonotactic quest of the parasitoid fly Ormiaochracea in response to broadcast cricket calls, presentedboth simultaneously (choice paradigm) and sequentially (no-choiceparadigm). Flight trajectories were recorded in darkness usingthree-dimensional active infrared video tracking. The flies showed remarkable phonotactic accuracy by landing directly onthe loudspeaker. The introduction of acoustic fragmentationthat resembles calls of many chirping crickets altered theflies' phonotactic accuracy only slightly. Our results documentdifferential attraction between trilling and chirping cricket songs and quantitatively demonstrate that chirping songs, ifpresented alone, do not impair the efficiency (temporal investmentand landing accuracy) of the flies' phonotactic quest. Thisstudy shows that song fragmentation is no safeguard againstacoustic parasitism. We conclude that, in general, a cricket may reduce predation only if its neighbors are acousticallymore conspicuous, chiefly by amplitude.     

Phonotaxis of the female parasitoid Emblemasoma auditrix (Diptera, Sarcophagidae) in relation to number of larvae and age

The dipteran parasitoid Emblemasoma auditrix locates its host acoustically. Analysis showed that phonotactic female flies usually carry fully developed larvae within their uteri. The mean number of larvae per female at the beginning of the season was 37.9 (range from 10 to 50). The number of larvae decreased rapidly with increasing singing activity of the host cicada (Okanagana rimosa). In high-density host populations the parasitoid is likely to become egg-limited. A possible selective phonotactic responsiveness depending on the number of larvae or the age of the female was tested with song models. Phonotaxis depended on both the temporal structure and the frequency content, but in the field no correlation was found between the number of larvae and the preferences for the acoustic signal. Experiments in the laboratory showed that flies without host contact broadened their phonotactic stimulus range with age.     

Host preferences in a phonotactic parasitoid of field crickets: the relative importance of host song characters

Abstract.  1. Predators, including insect parasitoids, often eavesdrop on prey signals, and as a result, predation can have important effects on the evolution of prey signalling behaviour.
2. The phonotactic parasitoid fly, Ormia ochracea , uses the calling songs of male field crickets to locate their field crickets hosts. In the western USA, this fly parasitises the variable field cricket, Gryllus lineaticeps . Previous work with one fly population suggested that female flies, like female field crickets, preferentially orient to male songs with higher chirp rates and longer chirp durations, although a limited range of male song types was used in this previous study. The current study, with a different fly population, used field-based, two-speaker choice tests to examine: (1) the effect of male chirp rate and chirp duration on fly attraction, using a natural range of song types; and (2) the relative importance of these song types in host selection by the flies.
3. Three lines of evidence suggested that chirp rate is more important than chirp duration in host selection. (a) The flies consistently preferred higher chirp rates but only sometimes preferred longer chirp durations. (b) The flies consistently preferred higher chirp rate/shorter chirp duration songs to lower chirp rate/longer chirp duration songs. (c) Preferences for longer chirp durations could be eliminated by increasing the amplitude of the less attractive song type, while preferences for higher chirp rates could only sometimes be eliminated by increasing the amplitude of the less attractive song type.
4. Fly predation may favour lower chirp rates and shorter chirp durations in G. lineaticeps , and may have resulted in stronger selection on chirp rate than on chirp duration.     

Spatial and temporal variation in the parasitoid assemblage of an exophytic polyphagous caterpillar

Abstract 1. Over 3400 larvae of the polyphagous ground dwelling arctiid Grammia geneura were sampled and reared over seven generations in order to characterise its parasitoid assemblage and examine how and why this assemblage varies over time and space at a variety of scales.
2. The total parasitoid assemblage of 14 species was dominated both in diversity and frequency by relatively polyphagous tachinid flies.
3. Both the composition of the parasitoid assemblage and frequency of parasitism varied strikingly among and within sampling sites, seasons, and years.
4. Overall rates of parasitism increased consistently over the duration of caterpillar development.
5. Within sampling sites, parasitism rates were non-random with respect to habitat structure and caterpillar behaviour for the most abundant parasitoid species.
6. The large variability in parasitoid assemblage structure over space and time in this system may be a function of local host population abundance, habitat-specific parasitism, and indirect interactions between G. geneura and other Macrolepidoptera through shared oligophagous and polyphagous parasitoids.     

Preference and performance of the hyperparasitoid Syrphophagus aphidivorus (Hymenoptera: Encyrtidae): fitness consequences of selecting hosts in live aphids or aphid mummies

Abstract.  1. Theoretical models predict that ovipositional decisions of parasitoid females should lead to the selection of the most profitable host for parasitoid development. Most parasitoid species have evolved specific adaptations to exploit a single host stage. However, females of the aphid hyperparasitoid Syrphophagous aphidivorus (Mayr) (Hymenoptera: Encyrtidae) display a unique and atypical oviposition behaviour by attacking either primary parasitoid larvae in live aphids, or parasitoid pupae in dead, mummified aphids.
2. In the laboratory, the correlation between host suitability and host preference of S. aphidivorus on the host Aphidius nigripes Ashmead parasitising the aphid Macrosiphum euphorbiae (Thomas) was investigated.
3. The relative suitability of the two host stages was determined by measuring hyperparasitoid fitness parameters (survival, development time, fecundity, sex ratio, and adult size of progeny), and calculating the intrinsic rate of population increase ( r _m). Host preference by S. aphidivorus females and the influence of aphid defence behaviour on host selection was also examined.
4. Hyperparasitoid offspring performance was highest when developing from hosts in aphid mummies and females consistently preferred this host to hosts in parasitised aphids. Although aphid defensive behaviour may influence host selection, it was not a determining factor. Ecological and evolutionary processes that might have led to dual oviposition behaviour in S. aphidivorus are discussed.     

Consequences for a host–parasitoid interaction of host-plant aggregation, isolation, and phenology

Abstract.  1. Spatial habitat structure can influence the likelihood of patch colonisation by dispersing individuals, and this likelihood may differ according to trophic position, potentially leading to a refuge from parasitism for hosts.
2. Whether habitat patch size, isolation, and host-plant heterogeneity differentially affected host and parasitoid abundance, and parasitism rates was tested using a tri-trophic thistle–herbivore–parasitoid system.
3.  Cirsium palustre thistles ( n = 240) were transplanted in 24 blocks replicated in two sites, creating a range of habitat patch sizes at increasing distance from a pre-existing source population. Plant architecture and phenological stage were measured for each plant and the numbers of the herbivore Tephritis conura and parasitoid Pteromalus elevatus recorded.
4. Mean herbivore numbers per plant increased with host-plant density per patch, but parasitoid numbers and parasitism rates were unaffected. Patch distance from the source population did not influence insect abundance or parasitism rates. Parasitoid abundance was positively correlated with host insect number, and parasitism rates were negatively density dependent. Host-plant phenological stage was positively correlated with herbivore and parasitoid abundance, and parasitism rates at both patch and host-plant scales.
5. The differential response between herbivore and parasitoid to host-plant density did not lead to a spatial refuge but may have contributed to the observed parasitism rates being negatively density dependent. Heterogeneity in patch quality, mediated by variation in host-plant phenology, was more important than spatial habitat structure for both the herbivore and parasitoid populations, and for parasitism rates.     

Superparasitism in Cotesia glomerata: response of hosts and consequences for parasitoids

Abstract.  1. Superparasitism occurs in Cotesia glomerata (Hymenoptera: Braconidae), a gregarious endoparasitoid of Pieris spp. (Lepidoptera: Pieridae). The response of P. brassicae larvae to superparasitism and the consequences for the parasitoid were examined in order to elucidate the ecological significance of this behaviour.
2. Field surveys of a Swiss population revealed that C. glomerata brood sizes from P. brassicae larvae ranged from three to 158, and both the female ratio and the body weight of emergent wasps correlated negatively with brood size. In the laboratory, single oviposition on P. brassicae larvae did not produce any brood size larger than 62, but brood size increased with superparasitism.
3. Laboratory experiments demonstrated that both naive and experienced female wasps were willing to attack hosts that had been newly parasitised by themselves or conspecifics. Superparasitism reduced survivorship but increased food consumption and weight growth in P. brassicae larvae. Superparasitism lengthened parasitoid development and prolonged the feeding period of host larvae.
4. Despite a trade-off between maximising brood size and optimising the fitness of individual offspring, two or three ovipositions on P. brassicae larvae resulted in a greater dry female mass than did a single oviposition on the host. Thus, superparasitism might be of adaptive significance under certain circumstances, especially when host density is low and unparasitised hosts are rare in a habitat.     

Limited flexibility in female Pacific field cricket (Teleogryllus oceanicus) exploratory behaviors in response to perceived social environment

How populations adapt, or not, to rapid evolution of sexual signals has important implications for population viability, but is difficult to assess due to the paucity of examples of sexual signals evolving in real time. In Hawaiian populations of the Pacific field cricket (Teleogryllus oceanicus), selection from a deadly parasitoid fly has driven the rapid loss of a male acoustic signal, calling song, that females use to locate and evaluate potential mates. In this newly quiet environment where many males are obligately silent, how do phonotactic females find mates? Previous work has shown that the acoustic rearing environment (presence or absence of male calling song) during late juvenile stages and early adulthood exposes adaptive flexibility in locomotor behaviors of males, as well as mating behaviors in both sexes that helps facilitate the spread of silent (flatwing) males. Here, we tested whether females also show acoustically induced plasticity in walking behaviors using laboratory‐reared populations of T. oceanicus from Kauai (HI; >90% flatwings), Oahu (HI; ~50% flatwings), and Mangaia (Cook Islands; no flatwings or parasitoid fly). Though we predicted that females reared without song exposure would increase walking behaviors to facilitate mate localization when song is rare, we discovered that, unlike males, female T. oceanicus showed relatively little plasticity in exploratory behaviors in response to an acoustic rearing environment. Across all three populations, exposure to male calling song during development did not affect latency to begin walking, distance walked, or general activity of female crickets. However, females reared in the absence of song walked slower and showed a marginally non‐significant tendency to walk for longer durations of time in a novel environment than those reared in the presence of song. Overall, plasticity in female walking behaviors appears unlikely to have facilitated sexual signal loss in this species.     

Avoidance of the host immune response by a generalist parasitoid, Compsilura concinnata Meigen

Abstract.  1. Ecological interactions between parasitoids and their hosts are extremely strong as parasitoid offspring rely entirely on an individual host to complete development. The ability of a parasitoid to use a host is influenced directly by the degree to which the parasitoid can overcome host defences and grow within the host.
2. Hymenopteran parasitoids have evolved different host-specific strategies to defeat the host immune system, such as the use of venom, endosymbiont virus, or mimicking the host tissue. Dipteran parasitoids from the Tachinidae family do not use these subterfuges and rely mainly on avoiding the host immune system by hiding in specific tissues.
3. Little is known of the effect of this strategy on the host immune system, the absorption of nutrients by the parasitoid larvae, or the implications for parasitoid host range.
4. In this study, the impact of a polyphagous tachinid parasitoid Compsilura concinnata Meigen on a pest lepidopteran Trichoplusia ni Hübner are assessed. Phenoloxidase levels and haemolymph proteins were measured in parasitised T. ni as a function of host immune response.
5. Haemolymph phenoloxidase in the host did not vary with parasitisation but was triggered when a piece of monofilament was implanted in the haemocoel. Haemolymph proteins were depleted in heavily parasitised T. ni .
6. These results indicate that C. concinnata has a strategy that avoids the host immune system, and accesses the necessary nutrients for larval growth. This strategy could explain the success of this tachinid and its wide host range.     

Behaviour of a specialist parasitoid, Cotesia melitaearum: from individual behaviour to metapopulation processes

Abstract. 1. Foraging behaviour and movement within and among host patches of the specialist parasitoid wasp Cotesia melitaearum (Braconidae) attacking the larvae of Melitaea cinxia (Nymphalidae) were studied in the field and in the laboratory.
2. In the field, female wasps aggregated in large host groups in the autumn and caused positive spatial density-dependent parasitism in the field. Wasps stayed longer with groups of pre-diapause caterpillars than with post-diapause caterpillars, but attacked them less frequently.
3. In the laboratory, wasps attacked larger larvae more readily than smaller larvae. Also in the laboratory, wasps exposed to larvae outside their protective webs showed differences in the rates at which they attacked larvae fed different diets, implicating host plant-derived chemicals as proximate cues for foraging wasps.
4. Mark–recapture studies indicated that there was a low rate of successful movement of wasps among groups of young larvae within a habitat patch in the autumn and no successful movement of wasps across non-habitat. In contrast, wasps moved frequently among groups of late-instar caterpillars in the spring.
5. Host caterpillars of different ages responded very differently to wasp attacks. Pre-diapause larvae remained in groups and used collective head-jerking behaviour to defend themselves, whereas post-diapause larvae dispersed away from the group immediately after being attacked.
6. Population and metapopulation level dynamics of the host–parasitoid interaction are discussed in light of these observations of the behaviour of individual wasps.     

Searching and oviposition behaviour of Amitus fuscipennis, a parasitoid of the greenhouse whitefly

Abstract: Amitus fuscipennis MacGown & Nebeker (Hym., Platygasteridae) is a parasitoid of the greenhouse whitefly, Trialeurodes vaporariorum (Westwood) (Hom., Aleyrodidae) on some crops as bean and snap bean (both Phaseolus vulgaris L.) in Colombia. The searching and oviposition behaviour of A. fuscipennis was studied in the laboratory, using T. vaporariorum as a host on bean leaves. The parasitoid's basic search pathway consisted of walking, encountering the host, and drumming and probing it. While walking, the parasitoid stopped for short periods of time, partly to preen itself. Walking while searching comprised 61% of the adults' time budget and probing hosts represented 16%. After a host nymph was parasitized by A. fuscipennis , the parasitoid remained close by and continued searching for new hosts in the immediate vicinity. Such behaviour suggests area-restricted searching. The parasitoid preferred first instars of T. vaporariorum for oviposition, resulting in long developmental times. Amitus fuscipennis had a high percentage of host acceptance resulting in a high percentage parasitism (60%) of all encountered hosts. Amitus fuscipennis , on average, walked faster before an oviposition (1.4 mm/s) than other whitefly parasitoids. The implications of these findings for the control of T. vaporariorum are discussed.     

Dispersal and foraging behaviour of Platygaster californica: hosts can't run, but they can hide

Abstract.  1. Host–parasitoid models often identify foraging behaviour and dispersal distance as important for system persistence.
2. Laboratory observations and field trials were used to characterise foraging behaviour and dispersal capability of Platygaster californica Ashmead (Platygasteridae), a parasitoid of the gall midge Rhopalomyia californica Felt (Cecidomyiidae).
3. Although foraging parasitoids meticulously searched plants in laboratory observations, none of the laboratory trials resulted in 100% parasitism, and the proportion of parasitism declined as midge egg density increased.
4. The field trials showed that the distribution of parasitism over distance from a central release point was hump-shaped, as predicted by a simple diffusion model. Mean parasitoid dispersal distance was 4.5 m, considerably farther than the 1.7 m mean midge dispersal found in previous work.
5. Although the parasitoid appears to search thoroughly for midge eggs and to disperse farther than the midge, the results of this study show how this host–parasitoid system may persist due to spatially variable incomplete parasitism.     

Non-linear transmission and simple models for bovine tuberculosis

1.  A new model is presented for a possum–tuberculosis (TB) system ( Trichosurus vulpecula – Mycobacterium bovis ) that is both realistic and parsimonious. The model includes a phenomenological treatment of heterogeneity of risk for susceptible hosts, similar to that used in insect host–parasitoid systems.
2.  Parameter values for the model reflect current knowledge and differ significantly from those in other recent models of this system. Associated with these structural and parametric changes are substantially different predictions for the dynamics and control of TB in possums.
3.  The model predictions include (i) only limited host suppression due to the disease (< 10%, cf. several earlier simple models for TB in both possums and badgers); (ii) asymptotically stable disease dynamics (cf. homogeneous-mixing models that predict either extremely weak stability such that disease fails to recover when host density is temporarily reduced, or oscillatory behaviour and potential elimination of disease following such a perturbation); (iii) TB that is harder to control than in the homogeneous-mixing model equivalents, in line with practical experience; and (iv) a threshold host density for disease elimination that differs substantially from the host equilibrium density in the presence of disease.
4.  Homogeneous-mixing models are unable to reproduce this behaviour, whatever parameter values are chosen. Heterogeneous-mixing models with non-linear transmission may therefore be worth consideration in other endemic wildlife disease systems, as is now commonplace for insect–parasitoid and insect–pathogen ones.     

Dispersal between host populations in field conditions: navigation rules in the parasitoid Venturia canescens

Abstract. 1. Dispersal is a life-history trait that can have great ecological and evolutionary consequences, however understanding of how insects disperse is limited.
2. Navigation rules of the solitary koinobiont parasitoid of the pyralid moth larvae Venturia canescens (Gravenhorst) were studied in conditions that it is likely to meet when dispersing between host populations and in the absence of cues related directly to the presence of hosts.
3. Mark–release–recapture experiments were conducted in a natural host-free habitat, and letting the animals disperse for different periods.
4. In the presence of vegetation, wasps seemed to disperse rapidly (1 h for an area of ≥ 1 ha) and capture rates were independent of both dispersal time and distance from the release point.
5. The navigation rules of V. canescens during dispersal between tree stands can be summarised as: move up- or down-wind, avoid or pass through open, sunny areas, and go for shady and dense vegetation.
6. The consequences of the navigation rules for host–parasitoid dynamics are discussed in relation to different spatial scales.     

Host plant finding in the specialised leaf beetle Cassida canaliculata: an analysis of small-scale movement behaviour

Abstract.  1. Host plant finding in walking herbivorous beetles is still poorly understood. Analysis of small-scale movement patterns under semi-natural conditions can be a useful tool to detect behavioural responses towards host plant cues.
2. In this study, the small-scale movement behaviour of the monophagous leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) was studied in a semi-natural arena ( r = 1 m). In three different settings, a host ( Salvia pratensis L., Lamiales: Lamiaceae), a non-host ( Rumex conglomeratus Murr., Caryophyllales: Polygonaceae), or no plant was presented in the centre of the arena.
3. The beetles showed no differences in the absolute movement variables, straightness and mean walking speed, between the three settings. However, the relative movement variables, mean distance to the centre and mean angular deviation from walking straight to the centre, were significantly smaller when a host plant was offered. Likewise, the angular deviation from walking straight to the centre tended to decline with decreasing distance from the centre. Finally, significantly more beetles were found on the host than on the non-host at the end of all the trials.
4. It is concluded that C. canaliculata is able to recognise its host plant from a distance. Whether olfactory or visual cues (or a combination of both) are used to find the host plant remains to be elucidated by further studies.     

Intrinsic, inter-specific competition between egg, egg–larval, and larval parasitoids of plusiine loopers

Abstract.  1. Intrinsic, inter-specific competition between parasitoid wasp species is a key factor in ecological community dynamics and is particularly important for application in biological control. Here three parasitoid wasp species with overlapping host ranges and differing life history strategies were chosen to examine parasitoid–parasitoid interactions: the egg parasitoid Trichogramma pretiosum, the egg–larval, polyembryonic parasitoid wasp Copidosoma floridanum, and the gregarious larval parasitoid Glyptapanteles pallipes , with the plusiine loopers Acanthoplusia agnata and Trichoplusia ni as hosts.
2.  Copidosoma floridanum has been shown to be an intrinsically superior competitor against larval parasitoids because of their production and increased investment in a soldier larval caste during development, but little is known of their interactions with egg parasitoid species. Trichogramma pretiosum completely dominated intrinsic competition with C. floridanum regardless of oviposition order or sex of the C. floridanum egg.
3. Competition between C. floridanum and G. pallipes , however, depended on the host stage at which parasitism occurred, the sex of the C. floridanum egg, and parasitoid development time. Copidosoma floridanum outcompeted G. pallipes overall, despite the fact that G. pallipes injects a polyDNA virus into the host.
4. The sex of the C. floridanum egg was a significant factor in its ability to shift caste ratios to produce more soldiers in response to G. pallipes competition.
5. Only developing female C. floridanum responded to competition with G. pallipes by increasing the ratio of soldier to reproductive larvae, and this happened only when multiparasitism occurred in the host's 1st and 2nd instar.     

Bottom-up cascading effects in a tritrophic system: interactions between plant quality and host-parasitoid immune responses

Abstract.  1. Little is known about underlying mechanisms by which plants indirectly affect parasitism success in hymenopteran endoparasitoids. The hypothesis that host-plant effects can challenge the innate immune system of an insect host was experimentally tested in this study using a model tritrophic, crucifer – lepidopteran [ Plutella xylostella (L.)] – parasitoid [ Cotesia plutellae (Kurdjumov)], system.
2. The effects of host-plant suitability on herbivore performance and parasitism were examined. The bottom-up effect of plant suitability on host-parasitoid immune responses was then evaluated using measures of cellular and humoral effectors.
3. Host-plant quality showed a significant effect on the encapsulation response of P. xylostella to first instar but not to second instar parasitoid larvae. Encapsulation was never sufficient to prevent parasitoid emergence.
4. Poor host-plant suitability suppressed phenoloxidase activity in the absence of the parasitoid. The suppressive effect of C. plutellae on phenoloxidase activity was much greater and no plant effects were detectable after insects had been parasitized.
5. Despite strong plant effects on parasitism, those on immune effectors of the host were transitory or overwhelmed by the effect of the parasitoid.
6. These results demonstrated that plant-mediated variation in parasitism success by C. plutellae were not as a result of plant nutritional status or other attributes affecting the immune function of P. xylostella , nor to host-plant effects on superparasitism.
7. In these experiments, P. xylostella was a fully permissive host to C. plutellae and host-plant-mediated effects on the innate immune response appeared to play no part in parasitoid survival within hosts.     

Bottom-up and top-down effects in a tritrophic system: the population dynamics of Plutella xylostella (L.)–Cotesia plutellae (Kurdjumov) on different host plants

Abstract.  1. The effects of host-plant resistance on the population dynamics of the Diamondback moth, Plutella xylostella L., and its solitary parasitoid, Cotesia plutellae (Kurdjumov), were studied in replicated time-series experiments.
2. Host-plant resistance did not affect the equilibrial abundance of the Diamondback moth, but it affected the dynamics of Diamondback moth populations.
3. The mean population size of Diamondback moth showed no significant difference between Brassica rapa (a susceptible host plant) and Brassica napus (a partially resistant host plant) either in the presence or absence of the parasitoid.
4. Time-series analysis suggests that the dynamics of Diamondback moth on B. rapa were underpinned by delayed density-dependent processes. In contrast, the dynamics of the moth on B. napus were influenced by a direct density-dependent process.
5. Although measures of short-term parasitism showed a significantly higher rate of parasitism by C. plutellae on Diamondback moth feeding on B. napus compared with B. rapa , this individual performance does not translate into differences in the population dynamics. Analysis shows no significant difference in the persistence time of the population-level interaction between the host and parasitoid on the two different host plants.     

Changing dynamics of the pine beauty moth (Panolis flammea) in Britain: the loss of enemy free space?

1 The pine beauty moth Panolis flammea has two main host plants in Britain: Pinus sylvestris (Scots pine), which is the ancestral food plant where the insect is never abundant enough to cause tree mortality, and Pinus contorta (lodgepole pine), an introduced host tree that has experienced periodic widespread tree mortality due to this pest.
2 We review the recent literature, published mostly after the year 2000, regarding the impact of natural enemies on the population dynamics of P. flammea in Britain.
3 The natural enemies of P. flammea are more diverse and abundant in Scots pine habitat than in lodgepole pine habitat and some of them show differential selection for P. flammea larvae in Scots pine habitat over those located in lodgepole pine habitat.
4 It is concluded that the difference in the population dynamics of this insect in the two different habitats was probably the result of the P. flammea finding enemy-free space in lodgepole pine habitat.
5 Recent evidence on the diversity and impact of natural enemies on lodgepole pine has demonstrated that they currently have a much more significant impact on this pest than they did in the 1970s and 1980s, when outbreaks were frequent.